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J. Grin. SOC. (Bot.), 57, 369, p. 1 With 67 figures and 51 maps Printed i n Great Britain Berberis and Mahonia A taxonomic revision 1 By LESLIE WALTER ALLAM AHRENDT, M.A. (Cantab.), M.A., D.Phi1. (Oxon.), F.L.S. Sometime Scholar of Peterhouse, University of Cambridge, Wrangler in the Mathenaatical Trips; Merton College, University of Oxford ‘The time is fast coming when gardens of any pretensions to beauty will be judged by their collections of Berberia, for there is not any other class of evergreen shrub which affords so many points for interesting observation.’ SHIRLEY HIBBERD, a collector of Berberis, writing under this generic heading on both Berberia and Mahonia, in The Pbrd World and Garden Guide, Vol. v, p. 163 (1862). PREFACE The results presented in the following pages are the accumulation of studies covering the years 1931 to 1956. During this period, I have described a number of new species; and certain others, hitherto undescribed, are now included. The very great majority of other species have been re-examined, and described afresh. This has generally been done from type specimens, with the inclusion of many characters omitted from the authors’ original descriptions. Amplifkation has included numerical precision. Thus, instead of ‘finely, closely, or coarsely, toothed leaves’ (which phrases leave an undue burden of interpretation on the reader), the range of number and sue of spinules and serrations, and their distance apart are given. This revision provides also, for most species, detailed analysis of flower structure, a character hitherto largely overlooked. In a considerable number of species, living material has been studied over a number of years, to augment the knowledge provided by the dried specimens; but, in all such cases, the greatest caution has been exercised to confirm that the plants so used do represent the true species, before any information gathered from them has been incor- porated. Much of the living material of the many recently introduced species is that which has been grown from seed from the collector’s type number, and corresponds completely with its dried counterpart. The very sound division of Berberis into two groups by C. K. Schneider in 1905, and of Mahonia into two sections by Fedde in 1901, remains fundamental. Schneider further subdivided Berberis into sections, the general structure of which remains, although he himself at times revised their content. A further definition of these is now provided. Although the general character of these sections has become increasingly clear, and their use justified, the boundary between them is not always obvious. Very many keys have been provided, and these include, not only those to sections, but additional ones covering two or more related sections. This is of real value for determination when there is uncertainty, at first sight, in which of such related sections a given species belongs. The large Berberis section Wallichianae was more fully analysed into subsections by Schneider in a paper published in 1942; and, independently, and just previously, by myself in 1941. The broad separation of the Mahonias into four sections has remained for u t y years without further analysis, although much valuable work was done by Takeda in separating confused species. Sufficient information is now available to provide a further analysis 1 JOURN. LINN. S0C.-BOTANY, VOL. LVII

Berberis and Mahonia : A taxonomic revision

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Page 1: Berberis and Mahonia : A taxonomic revision

J . Grin. SOC. (Bot.), 57, 369, p . 1

With 67 figures and 51 maps Printed in Great Britain

Berberis and Mahonia A taxonomic revision

1

By LESLIE WALTER ALLAM AHRENDT, M.A. (Cantab.), M.A., D.Phi1. (Oxon.), F.L.S.

Sometime Scholar of Peterhouse, University of Cambridge, Wrangler in the Mathenaatical T r i p s ; Merton College, University of Oxford

‘The time is fast coming when gardens of any pretensions to beauty will be judged by their collections of Berberia, for there is not any other class of evergreen shrub which affords so many points for interesting observation.’ SHIRLEY HIBBERD, a collector of Berberis, writing under this generic heading on both Berberia and Mahonia, in The P b r d World and Garden Guide, Vol. v, p. 163 (1862).

PREFACE

The results presented in the following pages are the accumulation of studies covering the years 1931 to 1956. During this period, I have described a number of new species; and certain others, hitherto undescribed, are now included.

The very great majority of other species have been re-examined, and described afresh. This has generally been done from type specimens, with the inclusion of many characters omitted from the authors’ original descriptions.

Amplifkation has included numerical precision. Thus, instead of ‘finely, closely, or coarsely, toothed leaves’ (which phrases leave an undue burden of interpretation on the reader), the range of number and sue of spinules and serrations, and their distance apart are given.

This revision provides also, for most species, detailed analysis of flower structure, a character hitherto largely overlooked.

In a considerable number of species, living material has been studied over a number of years, to augment the knowledge provided by the dried specimens; but, in all such cases, the greatest caution has been exercised to confirm that the plants so used do represent the true species, before any information gathered from them has been incor- porated. Much of the living material of the many recently introduced species is that which has been grown from seed from the collector’s type number, and corresponds completely with its dried counterpart.

The very sound division of Berberis into two groups by C. K. Schneider in 1905, and of Mahonia into two sections by Fedde in 1901, remains fundamental. Schneider further subdivided Berberis into sections, the general structure of which remains, although he himself at times revised their content. A further definition of these is now provided. Although the general character of these sections has become increasingly clear, and their use justified, the boundary between them is not always obvious. Very many keys have been provided, and these include, not only those to sections, but additional ones covering two or more related sections. This is of real value for determination when there is uncertainty, a t first sight, in which of such related sections a given species belongs.

The large Berberis section Wallichianae was more fully analysed into subsections by Schneider in a paper published in 1942; and, independently, and just previously, by myself in 1941.

The broad separation of the Mahonias into four sections has remained for u t y years without further analysis, although much valuable work was done by Takeda in separating confused species. Sufficient information is now available to provide a further analysis

1 JOURN. LINN. S0C.-BOTANY, VOL. LVII

Page 2: Berberis and Mahonia : A taxonomic revision

2 L. W. A. AHRENDT of this genus into two main groups, corresponding in nature to those of Schneider for Berberis, and many subsections.

I must record sincere acknowledgement of the facilitiea aEorded to me for the thorough examination of dried specimens at the Royal Botanic Gardens at Kew and the Royal Botanic Garden, Edinburgh, and the British Museum (Natural History), and the depart- ment of Botany of the University of Oxford. And I must express thanks for the generous supply of living material afforded me from Kew, Edinburgh, Wisley, and from numerous private gardens, to which reference is made in the text in connection with cultivated material derived from them. However, the greater part of the cultivated material examined came from my own collection of typical plants grown from 1934 to 1945, on chalky soil, in the garden of my mother and father a t Watlington in Oxfordshire. All cultivated specimens cited without source of origin came from this source.

Duplicates of this living collection are to be found in the University Botanic Garden at Oxford, most of them having been transferred there from 1942 to 1945; and many in the Botanic Garden of the University of St Andrews, having been sent there in 1956. During the same period a large dried collection from the same plants was deposited in the Fielding Herbarium at Oxford. A duplicate of this dried cultivated collection was retained by myself for some years, but transferred to the British Museum Herbarium in November 1953.

References to dried and cultivated material

Nearly all the dried material cited comes from the herbaria of Kew, the British Museum, Edinburgh, or Oxford, which have been designated respectively by the abbreviations, K, BM, E, 0. The reference to cultivation is almost always to Britain, and provides information as to whether the species is there grown, and, in such cases, when and where it has been found and examined. In the majority of cases, where no place is named, the plants were at Watlingtonl, Oxfordshire; in other cases, the place is mentioned as well as the time. Where it is stated that a species is not in cultivation, the indication is that it has not been found in cultivation in Britain, after a quarter of a century’s persistent search, with the reliable inference that it is not growing there. In a few exceptional caaes it has been necessary to refer to cultivation elsewhere; these concern hybrids whose origin was in other countries.

PART I INTRODUCTION TO THE GENERA BERBERIS ANDdfALTONIA

DIAGNOSTIC CHARACTERS OF BERBERIS AND MAEONIA I. Stem characters (Fig. 1)

In HahOnia generally speaking, and in contrast to Berber&?, the character of the stem is so indefinite aa to provide but few data of diagnostic value. The stems are generally scaly, and green at first, though sometimes slightly purplish, ripening to a brownish grey. Smoother, dark red stems, such as are familiar in Berber&?, appear in the few HahniaC species of the unusual section Horridae; this being one of several characters which plaoe these Mahonias neaxer to Berber& than the others.

In Berberis, the young shoots of the current year are generally greenish, often flushed with a slight reddish tinge on the side facing the sun. Sometimes, after a few weeks’ development, and before first year maturity, the stems w u m e a vivid red colour, which changes fairly quickly to the more enduring colour of the mature stems. It is this latter which has the most marked diagnostic value. The occasional, and transitory, distinctive colour of the young shoots of certain species is mainly of ornamental garden

After 1945, Cultivated specimens without mrd of place, were from Broughton, North Oxfordshire.

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Berberis and Mahonia 3

value: in the case of cultivated plants, it may suggest or confirm an identity. It is not of more general diagnostic value since it is apparent only for a short period, and may therefore not be present a t the time of examining a living plant, or gathering material for the herbarium specimen. When this transitory colouring of the shoots is present in dried material, care must be taken not to assume that it represents the colour ofthe mature stems, from which it often differs, and about which accurate observation is essential. Thus, species with bright red shoots will sometimes have dark red mature stems, e.g. B. sargentiann. It may be quite otherwise, however, and bright red shoots may become pale yellow when mature, as in B. bergmunniae, B. veitchii, B . spragwi and B. forrestii. Then again, as B. pruinosa, purple shoots may become pale yellow.

In Berberis, then, the mature first-year stems are important for observation. These may either be: (a ) pale-yellow, yellowish brown, or yellowish grey; or ( b ) dark-reddish brown, dark red, dark purple, or almost black. So far, 163 Berberis species have been recorded with dark stems, and 227 with pale stems.

Old Berberis stems of the second year, and after, tend to become an ashen grey colour. Care must be taken in determining herbarium specimens which present only this old wood; and also with some living plants, such as big transplanted bushes, which take a few years to recover, and present no new or first-year stem growth. In such cases, the observation of the old grey wood must not be taken to indicate that it is from a pale- stemmed species.

A very large number of Berberis species exhibit verruculose characters on the shoots and mature stems. These vary from very dense and conspicuous to very sparse and in- conspicuous, and, when sparse or inconspicuous, they may be found only on certain parts of the stem. In many species vemculose stems are unknown. This character has diagnostic value for discrimination between species with densely verruculose stems, and those with smooth stems. But, where the character is manifested sparsely and variably it should be disregarded. Vemculose stems are seldom worth mentioning in taxonomic keys, but are worth recording for their confirmatory diagnostic value.

In some species, the young shoots and first-year stems are hairy, the hairiness varying from thinly puberulous to a dense pubescence. However, the degree of pubescence is not necessarily the same on young shoots and mature stems in any one species, as shown by the following examples: (a) shoots thinly and persistently puberulous, e.g. B. mitij'olia; (b) shoots thinly pubemlous when young, but glabrous when mature, e.g. B. w r o s e p d u ; ( c ) shoots densely and persistently pubescent, e.g. B. verruculosa, and B. dam%; (d) shoots densely pubescent when young, but glabrous, or perhaps slightly pubemlous, when mature, e.g. B. brachypoda; ( e ) shoots lightly puberulous when young but densely pubescent when mature, e.g. B. buxifolia.

The stems of 79 species are recorded as a t least slightly pubescent, and 321 as com- pletely glabrous.

In Berberis spp. the stems are either terete, angled, or sulcate (Fig 1, a-d), according as their cross-section presents a circle (or approximately circular smooth outline), a convex polygon (whose corners will be somewhat softened), or a re-entrant figure, so rendered by grooves running along the stem. Occasionally, the last or sulcate, character is so marked aa to leave a sharp thin, demarcation between the grooves, e.g. in B. sublevis, and B. submulialata. The outline of the cross-section is not necessarily the same in young and mature stems of any one species. In young stems the outline varies so much that it is of no diagnostic value, and indeed it may even be changed when the specimens are dried. The outline is also variable on old grey stems. On the other hand, it is a quite valuable diagnostic character for mature stems and is worth using in keys. Terete and very sulcate stems stand out in marked contrast to one another, but the cross-section is of less diagnostic value when the stems are slightly or very Gnely sulcate. Forty-two species have been recorded with terete or subterete stems, 158 species when they are angled or sulcate and 49 when they are markedly sulcate.

1-2

Page 4: Berberis and Mahonia : A taxonomic revision

4 L. W. A. AHRENDT

11. The spines

Spines never occur in Mahonia, so that all remarks under this h e h g apply only to B e r b h . Two plants, described by De Cmidolle as being Mahonias with spines on the stems, namely his Malumk caragawfolia and M . tragacanthokh, appear to belong to the genus Caragana.

(1) Spines absent. Spines are rarely absent from the Asiatic species, except for the four members of

Sect. Wallichiame, Subsect. Insignes. In all, 17 species of Berberk are recorded a having genuinely spineless stems, the remaining 13 being South American.

(2) Spine &-nt weak. Spine development may be described as weak where they are often solitary, though

sometimes threefold, slender, and only 1-9(-12) mm. long. Sometimes, the spines may be absent from some parts of the stems. Weak spines am characteristic of certain species in the Asiatic sections, amongst which they are more predominast. in the ‘ S i n e m and the Tschonoskyana.8. The weak spined character is still more predominant in the species from South America. One hundred and thirty-six species with weak spines have been recorded of which about 75 come from South America.

(3) standard spine aelhdqwment. When spines are only occasionally solitary, mostly threefold, and more or less

moderately stout, i.e. up to 1-1.5 mm. broad at base, and (1-)1-5-2-5 cm. long they may be described as exhibiting the standard development. This occurs in most (195) of the Asiatic species. This type of spine development is 80 rare in the South American group that it may be counted aa exceptionally strong spine development for that region, appearing only in some 15 species, i.e. those of the Sect. Uicifoliae, Subsect. Flexuosm.

(4) spine Avelqnnent strong.

Spine development may be described as strong in species that are at once conspicuous in having exceptiody developed stout large spines, i.e. 2-5 mm. broad at the base, and 3-6 om. long. This applies in certain species of the Sect. W a l l i c W , e.g. B. soulieana, B. &inmuntha, and generally, though not invariably, marks the Sect. Integerrimae, e.g. B, ja&na. Strongly developed spines are recorded in 34 species of which only seven come from South America.

(5) Spines manifold> palmate OT foliaceous There are certain species in the Asiatic group, with spines having a 5 to 11-fold

character, e.g. B. sibirica, B. tsarica, B. erythroclada, of the Sect. Angulosae. This diversion from the general threefold spines is more marked in the South American Sect. Actinacanthae. In its Subsect. Euactinacanthae, the spines, occasionally threefold, but generally manifold, are leaf green, and broadly flattened (palmate). In the Subsect. Congestiflorae, the final stage of the passage back along the path of leaf metamorphosis into spine is found. Here, the arms of the spinea are not separate, but joined so that the spine is of a more genuinely foliaceous nature; when this is so it differs from the leaf proper in that it is foreshortened with a cordate base, partially, or almost completely, encircling the stem, and margined with spinous teeth like the leaves themselves. The Subsect. Congestifloraa presents the South American species possessing this character,

1 Note. The abbreviations ‘3-fld.’, ‘3-5-fld.’ etc. used, in reference to the spine.s mean ‘threefold’ (i.e. ‘tri6d’ or ‘tripartite’ofmost authors), ‘3-5-fold’ etc. The same abbreviations used i n reference to the injEoreaence mean ‘three-flowered’ or ‘3-5-flowered’ etc.

Page 5: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 5

which is shared by one only from Asia, namely B. koreanu. It will be seen that only 24 Berberis spccies differ from the more customary arrangements of spines with which we are familiar.

Tracing the evolutionary development of spine leaves, we shall have to consider the types described above in the following order: spineless ( 5 I), foliaceous ( 5 5 ) , palmate ( 5 5) , manifold ( $ 5 ) , weak ( 4 2), standard ( 5 3), strong (0 4).

(6) Spine c o h r A further complexity is introduced by the colour of the spines on the mature h t - y e a r

stems. If the latter are pale (yellow, yellow-brown, or grey), the spines are invariably more or less concolorous. If the stems are dark (red-brown, dark red, or dark purple), the spines may be either: (a) concolorous, as in B . jamesiana; or, ( b ) discolorous, and yellowish orange, or even sometimes most conspicuously different and pale yellow, e.g. B. gyalaicu. This feature is immediately obvious, but it should be relied on only aa a confirmatory diagnostic character.

( 7 ) Spine section Spines may be either (a) terete (more or less circular in cross-section), (b) flattened

below, or (c) sulcate below. The cross-section of the spine is not necessarily the same aa that of the stem, and, while generally recorded, no critical use has been made of it.

( 8 ) Disposition of spines Spines are set mutually, sometimes at right angles, sometimes a t acute angles. Some-

times the laterals of a tri6d set are as long aa, rarely longer than, and generally shorter than, the central one. These characters are more or less constant in a species, but, again, have not been used for separating species.

(9) Spine development and habitat The spineless or weak-spined species are generally natives of moist regions, e.g.

B. insignis from Sikkim. Those with the very strong spines on the other hand, are from dry regions, e.g. B. smlieana from Kansu and Shensi, and the Sect. Integerrimae from Persia and Central Asia. The extent to which spines are developed, however remains constant in cultivation. A variety of B . insignis has been cultivated for over 100 years.

111. The leaves (Figs. 2-5 and 14) (1) Pinnate and simple

Whether a species possesses simple or pinnate leaves is the only discrimination that is invariably reliable for the separation of Berberis from Mahonia. I n Berberis the leaves, are always simple; in Mahonia they are always imparipinnate.

In Mahonia, the number of leaflets comprising the pinnate leaf is an easily recognizable diagnostic character. Although, as just stated, typical pinnate leaves are absent from Berberis, it must be remembered that in most species of Berberis the leaves are arranged in fascicles or whorls, each of which is attached at a node and protected by the spines. The whorl of Berberis leaves may be regarded as a pinnate leaf in which the leaflet inter- nodes of the rachis have been eliminated, thus bringing leaflets, now regarded as simple leaves, into a cluster a t the stem node. Similarly, further metamorphosis of these simple leaves has been regarded as producing, first the foliaceous spine, then the palmate spines, then the manifold, and finally the trifid group of spines, sometimes called spine leaves. Thus, the number of leaves, or the number of leaves and spines taken together, in each whorl a t a Berberis stem node corresponds to the number of leaflets in the pinnate leaf of a M a h i a . However. the number of leaves and spines that occur a t a node in

Page 6: Berberis and Mahonia : A taxonomic revision

6 L. W. A. AHRENDT Berbeods has seldom been recorded. In a few species, e.g. B. i&gnis, the leaves are actually solitary, but in most species there are quite a number of leaflets in each whorl, but lack of information precludes further analysis on this point. The genus Berberis presents such a great number of feature% for observation, and consequently so many possible combinations of these in its very numerous species and varieties, that botanists have inevitably ignored some of them. It is desirable, as a scientific principle that such data as these should be recorded, but it is doubtful whether, with this done, much use would be made of this particular character for diagnostic purposes. The general aspect of this matter is more important. Some species have been founded upon a selection taken from the total of possible characters, with complete lack of reference to others. Conversely, other species have been founded on the very characters that have been ignored for species in the first group. No rational concept of the genus can be gained on such a variable foundation. It is for this reaaon that I have found it necessary to collect, review and here present, all the charahrs to be considered, and their possible variations. The number of variable featurea that are easy to observe is less in MaOnia than in

Berber&; and consequently the number of possible combinations is much less. So, it is not s u r p r i s i i that 497 Berberis species are recorded and only 110 of M b k . The number of ledeta in a M a h k leaf is the only character in which this genus provides an extended range of diversity not found (or a t any rate, used) in Ber6eri.s. Under this heading four ranges of leaf character may be observed.

(a) Species with trifoliate leaves consisting of only one pair of leaflets and a terminal leaflet exist. This rare character, occurs in only four species, three Asiatic and onc American.

(b) Species with 2-3 (4) pairs of leaflets. "his type is rare in Asia, where it occurs in only four out of 66 species; but it becomes just predominant in the American Mahonias where it occurs in some 19 out of 43 species.

(c) Species with 5-9 (-ll), average 5-9, pairs of leafleta. This type occurs in 46 out of 66 Asiatic, and in 15 out of 43 American species. Since leaves of this type occur in 61 species out of 109, this type is the most common for the whole genus.

(d) Species with (9-) 10-20, average well over 10, pairs of leaflets. This type is re- stricted to nine Asiatic species.

(2) Leaf persistence All Mahonias are completely evergreen. Comment under this heading is, therefore,

confined to Berberis, for which the distinction between the evergreen and the deciduous species has been used as a primary taxonomic character. Undoubtedly there is a marked gulf between, on the one hand, (a) the 91 evergreen Wallichianae of Asia, and the vast evergreen majority of American Berberis spp. (probably about 170 out of 178) ; and, on the other hand, (b) the many deciduous species which comprise 186 out of 319 from Eurasia. If these species alone were to be considered, the demarcation on these lines would be completely clear. But some of the few deciduous species from South America, such as B. mrmtana, are obviously related to B. buxiyolia. More confusing still is the existence of some 42 species of the Asiatic sections Asiaticae and Tinctoriae. There is some uncertainty as to whether these should be classified as evergreen or deciduous since they have been variously described. Three pointa must be emphasized. (a) Herbarium specimens provide little or no evidence of whether the leaves are evergreen or deciduous, and collectors rarely say which they are. This information would, however be, much more valuable to the student than many facts recorded in field notes. (b) Cultivated plants of many species of the sections Asiaticae and Tinctoriae, e.g. B. lyciurn, B. parkeriana, B. fl0l.abund.a and B. umbeuata remain evergreen in rare thoroughly mild winters, but are defoliated in the more severe English winters; thus, it would be easy, when observed in cultivation, to regard them as deciduous when they are evergreen in their native habitats. (c) the 42 species of Asiaticae and Tinctoriae lie to some extent botanically

Page 7: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 7

between the 91 genuinely evergreen Wallichianae, and the 178 Eurasiatic species which are, without doubt, deciduous.

Hitherto, I have regarded the Asiaticae and Tinctoriae as truly evergreen species by nature. On this basis, it is easier to provide a simple key to the major divisions of the genus Berberis, as I attempted to do in 1941 (J. Bot., Limd. (Suppl.)). Later in this monograph, I have provided a key without making this assumption (Key 1); for, if there is uncertainty about this matter, there will be confusion of the Tinctoriae with the Vulgares and Angulosae.

Reference must also be made to a few other species which also cloud this issue: these belong to deciduous sections, but are themselves distinctly sub-evergreen, as they retain most of their leaves throughout the winter. I refer to B. wilsonae and its varieties (pami- folia, subcaulialutu, stupjiana) among the deciduous Polyanthae; and B. concinna, B. tsangpoensis and B. kumaonensis amongst the deciduous Angulosae. All these minor exceptions, while they do not destroy but rather underline the main demarcations, add considerably to the difficulty of constructing keys.

(3) Texture of evergreen leaves Evergreen leaves are more or less coriaceous. Sometimes they are markedly thick and

rigid, and a hypoderm is then present, which is sometimes easy to discern, but sometimes rather indistinct. Alternatively, they may be without hypoderm, when they are generally noticeably thinner, and more flexible or slightly stiff. In M a h i a , all the Asiatic species and one abnormal American one-M. nemsa-have more or less thick leaves with hypoderm and form a natural group, while nearly all the remaining American species have thinner leaves and lack hypoderm. In Berberis this character occurs in a number of American species, but otherwise only in some species of the Sect. Wallichianae. Schneider, in 1905, used this character for diagnostic purposes; and in 1913 and 1917, divided the Wallichianae, in his key, into three parts characterized respectively by the hypoderm being distinct, indistinct or absent. Since then, the presence of hypoderm has c e w d to be used as a diagnostic character, partly owing to the confusion introduced by the existence of the second group including species such as B. sargentiana, and partly because the determination of the hypoderm is too intricate an anatomical process to be accept- able to one examining large numbers of specimens for identification. However, there is a clear and obvious difference between the thick and rigid leaves of B. pruinosa and B. soulieam, on the one hand and the thin, flexible ones of B. mnipurana and B. sublevis on the other, and this is still a valuable diagnostic character.

In certain South American species, i.e. those of Sect. Actinacanthae, the leaves, with or without hypoderm, have a peculiar stiff, dry, chartaceous texture.

In Mahonia, there are 74 species with thick leaves with hypoderm. These are the 66 comprising the group of Asiatic Mahonias together with the American M . nervosa, and five other American species (Horridae). The species with thin leaves lacking hypoderm number only 42 and are all American.

(4) Basal lea$ets of Mahonia In some 67 species, the basal pair of leaflets are attached only 3-10 (-15) mm. above

the base of the rhachis. They are thus so close to the base that at a casual glance the leaves appear sessile. In these leaves which may, conveniently, be termed subsessile, the petiole is only one-twentieth or less of the length of the leaf. This character occurs in 57 out of 66 Asiatic, and in 10 out of 43 American species. The remaining minority consisting of only 10 Asiatic, and 33 American species, have petiolate leaves. In these leaves there is either (a) a short, but distinct, petiole of 2 4 cm., as in most of the American species, and the Asiatic M . gracilipes and M . hancockiana, or (b) a conspicuous petiole, one-third to one-half as long as the leaf, e.g. M . fortunei and three related Asiatic species.

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8 L. W. A. AHRENDT Thus, the Asiatic species have leaves which are predominantly subsessile, rarely

markedly and very rarely shortly, petiolate; while the American species have leaves which are predominantly shortly petiolate, with, however, a considerable res iduk of species with leaves that are subsessile.

In the species where the basal leaflets are virtually a t the base of the rachis, these are generally very much reatricted in size, being conspicuously shorter, and proportionately broader than the others. They are mostly, then, as broad as long, and angular. This character predominates in those Asiatic species with numerous leaflets referred to under 5 2 (c) and (d) on p. 6, where it is recorded in 42 out of 56. In the remaining Asiatic species with fewer leaflets, and most of the American ones, i.e. those dealt with under §2(a) and (b) onp. 6, there is a general, though not invariable, association of some degree of petiole with basal leaflets either subequal to the others, or only smaller in such degree aa is consistent with their general gradation of size :

(5) LeaJet internodes in Mahonia In the large number of Asiatic species for which there are accurate records, the inter-

nodes between the leaflets almost invariably decrease in length, sometimes slightly, sometimea conspicuously, in passing from the base of the rachis to the apex. Only two exceptions, M. jlavdu, and M. bealei, appear to have been recorded, in which the leaflets are roughly equidistant. M . fargmii is abnormal because the second pair of leaflets is exceptionally clo& to the basal pair, so that the decreasing series of internodes does not begin until after the second pair. This feature, of decreasing leaflet internodes, is less marked in the American Mahonias, because the fewer pairs of leaflets which are generally present scarcely provide a suflicient range for the character to become apparent.

(6) Size of leaflets in Mahonia

It is interesting to observe 10 different arrangements of gradation of size in the leaflets of the Asiatic Mahonias:

(i) Leaflets all more or leas equal, the difference in size being slight-seven species, i.e. M. weitchimum, M. lmgibracteata, M . b r - k , M . fortunei, M . pyomphylla, M . gracilipes and M . sumatrensis.

(ii) Length and breadth of the leaflets increasing steadily from the base to the apex of the rachis, e.g. M . h y p l e w , M . cleciyviens, M. sikkimmb, M. boredis and M.$uvdu.

In (i) and (ii) the shape of the leaflets from the base to the apex of the leaf does not appreciably vary.

(iii) Length and breadth of the leaflets gradually decreasing from the base to the apex of the leaf. Here, also, the shape of the leaflets is similar.

(iv) Length and breadth of the leaflets increasing from the base of the leaf to the mid-point, and then decreasing in pwing from there to the apex+.g. M . &micudia, M. leveilleum, M . fargesii and M . t ihhiensis . Here, also, some similaxity of leaflet shape is preserved, as length and breadth change together.

(v) Length of leaflet increasing but breadth fairly constant from the base to mid- point of leaf; then, from mid-point to apex, both length and breadth decreasing-e.g. M. siamemis and M. Bimonsii. An element of changing shape is here introduced into the lower leaflets.

(vi) Length of leaflet increasing from base to the mid-point of the rachis, but from there to the apex decreasing; leaflet breadth decreasing gradually, the changing shape on the lower part being thus further emphasized+.g. M. lesehenaultii and M. lomariifolia.

(vii) Leaflet length increasing to the middle of the rachis, then decreming ; breadth constant; the change of shape now extends the whole length, but with a reversal midway which restores the leaflets near the apex to the shape of the b a d 0nes-e.g. M. nupau- lensis and M. schochii.

Page 9: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 9

(viii) Length of leaflets fairly constant throughout the length of the leaves, but breadth steadily decreasing from base to apex4.g. M . manipurensis.

(ix) Length of leaflets increasing from the base to the apex of the leaves, but breadth c0nstant-e.g. M . philippinemis, M . sheridaniana and M . wnfusa. With (viii) and (ix), there is a steady change in leaflet shape in passing along the leaf rachis. (x) Length of leaflets increasing, and breadth decreasing from the base to the apex of

the leaf, so that the change in the shape of the leaflets is emphasized and is very marked- e.g. M. conferta, M. japonim and M . bealei.

Gradations in the size and shape of leafleets is scarcely noticeable in the American Mahoniaa, but it adds great diversity to the appearance of the leaf in the Asiatic ones, for which it should be accurately recorded, though it has often been omitted by authors who content themselves with giving the limiting leaflet measurements.

(7 ) Base of Berberis leaves and Mahonia leajlets (a) Mahonia. In many of the Asiatic Mahonias, the base of the leaflets is often sub-

cordate (slightly overlapping the rachis); or truncate (lying along the rachis); or so broadly rounded or broadly cuneate as to be almost subtruncate. In others, and many of the American species, the base is more narrowly cuneate, so that there is little more than a point of contact between the leaflet base and the rachis. There is only one Asiatic species (M. bodinieri) recorded with a very short stalk (petiolule) to the leaflets; it is only 2 4 mm. long; but this feature appears in a small minority of American Mahonias.

(b) Evergreen species of Berberis. The base of the leaf is generally sessile, with, on occasion, a short decurrent petiole, up to 3 4 mm. long. In a few South American species there is a longer, true petiole with conspicuous articulation.

(c ) Deciduous species of Berberis. These mostly have sessile or very shortly petiolate leaves; a decurrent petiole, up to 1-2 cm. long is present in certain northern species (Sects. Dasystachyae and Tschonoskyanae).

(8) Margins of Berberis leaves and Mahonia leaflets

The margins of Berberis leaves and of Mahonia leaflets are often spinose-toothed, and may be classified in three groups.

(a) Margins quite entire, or at least typically so. In Mahonia, this occurs in only five out of 110 species; of these one, M . shenii, is Asiatic, the other four being American species of the Sect. Paniculatae.

In evergreen species of Berberis entire leaves (though not quite constant) appear in only seven out of 91 Wallichianae species, but generally characterize 25 out of 42 Asiaticae and Tinctoriae. About 65 out of 178 American species are entire-leaved. In deciduous species of Berberis, some 75 out of 186 species have entire leaves.

(b) Margins sparsely or remotely toothed; i.e. the spinose teeth numbering 1-8 (-18) according to the size of the leaf, and being set 3-6 (-10) mm. apart.

In Mahonia, (b) is the commonest type, occurring in about 94 species out of 112, i.e. 58 out of 66 Asiatic and 36 out of 43 American. In evergreen species of Berberis, (b) also predominates. appearing in some 78 out of 91 Wallichianae species, and 79 out of 178 American. In deciduous species of Berberis, (b), which occurs in 77 species is hardly more common than (a ) .

(c) Margins densely spinose; i.e. the teeth numbering (lo-) 15-50 (-150) according to size of leaf, and being only 1-2 mm. (-2.5 mm.) apart. This is a minority character appearing in only six Wallichianae Asiatic evergreens, 10 South American, and some 20 Eurasiatic deciduous Berberis, and in only two (Asiatic) Mahonias.

Summing up we see that entire margins occur in 172 species, confined almost ex- clusively to deciduous Asiatic and evergreen South American species of Berberis ;

Page 10: Berberis and Mahonia : A taxonomic revision

10 L. W. A. AERENDT remotely toothed margins occur in 247 species, predominating heady in evergreens (both M a h i a and Berberis), but they are to be seen in only some two-fXths of the deciduous species of Berberk. Densely spinose margins occur in only 36 species of Berberis, just over half being deciduous (Vdqarea and relahd sections).

For diagnostic purposes, classification in these three groups is necessary. Vague descriptions, aa given for some older species in which mmgind serration are cited merely as remote or close, are not reliable, depending upon individual judgement. For the accurate description of leaf margins, whether for determination of specimens, or descrip- tion of species there should be metrically cited, (a) range of number of teeth; (b) range of distances between the teeth; (c) length (or depth) of actual serration, which varies from 0.25 to 5-8 mm. rarely more; (d) length of apinde surmounting tooth, and varying from 0-25 to 2-3 mm.

The length of spinule does not correspond to that of serration. In different species varying lengths of spinule surmount varying depths of tooth; accordingly, apinules or serrations may be the more conspicuous, or they may be equally so.

(9) Apex of Berberis leaves and, Mahonia leajkta The apices of the leaves or leaflets may be classified as follows: (a) Obtuse or raunded. This is common in South American and deciduous Asiatic

Berberis; in M a h i a only in six American species, e.g. M. repem. (b) Acute. General in the Asiatic evergreen Berberis. In Mahimia, on account of the

prevailing coarser toothing, the acute apex is generally associated with a somewhat inconspicuous cusp; this cusp continues approximately the outline of the leaf margin, is proportionately broad, 1 : 16-26 (-3) and fairly short, 0-61 .5 (-2.5) cm. Twenty- three Asiatic Mahonias with this character are recorded (e.g. M . lomariifolia and M. healed). The acute apes is general in the American Mahonias.

(c) Acuminate. This appears in moderate degree in some of the evergreen Wallichianae, e.g. B. muminah, and in a very few deciduous species of Berbehs, such as B. salicuria and B. dielsiana. In Mahonia, this character is conspicuous in eight species from Asia, e.g. M . w e h h k m . In these species the cusp is so slender that it abruptly changes the inclination of the leaf outline above the final pair of teeth; the cusp itaelf (20-25 x 4- 8 mm.) is longer and proportionately narrower (1 :3-5) than in (a) . (10) Venation of Berberis leaves and Mahonia h @ t s

The three following classes may be recognized : (a) enervate or subenervate : veins if present very few, and faint; (b) openly veined: the veins being distinct, more or leas numerous, with the lateral veins often branched into veinlets, but with these not suffi- ciently developed and joined to form a network; (c) reticulate: the branching of the veinlets extending and joining to form an open or close network.

Enervate, or subenervate leaves are recorded in some 140 out of 497 Berberis spp.: i.e. in 85 out of some 311 evergreen, and 15 out of 186 deciduous species. In M a h i a , the same character occurs, in conjunction with the thick nature and the presence of a hypoderm, in the 67 species of the Asiatic Longibracteatae, and the 7 of the American Horridae-i.e. in 74 out of 109 species. Although distinctly a minority character in Berber&, it is dominant in M a h i a .

Loosely veined leaves occur in 67 out of 311 evergreen, and 68 out of 186 deciduous species of Berber+ i.e. in 135 out of a total of 497 species. This form of venation is almost unknown in Mahonia, but in some of the Asiatic forms the few veins that are present are sometimes sufficiently clear for them to be considered under this heading. Occasionally the reticulation in the American Mahonias is a little indistinct, but a clear example of a leaf which is neither subenervate, nor subreticulate, but possessing a well- developed loosely branched venation does not often occur in M a h i a .

Page 11: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 11

Reticulate leaves occur in 94 evergreen, and 98 deciduous species of Berberis, the latter comprising, among others, most of the Sects. Integerrimae and Polyanthae where this character predominates; 192 in all. Reticulate leaves occur in a great majority of Mahonia spp. namely 37, of the American species. Thus reticulate leaves are the com- monest, although they are not predominant in Berberis, in which genus distinctly veined leaves are clearly the most frequent, while in Mahonia the more or less enervate character of the Asiatic species is dominant.

(11) Undersurface of leaves (i) Pruinose characters. (a) Epruinose, green ; concolorous, or generally paler-in 149

evergreen and 87 deciduous (i.e. 236 out of 497) species of Berberis; and in an over- whelming majority of Mahonias (100). (b) Slightly pruinose and greyish-in 133 ever- green, and 49 deciduous species of Berberis and in 8 Mahonias. (c) Heavily pruinose and conspicuously white. This occurs in only 23 evergreen and eight deciduous or semi- evergreen species of Berberis, and one species of Mahonia.

(ii) Papillose characters The undersurface of Berberis leaves may be either papillose or epapillose, the former

character frequently, but not invariably, concurring with a pruinose condition, and the latter is similarly related to the epruinose.

(12) Revolute leaves (Fig. 14) In a minority of some 21 evergreen species of Berberis, the leaf margin is recurved to

such an extent that the margin is quite hidden when viewed from above, e.g. B. replicata and B. taliemis from Asia, and B. empetrifolia from South America. This is a useful criterion for a first step in determination.

(13) Size of leaves

In certain of the species with the smallest leaves, these are only some 3-12 mm. long, e.g. B. amoenu and B. wikonae. In most deciduous species of Berberis the leaves are 1-3 cm., sometimes up to &6 cm. long; very rarely, as in B. pachyacantha, or B. amuren- sis, up to 10 cm. long. In the evergreen Wallichianae species of Berberis the length of leaf varies from 1 to 2 cm. (e.g. B. vewuculosa) to 10-22 cm. as in B . i w i g n i s , B. Willearn; mostly they are 3-8 cm. long. In Mahonia, the leaflets are (4) 6-10 (-15) cm. long in the Asiatic, but mostly only (2-) 3-6 cm. long in the American species. The complete Mahonia leaf measures, for the Asiatic species 20-40 (-60) cm., and for the American ones mostly 10-20 (-30) cm. Consequently, a herbarium specimen of Mahonia may only include one or two leaves.

IV. The inJIorescence (Fig. 6)

(1) Berberis ; nuture of injZorescence The following forms may be distinguished : (a) Flowers solitary. Sometimes there are occasional pairs or threes. (Sects. Walli-

chianae and Angulosae, from Asia ; Sects. Microphyllae, Buxifoliae, Montanae, Vir- gatae and Confertae from America.)

(6) Flowers fascicled. (Mostly from the same Sects.). (c ) InjZorescence umbellate. The perfect umbel rarely occurs, but there is often a

subumbellate form in which the pedicels spring from a short length of the stem, leaving a distinct peduncle below. (Sects. Angulosae, Franchetianae, some Tinctoriae, etc.)

( d ) InJEorescence racemose-pseudumbellate. Here the rachis from which the flowers spring is longer, the number of flowers increased, and the lower pedicels lengthened. The flowers having become pendulous, the inflorescence tends towards a raceme while still retaining the appearance of an umbel-like form. (Same Sect. as c).

Page 12: Berberis and Mahonia : A taxonomic revision

12 L. W. A. AQLENDT

(e ) Injlorescence racemose. The formal raceme, sometimes slightly compound below, characterizes certain sections, namely : Asiatic-Vulgaxes, Dasystachyae, Brachypodae, Integerrimae, Asiaticae and some Tinctoriae. South American-La-, Truxillenses, Goudotiae and Quindiuenses.

(f) InJlorescence paniculate. This appeaxs in the Asiatic deciduous Sect. Polyanthae, Sherr i5aw and the South American Paniculatae. An interesting, distinct form is presented by B. chitria, where the inflorescence is paniculate in its branching, but, &8

in (d) , presents an umbellate-corymbose appearance.

(2) Mahonia; nature of inflorescence

M a h i a spp., namely:

Horridae.

Paniculatae.

The forms of inflorescence which occur in Berberis appear in ody 23 out of 110

(a) A few-flowered umbellate form, in the seven species of the North American Sect.

(b) A loose-flowered simple raceme, in five species of the North American

(c) A panicle, in the remaining 11 species of the Paniculah. (d) In the remainder, which form the great majority of M a h i a spp., there is a form

of inflorescence quite unknown with Berberis. This is a fascicled group of narrow formal racemes, there being (2-) 3-20 racemes in the inflorescence. Thus for the greater part of the genus Mahania, the leaf distinction from Berberis is supported by this unique form of inflorescence.

(3) Bracts at the base of t h i n . a c e n c e in Mahonia The bracts to be found at the baae of the inflorescence have a high diagnostic value,

and may be either (a) persistent and large, (1-) 1-5-3 (4) cm. long, as in the 66 Asiatic Mahoniaa, which, together with one from North America ( M . nervosa), form the Sect. Longibracteatae or (b) deciduous and small, i.e. 2-6 (-8) mm. long, aa in 42 American species, namely all except M . nervosa.

(4) The pedumk of the injlorescence In Berberis a peduncle occurs in umbels, subumbels, pseudumbels and some racemes

and panicles, and is sometimes, though not often, a specific diagnostic character. With M a h i a the peduncle is practically absent or inconspicuous, except in the Sect. Pani- culatae.

( 5 ) The number of jlowers i n the injbrescence The number of flowers in the inflorescence varies from 1 to about 250. The four

following categories concur with different forms of inflorescence and other characters, and are often typical of sections. (a) Flowers solitary (Berberis). (b) Inflorescence, 2-6 (-8)-flowered, the average not over 6, occurrjng in a majority of the forms in (1, b-d), and (2, a, b) , on pp. 11,12. (c) Inflorescence (8-) 10-14 (-15)-flowered, occurring in the remainder of these and part of (1, e ) and (2, c) , on p. 12. (d) Inflorescence (lo-) 15-40 (-25O)-flowered, including a few fascicled forms (1, b) but mainly found in the racemose and paniculate species of Berberis (1, e, f ) , on p. 12, and the multiracemose typical Mahonias (2, d ) on p. 12.

(6) Length of pedicels of individual j lmers

be classified as follows. The pedicels of the individual flowers are of varying length. For Berberis they may

Page 13: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 13

(a) Pedicels, 1-9 mm. long, recorded in 232 Berberis spp., included almost entirely amongst those with many-flowered racemes and panicles. There are a few exceptions when these short pedicels occur with solitary flowers (e.g. B. candidula and B. multiflora), with fascicles (e.g. B. wilsonae),’and with umbels (e.g. B. amoena). ( b ) Pedicels, 1-14 (-2) cm. long. Recorded in 122 species of Berberis, mostly associated with inflorescence forms (1, a-d) on p. 11. (c) Pedicels, 143-2-3-5 (-5) cm. long, recorded in 52 species mostly associated, with (1, a, b) .

With Mahonia the pedicel length covers a shorter range which may be classified thus. (d) Pedicels, 1 4 (-5 mm.) long. Occurring in 39 species, 30 of them Longibracteatae,

nine from the remaining, American, sections. (e ) Pedicels, 5-8 (-10) mm. long, occurring in 30 species, 14 of them Longibracteatae and 16 of the American species. (f) Pedicels, (8 mm.-) 1-2.5 cm. long. Seen in only nine species, two Asiatic (Subsect. Dolichopodae), and seven American species.

(7) Thickness of pedicels Some pedicels are conspicuously slender (e.g. B. leptopoda), others noticeably stout

(e.g. B. glaucocarpa and B. miaria). In Mahonia, the pedicels tend to be more often stout, but there are slender-+celled species. Generally the pedicel is slightly thicker at the apex. An unusual example is B. incrrnsata whose pedicels, remarkably thin and thread-like at the base, are steadily broadened until they become very thick at the apex.

(8) Pubescence of Berberis pedicels Berberis pedicels may be glabrous, puberulous, or pubescent. These conditions do not

necessarily coincide with similar conditions of stem. Thus, in B. angulosa, stems and pedicels are both pubescent. In B. ludlowii, the stems are equally pubescent, but the pedicels glabrous. In B. hirtellipes the stems are glabrous and the pedicels puberulous. About 35 species are recorded as showing some degree of pubescence on the pedicels.

(9) Bracts of the jlowers The bracts of the flowers in Berberis are to be found at the bases of the pedicels, of the

subpeduncles of a paniculate inflorescence, and of the main peduncle of any inflorescence. They are mostly reddish, triangular, and vary from 1 to 4 mm. in length. The bracts are usually very much shorter than the pedicel they support, being only one-twentieth to one-third or rarely one-half as long. In certain outstanding species the bracts are of diagnostic value, where they are nearly, or quite, as long as, or even slightly longer than, their pedicels, which in such species are always short. This character appears in certain species of the Sect. Polyanthae (with paniculate inflorescence), e.g. B. edgeworthiam, B. aggregata; and in most of the Brachypodae, e.g. B. giraldii, B . brachypoda, which, on account of their short pedicels, have spike-like racemes which possess a prickly appearance in bud due to the protruding points of the bracts.

The bracts at the bases of the pedicels to the flowers of Mahonia may be divided into three categories. (a) Bracts distinctly shorter than the pedicels, i.e. 0-14 .5 (-0.6) x pedicels ; this feature appears in a majority of Berberis spp. and about half of the Mahonias (22 Asiatic and 27 American). ( b ) Bract,s about as long as the pedicel, i.e. 0.75-1.25 x pedicels, as in the few Berberis spp. with long bracts, and in 19 Mahonias (17 Asiatic, 2 American). (c) Bracts conspicuously long, i.e. 2 to 3 times as long as the pedicels, a feature which is never seen in the genus Berberis, and which appears in eight Asiatic M a h i a spp. (Eulongibracteatae).

It will thus be seen that the Asiatic Mahonias tend towards long flower bracts and the American species towards short flower bracts.

Page 14: Berberis and Mahonia : A taxonomic revision

14 L. W. A. ~ E N D T

V. TheJl0Wer.s (Figs. 7, 11 and 13)

The flowers consist of prophyls (or bracteoles), sepals, petals, stamens and ovary.

(1) F h r size

Three sizes, partly concurrent with other characters may be recognized. (a) Flowers abnormally small, 3-5 mm. diam., with the longest component 2-3 (-3.5 111111. long). (b) Flowers of normal size, (6-) 7-10 (-12) mm. diam., with’ the longest component 4-66 (-7) mm. long. (c) Flowers abnormally large (1.S) 1.4-2 (-3) cm. diam., with the longest component (7-) 76-10 (-15) mm. long.

Small-flowered species occur, particularly in the Sect. Integerrimae, Dasystachyae and Subsect. Keissleriam.

Large-flowered species occur in a small subsection of the Wallichianae (Hookerianae), generally in the most typical of the Angulosae, and in the South American Montanae, Buxifoliae and Quindiuenses.

( 2 ) Plower w h r In all the Asiatic species of Berber& and generally in Mah4mia the colour is a fairly

pure yellow, pale or deep in the range H.C.C. 1 to 3 : occasionally a pale greenish yellow as in B. virescens and B. phwnsh. Where recorded in the South American species, it is golden yellow, orange, apricot, or reddish yellow, H.C.C. 5-14. B. hispniea from South Spain, and Sect. Horridae of M a h i a tend to show the reddish colouring of the latter group.

( 3 ) The bracteoles or pophylls in Berberis At the apices of the pedicels in Berberis and considered to be part of the flower, 1-2

or occasionally 3 bracteoles are appressed to the calyx. The bracteoles are lanceolaie, or more shortly triangular, and acute or acuminate: generally reddish or greenish, or partly yellow. Most species possess bracteolate flowers. Sometimes the bracteoles are purely yellow and sepaloid in nature, though still very small and prophylloid in form. When this is so the flowers appear quite yellow beneath, a t Grst sight as if they were without bracteoles, and may be called pseudobracteolate. There are, hally, certain species with genuinely ebracteolate flowers, a character generally associated with the large-flowered species of the A n g u l o ~ , but found elsewhere as well. Ebracteolate flowers present, when viewed from below, or in bud, a clean, unbroken, yellow aspect, and the outer components are either (a) small sepals (e.g. B. concinna) which are yet larger, as well as different in shape from, pseudoprophylls, or (b) large sepals, often almost as large as the inner ones (e.g. B. unquha, B. nmrosepda). When, however, the flowers are truly bracteolate, the yellow appearance from below is broken by the green or red colour of the prophylls, which is often ornamental when the flower is in bud, and the prophylls are prominent.

(4) The bracteoles in Mahonia Bracteoles are absent from the Asiatic species, apart from M . mairei, and M . bracteo-

lata. In the American species, two-thirds are recorded as possessing prophylls. The prophyls of M h i a may be (a) detached, or (b) appressed. The former, detached from the flower, generally appear near the mid-point, but sometimes towards the base, of the pedicel, a disposition which is uncharacteristic of Berberis but which has been recorded in 23 Mahonias. In the second type, the prophylls a t the apices of the pedicels are appressed to the flowers, as a kind of epicalyx. This occurs in the majority of the genus Berber& but is recorded in only two (American) Mahonias, M. eutriphyUa and M . nevi&

Page 15: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia

(5) Sepals in Berberis

15

The sepals are generally in two series, or whorls, of three each. In a number of species, there are three such series, each of three sepals. There are 215 species recorded with two series, and 87 species with three series, of sepals. Rare exceptions are two species with only one series of three sepals, namely B. oblonga, and B. lepidifolia, and one species, B. retinervia, with four series of sepals. Generally the presence of two or three series is a specific character, the three series often being associated with the large-flowered species. There are, however, species with varieties, of which some have two and some three sepals, e.g. B. circumserrata.

In most species of Berberis the inner sepals are longer than the petals, and it is then generally found that the outer are between one-half and two-thirds aa long as the inner sepals. This structure occurs in a series of 144 species which may be termed Usitatae. Thirty-eight of these species have three whorls, and 92 species have two whorls, of sepals. Occasionally, the outer are less than half the size of the inner sepals (series Parvisepalatae), and this occurs in 53 recorded species (31 with three whorls of sepals, 17 with two whorls and one with four whorls). On the other hand, in some species, very often the large-flowered ones of the Sect. Angulosae, the outer are almost, or quite, as long as, though never longer than, the inner sepals. This, the series Grandisepalatae, numbers 29 recorded species, eight with three and 21 with two whorls. The outer, although they are in this series as long as the inner sepals, may not be identical owing to variations in breadth and shape. Thus in B. angulosa the outer are little more than half as broad as the inner sepals, while in B. "Ccrosepah they are as broad, and more or less identical with them. The nature of the median sepals, where there are three whorls, varies, apart from the above distinctions. Thus the median may be just duplicates of the inner sepals, or, subequal to them when this is so, the outer sepals may also be equal, giving three equal whorls as in B. parisepah. Secondly, the sepals of the outer may be distinctly smaller than those of the two equal inner whorls, aa in B. concinna. Then again, the median sepals may be intermediate in size between those of the outer and inner whorls, so that the three whorls form a regular gradation in sue, e.g. in B. kuma- onensis.

(6) Sepals in Mahonia These are, as Takeda has pointed out, almost invariably in three whorls. An odd

example, with only two whorls, has been recorded, but not verified. One exception has been vefied, M . nevinii with only one whorl of three sepals.

(7) Size of Berberis petals In the Usitatae and Parvisepalatae, the petals, though always shorter than the inner,

are longer than the outer sepals. In the Grandisepalatae, the petals, by virtue of being shorter than the inner, are also shorter than the subequal outer, sepals. The relatively large size of the sepals, which the name Grandisepalatae implies, is due to the large outer sepals, and not to any inherent smallness of the petals which bear a normal re- lationship to the inner sepals. There are, however, certain species in which the petals are shorter than the outer sepals, these, in turn, being normally smaller than the inner. In this series, Parvipetalatae, the petals are the smallest components by virtue of their own inherent smallness (e.g. B. micropetala). Fourteen such species are recorded, four with three, and 10 with two, whorls of sepals. Finally, there is the series Longipetalatae, where the petals are equal to, or are longer than, the inner sepals. This series in which the petals are the longest components, includes a considerable minority, namely 90 species, of which 10 have three whorls of sepals, 78 have two whorls, and two are the only species with but one whorl.

Page 16: Berberis and Mahonia : A taxonomic revision

16 L. W. A. ~ E N D T

(8) Relatkve size of sepah and petuls, in Berberis and Mahonia

in different proportions, in M a h i a , as shown by the following table: The five seriea of flower structure just described for Berberis may be discerned, though

Berberis M a h i a Usitata-s 144 Longiptahtas 90 Parviaepelatae 54 cfrandisepalat&3 29 Parvipetalet.e 14

9 8

41 3

These characters, described as series of flower structure, run as threads connecting otherwise widely distinct species belonging to different sections (or blocks of species) whose association depends on other concurrent characters, and within which the flower structure is a useful constant discriminant between species.

(9) Confwim between bractwka and outer sepals Sometimes, in referring to Berberis species, Merent authors, or even the same author

at different times, have referred to the outer flower components of the same species indiscriminably as brmteoles and outer sepals. Thus one and the same species has been described, at one time as powxising prophylls and two whorls of sepals, and elsewhere as having no prophylls and three whorls of sepals. If, for example, the generally small outer sepals of h€ahonia were classed as prophylls, the majority of Mahonias would then fall into line with Berbe& by appearing in the first s e r i ~ of flower structure. Some botanists might consider this a reason for regarding the outer components of these &labia flowers as prophylls. However, although some outer sepals are somewhat prophylloid in nature, it is generally possible, after experience in examining the flowers, to make a satisfactory distinction. Whereaa the prophylls a m nearly always 1 or 2 in number, and triangular and acuminate, the small outer sepals in a whorl of three, are ovate, and acute.

(10) Base of petals: Berberis (Fig. 8) The bases of the petals may be either cuneate and truncate, or clawed. Those that are

clawed may be slightly, broadly and shallowly, or narrowly and conspicuously so. This feature, though recorded wherever possible, and generally constant within a species, has not been used in diagnosis. At the base, there are always present two glands, which easily distinguish the petals from the sepals. The glands are either subconcolorous (at most a slightly deeper yellow than the petals), or more or less conspicuously discolorow (orange). The shape of the glands varies from narrowly lanceolate, through normal oblong or elliptic, to orbicular shapes. Their length, 0-3-1.5 mm., is from one-tenth to one-fifth the length of the petal. In extreme examples they are contiguous and centrally placed, or widely separated and actually on the margin of the petal. Generally they are moderately separated and non-marginal. They may be either within the basal claw, if this exists, or above it, but are always within the basal third of the petal. The glands and basal character of the claw, though fairly constant, have rarely been used as a specific discriminating character, an exception being B. avpproximata (glands approxi- mate).

(1 1) Base of petu& : Mahonia In nineteenth-century writings, it has been stated that the petals of Nahonia have no

glands, and this was cited as one of the bases of discrimination between the genera Berberis and Nahunia. This is far from accurate. The origin of the misconception appears

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Berberis and Mahonia 17

to be that, in many species, the glands are very indistinct and therefore apt to be over- looked. Many species, however, have quite distinct, and some conspicuous, glands, as in Berberis.

(12) Apex of petals The apex of the petals is entire in no more than 66 Berberis species. It is recorded as

emarginate in 266 species. The form of the emarginate apex of these varies. It may be acutely emarginate, shallowly retuse, or narrowly incised. In each case the surrounding protecting lobes of the petal may be either acute (apex biapiculate), or obtuse. The combination of these characters provides six different kinds of emarginate apex. The form of the apex is fairly constant, and the general distinction between entire and emarginate apices is sometimes used in combination with other characters, to separate species. In the genus Mahonia, the emarginate petal is even more predominant, there being recorded only four species with entire petals.

(13) Pentamerous Jlozvers A few abnormal species are recorded in which the components occur in the flowers in

whorls of five sepals and petals (instead of six or possibly nine for sepals). Examples are B. comberi and B. virgatu.

(14) Shupe of sepals and petals The outer sepals are generally ovate and acute or subacute, and the inner sepals

obovate and rounded. Petals are generally obovate and rounded, regarding the outline apart from the break therein in emarginate petals. There are exceptions in certain species. In rare and extreme examples both outer and inner sepals may be very acute or even subacuminate, as in B. kuwakumii, where the flower buds present a prickly appear- ance due to the slightly unfolding points of these components. A similar appearance ia sometimes caused by the bracts; see (IV, 9) on p. 13.

(15) The stamens in Berberis The stamens and the petals are attached at their bases. The stamens are generally

about two-thirds as long as the petals, and their length is rarely used in specific dis- crimination. However, the possession of stamens almost as long as petals haa been one of the several foundations for the separation of B. chillanensis from B . montuna, while there is recorded one species, B. dolichostemon, with stamens conspicuously longer than the petals.

(16) Dentate stamens in Berberis There are in a few exceptional species, of which about 10 are recorded for certain,

teeth on either side at the apices of the flaments, below the anthers. These teeth may be short and acute, (rarely obtuse), or long and acuminate, and these variations are very distinct and most valuable for diagnostic purposes. Teeth occur in some species of the South American sections Buxifoliae and Actinacanthae. All other species of Berberis have stamens edentate below the anthers. Often, in the Asiatic sections Asiaticae and Tinctoriae the stamens are somewhat swollen, in a slight, gentle curve, below the anthers. This presentation, scarcely worthy to be called subdentate, indicates a little connexioii with the South American species just mentioned. The fact is interesting, as it is one of several characters which place these two sections closer to the South American species than are the other Asiatic sections, but the other features relate the Asiaticae and Tinctoriae to the non-dentate South American sections.

2 JOURN. L INN. S0C.-BOTANY, VOL. LVII

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18 L. W. A. ~ E N D T

(17) A p k s of stamens in Berberis The apices of the stamens may be either distinctly produced beyond the anthers; or,

on the contrary, possess no such connective production, or one that is scarcely dis- cernible. 120 species are recorded with stamens not, or scarcely, produced; and some 150 species with stamens definitely produced. Of these species there are 86 where the apex is conical-apiculate, sometimes shortly so, sometimes with a conspicuously elon- gated acuminate connexion. There are a h some 64 species where the stamens, though distinctly produced, have an apex rounded, obtuse, truncate, or sometimes slightly retuse-truncate. This character has some diagnostic value. The conspicuously and slenderly apiculate W e n s are constant in a species, and a firm distinction can bemade between species having W e n s distinctly produced, or not so. It is, however, often difhult to decide between subsidiary Werencea such as whether the apex is shortly apiculate, subapiculate, or obtusely-subconical, since the appearance may vary to this extent in Merent flowers. Four categories may thus be established: (a) apex distinctly produced, slenderly acuminate ; (b) apex distinctly produced, shortly conical-apiculate, obtusely subconical, etc.; (c) apex distinctly produced but rounded, or truncate; (d) apex not, or scarcely, produced.

(18) Dentate stamens in Mahonia The stamens of M h i u may be classified in the same way as those of Berberis.

Dentate stamens appear in only three Asiatic, but are recorded in 21 American, species. Probably more American species have dentate stamens, but, in a number, the flower is not known. Only three American Mahonias are recorded with dentate stamens, namely M. hartwegii, M. arguta and M . trifolwlutu. The dentate stamen is thus pro- portionately much more common in M h i u than in Berberk

The presence of dentate stamens in so many American Mahonias gave rise to the name Ohternon Rafin., for the genus. This name has been, on one occasion, cited by Fedde, in Engler, Bot. Jb. 31,1901, as 0dOntoetem-m. The name implied that dentate stamens are characteristic of M h i u rather than of Berberis. Hutchinson, in G a d Chron. 43, 82, 1908, pointed out that this distinction is not complete, on account of the existence of a few South American species of Berberis with this character. In doing this, he refers to the species of Berberis with dentate stamens as the Sect. Odontostemones, but without giving any authority or reference. "he Odontostemones is not one of the recognized sections, but is in the nature of a series of species belonging to different sections but which happen to have one particularly outstanding character in common.

(19) A m of stanzena in Mahonia Classifying the apices of the stamens in the same way as those of Berberis we find the

following typea: (a, b), apex apiculate (long or short) in 12 species, all Asiatic and pre- dominantly Himalayan and Indian; (c) apex produced but flattened, in nine species, all Asiatic; (d) apex not, or scarcely, produced, in 24 species (18 Asiatic, six American).

(20) T h m r y and oprulee

The ovary contains 1-12 (-15), or occasionally, more ovules. While the number of ovules is not actually constant in a given species, it varies within limits which allow four grades to be recognized. Any doubt that may arise can be settled by the examination of a considerable number of ovaries, in order to find the average number of ovules en- countered. The four grades are as follows: (a) Uniovulate. ovules nearly always solitary, rarely 2: average very close to 1. (b) Pauciovulate. Ovules generally 2, occasionally 3, rarely 1 : average 2, or just above, always legs than 3. (c) Pluriovulate. Ovules rarely 3, generally 4-5, rarely 6 : average 44. (d) Multiovulate. Ovules occasionally 5, mostly

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Berberis and Mahonia 19

6-15: average above, and generally well over, 6 . In this grade all species have between 6 and 12 ovules, except B. d ian tha and B. tsangpoen&s with 12-15 ovules, and Mahonia huemutocurpa with 12-15 (-18) ovules.

The number of ovules per ovary, in Berberis and Mahonia, may be summarized as follows :

BerberCs M a h k (species) (species)

U n i o vul a te 41 3 Pauciovulata 180 25 Pluriovulata 130 21 Multiovulata 51 16

VI. The fruit (Fig. 12)

(1) Shpe and size of the berry The berry is sometimes globose, varying from 4-8 (-11) mm. in diameter: but far

more often ellipsoid, oblong, ovoid, or obovoid measuring from 4 mm. to 2 cm. in length. The small sizes are found in a small minority of species. Normally, the non-globose berry is (6) 7-10 (-12) mm. long. The larger berries, 12-18 (-20) mm. long are found in many members of the Sect. Angulosae, generally in association with the large flowers. These are red-fruited, e.g. B. ludEowi, B. concinna, B. circumerrata and B. aernulans. Blue-black berries of a similar sue are found in a few species of Sect. Wallichianae, e.g. B. hoolceri and B. mlliantha.

In Mahonia, the berry is very frequently, though not invariably, globose, a shape which occurs here more often than with Berberis.

(2) Colour of the berry in Berberis The following colour variations may be recognized. (a) Species with red, dark red, or

reddish purple berries of which there are recorded some 183. These are all Asiatic and nearly all deciduous, except for the Sect. Tinctoriae. (b) Species with black berries, with or without bloom, of which there are over 300, the great majority evergreen.

In Mahonia all the species appear to have black fruit (mostly pruinose) excepting only the three (American) members of the Subsect. Haematocarpae.

( 3 ) Pruirwse berries Red berries are predominantly epruinose, but there are a few species where red berries

are cwered with bloom, e.g. B. humdo-umbrosa, B. aristata, B.Wibu& and B. beaniana. The black berries of the South American Berberis species seem generally to have a dis- tinct blue bloom upon them, but the berries of the Asiatic forms vary as regards this character. Some black-berried, Asiatic species of Berberis show a conspicuous white bloom, e.g. B. pruiwsa, B. julianae and B. lempergiana, but others have a conspicuous violet or mauve bloom, e.g. B, soulieana and B. lcumkamii var. fomnosana. Others again have a less conspicuous, but distinct, blue bloom, e.g. B. gagnepainii varieties and B. verruculosa. In some species, e.g. B. hookeri, this blue bloom becomes very slight. Finally there are some species with berries quite black, and epruinose, e.g. B. atromrpa, B. panhnensis and B. sublevis.

With Mahonia, so far as may be gathered from written records and herbarium specimens and in the absence of much living material, it seems that the berries, almost all black, are also predominantly heavily pruinose, only a few exceptions having been recorded.

2-2

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20 L. W. A. AHRENDT

(4) The textwe of the bewy The berries of some species of Berber& are very soft, and of a juicy texture Within, e.g.

B. Wilsonae. Others have a somewhat more solid, but still somewhat juicy, fleshy sub- stance. Others again have a berry in which the seeds are set in a dry pithy interior, e.g. species of the Sect. Wallichianae. The second type is the most common in both Berbehs and M a h i a .

(5) The style in Berberis and Mahonia The style, which may be absent from or present on the ovary, is persistent, and more

easily discernible in the berry. Three types may be distinguished. (a) Style absent, or so short, indekite and indistinct that it may be disregarded. This type occurs in both Asiatic and South American species with both red and black fruit. (b) Style distinct but not more than ( 0 - S ) 0-5-0.75 (-1) 111111. long. With Berber&, this type occurs most often in the Asiatic species. (c) Style elongated (1-) 1-5-3 (-5) mm. long. This type occurs almost exclusively in about one-quarter of the South American species of Berberis. Styles that are 2-5 mm. long when fdly extended do not occur in the Asiatic species which, however, include a few in which the style is as much as 1.5mm. long, e.g. in (i) B. oemaetonii, which, among the Wallichianae, is unique in this as well as in being the weaternmost member of the section; (ii) B. polyanthu which difFers in this respect from all other deciduous species from Asia; (iii) certain species of the Sect. Asiaticae (viz. B. lycioidm);and "inctoriae (e.g. the south Indian B. tinctoh and B. wightiam; the Himalayan B. petwhris, B. chit&, B. sikkimnsis and B. &ria). It has already been seen that of the Asiatic species of Berberis, these two sections approach most nearly to the South American.

The following table gives a comparison of the style in Berbe9-i.s and Mahonia.

Berber& M&nk (species) (species) Diagnostic status

Style indistinct or absent 167 25 Insignificant Style short ( 0 . S ) 06-4-75 (-1) mm. 109 33 Significant in Mahonk and

Style elongated (1-) 1-54 mm. 61 9 Sieslificant in Mahonk and Asiatic Berber&

South American Berberia

On the whole, it appears that a style is more often present in X a h i a than in Berberis. An elongated style occurs in seven Asiatic Mahonias, but only two American, a geo- graphical emphasis which is the converse of that in Berberis. Nearly all the Mahonia species with a short style are Asiatic (26), only seven being American. Of the estylose species, 11 are Asiatic and 14 American. Thus we may say that in M a h i a , the style is most prominent in the Asiatic, and the wtylose character in the American species.

(6) The seeds The seeds are from 1.5 to about 7 mm. (mostly about 3 4 mm.) long. With reference

to their colour, the seeds may be either (a) dark, i.e. dark red, red-purple or black, or (b) pale, i.e. pale yellow, yellow-brown, or occasionally pale reddish yellow. The colour distinction between the pale yellow and the dark red is the same as that which has already bean noted in the mature stems of Berberis, but there seems to be no correlation between the pale and dark colour categories of stem and seed. Generally speaking, the dark seeds belong to the evergreen species (e.g. Wallichimae which are very pre- dominantly pale-stemmed, and South American Sect.), and the pale seeds to the deciduous and red fruited species. Only dark seeds have been recorded in M a h i a .

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Berberis and Mahonia 21

(7) Diagnostic characters which distinguish the Eurasiatic from the South American groups

Certain sets of characters distinguish largely, though not in complete detail, between the Eurasiatic and the South American species of Berberis : the South American species are characterized by deep orange flowers, foliaceous spines in a few species, dentate stamens in others, a very elongated style in a considerable number, or evergreen leaves associated with long paniculate inflorescence. The Eurasiatic species have paler yellow flowers, deciduous leaves, or red berries, or very rigid evergreen leaves with a fascicled inflorescence. This being so, Schneider, in 1905, made a primary division of the Berberis into two botanico-geographical groups. His first group, the Septentrionales, includes all the Eurasiatic species and the only two Berberis species from North America. This group is therefore of the eastern hemisphere apart from these two species. It is pre- dominantly Northern, since only three of its species lie just south of the Equator (two from East Africa, and one from Java and Sumatra). His second group, the Australes, includes all the species of Berberis from South and Central America. It is entirely of the western hemisphere and predominantly Southern, since only some 20 of its species come from north of the Equator, in Colombia, Venezuela and Costa Rica. These two Berberis groups thus occur in the north-east and south-west respectively.

I have found it convenient to divide M a h i a into two analogous groups, (i) the Orientales, confined to Asia (with the addition of M . nervosa from America), extending only just south of the Equator in Sumatra, and (ii) the Occidentales, extending through west North America and Central America. The two Mahmia groups thus occur in the north-east and north-west respectively.

of Berberis

SURVEY OF GEOGRAPHICAL DISTRIBUTION

(1) Berberis. Septentrionales LATrruDmAL-Australes LONGITUDINAL (Maps 1 and 13)

Most species of the Berberis group Septentrionales lie between latitudes 22" and 34" N. Although so confined, they are scattered over a wide range from east to west, namely 50" to 135" (-140") E. longitude (a few extending south to the equater or to 20" W., and the two North American species excepted). The distribution in the group Australes is of the reverse nature, namely a wide range of latitude, 10" N., to 56" S., combined, for the great majority of the species, with a narrow range of longitude, 65" to 80" W. The main body of Sect. Laurinae, lying a little further east, in Brazil, Paraguay and Uruguay form a small exceptional set.

(2) Berberis, Group SEPTENTRIONALES, and Mahonia, Group OI~IENTALES (Maps 1, 13, 46)

(a) Mediterranean Region (Maps 3, 13, 39) Madeira, N.W. Africa (Morocco, Algiers), S. Spain, Mediterranean Islands (Corsica,

Sardinia, Sicily, Crete, Cyprus), S. Italy, Greece, Asia Minor. The home of Berberis of Sect. Crataeginae. Mahmia absent.

(b) Western and Central Asia (Maps 2, 3, 13, 30, 39) Passing eastwards, we meet Berberis, Sect. Integerrimae, centred in Armenia, Trans-

Caucasia and Persia, but extending slightly into Afghanistan and Baluchistan, and Turkestan whence also come the eastern spp. of Sect. Crataeginae. Mahmia absent.

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22 L. W. A. AERENDT

(c) We.9kwn H i d p (Maps 2, 3, 4, 5, 12, 13, 17, 35, M) Hazm, Kaahmir, Punjab Himalaya, Jaunsctr, Garhwal, Kummn, and W. Nepal.

Here the sub-evergreen Be&& spp. begin, particularly Sects. Asiaticae and Tinctoriae. These extend slightly eastwards into the next region, but much farther southwards into India and Ceylon, and south-west to Arabia and N.E. Africa. In this west Himalayan region, the Sects. Angulow and Polyanthae begin to appear; the former continues to the fax eastern range of the Septentriodes in the Pacific Islands; the latter extends only as fax east aa west and west-central China. Mahonk, Group Orientales, haa its western limit in the western Himalayas.

(d ) Ea&m Himalayas (Maps 2, 3, 4, 5, 12, 13, 17, 18) E. Nepal, Sikkim, Bhutan, Assam, S.E. Tibet, N. Upper Burma. This marks the

eastern end of the fairly restricted Berberie, Sects. Asiaticae and Tinctoriae. The Sect. Angulosae flourishes here, and Sect. Wallichianae (apart from its unique westerly member, B. asmastonii from Garhwd) begins here and extends to the eastern limit of the Septentrionales.

Concentrated in this axea are certain very distinct Berberis spp., namely the unarmed Subsect. Insignes of the Wallichizmae; also Wallichianae, Subsect. Hookerianae and Angulosae, Subsect. Euangnlow; these subsections, the former evergreen, the latter deciduous, are characterized by their very large multiovulate flowers. The range of M a h i u continues through this region, side by side with Berberi.9, Sect. Wallichianae, but, instead of possessing such an extensive easterly range, takes a southerly short cut to meet the Wallichimae once more.

(e ) North- We& China (Maps 2, 4, 13, 17, 19, 28, 32, 34) N.W. Szechuan, N.E. Tibet, Kansu, Shensi. This is the home of the Berber&, Sect.

Brachypodae, which is c o h e d here, and is the centre of the Sect. Daaystachyae which extends S.W. to the Himalayas in one direction, and to N. and E. China in the other, The northern subsection Diaphanae of the Angulosae is found here, whence the Angu- losae extend northwards to Siberia. B. soulieana, the outlying northernmost alp. of Sect. Wallichianae belongs to this region from which the genus M a h k is absent.

(f) We& China (Maps 3 and 23)

Sect. Eranchetimae, and of some spp. of M h k . The borders of Tibet (Prov. Mekong), Szechuan, N. Yunnan. The home of Berberis,

(9) South-wt China (Maps 2, 12, 13) West Yunnan, bordering on N. Burma, S.W. Y m a n , Kweichou, Kwangsi. Whereas

the Berberis, Sects. Tinctoriae and Asbticae extend far south and farther westwards, in India, it is here that the Wallichianae find a southward (though smaller) extension of their range. Mahonk appears here with a range following S.W. with that of the Wallichimae.

(h) Cen4ra.l China (Maps 2, 13, 33) E. Szechuan, W. Hupeh, N. Hunan, provides some spp. of Berberis, Sect. Wallichianae.

The Sect. Vulgares with its range b e m g with the large area inhabited by B. vulgaris in Central Europe, reappears here, being represented by B. henryam. Although N . of the general M a h i a area, M . bealei is found here.

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Berberis and Mahonia 23

(i) North-east China (Maps 2, 3, 13, 23, 27, 33) Shansi, Shantung, Chihli, Honan, Kiangsu, Manchuria, Korea, and the adjacent

Amurland of S.E. Russia. The Sect. Vulgares, distinctly northern, and stretching widely from west to east, is represented here by B. amwemis, B. bretschneideri and B. km-eana. This region is the centre of Sect. Tschonoskyanae. Mahonia is absent from this region.

(j) Japan and the Korean archipelago (Maps 2, 3, 13, 23, 27, 33)

home of the Tschonoskyanae, M a h i a being absent. This region contains some endemic Berberis spp. of the Vulgares, and is another main

( k ) East China (Maps 2, 13, 32) Anwhei, Chekiang and Kiangsi. The Berberis, Sect. Daaystachyae, extending south

from N.E. China is represented here by B. chekiungewis and B. anwheinensis. This region includes part of the far eastern range of the widespread southerly section Wallichianae (B. lempergiana, B. chingii) with a few spp. of Mahonk, both of which are similarly represented in the next and less botanically explored region of S.E. China

( 1 ) South-east China Fukien, Kwangtung. Mahonia fordii occurs here.

(m) The Paci$c Islands (Maps 1, 2, 13) Formosa, The Philippine Islands, Java and Sumatra. The most easterly and southerly

extension of the Berberis, Sect. Wallichianae, represented here by various interesting spp. B. xanthoxyh from Java is one member of the Septentrionales, which just over- laps into the southern hemisphere. Its range having previously taken a short southern route, Mahonia meets Berberis again here.

(n) Siam, Indo-China It is here that Malwnia appears alone, its range having taken the southerly short cut

just referred to. It will be remembered that its companions of Berberis, Sect. WaU- chianae have a range proceeding more south-easterly through China (whence it is accom- panied part of the way by Mahonia) and curving round to meet the same southern extremity of distribution in Java and Sumatra.

(0) India, Ceylon (Maps 2 and 13) The southerly extension of Sect. Tinctoriae.

( p ) East Africa (Maps 1, 2, 13) From Abyssinia to Kenya there are a few spp. of Berberis, Sect. Tinctoriae, which, it

will be remembered has a long, though sparsely populated, south-westerly extension of its range from the western Himalayas, through Arabia, to these parts. Here are the other two members of the Septentrionales found just south of the equator.

(q) North America (Maps 1, 2, 47) The Septentrionales, though belonging so characteristically to the north and east,

have not only the three spp. mentioned in the southern hemisphere, but also two exceptionally placed species from the western hemisphere, namely B. fendkri and B. canadensis, from the U.S.A., in Virginia, Missouri, New Mexico and S. Colorado. The first two regions provide only B. canadensis. The last two are the habitat of B. fendleri which just verges on the west U.S.A. range of Mahonia, Group Occidentales.

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24 L. W. A. ~ E N D T

(3) Berberis, Group AUSTRALES (Maps 1 and 42-45)

The longitudinal distribution of the Australes provides botanical and geographical divisions more mutually concurrent, and leads to the conception of two subgroups. Them are: (i) the EUAUSTLGES, lying in the latitudes 30'46" S., and (ii) the AEQUINOC- TIALES, ranging from 10' N. lat. to 23" S. ht. These subgroups lie almost completely in the west of South America. Latitudinally between them, but longitudinally farther east, is the Sect. Laurinae, which is botanically related to (i), but, extending north-west, overlaps the area of (ii). It has seemed most natural to treat this as an intermediate section between the subgroups.

(3A) Subgroup EUAUS-s Chile, Argentine, with a slight extension into Uruguay.

(3B) Intermediate Sect. Laurinae Brazil, Paraguay, Uruguay, with a north-westerly extension as far as Ecuador.

(3 C) Subgroup AEQUINOCITALES (a) Southern tropical region. North Chile, Bolivia, Peru. The home of Sects. Aga-

patensas, Virgatae, Confertae, !Lhxillem (Subsects. Pichinchensea and Keisslerianae), Quindiuensea, and most of Paniculatae.

(b) Northern tropical region. Ecuador, Colombia, Venezuela, Costa Rica, Nicaragua. Here belong other spp. of the Sects. Confertae, Quindiuensea and Paniculatae; Sect. Trudenses (Subsect. Eutruxillenses) ; and particularly Sects. Latifoliae and Goudotiae.

(4) Mahonia, Group OCCIDENTALES (Maps 1,47,49, 51) These are found in Central America, Mexico, and western U.S.A. with one species in

S.W. Canada. (a) Central America. Malumiu, Sect. P a n i d a h , just overlapping the most northerly

spp. of the Berberi-s of Group Septentrionales. (b) Mexico, and reaching into the south of U.S.A. kialumiu of Sects. Paniculatae,

and Horridae, which, together, form a small exceptional set of Mkhoniaa with a Berberis inflorescence. It should be noticed that passing northwards the true Berberia of S. America first gives way in Central America to Mahonia, in the leaf character which is the fundamental distinction. But the Berberi-s kind of inflorescence persists much farther north, until there is found the distinctive Malumiu kind of inflorescence which unites the main body of American Nalumiu spp. with all those from Asia.

(c) West of U.S.A. Malumiu, Subsect. Euaquifoliatae, with the typical Mahonia inflorescence. One sp., M. uquifoZiurn, extends aa far north as British Columbia.

(5) General concurrence of botanical and geographical characters in Berberis, Group AUSTRALES (Maps 1,42, 44,45)

(i) Subgroup EUAUS-S. Generady, a few-flowered inflorescence combined with an insignificant style; or a many-flowered inflorescence combined with a signikant style.

(ii) Subgroup AEQUINOCTIALES. Generally, a few-flowered inflorescence combined with a s i d c a n t style; or a many-flowered inflorescence combined with an insigniscant style.

THE USE AND NATURE OF THE KEYS

Nature of the Keys A taxonomic key may serve two purposes, either singly or combined. It may indicate, or it may define. If an unknown specimen is to be determined, application of one, or more, keys, may result in the indication of a species which, the key suggests, may be

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Berberis and Mahonia 25

the desired determination. On the other hand, study of a key may provide considerable information concerning the nature of a species and the limiting characters of its popu- lation, together with the features which separate it from a closely related species.

Probably, all keys combine both properties, but in considerably varying proportions. In the many keys to be found in the following pages, some deal with the whole of a genus, or a considerable portion of it; these are the most difficult to construct, and appear most complex; their nature is predominantly indicative; examples of these are Keys 1-3, where, as often in such cases, the indications have to be given sometimes to sections, sometimes to species. When a smaller range is treated, it8 in Key 4, which indicates the subsections in a large section, the degree of definition offered by the key is increased; and, when, in Key 5, only a few spp. in a subsection are considered, still more definition is offered, and the content and nature and separations of this portion of the genus can be well appreciated before reference to descriptions is made. When, at the end of a section, it is again necessary to treat together a much larger number of spp., the key provided will necessarily offer much less definition, and be more indicative, as when we proceed to Key 27 ; subsequently the development just described will re-occur, with increasing definition allied to the indicative quality which is always present. Sometimes, the degree of definition afforded is very high indeed, a.s with Key 92.

Use of the Keys Key 1 is, as far as I am aware, unique in being a treatment of the whole genus Berberis

considered together. For the student, it shows, as well aa I can manage, the purely botanical relationships which are necessarily ignored by Schneider's botanico-geographical separation into groups. For purposes of determination, Key 1 isessential, if the investigator is examining material about which he has no previous knowledge, that is, begins un- aware even whether it is of Asiatic or American origin. If the material belongs to one of the many species subsequently described, Key 1 should give a valuable indication which can be followed in the later keys and descriptions. But, it should be remembered that, if this key is applied to what may prove to be a new species or variety-or, with cul- tivated material, to one of the numerous casual hybrids which abound-Key l is unlikely to give an effective result.

Fortunately, for the investigator, it should rarely be necessary to use Key 1. Whether the material is dried or living, the group to which it belongs is nearly always known, and it is possible to begin at a later stage, e.g. Keys 2 or 93. Continued investigation will lead, in due course, through the keys for sections and subsections to the species sought, and, if need he, to the further keys defining varieties within the species.

It may be noted that in some cases alternative methods of determination are provided, e.g. a general key to Berberis, Sect. Wallichianae, as well as a key to its sections followed by a series of keys to the spp. within the sections; again a combined key to the Berberis, Sects. Asiaticae and Tinctoriae, with included references to some species of the Angulosae ; the general keys, in such cases, will be mainly indicative, and the particular ones will offer more definition.

Often an investigator will already have a good idea concerning the section to which a specimen belongs ; then, passing over earlier keys, he will consult directly the one appro- priate to the section with which he has decided that he is dealing. Sometimes however, owing perhaps to the complexity of the genera, or to a little uncertainty in the mind of the investigator about characters, he may confidently be able to decide that his material belongs to one of two sections, but without knowing which. In such cases, the alter- natives just mentioned should be of great value, since he can confidently begin by con- sulting the more general of the two, without having recourse to those which precede.

Page 26: Berberis and Mahonia : A taxonomic revision

26 L. W. A. ~ E N D T

THE GENUS BERBERIS L. [The Barberry or Pipperidge Bush, Miller, Gard. Dict., 1769.1

KEY 1. GENEBAL KEY TO TEE GENUS B E R B ~ Inflorescence not panicdate, but solitary, fascicled, subumbellate or racemose.

Style conspicuous, (I-) 1 - 6 5 nun. Inflorescence 1-6 (-8)-fld., solitary, fascicled, or subumbellate. A

fascicled. B (cf. G)

C (cf. F)

subfascicled, rarely fascicled. DandG

Inflorescence (8-10-30-fld., racemose, occasionally racemose-subfascicled, rarely

Style absent, or short, to 0.5 (-0.75) mm. Inflorescence 1-6 (-8)-fld., solitary, fascicled, or subumbellate. Inflorescence (8-) 10-30-fld., racemose, occasionally racemose-subumbellate, or

Inflorescence panicdate. E

A. Flowers solitary. Leaves broad, or moderately narrow, 1 : 1-5-3 (4).

Ovules 2-6; leaves evergreen. Ovules 6-9; leaves deciduous.

Sect. VIRGATAE (Keys 79-81) Sect. MONTANAE (Key 66)

8. B. osmaston;ii,1 of Sect. WBLLICHIANAE

Sect. MONTANAE (Key 66)

Leaves very narrow, 1 : 5-6, evergreen; ovules 6-9.

Flowers not solitary. ovules 6-9. O d e s 2-5.

Leaves evergreen. Leaves 6-20 (-30) IIM. Leaves 3-8 cm.

Sect. VIBGATAE (Keys 79-81)

Sect. TRIGONAE- (Key 72) Sect. ILICIFOLIAE (Subsect. ETJILICIFOLIAE) (Key 70)

Leaves entire. Leaves dentate.

Leaves deciduous. 309. Bc?&& cd2iobow8 (k t . HETEBOPODAE)

6. Leaves deciduous. Fruit, red, or dark red. Fruit black, pruinme grey or blue.

Sect. TINCTORXAE (Keys 19, 21-26)

ovules 3-5 (-7). Sect. ASIATICAE (Keys 19, 20) ovules 2. 307. B. o b w a (Sect. HETEROPODAE)

Sect. TINCTORIAE (Keys 19, 21-26) Leaves evergreen.

Fruit red, or dark red. Fruit black, mostly pruinose. Stems pubescent. Stems dark, red-brown, terete.

Sect. ILXCIFOLIAE (Subsect. DARWI"AE)

Sect. LAWRINAE (Keys 73-76) (Keys 68, 69)

113. B. chitriCr (Sect. T ~ ~ O I U A E ) Sect. ' l h m R & i E (Key6 19, 21-26)

i Inflorescence racemose.

Infloreamnce corymbose-panicdate. Stems pale yellow, very sulcate.

Stems dark red. Stems glabrous.

spines absent or Weak. 105. B. petiolaris (Sect. TINCTORIAE) Sect. ILICIFOLIAE (Subsect. CEIILENSES (Key 71)

(Sect. LAURINAE (Subsect. FLEXTJOSAE) (Key 76) Spines strong

stems pale yellow. Leaves thin, flexible, subcorheous. 98. B. lycwides (Sect. ASIATICAE) Leaves thick, rigid, coriaceous. Sect. LA-AE (Keys 73-76)

1 B. o m t o n i i from Garhwal is remarkable in being both the westernmost species of the Asiatic Sect. Wchianae, and botanically the closest of this section to the South American Group Australes, with which it forms a link through the Sect. Virgatas.

Page 27: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 27

C. Leaves deciduous.

Fruit red, dark red or red-brown. Sect. TLNCTORIAE

Ovules (3-) 4-15. Sect. ANGULOSAE Sect. POLYANTHAE (Subsect. PSEUDOPOLYANTHAE) J Sect. FRANCHETIANAE

Sect. POLYANTHAE (Sukct . PSEUDOPOLYANTHBE)

\(Key 27)

(Keys 35, 53) I 1 1 Ovules 1-2 (-3). Sect. TSCHONOSKYANAE

Sect. HETEROPODAE (Key 55) Sect. CRATAEGINAE

Sect. ULICINAE

(Key 27)

I i h i t black, mostly pruinoee.

I Sect. T I N ~ R I A E (Subsect. UMBELLATAE) sect. ANQULOSAE

Fruit red, dark red or red-brown. Sect. POLYANTHAE (Subsect. PSEUDO-

Fruit black, often pruinose. POLYANTHAE)

Sect. ACTINACANTHAE (Keys 60-62)

i Leaves evergreen.

Spines foliaceous, palmate or manifold. Spines 1-3l-Bd., or absent.

Leaves broad, 1 : 1-1.3. Leaves narrower 1 : (1.5)- 2-3 (4).

ovules 6-12. Sect. AGAPATENSES (Key 83)

Flowers deep orange: fruit globose. Sect. BUXIFOLIAE (Key 65) Flowers yellow; fruit oblong. Sect. WALLICEIANAE (Subsect. HOOKERIANAE)

(Key 6) Sect. WALLICHUNAE

Ovules 1-5.

D. Leaves deciduous. F- { Sect. C O N ~ U T A E

Fruit red, dark red, or red-brown.

Inflorescence racemose.

Inflorescence fascicled, or subfaacicled.

Ovules 3-4; leaves entire, rarely with a few spinules.

}(Key 41) Sect. CANADENSES { Sect. INTEGERRIMAE

Sect. POLYANTHAE (Subsect. PSEUDOPOLYANTHAE) (Key 53) Ovules 1-2.

Inflorescence subfascicled, or subumbellate. Inflorescence subfascicled ; leaves closely reticulate.

Inflorescence subumbellate; leaves slightly or openly veined. Sect. POLY~LNTH~E (Subsect. PSEUDOPOLYANTHAE) (Key 53)

sect. SINENSES (Keys 41, 43) Inflorescence formally racemose ; leaves mostly multi-spinose.

(Keys 45-47) Sect. DASYSTACHYAE Leaves glabrous. Sect. VULGARSS Leaves Dubescent. Sect. BRACHYPODAE

Fruit black. Sect. CRATAEQINAE Sect. HETEROPODAE

Leaves evergreen. Spines foliaceous palmate, or manifold. Spines 1-3-fld., or absent.

Infloreacence solitary or fsscicled. Stems pubescent. Stems glabrous.

Sect. ACTINAOAWIT~AE (Keys 60-62)

92. B. &at& (Sect. ASIATICAE) Sect. WALLICHIANAE (Key 3)

Inflorescence racemose, subumbellate, or racemose-subfascicled. Fruit red.

Inflorescence racemose-subfascicled; leaves 8-20 m., closely reticulate. Sect. POLYANTHAE (Subsect. PSEUDOPOLYANTHAE) (Key 53)

Sect. TINCTOR- (Keys 21-26) Tdorescence racemose or racemose-subumbellate ; leaves larger, more

openly veined. 1 See page 4 for footnote.

Page 28: Berberis and Mahonia : A taxonomic revision

28 L. W. A. AHRENDT D. Leaves evergreen (cmtinued)

Fruit black.

E. Leaves evergreen.

Sect. ASIATICAE sect. TINCTOR-

Sect. TRUXIS.LENSES Sect. QUINDIUENSES

Fruit black. Fruit, red, or dark red, or red-broun.

Inflorescence corymbose-panicdate. Inflorescence normally paniculate.

Sect. PANICULATAE (Key 90)

113. B. &itria (Sect. TINCTORIAE) Sect. POLYANTHAE (Keys 49-52)

Leaves deciduous. Fruit black, pruinose blue. Fruit red.

Sect. SHERRIFFIANAE (Key 54) Sect. POLYANTHAE (Keys 49-52)

F. In effect a distinction between the Asiatic section Wallichianae, and the 5. American section Confertas, which in parte have a similarity. Stems pubescent.

Stems dark red. Leaves 1-3 cm. Sect. CONFERTAE (Key 82) Leaves P-10 cm. 1. B. chsycluda (Sect. WALLICHIANAE)

Flower diameter 5-10 mm. Sect. CONFERTAE (Key 82) Flower diameter 14-18 mm. 18. B. v e m l o s a (Sect. WALLICHIANAE)

Sect. CONFERTAE (Key 82)

101. B. kertdana (Sect. ASIATICAE)

Sect. CONFERTBE (Key 82)

Stem pale yellow or yellow-brown.

Stems glabrous. Pedicels pubescent. Pedicels glabrous.

Leaves narrow (1 : 36) , 26-4-5 cm. long. Leaves broader, (1 : 1.25-2.5, rarely 1-3 cm.), much shorter.

G. In effect a distinction between the S. American Sects. Quindiuenses, Goudotiae, and Truxillenseg, and the many-flowered species of the Asiatic Sects. Asiaticae, Tinctoriae, which approach them.

Fruit red. Fruit black, often pruinose.

Sect. TINCTORIAE (Keys 21-26)

Sect. QUINDIUENSES (Key 84)

Sect. GOUDOTULE (Key 85)

Sect. TRKJXILLENSES (Keys 86-89) Sect. ASIATICAE (Key 20)

Flower diameter 14-20 mm. Flower diameter 4-10 (-12) mm.

h v e s with base contracted to articulate petiole. Leaves with base cuneate to decurrent petiole or sessile.

Style nil or very short and indistinct. Style distinct, 0.5-1 (-1.5) 111l11.

Group Septentrionales Schneid., Bull. Herb. Bodss. (2 ) , 8, 264, 1908 Sectionibus (Wallichianae, Asiaticae, Tinctoriae, Angulosae, Franchetianae, Tschono-

skyanae, Canadensee, Sinensea, Integerrimae, Dasystrtchyae, Vulgares, Brachypodae, Polyanthae, She&anae, Chhginae, Hetempodae, Ulicinae) ut in clavibus 2-58, et in descriptionibus 1-319, circumscriptis; ab Australibus ut in clavi 1, distinguenda.

Stem variable. r9pinea nearly always present, generally moderate, sometimes weak, rmely strong, 1-3 (&)-fold, scarcely ever foliaceous. Leaves deciduous or evergreen. InJlorescence solitmy, fascicled, umbellate, subracemose-pseudumbellate, racemose or paniculate. P e d i d h m 1 mm. to 5 cm. long. Bracts generdy shorter than, but some- times aa long as or longer than, their pediceh (the latter casea occurring ody when the pedicels a m very short, 1-4mm.). Fluwer8 yellow (not orange or reddish). ProphylLs present or absent. Se+ in (1-) 2 (-3) series. Petals generdy shorter than, often longer

Page 29: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 29

than, the inner sepals; apex entire or emarginate; base cuneate or clawed, with glands indistinct or conspicuous. Stamens almost invariably shorter than petals; below the anthers edentate (except in one case of doubtful origin) ; apex often not produced, other- wise produced, and truncate, obtuse or apiculate. Ovules 1-15; mostly sessile or sub- sessile; often stipitate with stalk not longer than the ovule; rarely long stipitate. Berries red or black, pruinose or epruinose. Style nearly always insignificant, i.e. absent, or short, measuring up to 0.5 (-0.75) mm.; rarely longer, up to 1 (-1.5) mm., and then separated from the significant styles (2-5 mm. long), of some species of the Australes, a8 indicated in the keys.

HABITAT: Europe and Asia, with only 4 species in Africa (Morocco, Algeria, and from Abyssinia to Kenya), and only 2 species in U.S.A. (Map 1).

The Septentrionales belong, therefore, almost completely to the Northern hemisphere, the exceptions being only three species which just overlap the Equator-one from Java and Sumatra, and two from E. Africa.

The Septentrionales belong, also, almost completely to the East, the two (more notable) exceptions being the N. American species.

KEY 2. GENERAL KEY TO THE GROUP SEPTENTRIONALES Leaves evergreen.

Fruit black, often pruinose. Flowers fascicled. Flowers racemose-umbellate, rarely panicled.

Fruit red or red-purple. Leaves over 2.5 cm. long; fruit 8-12 mm. long, not globose; ovules 2-5.

Leaves 5-15mm. long: fruit either larger with ovules 6-15; or smaller and sub-

Sect. WALLICHIANAE (Key 3) Sect. ASIATICAE (Key 20)

Sect. TINCTORIAE (Key 21)

globose. Fruit 13-18 mm. long, oblong; ovules 6-15. Fruit 5-7 mm., globose; ovules 2-5.

Fruit red, or red-purple, rarely black-purple.

Cf. Sect. ANGULOSAE (Key 27) Cf. Sect. POLYANTHAE (Keys 49-52)

Leaves deciduous.

Inflorescence 1-8 (-12)-fld., and either ovules 6-15; or flowers 12-15 mm. diameter; or fruit 12 -18m. long; or any combination of these characters. Sect. ANOULOSAE (Key 27)

Inflorescence 1-8 (-12)-fld., with ovulo~ 2-5; and flowers and fruit smaller; or inflorescence lO-mny flowered.

Inflorescence 1-8 (-12)-fld. Ovules 1-2 (-3) ; leaves not, or but slightly and openly reticulate.

(Key 38) Sect. FRANCHETIANAE (Keys 36-39) \ Sect. TSCHONOSKYANAE (Key 40)

Sect. POLYANTHAE (Key 49) {

Ovules (2-) 3-5 ; leaves closely reticulate. Inflorescence 10-50 (-more)-fld.

Leaves glabrous. Inflorescence not paniculate.

Stems dark, red or purple; leaves entire.

(Key 41) I Sect. CANADENSRS (Key 42) ISect. SINENSES (Key 43) 1 Sect. INTEGE~RIMAE (Key 44)

Stems dark with leaves spinose; or stems pale yellow. Stems dark; leaves spinose.

Spines foliaceous; fruit globose, lustrous. 250. B. koreana Spines not foliaceous; fruit oblong or dull.

Leaves green, epruinose below. Inflorescence racemose.

Spines mostly 1-5-3 em., stout; leaves 2-3 cm. 233. B. turcomnnica

Page 30: Berberis and Mahonia : A taxonomic revision

30 L. W. A. AHRENDT

KEY 2 (colatin&)

Spinea 8-15 mm., weak; laves 4-10 ~ m . 254. B. pachyacadm (cf. B. x alk.Suthkd)

237. B. franciaei-ferdiniandii hfloreecence compound below.

Petals shorter than inner sepals. Fruit luetroue; ovules 2-4. Fruit dull; ovules 2.

Leaves grey pruinose below.

222. B.cawdmwb

255. B. hnryaaa

238 b. B. jaw8iana vm. kzccomp

Flower diemeter 9-10 mm. ; spines weak. Flower diameter 6-6 mm. ; spines strong.

Petals 88 long 88 inner sepals. Stems not terete; lavea 2-4 om.; fruit bright red.

Racemes 4-7 om.; fruit 8 x 6 mm. ReCemes2--4cm.;fruit 1Ox6mm.

239. B . k a m &

252 b. B. lamma var. hngeaw

253. B.bre&cJbm&& Stems terete; leaves 4-6 cm.; fruit dark red, 10-12 mm.

Stems pale yellow. Leaves entire. Stem subterete. or slightly angled; leaves green below.

Leaves 3-6 cm. ; pedicels 7-10 mm. ; fruit bright red, estylose.

Leaves 1-3 cm.; pedicels 3-7 mm.; fruit dark red, stylose. 248. B.vrimtalis

249. B. concolor 252. B.xluxijkm stems sulmte; leaves grey, p.uinose, below.

Leaves clof3eJly spindose Sect. DASYSTACHYAE (Key 46)) (Key

{Sect. V m m s (Key 47) Leaves more or less pubescent. Sect. BBACEYFODAE (Key 48)

Sect. POLYANTFUE (Key 49)

Sect. SHEBILIFFIANAE (Key 54)

Infiorescence panicdate. Fruit black, mostly pruinose. Inflorescence paniculate. Inflorescence not paniculate. Leaves fairly broad, or shrubs large, generally both.

Sect. CRATAEGINAE (Key 56) Sect. HETEROPODAE (Key 57)

Sect. Ur.IcrN& (Key 68)

Generally speaking the sections comprise very clearly deked sets of species. However, a few species do not conform to the generally s h q demarcation. It is for this reason that these few species, aa well aa the sections, have to be shown in the preceding key, for complete determination of an unknown specimen. If these species are omitted, the key will indicate the more clearly the character of the sections.

r Leaves linear; shrubs very mall.

Section Walllchianae Schneid., BUU. Herb. B&skr (2), 5, 400, 1905 (Map 2) Leaves evergreen, with or without hypoderm, thick or thin, coriaceous, more or less

stiff, more or less narrow and lanceolate, 3-22 em. long, margins spinose; rarely shorter, broader or entire. P ~ T S solitary, or fascicled 2-25. P e d i A 3 mm. to 3 em. Stantens edentate. O& 1-5 (-15). Fruit black, p ~ i n o ~ e or epruinose (white, grey, or blue). Style often nil, aometimes distinct and up to 0.5 mm. in length, occasionaUy to 1 mm., but &s long aa 1.5 mm. only in B. osnzastonii.

Garhwal, Nepal, Sikkim, Bhutan, Tibet, Assam, Manipur, Burma, China (S.W., W, S, N. and S.E., but excluding N.W., N.E. and extreme N.), Fomosa, Java, Sumatra and Philippine Islands.

Page 31: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonis

BY 3, to Sect. Wallichianae

31

Spines absent. 1. Subsect. INSIGNES Spines present. Ovules 6-15. 2. Subsect. HOOKERIANAE ovule^ 1-5.

KEY 3A Mature stems, dark red. Mature stems, not dark red, glabrous or non-glabrous.

Mature stems brown, pubescent, veruculoee. 18. B. verruculosa Mature stems pale yellow, glabrous.

41. B. &levis

Leaves linear oblong, 1 : 12-20. 36. B. insolita Leaves broader, 1 : 2-10 (-12).

Ovule 1, with stalk 3-6 timm as long as the ovule. Ovules semile or with stalk less than twice as long as ovule.

Pedicels 3-4 (-5) mm. Leaves 1-3 cm., very revolute; ovules 2-3. 16. B.?UZWe?&8 Leaves over 3 om. ; ovules solitary.

Leaves revolute, thin, green below; fruit stylose.

Leaves not revolute, thick, below grey, p ~ i n o ~ e ; fruit estylose. 15a. B. grodtmnniana var. @vorama

61. B. micropetah Pedicels 5 mm.-4 cni.

Leaves entire. Leaves not entire.

KEY 3B

Stamens longer than petals. 42 B. dolichostenwn Stamens much shorter than petals. Leaf margins with ( 2 5 ) 30-60 spinules distant 1-2.5 mm.

Flowers fascicled %5. 47. B. f a l h w s a Flowers fascicled 10-25.

stams terete. Leaves thick, elliptic, 1 : 3-3.5. 66. B. durnicoh Leaves thin, lanceolate, 1 : 3 - 5 5 . 46. B. argutn

Leaves thick. 67. B. ,ferdinundii-coburgii Leaves thin. 48. B. a r i a t a t o - a e d t a

Leaf margins with (1-) 3-20 (-25) spinule~, more remote. Leaves more veined below than above. 52. B. silvicola Leaves more veined above than below, or enervate.

Flowers fascicled, 15-30; pedicels 25-30 cm. Cf. 80. B. m i r e i Flowers fascicled 1-10, or pedicels less than 2 cm.

Stems sulcate or angled.

Leaf margins conspicuously revolute, hiding serrations as

Leaves plane, subrevolute, or undulate between serrations viewed from above. K E Y 3c

visible from above. Leaves narrow, 1 : (6) 4-5-7 (-12). K E Y 3u Leaves broader, 1 : 2-4.

Stems terete or subterete. Leaves white below, pruinose. Leaves green below, epruinose.

KEY 3E KEY 3F

Stems sulcate or angled. Leaves below white or grey, pruinose.

Leaves thin; ovules 2-5. Stems very sulcate; spines 1-3 cm. ; leaves fairly

Stems angled; spinea nil, or 2-9 mm.; leaves narrow, 1 : 3-3-5. 27. B. mingetsensi.9

broad, 1 : 1.5-2.5. Ovules 4-5; fruit estylose. 28. B. impedita Ovules 2; fruit stylose. 40. B. petrogena

Page 32: Berberis and Mahonia : A taxonomic revision

32 L, W. A. AHRENDT

KEY 3 (continued)

Leaves thick; ovules solitary; fruit pruinose, stylose.

Leaves grey below, margins with 7-16 spinul~. 85. B.~umwroaa

Leaves white below, margins with 1 4 spinulw. 86. B. &in&

Leaves below green, epruinw. Leaves enervate or slightly veined.

Fruit epruinOs0, stylose. Leaves thick, coarsely dentate; pediaels 18-

Leaves thin, finely serrate; pedicels 5-10 mrn. ; 25 111111. ; ovules 4. 58. B. barandana

ovules solitmy. 43. B. emilii Fruit pruinose.

Leaves thin, 1-3 cm.; ovules 4.

Leaves thick, 4-10 cm. ; ovules solitary. 19. B. aamyana

Stems angled; petals shorter than inner

Stem sulcate; petals longer than inner sepals; fruit stylose. 63. B. be-nnh

91. B. ca&rki Leaves reticulate or distinctly veined. KEY 3Q

sepals; fruit estylose.

KEY 3 A (stem dark red, glabrous) stems tereh.

Leaves 1-5 om., pruinose below; spines 1-2 cm. slender. Leaves narrow, 1 : 8-10; ovules 4-5. Leaves broader, 1 : 3-4; ovules 1-3.

30. B. t r k w n t h u p b a

Outer sepals and emarginate petals both 2.5-3 mm. long, and only half as long as

Outer sepals, inner sepals and entire petals all 5-6 mm. long; leaves thin; O v u l e s inner sepals; leaves thick; ovules mlitary. 72. B. brevisepdu

2-3. 29. B. uenuata

69. B. aargedanu

44. B.deinacantha

15. B. grodtmnnhrm

24. B. bullata 25. B. w d i i

Leaves 5-12 mn. long, epruinose, green below; spines 2-6 cm., stout. Leaves thick, closely reticulate; flowers pale; fruit epruinose. Leaves thin, openly or scarcaly reticulate; flowem deep yellow; fruit p r u i n o ~ .

Stems sulcate or angled; leaves 2-5 cm., slightly veined, below grey prUinose. Pedicels %4 111111. ; ovules solitary ; leaves very revolute. Pedicels 10-25 mm.; ovules 4-5; leaves not or scarcely revolute.

Fruit narrow, 11 x 5 mm., estylose, epruinose. Fruit broad, 8 x 5 mm., stylose, pruinose.

KEY 3 B ( s t e m yellow; h v m entire) Leaves white below, pruinose.

Leaves not or scarcely revolute; ovules 2-3; fruit pruinose. 89. B . p i m 8 a Leaves revolute; ovules 1-2; fruit epruinose.

Spines nil or 4-7 mm. ; leaves broad, 1 : 3-4; fruit estylose. Spines 1-2 om. ; leaves narrow, 1 : 6-7; fruit stylose.

77. B. hoberaapedon 13. B. ~eplicata

Spines 3-8 mm. ; flowers fascicled 3-86. 9. B. eolktii Spines 1-5 cm. ; flowers fascicled 1 W O .

Leaves green below, epruinose.

Leaves enervate; o d e s solitary. Leaves plane; spines 2.5-5 cm., stout. Leaves revolute ; spines 1-2.5 ~ m .

78. B. lev& 10. B.prmcil>ua

havf3S veined; ovules 4. 57. B. xanthxylan

Page 33: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 33

KEY 3c (stems yellow; leave with margins spinose and very revolute)

Leaves permanently white, pruinose below. Stems very sulcate ; flowers fascicled 2-5: fruit estylose. Stems terete.

14. B. taliensis

Flowers fascicled 6-25; fruit stylose. Leaf margin h e l y spinose; leaves light green above; fruit epruinose.

Leaf margin coarsely dentate; leaves lustrous above or fruit pruinose. 13. B. replicatu

Leaves lustrous; petals as long as inner sepals; fruit epruinose.

Leaves dull; petals shorter than inner sepals; fruit pruinose. 12. B. amabilis

88. B. hibberdiana Flowers solitary; fruit estylose. 17. B. candid&

Leaves always, or ultimately, green, epruinose below. Stems sulcate; leaves 1-3 cm., reticulate, somewhat narrow, 1: 3-4; flowers fascicled

Stems terete or angled; leaves 3-10 cm., not reticulate, narrower, 1 : (6) 5-6; flowers 2-3; pedicels 4-7 mm. 16. B. nuntoenab

fascicled 4-25; pedicels 8-22 mm. Leaves veined, $10 cm.; o d e s 1-2. Stems terete; flowers fascicled 5-10; fruit epruinose, estylose. 70. B. recurvata Stems angled; flowers fascicled 15-25; fruit pruinose, stylose. Cf. 10. B. p r . e c i ~

Ovules 4 ; fruit stylose. 11. B. gri$thiam Ovules 2; fruit stylose. 10. B.prwipua

Leaves enervate, 3-6 cm.

KEY 31, ( s t e m pale yellow, glabrous; h u e 8 not revolute, m o w , 1 : (6) 4.5-7 (-12)

Leaves very narrow, 1 : 7-12. Leaves leas narrow, 1 : P 7 .

margins with 1-25 spindes, distant 26-10 mm.) 62. B. mnlanschnenab

Leaves grey, pruinose below ; stems terete ; flowers fascicled 2-6 ; fruit pruinose. Pedicels 1-2 cm.; ovules 4-5; fruit estylw. 34. B. wisleyemis Pedicels 2-3 om.; ovules 2-3; fruit stylose. 87. B. tarone&

Leaves green, epruinose below. Leaves very lustrous above.

Leaves thin; pedicels 6-10 111111. Leaves thick; pedicels 2-3 om.

Fruit stylose; stems angled, or sulcate; leaves thick. Leaves not, or scarcely, lustrous above.

Leaves enervate or subenervate. Ovules 2-3.

Leaves stiff, rigid; fruit pruinose. Leaves flexible; fruit epruinose.

Leaves veined or reticulate; ovules 1-2.

Petals longer than inner sepals. Petals shorter than inner sepals.

Leaves reticulate, rigid. Leaves openly veined, flexible.

Ovules solitary.

Fruit epruinose.

35. B. lubrica 80. B. m i r e i

83. B. soulieana 82. B. atrocarpa 74. B. poluninii

68. B. wallichiana

67. B. ferdinandi-coburgii 79. B. aubcwiacea

Fruit pruinose. Fruit with slight blue bloom. 65. B. willeanrz Fruit with thick white bloom. 64. B. julianae

Fruit estylose. Ovules 1-2; fruit epruinose.

Leaves 1-3 cm. long, with marginal spinules distant 1.5-2.5 mm. 38. B. davidii

3 JOURN. LJNN. S0C.-BOTANY, VOL. LVII

Page 34: Berberis and Mahonia : A taxonomic revision

34 L. W. A. AEBENDT

Leaves 4-10 cm. long, with marginal spindes distant 3-8 111111. stems temte. 78. B. levis stems very sulate.

h v e s thick; pedicels 2-2.5 cm. 73. B..wbpteroclada Leaves thin; pedicels 3-20 111112. Ovulea subsessile; flowers fascicled 1-6; petals shorter than inner sepals.

Flowers solitery, greenish; pedicsls 3-7 1111~1. Flowem fascicled 2-4, yellow, red Without; pedicels 7-20 II11Z1.

37. B . p & w @

39. B.sangzli?tea

Sepals. Cf.41. B.suble&

31. B. veitchii

Ovules long-stipitate; flowers fascicled 10-50; petals longer than inner

ovules 4-5; fruit pruinose. Habit open, arching; shoote red; flowers pale ba. Habit dense; shoots &reen; flowers deep yellow.

Habit erect; leaves ovete to lanmlate, acUmiaate, mostly enervate.

Habit suberect; leaves elliptic, acute, veined. 32. B. g w p a i n i i

33. B.phamra

-Y 3~ ( h u e 8 broad, 1 : 2-4, white, pruhose below; stem8 hreh) Leaves reticulate; fiwit epruinose, stylose. Leaves enervate, or slightly veined; fruit pruinose.

71. B. bh&wnsis

Fruit stylose. 76. B. wangii Fruit estylose.

Petals entire; ovules 2-5. Leaves thin, flexible; petals shorter than inner seM. Leaves thick, rigid; petals longer than inner sepals.

BY 3r (a8 3 ~ , but with hm green below, epruinose) I a v e s enervate or subenervate ; petals emarginate ; fruit pruinoee.

26. B. GOXG 89. B . p i r w s a

88. B. hibberdiana: Petals emarginate; Ovulea solitmy.

ovules 4-5. 32. B. gagnepainii Ovules 1-3.

Leaves dull; pet& shorter than inner sepals; ovules solitary; style 1 nun.

Leaves lustrous; petals longer than inner sepals; ovules 2-3; style nil. 84. B. kmpergiam

90. B. ce?%ti@a

ovules 4-5. 33. B.phawra Leaves reticulate or distinctly veined.

Ovules 1-3. ovules solitmy; fruit pruinose.

Petals longer than inner sepals; leaves thick; spines not stout; fruit stylose.

Petals shorter than inner sepals; leaves thin; spines stout; fruit estylose. 66. B. dumkola

45. B. Val& Ovules 2-3; fruit epruinose or estylose or both.

Leaves thick, not reticulate. Fruit estyloee. Mt stylose.

70. B.reourvata 81. B. schnederkzw

Leaves thip, reticulate. 49,50,51,53,55, distinguished under Subsect. ACUMINATAE

BY 3a (stmw angled or sulcate; leaves moderately broad or m w , 1:2-4, green, epruinose below, reticulate, or distinctly veined)

Spines often absent, if present weak, 3-8 mm.; leaves thin. Leaves broad, 1 : 2.5, 3-6 cm. long; ovules 6 5 . Leaves narrower, 1 :3-4,6-9 cm. long; ovules 2-3.

28. B.&mpedita 54. B. submuminiztu

Page 35: Berberis and Mahonia : A taxonomic revision

Berberis and Mithonia 35

Spines present, not weak, often stout (1-) 1.5-4 cm. long. Fruit stylose.

Fruit epruinose. Flowers solitary; ovules 4. 60. B. alpicola Flowers fascicled ; ovules 2-3.

Leaves thin. 56. B. xanthodada Leaves thick. 59. B. kawakamii

Ovules 2-3. 59a. B. kawaleamii var. f m o s a n a Ovules solitary.

Stems very sulcate; leaves reticulate. 75. B. liophyllu Stems angled; leaves openly or but slightly veined. Cf. 64. B. julianae

Fruit estylose; ovules 4-5. Flowers solitary. 20. B. paraspecta Flowers fascicled 2-15.

Fruit epruninose.

Fruit pruinose.

Leaves thin; stems very sulcate; flowers fascicled 3-5. 23. B. f a l k Leaves thick; stems angled; flowers fascicled 7-15. 57. B. xanthoxybn

Leaves thick, rigid. 57. B. mnthaybn var. aumdra?tica Leaves thin, flexible.

Leaves closely reticulate ; flowers fascicled 6-16. 21. B. manipram Leaves not reticulate; flowers fascicled 2-5. 22. B. buchawnii

Fruit pruinose.

The Sect. Wallichianae is complex, and the largest in the genus, comprising nearly a quarter of it. Some species are known only in flower, or only in fiuit, and a few without either. So, it is desirable to give this direct key baaed on featurea easily discerned, as well as the following key which gives a general guide to the content of the subsections, and the succeeding keys to the species within these. With the aid of both methods it should be possible to identify any species. The second form of analysis also indicates the hybrids.

KEY 4. Guide to the subsections of the Sect. WALLICHIANAE Stems always without spines. Stern always with spines.

O d e s 6-15. O d w 1-5.

Leaves very revolute.

INSIGNES

HOOKERIANAE

REPLICATAE Leaves not, or scarcely, revolute.

Ovules (3-) 6 5 . Leaves thick, rigid or stiff, mostly enervate and more than 2 cm. long.

Leaves thin, flexible (occasionally stiff), but less than 2 cm. long. BARAND AN AE

MANIPURANAE

VERRUCULOSAE

TRJAC ANTHOPHORAE

Leaves reticulate or distinctly veined. Leaves enervate or almost so.

Leaves 1&25 mm. long, broad (1:2-3); flowers solitary. Leaves %12 cm. long, narrow, (1 :6-12); flowers fascicled.

Ovules 1-3. Ovules 2-3.

Leaves thick, enervate or subenervate. Fruit estylose. Fruit stylose.

Leaves reticulate, broad, 1 : ( 2 . 6 ) %-4 (-5). Leaves thin.

PRUINOSAE SOULIEANAE

Page 36: Berberis and Mahonia : A taxonomic revision

36 L. W. A. AERENDT &Y 4 (crmta'7bued)

Leaves enervate, narrow, 1 : (&) 6-10 (-20). Stems very sulcate or angled. stems terete.

Ovulea solitary (rarely 2). Leaves thin, flexible.

stem very sulcate. stems terete.

Leavea reticulate or distinctly veined. Leaves enervate or subenervate.

Leaves thick, rigid or stiff.

Fruit black, epruinose. Fruit pruinose.

Fruit stylose. Fruit estylose.

SANUmruEAE TRIACANTHOPHOR~E

SWLEVES ACUMII~ATAE

EUWALLICHIANAE

LEVIES

SOULIEANAE PRUINOSAE

Subsect. Insignes Schneid., BuU. Herb. Bokzier (2), 5, 401, 1905 Stems terete. Spine8 absent. Leave.9 evergreen, 6-22 x 1-5 om. F k r 8 10-14 mm.

diam. Pet& shorter than inner sepals. Anthers dentate. ovzlles (3-) 4-5 (-7). Fruit black, shortly stylose; rarely pruinose.

Sikkim, Bhutan, Assam, S.E. Tibet, N. Burma.

KEY 5, to Subsect. Insignes (Map 6) Sterne pubeeoent ; stamens apiculate. Stems glabrous; staznena obtuse or truncate.

I. B. daaydadu

Leaves blow green, epruinose; fruit black, epruinose. Leaves with hypodem; petals em-te; ovule8 4; f i t oblong, 9 x 6 mm.

Leaves without hypoderm; petals entire; ovulea 6-7; fruit globose, 5 mm. 2. B.in&gnW

3. B. incraeeata; Leaves white below, pruinoae; fruit pruinose white. 4. B . h y p k r i m

(1) Berberis dasyclada Ahrendt, J . Bot., M . 79, 12, (Suppl.), 1941 Sterne dark red. h u e s 4-9 (-11-5) x 1-2-2-6 (-3.5) cm., with hypoderm; narrowly

elliptic, both ends subacuminate; margins with 12-18 spinose (1-2 mm.) serrations (2-3 mm.), distant (3-) 5-20 mm.; above dull grey green with few lateral veins; below paler, yellow-green, sublustrous, with veins slightly elevated. Fbwers fascicled 3-6 (-8). Pedieels slender, 10-12 mm.; bracts lanceolate, acullljll&te, 3 - 5 4 x 1 mm. Prophylls c. 4 x 1.25 mm., lanceolate, acuminate. Outer sepals 5.5 x 3 mm., elliptic, obtuse. Medaan and inner 8epa.k 7 x 6 mm. ; obovate-elliptic. Petals 4 5 x 4 nun., obovate; base clawed with separate lanceolate glands, 1 x 0-35 mm. ; apex entire. Stamens 4-25 mm., filaments pubescent, apex produced and apiculate. Ovules 34, with stipes equal to or slightly longer than themselves. Be& unknown.

Assam Himalaya: Poshing La, 10 ,W ft., 25 May 1938, Kingdon-Ward 13700 (Type, BM). 'A low growing gregarious shrub forming denae thickets not above 2 ft. high; leaves dark gmen ; flowers golden yellow, fragrant.' Apparently not in cultivation, though seed was collected.

(2) Berberis insignis Hook. f. et Thorns., Fl. Ind., 226,1855

tions (2-4 mm.) ; sparsely veined. Fbwer8 fascicled 10-25. Pedkwh thick, 5-10 mm. Stenzs dark red. Leaves 8-16 x 2 4 om. ; margins with 10-20 spinose (1-2) mm. serra-

Page 37: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 37

G Y 5A Pedicels 5-15 mm. Stems dark, red or purple. Shrub 5-10 ft. ; flowers fascicled 8-25; petals emarginate.

Leaves coarsely aristate-dentate. 2a. B. insignis vm. insignis Leaves finely spinose-serrate. 2 b. var. elegantifolia

2 c . var. zelaica Shrub 2-4 ft. ; flowers fascicled 2-8; petals entire. Stems pale, yellow or grey.

Leaves thick; flowers fascicled 9-25; pedicels stout, 5-10 mm. ; petals emarginate. 2d. var.tongEoensis

Leaves thinner; flowers fascicled 5-9 ; pedimls slender, 9-14 mm. ; petals entire. 2e . var. slaergaomnais

Pedicels 3-4 cm. 2f . var. gouldii

(2a) Berberis insignis var. insignis Sikkim: Hooker, 7000-10,OOO ft., 23 May 1849 (Type, K). Bhutan: 8000-9000ft., Gri5th, 1838, (K); Dengchung, Khoma Chu, 2 May 1949,

8500 ft., Ludlow, Sherriff & Hicks 18807 (BM); Gamble, 1875 (K). Not in cultivation.

(2b) Berberis insignis var. elegantifolia Ahrendt, var.nov., foliis graciliter spinoso- serrulatis

Sikkim: Yaldong, 7 Dec. 1849, Hooker (Type, K). Bhutan: 1838, Griffith 658 (K). Not in cultivation.

(2c) Berberis insignis var. zelaica Ahrendt, J. Bot., Lond. 79, (Suppl.), 9, 1942 Spinas mostly absent, rarely 3-fld.) weak or subfoliaceous. Flowers fascicled 2-6 (-8) ;

pediceh thick, 6-9 mm. ; bracts 2.5-3 mm. Prophylls 4 x 2 mm. Outer sepals 6.5 x 4 mm. Petals entire. Ovules 3. Otherwise as in (2d).

Assam Himalaya: Zela, common at 9000-10,OOO ft., 1935, Kingdon-Ward 11644, (Type, BM) . Not cultivated.

( 2 4 Berberis insignis var. tongloensis Schneid., New Flora and Silva, 10,256, 1938;

Leaves 8-22 x 2-5 cm., contracted to petiole 2-4 mm. ; elliptic-lanceolate, acuminate ; margins with 10-20 spinose (1-2-5 mm.) serrations (1-2-5 mm.), distant 5-9 mm. ; lateral and marginal veins and veinlets visible either side, not reticulate; slightly lustrous deep bright green; below much paler yellow-green. Flcnvers fascicled (8-) 10-25. Pedicels stout, 5-10 (-14) mm. Prophylls c. 2 x 2 mm. Outer sepals 4 x 3 mm. Median sepals 7 x 4 mm. Inner sepals 8 x 5 mm. Petals 7.5 x 4 mm., obovate; base slightly clawed with ovate, obtuse glands, 1-5 x 0.8 mm. : apex acutely emarginate with acute lobes. Stamens 4 mm., produced, apex rounded-truncate. O& 4. Berries black, epruinose, oblong to oblong-ovoid, 8-9 x 5-6 mm., style short.

Mitt. Dtsch. Dendr. Ges. 55, 45, 1942. (Fig. 22)

Sikkim: 10,OOO ft., 1849, Hooker (Type, K), 1879, Gamble (K). Nepal: 1849, Hooker. Bhutan: 8000-9000 ft., Mara Chu valley, 28 May 1937, Ludlow & Sherriff 3127 (BM);

Cultivated: erect, tender, 6-9 ft. high, valuable for its foliage. Tunle La near Kinga Rapden, 9 Apr. 1949, Ludlow, Sherriff & Hicks 18659 (BM).

(2e) Berberis insignis var. shergaonensis Ahrendt, J. Bot., Lond. (Suppl.), 79,8, 1941 Internodes 3-8 cm. h w e s with 20-30 spinose (0.5-1 mm.) serrations (0.5 mm.) each

side. Outer sepals 3 x 2 mm. Median sep& 5 x 3 mm. Inner sepals 6 x 4 mm. Petals 4 x 2 mm. Otherwise as in ( 2 4 .

Page 38: Berberis and Mahonia : A taxonomic revision

38 L. W. A. AHRENDT

shrub in the forest amongst much undergrowth. Not cultivated. h a m Himalaya: 8000-9000 ft., 1938, Kingdon-Ward 13617 (Type, BM). An erect

(2f) Berberis insignis var. gouldii Ahrendt, var.nov., pedicellis elongatis I d 4 c. 8 cm. Leave.s c. 7.5 x 2-5 cm. ; margins with 10-15 spinose (1 mm.) serra-

tions (1.5-3 mm.) distant 5-8 mm.; very lustrous above, dull deep green below. Fkwera fascicled 3-5. Pediwls very slender, 3-4 cm.

Bhutan: 7000-11,OoO ft., 1938, Gould 243 (Type, K). Not cultivated.

(3) Berberis incrassata Ahrendt, Card. Chron. (3) 185, 371, 1939: J . Bot., Lond. (Suppl.), 79, 10, 1941; Mulligan, Card. Chron. (3) 108, 41, 1940 (fig. 19); Schneid., Mitt. Dtsch. Dendr. Gee. 55, 45, t. 6, 1942

Stem dark red, glabrous. Internodes 3-5 cm. Leuw 7-15 x 1.5-3 cm.; lanceolate to elliptic-lanceolate; apex acuminate; margins with 12-24 spinose (1-5-3 mm.) serrations (16-3 mm.) distant 5-8 mm.; lateral veins and few veinlets visible on both sides, not reticulate; dull grey-green; below slightly lustrous paler yellow-green. F k r s faacicled 15-30. Pedicels 10-15 mm., very slender a t base thickening to 1.5 xnm. at apex. ODcter se@ 4 x 3 mm., ovate, acute. Hedkn sepals 6 x 5 mm., ovate-elliptic, obtuse. Inner se@ 7 x 6 mm., obovate. Petals 6 x 4 mm., obovate, entire; base cuneate with glands 0.8 xO.3 mm. Stamem 4 nun. O& (4-) 5-7. Berries black, epruinose, globose, 4 - 6 5 mm., with short style.

KEY 5% Leaves narrow, 1 : 5-6; flowers fascicled 15-30. Leaves broader, 1 : 3.6-4-5; flowers fascicled 4-8.

3a. B. immecNcta, var. incmesata 3b. var. bv.cahumagunaia

(34 Berberis incrassata var. incrassata N. Upper Burma: 7000ft., 1931, Kingdon-Ward 9358 (Type, BM). ‘A small shrub,

not abundant, conked to limestone’; 1931, Tibet frontier, Kingdon-Ward 10148; Nam Tamai valley, 1937, Kingdon-Ward 13447 (BM).

Cultivated: fl. 4 Apr. 1939, fr. 2 Dec. 1956 (in Oxfordshire). Shrub 2-3 ft. high, with long arching stems. No. 10148 is described aa reaching 5-5 ft., and no. 13447, aa up to 10 ft. or more, but, these numbers do not appear to be cultivated.

(3b) Berberis incrassata var. bucahwangensis Abrendt, J. Bot., Lond. 79, (Suppl.),

Leaves to 12.5 x 3.5, or 13 x 3, or 14 x 3.5 cm. Peclkds 17-24 mm., thickened aa in

S.E. Tibet: Rov. Z q d , c. 5OOOft., 1938, Yu 21318 (Type, E). Cultivated: from seed sent by Kingdon-Ward in 1939, vide Osborn, Card. Chron. 259,

11, 1941.

type. Fruit globose, 5-6 xnm.

fig. 126, 24 June 1944.

(4) Berberis hypokerina Airy-Shaw, Kew Bdl . 1930,208,

S t e m dark red, sparsely and h e l y verruculose. Internodes (3-)5-?cm. Leavap 8-12 x 3-6.5 cm., without hypoderm; elliptic to oblong-elliptic; margins 6-10 (-15) spinose (2-3 mm.)-dentate (3-7 mm.), the teeth distant 10-18 mm.; lateral veins in- distinct; pale grey-green; below white, pruinose. F k r s H.C.C. 3/1, fascicled 6-12. P e d k l s 7-11 mm. Prophylls concolorous, triangular, 2 x 1.5 mm. Outer sepals 2.5 x 2 mm., ovate, acute. Median s e e 4.5 x 4 nun., ovate, obtuse. Inner sepals 7 x 4.5 mm., obovate. Petals 5-5 x 3 mm. acutely emarginab, with acute lobea; base

Page 39: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 39

cuneate with separate non-marginal glands, 0.7 x 0-4 mm. Stamem 2.5 mm. Ovules 4. Berries black, pruinose white, ellipsoid, 7 x 4 mm., with short style.

N.E. Upper Burma: 8000-10,OOO ft., 1926, Kingdon-Ward (Type, K); 1931, Kingdon- Ward 10151, abundant on granite cliffs; 1937, Kingdon-Ward 13479. In cultivation.

Subsect. Hookerianae Ahrendt, J. Bot., Lond. 79, (Suppl.), 12, 1941 Stem glabrous, mostly angled. Leaves evergreen, lustrous above, reticulate and

pruinose white below (unless otherwise recorded). lrlowers fascicled 1-6, diameter (I-) 1.5-2.5 cm. Ovules 6-15. Berries black with blue bloom (unless otherwise recorded).

Garhwal, Sikkim, Bhutan, S.E. Tibet.

KEY 6, to Subsect. HOOKEWAE (Map 5) Stems distinctly verruculose; flowers solitary; petals entire; fruit 8-10 mm. long.

Stems not, or scarcely, and even then partially verruculose; fruit (lo-) 12-18 mm. long;

Leaves spinose, moderately narrow, 1 : 3-4; pedicels 15-35 mm. ; inner sepals

Hybrids, 5e, 6a

flowers fascicled or petals emarginate.

9-12 mm. ; stamens truncate; fruit estylose. Leaves reticulate, not revolute; flowers fascicled; petals entire.

Leavea without hypoderm; ovules (5-) 6-9; mature stems yellow.

Leaves with hypoderm; ovules 12-15; mature stems red brown. 5. B. hookeri

6. B. culliantlm Leaves enervate, revolute; flowers solitary; ovules 9-12. 7. B. chrysosphaera

8. B. omnaatmii Leaves entire, narrow, 1 : 5-7 ; pedicels 3-5 mm. ; stamem apiculate; fruit stylose, with

long style (1.5 mm.).

(5) Berberis hookeri Lemaire, IUmtr. Hort. 6. 207, 1859; syn. ‘B. d i c h i a m ’ ,

Stem, mature yellow, not verruculose, angled or sulcate. Spines 3-fld., 10-25 mm., sulcate. Leaves 3-6 cm., x 8-18 mm. ; oblong-elliptic; margins with 7-15 spinose (16-2.5 mm.) serrations (1-2 mm.), distant 3-5 mm. ; reticulate or subreticulate, lustrous deep green, below pruinose white. Flowers fascicled 3-6. Pedicels 15-25 mm., stout. Prophylls red, ovate, acute, 2.5 x 1.5 mm. Outer sepals 6 x 4 mm., oblong-ovate. Median sepals 8 x 5 mm., oblong-elliptic. Inner sepals 10 x 7 mm., obovate. Petals 7 x 6 mm., obovate, entire; base cuneate, glands 1 x 0.6 mm. O w u h (5) 6-9. Berries 12-15 x 6- 8 mm., oblong, estylose, epruinose or almost so.

Hook. f., Bot. Mag., 78, t. 4656, 1852, non DC.

KEY 6a Leaves white, pruinose below. Leaves epruinose, subconcolorous green below. Leaves coarsely spinom-dentate, narrow, 1 : 3-5.

Fruit 12-15 mm. long. Fruit 9-11 mm. long.

Leaves finely spinulose-serrulate, broader.

5a. B. hookeri var. hookeri

5b. var. w i d i n 5c. var. microcarpa 5d. var. platyphylla

(5a) Berberis hookeri var. hookeri Sikkim: 1848, Hooker 255, (Type, K). Bhutan: 1838, G a t h 1740. Cultivated, but not a t all common, the variety 5 b being more frequent.

Page 40: Berberis and Mahonia : A taxonomic revision

40 L. W. A. ~ E N D T

(5b) Berberis hookeri var. viridis Schneid., Bull. Herb. BoisSieP (2), 8, 197, 1908 Bhutan: 1838, GrifEth; 11,500 ft., 1 Oct. 1949, Ludlow, S h e d & Hicks 17374 (BM). aeSam: ridge, w e t of Dirang Dzong, SOOOft., 10 Oct. 1938, Kingdon-Ward 14330

Cultivated: fl. June 1936; fi. 21 Oct. 1940,14 Oct. 1943. Very common, and generally (BM). A stout bush 2-3 ft. tall, on rocky, S. facing, shrub and tree-covered slope.

distributed as B. h k e r i , ‘ B. dichiana ’, or ‘ B. jamamii ’.

(6c) Berberis hookeri var. microcarpa Ahrendt, varaov., fructibus minoribua Sikkim: 25 Oct. 1874, Clarke 25561 (Type, K). Cultivated: at Kew, fl., fr., 1938.

(54 Berberis hookeri var. platyphylla Ahrendt, nom.nov.: syn. B. Zoawichiana var. ldifolia Hook. f. et Thorns., Fl. Ind. p. 226, 1855, non B. hookeri var. latifolia Bean, Trees &.Shrubs, 1,243, 1914

Sikkim: Hooker 40 (Type, K); 10,50Oft., 10 Oct. 1870, Clarke 12768; Nov. 1874, Gamble 1030; Nov. 1879, Gamble 7443 (all K).

Not cultivated: to be distinguished from Bean’s plant cited above, very common in gardens under this name, sometimes even called ‘B . knightii’, but properly named B. mnipuranu. This comes from a merent region, has flowers fascicled 6-15, emarginate petals and 3-5 ovules.

The hybrids placed in Key 6 belong here and may be distinguished as follows.

KEY 6 b Leaves pruinose white below; pedicels 15-18 nun. Leaves pruinose, grey-green below; pedicels 8-14 111111.

6 a. B. x ~lristoleneie 5e. B. x intepsita

(5e) Berberis x interposita Ahrendt, Gard. Illustr., 64, 1084, July 1947 Leavu 20-30 x 8-13 mm., elliptic; margins with 24-spinose (0-7-1-2 mm.) serrations

(0-5 mm.), distant 6 7 mm.; a few veins above, subenervate below. Flowers solitary. Pedicels stout. Outer sepals 5 x 3 mm. Median se@ 7 x 5 mm. Inner sepals 8 x 6 mm. Petals 7 x 5 mm., entire; base slightly clawed, g h d s 1-1 x 0.9 mm. Berries 9 x 4.5 mm.

Cultivated: fl. 13 June 1939, fr. 21 Dec. 1944 (Type, 0). Probably B. hook& virddis x B. ver rucha .

(6a) Berberis x bristolensis, Ahrendt, 1.c. Leavu to 30 (-40) x 14 (-18) mm.; elliptic; margins with 1&14 spinose (1-2 mm.)

serrulations. Prophyls 3 x 2.5 mm. Outer sepals 5-5 x 5 mm. Median sepals 8 x 7 mm. Inner s e p k 11 x 10 mm. Petals 8 x 8 mm., base clawed with widely separated sub- marginal glands, 1-6 x 0.6 mm., widely separated. Berries 9 x 4.5 mm., oblong-ovoid.

Cultivated: fl. May 1939,5 May 1944, fr. 21 Dec. 1944 (Type, 0). Origin in a garden near Bristol, as a seedling. Probably B. caUknthax B. vemuculosa.

(6) Berberis calliantha Mulligan, Gard. Chron. (3), 97,394, 1935. (Fig. 20) Intemzodes 2.54 cm. Spines 1-2 em., sulcate. Leavu 2-6 x 1-2-2 cm.; oblong-elliptic;

margins with 10-20 spinose (1.5-2.5 mm.) serrations (1*54mm.), distant 3-5mm.; reticulate; lustrous deep green; below pruinose white, less veined. Flowers fascicled (1-) 2-3. Pedicels slender, 2 4 cm., bracts 4-5 mm. Outer sepals (H.C.C. 3/1), 7 x 6 mm. Median se& 1Ox8mm. Inner sepals 12xlOmm. Petals 8x6.5mm., obovate, entire; basal glands 1 x 0.4 mm., ovate, separate, marginal, situated towards apex of claw. Otmkz 12-16. Be- 11-14 x 6-9 mm., ovoid.

Page 41: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 41

S.E. Tibet, Prov. Pemako: 8OOO-9oOO ft., 21 Nov. 1924, Kingdon-Ward 6308 (Type, K); 2 May 1947, Ludlow, Sherriff & Elliot 13605 (BM). Both collections are from Pemakochung, Tsangpo gorge; the type is described as a scrub plant 1 ft. high growing in maases with RJuAodendron venator; the second collection is from a bush 2-3 ft. high crawling over the ground in shade in the forest.

Cultivated: from the type, fl. 11 May 1944, fi. 28 Nov. 1944, a compact bush about 2-3 ft. high with sturdy stems aa in B. hokeri, though of different colour. Photographs: Card. Chron., (3), 97, 390, 391, figs. 168, 169, 1935.

(7) Berberis chrysosphaera Mulligan, Kew Bull. 1940, 77; J . Roy. Hort. SOC. 65,

Stem mature dark red, very sparsely verruculose. Internodes 1.5-3 cm. Spines 3-fld., 1-2 cm., sulcate. L m v ~ with hypoderm, 15-40 x 6 1 0 mm.; elliptic to ovate; margins very revolute, with 5-12 spinose (1-2 mm.) serrations (1 mm.), distant 2-5-3 mm.; dark very lustrous green, subenervate, or with, at most, very few lateral veins and veinlets: below pruinose, white, enervate, closely, finely papillose. Flowers solitary. Pedicels 18-25 mm. Outer sepals 4 x 3 mm., ovate, acute. Median sepals 7 x 6 mm., obovate- elliptic. Inner sepals 10 x 8 mm. Petals 8 x 6 mm., obovate, slightly emarginate; basal glands oalong. Stawns slightly produced. Ovules 9-12. Berries 10 x 6 mm., ovoid.

S.E. Tibet, Prov. Zayul: Rongto valley, granite cliffs, one single clump, fr. Dee. 1933, Kingdon-Ward 11036 (Type, BM).

Cultivated: from type, and only, introduction: fl. 13 May 1942, fr. 26 Dee. 1943. A compact bush, 1-2 ft., with very handsome foliage.

(8) Berberis osmastonii Dunn, Kew Bull. 1920, 335; Ahrendt in J . Roy. Hort. SOC.

Stem terete or subterete : shoots puberulous ; mature yellow, lustrous, glabrous. Intern& 5-10 mm. Spines 3-fld., slender, 1-2 cm. Leaves 10-20 x 2-3 mm. ; linear- oblong, or very narrowly elliptic; with hypoderm; margins entire, very revolute; ener- vate ; above lustrous ; below pruinose white, papillose. Flowers solitary. Pedicels 3-5 mm.; bracts 1 mm. Prophylls 3 x 0.4 mm., linear-lanceolate, acuminate. Outer sepals 3.5 x 1 mm., oblong-lanceolate. Inner sepals 7 x 3.5 mm., narrowly obovate. Petals 6 x 2.5 mm., similar, entire; base slightly clawed attenuate, with glands separate, orbicular, 0-4 mm. Stamens 5 mm., produced, apiculate. Ovules 6-8. Berries 10 x 6 mm., ovoid, excluding style 1.5 mm.

281 figs. 77, 78, 1940

74, 406, fig. 146, 1949

Garhwal: 9500 ft., 1915, Osmaston 225; 1918, Osmaston 919, 894 (K). Cultivated: from collector’s type seed, by Mr Oliver E. P. Wyatt, at Maidwell Hall,

Northampton, 20 Sept. 1947. This appears to be the only plant in cultivation of this species which is remarkable geographically for being the westernmost of the section Wallichianae; and botanically for the very long style, generally found only in S. American species; amongst these, it tends generally to be a link with the subtropical section Virgatae; in its leaves it is reminiscent of the subantarctic species B. empetriiolia, which has estylose berries. Its very narrow flower components are also remarkable. Summing up all its characters, it is a species much more distinct and separate from all others than is generally found in the genus. It is a subprostrate shrub, rising to about 8 in.

Subsect. Replicatae Ahrendt, subsect. nov. foliis multo revolutis Stems either terete or subterete, or conspicuously sulcate; mature yellow (except only

in grodtmnniana, purple), always glabrous. Intemzodes (1-) 1-5-2.5 cm. (though to 34, in grodtmunniana and amubilis). Leaves enervate or subenervate, very revolute. Flowers fascicled 3-12 (-25). Ovules 1 4 .

Bhutan, Assam, Burma, S.E. Tibet, W. Yunnan, S.W. Szechuan, Formosa.

Page 42: Berberis and Mahonia : A taxonomic revision

42 L. W. A. AHEENDT

KEY 7, to Subsect. Replicatae (Mipa 5 and 6) Flowers fascicled.

Stems tereta or subterete, yellow; fruit epruinose, or almost; leaves with hypodem. Leaves below, always, or finally, green, epruinose. Leaves always entire, broad, 1 : 2-4. Leaves with spinules, narrow, 1 : 5-8.

9. B. wUetii

Leaves lustrous; flowers fascicled 6-15; ovules 1 (-2); fruit stylose.

Leaves dull; flowers fascicled 6 8 ; ovules 3-4; fruit stylose. 10. B . p w i p a

11. B.&t?ithiana Leaves below permanently white, pruinose; o d e s 2; fruit stylose.

Leaves lustrous; flowers fascicled 12-25; petals longer than inner sep&.

Laves dull; flowers fascicled 5-12; pet& shorter than inner sepal8. 12. B. amubilb

13. B. replicuta Stem very sulcate. Leaves below pruinose white, with hypoderm; pedicels 12-20 Hm. ; fruit estylose.

Leaves below epruinose, green; pedicel8 3-6 mm.; fruit stylose. 14. B. talieneiS

Leaves with hypoderm; margins with 7-15 spinules; flowers fascicled 5-10;

Leaves without hypoderm; margins with 1-5 spinulea; flowers fascicled 2-3; ovules solitary; fruit epnrinoss.

ovules 2-3; fruit pruinw.

15. B.grodtwnnh

16. B.mntoen& Cf. 17. B. c(lndicIuLQL Flowers solitary.

(9) Berberis collettii Schneid., Ost. Bot. 2. 67, 140, 1918; syn. B. WaUichiuw var.,

Spim weak, 3-fld., 3-8 mm. terete. Leaves thick, stiff, 2-6 cm. x 5-15 mm. ; namowly obovate; apex subobtuse; aristate, gradually cuneate, sessile, entire, slightly lustrous; below epapillose, paler dull green. Pedimh 12-20 mm., fascicled 3-6. F k r s and Fruit unknown.

Upper Burma: Shan Hills, W f t . , May 1888, Collett 787 (Type, R), and possibly also GrifEth 787 of Jan. 1838 from Bhutan. According to the authors, B. w W i i is also to be found in Nepal, Bhutan and Khasia Rills; but they may be referring to the fol- lowing two species which they may have included with their own.

(10) Berberis praecipua Schneid., Fedde Rep. Sp. Nov. 46, 248, 1939; blitt. Dtsch.

Stem not verruculose. Leaves 15 x 3 mm., to 40 x 7-8 mm. : oblong-elliptic, to elliptic- lanceolate; acute, mucronate; margins 3-9 spinose (1-1-5 mm.)-serrate (0.5 m.), dietant (2.5-) 4 4 mm., slightly lustrous; below paler green. F k r s fascicled (3-) 5- 10 (15). PediceZa (7-) 10-15 (-18) mm. O& 1 (-2). Fruit 7-8 x 3-5-4 mm., with short style, almost epruinose.

KEY 7a

Collett and Hemsley, J. Linn. Soc. (Bot.), 28, 17, 1890

Dendr. Ges., 55, 14, 1942

Leaves 1-4 cm. ; flowers fascicled 3-15. Leaves 5-7 cm. ; flowers fascicled 10-25.

10a. B. padpa, var. paeeipua lob. var. major

(10a) Berberis praecipua var. praecipua

valley 10,OOO ft., 2 Oct. 1949, Ludlow, S h e d BE Hicks 17380 (BM).

cultivated.

Bhutan: 13 Aug., 1914, Phajudin Jimpu, Cooper 2524, (Type, E) ; Gyasa Dzong, MoChu

Assam: 7000-8000 ft.. 1938, Kingdon-Ward 13688, A compact bush of 3-5 ft., not

Page 43: Berberis and Mahonia : A taxonomic revision

Berberis and Mrthonia 43

(lob) Berberis praecipua var. major Ahrendt, var.nov. ; foliis fasciculisque majoribus Yunnan: Yunnan-sen Maire 1998 (Type, K). Not cultivated.

(11) Berberis griffithianaschneid., Bull. Herb. Boissier (Z), 5,403,1905; s p . B. asiatica Griffith non Roxb. ; syn. B. replieata var. dispar Ahrendt, J. Bot., h n d . 79, (Suppl.), 20, 1941.

Stem shoots slightly sulcate ; mature terete, slightly lustrous. Leaves 25-65 x 5-8 mm. ; elliptic-lanceolate ; acute, mucronate ; cuneate, sessile ; margins with 3-6 appressed spinules (0.5 mm.), subserrulitte, distant 3 4 mm.; dull grey-green; below, at first slightly whitish, finally lustrous paler green. Flowers fascicled 4-8; pedicels 16-22 mm. Prophylls 2 x 1 mm., acuminate. Outer sepals 5 x 3.5 mm., ovate, acute. Inner sepals 7.5 x 6 mm., elliptic. Petals 7-5 x 4 mm., emarginate; glands 2 x 0.8 mm. Ovules 3 4 . Stamens 5 mm., produced obtuse. Fruit epruinose, estylose.

Bhutan: 1838, Griffith 125 (Type, Vienna); Grif6th 1742 (K); 7500 ft., 1936, Ludlow & Sherriff 1187 (BM); 6500 ft., Ludlow & S h e d 1221 (BM).

S.E. Tibet, Prov. Monyul: 7500 ft., 1936, Ludlow & Sherriff 1187 (BM). Assam: Khasia, 1838 GriEith 858. Not cultivated. Shrub 4-6 ft. Depicted in Griffith, Icon. P1. As., t. 648. The name B. asiatica Griffith

is given in Posth. Pap. J. Travels, 211, 216, 1847, and Itin. Notes, 11, 122, no. 38, 1848. On p. 122, Griffith refers to this species, and also no. 381 from Khegumpa with deflexed branchlets. On p. 211 of the Travels he refers t o it at 6900 ft.; on pp. 245 and 247 a t Sanah a t 8000 ft. : on p. 266 at Jaiwa; on p. 268 a t Tongsa. In the Khasia Flora, Itin. Notes, p. 55, he refers to his asiatiea no. 858, recording another specimen 4-5 ft. high. The herb. sheet of no. 124 records oblong blaok fruit.

(12) Berberis amabilis Schneid., F& Rep. Sp. Nov. 46, 257, 1939 Leaves 4-8 x 0.7-1.2 cm. ; oblong-elliptic to oblong-lanceolate; petiole 1-2mm.;

margins with 4-9 spinose (2-3) mm. serrations (1.5-2.5 mm.), distant P 9 (-12) mm.; acute ; lustrous ; below pruinose white. Flowers fascicled 10-25. Pedicels slender, 12-18 mm. Prophylls 2 x 1 mm. Outer sepals 5 x 4 mm., ovate. Inner sepals 7 x 4-5 mm., oblong-obovate. Petals 7 x 4 mm., obovate, emarginate; base clawed; glands 1-2 x 0.7 mm. Stamens 5 mm., produced, apiculate. O& 2, sessile. Fruit black, epruinose, ellipsoid, 8 x 5-6 mm., style short.

N. Upper Burma: Tibet frontier, 7000-8000ft., 27 Mar. 1931, Kingdon-Ward 9324 (Type, BM); Kingdon-Ward 9299 (BM).

Yunnan: 10,500 ft., 1938, Yu 17208. A common shrub, 3-5 ft. Not cultivated.

(13) Berberis replicata W. W. Smith, Not. Rot. G a d Edinb. 11, 200, 1920; Stapf, Bot. Mag. 151, t. 9076, 1925

Stems somewhat thin, terete, slightly, distinctly verruculose. Spines sulcate. Leaves 20-35 (-45) x 3-5 (-8) mm. ; oblong-elliptic to oblong-lanceolate ; petiole 1-2 mm. ; margins entire, or with 1-3 (-6) spinose (1-1.5 mm.) serrations (0-5 mm.), distant 6 7 mm.; dull, light, green; below pruinose white. Flowers fascicled (&) 8-12 (-15) (6-) 8-12 mm.; bracts 1.5 mm. Prophylls red, 2 x 1.5 mm., acuminate. Outer sepals 4 x 3 mm., subacute. Inner sepals 5.5 x 3.5 mm., obovate. Petals 5 x 3.75 mm., obovate acutely emarginate with 2 rounded lobes; base subcuneate; glands 0.5 x 0.3 mm. Stamem produced, crenate-truncate. Ovules 2, sessile. Berries 6-8 x 3-5 mm., oblong; style scarcely 0.25 mm. ; epruinose or almost so.

W. Yunnan: open situations around Tengyueh, 5000ft., 1919, Forrest 7785 (Type, E, K). Also Forrest 8782, 9457, 16030, 24682.

Page 44: Berberis and Mahonia : A taxonomic revision

44 L. W. A. AHRENDT Cultivated: fr. 1 Nov. 1938 (F. 16030); fl. 20 May 1943, fr. Dec. 1944 (F. 24682). A

slightly tender shrub with arching branches, often of sprawling habit, and somewhat stoloniferous, valuable for its graceful foliage, and its particularly pretty appearance when in f d flower.

(14) Berberis taliensis Schneid., F& Rep. Sp, NOW., 46,252,1939 Stems thick, very sulcate, distinctly concolorous verruculose. Spines 5-7 (-12) m.,

sulcate. Leuve.9 25-40 x 4-8 (-10) mm. ; elliptic, cuneate, sessile, acute, mucronate; margins occasionally entire, generally 1-3 (-6) spinom (1-1-5 mm.)-subsedate, distrtnt 5-12 mm.; lustrous dark green; below pruinose, white. Flowers fascicled 2-5. Pedhk (1%) 15-20 mm. Prophy& 3 x 2 mm., ovate, acute. Outer s e e 4.5 x 3 m., oblong- elliptic. Inner 8e+ 6 x 3.5 mm. Pet& 6-5 x 4 mm., obovate, emarginate; base cuneate, glands 1 x 0-4 mm. Stamns 3.5 mm., not produced. O d e s 4, sessile. Berrim 11 x 5 mm., oblong, estylose, pruinose blue.

N.W. Yunnttn: Chien-chum-Mekong divide, side vdeys of the Chien-chuan, fr. Aug. 1922, Forrest 21968 (Type, K); July 1923, Format 23449 (K); E. flank of Tali shan, 9OOO ft., May 1921, Forrest 19417 (K).

Cultivated: fl. May 1938; fr. Nov. 1938, from type collection. A small shrub naturally growing in ledges of cliffs and rocky slopes in ravines. In cultivation, a dense and com- pact bush, scarcely over 2ft. high, though more across, with very stiff habit, thick, sulcate, erect stems, dark lustrous leaves, few greenish flowers with long stalks, and large pruinose berries. In contrast, the more familiar B. replicata has a sprawling habit with slender, arching, terete stems, dull, light green leaves, more numerous bright yellow flowers, and small black fruits without bloom.

(15) Berberis grodtmannia Schneid., Ust. Bot. 2. 32, 1918 Shrub erect, to 4 ft. Stem thick, very sulcate. A’pines plane. Leaves 30-50 x 5-9 mm. ;

lanceohte, cuneate, Sessile, acute, mucronate; margins 7-12(-15)-spinose (1-1.5 mm.)- subsemdate, distant 2.5-4mm.; deep green; paler green below. Flowers fascicled 5-10. Pedk& 3-4 mm. Prophy& 1 x 0-5 mm., acuminate. Outer sepals 2 x 1 mm., ovate, acute. Inner sepal8 5.5 x 3 mm., ovate-elliptic. Pet& 4.5 x 5 x 2-5-3 mm., obovate, emargimte. Stamem 3 mm., not produced. Ovules solitary, sessile. Berries immature, 5.5 x 3 mm., epruinose, shortly stylose.

KEY 7 b Stems when mature dark purple. Stems when mature bright yellow.

15a. B. gtodtrnannia var. grodtmnnia 15b. var. jlavoramea

(15a) Berberis grodtmanm ‘a var. grodtmannia S. Szechuan: fl. 1914, Schneider 1268 (Type, K). Not cultivated.

(15b) Berberis grodtmannia var. flavoramea Schneid., Fedde Rep. Sp. Nov. 46,

S . Szechuan: Yung-ning, 22 Apr. 1922, Kingdon-Ward 5078 (Type, E). Not cultivated. 256,1939

(16) Berberis nantoensis Schneid., I.c., 252, 1939; syn. B. &ns~dia Byhouer, J. Am.

Shrub 3-6 ft. Sterns very sulcate. Spines absent or weak, 1-3-fld., 8-12 mm. Leaves 10-30 x 5-8 mm. ; attenuate to petiole 1-2 mm. ; narrowly obovate, acute; margins to- wards apex 1-3-spinose (1-2 mm.)-subsedte, distant 4-7 mm. ; subenervate, rarely with a few lateral veins visible above; below at firat slightly pruinose, finally green,

Arb. 9, 133, 1928; non Rusby

Page 45: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 45 epruinose, epapillose. Flowers fascicled 2-3. Pedicels 3-5 (-7) mm. Ovules 2-3. Berries 6-7 x 4-5 mm., ellipsoid pruinose, and with short style.

Formosa: Prov. Nanto, 1918, Kiraishu, fr. 6 Mar. 1918, Wilson 10074 (Tgpe, K). Not cultivated.

Subsect. Verruculosae Ahrendt, J . Bot., Land. 79, (Suppl.), 40, 1941 ; syn. subsect. Candiduh Schneid., Mitt. Dtsch. Dendr. Ces., 55, 52, 1942

Shrubs 2 4 f t . Stems terete (except in asnzyana), yellow and distinctly verruculose. Internodes 7-12 (-25) mm. Spines mostly 1-2 cm., 3-0d. Leaves without hypoderm, enervate or subenervate; margins with 1-6 spinules. Flowers solitary. Petals shorter than inner sepals. Stamens produced (unleas noted). Oprules (3-1 4-5, sessile or subsessile. Berries estylose, pruinose blue.

W. Szechuan, Hupeh.

KEY 8, to Subsect. Verruculosae (Map 6) Stem terete; flower diameter (lo-) 12-18 mm.

Leaves green, epruinose below; dull, bright green above. Leaves white, or grey, pruinose below; lustrous dark green above.

Leaves revolute, narrow, 1 : 3, white below; fruit 8-9 mm. long.

Leaves plane, broad, 1 : 2, greyish below; fruit 10-12 mm. long.

l7a. B. hybrido-gagnepainii,

17. B. mndidula

18. B. verrucdosa

19. B. m y a m

Cf. 39. B. alpicola

Stem sulcate; flower diameter 7-10 mm. Stem verruculose; leaves paler below; fruit pruinose, estylose. (Stem not verruculose ; leaves concolorous below ; fruit epruinose, stylose).

(17) Berberis candidula Schneid., Bull. Herb. Boissier (2), 5, 402, 1905; syn. B . hookeri var. candidula Schneid., 1904; syn. B. d i c h i a n a var. @ida Bois., F M . Vilm. 15, 1905, non Hook. f. et Thorns.

Stems, shoots green, mature brown-verruculose. S p i m subterete. Leaves 12-26 (-30) x 6 8 (-10) mm., elliptic to ovate; margins very revolute, 3-4-spinose (0.5-1 mm.)- serrulate, distant 4-7 mm.; very lustrous dark green; below prUinose white. Pedicels 4 8 (-10) mm. Outer sepals reddish, 4 x 2 mm., oblong-ovate, acute. Median sepals 7 x 5 mm., oblong-obovate. Inner sep& 10 x 8 mm., obovate. Petals 8 x 6 mm., obovate, entire, cuneate; glands ovate, 0-8 x 0.5 mm. Sturnem 5 mm. Berries 8-9 x 4-5 mm., pruinose pale grey.

W. Hupeh: 7500 ft., 1907, Wilson 2880; Farges 637, seed ody. Cultivated: type plant cultivated by Vilmorin. This is a dense compact shrub, with

s t 8 arching branches forming a mound 2-3 ft. high and 4-5 ft. through. It is one of the commonest and most charming species in gardens. Examined, fl. 21 May 1942; fr. 17 Oct. 1940, 5 Dec. 1944.

(17a) Berberis x hybrido-gagnepainii Suringar, Jaarb. Ned. Dendr. Ver. 146, 1929; syn. B.xwokingensis Ahrendt, Card. Chron. (3) 108, 56, 1940 nomaud; ayn. B. gagnepainii, Tottenham variety, Catalogue Jackman

Stems stout. Spines sulcate. Leaves 20-40 x 7-15 mm., ovate; margins revolute, 6-9-spinose (1 mm.)-sedate , distant 3-5 mm. ; dull light green; below slightly lustrous, concolorous. Pedicels 3-7 mm. Outer sepals 2.5 x 2 mm., ovate, subacute. Median sepals 6 x 4.5 mm., similar. Inner sepals 8 x 5 mm., obovate. Petals 6-5 x 5 mm., obovate- spathulate, slightly emarginate; base cuneate with ovate, obtuse glands, 1 x 0.7 mm. Berries 7 x 4.5 mm., pruinose grey.

Page 46: Berberis and Mahonia : A taxonomic revision

46 L. W. A. AERENDT

gemis, 0).

has been given in cultivation to a hybrid between B. vemwhsa and B. candid&.

(18) Berberis verruculosa Hemsl. et Wilson, Kew BuU. 1906,151 ; Sprague, Bot. Mag.

Stem densely pubescent and verruculose; shoots green. Sph8ea slender. Leavm 12-22 x 6-11 mm., obovate-elliptic; margins 2-4-spinose (1-1-5 mm.)-serrulate; acute; lustrous dark green; below papillose, pruinose, greyish white, becoming gre*h green, and occasiody green as the bloom completely disappecL1.s. Pedkeh 4-7 (-9) mm. Outer s e p h 4x3mm. Median sepah 6x5mm., ovate. Inner se@ 1Ox8mm., obovate. Petals 8 x 7 mm., acutely emarginate, with rounded lobes; base cuneate with submarginal glands 1.5 x 0.6 mm. Berries oblong-ovoid, 1Cb-12 x 6-7 mm., pruinose blue.

W. Szechuan: Tachien lu, 1903, Wilson (Veitch) 3150, 3150A (Type, K). Collected from 1903 to 1910, at heights from 4200 to 13,300 ft.

Cultivated: fl. 21 May 1942. The large flowers are subglobose, not fully opening; the shrub is 2-3 ft. high, possessing a great beauty of form from its rigid arching branches. Like B. candid&, which is also most valuable for its more dense beauty of form, this is a species which is common in cultivation. The bloom on the leaf undersurface is less dense than with B. candid&, and becomes less with the age of the leaf; it only very occasionally disappears to leave a green undersurface. It is to this, I believe, that Dr Schneider refers when he described a form vkridescens, Fed,de Rep. 8 p . Nov. 46,266,1939 ; Mitt. Dtsch. Dendr. 55, 55, 1942, which I do not consider to be distinct from the type.

(19) Berberis asmyana Schneid., Sargent, PZ. Wils. 1, 357, 1913 Shrub 3 4 ft. Stems sul~ate, verruculose. S p * w slender, 2-6 (-9) mm. Leaves very

thin, 12-25 (-30) x 6-8 (-9) mm. ; elliptic to obovate-elliptic; margins occasionally entire, otherwise 1-3-appressed spinulose (0-25-0.5 m m . ) - s u b d t e , distant 7-12 (-15) mm. ; lustrous; below paler green. Pedicels 2-3 cm. Prophylls 1-75 x 1.25 mm., ovate. Outer eepah 3 x 2.5 mm., ovate. Inner s e p k 5 x 4-5 mm., obovate-orbicular. Pe.kda 4.5 x 3.25 mm., obovate, shortly emarginate; base clawed with approximate submarginal glands, 1 xO.5 mm., placed 1 m. above the base of the claw. Stamem 3.25 mm., apex crenulate-truncate. Bewia (immature), 6.5 x 5 mm.

cultivated: origin at Aaismeer. F1. Apr. 1939; fr. 29 Oct. 1940 (Type of B. ~ltokin-

C. K. Schneider, Yitt. Dbch. Dendr. Ces. 55,58,1942, records that the name B. frihrti i

138, t. 8454, 1912. (N. 23)

W. Szechuan: Mupin, June 1908, Wilson 2873 (Type, K). Not cultivated.

Subsect. Manipuranae Ahrendt, J. Bot., Lond. 79 (Suppl.), 36, 1941 Stew glabrous; mature, yellow (except only in B. mrdi i , purple). I&& (1.5)-

2-3 cm. (or to 5 cm. where noted). Leavea evergreen, fairly thin, without hypoderm, mostly reticulate (unless otherwise noted). Stwnens not, or scarcely produced, not apiculate. Ovules (3-) 4 4 . Berries estylose (except only in B. wardii), and nearly always pruinose blue (except where otherwise noted).

Assam, Manipur, Burma, Yunnan, Kwangsi, Formosa.

KEY 9, to Subsect. Manipuranae (Maps 5 and 7) Flowers solitary; pedicel8 3-4 cm. 20. B.paraapecta Plowers fascicled; pedicel8 6-25 mm.

Ovules seesile or subseaeile. Spines, 1-3 cm., 3-fld.

Leaves below green, epruinose. Leaves reticulate; flowers fascicled f5-15; inner sepals 9-10 mm.

21. B. wnipurana

Page 47: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 47

Leaves scarcely reticulate; flowers faacicled 2-5; inner sepals 6-7 mm. Stems subterete; petale subequal to inner sepals. 22. B. bucharutnii Stems very sulcate ; petals much shorter than inner sepals. 23. B. f a k

Fruit epruinose, narrow, 10 x 4 mm. 24. B. bulhtu Fruit pruinose, proportionately broader.

Stems purple, at any rate at first; fruit stylose. 25. B. wardii Stems green at first, mature yellow. Stems terete; leaves 3-7 cm. 26. B. cozii Stems sulcate-angled; leaves 10-25 mm. 21a. B. x watlingtonemk

Ovules distinctly stipitate. 27. B. mingetaenak Spines absent or very weak. 28. B. irnpeditu

haves below white, or grey, pruinose.

(20) Berberis paraspecta Ahrendt, sp.nov., aff. B. mnipranue a qua foliis angustiori-

Shrub to 6 ft. Stems sulcate. Spines concolorous, stout, 2.5-3.5 cm. Leaves 3-6.5 x 0.7--1.2 cm., lanceolate, acuminate; margins with 7-15 spinose (1.5-2 mm.) serrations (1 mm.), 2.5-4 mm. distant; both sides similar, concolorous, with lateral and marginal veins and a few veinlets elevated, closely and finely reticulate. Flowers solitary. Pedicels 3 4 cm. Berries (immature) 6 x 4 mm., estylose, pruinose blue.

Y-unnan: Lichiang fu region, by Ngu-bah-kah, 9500 ft., 27 July 1914, Schneider 2028 (Type, K). Not cultivated.

bus, floribus solitariis pedicellis longioribus inter alia distinguenda

(21) Berberis manipurana Ahrendt, Kew Bull. 1939, 262; Schneid., Mitt. Dtsch. Dendr. Ges., 55, 18, 1942

Stem angled, not verruculose. Internodes 2-5 cm. Spines concolorous, sulcate. Leaves 6-8-5 x 1.5-3 cm. ; oblong-elliptic to ovate-elliptic; acute, mucronate; base cuneate to petiole 2 4 mm. ; margins subappressed 10-20-spinose (0.5-1 mm.)-subserrulate, distant 3.5 (-7) mm. ; closely reticulate with a distinct marginal vein on the lustrous deep green upper surface; below paler yellow-green. Flowers fascicled 6-12 (-15). Pedicels slender, 10-18 mm. Prqhylls 3 x 1.5 mm., red, lanceolate. Outer sepaEs 5 x 3 mm., oblong- obovate, subacute. Median sepals 8 x 6 mm. Inner sepals 10 x 8 mm., obovate. Petals 8.5 x 7.5 mm., spathulate-obovate, acutely emarginate with rounded lobes; base cuneate, with oblong, separate non-marginal glands, 0.9 x 0-5 mm. Stanzens 5.5 mm. Berries 10-10.5 x 56-6 mm., pruinose blue.

Manipur: Apr. 1882, Ching Sow, N.E. Ranges, 8000 ft., Watt 6533 (Type, K, distri- buted erroneously in seed as 6428 and labelled B. Wauichiana microcarpa, which more recently has been raised to specific rank as B. sublevis W. W. Sm.); Apr. 1882, Sirvhi- furar, 7000 ft., Watt 7206 (K, seed distributed in error as 6533, and labelled B. walli- chiana htijolia; cf. 5d, B. h k e h var. latifolia) ; 7 Apr. 1948, Sirvhifiwar, on the fringe of the forest on the lee side of the ridge, 7000-8000ft., Kingdon-Ward 17220; this has leaves with somewhat less venation, and flowers fascicled 20-30, and may be a variety.

Cultivated : probably the most common Asiatic evergreen Berberis in cultivation, being slightly tender, and forming an erect bush, 6-9 ft. high, with very handsome foliage. In gardens it seems generally to have been called B. wallichiana latifolia, doubtless due to the second error referred to. When it became realized that cultivated plants of B. wdichiana Hook. non DC. were properly to be named B. hookeri Lemaire, the varietal epithet was caaually transferred and Watt’s plant was called B . hookeri var. latifolia Hort., by which name it is still very commonly known. It will be seen (cf. 5 4 that the true B . h h r i var. 2atifolia differs in its many ovules, few flowered fascicles and emarginate petals. More recently still, B . mnipurana has been confused with, and called by, the name of the Javan B. xanthxylon, which, though possessing a similar inflorescence of smaller flowers, has very thick, entire, leaves with hypoderm, and black epruinose

Page 48: Berberis and Mahonia : A taxonomic revision

48 L. W. A. AHBENDT berries. Finally, it must be recorded that B. munipram has, in gardens, for a long time been frequently distributed as B. high&, a name belonging to a very different, S. American, species.

(21a) Berberisx watlingtonensis Ahrendt, hybr.nov. (B. v e m h a x 1 B. m n i - prana) , a B. mnipurana foliis minoribus, venis paucis, subtus pruinosis, floribus fasciculatis tintum 2-3 dif€ert

Sterns sulcate, verruculose. Leaves to 25 x 13 mm.; elliptic-ovate; margins 2 4 - spinose (1 mm.)-serrate (0.25 mm.), distant 6-10 mm.; lustrous yellow-green, with few veins; below pruinose, white, finally greyish. Flowers fascicled 2-3. Pedictd.3 slender, 15-20mm. Outer se@ 3x2.5mm., elliptic, rounded. Hedian sepals 5 ~ 5 m m . , elliptic-suborbiculm. Inner sepals 8 x 5 mm., obovate. Petals 6-5 x 5.5 mm., or 6 x 6 mm., broadly obovate, shortly incised; basal glands 0.8 x 0.5 mm., ovate, subacute, separate, non-marginal. Stccnzens 4.5 mm., produced, obtuse. Be& 9 x 6 mm., oblong, pruinose blue.

Cultivated: origin at Watlington, Oxfordshire; fl. May 1940; fr. 29 Oct. 1940 (Type, 0). A dense compact shrub, with stiff arching branches rising to 3 ft.

(22) Berberis buchananii Schneid., Fedde Rep. Sp. Nov. 46,263,1939 Stems slightly or subangled. Spines sulcate. Leaves 20-40 (30) x 7-13 (-20) mm. ;

oblong-elliptic to oblong-ovate; petiole 1-2 mm.; margins scarcely revolute, with 6-12 spinoss (1 mm.) serrations (1 mm.), distant 2-5-3.5 mm. ; venation impressed, not reti- culate; above very lustrous; below subconcolorous or scarcely paler yellow-green. Flowers fascicled 26. Pedictd.3 12-22 mm. ProphyywS 2.25 x 1.25 mm., ovate. Outer sepals 4 x 2 111111. Inner sepals 7 x 3 mm., a.ll oblong-obovate. Pefale similar, 7 x 2.75 mm., eub- entire.

Petals subequal to inner sepals, both m o w ; stamens not produced.

Petals shorter than inner sepals; strunens produced. 22a. B. b&mnii vm. buchananii.

22 b. var. tamwed.

(22a) Berberis buchananli var. buchananii

cultivated. Upper Burma: Ruby Mine district, 23" N., 96" 20' E., Buchanan 165 (Type, K). Not

(22b) Berberis buchananii var. tawangensis Ahrendt, J. Bot., Lond. 79, (Suppl.),

Stems more angled sulcate. Outer sepals 5 x 4-5 mm. Inner s e p h 8 x 6 mm. Petals 6.5 x 5 mm., entire; all obovate. Stanzens 5 mm., produced. Berries ellipsoid, 10 x 5 mm. Sea& dark purple.

S.E. Tibet: Prov. Monyul, Tawang, 9500 ft., 1934, Ludlow & Sherriff 1089 (Type, BM) ; Prov. Pome, Tongkyuk Dzong, 10,OOO ft., 22 Dec. lW, Ludlow, S h e d & Elliot 12026; 9OOOft., 20 May 1947, Ludlow, S h e d & Elliot 13713; Deru, Po-tsangpo, IO,MH)ft., fl. 6 June 1947, Ludlow, Sherriff & Elliot 13080 (all, BM).

Cultivated: from type collection, fl. 29 Mar. 1948; fi. Nov. 1948; a valuable foliage evergreen of 3-5 ft. high, supeficially similm to B. hoolceri, but with more finely toothed, much less veined and even more lustrous and non-reticulate leaves, with smaller flowers and berries, and only 3-5 ovules.

37, 1941

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Berberis and Mahonia 49

(23) Berberis fallax Schneid., Fedde Rep. Sp. Nov. 46, 260, 1939

Shrub to 8 ft. Stem very sulcate, not verruculose. Spines concolorous, 1.5-2 cm. terete. Leaves 20-60 x 8-16 mm.; oblong-ovate, acute, mucronate; margins with 7-15 spinose (1 mm.), serrations (1 mm.) ; openly veined, lustrous; below slightly paler yellow. Flowers fascicled 3-5. Pedicels 1-2 cm. Prophylls 1-2 mm. Outer sepals 4.5 x 3 mm., ovate, subacute. Inner sepals 6 x 4 mm., elliptic. Petals 4 x 2-5 mm., obovate, acutely emarginate; base conspicuously clawed, with oblong glands, 0.8 x 0-3 mm., situated above the claw. Stanzens 2.5 mm. Berries 7-8 mm. (immature), epruinose.

N.W. Yunnan: Chetse-lo, 1934, Tsai 58521 (Type, Am. Arb.); N. of Yunnan-fu, be- tween Hsaiodsang and Loheitang, Schneider 388 (K). Not cultivated.

(24) Berberis bullata Ahrendt, J. Bot., Lond., 79 (Suppl.), 38, 1941

Stem very sulcate; shoots green becoming reddish; mature red-brown, becoming yellowish, sparsely, finely, verruculose. Spines sulcate, discolorous, yellow. Leaves 2 0 4 0 x 7-13 mm. ; elliptic to oblong-elliptic, acute, mucronate; base cuneate, subsessile; margins 3-5-spinose (1 mm.), serrulate (0.5-1 mm.) distant 3-5 (-7) mm.; slightly sub- revolute; with marginal and 3-6 lateral veins impressed; dull green; below pruinose grey. Flowers fascicled 2-6 (4). Pedicels 10-18 mm. Prophylls 3 x 1-1-5 mm., ovate. Outer sepal8 5 x 2.5 mm., oblong-ovate, subacute. Median sepals 6-5 x 4.5 mm., oblong- ovate to elliptic. Inner sepals 7 x 6 mm., obovate. Petals 6 x 5 mm., obovate; shortly emarginate; base cuneate with separate non-marginal glands, 1 x 0.6 mm. Stamens 4 mm., slightly produced. Ovules subsessile. Berries black, epruinose, 9-11 x 4-5 mm., narrowly oblong, often bent, estylose.

Cultivated: fl. 21 May 1942; fr. 19 Oct. 1940, 13 Jan. 1943 (Tgpe, 0). A dense erect shrub, P 5 ft.

(25) BerberL wardii Schneid., Fedde Rep. Sp. Nov. 46, 262, 1939

Stem angled; shoots purple; mature a t ikst purple, finally yellowish. Leaves 20- 30 x 10-15 mm. ; elliptic to elliptic-ovate; margins with 5-9 spinose (1.5-2 mm.) serrations (0.5 mm.) ; deep grey-green; lateral and marginal veins and a few veinlets elevated; below pruinose white, finely papillose, less veined. Flowers fascicled 2-5. Pedicels red, 15-25 mm. Prophylls small. Outer sepals 4 x 3 mm., ovate, obtuse. Median and Inner sepals 7-8 x 4-5 mm., oblong-elliptic to oblong-ovate, subentire ; base clawed, with oblong glands. Stamens slightly produced, apex crenulate with centre shortly apiculate. Ovules sessile. Berries shortly stylose.

Assam: Naga Hills, Japvo, fr. 2 Dec. 1927, Kingdon-Ward 7729 (Type, K); 10,OOO ft., 1935, Kingdon-Ward 12573 (BM); 24 Nov. 1949, Kingdon-Ward 19086 (BM). A thick compact, small, scrubby bush, on the S. face in open places amongst grass, Ilex, Rubus. A small plant of this species was in cultivation at Wisley in 1938-9, but it died in a succeeding winter, and I have no record that it is now in cultivation.

(26) Berberis coxii Schneid., New Flora and Silva, 10, 257, 1938, anglice; Fedde

Stems terete, very stout, finely brown verruculose. Spines plane. Leaves (3-) 4-6 (-7-5) x (1.2-) 1-6-2.6 (-3) cm.; elliptic to ovate-elliptic; petiole 3-10 mm.; margins scarcely revolute, with 6-11 appressed spinose (0.5 mm.) serrations (0.5 mm.), distant 6 8 mm., with but few veins, impressed on the very lustrous upper, and elevated on the pruinose white lower, surface. F h e r s fascicled 3-6. Pedicels 10-15 (-20) mm. Outer sepals 4 x 2.2 mm., oblong-ovate, acute to subacuminate. Median sepals 6-7 x 3.5 mm, oblong-ovate to oblong-obovate. Inner sepals 8-9 x 5-7 mm., obovate. Petals 6.5 x

Rep. Sp. Nov, 46, 261, 1939, latine.

4 JOURN. LTNN. S0C.-BOTANY, VOL. LVII

Page 50: Berberis and Mahonia : A taxonomic revision

50 L. W. A. &IRENDT

5.5 mm., obovate, entire; base clawed with oblong approximate glmds, 1.2 x 0.5 mm. Berries 11-12 x 5-6 mm., estylose, pruinose blue.

Upper Burma: Hpimaw, 1919, Farrer 1030 (Type, E). cultivated: from type collection, fl. 9 May 1939, 3 June 1943; fr. 17 Nov. 1943. A

magnificent dense, compact, large evergreen shrub, growing to 6-7 ft., valuable for its foliage.

(27) Berbris mingetsensis Hayata, IG. PI. Fomos., 5,4, 1915 Sterne slender, very suImte. Internodes 3-5cm. Spines stout to 13mm. Leava

6 - 6 9 x 1.3-2-5 cm., lanceolate, acumin&te, aristate; cuneate to petiole 2-3 mm.; margins (7-) 9-15 (-2O)-spinose (1-2 mm.)-sedate (0.25 mm.) distant, 3-6 mm.; lustrous; be- low pruinose white. Flowers fascicled 3-5. Pediceh 6-9 mm. Prophylh 1.25 mm., ovate. outer 8t?+ 3 mm. Medkn se@ 4 mm. Inner se+ 5 mm. Petals 3 x 2.5 mm., obovate, entire; base cbwed. ovules with s t i p as long as, or up to twice as long m, the ovule. Formosa: 1914, Mt. Arism, Mingetsuki, Hayata, s.n., Type. Not cultivated.

(28) Berbris impedita Schneid., F& Rep. 8p. Nou. 46, 263, 1939 Stems slightly angled. Spins absent, or very weak. Leaves 4 4 . 5 x 1-8-2-5 cm.;

elliptic to obovate-elliptic; petiole 5-8 mm.; margins by apex, 8-15-spinose-serrate (2-2.5mm.); both sidea clearly veined, above impressed reticulate and deep green; below pruinose, grey, h d y almost green. Bewk (immature) c. 8 mm., estylose.

Kwangsi: near Kweichou border, Min Shan, N. Luchen, 17 June 1928, Ching 6053 (Type, Vindob); Wang 4009p. Descriptions and citations according to C. K. Schneider. Not cultivated.

Subsect. Triacmthophorae Schneid., Matt. %ch. Dendr. Ges. 55,33,1942 Stem glabrous, terete (except in B. lubriia). I&& 2-4 (-6) cm. Spines 3-0d.,

1-2 (-2.5) cm. (unless noted). k v e s without hypoderm, thinly cori&c8ous, nmow ( 1 : b l I ) ; enervate (unless noted), rarely slightly openly veined. F h r 8 fascicled 2-5 (-15). Pe.dkz2.s (1-) 1-5-3-5 cm. Petals shorter than inner sepals. Ovules 23, sessile or subsessile. Be& estylose, pruinose blue.

W. Hupeh, W. Szechuan, Yunnan, N. Burma.

KEY 10, to Subsect. Triacanthophorae (Maps 5 and 7) Stems dark, red or purple, terete, when mature; o d e a 2-3; leaveg below pruinose,

Leaves broad, 1:4, enervate; flower diameter 13 mm.; pedicels 8-12 mm.; stamens

Leaves narrow (1 : S), veined ; flower diameter 7 mm. ; pedicels 15-30 mm. ; stamens

grey; stamens produced.

apiculate. 29. B.venwta.

trunmte. 30. B. triacanthopka. Stems pale yellow when mature; ovules P 5 . Stems terete; leavea typically dull above.

Leaves below epruinose, yellow-green; stems verruculose. Leavea ovate to lmceolate, scuminate; enervate or almost so.

Habit open, arching. Shoots red; leavea green below; petals emarginate. 31. B. ueifchii. Shoots green; leaves at first pruinose grey below; petals subentire.

32g. B. x chewdtii . Ysbit dense, erect; shoots green; pet& emarginate. 32. B. gapepainii.

lleaves elliptic, acute, veined. 33. B.phamra.

Page 51: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 51 Leaves below pruinose grey.

Leaves enervate. Stems not verruculose ; shoots red; leaves dull, narrow (1 : 5-7).

Stems verruculose ; shoots green ; leaves lustrous, broader (1 : 4-5). 34. B. wisleyensia, p.

31a. B. x chilternemk, p. 32a. B. gagnepinii var. subovata.

Stems sulcate ; leaves lustrous above. 35. B. lubrica. Leaves veined.

(29) Berberis venusta Schneid., Fedde Rep. Sp . Nov. 46, 256, 1939; Mitt. Dt.sch.

Spines yellow, sulcate, slender. h v s 15-48 x 4-12 mm., oblong-elliptic to oblong- lanceolate; petiole 1-3 mm.; margins with 3-10 spreading spinose (1-2 mm.) serrations, distant 4-8 mm. ; below pruinose grey. Flowers fascicled 2-5. Pedi& 8-12 mm. Prophylla oblong, acuminate. Outer sepals 5-6 mm. Inner sepals 7-8 mm. Petals 5- 6 x 4 4 mm., obovate, subentire; base clawed with elliptic glands. Ovules 2-3. Sturnens apiculate.

North Burma: above Nyitadi, below Mokuji Pam, 1920, Farrer 1579 (Type, E). An erect shrub, 3 4 ft. high, not in cultivation.

Dendr. Bes. 55, 33, 1942

(30) Berberis triacanthophora Fedde, Bot. Jb. 36, (Beibl. 82), 43, 1905 Leaves 20-60 x 2.5-8 mm., narrowly elliptic to oblong-lanceolate; margins by apex 2 4 -

spinose (0-5-1 mm.)-serrulate, distant 4-7 mm. ; both sides openly veined; above lustrous dark green; below pruinose grey, papillose. Flowers fascicled %6. Pedicels 15-30 mm. Prophylls red, ovate, 1 x 0.8 mm. Outer sepals 2 x 1-75 mm., ovate-orbicular, rounded. Median sepals 3-5 x 2.5 mm., ovate, obtuse. Inmr sepals 4.5 x 4 mm., obovate. Petals 3.75 x 2.5 mm., obovate; glands 0.8 x 0.4 mm., oblong-ovate. Stamens 2 mm., produced. Ovules 2. Berries 6-7 x 4-5 mm., ellipsoid.

E. Szechuan: fl. Changyang, 1885-8, Henry 5861 (Type, K). W. Hupeh: Kienshi, 4000-6000 ft., 1900, Wilson (Veitch) 952 (K). An apparently attractive species not in cultivation; €or plants bearing this name see

B. wideyewis.

(31) Berberis veitchii Schneid., Sargent, PI. Wik. 1, 363, 1913; syn. B. acurninata

Stems shoots bright red; mature pale yellow, concolorous-verruculose. Spines 3-3d., concolorous, stout, 1.5-3.5 cm., sulcate. Leaves 5-11 x 1-2 cm., lanceolate, acuminate; margins undulate with 10-24 somewhat spreading spinose (16-2 mm.) serrations (2-3 mm.), distant 3-9 mm. ; above dull grey-green; below slightly lustrous paler, yellow- green. Flowers fascicled (2-) 4-8 (-10). Pedicels (1-6)-2-3-5 cm. Prophylls 2 x 2 mm., ovate, subacute, these and the outer sepals (which are 3.5 x 3 mm., oblong-ovate and truncate) being much suffused with pink, reddish brown, and green. Median sepals (5 x 4 mm.) and inner sepa2s (7.5 x 5.5 mm.) all obovate, concave, pale straw colour, also often similarly suffused and flushed. Petals (H.C.C. 403/1), obovate with apex rounded and narrowly incised ; base slightly clawed with approximate non-marginal subcon- colorous glands, 1.5 x 0-8 mm. Stamens 4 mm., slightly produced, obtuse. Berries 9 x 6 mm., ovoid to ellipsoid, estylose, pruinose blue.

W. Hupeh: 1900, Wilson (Veitch) 1138 (Type, K); Ichang, Patung district, 1886, Henry, s.n.

Cultivated: fl. May 1936, May 1940 (flowers mostly pink without), 17 May 1946 (flowers mostly red and brownish green without) ; fr. Nov. 1936. Widely distributed in gardens, this forms a most attractive shrub €or: (a) habit, with its openly arching stems,

Veitch, Hort. Veitch, 391, 1906, non Ranch., nec Stapf.

4-2

Page 52: Berberis and Mahonia : A taxonomic revision

62 L. W. A. AHRENDT bright red when young, some 3 4 ft. high; (b) bold and striking foliage; and (c) unusual flower colour. ,There are many inferior garden hybrids of this and the next species; even with these a distinction c8n generally be effected by observing the red shoots, pale buff flowers and loose habit of B. veitchii, in contrast. with the green shoots, bright yellow flowers, and deme habit of B. gagnepinii. B. veitchii is often distributed as ‘B. mumi- natac’, an error which probably originated in 1906.

(31 a) Berberis x chilternensis Ahrendt, hybraov. (B. veitchii x probably B. werm- culosa), aff. B. veitchii, sed foliis minoribus nitidis, subtus cinereo-prUinosiS distinguenda

Stem verruculose. S p i m sulcate. h u e s to 45 x 10 mm., elliptic-lanceolate; margins slightly undulate, with 5-8 spinose (2 mm.) serrations (1 mm.), distant 3-5 mm.; above lustrous, below pruinose grey. F h r 8 fmcicled 3-5. Pdi& 15-22 mm. Flozvers and fruit aa in B. veitchii.

Cultivated: origin at Watlington, Oxfordshire; 0. 25 Apr. 1939; fr. 1 Nov. 1938 (Type, 0). Grown from seed of B. veit&ii near a bush of B. v e m h a , this seems most probably a hybrid between the two species; it presents an intermediate habit, 3-4ft. high with arching branches, h d y becoming more denae and compact ; leavea n m w as in B. ueitchii, but smaller and lustrous and pruinose below as in B. verruculosa.

(32) Berberis gagnepainii Schneid., Bull. Llerb. B&&r (2), 8, 196, 1908 KEY 10a Leaves broad, 1:3-8.

Flower diameter 14-17 mm. 32h. B. x chenacultii. Flower diameter 9-12 mm.

Leaves dull above; marginal spinules P 7 mm. apart. Leaves green below, subenervate. 32a. B. gagnepainii var. gcdg.nepinii. Leaves at first pruinose, grey, below, veined. 32 b. var. SzLbOvata.

32c. var. omeie&. h v e a lustrous above; marginal spinules 1-2 mm. apart.

Leaves very undulate-dentate, the spinulw and serrations each measuring 2 nun. Leaves narrow, 1:(5-) 7-9 (-11).

Pedicels 10-20 (-25) mm. Flowers fascicled 3-7. 32d. var. lanceifolfolia. Flowers fascicled 8-15. 32e. var. h ~ i f o l i a ; f. pluri&ra.

Pedicels (2-) 2.5-4 cm. 32 f. vax. filipees. Leaves scarcely undulate, with margins finely pi nu lose or subentire.

32g. var. pmeat4ans.

(32a) Berberis gagnepainii var. gagnepainii Leuwm 20-30 (40) x 8-11 (-13) mm., narrowly ovate, acuminate, mucronate, cuneate,

sessile; margins undulate 6-10-spinose-serrate. li’knmrs fascicled 3-5. Pedkls 10-15 nun. Flowers 11 mm. diameter, similar to, but slightly smaller than in the variety lanceifolia, from which, otherwise, it differs only in the leaves.

W. Hupeh: fl. lO,OOOft., July 1903, Wilson (Veitch) 3148 (Type, K). Not in culti- vation.

(32b) Berberis gagnepainii var. subovata Schneid., Fedde Rep. Sp. Now. 46, 264, 1939; Mitt. Dtsch. Dendr. Ges. 55,53, 1942

Internodes 3-4 cm. Spines 2-4.5 em., sulcate. Leaves 3-5-6 x 1-2.2 cm., narrowly ovate, subacuminate; margins scarcely unddate, 5-8-spinose (1-2 mm.)- serrulate; above dull green, with some lateral veins and veinlets distinctly visible; below at first pruinose grey, fairly soon becoming deep yellowish green and epruinose. Flowers fascicled 2-6.

Page 53: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 53

Pedicels 1-2 om. Inner sepals 5 . 5 ~ 5 mm. Petals 5 x 3.5 mm., slightly emarginate. Fruit, 8.9 x 5.6 mm., oblong-ovoid, pruinose blue.

W. Szechuan: Ching-chi-hsien, near Wa-wu-shan, 8500 ft., 16 Oct. 1908, Wilson 2874

Cultivated: I fist discovered this plant on 20 Sept. 1947, at Maidwell Hall, Northamp- ton, in the grounds of Mr Oliver E. P. Wyatt, where it had probably originated (as in the cam of B. polyantha, q.v.) from Wilson’s seed; it suckers freely; fl. May 1957, fr. Oct. 1956 (Broughton, Oxfordshire).

(Type, K).

(32c) Berberis gagnepainii var. omeiensis Schneid., I.c., 1939, 1942

1-2 mm.

cultivation.

Leaves very lustrous, 3-7 x 1-14 cm., with 20-40 marginal spinose serrations distant,

S. Szechuan: Mt. Omei, 8 ~ ~ f t . , 13 Bug. 1929, Fang 2916 (Type). Not in

(32d) Berberis gagnepainii var. lanceifolia Ahrendt, J. Bot., Lond. 79 (Suppl.), 39, 1941; syn. ‘ B . acurninuta’ Stapf, Bot. Mag. t. 8185, 1908, non Franch, nec.

Stems shoots green; mature yellow, black-verruculose. Spines plane. Laves 36x 4 mm. to 9-11 x 1-2 cm., lanceolate, acuminate, aristate; margins very undulate with 10-20 spinose (2 mm.) serrations (1-2 mm.), distant 5-7 mm.; above dull, pale, grey- green, almost enervate; below paler slightly lustrous yellow-green. Flowers (H.C.C. 2) fascicled 3-7. Pedicels (6-) 10-15 mm.; bracts 1.6 mm. ProphyUs 3 x 2.5 mm., oblong- ovate. Outer sepals 4.5 x 4 mm., oblong-ovate, acute. Median sepals 6-5 x 5.5 mm., elliptic to ovate. Inner sepals 8 x 7 mm., obovate. Petah 7 x 6 nun., obovate, acutely emarginate with rounded lobes ; base cuneate with widely separated submarginal glands, 1.2 x 0.6 mm. Berries 9-10 x 6-66 mm., oblong-ovoid, thickly pruinose blue.

W. Hupeh: 5000-6000ft., 16 May 1907, Wilson 1503 (Type, K); IO,OOOft., 1903, Wilson (Veitch) 3148B.

W. Szechuan: 6000-8000 ft., Mupin, 1908, Wilson 1137 (in part) (BM); Washan, 1908, Wilson 1344 (K).

Cultivated: fl. 3 June 1942; fi. Dec. 1938, 22 Oct. 1940. A compact shrub, 3-5ft. high, with noticeably erect branches. This is a shrub handsome in habit, leaf, flower and berry, and is widely distributed, often merely as ‘B. gagrnpainii’.

Veitch.

(32e) Berberis gagnepainii var. lanceifolia f. pluriflora Ahrendt, forma nov.,

W. Szechuan: Mupin, 6500-8000ft., fr. Wilson 1137 (in part) (Type, K). A shrub floribus fasciculatis 8-15, cetera ut in (3%)

4 4 ft. high in thickets, sometimes to be found in cultivation.

(32f) Berberis gagnepainii var. Wipes Ahrendt, J. Bot., Lond., 79 (Suppl.), 39 1941 Leaves 45-75 x 6-10 mm., lanceolate; margins undulate with 7-15 spinose (1.5-2 mm.)

serrations (1-1.5 mm.) ; above darker green than in variety hwifoll is . Flowers fascicled 5-8. P e d i A very slender, (2-) 26-34 (4) cm. long. Otherwise aa in (32c).

W. Szechuan: S.E. of Tachien lu, 7000-9000ft., 1908, Wilson 1137A (Type, K); cliffs, 10,000 ft., 1903, Wilson (Veitch) 3148A; Kuan Hsien, 1928, 3000-4000 ft., Fang 2372 (K). Fang’s specimen has leaves slightly veined.

cultivated: fl. June 1939; fr. 9 Oct. 1939, by Mr R. D. Trotter; fl. May 1957, fr. Oct. 1956 (Broughton, Oxfordshire).

Page 54: Berberis and Mahonia : A taxonomic revision

54 L. W. A, AERENDT

(329) Berberis gagnepainii var. praestans Ahrendt, 1.c. Leaves to 7.7 x 1-4 cm., or to 8 x 1-2 cm.; margins subentire or 12-16 spinulose

(0.5 mm.)-subsemulate, distant 3-5 mm. FZowers fascicled 3-45. Ped&L 10-16 mm. Otherwise as in (32c), except that the sepals and petals are slightly narrower.

Cultivated: fl. June 1939; fr. 20 Oct. 1939, Wisley; fl. May 1957, fr. Oct. 1956 (Bronghton, Oxfordshire).

(32h) Berberis x chenaultii (Hort. ex Cat. L. Chenault, Orldans, France, 19334), Ahrendt, &rd. Chron. (3) 108,56,1940, nomaud. (B. verrucuha x B. gagnepainiii), aff. B. gasnepainii sed foliis nitidis, ramulis arcuatis, petalis subintegris dis- tinguenda.

Stem arching, black-verruculose. Spines 10-16 mm. h u e s 15-40 x 4-11 mm., ovate- lanceolate, subacuminate ; margins undulate, 5-8-spinose (0.8-1.5 mm.)-serrulate, sub- enervate. F k r s fascicled 24, with pedicels 7-18 mm., and components slightly larger than in (32c), and with petals subentire.

Cultivated (origin with Measrs Chenault, distributed by Mess~3 Hillier, Winchester): fl. 16 May 1939; fr. 3 Oct. 1940 (Type, 0).

(33) Berberis phanera Schneid., Ost. Bot. 2. 67, 22, 1918; Mitt. Dtsch, Ges. 55, 54,

Internmh 2-46 cm. 8pines fairly stout, plane, 1-26 cm. Leaves 30-45 x 8-10 mm., or to 55 x 17 mm., ovate, acute; margins undulate with 8-12 spinose (1.5 mm.) serra- tions (1 mm.), distant M m m . , subrevolute; above dull deep green; below slightly paler lustrous deep green; openly veined. F k r s (H.C.C. 1/2), fascicled 2-6. Pedhls slender (1-) 16-26 (-3) cm., green in flower, red in hit . Outer s e p h 3 x 1.5 mm., ovate, obtuse. Median sepak 5 x 4 mm., elliptic to oblong-ovate. Inner se& similar, 7 x 5.5 mm. Petals (H.C.C. l/l), 5-5 x 4-5 mm., oblong-obovate, rounded, acutely emarginate to subentire; barn clawed with separate non-masginal glands, 0.8mm. Stamens 3-5 mm., slightly produced. Berries 10-11 x 6 mm., estylose, thickly pruiuose blue.

S. Szechuan: woods between Ouentin and Wo-lo-ho, and Ka-la-pa, fl. 6 June 1914, 9OOO ft., Schneider 1460 (Type, K).

N.W. Yunnan: Chungtien, Haba, under forest, 9100 ft., fr. 23 Nov. 1937, Yu 14944 (E). A casual shrub, 6 ft. high.

Cultivated: from Yu 14944, fl. 14 May 1945; h. 26 Nov. 1950. A very handsome shrub in foliage and fruit; a few specimens were wrongly distributed as B. gagnepainii subovata under which I placed Yu’s specimen 14944 in J. Bot., Lond. 79 (Suppl.), 39, 24 July 1942.

Slightly Mering from the above is Schneider’s no. 1535, collected on 13 June 1914, from near the same locality a t 10,000-12,000 ft. This has leaves to 60 x 16 mm., margins 4-8-spinose (1-1-5 mm.)- serrulate; fascicles of 10-14 flowers, with pedicels only 6-9 mm. long, and much smaller flDwers (7-8mm. diameter). This was gathered in mountain forests between Hidm and Wo-lo-ho.

1942

(34) Berberis wisleyensis Ahrendt, Gum?. Chron. (3) 105, 372, 1939 Skmw shoots red; mature yellow, subangled, not or scarcely verruculose. Spines

16-36 cm. sulcate. Leaves 20-55 x 4 4 mm., linear-lanceolate; margins 2-5-spinose (1.5 mm.)-serrulate, distant 8-12 mm.; above dull grey-green; below pruinose grey. Pluwers (H.C.C. 2), fascicled 2-6. Pedicels 10-17 mm. 0uter~sepl-s reddish 4 x 3 mm., ovate, base truncate, apex acuminate. Median sepals 6 x 4 mm., ovate, acute. Inner

Page 55: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 55

sepals 8 x 5-5 mm., obovate. Petals 6-5 x 5-6 mm., obovate, subentire; base clawed with lanceolate acuminate glands, 1.6 x 0-4 mm. Stamens 5 mm., not produced. Berries 9 x 4 mm., oblong-ovoid, estylose thickly pruinose blue.

Cultivated: a t Wisley, 0. June 1938; fr. Oct. 1938. Also a t Watlington, fl. 3 June 1942; fr. 28 Oct. 1943 (Type, 0). A very handsome evergreen in leaf and berry. Originally erroneously distributed as B . triacanthophora.

Stems scarcely verruculose; leaves dull grey-green; pedicels 1-2 cm.; flowers 14 mm.

Stems very verruculose; leaves lustrous yellow-green; pedicela 2-4 cm.; flowers 10 mm. diameter; fruit 9 x 4 mm., pruinose blue.

diameter; fruit 9 x 7 mm., pruinose mauve.

34. B. wisleyenaia.

34a. B. helenae.

(34a) Berberis helenae Ahrendt, J . Bot., Lond., 80 (Suppl.), 95, 1942 Shrub with arching branches, 4-5 ft. Stem terete, pale yellow. Spines concoloroua,

10-17 mm. Leavea 20-50 x 5-13 mm., narrowly elliptic, subacuminate; margins 6-9- spinose (1-1.5 mm.)-subserrulate; lateral and marginal veins visible; below grey, pruinose, densely papillose. InJEarescence (2-) 4-6 (-10)-fld., mostly fascicled, occasion- ally partly umbellate, 3-5 cm. long. Pedids red slender (2-) 3 4 cm. ; bracts 1.5 mm. Outer sepals 3 x 2 mm., oblong-ovate, acute. Median se+ 5 x 3 mm., oblong-elliptic. Inner sepals 6 x 4.5 mm., obovate. Petals 5 x 3 mm., oblong-obovate, subentire; base cuneate with separate submarginal, ovate, obtuse glands, 0.9 x 0.6 mm. Stamens 3.5 mm., slightly produced, rounded. Ovules (3-) 4, sessile or substipitate. Berries 8-10 x 6-7 mm., globose-obovoid, thickly pruinose mauve.

Cultivated: at Wisley (no. 41), fl. June 1938; fr. 28 Oct. 1938. B. helenue var. heteranth Ahrendt, Lc., differs only in its more subumbellate inflorescence of 10-25 flowers with pedicels only 1-2 cm. Both these plants, of which the former is the more attractive, are valuable for their mauve bloomed berries, and, together with B . wisley- emis, may well be of hybrid origin. All grow p5 ft. high.

(35) Berberis lubrica Schneid., Fedde Rep. Sp. Nov. 46, 265, 1939; Mitt. Dtsch.

Shrub 3 4 ft. Stem angled-sulcate, verruculose. Spines concolomus, plane. Leaves 6-9 x 1-16 cm., lanceolate, acuminate; petiole 1-3 mm.; margins with 12-24 spinose (1-1.5 mm.) serrations (1-1-5 mm.), distant 2-5-5 mm. ; lustrous both sides, paler below. Flowers fascicled 6-10. Pedicels 5-10 mm. Prophylls 2.5 x 1.5 mm., ovate, acute. Outer sepals 3.5 x 2 mm., ovate, acute. Inner s e p h 6-7 x 4.5 mm., obovate. Petals 3.5 x Zmm., oblong-obovate, subentire; base cuneate with separated marginal glands, 0.8 x 0-4 mm. Stamens much produced, truncate, 3.5 mm.

S. Szechuan: N. of Kuah Pieh, S. of Molien, between Huali, Molien and Yalang, 9OOO ft., fl. 25 May 1914, Schneider 1384 (Type, K). Not cultivated.

Dendr. Ges. 55,54, 1942

Subsect. Sanguineae Schneid., BuU. Herb. Boissier (2), 5, 403, 1905 Stems yellow, very sulcate, or a t least distinctly angled. Internodes (I-) 1-5-2-5 (-3) cm.

Spines 3-fld., 1-2 cm. (except in B. petrogena). Leaves thin, without hypoderm, narrow, 1 : 5-20 (except in B. petrogena), always green, epruinose, below; enervate or at most with a few faint occasional veins. Flauers fascicled 1-6, diameter 6-1 1 mm. Pedicels 3-20 mm. O v u k 2-3, sessile or subsessile. E’ruit 5-8 (-9) mm. long.

W. and S.E. Szechuan, W. Yunnan.

Page 56: Berberis and Mahonia : A taxonomic revision

56 L. W. A. AEIBENDT

KEY 11, to Subsect. Sanguineae (Map 8) h v e s narrow, 1 : 5-20. Leaves very narrow, 1 : 10-20. 36. B.Crmolita. h v e s lass narrow, 1 : 6-7 (-10).

Flowers fascicled 1 (-2); pedicels 3-5 (-7) mm.; flower diameter 9-11 mm.; o d e s 3.

Flowers fascicled 2-6; pedicels 10-20 mm.; flower diameter 6-8 mm.; ovules 2. 37. B . p h & .

Leaves 1-2 em., with 3-6 a p p d spinules. 38. B. dccvidii. Leaves 3-6 em., with 7-14 spreading spinules. 39. B.aanguin~a.

Leaves broad, 1: 1-52. 40. B. ptrogena.

(36) Berberis insolita Schneid., Fedde Rep. Sp. Nou. 46, 257, 1939. (Fig. 21)

Shrub 6-10 ft. Stem slender angled, sparsely vermcdose. Leaves 3 - 5 4 cm. x 3 4 mm., linear-oblong; margins with 5-10 appmased spinules (OCimm.), subsedate, distant 6-12 mm.; dull, papiuose, above; below dull, paler yellow green, epapillose. Flowers and

S.E. Szechuan: E. of Ning Yuan fu, 25 Apr. 1914, Schneider 1029 (Type, K). This species is not in cultivation, but the herbaxium sheet suggests that its remarkable foliage would make it a very striking plant in gardens.

(37) Berberis panlanensis Ahrendt, Kew Bu.U., 1939 265, Schneid., Nitt. Dtsch. Dendr. Ges. 55,26, 1942; syn. B. sanguinea var. microphylla Hort.; apud Creaay, Card. Chron. (3) 98, 11, 1935. (Fig. 19)

Sterns very sulmte, finely concolorous verruculose. Leaves 10-35 x 2-5 111111. (or, on strong new shoots, to 65 x 9 mm.), narrowly oblong, acute, mucronate; petiole c. 1 mm.; margins 3-8 (-14) appressed spindose (0-25.0-5 mm.)-semuhte, distant 2-4 mm. ; above dull grey green; below lustrous deep yellow green. ~'loecners (H.C.C. 63/2), fascicled 1 (-rarely, 2). Pedkels 3-5 (-7) mm. Prophylk red, 1.75 x 1 mm. ozlter sepal8 3 x 2 mm., ovate, obtuse. Medkn sepals 5 x 4 mm., oblong-elliptic to ovate. Inner sepal8 6-5 x 5 mm., oblong-elliptic. Pe.tds 5.25 x 4.75 mm., obovate, emarginate with rounded lobes; baaal glands 1 x 0.5 mm. Stamens 4 mm., not or scarcely produced. Oudes 3. Berries black epruinose, 5-6 x 3-36 mm., oblong, estylose.

frui t unknown.

W. Szechuan: Pan lan shan, 9750 ft., rare, 1908, Wilson 2875 (Type, K). Cultivated: fl. 27 Apr. 1941, 13 May 1942; fr. 7 Nov. 1940,31 Dec. 1942,6 Jan. 1944.

The small black berries hidden under the leaves are rarely seen, and need to be carefully sought; they rarely ripen until Christmas, although the 00wering seaeon is not late. The pale greenish flowers, devoid of any red marking, axe not sufficiently numerous to be of garden value. Nevertheless, this speciea, though rare in its habitat, is fairly common in cultivation and deserves to be still more used on account of its graceful leaves and dense compact habit, with which it grows slowly up to 6, or even 7 ft. Originally distributed aa B. sarquinea mkrophylla, it should be noted that Franchet's species, B. sawinea, differs in its larger leaves, with serration coaxser and more spreading, longer pedicels in larger fascicles bearing smaller flowers which are deep yellow and conspicuously red without.

(38) Berberis davidii Ahrendt, nommov.; syn. B. Lnsa Schneid., Fedde Rep. 8p. Nov. 46, 254, 1939; Mitt. W c h . Dendr. Ces. 55, 31, 1942, non Tr. et Pl.; s p . B. dJichiam parvifolia Franch.

Shrub dense, 2-3 ft. I&r?wdes 1-2 cm. Stem angled. Leaves 10-2Ox 2-34 mm., lanceolate, acute, mucronate; petiole c. 1 mm.; margins with 3-6 a p p r d spinules (0.5-1 mm.)-sedate, distant 1.5-3 mm.; above dull; below slightly paler, and sub- lustrous. Ovules 2. Berries 8-9 x 7 mm., ellipsoid, estylose, pruinose grey.

Page 57: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 57 W. Yunnan: Mt. Tsang shan above Tali, 28 Mar. 1884, Delavay 1124 (Type, K).

E. Flank of Tali range, 25" 40' N., 100" 10' E., 1910, Forrest 7276 (K); Chien-Chuan- Mekong divide, 28" 3O'N., 1923, Forrest 23599 (K); 1929, Forrest 28167 (K). Not in cultivation.

(39) Berberis sanguinea Ranch., Nouv. Arch. Mus., Paris, (2), 8, 194 t. 5, 1886; Pluntae Davidianae, 2, 12, t. 5, 1888

Shrub 6-10 ft. Stems very sulcate. Leaves 3-6 cm. x 3-6 mm., linear-lanceolate; acute, mucronate, subsessile; margins with 7-14 spreading spinose (1-1-5 mm.) serrations, slightly revolute ; above dull grey-green ; below slightly paler, sublustrous yellow-green. Flowers 6-8 mm. diameter, fascicled 2 4 (-7). Pedkls (7-) 10-15 (-20) mm. Prophylls minute, red. Outer sepals 3 x 2 mm., mostly red, ovate, acute. Median and inner sepals deep yellow, 5 x 4.5 mm., elliptic. Petals 4 x 3 mm., obovate; apex shortly retuse with rounded lobes; basal glands lanceolate, separate, non-marginal, 1 x 0.3 mm. Stunzens 3 mm., not produced, apex rounded-truncate. Oerules 2. Style absent.

W. Szechuan: Mupin, Apr. 1869, David, s.n., Type. Sa-washan, Henry 3763 (K). Plants distributed with this name are occasionally B. recurvatcc, but generally B.

panlunensis which has smaller leaves with fewer appressed spinules, solitary larger short pedicelled pale green flowers, and 3 ovules.

(40) Berberis petrogena Schneid., Fedde Rep. Sp. Nov. 46, 253, 1939; Mitt. Dtsch.

Shrub2-3 ft. Stemssdcate. Spines often absent, otherwise weak, 3-fld., 6-12(-15) mm. Leaves 12-30 x 8-17 mm., ovate, obtuse, subsessile; margins 6 8 (-12) spinose (0.5- 1.5 mm.)-subsermlate, distant 2-6 mm. on the newer leaves, but 2-2.5 mm. on the others; both sides dull, concolorous, dull, enervate or with 1-2 rectangular lateral veins. Flowers fascicled 2-3. Pedicels 7-10 mm. Ovules 2. Berries 7 x 4 mm., obovoid, slightly pruinose blue, with style short or indistinct.

W. Yunnan: N'Markha-Salween divide, 26" 30' N., 1919, Forrest 18195 (Types, K, E); Yunnan-sen district, Pin-fa, Cavalerie 7854. The latter specimen has leaves somewhat paler below, and slightly narrower (1:2-3), and flowers in larger fascicles of 10-15. The species is not cultivated.

Dendr. Ges. 55, 31, 1942

Subsect. Subleves Ahrendt, J. Bot., Ld., 79 (Suppl.) 31, 1941; Schneid., Mitt. Dtsch. Dendr. Ges. 55, 29, 1942

Stems very sulcate, yellow when mature. Intern4mk.s (1-) 15-26 cm. Spines 3-fld.) concolorous. Leaves thin without hypoderm. Flowers 6-9 mm. diameter, fascicled 5-50. Pedkls 5-25mm. Petals emarginate. Stamns distinctly produced, obtuse to sub- apiculate. ovules solitary. Berries black, epruinose.

h a m , Manipur, W. Yunnan, Kweichou.

KEY 12, to Subsect. Subleves (Maps 5 and 8) Ovules with stipe 3-6 times as long as ovule; petale longer than inner sepals.

Ovules subsessile or with stalk scarcely longer than ovule; petala shortar than inner 41. B. 8 U b k V i 8 .

sepals. Leaves subconcolorous below ; ovules stipitate. 42. B. dolichoetemon. Leaves at fkst pruinose, finally paler green below; ovules subsessile. 43. B. ernilii.

Page 58: Berberis and Mahonia : A taxonomic revision

58 L. W. A. AERENDT

(41) Berberis sublevis W. W. Sm., Not. Bat. Qard. Edinb. 9, 83,1916 Stew slightly verruculose. Spines 1-2 cm., plane. Internodes 2-5 cm. h v e s 2.5-

8 x 0-5-4 cm., lanceolate, acuminate; petiole 1 4 mm. ; margins 10-18 subappressed spinulose (0-25-0-75 nun.)-sedate, distant 3-7 mm., sometimes entire or subentire; dull green with lateral and marginal veins visible, indistinctly openly reticulate; below slightly paler green, enervate or slightly veined. F k r s (H.C.C. 2/1), fascicled (lo-) 15-30 (-50). Pedkds slender (5) 7-10 (-25) mm. ProphyZk 1-5 x 1 mm., ovate, acute, red. Outer s e p h 2-5 x 2 mm., reddish, ovate, subacuta- I n w se@ 5 x 3 mm., obovate to oblong-elliptic. Petals 5 3 . 5 x 3 mm.; base clawed, glands 1-5 x 0.5 mm. Stamens 3.5 mm., obtuse. Berries 6-7 x 3-34 mm., estylose, black.

I(EY 12A Leaves P 8 cm., nmrrow, 1 : 45-6.

Pedicels 5-16 mm. Leaves entire or with up to 18 spindes ciistant 3-7 mm.; flowers fascicled 10-60.

Leaves with 15-30 marginal spinules distant 1-3 mm.; flowers fascicled 3-10. 41a. B. sublevis var. 8.ublevks.

41 b. var. microcarpa Pedicels 1-5-2.5 cm. ; flowers fascicled 5-8. 41c. var. exquiaita.

41d. vw. grandvolia. Leaves 8-12 cm., broader, 1 : 3-5, densely spinulose.

(41 a) Berberis sublevis vm. sublevis W. Yunnm: around Tengyueh, 6000-7000 ft., Forrest 9560 (Type, E), 7261,18775. Manipur: 8OOO ft., Khaiyrtng, 4 May 1948, Kingdon-Ward 17381 (BM); Sirhai, 7 Apr.

1948, Kingdon-Wasd 17220 (BM). Cultivated: fl. 9 May 1941; fr. Dec. 1938. Photographs of this variety are in Qard.

IZZust. p. 456, Oct. 1944 (erroneody entitled B. d&kna), and p. 617, 28 Nov. 1941.

(41 b) Berberis sublevis var. microcarpa (Hook. f. et Thorns.) Ahrendt, var.nov., folik mmginibus spinulis pluribus, fasciculis paucif3omtis distinguenda; syn. B. prainiana: Schneid., ex Stapf in Bot. Hag. 157, sub. t. 9153 (1928), in obs.; syn. B. wauichiarm: var. microca?ywl Hook. f. et Thorns. li’l. Ind. 226, f. 1855, Hooker f., Pl. Br. Id. 1, 111, 1872; syn. B. d i e h i a m var. gradi- Ahrendt, J. Bot., Lonal. 79 (Suppl.), 17, Ildar. 1941.

Assam: Khasia Rills, Motling, Feb. 1850, Simons 114 (Type, I(); July 1850, Simons 119 (K); 5000-6000ft., Hooker, s.n.; Kalapanee river valley, 17 June 1850, Hooker 1237 (0).

Mmipur: Sirwhifurar, 7000-8000 ft., Apr. 1882, Watt 6428, 6449 (K). W. Yunnan: Tengyueh, Forrest 8636, 17791 (K). Cultivated: from Forreat 8635, fk. Dec. 1938 (Kew).

(41 c) Berberis sublevis var. exquisita Ahrendt, var.nov., pedicellis mulb longioribus, fasciculis paucifloratis.

Leawea 2-6-3 x 0.6-1.4 cm. ; margins 18-28 appressed spinules distant 2-4 mm. W. Yunnan: Hills, N.W. of Tengyueh, 6000-7000ft., fl. Feb. 1925, Forrest 26196

(Type, K). ‘In dry situations amongst scrub, and in open drg pine forests; flowers fragrant, pale yellow.’ Not in cultivation.

(4ld) Berberis sublevis var. grandifolia Schneid., Fedde Rep. Sp. Nov. 46,253,1939 Leaves to 12 x 3-4 cm., margins with 25-50 spinulw, distant 1-2 nun. W. Yunnan: by Tengyueh, 1923, Rock 7930 (Type, Am. Arb.); Tsai 53605. Not in

cultivation.

Page 59: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 59

(42) Berberis dolichostemon Ahrendt, sp.nov., staminibus petalis longioribus. Stems brown-verruculose. Spines sulcate, stout, 2-3 cm. Leaves to 4 x 1.3 cm., oblong-

elliptic to oblong-obovate, subacute, mucronate, subsessile ; margins subrevolute with 7-11 spinose (0-5-1 mm.) serrations (0.5-1 mm.); subenervate or with few indistinct lateral veins ; both sides dull subconcolorous green. Flowers fascicled 10-20. Pedicels 8-12 mm. Outer sepals 2 x 1 mm., ovate, acute. Inner sepals 4.5 x 2 mm., obovate. Petals 3 x 1-5 mm., obovate; glands 0.5 x 0.2 mm., separate. Stamens 4 mm., produced, obtuse. Ovules solitary, shortly stipitate.

Yunnan : Yunnan-sen, Cavalerie 7852 (Type, K). Not cultivated.

(43) Berberis emilii Schneid., Fe& Rep. Sp. Nov. 46, 255, 1939 S p i m 1-2 cm., plane. Leaves 10-30 x 6 1 0 mm.; ovate to obovate; petiole 1-2 mm. ;

margins 3-10 spinose (0.5-1 mm.) serrate, distant 3-5-10 mm.; openly veined; dull above; below a t b t slightly pruinose, finally epruinose and very pale green. Flowers fascicled 5-13. Pedicels 5-10 mm. Prophylls lanceolate, acuminate. Outer s e w 5 mm. Inner sepals 6 mm. Petals 5 mm., all obovate, and petals slightly emaxginate, their baaes clawed with oblong glands. ovules solitary, shortly stipitate. Berries (immature) epruinose, with short style.

Kweichou: near Kouy-yang, 30 Mar. 1898, Bodinier 2145 (Type, E). Not cultivated.

Subsect. Acuminatae Ahrendt, J. Bot., Lund., 79 (Suppl.), 32, 1941; Schneid., Mitt. Dtach. Dendr. Bes. 46, 36, 1942

Stems terete (unless noted), yellow (except only in B. d e i m w d b ) , always glabrous. Iternodes 1.5-3-5 cm. Spines generally 3-0d., (1-) 1-5-2 (-2-5) cm. (unless noted). Leaves more or less thinly coriaceous without hypoderm, always green, epruinose below, generally concolorous (but paler when so noted), mostly reticulate (unless otherwise recorded), always much and clearly veined; margins multispinose. Flowers 8-10 (-13) mm. diameter. Fruit scarcely known.

S.E. Tibet, Yunnan, S.W. and S.E. Szechuan, W. Hupeh, Kweichou.

-Y 13, to Subsect. Acuminatae (Maps 5 and 9) Spines %6 cm. long, conspicuous and stout, very broad at base; ovules solitary.

stems purple. 44. B.&inucanthct. Stems yellow. 45. B. valida.

Leaves very densely spinose, distant 1-2 (-2-5 mm.). Spines 1-3 cm., or less, ordinary or weak; ovules 2-3.

Leaves narrow, 1 : 4-5, mostly 7-13 cm. long. Leaves broader, 1 : 3-4, mostly 2-5-6 cm. long; stamens produced; fruit estylose.

46. B. argutcl.

47. B. fallacioaa.

48. B. ariatato-aerrulata.

Spines 7-1 3 mm. ; flowers fascicled 2-5; ovules subsessile. Spines 2-4 mm. ; flowers fascicled 10-15; ovules stipitate.

Lertves less densely spinose, distant (2-5)- 3-5 (-8) 111111.

Flowers fascicled 6-15; stem terete; fruit estylose. Leaves reticulate, concolorous below.

Spines absent or 2-8 mm.; leaves 7-19 cm. Spines 1.5-3 cm. ; leaves 3-6 cm.

Flowers fascicled 2-6. Stems terete; pedicels 5-15 (-20) mm.

49. B. muminab. 50. B. delavayi.

Outer sepal4 (2.5-3 mm.) only half as long as inner sepals; leaves 6-10 cm.

Outer sepals (5-5 mm.) more than half as long as inner sepals. 51. B. simulans.

Leaves 2-5 cm., slightly veined above, reticulate below; pedicek 5-9 mm. 52. B. silzricola.

Page 60: Berberis and Mahonia : A taxonomic revision

60 L. W. A. AHRENDT

KEY 13 (continued) Leaves 7-10 cm., reticulate both sides; pedicels 10-15 mm. 53. B. &&lor.

54. B . s u b a c u m i ~ . Stems date; leavee 6-9 cm.; pedicels 1.5-2-5 cm. Leaves not reticulate, paler below.

Leaf margins with 4 4 spinulee distant 4-8 mm. ; flowers fescicled 15-40; pedicels

Leaf -gins with 12-25 spinules distant 2-3 mm. ; flowers fascicled 2-6; pedicels 15-28 Lnm. 55. B. kptopoda.

6-10 mIn. 56. B. rnrahdudu.

(44) Berberis deinacantha Schneid., Fedde Rep. Sp. Now. 46, 259, 1939; Mitt. Dtsch.

Shrub 4-6 ft. Stem very stout, not venuculose. Iraternodes (2-) 3-5 (-7) cm. Spines sulcate. k w m 4-12 x 1-2-3 cm. ; oblong-elliptic, acute; petiole 2-4 mm.; margins 20-30 spinose (0.5-1-5 mm.)-serrulate (0.5 mm.), distant 3-5 mm.; openly reticulate; slightly lustrous; slightly paler below. Flowers deep yellow, fascicled 6-12 (-15). Rdkek 12-15 (-20) mm. Prophylls 2 x 2 mm. outer sepals 4 x 3.5 mm., ovate. Inner sepal8 5.5 x 5 mm., obovate-orbicular. Petals 4-5 mm., oblong-obovate emarginate. &wit (immature) 7 x 2.5 mm., ellipsoid, pruinose.

W. Yunnan: Chien-chuan-Melong divide, 9O00-10,OOO ft., 1923, Forrest 23556 Type, K); Mar. 1923, McLaren D 50 (K). Not cultivated.

Dendr. Qes. 55, 39, t. 5, 1942. (Fig. 24)

(45) Berberis valida (Schneid.) Schneid., 1.c. 55, 40, 1942; syn. B. d e i v n t h u var.

Stem not, or scaxcely, verruculose. Irate& 4 4 cm. Leaves with margins 18-26- spinose (1-1.5 mm.)-serruhte, distant 2 - 5 4 mm. ; oblong-elliptic, acute, to 7 x 2-25 cm., fairly closely reticulate. Pediceh 6-12 mm. Otherwise as in (44).

S.E. Szechuan: Ning Yuen fu, fl. 16 Apr. 1914, Schneider 918 (Type); Huili chou, 25 Mar. 1914, Schneider 517 (K); 1932, Tsai 50684. Not cultivated.

valida Schneid., F& Rep. Sp. Nov. 46,260,1939

(46) Berbeds arguta (Franch.) Schneid., B d . Herb. Boissier (2), 8, 197, 1908; syn. B. d d k m forms arguta Ranch., BuU. Soc. Bot. Fr. 33, 388,1886; P1. Delavay. 38, 1889

Lmwm (34%) 6-13 x (1-) 1-2-3 cm., lanceolate, acuminate, subsessile; margins 30-60 spinose (0-5-1-5 mm.)-subserrulate, distant 1-1-5 (-2-5) mm.; both sides closely reti- culate, concolorow. Peokels fascicled 10-15, and 12-15 nun. long. Flowers and fruit unknown.

Yunnan: Mt. Tsang-chan, and mountain woods at Tchenfongchan by Takouan, 4 May 1882, Delavay 2354 (Type, K). Not cultivated.

(47) Berberis fallaciosa Schneid., Fedde Rep. Sp. Nov. 46, 258, 1939 Leaves 3-7 x 1-14 cm., elliptic-lanceolate, subwuminate; petiole 2-4 mm.; margins

s u b a p p r d 15-30 (-40)-spinulose (0-5 mm.)-sedate (0.5 mm.), distant 1-5-2.5 mm. ; closely reticulate; paler below. Flowers fascicled 2-5. Pedkek 10-15 mm. Prophylls lx0*7mm., ovate, subacute. Outer sepals 3-5x3mm. Inner sepals 5x4mm., all obovate. Petala 4 x 2.25 mm., oblong-obovate, emarginate; base slightly clawed with lanceolate, approximate, or even partly contiguous, &;lands, 0.8 x 0.4 mm. Stan~en~ 3 mm., slightly produced, subapiculate. 0tm.b 2. Berries 8 mm., obovoid.

S.E. Szechuan: Nanchuctn, S.E. of Chunking fu, fr. Rosthorn 68 (Type); fl. 20 May 1928, Fang 842 (K). Not cultivated.

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Berberis and Mahonia 61

(48) Berberis aristato-serrulata Kayata, Ic . PI. Forms., 3, 13, fig. 5, 1913 Stem slender, sulcate. Internodes 5-6 cm. Leaves 2.5-5 x 04-1.7 cm. ; rhombo-

elliptic to lanceolate, acute, aristate ; petiole 2 4 mm. ; margins 2040-spinose (2-2.5 mm.)- serrulate, distant 1-5-2-5 mm. ; reticulate, unobtrusively above, clearly elevated below. Flowers fascicled 10-15. Pedicels c . 1 cm. Prophylls 2.5 mm., oblong-ovate, acuminate. Outer and inner sepals equal, 5-5 x 4 mm., obovate with base truncate and 1-5 mm. broad. Petals 5 x 2.5 mm., obovate-spathulate, apex incised; basal glands 1 mm. Ovules 2, with stipes as long as, or up to twice as long as, the ovule.

Formosa: Central Mts., Apr. 1910, Hayata, s.n. (Type). Not cultivated.

(49) Berberis acuminata Franch., Bull. Soc. Bot. Fr., 33, 387, 1886, non Veitch, nec

Internodes 3-5-8 cm. Leaves 7 x 1-2 cm. to 15x 3.5 cm., or 18x 3-3.3 cm.; narrowly elliptic, slenderly acuminate; petiole 2-5 mm. ; margins 24-40-spinose (0.5-1-5 mm.)- subserrulate, distant (2-) 3-5 (-7) mm. Flowers fascicled 6-10 (-15). Pediceb stout (1-) 1.5-2-5 (-3) cm. Prophylb lanceolate, acuminate. Outer sepals 4-5 mm., ovate. Inner sepals 5-6 mm., obovate. Petals 4-5 mm., narrowly obovate, emarginate. Owuks 2-3, shortly stipitate. Berries black, epruinose or scarcely pruinose, 9 x 6 mm.

Yunnan: woods, Tchen-fong-chan, fl. May 1882, Delavay 494 (Type); Mt. Omei, 7500ft., Wilson (Veitch) 3158 (K). Not cultivated. Plants so named are B . veitchii, which differs in its coarsely toothed grey-green enervate leaves, which are paler below, its few-flowered fascicles, its more numerous ovules (M), and its conspicuously pruinose berries. B . acuminata superficially resembles the Sikkim B. insignis tong2oensis which differs in its complete lack of spines, its non-reticulate leaves which are paler below, its much larger fascicles of short pedicelled flowers, its more numerous ovules ( P 5 ) , and its shortly stylose berries.

(50) Berberis delavayi Schneid., Sargent, PI. Wils. 1, 364, 1913 Stems slightly verruculose. Spines slender, 3-fld. Leaves 3.3-5.5 x 1-14 cm., ovate to

oblong-ovate, acute, mucronate; petiole 2-5 mm. ; margins 7-1 1-spinose (1 mm.)-serrate (1 mm.), distant 4-5 mm.; openly reticulate, lustrous; concolorous below. Ovules 3. Berries ovoid, 9 x 6 mm., pruinose blue, eatylose.

Stapf

GY 1 3 ~ Flowers fascicled 6 1 5 ; pedicels 1-2 cm. Flowers fascicled 2-5; pedicels 1.5-2.5 cm.

(50u) Berberis delavayi var. delavayi

50a. B. delavayi var. delavayi. 50b. var. wachinenais.

Yunnan: Houang-le-pin, 12 Mar. 1883, Delavay 485 (Type) ; Fang-yang-chang, fr. 10 Oct. 1887, Delavay, s.n. (K). According to Dr Schneider, Lkiang-fu, 6 Apr. 1922, Kingdon-Ward 5023. Not cultivated. For plants in cultivation named B. delauuyi, see B . dumicolu which differs in its closely spinose thicker leaves with hypoderm and close reticulation, its fewer, 1 (-2), ovules, and stylose berries. B. delavayi was originally associated by its author with B. hookeri wiridis from Sikkim, but this has very many ovules (6-9), much larger berries (12-15 mm.), and sulcate stems.

(50b) Berberis delavayi var. wachinensis Ahrendt, J. Bot., Lond. 79 (Suppl.), 33,

Leaves with 4-7 marginal spinose (15-2 mm.) serrations (0-5 mm.), distant 3-6 mm. W. Szechuan: Muli, Wachin, 10,OOO ft., Yu 14401 (Type, E). Common shrub, 2-3 ft.,

1941

with berry greyish black. Not cultivated.

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62 L. W. A. AERENDT

(51) Berberis simulan~ Schneid., Fedde Rep. Sp. Nov. 46, 258, 1939 Shrub erect, 10 ft. Spines weak, 5-10 mm. Leaves 6-10 x 2-4 cm.; ovate-elliptic, sub-

acuminate; petiole 2-6 mm.; margins with 15-30 spreading spindose (0-250-5 =.)- semtions (0.5 mm.), & a t 2-5 mm.; below yellow-green. Flowers fascicled 2-3. P d k e k slender, 12-20 mm. ProphyuS ovate, acute. Outer se@ 3 x 1-5 mm., elliptic. Inner sepals 6 x 4 mm., elliptic-obovate. Petals 5 x 2.75 mm., oblong-obovate, slightly emaqimte; baae clawed. 0vd.a 2. Style absent.

S.W. Szechuan: Mt. Omei, 19 Apr. 1922, Yu 414 (Type, EL Not cultivated.

(52) Berberis silvicola Schneid., Sargent, P1. Wils. 3, 438, 1917 Shrub 1-2 ft. Intenodes 36-55 cm. Spines absent or very weak, 3-6 (-10) mm., very

slender. Leaeres with petiole 1-3 mm. ; maxghs 12-16-spinose (1-1-5 mm.)-serrate (0-5 mm.), distant 2-4 (-6) mm.; above subenemate or with very few faint lateral veins; 5-7 (-9) mm. ProphyUs 2-5 x 1 mm., ovate, acute. Stumm not or scarcely produced, below distinctly veined to subreticulate. Flowers fascicled 2-5. Pedicels slender. Style short.

KEY 1 3 ~ Leavea broad, 1 : 2-2.5; mar@ with 12-16 spbules; pedicels 5-8 nun.

Leaves narrow, 1 : 5-6; margins with 5-10 spinulea; pedicels 8-12 mm. 52a. B. BilwicOla var. silvicola.

528. var. amgwtata.

(52a) Berberis silvicola var. silvicola Leaves 2247 x 10-18 mm., elliptic, acute. Outer sepals 4 x 1-75 mm., ovate, obtuse.

Inner sepals 6 x 3-5 mm., obovate. P& 6 x 3 mm., obovate, incised, cuneate. Ovules 2. W. Huph: King-shan-hsien, fl. 31 May 1907, Wilson 2879 (Type, K). In woods,

6000-8000 ft. Not cultivated.

(52b) Berberis silvicola var. angustata Ahrendt, var.nov., f o b multo angustioribus Sterns tinely sulcate to subterete, not verruculose. I&& 2-3 cm. Spines 3-fld., to

11 mm. h u e s 25-55 x 5-10 mm., lanceolate to narrowly oblong-elliptic, subacuminate. Outer sepals 3-5 x 1.5 mm., ovate, acuminate. Inner sepals 6 x 3 mm., oblong-elliptic. Petals 4 x 2-75 mm., oblong-elliptic. Ovules 3.

Szechuan: Tien-chuan-hsien, 6300ft., fl. 20 Apr. 1936, Chu 2368 (Type, E). Not cultivated.

(53) Berberis bicolor Uveill6, Fedde Rep. Sp. Nov. 9, 454, 1911 Shrub 3-5ft. Internodes 2-5-3-5 cm. Spines 3-fld., slender, 10-15mm. Leaves

7-10 x 2-26 cm., elliptic to elliptic-lanceulate, acuminate, subsessile; margins 15-20- spinose (1 mm.)-subsemulate, distant 3-5 mm. ; both sides concolornus green, with lateral and marginal veins visible, and elevated below, finely and leas conspicuously reticulate. Flowers fascicled 3-6. Pedicels 10-12 mm. Prophylls 2 x 1 mm., acute. Outer se& 3 x 1.75 mm., ovate, acute. Inner sepals 4-5 x 3-5 mm., obovate. Petals 3.75 x 2 mm., obovate, entire. StamRns 3 mm., not or scarcely produced, truncate. Ovu2es 2.

Kweichou: Ma-jo, fl. with flowers red without, white within, very rare, May 1908, Cavalerie 3043 (Type, K). Not cultivated.

(54) Berberis subacuminata Schneid., Sarg. PI. Wilson, 1, 363, 1913 Shrub 3-5 ft. Sterns sulcate. Intern& 3-6 cm.; Spines absent, or 1-3-fld., 4-9 mm.

Leaves 4.5-9 x 1.4-2.2 cm., elliptic-lanceolate, subacuminate; petiole 2-5 mm.; margins

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Berberis and Mahonia 63

8-15-spinulose (0-25-0.5 mm.)-subserrulate, distant 3-5 (-8) mm. ; fairly closely finely reticulate; concolorous below. Fl4nuers fascicled 2-6. P e d i d (I-&) 2-2-5 cm. Prophylls 2 x 1.5 mm. Outer sepals 4 x 2 mm. Inner sepah 7 x 5 mm., obovate. Petals 5 x 2.5 mm., obovate, emarginate; base clawed with glands 1.5 x 0.5 mm., lanceolate, acuminate, approximate, submarginal. Sturnem 4.5 mm., not or scarcely produced, truncate. ovules 2-3.

Yunnan: 5000 ft., Henry 13267 (Type, K). Not cultivated.

(55) Berberis leptopoda Ahrendt, J. Bot., Lond., 79 (Suppl.), 33, 24 July 1941 Stems shoots subsulcate; mature terete. Intermodes 2 4 (-7) cm. Spines 3-fld., deep

yellow, stout, 1.4-2.3 cm. Leaves 3.3-6.4 x 0.9-1-8 cm. ; oblong-elliptic to ovate, acute, sessile; margins 4-9 spinose (1-1-5 mm.)-serrate (1-1-5 mm.), distant 4-8 mm. ; both sides openly veined, not reticulate; below much paler green. Flowers fascicled (S-) 8-18 ( 4 0 ) . Pediuds red, very slender, 16-2-5 cm. Prophylls 2-5 x 2 mm., ovate, subacute. Outer se& 4 x 3.25 mm., oblong-elliptic. Inner se& 6 x 4-25 mm., obovate. Petals 4.5 x 3 mm., spathulate-obovate, entire or slightly retuse; base cuneate with oblong, approximate or contiguous, glands 1.2 x 0.5 mm. Sturnem 3 mm., produced, apiculate. ovules 3, shortly stipitate. Xtyb absent.

S.E. Tibet: Pachakshiri district, Lalung, 7000ft., fl. 29 Apr. 1938, Ludlow, Sherriff & Taylor 3697 (Type, BM). ‘In clearings of forest and along river banks, 3 4 ft. ; flowers bright yellow, calyx red; leaves pale green below.’ Not cultivated.

(56) Berberis xanthoclada Schneid., Fedde Rep. 8 p . Nov. 46, 261, 1939 Stem angled. Spines concolorous, terete. Leaves 6 7 - 5 x 1-5-2-9 em., ovate to elliptic,

acute, mucronate ; margins with 12-25 spreading spinose-serrations (0-5-2 mm.), distant 2-3 mm.; distinctly openly veined both sides; much paler below. P e d k h fascicled 2-6, thick, 7-10 mm. Ovules 3. Berries 8-9 mm., ellipsoid, black, epruinose, shortly stylose.

Kweichou: Lao shan, Fang Ching shan, 5-7 ft., 30 Sept. 1931, Stewart, Chiao t Cheou 482 (Type, Am. Arb.). Not cultivated.

Subsect. Barandanae Schneid., Bull. Herb. Boissier ( 2 ) , 5 , 401, 1905 Stems very sulcate (except in B. xanthoxyh), glabrous; mature yellow. Intermodes

(1.5-) 2-5 em., (except in B. alpiwh). Spines 3-fld. (I-) 1.5-2 (-3) cm. Leaves enervate, slightly veined, or a t most openly and indistinctly reticulate; below epruinose, green, paler except where noted. Ovules 2 4 . Fruit stylose (except in B. xanthoxylon).

Philippine Islands, Java, Sumatra, Formosa.

KEY 14, to Subsect. Barandanae (Maps 9 and 13) Flowers fascicled 3-20; leaves 3-10 cm., thick, with hypoderm.

Stems slightly angled to subterete; leaves 5-10 cm.; ovules 4; fruit estylose.

Stems very sulcate ; leaves 3-6 cm. ; fruit stylose. 57. B. xanthoxyylon.

Stems not verruculose ; flowers fascicled 3-9 ; pedicels 18-25 mm. ; prophyls and outer sepals acute to subacute; petals longer than inner sepals (4 mm.); stamens apiculate; ovules 4; fruit 9-10 mm. 58. B. barandann.

Stems verruculose; flowers fascicled 10-20; pedicels 5-12 nun.; prophyls and outer sepals acuminate; petals shorter than inner sepals (7-9) mm. ; stamens truncate ; ovules 2-3 ; fruit 6-7 nun. 59. B. kawakamii.

60. B. alpicola. Flowers solitary; leaves 1-3 cm., thin, without hypoderm.

Page 64: Berberis and Mahonia : A taxonomic revision

64 L. W. A. AHRENDT

(57) Berberis xanthoxylon Haaskarl ex Schneid., Oat. Bot. 2. 67, 27, 1918; Ahrendt, J . Bot., M., 79 (Suppl.), 26, 1941

Stems not verruculose. Leaves thick, below at fht p&ose white, finauy green, epruinose. Pedi& 1-2 cm. Owda 4. Fruit estylose. According to Lindley, this species was introduced to cultivation by Lobb and stood three winters before 1850, without shelter, growing to 4-5 ft., and being expected to reach 10 ft. B. zanWlozylon haa been confused with, but is quite distinct from: (1) B. wauichiana DC., from Nepal, which has solitary ovules, larger fascicled, short pedicels, petals longer than inner sepals and short style; (2) B. m n i p r u m Ahrendt, from Mmipur, which has a similar infiort?scence, but M e r s in its thin leaves, without hypoderm and with close reticulation, its large flowers and large blue pruinose fruit; and (3) B. jdhm Schneid., from Hupeh, which M e r s in its scarcely veined leaves, solitary ovules, and fruit with style and conspicuous white bloom. The plate, Bot. Mag. 9283, entitled B. x a n t h ~ ~ ~ l o n was actually made from material of the type plant of B. jdknue which was erroneously described aa having ‘about 3 odes’, instead of 1 ovule. B. xun$?wxybn is not in cultivation in any of its varieties. It is the southernmost species of this section.

KEY 148, Leaves typically entire, rarely with 2-7 spindes. Leaves typically with 8-24 sphdes; fruit obovoid, 8 x 4 nun.

57a. B. xuntlwxylorr vm. xunthoxybm.

Leaves densely spinom, distant ( 1 . k ) 2-3 (4) mm. ; fruit black, epruinoSe.

Leaves sparsely spinose, distant 3-6 mm.; fruit pruinose, blue. 57 b. var. junghuhnhna.

570. var. au-nb.

(57 a) Berberis xanthoxylon var. xanthoxylon h u e s with thick cuticle, and hypoderm of small thin-wded cells; (3.6) 5-9 x (I-%)

1-8-2.5 cm. ; oblong-elliptic, acute, mucronate; margins subrevolute, entire, or occasion- d y 2-7-spindose (0.25 mm.)-subsedate; above distinctly veined, scarcely (or very openly and indistiiictly) reticulate; below with lateral veins (and often open reticu- lation) elevated, and more finely closely reticulate. Pediah 10-17 mm., fa-acicled 7-16.

Java : Zollinger 2325 (Type, BM).

(576) Berberis xanthoxylon var. junghuMana Ahrendt, J. Bot., M., 79 (Suppl.), 26,1941 ; syn. B. holyida Junghuhn, non Gay; syn. B. jummicu Miq., Ms.

Stem subterete pale reddish yellow. Leaves to 10 x 2.5 om., acute; petiole 4-6 111111. ; margins 15-24-spinose (1-1-5 mm.)-sedate; above openly, often indistinctly reticu- late; below with lateral veins and often open reticulation clear, closer reticulation fine and faint. Flowers 8-15-fld.

Java: 2000-3000 ft., according to Junghuhn; 8000 ft., fr. 30 Dec. 1912, Leeuwen- Reijnvaan 1180 (Type, K); Paaoeran, 1935-7, van Steenis 7141 (K); 1802-18, Horsfield I l l 6 (K).

(57c) Berberis xanthoxylon var. sumatranica Ahrendt, 1.c. 79, (Suppl.), 27, 1941 Stems slightly angled-sulcate. Leaves 5-7 x 1.7-2-2 cm., oblong-elliptic; petiole

c. 3 mm. ; margins 8-11-spinulose (0.5-1 m m . ) - s e d t e (0-25-0.5 mm.) ; above in- distinctly veined; below with distinct lateral veins, often openly reticulate. Fluwers fascicled (7-) 10-15. P d W 15-20mm. Prcvphylds 1 . 5 ~ 1 mm. Outer sepals 3 . 5 ~ 2.25 mm. Inner 5epal.s 5-6 x 4 mm., obovate. Pet& 4.5 x 3 mm., obovate, scarcely or shortly incised; glands separate, oblong-elliptic, placed near the base of the claw. Stumm 3.5 mm., produced, retuse-truncate.

Page 65: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 65

Sumatra: Korinchi Peak, fl., fr. 10 May 1914, Robinson & Kloss, s.n. (Type, BM). ‘Thorny shrub with pale yellow flowers.’

(58) Berberis barandana Vidal, Rev. P1. Vmc. Filip. 45, 1886 Stem not, or scarcely verruculose. Leaves 2-5 x 0.7-1.7 cm. ; elliptic, acute, aristate;

margins slightly undulate and revolute, 6-9 aristate (1-3 mm.)-dentate (1-3 mm.) ; in- distinctly veined, the lateral and marginal veins being sometimes slightly visible: above lustrous; below dull subconcolorous epapillose. Fbwers fascicled 3-7 (-9). Pedkd.8 18-25 mm. Prophylls 2.5 x 1.5 mm. Outer sepaI.8 3 x 2 mm. Inner sepals 4 x 2-5 mm., oblong-obovate. Stamens 3 mm. produced, apiculate. Berries black, epruinose, 9-10 x 7-8 mm., excluding style 0.5-0.75 mm.

Philippine Islands: Prov. Benguet, Lepanto district, Luzon, fl. Vidal, s.n. 1911 (Type, K); Luzon, Mt. Data, Loher 59 (K); Mt. Data, Merrill4607 (K). Not cultivated.

(59) Berberis kawakamii Hayata, Materials for a Flora of Formosa ( J . Imp. Coll. Sci., Tokyo, 30, Art. l ) , 24, 1911; Schneid., 0st. Bot. 2. 67, 140, 1918.

KEY 1 4 ~ Leaves reticulate; inner sepals lanceolate, acuminrrte; petals entire, duplicate; fruit

Leaves not reticulate; inner sepals elliptic, obtuse; petals alternate, incised; fruit epruinose. 59a. B. luamkamii var. hwkamii.

pruinose. 59a. var. j m s a n a .

(59a) Berberis kawakamii var. kawakamii Shrub M f t . , erect, densely branched. Stem very pale whitish yellow. Spines

1-3 cm., not sulcate. Leaves 3-3-5 x 1 cm. to 5-6 x 2-2.7 cm., elliptic, ovate, or obovate; margins (%) 6 7 (-12)-spinose (1-2 mm.)-serrulate (0.5 mm.), distant 3-6 mm.; openly reticulate both sides, with veins impressed above and prominent below ; veinlets pro- minent on both sides; paler green below. F h e r s fascicled 10-15.. Pedicek 5-12 mm. Prophylls, outer sepals and inner sepals (the latter 9 mm. long) all lanceolate, acuminate. Petals 4.5 mm., oblong. Ovules shortly stipitate. Stcclnens slightly produced, truncate. Berries 6 x 4.5 mm.

Formosa: Prov. Kagi, Mt. Morrison, Oct. 1 W , Kawakami 1941, Type; Arisan 7600 ft., 1918, Wilson 9720 (K); 8100 ft., 1914, Faurie 447 (BM); Prov. Nanto, Muska, 7000-10,000 ft., 1918, Wilson 10060 (K). I have not seen this variety in cultivation.

(59 b) Berberis kawakamii var. formosana (Ahrendt) Ahrendt, sepalis interioribus obtusis, petalis longioribus, fructibus valde pruinosis; syn. B . f o m s a n a Ahrendt, J . Bot., Lond. 79 (Suppl.), 24,1941; ‘B. kuwakumii’ Seals, Bot. Mag. 163, t. 9622, 1942

Spines sulcate, stout. Leaves 30-50 x 7-12 mm., elliptic to obovate-elliptic; with slight elevated venation both sides, often subenervate, sometimes with the lateral and marginal veins visible; slightly lustrous deep green; below paler yellow-green. Flowers fascicled (&) 14-20. Pedicels 5-9 mm. Prophylk pale lanceolate, acuminate, 2.5 x 1 mm. Outer sepals 3.5 x 1.75 mm., ovate, subacuminate, reddish. Inner sepak 8 x 3.5 mm., narrowly oblong-elliptic, obtuse. Petals 7 x 3-25 mm., narrowly oblong-obovate, apex narrowed and incised; base cuneate with separate ovate acute glands, 1 x 0.5 mm. Berries thickly pruinose mauve; style short.

Formosa: Prov. Kagi, Arisan, fr. 25 Oct. 1918, 2-6 ft., on grassy mountains, Wilson 10910 (Type, K); 1933, seed only, Yashiroda 100127.

Cultivated (from Yashiroda’s seed) : fr. 18 Nov. 1939. 5 JOURN. LINN. S0C.-BOTANY, VOL. LVII

Page 66: Berberis and Mahonia : A taxonomic revision

66 L. W. A. AERENDT A very handsome compact evergreen shrub of moderate size, remarkable for its unusual

flower structure; the flower buds preaent an UDuSually prickly appearance due to the points of the unfolding acuminate prophyls and sepals; on account of the thick mauve bloom, this is an ornamental benrying plant.

(60) Berberis alpicola Schneid., Fedde Rep. Sp. Nov. 46, 253, 1939; Mitt. Dtsch.

Shrub small, compact. Intemwde.~ 7-10 (-12) mm. Stem not or scarcely verruculose. Spines concolorous, terete. L a v e s 10-20 (-25) x 5-8 (-9) mm. ; elliptic to oblong-elliptic; margins with 2-8 spinose (1 mm.) serrations (1 mm.), distant 2-3 mm.; lateral veins visible above, distinctly elevated below; both sides concolorous lustrous green, finely papillose. Flowers solitmy. Pedicels 12-16 mm. owtdes 4, sessile. Berries 6 x 4 mm., ellipsoid, black, epruinose; style 0.3 mm.

Formosa: Prov. Kagi, Mt. Morrison, 12,OOO ft., fr. 24 Oct. 1918, Wilson 10952 (Type, BM, K). Not cultivated.

Den&. h. 55,26,1942

Subsect. Euwallichianae Schneid., BuU. Herb. Boissier (Z), 5, 402, 1905

Internodes 2-5 (-8) cm. Stem mature yellow (except only in B. sargentiana). Spines 3-fld.; occasionally only 6-15 mm.: mostly 1-5-2-5 cm. (unless otherwise recorded), sometimea stout, 3-6 cm. Leaves with hypoderm, thick, often rigid; (2-) 5-12 cm. long; mostly with 10-40 marginal spinules; venation nearly always distinct, and mostly reti- culate (unless otherwise recorded) ; below epruinose green (except only in B. bhutunensis). Fbwers 7-10 (-12) mm. diameter (except only B. mdcrropetah, 12-15 mm. diameter). Stanzens generally (and unless otherwise recorded), not, or scarcely produced, truncate. Ovules 1 (-2), rarely 3, generally (and unless otherwise recorded) sessile or subsessile. Fmit 7-10 mm., mostly (and unless recorded otherwise), stylose.

Nepal, Bhutan, Yunnan, Huph.

KEY 15, to Subsect. Euwallichianae (Maps 5 and 10)

Pedicels 3-4 mm. ; petals shorter than outer sepals. Pedicels (5) 8-20 mm. ; petals longer than outer sepals.

61. B. micropetala.

62. B. zanlanscianenais. Leaves very m o w , 1 : 9-12. Leaves narrow leas, 1 : 3-6.

Fruit conspicuously pruinose; flowers fascicled 8-20; pedicels (5-) 9-15 mm. ; leaves green below.

veined; petals shorter than inner sepals; stamens not produced. Leaves with 5-20 marginal spinules, distant 3-5 mm.; subenervate or openly

F h i t eatylose; petals entire; stems subterete. 63d. B. x omienaia. Fruit stylose; petals emarginate; stems angled.

Leaves 3-7 cm.; with 5-10 marginal spinules; fluit pruinose blue. Leaves m o w , 1 : 5-6, Gnely toothed; pedicels 5-10 mm.

Leaves broader, 1 : 3, coersely toothed; pedicels 10-15 mm. 63c. B. x wintone&.

63. B. bergmunniue. Leaves 6-10 cm., with 12-20 marginal sphules.

Leaves obovate, elliptic or oblong; acute; fruit pruinose white. 64. B. jd iaw.

Leaves lrtnceolate, muminate; fruit pruinose blue. 65. B. willearn. Leaves with 2 0 4 0 marginal spinules, distant 1-5-2-5mm.; closely reticulate;

petals longer than inner sepals; stamens produced, subapiculate. 66. B. durnkolu.

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Berberis and Mahonie 67

Fruit black, epruinose. Leaves green, epruinose below.

Flowers fascicled 10-25; fruit stylose; stems angled, Petals emarginate, shorter than inner sepals; ovules sessile.

Petals entire, longer than inner sepals; ovules stipitate. 67. B. jerdinandi-whrgii.

68. B. wallichiunan.

69. B. aargentiana. 70. B. recurvatu.

Leaves white, pruinose below. 71. B. bhutanenaie.

Flowers fascicled 3-10; fruit estylose; stems tmete. Stems dark red; leaves closely reticulate; ovules 1-2. Stems pale yellow; leaves openly veined; ovules 3.

(61) Berberis micropetala Schneid., F& Rep. Sp. Nou. 46, 266, 1939 Internodes 2-5-3.5 cm. Spines sulcate. Leaves 2.54.5 x 0.9-1.8 cm.; oblong-elliptic,

acute; margins 14-spinose (1 mm.)-serrate (1 mm.), distant 5-12 mm. ; openly veined; below at first grey, pruinose, finally brownish green and epruinose. Plowers fascicled 4-5. Pedicels 3 4 m m . Outer sepals 4 . 5 ~ 3 mm. H e d h sep& 7-8x5mm. Inner sepals 8-9 x 5 mm. Petals 4 x 2.5 mm., very pale or whitish, emarginate. Stamem apiculate. Style absent. Fruit unknown.

Assam: Naga Hills, Saramati ridge, 9500 ft., fl. 2 Mar. 1935, Bor 2921 (Type, K). Not cultivated.

(62) Berberis zanlanscianensis Pamp., Nuovo Ciorn. Bot. Itccl., N.S., 22, 293, 1915 I~eaues 6-7 cm. x 5-8 mm., or 4-9 cm. x &12 mm.; narrowly lanceolate; margins

10-15-spinose (1-5-2 mm.)-serrate (1 mm.), distant 2 - 5 4 mm.; venation slight; below paler green. Flowers fascicled 12-30. Pedicels (8-) 10-20 mm.

Hupeh: Zanlanscian, Apr. 1912, Silvestri 4110 (Type), 4110a. Yunnan: Imen district, 6-15 ft. tall, fl. Henry 10618 (K). Not cultivated.

(63) Berberis bergmanniae Schneid., Sargent, PI. Wils. 1, 362, 1913 Shrub erect, 5-7 ft. Stem angled, sparsely minutely brown-vermculose. Spines stout,

rigid, subconcolorous, subterete or plane. Leaves very thick, rigid ; oblong-elliptic to elliptic; acute, mucronate, subsessile; subenervate or with 3-7 faint lateral veins; below very lustrous, enervate much paler than above. Flowers fascicled (&) 10-15 (-20). Pedicels (7-) 10-15 mm. Prophylls 1.5 x 0.75 mm., acuminate. Outer sepals 5.5 x 3-5 mm., ovate. Inner sepals 7 x 5 mm., obovate. Petals 6 x 5 mm., obovate, apex rounded but narrowly incised; base slightly clawed, glands 1 x 0.5 mm. Stamens 4-5 mm. Berries 8 x 6 mm., oblong-ellipsoid, pruinose blue, shortly stylose.

KEY 15a Leaves lustrous above. Leaves dull above.

63a. B. bergmanniae var. bergmanniae. 63 b. B. bergmnniae var. acanthophylla.

(63 a ) Berberis bergmanniae var. bergmanniae

serrate, distant 3-7 mm. Leaves (3-) 5-8 (-10) x (1-) 1.6-2.5 (-3) cm.; margins 5-10 (-15)-spinose (1-2 mm.)-

W. Szechuan: Mountain sides, Chingchihsien, 5000ft., Nov. 1908, Wilson 2876

Cultivated: fl. 13 May 1942; fr. 13 Jan. 1943. A large evergreen with stout, s t 8 branches, valuable for its remarkable rigid leaves; but somewhat tender in Oxford- shire.

(Type, K).

5-2

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68 I,. W. A. ~ E N D T

(63b) Berberis bergmanniae var. acanthophylla Schneid., 1.c.

6-10 mm. h u m to 6.5 x 2.3 cm., margins 6-10-spinose (2-3 mm.)-denhte (2-3 mm.), distant

W. Szechuan: west of, and near Wenchuanhsien, 6000-8OOOft., 1910, Wilson 4149

Cultivated: without flowers, or fruit, Nov. 1936. A more tender variety than the ( T w , K).

former, and one which seldom flowers, but has even more rigid leaves.

(63c) Berberis x wintonensis Ahrendt, Card. Chron. (3) 108, 56, 1940, nom.nud. ; hybr. aff. B. bergmanniue, sed foliis multo angustioribus

Stem partly sulwte, partly terete, stout, black-vemculose. S p i m 1-2.5 cm. h u e s 30-60 x 6-12 mm. ; oblong-elliptic; margins 9-13-spinose (0-5-1 mm.)-sedate (0.25 mm.), distant 2-3 mm.; dull grey-green; lateral veins and few veinlets visible; below subenemate, pale lustrous yellow-green. Flinuers fwcicled 10-20. Pedicel8 6-12 mm. Prophyh 2 x 1.5 mm. F’lower structure as in no. 63, but with the components alightly smaller. Berries 9 x 4 nun., pruinose blue.

Cultivated: origin in Hilliers nursery, Winchester, probably from seed of B. berg- manniue; fl. 11 Oot. 1938; fr. 18 Nov. 1940. An evergreen of dense habit, growing slowly to 6 ft., quite h d y , and very early in flower.

(63d) Berberis x oxoniensis Ahrendt, hybr.nov., aff. B. bergmanniae et B. julianae sed foliis flexibilibus acuminatis

Stem subterete to subangled. h u e s thick, flexible, 3-7 x 0-7-1.6 cm. ; lanceolate to elliptic-lanceohte, muminate; margins 10-14-spinose (1-1-5 nun.)-serrulate ; above very smooth, dull deep green, with lateral and marginal veins very faint and not elevated or impressed; below paler yellow-green, subenervate. F’lowers fascicled (8-) 10-20. Pedicel8 10-15 mm. ProphyyuS greenish, 2 x 1.5 mm. Outer sepals 6 x 4 mm., oblong-ovate. Nedkn se@ 7 x 6 mm., obovate. Inner sep& 8 x 6.5 mm., obovate. Petals (H.C.C. 2) 6 x 5 mm., obovate, entire. Stamns 4 111113. Berries ovoid, 10 x 6 mm., thickly pruinose bright blue.

Cultivated: origin at Watlington, Oxfordshire, aa a seedling of unknown antecedents: fl. 6 June 1942; fr. 6 Nov. 1940 (Type, 0). preserved out of many seedlings on account of its ornamental appearance, due to its dignified foliage and bright abundant berries. The leaves somewhat resemble those of B. imignit~ tongloensis, but m smaller and much smoother.

(64) Berberis julianae Schneid., Sargent, P1. Wils. 1, 360, 1913 Shrub dense to 7 ft. Stem angled, minutely black-vemculose. h u e s thick, rigid;

above deep, scarcely lustrous green; with lateral veins faint, veinlets few; below paler, lustrous, subenervate. Plowers fascicled 15-30. Pedi& 8-15mm. O& solitary. Fruit 7-8 x 3-5-4 mm., oblong, heavily pruinose white. Style 0-5 mm.

KEY 1 5 ~ Leavea somewhat m o w , 1 : 36-4-5. Leaves obovah, broadest at apex. Leaves oblong, broadest at middle.

Leaves broader, 1 : 2-26.

64a. B. j d k m var. jdianae. 64b. var. oblongqolia. 64c. var. patunge&s.

(64a) Berberis julianae var. julianae Internooh 2-5-5 cm. Spines concolorous, sdcate, 2-3-5 cm. Leaves 4-10 x 1-26 em.,

obovate, subacute; petiole 1 4 mm. ; margins 10-20-spinose (1-1.5 mm.)-serrate (1- 1.5 mm.), distant 3-6 mm.; dark green. Prophylk 2.5 x 1.5 mm., acute. Outer sepals

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Berberis and Mahonia 69

4-5 x 2.5 mm., ovate, acute. Inner sepals 6.5 x 3.5 mm., oblong-elliptic. Petals 5.5 x 2.75 mm., oblong-elliptic, emarginate; base clawed with oblong glands 1 x 0.4 mm.

W. Hupeh: Ichang, 3000-4000 ft., 1907, Wilson 417 (Type, K). Cultivated: the plant a t Kew, no. 285, and certain others were grown from seed of the

type collection. No. 285 was inspected and approved by Wilson (and was the actual plant figured in Bot. Mag. as B. xanthoxylon, g.v., with its undoubtedly solitary ovules described and figured as c. 3; this inaccuracy led Schneider to refer to the plant of t. 9283 as a possible hybrid, since he knew it was not B. xanthoxylon, and, having no access to the material used, was unaware of its uniovulate character). Mr W. J. Marchant distributed plants of this typical collection, but, in the first instance, wrongly named as B. xant7wxyZcna and B . wallichiana. F1. May 1939; fr. 13 Oct. 1939.

(64b) Berberis julianae var. oblongifolia Ahrendt, var.nov., foliis oblongo-ellipticis

and marginal spinules more spreading than in 64.

3170 (K).

Stems black-verruculose. Leaves to 9 x 2-2-5 cm., with lateral veins more rectangular

Hupeh: Chang-yong, 3-6 ft., 1900, Wilson (Veitch) 535 (Type, K); Ichang, fi. Henry

Cultivated: fl. May, 1940; fr. 20 Oct. 1940.

(64c) Berberis julianae var. patungensis Ahrendt, var.nov., foliis latioribus Leaves to 4.8 x 2.1 cm., margins 5-7-slenderly spinose. Hupeh: Patung hsien, thickets, 4200 ft., 1907, Wilson 2878 (Type, K). Not cultivated.

(65) Berberis willeana Schneid., Ost. Bot. 2. 67, 141, 1918 Shrub 6-8ft. Stems angled or subangled, or subterete, verruculose. Intermodes

3-8 cm. Leaves narrowly elliptic to lanceolate, subacuminate; base attenuate to petiole 3-8 mm. ; lateral veins not distinct ; above slightly lustrous grey-green ; below scarcely lustrous paler yellow-green. Pedicek 5-10 mm., slender. Ovules 1 (-2). Berries ovoid to ellipsoid, 7 x 3 mm., pruinose blue. Style short.

KXY 150 Leaves with 5-20 spinules, distant 3-7 mm. Leaves with 20-30 spinules, distant 14-26 mm.

65a. B. wiUeana var. wiUeana. 65b. var. senulata.

(65a) Berberis willeana var. willeana Stems more or less angled. Spines 1-2-3 cm., terete. Leaves (6) 6-10 (-14) x (1.2-)

1.7-2.6 (-3) cm.; margins (5-) 12-20-spinulose (0.5 mm.)-serrulate, distant 3-7 mm. Flowers fascicled 10-20.

Yunnan: 1914, Schneider 1763 (Type), 2290 (both, K).

(65b) Berberis willeana var. serrulata Schneid., Fedde Stems subterete. Spines 15-18mm.) plane. Leaves

possibly fascicled only 3-5. N. Yunnan: Pinchuan hsien, 1933, Tsai 53000, Type;

9100 ft., 1922, Rock 3214. Not cultivated.

(66) Berberis dumicola Schneid., New Flora & Silva,

Not cultivated.

Rep. Sp. Nov. 46, 245, 1939 3-2-5 x 0-8-1-3 cm. Fltnuers

between Talifu and Likiang,

10, 259, 1938; Fedde Rep. . I

Sp. Nov. 46, 249, 1939; Mitt. Dtsch. Dendr. Ges. 55, 17, 1942 Shrub 4-5 ft. Stems terete; shoots bright red; mature pale yellow, finely verruculose

(dark brown). Internooks 3-7 cm. Spines 6-15 mm., plane. Leaves fairly thick, stiff;

Page 70: Berberis and Mahonia : A taxonomic revision

70 L. W. A. AERENDT 5-9.5 x 1-14 cm. ; elliptic ; acuminate ; petiole 2-4 mm. ; margins 20-40-spinose (1.5- 2.5 mm.)-serrate (1 mm.); above convex, slightly lustrous with close impressed reti- culation; below much paler green, scarcely lustrous, less visibly reticulate. Plowers fascicled (8-) 10-20-30. Pedicels 8-15 mm., slender. PrqvhyuS red, 1-2 x 0-7-1.5 mm., acute. ozlter 8epL.9 reddish, 3 x 2-5 mm., oblong-ovate, obtuse. Inner sepals (H.C.C. 63/1) 6 x 5 mm., oblong-obovate. Petals 6-5 x 5 mm., oblong-obovate (H.C.C. 1/1-2/1), apex rounded and incised-emarginate; base conspicuously clawed with ovate, separate, marginal glands 1-5 x 0-7 m., situated near the base of the claw. Stanzens 4.5 mm., produced, subapichte. ovules (1-2), with stipes as long as the ovule. Bewie.s 7 x 4 mm., pruinose blue; style 0-5 mm.

N.W. Yunnan: Mekong-Yangtze divide, E. flank of Li-ti-ping, 10,000ft., Forrest 19474 (Type, K); Mekong-Salween divide, 1914, Forrest 13295 (K).

Cultivated: (i) from Forrest 19474, Type, erroneously distributed as ‘B. Perdinandi- Coburgii’; fl. 15 Apr. 1943; fr. 26 Oct. 1938; (ii) from Forrest 13295, erroneously distri- buted aa ‘B. dela0ra;Yi’; fl. 21 May 1942, 18 May 1941; fr. 20 Oct. 1939. A small or moderate-sized handsome leaved evergreen, particularly attractive in spring and early summer when the shoots and new leaves are bright red.

(67) Berberis ferdinandi-coburgii Schneid., Sargent, PI. Wils. 1,364, 1913 Shrub 3-5 ft. Stem angled-sulcate, verruculose. Intemdes 2 - 5 4 cm. Spines 3-fld.

Leaves thick, rigid; above lustrous, closely reticulate; below sublustrous, paler with scarcely elevated open reticulation. h r s fascicled (8-) 12-25. Pediixh slender, 1-2 cm. Prophylls red, 1-5 x 0.75 mm. Stamens 3 mm. Ovules solitary, subsessile.

KEY 1 5 ~ Leaves with 35-60 marginal spinulm, 0.5-2 mm. distant, oblanceolab.

Leaves with 10-35 spinules distant 2 - 5 4 mm.; 1anceolat.e. 67a. B. ferdimndi-coburgii var. ferdinandi-wburgii.

67 b. B. ferdimndi-wburgii var. vernalis.

(67 a) Berberis ferdinandi-coburgii var. ferdinandi-coburgii Spines weak, 3-fld., 6-10 mm., sulcate. Leaves 5-13 x (1-) 1-5-2.5 (-3) cm., elliptic-

oblanceolate, acute; petiole 2-4 mm. ; margins 35-6O-spinose (0.5-1 nun.)-sedate, distant 1-2 (-2.5) mm. Outer sepals 2.25 x 0.75 mm., lanceolate acute. Inner sepals 5 x 3 mm., ovate. Petals 4.5 x 2.75 mm., narrowly obovate, emarginate; base clawed with oblong glands 0.9 x 0-5 mm. Bewies black, epruinose, 8 x 5 mm., ellipsoid, or ovoid, stylose.

Yunnan: Mengtze, woods, 5500ft., fr. Henry 10257 (Type, K); Mengtze, Kochin range, 6000-7000 ft., 24 Dec. 1895, Hancock 448 (K). Not cultivated.

(67b) Berberis ferdinandi-coburgii var. vernalis Schneid., FecZde Rep. Sp. Nov. 46,

Spines 15-25 mm., stout, terete. Leaves lanceolate to elliptic-lanceolate, or oblong- elliptic; 3-12-5 x 1-26 (-3-5) cm.; margins (10-) 16-24 (-35)-spinose (0.6-1 mm.)-ser- d a t e (0.5 mm.), distant (2-) 2-5-3-5 (4) mm. Outer sepals 3 x 1.3 mm., ovate, acute. Inner s e p h 4 x 2.5 mm. Petals 3.25 x 1.75 mm. Berries black, epruinose, 5 x 4 mm., ellipsoid.

Yunnan (central to east): 1914, Schneider 226 (Type); Maire 1738; 9 Oct. 1938, Yu 17968 (E). Common at 9ooo ft. Shrub 3-5 ft. Not cultivated.

249, 1939

Page 71: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 71

(68) Berberis wallichiana DC, Prodr. 1, 107, 1824; Wall, PI. As. Rar. 3, 23, t. 243,

Shrub erect, 6-10 ft. Stems subangled. Spines slender, slightly sulcate. Leaves (3.5)- 6-12 x (0*7-) 2-24 cm.; oblong-lanceolate; acute to subacute, or even obtuse; subsessile ; margins 12-25-spinose (05-1 mm.)-subsedate, distant 3-6 mm. ; above sublustrous; with 5-9 lateral veins and part of the marginal vein distinct, but with reticulation indistinct ; below paler, lustrous, epapillose. Flowers fascicled (8-) 10-20. Pedicels stout, 6-10 mm. Prophylls 1 x 1 mm., acuminate. Outer sepals 3 x 1.6 mm., oblong-obovate. Inner sepals 4-5-5 x 3 mm. Petals 4.75-5-25 x 2.5 mm., obovate, entire; basal glands separate lanceolate. O v d a solitary, shortly stipitate. Stamens 3.5-4 mm. Berries 8-9 x 4-5 mm., excluding style 0-5-0.75 mm., ovoid, black, epruinose.

1832

Nepal: Wallich, Mt. Sheopur, 1819 (Type, K). Not cultivated. An interesting species, being the fist discovered of this section, and lying towards its

western range; furthermore, the type seems to be the only collection. I n cultivation, the name is commonly misused for (i) B. hookeri, from Sikkim, which differs in its smaller, thinner leaves and small habit, its few-flowered fascicles, and very large flowers and estylose berries with 6-9 ovules, and also, but much less frequently for (ii) B . julianae which has estylose berries, heavily pruinose white.

(69) Berberis sargentiana Schneid., Sargent, PI. W i b . 1, 359, 1913 Shrub dense, 6-7 ft. Stems terete; mature dark red, mostly not, but in parts sparsely

and distinctly, black-verruculose; shoots (and young leaves) bright red. Internodes 3-6 cm. Spines stout, sulcate, (2-) 3-6 cm. Leaves thick, but flexible, 4-10 x 1.5- 2.5 cm. ; oblong-elliptic, subsessile, acute ; margins 15-25-spinose (1-2 mm.)-serrate (1-2 mm.), distant 2-5-4 mm.; above lustrous dark green with close impressed reticu- lation; below much paler dull yellow-green, with much slighter venation. Flowers (H.C.C. 64/2), fascicled (3-) 4-8 (-10). Pedicels 1-2 cm. Prophylls red, margined white, 2 x 2 mm. Outer sepals (H.C.C. 663/1), 3.5 x 3 mm., ovate, subacute, with a red band along the middle. Median sepals 5 x 4-5 mm., rhombo-elliptic. Inner sepals 6.5 x 5 mm., obovate. Petals (H.C.C. 64/3), 6 x 4.5 mm., obovate: acutely emarginate with rounded lobes: base cuneate with contiguous discolorous, orange glands, 1.75 x 1 mm. Stamens 4-5 mm. Ovules 1-2. Berries 6-5-8 x 4.5-6 mm. black, epruinose, estylose, oblong or oblong-ellipsoid.

W. Hupeh: Hsing-shan-hsien, woods, 4000-5000 ft., 1907, Wilson 564 (Type, K). Cultivated: 0. 17 May 1941, 15 May 1943,4 May 1944; fr. 6 Jan. 1944. An evergreen with very h e , dignified, foliage, and rather unusual pale greenish

flowers; it is slightly tender in Oxfordshire.

(70) Berberis recurvata Ahrendt, &rd. Chron. (3) 124, 175, 1948 Shrub 7-8 ft. Stems black-verruculose. Internodes 3-6 cm. Spines concolorous, stout,

3-6 cm. Leaves thick, flexible, convex; 4-10 cm. x 8-18 mm.; elliptic, oblong-elliptic, or ovate-elliptic; petiole 1-2 mm. ; margins 10-20-spinose (1-1.5 mm.)-serrate (1-1.5 mm.), distant 2-5 mm.; above dull light green, with lateral and marginal veins indistinct; below dull paler yellow-green, subenervate. Flowers fascicled 5-10. Pedicels 8-15 mm. Prophylls 3 x 2 mm., red, acute. Outer sepals 3-5 x 2.5 mm., red. Median sepals 5.5 x 3-5 mm., ovate-elliptic obtuse. Inner sepds 6 x 4 mm., obovate. Petals 5 x 3-25 mm., obovate-cuneabe, acutely emarginate ; basal glands oblong, separate, concolorous, in- conspicuous 0.6 x 0.5 mm., non-marginal. Stamens 3.75 mm. Ovules 3. Berries black, epruinose, estylose, ellipsoid, 7 x 4 mm.

W. Hupeh: Fang hsien woodlands, not common, 5200ft., 1907, Wilson (Veitch) 555 (K).

Page 72: Berberis and Mahonia : A taxonomic revision

72 L. W. A. AHBENDT Cultivated: fl. 14 Apr. 1939, 6 Apr. 1945; fr. 15 Nov. 1938, 21 Dec. 1944 (Type, 0).

In cultivation more common than B. sargentiana, under which name it is often in- accurately distributed, but from which it differs in its mature pale yellow stems; dull, slightly veined, convex leaves, and inconspicuous glands.

(71) Berberis bhutanensis ahrendt, J. Bot., Lond., 79 (Suppl.), 17, 1941; s p . B.

Stem subterete, sparsely verruculose. Leaves 3-6 x 0.7-1.5 cm., elliptic-oblong, acute; petiole 1-2 mm. ; margins 4-8 (-10)-spinose (1-3 mm.)-serrate (1-2 mm.), distant (3-) 4-6 (-8) mm.; above slightly lustrous with lateral veins distinct, but reticulation indistinct; below pruinose white with lateral veins elevated. Flowers fascicled 3-6. Pedicels 12-20 mm. ProphyUs 1.5 x 1 mm., ovate, acute. Outer s e w 3-5 x 2 mm., ovate, obtuse. Inner sepals 6-5 x 4 mm., elliptic-obovate. Pet& 5 x 3 nun., obovate. XtacmRn~ 4 mm. O& 1 (-2). Berries black, epruinose, obovoid, 8-9 x 4 - 5 4 mm., with short distinct style.

Bhutan: 1838, GrifEth 1741 (Type, K); SO00 ft., shrub 4-6 ft., in dense rain forest, 16 Apr. 1949, Ludlow, S h e d & Hicks, 18712 (BM).

N. Burma: Nam Tam& valley, 7000-8000ft., 14 Oct. 1937, Kingdon-Ward 13421 (BM). Not cultivated.

dichiancl var. @& Hook. f. et Thorns. 1855, non Boissier

Subsect. Leves Ahrendt, J. Bot., M . 79 (Suppl.), 18, 1941 Stem mature yellow (except only in B. bre&epda), sparsely, distinctly verruculose.

Internodes (1 .6) 2-3 (4) cm., rarely 4-7 cm. S p i w generally 3-0d., (1-) 1-5-2-5 (-3) cm., rarely very weak and only 2-8 mm., or very strong and 3-5 cm. Leaves thick, with hypoderm, enervahe or subenervate. Fhoers fascicled 4 4 0 . ovules solitary (very rarely 2). Be& mostly black and epruinose (unless otherwiee recorded).

Bhutan, Szechuan, Yunnan, Form-.

ICEY 16, to Subsect. Leves (Maps 5 and 10) Mature stems dark red. 72. B.bre&e@. Mature stems yellow.

Stems very sulcate. Leave3 below concolorous green.

Fruit pruinom; style distinct (0.5 m.). 73. B. pluninii. Fruit estylose. 74. B. aubpteroclada.

75. B.liophylla. Leaves below much paler green. stems terete.

Leaves pruinose, white, below. Fruit pruinose, stylose. 76. B. zuangii. Fruit epruhose, estylose. 77. B.hlocmspedon.

Leaves epruinose, green, below; fruit black, epruinose. Spines 1-5-5 cm. ; inner sepals narrowly lanceolate; fruit estylose. 78. B. levis. Spines P 8 nun. ; inner sepals obovate ; fruit stylose. 79. B.aubcoriacea.

(72) Berberis brevisepala Hayata, Ic. PI. F'mnzos., 3, 14, 1913; Schneid., oat. Bot. 2. 67,136,1918

Stem subterete. Spines c. 1 cm. Leaves to 5-5 x 1-5 cm., narrowly obovate, acute, mucronate; attenuate-cuneate at baae; margins with 3-6 spinose teeth, distant 6 1 0 mm.; below pruinose grey. F'lowes fascicled 7-8. Pedimls c. 1 cm. Outer sepals reddish 2-2.5 mm., ovate, acute. Inner sepals 5 mm., ovate. Petale 2 - 6 4 mm., elliptic, emar- ginate; base scarcely clawed, glands small. Ovules solitary, sessile. Stamens c. 1-5 mm., produced, apiculate. Bemdea estylose.

Formosa: Mt. Morrison, Mori, 8.n. 1910 (Type). Not cultivated.

Page 73: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 73

(73) Berberis poluninii Ahrendt, sp.nov., aff. B. soulianue sed ramulis magis sulcatis,

Stem sulcate, mature yellow. Internodes 1.5-2-5 cm. Spines 3-tld., sulcate 7-16 mm. Leaves to 47 x 12 mm., oblong-elliptic; margins 5-9-spinose (1-2 mm.)-serrate (1-2 mm.), distant 3-5 mm., subrevolute; thick, rigid; above almost dull; below epruinose, con- colorous; subenervate, or with very few parts of veins slightly elevated both sides. Flowers fascicled 3-7. Pedicels 5-7 mm. Ovules solitary. Flower structure unknown. Berries pruinose blue. Style 0.5 mm.

Central Nepal: Compaling, 3 miles east of Timure, 10,500 ft., 2 July 1949, Polunin 746 (Type, BM). Not cultivated. To be compared with the Kansu B. soulieam which has less angled stems, longer leaves and spines and 2-3 ovulea; with the Assam B. micro- petah which has leaves broader, openly veined and partly pruinose below ; and with the Nepal B . Wallichianu with angled stems, much larger veined leaves, larger fascicles of flowers, and black epruinose fruit.

spinis foliisque brevioribus, ovulis solitariis distinguenda

(74) Berberis subpteroclada Ahrendt, J. Bot., Lond. 79 (Suppl.), 21, 1941

Style absent. Stem verruculose, yellow, very sulcate. Leaves thick, subenervate. Ovules solitary.

KEY 168, Pedicels S 2 . 5 cm.

Leaves 3-6 cm., lanceolate; flowers fascicled 44.

Leaves 1.5-4 (-6) cm., oblong-elliptic ; flowers fascicled 9-20. 74a. B. mbpteroclada var. subpteroclada.

74b. B. aubpteroclada var. h p a r . Pedicels 5-10 mm. 74c. B. subpteroekcda var, m i d p e s .

(74a) Berberis subpteroclada var. subpteroclada Spines 1-25 cm., plane. Leave.!? 3-6 x 0-3-1.3 cm., lanceolate, acuminate, sessile;

margins 6-IO-spinose (1-1.5 mm.)-serrulate (0.5 mm.), distant 3-8 mm. Plowers fascicled 4-9. Ped* slender. Prophylls 1.5-2 x 1.5 mm. Outer sepals 4 x 3 mm., ovate to obovate. Inner sepale 7 x 4 mm., obovate. Petub 5.25 x 3 mm., obovate, apex retuse with acute lobes; glands 1.5 x 0.5 mm., lanceolate, acuminate, 1.5 x 0-5 mm.

Central Bhutan: Chendebi, 9500 ft., 14 May 1937, Ludlow & SherriE 3045 (Type, BM). Shrub 5-8 ft., in rhododendron and conifer forest; buds tinged red; 1949, from central and east Bhutan, Ludlow, S h e d , & Hicks 18939, 18916, 20532, 20585.

(746) Berberis subpteroclada var. impar Ahrendt, 1.c. 21, 1941

elliptic. Flowers fascicled 9-20. Otherwise as in 73.

1949, 8000-9000 ft., Ludlow, SherriE & Hicks 16053, 18654 (BM).

Leaves 1 - 5 4 (-6) cm. x 0-3-0.6 cm., margins with spinulw distant 2 - 5 4 mm., oblong-

Central Bhutan: Chendebi, 9500 ft., 14 May 1937, Ludlow BE SherriE 3046 (Type, BM) ;

(74c) Berberis subpteroclada var. minoripes Ahrendt, var.nov., pedicellis multo

b%rns partly sulcate, partly terete. Spines 2-3 cm., stout, 1-2 mm., broad a t base, shallowly sulcate. Leaves 25-42 x 7-11 mm., narrowly obovate margins 10-15-spinose (1.5-2 mm.)-serrulate (0.5 mm.), distant 1.5-2 mm.; almost dull above, enervate; below with occasionally a few veins and veinlets elevated. Flowers fascicled 3-10. Pediceb 5-8 (-10) mm., slender. Outer sepals 2.5 x 1.5 mm., ovate, subacute. Inner sepals 4.5 x

brevioribus

Page 74: Berberis and Mahonia : A taxonomic revision

74 L. W. A. AHILENDT 3.25 mm., obovate. Petals 3-5 x 2 mm., obovate, entire. Stamens 3 m., produced,

Bhutan: Ha Dzong, 9OOO ft., 4 Apr. 1949, Ludlow, S h e d & Hicks 16015 (Type, BM). ‘Shrub 3-4ft., growing on dryish hill slopes in Piaw forest. I think this is the most viciously spiny Berberis I have every pricked myself with’; 8OOO ft., Ludlow, S h e d & Hicks 16203 (BM). !Chis species is not cultivated in any of its varieties. It may be noted that some of the species from N. and N.W. China have spines longer and as rigid and stout, or even more so, e.g. B. iwndkm, B. &inuwntha and B. circumewata.

truncate. ovules solitary.

(75) Berberis liophylla Schneid., F& Rep. Sp. Nov. 46,247,1939 Shrub 4-5 ft. Spines plane below, 3-fld., 1-3 cm. Stem markedly sulcate.

KEY 1 6 ~ Flowers fascicled 5-10. Flowers fascicled 10-25.

75a. B. Ziophylla var. lwphylla. 75b. B. liophyUa vm. wnglobatu.

(754 Berberis liophylla var. liophylla Leaves 2-6 x 1-2 cm., elliptic to lanceolate, acute; base attenuate; margins 10-25-

spinose (0.5-1.5 mm.)-serrulate, distant 2-3 mm., lustrous above. Flowers fascicled 5-10. Pedida 8-12 mm. Prophylb ovate, acute. outer sepals ovate, elliptic, subacute. Inner sepals 4.5 mm., obovate. Petals 4 mm., obovate, emargjnate, clawed. Ovuk8 solitary. Fruit (immature) 5-6 mm., epruinose, estylose.

S.E. Szechuan: between See-guei and Tschankio, fl. 22 Apr. 1914, Schneider 969

N.E. Yunnan: 1932, Tsai 50979. (TYPf-4.

(75b) Berberis liophylla var. conglobata Ahrendt, varnov., fasciculis multifloratis Stem with young shoots very sulcate ; mature less sulcate. I?eternodes 3-5 cm. Leaves to

4.5 x 1.5cm.,namwlyelliptic to oblong-elliptic;margjns 15-20-spinose (l-l-5mm.)-serru- late (0.5 mm.), distant 14-26 mm.; venation above, branched to reticulate; below dull paler yellow-green, with lahral veins clearly elevated, and reticulation close but in- distinct. Flowers fascicled 10-25. Pedicels 6-15 mm. Prophylls ovate, subacuminate, 1-5-2 x 0.74 mm. Outer sepals 3 x 1.3 mm., ovate, acute. Inner sepals 4 x2-25 mm., obovate. Petakr 3 x 1-75 mm., obovate. Stanaens 2-75-3 mm., slightly produced, retuse- truncate. ovules solitary, subsessile. Style in ovary, apparently very short.

Yunnan: vicinity of Yunnan-sen, Nov. 1906, Maire 1738 (Type, K). The specimens of this number in the herbarium of the British Museum differ in having their 7 x 2 cm., elliptic leaves entire (rarely with 1-2 spinules). Accordingly, the above description is based entirely on the Kew specimen. The species is not in cultivation in either of its varieties.

(76) Berberis wangii Schneid., Fedde Rep. Sp. Nov. 46,246,1939 Spines concolorous, terete. Leaves to 5 x 1.2 cm., elliptic to elliptic-lanceolata, acute;

cuneate, subsetwile; margins 5-8-spinose (0-5-1 mm.)-sedate (0.5 mm.), distant 3-5 mm. ; above slightly lustrous. Flowers fascicled 8-12. Pedicels 12-18 mm.

Yunnan: Kunming, Apr. 1935, Wang 62639 (Type). Not cultivated.

(77) Berberis holocraspedon Ahrendt, J. Bot., Lond. 79 (Suppl.), 22, 1941 Internodes 4-7 cm. Spines often absent, always weak, 4-7 mm., 3-fld. Leaves 4-8 x

14!-2.5cm.; oblong-elliptic; margins very slightly revolute, entire; above pale grey

Page 75: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 75 green, subenervate ; below densely papiUose, with at most 5-8 lateral veins and part of the marginal vein faintly visible. Flowers fascicled (&) 8-16. Pedicels red, stout, 8-13 mm. Berries 8-9 x 5-5.5 mm., oblong-ovoid, black, epruinose. W. Yunnan: Shunning, among thickets on open slope, 9750 ft., fr. 22 Nov. 1938, Yu

18228 (Type, E). ‘Common spiny shrub, 5-8 ft., fruit black.’ Not cultivated.

(78) Berberis levis Franch., Bull. SOC. Bot. Fr. 33,386, 1886; PI. Delavay. 37, 1889

below. Flowers fascicled (1%) 15-25 (-40). Ovules solitary. Spines stout, 2-5-5 em., plane below. Leaves subenervate, dull above, paler green

KEY 16c Leaves narrow (1 : 6-10, 10-12 x 1-14 cm. ; pedicels 15-20 111113.

Leaves broader (1 : 4-6), 3-6 cm. x 6-9 mm.; pedicels (4-) 6-10 (-12) 111111. 7 8 a . B . levis var. levis.

78b. B. le& vsr. brachyphylla.

(78a) Berberis levis var. levis

s e d a t e . Flowers 6-8 mm. diameter. Pedicels 1-5-2 cm.

cultivated. For plants so called see B. atrocurpa, or rarely B. soulieuna.

Leaves lanceolate, acuminate, mucronate ; entire or sparsely finely spinulose-sub-

Yunnan: Mao-kou-chan, above Tapintze, fl. 23 Apr. 1883, Delavay 495 (Type). Not

(78 b) Berberis levis var. brachyphylla Ahrendt, var.nov., foliis conspicue brevi-

Inte.rno&s 2 4 cm. Leaves lanceolate subacuminate subsessile ; margina 5-10-spinose (1 mm.)-subsermlate, distant 3-6 mm. Flowers 5-7 mm. diameter. Prophylls narrowly lanceolate. Outer sepals lanceolate. Inner sepals 5 x 1.5 mm. Petals 5-6 x 2.5 mm., cuneate, apex incised. Ovules solitary, stipitate. Stamens 4-5 mm., slightly produced, obtuse.

Yunnan: Pi-iou-se, above Tapintze, near Tali, 7200 ft., fl. 14 Apr. 1884, Delavay 893; Hee-chan-men, fr. 12 Oct. 1885, Delavay 993, both cited by Franchet as ‘as second form with leaves shorter and relatively broader’; Sungkwei Pass from the Langkong valley, 9ooO-10,OOO ft,, fl. Apr. 1906, Forrest 2012 (Type, K): ‘Shady situations amongst scrub on ascent’; west flank of Sungkwei Pass, 9OOO ft., fl. 21 May 1921, Forrest 19393 (K): ‘2-3 ft., in open situations amongst scrub’. Not cultivated.

oribus

(79) Berberis subcoriacea Ahrendt, Gard. Chron. (3), 105, 371, 1939 Shrub 4-6 ft. Stems variable, subterete to subsulcate. Internodes 2-3 cm. Spines

weak, 4-8 mm. Leaves 3-3-6.5 cm. x 6-13 mm.; lanceolate to oblong-lanceolate; acumi- nate, mucronate; margins 8-13-spinose (1-1-5 mm.)-subsedate, distant 4-7 mm. ; with only lateral veins and few veinlets visible; dull green; slightly paler below. Flowers 10-12 mm. diameter, fascicled 10-24. Pedicels slender 1-2 cm. ; bracts 2 x 0.5 mm. ProphyUs 2.5 x 1 mm., oblong-ovate. Outer s e p k 4-5 x 3 mm., obovate. Inner sepals 7 x 4 mm., obovate. Petuls 5.5 x 4 mm., obovate, incised; basal glands 1-5 x 0.5 mm. Stamens 5 mm., not produced. Ovules 1 (44, mostly stipitate, occasionally subsessile. Berries 9-10 x 5.5-6 mm., with style 0.25 mm.; oblong or oblong-obovoid, black, epruinose.

Yunnan: Fang-yang-Tchang, by Mo-so-yn, fr. 21 Oct. 1887, Delavay s.n. (K). Cultivated (from reputed Forreat seed): fl., fr. 1938, Wisley (Type). Plant dead by

1946.

Page 76: Berberis and Mahonia : A taxonomic revision

76 L. W. A. AHRENDT

Subsect. Soulieanae Ahrendt, J. Bot., Lond. 79 (Suppl.), 23, 1941 Stems glabrous, mostly terete (unless otherwise recorded) ; mature always pale yellow.

Internodes (1-) 2 4 (-7) cm. Slyines stout, mostly 3-6 cm., rarely 1-26 em., very rarely weaker and to 4-9 mm. Leaves thick, with hypoderm, generally flexible (but very rigid in B. 8 O d k M ) ; narrow (1: (2.7%) 443; generally enervate (unless otherwise recorded). Fi2rux?r8 10-14 mm. diameter (where known), and with pet& shorter than inner sepals. Shmw generally not, or scarcely, produced (unless otherwise recorded). Ovules (1-) 2-3, sessile or subsessile. Fruit 5-7 (-10) mm. long, always stylose, pruinose (white, mauve or blue) except only in B. atrocarpa; and B. 8chneideriana. Style 0.5-1 mm. S.E. Tibet, Y~nnan, N.E. Szechuan, N. Hupeh, Shensi, Chekiang, Anhwei.

KEY 17, to Subsect. Soulieanae (Maps 5 and 11) Flowers fascicled 12-25; pedicels slender, 2-3 cm.; ovules 2-3. Flowers fascicled 2-12 and/or pedicels 4-15 mm.

80. B. mairei.

Fruit black, epruinose; pedicels 5-10 mm.; ovules 2-3. Stems terete; spines 3-10 mm.; leaves broad, 1:2-5-3; flowers fascicled 15-25.

stems sdcete ; spines 2-5-6 cm. ; leaves narrow, 1 : 4-5; flowers faecicled 5-9. 81. B. achneid&a?la.

82. B. atrocarpa. .h& p&OSe.

Leaves below green, epruinose; ovules 2-3. Stems sulcate; leaves narrow, 1 : 5-8, rigid deep yellow-green, slightly lustrous

above; flowers fascicled 8-20; pedimls 5-9 mm.; fruit pruinose mauve;

Stems terete; leaves broader, 1:3-5-4, more flexible dull, grey-green; flowers fascicled 3-8; pedicels 10-15 mm.; fruit pruinose white; style 1 mm.

StyIe 0-5 mm. 83. B. 8&&3?la.

84. B. lempergimm. Leaves below pruinose, white or grey.

Leavea plane; margins with 2-16 fine teeth. Leaves 2 4 cm. ; broad, 1 : 2-3-5; fruit pruinose white.

Leaves grey below; mar+ with 7-16 spinules; ovules solitary.

Leaves white below; margins with 2-4 spinules; ovules 2-3.

Leaves 4-7 cm., m o w , 1:5; ovules 2-3; ii-uit pruinose blue.

85. B.nenwrosa.

86. B. chingii.

87. B. W d . 88. B. hibberdha. Leavea revolute; margins with 1-2 coarse teeth.

(80) Berberie mairei Ahrendt, sp.nov., fasciculis multifloratis, pedicellis gracilibus

Internodes 3-4cm. Spines stout, 3-0d., 12-22mm., plane or subterete. Leaves 3-6.3 x 0-6-1-3 cm., scarcely rigid, with distinct hypodem of thick-walIed cells in two series; narrowly elliptic; apex and base equally acute; margins 10-18spinose (1.5- 2 mm.)-serrate (1 mm.), distant 3-6 mm.; lateral veins and veinleta elevated on both sides; above very lustrous bright green; below dull paler green.

Yuman: Yunnan-sen, Maire 1999 (Type, K). Not cultivated.

elongatis, ramulis teretibus, foliis crassis

(81) Berberis schneideriana Ahrendt, sp.nov., aff. B. at--, sed ramulis tere- tibus, spinis brevioribus gradioribusque, foliis latioribus, floribus pluribus dis- tinguenda

Spines 3-fld., weak, 3-10 mm. Leav,es with hypoderm of continuous thick-walled cells; 2-3.2 x 0-7-1.3 cm.: elliptic to oblong-elliptic; acute, subsessile; margins 10-16-spinose (1 mm.)-subsemdate, distant 2-3 mm.; subenervate, or at moat a few lateral veins, in-

Page 77: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 77

distinct above, below scarcely elevated; above dull grey-green; below at first slightly pruinose, ultimately dull paler yellow green.

Yunnan: 21 Feb. 1914, Schneider 164 (Type K). Not cultivated.

(82) Berberis atrocarpa Schneid., Sargent, PI. Wils. 3, 437, 1917 Shrub dense, 6-8 ft. Stem finely black-verruculose, angled to sulcate. Internodes

(2-) 3 4 (-5) cm. Spines concolorous, plane, 3-0d. Leuvea thick, flexible; above snb- enervate, or at most a few lateral veins; oblanceolate to narrowly oblong-elliptic; above scarcely lustrous green; below much paler, dull.

KEY 17a Leaves bright green; pedicels 5-9 mm.

Leaves with spreading maxginal spinules. 82a. B. atrocurpa var. atrocarpa. Leaves entire or with appresed spinules. 826. B. atrocurpa var. subintegra.

82c. B. atroeccrpa var. bngipes. Leaves dark green; pedicels 1-2 cm.

(82a) Berberis atrocarpa var. atrocarpa Spines stout (2-) 3-5 (-6) cm. Leaves 3 x 0.7 cm. to 7 x 1.2-1.4 cm.; margins 5-10-

apinose (1-2 mm.)-serrate (1-2 mm.), distant 4-8 mm. F k r s fascicled 5-10. Pedicels 5-10 mm. Prophylls 2 x 1.5 mm. Outer sepals 4 x 2 mm., oblong-obovate. Inner sepals 7 x 4 mm., obovate. Petals 6 x 4-5 mm., obovate, rounded, deeply narrowly incised; base cuneate with glands 0.8x0.5mm. Stamens 4mm. Ovules sessile, or very shortly stipitate. Berries black, epruinose, ovoid, 5 x 4 mm. Style 0.5 mm.

W. Szechuan: Mupin, fr. Nov. 1908, Wilson 1284 (Type, K). Cultivated: from type collection, fl. May 1938; fr. Dec. 1938 (K).

(82b) Berberis atrocarpa var. subintegra Ahrendt, var.nov., foliis subintegris, vel

Spines 2-3 cm. h v w c. 5 cm. x 8.5 mm., narrowly oblong-elliptic; subentire or

Szechuan: Lushan hsien, 3600 ft., among bushes, fr. 18 Oct. 1936, Chu 4040 (Type, E).

Cultivated: 0.20 May 1942,6 Apr. 1945; fr. 23 Nov. 1940,4 Nov. 1942,26 Nov. 1943

spinulis pauculis appressis

margins 6-9-spinose (1 mm.)-serrate (0.25-0.5 mm.), distant 4-6 mm. Style 0.75 mm.

‘Shrub 6-12 ft. ; leaves light green beneath.’

(Type, 0).

(82c) Berberis atrocarpa var. longipes Ahrendt, var.nov., a var. utrocarpa foliis

Spines 1-2.2 cm. Flowers fascicled 8-18. Pedicels 1-2 cm., very slender. Berries

Cultivated: fl. Apr. 1944; fr. 26 Dee. 1944 (Type, Oj.

atroviridibus, pedicellis multo longioribus distinguenda

5-6 x 3-3.5 mm.

(83) Berberis soulieana Schneid., Sargent, PI. Wils. 3, 437, 1917; syn. B. stemphylla

Shrub erect, 4-5 x 2-3 ft. Stems sulcate, concolorous-vermculose. Spines 3-fld., con- colorous, stout, plane. Leaves thick, very rigid, 3-8 cm. x 6-9 mm., or to 7-5 x 1.3 cm., oblong, oblong-ovate, oblong-elliptic, or oblong-obovate ; petiole 1-2 mm. ; margins (P) 6-10 (-18)-spinose (1-14 mm.)-serrate (1 mm.), distant. 3-7 mm. ; enervate; above scarcely lustrous deep bright green ; below much paler creamy green, epruinose, Snely papillose. Pedicek slender, 5-9 mm. Berries pruinose, mauve. Style 0.5 mm.

Hance, J. Bot., Lond., 30, 257, 1882, non Lindl.

Page 78: Berberis and Mahonia : A taxonomic revision

78 L. W. A. AERENDT

&Y 1 7 ~ Leavea enervate or subenervate; flowers fascicled (6) 10-20; h i t obovoid, 7 x 5 mm.

Leaves with lateral veins and veinletr, clear; flowers fascicled 2-6; fruit subglobose, 83a. B. sodieuw var. aoulieana.

5-6 x 5 mm. 83 b. €3. sodieana var. pzueimruata.

(83u) Berberis soulieana var. soulIeana Spines 2-5 cm. long, 1.5 mm. broad at base. Pwphylk red, 2.5 x 1.2 mm. Outer sepals

5 x 3.5 mm., oblong-elliptic. Inner s e p h 7 x 5 mm., obovate. Petals 6 x 5 mm., obovate; acutely emarginate with rounded lobes; base cuneate with glands 0.8 x 0-6 mm. O d e s sessile.

N.E. Szechuan: 1897, seed, Farges, s.n.; 1881, Parker 21774 (Type of B. stemphylla Hance) .

N. Hupeh: Mt. N g m , July 1899, Hugh, s.n. (K). Cultivated: Vilmorin 1005 (from Farges’s seed) (Type); ah, fl. 13 May 1942; fr.

23 Nov. 1940.

(83 b) Berberis soulieana var. paucinervata Ahrendt, va,r.nov., folk venis lateralibus venulisque distincte visibilibus, fasciculis f l o m tantum 2-6, fructibus oblongo- subglobosis distinguenda

Spines 1-2 cm. Otherwise as in (83u), except as distinguished in Key 1 7 ~ . Central Shensi: 1916, Licent 2588 ( T e e s , BM, K). Not cultivated.

(84) Berberis lempergiana Ahrendt, &rd. Chrm. (3) 109, 101, 1941; Sealy, Bot.

Stems sparsely black-verruculose. Spines concolorous, subterete, stout, 2-4 cm. Leaves thick, slightly rigid, 4 - 8 ~ 1-3-2 cm.; oblong-elliptic; margins 6-10 (-14)-spinose (1 mm.)- serrate (1 mm.), distant P10 mm.; above d d , grey-green or blue-green, with 4-5 lateral veins and part of the marginal vein indistinctly visible; below scarcely lustrous paler yellow-green, subenervate. Flowers H.C.C. 211. PmphyUs 1-3 x 0.6 mm., ovate, red. Outer sepals 2.5 x 2 mm., ovate-elliptic. Median sepals 5-5 x 4 mm., similar. Inner sepals 7 x 6 m., obovate. Petals 6 x 4 mm., oblong-obovate, acutely emarginate with rounded lobes; base cuneate with contiguous glands 1 x 0.8 mm. Stumm 4-4-5 mm., much produced, truncate. Ovules subsessile. Berries 9 x 5.5 mm., excluding style 1 mm., oblong-ellipsoid, pruinose white.

Mug., 167, N.S., t. 90, 1950

Chekiang: Prentai mountains, 3000 ft., 1889, Faber 179; Hickin, s.n. (both, K). Cultivated: fl. 9 May 1939, 27 Apr. 1941, 13 May 1942; fr. 26 Oct. 1940 (Type, 0).

Raised at Hillier’s nursery by seed sent to Dr Fritz Lemperg from Nanking Botanic Garden; erroneously distributed as ‘B. eudr ie i ’ , which differs in having stems very sulcate, larger fascicles of flowers with narrower components, solitary ovules and estylose berries.

(85) Berberis nemorosa Schneid., Fedde Rep. Sp. Nov. 46, 1939 Stems angled. Internodes 1.5-3 cm. Leaves 2-4x 1-16 cm., oblong-elliptic; obtuse;

petiole 1 4 mm. ; mar* 7-16-appressed spbulose (0-5 mm.)-sedate, distant 2-3 mm.; subenervate; above lustrous; below pruinose grey. Flowers fascicled 5-7. Pedicels 12-15 mm. Ovules solitary, sessile. Berries 6-7 mm., ellipsoid, pruinose white, with short style.

E. Kwangsi: Dar Siar Ping, Miu shan, fr. 29 June 1928, Ching 6192 (Type). Wide- spread in this district, in thickets at 4500 ft. Not cultivated.

Page 79: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 79

(86) Berberis chingii Cheng, Contr. biol. Lub. Sci. SOC. China (Bot. Ser.), no. 3, ix, 191, 192, fig. 17 (Dec. 1934); syn. Berberis sp. Rehder, J . A m . Arb. 8, 108, 1927; syn. B . cuvaleriei var. pruinosa Byhouwer, J . Am. Arb. 9, 132, 1928.

Stems subangled to subterete, closely, finely, verruculose. Spines concolorous, stout, 3-fld., 1-2.5 cm. Leaves rigid, 2-5 x O + l cm. ; oblong-elliptic; margins 24-spinose (0-5-1 mm.)-subserrulate, distant 3-6 mm., rarely entire; but few lateral veins visible; above sublustrous; below pruinose white. Flowers fascicled 8-12. Pedicels 9-12 mm. Berries ellipsoid to obovoid, pruinose white, 7 x 4 mm., excluding style 0.5 mm.

S. Anhwei: Chang-gon-shan, Wu Yuan, fr. 17 Aug. 1925, Ching 3248 (Type, K). ‘Leaves dull shining green, in thickets.’ Not cultivated.

(87) Berberis taronensis Ahrendt, J. Bot., Lond. (Suppl.), 23, 25 Apr. 1941 Stems not verruculose; shoots ihely sulcate; mature subterete. Spines 3-fld., sulcate,

5-15 mm. Leaves narrowly oblong-elliptic ; above dull bright green, subenervate. Flowers fascicled 2-8 (-15). Petals with contiguous glands at base of claw. Stamens distinctly produced, obtuse.

KEY 17c Leaves below, pruinose grey, densely papillose, with only few lateral veins visible.

Leaves below, epruinose, pale green, with few clear lateral veins, and fine reticulation.

(87 a) Berberis taronensis var. taronensis Leaves 3-7.5 x 0.6-1.5 cm. ; margins 4-9-spinose (1-2 mm.)-serrate (1 mm.), distant

4-9 mm. Pedicek 2-3 cm., slender, red. Prophylls red, 1 x 0.5 mm., ovate, acute. Outer sepals 4 x 2.5 mm. Inner sepals 6 x 4 mm., obovate. Petals similar, 5.5 x 3 mm., ovate, entire, broadly clawed. Stamens 4 mm. Berries obovoid, pruinose blue, 7 x 4 mm., ex- cluding style 0.75 mm.

S.E. Tibet: Prov. Zayul, Taron valley, Chilung, 8500 ft., fr. 5 Aug. 1938, Yu 19658 (Type, E). ‘Margin of mixed forest, casual.’

Prov. Pome, Trilung, Po-tsangpo, valley 7000 ft., 8 Apr. 1947, Ludlow, Sherriff & Elliot 12353 (BM) ; very common, 2-6 ft. Not cultivated.

87a. B. taronensis var. taronensis,

87 b. B . taronensis var. trimensis

(87 b) Berberis taronensis var. trimensis Ahrendt, var.nov., foliis subtus viridibus epruinosis, floribus majoribus

Leaves 20-50 x 6-11 mm.; margins 2-6-spinose (1 mm.)-serrulate, distant 6 1 0 mm. Prophylls 1.5 x 1 mm., ovate, acute. Outer sepals 2.5 x 1-5 mm., ovate, subacute, red. Median sepals 5 x 3 mm., oblong. Inner sepals 8 x 5 m., obovate. Petals 6.6 x 5 mm., with lanceolate glands. Stamens 5.5 mm.

S.E. Tibet: Prov. Takpo, Trimo, Nyam, Jang Chu, 11,500 ft., fl. 23 May 1947, Ludlow, SherriE & Elliot 12518 (Type, BM). Shrub, fairly common in dense wet forest. Not cultivated.

(88) Berberis hibberdiana Ahrendt, sp.nov., foliis marginibus valde revolutis e t

Stems terete, not verruculose. Internodes 2-5 cm. Spines 3-0d., 2-35 cm., fairly stout, plane below. Leaves thick, rigid, 2-3-7 x 0.8-14 cm., elliptic; margins very undulate and revolute, 1-2-spinose (2-3 mm.)-dentate (2-3 mm.), distant 8-12 mm. ; subenervate; above dull grey green, below pruinose white. F’lowers fascicled 6-10 (-12). Pedicels 12-20 mm. Prophylls 2 x 2 mm., ovate, acute. Outer sep& 2.5 x 2 mm., ovate, obtuse.

undulatis, crasse dentatis, fructibus albo-pruinosis stylosis distincta

Page 80: Berberis and Mahonia : A taxonomic revision

80 L. W. A. ~ N D T

Inner sepal9 6 x 3 mm., obovate. P e a 5.5 x 4 mm., obovate-elliptic; base clawed with ovate widely separated glands 1-2 x 0-5 mm., situated near the base of the claw. Stumem 5 mm., not produced, truncate. 0vu.b 1-2, stipitate, with stipes about half as long as the ovule. Be& pruinose white, with style 0-5 mm.

Yunnan: immature fruit, McLasen C. 103 (Type, I(). Not cultivated.

Subsect. Pruinosae Schneid., BuU. Herb. BoisSier (2), 5, 398, 1905 Stems glabrous ; mature pale yellow. Intenodes (1-5-) 2-3 (-4) cm. Spines concolorous,

more or lese stout, 1-3 cm. Leaves thickly co~ceous , with hypoderm; enervate, or with few somewhat indistinct veins. O& 1-3. Petals longer than inner sepals. Fruit estylose, heavily pruinose white.

Yunnan, Kweichou.

Stems terete; ovules 2-3. ~ ( E Y 18, to Subsect. Pruinosae (Map 11)

Stems not, or scarcely, verrucdose. Leaves above dull grey-green; margins entire, or with 1-6 spindes; outer sepals

much shorter than inner sepals; stamens not, or scarcely, produced, truncate.

Leaves lustrous yellow-green; margins with 10-20 spinules; outer sepals e q d to 89. B.pminosa.

inner sepals; stamem conspicuously produced and apiculate. 90. B. centijora.

89h. B. pminosa var. punctata. stems very sulcate; ovules solitmy. 91. B. cavakriei.

Stems conspicuously verruculose; stamens much produced.

(89) Berberis pruinosa Ranch., BuU. Soc. Bot. Fr. 33,387, 1886

estylose and heavily pruinose white. Otherwise variable. Shrub with terete stems, thick rigid leaves, pet& longer than inner sepals, and fruit

KEY 18a Stems not or scarcely verrucdose; stamens not produced.

Leaves with (0-) 1-6 m g b l teeth. Leaves white, pruinose below.

Pedicels 10-18 nun.; fruit to 7 x 3.5 mm. 89a. B. pminosa var. p i n o s a . Leaves broad, 1 : 2-26 Leaves narrow, 1 : 4.5. 89b. B. pminosa var. hgifolia.

89c. B. p i n o s a vm. brevipes. Pedicels 5-7 (-9) mm.; fruit to 7 x 5 mm. Leaves i3rmlly green, epruinose below.

Flowers fhscicled 10-20; pedicels 10-18 mm.; petals incised.

Flowers fascicled 3-9, or petals entire. 89d. B. p i n o s a var. viridifolia.

Pedicels 1-3 cm.; petals entire. Pedicels 10-18mm. 89e. B. pminosa var. b a r r k m . Pedicels 2-3 cm., slender. 89f. B. pminosa. var. tenuipw.

Pedicels 6-10 mm.; petals incised. 89g. B. pminosa var. brevifolia. Leaves with 10-20 marginal spinules. 89h. B. pminosa var. serratifolia.

89j. B. p i n o s a var. punctata. Stems conspicuously verruculose ; stamens conspicuously produced.

(89a) Berberis pruinosa var. pruinosa Shrub 3-5 ft. Spines stout, sulcate, 1-3 cm. Leaves 16-45 x 0.7-2-5 cm.; elliptic to

obovate; margins rarely entire, mostly 1-6-spinose (1-1.5 mm.)-serrate (1-2-2 mm.), distant 5-12 mm. ; subenervate, dull pale grey-green becoming slightly sublustrous with age; below pruinose white, epapillose. F k r s fascicled (%) 10-25. Pedicels 10-15 mm.

Page 81: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 81

Prophylk lanceolate, acuminate, 2 x 1 mm. Outer sepals 4 x 2 mm., oblongelliptic. Inner sepals 6.5 x 5 mm., obovate. Pet& 7 x 4 4 mm., obovate, deeply incised; base cuneate with completely basal glands, 0.9 x 0.5 mm., ovate, submarginal and contiguous. Sturnens 6 mm. Berries 6-7 x 3-3.5 mm.

Yunnan: Mo-so-yn, near Lankong, fl. Feb. 1883, fr. Nov. 1883, Delavay 493, 1861 (Type, K); 2 Mar. 1887, fl. Delavay, s.n. (K).

Cultivated: fl. 5 May 1944; fr. 28 Nov. 1944. Somewhat tender.

(89b) Berberis pruinosa var. longifolia Ahrendt, var. nov., foliis angustioribus

distant 4-7 mm. Berries with a hint of a style.

Fairly common, hardy.

Leaves 5-9 x 1-1.8 cm.; margins 4-7 (-10)-spinose (1-2 mm.), serrate (1-2 mm.),

Cultivated: origin possibly hybrid. Fl. 13 May 1942; fr. 12 Nov. 1940 (Type, 0).

( 8 9 ~ ) Berberis pruinosa var. brevipes Ahrendt, J. Bot., Limd., 79 (Suppl.), 15,26 Feb.

Spines 2-3, 5 cm., 15-2 mm. broad a t base, plane. Leaves 2-4 x 04-14 cm., elliptic; margins 2-4-spinose (2.5-3.5 mm.)-serrate (1-5-2 mm.), distant 7-12 mm. ; above slightly lustrous dark green; below pruinose white. Fknuers fascicled 10-16. Pedicels 5-7 (-9) mm.

N.W. Yunnan: Atyntze, Hungpoh, among thickets, 850-9500 ft., casual 2 ft., leaves shining above, Nov. 1938, Yu 15662 (Type, E); Plaine et colline a Tongchuan, 8000 ft., Maire 148 (K)-Arbuste epineux, buissonnent. Baiea farineuses.

1941

Cultivated : from the type collection; hardy.

(89d) Berberis pruinosa var. viridifolia Schneid., Fedde Rep. Sp. Nov. 46, 250, 1939 Spines 3-fld., sulcate, 1.2-2 cm. Leaves 2 - 4 ~ 1-2 cm.; elliptic; margins (0-) 2-4-

spinose (0.5-1.5 mm.)-serrate (0-5-1.5 mm.), distant 5-10 mm. ; above dull grey-green, subenervate ; below enervate, slightly paler yellow-green, slightly papillose. Flowers fascicled 8-15. Outer sepals 2 x 1.5 mm. ovate, acute. Inner sepals 6-5 x 4 mm. Petals 7 x 4.5 mm. ; glands ovate, acute, 1-25 x 0-5 mm. Sturnem 5 mm., not produced, rounded- truncate. Berries ellipsoid, 6 x 4 mm.

Yunnan: Plaine et coteaux a Tong chouan, 8100ft., common, Maire 398 (K). Not cultivated.

(89e) Berberis pruinosa var. barresiana Ahrendt., Kew BuU. 1939, 266

Spines stout, 2-3.5 cm. ; to 2 mm. broad at base, plane. Leaves to 7 x 2.6 cm. ; marghs S7-spinose (1 mm.)-serrate (1 mm.), distant 5-8 mm. Flowers fascicled 6-9. Outer sepals 3.5 x 2 mm. Inner sepals 5 x 4 mm. Pet& 5.25 x 2.5 mm., with narrowly obovate, acuminate glands 1.5 x 0-5 mm. Berries 6 x 4 mm.

Yunnan: near Yunnan-fu, between Shinling and Siae Ma Kiu, 1-2 ft., Schneider 82 (K), (‘seed number 18’).

Cultivated (from Kew 4775 M V , Banes): fl. May 1943; fr. Sept. 1943.

(89f) Berberis pruinosa var. tenuipes Ahrendt, var.nov., pedicellis elongatis

Spines 3-fld., concolorous, stout, to 2.5 cm. Leavu to 4 x 1.6 cm.; elliptic; mar@s 3-5-spinose (1.5 mm.)-serrate (1 mm.), distant 4-9 mm. ; lateral veins slightly elevated both sides; above slightly lustrous ; below paler yellow-green, epapillose. Flowers fascicled 4-8. Pedicells 2-3 cm., very slender. Prophylls 2 x 0.75 mm., ovate, acuminate.

gracillimisque distincta

6 JOURN. UNN. SW.-BOTANY, VOL. LVII

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82 L. W. A. AHI~ENDT

Outer 8epals 4.5 x 2.5 mm., ovate, obtuse. Inner sepals 6.5 x 3 mm., oblong-elliptic. Petals 7x3mm., entire, acute; base clawed with Oonspicuous discolorous separate marginal lanceolate glands 1.75 x 0.4 mm. Stamens 5 mm.

Yunnan: north of Yunnan-fu, between Shinling and Da-sung-shu, fl. 10 Mar. 1914, Schneider 342 (Type, K). Not cultivated.

(899) Berberis pruinosa var. brevifolia Ahrendt, Kew BuU. 1939,267, Spines weak, 1-3-fld., 5-10 mm. k z v e s 16-32 x 8-15 mm.; obovate; margias 2-7-

spinose (1 mm.)-serrate (1 mm.). Outer sepals 3.5 x 2 mm. Inner sepah 5 x 4 mm. Petals 5 x 3.5 mm. ; glands acuminate 1.2 x 0-4 mm. BerriRp 6 x 4 mm.

Cultivated: fl., fr. 1939 (K). Origin unknown. Of little intereat and pomibly hybrid.

(89h) Berberis pruinosa var. serratifolia Ahrendt, J. Bot., Lond., 79 (Suppl.),

Spines 1.5-3 cm. Leaves to 5-5 x 1-5-2-5 cm.; elliptic, subacuminate; margins 15-20- spinose (1-2 mm.)-serrate (1-2 mm.), distant 1-54 mm.; with distinct branched vena- tion. Flowers fascicled 3-7. Pedbls 10-15 mm. Prophylk 1.2 x l mm. Outer sepa.5 3 x 2 mm., oblong. Inner sepals 4 x 2 mm. Petals 4.5 x 3 mm., rounded, entire; glands lanmlate, acuminate, 1-8 mm.

15, 1941

Cultivated: fl., fr. 1939 (K). Possibly a hybrid.

(89j) Berberis pruinosa var. punctata Ahrendt, vamov., a ceteris ram& valde

Stems conspicuously verruculose. Inter9wdes 2-4 cm. Spines stout, sulcate, 12-22 mm. Leaves 3-4-5 x 1-1.5 cm. ; obovate; margins subrevolute, 3-5-spinose (2 mm.)-serrate (1-1.5 mm.), distant 5-8 mm. ; above slightly veined bright green; below pruinose white; texture thinner than in the other varieties. Flowers H.C.C. 3, fascicled 2-6. Pedi& 1-2 cm. ProphyuS yellow 1-75 x 1 mm. Outer s e p h 3 x 2 mm., oblong-ovate, acute. Inner sepah 6.5 x 5 mm., obovate. Petals 7 x 5-5 mm., obovate. Stamens conspicuously p r o d u d , 6.5-7mm., slightly swollen below the anthers, apex obtuse. Ovules 2-3. Berries 7 x 5 mm., oblong-ellipsoid.

verruculosis distinguenda

N.W. Yunnan: seed only, without locality or herbarium sheet, Yu 14938. Cultivated: (Yu 14938), fl. 7 Apr. 1943,4 May 1944; fr. 20 Nov. 1944 (Type, 0). A

very interesting but tender plant, which seems suitable for the south-western counties. My own planta died several years ago in the very harsh Oxfordshire climate.

(90) Berberis centiflora Diels, Not. Bot. Card. Edinb. 5,167, 1912 Stem not verruculose. h u e s 3-7 x 1-3 cm.; oblong-obovate; petiole 1-4 mm.;

margins 10-20-spinose (1-1.5 mm.)-serrate (1 mm.), distant 2.5-4 mm. ; lustrous yellow- green; below paler green, sublustrous. Flowers fascicled (1%) 2030. P d i d 15-25 mm. Prophylk 2 x 1-5 mm. Outer and inner sepals equal, 5.5 x 4 mm., obovate. Petals similar, 6.5 x 5 mm., slightly retuse: base slightly clawed with lanceolate acuminate glands, 2x0-5mm. Stamens 5mm., produced and shortly apiculate. ovules 2-3. Berries oblong, 7 x 6 mm.

Yunnan: E. flank of Tali range, 90oO-11,OOO ft., 1906, Forrest 4689 (Type, E). 'Shrub, 2-5 ft., in open situations in pine and rhododendron forest.'

Cultivated: fl. 12 Apr. 1939; fr. 20 Oct. 1938, 17 Nov. 1939. An attractive evergreen, sometimes confused with B. pruinosa var. viridifobia, from

which it is eaaily distinguished (as from all varieties of B. p i n m a ) by its larger outer sepals. The apicdate sfamens, and somewhat veined lustrous bright yellow-green leaves are further distinctions from B. puinosa var. viridifolia.

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Berberis and Mahonia 83

(91) Berberis cavaleriei Uveilld, Fedde Rep. 8p. Nov. 9, 454, 1911 Stem very sulcate. Internodes 2-3 cm. Spines 3-fld., concolorous, plane, 1-26 cm. ;

oblong-elliptic; apex subacute to obtuse, aristate; petiole 1-2-5 mm.; margins 6-12- spinose (0-5-1 mm.)-seda te (0.25-06 mm.), distant 2-4 mm. ; both sides subenervate, sublustrous, concolorous. Fknuers fascicled 8-20. Pedicela slender, 8-20 mm. Prophylle 2.5 x 0.6 mm., ovate, acuminate. Outer sepah 3 x 1 mm., narrowly ovate, obtuse. Inner sepals 4.5 x 1.5 mm., narrowly ovate, subacute. Petah 5.5 x 2.75 mm., ovate, obtuse, incised; basal glands lanceolate, 1.5 x 0.5 mm.

Kweichou: 1907, Cavalerie 3209 (Type, K). Not cultivated; for plants so named see B. lempergiana, which species was inaccurately distributed as B. cavakriei. Our present species is one of the few with conspicuously narrow flower components.

KEY 19, to Sects. Asiaticae and Tinctoriae combined (Maps 2, 3, 12, 13)

Internodes mostly 4-10 mm. Internodes (1-) 1.5-7 cm.

Stems pubescent, or at any rate puberulous at first. Stems always glabrous.

132. B. forskaliana

A

Stems, when mature, dark red, brown, or purple; stamens mostly not produced and

Stems, when mature, pale yellow. truncate. B

C Style distinct (0.26) 1-1.5 mm. Style absent. D

A. Leaves very rigid; ovules 5-7; stamens not produced. 92. B. asiatiCa Leaves flexible ; ovules (2-) 4 4 ; stamens mostly produced, apicdate.

Mature stems dark red; style 1-1.5 mm. Leaves lustrous green, epruinose, below; inflorescence umbellate-paniculate.

Leaves grey, pruinose below ; inflorescence racemose or racemose-subpanicdate. Leaves 2.5-5 cm. ; inflorescence 5-7 cm. ; pedicels 12-20 mm. ; ovules sessile;

Leaves 1-2 cm. ; inflorescence 2-4 cm. ; pedicels 3-7 mm. ; ovules stipitate ;

113. B. chitria

fruit 14 x 6 mm. 113c. B. x parsomii

fruit 10 x 6 mm. 112. B. micrantha Mature stems pale yellow.

Style distinct ( 0 . 5 ) 1-16 mm. Leaves always (or ultimately) green, epruinose, below; ovules 4-45. Stems very sulcate; leaves 15-27xS12mm.; pedicels PlOmm.; fruit

Stems subangled; leaves 20-60 x 5-12 mm. ; pedicels 10-20 mm.; fruit 15 x 5 mm., dark red, epruinose. 114. B. sikkimnsis

9 x 7 mm., black, pruinose blue. 97. B.prke&nu Leaves below always pruinose, grey or white.

Ovules 4-5. Stems terete or subangled; leaves 4-7 cm. 95. B. lycium Stems sulcate; leaves 1-2 cm. 96. B. khasianu

Stems terete or subterete; leaves 15-25 mm. 111. B. ufshanica Stems sulcate, or angled; leaves 2-4 cm.

Ovules 2-3.

Leaves mostly entire; petals shorter than inner sepals; fruit 9 mm.; style 1 mm. 109. B. wightiana:

Leaves mostly spinose; petals as long as inner sepals; fruit 12 mm. ; style 0.5 mm.

Inflorescence 5-10-fld., subumbellate. 107. B. ceylanica Inflorescence 10-20-fld., racemose-paniculate. 106. B.huegeliana:

127. B.duthieanu 128. B. thomsoniana

Style absent. Stems subterete ; peduncle 2-3 cm. ; pedicels 5-9 mm. Stems sulcate; peduncle 10-15 mm.; pedicels 10-15 mm.

6-2

Page 84: Berberis and Mahonia : A taxonomic revision

84 L. W. A. AHRENDT KEY 19 (continued)

B. Style distinct (0.25) 1-1.5 m. Leaves grey, pruinose below; fruit black, pruinoee; style 0-25 mm.

Leaves narrow, 1 : 3.5, oblanmlate; mar& with 3-6 spinules; inflorescence

Laves broad, 1 : 2.6, ovate-elliptic; margins with 6-10 spinules; inflorescence 6-12-fld., panicdate. 101. B. kerriana

16-20-fld., racemose. 95e. B. x kern& Leaves green, epruinose, below; style 0-5-1.5 mm. Stems sulcate; fruit pruinose blue.

Leaves thick, rigid; infloreacence rigidly racemose; bracts 1-1-5mm.; fruit

Leaves thin, flexible ; inflorescence corymbose-paniculate ; bracts 3-3,5 mm. ; black. 94. B. baluehktanica

120. B. hoktii fruit dark red. Stems terete; fruit red, epruinose.

Leaves thin, flexible, reticulate ; fruit dark red. Inflorescence corymbose-panidate; petals shorter than inner sepah.

Inflorescence umbellate-racemose; petals as long as inner sepals. 121. B. grantii

105. B. petwkwk Leaves thick, rigid, enervate, or slightly veined; fruit pale red.

Sect. TINCTORIAE, Subsect. Potminkme Style absent. Inflorescence 2-10 (-15)-fld., 2-4 cm. long, fascicled, umbellate, or umbellate-

racemose. Leaves green below, crinkled; pedicels 15-30 mm. Leaves grey pruinose below, plane; pedicels 8-16 mm.

Leaves reticulate ; petals longer than inner sepals. Leaves slightly veined; petale shorter than inner sepals.

126. B . d t e r a n a

Stems sdcate; petiole 2-8 mm. 124. B. umbellata

124c. B. campbellii 125. B. brandiaiana Stems more or less terete; petiole 8-20 mm.

Inflorescence (lo-) 12-26-fld., formally racemose.

pedicels 1-3 ~ m .

pedimls 5-10 IIM.

Leaves narrow, 1 : 6 ; outer sepals 6 mm. ; inner sepals 8.6 mm. ; petals 4.6 mm. ;

Leaves broad, 1 : 2.5; outer sepals 2-6 mm. ; inner sepals 6 mm. ; petals 5.6 mm. ; 131. B. kongboeneis

132. B. lindkyama

C. Inflorescenoe 2-6 (-10)-fld., fascicled, umbellate or pseudumbellate, 1.5-4 cm. Stems terete.

Leaves reticulate, below pruinm grey. Leaves not reticulate, below epruinoee green.

Infloreamnce (lo-) 12-26-fld., racemoee or paniculate.

122. B.petithna 93. B. heteracantha

114c. B. t3ikkimwia var. glubramea

118a. B. x macracantha

Stems very sulcate.

Leaves with 5-15 marginal spinules. Leaves entire, rarely 1-4 marginal spinules.

stems very sulcate. Petals longer than inner sepals; stamens not produced, truncate.

Petals shorter than inner sepals; stamens produced, apiculate. 108. B. nilghirknsis

110. B. t inhria 115. B. cerabphylla

Leaves below grey pruinose. 113e. B. x epaethii Leaves below green epruinose.

Leaves below, pruinose white; ovules 2. Leaves below epruinose, green; o d e s 4-5.

stems terete.

Leaves narrow, 1 : 4.5. Pedicels 3-7 mm.; spines 4-6 mm.; fruit red. Pedicels 7-13 mm.; spines 1-2 cm.; fruit b h k , pruinose white.

116. B. aflnie

98. B. lycwidee

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Berberis and Mahonia 85

Leaves broad, 1 : 2-2-5 (-3). Flower diameter 10-12mm.; fruit 7-9mm., dark red, or black and

pruinose; spines weak 4-10 mm.; petals equal to, or longer than inner sepals.

Leaves entire; petals entire, equal to inner sepals in length; stamens not produced, truncate; style 0.5 mm. ; fruit red, pruinose blue.

117. B. jloribumda Leaves dentate ; petals emarginate, longer than inner sepals ; stamens

produced, apiculate; style 1 mm. ; fruit black, pruinose white. 99. B. glaucocarpn

Flower diameter 12-17 mm.; fruit 11-13 mm., bright red; spines 1.5- 3 cm., stronger ; petale shorter than inner sepals ; stamens apiculate.

Flowers 12-14mm. diameter; petals entire; fruit 11 mm. long, pruinose blue. 118. B. aristatn

Flowers 15-17 mm. diameter; petals emarginate; fruit 13 mm. long, epruinose. 119. B. c&ricr

D. Spines absent; flower diameter 4-6 mm. 123. B. africanci Spines (6 mm.-) 1-3.5 cm. ; flower diameter 8-12 mm.

Leaves below green, epruinose ; petals shorter than inner sepals. Pedicels 2-5-4 nun. ; stamens apiculate; fruit bright red. Pedicels 1-3-5 cm. ; stamens not produced; fruit dark red.

130. B. decandolleana

Stems terete; leaves subacuminate, concolorow below, and with 15-25 marginal spinules, reticulate ; inflorescence racemose.

129. B. kaahmiraim Stems sulcate; leaves obtuse to subacute, paler below, and with 3-10 marginal

spinules, openly veined ; inflorescence umbellate to fascicled. 126. B. walterami

Stems terete; fruit conspicuously pruinose. 100. B. hainesii Stems very sulcate; fruit epruinose.

Inflorescence pseudumbellate, 3-9-fld. ; ovules 4-5. 124. B. urnbellata Inflorescence 10-20-fld., racemose ; ovules 1-2. 113c. B. x hauniensis

Certain species of the Sect. Asiaticae, and of the Sect. Tinctoriae, Subsects. Eutinc- toriae, Chitriae, which possess conspicuous styles 1-2 mm. long should be compared with the long styled South American Group Australes, especially with its sections Ilicifoliae and Laurinae (see Key 1).

Leaves grey pruinose below.

Sect. Asiaticae Schneid., Bull. Herb. Boissier, (2), 5, 456, 1905

Shrubs &12 ft. Stems nearly always terete, pale yellow. Internodes 2 4 (-7) cm. Spines (0-,5-) 1-2 (-5) cm. Leaves evergreen, without hypoderm, thinly and flexibly coriaceous (except only in (92)); margins entire or 2-5 (-8)-spinose; openly veined or reticulate. Injlorescence (7-) 10-20 (-30)-fld., racemose, umbellate, fascicled, or rarely paniculate. Pedicels 4-9 (-15) mm., rarely 15-25 mm. Flowers 6-14 mm. diameter. Ovules (3-) 4-6 (-7). Fruit blue-black, 8-10 (-12) x 4-7 (-11) mm., mostly pruinose and stylose.

Bengal, Assam, Bhutan, Sikkim, Nepal, Kumaon, Garhwal, Jaunsar, Punjab-Hima- laya, Kashmir, Hazara, Wazaristan, Afghanistan, Baluchistan, Central India.

KEY 20, to Sect. Asiactiae (Maps 3 and 12) Inflorescence paniculate. 101. B. kerriana Inflorescence not paniculate.

Leaves thickly, rigidly coriaceous, with conspicuous venation, reticulate, or a t least openly veined.

Stems yellow, terete; infiorescence umbellate subfaacicled; pedicels 15-25 mm. Leaves pruinose white below. 92. B. aaiatb

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86 L. W. A. AHRENDT KEY 20 (continued)

h v e s epruinose green below. 93. B.h&raccsntha

94. B. balocchietartica Stems red, sulcate; idoscence rigidly racemose; pedicek 4-6 mm.

Leaves M y subcoriaceous; venation less conspicuous. Moreaceme 10-30-fld.; style distinct (0-25-) 04-1-25 mm. Stems pubescent, at any rate at h t .

Leaves above dull grey-green, below pruinose grey-white.

Stems mostly terete; leaves P 7 cm., mostly spinose. 95. B. lyciurn Stems sulcate; leaves 1-2 cm., entire. 96. B. khaeianu

Stems dark purple. 956. B . x h &

pruinose, deep blue. 97. B.pkeri.ana

95. B. ly&um varieties

Stems pale, yellow-grey.

Leaves bright green, slightly lustrous above, subconcolorous below; fruit

Stems always quite glabrous. Stems sulcate or leaves below prujnose grey, papillose. Stems terete, and leaves below epruinose, subconcolorous green.

Leaves narrow, 1 : 4, obovate, entire; racemose loose, 6-9 cm.; pedicels

Leaves broader, 1 : 3, elliptic, spinose-dentate; racemes rigid, 2-5 cm.; 7-9 mm.; petals entire and shorter than inner sepals.

pedicek 5-7 mm.; petals emarginate and longer than inner sepals.

98. B. lycioidee

99. B. gbwmrpa

Stems terete; fruit oblong, pruinose. 100. B. hiMi Stems very sulcate; fruit globose, epruinose. Cf. B. roxburghianu

Inflorescence M-fld., fascicled to subumbellate; style absent.

(92) Berberk asiatica Roxb. ex DC., Syst. 2, 13,1821, non G S t h Stem terete; shoots puberulous, mature glabrous. Spines 3-fld., 1-3 cm. h u e s thick,

rigid, 1-8-7-5 x 1 - 6 3 cm., obovate, oblong-obovate, or elliptic; acute, mucronate; petiole 3-8 mm. ; very slightly lustrous dark green with pder reticulation; below white, pruinose, papillose, with leas conspicuous reticulation. InJlorescence 15-25-fld. Pedicels glabrous, slender, red, 15-25 mm.; bracts 2-5 x 2.5 mm. Prqhylb 2-25 x 2 mm. Outer s e p b 5 x 4 mm. Inner aepd-3 8 x 5 mm. Petah 7 x 5-5 mm., obovate, ernaxginate, cunea.te; glands obovate, separate 1.2 x 0.8 mm. Stamens 5.25 mm., not produced, truncate. O& 5-7. B e d black, pruinose white, oblong-ovoid, 8 x 7 mm. Style 0 . 6 1 mm.

KEY 20A Leaves typically with margins (6) 2-5-8pinose (1-2 mm.)-dentate (1-2 mm.), distant

Leaves typically entire; flowers fascicled. 5-10 nun. ; inflorescence umbellate-racemose. 92a. B. asiatiCa; var. d i m

92 b. B. as&im var. &rkeana

(9%) Berberis asiatica vm. asiatica Nepal: 1819, Wallich 1477, part (Type I(); Sept. 1935, Bailey I (BM). W. Nepal: Balle, 5200 ft., 27 Mar. 1952, Polunin, Sykes & Williams 622 (BM); same

loc., 5800 ft., shrub to 8 ft., 27 Mar. 1952, Polunin, Sykes & Williams 625 (BM); Khater Dam, 7000 ft., shrub 5 ft., 24 Apr. 1952, Polunin, Sykes & Williams 825 (BM); Karkhan Khater Dam, 7000 ft., shrub 4-8 ft., with other shrubs, 24 Apr. 1952, Pol-, Sykes & Williams 836 (BM) ; Raintakura, N.W. of B i j a ~ , 4OOO ft., shrub to 5 ft., 26 Mar. 1952, Polunin, Sykes BE Williams 3673 (BM). These are described variowly as growing on dry grassy banks (622), on shaded rocky slopes (625), by stream sides in heavy forest (825), on north facing slopes (836), and on open hillside (3673).

S. Bhutan: Kinga Rapden. Mangde Chu, common on open hiUside, 5500 ft., 2 Apr. 1949, Ludlow, S h e d t Hicks 18605 (BM); 1838, G S t h 123 (K).

Page 87: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 87

Garhwal: Dehra Dun district, Paled, 4000 ft., Feb. 1892, Gamble 23440 (K); Malkot, 3000 ft., Feb. 1894, Gamble 24537 (K).

E. Kumaon: Feb. 1885, Hearle, s.n. (0); Naini Tal, Apr. 1884, Thomson, s.n. (K, sheet 1127).

Cultivated: fl. 5 May 1939 (Par, Cornwall). According to Lindley, it ‘was grown in quantity a t Killerton by Sir Thomas Dyke-Acland who grew 100 plants from seed received about 1832, and thousands for subsequent use in hedges, where they are free from mildew and can be kept clipped. It is hardy enough to defy any frost south of the Humber’. Personally, I find that this, and other tender species, will not withstand the harsh climate of Oxfordshire.

(92b) Berberis asiatica var. clarkeana Schneid., BUU. Herb. Ikss ier , (Z), 5,457, 1905; syn. B. hypoleuca Lindl., J . R. hort. Soc. 2,246,1847, and 5,9,1850

Bengal: Clarke 3417B (Type, K); Parasnath, 3300 ft., Hook., 8.n. (K). Bhutan: Hinglai-la to Nahi, 11,300-650Oft., 28 May 1938, Gould 275 (K); 1838,

Griffith 747 (K); Chunzu Gompa, 6 ft., open hillside, 7000 ft., 21 Apr. 1949, Ludlow, Sherriff & Hicks 16073 (BM).

Assam: Rufa, 5000-6000 ft., 9 July 1938, Kingdon-Ward 13883 (BM). A large bushy shrub, in thickets along the streams in limestone country.

Cultivated: according to Lindley, seed waa collected from Nepal before 1847, pro- ducing a hardy shrub injured only by severe winters.

(93) Berberis heteracantha Ahrendt, sp.nov., aff. B. asiaticae, sed spinis, brevior-

Sterns stout. Internodes 1.7-3.2 cm. Spines 3-fld., stipitate, the lateral parts, which are 6-10mm. long and 1-1.5mm. broad, diverging 2-3mm. from the stem. leave^ 24- 40 x 6-8 (-12) mm. ; narrowly obovate, cuneate, sessile; margins slightly revolute, 1-3- spinose (1-1-5 mm.)-dentate (2-3 mm.), distant 3-10 mm.; veins and veinlets branched, elevated, not reticulate ; below paler green, epruinose. InfZorescence 6-10-fld., umbellate or umbellate-racemose. Pedicels slender, 8-12 111111. Flowers 5-6 mm. diameter.

ibus et foliis subtus viridibus distinguenda

Afghanistan: fl., immature, G d i t h 1424 (Type, K). Not cultivated.

(94) Berberis baluchistanica Ahrendt, J . Asiat. SOC. Beng. (Sci.), (3) 11, 1, 1945 Internodes 1-2 (-3) cm. Sterns red, sulcate. Spines 3-fld., stout, 1-2 cm., plane.

Leaves thick, rigid ; margins mostly entire, occasionally 34-spinose (0.5 mm.)-subserru- late, distant 3 mm. ; closely elevated-reticulate. Inflorescence 10-20 (-25)-fld., rigidly racemose, 3-5 cm., including peduncle, 4-10 mm. Pedicels fairly slender, 4 4 mm.; bracts 1-14 mm. Berries black, pruinose blue, ovoid, 6 x 3 mm., excluding style c. 0.5 mm.

Baluchistan: Larghun, immature fruit, Aug. 1885, Gamble 2 (Type, K); fl. 1851, Stocks 931 (K). Not cultivated.

(95) Berberis lycium Royle, Trans. Linn. Soc. Lond. 17,94, 1834; J. D. Hooker, Bot.

Shrub suberect, 9-12 ft. Internodes 16-35 cm. Spines 6-20 mm., subterete. Leaves oblanceolate, acute to subacuminate ; attenuate subsessile ; margins mostly entire, rarely with 1-2 spinules; openly veined; above dull grey-green. Pedicels slender, glabrous (6-) 8-12 (-15) mm., occasionally to 2 cm. near base; bracts 2-5 mm. Flowers 7-8 mm. diameter. Prophylls 1 x 0-6 mm., red, ovate, acute. Outer sepah 2 x 1 mm., yenow, ovate, obtuse. Median sepals 3.5 x 2 mm., ovate, obtuse. Inner sepals 5 x 3 mm., obovate. Petals 4.25 x 2-25 mm., obovate, emarginate with acute lobes; glandslanceolate, separate,

Mag. 115, t. 7075 1889; Parker, Forest Fl. Punjab with Hazara and Delhi, 14,1918

Page 88: Berberis and Mahonia : A taxonomic revision

88 L. W. A. AERENDT submarginal, 1.2 x 0.3 nun. Skronerrs 3-25 mm., produced, apiculate. Ovules 4, shortly stipitate. Berries 9-10 x 4 .56 mm., black heavily pruinose grey-white, including style, 1 mm.

KEY 20B Stems terete; leaves 3.5-6 x 0.7-1 (-1.2) em., whitish and conspicuously papillose

Stems glabrescent (i.e. with the pubescence reducing, so that the mature stems are below; infloreacence 10-25-fld.

puberulous or glabrous). Inflorescence racemose, 5-9 cm. Inflorescence subfascicled, 2-26 cm.

95a. B. lycium var. lycium 95 b. B. lycium var. subfascioularie

pubescent); inflorescence racemose, 2-5 cm. 95c. B. lycium var. Sintkn~is

lose, to 4 x 1 cm.; inflorescence 6-16 fld., with pedicels 5-10 mm.

Stems pubescent (i.e. with the pubescence persisting, so that the mature stems are also

Stems slightly sulcate; leaves becoming dull greenish below, and not or scarcely papil-

95d. B. lycium var. subvireamne

(954 B. lycium var. lycium Kashmir: Royle, 8.n. (Type, K); Achaba, 5000ft., fl., 13 May 1940, Ludlow &

SherriE 8070 (BM), common on open hillsides; Chmgha, 2250 ft., fr. 1 July 1876, Clarke 28170 (K); according to Parker, common in Kaahmir, 3000-7000 ft.

Garhwal: Nmda,kin 3500 ft., Strachey & Winterbottom 7 (K); Dehra Dun district, below Muaciooree, 6000 ft., Apr. 1892, Gamble 23759 (K).

Jaunsar, Mandhali, 5OOO ft., June 1892, Gamble 23502 (K). According to Royle from Ragpore to Mussooree, 3000-7000 ft., and from Mehri to Choor.

Cultivated: fl. 26 June 1939 (K).

(95b) Berberis lycium var. subfascicularis Ahrendt, var.nov., hflorescentiis

Kashmir: Kishtwar district, Tsingam, 6500ft., fl. 20 June 1943, Ludlow t Sherriff

Cultivated: fl., fr., Wisley, 1939. Often distributed merely as B. lyciurn.

brevioribus et racemoso-subfasciculatis

9105 (Type, BM), in foreat clearings.

(95c) Berberis lycium var. aimlensis Ahrendt, J. Asiat. SOC. Beng. (Scd.), (3) 11, 1, 1945

Punjab Himalaya: Simla, 7000 ft., July 1885, COlIett 234 (Type, K) ; Collett 28 (K); Khasauli, near Dhanapur, 4000 ft., fl. 14 Apr. 1877, Gamble 4126A (K); Simla, Pagog temple, 5700 ft. 28 Apr. 1887, Gamble 41598 (K); Simla, Jaru valley, 5000 ft., 4 July 1877, Gamble 4501A (K).

Garhwal: Parewa, Kotah range, Inayat, fl. 15 June 1902, Duthie 25853a (K). Jaunsar: Kathyou, Tonso basin, 7000 ft., May 1891, Gamble 23070 (K). Cultivated: fl. June 1939 (Wisley).

(95 d) Berberis lycium var. subvirescens Ahrendt, var.nov., ram& leviter suloatis, foliis ultimo subtus viridibus, vix vel non papillosis

Stems slightly sulcate, glabrous. Leaves below pruinose at h t , but becoming dull

Kashmir: Gandarbal, Sind valley, 5200 ft., fl. 31 Ma,y 1940, Ludlow & S h e d 8095

Cultivated: often merely as B. lycium.

yellowish green below.

(Type, BM), on banks of streams.

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Berberis and Mahonia 89

(95.e) Berberis x kewensis Schneid., J . Am. Arb. 4, 197, 1923 Stems glabrous, sulcate, purple-brown. Internodes 1.5-3 cm. Spines yellow, stout,

6-25 mm. Leaves 25-50 x 9-20 mm., ovate-elliptic, obtuse to subacute; petiole 2-4 mm. ; margins 5-10-spinose (1-1-5 mm.)-serrate (but on the new shoots longer, 5.5 x 1.5 cm., or broader 5 x 3 cm., and subentire) ; openly reticulate; below grey pruinose. InJlores- cence 15-20-fld., 4 4 cm., racemose, subsessile. Pedicds 5-10 mm., stout; bracts 2- 2.5 mm. Prophylls 2 mm. Outer sepals 3 mm. Inner sepals 6 mm. Petals 6.5 mm., sub- entire. Stamens 5 mm. Ovules 2-3, subsessile. Berries 10-12 x 5-6 mm., black, pruinose. Style short.

Cultivated: origin in U.S.A., from seed derived from Kew and reputed to be that of B. lycium. Not cultivated in England.

(96) Berberis khasiana Ahrendt, J . Bot., Lond. 80 (Suppl.), 100, 1943 8tem sulcate, mature, yellow, pubescent. Internodes 1.5-2.5 cm. Spines 1-1.5 cm.,

plane. Leaves 15-20 x 5-7 mm., obovate, sessile, acute, mucronate, entire; above dull, with lateral veins and few veinlets elevated. InJlwescenw 2-3 cm., partly umbellate, partly fascicled, occasionally partly loosely racemose, (7-) 12-20-fld. Pedieels 8-13 mm. Berries ellipsoid to oblong-ovoid, 5-7 x 3 4 mm. ( ? immature), excluding style, 0.5 mm.

h a m : Khasia Hills, 1844, GrifEth s.n. (Types, BM, K). Not cultivated.

(97) Berberis parkeriana Schneid., Fedde Rep. Sp. Nov. 11, 162, 1912; Ahrendt, J . Bot., Lond., 80 (Suppl.) 105, 1944; Parker, A Pore& F b a for the Punjab with Hazara and Delhi, 14, 1918

Shrub 7-9 ft. Stems sulcate; shoots puberulous. Internodes 1.5-3 cm. Spines slender, 1-2 cm., sulcate. Leaves 20-60 x 5-12 mm.; narrowly obovate, acute, mucronate, entire; petiole, 1 4 mm. ; both sides openly veined lustrous concolorous green, epapillose. Racemes subfascicled, 3 4 cm., (7-) 10-14 (-18)-fld. Flowers (H.C.C. 2/1). Prophylb 0.6-1.5 x 0.2-0.7 mm. Outer sepals 2.5 x 1.5 mm., reddish, oblong-elliptic. Inner sepals 4.5-6 x 2 5 4 . 5 mm., obovate-elliptic. Petals 4.5-5 x 2-3 mm., narrowly oblong-elliptic, subentire; cuneate, with separate glands, 0.8 x 0.2 mm. Stamens 4 mm., produced, shortly apiculate. Ovules 4. Berries 8-9 x 6-7 mm., excluding style, 1 mm., pruinose bright blue.

Hazara: above 3000 ft., Hooker 2371 H (K); Deru Lal, Khan, 8500 ft., 9 June 1896, Parker 4435 (Type); Lahore, Bagnolar Reserve, 4100 ft., July 1910, Parker s.n. (K).

Wazaristan: 6000-7000 ft., according to Hooker on the herbarium sheet previously cited, though material wa9 not preserved from here.

Kashmir: Chenari, Jhelum valley, 3000-3500 ft., 8 May 1940, Ludlow & Sherriff 8063

According to Parker, this species replaces B. lycium in Hazara and Rawalpindi, the leaves persisting over the winter.

Chltivated: fl., 12 May 1939 (K); fr. NOV. 1939 (K); fl. 7 Apr. 1943. This species is often defoliated in an English winter. The Kew plant, some 7 ft. high, was cut to the ground, but not kitled in Jan. 1940.

(98) Berberis lycioides Stapf, Bot. Mag. 151, t. 9102, 1926; Ahrendt, J . Bot., Lond., 80 (Suppl.), 102, 1943, non Linden 8z Planch., Trois Voy. Lind, Bot. PI. Columb. 1, 9, 1863. (Fig. 25)

Shrub 9-12 ft. Stem glabrous, terete, pale yellow. Internodes (2-) 3-5 (-7) cm. Spines concolorous, 1-2 cm. Leaves 3-7 x 0-8-1.8 cm.; narrowly obovate, subacute, mucronate; attenuate, sessile, entire ; venation clearly branched, finely reticulate ; above sub- lustrous, deep green, below slightly paler, lustrous, epruinose. In$orescem loosely

(BM).

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90 L. W. A. ~ N D T

racemose, sometimes compound below, 10-20 (-25)-fld., 6-9 (-12) cm. long, including peduncle, 1-2 (3 .5) cm. Pedicels 7-10 (-13) mm.; bracts, 2-3 mm. F h r s 12-14 mm. diameter. Ptvphylb 1-25 x 1 mm. Oecter 8epa.l.s 2-75 x 1.75 mm., ovate, acute. Hedicln se@ 6 x 2-5 nun., fjhdax. Imer s e p h 7-8 x 4 - 5 4 mm. Pef& 6.5 x 4 mm., obovate- elliptic, obtuse, truncate, entire; base broadly clawed, with subacumjnate, separate marginal glands, 2 x 0.5 mm. Stamew 5 mm., produced, apiculate. Cbule.8 3-4, shortly stipitate. Be& black, pruinose grey-white, oblong-ovoid, 11-12 x 6-7 mm., excluding style 1-1-26 mm.

Jaunsar: seed, Gamble, s.n.; Chitchfield 9 (0). Cultivated: fl. June 1939; fr. 27 Sept. 1939 (Type, BM), from Gamble’s original plant

a t Liss, Hampshire, by courtmy of Mr G. D. H. Coates. The flowers of this species are remarkably fragrant, and one large plant will fill the air with a heavy scent for a long distance around.

(99) Berberis glaucocarpa Stapf, Bot. Mag. 101, sub. t. 9102 in obs., 1926; Ahrendt, J. Bot., M., 88 (Suppl.), 101, 1943, descr. ampl.; syn. B. Coriacea Brandis in herb. non St Hil.; ‘B. aristda’ Parker, Forest Fl. Punjab with Hamra and Delhi, 15, 1918, non DC.; ‘B. miar ia ’ Collett, FZ. X i d . 21, 1902, non Royle. (Fig. 26)

ahrub 9-12 ft. Stems glabrous, pale yellow, terete, very minutely verruculose. Ider- nodes 2-5-4-5 cm. S p i m mostly solitary, 6-9 mm. Leaves 3-6 x 1-2.3 cm., elliptic, acute, mucronate; petiole, 2-6mm.; margins with 4-8 spreading spinose (1-2mm.), teeth (1-2 mm.), distant (3-) 6-10 mm.; both sides sublustrous concolornus green; scarcely reticulate but with veins and veinlets visibly branched and elevated below. Racemes rigid, 2-4 cm.; 15-25-fld., occasionally slightly compound below. Flowers 10-12 mm. diameter. Prophylls 1 x 0.5 mm. Outer sepals 2 x 1-6 mm., ovate, subacute. Median sepals 4.5 x 3 mm. Inner sepa.l.s 6-66 x 5 x 5 mm., obovate. Petale 7 x 4.5 nun., obovate; apex retuse with rounded lobes; base clawed with separate marginal glands, 1-8 x 0.6 mm. Stanzens 5.5 nun., produced, obtusely subapiculate. Cbule.8 4, shortly stipitate. Berries oblong-globose, black, pruinose white, 8-9 x 6-5-7 mm., excluding style 1 mm.

Jaunsar: hills between Tons and Gisi rivers, 8000ft., immature fruit, May 1875, Brandis 746 (Type, K); Dussatin, Sangri, 8000ft., fl. May 1881, Brandis, 8.n. (K); Mandhali, fl. Apr. 1898,7000-8000 ft., Duthie, 8.n. and fr. 4 June 1898, Duthie 2190 (K).

Punjab Himalaya: S a a , near Narkanda, fk., 9OOO ft., 30 Sept. 1876, Gamble 1484B (BM, K) ; Chamba, above Ulwass, Ravee basin, 8OOO ft., 7 June 1878, Watt 957 (K) : ‘flowers with a very strong smell’.

Tehri: Lambatach, 8000 ft., fl. May 1891, Gamble 22722 (K). Kashmir: Kishtawar district, fl. 8000-10,000 ft., immature fruit, Aug. 1943, Ludlow &

SherriB 9289 (BM), 7-8 ft., in open conifer forest. According to Collett, this species occurs from Kumaon to Simla, 7000-10,000ft.

Accodmg to Parker, it occurs from Chota Bengal to Nepal, in open shrubby places, but is not common west of Sutlej. However, I have seen no herbarium specimens from east of Tehri, and the references to Kumaon and Nepal may introduce confusion with B. coriaria and B. ar48tatac respectively.

Cultivated: fl. June 1939; fi-. 6 Nov. 1939 (N. G. Hadden, Porlock, Somerset); fr. 21 Sept. 1942 (W. J. Marchant, Wimborne, Dorset).

(100) Berberis hainesii Ahrendt, J. Bot., Land., 80 (Suppl.), 105, 1944 Stems terete, mature pale yellow. I n t e d 1-54 cm. Spines 6-12 mm. Leaves thin,

4-6 x 2-3 cm. ; broadly oblong-elliptic, excluding petiole, 5-10 (-15) mm. ; margins en- tire, rarely with 1 4 spinules (but on new shoots often with truncate base, rounded apex, and margins coarsely toothed with 3-5 spinules, 1.5mm. long, surmounting teeth, 5-7 mm. deep, distant 7-12 mm.); veins and veinlets branched, not reticulate; above

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Berberis and Mahonia 91

dull grey green; below pruinose grey, papillose. Inflorescence umbellate-racemose to subfaslcicled, 3-6-fld. F’lowers 8-10 mm. diameter. Petah 6 mm., narrowly obovate, obtuse, entire. Berries oblong-ellipsoid, heavily pruinose.

KEY 20c Pedicels (1-) 1.5-2 cm; fruit estylose Pedicele 6-9 mm; fruit shortly stylose

100a. B. huineaii var. huinesii 100 b. huineaii var. brewifilipes

(100 a) Berberis hainesii var. hainesii India: Central Provinces, Pachinaki, June 1911, Haines 2905 (Type, K), 2906 (K)

23 June 1911, Hainea 2 (K): ‘Frequent on the sandstones above ZOO0 ft.; in previous years, fruit seen on one specimen. Erect 5-7 ft.’ Not cultivated.

( l00b) Berberis hainesii var. b r e w p e s Ahrendt, varaov., pedicellis, brevioribus

India: Central Provinces, near Bes falls, 18 Feb. 1891, Duthie 13202 (Type, K); Hole 678,856 (K). Not cultivated.

This remarkable species is interesting in its unusual habitat, and its character; it has some aflinity with Sect. Tinctoriae, Subsect. Umbellatae, but even more with Sect. Heteropodae.

gracilioribusque

(101) Berberis kerriana Ahrendt, sp.nov., foliis ut in Sect. WATLICIILBNAE, sed

Stems glabrous; shoots reddish; mature deep red, becoming red-brown, and quickly yellow-brown. Interrwdes 1-2 cm. Spines discolorous, pale yellow, slender, 2-3-5 cm. Leavea slightly s t a y coriaceous, 25-45 x 7-13 mm.; oblanceolate attenuate; acute, entire; margins towards apex 3-5-spinose (1-2 mm.)-serrate (1-2 mm.), distant, 5-8 mm. ; venation branched, scarcely subreticulate, on the slightly lustrous upper surface ; below pruinose white, with venation more distinct. Injbrescence panicdate, 6-12-fld., 3-5 cm. Pedicels (7-) 10-14 mm.; bracts, 1 mm. O& 2-3. Berries black, pruinose grey, oblong, 9-10 x 5-6 mm., excluding a very short, but distinct, style.

inflorescentiis paniculatis

China: seed, 1935, Kerr 39. Cultivated: from abqve collection, fr. 9 Oct. 1943; fl. May 1944 (Type, BM). This is a remarkable plant, cultivated at Wisley. Its leaves are quite typical of those

of many species of the Sect. Wallichianae, but the inflorescence is as far apart from what is constant in that section as could be. It does not really fit into any section happily, and I should have been inclined to regard it as a hybrid, if it were not that that I had the above information as to its origin supplied to me by Mr B. 0. Mulligan, from the records at Wisley where it grew. B. kerriana should also be compared with Sect. Tinctoriae, Subsect. Potaninianae.

Sect. Tinctoriae Schneid., Bull. Herb. Boissier, (Z), 5, 450, 1905

Leaves evergreen (but in cultivation often sub-evergreen or defoliated in winter). Injlorescence racemose, umbellate, pseudumbellate or fascicled or occasionally corym- bose-paniculate. Ovules 1-5. Stamens dentate. Fruit pale, bright or dark red, often epruinose, sometimes with bloom.

E. Africa, Arabia, Ceylon, South India, Afghanistan, Ka~hmir, Punjab, Garhwal, Kumaon, Nepal, Bhutan, Sikkim, Tibet (Kongbo). One variety extends to W. Yunnan. The exceptional Subsect. Potaninianae belongs to N.W. China (Kansu, Shensi, N.W. Szechuan).

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92 L. W. A. AHRENDT

Iclcy 21, to Subsects. of the Sect. Tinctorha (Maps 2 and 13)

Leaves stiff, rigid, mostly with hypodem; style distinct, 0.6-0-76 111l11.

Leaves flexible, thin, without hypodem. Subsect. POTANINIANAE

Subsect. EWTINCTORIAE

subsect. CHZTBIdE

Subsect. U ~ E L I ~ T A E

Ovules 1-2 (-3); style conspicuous ( 0 . 6 ) 1-1-6 mm. Ovules (3-) 4-6.

Style absent. Style conspicuous ( 0 . 6 ) 1-1-5 mm.

Inflorescence 2-1O-fld., subumbellate, or subfescicled. htlomcence 10-25-fld., racemom. Subsect. LINDLEYANAE

Subsect. Potaninianae Ahrendt, J. Bot., LoluE., 79 (Suppl.), 79, 1942 Stem ghbrous, mature, dark red, terete or subangled. Internodes 1.5-3 cm. Spines

subconcolorow, 3-0d. Leaves evergreen, thick, rigid; below h d l y paler yellow-green, subenervate. Fruit pale red, epruinose. Style 0.5-0-75 111111.

Kansu, Shensi, N.W. Szechuan.

Iclcy 22, to Subsect. Potaninianae (Map 13)

Ovules 1-2; infloscence racemose or umbellate; petals shorter than inner sepals; sttunens not produced.

Leaves narrow, 1 : 5-6; margins each with 1-2 sphulea; fruit subglobose.

h v e a broader, 1 : 2-5-4; margins each with 3-6 spinules; fruit ovoid. 102. B. potaninii

103. B. liechtensteinii

104. B. anniae Ovules 3-4; infloreacenca fascicled; petals longer than inner sepels; stamens produced.

(102) Berhris potanid Maxim., A& Elorti. Petrop. 11, 41, 1891; s p . B. sphalera

Leavea 10-30 x 2-5-5 mm., narrowly obovate, acute, mucronate, attenuate, m i l e ; margins l-2-spinose (1-2 mm.)-serrate (1-2 mm.) ; paler below. Inflorescence racemose. Pedids c. 5 mm. Bewies oblong-globose.

Kansu: Heishu, fl., and immature fr. 2 June 1885, Potanin, s.n. (Type); Tshung-sha- wan, ad rivulam, immature fr., fi. 11 Sept. 1885, Potanin 8.n.; seed only Farrer 358; near Chu Kun, 18 Oct. 1914, Meyer 1984.

Shemi: Mts., by Lun san hao, immature fi., May 1895, Giraldi 62 (Type of B. s p w r a ) ; Kisan, Giraldi 61; near Shan lan hi, 17 Sept. 1914, Meyer 1926. Described by Meyer as a shrub 3-5 ft., with bright red fruit.

Fedde, Bot. Jb., 36, Beibl. 82, 44, 1905

Cultivated: from seed of Farrer 358.

(103) Berberis liechtensteinii Schneid., Sargent, PI. Fib. 1, 377, 1913 Shrub 3-4 ft. Stem not verruculose. Spines slightly sulcate (2-)3-6 cm.) Leaves

26-35 (4) x 10-15 (-18) mm.; obovate, acute, mucronate, cuneate, sessile; margins by apex (2-) 3 4 (-6)-spinose (0.5-1.5 mm.)-dentate (2 mm.); above dull dark grey-green; below at Grst pruinose, Snally paler yellow-green. InJlorescence 6-12-fld., umbellate- racemose, 2 4 cm. Pedieels 5-10 mm. Flowers H.C.C. 211. Prophy& concolorous, 2-5-3 x 1-1.5 mm., lanceolate. Outer sepals 6 5 x 3-4 mm., elliptic to obovate, obtuse. Inner se& 5-6 (-7) x 4.25-5 mm., obovate. Petah 4-25-5 x 3-3-5 mm., obovate, entire; base truncate, with lanceolate separate marginal glands, 0.5-0-7 x 0.2-0.3 mm., situated 0-5 mm. above the base of the petal. Stamens 4-5 mm. Ovules sessile. Berries ovoid, 7-8 x 6 mm.

Page 93: Berberis and Mahonia : A taxonomic revision

Berberis and Mtthonia, 93

N.W. Szechuan: Min river valley, near Mao Chu, arid region, 26 May 1908, Wilson

Cultivated (Wilson 4154): fl. 12 May 1942, 21 May 1944; fr. 9 Dec. 1942. A species 2871 (Type, Am. Arb.); Ma0 chu, 6500-9000 ft., Wilson 4154 (K).

tender in Oxfordshire, where the fruit ripens slowly and late.

(104) Berberis anniae Ahrendt, J. Bot., Lond., 79 (Suppl.), 79, 1942 Stem verruculose. Spines stout, 1-26 cm., plane. h u e s slightly less rigid than in the

preceding species, oblong-elliptic, rarely obovate-elliptic; 2 0 4 5 (-50) x 8-14 (-20) mm. ; acute, mucronate, cuneate, sessile ; margins (3-) 5-8 (-10)-spinose (1-1.5 mm.)-serrate (0.5-1 mm.), distant (3-) 4-6 (-8) mm.; above slightly lustrous light green, loosely veined with lateral veins, few veinlets, and part of the marginal vein visible ; below much paler, slightly lustrous, papillose, subenervate. Flowers fascicled 2-6. Pedicels 10-15 mm. Prophylls 1-1.5 mm. Outer sepals 4 x 3 mm., elliptic. Inner sepals 6 x 4 mm., obovate. Stamens 4.5 mm. Berries 10 x 6.5 mm., oblong, pale red.

Cultivated: origin unrecorded, but found growing at Wisley; %. 12 Apr. 1939 (Wi.); fr. 11 Nov. 1938 (Wi.); fr. 28 Oct. 1942 (Types, BM, 0). A tender species which, however, grew to over 6 ft. at Wisley until badly damaged in January 1940. It has been named after my mother, Annie Rachel Ahrendt. When C. K. Schneider first referred to B. liechten- steinii, he remarked that it w&s difficult to place in the genus. This still applies to these three species. The Sect. Tinctoriae is a predominantly southern one, lying outside China. Yet there seems no other reasonable place for these than as an extraordinary subsection of the Tinctoriae; one which lies, not only in China, but in the northern regions thereof, and one whose species are remarkably tender despite this fact. Their rigid leaves are reminiscent of the Wallichianae, but their inflorescence and fruit of the Tinctoriae.

Subsect. Eutinctoriae Schneid., Bull. Herb. Bossier, (2)) 5, 451, 1905 h u e s evergreen, coriaceous, without hypoderm, generally (and unless otherwise

noted) pruinose grey-white below. Stumem produced (unleas the opposite is stated). Ovules 1-2. Berries red, dark red, or red-purple. Style mostly 1-1-5 mm.

Ceylon, South India, Afghanistan, Nepal, Garhwd, Kumaon.

KEY 23, to Subsect. Eutinctoriae (Maps 13 and 14) Stems dark red, when mature, and terete; leaves green, epruinose below.

Stems pale yellow, sulcate, or angled; leaves below pruinose, grey-white. 105. B. petiolaris

Inflorescence racemose or pseudumbellate, 3-8 cm. long. Petals longer than, or as long as, inner sepals.

Petals entire and as long as inner sepals; stems puberulous. Inflorescence 10-20-fld., racemose-paniculate. 106. B. huegeliana Inflorescence 5-10-fld., subumbellate. 107. B. ceylanica

108. B. nilghiriensis

Stems puberulous; racemes 6-12-fld.; pedicels 8-18 mm. 109. B. wightiana Stems glabrous; racemes 12-20-fld.; pedicels 5-10 mm. 110. B. tinctoria

Inflorescence subfascicled, 1-2 cm. long. 111. B. ajgbnica

Petals emarginate, and longer than inner sepals; stems glabrous.

Petals shorter than inner sepals; stems very sulcate.

(105) Berberis petiolaris Wall. ex G. Don, Gen. Syst. 1, 116, 1831; Ahrendt, J. Bot., Lond., 80 (Suppl.), 82, 1942

Stem glabrous, terete, sublustrous, mature deep red to red-brown. Internodes 3 4 cm. Spines often absent, occasionally solitary, 3-7 mm. long. Leaves both sides concolorous green, epruinose. Pedicels glabrous, 7-15 mm. long. Ovules 2, subsessile.

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94 L. W. A. ~ R E N D T

KEY 23a Leaves entire; racemes %lS-flowered, 3-6 cm. long; style 1-1-6 ILM. long.

105a. B. petwikk var. p e t w h 6 Leaves spinose; umbels %'i'-flowered, 2-3 cm. long; style 0.6 mm. long.

105b. B. pt?ttidcM.is var. g a r h w w

(1050) Berberis petiolaris var. petiolaris Leaves obovate to elliptic, 4-8 x 1.4-3 cm., including petiole 1-2 om. long, subacute,

entire, rarely with 1-2 marginal spinules, finely, loosely reticulate both sides. RarcemRs 8-15-fld., 3-6 cm. long, including peduncle 6-10 mm., and 1-16 cm. broad. Outer sezkzh 3.5 x 2 mm., ovate. Inner sep& 6-5 x 5 mm., obovate. P& also 6-5 x 5 mm., obovate, entire; base clawed with hcmlate, acuminate, separate, non-margkd g h h , 1 . 2 ~ 0.3 mm. Sta- 5.5 mm., not produced. Style 1-1.5 mm.

Nepal: 1818, fl., Wrtllich 1475, part (Type, BM). Not cultivated.

(105b) Berberis petiolaris var. garhwalana Ahrendt, J. Bot.LOnd.,8O(Suppl),82,1942 Leaves obovate, acute, with petiole, 1-5-2.7 cm.; margins with c. 14 spinose (1.5 mm.)

teeth (1-2 mm.), distant 3-9 mm.; both sides loosely veined to subreticulate; 5-10 x 1.8-3.7 cm., including petiole. Umbek with peduncle 7-15 mm. long. Berries ellipsoid, or oblong, 7.5 x 3.5 cm., excluding style, 0.5 mm. long.

Garhwal: 8000 ft., Tunghasi, Strachey & Winterbottom 5 (Type, BM). Kummn: Blinkworth, s.n. (BM). Not cultivated. This species is often confused with B. pachyacantha;, which has deciduous, closely

spinulose leaves, many-flowered racemes, petals emarginate and shorter than the inner sepals, and estylose berries.

(106) Berberis huegeliana Schneid., BuU. Herb. BoisSier, (2), 5, 451, 1905 Stems angled; shoota puberulous; mature glabrous or subglabrous. I&& c. 2 cm.

Spines concolorous, 1-2 cm., slender. Leaves subcoriaceous, 3-4-5 x 1-2-1-7 cm., oblong- obovate, subacute; cuneate to decurrent petiole 3-7 mm.; margins occasionally entire, mostly with 1-4 spinules towards apex; reticulate; below pruinose grey, papillose. Racemes sometimes compound below, 10-18-fld., 3-6cm. long, including peduncle, c. 1 cm. Pedicda 5-10 mm.; bracts c. 2 mm. Inner sep& and entire p& both 6 mm. long. ovules stipitate. Style distinct.

Kashmir: Huegel 4030 (Type, Vienna). Not cultivated. Known only through C. I(. Schneider's description.

(107) Berberis ceylanica Schneid., BUZZ. Herb. B&sier, (2), 5, 453, 1905 Stems angled, puberulous. Spines concolorous, sulcate, 1-2 cm. h v e s thick, 15-40 x

10-25mm., ovate, elliptic or obovate; petiole, 2-4mm.; margins 1-3-spinose (0.5- 1 mm.)-subserrulate, distant 4-5 mm. ; subreticulate; below grey, pruinose. In$orescenee 5-10 (-15)-fld., umbellate-racemose, sometimes compound below, sometimes corymbose- subpaniculate, c. 3-5cm. long. Pedice.?a 7-16mm.; bracts 1-2mm. Outer sepals 3-25 x 2.5 mm., oblong-ovate. Inner sepals 6-5 mm., ovate. Petals 6 x 4-5 mm., obovate, entire. Berries ellipsoid to obovoid, 12 x 5 mm., excluding short style.

Ceylon: Pidm Talgalla, 1886, Meyer, s.n. (Type, Vienna); 1857, Hooker, s.n. (K). Not cultivated.

(108) Berberis nilghiriensis Ahrendt, J. A&. Soc. Beng. (Sci.), (3), 11, 1, 1945 Stems glabrous. I&& 1-5-3 cm. Spines 15-25 cm. h v e s 20-40 x 10-15 mm.,

oblong-obovate, entire, reticulate, below grey-pruinose. Racemes 10-15-fld., including peduncle, 3-4 cm., 5-8 cm. long. Pedicel8 5-10 mm. Prophyh 3 x 1-75 mm., red, ovate,

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Berberis and Mahonia 95

obtuse. Outer sepals 4.25 x 2.5 mm., elliptic, yellow flushed red. Inner sepals 6 x 4 mm., obovate. Petals 7 x 4 mm., obovate; acutely emarginate with rounded lobes; base clawed with linear-lanceolate glands, 2-3 x 0.5 mm. StamRns not produced.

South India: Nilghiri Hills, 7000ft., fl. 7 Mar. 1870, Clarke 10504 (Type, K). Not cultivated.

(109) Berberis wightiana Schneid., Bull. Herb. Boissier, (2), 5, 452, 1905 Stems very sulcate, puberulous. Internodes 1-5-2-5 cm. Spines concolorous, slightly

sulcate, 1-5-3 cm. Leaves thick, 2 4 x 1-1.4 cm., oblong-obovate obtuse, subsessile, entire, rarely with 1-2 marginal spinules; reticulation openly elevated; below grey, pruinose. Inflorescence 7-12-fld., racemose to subumbellate-racemose, 2 4 (-5) cm. Pedicels 8-18 mm.; bracts 1-1-5 mm. Prophylls 1.5 x 0.5 mm., acuminate. Outer sepals 3 x 1.25 mm., oblong-ovate. Inner sepals 5 x 3-75 mm., obovate. Petals 4 x 2.5 mm., obovate. Stamens 3 mm., shortly apiculate. Ovules 2 ( 3 ) , shortly stipitate. Berried narrowly obovoid, epruinose, 9 x 4 mm., excluding style, 1 mm.

South India: Nilghiri Hills, Wallich 1476 (K); Wight 50 (Type, K). Ceylon: 6000 ft., Hooker s.n. (K). Not cultivated.

(110) Berberis tinctoria Lesch., MJm. Nus., Paris, 9, 306, 1822 Stems very sulcate. Spines 3-fld., concolorous, 1.5-3 cm. Leaves thick, 1.5-3.5 (-5) x

0.7-1-3 (-2.3) cm., obovate, obtuse, mucronate; petiole 2-4 (-6) mm.; margins entire, rarely with 1-2 spinules; venation branched, not reticulate; above dull; below pruinose, papillose, white or greyish. Racemes 10-20-fld., 3 - 5 4 cm. Pdi& 5-10 mm. Prophylls 1 x 0.5 mm. Outer sepals 2.5 x 1 mm., ovate, acute. Median sepals 4 x 2.5 mm., ovate, obtuse. Inner sepals 7 x 5 mm., obovate. Petals 6 x 4.5 mm., obovate; clawed, with glands 1 x 0.4 mm. situated towards the base of the claw. Stamens 5 mm., shortly apiculate. Ovules stipitate. Berries slightly pruinose blue, 9-10 x 5 mm., excluding style 1-1.5 mm.

South India: Nilghiri Hills, Gardner, s.n. (K); Ootacamund, 7000ft., June 1884, Gamble 14280 (K).

Ceylon: Thomson, s.n. (K). Cultivated: Lindley, J . R. h t . SOC. 2, 246, 29 May 1847; 1.c. 5, 9, 1850, records this

species as being then in cultivation. It is not so now.

(111) Berberis afghanica Schneid., BUZZ. Herb. Boissier, (2), 5, 455, 1905 Stems at first puberulous, finally subglabrous; subterete. Internodes 1-2.5 cm. Spina

concolorous, subsulcate 10-18 mm. Leaves 15-25 x 5-12 mm., obovate; margins with 2-3 spinules; openly reticulate; below grey, pruinose, papillose. In.orescence 8-12 (-18)- fid., racemose-subfascicled, 1-2 cm. long. P d k l s slender, 8-12 mm., slightly puberu- lous, but glabrescent. Ovules 3, sessile. Style distinct.

Afghanistan: Griffith, 121 (Type). Not cultivated.

Subsect. Chitriae Schneid., Bull. Herb. Boissier, (2), 5, 453, 1905 Internodes (1-) 1-1.5-3.5 (-6) cm. Spines 0-5-(1-3) cm. Leaves mostly evergreen, sub-

coriaceous, without hypoderm (1 .S ) 3-7 (-12) cm. long; margins entire or with 1-9 spinules. InJEorescence 3-25-fld., racemose, umbellate, subracemose-pseudumbellate, rarely corymbose-paniculate. Stumens with apex mostly produced, obtuse or apiculate. Ovules (3-) 4-5. Fruit bright or dark red, pruinose or epruinose, 8-15 mm. long, always stylose, the style being 0.5-1.5 mm.

East Africa, Punjab Himalaya, Garhwal, Kumaon, Nepal, Sikkim, Bhutan, with one variety from W. Yunnan.

Page 96: Berberis and Mahonia : A taxonomic revision

96 L. W. A. AERENDT

KEY 24, to Subsect. Chitriae (Maps 13 and 15) Stems pubescent or puberulous, at any rate at k t . Stems pale yellow, subterete; idoremmce fascicled, 1-2 cm. Stems dark red (or pale yellow, and very sulcate); infloescence umbellate-racemose,

or panicdate, 3-12 cm. Stems dark red, terete to subq led ; leaves distinctly veined; inflorescence race-

mose or paniculate. Leavea 1-2 cm. long, closely reticulate; inflorescence racemose, 2-54 cm., dense;

pedicels 4-7 mm.; fruit 8-10 x 5-6 mm. h v e a 3-12cm. long, openly veined; inflorescence paniculate or compound

below, 5-13 cm., loose; pedicels 8-20 mm.; fruit 12-14 x 4-5 mm. Inflorescence corymbose-paniculate; leaves green, epruhose below; inner

sepals 9-11 mm. 113. B. chitria Infiorescence racemose, compound below; leaves grey, pruinose below; inner

Stems yellow, very sulcate; leaves subenervate; idoreecence subumbellate (oc-

Cf. 111. B. afghnicu

112. B. mimanth

sepals 6-7 mm. 1130. B. X p 8 d i

casionally subpanicdate). 114. B.eikkintensis Stems always quite glabrous. Stems very sulcate, pale yellow.

Inflorescence umbellate, 3-12-fld. 1140. B. sikkinzensis var. glabramm Inflorescence racemose, compound below, or paniculate. 115. B. ceratophylla

Inflorescence formally racemose, mostly 15-25-fld. Stems terete or slightly angled, OT dark purpIe.

Petale entire; inner eepals 4-5-8mm.; racemes open; pedicel6 6-1Omm.; fruit 9-11 mm., dark red or with blue bloom.

Leaves narrow, 1 : 4; Spines 4-6 mm., solitary. Leaves broader, 1 : 2-2.5.

116. B. a&&

Spines solitary, 6-10 mm. ; inner sepals 5-6 mm.; fruit oblong, dark red,

Spines 3-fld., 1-3 em.; inner sepals 7-8 mm.; fruit ovoid, 9-11 mm. pruinose blue. 117. B.Mbundu

Shoots green; leaves entire (or with 3-8 marginal apinules), mostly 5-9 cm.

Shoots purple; leaves with 8-15-spinules, mostly 24-5cm. long; fruit

Petals emarghte; inner sepals 8-10 mm.; racemes stiff; pedicels 2-5 mm.;

long; fruit bright red, pruinose blue. 118. B.aristata

dark red, pruinose blue. 118a. B. x mac*acantha

fruit bright red, epruinose, 12-13 mm. long. 119. B. &ria Intlorescence subumbellate or corymbose-paniculate, 2-10 (-lS)-fld.

Stems dark purple. Stems distinctly sulate; leaves entire; petale longer than inner sepals.

Stems subterete; leaves coarsely dentate; petals shorter thm inner sepals. 120. B. holetii

121. B. grantii

Spines present ; inflorescence subfascicled. 122. B. petitiana Spines absent ; infloscence racemose to paniculate. 123. B. afrimna

Stems pale yellow.

(112) Berberis micrantha Ahrendt in chrrd. IUust. 63, 618, 1841, 64, 426, 1944, anglice; chatterjee in Recorcls Bot. Survey of I d & , 16, nQ. 2 (studies in Indian Berberid.) 17, 1953, anglice; syn. B. aristata vm. mimanth Hook. f. et Thorns., Fl. Id. 1,222,1855, in part, a far a the specimens from Bhutan are concerned

Sterne subterete, to subaqled, not vemculose. Interraodes 1-2.5cm. Spine0 sub- concolomus, slender, plane, 3-fld., 1-5-2.5 cm. Leaves 10-20 x 5-10 mm., obovate- elliptic ; acute, mucronate, cuneate, subsessile ; margins 24-spinulose-subserrulate ; above slightly lustrous, deep yellow green, conspicuously reticulate; below pruinose grey, not, or scarcely, papillose. Race= 8-15-fld., 2-5-4 cm., including peduncle 2-4 mm., some-

Page 97: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia, 97

times slightly compound below. Pedicels 3 - 5 4 (-7) mm., glabrous; bracts puberulous, 2-2.5 mm. Flowers 10-12 mm. Prophylls red, acute, 2 x 0.9 mm. Outer sepals (H.C.C. 2), 3 4 x 2-2-5 mm., prophyl shaped, ovate-triangular with a truncate base. Median sepals 6-6 x 3.5405 mm., ovate-elliptic, obtuse. Inner sepah 65-76 x &5 mm., obovate. Petals (H.C.C. 3); 6-7 x 3 4 mm., obovate; apex rounded, shortly acutely emarginate with 2 obtuse lobes; cuneate with oblong, separate, non-marginal glands, 0.8 x 0.5 mm. Sbmm 4-4.5mm., produced, subobtuse to slightly apiculate. Ovules 34, shortly stipitate. Berries dark red, epruinose oblong, 8-10 x 5-6 mm., including style 1-1-25 mm.

Bhutan: fl., 1838, G S t h 1744 (Type, K); G a t h 121, in part (K). Cultivated: fl., 6 July 1939, 6 July 1940, 17 June 1942; fr. 7 Nov. 1940. A suberect

shrub reaching 6-8 ft., and quite hardy.

(113) Berberis chitria Lindl., Bot. Reg. 9, t. 729, 1823; syn. 'B. aristakc' Collett, Fl. Simlensis, 21,1902; Sims, Bot. Mag. 52, t. 2549; and many other authors; non DC.

Stems terete; shoots pubescent; mature puberulous or subglabrous. Internodes 3-5 cm. Spines concolorous, mostly 3-fld., plane, 1-2 (-3) cm. Leaves ovobate or elliptic, sub- sessile; both sides with loosely elevated venation and h e reticulation; above dull green; below slightly lustrous paler yellow green, sparsely indistinctly papillose. In$mescence 10-20-fld., loosely corymbose paniculate, with flowers formally in groups of three; (5-) 8-13 cm. long, including peduncle (1.5-) 4-6 (-8) cm.; subpeduncles 10-18 mm., each bearing 3 flowers. Pedicels glabrous, 8-15 mm.; bracts 2.5-4 mm. Flowers (H.C.C. 2/1), 1.6-2 cm. diameter. Prophylls 1.5 x 1 mm., either appressed to the flower or at the mid-point of the pedicel, truncate, acuminate. Outer sepals 6-7 x 5-6 mm., obovate. Inner sepals 9-10 x 7-5-5-9 mm., obovate. Petals broadly elliptic, 8-9 x 6-7 mm.; apex subacutely emarginate with rounded lobes ; base cuneate with separate oblong-elliptic glands, 1.2 x 0.6 mm. Stamens 7 mm., below the anthers thickened but not dentate; apex produced, subapiculate. Ovules 4-5, with stipes longer than themselves. Berries dark red-brown, epruinose, narrowly ovoid, or oblong-ellipsoid, excluding style 1-1-5 mm.

KEY 24a Leaves 4.5 (-7-12) x 2-3 (-4.5) cm.; margins mostly 3-9-spinose (1-2 mm.)-serrate

(1-2 mm.), distant 3-7 mm. ; fruit 12 x 4 mm. Leaves 2-4.5 om., mostly entire; fruit 12 x 6 mm.

113a. Berberis chitria var. chitria 113b. B. chitria var. occidentalis

(113n) Berberis chitria var. chitria. (Fig. 15) Nepal: 1821, fl., Wallich 1474 (1) A (K). Kumaon: fl., Wallich 1474 (2) A (K); Naini Tal, 7500 ft., fl., immature fruit, Strachey

& Winterbottom 2, in part (K); fl., Apr. 1844, Eooker & Thomson 718 (K); fl. 12 June 1847, Hooker & Thomson 626 (K).

Tehri: between Gundiard and Oura, 5000-7000 ft., fl. May 1879, Gamble 1014 (K); below Mussoorie, 30" 27' N., 78" 4' E., 5000-8000 ft., fl., 8 June 1897, Duthie 19860 (BM, K); 9OOO ft., May 1892, Gamble 23607 (K); Doota, Gamble 23662 (K).

Jaunsar: Mandhali, 8000 ft., fl. June 1893, Gamble 24405 (K); Lokar, 7500 ft., June 1891, Gamble 22800 (K); Kaltryan, 7000 ft., Gamble 23794 (K).

Cultivated: fl., 26 June 1939; fr. Nov. 1939 (K), 28 Nov. 1941. A shrub 10-12 ft., of which there is an illustration in Card. Illust. p. 455, Oct. 1944.

(113b) B. chitria var. occidentalis Ahrendt, J . Bot., Lond., 80 (Suppl.), 85, 1942 Jaunsar: Chakrata, 30" 45' N., 77" 50' E., 8500 ft., fl. 1 June 1897, Rich 2 (32B) (K);

Lokar, 8000 ft., fl. May 1892, Gamble 23789 (K); Mundali, fl. May 1929, Kharyal, 8.n. (BM).

rl JOURN. LJNN. S0C.-BOTANY, VOL. LVII

Page 98: Berberis and Mahonia : A taxonomic revision

98 L. W. A. AHRENDT Punjab: Simla district, 7000 ft., fl. 6 June 1877, Gamble 43298 (K); 7000 ft., fr. oct.

1877, Gamble 59468 (K); valley below Narkanda, 31" 15' N., 77" 20' E., 8OOO ft., 1 June 1873, Gamble 6127A (K); Schlich 26 (2), (0); 1 June 1878, Collett 1902 (K).

Chamba district: Alwaa, 8OOO ft., 27 June 1917, Stewart 2459 (K). Kadmir: 8500 ft., fl. 16 June 1939, Ludlow & S h e d 7309 (BM). Cultivated: fl. 25 June l942,lO June 1943; fi. Nov. 1940. This variety is a constituent

of 'the small-leaved specimens from Simla' of B. aristata var. mmuz.Ji.9 Hook. f. et Thoma. non DC. It represents B. chitria at the extreme western end of its distribution range, Jaunsar, Punjab and I(ashmrr * , and, mcordjng to Parker, is common at 6000-9000 ft., in the Simla and Parbar valleys; the previous variety belongs more to Kumaon, but extends &s far west as J a m where the two varieties meet.

KEY 2 4 ~ , to the hybrids connected with B. chitria Racemes open and loose.

Fruit 14 mm. long; pedicels 1-2 an.; style 1 mm. Fruit 10 nun. long; pedicels P 6 mm.; style short.

Racemes dense; style short.

113c. B. xpa~sOneii 113d. B. x ItaunknSie

113e. B. x spaethii

(113c) Berberis x parsonsii Schneid., J. Am. Arb. 4,202, 1923 Stems angled, mature red-brown to purple. Spines fairly stout, 6-22 mm., the later&

components slightly longer, 3-fld., plane to subsulcate. Leaves 25-50 x 9-20 mm., ovate- elliptic, obtuse to subacute, mucronate; cuneate petiole 2-5 mm.; margins subentire or 2-6-spinose-serrate (1-3 mm.) ; very openly reticulate; bright green; below grey prUinose scarcely papiuose. InJlorescence very loose, racemose or pseudo-umbellate-racemose, sometimes compound below, 5-7 cm., including peduncle 2-3-5 cm. Pedicels 1-2 cm.; bracts 2 - 5 4 mm. Pr0phyll-s red, acute. Outer sepah ovoid, twice aa large aa the bracts. Inner sep& 6-7 x 5-6 mm., elliptic to obovate. Petals 7 x 5-6 m., obovate, emarghate. Stanzens 5 mm., truncate. ovu2es 3-4, sessile. Berries oblong-obovoid to oblong-ellip- soid, dark red, slightly pruinose, 14 x 5-6 mm. Style, 1 mm.

Cultivated: in U.S.A. (Type, Herb. Am. Arb.). Reputed to be B. chitria x B. lyciurn? However, both these species have stipitate ovules, and their petals are shorter than their inner sepals. Not cultivated in England.

(113d) Berberis x hauniensis Schneid., I.c., 199, 1923 Stem sulcate, yellow red. Spines solitary, 1-2 cm., sulcate. Leaves 20-40 x 9-17 mm.;

oblong-obovate; obtuse; petiole, 5 mm.; margins with 4-5 spinules, openly reticulate; with the texture of B. chit&, but below pnzinose grey. Racemes 10-20-fld., compound below, 2-3.5 cm. Pedi& 4-6 mm., slender; bracts, 2-3 mm. Outer sepal8 3.5 mm. Inner sepals 5 mm. Pet4d-s 6 mm. Stamens 3 mm. Berries dark red, 10 x 5 m., ellipsoid almost estylose.

Cultivated: in U.S.A. (Type, Herb. Am. Arb.). Not cultivated in England.

(113e) Berberis x spaethii Schneid., l.c., 203, 1923 Stem glabrous, subterete. Intemtodes 2-36 cm. 8p.m solitary, 5-20 mm. Leaves

35-60 x 15-23 mm., oblong-elliptic to -ovate or -0bovate; petiole, 5 mm.; openly reticu- late; acute; dull green. Racemes dense, compound below, over 6 cm. Berries red-purple, pruinose, 11 x 6 mm. Style very short.

Cultivated: in U.S.A. (Type, Herb. Am. Arb.). Not cultivated in England. The information given about the preceding three hybrids depends on D r Schneider's original descriptions.

Page 99: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 99

(114) Berberis sikkimensis (Schneid.) Ahrendt, J . Bot., Lond., 80 (Suppl.), 85. 1942; Sealy, Bot. Mag., 168, N.S., t. 173, 1951; syn. B. chitria var. sikkimnsis Schneid; syn. B. aristata var.parvifira Hook. f. et Thorns., MS., in Herb. Kew; syn. B. aristakc var. rnicranthu Hook. f. et Thorns. in part

Stem very sulcate, always pubescent a t first on the shoots; mature pale yellow-grey, not verruculose. Internodes 6-25 mm. Spines 3-fld., h e l y sulcate, 1-2 cm., concolorous or slightly yellower than the stems. h u e s fairly coriaceous, obovate, acute, mucronate; 15-27 x 5-10 mm. ; cuneate, subsessile, margins slightly subrevolute, often entire, some- times 1-7-spinose (1-5 mm.)-serrate (1 mm.) distant 3-5 mm.; above deep bright green, dull or very slightly lustrous; below a t h s t pruinose grey; ultimately lustrous yellow- green, epapillose, or scarcely papillose; both sides subenervate, or at most with 2-3 lateral veins visible. Pedieels glabrous, 4-7 (-10) mm.; bracts red, ovate, acute, 1.5 x 0.9 mm. Prophylls yellow, sepaloid, 2 x 1.6 mm., acute. Outer sepals (H.C.C. 2) 5.5 x 4 mm., ovate, acute. Inner sepals l o x 7 mm. broadly obovate. Pet& (H.C.C. 3) 9 x 6.5 mm., subacute, entire or with apex slightly crenulate; base scarcely clawed, with subconcolorous, oblong, separate, submarginal glands 1.7 x 0.7 mm. Stamens 6 mm., produced, subapiculate. Ovules 4. Berries dark red, epruinose, narrowly ovoid, often curved, to 15 x 4-5 mm., excluding style 1.5 mm.

%Y %B

Stem with shoots pubescent, and the mature stems puberulous. Inflorescence 3-12-fld., umbellate. 114a. B. sikkimnais var. sikkirnensis Inflorescence 8-20-fld., subpaniculate. 114b. B. sikkimnsis var. baikyi

Stems with shoots puberulous, or glabrous; mature stems always glabrous. 114c. B. sikkimensis var. glubramea

(114a) Berberis sikkimensis var. sikkimensis Injlorescence 3-12 ft., umbellate or subumbella,te, occasionally racemose, 3-4 cm.,

including red peduncle 12-22 mm. Sikkim: Larhoery, 9OOO ft., 27 Aug. 1849, Hooker & Thomson, 8.n. (Type, K)-labelled

‘ aristata parviflora’, but probably meaning ‘aristata parvifolia’. Bhutan: Chembi, Rongang, Kuru Chu valley, Kurted, shrub 6 1 0 ft. ; flowers bright

yellow, 5500 ft., 3 May 1949, Ludlow, SherrifT & Hicks 18814 (BM). Cultivated: (L.A. 283, from Sikkim seed), fl. 17 June 1942, 25 May 1943; fr. 8 Nov.

1943. Also, a t Wisley, fr. 18 Oct. 1939; 2 Nov. 1930. A shrub 3-5 ft. This is part of B. aristata var. micrantha Hook. f. et Thorns., non DC, aa far aa ‘the

small-leaved specimens green below from Sikkim’ are concerned. Its tenderness in cultivation suggests relationship with some of the Ceylon and South Indian species, B. tinctoria, B. wightiana and B. ceylaniea, with which it should be compared. However, these have thick leaves which are pruinose and white or grey below, while B. sikkimnsis possesses thin green leaves.

(1144 Berberis sikkimensis var. baileyi Ahrendt, var.nov., inflorescentiis longiori- bus et subpaniculatis

InJZorescence 5-8 cm. long, 8-20-fld., compound below, or subpaniculate. Nepal: Gyang, fl. 3 June 1935, Bailey s.n. (Type, BM); Bailey s.n., fl. 4 June 1935

(BM); Nagarkot, 7OOOft., fr. 7 July 1935, Bailey s.n. (BM). This variety is not in cultivation.

(114c) Berberis sikkimensis var. glabramea Ahrendt, J. Bot. Lond., 80 (Suppl.),

Stem at first puberulous or glabrous; fhally glabrous. InJlorescence as in 114a, var. 87, 1942

sikkimnsis. 7-2

Page 100: Berberis and Mahonia : A taxonomic revision

100 L. W. A. ~ E N D T

Mid-weat Yunnan: Shunning, 24" 36' N., 99" 52' E., Wumulung, among thickets, 7500 ft., fr. 8 July 1938, Yu 16607 (Type, E). Not in cultivation.

We thus have a western variety from Nepal, with slight leanings in the compound infiorew3nce fo the m h t d and still more western B. chitria and still closer relationship to the following, and also Nepal-, speciea B. ceratqvhylk; and an eastern variety from extreme western China, still farther removed from B. chit& in character by its com- pletely glabrous stems. In dl cases the difference between the terete dark red stems of B. c h W , and the very sdcate pale yellow stems of B. &kkimwk maintains the specific distinction very clearly.

(115) Berberis ceratophylla G. Don, Syst. &rd. 1, 115, 1831; Ahrendt, J. Asiat.

Stems glabrous, very sulcate, pale yellow-grey. Internoda 1-2.5 om. Spine% 1-3-fld., fairly stout, 6-14 mm. Learn 2-46 x 1-2 cm., cuneate, s u k i l e , submute, entire; r m l y margins sparsely sphose (1 mm.)-senuliLte, distant 3-6 mm.; both sides with dis- tinct branched venation, finely reticulate; below at Grst slightly pruinose grey, finally epruinose, concolornus green. Racemes 1&25-fld., below compound, sometimes pani- c h t e , 5-8 cm., hcluding peduncle, 1-2 cm. Pmphylylls 1.5 x 1 mm., acute. Outer sepals 2.5 x 1.75 mm., ovate, subacute. &€&.an se@ 4 x 2 mm., oblong-elliptic. Inner sepals 7 x 3.5 mm., narrowly obovate. Petals 6 x 3 mm., obovate. Stanzens 4 mm., distinctly produced, shortly apicuhte. ovules 43. B e d oblong-ovoid, very dark red or purple, epruinose, c. 12 x 6 mm., excluding style, 1-1.5 mm.

Nepal: Wallich, s.n. (Type, 0); Foketey, 13,000-14,000ft., 1930, Dhwoj 492 (BM); on route, Langtang Khola, forest region, 15 ft,; flowers clear yellow, 9OOO ft., 6 June 1949, Pol@ 161 (BM); Thare, open scrub, 15 ft., a t 7000 ft., 2 June 1949, Polunin 60 (BM). En descendant de Namche Bazar, rive gauche de la Dudh Kose, 9500ft., fr. 26 oct. 1964, Zimmermann 1835 (Geneva).

Bhutan: (central) Chendebi, 7500 ft., fl. 12 May 1937, Ludlow & S h e d 3031 (BM). Shrub 4 4 ft., on open grassy hillaides. Not cultivated.

Further investigation is desirable, preferably with cultivated material aa well, to determine the relationship of B. ~rcubphylla with B. sik;kimen8is which, although other- Wise distinct, approaches close to the former in its variety baileyi. We may note that B. sikkimmk has larger flowers with broader components.

(1 16) Berberis affinis G. Don, 1.c. ; Ahrendt, 1.c. ; syn. B.jbdmnda var. .fink Ahrendt,

Stem glabrous, terete to subangled, pale yellow. Internodes 1-5-3 (-5) cm. Spines absent, or concolornus and weak, 4 4 mm. Leaves 4 - 8 ~ 1-2 om., or to 5 x 1 cm., narrowly obovate to oblanceolate; apex narrowly acuminate or subacuminate; base attenuate, subsessile or with petiole 1-3 mm. ; margins entire, rarely with 2 4 spinose teeth ; both sides openly reticulate; above dull; below paler sublustrous green. Jh-ctma 10-25-fld., 4-8 cm., including peduncle, 1-2 cm. P e d W 6-8 mm.; bracts 2-3.5 mm. Flowers 8-9 mm. diameter. Berries ovoid, 1 0 ~ 4 . 5 mm., excluding style, 1 mm.

Kumwn: Namik, 30" 7' N., 80" 7' E., fl. 7700 ft., Strachey & Winterbottom 4 (BM). Not cultivated.

Soc. Beng. (Sci.), (3) 11, 1, 1945

J . Bot., W., 80 (Suppl.), 89, 1942

(117) Berberis floribunda Wall. ex G. Don, Syst. Card. 1,115,1831; Ahrendt, J . Bat., Lond. 80, (Suppl.), 87, 1942

Shrub with arching branches. Stems shoots green, glabrous; mature pale yellow, terete or subterete; sparsely distinctly verruculose. Internodes 2 - 6 5 cm. Spines absent, or 1-3-fld., 4-10 mm., with shorter lateral parts. Leaves 2.5-5 x 1-2-2.4 cm., elliptic to

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Berberis and Mahonia 101

ovate-elliptic; base contracted to petiole 1-3 mm. ; apex obtuse; margins entire, rarely with a few spinules (except on the coarsely serrate leaves of new summer shoots) ; both sides with venation openly branched, but with open reticulation scarcely visible or very indistinct; above dull; below sublustrous paler yellow-green, papillose. Racemes 10- 25-0d., mostly sessile. Pedicels 4-7 mm. (rarely, basally, to 12 mm.); bracts 1-2 mm., acuminate. Flowers H.C.C. 3. Outer sepals 2.5 x 1.5 mm., oblong-ovate, subacute, pro- phylloid. Median sepals 4 x 3 mm., elliptic. Inner se@ 6 x 4-5 mm., obovate, oblong- elliptic, entire; base clawed with ovate, subacute, separate, non-marginal glands, C. 0.7 x 0.3 mm. Stamens 3.4 mm., not or scarcely produced, truncate or rounded- truncate. O& 4 4 . Berries dark red, ha l ly almost purple, pruinose blue, 6.5-7.5 x 4.5-5 mm., broadly oblong-ellipsoid, excluding style 0.5 mm.

Nepal: Wallich (Type); between Langtang and Birdim, in Juniper scrub, with small flowers in long racemes; shrub up to 20 ft., 9500-11,OOO ft., 28 June 1949, Polunin 704 (BM).

Cultivated: L.A. 205,0. 9 May 1938, 21 May 1942; fr. 14 Nov. 1943. This has been generally distributed in cultivation, erroneously as ' B . aristata', or ' B . nzaccrO(phy1lla'. It is much more common in cultivation than B. aristata which is very rare.

(118) Berberis aristata DC., Syst. 2 , 8 , 1821; Ahrendt, J . Bot., Lund., 80 (Suppl.),

Stems glabrous, pale yellow, terete. Internodes 3-6 cm. Spines fairly stout, 1-3 cm., solitary towards apex of stems, threefold towards their base. Leaves (2 .6 ) 4-6 (-9) x (1-2-) 2-2.6 (3.7) em., obovate to obovate-elliptic; subacute to obtuse; cuneate, sub- sessile; entire, rarely with 3-5 (-8) marginal teeth; above deep green, sublustrous, fairly openly reticulate ; below slightly paler lustrous yellow green, openly veined, scarcely reticulate. Racemes 8-25-fld., 4-6 cm., including peduncle, 1-35 cm. Pediceb 6-10 mm., with greenish lanceolate bracts 3 x 0.75 mm. Flowers 11-14 mm. diameter. Outer sepals yellow, lanceolate, 3 x 1-1.5 mm. Median sepals narrowly ovate to elliptic, 5-6 x 2.5- 3.5 mm. Inner sepals 7-8 x P 5 mm., obovate. Petals 6-7 x 4 mm., obovate to oblong- obovate; entire; base cuneate with separate non-marginal glands, 1.2 x 0.4 mm. StamRns 4.5-8.5 mm., produced, shortly apiculate or obtusely subconical. 0wd.a (3-) 4-5. Berries ovoid to oblong-ovoid, bright red, slightly pruinose blue, 9-11 x 6-7 mm., in- cluding style 1 mm.

Nepal: 1818, Wallich 1474 (1) B (BM); Buchanan-Hamilton s.n. (Type); Matia, 28" 3' N., 82" 6' E., 25 May 1886 (BM).

Cultivated: (Kew 281), fl. 19 May 1939; fr. 19 Oct. 1939 (K). In cultivation, an erect shrub to 12 ft. I know only this one cultivated plant; it has defied all attempts to propagate it by cuttings. There is a photograph in Cnrd. Illust. p. 455, Oct. 1944. Most plants passing by this name in cultivation are B. floribunda; some may be B. chitria.

91, 1942

(118a) Berberis x macracanthaschrader, Linnuea, 12,366,1838; Schneid., Mitt. Dtsch.

Shrub 10-12 ft. Stems sulcate to subterete; shoots purplish; mature yellow grey. Internodes 16-3 cm. Spines on older stems weak, otherwise fairly stout, 1.5-3 em., sul- cate. Leaves 2-54.5 x 1-2 em., including petiole to 1 em.; obovate, obtuse, mucronate; margins with 5-15 spinose serrations (0.5-1 mm.); venatioqas in B. chitria: below finally bright green. Racemes loose, 8-25-fld., with peduncle, or added basal pedicel, 1-2 cm. Pedicels 5-10 mm. Prophylls 1.5-2-5 mm. Outer sepals 3-4 x 2-3 mm., ovate. Inner sepals 6 x 4.5 mm., obovate. Petals obovate-elliptic, entire or slightly emarginate. Stamens from one-half to three-quarters as long as the petals, scarcely apiculate. Ovules 2 4 (-5). Berries dark red, slightly pruinose purple, ellipsoid to ovoid, 8-10 x 5-6 mm. Style more or less short.

Dendr. Ges. 14, 176, 1907; Schneid., J . Am. Arb. 4, 200, 1923

Page 102: Berberis and Mahonia : A taxonomic revision

102 L. W. A. AERENDT

in England now.

KEY 240 Leavea broad, 1 : 2-2.6, openly veined; raeemea 8-20-fld.; pedimls mostly 7-10 1z~. ;

1lSa. B. x macracarrtha Leaves narrower, 1 : 2.63, closely reticulate; racemea 6-8-fld. ; pedicels mostly 5-7 mm. ;

118b. B. x nuzcracunthu var. &hra

(118b) Berberisxmacracmtha var. pulchra Schneid., J. Am. Arb. 4, 201, 1923.

(119) Berberis coriaria Royle ex Lindl., Bot. Reg. 27, t. 46, 1841; Ahrendt, J. Bot.,

Stem glabrous, h e l y sulcate, or slightly angled, sometimes terete; mature pale yellow. I&& (2-) 2.5-4 cm. S p * w fairly stout, mostly 3-fld., plane (1-) 1-6-26 (-3) om. Leaves 2-5xO.7-1.8 om.; fairly narrowly obovate; apex acute to subacute; cuneate, s u b s d e ; margins entire; above dull green with venation openly elevated; below concolorous lustrous green, finely reticulate. Racem~s peduncled, but with 1-2 additional flowers a t the base, 10-24-fld. Peduda 1-2cm. Pedicel-s with bracts, 1-5-2 mm. Flowers 1.4-1-7 cm. diameter. Outer s e p b 6.5-7 x 36-4 mm., ovate, sub- acute. Inner 8 e p b 8-9 x 7-7-25 mm., obovate, subacute. P& 8 x 4.5 mm., oblong- elliptic; acutely emarginate with acute lobes; base broadly clawed with separate oblong glanh, 1.6 x 0.8 mm. Stamem 5.5 mm., produced, a p i d t e . ovules 4-5. Be& bright red, epruinose, oblong, 12-5-13-5 x 6-76 mm., excluding style 1-16 mm.

Cultivated: reputed to be B. chitria x B. tdgark. As far aa is known, not cultivated

style short.

style longer.

Also of hybrid origin; not cultivated in England

Led., 80 (Suppl.), 89, 1942. (Fig. 16)

KEY 2 4 ~ Racemes dense, rigid, 6 5 cm.; pedicels 3-5 111111. Racemes lax, 6 8 cm.; pedicels 6-12 mm.

(119a) Berberis coriaria var. coriaria Kumaon: 1835, Royle, s.n., seeds only; 1821, W&ch 1474B (0, K); Dwali, Strachey

& Winterbottom 3 (K); Hooker & Thomson, s.n. (K); Almora, Dhakari, lO,OOOft., July, Oct. 1910, Lambert, s.n. (K); Almora, Rur, 10,OOO ft., fr. Oct. 1920, Lambert 8.n. (K); Jullut, 10,OOO ft., fr., Strachey & Madden 2103 (0); 1933, seed o.nly, Osmaaton 51,

119a. B. &ria var. &ria 119b. B. &ria var. p a t h

in part. Tehri-Garhwal: above Deota, 8000-9000 ft., fl. 20 May 1893, Duthie, s.n. (BM). Cultivated in 1841, from seed collected by Royle (Type). Also from 1933 seed, col-

lected by Osmaston (51) (Kew 28), fl. 26 June 1939, fr. 18 Od. 1939 (K). A magni6cent erect shrub to 10 ft.

(119b) Berberis coriaria var. patula Ahrendt, J. Bot., Lond., 80 (Suppl.), 91, 1942 Kumaon: Naini Td, 7500 ft., Strachey & Winterbottom 2, in part (BM). Cultivated: a t Wisley, fl. 19 May 1939; fr. 10 Nov. 1938 (Type, BM).

(120) Berberis holstii Engler, Abh. p e w 8 . A M . Wbs . 1894,64,1894; Pfinzenw. 0st.- Afr., C, 181, 1895; Spra.gue in Hook., Ic. PI. 31, t. 3021, 1915.

8tem.s glabrous, sulcate, dark red-purple. I&& 1-5-2 cm. S p i m 1 (-3)-fld., 2-3 cm., very sulcate. Leaves 1-36 x 0.4-1-1 cm., obovate or narrowly obovate; acute to subacuminate; entire, rarely with 1-2 spinules; above dull green with elevated reti- culation (but on the older leaves sometimes subenemate); below concolorous green, or scarcely paler, indistinctly reticulate, not or scarcely papillose. Inflorescence 8-12-fld., racemose, or partly umbellate, pseudumbellate, or pseudo-fascicled, often with the lower

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Berberis and Mahonia 103

part compound; 2-3 (-5) cm. long. Pedicels glabrous, 7-8 mm.; bracts lanceolate, 3-3.5 mm. ; but sometimes foliaceous, obovate-spathulate, cuneate, entire, 20 x 6 mm. Prophylls sepaloid, 3 x 1.5 mm., ovate, subacute. Outer sepaih 4 x 3 mm., ovate, subacute. Inner sepals 6 x 3.5 mm., obovate, subacute. Petals 6.5 x 4 mm., obovate, entire; base truncate, not clawed, with lanceolate separate non-marginal glands, 1.5 x 0-3 mm. Stamens 5-5 mm. ; below the anthers distinctly thickened, sometimes almost rounded- subdentate; apex not produced, rounded. OvUEes 4-5, stipitate. Berries dark red, pruinose-blue, oblong-ellipsoid, 7-8 x 4-5 mm., excluding style 0-5-0.75 mm.

East Africa : Somaliland Protectorate: Gar [Mt.] Libah, 7900 ft., on summit on edge of escarpment

Uganda: summit of Mt. Moroto, 9700 ft., Feb. 1936, Eggeling 2890 (K) Kenya: Aberdare Mts., Kinangop forest, 8000-10,000 ft., flowers yellow, fruit purple,

Munro 101 (0). Mt. Kenya, 9O00-10,000 ft., fl. 24 June 1909, Battiscombe 92 (K). Tanganyika: Masai district, Longido Mt., 8500 ft., growing in association with Pteri-

dium, I k , Eheodendron, Myrsine in clearings in T e c h , Cassipourea, Lachnostylis mist- forest, locally scattered, 10 ft. tall, 16 Jan. 1936, Greenway 4377. Mbulu district, west slopes of Mt. Hanang and foot-slopes of Mt. Gendabe, rare among rocks in Dombeya, Combreturn gueinzii, A m i a hockii, Hetermnmpha arborescem, open bushland with Euclea spp., Osyris abyssinim, Protea angolensis, in a reddish-orange loam of gneiss origin on stony rocky slope, a much-branched spiny shrub up to 5 ft. tall with bright yellow flowers, 11 Feb. 1946, Greenway 7693 (K, 0). Mbulu district, Mt. Hanang, 8500 ft., on lava outcrop in Juniperus procera zone on south-west crater-rim bordering grassland, without flower or fruit, 2 Sept. 1932, B. D. Burtt 4020 (K, 0). Kilimanjaro, north slope above Laitokitok, 7100 ft., in open scrub-forest above evergreen high forest, with Pteridium and Hypericum, an erect little-branched deciduous shrub up to 8f t . high, flowers lemon yellow, 15 Feb. 1933, Gilbert Rogers 415. Kilimanjaro, east side, Keni, 4000ft., May 1927, Haarer 688 (K). Usambara Mts., Holst 427 (K, isotype). West Usambara Mts., Malindi to Lushoto road near Cligg’s House, almost a t bottom of valley, in thickets, shrub, sometimes scrambling, stems up to 1 in. in diameter, bark smooth grey, wood bright yellow, Kishambaa names Kilasho, Kidangadwa, roots used as a stomach medicine, 19 Feb. 1947, Hughes 35 (K, 0). Uluguru Mts., Lukwangule, 8000 ft., bush, flowers yellow, 2 Jan. 1934, Michelmore 877 (K). Lukwangule plateau, 8200 ft., two patches in grassland on summit near marsh, 3ft. spiny shrub, flowers yellow, 30 rJan. 1935, E. M. Bruce 708 (K). Iringa district, Dabaga, 7 Feb. 1932, Lynes 14 (K).* This species is not in cultivation.

(121) Berberis grantii Ahrendt, sp.nov., aff. B. Jwlstii, sed ramulis teretibus, foliis crasse dentatis, petalis sepalis interioribus multo brevioribus distinguenda

Stems glabrous, slender, terete or subterete, dark purple. Internodes 1-2 cm. Spines to 1 cm., fairly stout. Leaves to 4 x 1.5 cm., or 3 x 1.4 cm.; obovate; base cuneate, sub- sessile; margins with 4-14 conspicuous aristate (2-3 mm.) teeth (1-2 mm.), distant 2-5 mm. ; open reticulation conspicuous and elevated, close reticulation more fine; above somewhat lustrous green ; below paler yellow-green, dull. InJEorescence 12-18-fld., corymbose-paniculate, 3.5-6 cm., including peduncle 1-3 cm. long. Inner sepals 6 x 2.5 mm., oblong-elliptic. Petals 4.5 x 2 nun., obovate, entire. Berries 7 x 5 mm., obovoid. Style, c . 0.3-0.5 mm.

facing Heis, 25 Jan. 1945, Glover & Gililand 536 (K, 0).

East Africa : Kenya: Slopes of Mt. Longonot, 9OOO ft., shrub 3 ft., Mar. 1922, Dummer 5152 (K). Tanganylka: West Usambara Mts., 7000 ft., shrub 4-6 ft., Sept. 1922, D. K. S. Grant

19 (K, holotype). West Usambara Mts., Mgamba, Pitt Schenkel449 (0). * B. ho&iiEngl. isnowrecorded from N y h Plateau, Nyasaland (see Mem. &V.Y. Bot. Gard. 8, 215,

1953).

Page 104: Berberis and Mahonia : A taxonomic revision

104 L. W. A. ABRENDT Of the three species of the group Septentrionales which occur south of the Equator, it

may be noticed that B. laolstii comes from between 11" N. and 11" S. latitude, B. grantii from 1" to 5" S., and the Javan B. mmthzgh from 8's.

(122) Berberis petitiana Schneid., BuU. Herb. Boissier, (2), 5,455, 1905 Stem glabrous, terete; pale yellow to grey. S p i w concolorow, slender, sulcate,

1-8-3 cm. Leaves 1-2-3 x 0 4 1 . 2 cm., oblanceolate, obtuse, subsesaile, entire, mucronate, closely reticulate both sides; below papillose, grey, pruinose. InJlorescence 24-0d., pseudumbellate or subfaacicled with a short peduncle. Pedicel.s 12-15 mm. Flowers and

Eaat Africa: Ethiopia: Menisa, Dillon & Petit (Type). Not cultivated. This species

Apparently related here, and, presumably, with pale yellow stems is:

fruit unknown.

comes from 5 or more degrees north of the Equator.

(123) BERBEBIS APEICANA Hebenatr. et Ludw. ex Schultea f., Syst, Veg., 7 , 11, 1829 This is described as 'unarmed', but with 'stems otherwise like B. vulsar;s; leaves 1 in.

long, attenuate to petiole, apex apiculate; idloreacence subpaniculate, with racemes almost twice aa long as the leaves; flowers about half aa large aa those of B. mlgarh, but otherwise similar; petals white'. Almost certainly Nitraria retusa (Forsk.) Aschers. collected by Hebenatreit and Ludwig in the neighbourhood of Carthage.

Subsect. Umbellatae Schneid., Bull. Herb. B&skr, (2), 5, 45, 1905 Stem diverse. Inter?wde.s (4 mm.) 2-5-3.5 (4 .5) cm. Spines (0.7-) 1-2 (-2.5) cm.

Leaves evergreen, subcoriaceous, otherwise diverse. InJloesmence mostly 2-10-fld., sub- umbellate or subfaacicled; occaSiody 10-25-fld., and racemose. Inner sepal8 6-86 mm., always longer than the petals. Stamem not, or scaxcely produced, truncate. Ovules 6 5 , sessile (unless stated), or shortly stipitate. Be& dark red, red-brown, or bright red, nearly always epruinose (unless stated), (8-) 9-10 (-11) mm. long.

Punjab, Nepal, Sikkim, Bhutan.

%Y 25, to Subsect. Umbellatae (Maps 13 and 16) Internodes 1-2-5 em. Stems glabrous.

Leaves grey, pruinose below. Flowers !+I0 mm. diameter.

Leaves closely reticulate, margins spinulose; subsessile or with petiole 5-8 mm. 124. B-urnbellata

Leaves openly reticulate, entire; petiole 8-20 111111. 125. B. brandkiana 124c. B. mmpbellii

Leaves green, epruinose below. 126. B.walterana

Stems subterete; peduncle 2-3 cm. ; pedimls 5-9 mm. 127. B. duihieana Stems sulmte; peduncle 10-15 mm.; pedicels 10-15 mm. 128. B. tlunnaonkna

Cf. 133. B. forshxdkzna

Flowers 14 mm. diameter; leaves slightly veined, subseasib.

stem puberulous.

Internodes 479 (-12) mm.

(124) Berberis umbellata Wall. ex G. Don, @en. Syst. 1, 116,1831; Ahrendt, J. Bot., Lvnd., 80 (Suppl.), 93 1942; 8ealy, Bot. Mag., 168, N.S. t. 145, 1951. (Fig. 17)

Stems with young shoots deep red, sulcate; mature red-brown, sparsely, coarsely (though partly he ly) verruculose (black), older yellow-brown to grey, subsulcate to sub- terete. Spines sometimes absent, mostly 3-0d., 7-17 mm., sulcate with the lateral parts shorter. Leaves (2-) 3-5-44 (-5.5) x (0-7)- 1.2-1.6 (-2.5) cm.; subacute to obtuse, mucro-

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Berberis and Mahonia 105

nate; shortly cuneate to decurrent petiole 5-8 mm.; margins 5-12 (-20)-spinulose (0.25-1 mm.)-subserrulate, distant (1-) 2 4 mm. (but on the shoots entire) ; slightly lustrous above ; below grey, densely papillose ; both sides conspicuously and fairly closely reticulate, with the venation impressed above, and the younger leaves showing less venation. Inflorescence 3-6-fld., rarely to 10-fld., subumbellate or fascicled, 1-5-3.5 cm., including peduncle 1-1-5 cm. Pedicels glabrous (0-5-) 1-2 cm. ; bracts, 2 x 1.8 mm., ovate, acute. Flowers H.C.C. 2. Prophylls 2.5 x 1.75 mm., red, triangular. Outer sepals 3.5-4-5 x 2.5-3 mm., ovate, acute. Inner sepals 5-6 (-7) x 3.25-4 ( 4 - 5 mm.) obovate. Petuls 5-5-6.5 x 4-5 mm., obovate, entire; base scarcely clawed with lanceolate glands, 1 x 0.4 mm. Stamens 3 mm. Ovules stipitate. Be& dark red, oblong, estylose, 1041 (-12) x 4-5-5.5 (-6) mm., pruinose purple. Seeds dark brown.

KEY 258. lnternodes 2-24 cm.; shrub 5-7 ft.; leaves dark green, slightly lustrous; stems not

Internodes 1-2 cm.; shrub 2-3 ft.; leaves very lustrous bright green; stems distinctly stoloniferous. 124a. B. urnbellata var. urnbellata

stoloniferous. 124b. B. umbellata var. brianii

( 1 2 4 ~ ) Berberis umbellata var. umbellata Nepal: 1818, Wallich 1475, in part (Type). Garhwal: Rikshin range, 10,000 ft., 13 May 1898, Duthie 21040 (BM). Bhutan: Ringshi Timpu, 13,000 ft., 29 July 1919, Cooper 1804 (BM). Kumaon: Blinkworth s.n. (BM). Cultivated: L.A. 80 (source of material for Bot. Nq. t. 145), fl. 18 May 1938; 21 May

1942; fr. 31 Oct. 1939, 20 Oct. 1940. A good garden plant, but partly defoliated in ordinary winters, and completely so in severe ones.

(124b) Berberis umbellata var. brianii Ahrendt, J. Bot., Lond., 80 (Suppl.), 94,

Stems densely, coarsely concolorous-verruculose. Leaves smaller than in the preceding variety, less toothed, less veined, bright green, more lustrous. Prophyll-s 2.5 x 1.5 mm., red, triangular. Outer sepals 4-5 x 3 4 mm. Inner sepals 7-8 x 6-7 mm., all H.C.C. 2. Petals, H.C.C. 3, 6-7 x 6-7 mm., broadly obovate, entire, cuneate. Stamens 5 mm.

Sikkim : Hook. f. et. Thoms, s.n., 8000-10,ocH) ft. (0), labelled B. urktatu var. umbeZhta Hook. f. et Thoms.; Rhenek, about 1932, Pradhan, seed only.

Cultivated: L.A. 113, from Pradhan’s seed, fl. 21 May 1942; fr. 20 Oct. 1940 (Types, 0, HM).

1942

(124c) Berberis campbellii Ahrendt, Gard. Illust. no. 3307,64,425,426,20 Sept. 1944 Stem with young shoots green; mature dark red to red-brown, partly sparsely verrucu-

lose. Spines 3-fld., 4-14 mm. Leaves to 4 x 1.7 cm., obovate-elliptic, subacute; margins to- wards apex 6-10-spinose (1.5 mm.)-serrate (1.5 mm.), distant 2 4 mm. ; above dull with few lateral veins ; below sparsely papillose. InJorescence 3-6-fld., fascicled. Pedicels 11-16 mm. Outer sepals 6-7 x 4-6 mm. Mediun sepals 8 x 7 mm. Inner sepal8 9-10 x 7-9 mm., all broadly elliptic obovate, and of colour H.C.C. 2/1. Petals, H.C.C. 212, 6-7 x 5-6 mm., spathulate-obovate. Berries dark red, oblong, 11 x 6 mm.

cultivated: L.A. 345, fl. 18 June 1941,20 May 1942; fr. 13 Jan. 1943. The plant came from Kew labelled ‘B. mucrosepala’, and is named in honour of the Curator. It is not very profitable to surmise whether it might be -a hybrid between B. macrosepalu and B . urnbellata; from the latter it differs in its narrower leaves and very large flowers; from the former in its evergreen leaves and smaller dark red berries.

Page 106: Berberis and Mahonia : A taxonomic revision

106 L. W. A. AEU~ENDT

(125) B. brandisiana Ahrendt, J. Asid. Soc. Beng. (Sci.), (3) 11,2, 1945 Stem h e l y subsulcate to subterete, dark red, lustrous. Internodes 1-5-2-5 cm. spines

absent, or solitary, 3-10 mm. Leaves 2-4 x 1-2.2 cm., excluding petiole 0-5-2 cm. ; base contracted; mar@ entire, m l y 8-15 spinules; apex obtuse, mucromte; both sides d d , with venation branched but not reticulate. Injikescmm W-fld., umbebte-racemose, 2 - 5 4 cm. Ped& 1-1-5 mm. Berries red, firm, obovoid, 9 x 6 mm. Style absent, or rarely extremely short and indistinct.

Funjab Himahya: Narkanda, fr. 1876, Brandis a.n. (Type, K, Sheet 1229); Murree, 5000 ft., 18 Apr. 1917, Stewart 1554, in part (K). Not cultivated.

(126) Berberis walterana Ahrendt, J. Bot., Land., 80 (Suppl.), 96, 1942 Stem rigidly arching, spreading, thick; shoots very sulmte, yellowish green to reddish ;

mature very sulcate, red-brown not verruculose. I n t e d 2-4 cm. Spines 3-fld., stout, sulcate, 1-24 cm., 1 - 5 4 mm. broad at the base. Leaves 2-4.5~0.4-1.3 cm., elliptic, obtuse, subwile; margins revolute, 7-ll-spinose (1-2 mm.)-subserrulate, distant 3 4 mm.; above lustrous deep green with lateral veins indistinct; below slightly paler lustrous yellow-green, venation openly branched and subelevated. In@escence 6-10 (-15)-fld., fascicled to umbellate-subfascicled, 2 - 5 4 cm. Pedicels (1 .5) 2 3 . 5 cm. F b r s H.C.C. 2. Prophylls 2 x 1-5 mm., red, trianguh. Outer sep& 4-5 x 3-46 mm., suborbicular. Inner sepals 6 x 4 mm., obovate. Petals 3-4 x 2-3 mm., obovate, rounded, entire; base scarcely clawed, with narrowly ovate, acute glands, 0-7 x 0-25 mm. Stamna 2.5 mm. Ovules subseasile. Berries red-brown, oblong, 9-10 x 5-6.5 mm.

Nepal: seed probably sent (though there is no certainty on this matter) with seed of B. umbellata, before 1900, and labelled ‘B. macrophylla’.

Cultivated: LA 62 derived from seed cited-fl. 3 June 1942,20 June 1945; fr. not quite mature, 14 Nov. 1939,8 Nov. 1940 (Type, BM). A most attractive plant for its unusual crinkly green leaves.

(127) Berberis duthieana Schneid., Bull. Herb. Boissier, (2), 8, 200, 1908 Stem puberulous, aubangled, pale yellow-grey. Spines solitary, mostly 1-2 cm. Leaves

20-30 x (4) 10 mm., entire, green both aides, papillose below. In@escence !2-7-fld., sub- umbellate, W c m . , including peduncle, 2-3cm. Ped& 5-9mm. Be& oblong, 8-10 mm.

Nepal: Duthie 2696 (Type). Not cultivated.

(128) Berberis thomsoniana Schneid, l.c., 1908 Stem with young shoots green, puberulous ; mature yellow,glabrous, verysulcate. Inter-

nodes 1 - 5 4 cm. Spines 3-fld., 6-13 mm. Leaves 2-4 x 1-1.5 cm., narrowly oblong-obovate ; base cuneate to deckent petiole 1-3 mm.; margins entire; apex obtuse, mucronate; venation branched on both sides; below paler, papillose. I?$omsct?cence 5-8 (-12)-fld., um- bekte-racemose, 3-5 cm., including peduncle 1-1-5 cm. P&& 1-2 cm. Outer sep& 3.5 x 1-5 mm., ovate, acute. Mediun sep& 4.5 x 2-5 mm., ovate-elliptic, obtuse. Ovules sessile. Be& oblong, 8-9 x 5-6 mm.

Sikkim: Tonglo, lO,OOOft., Hooker & Thomson, 39 (Type, K); Dmjeeling, 12 Sept. 1875, Clarke, s.n.

E. Nepal: Nov. 25, Wdich, s.n. Not in cultivation.

Subsect. Lindleyanae Ahrendt, subsect. nov., infloreacentiis elongatis, racemosis, hct ibus eatylosis

Kashmir, Nepal, S.E. Tibet.

Page 107: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonie 107

KXY 26, to Subsect. Lindleyanae Internodes 4-9 (-12) mm. Cf. 133. B. forskalhna Internodes 1.2-5 cm.

Stems yellow, subterete; leaves green, epruinose below; inflorescence 2.5-4 cm. Leaves closely spinose; pedicels 8-17 mm.; stamens truncate. Leaves entire ; pedicels 3-5 mm. ; stamens apiculate.

Leaves very narrow, 1 : 5; outer sepals 6 mm. ; inner sepals 8.5 mm. ; petals 4-5 mm. ;

Leaves broader, 1 : 2.5; outer sepals 2.5 mm.; inner sepals 6 m.; petals 5.5 mm.;

129. B. kashmirana 130. B. deeandolhna

Stems purple, sulcate; leaves grey, pruinose below; inflorescence 5-8 cm.

pedicles 1-3 cm. 131. B. kongboensia

pedicels 5-10 mm. 132. B. lilzdleyana

(129) Berberis kashmirana Ahrendt, J. Bot., Lond. 80 (Suppl.), 97, 1943 Stems glabrous, terete to subsulcate; mature yellow. Spines yellow-brown, 3-fld.,

1-1-5 cm. Intewwda 2-3 cm. Leaves 3-6 x 1-14 em., including decurrent 2-6 mm.; narrowly obovate; apex acute to subacuminate; margins 15-25-spinose (1.5 mm.)-ser- rate, distant, 1-5-3 mm. ; both sides with closely elevated reticulation; above dull green; below concolorous, not or scarcely papillose. Racenzes 8-10-fld., 3 4 cm. Pedicele glabrous, 8-17 mm. Outer sepals 5.5 x 3.5 mm., ovate, acute. Inner sepals 6-7 x 5.5- 6 mm., obovate. Petals 4.5 x 4 mm., broadly elliptic; apex subentire; base not clawed, with separate elliptic glands, 0.9 x 0.6 111111. Stamens 4 mm. ovu2es stipitate. Berries oblong-ellipsoid, 9-10 x 4 - 6 5 mm.

Kashmir: Mantnar valley, near Desu, 11,500 ft., fl. 22 June 1939, Ludlow 122 (Type, BM). ‘Rather a fine Berberis, about 4 4 ft., amidst Juniper scrub, on south facing slope of valley’; Mantnar Nullah, Bringhi valley, 11,500ft., fr. 3 Sept. 1940, Ludlow & SherrifT 8047 (BM). Not cultivated. Seeds labelled ‘Berberis hashmiram, Ludlow & SherriE 8281 ’ have been distributed. But the number should be Ludlow & SherrifF 8181, and the species is B. pzchyacantha which differs conspicuously in its dark red stems, broadly obovate, rounded leaves, and ovaries containing ody 2 ovules.

(130) Berberis decandolleana Ahrendt, sp. vel hybr. nov., aff. B. coriarh a qua inter alia fructibus estylosis distinguenda

Stems glabrous; mature pale yellow, not or scarcely verruculose, at &st slightly sulcate, finally subterete. Internodap 3 4 cm. Spines concolorous, 1 (-3)-fld., stout, 1-2 cm. Leaves obovate, to 5 . 5 ~ 2 cm., including petiole 4 4 mm.; margins entire; apex sub- acute; branched venation clearly visible, reticulation h e ; above dull bright green ; below at f ist pruinose grey, finally green, epruinose, papillose. InJlorescence 10-16-fld.) rigidly racemose, 4-5 cm., including peduncle 1-2 cm. Pedicels stout, 2.54 mm. ; bracts 1-2 mm. Outer sepals 3 4 x 2-2.5 mm., ovate, acute. Median sepd.9 4.5 x 2.5 mm., ovate, acute. Inner sepals &6 x 4.5-5 mm., broadly obovate. Petals 4.5 x 2.5 mm. Starnew 4 mm., conspicuously produced, apiculate. Berries bright red, ovoid, epruinose, 12-13 x 6-7 mm. Seeds dark purple-brown.

Cultivated (Kew DN 31): fl. 1938; fr. 1939 (K) (Type, BM). This plant very much resembles B. coriaria which has very much larger flowers, and a conspicuously long style. Like B. miaria, it is very ornamental in berry.

(131) Berberis kongboensis Ahrendt, J. Bot., Lond., 80 (Suppl.), 97, 1943. (Fig. 18) Xtems glabrous, sulcate ; shoots red; mature slightly lustrous, deep red-purple. Inter-

d s 2 4 c m . Spines 3-fld., discolorous, yellow, sulcate, stout, 1-2.5cm. Leaves 2-5 cm. x 6 1 0 mm. ; oblanceolate; slenderly cuneate, subsessile; entire; acute, mucro- nate ; both sides distinctly veined, scarcely reticulate; below pale, grey, pruinose. Inflorescence 15-25-fld., loosely racemose, (4-) 6-8 cm. long, including peduncle 1-2 cm.

Page 108: Berberis and Mahonia : A taxonomic revision

108 L. W. A. ABRENDT Pedicelsglabrous, slender, 1-2-7 cm. ; bracts acuminate, 3-5 mm. Outer sepals 6 x 2.5 mm., oblong-ovate, acute. Inner s e p h oblong-elliptic, 8.5 x 4 mm. Pet#& 4.5 x 3 mm. ; apex rounded, shortly ernargimb; basal glands lanceolate, 1-2 x 0-4 mm. Stanzens 3 m., slightly produced. O w d a subsessile.

S.E. Tibet: Prov. Kongbo, valley of the Lilung Chu, west bank of river, 10,300- 10,600 ft., fl. 24 May 1938, Ludlow, S h e d & Taylor 4424 (Type, BM). 'A shrub of about 6 ft., with sepals and petals greenish yellow without, yellow, within; young twigs reddish brown; in cleasings in &ue.eus I& forest.'

Cultivated: 1938-9, at Wisley.

(132) Berberis lindleyana Ahrendt, J. Asiat. Soc. Beng. (Sci.), (3) 11, 3, 1945; syn. 'B. umbellata' hdl . , Bot. Reg., 28 Mhc. 41,1842; Bot. Reg. 30, t. 44,1844; non Wall. ex Don

Stems very slender; mature dark red. Leaves obovate; to 4 x 1.7 cm.; entire; lateral veins visible, not reticulate; below pale, grey-pruinw. I n , a c e n a 10-20-fld., racemose 5-7 cm., including peduncle 1-2-5 cm. Pediceh 5-11 mm.; bracts 2-2.5 mm. Outer 8epals 2-53x1-75-2mm., reddish, ovate. Inner sepals 6x4mm., obovate. Petals 5 x 3.5 mm., obovab, entire. Style absent.

Nepal: seed only, c. 1835. Cultivated from the Nepalese seed, in the garden of W. Web, of R d e a f : fl. c. 1843,

and observed and described (m B. umbellata) by Lindley.

(133) Berberis forskaliana Schneid., Bull. Herb. Boissier, (2) 5, 456 (1905) Stem glabrous; shoots k e l y sulcate; mature slightly angled, red-brown. Internodes

(6) 6-9 (-12) mm. Sp'na stout, 3-fld., sulcate, 1-1.5 cm., discolorous, yellow. Leaves 7-17 x 5-9 mm.; ovate; margins with 2-3 spinules (1-1.5 mm.)-serrulate, distant 5-8 mm. (but on the new shoots sometimes entire, orbicular, 1-2 cm. across excluding petiole 1-2 om.); above dull grey green; pale green with elevated reticulation; below papillose, a t h t pruinose and greyish, finally yellow-green and scarcely paler than above; thickly coriaceous. InfEorescenee, flowers and fruit unknown.

Arabia: Schibam above El Ejan, Schweinfurth 1682 (Type, K). This species is not in cultivation, and is, indeed difEicult to place owing to the lack of so

many important characters. Its geographical position suggests the Tinctoriae, and it may belong to thie subsection, where Schneider origimlly placed it, and where, owing to lack of any further material, I am compelled to leave it. The thickly coriaceous leaves suggest that it may belong to the more southern Subsect. Eutinctoriae of South India and Ceylon; only further knowledge, in particular of the nature of the style and inflorescence, can decide this.

Sect. hgulosae Schneid., Bull. Herb. Boissier, (2) 5, 396, 1905

Stem, spines, inkrnooks, diverse. h u e s nearly always deciduous, rarely persistent to semi-evergreen. InJrorescence 2-8 (-15)-fld., solitary, fascicled, umbellate, subumbellate, sometimes pseudumbellate-subracemose. Flowers mostly large, 1 - 6 2 cm. diameter with inner sepals 8-11 mm.; sometimes (6-) 10-12 mm. diameter, with inner sepals (6) 6-7 mm. Petals always shorter than inner sepals (except only in B. ambrozyam, where they are aa long). O& (3-) P 1 2 (-15). Berries mostly red, occasionally dark red, mostly epruinose, Sometimes pruinose white or blue : occasionally purple, and epruinose ; globose, 7-9 mm., or elongated, 7-17 mm. long, estylose or with very short style.

Afghaniatan, Sinkiang, Kashmir, Punjab, Jaunaar, Garhwal, Kumaon, Nepal, Sikkim, Bhutan, S.E. Tibet, Yunnan, Szechuan, Chihli, Siberia, Formosa.

Page 109: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 109

KEY 27. General Key to Sect. Angulosae (Maps 4 and 17) (N.B. Certain species of Sect. Tinctoriae, Subsect. Umbellab, and of Sect. Polyan-

thae, Subsect. Pseudopolyanthae, are sometimes in dried form, or from brief description, difKcult to separate with certainty from the Angulosae. Accordingly, indications to these have been included in the large key which follows. Further, this key contains indications to the hybrids, which are omitted from the keys to the sections.) Stems pubescent, or at any rate puberulous at first.

Leaves always, or ultimately, green, epruinose below. Leaves always grey, pruinose, below.

Ovules 6-15. Stems always glabrous.

Stems dark red. Stems pale yellow.

Mature stems dark red. Ovules 3-5.

Flowers solitary, or nearly always so. Flowers fascicled or subumbellate, 3-8 (-10).

Fruit stylose. Fruit estylose.

Fruit dark red or purple, or brown. Fruit pale or bright red.

Leaves mostly or always entire. Leaves mostly or always with spinom margins.

Mature stems pale yellow. Fruit very narrow (1 : %4.5), 3-4 mm. broad. Fruit broader (1 : 1.5-2), (6) 5-7 mm. broad.

Bracts 4 4 mm., subequal to or longer than the pedicels. Bracts 1-2 mm., much shorter than pedicels.

Leaves narrow, 1 : 5-6, revolute, enervate. Leaves broader, 1 : 1.5-3, plane, veined.

Leaves green, epruinose below. Leaves grey, pruinose below.

Fruit stylose. Fruit estylose.

A B

C D

E

F

G

H J

177. B. johannis

163. B. burmnica

167. B. lambertii

K

L M

A. Flower diameter 14-18 mm.; flowers solitary; pedicels pubescent; outer sepals as

Ovules 6-12; outer sepals much narrower than inner sepals; petals entire; fruit

Ovules 4; outer sepals as broad as inner sepals; petals emarginate; fruit stylose.

Flower diameter (&) 9-11 mm.; either flowers not solitary; or leaves with spinose

long as inner sepals; leaves entire.

estylose. 134. B. angulosa

135. B. parisepala

margin; and fruit estylose. Berries not globose, estylose, or with very short style.

Flowers solitary ; ovules 5-7 ; spines manifold. Flowers fascicled 2-6; ovules 4-5; spines normal.

Stamens not produced, truncate. 160. B. sibirica

Cf. 127. B. duthieana Stems subterete, or subangled; pedicels 5-9 mm. Stems very sulcate; pedicels 8-20 mm.

Leaves 6-15 mrn. Pedicels &lo mm. 169. B. hobsonii Pedicels 2-3 mm. Cf. 292. B. beesiana

Leaves 2-4 cm. Cf. 128. B. thornsoniana

Leaves below always concolorous green; pedicels pubescent. 164. B. lasioclema Leaves at fist pruinose grey below; pedicels glabrous. 173. B. jaeschkeana

Cf. 168. B. muerqolia

Stamens produced, more or less apiculate.

Berries globose, w;th conspicuous style (1-2 mm.).

Page 110: Berberis and Mahonia : A taxonomic revision

110 L. W. A. AHRENDT

KEY 27 (conlinued)

B. Ovules 6-10; flowers sofitary, 14 mm. diameter; fruit 12-18 mm. long; outer sepals

Stems purple; laves, and often petals, entire; stem very sulcate; pedicels

Stems yellow, subsuloate to subterete; leaves, and petals not entire; pedicel8

Ovules 4-5; flowers fascicled or umbellate (1-) 3-10; diameter 7-9 (-13) mm.;

as long as inner sepals.

glabrous, 2-5-6 cm. 136. B. ludlowii

1-2 cm. 137. B. macrosepala

outer sepals shorter than inner sepals. Flowers solitary.

Fruit globose, with conspicuous style. 167a. B. mucrifolia Fruit obovoid with very short style. 162. B.i%a&a

Leaves openly veined; pedicels (7-) 12-18 mm. Flowers not solitary.

Stems dark red; podicels 12-18 mm. 176. 3. trichohaenZato&S stem pale yellow; pediwls 6-12 mm. 172. B. jaechkeana

Leaves reticulate; pedicel8 2-6 mm. Fruit 8-12 mm. long.

Leaves spinom; petals emarginate; style short. Leaves and petals entire; style nil.

183a. B. x Emperor 185. B. x Cherry Ripe

Fruit P 6 mm. Cf. 293. B. wikonae

C. Leaves m o w (1 : 4) ; outer sepals lanceolate, rtcuminate ; fruit attenuate-stylose.

Leaves broader (1 : 2-2.5) ; outer sepals obovate, elliptic, or ovate, rounded or obtuse, 147. B.numisone&

rarely acute. Leaves reticulate; fruit attenuate-stylose.

Flowers fascicled 2-4; stamens apiculate; leaves 8bOVe @;reen, below grey. 146. B. mmduna

143. B.temolaica: Flowers solitary (-2) ; &=ens trunwte ; 1 - v ~ above bluish, below white.

Leaves not reticulate; stamens not apiculate. Leaf margins with 2-7 (-15)-sphulea; outer sepk shorter than inner.

Fruit stylose. Fruit pruinose, ovoid-attenuate, with thick bent style; flowers solitary.

Fruit epruinose, oblong-elliptic, with short thin style; inflorescence 4-8-fld. Cf. 143. B. tentolaica

183. B . dczrecta Fruit estylose.

Leavea pruinose white below; margins with 2-6 spinose teeth.

Leaves below ultimately epruinose, green; margins with 6-15 spinose teeth. 140. B.concinna

139. B. ergthroclada Cf. 136b. B. ZucUOwii vm. capillaris

144. B. circumserrata

Leaves entire; outer sepals as long as inner.

D. Leaves broad (1 : 1.5); margins with 15-40 spinules. Leaves narrower (1 : 2-4); margins with 1-12 spinules or entire.

Fruit ovoid-attenuate, with thick bent style. Leaves mostly 16-40 mm., deciduous, reticulate; pedicels 12-20 mm.; stamens

Leaves mostly 6-15 mm., semi-evergreen, not reticulate; pedicels 5-9 mm.; truncate; internodes 1 5 3 0 mm. 145. B. diaphana

stamens apiculate; internodes 5-15 mm. 142. B. kumaonertffis Fruit oblong or elliptic, estylose.

Flowers solitary; leaves white below; stamens truncate; o d e s 12-15.

Flowers 3-11-fld.; leaves grey or green below; stamens apiculate; ovulea 6-9. 141. B. tsangpoelzsis

Internodes 5-1Omm.; stems subterete; leaves m o w , (1 : a), openly veined; pedicels 5-9 mm. 165. B. apiculata

Page 111: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 111 Internodes 18-35 mm.; stenzs sulcate; leaves broader (1 : 1.5-2.5, -3);

reticulate; pedicels 10-25 mm. Leaves below, green, epruinoses spines absent or to 8 mm. long.

Leaves below, grey, pruinose; spines normal, 10-25 mm. long. 166. B. zayulana

Leaves entire. 151. B. faberi Leaves with 6-18 marginal teeth. 173. B. x vi lmrini i

E. Fruit estylose, 15-18 mm. 156. B. rnuliensis Fruit stylose, 6-10 (-12) mm.

Leaves closely reticulate. Leaves with spinose margins. 152. B. approximata Leaves entire.

Stems and undersurface of leaves epruinose. Spines absent or weak, 5-10 mm.; petals as long as inner sepals.

Spines strong, 1-3 cm. ; petals shorter than inner sepals. 154. B. ambrozyana

153 b. B. dictyophylla var. epruinosa Stems and undersurface of leaves pruinose, white. 153. B. dictyophylla

Leaves scarcely veined. 155. B. stiebritzyana

Leaves below epruinose, green. cf. 179e. B. orthobotrys var. rubicunda Leaves below, pruinose, grey.

Fruit red, ovoid-attenuate; style bent. 149. B. virescens Fruit purple, oblong; style very short, straight. 189. B. shensiana

Cf. 293. B. wilsonae, varieties

G. Flowers 9-10 mm. diameter; fruit 9 x 10 mm., purple-black. 188. B. paravirescens

Flowers subumbellate; pedicels 6 8 mm. ; stamens apiculate; fruit purple,

Flowers fascicled; pedicels 1-3 cm.; stamens truncate; fruit dark red-brown,

F. Leaves deciduous, openly veined; fruit 8-10 mm., oblong or ovoid.

Leaves semi-evergreen, closely reticulate ; fruit 4-5 mm., globose.

Flowers 15 111111. diameter; fruit 11 x 9 mm.

1 1 X 9 m m . 184b. B. rubrostilla x var. chealii

l l x 6 m m . Cf. 124c. B. campbellii; 126. B. walterana

broad. 177. B. johannis H. Flower diameter, 7-8 mm.; stamens truncate; fruit bent, narrow, S 3 . 5 (-4.5) 111111.

Flower diameter 12-16 mm.; stamens apiculate; fruit 6-7-5 mm. broad. Leaves broad (1-1.5 (-2), openly veined, below grey, pruinose.

Stems sulcate ; leaves sessile ; inflorescence 15-25 mm. Stems terete ; leaves petiolate ; inflorescence 2-4 cm.

Leaves scarcely reticulate above; pedicels 12-18 mm.; flowers 14-16 mm.

182. B. platyphylla cf. 125. B. brandishxna

Leaves 1 : 2.5, reticulate, below white, pruinose.

diameter, with 3 series of sepals; stems and pedicels epruinose. 157. B. valdisepala

Leaves closely reticulate above; pedicels 5-10 mm. ; flowers 12 mm. diameter, with 2 series of sepals; stems and pedicels pruinose, white.

158. B. calcipratorum

Fruit globose, 7 mm., 6rm, lustrous. 178. B. cooperi Fruit oblong, 9-10 x 5-6 mm., soft, dull. 179. B. orthobotrys

Leaves narrow (1 : 4); pedicels P 7 mm.; stamens truncate; fruit 15-17 x 5-9 mm.

Leaves broader (1 : 2-3); pedicels 7-12mm.; fruit 9-12 x 5-8mm; stamens apiculate.

J. Leaves green, epruinose below.

Leaves grey, pruinose below.

184. B. rubrostilla

Stamens apiculate. Leaves reticulate. 179f. B. orthobotrys var. canescens Leaves scarcely veined. 181. B. ignoratu

Stamens not produced, truncate. 180. B. ambigua

Page 112: Berberis and Mahonia : A taxonomic revision

112 L. W. A. AHBBINDT

KEY 27 (continued) I(. Leavesentire.

Fruit eetylose; staJnenS trwlate. Internodes 5-10mm.; leaves 6-12x2-4mm., openly veined; flowers s o l i t q

14 mm. 138. B. everestiana

10 mm. diameter. 170. B. kartanica Fruit stylose. 150. B. ellwtii

Leaf margins with 1-2 spinules; 8-15mm. long; inflomnca 3-6-fld.; fruit

Internodes 12-20 mm.; leaves 12-30 x 6-12 mm., reticulate; flowere 5-7-fld.,

Leavee spinose.

10-12 mm. Cf. 173 b. B. jaeschkearra var. usteriana

8-9 mm. 161 c. B. x provincialis Leaf margins with 5-15 spinules; 15-25mm.; infl0-m 6-12-fld.; fruit

L. Fruit attenuate-ovoid, 10-12 x 5-7 mm.; style thick, bent. Leaf margins with 10-40 apinules; flower b e t a r 10-14 mm.; inner sepals

Leaf margins with 3-7 apinules; flower diameter 9-10 mm.; inner sepals narrow. broad. 144a, b. B. cirzcmserrata varieties

148. B. tiechleri Fruit oblong-elliptic; style straight, thin.

Fruit dark red to purple; leaves with 5-15 marginal apinules. Flower diameter 10-12 mm.; pedicels 7-13 mm. Flower diameter 8 mm.; pdicels 3-8 mm.

Fruit bright red; leaves with 0-4 marginal spinulea.

Flower diameter 6-6 mm.; pedicels 18-25 mm.; fruit 12 x 6 mm., ovoid.

Flower diameter 13-15 mm.; pedicels 5-9 mm.; fruit 9 x 6 mm., obovoid.

187. B. fmoniana 161. B. emurginuta

173. B. j a e s c h h m varieties

M. Leaves entire.

171. B. hinzakcica

172. B. hcrmiltoniana Leaves spinose.

Leaves with 1-12 marginal spinules. Petals shorter than inner sepals.

Fruit bright red; leaves 8-16 mm. Fruit dark red, or purple; leaves 2-5 cm.

172b. B. jaeschkeanu var. wrtericcna

174,. B. x vanfleetii 186. B. consirnilis

Cf. 124. B. umbellata

Flower diameter 6-7 mm.; fruit dark red. Flower diameter 12-14 mm.; fruit purple.

Petals as long aa, or longer than, inner sepals. Leaves with 15-25 marginal spindee.

Fruit dark red-brown, 10-12 111111. Fruit bright red, 7-8 mm.

Cf. 124. B. umbellatcl 175. B. x regleriana

Subsect. Euangulosae Schneid., Bull. Herb. B0is&r, (Z), 5, 396, 1905 S t e m mostly markedly sulcate. Leaves deciduous (except only in B . m i n m ,

B. tsangpoensis and B. kumaonensis, where they are semi-evergreen). F h e r s large (12-) 1 6 2 0 (-25) mm. diameter, and solitary (except only in rare varieties of B. angulosa, B . 1uIuOwii var. cupihrh where they are few-fascicled). Outer sepals often as long as inner. Inner sepals (7-) 8-12 (-14) mm. long. Petals 5-8 mm., always much shorter than inner sepals. Stamens mostly not, or but slightly, produced, more or less truncate (except only in B. kumamenah where they are apiculate). Ovules 6-12 (except only in B. pari- sepala and B . everestiana, where they are 445, and in B. tsangpoenais where they are 12-15). Berries bright red, varying from globose-ellipsoid, 10-12 x 9-12 mm. (in B. angulosa and B. parisepala), to broadly oblong, 15-18 x 10-12 mm. (in B. mucrosepala), or narrowly oblong, 13-16 x 6-8 mm. (in B . concinna), estylose, or very shortly stylose; the only exceptions being B . kumaonensis and B. tenzolaica, with berries ovoid with bent stylose apex.

Kumaon, Nepal, Sikkim, Bhutan, Tibet, Burma.

Page 113: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 113

KEY 28, to Subsection Euangulosae (Maps 17 and 18) Pedicels and stems pubescent ; fruit globose-ellipsoid; stamens not produced, truncate. Ovules 6-10; outer sepals much narrower than inner sepals; petals entire; pedicels

Ovules 4; outer sepals as broad as inner sepals; petals emarginate; pedicels 5-12 mm. ; 12-18 mm.; fruit estylose. 134. B. angdoaa

fruit shortly stylose. 135. B. parisepala Pedicels glabrous; stems pubescent or glabrous; fruit narrower.

Stems pubescent, a t any rate at first; leaves grey, pruinose, below; stamens produced, truncate ; fruit oblong.

Stems dark, purple, very sulcate; leaves entire. 136. B. IwEbwii Stems pale, yellow, slightly sulcate to subterete; leaves spinose. 137. B. macrosepala

Fruit oblong, estylose. Stems always glabrous.

Leaves green, epruinose, below; ovules (P) 6-9. Stems pale, yellow; pedicels 5-9 mm.; leaves entire; ovules 4-5.

Stems dark, red; pedicels 15-25 mm. ; leaves spinose; ovules 6-9. 138. B. evermtianu

139. B. erythroclada Leaves grey or white, pruinose, below; ovules 6-15.

Stems dark, red or purple; leaves openly veined; ovules 6-9. Outer sepals as long as inner sepals; leaves entire.

Outer sepals only half as long as inner; leaves spinom. Cf. 136 b. B. ludbwii var. eapilluris

140. B. concinna

141. B. tsangpoensis Stems pale, yellow; leaves reticulate; ovules 12-15; habit prostrate.

Fruit ovoid, with bent stylose apex. Stems slender, sulcate, yellow; leaves &20mm. long, with spinose margins;

pedicels 5-9 mm.; fl. diameter c. 2 cm., with narrowly obovate inner sepals, 12 mm. long; stamem apiculate; fruit 8-10 mm. long.

142. B. kumaonensis Stems stout, terete, purple; leaves 20-45 mm. long, entire; pedicels 10-15 mm.;

flowers c. 14 mm. diameter, with orbicuh inner sepals c. 8 mm. ; stamens truncate ; fruit 11-14 mm. 143. B. temolaica

(134) Berberis angulosa Wall. ex Hook. f. et Thorns., Fl. Ind. 1,227, 1855; Bot. Mag. 115, t. 7071, 1889

A rigid, stiff, stout stemmed shrub, about 3 4 ft. high. B t e m conspicuously hairy and dark brown. Spines slender (1-) 3 (-5)-fld., often puberulous below, 4-16 mm. Leaves 10-40 x 6-15 mm., obovate, base cuneate, margin entire, apex rounded; above lustrous deep yellow-green ; below concolorous, lustrous, epapillose. Pedicels 12-25 mm. Outer sepals 8-10 x 4 4 mm. Inner sepals 8-10 x 7-9 mm. Petals paler, 7-8 x 5-6 mm., apex rounded, entire; basal glands separate, oblong, 1.2 x 0.6 mm. Berries fim, red, very lustrous, ellipsoid-subglobose, 10-12 x 9-12 mm. Style absent or scarcely distinct.

KEY 28a

Flowers solitary. Flowers fascicled.

134a. B. angulosa var. angulosa 134b. B. anguloaa var. fasciculuta

(1344 Berberis angulosa var. angulosa Sikkim: Yeumting, 6 Sept. 1848, Hooker & Thomson, s.n. (K). Lachen, 11,000-

12,OOOft., 14 July 1849, Hooker & Thomson, s.n. (K). Darjeeling, Subuikuiaci, Oct. 1875, Gamble 90 (K).

Nepal: Wallich 1475, in part (Type, K). Central Chilime Kharka, in Juniper scrub, 14,5OOft., July 1949, Polunin 1269 (BM); between Namche Bazaar and Tesingar,

8 JOURN. LIKN. S0C.-BOTANY, VOL. LVII

Page 114: Berberis and Mahonia : A taxonomic revision

114 L. W. A. AHRENDT 3-10 ft., 11,350 ft., 13 May 1952, Zimmermann 494 (Geneva); between Chuli et Thiimi, 13,300 ft., 11 O d . 1954, Zimmermann 1619 (Geneva).

(134b) Berberis angulosa var. fasdculata Ahrendt, J. Bot., Led. 79 (Suppl.), 42, 1941; syn. B. a q h a var. of Hook. f. et Thorns., 1.c.

Sikkim: 10,700 ft., Hooker & Thomson (Type, K). S.E. Tibet: Mago district, Mago Chu, open places in forest, and among rocks in river

Cultivated: fl. 14 May 1940; fr. 20 Oct. 1938. It is valuable for its large, very shining, bed, 11,000-12,OOO ft., 9 June 1935, Kingdon-Ward 11653, part (BM).

bright vermilion berries.

(135) Berberis parisepala Ahrendt, Card. Chron. (3) 109, 100, 1941; J . Bot., Lond.

Stems and leaves as in B. angulosa. Spines stout. Fl4nuer.s solitary. Pedicela 5-9 (-13) mm. Prophylls 1-25 x 0-75 mm., yellow, ovate, acute. Sepals in three series, outer, median, inner, all 8-9 x 6-5-7-5 mm. Petals 7.5 x 4 mm., acutely emarginate. Ovules 4. Berries 10 x 8 mm., with &tinct short style.

S.E. Tibet: Prov. Zayul, Assam-Tibet frontier, &hmi hills, &lei valley, 12,000 ft., 21 June 1928, Kingdon-Ward 8350, seed number only.

Bhutan: in many places, 11,OOo-13,000 ft., May to Oct. 1949, Ludlow, Sherriff & Hicks numbers 16328, 18975, 19058,19683, 16611, 16413,21269, 17441, 17453,20703 (BM).

Nepal: (east to central), east of Dhorpatan, 5 ft., on open slope at 11,500 ft., 21 Oct. 1954, Stainton, Sykes & Williams 9109 (BM); Annapura Himal Seti Khola, 3 4 ft., at 12,500 ft., 3 Aug. 1954, Stainton, Sykes & Williams 6656 (BM); Samargaon (north of Tukucha), Kali Gmdaki, 5ft., amongst shrubs on steep hillside, 5ft., a t 13,000ft., 10 Oct. 1954, Stainton, Sykes & Williams 8099 (BM); Muktinath, 4 ft., open rocky slopes at 14,000 ft., 1 Oct. 1954, Stainton, Sykes & Williams 8062 (BM); Rambrong, Lamjung Himal, 2-3 ft., at 13,000 ft., fl. 3 July 1954, Stainton, Sykes & Williams 6080 (BM), and 3 4 ft., at 12,000 ft.; fr. 29 June 1954, Stainton, Sykes & Williams 6047 (BM); south of Gurjakhani, 8 June 1954, Stainton, Sykes & Williams 3062 (BM).

The Bhutan and Nepal specimens probably contain a number of variations. No. 3062 haa the charactektic equal sepals, but the stems have become nearly glabrous; 6047, 6080, possess 4-5 ovules, but have reddish stems; 8099 and 9109, also have 4-5 ovules and almost glabrous yellow-brown mature stems, the former short (4-7 mm.) glabrous pedicels, the latter long (1-5-2cm.) puberulous pedicels, and in both an extremely short style, which seems lacking in the other h i t i n g specimens; 6656 has 5-7 ovules, short pedicels and the persistently pubescent stems.

Cultivated (from seed of KW 8350): fl. 28 June 1939; fr. 5 Oct. 1939 (Type, BM). The herbarium specimen of 8350 represents a quite difFerent species. B. parisepala is

very similm in appearance. to B. angulosa, from which it differs in the botanical characters cited, and its greater height (4-10 ft.).

79 (Suppl.), 42, 1941; Sealy, Bot. Mag., 147, N.S., t. 119, 1950

(136) Berberis ludlowii Ahrendt, J. Bot., Lond., 79 (Suppl.), 43, 1941. (Fig. 27) A tall shrub, 6-10 ft. Stems pubescent, with reddish shoots, when mature, dark purple,

sulcate, with internodes 3-8 cm. Spines absent or weak and solitary towards apex of stems, but 3-fld., 3-9mm., below, though occasionally stouter and to 18mm. Leaves oblong-obovate, with cuneate, sessile base, entire margin and rounded apex; above dull grey-green, below grey-pruinose; both sides with loosely branched vcnation, 20-40 x 9-19 mm. Flowers solitary. Pedicels glabrous, 2-5 cm. Outer sepals 9 x 4.5 nun. Inner sepals 9.5 x 5.5 mm. Petals 6 x 4 mm., subentire; their basal glands separate, narrowly oblong-ovate, subacute, 1-5 x 4 mm. B e d dull red, estylose, 15-18 x 6-9 mm.

Page 115: Berberis and Mahonia : A taxonomic revision

Berberia and Mahonia 115 KEY 28b Leaves entire; pedicels 2-5 cm. Mature stems pubescent; shrub 6-12 ft. ; berries narrow, 15-18 x 6-9 mm., epruinose.

Mature stems glabrous; shrub 1-3 ft. ; berries broader, or shorter, often with blue bloom. Outer sepals as long as, but narrower than, inner sepals; petals entire; fruit 15-18 mm.

Oiiter sepals shorter than, but proportionately as broad as, inner sepals; pet& emar-

136a. B. ludlowii var. ludlowii

long. 136 b. B. ludlowii var. capillaria

ginate; fruit 10-14 mm. long. 136c. B. ludlowii var. deleica 136d. B. ludlowii var. a&liwicOla Leaves spinose; pedicels 6-10 mm.

(136~) Berberis ludlowii var. ludlowii S..E. Tibet, Tsari district, Singo Samba, fr. 10 Oct. 1938, shrub 8-12 ft., Ludlow, Sherriff

& Taylor 6589 (Type, BM). Also 11,000-13,OOOft., under numbers 3851, 6623, 6337, 1626. This came into cultivation through number 6337.

(136 b) Berberis ludlowii var. capillaris, stat.nov. ; syn. B. cupillaris Cox ex Ahrendt, J . Bot., Lond. 79 (Suppl.), 47,1941 ; Cox, nomen nudum, in Farrer’s Last Journey, 121, 1926

Upper Burma: Hpimaw, Farrer 1399, seed number. N.W. Yunnan: Atuntze, Doker La, 12,500 ft., 11 Nov. 1937, Y u 7916 (E). Mekong-

Cultivated: (from seed of Farrer 1399) fr. Aug. 1939; and (from seed of Yu 7916), Salwin divide, Sila, 11,700 ft., 10 Oct. 1936, Yu 22790 (E).

fl. 16 Apr. 1945.

(136~) Berberis ludlowii var. deleica, comb.nov.; syn. B. macrosepala var. dekica Ahrendt, J. Bot., Lond., 79 (Suppl.), 46, 1941

S.E. Tibet: Prov. Zayul, Assam border, Mishmi hills, Delei valley, Dri La, 12,000 ft.,

Upper Burma: Nam Tamai valley, 11,OOO ft., 4 Oct. 1937, Kingdon-Ward 13364. Also

N.W. Yunnan: Mekong-Salween divide, Sewalongba, 12,650 ft., Yu 22170 (E). Cultivated: (from KW 11044), fl. May 1939; and (from KW 13364), fl. 3 May 1944. The typical tall var. ludlourii grows in Abies, Rhododendron and mixed forest, generally

by river banks. The variety capillaris has stems only slightly puberulous or often quite glabrous at first, and always quite glabrous very soon. The sepals are all 8-10 mm. long, the outer 3 4 mm., and the median and inner 5-6 mm. broad. The variety deleicu has outer sepals 7 x 5 mm., and median and inner 8-9 x 6-7 mm., of colour H.C.C. 1. These two low growing varieties come from forest clearings, amongst scrub, or grassy mountain slopes. Related is Ludlow, Sherriff & Elliott 15758, from S.E. Tibet. Prov. Kongbo, Chachima, Lotu Chu, at 12,500 ft., on 22 Sept. 1947. This has discolorous yellow spines, flowers in subfascicled groups of 1-5 with short (8-12 mm.) pedicels, and short style.

(1364 Berberis ludlowii var. saxiclivicola stat.nov. ; Ahrendt, l.c., 43, 1941

pedicels, and petals slightly incised.

SherrifF & Taylor 3968 (Type, BM). Common on open stony hillsides.

12 Dec. 1938, Kingdon-Ward, 11044, (Type, BM), Sept. 1931, Kingdon-Ward 10006.

in 1925, Forrest 26945, 27519.

This differs in its leaves with 2-6 marginal spinules (1-1-5 mm.), and short (6-10 mm.)

S.E. Tibet: Tsari district, Langong, Chubumbu La, 13,000 ft., 6 June 1938, Ludlow,

(137) Berberis macrosepala Hook. f. et Thorns., F1. Ind. 228, 1855 A spreading shrub up to 6 ft. Stems with young shoots green, puberulous and often

somewhat sulcate ; mature glabrous or subglabrous, subterete, yellow-brown, sparsely 8-2

Page 116: Berberis and Mahonia : A taxonomic revision

116 L. W. A. ~ E N D T

black-verruculose. Intern& 1-3 cm. Spines 1-2 cm., 3-fld., plane subconcolorous. Leaves 20-45 x 10-20 mm.; obovate; base contracted to decurrent petiole, 2-7 mm. ; margins with (2-) 4-8 (-12) spinose (1-5-2 mm.) serrations (1 mm.), (2-) 4-7 mm. apart; above slightly lustrous deep yellow-green; below grey, pruinose. Plowers solitary. Outer and inner s e p b both 8-5-10 x 5-6 mm. Petals 6-7 x 3.5-4-5 mm. ; obovate; apex emarginate with two acute lobea; base not clawed, with separate lmceolate glands 1.5 x 0.3 mm. Stanzens 4-4.5 nun., produced, rounded-truncate. Bepries oblong, 12-16 x 7-10 mm.

KEY 28c Stems subangled or terete; leavea with apices rounded.

Stems sulcate; leavea with apices acute. 137a. B. macrosepala var. macrosepala

Leavea with 2-7 spinulea to each margin; pedicels 8-15 mm.

Leaves with 8-14-spinulea to each margin; pedicels 26-30 mm. 137 b. B. nzacrosepala var. sakdenen.m:e

137c. B. macrosepala var. setqolia

(137a) Berberis macrosepala var. macrosepala Sikkim: Hook., s.n. (Type, K). Assam Himalaya: 10,000-14,000 ft., Kingdon-Ward 13713, 11562. Bhutan: 12,500-14,500ft., 1934, Ludlow & Sherriff603; 1949, Ludlow, S h e d & Hicks

16856,21461. S.E. Tibet: Tsari and Pachakshiri districts and Prov. Kongbo, a t 10,500-13,500 ft.,

1936, Ludlow & Sherriff 1652, 2647; 1938, Ludlow, S h e d & Taylor 4819. Cultivated from seed of LS 2647. In the Mago district, a t 11,000-12,000 ft., Kingdon-

Ward 11563, in part, appears aa a form with extremely large flowers, having outer sepals 7-9 x 46-56 mm. ; median sepals, 9-5-11 x 6 mm. ; inner sepals 11-14 x 5-56 mm., and petals 6.5 x 4.25 mm. (BM).

(1374 Berberis macrosepala var. sakdenensis (Ahrendt), Ahrendt, comb.nov. ; syn. B. ludlowii var. sakdenemis Ahrendt, J. Bot., Lond., 79 (Suppl.), 44, 1941

Bhutan: Sakden, amongst rhododendron scrub at Nyuksang La, 12,500ft., 6 July 1934, Ludlow & Sherriff 604 (Type, BM). Also, from N.E. and Central Bhutan, 13,000- 14,000 ft., 1949, Ludlow, S h e M & Hicks 20337, 16650, 17287. A shrub of 2-3 ft., not in cultivation.

(137 c) Berberis macrosepala var. setifolia Ahrendt, I.c., 46, 1941. Leaves ovate, acute, margins with 8-14-spinose (2 mm.) serrations (1 mm.), only

2 mm. apart. PedicelS 2-3 cm. Ovules 9-10. Berries 13 x 7 mm. Bhutan (N.E.): Me La, 14,5OOft., 5 Oct. 1934, Ludlow & Sherri€f 1025 (Type, BM).

Open rocky hillsides above tree-line. All plants cultivated from this number have borne a many-flowered racemose inflorescence, which places them in a different section (Vul- gares), characterized by having small flowers with only 1-2 o d e s .

(138) Berberis everestiana Ahrendt, sp.nov., ramulis glabris flavis, foliis subtus viridibus, floribus fructibusque satis parvis solitariis, OW 4-5, distinguenda

Stem glabrous; mature, sulcate yellow. I n t e d 6-10 (-20) mm. Spines 8-12 mm., subterete with shorter laterd parts, 1-3-5-fld. Leaves to 12 x 4 mm., obovate, mostly entire, mucronate ; below paler yellow-green. Flowers solitary. Prophylls lanceolate, 4 x 1 mm. Outer sep& 6-7 x 44.5 mm. Inner s e p d 7-8 x 5-6 mm. Petals 6-65 x 3- 3.5 mm., entire. Stanzens 3 mm., scmcely produced. Ovules 3 4 . Bewies (7-) 9-10 x (6) 5-5-6 mm., ovoid, estylose, or with style scarcely distinct. Seeds purple.

Page 117: Berberis and Mahonia : A taxonomic revision

Berberis und Mahonia 117

KEY 28d Leaves always entire; stamens truncate; fruit 9-10 mm. ; seeds purple.

Shrub 6-9 in. high; internodes mostly 5-10 mm.; leaves very slightly veined.

Shrub 2-4 ft. high; internodes 10-20 mm.; leaves distinctly veined or subreticulate. 138a. B. everestiana var. everestianu

138 b. B. everestiana var. numbuensis

138c. B. everestiana var. ventosri Leaves sometimes epinose; stamens apiculate; fruit 7 mm.; seeds yellow-brown.

(1384 Berberis everestiana var. everestiana. (Fig. 28) South Tibet: Prov. U, Mt. Everest expedition, stony soil north of Kharta Chu, fl.

11 June 1922, no. 77 (Type, K). Rongsha valley, 12,000ft., common in the beds and sides of the gorge, fl. 19 June 1924, Hingston 4 (K), and 13,000 ft., in the stony bed of the gorge, Hingston 95 (K); Kharta, 12,0OOft., fl. 11 June 1922, Norton 90 (K). Above Singma, Klangchung, 11,500 ft., fr. 21 Aug. 1936, Chapman, s.n. (K).

8.E. Tibet: Prov. Pome, Nyima La, 14,000-15,000ft., fl. 20 June 1924, Kingdon- Ward 5811 (K).

Nepal: Chilang Pati, Bailey 46, 25 Oct. 1935 (BM). This dwarf or subprostrate shrub is described by Kingdon-Ward as ‘a matted under-

shrub rising only 6-9 in. above the ground on the open hillside with dwarf rhododendron, Lonicera and Potentilla fruticosa. It is not in cultivation, but may be compared with B. tsangpoensis, also subprostrate, but having reticulate leaves, white below ; ovules 12-15; and much larger berries, with much longer pedicels.

(138b) Berberis everestiana var. nambuensis Ahrendt, var.nov., elatior, internodiis longioribus, foliis distincte venatis

Leaves to 27 x 9 mm. Pedicels stout. Outer sepals 6.5 x 3 mm. Inner sepals 7.5 x 4 mm. Petuls 5 x 2.5 mm. Stamens 3-5 mm.

S.E. Tibet: Prov. Pome, Nambu La, 14,5OOft., 7 July 1947, dry ground; Ludlow, Sherriff & Elliott 15385 (Type, BM), also, same loc., on 12 June 1947, 13,884 (BM). Not in cultivation.

(138c) Berberis everestiana var. ventosa Ahrendt, var.nov., a typo foliis interdum dentatis, staminibus apiculatis, fructibus brevioribus, seminibus flavo-brunneis distinguenda

Stem very sulcate; shoots reddish; mature pale yellow. Internodes 5-12 mm. Spines 3 (-p>)-fld., 5-12 mm. Leaves 5-15 x 3-8 mm., entire, or with margins with 2-3 spinose (1 mm.)-serrations (0.5-1 mm.), distant 2 4 mm. ; venation branched; below slightly greyish a t &st, ultimately lustrous paler green. Plowers solitary. Pedicels 5-8 mm. Outer sepals 6 x 3-3.5 mm., ovate-elliptic, subacute. Inner sepals 7-76 x 4 4 . 5 mm., obovate. Petuls 5 x 3 mm. Stamens 4 mm., produced, apiculate. Berries 7 x 4 mm., oblong-ovoid, estylose, or with an extremely short, thick, scarcely distinct style.

Nepal: between Pudamigaon and Ringmigaon, 14,000 ft., 22 Sept. 1952, Polunin, Sykes & Williams 3554 (Type, BM). Phoksumdo Tal, 12,500 ft., 11 June 1952, Polunin, Sykes & Williams 2202 (BM). Barbung Khola, Pemriggaon, 15,000ft., 9 June 1952, Polunin, Sykes & Williams 1104 (BM).

Cultivated: seed was collected of no. 3554, so the variety may ultimately appear in cultivation. It forms a bush 1-3 ft. high growing on dry, stony, exposed slopes and cliffs, and is a frequent component of open scrub a t the higher altitudes.

Page 118: Berberis and Mahonia : A taxonomic revision

118 L. W. A. AERENDT

(139) Berberis erythrodada Ahrendt, J. Bot., Lond., 79 (Suppl.), 49, 1941 Stems stout, glabrous, sulcate; mature lustrous dark red. l a t e 4 8-11 (-25) mm.

h'phe.~ (3-) 5-7 (-9)-fld., orange red, 5-10 mm. Leavee 10-2Ox 3-7 nun., or 12-28 x 8-15 mm.; obovate; base cuneate, s u b d e ; margins with 5-15 spinose (1-2) mm. semtiom (0.5-1 mm.), spreading, 1-2 (4) mm. apart; apex rounded to obtuse; vena- tion clearly branched or openly reticulate; below, at first slightly greyish pnrinose, dtimately pale dull green. Fbwer8 solitary. Pedicels very slender. Outer sepals oblong- elliptic, 6.5-6-5 x 3-3-75 mm. (or as large as inner). Inner se& 7-8 x 4.5-5 mm. Petah 5-64 x 2-6-44 mm., subenthe; glands 0.8 x 0.2 mm., narrowly oblong, sepwate. Stanaens 3 4 mm., scarcely produced, subretuse. 0vuk.a 6-9, often stipitate. Fmit un- known.

KEY 2 8 ~ Internodes mostly 8-1 1 mm. ; shrub to 1 ft. ; outer sepals shorter than inner.

Internodes 10-26 mm.; shrub 4-6 ft.; outer sepals BB long as h e r . 139a. B. eryt?&ro&da var. esythrocladar

139 b. B. esythroclada var. t d u n g e d

(1394) Berberis erythroclada var. erythroclada S.E. Tibet: Prov. Zayul, Assam-Tibet border, Chutang camp, Atakmg La, 13,000 ft.,

6 Aug. 1933, Kingdon-Ward 10562 (Type, BM): Kase, Delei valley, 1928, Kingdon- Ward 8427 (K).

Prov. Pemako, Doshong La, 12,000 ft., 1924, Kingdon-Wad 6233 (K). This species is not in cultivation. The above type is described aa a small bush or under-

shrub, growing on south facing turf slopes; the flowers pale yellow, and the berries coral pink.

(139b) Berberis erythroclada vw. trulungensis Ahrendt, var.nov., interno&

S.E. Tibet: Prov. Pome, above Trulung, 12,500 ft., 26 June 1947, Ludlow, She& &

longioribus, sepalis exbrioribus interioribusque aequalibus

Elliot 1319 (Type, BM).

(140) Berberis concinna Hook. f. et Thorns., Bot. Mag. 79 t. 4744, 1853. (Fig. 30) Stems glabrous, stout, very sulcate; shoots green; mature lustrous dark red, not or

scarcely verruculose. Internodes 1-2 cm. Spines orange, sulcate, 3-fld., 1-26 cm. Leaves semi-evergreen, 10-30 x 5-14 mm. ; oblong-obovate; base cuneate to decurrent petiole, 1-5 mm. long; margins with 3-5 spinose (1-2 nun.)-serrations (1.5 mm.), 3-6 mm. apart; apex rounded, mucronulate; somewhat lustrous deep green with few lateral vehs; below white, pruinose, densely finely papillose. Pediceh (5 mm.-) 1-5-2 cm. Prophylls 2-24 x 1-5-2 mm., red, oblong-triangular, acuminate. Outer sepals 4-6 x 3 4 mm., oblong-ovate, subacute. Median and inner sepah equal, 4-10 x 6-7 mm., obovate. Petals 6-5 x 45 mm., obovate, apex acutely emarginate with 2 rounded lobes; basal glands concoloous, C. 1 x 0.7 mm. Stamens 4-46 mm., not produced, truncate. Ovules 6-8. Be& 13-16 x 6-8 mm., oblong, dull red.

KEY 28r Flowers solitary.

Pedicels 15-20 mm.; fruit 13-16 x 6-8 mm. Pedicels 5-10 mm.; fruit 10-11 x 7-8 mm.

Flowers umbellate.

140a. B. concinna var. concinnu 140b. 33. wncinna var. bre&or

140~. B. concinnu var. atsm@ora

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Berberis and Mahonia 119

(1404 Berberis concinna var. concinna Sikkim: Lachen valley, 12,000-13,000ft., fl. 31 July 1849, Hooker, s.n. (Type, K).

Tonglo, 10,000 ft., Nov. 1874, Gamble 1029 (K). Nepal: (east to central); B h m g d i , Khola, 1-2 ft., at 9500 ft., in flower bud, 15 June

1954, Stainton, Sykes & Williams 5811 (BM); Lete, Kali, Gandaki valley, l-zft., a t 11,500 ft., flowers more than 2 in. diameter, 4 June 1954, Stainton, Sykes & Williams 5595 (BM) ; above Ranmagaon, 12,000 ft., flowers lemon yellow, 2 July 1954, Stainton, Sykes & Williams 3352 (BM) ; west of Lumsum, 3 ft., among shrubs upon ridge, 11,000 ft., 22 Oct. 1954, Stainton, Sykes & Williams 9112 (BM). The last specimen possesses bracts (1-16 mm.) on the pedicel, about two-thirds up from the base.

Cultivated: fl. 2 June 1934,4 June 1942: and fr. 4 Oct. 1934,7 Oct. 1943. Apparently very rare in the wild state, this has been long in cultivation and forms a compact bush some 1-2 (-3) ft. high. There is a form growing 5-7 ft. high, namely Ludlow & SherriE 1086; from S.E. Tibet, Prov. Monyul, Milakatong La (Tawang), 12,000ft., 22 Oct. 1934 (BM). This has been in cultivation at Wisley under this number.

(140b) Berberis concinna var. brevior Ahrendt, J . A d . SOC. Beng. (Sci.), (3) 11, 3, 1945

Nepal: Namlang, 9OOO ft., 28 Oct. 1931, Sharma E 269 (Type, BM); (central), 5 miles east' of Timure, 13,5OOft., 3 July 1949, Polunin 782 (BM); Langtang, 12,500ft., 21 June 1949, Polunin 482 (BM); Rambrong, Lamjung Himal, 1-2 ft., at 12,000 ft., 26 June 1954, Stainton, Sykes & Williams 6046 (BM); Rambrong, Lamjung Himal, in rock crevices a t 14,500 ft., 7 July 1954, Stainton, Sykes & Williams 6222 (part) (BM).

A. shrub 1-2 ft. high, which has been cultivated at Kew, where it was examined in fruit on 18 Oct. 1939, grown from seed of the type number (Kew 33/269-Hay 322132).

(14Oc) Berberis concinna var. extensiflora Ahrendt, var.nov., inflorescentiis 2-5- floratis, umbellatis, 2 4 cm. longis

Stem mature dark red, sulcate. Leaves with pinnate veins, rarely slightly subreti- culate, pruinose white below. InJIorescence 2-5-fld., 2-6 cm. long, umbellate, with peduncle 1.5-4 cm. long, a t the base of which there is sometimes an extra flower. Pedicels 7-10 mm., in the umbellate part, 1-2 cm. below; bracts, 1-16 mm. Berries oblong to oblong-ovoid, red, 16 x 7 mm., estylose. Seeds 6-9 in number, yellow.

Nepal (east to central): Larjung (south of Tukucha), Kali Gandaki, among hillside shrubs, about 4 ft., fr. 17 Oct. 1954, Stainton, Sykes & Williams 8175 (Type, BM); Lam- jung Himal, 2 ft., a t 11,000 ft., fr. 27 Oct. 1954, Stainton, Sykes & Williams 9352 (BM): Rambrong, Lamjung Himal, 2-3 ft., at 10,500 ft., fl. 27 June 1954, Stainton, Sykes & Williams 5964 (part) (BM).

There was once in cultivation a plant of B. coneinna of this variety with a 3 4 - flowered umbellate idlorescence whose long pediceh were supported by perfectly foliaceous bracts. These came from seed gathered by Gamble in Sikkim, and were a t his home a t Liss, Hampshire, but could not be found when I sought there in 1938. Specimens are preserved (K), flowering in 1909 (Gamble 29055), and 1910 (Gamble 29883), and fruiting in 1914 (Gamble 30320).

(141) Berberis tsangpoensis Ahrendt, Card. C~TOTZ. (3) 109, 101, 1941; J. Bot.,

8tem glabrous, slender, very sulcate : shoots green; mature lustrous bright yellow, not verruculose. Internodes 1.5-4 cm. Spines 3-0d., concolorous, 10-18 mm. Leaves semi-evergreen, 13-21 x 6-9 mm. ; obovate; base cuneate, sessile; margins with 2-5

Lond., 79 (Suppl.), 50, 1941. (Fig. 29)

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120 L. W. A. ABRENDT spinules (1 mm.), 1 4 mm. apart, but scarcely serrulate; apex rounded, mucronate; above dull yellow-green, closely reticulate; below white, pruinose. Flouters solitary. Pedicels slender, 2-5-3 cm. Prqhylls nil, or 1-2, concolomus, oblong, acute, 4.5 x 2.25 mm. Outer sepals 7-9 x 5-56 mm., elliptic, obtuse. Median and inner sepals both 9-1Ox6-7mm., obovate. Peh% 6-7x4.5-5-25mm., obovate; apex entire, or very slightly incised; base scarcely clawed, with approximate elliptic glands, c. 1-5 x 1 mm. Shmns 4-5mm., apex not or scarcely produced, truncate. Ovules 12-15 sessile, or shortly stipitate. Berries 12-13 x 9-11 mm., oblong, estylose, a t h t partly whitish, ultimately pale to bright red.

S.E. Tibet: h v . Pemako, Tsangpo gorge, near Pemakochung, 7000 ft., fr. 29 Nov. 1924, on steep sunny slopes in gritty soil, Kingdon-Ward 6326 (K) (Type).

Cultivated from seed of the type collection; examined in flower June 1938, 3 June 1942 ; and in fruit, Oct. 1938, the earlier specimens being from Exbury. A very attractive subprostrate species, spreading over several feet, rising 6-12 in. from the ground. Easily distinguished from B. concinnu by this habit, its slender yellow stems, reticulate leaves and 12-15 ovules. Described as growing in long paas, it has apparently not been found elsewhere, or gathered apart from the type collection.

(142) Berberis kumaonensis Schneid., Bull. Herb. Bohsier, (2 ) 5 , 397, 1905; syn. B . caespitosu Madden MS. (Fig. 31)

Stems glabrous; shoots slightly reddish; mature yellow sulcate, not verruculose. Inter- & 5-12 (-20) mm. Spines concolorous, 3-fld., finely sulcate, 15-25 mm. Leaves semi- evergreen, 8-20 x 4-9 mm., obovate; base cuneate, subsessile; margins with 1 (-2) spinose (1.5-2 mm.) serrations; venation branched, not reticulate; above dull green; below pruinose, grey. Pedicels 5-9 mm. Flowers H.C.C. 60111, solitary. Prophylls con- colorous, triangular, acumhte, 2 x 1.5 mm. Outer se@ 7-76 x 3.5-4 mm., ovate, acute. Median sepals 8-10 x 5-6 mm., oblong-obovate, obtuse. Inner sepah 11-12 x 7- 7-5 mm., obovate. Petals (H.C.C. 2), 8 x 5-5 mm., obovate; acutely emarginate with subobtuse lobes ; base scarcely clawed, with oblong, subconcolorous, separate, non- marginal glands, 1-5 x 0.8 mm. Stamens 4-5 mm., produced and apiculate at the centre of their apices. Ovules 6-8. Berries 8-10 x 5-6 mm., red, ovoid, apex attenuate and bent with short thick style.

Kumaon: rocks near Garbyang, Kali valley, 13,OOOft., 15 Sept. 1894 Duthie 2697 (Type, K). Chychna, Bincala, 9OOO-11,OOO ft., Strachey & Madden 3012 (0). Seed only, Osmaston 50.

Garhwal: hiUs above Ramri, 9OOO ft., 4 Sept. 1885, Duthie 3816 (K). Nepal (west) : Ghurchi Lekh, between Lumsa and Murma, 11,OOO ft., growing from dry

crevice at foot of cliff. Prostrate shrub, Sin. high, the only one seen, 11 Aug. 1952, Polunin, Sykes & Williams 5124 (BM).

Cultivated: from Osmaaton’s seed, no. 50, this species waa examined; fl. 17 June 1942; fr. 18 Oct. 1940,s Oct. and 13 Nov. 1943. It forms a compact bush about 1 ft. high and through. The Nepal specimen appears to be a variant with stems more markedly red- purple before becoming mature yellow, and of a prostrate habit. The fruit character of this species links it with Subsect. Diaphanae.

(143) Berberis temolaica Ahrendt, Curd. Chron. (3) 109, 101, 1941; J . Bot., Lond., 79 (Suppl.), 54, Sept. 1941

Stems arching, stout, glabrous; mature dark purple, not verruculose. Internodes 1.5-3 cm. Spines 3-fld., 5-10 (-15) mm., concolorous, slightly sulcate. Leavas 20-45 x 8-21 mm., oblong-obovate; base cuneate to decurrent petiole, 1 4 mm. ; margins with 3-9 spinose (1-2 mm.) spreading teeth (1-2 mm.), 3-8 mm. apart, but sometimes (and on the new shoots always) entire; above dull blue-green, slightly pruinose, with venation

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Berberis and Mahonia 121

branched but scarcely reticulate ; below pruinose, white. Flowers solitary. Stamem 4 mm., slightly produced, apex truncate or obtuse. Ovules 6-11. Berries red, pruinose white, 11-14 x 6-7 mm., oblong-ovoid, with attenuate bent apex and thick short style.

KEY 28a Sepals broad; pedicels 10-15 mm. Sepals narrow; pedicels 5-10 mm.

143a. B. temlaica var. temolaica 143 b. B. t em la i ca var. artisepala

(143 a) Berberis temolaica var. temolaica Pedicels glabrous, pruinose white, 10-15 mm. Outer sepals pruinose white, 5 x 3-25 mm.,

ovate, acute. Median sepals 7 x 5 mm., obovate. Inner sepals 8 x 7-5 mm., obovate- orbicular, concave. Petals 7 x 6 mm., broadly obovate; apex rounded, shortly incised; base broadly clawed with separate submarginal glands, 1.3 x 1.1 mm.

S.E. Tibet: Prov. Pome, Temo La, 13,000-14,OOO ft., fr. 13 June 1934, Kingdon- Ward 5773 (Type, K). Nambu La, 12,0OOft., 3 June 1947, Ludlow, Sherriff & Elliot 13831 (BM).

Prov. Kongbo, Pasum Chu, 13,500ft., fl. 22 June 1947, Ludlow, Sherriff & Elliot 13965 (BM); a t 12,500ft., 10 July 1947, no. 15375; a t 14,800ft. on 14 July 1947, no. 15449; on 27 Sept. 1947, no. 15973.

F’rov. Zayul, Burma-Tibet border, 12,OOOft., 2 Sept. 1931, Kingdon-Ward 10008

This species forms a link between this Subsect. and the Subsects. Dictyophyllae and Diaphanae. It is a straggling bush 6-8ft. high (originally erroneously distributed as ‘ B . melcongemis’) cultivated from KW 5773, examined in flower 30 May 1939, and in fruit, 16 Sept. 1939. It is interesting for its bluish leaves contrasting with purple black stems, and greater abundance of white bloom (which covers all its parts) than on any other species.

(BM) .

(143 b ) Berberis temolaica var. artisepala Ahrendt, J . Bot., Lond., 79 (Suppl.), 55, 1941

Outer sepals 7-5 x 2-75 mm. Median sepals 9-5 x 3 mm. Inner sepals 9.5 x 4.5 mm. Petals 6-5 x 4 mm., submarginate, with lanceolate, separate acuminate glands, 1.5 x 0.35 mm.

S.E. Tibet: Tsari district, Langong, 12,00Oft., 28 May 1938, Ludlow, Sherriff & Taylor 3911 (Type, BM): also 13,000-13,500 ft., 7 June 1936, Ludlow & Sherriff 1785; 21 ,June 1936, Ludlow & Sherriff 1850.

Cultivated: a t Wisley from no. 3911. Berberis temolaica is described as growing in more or less open situations-on grassy

hill slopes, on shrub-clad slopes, or amongst rhododendrons.

Subsect. Diaphanae Ahrendt, J . Bot., Lond., 79 (Suppl.), 55, 1941 Stem glabrous, sulcate (except only in B. virescens where they are terete). Internodes

( l o t i + ) 2 4 (-5) cm. Spines 1-5 cm. Leaves deciduous. InJorescence (1-) 2-6 (-15)-fld., fascicled or subumbellate, rarely elongated and subracemose. Pedicels (6-) 15-40 mm. Flowers (lo-) 13-18 mm. diameter. Inner sepals ( 6 . s ) 76-10 mm. Petals (P) 5-7 mm., always shorter than inner sepals. Ovules (4-) 6-12. Fruit (8-) 10-15 (-17) x 5-8 mm., ovoid, with bent attenuate apex and short thick style.

Kansu, Shensi, Honan, W. Szechuan, Sikkim, Formosa.

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122 L. W. A. AHRENDT

KEY 29 to Subsection Diaphanae (Map 17 and 19) Leaves with 20-40 marginal spindes, b d y elliptic to suborbicular (1 : 1-16).

Leaves entire or with 1-7 (-12) marghd spinules, m o w e r (1 : 2-4). 144. B. Cdrcumewda

ovules 6-12. Mature stems yellow. 145. B.di5phanu Mature stems dark, red or purple. Leaves 1 : 2-2-5, pruinom grey below; stamens apiculete; f i t 16-16 x 7-8 mm.

Leaves 1 : 4-6, epruinose green, below; stamem truncate; fruit 8-9 x 6-7 mm. 146. B. aernulana

147. B.v.nowisomW ovules (3-) 475. Leaves grey, pruinom below; fruit 10-12 mm.

Inner sepals narrow, mute; petals emarginate; leaves spinom; inflorescence

Inner sepals broad, rounded; pet& entire; leaves entire; intloreacence 1-3 cm., 4-9 cm. (4-) 6-16-fld. 148. B. tischleri

3-8-fld. 149. B. wireacem 150. B. elliotii

Probably hem, but indeterminate. 151. B. faberi

(144) Berberis circumserrata Schneid., Sargent, PZ. Wils. 3, 435, 1917; syn. B. d k -

Stems stout, yellow, sparsely black-vemculose. Internodes 1.5-4 cm. Spines stout, concolorous, 2 4 cm. Lea;ves 1.P2-8 (3 .4 ) x 0-6-1.8 (-2-5) cm., oblong-orbicular ; baae contracted to petiole, 2-5 mm. ; margins with 15-40 spinules (1-1-5 mm.) and serrations (0.5-1 mm.), distant, 0-75-2 mm.; closely reticulate, dull; below grey, pruhse. F k r s H.C.C. 2, fascicled 2-5. Pedkls 1-5-3 (4) cm. Outer selm;ls 7-8 x 46-55 mm., oblong- elliptic. Inner sepds 9-10 x 6-7 mm. Petals 7-7-5 x 4-25-4-5 mm., obovate, entire; slightly clawed with glands, 1x0-5mm. Stamens 4-54mm., scarcely produced, rounded-truncate. ovules (3-) 6-7 (-8). Fruit 13-15 x 5-6 mm.

KEY 29a Petah entire; ovules 6-8; pedicels 2-4 cm; M t 13-15 mm. long.

Petals emarginate; ovules 3-5; fruit 9-12 mm. long.

Leaves green, epruinom below; fruit 1P17 mm.

p h m var. circunwewata Schneid., I.c., 1, 354, 1913. (Fig. 32)

144a. B. cirmmerrata var. circunzserrata

144R B. ciroumeenata; var. occidentalim 144 b. B. circzlmserratcc var. subamzata

Spines 15-35 mm.; pedicel8 8-15 mm. Spines 2-10 mm.; pedicel8 15-25 mm.

(144a) Berberis circumserrata var. circumserrata

seed numbers. Shensi: Tai-pei-shan, 1910, Purdom 4 (Type); 8 Feb. 1911, M o m 604, 6048, a l l

Honan: Lao-gum-shan, 21 Sept. 1919 (K), Hers 1202,5000-6000 ft. Cultivated: from Purdom 604A, fl. 21 May 1938; 27 May 1943,3 June 1942. A shrub

of 2-3 ft. in height, with very stout, rigid, yellow stems which are both conspicuous in themselves, and give to the plant a firm shape lacking in so many Berberis species whose slender stems produce a loose, sprawling, ungainly habit. The large flowers, and very long pale red berries are ornamental.

(144 b) Berberia circumserrata var. occidentalior Ahrendt, var.nov., ovulis tantum

Spines stout. Petioles 6-12 mm. Outer sepals 5-5.5 x 3-3.25 mm., oblong-ovate, trun- cate. Median sepals 6.5-7 x 4 - 5 5 mm., elliptic. Inner sepals 7-8 x 5.5-6 mm., broadly obovate. Pekd.9 6.5 x 3.5 mm., emarginate, with obtuse lobes; base cuneate, with elliptic

3-5, fructibus pedicellisque brevioribus distinguenda

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Berberis and Mahonia 123

separate marginal glands, 0.7 x 0.4 mm. Stamens 4.25 mm., slightly produced sub- apiculate to subobtuse. Ovules 3-5. Fruit 10-12 x &-7 mm. Style 0-25-0.4 mm. El. Tibet: Kokonor region, Koko gorge, Sept. 1925, M f t . , Rock 13272 (Type, K),

Rock 14108. S.W. Kansu: Tao river basin, ridges beyond Adjuan, July 1925, Rock 12364 (K); and

E. Tebbu land, 10,500 ft., Rock 13529. Cultivated: from Rock 13272, fl. 21 Apr. 1939 (K).

( 1 4 4 ~ ) Berberis circumserrata var. subarmata Ahrendt, J. Bot., Lond., 79 (Suppl.), 56, 1941

Spines absent or solitary, 2-5 mm., occasionally 3-fld., 5-10 mm. Leaves lustrous above; margins with 15-25 spinules. InJEOrescence 3-5 (-8)-fld., fascicled or partly um- bellate, 3 4 cm. Outer sepals 5.5 x 3 mm. Inner se& 8 x 6 mm., all obovate. Petals 6 x 3.25 mm., elliptic-obovate, rounded, shortly incised; base cuneate, with separate ovate glands, 1.3 x 0.7 mm. Sta7nens 4 mm., truncate. Oeyules (3-) P5. Berries 9-10 x 46-5.5 mm., including style, 0-5-0.75 mm.

Kansu: Farrer 238, seed number. Cultivated: from Farrer 238, fl. Apr. 1938; fr. Oct. 1938. (Type, BM). This plant was

originally cultivated a t Wisley (under the name ‘ B . tischleri var. ’) whence it has been distributed.

(145) Berberis diaphana Maxim., Bull. A d . Sci. St-P&mb. 23, 309, 1876 Stem with young shoots green; mature pale yellow, verruculose. Internodes 1-2 cm.

Spi,nes 3-fld.,concolorous, plane, stout, 1-2 cm. Leaves 16-40 x 5-16 mm. ; petiole 1-3 mm. ; margins with 4 1 2 spinose (0.5-1 mm.), sermlations, 3-6 mm. apart; above dull grey green with veins elevated, reticulate ; below pruinose, grey. InJlorescence (1-)%5-fid., fascicled or subracemose. Pedicels 12-22 mm. Plowers H.C.C. 2. Outer sepals 8 x 5.5 mm. Inner sepals 9 x 6 mm. Petals 7 x 5.5 mm., ovate-elliptic, acute, narrowly incised; base clawed, with ovate, separate non-marginal glands, 1.2 x 0.4 mm., situated just above the base of the small claw. Stamens 4-5mm., scarcely truncate. Ovules 6-10. Berries 10-12 x 6-7 mm. Style 0-25-0.5 mm.

KEY 2 9 ~ Flowers not solitary; fruit bright red.

Flowers fascicled. Flowers subracemose.

Flowers solitary : fruit dark red.

145a. B. diaphana var. diaphana 145 b. B. diaphana var. tachiensis

145c. B. diaphana var. uniflora

(146a) Berberis diaphana var. diaphana Kansu: Przelwalski, s.n. 1872 (Type, K). Cultivated: Vilmorin 50, type of Bot. Mag. t. 8224. L.A. 204, fl. 20 May 1942,25 May

194t; fr. 29 July 1943,24 Aug. 1944. Similar in habit to B. circumerratu, this is a shrub 2-3 ft. high with stout rigid, horizontally arching branches; the berries are shorter and paler, and in both species ripen very early.

(145 b) Berberis diaphana var. tachiensis Ahrendt, var.nov., idorescentiis sub- racernosis; syn. ‘B. diaphana’ Rehd., in Sargent, Trees and Shrubs, 2, 19, t. 119 (19077, non Maxim.

InJlorescence 3-5-fld., pseudumbellate-subracemose, 2-5-3-5 cm. long, else as in type. W. Szechuan: Kangtin Hsien, Tachien lu, 28 Sept. 1928, Fang 3655 (Type, K). Omei

shall, Fang 7711 (K). Not in cultivation.

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124 L. W. A. ~ E N D T

(145c) Berberis diaphana var. uniflora Ahrendt, vwnov., floribus plenunque

Stems suberect. P’lowers mostly solitary. Pedhls 12-22mm. Sepals H.C.C. 1-3/1, outer 5.5 x 4 mm., median 8 x 5-5 mm., inner 8-9 x 6 mm. Petccls 6-5 x 5 mm., incised, or acutely emarginate with narrowly obtuse lobes; base clawed, with oblong, separate, non-marginal glands, 1 x 0-5 mm. r9tanzens 5 mm., produced, obtuse. O& 7-9. Berries dark red, epruinose, 10 x 6 mm.

W. Szechuan: Pan lan shan, W. of Kuan Hsien, 13,000-15,000 ft., 2-4 ft., 24 June 1908, Wilson 2865 (Type, K).

Cultivated: L.A. 444, fl. 29 Apr. 1943; fr. 16 Sept. 1943.

(146) Berberis aemulans Schneid., Sargent, PI. Wib . 3, 434, 1917; Ahrendt, Bot.

Stems with young shoots e n ; mature dark purple, verruculose. Spines orange-red, sul- cate, weak, 3-fld., 6-11 (-15) mm. h v e s 2-4 (-5) x 1-2 cm., oblong-obovate; base cuneate to decurrent petiole, 2-5 mm.; margins with 5-12-spinose (1 mm.) serrations, distant 44mm.; above reticulate, dull green; below grey, pruinose. InJlorapeence 2-4-fld., fascicled. Ped& 2-3 cm. Outer sepals 7-7-5 x 4-4.5 mm. Inner sepals 7.5-8 x 4-5- 5 mm., all oblong-elliptic. Petals 5 x 3-75 mm., oblong, entire; glands oblong-elliptic, separate, non-marginal, 0.6 x 0.3 mm. Stamens 4 mm., produced, apiculate. Ovules 7-11. Berries 15-16 x 7-8 mm.

W. Szechuan: near Washan, thickets, 1908, Wilson 930 (Type, K). Mt. Omei, thickets, 1928, Fang 2977 (K) ‘fruit with a long point ’.

Cultivated: L.A. 200, fl. 12 May 1942; fr. Oct. 16, 1942. The Bot. Mug. specimen is atypical, with shorter pedicels.

solitariis.

Mag. 169 N.S., t. 179,1952

(147) Berberis morrisonensis Hayata, J. Coll. Sci., Tokyo, 30, Art. 1, 25, 1911,

Stems with young shoots green; mature dark red, scarcely verruculose, #pines 34d., slender, 1-2 cm., slightly paler, plane or subsulcate. Leaves 10-30 x 3-7 mm., oblanceolate ; base attenuate to decurrent petiole, 2-5 mm. ; margins with P 7 spinose (1 mm.), serrations (0.5 mm.), distant, 2-4 mm.; above openly reticulate, lustrous deep yellow-green; below at Grst grey, pruinose, often ultimately epruinose and pale green. Flowers H.C.C. 2, fascicled 2-5. Pedicels 12-25 mm. Outer sepals 4 x 1.5 mm., lanceolate, or 4.5 x 2.5 mm., narrowly ovate, subacuminate. Necliun sepah 5.5-6.5 x 3-3.5 mm., oblong-elliptic. Inner sepals 7.5 x 4 mm., narrowly obovate. Perk& 5-6 x 3.54 mm., elliptic; acutely emar- ginate with subobtuse lobes; basal glands elongated. Shmm 34.5 mm., scarcely pro- duced, rounded or truncate. OvuZm (&) 5-7. Berries 8-9 x 6-7 (-8) mm.

Formosa: 1906, Kawakami t Mori 2289,2297, Types. Yashiroda 8, seed only. Prov. Kagi, Arisan to Mt. Morrison, 11,500-13,500 ft., 25 Oct. 1918, Wilson 10912 (K).

Cultivated: L.A. 99-, fl. 3 June 1942; fr. 7 Oct. 1943. Also at Kew, from Yashiroda 8, fl. 23 May 1940; fr. 18 Oct. 1939; also, type of Bot. Mug. t. 9017, from Wilson 10912. Described as an abundant shrub, 2-6 ft. high.

Bot. Mug. 149 t. 9017, 1924

(148) Berberis tischleri Schneid., BuU. Herb. Boissier, (2) 8,201,1908; Sargent, PI. Wils. 1, 355, 1913

Sterns shoots reddish; mature yellow-brown. Spines 1-2-5 cm., sulcate. Leaves (1.6) 2.7-5 (-6) x ( 0 . s ) 1.2-2.4 cm., including petiole 2-3 mm.; oblong-obovate; mar- gins with 4-7 spinose (1 mm.), serrulations; occasionally entire; apex rounded or obtuse; below grey, pruinose; venation branched. Inflorescence (P) 8-12 (-15)-fld., loosely um- hellate-racemose, 3-9 cm. long, including peduncle, 1-5-3 cm. Pedicels (5 mm.-) 1.5-

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Berberis and Mahonia 125

2 cm., towards apex of inflorescence, but a t base, 2.5-3.5 em. ; bracts lanceolate, 2-3 mm. Outer sepals 5 x 2 mm., narrowly ovate, acute. Inner sepals 6.25 x 3-25 mm., narrowly oblong-obovate, acute. Petals 4 x 2mm., emarginate; basal glands oblong-ovate, separate. Stamns 3.5 mm., shortly produced, apiculate. Ovules (3-) 4. Fruit 10-12 x 5-6 mm.

KEY 29c Inflorescence 8-15-fld., 6-9 em.; pedicels 15-35 mm.

Inflorescence 68-fld., 3-6 em.; pedicels 5-15 mm. 148a. B. tischleri var. tischleri

148b. B. thchleri var. abbrewkta

(148 a) Berberis tischleri var. tischleri W. Szechuan: Nereku, fr. 26 July 1885, Potanin, sm., Type. Also Pan lan shan, 1910,

Wilson 2859 (K), Min valley, Mao-chu, fr. 1908, Wilson 1177 (K). Tachien lu, 1904, seed only Wilson (Veitch) 1731.

Cultivated: from Wilson 2859, fl. May 1939; and from Wilson (V) 1731, fl. May 1938; fr. 26 Oct. 1938. A tall shrub, 6-8 ft. high.

(148b) Berberis tischleri var. abbreviata Ahrendt, var.nov., inflorescentiis pedi- cellisque brevioribus.

W. Szechuan: Tachim lu, 8000-12,000ft., 1908, Wilson 2854 (Type) 2855; Pan lan shan, Wilson 2856 (all, K).

(149) Berberis virescens Hook. f. e t Thorns., Bot. Mug. 116, t. 7116, 1880 ~Ytt?m mature dark red, scarcely verruculose, lustrous, terete or almost so. Spines

concolorous, subsulcate, 3-fld., 6-15 mm. Leaves 10-30 x 4-12 mm. ; oblong-obovate; base cuneate to decurrent petiole, 2-6 mm. ; margins entire; apex obtuse, mucronate (1.5) mm.; openly veined, not reticulate; above dull green; below grey, pruinose. In- jlorescence P8-fld., umbellate-racemose, 1.5-3 em., including peduncle, 6-10 mm. Pedicels 8-12 mm., slender. Plowers pale yellowish green, H.C.C. 66312-311. Prophylls con- colorous, 4 x 1.5 mm. or 4.5 x 1 mm., oblong-lanceolate. Outer sepals 6-25 x 4 mm., obovate. Inner sepals 7.25 x 5 mm. Petals 6 - 3 4 mm., obovate, entire; base clawed, glands greenish, subapproximate, non-marginal, 1 x 0.6 mm., situated above the claw. Sta,mens 5 mm., produced, apex couical-apiculate. O& (3-) P 5 . Fruit 8-10 x 3.5- 4.5 mm., with style 0.5-0-75 mm.

Sikkim: 1849, Hooker s.n. (Type, K). 8.E. Tibet: Prov. Kongbo, 10,000-15,000 ft., 23 May 1938, Ludlow, S h e d & Taylor

4394; 24 May 1938, Ludlow, S h e d & Taylor 3834; 23 May 1938, Ludlow, S h e d & Ta;ylor 3831.

Tsari district, 2 June 1936, Ludlow & S h e d 1758. W. Bhutan: Ha valley, Dumthang, 1 June 1933, Ludlow & Sherriff 48 (all BM). Nepal: Khyangshar Khola, rive droite, fleurs blanches, 6-12 ft., 9350 ft., fl. 10 May

Cultivated: L.A. 258, from seed from Sikkim, fl. 19 May 1943; fr. 22 Sept. 1943. B. virescens has, in cultivation, been much confused, with two other species which are

superficially similar in habit, stem, leaf and inflorescence. Thebe are: (1) B. paravirescem, which is identical in the above characters, and in its apiculate stamens, but has a different flower structure, and oblong black, estylose fruit. Hooker confused these two, and in his Bot. Mag. description stated ‘fruit red or black’; his figure is a mixture of the two species, showing the flower of B. paravirescens, and the bright red attenuate-stylose fruit of B. virescens. (2) B. ignorata, which has sulcate stems, spinose leaves, bright yellow flowers (not pale green as in B. virescens and B. paravirescens), obtuse stamens

1952, Zimmermann 480 (Geneva).

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126 L. W. A. ~ E N D T

and shorter, red, oblong, estylose fruit (cf. J. A&. Sm. Beng. (Sci.), 4, 19 Nov. 1945, where I Grst made theae points clear). B. p r a v i r m w is the most common in culti- vation (but as B. virescew). The true B. virescew is very ram, and forms a spreading shrub, 6-10 ft. high, whose red stems are attractive in the winter.

(150) Berberis elliotii Ahrendt, sp.nov., aff. B. tiscfiri, sed foliis plerumque integri~3, subtus viridibus, epruinosis; infloreacentiis paucitloris pedicellisque brevioribus, fructibusque multo longioribus distinguenda

Stemwith youngshootsred; matureyellow-brown,lustrous. Spines3-fld.,to 1 cm. Leaves obovste, to 35 x 15 mm., entire (rarely with few spinules), venation branched, scarcely even faintly reticulate; above dull; below dull green, epruinose. I n . e s c e m 5-8-fld., umbellate-racemose, 2-5-4-5 cm. long. PediA 8-12 mm.; bracts 1.5 mm. Ovules 4 4 . Berries red, epruinose, 13-17 x 6-8 mm., ovoid, apex attenuate to a very short style. S d pale red.

S.E. Tibet: Prov. Kongbo, Lusha La, 12,000 ft., 1 Oct. 1947, Ludlow, Sherriff & Elliot 13293 (Type, BM).

Cultivated: from the type collection. The following specimen from Nepal is apparently related to B. eUiotii : Okhaldhunga-

gaon, south of Dhorpatan, amongst hillside shrubs, a t 10,000 ft., 1 May 1954, Stainton, Sykes t Williams 391 (BM), which has stout yellow subglabrous, very sulcate, slightly verruculose stems; 3-fld. spines, the lateral p& equal to the centre, 1-5 cm., and inter- nodes (1.5-) 2-3 (-4) cm.; obovate leaves to 3-5-1-7 or 3 x 1-8 cm., thin, entire, dis- tinctly reticulate, subconcolorous green below ; racemes 4-8-fld., subumbellate and 1-63 cm. long with glabrous pedicels, 5-10 mm., and bracts 1.5 mm.; flowers 16 mm. diameter, with outer sepals 6 x 3 mm., elliptic, rounded; inner sepals 9 x 7 mm., obovate; petals 7 x 4 mm., subentire, rounded, cuneate with marginal separate glands, 1 x 0.5 mm., and distinctly produced, subapiculate stamens. As this specimen is only in flower, and B. eUwtia' is known only in fruit, and their habitats are Merent, no closer association can be made.

(151) Berberis faberi Schneid., Ost. bot. Z., 67, 215, 1918 Stems glabrous. Intf?& 2-2.5cm. Spines 3-fld., subsulmte, 1-1-5cm. Leaves

15-32 x 10-16 111111. ; base contracted to decurrent petiole 1.2 mm.; margins entire; closely reticulate; below grey, pruinose. Injlmescem 3-6-fld., fascicled or subfascicled. Pedicels 1-8-2-5 cm. ; bracts 1-1-5 mm. O m l a 6-7, sessile. Otherwise unknown.

W. Szechuan: Mt. Omei, 9800 ft., 1887, Faber 229 (Type, K). Not in cultivation.

Subsect. Dictyophyllae Ahrendt, J. Bot., LonrE., 79 (Suppl.) 51, 1941 Stem glabrous; mature always dark, red, purple or black, mostly pruinose white.

Leaves deciduous, 5-20 mm.; margins entire or with 2-6 spinules; mostly closely reti- culate. P'lowers solitary, or fascicled, 2-8 x 10-18 mm. diameter. Inner sepals (6) 7- 10 mm. Petals emargimte, 4-8 mm., nearly always shorter than (but in B. ambrozyana equal to) inner sepals. PediA glabrous, 1 mm. to 4-5 cm. Stamens always produced, truncate or apiculate. 0wu.h 3-5 (-6). Fruit red, always stylose and pruinose white, globose, 6-9 mm., or elongated, 9-17 mm.

Yunnan, Szechuan.

KEY 30, to Subsection Dietyophyllae (Maps 17 and 19) Flowere solitary; h i t stylose.

Leaves cloeely reticulate. Leavea ephose; ovules 44. 152. B. a p p r o x i t a

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Berberis and Mahonia 127

Leaves entire; ovules 3-4. Stems and undersurface of leaves pruinose white; petals shorter than inner

sepals. 153. B. dictyophylla Stems red, undersurface of leaves green, both epruinose; petals as long as inner

sepals. 154. B. ambrozyana Leaves subenervate, or slightly veined. 155. B. atiebritziana

Flowers solitary, 1C20 mm. diameter; fruit 14-17 mm. long. 156. B. rnwliensis Flowers subumbellate, 2-8, diameter 12-16 mm.; fruit 10-12 mm.

Flowers fascicled (1-) 2-8; fruit estylose; ovules 3-4.

Leaves scarcely reticulate above; pedicels 12-18 mm.; flowers 14-16 mm. diameter;

Leaves closely reticulate above; pedicels 5 - 1 0 mm. ; flowers 12 mm. diameter ; fruit and stems and pedicels typically epruinose. 157. B. validbepala

stems and pedicels pruinose white. 158. B. eakipratorurn

(152) Berberis approximata Sprague, Kew Bull. 1909, 256, syn. ‘B. dictyophylla’ Hook. f. Bot. Mag. 228, t. 7833, 1902 non Franch.

f3%m shoots red. Spines slender, 3-0d., 8-18 mm. Leaves 6-20 x 3-10 mm., obovate, cuneate, sessile, margins with 2-6 spinose (0.5-1 mm.), s e d a t e , distant 2 4 mm.; apex rounded; dull bright green; below pruinose, white. Pedicels 4-6mm. Outer sepals 4 2: 3 mm., elliptic. Inner sepals 6-7 x 3 - 5 4 mm., obovate. Petals 4 x 3 mm., shortly acutely emarginate, with 2 acute lobes; glands approximate, 1 x 0-5 mm. Stamens 3.5 mm., truncate. Ovules 5-6. Fruit subglobose, 7-9 mm., excluding style, c. 1 mm. ; red, pruinose white. E. Szechuan: Farges, s.n., seed only. Cultivated : Vilmorin 572, from Fargea’s seed, 1897 (Type, K). Not now in cultivation

in England.

(153) Berberis dictyophylla Franch., Plant. Delavay. 39, 1889 Stems with young shoots subterete to subsulcate; mature terete. Internodes 1-2 cm.

Spines stout, 1 . 6 3 cm. Leaves 10-20 x 4-10 mm., obovate, cuneate, sessile, entire, obtuse, dull, below pruinose white. Pedicels 6-15 mm. Outer sepals 6 x 2.5 mm. Inner sepals 8-9 x 4mm., all narrow (except in 153c). Petals 8 x 3 (-6) mm., narrowlyobovate, entire or broadly incised; base clawed, withwidelyseparatedglands, 1.2 x 0.6 mm. situated towards base of claw. Stamem 4 - 6 5 mm., slightly produced, shortly apiculate to subobtuse. .Fruit red, pruinose white, ovoid, (9-) 10-12 (-13) x (6-) 7 (-8) mm. Style 0-25-0-5 mm

KIGY 30a Leaves narrow (1 : 2.5) ; pedicels 6-15 mm. ; fruit ovoid with straight style.

Stem subterete, pruinose white; leaves below pruinose white; flowers pale yellow.

Stems sulcate, epruinose; leaves below green, epruinose; flowers deep yellow. 153a. B. dictyophylla var. dictyophylla

153 b. B. dictyophylkc var. epruinoaa

153c. €3. dictyophylla var. campylogyna Leaves broad (1 : 1-75) ; pedicels 1-3 nun. ; sepals broad; fruit globose with bent style.

(153a) Berberis dictyophylla var. dictyophylla Yunnan: Yen-tze-hay, above Lankong, and above Mo-so-yn, 0. 31 May 1885; fr.

14 Oct. 1887, Delavay, s.n. (Type, K). Tali, Yang-pi, 1914, Schneider 2625 (K). Chung- tien, Lichiashica, 10,400 ft., 16 Nov. 1937, Yu 10971 (E).

S.W. Szechuan: Yetsi Mts., 11,OOO ft., 1932, Rock 24416 (BM, K). Cultivated: from Yu 10971, fl. 5 May 1944. The cultivated plants have somewhat

wider flower components, outer sepals 7.5 x 4 mm., or 7 x 5 mm., inner sepals 8-10 x 6-

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128 L. W. A. ~ E N D T

6.5 mm., and petals 7-8 x 5-6-25 mm. Described as casual in its habitat, this 6 ft. high, very loosely spreading shrub is very rare in cultivation, where it is mostly represented by the variety cam~bgyna.

(153b) Berberis dictyophylla var. epruinosa Schneid., S q e n t , PI. Wils. 1, 353,

Stem red, epruinose, sparsely verruculose. Spines yellow, 3-fld., 1-2 cm., sulcate. Leaves to 16 x 6 mm., acute, mucromte; below green, epruinose. Pedicds stout. Outer sepals 6 x 3 mm., ovate, acute. Inner sepals 7.5 x 4 mm., obovate. Petals 5 x 3.5 mm., slightly emarginate with two acute lobes; glanb narrowly obovate, 1.2x0.6mm. StamRnS 3-75 mm.

W. Szechuan: Ta-pm-shan, N. of Tachien-lu, thickets, 1 2 , ~ 1 4 , 0 0 0 ft., July 1908, Wilson 2866 (Type, K).

S.E. Tibet: Tsari district, Kyimdong Chu, 13,700 ft., Ludlow & S h e d 1808 (BM). Prov. U, Reting, 60 miles N. of Lhasa, 13,500 ft., 12 July 1944, Ludlow & S h e d

9974 (BM). Not in cultivation. Described aa bearing golden yellow flowers, and growing on open grassy valley beds.

1913

'

(1534 Berberis dictyophylla var. campylogyna (Ahrendt) Ahrendt comb.nov., ; s p . B. approximata var. canvpybgyna Ahrendt, J. Bot., M., 79 (Suppl.), 53, 1941. (F'ig. 33)

Stem shoots bright red, not verruculose, all pruinose white. Spines 1-3-fld., 5-9 mm., finely bisulcate. Leaves 10-16 x 6-9 mm., entire, subobtuse, bright, yellow-green, below pruinose white. Flowers globose, not fully opening. Pediceh 1 3 m m . Outer sepals 5 x 4 mm. Median sepals 7 x 5.5 mm. Inner sepals 8 x 6 mm. Petah 7 x 5 mm., narrowly incised; base clawed with approximate glands, 1.2 x 0-6 mm. Berria pale red, pruinose white, globose, 7-8 mm., with apex attenuate into a thick bent style, 0.5-0.75 mm.

Yunnan: Forrest 13224, seed number. Rock 24276, seed number. E. flank of Lichiang range, 11,OOO ft., May 1910, Forreat 5555 (BM, K). N. end of Hockin valley, 1913, Forrest 10029 (K).

W. Szechuan: Muli, Wachin, near Lamasery, 2 Oct. 10,200 ft., Yu 14412 (E). Cultivated: from Forrest 13224, fl. 30 May 1939, 18 Bhy 1941, 20 May 1942, 3 June

1942; fr. 27 Dec. 1939 (Type, 0). Also from Rock 24376, fl. 1939 (K); and from Yu 14412, fl. 4 May 1944.

Described as a common shrub in open mountain pasture, this variety is also wide- spread in cultivation; growing to 6 ft., with a very lax habit, and generally passing as ' B. dictyophyh ', it is valuable for the white bloom on its shoots and leaves and the red colour of its leaveg each autumn.

(154) Berberis ambrozyana Schneid., Sargent, PZ. Wils. 1, 356, 1913 Stemwhen mature daxk red, epruinose, finely sulcate. Spines often absent or subterete,

1-3-fld.,5-10mm. Leaves8-22 x 1.5-8mm.,obovate, cuneateseasile entire(rare1ywith 3-6 spinules), obtuse, mucromte; below at first slightly greyish pruinose, ultimately eprui- nose, green. Pedkek 2-3 mm. Outer s e p h 6 x 3 mm. Inner s e p h 7 x 3-75 mm. Petals 7 x 3-75 mm., base slightly clawed with oblong, separate, approximate glands, 1.5 x 0.5 mm. Stamens 4 mm., apiculate. Be& subglobose, 6-7 mm., shortly stylose.

W. Szechuan: 10,400-11,OOOft., fl. June 1904, Wilson (Veitch) 3146a (Type, K), 3146; Kangtu Hsien, Tachien lu, 9500 ft., 27 Sept. 1928, Fang 3645 (K). Not in culti- vation.

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Berberis and Mahonia 129

(155) Berberis stiebritziana Schneid., Ost. Bot. Z., 66, 320, 1916 #pines 3-fld., 1-2.5 cm. Leaves 6-20 x 3-7 mm., obovate, subobtuse, sessile, entire

(rarely margins with 1-4 spinules) ; enervate or but shghtly veined; below grey, pruinose. Flowers subglobose, not fully opening, H.C.C. 312-311. Outer sepals 8-5 x 6-5 mm. Inner sepals 9 x 7 mm., all oblong-obovate. Petals 7 x 5 mm., acutely emarginate, with acute lobes; base slightly clawed with approximate submarginal glands, 1.5 x 0.8 mm. Stamens 5 mm., not produced, truncate. Ovules 3-4. Berries red, slightly pruinose, white, ovoid, 10-12 x 6-7 mm., with short style. Pedicels 12-23 cm.

Jlunnan: E. of Lichiang, thickets 9800 ft., 16 Sept. 1914, Schneider 2908 (Type, K). Hungpoh, Bengah, 11,000 ft., 19 Nov. 1937, Yu 7999 (E). Weihi, N. of Tung chuling, 12,300 ft., 5 Nov. 1937, Yu 10625. Bei-ma-shan, 12,300 ft., 26 Nov. 1937, Yu 10798 (all E).

S.W. Szechuan: Muli, Wachin, Jin-chang, 12,000 ft., fr. 22 Oct. 1937, Yu 14611 (E). S.E. Tibet: Prov. Kongbo, Lotu, 12,000 ft., 21 Sept. 1947, Ludlow, Sherriff & Elliot

This is a variable species within which may be noticed four forms: (A) The type, with pedicels 12-25 mm. ; style very short, often indistinct shrub, 2-3 ft.,

(B) Pedicels short, 6-11 mm.; style more distinct, Schneider 1922, LSE 15755, not

((7) Pedicels long, 12-20 mm.; style distinct, shrub 5-7 ft., Yu 10978. (ID) Pedicels short; style scarcely distinct; shrub 5-7 ft., Yu 14611 (cult.). In all its forms this is described as a common shrub among thickets (casual only in

15755 (BM).

Schneider 2908, Yu 7999 (cult., fl. 3 May 1944), 10625 (cult.).

CUltL

10625).

(156) Berberis muliensis Ahrendt, Kew Bull. 1939, 268, 1939 Stems shoots bright red, mature dark red, epruinose, sulcate. Internodes 2-5 cm.

Spines sometimes absent, sometimes stout, 2-3.5 cm., plane below. Leaves narrowly obovate; base cuneate to decurrent petiole, 2-3 mm. ; entire (or spinose-serrate), obtuse; dull above; below grey, pruinose. Pedicels stout. Outer sepah 7-8 x 4-5 mm., ovate, acutB. Inner sepals (8-) 9-10 x (6) 6-7 mm., obovate. Petals 6.5 x 5.5 mm., obovate, shortly emarginate; glands narrowly elliptic, acute, 1 x 0.2 mm. Stamns 5 mm., apicu- late. Berries narrow, ovoid or oblong-ovoid, 14-17 x 6-9 mm.

KEY 3 0 ~ Leaves entire. Leaves spinose serrate.

Lftaves 1 : 2-2.5; pedicels 2.5-4.5 111~~1. Leaves 1 : 5; pedicels 18-22 mm.

156a. B. muliensis var. mulien&

156b. B. muliensis var. atuntzeana 156c. B. muliensis var. beirnanica

(156 a) Berberis muliensis var. muliensis Leaves 15-30 ( 4 5 ) x 3-7 (-11) mm., entire. Pedicels (I-) 1.5-3 cm. Prophylls yellow,

broadly obovate 2-5 x 2.5 mm., acute. S.W. Szechuan: Muli Mts., 11,700ft., 1921, Forrest 20431, and fr., Forrest 20633

(Type, K). Muli territory, Muli Konka, snow range, east of the Yalung, Mutimong, 11,500 ft., MayJune 1932, Rock 23738 (BM, K). (The latter specimen hm lanceolate prophylls, and equal outer and inner sepals, 9-10 x 4-5 mm., but conforms otherwise).

Cultivated: fr. 10 Sept. 1947, from E. H. M. Cox, Glendoick, who states that the species is remarkable for its lateness in coming into growth (mid-June), and for the long whipping. young growths with no side shoots. Described as coming from open alpine meadows and Spruce forests.

9 JOURN. LIEN. S0C.-BOTANY, VOL. LVII

Page 130: Berberis and Mahonia : A taxonomic revision

130 L. W. A. AHRENDT

(1566) Berberis muliensis var. atuntzeana, Ahrendt, Is., 1939,269.

ProphyUs red, 3 x 1 mm. Inner s e p b 8 x 5-6 mm. Leaves 2-5-3 x 1-16 cm., obovate; contracted, sessile; margins with 3-7 s p h d e ~ .

Yunnan: N.E. of Atuntze, Sept. 1921, Forrest 20713 (Type, K). S.E. Tibet: Prov. Tsmng, Forrest 20624 (K). Mts. of Tjonatong, Upper Salween

river, 13,000 ft., June 1932, Rock 22350 (BM, K). Not cultivated. Described as 3-5 ft. high, with oraage flowers.

Kew Bull. 1939, 270 (156~) Berberis muliensis var. beimanica Ahrendt, 1.c. 1939

distant.

Described aa a shrub of 2-3 ft., on ledges and clifFs on the stony s l o p of its habitat.

(157) Berberis validisepala Ahrendt, Kew Bug. 1939,270,1939 Internodes 14-36 cm. S p i w 3-fld., plane, 10-14 mm. Leaves 10-30 x 4-10 mm.,

obovate; base contracted to petiole, 2-7 mm. entire; rounded, mucronulate; above dull grey green, with only a few lateral veins; below white, pruinose, reticulate. InfEoresCence fascicled, rarely umbellate with peduncle, 4-8 mm. Fknmrs H.C.C. 3. Pedicels stout, 12-18 mm. ProphyUe yellow, oblong, acute, 3.75 x 1-75 mm. Outer sepal8 7 x 5 mm., oblong-ovate, acute. N e d k n ee+ 7-8 x 4.5-6 mm., obovate. Inner sepals 9 x 7 mm. Petals 7-8 x 4 5 mm., obovate, shortly emarginate with acute lobes. Base cuneate with broadly elliptic glands, 1 x 0-6 mm. Stanzens 4 5 mm., apex produced 0.5 mm., apiculate or subapiculate. Ovule8 3 4 . Be& broadly ellipsoid, 8-9 x 6-5-76 mm.

Leaves oblong-oblanoeolate, 20-30 x 4-6 mm., margins with 5-7 spinules, 2-5 mm.

N.W. Yunnan: Bei-ma-shan, 1921, Forrest 21002 (Type, K). Not in cultivation.

This species is related to Subsect. Euangulosae.

KEY 300 Stem and fruit epruinose. 157a. B. waMi.sepakz var. validhepala Stem and fruit at first pruinoae, white. 157 b. B. d k h e p d a var. p r i n w g h w

(157 a) Berberis validisepala var. validisepala Yunnan: seed, Forrest, s.n. Cultivated: fl. 20 May 1938 (K), 19 May 1943, 5 May 1944; fr. 26 Oct. 1938, 13 oct.

1939 (K). The type is Kew no. 152 (K)

(1576) Berberis validisepala var. primoglauca Ahrendt, J. Bot., Lond. 79 (Suppl.), 62, 1941.

Yunnan: 1932, seed, Forrest (native collectors after his death). Cultivated: fr. 15 Oct. 1939 (Goring, Sir Frederick Stern, Type BM).

(158) Berberis calcipratorum Ahrendt, sp.nov., aff. B. dictyophyllae, sed inflores-

Stems very sulcate, thick, dark red, pruinose white. I&& 2-3 cm. Spines orange, stout, 1-3-fld., 8-15 mm. Leaves 10-22 x 4-9 mm., narrowly obovate; base cuneate, sessile; entire, obtuse, mucronate. Inflmescence 2-2.5 cm., 2-5-fld., with peduncle subnil. Pedicels 5-10 mm., pruinose, white. Bracts c. 2 mm. Outer sepals 5.5 x 4 mm., elliptic. Inner sep& 7.5 x 4 mm., obovate. Petals 5.5 x 3.5 mm., oblong-obovate; glands elliptic, separate, non-marginal, 1 x 0.4 mm. Ovules 3-4, stipitate. Stanzen~ 3.5 mm., apiculat e.

centiis pseudumbellatis, ramulis valde sulcatis distinguenda

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Berberis and Mahonia 131

Punnan: E. flank, Tali Range, 11,000-12,OOO ft., May 1910, Forrest 5552 (Type, K). Not cultivated. A shrub 5-8 ft., with deep yellow flowers growing in open situations on limy pasture.

Subsect. Sibiricae, subsect. nov., aff. Subsect. Angulosae, sed floribus multo minoribus, ovulis plerumque tantum 3-5 (Maps 17 and 19)

Spines 3-5 (-7)-fld. h v m deciduous. Plowers solitary, in diameter 6-8 (-10) mm. .Pedz'cels &9 (-12) mm. Inner sepals P 5 (-6) mm. Petals 3.5-4-5 (-5) mm., always shoi%er than inner sepals. Stamens 2.5-3 mm., always shorter than petals, and with apex not or scarcely produced, truncate. Ovules 3-5 (-8). Berries red, 7 4 - 9 ) x 6-7 mm., estylose or with extremely short style.

Siberia, Northern Mongolia, S.E. Tibet (Tsari district), North-east, and Central Bhutan.

BY 31 to Subsect. Sibiricae (Maps 17 and 19) Stems glabrous, subterete; leaves broad (1 : 1.5). Stem pubescent, sulcate; leaves narrow (1 : 2.5-4).

159. B. boreali-sinenais

Leaves 1&24 mm., with spinose margins, below subconcolorous lustrous green;

Leaves 5-12 mm., entice, below pruinose and grey or white; ovules 3-4; fruit stylose. ovules (5-) @3; fruit estylose. 160. 3. sibirica

162. B. tsarica

(159) Berberis bored-sinensis Nakai, J. Jap. Bot. 15,528, 1939 Stems mature red or orange red, quite glabrous. Spines 4-7-fld., 9-12 mm. Leaves

10-18 x 6-11 mm., obovate or orbicular-obovate, openly veined. Pedicels 6-7 mm., glabrous, red, 2 mm. broad a t apex. Berries red, ellipsoid, 6-8 x 4 mm.

Hnpeh: in Monte Hsiao-wu-tai-shan, Yo Takenaka 38, fr. Aug. 1938 (Type, in Herb. Univ. Imp. Tokyo). Not in cultivation.

(160) Berberis sibirica Pallas, Reise Prov. Russ. 2, Anh. 737, 1773, F1. Ross. 1 pars

Stem very sulcate; shoots puberulous, mature glabrous and pale yellow brown. Inter- nodas 6-10 (-15) mm. Spines 3-7-fld., slender, 3-8 (-11) mm., spreading a t base to 2 mm. broad, or partly foliaceous. Leaves 10-24 x 4-6 mm. ; base cuneate to decurrent petiole, 3-5 mm. ; margins with 4-7 spinose (1-2) mm. spreading serrulations (0.5 mm.), 2-3 (4) mm. apart, above bright green; below subconcolorous lustrous green, epruinose, venation on both sides openly elevated, sometimes subreticulate. Flowers 6-7 mm. diameter. Pedicels 7-10 (-12) mm. Outer sepals 4 x 2 mm., oblong-ovate. Inner sepals 4.5 x 2.5 mm., obovate. Petals 4.5 x 2.5 mm., obovate, shortly emarginate. Fruit red, obovoid, estylose, 7-8 (-9) x 6 (-7) mm.

Siberia : in cacumine alpis Serschenskei prope munimentum Feklistovskei inter saxa, Karelin & Kiriloff 606, and 1840 (K).

N. Mongolia: 1877, Potanin, s.n., and 1910, Pride 335 (both K). A densely branched shrub to 2 ft., apparently not now in cultivation, though it was recorded in gardens in the nineteenth century. In PI. URSS 7 , 555,1937, it is stated to grow in the Altai, Angaro- Sayan, Dauria and Dzungaro-Tarbagatai floristic regions. Certain hybrids are recorded below.

2, 42, t. 67, 1788

KEY 31a Style short; leaves grey balow.

Spines weak, &12 mm.; leaves with &12 marginal spinules; racemes 1-3 cm. stems red brown at first; flower diameter 7-8 mm.; inner sepals 5-5 mm. ; fruit

9-2 ellipsoid, 9-11 mm. 161. B. x ernarginata

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132 L. W. A. AERENDT

KEY 31a (continued) Stems yellow-brown; flower diameter 10 mm. inner sepale 6.5 mm. : fruit obovoid,

Spines 1-2 cm.; leavea with 20-40 marginal spinules; racemes 3-4 cm. long. 161a. B. x lemoinei

161 & B. x emarghata var. britzensis 161c. B. x prowincialis

9 mm.

Style absent, leaves green below.

(161) Berberis x emarginata Willdenow, Enum. PI. Hort. Berd. 395,1809; Schneid., BuU. Herb. B&&er, (2), 5, 663, 1905; DC., Syst. 2, 7, 1821; syn. B. pylllgaris var. ernarginata (Wad.) Gordon, Qard. Mag., 16,2,1840

Shrub 2 4 ft. Stem red-brown, becoming yellowish. I & d 1-2 cm. Spines weak, 8-12 mm., sulcate (1-) 3 (-7)-fld. hve.9 13-40 x 5-15 mm., oblong-obovate (but on the young shoote more ovate-elliptic and to 4 x 2 cm.) ; margins with 3-8 spinose teeth, 2-3 mm. deep; below paler and slightly pruinose. Inflorescence racemose-subfaacicled, 1-2 cm. long, densely 5-15-fld. Pedicels 5-8 mm.; bracta 2-3 mm. Outer sepals 4 mm. Inner se@ 5-5 mm. Petala 5 mm. Stamens subapicdate. ovules (2-) 3-4. Be& deep red, slightly pruinose, ellipsoid, 9-11 x 4-5 (-6) mm., excluding short style.

Cultivated: reputed to be B. vdgaris x B. sibirica, but we should notice the discre- pancies in the under surface of the leaves, the apiculate stamens and presence of style. Not cultivated now.

(161a) Berberis x lemoinei Ahrendt, Card. Illust. 44 (3306), 411, 1944 Stem shoota reddish, slightly pruinose; mature pale yellow. Internodes 2-3cm.

Leaves 10-30 x 4-12 mm. ; base contracted to decurrent petiole, 1-2 mm. ; margins with 6-10 spinose (1-2 mm.) sedat ions 2-3 mm. apart; closely reticulate, dull grey-green, grey and pruinose below. Inflorescence &7-%d., umbellate racemose, 2-3 cm., including peduncle 8-12 mm. Pedicds red, pruinose, 3-8 mm. ProphyyWS acuminate, 2.5 x 1.5 mm. Outer sepals 3.25 x 1-25 mm., ovate, obtuse. Inner sepals 6 x 5 mm., obovate. Petals 5-75 x 3-75 mm., apex conspicuously emarginate with two acute lobes; base cuneate, with oblong glands, 1.2 x 0-6 mm. Stanzens shortly apiculate. O& (3-) 4-5 (-6). Fruit red, obovoid, 9 x 6 mm. Style short.

Cultivated: LA. 122, fl. May 1940, 16 May 1944; fi. 11 Oct. 1938, 20 Oct. 1940 (Type, BM). A shrub 5-6 ft., with arching branches, originating from Messm Lemoine’s nursery, and distributed inaccurately aa ‘B. edgeumthiana’.

(161b) Berberis x emarginata vm. britzensis Schneid., J. Arn. Arb. 4, 195, 1923 Innm sepals 4.5 111111. Petah 5 mm., emarginate. Stamens truncate. Reputed to be B. x emrginata x B. amuremi8, but note the discrepant petals longer

than the inner sepals, which might, however, be derived from the supposed B. vulgaris parentage of number 161.

(161c) Berberis x provincialis Audibert ex Schrad., Linnaea, 12,381, 1838; syn. B. vulgaris var. provincialis Audibert cat. 1831-32, 2, 6, nom. nudum

Stem sulcate, yellow. Spines sulcate, 10-15mm. Leaves openly reticulate; below slightly pruinose at h t , ultimately epruinose, slightly paler green. Fruit obovoid, 8-9 x 5 mm., estylose. Otherwise as in B. x emrginata.

Cultivated origin: reputed to be B. vulgaris x B. sibirica. Not now cultivated.

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Berberis and Mahonia 133

(162) Berberis tsarica Ahrendt, J . Bot., Lond., 79 (Suppl.), 48, 1941 Stems slightly sulcate or angled; young shoots pubescent or puberulous; mature dark

(red purple or black), or pale (yellow). Intern& 5-10 mm. Spines 3-5-fld., slender, 3-8 (-11) mm. Leaves 5-12 x 2-5 mm., obovate; cuneate, sessile; entire; obtuse, mucro- natle; below grey (or white), pruinose. Pedicels glabrous or sparsely puberulous, 4-7 mm. Prophylls red, ovate, acute, 2.25 x 1 mm. Outer sepals 3 - 5 5 x 2-5-3 mm., ovate, acute. Inner sepals 5-66 x 3.5-4 mm., oblong-obovate. Petals 3.5-4 x 2-2-25 mm., oblong- obovate ; apex emarginate with 2 acute lobes ; base cuneate with oblong-elliptic glands, 0-7 x 0.4 mm. Stamens 2.5-3 mm., slightly produced, rounded-truncate. Ovules 3 4 , shortly stipitate. Berries obovoid, 8-9 x 6 mm., with short style.

KEY 3 1 ~ Sterns puberulous; mature dark purple ; spines 5-fld. ; leaves grey below.

Sterns pubescent, mature yellow; spines 3-fld., leaves white below. 162a. B. tsar& var. tsarica

162b. B. tsarica var. ritangensk

( 1 6 2 ~ ) Berberis tsarica var. tsarica S.E. Tibet: Tsari district, near Langong, Chianang, 14,500 ft., fl. 5 June 1938, Ludlow,

Sherriff & Taylor 3961 (Type, BM). Tsari Chu, Chusam, 17 Oct. 1938, Ludlow, Sherriff & Taylor 6376 (BM). Bimbi La, 14,000 ft., 3 June 1936, Ludlow & SherriE 1763 (BM).

Prov. Kongbo, Sang La, 14,000 ft., Ludlow, Sherriff & Taylor 5050 (BM). Bhutan (N.E.): Shingbe, Me La, 14,000 ft., 9 June 1949, Ludlow, S h e d & Hicks

20718; Lingshi Dzong, 13,000 ft., Ludlow, SherrifF & Hicks 16303 (BM). Nepal: Alpage de Pheriche, shrub 1-lgft., with petals slightly red outside, a t

14,200 ft., 2 June 1952, Zimmermann 593 (Geneva). This specimen has only old puberu- lous stems.

A small shrub, the type described as 1 ft., the other specimens as 13-3 ft., growing on open hillsides among dwarf junipers. It has been cultivated a t Wisley from no. 6376.

(162b) Berberis tsarica var. ritangensis Ahrendt, 1.c. 79, 49, 1941 Sterns sulcate, very pubescent, pale yellow. Spines 3-fld., plane 6-9 mm. Leaves

c. 6 ;i 3 mm., elliptic, openly veined, below white. Perlicels glabrous, 5-8 (-12) mm. Outer sepals 5.5 x 3 mm. Inner sepals 6.5 x 3 mm. Petals 4 5 x 3 mm., with lanceolate acumi- nate glands, 0.9 x 0.4 mm.

Bhutan (Central): Tang Chu, Ritang, 14,000 ft., fl. 8 June 1937, Ludlow & Sherriff 3220 (Type, BM). Common amongst dwarf juniper and rhododendron; flowers bright yellow. Not cultivated.

Subsect. Jaeschkeanae Ahrendt, J . Bot., Lolhd. 79 (Suppl.), 63, 1941 Stems; mature yellow, otherwise variable. Leaves deciduous; margins entire, or with

1-6 (-8)-spinules; up to 15-45 x 5-19 mm. InJlorescence 2-10-fld., fascicled, subumbel- late or racemose-pseudumbellate. Flowers diameter (6-) 8-10 (-12) mm. Inner sepals 4-7 (-8) mm. Petals 3-6 mm., always shorter than inner sepals. Stamens with apex variitble. Ovules (3-) 6 5 (except in apiculata, zayulana and lasioclem where there are 6-9 ovules). Style mostly absent (distinct only in laswclem, sometimes very short in joceschkeana).

Kashmir, Punjab, Kumaon, Nepal, Sikkim. Bhutan. 8.E. Tibet.

Page 134: Berberis and Mahonia : A taxonomic revision

134 L. W. A. AHRENDT

KEY 32 to subsect Jaeschkeanae ( M a p 17 and 20) Brmb longer than or equal to pedicels. Bracte much shorter than pedicels. Ovules 6-9; stamens apiculate. Stems pubescent; fruit stylose. Stems glabrous; fruit estylose.

163. B. b u m n z c a

164. B. lasioclenw

Leaves entire or with 2-3 marginal spinules. Leaves spinose, 10-25 mm. long; stems more or less temte; pedicel8 5-9 mm.

Leaves entire, 25-45 mm. long; stems sulate; pedicel8 10-18 mm. 165. B. cupiculula

166. B. mydana Cf. 174. B. x Pvilmminii Leaves with 6-12 maxgimd spinules.

O d e s (3-) P 5 . Leavea very nasl'ow, 1 : 5-6. Inflorescence P7-fld., elongated; berries oblong, estylose. Infloresoence 1 (-2)-fld., mostly solitmy; berries globose, long stylose.

167. B.hmbertii

168. B.mucnfo1i.a Leaves broader, 1 : 2-3 (-4). Leaves entire. Stems and pedicels pubesceozt ; leaves green below. Stems and pedicels glabrous.

169. B. hobsonii

170. B. lcartanica

171. B. himidaka stems terete. stems sulate. 172. B . h W & n a

Leaves green below. Leaves grey below.

Leaves spinose. Leaves 8-25 mm. long, with 1 4 (-8) marginal spinules; style often present,

Leaves 2 - 6 6 cm. long, with 8-25 marginal spinules; grey, prubose below; very short. 173. B. jaeschkeuna

stems glabrous; Guit estylose. Marginal spinulea 8-12; stems subterete; fruit 10 x 7 111111.

Merginal sphules 15-25; stems sulata; fruit 8 x 4 mm.

(163) Berberis burmanica Ahrendt, J. Bot. Lvnd., 79 (Suppl.), 69, 1942 Stems sulcate; mature pale yellow brown. I&& 2-45 cm. Spines absent or weak,

3 (-7)-fld., 5-10 mm. Leaw 2-4-3-8 x 1-3-1-9 cm.; oblong-obovate; base contracted to petiole 2-7 mm. ; margins subentire, or with 3-6 (-10) spindose, (1-1.5 mm.) serrations (1-5 mm.), distant 3-5 (-8) mm.; venation openly retidate; below grey, pruinose. [email protected] 3-6 (-10)-fld., racemose-subfascicled, 1-2 cm. long. Pedicela puberulous, 3-4 (-8) mm.; bracts (3-) 4-5 mm., nearly always longer than or equal to the pedicels in length. Berries oblong-obovoid, estylose (7.54 8-9 x (4-) 5-6 mm.

Upper Burma: Nam Tamai valley, Adung W a n g - G d a q W a q , fr. 4 Oct. 1937, Kingdon-Ward 13363 (Type, BM). Growing in thickets along the torrent bank, scattered and not very abundant. In cultivation from the type collection.

(164) Berberis laaioclema Ahrendt, J. Bot., Lond. 79 (Suppl.), 57, 1941 Stenas stout. I&& 1-5-2cm. Spines deeper yellow, broad, 3-fld., 7-10mm.

Leaves obovate, 10-20 x 5-9 mm.; baae cuneate, sessile or with petiole only 1-2 mm. ; mmgins entire, or occasionally with 1-4 marginal spinulea (1-1-5 mm.) towards apex; above deep green, with veins very indistinct ; below epruinose, subconcolorous green, with loosely branched venation. InJEorescence umbellate or subumbellate, some 2 cm. long, including peduncle about 6 mm., and about 4-fld. Pedicela slender, pubescent, 9-12 mm. ; bracts C. 2-5 mm. Prophylk ovate, 1-75 x 1-25 mm. Sepal% obovate-elliptic, all similar

Cf. 174 a. B. x vanfietii

Cf. 175. B. x regle.iana

Page 135: Berberis and Mahonia : A taxonomic revision

Berberis a d Mahonia 135 with apex obtuse or subacute, the outer 4.5-5 x 3-3.25 mm., the inner 5-5-6 x 4 mm. Petals obovate 3-35 (4.5) x 25-2-75 (-3) mm. ; apex very shortly, acutely, emarginate; basal glands separate, narrowly ovate, acuminate, 1 x 0.3 mm. Stamem 2.6 mm., with apex produced and shortly conical-apiculate. Ovary stylose with (41-) 6-8 ovules. Berries unknown.

West Bhutan: Charithang, 11,500 ft., 31 May 1933, Ludlow & Sherriff 41 (Type, BM). Not in cultivation. A shrub with bright yellow flowers, growing on open hillsides.

(16!5) Berberis apiculata (Ahrendt) Ahrendt; syn. B. usteriana var. apiculata Ahrendt.

Stem subterete, yellow. Intemzodes 5-10 mm. 8pine.s 3-fld., 8-15 mm. Leaves to 23 >: 6 mm., narrowly obovate, rounded, mucronate; margins with 2-3-spinose serrations (1 mm.), distant 2-4 mm.; openly veined, not reticulate; below ultimately green, epruinose. InJEorescence 5-8-fld., subumbellate, 2 - 5 4 cm. long. Pedicels 5-9 mm. Prophyb yellow, 3 3 . 5 x 1-16 mm. Outer sepals 5 x 3 mm., elliptic. Inner sepals 8 x 5.5 mm., obovate. Petals 6.5 x 3.75 mm., subacute to subobtuse, entire; base clawed, with separate, oblanceolate glands, 1 x 0.24 mm. Stccmens 5 mm., produced, con- spicuously apiculate. Ovules (&) 6-7, stipitate, with stipes from one-half the length of, to a5 long as, the ovule.

(166) Berberis zayulana Ahrendt, J . Bot., Loncl. 79 (Suppl.), 64, 1941 Stem glabrous; shoots red, subsulcate; mature pale yellow, h e l y sulcate. Intermodes

1-7-3-5 cm. Spines absent, or 1-3-fld., 4 4 (-12) mm. Leave-3 24.5 x 0.7-1-3 cm.; base cuneate to decurrent petiole, 1-8 mm. ; margins entire, rarely with 1-7 spinules; obovate; apex rounded; closely reticulate; below paler green. Inflorescence 5-1 1-fld., pseudumbel- late-subracemose, 2.5-4 cm. Prophylls acuminate, whitish, 1-5 x 1.5 mm. Outer sepab 6 x 3-33 mm., oblong-elliptic to -ovate. Inner s e w 7-5 x 5 mm., obovate-elliptic. Petals 4.5 x 3-5 mm., entire or subemarginate; base scarcely clawed, with separate oblong-obovate glands, 1 x 0.4 mm. Stamens 3 mm., apex produced, truncate at the edge, but shortly apiculate in the centre. Ovules 6-9, often stipitate. Berries estylose, oblong, red, slightly pruinose blue, 10-11 x 5 mm. S.E. Tibet: BOP. Zayd, ham-Tibet border, Rong To-Dibang divide, 12,000 ft., fr.

24 Nov. 1933, Kingdon-Ward 11017 (Type, BM). ‘In thickets in the open; berries scarlet; a very similar species down in the valley has the berries blue-violet.’ Atakang La, Chutang Camp, 12,000-14,000ft., fl. 30 June 1933, Kingdon-Ward 10548 (BM). ‘Flowers bright yellow.’

(167) Berberis lambertii Parker, Kew Bull. 1921,307, 1921 Stems glabrous, angled and h e l y sulcate, pale yellow. Spines slender, 1-3-fld.,

5-8 1(-15) mm. Leaves 12-28 x 2-5 (-7) mm., oblanceolate, sessile, entire, revolute, mucronate; above with but few lateral veins; below much paler, whitish. InJlorescence 4-7-fld., pseudumbellate-subracemose, 26-55 (-7) cm. long, including peduncle 1.5- 4.5 cm. Pedicels fairly stout, (5) 7-10 (-15) mm. Ovules (3-) 6 5 (-6). Berries red, estylose, oblong-ovoid, 10 x 6 mm.

Kumaon: Almora, Lambert, s.n. (Type, K). ‘Between Humidhura and Ratapani, 8100 ft., found nowhere else.’ According to Parker, at Mdba, Ratapani, Kalimundi, 8500 ft., a small shrub, 2-3 ft., growing with normal B. chitria.

(168) Berberis mucrifolia Ahrendt, J . R. Hort. Soc. 81, 135, 1956 A low-growing shrub, less than 1 ft., or up to 2 ft. high. Stem, when mature, yellow,

pubescent, but becoming finally puberulous or partly glabrous ; angled and finely sulcate.

J . Asiat. SOC. Beng. (Sci.) (3) 11, 3, 1945

S h l a Hill States: 1939, Ludlow & Sherriff 7356 (Type, BM). Not in cultivation.

Cultivated at Wisley from the type number 11017.

Page 136: Berberis and Mahonia : A taxonomic revision

136 L. W. A. AHRENDT I n t e 4 5-9mm. S p i m slender, below almost plane, 3-fld. with the centre part longest, 7-2Omm. Leaves narrowly elliptic to narrowly obovate; apex acute to sub- acuminate and conspicuously mucronate (1-2 mm.) ; 6 x 2 mm. to 25 x 3-3.5 mm., or 16 x 3-3-3 mm.; above slightly lustrous green, subeneravate, or with a very slight and indistinct venation; below epruinose, dull, very pale ivory-green; texture some- what stiff. Fluwers mlitary (rarely fascicled 2); 9mm. diameter. Pedicek 5-9mm., glabrous, fairly slender, though sometimes somewhat thickened towards apex ; bracts c. 0.5 mm. Outer se@ 3 x 2 mm., ovate. Inner sep& 5.5 x 3 mm., oblong-obovate. Petals 3.5 x 2-5 mm., oblong-obovate; base cuneate; apex entire. Stamens c. 3 mm., apex slightly produced and rounded. 0mde-s 4-5. Berries bright red, globose or oblong-globose 6-7 x 6-7 mm., excluding a conspicuous style, 1-2 mm. long. Seeds dark

Nepal (east to east central): Tegar, north of Mustang, 14,500 ft., fr. 8 Oct. 1954, Stainton, Sykes & Williams 8111 (Type, BM). ‘On steep graas slopes; shrub 2 ft., berries red. Seed collected.’ Okhaldhungagaon, south of Dhorpatan, 11,OOO ft., fl. 1 May 1954, Stainton, Sykes & Williams 367 (BM). ‘On open grass and rocks; flowers yellow; leaf glossy.’ Above Gurghakhani, 95OOft., immature fr. 21 July 1954, Stainton, Sykes & Williams 3567 (BM). ‘On steep, exposed, earth and rock slopes: leaves whitish green beneath; young fruit reddish.’

Nepal (West): Munigaon, S.E. of Jumla, 9500 ft., fr. 28 Sept. 1952, Polunin, Sykes & Williams 5444 (BM). ‘A dwarf shrub of less than 10 in. on open slopes. Fruit red. Seed collected.’ Sinja, Pulanto Dam, 9OOO ft., fl. 29 Apr. 1952, Polunin, Sykes & Williams 876 (BM): ‘Growing on dry, g r m y hillsides; flowers yellow.’

B. mucrtfdia should be compared with the Kumaon species B. hmbertii Parker, which has similar though larger leaves, and is also a low-growing shrub. B. hmbertii differs in its elongated subumbellate, 47-fld., inflorescence, and in its long (10 x 6 mm.) berries which are quite estylose, and its glabrous stems.

Comparison may also be made with the Garhwal species, B. osnzastonii Dunn, the extreme westernmost representative of the Sect. Wallichianae. Its leaves are like those of B. rnwri,folia, but evergreen and very revolute; it has also solitary flowers and the very long style, and is of a low prostrate habit. But B. osmastonii M e r s in its many ovules and black fruit.

The specimen 3567, cited above, has fruits which appear darker red, and ovoid; the shape may, however, be due merely to immaturity. mere is evidence, however, of a series of plants which lie between B. mwrifolia and B. chitria (with its tall stature, paniculate inflorescence, and narrow elongated dark red berries, also with a long style), and which may be natural hybrids between these, or between B. mucrifolia and B. sikkimenais.

The plants of this variant series were collected in West Nepal by Polunin, Sykes & Williams in 1952 under their numbers 5443,1052, and 4103, the last showing a paniculate inflorescence.

A similar case of natural hybrids presents itself in mnnexion with B. osmastrmii. The collector, after whom it is named, has shown me a plant gathered with it which has the same many-omled, long-stylose, solitary, black berries, but a taller upright habit with larger, broader leaves; this could be a hybrid with B. lycium.

These two -8, suggesting hybrids between (a) low-growing, solitary-flowered species of the Sects. Wallichianae and Angulosae, and (b) tall species of the Sects. Asiaticae or Tinctoriae, possessing racemes or panicles, are somewhat paralleled in Parker’s statement that B. hmbertii was found growing with ‘normal B. chitria’. But, here, there are two differences; no possible intermediate hybrids have been collected; and B. hmbertii possesses no style. In view of the further fact that B. hmbertii has not solitary flowers, we cannot regard it aa the extreme end of the Nepal series of plants with B. mucr$olia (LB an intermediary.

Purple-

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Berberis and Mahonia 137

(169) Berberis hobsonii Ahrendt, sp.nov., aff. B . angulosae, sed inilorescentiis pauci-

A'tems very sulcate; mature pale yellow, puberulous. Spines 10-14 mm., 1 (-3)-fld. h u e s to 15 x 6 mm., obovate, entire, acute; venation branched, not reticulate; lustrous both sides; above deep green, below paler green. Injlwescence 2-6-fld., fascicled or subfascicled. Pedicels puberulous, slender, 6-12, (-16) mm. Flowers 8-9 mm. diameter. Outer sepals 3.5 x 2 mm. Inner sepals 5 x 3 mm. Petals 4-54 x 3 mm., convolute; basal glands 0.8 x 0.5 mm., separate, non-marginal. Sturnens 3 mm., not produced, truncate. ovules (3-) 6 5 . Berries red, oblong, estylose, 8 x 4.5 mm.

South Tibet: Yatung (between N.W. Bhutan, and N.E. Sikkim), fl. 1897, Hobson, s.n. (Type, K).

Cultivated: fl., fr. 1944.

(170) Berberis kartanica Ahrendt, J . Bot. Lond. 79 (Suppl.), 68, 1942 Stems; mature, yellow, subterete, not verruculose. IntemZ0de.s 10-17 mm. Spines 3-fld.,

concolorous, stout, plane, 15-25 mm. Leaves 13-29 x 6-12 mm., obovate, cuneate, sessile, entire, rounded, mucronulate ; branched venation distinct, reticulation fine ; below sub- concolorous green, lustrous. InJEMescem 5-7-fld., racemose-subumbellate, 1 . 8 3 cm., inchiding peduncle 7-14 mm. Pdicels glabrous, 5-10 mm. ; bracts, c . 2.5 mm. Outer sepals 3.5-4 x 1.5-2 mm. Inner sepals 5 - 5 4 x 3.25 mm., all oblong-elliptic. Petals 3.75 x 3 mm., broadly obovate; apex emarginate with acute lobes; base scarcely clawed, with. separate oblong-obovate glands, 0.8 x 0-5 mm. Stamens 2.8 mm., scarcely pro- duced. Ovules 4-5, often stipitate.

S.E. Tibet: Mago district, Karta, 14,OOOft., fl. 15 June 1935, Kingdon-Ward 11716 (Type, BM). 'In the arid valley above Karta, one of the first shrubs to appear, erect, with upright yellow flowers.' Not in cultivation.

(171) Berberis himalaica Ahrendt, J . Bot., Lond. 79 (Suppl.), 68, 1942 Stems subterete or subsulcate. Internodes 2-1; cm. Spines 3-fld., 7-11 mm. Leaves

1.3-38 x 0-5-1-5 em., obovate; cuneate to decurrent petiole, 3-7 mm., entire, obtuse, mucronulate ; venation branched, scarcely reticulate ; dull green ; below pruinose, paler, greenish grey. InJloTescence 34-fld., fascicled. Pedicels 1.8-2.5 cm. Prophylls 2, yellow, triangular, 2-5 x 1 mm. Outer sepals 2.75 x 1.5 mm., oblong-elliptic, rounded. Inner sepal8 4 x 3 mm., obovate, rounded. Petals 3 x 2.75 mm., obovate, cuneate, entire; glands contiguous. Stamens 2.75 mm., scarcely produced. Berries ovoid, estylose, 12 x 6 mm.

Bhutan (N.E.) : Me La, 12,500 ft., fr. 5 Oct. 1934, Ludlow & Sherriff 1013 (Type, BM). (E.), Singhi Dzong, 7500 ft., 27 Bept. 1949, Ludlow, Sherriff & Hicks, 21476 (BM); Guni Sa, 8 miles upstream of Drugye Dzong, 8500 ft., fl. 14 May 1949, Ludlow, Sherriff & Hicks 16229 (BM). Barshong, Thimbu Chu, 11,500 ft., fr. 15 Oct. 1949, Ludlow, Sherriff & Hicks 17524 (BM).

Cultivated: from numbers 17524, 21476; a rather leggy shrub, 4-6 ft. b h , growing in open scrub jungle and Pinm excelsa forest.

(172) Berberis hamiltoniana Ahrendt, Card. IUust. 64, 426, 1944 Stems glabrous, angled or sulcate; mature yellow, not verruculose. Internodes 1-2 cm.

Spini?s 3-fld., 6-15 mm. Leaves obovate, to 3 x 1.3 cm. ; sessile, entire, subacute; lustrous deep yellow-green, with few lateral veins; below grey, pruinose. Inflorescence SlO-fld., pseuclumbellate-subracemose, somet,imes with all, or part, fascicled, ( 1 . k ) 3 4 cm. long. Pedicels 5-9 mm., but 10-16 mm. in the fascicled parts; bracts 1.5-2 mm. F h e r s H.C.C. 2 to H.C.C. 601. Outer sepals 4-5 x 2-3 mm., oblong-ovate or prophyll-like, with a truncate base, acute. Median sepals 4.5-6 x 3-5 mm., obovate. Inner sepals 6.5- 8 x 4-43 mm., concave. Petals 56 .25 x 44 .25 mm., obovate; emarginate with acute

floris, owlis tantum 4-5, floribus fructibusque multo minoribus

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138 L. W. A. ~ E N D T

lobes; base cuneate, with sepmate, oblong, non-mmgid glands, 0.7 x 0.4 mm. Stamens 2.5-3.5 mm., apex smcely produced, subtruncate. Chndea 4-5. Be& very bright red, dry, somewhat crinkled, estylose, 9-10 x 6 mm.

Nepal: Jey-Jey, 12,500 ft., 1934, seed only, 19 Oct. ; Kmka Dam, fr. 13,000 ft., 14 Oct. 1935, Bailey 42 (BM); Mtmiandi, 11,5OOft., fl. 5 June 1950, hwndes L. 941 (BM); (Western), between Pudamigaon and Ringmigaon, 14,000ft., 22 Sept. 1952, P o l e , Sykes & Williams 3556 (BM). Chutta, S.E. of Jumla, lO,000 ft., 28 Sept. 1952, P o l e , Sykes & Williams 5480; Bmbmg Xhola, Pemringgaon, 14,000 ft., 7 June 1952, Polunin, Sykes & Williams 1100; Bhurchula Lekh, near Jumla, 12,50O,ft., 12 July 1952, Pol&, Sykes & Williams 4579; Sialgmhi, 9600 ft., 22 Bhy 1952, Polunin, Sykes & Williams 989 (all BM) .

Cultivated: L.A. 347 (from seed from Jey-Jey, Kew 1056/34), fl. 3 June 1942,lO June 1943; fr. 15 Oct. 1943 (Type, BM).

The OW collector gives the mtive name as Chutra.43, these seeds having been received with those of Prinaukc, G&iuna, and Me.mwp&. Later collectors describe it as growing on stony hillsides, dry s l o p where there is a little surfme water, among boulders on river banks, and generally on dry or stony s l o p . This erect, compact shrub, is attractive for its leaves, flowers large for this subsection, and peculiar, vivid berries.

(173) Berberis jaesebkeana Schneid., Bull. Herb. B&&r, (2), 5, 399, 1905; Parker,

Stem stout; young shoots green; mature at h t slightly reddish, soon yellow-brown; angled. Internodee 5-10 (-14) mm. 8p.w 3-fld., concolorous, sulcate, 10-15 mm. b v @ below paler; at h t greenish grey, slightly pruinose ; generally ultimately green, epruinose. InJEoIeseence 3-8-fld., umbellate to subumbellate, 2 4 c m . long, including peduncle 1-2-5 cm. Pedi& glabrous.

A Pore& Flora for the Punjab with Hau;cra and Delhi, 14,1918

KEY 3 2 A Stems at first puberulous; petals entire. Stems always glabrous; petals emarginate.

173a. B. jaeachkeam var. jmchkeancc

173b. B. jaeachkeana var. usteriana 1730. B. jmchkeam var. bhnbilaica

Pedicels 3-6 mm. long; berries epruinose. Pedicels 6-12 mm. ; berriw pruinose blue.

(173a) Berberis jaeschkeana var. jaeschkeana Leava 10-23 x 6-8 mm., oblong-elliptic; sometimes entire, mostly with margins with

(1-) 2-4 spinose (1 mm.) sedat ions distant, 4-6 mm. ; both sides openly veined and reticulate. P&i& 6 9 (-12) mm. Flowers H.C.C. 602/1. Outer sepals 4-4.5 x 3 - 5 4 mm., obovate-elliptic, subacute. Median sepals 5.5-6 x 4 4 . 5 mm., obovate, subobtuse. Inner sepal8 7-8 x 5-6 nun., obovate, concave. Petals 6.5 x 5.5 mm., obovate; base clawed, with concolorous approximate glands, 1.2 x 0-5 mm. 0wu.k~ 4-5 (-6). Stamem 4 mm., with apex slightly produced, truncate. Bewia red, oblong-ovoid, very shortly stylose, 9-10 x 5-6 mm. Seeds dark purple. Kashmir: Falconer 97 (Type, Vienna); Putakhan Cali, near Lolab, lO,OOOft., fr.

11 Oct. 1940, Ludlow & Shemiff 8265 (BM). Punjab Himalaya: Alundhar, Lahul, Kangra Thal, Jawchke, s.n. (K). Chamba State,

Pa@, Chota Brunbal reserve, 11,7OOft., fl. 18 July 1920, Parker 33 (K). ‘A shrub of 2 ft., in patches on dry rocky ridges, and only once seen in Pangi.’ This diverges in the following particulars: leaves to 30 mm., with 6-8 marginal spinules, shorter petals, and pedicels 12-18 mm.

Nepal: near Seng Khola, fr. 6 Oct. 1954, open ridge top, 11,000 ft., Stainton, Sykes &

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Berberis and Mahonia 139

Williams 4706 (BM); Rambrong, 5-6 ft., at 11,500 ft., 27 Oct. 1954, Stainton, Sykes & Williams 9353 (BM). These Pr’epal specimens have larger leaves, and a looser habit with longer internodes.

According to Parker, distributed from Kashmir to Kumaon, 900&13,500 ft., particu- 1arl;y in Kagan, Lahul and Kunawur, but not common. His Kumaon reference probably includes the variety usteriana.

(1736) Berberis jaeschkeana var. usteriana Schneid., 1.c. 1905; syn. B. usterianu (Schneid.) Parker, Indian Forester, 50, 399, 1924

Leaves 8-16 x 4-7 mm., with 1-2 marginal spinules. Prophylls lanceolate, acute, 1-2 x 0.5-1 mm. Outer se@ 4.5 x 3 mm., elliptic. Inner sepals 5-6 x 4-45 mm. Petals 4.5 x 3 mm., obtusely emarginate with obtuse lobes; base cuneate, with separate, non- marginal glands. Stantens slightly produced, retuse-truncate. BemieS oblong-obovoid, 9-12 x 6-7 mm. Style very short or absent.

Kumaon: Chaleb, Byans, Duthie 5307 (Type, K). Almora, 9250-10,000ft., 1923, Parker, 2075, 2078 (K). Schlich 27 (I), as ‘umbellata’ (0).

Kashmir: Mantnar Nullah, Bringhi valley, 12,000 ft., 18 Oct. 1940, Ludlow & Sherriff 8273 (BM). Sinthan pass, Bringhi valley, 12,000ft., 24 Oct. 1940, Ludlow & Sherriff 8283 (BM).

Cultivated: (Ludlow & S h e d 8265), fl., fr. 1944.

Nepal: east to east central. Thinigmn, Muktinath Himal, growing 4 ft. high among scattered shrubs, with under-

surface of leaf light green, 20 June 1954, Stainton, Sykea & Williams 1225 (BM). Above Sauwala Khola, 10,500 ft., shrub to 8 ft., in open forest with fruit orange red,

19 Sept. 1954, Stainton, Sykes & Williams 4376 (BM). Near Sirtibang Lekh, growing 5 ft. high, above forest limit, with fruit bright red, a t

11,OOO ft., 14 Oct. 1954, Stainton, Sykes & Williams 9024 (BM). Muktinath, 3 4 f t . , flowers yellow, at 12,500ft., 9 June 1954, Stainton, Sykes &

Williams 5694 (BM). Muktinath, growing 1-2 ft., on open stony slopes at 12,500 ft., 9 June 1954, Stainton,

Sykes & Williams 5705 (BM). Between Namche Bazar and Pangboche, 11,700 ft., 20 Oct. 1954, Zimmermann 1758

(Geneva). The Nepal specimens exhibit some variation. Numbers 5694, 5705 and 1225 have

small (1-2 cm.) leaves, entire or with 1-2 spinules each side. Number 9024 has larger ( 2 3 . 5 x 0.8-1-5 cm.) leaves with 5-10 very inconspicuous h e spinules each side, but entire on the shoots. Number 4376 is similar with still larger (5 x 2 cm.) leaves. The inflorescence is short (1-2-5 cm.) in the first three but longer in the last two (2-4 cm.). The 6rst three seem the usual compact dense shrubs. The last two of looser habit with longer internodes (1.5-26 cm.) and stouter spines to 1.5 cm. All have short pedicels, 3-7 (-9) mm. long and leaves which (at any rate finally) are clearly green below; the latter of these two characters distinguishes these from the Nepal B. hurniltonianu, and both separate them from the Bhutan B. himuhim. However, further investigation is necessary amongst these three species. Zimmermann 1758 has large (13 x 7 mm.) berries.

(173c) Berberis jaeschkeana var. bimbilaica Ahrendt, J . Bot., M . 79 (Suppl.), 65, 1941

Stems sparsely verruculose. Leaves 10-28 x 3-7 mm., oblanceolate, to narrowly obov:tte; acute to subacute; entire or with margins with 1 4 spinose (1-1-5 mm.) serru- lations; more persistently greyish pruinose below. outer sepals 3.- x 1-5-2 mm. obloIig- ovate, acuminate. Nedian se@ 4-5-5 x 3-3-5 mm., obovate. Inner sepals 5-6 x 4 mm., obovate. Petals 4-5 x 3 mm., apex acutely emarginate with acute lobes; glands separate.

Cultivated: (Ludlow & Sherriff 8273).

Page 140: Berberis and Mahonia : A taxonomic revision

140 L. W. A. AHRENDT Stamem scarcely produced, truncate. Berries oblong-ovoid, 9-10 x 4-46 mm., with short style.

S.E. Tibet: Tsari district, Bimbi La, 11,000-13,000ft., fr. 13 Oct. 1938, Ludlow, Sherri€€ & Taylor 6305 (Type, BM). ‘Scattered over hilleides, brilIiant autumn colourer.’

Cultivated: L.A. 274 (seed of type), fl. 5 May 1944, 16 Apr. 1945. A shrub of 1-3 ft. Three hybrids between diverse sections fall here on account of their combined

characters, but they have no connexion with the preceding species:

(174) Berberis x vilmorinii Schneid., J. Arn. Arb., 4, 231, 1923 Stem glabrous, sulcate; young shoots red; mature yellow. Spines 3-fld., sulcate, 10-

23 mm. Leaves to 5 x 3 cm., oblong-elliptic; base c o n t r d to petiole 3-7 mm. ; margins with 6-12 spinose serrations (1-2.5 mm.), openly reticulate. In,escence unrecorded. Pedicekr 1-2 cm. Inner sepal8 8 mm. Petals 7 mm., emargiuak. 8tamen.s apiculate. Ovdm (6) 7-8, seaaile. Berries dark red, 11 x 5 mm., estylose.

Origin: cultivated at Les B m s , 1008Y, seed planted 30 Mar. 1905; fl. 10 May 1908; fr. 21 Sept. 1910 (Type, Am. Arb.). Reputed B. diuphana x B. p i m a , in which cwe the flowers should be fascicled. Not cultivated in England.

(1744 Berberis x vanfleetii Schneid., 1.c. 230, 1923 Stem subterete, yellow. Spines 3-fld., sulcate, to 2.5 cm. h u e s elliptic to obovate,

acute; petiole 1-5 mm.; margin subsinuate with 8-12 spinose serrations (1-1.5 mm.), openly reticulate deep green, grey pruinose below. In.eswnce 4-9-fld., umbellate- racemose, with added basal flowers, 2.5 cm. long. Pedkeb 5-10 mm. ; bracts 1-16 mm. Berries ellipsoid, estylose, dark red, scarcely pruinose, 8-lox 6-7 mm. Seeds dark purple.

Origin: B. veitchii x B. vulgaris (Type, Am. Arb.). Not cultivated in England.

(175) Berberis x regleriana Notcutt, Catalogue of Trees and Shrubs, p. 25, 1938 Shrub 5-8 ft. Stemwith young shoots bright red; mature yellow. Spines 4-8 mm. (but on

shoots 1-1-5 cm.), subterete 1-3-fld. Leaves obovate, to 5 x 2.4cm. ; base contracted to short petiole; margins with 15-25 spinose (1.5-2 mm.), sedat ions (0.5 mm.), distant 1-2 mm. (but larger and subentire on shoots), dull, below grey. InJlorescence 6-10-fld., 2 4 cm., pseudumbellate-subracemose. Pedicds 7-15 mm.; bracts 1-2 mm. E”l0Zuers H.C.C. 601. ProphyUs red, ovate, acute, 1.5 x 1 mm. Outer sepals 3.5 x 2.5 mm., oblong-elliptic, rounded. Inner sepals 5-6 x 3-4 mm., obovate. Petals 5-5-6 x 3-54 mm., entire; base cuneate, with sepmab, non-marginal glands, 0.9 x 0.7 mm. Stamens scarcely produced, rounded. Ovules (3-) 4. B e d 7-8 x 3-4 mm., narrowly oblong-obovoid, red, epruinose,

Origin: raised by R. C. Notcutt, Woodbridge, Suffolk, from seed of B. vulgaris (prob- ably misnamed B. regeliana, and gathered at Wisley), and presumably a hybrid with a species of Sect. Angul0sa.e.

estylose.

Cultivated: L.A. 78-4. 25 Apr. 1939, 16 May 1941; fr. Oct. 1938, (Type, 0).

Subsect. Subangulosae Ahrendt; syn. Subsect. Yunnanema Ahrendt, J. Bot., Lond.

Stems always dark, red or purple; when mature, glabrous and sulcate (except only in B. trichohmmatoi&s, where they are pubescent and subterete). Leaves deciduous, up to 18-43 x 6-15 mm.; margins mostly with 1-7 (-9) spinules (there being more teeth only in B. Orthobotrys, and the leaves being entire only in B. t r i c h k m , B. johnnk and B. platyphyh) ; .below mostly pruinose grey, sometimes epruinose green; venation mostly slight or branched, occasionally openly reticulate. Injhescew 1-4 cm. long,

79 (Suppl.), 58, 1941, excluding only B. yunnanensiS and B. Veracem

Page 141: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 141

2-8 (-14)-fld., fascicled, umbellate or pseudumbellate-subracemose. Pedicels glabrous, 4-20 mm., with much shorter bracts (1-2, rarely to 4 mm.). Inner se@ (51) 6-8 (-10) mm. Petals always shorter than inner sepals. Stamens mostly not produced, some- times much produced, apex retuse or truncate or obtuse (apiculate only in B. platy- phylla). Ovules (3-) 4 4 (but 6-8 only in B. suberecta). Berries oblong, 10-12 (-17) mm. (but globose, 7 mm., only in B. mope&), estylose (except only in B. suberecta where the style is very short).

Afghanistan, Sinkiang, Kaahmir, Punjab Himalaya, Nepal, Sikkim, Bhutan, S.E. Tibet (Provinces U, Zayul, Kongbo, Pome), W. Yunnan.

K.EY 33, to subsect. Subangulosae (Maps 17 and 21) Flowers 7-12 (-14) mm. diameter; fruit 7-12 (-14) mm. long.

E’ruit estylose; ovules 3-5. Stems pubescent, subterete. 176. B. trichohaematoides Stems glabrous, sulcate.

177. B. johannis

Fruit lustrous, tkn, globose, 7 mm. 178. B. cooperi Fruit dull, softer, oblong, 1&12 mm. long.

Leaves spinose; stamens not apiculate.

Fruit bent, narrow, c. 12 x 3 mm. Fruit broader, globose, or 10 x 5-7 to 12 x 8 mm.

Leaves green, epruinose below. 179. B. orthobotrys Leaves grey, pruinose, below.

Leaves narrow (1 : 3-4), with 8-15 marginal spinules, reticulate; petals

Leaves broader (1 : 2-3), with %S marginal spinules, scarcely reticulate;

Leaves to 23 x 11 mrn.; stamens not produced, truncate; fruit

Leaves to 33 x 11 mm.; stamens produced, obtuse; fruit oblong to

entire. 179 f. B. orthobotrys vm. caneacens

petals emarginate.

obovoid, 10 x 7 rnm. 180. B. ambiguu

13x7rnm. 181. B. ignorata Leaves entire ; stamens apicdate. 182. B. platyphylla

Fruit shortly stylose; ovules 6-8. 183. B. auberecta 184. B. rubrostilla Flowers 16-18 rnm. diameter; fruit 15-18 nun. long.

(178) Berberis trichohaematoides Ahrendt, J . Bot., Lond. 79 (Suppl.), 62, 1941 I,nternodes 1-5-3 cm. Spines absent, or 1-3-fld., 5-10 mm. Leaves 1.63 .5 x 0.6-

1.5 em.; cuneate, sessile or with petiole 1-2 mm.; entire, obtuse or rounded; venation indistinctly branched, not reticulate; below pruinose grey. InJlorescence 5-10-fld. (fascicled, rarely fascicled-subracemose, 2-3 cm., including glabrous peduncle, 2-7 mm.). Pedicels glabrous, slender, 12-17 mm.; bracts acuminate, 1.5-2 mm. Berries elliptic- obovoid, 12-13 x 4-4.5 mm. Seeds pale yellow-brown.

S.E. Tibet: Prov. Kongbo, Kulu Phu Chu, near Paka, 12,OOOft., fr. 22 Sept. 1938, Ludlow, SherrifT & Taylor 6513 (Type, BM). Fruit bright scarlet. In clearings in Abies forest.

Cultivated : from type collection.

(177) Berberis johannis Ahrendt, Curd. Chron. (3), 109, 101, Mar. 1941; J . Bot., Lond. 79 (Suppl.), 66, 1942; Sealy, Bot. Mag., 166, N.S., t. 57, 1949. (Fig. 34)

Internodes (6) 10-15 (-20) mm. Spines 3-fld., slender, plane, 7-14 111111. Leaves 15-20 x 8-11 mm., base con- tracted to petiole 1-7 mm.; obovate, margins entire (rarely 2-5 spinules); acute or sub- acute; subenervate or with few lateral veins a t most indistinctly openly veined; dull

Stems slender, shoots green, not or scarcely verruculose.

Page 142: Berberis and Mahonia : A taxonomic revision

142 L. W. A. AERENDT green; below grey, pruinose. In ,e scenw 3-7-fld., umbellate, 2-3 cm., including slender red peduncle 3-12 mm., or subracemose, 2-54 cm. Pedicels red, (P) 6-9 (-10) mm., below occasionally 12-17 mm. ; bracts 0.5 mm. Rowers H.C.C. 112, 7-8 mm. diameter. Outer sepals yellow, 2-5 x 1.5 mm., oblong-triangular, acute. Median sepals 4 x 2 mm., oblong-ovate, subobtuse. Inner sepal8 5 x 2.5 mm., elliptic. €'eta% 4.5 x 3 mm., apex acutely emarginate with remote mute lobes; basal glands approximate, submarginal, obovate, 0-8 x 0.4 mm., situated above short claw. Stamens 2.5 mm., scarcely produced, truncate. Ovdes stipitate with atipe aa long aa ovule. Bem'es bright red, namow, bent, oblong-ellipticoroblong-ovoid, 11-12.5 x 3-3-75 mm., or 13 x 3.5 mm., rarely 12 x 4.5 mm.

S.E. Tibet: Prov. Pome, Tumbatse, 11,OOO-12,OOO ft., 13 July 1924, Kingdon-Ward 5936; Sip valley nem Tongguk Dzong, 9500ft., 21 May 1947 Ludlow S h e M & Elliot 13731 (BM); Nambu La, Tongyuk river, 11,500 ft., 3 June 1947, Ludlow, Sherriff & Elliot 13832 (BM).

Cultivated: L.A. 252, from seed of Kingdon-Ward 5936; fl. 4 June 1942,4 May 1944; fr. 20 Oct. 1944 (Tylpe, BM). Erroneously distributed aa Kingdon-Ward 5962 (which is B. gyalakcu), see Report of R.H.S. Conference of Trees and Shrubs, 1938. Described as from 3 to 6 ft. high, growing in thickets, open jungle, or conifer forest.

(178) Berberis cooperi Ahrendt, Card. Chron. (3) 109, Mar. 1941; J. Bot., Lond. 79 (Suppl.), 61, 1941. (%. 35)

Intemzodes 1-2-2cm. S p i w 3-fld., P9mm. Leaves 15-35x7-15mm.; oblong- obovate; cuneate to decurrent petiole 2-5 mm.; margins with 6-10 spinose (1-2 mm.) serrulatiom, distant 3-6 mm.; venation branched, not reticulate; lustrous deep green; below at fkt shghtly greyish pruinose, hally lustrous green. InflM.escence fascicled, or partly umbellate, 5-10-fld. Ptdkls 10-18 mm.; bracts 1 mm. O u t e r s e p h 3.5 x 2.5 mm., ovate, obtuse. Median se@ 6 x 4 mm., oblong-elliptic. Inner sepals 7 x 4-5 mm., obovate-elliptic. Petals 5.5 x 4 mm., not produced, truncate. B e d red, lustrous h, globose, 7 mm.

Bhutan: 1914, seed, Cooper 2979. Cultivated: LA. 491, from seed of Cooper 2979, fl. June 1938; fr. 26 Oct. 1938 (both

Kew 74) (Type, BM); fr. 9 Oct. 1940 (E). Shrub 5-7 ft.

(179) Berberis orthobotrys Bienert ex Aitch., J. Linn. Soc. Bot. 19, 150, 1882;

Stems glabrous, mature dark red. Leaw spinose. Infl.rescence racemose, umbellate- racemose, or racemose-subfaacicled, with (%) 5-10 (-12) flowers. Pedkls 6 1 4 mm. O& 3-5. Berries estylose, red, 7-10 x 5-6 mm.

Schneid., Bull. Herb. Boissier (2), 5, 662, 1905

KEY 33A Leaves broad (1 : 1-5-2), below green, and epruinose.

Stamens with apex not, or warmly, produced; intloreacence subumbehte, 5-12-fld. Pdcels slender, 8-14 mm.

tam ens shorter than petals; fruit 9-10 mm. long.

Stamens longer than petals; fruit 7 mm. long. 179a. B. mtlwbotrys var. orthobotrya

179b. B. orthobotrys var. m w a y i Pedicels stout, 4-8 mm. 179c. B. orthobotrye vm. sinthamnsis

179d. B. orthobotrya vm. rwpmtria S-ens with epex much produced; pedicels stout, 8-10 mm.; flowers faecicled.

Leaves m o w (1 : 3-4); stamens conspicuously produced, obtuse. Leaves to 18 x 6 mm., green, eprukose below; lDarginal spinules 1-4 in number;

venation branched; inflorescence 2-4-fld. ; pedicels 6-8 mxn. 179e. B. orthobotrys var. ru&u&

Page 143: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 143

Leaves to 25 x 8 mm., grey, pruinose below, with 5-15 marginal spindes; venation reticulate; inflorescence 5-10-fld. ; pedicels 6-12 mm.

179f. B. orthobotrys var. canescens

(1794 Berberis orthobotrys var. orthobotrys hlhrub G. 4 ft. Internodes 1-2 cm. Spines 3-fld., orange-yellow, 10-15 mm. Leaves

(1-1 1.7-2.5 (-3) x (04%) 0.7-1.2 (-1.7) cm., including petiole 2-5 mm.; obovate, with contracted base; margins with 5-13 (-20) spinose (0.5-1 mm.)-serrulations, distant 2-4 (-7) mm., but on the new shoots sometimes entire; fairly closely reticulate; both aides concolorous green. Injiorescence 5-12-fld., racemose, racemose-subfascicled, or racemose-subumbellate, 2-3.5 cm. long. Pedkls 5-10 (-12) mm. Inner sepals 6 x 5 mm. Petals 5-75 x 3.5 mm., emarginate. Stanzens scarcely produced. Berries 9-10 x 5-6 mm., ddl red, oblong, estylose. Kashmir: 5500 ft., Thomson, s.n. (K); Bhujaz, foot of Umasi La, l1,oO ft., 14 July

1943, Ludlow & Sherrif€ 9190 (BM), shrub 2-3 ft., on grassy slopes. Afghanistan: Kurrum valley, Hills above Kaiwas, 11,OOO ft., fl. 3 July 1879, Aitchison

726 (K); Sikaram, June 1879, Aitchison 499 (Types). Nepal: Rambrong, Lamjung Himal, at 12,OOOft., 29 June 1954, Stainton, Sykes &

Williams 6049 (BM). A shrub of 6-8 ft., which seems to belong here; its mostly entire leaves, with only branched venation, may be of new shoots.

Cultivated: a t Kew.

(179 b) Berberis orthobotrys var. conwayi Ahrendt, var.nov., staminibus con-

Leaves to 3.5 x 1.9 cm. In$orescence 4-8-fld., 1 - 5 4 cm. long. Prophylls yellow, sub- acurninate, 25-1-25 mm. Outer sepals 4 x 2.25 mm., ovate, acute. Inner sepals 6-7 x 3.5 mm., oblong-obovate. Petals 5 x 2.5 mm., convolute. Stumm with apex scarcely produced, but 5 mm. long and protruding beyond the convolute petals. Berries 7 x 5 mm.

Sinkiang: North Kashmir border, Karakoram Mts., c. 36"N., 76"E., south side of Rash Pass, 15,000 ft., fl., fr. 1892, Conway 146 (Type, K). Not in cultivation.

spicuis petalis longioribus, fructibus brevioribus distinguenda

(1794 Berberis orthobotrys var. sinthanensis Ahrendt, var.nov., pedicellis

Apart from the stout pedicels only 4 4 mm. long, this is similar to the variety ortho-

Kashmir: Sinthan pass, head of Bringhi valley, 10,OOO ft., fr. 24 Oct. 1940, Ludlow &

Cultivated: from the type collection.

validis, brevioribus, distinguenda

botrys.

Shemiff 8279 (Type, BM).

(179 d ) Berberis orthobotrys var. rupestris Ahrendt, var.nov., staminibus elon-

Intwnodes 2.5-3.5 cm. Spines weak, to 1 cm. Leaves obovate, to 35 x 18 mm.; margins with 10-15 spinose (1.5 mm.) serrations (1-5 mm.), distant 1.5-3 mm. (although less spinose to subentire on the shoots) ; venation slightly branched; below slightly paler green. Pedicels stout, 8-10 mm.; bracts 4 mm. Outer sepals 7 x 3.5 mm., obovate, acute. Inruzr sepals 8 x 5.25 mm., obovate, rounded. Petals 6.5 x 3-75 mm., obovate, subretuse; glands obovate, bilobed, separate, non-marginal, 2 x 0-6 nun. Stamens 5 mm., con- spicuously produced, apex retuse.

S.E. Tibet: Prov. Zayul, Delei valley, 1ll000-12,000ft., 21 June 1928, Kingdon- Ward 8350 (Type, BM). Small shrub in gneiss cliffs in the more open parts of the Abies forest, and (4 July 1928) in the alpine region amongst dwarf rhododendrons. Not in

gatis, pedicellis validis, floribus fasciculatis tantum 3-5, distinguenda

Page 144: Berberis and Mahonia : A taxonomic revision

144 L. W. A. AERENDT cultivation. The seed bearing this KW number seems to have produced a very different species, B. parkepakz, with pubeacent yellow stems, entire leaves, larger solitary flowers, and truncate stamens not produced.

(1794 Berberis orthobotrys var. rubicunda Ahrendt, var.nov., foliis minoribus angustioribus, margine spin& tastum 1 4 , inflorescentiis tantum 24-floris,

Illternodes 1-14 cm. Spines 3-fld., 8-12 mm., plane below. Leaves narrowly obovate, to 18-20 x 5-6 mm.; base cunwte, margins with 1-4 spinose (1 mm.) serrulations (0.5 mm.) on the apex half; venation loosely branched; above dark green, paler below. In,escence, (1-) M-fld., umbellate-racemose 1-5-2 cm. Pedicek red, slender, 5-7 (-9) mm. Outer sepals 4 x 2.5 mm., ovate, acute. Inner 8epaJ.s 6 x 5.5 mm., broadly obovate. Pet& 5 x 3-75 mm., broadly obovate. StamRns 4-25 mm., much produced, apex obtuse.

Central Nepal: La&ang, on cliff sides above, 11,500-12,500 ft., Polunin 514, Type, and 483 (both BM). Not cultivated.

Nepal: Lete, south of Tukucha, 6ft., amongst scattered shrubs, 1OY500ft., 6 June 1954, Stainton, Sykea & Williams 975 (BM). Lete, KaL Gandaki valley 3-4ft., with flowers bright yellow, at 11,500 ft., 4 June 1954, Stainton, Sykes & Williams 5594 (BM). Rambrong, Lamjung Himal, shrub 8-10 ft., with flowers bright yellow, at 10,500 ft., 27 June 1954, Stainton, Sykea & Williams 5961 (BM). These three specimens have their dark red stems pubescent in paxts.

(179f) Berberis orthobotrys var. canescens Ahrendt, var.nov., f o b angustioribus, subtus canescentibus, staminibus valde productis distinguenda

Spines concolorow, sulcate, 1-26 cm. Leaves 10-25 x 3-8 (-10) mm. ; narrowly obovate; margins with (5) 8-12 (-15) spinose (1-2 mm.) serrulations, distant 2-3 mm., but entire on the new shoots; reticulate; dull grey-green; below grey, pruinose. In- Jlorescence 5-1O-fld., umbellate-racemose, 1 . 5 3 cm., including peduncle 1-5 mm. Pedicels 6-12 mm. Flowers H.C.C. 3. Outer sepals 3-6-4~ 1.3 mm., lanceolate, acumi- nate, yellow. Median se@ 6 x 3.5 mm., elliptic-obovate. Inner sepaLs 7 x 4.5 mm., obovate. Petals 7 x 4 mm., obovate, obtuse, entire; base scarcely clawed, with lanceolate glands, 0.8 x 0.3 mm. Stamens 4-46 mm., much produced, obtusely subconical. Berries red, 4-10 x 5-6 mm.

pedicellis 5-8 mm., distinguenda

Kashmir: seed received and raised by W. J. Marchant, Dorset. Nepal: Samargaon, north of Tukucha, Kali Gandaki, 4 ft., on open ground at edge of

hills at 13,000 ft., fr. 10 Oct. 1954, Stahton, Sykes t Williams 8100 (BM). This specimen has large berries, 14-16 x 5-6 mm.

Cultivated: LA. 390 (from above mentioned seed); fl. May 1940; fr. 13 Oct. 1940, 6 Nov. 1941,30 Sept. 1942. Distributed as ‘B. orthobotrys’, and also as Berberis unique. A valuable compact shrub for berry.

(180) Berberis ambigua Ahrendt, J. Bot., Lond. 79 (Suppl.), 60, 1941 Stems not verruculose. Inte?-nodes 15-20 mm. Spines 3-fld., 5-10 mm. Leaves 10-23

(-27) x 5-12 mm.; obovate, cuneate, sessile; margins with 6-9 spinose (1 mm.) s e m - lations, distant 3-6 mm.; apex rounded; openly reticulate; below grey, pruinose. In- Jlorescence 2-3-3d.. fmcicled. Pedicela 10-18 mm. ; bracts lanceolate, acuminate, 4 x 1-25 mm. Prophylb concolorous, 3.5 x 1.75 mm. Outer and inner sepals both 6 x 4 mm., obovate. Petals 5 x 3 mm., obovate-elliptic, shortly incised; base with widely separate glands, 0.7 x 0-3 mm. Stamens 4 mm., scarcely produced, truncate. Berries dull red, obovoid, estylose, 9-10 x 6-76 mm.

Yunnan: Tali Range, seed ody, 1917-19, Forrest, 8.n.

Page 145: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 145

Cultivated: L.A. 210, from above seed; examined in fl., and fr., 1938. The Yunnan seed was labelled ‘ B . dictyophylla, Tali Range’. Some packets produced this species, and others B. suberectu.

(181) Berberis ignorata Schneid., Bull. Herb. Boissier, (2), 5,661,1905; syn. B. virescens var. ignorata (Schneid.) Ahrendt, J . Bot., Lond., 79 (Suppl.), 60, 1941

Spines solitary, sometimes 3-fld., 7-13 mm. Leaves 14-32 x 4-11 mm. ; narrowly obovate; base contracted and then cuneate to short decurrent petiole; margins with 3-8 spinose (1-2 mm.) serrulations, distant, 2-4 mm. ; venation branched, scarcely reticulate ; dull green, below grey, pruinose. Inflorescence (3-) 5-8 (-10)-flowered, fascicled or race- mose-subfascicled, 1-5-3 cm. Pedicels (&) 9-13 (-15) mm.; bracts, 1.5 mm. Outer sepals 4-44 x 2-2.5 mm., oblong-ovate, obtuse. Median sepals 5 x 2.5 mm., elliptic, obtuse. Inmr sepals 5.25-6 x 3.75-4 mm., elliptic-obovate. Petals 4 - 6 5 x 3 mm., obovate; shortly emarginate with rounded lobes; glands 0.9 x 0.6 mm., separate, non-marginal. StanMns 4 mm., produced, obtuse. Fruit soft, oblong, 10-13 x 5-7 mm. Seeds pale yellow.

Sikkim: Lachen valley, 9750 ft., 4 June 1849, Hooker, s.n. (Type, K). S.E. Tibet: Prov. U, Tsela Tzong, 10,000 ft., 31 May 1924, Kingdon-Ward 5724 (BM);

Tsangpo valley below Tsetang, 1O,OOO-11,000 ft., 29 Apr. 1924, Kingdon-Ward 5632, 5633 (K); Tsetang, 3 May 1938, Ludlow, S h e d & Taylor, 4110 (BM); has , north of Lhama, 13,500 ft., 6 June 1943, Ludlow & S h e d 9556 (BM); Lhasa and surrounding hills. and 40 miles north of Lhasa, up to 15,000 ft., Apr.-June 1939, Richardson 4 (BM).

Prov. Kongbo, Lotu, Shoga Chu, 12,000 ft., 15 June 1947, Ludlow, S h e d & Elliot 13893 (BM) ; Kamchang, Tsangpo valley, 10,000 ft., 28 Apr. 1947, Ludlow, S h e d t Elliot 12414 (BM).

Cultivated: L.A. 462, from seed of KW 5724. The cultivated plant was erroneously distributed as KW 5734. The species is described

as having bright, or golden, yellow flowers, as being a big bush, growing in thickets, on shaded slopes, or by streams, and as being common among other shrubs by streams in the dry Tsangpo valley. Its sulcate stems, bright yellow flowers and oblong red estylose berries distinguish it from two other species with which it has been confused, namely B. vi:rescens and B. paravirescens; these both possess terete stems and pale greenish flowers; the former is further distinguished by its narrowly ovoid red stylose berries, and the latter by its oblong black estylose fruit.

(182) Berberis platyphylla (Ahrendt) Ahrendt, stat.nov. ; s.yn. B. yunnanensis var. platyphylla Ahrendt, J . Bot., Lond., 79 (Suppl.), 61, 1941

Internodes 2-6 cm. Spines yellow, 1-3-fld., 5-15 mm. Leaves 25-43 x 15-27 mm., broadly obovate ; contracted, subsessile ; entire ; rounded ; venation distinctly branched, scarcely reticulate ; below grey, pruinose. Inflorescence 3-7-fld., fascicled or partly sub- umbellate, 16-26 cm. Pedicels (7 mm.-) 1-1-5 (-2) cm. Outer sepals 6 x 2 mm., ovate, acute. Inner sepals 7.5 x 4 mm., oblong-obovate. Petals 6 x 3 mm., obovate; acutely emarginate, with acute lobes ; base scarcely clawed, with oblong, separate, non-marginal glands, 1.3 x 0.4 mm. Stamns 5 mm., much produced, conspicuously apiculate. Berries oblong, 10 x 7 mm.

N.W. Yunnan: Mekong-Salween Divide, Sewalongba, 10,500 ft., fr. 31 Aug. 1938, Yu 22618 (Type, E).

S.W. Szechuan: Yetsi Mts., N. of Kulu, Muli territory, 11,000ft., May 1932, Rock 23911 (BM, K). Not cultivated.

This species is described as a shrub 3 4 ft. high, common, growing in pine forest or under mixed forest.

10 JOUBN. LINN. S0C.-BOTANY, VOL. LVII

Page 146: Berberis and Mahonia : A taxonomic revision

146 L. W. A. ABRENDT

(183) Berberis suberecta Ahrendt, Kew Bull. 1939, 271, 1939; J. Bot., Lvnd., 79

Shoots green, scarcely verruculose. Intermda 1-4-2-2 cm. Spines orange-yellow, sul- cate, 3-fld., 15-28 mm. Leaves 12-27 (-32) x 7-10 (-12) mm., oblong-obovate; base cuneate 1-2 mm. to decurrent petiole, 1-5 mm. ; margins with 2-5 (-7) spinose (1-2 mm.) serrations (1-2 mm.), distant 3-4 (4) mm. ; indistinctly veined; dull bright green; below grey, pruinose. Injeorescelace (3-) 5-6 (+)-ad., pseudumbellate-subracemose, 2-3 (4) cm., including peduncle (2-) 6-10 mm. Pedicek 4-6 (-10) mm.; bracts ovate, acute, red, 1-1 x 0.9 mm. Flower8 H.C.C. 2. Outer sepds 26-36 x 1-5-2 mm., ovate, acute. Median sepals 6-7 x 3.54 mm., oblong-obovate. Inner s e p k 7-8 x 5-6 mm. P& 4.5-6 x 2.5- 3-5 mm., obovate, subobtusely emarginate with obtuse lobes; base clawed, with oblong, separate, subconcolornus glands, 0-5-0-9 x 0.3-0-6 mm., close to margin and situated above the claw. Stamem 4mm., scarcely produced, apex rounded. ovules 6-8, sub- seasile. Berries dull, oblong-ovoid, 10-12 x 7-8 mm., short but distinct style.

(Suppl.), 57, 1941

Yunnan: Tdi Range, seed only, Forrest, s.n., 1917-19. Cultivated: L.A. 50, from above seed, fl. 15 June 1938,25 June 1942; fr. 11 Oct. 1938

( T p , 0, BM). Distributed aa ‘the upright form of B. rubrostiua’. See note under B. ambigzra.

(183~) Berberis x ‘Emperor ’ Chittenden, n.n. Stem pubemlous. S p i w 8-13 mm., yellow, sulcate. Leaves to 25 x 16 mm., margins

with 2-7 spinose (1 mm.) s e d t i o n s (but entire on shoots); reticulate, indistinctly above, elevated below; bright green; below pruinose grey. InJ~oreecenee 3-6-fld., race- mose-subfascided, 1-2 cm. Pediceh 4 4 (-8, in fruit) mm. Outer sepals 3 x 2.5 mm., ovate, acute. Median 8epab 5.5 x 4 mm., ovate-elliptic, rounded. Inner sepals 6-5 x 4*5mm., obovate. Pet& 5-25x2-25mm., obovate, retuse with acute lobes; base cuneate with obovate glands, 1 x 0.7 mm. S m 3-5 mm., produced, truncate. ovules 3-4. Berries dull, oblong-ovoid, 10-12 x 9-10 mm. Styk short, distinct.

Cultivated: fl. 12 July 1938; fc. Nov. 1938 (Type, Wisley).

(184) Berberis ( ? x ) rubrostilla Chittenden, J. R. Hort. Soc. 42,353, t. 65, 1917 8 h d 3-4ft. Leaves 10-2Ox4-6mm.; oblanceohte; margins with 4-6 spinules;

reticulate; below grey, p d o s e . Injeorescence 2-4-fld., umbellate-racemose, 2-2-5 cm., including peduncle, c. 5 mm. Pediceh 4-7 mm. Berries ovoid, 15 x 9 mm., estylose.

Cultivated: origin at Wisley.

KEY 33B Fruit bright red, 15-18 mm.; petals entire.

Peduncle 2-6 mm.; fruit 15 x 9 mm. 184. B. x &08t&Z

Peduncle 16-30 mm.; h i t 16-18 x 6-6 mm. Fruit dark purple, 10-11 mm.; petals emarginate.

1840. B. x rubrmtillu var. crawleye& 184b. B. x rubrostiua var. chealii

(184~) Berberis x rubroatilla var. crawleyensis Ahrendt, J. R. Hmt. Xoc. 67, 133, 1942

Leaves 20-40 x 5-10 mm., attenuate, sessile, oblanceolate; rnazgins 4-8-spinules (1-1-5 mm.), and serrations (1.5 mm.); reticulate. In@es~nce 5-7-fld., umbellate or subumbellate, 3-6cm., including peduncle, 1.5-3cm. Pedicab P7mm. F h e r s H.C.C. 2/1. Prophylls 2-4 x 1 mm., concolorous, oblong, acumimte. Outer sepals 5-7 x 3-3.5 mm., oblong-ovate, subacute. Median and inner mpds 9-10 x 5-6 mm., obovate, rounded, entire; base cuneate, with separate, lanceolate, non-marginal glands, 1 . 5 ~

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Berberis and Mahonia 147

0.5 mm. Stanzens 3.5 mm., scarcely produced, truncate. Ovules (3-) 4.5. Berries lustrous, oblong-ovoid, estylose, 15-17 x 5-6 mm.

Origin: at Cheal’s Nursery, Lowfield Heath, Crawley, Sussex. Cultivated: L.A. 37; fl. 14 Aug. 1938, 17 June 1942; fr. 12 Oct. 1938 ( T y p 0, BM).

One of the finest Berberis for individual berry.

(184b) Berberis x rubrostilla var. chealii Cheal, Catalogue of Shrubs, 18, 1937 Outer sepals 6 x 5 mm. Median sepals 6-5mm. Inner sepals 7.5mm. Petals 5 x

3.75 mm., emarginate with acute lobes; basal glands, 0.7 x 0.6 mm., obovate, separate. Stamens 2.5 mm., produced, conical-apiculate. Berries purple, oblong-ovoid, estylose, 10-11 x 8-9 mm.

Origin: Cheal’s Nursery, Lowfield Heath, Crawley, Sussex. Cultivated: fl. 23 Aug. 1939, June 1943; fr. 23 Dec. 1943,12 Dec. 1944 ( T y p , 0, BM).

(185) Berberisx ‘Cherry Ripe’ A. R. Ahrendt, R.H.S. Dict. Card. 1, 269, 1951 Bhrub 5-6 ft. Stem sulcate; young shoots puberulous, green; mature dark red, closely

and Gnely black-verruculose. Internodes 1-2.5 cm. Sphe.~ brown, sulcate, 1-3-fld., 15-25 mm. Leaves 16-28 x 8-12 mm., obovate, contracted, subsessile, entire, obtuse; above dull bright green, indistinctly reticulate ; below grey, pruinose, closely reticulate. InJlores- cena 2-8-fld., subumbellate, 4-10 cm., including peduncle 1-6 cm. (on older plants some- times subpaniculate). Pedicels 2-5 mm.; bracts, 1-1.5 mm. Flowers H.C.C. 3/1. Pro- phylls red, oblong, acute, 2 x 1 mm. Outer sepals 3.5 x 2-5 mm., ovate, acute. Median s e p L 5 x 4 mm., elliptic. Inner sepals 6.5 x 4.5 mm., obovate. Petals 5 x 3.5 mm., obovate, entire; glands discolorous orange, separate, submarginal, 043 x 0.8 mm. Stamens 3 mm., not produced, truncate. ovules 4-5. Berriea lustrous, at first creamy white, becoming partly light red, and then cherry red, estylose, broadly ovoid, 8-9 x 6-7 mm.

Origin: at Stonefield, Watlington, Oxford, from seeds of B. rubrostih var. erawley- ensis, sown by my mother on 21 Oct. 1932.

Cultivated: L.A. 156A, fl. 23 July 1938, 17 June 1942; fr. Dec. 1938, 15 Nov. 1944 (Type, 0). Named by my mother and generay commended aa one of the finest hybrid Beriieris for gardens.

Subsect. Consimiles Ahrendt, J. Bot., Lond., 79 (Suppl.), 70, 1942 Stem glabrous. Internodm 1 . 2 4 cm. S p i w sulcate. Leavu deciduous, grey, pdnose

below. Injloreacence (2-) 4-8 (-10)-fld., umbellate, or pseudumbellate-subracernose, 1-5-6 cm. Pedicels glabrous, (6) 6-10 (-12) mm.; bracts, 1-1.5 mm. Ovules (3-) 4-5 (-6). Be& epruinose, purple, 8-11 x 4-6 mm., estylose or with very short style.

Bhutan, Sikkim, W. Szechuan, Shensi.

KEY 34 to Subsect. Consimiles (Maps 17 and 22) Stenls pale, yellow; flowers 11-14 mm. diameter.

Stems sulcate; leaves broad (1 : 1-5-2); infloreacence 3-6 cm.; ovules 4-6; fruit

Stems subterete; leaves narrow (1 : 2.5-3); inflorescence 1.5-3 cm.; ovules 3-4; estylose. 186. B. coneirnilis

fruit stylose. 187. B. famnkna

Leaves entire; stamens apiculate; fruit estylose. 188. B. (parrlwirescena Stenls dark, red; flowers 6-9 mm. diameter.

haves spinose; stamens truncate; fruit stylose. 189. B. 8 h e ~ ~ 10-2

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148 L. W. A. AHRENDT

(186) Berberis consimilis Schneid., Ost. Bot. 8. 66, 394, 1916 Shrub 6-7 ft. Spines 3-fld., 1-2 cm. Leaves 4.5 x 2.5 cm., including decurrent petiole

to 4 m. oblong-obovate; margins with 6-14 spinules, distant 1-5-2.5 mm. (on shoots entire); reticulate. Pedumle 7-15mm. Outer se@ 4-5x2.5mm. Median sepals 6 x 3.5 mm. Inner sepals 7-8 x 4.5 mm. Petals 5-5 x 3.5 mm., emargbate; cuneate, with oblong-elliptic, separate, non-marginal glands. Stamns 3 mm., produced, truncate. Berries epruinose, 9-11 x 4.5-5 mm.

W. Szechuan: seed, Type cultivated at Arnold Arboretum. Cultivated: fl. 4 June 1938; fr. Nov. 1938 (Kew, no. 262).

(187) Berberis faxoniana Schneid., 1.c. 325, 1916

Shrub 5 ft. Spines weak, 5-8 mm. Leaeres to 3 x 1 cm., narrowly obovate; margins with 4-10 spinose (0-75-1 mm.) serrulations, distant 2 4 mm., by apex; reticulate. Peduncle 9-18 mm. Outer se@ 4.25 x 2.5 mm., oblong. Median s e p h 5.5 x 3-5 mm., oblong-ovate. Inner se* 7 x 5.5 mm., obovate. Petals 6-6-5 x 4.5 mm., oblong- obovate, incised; base cuneate, with small glands. Berries epruinose, 8-10 x 65.5 mm.

W. Szechuan: seed, Type cultivated at Arnold Arboretum. Cultivated: Vilmorin 1189 (K). Rock 24804 (BM, K), from Yunnan, Haba Shan, north

of Yangtze, 3rd peak of Likimg snow range, Bardar, 12,000 ft., fl. 1932, growing in an open rocky region, may be a variety of this species from which it differs in its stems being red at first, its entire leaves, and apiculate stamens.

(188) Berberis paravirescens Ahrendt, nomen nudum, Gard. IZZust. 64, (3307) 426, 21 Sept. 1944; J. Asiat. Soc. Beng. (Sci.), (3) 11; 4, 1945; syn. B. virescens var. m r m r p a Bean, Trees and Shrubs, ed. 4,1, 251, 1925; syn. B. virescens Hook. f. Bot. Mag. 46, t. 7116, 1890, pro pazte

Shrub 6-9 ft. Sterna subterete, not venuculose. Spines concolorous, 5-15 mm. Leaves 10-30 x 5-15 mm., including petiole, 2-5 mm.; cuneate, entire, obtuse, mucronate; venation indistinctly openly branched. Inflorescence 4-10-fld., 1-2.5 cm. Pedicel8 4-10 mm. Plowers pale yellowish green. Outer se& 2 . 5 ~ 1.5 mm., ovate, subobtuse. Nedian se& 3-5 x 2-25 mm. Inmy sepals 5 x 3 mm., obovate. Petah 4.5 x 3 mm., obovate, entire; glands 0-6 x 0.3 mm. Stamens 3 mm., shortly conical-apiculate. Berries at first red, ultimately black-purple, oblong, estylose, 9 x 5 mm.

Sikkim: Hooker, s.n., Type. Bhutan: Shabjetmg Bumthang Chu, 9500ft., fr. 5 Sept. 1949, Ludlow, S h e d &

Hicks 19677 (BM). Between Laya and Gyasa Dzong, Mo Chu, 11,OOO ft., 10 June 1949, Ludlow, S h e d & Hicks 16482 (BM). Bumthang Tang, 9500 ft., 5 Sept. 1949, Ludlow, S h e d & Hicks 18903 (BM). Tsele La, Wangdipoang, 10,50Oft., 27 Aug. 1949, Ludlow, S h e d & Hicks 19653 (BM). Lao, %ashi Yangsi Chu, 9000 ft., 15 May 1949, Ludlow, S h e d & Hicks 20635 (BM).

Cultivated: L.A. 34, from Sikkim seed, fl. 21 May 1942; fr. 7 Oct. 1943 (0). Also from LSH 19677. Described having fruit, a t h t red, ultimately black-purple to black. Also grown at Kew, (fl. 12 May 1939; fr. 31 Oct. 1939) as ‘the black fruited species from Bhutan, received from Headfort’.

(i89) Berberis shensiana Ahrendt, Qard. Chron. (3) 112, 155, 1942 Shrub suberect, to 6 ft. Stem shoots red, partly venuculose. Spines 3-fld., 8-15 mm.

Leaves 2 0 - 4 0 ~ 6-13 mm., or on shoots to 6 x 2 cm.; obovate; cuneate to decurrent petiole, 2-9 mm.; margins with 4-12 spinose (0.5 mm.) serrulations (0-25 mm.); openly veined to subreticulate. Racemes 6-10 (-15)-fld. PedS (P) 6-10 (-12) mm. Prophylls 2 x 1.25 mm. Outer sep& 4 x 3 mm., ovate. Inner sepals 5.5 x 3-5 mm. Petals 4.25 x

See under B. vkreacens and B. igmrata.

Page 149: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 149

3 mm., apex rounded, slightly retuse; with rounded lobes; base cuneate with approxi- mate, non-marginal, concolorous glands. Berries oblong, 8-9 x 6 6 mm., with style 0.3 mm.

shensi: Beed, Purdom 543. Cultivated: from above, fl. 5 May 1939; fr. 3 Oct. 1939 (Type, 0). Distributed as

' B. diehiam'.

KEY 35, to Sects. Franchetianae and Tschonoskyanae combined (Maps 3, 23, 27) Stem pubescent, or puberulous, at any rate at first. A

Bracts 4-5 mm., subequal to pedicels. 209. B. bracteatrc Bracts 1-2 (-2.5 mm.) much shorter than pedicels.

Leaves very narrow, 1 : 8-10; fruit blue-black. Cf. 224. B. lepidifoliolicc Leaves broader, 1 : 1-5-3; fruit red.

210. B. sieboldit

Sterns always glabrous.

Leaves with 50-150 marginal spinules; fruit globose, 6 mm. Leaves entire, or with 1-40 spinules; fruit oblong, 8-11 mm. long.

Leaves below, green, epruinose. 13 Leaves below grey or white, pruinose.

Stamens acutely dentate below anthers. 219. B. miqueliann Stamens edentate.

Stems terete, purple. 221. B. tschonoskyanrr Stems sulcate and/or yellow.

Flower diameter 12-15 mm.; pedicels 2.5-4 em.; fruit 10-12 mm.,

Flower diameter 6 1 0 (-12) mm. ; pedicels 1-12 (-20) mm. ; fruit estylose. 206. B. yunnanensis

6-9 (-10) mm.

half their length; fruit estylose. 208. B. spraguei Outer and inner sepals acuminate and narrow, their breadth less than

Inner sepals always, outer sepals often, broader, obtuse. Leaves ( 3 . 5 ) 5-10 em., acuminate; flower diameter 4-6 mm.

Leaves (0 .k ) 1-3 (-5) em., obtuse or acute; flower diameter, 216. B. virgetorum

6-10 (-12) mm. c'

A. Leaves below grey, pruinose, entire or remotely spinose. Flowers subumbellate, 2-9.

Pedicels 1.5-2 mm. Cf. 282. B. a&o-calida Pedicels 4-12 mm.

Pedicels glabrous. Leaves dull, entire; spines often absent, otherwise 1-12 nun.

Outer sepals as long as inner sepals; petals shorter than inner sepals;

Outer sepals shorter than inner sepals; petals as long &s inner sepals; style short. 201. B. papillifera

style absent. 202. B. microtricha 203. B. mekongensis

Pedicels pubescent. Cf. 192. B. franchetiana Flowers solitary. 190. B. minutiflora

200. B. tomentulosa

Leaves lustrous, often spinose; spines stout, 10-25 mm.

Leaves below green, epruinose ; fairly closely spinulose.

B. Stems, mature dark, red or purple. Leaves entire, always or mostly (rarely 1-10) inconspicuous spinules; spines

absent or weak, 5-9 mm. Leaves closely reticulate. Cf. 284. B. hemleyana Leaves with venation slight, at most branched, not reticulate.

199. B. rufescens 215. B. nwuillacana

Leaves with margins with 6-15 close spinules. 212. B. dubia

Stems terete; petioles 3-8 mm. ; pedicels 14-20 mm. Stems sulcate; petiole 8-20 mm.; pedicels 4-15 mm.

Page 150: Berberis and Mahonia : A taxonomic revision

160 L. W. A. AHRE~NDT

KEY 35 (continued) Stems, mature pale yellow.

Leaves entire, or margins with 1-5 remote spinules. Leavea 15-35x8-15mm., reticulate; pedicels 7-15mm.; flower diameter

10-14mm.; outer sepals as long as inner sepals; petals shorter than inner sepals; fruit stylose, pruinose.

Leevea 15-25x5-9mm.. sarmly veined; pedioels 3-5mm.; flower diameter 8-9 mm. ; outer sepals shorter than inner sepals; petals as long aa inner sepals; fruit estylose, epruinose.

Leavea with margins closely (7-) 12-35-spinulom; pedioels 4-6 mm.; fruit 5-8 x 3-5 mm., estylose.

Leaves 10-25 x 4-10 mm., with 7-15 marginal spinules; petals entire, shorter than inner sepals.

Leaves 25-66x 10-25mm., with 20-35 mar@ spinulea; p e a emarmte , longer than, or as long 88, the inner sepals.

205. B.dawoeltgis

208 f. B. x brewkcqm

201. B. dietymura

214. B. qdpat?&&

c. Leaves fsirly cloeely 5-20 (-4O)-spinOSe; fruit estylose. Pedicels 10-15 mm. 213. B. oritrepha Pedimls 4-10 mm.; petals aa long as, or longer than, inner sepals.

Leaves lustrous; fruit 7 x 4 mm.; stems red-brown. 198a. B. X SeW&

Leaves dull; fruit 9-10 x 4-5 mm. ; stems yellow-brown. Leaves entire, or with 1-4 inconspicuous marginal spinules.

208c. B.xcEeclinatcc

Fruit subglobose, 4-6 x 4-5 mm. Stems dark red; fruit estylose. Stems pale yellow; fruit stylose.

Petals longer than, or as long as, inner sepals; stamens truncate; fruit estylose.

198. B. boschanii Cf. 283. B. heteropsia

Fruit oblong or ellipsoid, 6-10 x 3-6 mm., or 10 x 9 mm.

Pedimls 9-20 mm.; petals as long as inner sepals; fruit 9-14 mm., soft.

Pedimls 4-9 mm. ; petals longer than inner sepals; fruit 7-9 mm., firm, lustrous. 197. B. taarorqensis

217. B. thunbergii 218. B. x ottaweltgis

Stems purple; leaves always entire; petals emargimte. Stems yellow; leaves partly spinose ; petals entire.

Petals shorter than inner sepals. Flowers solitary; pedicels 1-7 mm.; fruit 6-7 mm.; leaves mostly spinose. Stems yellow; pedicels 5-7 mm.; fruit estylose; shrub erect.

Stems purple; pedicels 1-3 mm.; fruit stylose; shrub prostrate. 190b. B. rninuti$ha var. glabrannea

191. B. graminea Flowers fascicled or subumbellate ; leaves mostly entire. Stems yellow; flower diameter 5-6 mm. ; f i t stylose.

Stems red, or purple; flower diameter (6) 8-10 mm. 204. B. hurnido-umbrosa

220. B. eilva-tarmmna

Fruit estylose. 196. B. lecorntei Fruit stylow; style 04-0-8 mm.

Fruit broad, 9-10 x 7-9 mm., estylose. Fruit narrower, 6-10 x 3-6 mm.

Fruit bright red; leaves with much branched venation; outer sepals shortly acute or obtuse; inner sepals broadly obovate; stems sulcate.

Fruit oblong, 9-10 x 5 mm., with straight style. Pedicels 10-20 mm.; leaves 20-40 x 6-10 mm., grey below;

Pedicels 4-8 (-17) mm. ; leaves 10-16 x 3-4 mm., white below;

Fruit 10 x 7 mm., h, lustrous, ovoid-attenuate to bent style;

Fruit dark red ; leaves only slightly veined ; outer sepals acUminate ;

stamens truncate. 192. B. franchetiana

stamens apiculate. 193. B.amena

stamens apiculate. 194. B. thibetica

inner sepals narrowly elliptic; stems subqled . 195. B. stearnii

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Berberis and Mahonia 151

Sect. Franchetianae Ahrendt, stat.nov. ; syn. Sect. Sinenses, Subsect. Franchetianae

Anternodes 1-3 (-5) cm. Spina mostly weak, though on new shoots and in a few species stronger. Leaves deciduous; margins in most cases entire or subentire, though in a fow species spwsely or closely spindose. InJbreswnee 4-8 (-15)-fld., fascicled, umbel- late, or pseudumbellate-subracemose, rarely solitary. Flowers 5-9 (-12) mm. diameter. Pet& mostly emarginate, nearly always shorter than inner sepals. Ovules (1-) 2 (-3). Fruit red, occaaionally dark red, oblong, 6-9 (-11) 111111. long, estylose or with style up to 8 mm.

S.E. Tibet, Burma, Yunnan, Szechuan, Kiangsi, Mongolia, Japan, Quelpaert Island.

Ahrendt, J. Bot., Lond., 80 (Suppl.), 112, 1944

Subsect. Minutiflorae, Ahrendt, J . Bot., Lond., 80 (Suppl.), 109, 1944 sltenzs sulcate. Internodes 6-18 mm. Spines 4-10 (-12) mm. Leave-s narrowly obovate,

6-14mm. long; margins entire, or with 1 4 spinules; openly veined; below pruinose, grey or white. Flowers solitary, 7-11 mm. diameter. P e d k h 1-8 mm. Outer sepals comparatively large. P& emarginate, shortei than the inner sepals, but longer than the outer sepals. Stanzens truncate, or obtuse. ovules 2. Berries red, epruinose, 6-7 x 4 4 5 mm.

S.E. Tibet, Yunnan, Szechuan.

KEY 36, to Subsect. Minutiflorae (Map 24) Stems yellow; pedicels 5-8 mm. ; flowers 6-8 mm. diameter; stamens produced; fruit

Steras purple; pedicels 1-3 mm.; flower diameter 10-11 mm.; stamens not produced; e&ylose. 190. B. minutifiora

fruit stylose. 191. B. graminera

(190) Berberis minutiflora Schneid., IU. H a d b . Laubh. 2 , 914, 1912, bst. Bot. Z., 67, 221, 1918; Ahrendt, J. Bot., Lond., 80 (Suppl.), 110, 1944; syn. B. angulosa var. brevipes Franch., PI. Delavay. 39,1889; syn. B. brewips (Franch.) Schneid., Bull. Herb. Boissier, ( 2 ) 8, 194,1908, non Greene, Ottclwa Nat . 15,42,1901. (Fig. 36)

Sfem not verruculose. S p i w mostly 4-8 mm. Leaves 8-12 (-14)x2*25-3*5 (3.75) mm., acute or subacute; margins 2-5-spinose (1-2 mm.)-serrate (1 mm.) distant 2-3 mm.; below papillose. Peokeh slender. Prophylls red, acute, 1.4 x 1.2 mm. Outer sepals 4 x 2-2 mm., oblong-ovate, obtuse. Inner sepals 5.5 x 3 mm., oblong-obovate. Petals 4.5 x 2.5 mm., incised with acute lobes; base not clawed, with ovate, obtuse, separate glands, 1 x 0.7 mm. Stamens 2-2 mm. Berries ovoid.

KEY 35A Shrub t,o 1 ft. ; young shoots pubescent; mature stems subglabrous.

Shrub to 6 ft. ; stems always glabrous. 190a. B. minuti,flora var. minutiflora

190b. B. minutifira var. glabramea

(190o) Berberis minutiflora var. minutiflora Yunnan: ad oras silvarum prope collum Yen-tze-hay, baud procul a Lankong,

10,400 ft., fr. 30 Oct. 1885, Delavay. 1046 (Type); Keua lape, 11,400 ft., fl. 26 May 1884. Delavay, s.n. (K).

S.W. Szechuan: Siga shan, open scrub, 1 ft., 11,OOO ft., May 1932, Rock 243922 (K). Cultivated (from Rock 24392) : fl. 12 Apr. 1939. A very useful compact small Berberis

for the rockery.

Page 152: Berberis and Mahonia : A taxonomic revision

152 L. W. A. AERENDT

(190b) Berberis minutiflora par. glabramea Ahrendt, var.nov., ramulis semper

Spinee 6-15 mm. Leaves to 13 x 4.5 mm.; margins (0-) 1 (-2)-spinose; subreticulate. Outer sepale 2-75 x 1-5 mm., ovate. Inner sepals 4-25 x 3 mm. Petals 3 x 2 mm.

S.E. Tibet: Prov. Takpo, Tstlngpo valley, Lapso Dzong, 11,500 ft., fl. 5 Apr. 1947, Ludlow, S h e d & Elliott 12437 (Typ, BM). ‘Steep hillsides, 4 4 ft.’ Not cultivated.

(191) Berberis graminea ahrendt, J. Bot., Lond., 80 (Suppl.), 110, 1944. (Fig. 37) Shrub prostrate, spreading. Stem with young shoots green, scarcely verruculose. Spines

subconcolorous, 1-3-fld., 6 1 2 mm. h u e s 8-21 x 2-5-5 mm. ; acute, mostlyentire, some- timeswithmargins 2-3-spinose(l.5mm.)-sedate (0.5-1 mm.),distant 2-4mm.;venation branched, scarcely reticuhte ; below whitish pruinose, scarcely papillose. Flowers H.C.C. 212-211. Prophylls concolorous, 2.5-1-5 mm., oblong-elliptic. Outer sepals 5 x 2 - 6 3 mm., oblong-ovate. Inner sepals 6-6.5 x 54-25 mm., elliptic-obovate. Petals 5.25x445mm.; apex rounded, narrowly incised with rounded lobes; glands dis- colorous, orange, elliptic, separate, submarginal, 1-1-25 x 0.5-0.7 mm. Stamens 2 mm. Style 0.5 mm.

S.W. Szechuan: Muli, Wachin, near Lamasery, lO,OOOft., fr. 4 Oct. 1947, Yu 14429 (Type, E). ‘Small common shrub, growing 1 ft. high on grmy mountain slopes.’

Cultivated: from the type collection, 5.20 May 1942,5 May 1944. This is also a useful garden species for ground cover, in rockeriea, or elsewhere. Although it rises very little upwards, it covers more ground than the very compact, B. minu.ti$ora. Both species flower moderately well, but, a t any rate in Oxfordshire, rarely bear fruit.

glabris

Subsect. Eufranchetianae Ahrendt, subsect. nov., ramulis atrorubris vel atropurpureis; f o b integris vel spinulis pauculis remotis

Stem glabrous, sulcate, mature dark, red or purple. Internodes (0.S) 1-2.5 cm. (except as noted, in raxe cases, to 5 cm.). Spinee mostly weak, 2-9 mm.; occasionally 1-2.5 cm. h u e s 1-3 (-5-5 cm.), entire, rarely with a few marginal spinules; below greyish or whitish, pruinose ; always openly veined, scarcely even subreticulate. InJlorescence (2-) 4-8 (-10)-fld., fascicled, umbellate, subumbellate, or pseudumbellate-subracemose. PZower8 6-8 (-11) mm. diameter. Pedicels 3-10 (-18) mm. Petals emarginate, always shorter than inner, but rarely shorter than outer, sepals. Stamens not produced, trun- cate or retuse at apex (rarely apiculate, and then so recorded). O& 1-2 (-3). Berries red, 6-8 (-11) mm. long, estylose or with very short style.

S.E. Tibet (provinces Kongbo, Tsarong), Yunnan, Szechuan.

KEY 37, to Subsect. Eufranchetime (Map 24) Leaves broad, 1 : 2-3; fruit red.

Fruit stylose. Fruit bright red; leaves with much branched venation.

Fruit oblong, 9-10 x 6 mm., soft, dull. Pedicels 1-2 cm.; leaves 2-4 cm. 192. B. framhtkma Pedicels 4 4 mm. ; leaves 1-2 cm. 193. B. anzoena

Fruit ovoid-attenuate, 10 x 7 mm. 194. B. thibetica Fruit dark red; leaves only slightly veined. 195. B.stamaii

Stems sulcate; pedicels 4-10 (-14) mm.; flowers 6-8 mm. diameter. h i t 8StylOSe.

Fruit 8-10 x 6 mm. Leavea entire; pedicels P l O m m . ; petals shorter than, or as long as, inner

Leaves spinose ; pedicels 8-20 mm. ; petals as long as inner sepals. Sepals. 196. B. lecomtei

197. B. tSar0ngenai.q

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Berberis and Mahonia 153

Fruit 6 x P4.5 mm. Leaves entire. 198. B. boschanii Leaves spinose. 198a. B. x serrata

199. B. rufescens Cf. 224. B. lepid~olfolia

Stems subterete; pedicels 14-20 mm.; flowers 10-12 mm. diameter. Leaves narrow, 1 : 8-10, fruit black; pruinose blue.

(192) Berberis franchetiana Schneid., Ost. Bot. 2. 67, 223, 1918

obovate; entire. Berries 9-11 x 4.5-6 mm., oblong-ovoid. internodes 2-3 cm. Spines 3-fld., slightly sulcate. Leaves 2-36 x 0.6-1 cm. ; narrowly

KEY 37A

Peclicels pubescent. 192a. B. franchetiana var. franchetiam Pecli-els glabrous.

Inflorescence 3-8-fld., 2-4 cm. long. 192b. B. franchetiana var. glabripes Inflorescence 8-14-fld., P 7 cm. long. 192c. B. franchetiana var. macrobotrys

(192a) Berberis franchetiana var. franchetiana

(7-) 10-15 (-18) mm.; bracts 1.5 mm. Style short. 1nJlorescence 4-6-fld., fascicled or subfascicled, 1.5-2 cm. long, almost sessile. Pedicels

Yunnan: woods a t Kou Tui, fr. Delavay, s.n. (Type, K). Not cultivated.

(192b) Berberis franchetiana var. glabripes Ahrendt, J . Bot., Lmul., 80 (Suppl.), 114, 1945

f i t e m partly sparsely vermculose. I n t e r n o h 1 6 4 . 5 cm. Spines absent or 6-18 mm. Injlwescence 3-8-fld., 2 4 cm. long, including peduncle to 15 mm. Pedicels 7-17 mm.; bracts 2 mm. Flowers H.C.C. 1. Prophylls lanceolate, 2.75 x 0-75 mm. Outer sepals 4.5 x 1.1 mm., lanceolate, obtuse. Median sepals 6 x 3.75 mm., oblong-obovate. Inner sepzls 7 x 5.5 mm., obovate. Petals 5-75 x 3.5 mm., elliptic, acute; base broadly clawed with concolorous, lanceolate, separate, submarginal glands. Stamens 3-75 mm. Berries oblong-ovoid, sometimes with apex attenuate. Style 0-5-0.8 mm., sometimes bent.

Yunnan: Atuntze, Doker La, 4 Nov. 1937, Yu 7864 (Type, E). Also Yu 10713, 10819

Cultivated (from Yu 7864, seed) : fl. 5 May 1943,5 May 1944; fr. 8 Sept. 1943. Plants, dis6ributed as Yu 10819, were B . forrestii. Plants distributed as Yu 10713 were a variety of B. spraguei. S.E. Tibet: Prov. Kongbo, Pasum Chu, 13,000 ft., in meadow, 25 June 1947, Ludlow,

ShwrifF t Elliot 13993 (BM).

(E).

(192c) Berberis franchetiana var. macrobotrys Ahrendt, l.c., 1945 I’edicels above, 5-13 mm. ; below, 10-25 mm. ; bracts acuminate, 2-3 mm. Prophylls

s m d . Outer sepals 3 x 1-25 mm. Median s e p d 5.25x3.25mm. Inner sepals 6 6 x 4.5 mm. Petals 5 x 3 mm., acutely emarginate, with acute lobes; glands approximate, 0.8 x 0.5 mm. Fruit 9 x 4-5 mm. Style 0.5 mm.

Yunnan: Chungtien, Pica, grassy mountain slopes, 10,400 ft., 30 Oct. 1937, Yu 13903 (Type, E). Lichiang snow range, 9800 ft., 23 May 1937, Yu 15053 (E); common.

Cultivated: from both these numbers; examined (15053), fl. 16 Apr. 1945; fr. Oct. 1944. This variety somewhat resembles B . forrestii which differs in its larger, estylose fruit; its bright red shoots, and its mature yellow stems.

Page 154: Berberis and Mahonia : A taxonomic revision

154 ’ L. W. A. AIXRENDT

(193) Berberis amoena Dunn, J . Linn. Soc. (Bot.), 39,422,1911 ; syn. B. sinensis var. eLqam h c h . , PI. D&my. 35, 1889; B. w a r n (fianch.) Schneid., BuU. Herb. Boissier, (2), 8, 463, 1908, non Kirchner, non IRveilld et Vaniot; syn. B. lepi!ududa Diels, Not. Bot. Card. Edinb. 5, 167, 1912; syn. B. sc7lnei&eri Rehder, J . Am. Arb. 17,323, 1936. (Fig. 38)

InternaEap 8-18 mm. 8pk.s slender, P 1 0 mm. Leavea 12-16 x 3 4 mm., narrowly obovate, obtuse, mucronulate, cuneate sessile, entire, dull green, below white pruinose papillose; venation very alight. PrqhyUs lanceolate, 16-2-25 x 0.5 mm. Outer sepals 2-2-25 x 1.25-2 mm., obovate. Inner sepal8 44.5 x 3-3.5 111111. Petals 3.54 x 2.5- 2-75 mm.; apex retuse with two rounded lobes; base cuneate with separate, non-mar- ginal, narrowly oblong glands, 0.6 x 0.2 nun. &!mnm.s 2.5 nun., apex produced, apiculate. Bewies 6 x 3 mm.

In none of its varieties is this attractive species in cultivation. In appearance it bears some resemblance to the forms of B. Ouilsonae, from which it is easily distinguished by its scarcely veined leaves and apiculate stamens, and fewer ovules. See under (550).

KEY 37B Pedicels 4-8 mm.; bracta 1-5-24 mm.

Moreacence subracamuse, 3-5 cm. Inflorescence umbellate, 1.5-3 cm.

PediCel3 9-18 mm.

1930. B. anwena var. m e n u 193 b. B. amem var. urnbeUi,fEoa

193c. B. amoenu vm. moloenaia

(1934 Berberis amoena var. am~ena Yunnan: chalk hills at foot of mountains, Yang-in-chan, above Mo-so-yn, 8200ft.,

fl. 18 June 1887, Delavay, s.n. (type, of B. &newis elegam). On ledges of cliffs and stony slopes in side valleys on the Chienchum-Mekong divide, 11,OOO ft., July 1923, Forrest 23448 (K). ‘Low branched shrub, 1-2 ft., with orange yellow flowers’; dry rocky hill- sides on the descent from the Chungtien plateau, to the Yangtze, near Tang-tu, 10,OOO ft., Sept. 1904, Forrest 330 (Type of B. leptoduda) ‘Shrub 4 A ft.’.

(193b) Berberis amoena var. umbelliflora Ahrendt, var.nov., inflorescentiis um- bellatis

Inflorescence 5-7-fld., perfectly umbellate, or (rarely) nearly so, 15-25 mm., including peduncle, 13-15 mm.

Yunnan: fl., 1917-19, Forrest 16323 (Type, K): dry stony situations on the plain, north end of Lichiang valley, June 1906, Forrest 2233 (K). ‘Shrub 2 4 ft., with fragrant flowers’; 1O,OOO-11,OOO ft., Chungtien, 1932, Rock 24688 (a); Lichiang, 1932, Forrest 21116 (K). Both the Iatter 6-24 in. high; also Forrest 5632,5697,10282. Chalk hills, W. of Lichiang, July 1914, Schneider 1783 (K); Yunnan-fu, 18 &by 1916, Schoch 8 (K).

(193c) Berberis amoena var. moloensis Ahrendt, varaov., pedicellis multo lon-

In$wescenm 5-9-fld., 1.5-3 cm. Pedicels very slender. Outer sepah 3.25 x 1.25 mm., narrowly ovate, subacute. Inner sepals 4.5 x 3.75 mm., obovate-elliptic. Peta;ls 44.5 x 2-2.25 mm., narrowly obovate; clawed, with glands 0.9 x 0.4 mm. O m h solitary.

S.E. Tibet: Prov. Kongbo, W o n g Chu, Molo, 10,500 ft., fl. 24 May 1938, Ludlow, SherrH & Taylor 3835 (Type, BM). ‘Shrub 4 4 ft., with perianth bright yellow.’

(194) Berberis thibetica Schneid., Fedde Rep. Sp. Nov. 6, 268, 1909 Shrub sparsely branched, to 6 ft. Spines solitary to 7 mm., on shoots sometimes

3-fld., 7-12 (-25) mm. Leaves 1-26 (4) x 04-0-9 (-1.3) cm., obovate, subseasile, entire,

Possibly B. leptocIada Diels is a taller growing form of the dwarf variety anwe’lu16.

gioribus

Page 155: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 155

rounded, dull above, below scarcely papillose, venation branched. In$wescence (2-) 4 7 (-9;)-fld., racemose-subfaacicled to subumbellate, (1-) 2-3 (4) cm. Pedials 4-6 (-10, below) mm.; bracts 1-1.5 mm. Prophylls yellow, ovate, acuminate, 3.5 x 1.5 mm. Outer sepah 5.5 x 2.75 mm., ovate, acute. Inner sepals 6.5 x 3.5 mm., elliptic. Petals 4.5- 5 x 2.75-3.5 mm., obovate, emarginate; base cuneate with separate submarginal glands, 0 7 x 0-4 mm., situated about 1 mm. above the base of the petal. Stamens 3 mm., produced, apiculate. Ovuks 2-3. Berries firm, scarlet, lustrous, pruinose, ovoid, 9-10 x 6-7 (-8) mm., excluding style 0.75 mm.

S.E. Tibet: Prov. Tsarong, Tseku, on Mekong, 5000-8000 ft., seed, Soulie, 8.n. Cultivated: originally from Soulie’s seed. Examined L.A. 163, fl. 16 May 1939,6 June

1943, 16 May 1944; fr. 3 Oct. 1939,22 Oct. 1943,13 Oct. 1944. The berries are attractive, but otherwise, with its sparse straggling branches, this is an ungainly plant for the garden.

(195) Berberis stearnii Ahrendt, in Card. Chron. (3), 109, 101, 1941. 8hru6, suberect, to 6-7 ft. Stems with young shoots greenish, terete to subsulcate;

mature sublustrous, red-brown, not verruculose. Internodes 1-2.2 cm. Spines con- colorous, solitary or 3 fld., 8--10 (-12) mm., a t the bme nearly 1 mm. broad, subsulcate. Leaves 1.2-3.3 x 0.4-1 cm. ; obovate or oblong-obovate; rounded; attenuate, subsessile; mostly entire, sometimes with 1 4 spinules each side; openly veined, not reticulate; above bright green; below greyish pruinose, scarcely papillose. Inflorescence (3-) 4 8 (-10)-5d., mostly fascicled, sometimes racemose subfaacicled, or partly umbellate, 1-1.5 (-2) cm. Pedicels slender, 6-10 mm., with basal bracts scarcely 1 mm. Flowers, H.C.C. 3, red without, 9-10 mm, diam., with narrowly ovate, acuminate, truncate prophylls, 3.5- 4 x 1.5-1.75 mm. Sepals, outer 5-6 x 2-3 mm., narrowly obovate, acuminate; inner 5-6 x 25-3.5 mm., very concave, narrowly obovate, obtuse. Petals broadly obovate, 4-5-5 x 3-3.5 mm., acute, but narrowly incised, biapiculate; scarcely clawed, with glands separate, non-marginal, oblong ovate, obtuse, 1.5 x 0.5 mm. Stamens 2.5 mm., scarcely produced, truncate. Ovules 2 ( 3 ) , sessile. Berries narrowly obovoid, or oblong, more or less dark red, 6-5-7.5 (-8) x 3.5405 (-5) mm., excluding distinct style, c. 0.5 mm.

Yunnan: seed, 1932, F. 29042. Cultivated: LA 73 (F. 29042, type, from seed)-fl., 14 April 1938, 12 May 1939;

fr., 31 Oct. 1939, 7 Oct. 1942, (all S) (Type, 0). Also fl., 5 May 1944, (Wisley); fr., 26 Oct. 1938, (K). The bright red 5ower buds come out with the bright green leaves which are then mottled with red, and at this stage the species is unusually fine.

(196) Berberis lecomtei Schneid., Sargent, PI. WiL. 1, 373, 1913; syn. B. thunbergii var. glabra Franch., PI. Delavay. 35, 1889; syn. B. sinensis var. typica Franch,, 1.c. 1889, non Desf.

Shrub compact, densely branched, 6-7 ft. Spines sometimes absent, mostly 1-3-fld., 2-10 (-15) mm. Leaves 12-28 (-35) x 5-11 (-14) mm., obovate, obtuse, entire, base with petiole 2-5 mm., dull dark green. In.escence 44-fld., racemose, subfascicled, or sub- umbellate, (1-) 1.5-2.5 (-3) cm. long, including peduncle 1-10 mm. Ppdicels 4-10 mm.; bracts 1 x 0.7 mm. Flowers with sepals H.C.C. 212-211. Prophylls red, 1.5-2 x 0.8-1 mm. Outer sepals 2 - 5 3 x 2.5 mm., broadly ovate, acute. Inner sepals 4-4.5 (-5-25) x 2.5- 2.75 (-3.25) mm. Petals H.C.C. 2, obovate, 6 5 x 2-3 mm.; apex acutely emarginate with. acute lobes ; base cuneate, with separate to subapproximate discolorous orange glands, oblong, 0.6-1.2 x 0.3-0-5 mm. Stamens 2.5 mm., scarcely produced, truncate, or slightly obtusely subconical. Ovules sessile. Berries deep red, firm, lustrous, oblong, or oblong-obovoid, 8-10 x 4 - 6 5 mm., estylose.

Yunnan: in silvis ad Kou-tou, supra Mo-so-yn, and forests in the Heechan-men distriot, west of Yen-tze-hay pass, 9800 ft., fl. Delavay s.n. (Type, K, of B. lecomtei and

Page 156: Berberis and Mahonia : A taxonomic revision

156 L. W. A. AERENDT B. thunbergii glabra) ; Hokin, Mt. Houa la pa, 8 km. north of Yan-tze-hay p w , Delavay 1047 (Type of B. 8inemk var. typica Franch.) ; Lankong-Hokin divide, 9OOO ft., May 1913, Forrest 9991 (K) ; Mekong-Salween divide, Chingpoh, among thickets on open dopes, 98OOft., 14 June 1938, Yu 19089 (E) ‘common, 3-5ft.’; Chungtien Habe, 91OOft., 6 Sept. 1937, Yu 13511 (E); casual, 4ft.

S.W. Szechuan: open situations, inside valleys on the mountains south-east of Muli, 11,OOO-12,OOOft., Sept. 1922, Forrest 22395 (K); western slope of Mt. Mitzuga, Muli territory, rocks, 13,000 ft., 1932, Rock 24513 (K), Rock 24025 (BM). The specimens of Rock and Yu have a longer (3-5 cm.) subracemose idorescence; the Szechuan specimens come from a considerably higher altitude than the Yunnan ones.

Cultivated: L.A. 54 (derived by cutting from an old plant at Kew, no. 252), fl. 4 May 1944; fk. Nov. 1944. Also from Forrest 20292, fl. 12 May 1939 (K); and from Forrest 22395, fl. 26 June 1939 (K). This dense, compact species is certainly remarkably like B. thunbergii (from Japan) which differs in ita broader leaves, with longer petiole and which are a brighter yellow green in colour, its paler flowers with pet& distinctly longer than the inner sepals (a character which, however, is closely approached in B. kcomtei), and its smaller berries. The geographical distinction is very great, and presumably Franchet laid a smaller emphwis on this when, noting the close botanical resemblances between the Yunnan-Szechuan and the Japanese specimens, he made them conspecific. However, there is very little connexion between either of these species and ‘ B. sinensis ’, whichever interpretation be attached to this name (see under Sect. Sinenses).

(197) Berberis tsarongensis Stapf, Bot. Hag. 161, t. 9332, 1933 Stem with young shoots red; mature dark red. Injbrescence 4-9-fld., umbellate, or

pseudumbellate subracemose, 1-5-36 cm., including peduncle, 5-9 mm. Stamens not, or scarcely, produced. B e w k red, epruinose, oblong-ellipsoid.

ILEY 37c Fruit crimson, 8-10 x 5-7 mm. Fruit bright red, 12-15 x 4-5 mm.

197a. B. tsarongensis var. tsarongeneis 197b. B. taarongensis var. megacurpa

( 197 a) Berberis tsarongensis var. tsarongensis Spines conmlorous, 13-fld., slender, 1-2 cm. long. L m v a 10-18 x 5-8 mm., obovate

or elliptic; cuneate, subsessile; margins each, generally, with 1 4 spinules, occasionally entire; venation branched; above bright green, below papillose. Ped i~ l s glabrous, 8-15 (-20) mm. ; bracts 1-16 mm. Pmphylls acute, 2 x 1-25 mm. Outer sepals 3.75 x 2 mm., oblong-elliptic obtuse. Inner se& 5-5 x 4-5 mm., obovate. Petccls 5.5 x 3.5 mm., oblong-obovate ; apex emarginate with rounded lobes; base cuneate, with ovate, obtuse glands, 1 x 0.6 mm. Stamens 3-5 nun., rounded.

S.E. Tibet: Prov. Tsarong, Mekong-Salween divide, open scrub, alpine meadows or cliffs, 12,000-13,OoO ft., Sept. 1917, Forrest 14290 (Type, K). Also Forrest 22623.

N.W. Yunnan: MekongSalween divide, 11,OOO-13,OoO ft., Forrest 16554, 18992, 20621, 20802,20706.

Cultivated: L.A. 115, fl. 30 May 1939, 1 May 1942,4 May 1944, 16 May 1945.

(197b) Berberis tsarongensis var. megacarpa Ahrendt, varmov., fructibus multo longioribus, paululum angustioribus, scarlatinis

Stem sparsely verruculose. Interrwda 17-35 mm. Spines solitary, 5-10 mm., or 3-fld., to 17 mm. Leaves 20-35 x 6-12 mm., narrowly obovate; margins 1-3 (-5)-spinose (16-2 mm.)-serrate (1-1-5 mm.), though entire on the new shoots; venation branched to subreticulate; below scarcely papillose. Pedacels 6-12 mm.; bracts 1 - 6 3 mm. Outer

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Berberis and Mahonia 157

sepa.ls 3 x 1-5 mm., obtuse. Inner sepals 5.5 x 4 mm. Petals 5-5 x 3.5 mm., acutely emarginate ; base clawed, with oblong-elliptic, separate, subacute glands, 0.6 x 0.3 mm. Stamens truncate. Seeds pale yellow-brown.

S.E. Tibet: Prov. Kongbo, Tammyen Chu, Picea forest, 10,OOO ft., fl. 24 June 1938, Ludlow, Sherriff & Taylor 4976 (BM). ‘Shrub 6 ft.. with flowers butter yellow’; Molo, Lilung Chu, 10,5OOft., 30 Sept. 1938, Ludlow, Sherriff & Taylor 6544 (Type, BM). ‘Common, 5-8 ft.’

Ctdtivated: from Ludlow, SherrB & Taylor 6544, but with no record for the last 12 years of any flowers in cultivation.

(198) Berberis boschanii Schneid., Sargent, P1. Wils. 1, 369, 1913 Shrub 6-8ft. Stems with shoots red. Spines 1 (-3)-fld., 1.3-2.5 em. Leaves 1 P

35 x 8-16 mm., obovate, obtuse; contracted to a decurrent petiole, 2 4 mm.; entire; slightly veined. Inflorescence &8-fld., racemose-subfascicled, 1.5-3 cm. Pedicels 3-8 mm.; bracts 1 mm. Ovules 2. Berries broadly obovoid, 6 x 4-5 mm.

W. Szechuan: Mao-chou, Min valley, 6500ft., fr. Oct. 1908, Wilson 1166 (Type, K). Shrub 4-6 ft., with scarlet fruit.

Cultivated: a plant of the type was at Kew until about 1938, but has since died. I have no record of its distribution, or of any further cultivated plants.

(198a) Berberis x serrata Koehne, Deutsche Dendr. 170, 1893; syn. B. xprovinciu.lis var. serrata (Koehne) Schneid, J . A m . Arb. 4, 196, 1923.

Spines to 1 cm. Leaves to 10-23 x 3-6 mm., acute, with 6-12 marginal spinules sub- reticulate; bright lustrous green, below scarcely paler. Inforescence 6-15-fld., racemose, 2-3 em. Pedicels 5-10 mm. ; bracts 2-3 mm. Prophylls 2 mm. Outer sepals 4 mm. Inner sepals 5 mm. Petals 5-6 mm., emarginate. Stamens truncate. Ovules 2-3, subsessile. Berries red, ellipsoid, 7 x 4 mm., estylose. Seeds cinnamon coloured.

Cultivated origin : not cultivated in England now.

(199) Berberis rufescens Ahrendt, sp.nov., ramulis subteretibus, pedicellis longis, floribus maj oribus

Stems with young shoots green ; mature subterete, lustrous dark red. Ircternodev 2-5-5 cm. Spines nil, or 1 (-3)-fld., 5-9 mm. Leaves to 5-5 x 2.1 em., oblong-obovate, rounded; entire or with margins with 5-10 inconspicuous spinules; venation faint but branched; below, ultimately green, scarcely papillose. InJlorescence 4-7-fld., fascicled, or loosely racemose- subfascicled, 2-3.5 cm. long, including peduncle to 8 mm. Pedicels very slender, 14-20 mm. ; bracts, 1.5-2 mm. Outer sepals 4-5 x 2.5 mm., oblong-elliptic. Inner sepals 7 x 4.5 mm., obovate. Petals 4 x 3.5 mm., broadly obovate, subentire or slightly emar- ginate; glands, 0.8 x 0.5 mm. Stamens 3 mm., apex produced, acutely emarginate with two acute lobes. Ovules 2. Style absent.

N. Upper Burma: Adung valley, 9OOOft., fl. 19 May 1931, Kingdon-Ward 9497 (Type, BM) : ‘A big straggling shrub, the long loose branches drooping or clawing their way up through the thickets and reaching a height of 10 ft., or more; leaves red edged. In conifer forest, scattered or in thickets, along the river bank.’ Not in cultivation.

Subsect. Yunnanenses Ahrendt, J . Bot., Lond., 79 (Suppl.), 58, 1941, sensu emend. speciebus omnibus exclusis, sp. typica, B . pnmnensi, solum excepta, speciebus diversis additis

Stern sulcate; mature pale yellow. Intemzodes (1-) 1.5-3 (-7) cm. Leaves 1-3 (-6) cm., below pruinose, white or grey in most cases, occasionally green epruinose. InJEorescence (2-) 4-8 (-12)-fld., fascicled, umbellate, subumbellate, or pseudumbellate-subracemose.

Page 158: Berberis and Mahonia : A taxonomic revision

158 L. W. A. AHRENDT Pedicels mostly (3-) 6-12 (-20) mm., sometimes 2-5-4 cm. F k r s (6) 7-10 (-14) mm. diameter. Inner sepal8 nearly always shorter than, rarely subequal to, the petals. ovudes 1-2 (-3). BeMies bright or dark red, (6) 8-10 (-12) mm. long, estylose or with extremely short style not exceeding 0.3 mm.

S.E. Tibet (Prov. Monyul, Takpo, Kongbo, Pome, Zayul, Tsarong), Yunnan, Szechuan.

BY 38, to Subsect. Yunnanenses (Map 25) Stems when mature pubeacent or puberulous.

Leaves green, epruinose below; pedicels pubescent. 200. B. tonzentubau Leaves grey, pruinose below; pedicels glabrous.

Fruit stylose. 201. B. p p d @ f e r a Fruit estylose.

Spines weak, 1-7 mm.; leaves 1-2 cm., dull; petals as long as inner sepals; fruit

Spines stout, 10-25 mm. ; leaves 2-4 cm., lustrous; petals shorter than inner 6 mm. 202. B. rnicrotricha

sepals; fruit 9 mm. 203. B . m e k o n g e d Stems always glabrous.

Leaves quite entire. Fruit Stylose, 7-8 mm.

Leaves n8rrow (1 : 3-4), mute. Leaves broader (1 : 2-3), rounded.

204c. B. humido-u&om var. dispersa

Leaves below grey, pruinose ; flower diameter 6 mm. ; outer sepals shorter than

Leaves below finally green; flower diameter 12-15 mm. ; outer sepals as long inner sepals. 204. B. humido-U&OaU

as inner sepals. 205. B.&woensis 206. B.yocnlaaneneis Fruit estylose 10-12 mm.; flowers 15 mm. diameter.

Leaves always spinoee-serrate; flower diameter 7-10 mm. Br& 1-2 mm., leas than (t) t ae long ae pedicel. Petals shorter than inner sepals.

Leaves green, epruinose below; sepals broad, obtuse; pedicels 3-6 mm. 207. B. dictymura

208. B.qnagwi

208c. B. x declinata 208e. B. x brevbmpa

209. B.bacteata

Leaves @By, plYlhOS0 below; Sepals =OW, fMXldIl8k3; P e d i C e l S 6-14 Illm.

Petals as long as inner sepals. Leaves below grey, pruinose. Leaves below green, epruinose.

Brwte 4-5 mm., subequal to, or exceeding, pedimls.

(200) Berberis tomentulosa Ahrendt, J. Bot., M., 80 (Suppl.), 112, 1944 Stern very sulcate, very pubescent. Inter?wde.a 7-12mm. Spines 3-fld., 6-1Omm.

Laves to 14 x 5-5 mm., obovate, rounded, cuneate, d e ; margins towards apex 5-8- spinose (1-1.5 mm.)-serrate (0-5 mm.), distant 1-1-5 (2 mm.) cloaely; reticulate. In- jzorescence 5-10-fld., pseudumbellate-subracernose, 1-5-2 cm. long, including peduncle, 7 mm. Pedicel8 puberulous becoming subglabrous, 2 4 mm. ; b m t s 1 - 6 2 mm. Flowers 5-6 mm. diameter.

Yunnan: Upper Kiukiang valley (Chulung), Sangolila, mountain slopes and open rocky places, 8200 ft., 5 Aug. 1938, Yu 19640 (Type, E). ‘Shrub 2-3 ft., with pale yellow flowers, casual.’ Not cultivated.

(201) Berberis papillifera (Franch.) Koehne, Carten,. 48, 21, 1899; syn. B. thun- bergii var. papillifera Franch., PI. Delavay. 36, 1889; syn. B. jimtii Schneid., BuU. Herb. Boksier, (Z), 8, 203, 1908

Shrub 5-7 ft. Stern shoots pubescent; mature puberulous and partly venuculose. Spines absent, or 3-fld., 2-12 (-20) mm. Leaves 10-30 x 3-10 (-12) mm. ; narrowly oblong-obovate; attenuate to petiole 2-4 mm.; entire; above deep green, scarcely

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Berberis and Mahonia 159

lustrous ; below closely, finely papillose ; subreticulate. Inflorescence 3-7 (-9)-fld., €ascicled or pseudumbellate-subracemose, (1-) 1-5-26 (-3) cm., including peduncle 4-8 mm., or umbellate and partly fascicled, 3-6 cm., including peduncle 2-3 cm. Pedicels 3-6 (-9) mm., above, 8-12 (-20) mm. below: bracts 1.5-2 (-3) mm., acuminate. Flowers H.C.C. 1. Prophylls yellow, lanceolate, acuminate, 2.75 x 0.6 mm. Outer se@ 6 x 3 mm. Inner sepals 6 x 4 mm., all obovate. Petals 4.5 x 2.5 mm., elliptic, acute; apex narrowly incised with acute lobes. Stamens 2-5 mm., not produced, truncate. Ovules sessile. Berries red, oblong-ellipsoid, 8-85 x 4.5-5-5 mm. Style 0.3 mm.

Yunnan: woods on Mt. Hee-chan-men, by Lankong, 9100 ft., 23 Aug. 1884, Delavay 1087 (Type of all citations) (K); Fang-yang-tchang, above Mo-so-yn, 21 Oct. 1887, Delavay, s.n. (K) ; Chungtien, Lichiashica, grassy mountain slopes, 11,300 ft., fr. 16 Nov. 1937, Yu 10972 (E), ‘Common’.

Cultivated: from Yu 10972, fl. 19 May 1943; fr. 4 Nov. 1943.

(202) Berberis microtricha Schneid., &t. Bot. 2. 67, 223, 1918 Sikm.s shoots reddish. Spines absent, or 1-3-fld., &8mm., concolorous. Leaves

20-25 x 4-8 mm. ; narrowly obovate; subacute, attenuate, subwsile, entire, venation branched, scarcely subreticulate ; above dull; below papillose. InJlorescence 2-7-fld., fascicled or racemose-subfascicled, 1-2 cm. Pedicels (5) 7-9 (-12) mm. ; bracts 1.5 mm. Outer sepals 3-5 x 2 mm., oblong. Inner s e p d 5 x 2.5 mm., elliptic. Petals 5 x 2.75 mm. : oblong-elliptic ; glands oblong-obovate, separate, non-marginal, 0.8 x 0.3 mm. Stamens 3 mm., produced, truncate. Berries epruinose, ellipsoid, 6-7 x 4 mm. 8. Szechuan : in woods, near streams at Wo-lo-ho, between Yen-juan-hisen, Yung-

ning, 8500 ft., 23 June 1914, Schneider 1543 (Type, K). Yunnan: Lichiang snow range, 5-6 ft., common among thickets, 9700 ft., 0. 21 May

1937, Yu 15048 (E). Chltivated: from Yu 15048, fl. 9 June 1943; fr. 18 Nov. 1943; and, at Wisley, fl. 5 May

1944.

(203) Berberis mekongensis W. W. Sm., Not. Bot. Card., Edinb., 9, 82, 1916 Shrub 4-6 ft. Stem stout, rigid; shoots puberulous, mature glabrous, sparsely finely

vermculose. Spines 3-fld., slightly paler, sulcate, stout, 1-2-5 cm. Leaves 20-45 x 10- 20 inm., obovate, rounded; broadly cuneate, subsessile; entire or with margins 10-15- sphose (0.5-1 mm.)-serrulate; very openly reticulate; very lustrous deep yellow green; below scarcely papillose. Inflmescence 6-12-fld., subumbellate or pseudumbellate-sub- rac(:mose, mostly sessile, 3 6 cm., long, sometimes 4 7 cm. long, including peduncle 1-2.5 cm. P e d i d (P) 7-12 (-15) mm. ; bracts 1-16 mm. Fbwers H.C.C. 2. Outer sepuh lanceolate, concolorous, acuminate, 4 x 1-1.4 mm. Mediun sepah 5-55 x 2-3 mm., oblong-elliptic. Inner sepals 6-6-5 x 3 - 5 4 mm., obovate. Petals P 5 x 26-35 mm., obovate, subacute, narrowly incised ; base clawed, with separate non-marginal obovate glands, 0.8 x 0.6 mm. Shmew 3.25 mm., produced, truncate. Berries red, oblong, estylose, epruinose, 8-10 x 4-6 mm.

Yunnan : Mekong-Yangtze divide, open situations amongst scrub on the Bei-ma- shan, 12,OOO ft., 14 Aug. Forrest 13204 (Type, K), ‘shrub 6 ft.’; Atuntze, Doker La, 11,(H)o ft., open situations, 8 Nov. 1937, Yu 7925, seed only, differing from the herbarium sheet of this number which is B. yunnanensis.

Cultivated: from type, Forrest 13204, f l . 10 May 1940 (K); fr. 26 Sept. 1940 (K); L A . 91, fl. 6 June 1939, 5 June 1940, source unknown; also from seed of Yu 7925.

(204) Berberis humido-umbrosa Ahrendt, J. Bot., Lond., 80 (Suppl.), 115, 1945 Shwb 5-8 ft. Stems verruculose. Spines 1-3-fld., 1-2 cm. Leaves oblong-obovate;

bast: cuneate to decurrent petiole 1-2 mm.; entire with venation branched; below

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160 L. W. A. AHRENDT scarcely papillose. In,escence P8-fld., loosely racemose to racemose-subumbellate, (1-) 2-3 cm., including peduncle (3-) 8-15 mm. PediuA glabrous. Be+ red, obovoid,

This small-flowered species, has a long west to east distribution from the Bhutan 7-8 x 4 - 5 5 mm. Style distinct.

border to just within W. Szechuan.

KEY 3 8 ~ Leaves rounded, broad, 1 : 2-3, pruinose below; pedicels 6 9 mm.

Fruit pruinose. Fruit epruinose.

204a. B. hud-uTnbr08a vm. hurnido-umbrosa 204b. B. humid~-umb*o8a vm. inornata

204c. B. hmido-untbrosa vm. dispersa Leaves acute, narrow, 1 : 3-4, epruinose green below; pedicels 7-14 mm.

(204 a) Berberis humido-umbrosa var. humido-umbrosa Leaves rounded. Ped& (6) 6-9 mm.; bracts 1-1-5 mm. Outer sepal8 2-75 x 1-75 mm.

ovate, obtuse. Inner sepal% 3.5 x 2-5 mm., obovate. Petals 3.25 x 2.5 mm., obovate, subentire ; glands elliptic, sepaxate, 0.6 x 0.4 mm. O w d a 2. StQnzens 2.25 mm., produced, truncate. Be& heavily pruinose blue or violet.

S.E. Tibet: Charme district, Sanga Choling, 11,000-12,000ft., fr. 18 Sept. 1935, Kingdon-Ward 12357 (Type, BM). ‘Common in ravines or wherever there is water or shade; fruit either red, or blue violet, two such bushes growing side by side.’ Chayul Chu, 11,500 ft., fl. 5 May 1936, Ludlow & S h e d 1547 (BM). ‘Growing in damp places along water channel in irrigation zone.’

Prov. Kongbo: Deyng La, l0,OOO ft., 30 May 1947, Ludlow, S h e d & Elliot 15077 (BM), fl. in rain-forest, and fr. 17 Sept. 1947.

Prov. Pome: Lokmo, 10,500 ft., 1 June 1947, Ludlow, S h e d & Elliot 13802,13805, 13806 (BM).

Prov. Takpo: Kongbo nga la, Tsangpo valley, 11,500ft., 30 Apr. 1947, Ludlow, SherriE & Elliot 12425 (BM). ‘Flowers bright yellow, common.’

(204b) Berberis humido-umbrosa var. inornata Ahrendt, l a , 116, 1945 Berries red, epruinose. Otherwise as in 204a. S.W. Szechuan: Muli, Kulu, 10,40Oft., 17 Sept. 1937, Yu 14298 (Type, E). ‘Shrub

S.E. Tibet: Charme district, Sanga Choling, as previous variety. Prov. Tsarong: Mekong-Salween divide, Londre La, lO,OOo-l1,0o0 ft., May 1922,

Forrest 21609, and Oct. 1922, the same in fruit, F o m t 22760 (K). ‘Shrub 6-9 ft.; stems arching; flowers dark yellow.’

Cultivated: from F o m t 22760, fl. 12 Mar. 1939; fr. 25 Oct. 1939 (K). From Yu 14298, the type of this variety, fi. 20 Sept. 1947, at Maidwell Hall, Northampton.

5-7 ft.; fruit red.’

(204 c) Berberis humido-umbrosa var. dispersa Ahrendt, varnov., occidentalior

Leaves 13-33 x 5-8 mm. In.orescence 8-12-fld., 3 - 5 5 cm., including peduncle 1.5- 2.5 cm. P&i& very slender, 7-14 mm. Prophylk 1-5 mm., acuminate. Outer sepals 3-5-4 x 1-5-2 mm., oblong-ovate, subacute. Inner sepals 5 x 3.5 mm., obovate, rounded. Petuls 3.5 x 2-5 mm., obovate, entire. Shmm 2-75 mm., produced obtuse.

S.E. Tibet: h v . Monyul: Trimo, Nyam Jang Chu, M R . , flowers lemon yellow, common, 10,500 ft., 23 May 1947, Ludlow, S h e d & Elliot 12352 (Type, BM). Not cultivated. Thie variety is very distinct from the two preceding ones, in the leaves being quite green below as in the following speciea.

foliis acutis subtus viridibus

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Berberis and Mahonia 161

(205) Berberis dawoensfs K. Meyer, Fedde Rep. Sp. Nov. Beih. 12, 379, 1922 Stemswithyoungshootsred. Spines 3-fld.,6-15 mm.,subsulcate. Leaves 10-30(-35) x 7-

11 (-15) mm., oblong-obovate; attenuate to decurrent petiole 1 4 mm.; entire or with margins 3-7-spinulose (0.8 mm.)-subsedate, distant 3-5 mm. ; obtuse; openly reti- culate; below a t first grey, pruinose, finally green, epruinose. InJlorescem GlO-fld., subfascicled, or umbellate-subracemose, 1-5-3 (4) cm. Pediwls 7-12 (-16, below) mm., very slender, glabrous. Flowers H.C.C. 64. Outer sepals 6-8 x4.5 mm. Inner sepals 6.5-8.5 x 4-6 mm., all obovate. Petals 5-7 x 3-5 mm., acutely emarginate with acute lobes ; base cuneate, with approximate glands, 1.5 x 0-8 mm., obovate. Stamens 3 4 mm., produced, obtuse. Ovules 2. Berries oblong-ovoid, red, slightly pruinose blue, 9 x 5 mm. ; Style short.

S.E. Tibet: Prov. Tsarong, Dawo, by the temple Belo, west of Tschitse tsung, 13,000 ft., Limpricht 1936 (Type).

N.W. Yunnan: Atuntze, Mt. Miyetzimu, 11,200ft., fr. 18 Oct. 1937, Yu 10560 (E), shrub 4 ft.

Cultivated: from Yu 10560, fl. 7 Apr. 1943, Nov. 1943. Also, at Wisley, fl. 4 May 1944.

(206) Berberis yunnanensis Ranch., BuU. Soc. Bot. Fr. 33, 388, 1886; PI. Delavay. 38, 1889. (Fig. 39)

&‘terns mature very sulcate, finely verruculose. Intemzodap (1.6) 2.5-5 (-7) cm. Spina 3-fld., concolorous, stout, 1.5-2-5 cm. Leaves 3 4 (-6) x (1-) 1.4 (-2) cm., obovate, rounded; cuneate to decurrent petiole 2-5 mm.; entire, rarely 2-3 spinules; venation branched, not reticulate; below scarcely papillose. Flozoers fascicled 2 4 . Pedicels 2 - 5 4 em.; bracts 1-75 mm. Outer sepal8 5 x 2.75 mm., oblong-elliptic. Inner sepals similar 7-8 x 4-5 mm. Petals 5 x 3 mm., obovate, rounded; apex retuse with rounded lobes; glands separate, 1-25 x 0.5 mm. Stamem 4 mm., produced, subapiculate or sub- obtuse. Ovules (1-) 2 (-3). Berries epruinose, oblong-ovoid, 10-12 x (5-) 6-7 mm.

Yunnan: ad collum Yen-tze-hay, prope Lankong, 10,400 ft. (specimen received at Kew, 31 May 1888), Delavay 1660 (Type, K); Atuntze, Doker La, 11,400 ft., fr. 8 Nov. 1937, Yu 7025 (E). ‘ h i t bright scarlet.’

Cultivated (Kew 254): fl. 12 May 1939; fr. 26 Sept. 1940 (K). This is the o d y plant of this species that I have encountered in cultivation. Plants labelled Yu 7925 are B. rnekon- gen&; plants for long cultivated as ‘ B . yunnanensis’ are mostly B. forrestii, very occasionally B. diaphana.

(207) Berberis dictyoneura Schneid., Sargent, Pi. Wils. 1, 374, 1913. (Fig. 45) Shrub 4-5 ft., erect. Stems with young shoots green; mature sparsely verruculose. Spines

3-fld., 1-2 cm., sulcate. Leaves 10-25 x 4-10 mm. ; obovate; obtuse or rounded; cuneate to decurrent petiole 2-3 mm. ; margins 7-15-spinose ( 2 mm.)-serrulate (0.3-0.5 mm.), dis- tant 1-2 mm. ; closely reticulate both sides; above dull grey-green; below yellow-green. InJlorescem 7-14-fld., racemose-subfascided, 1-2 cm. long Pedicek glabrous, 4-6 mm. ; bracts c. 1 mm. Fknners H.C.C. 112. Prophylls acute, 2-5 x 1-25 mm. Outer sepals 4-4.75 x 2.25-2.75 mm., oblong. Inner sepals 5-5-6.5 x 3-54 mm., obovate. Petals 5-5-75 x 3-3.5 mm., obovate, obtuse, entire; base clawed, with discolorous, ovate, acute, separate, non-marginal glands, 1 x 0-5 mm. Sturnem 3-5 mm., produced, subapiculate or subobtuse. Ovules 1-2. Berries pink, or pale red, epruinose, estylose.

W. Szechuan: Min valley, near Sung pan, roadsides, 850&9800ft., 10 Aug. 1910, Wilson 4633 (Type, K).

Cultivated: L.A. 176 (from the type Wilson 4633), fl. 18 May 1938, 16 May 1939, 28 Apr. 1943, 16 Apr. 1945; fr. 1 Oct. 1942, 8 Oct. 1943.

A most attractive medium-sized to small bush, charming in its closely spinulose small grey leaves and pink berries. It may be remarked that botanically with its closely

11 JOURN. LINN. SOC. BOTANY-VOL LVII

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162 L. W. A. AHRENDT spinulose leaves, and geographically in its northern rather than western position in Szechuan, this species tends towards the Subsect. Sieboldiae; it may be regarded tw lying between that subsection and this.

(208) Berberis spraguei Ahrendt, h r d . Chron. (3), 109, 101, 1941 Shrub with a few straggling, sparsely branched stems, 6-8 ft. Stem at Grst green,

soon becoming vividly red; mature pale yellow. Spines 3-fld., concolorous, plane to subsulcate, 3-9 mm. Leaves 10-30 x 4-10 mm.; narrowly oblong-obovate; apex acute; base attenuate to decurrent petiole 2-4 mm.; margins 6-ll-spinoSe (1-1.5 mm.)-sem- late, distant 2-3 (4) mm.; venation branched, scarcely reticulate; above dull; below densely papillose. In$oreaxce (6) 7-12-fld. Pediceh glabrous, (3-) 5-10 (-14 mm.) ; bracts 1-25 mm.

The narrow, acumhte sepals, which mark B. sprqpei, me found in only a few species in widely Merent parts of the genus.

KEY 3 8 ~ Infloregoence 2-3 cm., sessile. Inflorescence P 8 an., peduncled.

208a. B. spmguei var. spraguei 208b. B. spmguei vax. pedu-

(2080) Berberis spraguei var. spraguei Prophylls acuminate, 2-25 x 1 mm. Outer s e p h 4.5 x 2 mm., ovate, mnspicuously acute

or subacuminate. Inner sepl.8 5 x 2-25 mm., ovate, conspicuously mute. Petals 3.5 x 1.75 mm., obovate, ernwginate; glands ovate, obtuse, 0.5 x 0.4 111111. Stanzens 2-75 mm., not produced, truncate. Ovuks 2. Be& oblong, estylom, bright red, epruinose, 8-8.5 x 4-5-5 mm.

Yunnan: seed only, 1932, Forrest 30613. Cultivated: L.A. 75 (from the seed cited), fl. May 1938,3 May 1939; fi. 12 Oct. 1938

(Types, BM3 0).

(208 b) Berberis spraguei vm. pedunculata Ahrendt, var.nov., inflorescentiis elon-

Leaves more often entire. F'2owers H.C.C. 1-211. ProphyyUe 3-3-5 x 1-1-25 mm. Outer sepals 5-6 x 2.5-3 mm., narrowly ovate. Inner se+ 5-6 x 3-3.5 mm., oblong-elliptic, all conspicuously acute or subacuminate. Petals 4-5 x 3 mm., glands oblong, approxi- mate, non-marginal, concolorous, 1 x 0.5 mm. Berries 7.5 x 4.5 111111.

gatis pedunculatisque

Yunnan: 1932, seed only, Forrest 30717. Cultivated: L.A. 107 (fkom seed cited), fl. 9 May 1939, 13 May 1942, 29 Apr. 1943

(Types, BM, 0).

KEY 38c Leavee obtuse to eubute, margins 8-2O-spinose; Momcence 3-8-fld.

Leaves subacute to acute, 25-40-spinos43; Moreaceme 8-18-fld. 20th. B . x & d

208d. B. x Mi- vax. oxyphylla:

(208c) Berberis x declinata Schrad., Linnaea, 12,368,1838 Shrub 7 ft. Stem verruculose. Spine.5 concolorous, sulcate, 1-3-fld., 5-15 mm. Leaves

2-4 x 1-2 cm. (on shoots to 6 x 2.5 cm.), obovate-elliptic; cuneate to decurrent petiole 2-10 mm.; obtuse or subacute; margins 8-15-spinose (0-5-1) mm.-BeITu1&te (0.5 mm.), diatant 3-6 mm.; venation branched; below grey, pruinose, subreticulate. In.e-sci?nce

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Berberis and Mahonia 163

3-!)-fld., 3-5 (-7) cm., including peduncle 1-2 cm., loosely subracemose. Pedicels glabrous, 5-7 (-11) mm.; bracts 1-5-2 mm. Flowers H.C.C. 3. Prophylls red, acute, 2 x 1.75 mm. Outer se@ 3.5 x 2-75 mm., oblong-elliptic. Inner sepah 6 x 5.25 mm., obovate. Petals 6 x 5 mm., obovate, subretuse to subentire; slightly clawed with approxi- make glands, 0.8 x 0.5 mm. Stamens 3-5 mm., not produced, truncate. Ovules 2. Berries epruinose, estylose, obovoid-ellipsoid, 9-10 x 4.5-5 mm. Seeds red-brown.

Cultivated: L.A. 203, fl . 13 May 1942, 16 Apr. 1945; fr. 21 Sept. 1944. This and the following are reputed to be hybrids between B. vulgaris and B. chinensis.

(208d) Berberis x declinata var. oxyphylla Schneid., J. Am. Arb. 4, 212, 1923 heuves 3-6 x 1.2-2.5 cm., or to 5-5 x 1.8 cm.; margins 20-40-spinose (1.5-2.5 mm.)

serrulate (1 mm.), distant 1-2.5 mm. Pedicels 4-7 mm. Flowers H.C.C. 2. Outer sepal9 5.5 x 3 mm. Inner sepals 6-5 x 4 mm. Petals 6.5 x 4 mm. ; glands separate, submarginal, discolorous, 0.8 x 0.4 mm.

Cultivated: L.A. 448, fl. 13 May 1942, 25 Apr. 1945; fr. 21 Sept. 1944. A shrub to some 4 ft., more attractive than the preceding hybrid.

(208e) Berberis x breviscapa (Ahrendt) Ahrendt, stat.nov.; syn. B. issarongensis var.

flpines 6-15 mm., plane. Leaves obovate, 15-25 x 5-9 mm.; margins 1-3 (-5)-spinose (1-1.5 mm.)-serrate (1-1.5 mm.), distant 3-4 mm. ; slightly lustrous, yellow-green; below paler green; venation slight. Inflorescence 443-fld., subumbellate, 1-2-5 cm., including peduncle 3-7 mm. Pedicels 3-5 mm.; bracts acuminate, 2 x 1 mm. Prophylb acute, 2 x 1.25 mm. Outer sepals 3.75 x 2 mm., oblong-elliptic, obtuse. Inner sepals 5.5 x 4.25 mm., obovate. Pettcls 5.5 x 3.5 mm., oblong-obovate, entire; base cuneate, with obtuse ovate glands, 1 x 0.6 mm. StamRns scarcely produced rounded. O& 2 (-3). Berries bright orange red, oblong-ellipsoid, c. 9 x 5 mm., estylose, with fruiting pedicels 5-7 mm.

Cultivated: from plant a t Wisley, inaccurately labelled ‘B. edgeworthiam’, fl. May 1938; fr. 28 Oct. 1938 (Type, BM).

(209) Berberis bracteata (Ahrendt) Ahrendt, stat.nov. ; syn. B. dictyoneura var. bracteata Ahrendt J. Bot., Lond., 80 (Suppl.), 111, 1944; bracteis pedicellis sub- guilongis distinguenda

S t e m very sulcate. Spines 3-0d., 10-15 mm. Leaves 1 7 4 0 x 7-17 mm., obovate, obtuse; cuneate with decurrent petiole, 1-3 mm.; margins 10-12-spinose (1.25-2 mm.)- serrate (1-1.5 mm.), distant 2-5-5 mm.; reticulation elevated; above lustrous green; below grey pruinose. InJlorescence 5-8-0d., subracemose or subfascicled, 16-2 cm. Pedicels glabrous, 5-6 (-9 below) mm. ; bracts lanceolate, acuminate, 4-5 mm. Flowers c. 7 mm. diameter.

S.E. Tibet: Prov. Zayul, Upper Kiukiang valley (Cldung), Choherton, 10,400 ft., 6 Aug. 1938, with flower nearly over, and berries scarcely formed, Yu 19690 (Type, E), open and rocky places and mountain slopes. Not cultivated.

breviscupa Ahrendt, Card. Chron. (3) 105, 372, June 1839

Subsect. Sieboldiae Ahrendt, subsectmov., f o b crebre spinulosis, deciduis, inflorescentiis paucifloratis plusminusve umbellath

Stems glabrous. Internodes (0.7-) 1-2 cm. Spines mostly weak, 6-15 (-20) mm. Leaves deciduous, 1-7 cm. long; margins (6) 20-100-spinose. InJlorescence 3-8 (-15)-fld., um- belhte, or subumbellate, rarely pseudumbellate-subracemose. Pedhls 2-8 (-15) mm. Plowers 5-12mm. diameter. Shmm not produced, truncate, except in B . dubiu. Ovules 1-2. Fruit globose, 6 mm., or mostly oblong, 7-10 mm. long, estylose.

Japan, Quelpaert Island, Mongolia, Shensi, Kansu. 11-2

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164 L. W. A. AHRENDT

KEY 39 to Subsect. Sieboldiae (Map 23 and 26) Mature stems dark, red or purple.

Fruit globose, 5-6 nun. ; leaves with margin ciliate, with 50-100 s p h u l ~ .

Fruit oblong, 7-11 mm.; leaves with 5-25 marginal spinules. 210. B. sieboldii

Leaves reticulate; pedicels 3-6 (-9) mm. Leaves grey, pruinose below; flower diameter 11 mm.; fruit 11 mm. long.

Leaves green, epruinose below; flower diameter 7 mm. ; fruit 7 mm. long. 211. B. reticulata

212. B. Leaves openly veined; pedicels 10-15 mm. 213. B. Oritrepha

214. B.quelpamte& Mature stems pale yellow.

(210) Berberis sieboldi Miq., Ann. M w . Lugd. Bat. 2, 69, 1865. (Fig. 44) Shrub compmt, 2-3 ft. Stem with young shoots green; mature lustrous, subangled.

Leaves 2 - 5 7 x 1-24 cm., rhombic-ovate; apex obtuse; base attenuate to decurrent petiole 2-8 mm. ; margins 50-100 (-150)-ciIiate-~pinulose (1-2 mm.)-subserrulate, distant 0.2- 0.5 mm. ; openly reticulate; above bright green with the young leaves suffused red, or with the veins marked in red; below much paler lustrous green, epapillose. Inji?ore-scem 3-6-fld., umbellate-racemose, 2-3 cm., including peduncle 1-14 cm. Pedkls stout 6 7 mm. Flowers H.C.C. 4/1 to 5/1. Outer se@ 5 x 3-5 mm. Inner sepals 6.5 x 5 mm., all obovate. Petals 5 x 3 mm., oblong-obovate; apex acutely ernargbte with rounded lobes; base clawed, with oblong, separate, margind, concolorouB glrtnds, 0-8 x 0.4 111111. 8tanzenS 3mm. (hula 2, sessile. Berries deep to dark red, W m m . , h, globose, lustrous.

Japan: without locality, Thunberg, (Type.) Hondo, and Hokkaido: on the Rehder Mts., Sargent, 1892. Cultivated: fl. 4 June 1942, 16 May 1940; fi. 12 Sept. 1940,28 Oct. 1942.

(211) Berberis reticulata Byhouwer, J. Am. Arb. 9,132,1928 Shrub 6-8 ft. Stems angled. Spines solitary, to 9 mm. Leaves 2 4 (-6) cm., 9-16

(-21) mm.; obovate; apex obtuse; base with decurrent petiole 2-12 mm.; margins very closely spinose (1 mm.)-denticulate. InfEorascem 5-8-fld., pseudumbellate-subracemose, 1-4-2 cm. Pedkls 6 7 mm.; bracts 1 mm. Inner 8epa.l-s 7 mm. Petah emarghate. Ovules 2, sessile. Berries red, ovoid, epruinose, 11 x 8 mm.

Shemi: seed only, M o m 644, Cultivated: in U.S.A from Purdom’s seed (Type, Am. Arb.). Not in cultivation in

England.

(212) Berberis dubia Schneid., BuU. Herb. Boissier, (2), 5, 663, 1905 Stems subsulcate, slightly lustrous. I & d 6-12 mm. Spines 1-3 (-5)-fld., 7-15

(-20) mm. Leaves 15-30 (40) x 5-15 (-18) mm., including petiole 1-3 mm. ; narrowly obovate, subacuminate; margins towards apex 6-10-spinose (0.51 mm.)-subserrulate, distant 1-2 mm. ; reticulate; both sides deep green and sublustrous, the lower sometimes being slightly paler and more yellowish. Infore-sce.ce 5-8-fld., racemose-subfascicled, 1-16 (-2) cm. Pedkel-s 3-5 (-9) mm. Prophylls 1-5 x 1 mm., acute. Outer sepals 2.5 x 1.5 mm., ovate. Inner 8epcd.s 4.5 x 3.5 mm., broadly obovate, subspathdate. Petah 3-5 x 2.5 mm. Stamens 2.5 mm., produced, shortly apiculate. Ovules 2. Berries 7 x 4 mm. red, estylose.

W. Kansu: 1885, Potanin, s.n. (Type). E. Mongolia: Mt. Alaschau, 1873, Przewalski, s.n. W. Mongolia: Montes Suma-nade, a Kalgen Occid., versus, fl. 1871, Przewalski,

an. (K).

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Berberis and Mahonia 165

Cultivated (Kew 257): f l . May 1939; fr. Oct. 1939. This, the only plant of its species which I have seen in cultivation, deserves the most careful preservation; the bush was already very ancient when I last saw it in 1950. I imagine that younger plants of this urmsual northern species would be quite attractive as garden plants. No. 257 has defied every attempt that I have made to propagate it vegetatively ; but I am very grateful to the authorities a t Kew for the generous material given to me for this purpose over many years.

(213) Berberis oritrepha Schneid., Ost. Bot. 2. 67, 293, 1918 Stems sulcate. Spines 1-3-fld., 1.5-2-5 cm. Leaves 12-25 x 5-10 mm., excluding

petiole, 8-13 mm. ; elliptic-obovate ; base attenuate ; margins densely and slenderly 5-15- spinulose; above openly veined; below openly reticulate, grey, pruinose (but on the new shoots subentire or remotely spinose, and to 3.5 x 1.5 cm.). Inflorescence 5-10 (-15)-fld., umbellate-racemose, 1-3.5 cm. Pedicels 10-15 mm. ; bracts 1-1.5 mm. 0vzrle.s 1-2. Berries 10 x 5 mm., obovoid-ellipsoid, red, epruinose, estylose.

Shensi: Tai-pai-shan, 8 Feb. 1911, seed only Purdom 592. Cultivated: in U.S.A. from Purdom’s seed (Type). Not cultivated in England.

(214) Berberis quelpaertensis Nakai, Bot. Mag., Tokyo, 27, 31, 1913 ,Shrub 5-6ft. Stems with young shoots red, becoming red brown, but when mature pale

yellow, angled or sulcate, scarcely vermculose. Spined concolorous, 3-fld., 5-15 mm., sul- cate. Leaves thick, to 6-5 x 2.5 cm., oblong-obovate; base contracted to decurrent petiole, 2-6 mm.; subacute to subobtuse; margins 20-35-spinose (1-1.5 mm.)-serrate (0.5 mm.), distant 1-16 (-2.5 mm.); closely reticulate; dull deep green; below much paler sub- lustrous green. In$brescence 8-12 (-15)-fld., loosely racemose or subracemose, with stout rachis. Pedicels 4-6 mm. Flowers H.C.C. 1/1-1/2. Prophylls red, truncate, acuminate, 1-5 x 1 mm. Outer sepals 2.5 x 1.5 mm., oblong-obovate, subacute. Inner sepals 5-5 x 3-5 mm., obovate. Petals 5.5-5.75 x 3-5-3.75 mm., obovate; apex acutely e m a r e a t e with acute lobes; glands orange, elliptic, separate, non-marginal 1 x 0.6 mm. Ovules 2. Berries oblong, red, epruinose, estylose, 7-7.5 x p 4 - 5 mm.

Quelpaert Island: Mt. Hallaisan, 3300 ft., and above, fl., Faurie 1674, and fr., Ishidoya, s.n., Types: 17 Aug. 1938, fr., Lecittaquel, s.n. (K); seed only Yashiroda, s.n.

Cultivated: L.A. 398 (from Yaahiroda’s seed, raised by W. J. Marchant, Wimborne, Dorset), fl. 28 Apr. to 6 May 1943, 16 Apr. 1945; fr. 14 Oct. 1943. This is an attractive species for cultivation, being stif€ and compact with stout stems. Its tendency to have a racemose inflorescence, and yellow mature stems associate this species with the Sect. Vulgares, and it may well be regarded as a link between that section and this sub- section. Since this subsection, as a whole, connects the Franchetianae (typically with entire leaves and few-flowered subumbels) to the Vulgares (typically with closely spinose leaves, and many-flowered racemes), B . quelpmrknsis is the extreme connecting species between the two.

Sect. Tschonoskyanae Schneid., Bull. Herb. Boissier, ( Z ) , 8, 215, 1908; syn. Sect. Heteropodae subsect. Tschonoskyanae Schneid., l.c., ( Z ) , 5, 458, 1905

Stems glabrous; mature, dark, red or purple (yellow only in B. virgetomm, B. mique- liama, and hybrids). Internodes (1-) 16-2.5 (4) cm. Spines mostly 3-10 (-25) mm. Leaves 0.5-10 cm. ; base conspicuously petiolate ; margins nearly always entire (rarely with 1-8 inconspicuous spinules). In$orescence 2-8 (-14)-fld., fascicled, umbellate or sub- umbellate, very rarely solitary. Pedicels 4-9 (-20) mm. Flowers 5-10 mm. diameter. 0vzr.k~ 1-2. Berries red, estylose, 7-10 mm.

Japan, Kiangsi, W. Szechuan.

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166 L. W. A. AERENDT

-Y 40, to Sect. Tschonoskyanae (Maps 3, 23 and 27) Leaves lustrous above; below concolorous green, epapillose. Leaves dull above; below pale, grey, pruinose.

215. B. nundhcam

Leaves 6 1 0 cm., longer than infloresoence (2-3.5 cm.); pedicels 3-6 (-8) mm.; petals shorter than inner sepals.

Leaves 2-4 (-5) cm., mostly shorter than the inflorescence; but if longer than in- florescence, then with petals longer than inner sepals; pedicels (&) 8-20 mm.

216. B.&g&wum

Inflomicence 1-2 cm. ; petala longer than inner sepals.

Stems dark red; leaves entire; petals emarginate. Stems pale yellow ; leaves sphose ; petals entire.

stemene dentate. 217. B. thunbergG

218. B. x ottiaweda Stamem dentate. 219. B.miqu&aw

Inflorescence (3-) 6 6 (-7) cm.; petals shorter than inner sepgls. Stem sulcate; leaves typically spinom; fruit 9-8 mm. 220. B. SilUa-Wmm Stems terete ; leaves typically entire ; fruit 9 x 6 mm. 221. B. tadwmakgana

(215) Berberis mouillacana Schneid., Sargent, PI. Wik . 1,371, 1913 Shrub 6-10 ft. Stem sulcate. Spines absent, or solitary, terete, yellowish, 3-12 mm.

Leaves 1-46 (-6) x 0.5-2.25 (-2.5) cm., or to 5 x 3.5 cm., including decurrent petiole 8-14 (-20) mm.; obovate; base contraded to petiole; margins entire, or 1-8 incon- spicuously spinulose; apex rounded; venation branched above, but below openly reti- culate. InJEorapcence 4-8 (-12)-fld., racemose-subfascicled, umbellate-racemose or loosely racemose (%) 3-5 cm. FEowers H.C.C. 602. Pedicds glabrous, 6-15 mm.; bracts 1.5- 2 mm. ProphgZk, lanceohte, acuminate 3 x 1 mm. Se@ (3), elliptic, acute, 4-5 x 1.5mm. Petals 4 x 2 mm., elliptic, acute, entire. Stanzens 3 mm., truncate. ovules 2. Be& ovoid-ellipsoid, epruinoae, 9-10 x 5-7 mm.

W. Szechuan: Tachienlu, thickets, 8800-11,700ft., 1908, Wilson 1039 (Type, K); Wilson 1041,1283 (K).

Cultivated: at Kew (52-12, Veitch), Wilson (Veitch) 1707, fr. Nov. 1938; fl. Apr. 30 1954; Kew 724, fl. 30 Apr. 1954.

(216) Berberis virgetorum Schneid., I.c., 3, 440, 1917. (F'ig. 46) Shrub to 6ft. Stems slightly angled, slender, pale yellow. Spines solitary, sulcate,

1-26 cm. Leaves 36-10 x 1.5-3-5 (4) cm., including petiole 1-2 cm.; oblong-rhombic; base attenuate to petiole; apex subacuminate, subobtuse; margins completely entire, sometimes slightly undulate; only a few elevated veins visible; above deep d d yellow- green; below epapillose. InJlorecew (%) 5-10 (-14)-fld., subracemose to subumbellate, 2-36 cm., including peduncle, 1-2 cm. PediceZa slender, p6 (-8) mm. ; bracts lanceolate, acuminate, 1-1.5mm. outep. sepals 1.5-2mm., oblong-ovate, acute. Inner sepals 4.2 mm., oblong-obovate, obtuse. Petals 3-36 x 1-1-75 mm., elliptic-obovate, obtuse entire; glands separate, oblong. Stanaens 3 mm., obtuse. Ovules solitary, sessile. Berries (8-) 9-12 x (3-) 4.5 mm., oblong-ellipsoid, occasionally slightly bent, firm, lustrous, quite estylose, a somewhat greyed shade of H.C.C. 016, or H.C.C. 014, ripening to H.C.C. 822.

Kiangsi: Khuling, thickets, 4600 ft., fi. 29 July 1907, Wilson 1517 (Type, Am. Arb.). Cultivated: L.A. 576,30 Bug. 1946,ll Mar. 1954, without flower or fruit; fl. May 1955;

fr. 28 Sept. 1955. !Chis most unusual species is valuable in gardens solely for its remark- able foliage.

(217) Berberis thunbergii DC., Syat., 2,9 , 1821 Shrub dense, compact. Stems with young shoots green, glabrous; mature dark red, sul-

cate, not verruculose. Internodes 7-15 mm. Leaves 5-20 x 2-16 mm., including decurrent petiole, 3-12 mm. ; rhombic-ovate; apex obtuse (rarely acute) ; base contracted to petiole; margins completely entire; subenervate; above dull bright yellow-green; below grey,

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Berberis and Mahonia 167

papillose (rarely green). In&rescence (1-) 2-5 (-lZ)-fld., umbellate-subfascicled, 1-2 cm. (rarely solitary). P e d i A 5-9 mm.; bracts 1-1.5 mm. Prophylls red, acute, 2-25 x 1 mm. Outer sepals 4 4 . 5 x 26-2-75 mm., ovate, subobtuse, partly red. Inner sepals H.C.C. 112, 5-5-5 x 3.25-3.5 mm. Peta2s 5.5-6 x 3 .754 mm., H.C.C. 1/2; oblong-obovate; apex slightly emarginate ; base scarcely clawed, glands approximate, non-marginal, 1-1-2 x 0*6-0-7 mm. Stctnaens 3-3-5 mm., not produced, truncate. O v d a 1-2, sessile. Berries firm, lustrous, ellipsoid.

KEY 4 0 ~ Skims spreading horizontally.

IAaves green, except in autumn; fruit deep red, 7-8 x 4-4-5 mm. (or somewhat amaller in var. minor).

Leaves obtuse, grey below. Height 3-5 ft. ; fruit 7-8 mrn. ; leaves mostly 1-2 ern.

Inflorescence 2-5-fld. 217a. B. thunbergii var. thunbergii Inflorescence solitary or 5-12 fld.

Inflorescence 5-12-fld. 217b. B. thunbergii var. pluri@ra Inflorescence solitary. 217c. B. thunbergii var. uniJora

217d. B. thunbergii var. minw Leaves acute, green below. 217e. B. thunbergii var. m x i m w i c z i i

Leaves on new shoots margined white ; berries larger and very bright red.

Leaves red or purple; fruit deep red, 7-8 x 4-44 mm.

Height 1 ft.; leaves and fruit smaller.

Leaves always otherwise coloured or variegated.

217f. B. thunbergii var. argenteo-margiwta

Leaves dark purple. 2179. B. thunbergii var. atropurpureu Leaves brighter red. 217h. B. thunbergii var. rubrjfolia

Sterns suberect. 218j. B. thunbergii var. erecta

(21'7 a) Berberis thunbergii var. thunbergii Japan: Yokohama, 1862, Maximowicz, 8.n. (K). Cultivated: one of the most widely cultivated of Berber&%

(21'7 b) Berberis thunbergii var. pluriflora Koehne Deutshe Dendr. 169, 1893; Rehd., Mitt. Dtsch. Dendr. Ces. 13, 33, 1898; Not cultivated.

(21'7c) Berberis thunbergii var. uniflora Koehne, 1.c. 1893; syn. B. leichtlinii Hort. Chltivated: L.A. 76; fl. 13 May 1942; fr. 26 Nov. 1940.

(21'1d) Berberis thunbergii var. minor Rehder, l.c., p. 330, 1898; syn. B. akwsonii, Bean, Trees and h'hrub8, 1, 250, 1914

(217e) Berberis thunbergii var. maximowiczii (Regel) Regel Acta. Horti. Petrop. 2 , 420, 1878; syn. B. maxinwwiczii %gel, aartenjl. 21, 238, 1872

Cultivated: distributed by W. J. Marchant, Wimborne, Dorset.

(21i'f) Berberis thunbergii var. argenteo-marginata Schneid., J. Am. Arb. 4, 221, 1923

Berries larger than in preceding varieties, 10 x 6 mm., and very bright red. Cultivated: L.A. 58, fr. 12 Sept. 1940. The variety argenteo-murginata is well worth

growing, in addition to the common one, on account of its brightly variegated new leaves and very bright berries.

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168 L. W. A. AETEENDT

(217g) Berberis thunbergii var. atropurpurea Chenault in Rev. Hort (N.S.), 20, 307, 1926, foliie atropurpureis

Cultivated: L.A. 33, fl., fr., 1937, A variety which deaerves its widespread cultivation.

(217 h) Berberis thunbergii var. rubrifolia Ahrendt, var.nov., foliis rubris Cultivated: L.A. 33A, fl., fr., 1937. This variety often passes under the name of the

preceding; but the two plants are very distinct in leaf colour, and planted side by side present a pleasing contrast.

(217j) Berberis thunbergii var. erecta (Rehd.) Ahrendt; syn B. thuderg& f erecta Rehder, J . Am. Arb. 20, 412, 1939; syn. B. thunbergii pluriJlora erecta Anon., in Horticulture, 13, 48 (fig.), 1935

Not cultivated in England. I have not seen this plant, but feel that its erect habit would spoil the characteristically attractive form of the species which lies in its dense habit composed of horizontal branches. For purposes of cultivation, I should recom- mend concentrating on numbers 217a, f, g , h.

This species is marked out from most other Berberis by the fact that garden seedlings, self-sown, even from gardens stocked with many Berberis, produce plants as character- istic of their parents as if vegetatively propagated. However, a hybrid has been artifi- cially raised.

(218) Berberis x ottawensis Schneid. in Silvlt Tarouoa, Schneid., Unsere Freihnd- Laubg., ed. 2, 116, 1922; J . A m . Arb. 4,221,1923

Shrub 2-4 ft., habit intermediate between its parents, B. thunbergii and B. vulgaris. Stem yellow. Leaves 1.5-3-5 x 0.6-1.5 cm. ; obovate-rhombic; apex obtuse; base con- tracted to petiole 1-6 mm.; entire or sparsely spinose; openly veined. Inflorescence 3-15-fld. ; subumbellate, 2-5 cm., including peduncle about 2 cm. Pedkd.9 4-6 mm. ; bracts 1-5-2 mm. ProphyUs small. Inner sepah 3-4 mm. Petals 4-5 mm., entire. Stamens 3.5 mm., truncate. ovules 2, sessile. Berries ellipsoid, 9 x 5 mm., estylose, bright lustrous red, persistent as in B. thunbergii.

Cultivated: by Dr Saunders of Ottawa Experimental Station, raised artificially.

(218a) Berberis x ottawensis var. purpurea Schneid. ex Rehder, Man. Cult. Trees and Shrubs, 246, 1927

As in (218), but wi6h purple leaves. Cultivated: by Dr Saunders of Ottawa. Neither of these hybrids is cultivated in

England. "he influence of B. dgaris is shown in the yellow stems, occasional leaf spinules and the bright berries. The influence of B. thunbergii is shown in the leaves having few spinules or none, and in the inflomcenee being fewer flowered and more umbellate than racemose.

(219) Berberis miqueliana Ahrendt, sp.nov., aff. B. silva-taroucam sed ramulis

Shrub spreading, to 6 ft. Sterns sulcate; young shoots red; mature pale yellow-grey. Xpine.9 absent, or weak, 3-8 mm. h u e s 10-20 (-25) x 7-12 (-15) mm., including petiole, 3-8 mm. ; broadly obovate-elliptic; margins 5-8-spinose; apex rounded; venation branched. Inflor~cenee 2-5-fld., subfascicled, 1-5-2 cm. P e d i d slender, 10-15 mm. Outer sepals 2-2.5 x 2-2.5 mm., suborbicular, reddish. Inner sepal8 6.4-5 mm., obovate, concave. Petab 7x4mm., convolute. Stamem 3.5mm., below the anthers conical- dentate; apex not produced, truncate; basal glands discolorous, orange, contiguous or approximate. Ovuks2.

pallide flavis, staminibus dentatis distinguenda

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Berberis and Mahonia 169

Chltivated: L.A. 294, fl. 6 Apr. 1945 (Type, 0). This remarkable plant was distributed as B. silva-taroucanu by W. J. Marchant, of Wimborne, Dorset; it differs in its pale yellow stems, petals longer than inner sepals, smaller leaves, and particularly in the dentate stamens, found otherwise in the S. American species.

(220) Berberis silva-taroucana Schneid., Sargent, PI. Wih. 1, 370, 1913 Shrub 5-loft. Stems sulcate. Spines absent, or solitary, 3-7 mm. Leaves (1-5-)

2-5-5 x (0.5-) x 0.8-3 cm., excluding petiole 1-2.5 cm. ; base abruptly contracted to petiole; oblong-obovate; apex rounded or obtuse; margins entire, or rarely with 3-10 inccrnspicuous spinules ; openly reticulate. Inflorescence 612-fld., loosely subracemose or subumbellate, 3-7 cm., with peduncle nil or short. Pedicels 8 mm. above to 2 cm. below; bracts 1-1.5 mm. Prophylls red, lanceolate, 2 x 0.8 mm. Outer sepals 4 x 3 mm. Inner sep& 6 x 4.5 mm., obovate. Petals 4.5 x 3.25 mm., obovate, subentire; glands separate, non-marginal, 0.7 x 0.4 mm. Stamens 3-5 mm., apiculate. Ovules 2, sessile. Berries ovoid-globose, 9-10 x 7-9 mm., estylose.

W. Szechuan: Chiu-ting-shan, thickets 6500 ft., 23 May 1908, Wilson 2860 (Type, K) ; west of Kuan hsien, summit of Niu tu shan, 10,OOO ft., Wilson 2857 (K) ; Wa-shan, thickets, 8500 ft., Wilson 2858 (K); west and near Wenchuan hsien, 7OOO-10,000 ft., Wilxon 1059 (K).

(221) Berberis tschonoskyana Regel, Acta Horti Petropol. 2 , 421,1873; syn. B. s i b -

Stems terete. Internodes 24-3.5 cm. Spines absent, or only 2 mm. Leaves to 5.5 x 1.2 om,, obovate; base attenuate to decurrent petiole, 4-7 mm.; margins entire; apex acute. Inflorescence 5-7-fld., loosely racemose, c. 5 cm., including peduncle 1.5-2 cm. Pedicels 12-17 mm. Otherwise unrecorded.

Japan: Nippon, in Alpe Niko, 1864, Tschonoski, s.n. (Type, 9); Maximowicz, s.n. (K). Not cultivated.

kiana Yatabe, Tokyo Bot. Mag. 5, 283, 1891

KEY 41. General Key to Sects. Canadenses, Sinenses, Integerrimae, combined Ovules 3-4.

Loaves below grey, pruinose; margins spinose. 222. B. canadensis Loaves below green, epruinose, entire.

Spines weak, 3-7-Ad.; leaves narrow, 1 : 3; fruit globose, scarlet, lustrous, epruinose.

Spines strong, solitary; leaves broad, 1 : 1.5-2; fruit oblong, dull, dark red, pruinose 223. B. fendleri

blue. 234. B. integerrim Ovules 1-2.

Loaves narrow, 1 : 4-10. Leaves very narrow, 1 : 8-10; fruit black, pruinose blue. Leaves less narrow, 1 : 4-6; fruit red.

224. B. lepidifolia

Pedicels 1-6 mm. Bracts longer than pedicels; leaf margins 5-15-spincde. 230. B. pUTdOT?&ii

Bracts shorter than pedicels; leaves entire. Pedicels 1-3 mm. ; bracts subequal ; fruit globose, 4 mm. Pedicels 3-6 mm. ; bracts 1-3 mm. ; fruit oblong, 7-9 nun. long.

229. 33. vemae

Stems sulcate; fruit estylose. 228. B. poiretii Stems terete ; fruit stylose. 231. B. caroli

Pedicels 12-25 mm. 225. B. pallens Leaves broader, 1 : (1-) 1.5-3 (-3.5).

Spines absent or weak, 1-9 mm.; leaves openly veined to subreticulate; inflores-

Spines stout, (1-) 2-5 cm.; leaves more or less closely reticulate; inflorescence cence subumbellate. Sect. SINENSES

racemose. Sect. INTEGERRIMAE

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170 L. W. A. AERENDT

Sect. Canadenses Ahrendt, sect.nov., aff. Sect. Sinemibus, sed ovuli8 3 4 , et fructibus solidis, nitidis, disthguenda; syn. Sect. S i n e m , Subsect. Eusinenses Schneid., BuU. Herb. Boissier, (2), 5, 463, 656, 1905, for the lesser past, compriaing those species already botanically distinguished, and native to N. America.

Stems glabrous, dark, red or purple. Intenzodes (1-) 1-5-2-5 (-4) cm. Spines con- colomus, 3-7-fld., 5-12mm. Leaves deciduous, 2-6cm. long; margins entire or with 1-6 spindei; subenervate or but slightly veined. In$oresmm 8-15-fld., racemose, 2-5 cm. long, including peduncle 1-3 cm. Ped+ P 1 0 mm. P k r 8 7-9 mm. diameter. Pet& entire or subentire, alwap shorter than inner sepals. Stumm not or scarcely produced, obtuse or truncate. ovules (2) 3-4. Be& firm, scarlet, lustrous, estylose, 6-9 111111. long.

U.S.A. : Virginia, Missouri, New Mexico, Colorado.

KEY 42, to Sect. Canadensea (Map 2) Stems angled; leaves with margin spinose, below grey and pruinose.

Pedicels 6-10 mm.; fruit ellipsoid, 8 x 6 mm. Pedicels 4 4 mm.; fruit subglobow, 6 x 5 mm.

222. B. c a d & 222 a. B. x rehderiana

223. B. fendhi

The hybrid, B. x notubilis, may be confused with this section on account of some of its characters, but is eady distinguished by its pale yellow stems and berries 15 mm. long.

Stems terete; leaves entire, below green and epruinoee; pedicels 4-6 mm.; fruit 6 x 5 mm.

(222) Berberis canadensis Mill., &rd. Did. ed. 8, Berberis no 2, 1768; Loud., Arb. et F W . Britt. 1,303, 1838

Shrub 4 4 ft. Sterns dull, densely verrucdose. Leavea 2-5 x 0.6-14 cm.; oblong- obovate; base attenuate to petiole, 3-6 mm.; apex rounded; margins with 1-9 spinules, distant 3-6 mm.; below epapdlose. InfloreLlcence racemose. Pedic4.A~ 5-10 mm., slender; bracts 1-1-5 mm. Prophylls ovate, acute, 1-25 x 1 mm. Outer s e e 2-2-5 mm., elliptic. Inner apah 4-5mm., obovate. Petals 3*5-4mm., oblong-obovate. Stumns 3mm. Ovules sessile. B e d 8-9 x 6-7 nun., ellipsoid.

Virginia: fr. 1850, Gray, s.n. (K). (S.W.), Smyth County, Middle Holston valley, bluffs on middle fork of Holston river,

near Marion, fl. 20 Jdy 1892; fr. 22 May 1892, Small, 8.n. (K). Missouri: (S.E.), Shannon County, high rocky cliffs, uncommon, fr. 21 Oct. 1893,

Bush 9 (K); Monteer Mts., high hills, f?. 18 Oct. 1905, Bush 3601 (K); fl. 12 May 1926, Russell 21,24,371 (K).

Nuttall, in Gen. Amr. PI., 1,211,1818, gives the distribution as AUeghany Mts. from Canada to Georgia; also in Tennessee where it appears almost semi-evergreen.

Cultivated: a t Kew, fl. May 1950; the true species is rarely to be found in cultivation now.

(222a) Berberis x rehderiana Schneid., Bull. Herb. Boissier, (2), 5, 659, 1905

Stem dull. Spines 3-5-fld. Leaves 1-18 (-3-3) x 0.4-0.8 (-1.4) cm.; oblong-obovate; contracted to petiole 2-8 mm. ; margins '7-11-spinose (1 mm.)-sedate, distant P 8 mm. ; below epapillose. Pedicels and fruit as in B. fendleri.

Cultivated in U.S.A., and stated by Rehder to be B. fendlerixB. canadensis. The stems and leaves are similar to B. canadensi-s, and the spines between the two species, so it bears strong evidence of being so. It is not cultivated in England.

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Berberis and Mrthonia 171

(223) Berberis fendleri Gray, Mem. Amer. A d . Arts & Sci. (N.S.) 3, 5, 1848 Shrub 3-5 ft. Stems lustrous. Spines slender, 3-7-fld. Leaves 2-5 x 0-6-1-6 cm. ; nar-

rowly oblong-obovate ; sessile, subacute ; entire ; above sublustrous bright green ; below palw green. Inflorescence subumbellate, or corymbose-racemose. Pedicels 4-7 mm. Fbmers H.C.C. 212. Prophylls red, ovate, acute, 2.25 x 1.25 mm. Outer sepals 2.25- 2.5 K 1.75-2 mm., oblong-elliptic. Inner sepals 4-5-5 x 2-5-3 mm., obovate; base slightly clawed, with discolorous orange glands, separate, marginal, 0.9 x 0.5 mm. Stanzens 2 6 - 3 mm. 0m.h stipitate, the stalks being from one half as long as, up to as long as, the ovule. Berries oblong-globose, 6 x 5 mm.

Kew Mexico: Santa Fe County, in canyon, 4 miles east of Santa Fe, 8000 ft., Heller 3607 (K). ‘Authenticated specimen from type locality’; Taos, 1869, Thomaa, s.n. (K); 1847, Fender 15 (Isotype, K).

South Colorado: La Plata County, Arborlea, 6000ft., June 1899, Baker 377 (K); Durango, 20 July 1898, Baker, Earle & Tracy, s.n. (K). ‘Common bush, 3-5 ft.; com- mon also at Mancos.’

Cultivated: L.A. 233, fl. 6 May 1942; fr. 30 Sept. 1942.

Sect. Sinenses Schneid., Bull. Herb. Boissier, (2), 5, 463, 1905 in part Stems glabrous, sulcate; mature dark red, purple, or red-brown (only in B. forrestii

are the mature stems yellow). Ink& (1-) 1.5-3 (4) cm. Spines nearly always weak, 4-9 (-14) mm., rarely on the shoots stout, and to 2-5 cm. Leaves 1 . 5 4 cm., entire, rarely with a few inconspicuous spinules ; venation loosely branched, occasionally incon- spicuously openly reticulate. Injlmeescence (6) lO-ZO-fld., more or less subumbellate- racemose, (2-) 3-8 (-12) cm. long, including peduncle (3 mm.-) 1-3 cm. Pedicels 3-17 mm. Flower diameter, 4-10 mm. Petals emarginate. Fruit bright or dark red and mostly estylose (only in B. kpidifolia black, pruinose blue, and shortly stylose).

Caucasus, Yunnan, Szechuan, Chihli.

KEY 43 to Sect. Sinenses (Maps 4 and 29) Leaves very narrow, 1 : 8-10; sepals in one series; fruit black, pruinose blue, shortly

Leaves less narrow, 1 : 3-7; sepals in two or three series; fruit red, mostly estylose. stylose. 224. B. lepidqolia

Leaves entire; petals shorter than, or aa long as, inner sepals. Leaves grey, pruinose below.

Leaves narrow, 1 : 5-6, reticulate; style short. Leaves broad, 1 : 3, not reticulate; style absent.

Leaves fairly narrow, 1 : 3-4; pedicels 7-15 mm., much longer than bracts; petals as long as inner sepals; fruit dark red; leaves paler below.

Leaves narrower, 1 : 4-7; pedicels 3-6 mm., not more than twice aa long aa bracts; petals shorter than inner sepals ; fruit bright red; leaves concolorous below.

225. B. pallens 226. B . forreatiit

Leaves green, epruinose below.

227. B . chinensis

228. B. poiretii 228a. B. x durobrivensia Loaves with spinose margin; petals longer than inner sepals.

N.B. If the leaves are broad and the red berries stylose, cf. B . franchetiam var. m r o b o t r y s . If the stems are terete, cf. B. bretschneideri.

(224) Berberis lepidifolia Ahrendt, Kew Bull. 1939, 269, 1939. (Fig. 41) Shrub suberect, 6-8 ft. Shoots green. Internodes 8-20 mm. Spines solitary, 3-6 mm.,

at base of stems sometimes 1-2 cm., often absent, Leaves 17 x 1.5-2 to 45 x 6 5 mm., narrowly oblanceolate, acute ; base slenderly attenuate, sessile ; margins entire, slightly

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172 L. W. A. AHRENDT subrevolute: lateral veins few: above dull deep green; below grey, pruinose, papillose. Inflorescence 5-8-fld., umbellate or subumbellate, 2-45 mm., including slender peduncle, 1.2-3 cm., with lanceolate, acuminate bracts, 1-5 x 0.7 mm. towards the middle of the peduncle. Pedicek (5-) 7-10 mm.; bracts ovate, 1 x 0-6 mm. Flowers c. 4 mm. diameter. Sepals in one series of three, ovate, subacute, 2-2-5 x 1.3-1.5 mm., red down the centre with a yellow margin. Pekcls 2.5 x 1-25 mm., oblong-elliptic; acutely emarginate with acute lobea; base cuneate with ovate-elliptic, obtuse glands, 0-5 x 0.2 mm. Stawns 1.75 mm. Ovules 2, shortly stipitate. Berries ovoid, 5-5-6 x 4-5-5 mm., black, slightly pruinose blue, with short style.

Yunnan: Chien-chum-Mekong divide, 10,700 ft., Forrest 23614 (Type, K). S.W. Szechuan: betweenyungpoh, and Yungning, 10,OOO ft., 1922, Forrest 22051 (K). Cultivated: from Forrest 23614, fl. 13 May 1938; fr. 26 Oct. 1938 (K). Also at Wisley,

fl. 4 May 1944; this specimen was labelled Yu 13744, but the herbarium sheet of this number is the very different B. m h q e n s i s . This is a most unusual species in flower structure, and a most ornamental garden plant on account of its very graceful leaves which are distinctly persistent in the autumn, but finally disappear by the new year.

(225) Berberis pallens Franch., Pl. Delavay. 36 1889; Schneid., 8st. Bot. Z. 67, 214,

Leave.9 3-5 cm. x 5-10 mm., oblanceolate, attenuate, subsessile, entire, obtuse, more or less reticulate, papillose below. Inflorescence umbellate racemose, 4-6 cm. Pedia3l-s 15 mm., above to 25 mm. below. Prophylls or outer sepals ovate-lanceolate, 4-4-5 x 1.5mm. (according to Franchet). Median and inner sepals equal, broadly ovate, 6-6.5mm. long (according to both Franchet and Schneider). Pekcls about 5*5mm., emarginate. Stamens produced, shortly apiculate. ovules 1-2. Style distinct, short.

Yunnan: ad collum Lo-pin-chan, supra Lankong, 10,400 ft., fl. 25 May 1886, Delavay, s.n. (Type). The species is not in cultivation.

The Kew herbarium specimen of this place and date has a fascicled 2-5-fld., in- florescence, with pedicels 3-6 mm. long, instead of a 10-20-fld. open raceme with pedicels 1-5-2.5 cm. as described. There has, obviously, been some confusion, and the description of this species must rest upon the description by Franchet amplified by that of Schneider. There has also been confusion between this species and B. fmmti i which comes from an adjacent region of Yunnan. B. fmestii has flowers of similar structure to those of B. pallens, but differs in its broad, slightly veined leaves, paler stems, and lack of style.

1918

(226) Berberis forrestii Ahrendt, Card. Chron. (3), 109, 101 Mar. 1941 ; J. Bot., Lond.

Stems loosely arching; young shoots bright red, becoming yellowish red, finally yellow when mature. Leave.9 3.5-6 x 1-14 cm., oblong-obovate; apex rounded; somewhat contracted to petiole 4 4 mm. ; venation slight, or a t most loosely branched. Infireseeme racemose subumbellate, (6-) 8-12 cm. long, including peduncle 1 4 cm. with stout rachis. Pedicels slender, 7-10 (-20, below) mm. ; bracts acuminate, 2-2.5 mm. Flowers 8-10 mm. diameter. Prophylls red, triangular, 3 x 1 mm. Outer sepals (H.C.C. 2), 4.5-5 x 1.5 mm., lanceolate. Median and inner sepals both 5-6 x 3 4 mm., oblong- obovate. Petals P 5 x 2-3 mm., obovate-elliptic; apex distinctly acutely emarginate with acute lobes; base cuneate, with glands 1 x 0.8 mm., approximate but not con- tiguous. Stamens 2.5-3 mm., apex produced, shortly apiculate. Berries bright red, oblong-ovoid, (9-) 10-11 (-12) x (6) 7-8 mm., epruinose, estylose.

Yunnan: on the west flank of the Lichiang range, Oct. 1918, Forrest 17143 (Type, K); 10,OOO ft., fl. June 1906, Forrest 2271 (K); and East flank, 10,00&12,OOO ft., May 1910, 10-15 ft., Forrest 5553, 5554 (K). Recorded by Forrest as growing 6-10 ft., and in the

80 (Suppl.), 108, 1944. (Fig. 40)

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Berberis and Mahonia 173

last two specimens 10-15ft., in open situations in meadows and pine forests and on Limy pastures; flowers canary yellow (2271), or golden yellow (5553, 5554).

Cultivated: from the type, Forrest 17143, fl., fr., 1939, at Wisley; L.A. 108, fl. 13 May 1942; fr. 12 Oct. 1941; from L.A. 237, a plant labelled Yu 10819, fl. 6 June 1941 (and inner flower components, especially petals recorded as of colour H.C.C. 4), 20 May 1942, 27 ]May 1943; fr. 7 Oct. 1943.

These collections of Forrest 2271, 5553 and 5554, have, at various times, been deter- mined on herbarium sheets, as B. pallens and B. lewmtei, while 17143 was originally misnamed B. yunnanensis. Accordingly, in cultivation, these plants have been distri- buted as ‘pallens’ or ‘yunnanensis’, the former name being more common. The late Mr W. J. Marchant informed me in a letter of 11 Dec. 1942 that: ‘the plants called pallens were distributed by Messrs Bees about 1911, raised from seed collected by Forrest on behalf of the late Mr A. K. Bulley. It would appear, therefore, that the original distribution was derived, probably, in connexion with the collections 2271,5553, 5554, and long predated the type collection. The original use of the name pallens for this species is clearly due to the uncertain nature of Franchet’s species, combined with marked similarities 60 some of its known features. However, the marked differences, and the confusion concerning the complete nature of B. pallens do not allow me to include Forrest’s specimens within the same specific boundaries as Delavay’s. To do so, in a case where doubt like this arises, would cause more confusion in the understanding of the genus than to allow a clearly understood unit, such as B. fmestii, to lie beside a confused conception, such as B. pallens, until such time has elapsed as to provide a great increase of relevant material which can resolve the problem.

(227) Berberis chinensis Poir., E m y d 8, 617, 1808, syn. B. sinensis DC., Syst. 2 , 8, 1821; Desf., Tabl. Eeole Bot. p. 150, 1804, et ed. 2, 175, 1815, nomina nuda, et Hist. Arb. Arbriss. 2, 27, 1809; Hook. f. et Thorns., Bot. Hag. 107, t. 6573 1881; syn. B. iberica Stev. et Fisch., ex DC., 1.c. ; syn. B. serotina Lange, I d . Sem. Hort. Bot. Haun., 1872, 5, 1872; Bot. Tidskr. (2) 2, 138, t. 4 (1879)

Shrub, fairly erect, 5-6ft. Stem slightly lustrous. Leaves 15-30 (40 ) x 5-10 (-13) mm., obovate, attenuate to petiole 1 mm., entire, obtuse. Inflorescence racemose-subumbel- late, 4-6 cm., including peduncle 5-10 mm. Pedicels 4-15 mm.; bracts 2-3 mm. Pro- phylls red, 1.5-2 mm. Outer sepals 3-3.5 x 1-16 mm., ovate, acute. Inner sepals 4-5 x 2 .753 mm., obovate. Petals 4-5 x 2-2.5 mm., oblong-obovate; shortly clawed, with dis- colorous orange glands. Stamens 3-36 mm., not produced, truncate. Ovules 1-2, sessile. Berries ellipsoid, 9-10 x 4-5 mm., dark red, epruinose.

KEY 43A Spines 5-9 mm., weak; pedicels 7-15 mm. Spines .%3 cm., stout; pedicels 4-6 mm.

227a. 33. chinensis var. chinensis 2276. B. chinemis var. paphhgonica

(227 a) Berberis chinensis var. chinensis Ptdicels 7 mm. above, to 15 mm. below. S.W. Russia: Caucasus, specimen labelled B. iberica (K). Cultivated : L.A. 500 derived from a plant at Wisley, where it was examined in flower

and fruit in 1939. There has been much confusion about the name of this species, as the earliest

name, sinensis, was not validly published. In Bull. Herb. Boissier (2), 5 , Schneider refers (i) to B. poiretii as 71 B. sinensis on page 655, and, (ii) to B. chinensis on page 656 as 7 2 B. iberica. This confusion he remedied in 1906, in the process of which he provided the :name B poiretii. But, as a result, the next variety must be renamed.

The name chinensis is misleading as the species does not occur in Eastern Asia..

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174 L. W. A. AEBENDT

(2273) Berberis chinensis var. paphlagonica (Schneid.) Ahrendt, comb. nov.; syn. B. iberieor var. papldagmictz Schneid., BUU. Herb. Boiaeier, 5 , 656, 1905; B. si& var. ppiczagoniCrr Schneid., I.c., 8, 259, 1908; a vax. dbinensi spinis longis validiaque, pedice& multo brevioribus distinguenda

Paphlagonia: Tea&, fl. 13 May 1892, Sintenis 3646 (Type), 4717 (both Fielding Herbarium, Oxford). Schneider citea these c3pecimene aa from Wilajet Kastambuli. Not cultivated.

(228) Berberis poiretii Schneid., Mitt. Dtsch. Dendr. p. 180, 1906; Bd. Herb. Boissier, (2), 8; B. Fedtschenko, in Kommv, Fl. U.R.S.S., 7,657, 1937; syn. B. sinensis var. angustifdia Regel, A& Hwt i Petropol., 2,416,1873

Shrub 5-6 ft. 8tem slightly lustrous, verruculose. Spines absent, or 6 9 mm. Leaves narrowly obhceolate, 20-40 x 3-6 (-7) mm.; attenuate, sessile; apex acute to sub- muminate. In,acence subumbellate, (%) 3-6 cm., including peduncle (3 mm.-) 1-2 cm. Pedicel.~ 3-6 mm.; bracts 23 mm. ProphyU.9 lanceohte, 1-5-2 x 0.5 mm. Outer se@ 2 x 1 mm., oblong-ovate; acute. Inner sepak 3-3.25 x 1-75-2 mm., oblong- elliptic. Petale 2.5 x 1.5 mm., obovate. S w 1.75 mm., not produced, truncate. ovules solitary. Berries bright red, oblong, 9 x 4.5 mm.

N. Chihli: Cehol, May 1862, David 1723,5363; Pekin, Brehchneider 16; Mt. Connolly, May 1888, not common, Bullock 223; W. Tomb, roadside, 15 May 1927, Liu 1467; Chiu- lung-shan, in small hill ravine, apparently rather rare, fl. 16 May 1880, Hancock 31 (all K). According to Fedtschenko, on sandy mountain slopes and rivers, in the Far East, Ibsuri, sands along the b u r river, and estuary of the Zungaxi. If this be so, the type habitat of B. piretii, mmely 41" N., 115" E., is extended through Mhnchuria to 53" N., 140" E., in S.E. Russia. However, Fedtschenko record~ his plant aa having only few flowers, and dark red berries. Accordingly, we have here no certain evidence for the existence of B. p i r e t i i outside Chihli. But Schneider cites:

S. Manchuria: Schinking, Ross 325; he includes David's numbers in E. Mongolia. Cultivated: I received, about 1950, seed of this species from the Great Wall, just N. of

Pekin, but, to my very great regret, although a few Eleeds began to germinate, I failed to raise any plant of this attractive species.

( 2 2 8 ~ ) Berberis x durobrivensis Schneid., J. Am. Arb. 4,206, 1923 Stems becoming terete and somewhat black. 8piw yellowish. Leaves 15-26x5-

13 mm.; oblong-obovate; attenuate to petiole 1-6 mm.; margins subentire or indis- tinctly spinulose; apex obtuse; the leaves on the new shoots being broader and larger, having a contracted base and more spinules. Inflorescence subumbellate, dense, pendent. Pedicela (3-) 4-6 (-8) mm. ; bracts 14-24 mm. ProphyuS 1:5 mm. Outer sepds 3 x 2 mm., ovate. Inner sepals 6 x 3 mm., oblong-ovate. Petds oblong-obovate, 5-5 mm. Btamns truncate or subemargimte. ovules 1 (-2), sessile. Be& obovoid to ellipsoid, slightly pruinose, 7-8 x 4-5 mm.

Cultivated: in U.S.A., B. poiretii x B. cana&ensis. Not in cultivation in England.

Sect. Integerrimae Schneid., BUU. Herb. Boissier, (2), 5,458, 1905 Skm-s glabrous; mature dark, red, purple or purple-brown; generally lustrous; terete

(except only in B. de&jZoru). Internoah (1-) 1 - 6 3 (-5) cm. Spines mostly con- spicuously discolorous, orange or yellow, generally solitary and conspicuously stout, 1-5 cm. long. k v e s deciduous, 1-10 cm. long; mostly entire, sometimes spino~e; mostly closely, sometimes openly, but always distinctly, reticulate. Inflot.escence (lo-) 15- 4O-fld., racemose, rarely compound below, (2-) 3-10 cm. long. Pedicels 1-10 mm. long. Flowers 3-7 (-10) mm. diameter. Petals aa far aa recorded, emarginate, and mostly

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Berberis and Mahonia 176

shorkr than inner sepals. Ovules 1-2 (4). Berries often dark red, sometimes bright or pale red. Style very short or absent.

Kansu, Shensi, Yunnan, Mongolia, Persia, Kurdistan, Armenia, S i a n g , Turkestan, Szechuan .

KEY 44, to Sect. Integerrimae (Maps 2 and 30) Leaves narrow, 1 : 4-6.

Pedicels 1-3 mm. ; bracts longer than, or equal, to pedicels; fruit estylose. Leaves entire. 229. B. vernae Leaves spinose. 230. B. p u r d m i i

231. B. caroli Pdcels longer ; bracts shorter than pedicels; fruit stylose.

Loaves below concolorous green, and epruinose. Leaves broader, 1 : 1-2-5 (-3).

Inflorescence simply racemose; fruit 6-9 x 4-5 (-6) mm. Stems distinctly sulcate. 232. B. densiflora Stems terete or subterete.

Leaves always spinose-serrate. 233. B. turcomnnica Leaves entire.

234. B. integerrim

235. B. iliensis 236. B. nummularia

237. B. francisci-ferdinandii

Stems sulcate; fruit etylose. 232e. B. &&$ora var. bungeana Stems terete; fruit estylose.

Fruit 6 mm. long. 236 b. B. nummularia var. pyrocarpa Fruit 9-10 mm. long. 238. B. jamsiana

(229) Berberis vernae Schneid., Sargent, PI. Wile. 1, 372, 1913; Stapf, Bot. Mag. 151, t. 9089, 1926; syn. B. curoli var. hoanghnsis Schneid., Bull. Herb. Boissier, (2), 5 , 459, 1905. (Fig. 43)

S.hrub 3 4 f t . Stems very slender, arching, glabrous, sulcate; shoots green; mature dark red, verruculose. Spines stout, solitary, 1 . 5 4 cm., yellow. Leaves 15-35 x 4 4 mm. ; oblanceolate; attenuate to petiole 2-6 mm. ; entire; subacute; above bright green; below subconcolorous green, epruinose, epapillose ; openly reticulate. Racemes 15-35-fld., 2 4 cm. long, including peduncle 5-10 mm. P e d k l s 1-3 mm. ; bracts subequal. Flozoers 3-3,,5 mm. diameter, H.C.C. 2. Prophylls red, lanceolate, 1 x 0.3 mm. Outer sepals 1.5 :K 1 mm. Inner sepals 2.5 x 1.5 mm., all obovate. Petals 2-25 x 1.5 mm., obovate- elliptic; apex subacute, entire; base truncate. Stamens 1.5 mm., not produced, truncate. Ovules 1-2, subsessile. Berries soft, pale red, globose, 3-5-4.5 mm., estylose.

Kansu: Minchow, 10,400-11,700ft., fl. Purdom, s.n. (Type, K), 1880, Tangut, Przelwalski, s.n. (type of B. carolihoanghensis); 1913, seed only Farrer 468, 486.

Cultivated: L.A. 206, fl. 5 May 1943; fr. 30 Nov. 1944; L.A. 92, fl. 25 May 1942. Also, at Kew: (i) from Farrer 468, fl. 12 Apr. 1939; (ii) labelled Rock 13830, fl. 12 Apr. 1939. In early leaf and flower this is one of the most beautiful species for the garden, but it is rardy cultivated. At Wisley, in flower in May 1938, was a form or hybrid with broader leaves (to 3 x 1-1.1 cm.), pedicels 3 mm., and bracts 1 mm.

Fruit dark red, pruinose, oblong; 8 x 4 mm.; ovules L4 . Fruit bright red, epruinose, ovoid to subglobose, 6 x 4 mm. ; ovules 1-2.

Leaves oblong ; pedicels 5-7 mm. ; bracts 1 mm. Leaves orbicular ; pedicels 3-5 mm. ; bracts 2 mm.

Inflorescence compound below; fruit 10-12 x 7-9 mm. Loaves grey, pruinose below.

(230) Berberis purdomii Schneid., Sargent, PI. Wile. 1, 372, 1913 Stem subsulcate to subterete. Spines yellow, 1-3-fld., 1-2 cm. h u e s 10-40 x

4 4 mm. ; narrowly obovate to oblanceolate; base attenuate, subsewile; margins 5-15- spinose (1 mm.)-serrulate; openly reticulate; concolorous green below. Racemes 15- 25-fld., immature, 2 - 5 4 cm., includmg peduncle 1-5-3 cm. Pedicek ( 1 immature), 1 mm. ; bracts 2 mm. Ovules 2. F h r s and fruit unknown.

Shensi: Purdom 3 (Type, K); Yenan fu, Purdom 45 (K); shrub 3 ft. Not cultivated.

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176 L. W. A. AHRENDT

(231) Berberis caroli Schneid., Bull. Herb. Boissier, (2), 5, 459, 1905; syn. B. integer- r i m var. skmphyllu Maxim., Fl. Tangut. 29, 1889

Stems terete. Leaves 30-40 x 6-8 mm., oblanceolate, openly reticulate; papillose

S. Mongolia: Ordos, 1884, Potanin, s.n., Type. Not cultivated. below. Racema 10-15-fld. Pedicels 5-6 mm. Berries 7-9 x 5-6 mm. Style short.

(232) Berberis densiflora Boiss. et Buhse, Nouv. Mdm. SOC. Nut. Nosmu, 12,9,1860; B. Fedtschenko, in Komarov, Fl. U.R.S.S., 7,559,1937; syn. B. integerrim var.

&?&@a Schneid., BUW. Herb. Boissier, (2), 5,461, 1905; syn. B. turunnannicu var. &?mi$ora Rehder, Manual of Cult. Trees and Shrubs, 2,236, 1940; syn. B. num- mulurk var. szmitzkm Schneid., 1.c. ; syn. B. pycnuphylla Bienert MS., in Herb., ex Schneid., 1.c.

Stems not vemculose. Spines solitary, stout, orange, generally 16-3 cm. h u e s fairly thick, 2-35 (-4-8) x 06-1.5 (-2-2 cm.), excluding petiole to 1 cm. ; elliptic to oblong-obovate, cuneate; apex rounded, fairly closely reticulate both sides. Racemes narrow, 1560-fld. Pedicels slender, 3-6 (-7, in fruit) mm.; bracts 1 mm. Outer sep& 2-26 x 1 mm., ovate, acute. Inner sepals 3-5 x 2-2-5 mm., obovate. Petah 2.5-3 x 1.5- 1-75 mm., obovate, entire; base slightly clawed with elliptical, approximate, submarginal glands, 0.3x0.15mm. Stanzens 2-2-25mm., not praduced, truncate. O& 1-2. Berries elliptic-obovoid, dark red, pruinose blue, 6-7 (-8.5) x 44.5 mm. Style short.

According to Fedtschenko, on the mountain slopes of U.R.S.S., and S.E. Trans- Caucasus, with general distribution, Turkish Armenia and Kurdistan. This may include several of the following varieties, of which only the last is in cultivation.

KEY 4 4 ~ Leaves entire.

Racemes 2-36 cm. Spinea 1 - 5 3 cm. Spines 3-5 cm.

Racemea 4-7 cm.

Leaves below green, epruinose. Leave8 spindose.

Leavea below grey, pruinose.

232a. B. &&$ma vm. den&@a 232 6. B. den&i@na vm. mawatantha 232c. B. &m@ra vm. macrobotlys

2323. B. &&$ma vm. aerratqolia 232e. B. M M a vm. bungeana

(232 a) Berberis densiflora var. densifiora Persia: Prov. Teheran, Elbun Mts., 5000-6000 ft., fl. May 1933, Trott 77 (bushes on

hillsides). Prov. Azerbaijan, hills south of Tabriz, Zindjanab torrent, immature fruit, 27 June

1929, GiUiatt-Smith 2573; Sahend hills, valley below Zindjanab, 5500ft., Cowan t Darlington, 1870; Ushnu, Mambrahim, 6500 ft., fl. 31 May 1929, Cowan & Darlingtan 1384. Karadagh district, fr., July 1847, according to Boiss. et Buhse.

Prov. Khorasan, above 3000 ft., fl. 18 June 1885, Aitchison 650, part; near Meshed, fl. Apr. 1936, Daly M12.

Prov. Kerman; Mts. between Chabbis and Kerman, Apr. 1859, Bunge, 8.n. Prov. M a n : Sihna, Kurdistan, fr., 1861, 6000 ft., Haussknecht, s.n. (all K).

(232b) Berberis densillora var. macracantha Boiss., Fl. Or. 1, 103, 1867 Persia: Prov. Farsistan, Shiraz, fr., 1926, Chick 12 (K). 'Called Zirishk, a thorny

bush, growing 7-8 ft., at 6500 ft., not under; found near streams in the hilly countrj round Shiraz, notably the Kuhgilu.'

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Berberis and Mahonia 177

(232 c) Berberis densiflora var. macrobotrys Ahrendt, var.nov., inflorescentiis elongatis

Persia: Prov. Yezd, 6800 ft., fl. 6 Apr. 1892, Bornmuller 2017 (0). Prov. Khorasan, fl., immature, fr. 16 June 1885, Aitchison 650, part (K). ‘Native

name Zihr-bar; common a t an altitude above 3000 ft., near water in Khorasan and on the Paropamisus range. The edible fruit is collected for sale in the markets.’

(232d) Berberis densiflora var. serratifolia Boiss., 1.c.; syn. B. integerrim var. eriwanensis Schneid., BuU. Herb. Boissier, (Z), 5 , 461, 1905

Armenia: Eriwan, Szovits, s.n. N.W. Persia: Prov. Azerbaijan, cultivated, N.E. of Tabriz, a tree, June 1928, Gilliatt-

S.E. Persia: Prov. Kerman, Rahbur, between Kerman and Bender-Abbas, 9700 ft., Smith 2393.

2 Aiig. 1892, Bornmuller 2018 (all K).

(232 e) Berberis densiflora var. bungeana Ahrendt, var.nov., foliis serrulatis,

Leaves sessile; margins 5-10-spinose (G. 1 mm.)-sermlate, distant 3-6 mm.; below grey, pruinose. Racenzes 10-15-fld., 2.5-3 cm. Pedkls 24-3 mm.; bracts 2-24 mm. Ovules 1-2. Berries dark red, epruinose, oblong-obovoid, 6.5-7 x 3.5 mm. Style short.

subtus cinereis, pruinosis

Cultivated: Kew no. 14, fr. 18 Oct. 1939 (K) (Type, BM). In the case of the last two varieties, compare B. turcomannica.

(233) Berberis turcomannica Karelin ex Ledeb., Fl. Ross. 1,79,1842; B. Fedtschenko in Komarov, Fl. U.R.S.S. 7 , 559, 1937; syn. B . integerrim var. turwmnnica Schneid., Bull. Herb. Boisaier, (Z), 5, 431, 1905; syn. B . turcomannica Rehder, Manual of Cultivated Trees and Shrubs, 236, 1940

Spines mostly solitary, 1-3 cm. Leaves to 2-3 by 1.3-2 cm., excluding petiole to 5 mm. ; margins fairly closely spinose-sermlate ; epapillose ; below almost concolorous green; fairly closely reticulate. Pedicles 4-7 (-10) mm. Berries 8-9 x 4-5 mm., dark red, scarcely, if a t all, pruinose. Style short.

I(EY U B

Racemes 12-24-fld., 2 . 5 4 cm. Racemes 6-12-fld., 1-2 cm.

233a. B. turcomannica var. turcomnnica 233 b. B. turcomannk var. buhseana

(233 a) Berberis turcomannica var. turcomannica Armenia: Karelin s.n., Type. According to Fedtschenko, endemic to the highlands of

Turkmenisten. Not cultivated. This species differs in its terete stems from B. &nsi$ora varieties serratifolia and buqeanu ; and also in its spinose leaves from other related species.

(233 b) Berberis turcomannica var. buhseana (Schneid.) Ahrendt, comb.nov., syn. B. integerrim var. buhseuna Schneid., Bull. Herb. Boissier, (2), 5, 462, 1905

Leaves to 3 x 1.4 cm. ; petiole to 5 mm. ; margins 8-lkspinose (0.5-1 mm.)-subserru- late. Pedicels P 5 (-7) mm. Berries ellipsoid or obovoid, 8-9 x 5-6 mm. Style short, distinct.

N.E. Persia: Tscheschme, 1847, fr., Buhse, s.n. (Type, K). Not cultivated. The record of the treatment of B . densijlora and B. turcomannica, and some of the species geo- graphically and botanically related, forms a particularly good example of the danger of complication and confusion when synthetic taxonomy is introduced prematurely.

12 JOURN. LINN. SOC BOTANY-VOL. LVII

Page 178: Berberis and Mahonia : A taxonomic revision

178 L. W. A. AERENDT

(234) Berberis integerrima Bunge, Linnam, 18, 149, 1844; Arb. Naturf. Ver. Riga,

Shrub to 12 ft. Stems lustrous. Spines solitary. Leuve 2 4 x 1-2 cm., including short petiole; oblong-obovate; cuneate; entire; obtuse to subacute; below papillose; oc- casionally spindose on leaves of shoots. &cewm 10-20-fld., 3-6 cm. Pedkd-s 7-10 cm. Ovules 3-4. Berries Ark red, slightly pruinose blue, oblong-obovoid, 8 x 4 mm. Style very short, scarcely distinct.

S. Turkeatan: Karatau, near Samarkand, in woods, Bunge 44, Type. According to Fedtschenko, on stony slopes, Dzungar Alahu , Thian shan, Pamir Ali. Generally in Sinkiang. Not in cultivation.

1, 129, 1847; B. Fedtschenko in Komarov, F1. U.R.S.S. 7,558,1937

(235) Berberis iliensis Popov, I& Seminum Horti Botank Almaatensis Academiw Sckntkrurn, 1936, no. 3, pp. 3-4 (Alma-Ata, I(clzakstan) ; syn. B. nurnrn&ria var. schrenkkm Schneid., Bull. Herb. B&&r, (2), 5, 460, 1905

Spines solitary (0%) 1.5-2.5 (-3) cm., terete or plane. h u e 20 x 8 mm., to 4-4.5 x 1.5 cm., including petiole (3-) 5-15 mm.; baae contracted to petiole; margins completely entire; apex rounded; closely reticulate. Rueem (lo-) 15-25-fld., 3-5 cm. long, in- cluding peduncle scarcely 5 nun. Pediceh 5-8 mm.; bracts 1 (-1.5) mm. O v u h 2, dipitate, with stipea aa long as the ovule. Be+ bright red, 5.5-6 x 3 - 5 4 m., ovoid- ellipsoid, epruinose.

N.W. Sinkiang: Zungaria, east side of the Tian-shan mountains, by the River Ili, 1600-2100 ft. Type; same locality Meyer, s.n. (0), which is the type of B. numrnuhria schrenkkna; same locality Schrenk, s.n.; Pamir and Tian-shan journey, 1906, Appleton 672 (K).

Cultivated: L.A. 449. This plant was reputed to have been raked from seed from Alma-Ah; it has not yet flowered, and the leaves are very spinose, so its correct identity is doubtful.

Popov records that B. ilk& lies eaat of the Tian-shan, in contrast to B. nummalark which lies west of the Tian-shan at the higher altitudes of 29004700 ft.

(236) Berberis nummularia Bunge, Linnaea, 18,149,1844; B. Fedtschenko in Koma-

Shrub to 12 ft. Stems lustrous. Spines solitary, stout, (1-) 2-4 cm. Leaves 2-5 x 2 4 cm., excluding petiole; orbicular or obovate-orbicular ; entire; fairly closely reticu- late; below, epapillose. Racemes 15-30-fld., P10 cm. long, including short peduncle. Peclkels 3-5 mm.; bracts 2 mm. Petals emmginate. Ovules 2, stipihte. Berries bright red, ovoid-subglobose, 5-6 mm.

rov, F1. U.R.S.S. 7, 559, 1937

KEY 44c Spines 2-3 cm. ; leaves euborbicular, green below. Spines 1-2 cm.; leaves obovate, grey below.

236 a. B. nu-&& var. numm&k 236 6. B. numm&ria var. pyrouwpa

(236 a) Berberis nummularia var. nummularia S. Turkesh: Ka,ra.hu, near Samarkand, woods, Bunge, s.n., Type. According to

Fedtschenko on the stony mountain slopes of the Tian-shan and Pamir Ali. Not in cultivation.

Page 179: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 179

(236 b ) Berberis nummularia var. pyrocarpa Schneid., Bull. Herb. Boissier, (2), 5,460, 1905; syn. B . integerrim var. pyrocarpa Regel, Actu Horti Petropol., 5, no. 1, 228, 1877

Spines solitary 1-2 cm., subterete. Leaves to 3.5 x 2 cm., excluding petiole to 1.5 cm. ; obovate, cuneate, entire, rounded, reticulate ; below grey, pruinose. Racemes 20-40-fld., 4-8 cm., including peduncle to 1-5 cm. Pedicels 3-5 (-6) mm.; bracts 1-2 mm. Berries obovoid, 6 x 4 mm.

Turkestan: Tschatkal, 6000-9000 ft., 1876, Regel, s.n. (Type, K). Not cultivated.

(237) Berberis francisci-ferdinandi Schneid., Sargent, PI. Wils. 1, 367, 1913 ; 1.c.

Shrub 8-12 ft. Stems with young shoots red; mature red-brown, subterete. Spines 1-3-fld., sulcate, 2-4 cm. Leaves 2-7 x 1-3.25 cm., excluding petiole 5-15 mm. ; ovate to elliptic ; base contracted to decurrent petiole; margins 15-30-spinose (0.5 mm.)-serrate (0.5 mm.), distant 1-2 mm.; acute or subacute; above sublustrous; below slightly paler green. Racemes 20-30 (40)-fld., compound towards the base, 5-12 cm. long, including peduncle 1-3 cm. Pedicels slender, 4-8 (-10) mm. ; bracts 1.5-2 (-3) mm. Flowers H.C.C. 2. Prophylls red, acute, 1.5-2 x 0.5-0.7 mm. Outer sepals 2-4 x 1.5 mm., ovate, acute. Median sepals similar, 3 x 2 mm. Inner sepals 3.25-4.25 x 2-2.5 mm., obovate. Petals 3 6 4 . 5 x 2-5-3 mm. ; apex acute, scarcely or very shortly incised; base slightly clawed, with discolorous, separate, oblong glands, 0.8 x 0.4 mm. Stamens 2-5-3.5 mm., produced, obtusely subapiculate. Ovules 2, sessile. Berriss scarlet, epruinose, ovoid-ellipsoid, 10-12 x 7-9 mm. Style short.

W. Szechuan: Mao-chou, thickets, 4200-7500 ft., 1908, Wilson 1180 (Type, K); 8000 ft., May 1908, Wilson 2869; 11,000-13,000 ft., May 1904, Wilson (Veitch) 3151.

Cultivated: L.A. 49, fl. 18 May 1938,4 June 1942,24 May 1944. Also, from the type, Wilson 1180, fr. 20 Sept. 1947, Maidwell Hall, Northampton.

(238) Berberis jamesiana Forrest et W. W. Sm., Not. Bot. Gard. Edinb., 9, 81, 1916; Stapf, Bot. Mag. 156, t. 9298, 1933; syn. B. integerrim Ranch., Bull. SOC. Bot. Fr. 33, 386, 1886 and PI. Dehvay. 36, 1889, non Bunge; syn. B. nummuluria var. sinica Schneid., Bull. Herb. Boissier, (2), 8, 202, 1908

Shrub 8-12 ft. Stems very stout; young shoots purple; mature lustrous red, not verru- culose. Spines 3-fld., stout, sulcate, 3-5 cm. Leaves thick, subcoriaceous; apex rounded; reticulate; above dull light green; below grey, pruinose, epapillose. Racemes 20-40-fld., sometimes compound below, 7-10 cm. long. Pedicels 6-10 mm., slender, glabrous; bracts 1-1.5 mm. Flowers H.C.C. 211, diameter 5-7 mm. Ovules 2. Berries translucent, a t first creamy white, finally light red.

3, 437, 1917; Stapf, Bot. Mag. 155, t. 9281, 1932

KEY 4411 Leaves entire, ovate, or elliptic; fruit globose.

Leaves broad, 1 : 1.5-2.2; fruit 9-10 mm. Leaves narrow, 1 : 3; fruit 6 7 mm.

238a. B. jamsiana var. jamesiana 238b. B. jamesiana var. saepium

238c. B. jam&anu var. leucocarpa Leaves closely spinulose, suborbicular; fruit ovoid.

(238a) Berberis jamesiana var. jamesiana. (Fig. 42) Leaves to 86-10 x 3.3-5 cm., including petiole l-fk2-5 cm., ovate: obtuse, margins

entire or subentire. Prophylls 2-2.5 x 1.5 mm., ovate, acute. Outer sepals 3 x 2 mm., oblong-obovate. Inner sepals 4-5 x 2-4 mm., narrowly obovate. Petals 4.5 x 2 mm., naxrowly oblong-elliptic to obovate ; apex shortly acutely emarginate with acute lobes ; clitwed, with conspicuous discolorous subapproximate marginal glands, 0-6 x 0.5 mm . ,

12-2

Page 180: Berberis and Mahonia : A taxonomic revision

180 L. W. A. AHRENDT situated within the long claw. Stamem 2.75mm., produced, subapiculate. Berries globose, 9 x 9-10 mm.

Yunnan: open situations amongst rocks on the mountaim N.N.E. of the Yangtze bend, 11,000 ft., fr. Oct. 1913, Forrest 11474 ( T y v , E, K). Also from Weishi, west of Tungchuling, 95OOft., side of forest, 19 Nov. 1937, Yu 10750; Atuntze, Hungpoh, 12,000 ft., 23 Nov. 1937, Yu 16653; Atuntze, Mt. Myetzimu, among forest, 11,400 ft., 10 Oct. 1937, Yu 10576 (all, E); Rock 22836; Forreat 25436.

Cultivated: L.A. 97, fl. 16 May 1944; fr. 21 Dec. 1944; also from many of the collectors’ numbers cited. A large ferocious plant, magnificent in berry, and autumn colour of leaf.

(238b) Berberis jamesiana var. saepium Ahrendt, var.nov., foliis minoribus angustioribusque, fructibus minoribus

Leave-9 3-6 x 1-1-7 cm., elliptic. Berries globose, 6-7 mm. W. Szechuan: forming 6-12 ft. hedges, 13,000-14,000 ft., Sept. 1903, Wilson (Veitch)

Cultivated : plant from Wilson’s seed, fr. 20 Sept. 1947, at Maidwell Hall, Northampton.

(238c) Berberis jamesiana var. leucocarpa (W. W. Sm.) Ahrendt, st&t.nov.; syn. B. le-rp W. W. Sm., I.c., 9, 82, 1916; a var. j a d m foliis orbiculatis crebre spinulosis, fructibus obovoideis distinguenda

Spines 1-24 cm. Law to p5 x 3-4 cm., excluding petiole, 2-3 cm.; suborbicular; margins 20-40-spinose (1 mm.)-sedate, distant 1-14 mm. Pmphylls 1 x 0.4 mm., acute, red. Outer se& 1-75 x 1 mm., oblong-elliptic. Inner se& 3.75 x 2.75 mm., obovate. Petals 3.5 x 1.75 mm., oblong-elliptic; apex entire; glands elliptic, contiguous, marginal, 0.5 x 0-3 mm., situated above the claw. 8tamem 2 mm., not produced, truncate. Bem.e-9 obovoid, 10 x 6-7 mm., a t first partly whitish, partly pale red, ultimately all red.

N.W. Yunnan: open scrub and amongst rocks, on the Mekong-Yangtze divide, 11,OOO ft., 1914, Forrest 12855 (Types, E, K). ‘Shrub 6-8 ft., h i t white.’ Chien-chuan- Mekong divide, dry stony hillsides, Forrest 23743; Atuntze, 8450 ft., 24 Sept. 1937, Yu 10423 (E), seed only Forrest 29085.

Cultivated: from Forrest 29085, fl. 12 Apr. 1939,21 May 1942; fr. 28 Oct. 1943. Also, at Kew, from the type, Forrest 12855, fl. 16 May 1944; fr. 12 Sept. 1940.

It will be noticed that, geographically, the section Integerrjmrte, falls into three portions: (i) a westerly one from Armenia and Persia, whose population may, with further data, be finally collected in one specilk unit, B. &&$ha. (ii) A central one, from Turkestan and Sinkiang, which also, later, may be justifiably regarded as a wide- spread speci6c unit, B. nummukcrria. (iii) An easterly set from N.W. China and adjacent southern Mongolia, which, in a smaller area, includes a greater diversity of species.

3157 (Type, K).

KEY 45, to Sects. Dasystachyae and Vulgares combined Stems dark, red or purple, when mature. A

Cf. 317. B. x notabilia Stems pale, yellow when mature.

Fruit 15 x 10 mm.; ovules 3 4 . Fruit 6-10 (-12) x 5-7 mm.; ovules 1-2. Leaf blade 12-15 cm., orbicular, excluding petiole 5-16 om.; f i t blue-black.

Leaf blade 3-10 cm., excluding petiole 1-4 cm.; f d t red. 241. B.d&dWbOtryS

B Leaves entire, or scarcely, inconspicuously, remotely apinose. Leaves closely spindose.

Leaves narrow, 1 : 3-4; acute, below grey, pruinose. Pet& longer than inner sepals.

252 b. B. x .?uxi$m vm. oblunwolata

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Berberis and Mahonia 181

Leaves broad, 1 : 1.5-2.5, obtuse to rounded; below green, epruinose. Stems very sulcate; fruit 5-7 mm. long pedicels 4-7 mm.

Petiole 2-6 mm. Cf. 214. B. quelpxrtensia Petiole 1-2 cm. Cf. 246. B. anhweinenais

251. B. vulgaris c

Stems angled; fruit 9-12 mm.; pedicels G 1 2 mm. Petals shorter than inner sepals.

A. Petals as long aa, or longer than, inner sepals. Leaves narrow, 1 : 3-4, entire; green below. Leaves broad, 1-2 (-2.5); grey, pruinose below.

240. B. dielaiana

Spines foliaceous; fruit 7-8 mm., lustrous, globose. 250. B. koreana Spines not foliaceous; fruit 8-12 x 5-7 mm., oblong, dull.

, Leaves broad, 1 : 1-1.5; margins 15-30-spinose. 239. B. kansuensia 252c. B. x laxiflora var. langeana

Stems sulcate.

Leaves narrower, 1 : 2.5, entire.

Leaves 40-60 x 18-30 mm.; racemes shorter than leaves; spines solitary.

Leaves 20-40 x 10-13 mm.; racemes longer than leaves; spines 5-7-fld.

Stems terete.

253. B. bretschneideri

252e. B. x ceraaina Petals shorter than inner sepals; spines mostly absent, or weak and 4-10 mm. long.

Stems terete or subterete. Petals 3-5 mm. broad, emarginate.

Leaves below, green, epruinoae; reticulate; 4-10 om. long. 254. B. pachyamntha Leaves below grey, pruinose; scarcely reticulate; 1-4 cm. long. 255. B. henryana

Petals 1.75 mm. broad, entire. 256. B. farreri

Leaves 3-6 cm. long, closely spinulose, reticulate; pedicels 7-10 mm.

Leaves 1-3 cm. long, entire, not reticulate; pedicels 5-7 mm.

Stems sulcate.

252 f . B. x dksuthiensh 257. B. agricola

B. Stems terate; flowers diameter 6-7 mm. Leaves 35-60 x 22-40 mm. ; fruit estylose. Leaves 10-25 x 3-8 mm. ; fruit stylose.

Leaves green, epruinose below. Stems sulcate.

Leaves paler below, acute; petiole 1@-15 mm. Leaves concolorous below, obtuse; petiole 1-4 mm.

Leaves grey, epruinose below.

C. Leaves green, epruinose below. Stems terete; petiole 1-4 em.; flower diameter 6 6 mm.

Petals entire; fruit 6 x 5 mm. Petals amarginate; fruit 10 x 5 111111.

Petiole 1-3 om. Stems sulcate, or angled.

Pedicels 4-7 mm.; fruit 5-6 mm. Pedicels 8-13 mm. ; fruit 9-10 mm.

Petals entire; pedicels 8-13 mm. Petals emarginate; pedicels P 9 mm.

Petiole 1-10 mm.; fruit 8-11 mm.

Spines 10-25 mm. ; leaves openly reticulate. Spines S 6 mm. ; leaves closely reticulate.

Leaves grey, pruinose below. Indeterminate, but belonging somewhere here.

242. B. feddeana 249. B. conwlor

245. B. chekiangenaia 248. B. orientalis

252. B. x h i$ora

243. B. daaystachya 244. B. zabeliana

Cf. 246. B. anhweiensis 247. B. homnensis

258. B. atroprasina

259. B. amuremia 260. B. hersii

261. B. regeliccna 262a. B. x meehanii

262b. B . x b e k

Page 182: Berberis and Mahonia : A taxonomic revision

182 L. W. A. AEIRENDT

Sect. Dasystachyae Scheid., &. Bot. 2.67,287,1918; syn. Sect. V~L~AFCES, subsect. Euvulgares Schneid., BUU. Herb. Bokier, (2), 5, 064, 1905

Stem glabrous; when mature mostly pale yellow, occasionally dark red. Ink?& (1-) 2-4 (-7) cm. 8p.m mostly weak, 2-10 mm.; sometimes 1-3-fld., and 1-2-5 cm. Leaves deciduous, petiolate; lamina (2 .6) 4-7 (-12) cm. long; petiole (Om&) 1-3 (-12) cm. ; margins mostly 15-30-spinose. Inflorescence ( 1 6 ) 15-30 (-lOO)-fld., fairIy narrowly racemose (2 .6) 3-5-7 (-28) cm. long. Peddcels &t3 (-12) mm.; bracts 1.2 (-2.5) mm. Flower8 ( 1 . 6 ) 4-7 mm. diameter. Petals mostly shorter than inner sepals. O w h 1-2. Be& wtylose, 6-8 (-11) mm. long, red (except, blue-bhk, only in B. doZichobat~8).

Kathnir, Kumaon, KZMSU, Shensi, Honan, Hupeh, C h e b g , Anhwei.

-Y 46, to Sect. DASYSTACHYAE (Maps 2 and 32) Leaves entire, acute to subacute; ultimately green, epruinose below. Stems dark red when mature. 240. B. diehiana Stems pale yellow when mature.

stems very sulmte. stems terete.

245. B. dwkiangensie 242. B.feddeana

Leaves with spinose margins. Stems dark red when mature; petals equal to or longer th&n inner sepals.

Leaves conspicuously spinose, with rounded apex, below pruinose grey; broad,

Leaves subentire or inconspicuously spinose, apex attenuate; below h l l y green, 1 : 1-2-1-6. 239. B. kanmmaia

epruinose; m o w e r , 1 : 3-46 Cf. 240. B. dielsiana Stem pale yellow when mature. Infloreecence 50-100-fld., 10-28 cm. long; leaves with petiole 5-16 cm.

Inflore9cence 16-30-fld., 3-7 om. long; leaves with petiole up to 4 cm.

Leaves with w g i n inconspicuously spinose; apex subacute. Cf. 242. B. f h m h v e a with -gin conspicuously spinose and apex rounded.

241. B. dolkhbOtqi8

stems terete.

Leaf -gins with 25-50 spinules dktant 1-1-5mm.; petals enth; fruit 7X5mm. 243. B.daeystrrciya

fruit IOx6mm. 244. B. zabeliana Leaf margins with 10-35 spinules distant 1.5-3mm.; petals ema&ate;

stems Sulc&te. h v e s green, epruinose below; fruit 10 x 5 mm.

Leave8 with base cuneate; spines 0-8 mm.; pedicels P 7 mm.; inner sepals

Leaves with base contracted; spines 16-25 mm.; pedicels 8-13 mm.; inner 3.25 mm. Cf. 244. B. zabeliana

sepals 7 mm. 247. B. honaneneie 246. B. anhmkd Leaves grey, pruinoae below.

(239) Berberis kansuensis Schneid., Ost. Bot. 2. 67, 288, 1918 stem angled or sulcate. Internoah 1 3 c m . #pines discolorous orange, sdcah,

1-2.4 cm. Leaves with blade orbicular-elliptic, 2.5-3-5 (-5) x 2-2-5 (-3) cm., excluding petiole 1-2 cm.; base contracted; apex rounded; margins 15-30-spinose (1 mm.)-serrate (1 mm.), distant 1-2 mm.; openly reticulate; above dull; below pruinose, grey. R a ~ e m e ~ narrow, 15-25-fld. (2 .6) 3-5-7 cm. Pedi& verticillate, 4 4 mm.; bracts 1-1.5 mm. Prophylk red, mumhate, 1-4 x 0.7 mm. Outer sepal3 2.5 x 1.5 mm., ovate, acute. 1nne.i- sepal3 4-5 x 2.5 mm., oblong-obovate. Petah 4-5 x 2 mm., oblong-elliptic; apex nar- rowed and acutely emaxginate with acute lobea; base shortly clawed with obovate separate g h d s , 0.7 x 0.5 mm. Stamma 3 mm., scarcely produced, rounded-truncate. Ovda 2, with a joint stalk leading to two short sepaxate stipee. Berries oblong-obovoid, 8 (-8.5) x 6 mm.

Page 183: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 183

KEY 4 6 ~ Shrub 3-5 ft. Shrub 12-15 ft.

239a. 3. kamuensis var. kanauenais 239 b. B. kansuensia var. procera

(239 a) Berberis kansuensis var. kansuensis

June 1911, seed only, Purdom 680. S. Kansu: Minchow and Choni district, 1910, fl. Purdom 1014 (Type, K); 8000ft.,

C’ultivated: Wisley, fl., fr. 1939. An attractive small shrub, very rare in cultivation.

(239 b) Berberis kansuensis var. procera Ahrendt, var.nov., multo elatior S t e m subterete to subsulcate; red-brown, ultimately paler. Leaves with blade

5 x 3 cm., excluding petiole 1 cm.; margins 12-20-spinose (0-5-1 mm.) s e d a t e ; reticu- lation openly elevated but conspicuous. Racem~a to 8 cm. Berries oblong, 8.5 x 6 mm.

W. Kansu: Kemang su, Oct. 1925,9800 ft., Rock 13290 (Type, K). ‘A shrub of 15 ft., with long whiplike branches’; summit range of Kemang su, N.E. of Tankar, 10,500 ft., Rock 13291 (K). Not cultivated.

(240) Berberis dielsiana Fedde, Bot. Jb. 36, Beibl. 82, 41, 1905 Shrub 6-9 ft. S t e m with young shoots green; mature dark red, finely angled-sulcate,

sparsely verruculose. Spines solitary, terete, 3-12 mm., though on new shoots to 2.5 cm. Leuoes 5-9 x 1-5-2 cm., including petiole, c. 1 cm. ; base contracted; margins entire, or with few inconspicuous spinules ; apex attenuate, acute to subacute ; venation openly branched, not reticulate, above dull deep green; below a t first grey, pruinose, finally green, epminose, scarcely papillose. Racemes 10-20-fld., 5-7 cm., sometimes compound below, reduced towards apex of stem. Pedicels 3-5 mm.; bracts 2 mm. Floluers H.C.C. 3/1. Prophylls red, lanceolate, acuminate, 2-2-5 x 0-5-0.75 mm. Outer sep& 2-2.5 x 0.75- 1 mm., oblong-ovate, acute. Inner sepals 4-46 x 3-25 mm., obovate. Petals convolute, 5-55 x 3 mm., acutely emarginate; glands discolorous, separate, non-marginal, 1 x 0.4 mm. Stamens 3 mm., not produced, truncate. O d e s 2. Berries red, epruinose, oblong, 8-9 x 4-5 mm.

Shensi: Sciu jan shan, near Kan y huo, fl. 13 May, Giraldi 2298, Type; south of Yenan fu, fl . 1910, Purdom 341 (K).

Honan: Sung hsien, Shih-tze-miac, immature fr., 26 May 1919, Hers 598 (K, a specmen with leaves 10-18-spinose (1 mm.), distant 2-4 mm., and pedicels 5-7 mm.). According to Rehder, also from the Yung-ning, and Lushih districts.

Cultivated: L.A. 441, fl. 29 Apr. 1943; fr. 6 Nov. 1941, 26 Nov. 1942, 14 Oct. 1943. This is a large, but unusual looking Berberis worth cultivation, where there is room, on account of its general character and attractive foliage.

(241) Berberis dolichobotrys Fedde, Bot. Jb. 36, Beibl, 82, 41, 1905; Diels, Jugend- f m n und Bliitenreife im Pjtanzenreich, 105, fig. 30 B, 1906; Schneid., Bull. Herb. Boissier, (2), 8, 1908, Illu-str. Handb. Laubg. 2, 922, 1912

Stem slender, pale grey. Spines 1-2 cm. Leaves thick with an orbicular or flattened orbicular blade 7-12 x 6-15 cm., excluding petiole 5-16 cm.; base contracted; apex rountied ; margins 20-50-spinulose-denticulate ; venation much branched, but not reti- culate; both sides lustrous green. InJlorescence 50-100-fld., in a spikelike raceme, com- pound or subpaniculate below, 10-28 cm. long, 1-2 em. broad. Pedicels 5-7-5 mm. ; bracts scarcely 1 mm. Flowers 1-5-2 mm. diameter. Prophylls lanceolate, acute. Outer sepah much smaller than inner sepals. Berries oblong, 6-7.5 mm., estylose (according to Fedde), or shortly stylose (according to Schneider), berries black, pruinose blue (ac- cording to Fedde).

Page 184: Berberis and Mahonia : A taxonomic revision

184 L. W. A. AEECENDT

Schneider c i h , also, Farges 742 from Tschen-keo-tin in E. Szechuan.

sirable to see in living form, but it is not in cultivation.

Shensi: Tai-pai-shan, fr. Aug., Giraldi 51, Type; Hua-tze-pin, fl. 20 June, Girddi 49.

This species, known only from deacription, sounds quite remarkable, and very de-

(242) Berberis feddeana Schneid., BuU. Herb. Boissier, (2), 5,665,1905; (2), 8,261,1908 Stem with young shoots slightly reddish ; mature terete, yellow. Spines 2 4 mm. Leaves

3 - 5 4 x 2-2-4 cm., excluding petiole 6-15 mm.; obovate to oblong-obovate; subacute to subobtuse; base contracted; margins entire or with many very inconspicuous spinules; both sides openly reticulate. Racenzes 18-24-fld., 3-7 cm., including peduncle, 1-3 cm. Pedicels 4-8 (-11, in h i t ) mm. Flinmrs 6-7 mm. diameter. Berries 8 x 6 mm., oblong.

W. Szechuan: N.E. of Tmhien lu, thickets, ~ l O , O O O f t . , fl. 9 July, Wilson 2862 (Type, K); Washtan, thickets, 52003800 ft., fr. May 1908, Wileon 2863 (K).

W. Hupeh: Patung district, Ichang, fl. May 1888, Henry 4937 (K). Shensi: Huan tou shan, 20 Oct. 1897, Girddi 2296, cited by author. Not in culti-

vation.

(243) Berberis dasystachya Maxim., Bull. A d . Sci. St-Pdtersb. 23, 308, 1877; Schneider, I.c., (Z), 8, 262, 1908

Shrub 3-5 ft. Stem with young shoots sometimes reddish; mature terete, pale yellow, very sparsely finely verruculose. S p i w absent, or concolorous, solitary, 5-12 mm. Leaves thick; blade oblong-elliptic 3-6 x 2-5-4 cm., contracted to petiole (1-) 2-3 (-4) cm. ; apex rounded; margins 25-50-spinose (1-1.25 nun.)-subsedate, distant 1-16 mm. ; both sides reticulate; above dull yellowish green; below epruinose, lustrous yellow-green. Racemes 15-30-fld., 3-5-55 (-7) cm., including peduncle 1-16 cm. Pedicels 4-7 mm. Flowers 6-6 mm. diameter. Outer s e p h 2 x 0 6 mm., lanceolate. Inner sepia 4 x 1-75 mm., obovate with base slightly clawed. Petals 3 x 2 mm., obovate, entire; base clawed with oblongelliptic, separate, submarginal glands, 0.5 x 0.25 mm., situated above the baae of the claw. Stamens 2.5mm., not produced, truncate. OvuIes 1-2. Berries red, epminose, 6-7 x 5-55 mm.

Kansu: immature, fl. 1873, Pnewalski, s.n. (Type, K); 1914, seed only, Farrer 80. W. Hupeh: Mar. 1889, Henry 6816 (K); Hsing shan hsien, woods, 6800 ft., 2-5 ft.,

Wilson 307 (BM). N. Hupeh: Mt. Lao-y san, Teiuz-saon, June 1897, Hugh s.n.; Chochi, Che Fangson,

3 June 1897, Hugh 53; Mt. Thae-pei-shan, Aug. 1899, Hugh, s.n. (all BM). Shensi: Kuiqua shan, 9 July 1897, Giraldi 2316; Tai-pai-shan, 20 Sept. 1897, Giraldi

2306 and Hum-tou-shan, 10 July 1900, Giraldi 7018 are all cited by Schneider. Cultivated: L.A. 389, from seed, Farrer 80, fl. 29 Apr. 1942; fr. 8 Sept. 1943.

(244) Berberis zabeliana Schneid., BuU. Herb. Boissier, (2), 5, 667, 1905; Parker, A Forest F h a for the Punjab, with Hazara and Ddhi, 13, 1918

Shrub 5-6ft. Stems sometimes subterete, mostly finely angled or sulcate. Spines concolorous, 1-3-fld., 3-8 (-12) mm., sometimes absent. Leaves 24-66 (-8) x 1.6- 2-7 ( 3 . 8 ) cm., including petiole, 0.5-2-2 cm.; blade oblong-elliptic; base cuneate; apex obtuse or rounded; margins 10-35-spinose (0.5-1 mm.)-sedate (0.5 mm.), distant 1.5-3 mm.; openly reticulate above, and dull light green; below with inconspicuous venation, a t h t grey, pruinose, Snally epruinose, subconcolorous green. Racemes 12-20-fld., 3-5 cm. Pedicels 4-7 mm.; bracts 16-26 mm. Flowers 6 5 mm. diameter. Prophylls 1-5 x 1 mm., ovate, acute. Outer s e p h 2.75 x 1-5 mm., elliptic-obovate. Inner sepals 3.25 x 2.5 mm., obovate. Pet& 3 x 1.75 mm., obovate; base scarcely clawed, with

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Berberis and Mahonia 185

elliptic glands, 0.7 x 0.4 mm., approximate or subcontiguous, but submarginal. Stamens 2.5 mm., not produced, truncate, or slightly retuse. Ovules (1-) 2. Berries oblong obovoid, 9-11 x 4-5 mm.

Kashmir: Falconer 95 (Type, K). Hazara: according to Parker, Upper Kagan valley, Battakundi, 9OOO ft., gregarious

in dense patches in dry open places. Afghanistan: a specimen from the woods of Shendtoi gorge, 23 June 1880, a t 9OOO ft.,

though without flowers or fruit seems to belong here. Cultivated: L.A. 410 (Kew 29/48&33, reputed from seed, Osmaston 51)-fl. May

1939; fr. 18 Oct. 1939 (K).

(245) Berberis chekiangensis Ahrendt, sp.nov., ramulis valde sulcatis, pallide flavis, foliis integris acutis subtus viridibus, idorescentibus racemosis, ovulis solitariis, distinguenda

Stems with young shoots slightly reddish ; mature pale yellow, very sulcate. Internodes (3-) 4-7 cm. Spinessolitary 1-2.2cm. Leaveswithrhombo-elliptic blade4-7 x 2-3 cm., con- tracted to petiole (0.5-) 1-16 cm.; entire; acute; lateral veins elevated, reticulation fine; below epruinose concolorous or scarcely paler green. Racemes 10-15-fld., 36-66 em. long, including peduncle 2-34 cm. Pedicels 7-12 mm. Flower diameter 5 mm. Outer sepals 2 x 1 mm., ovate, acute. Inner sepals 3-5 x 2.5 mm., obovate. Petals 3.5 x 2.5 mm., obovate. Stamens 1.75 mm., produced, apiculate. Ovules solitary.

Chekiang: Tientai Mts., 6500 ft., fl. 1889, Faber 260 (Type, K). Not in cultivation. The only other species recorded from this eastern province, and from the same mountain range is the very different B. lempergiana of the Sect. Wallichianae.

(246) Berberis anhweiensis Ahrendt, sp.nov., a B. chekiangensi foliis marginibus multis spinulosis, subtus pruinosis cinerascentibus distinguenda

Stems mature yellow, sulcate. Internoh 2 4 cm. Spines solitary, 1-16 cm. Leaves with blade 3-56 x 1.5-3 cm., contracted to petiole 1-16 cm.; apex rounded; margins 15-25-spinose (1 mm.)-serrate (1 mm.), distant 2-3 mm. ; venation elevated and reti- culated both sides; below grey, pruinose. Racenzes 10-20-fld., 3 4 cm. long, including peduncle 1-16 cm. Pedicels glabrous, 4-7 mm.; bracts 1 mm. Ovules 1-2. Berries obovoid, 5-6 x 33.5 mm.

Anhwei: Wang-shan, back of Lion Ridge, in dense woods near ridge, 4300ft., fr. 12 July 1925, Ching 2981 (Type, K). ‘Spreading shrub, 6 ft.’ Compare with B. regeliana. Not in cultivation.

(247) Berberis honanensis Ahrendt, Card. IUust. 64, 426, 1944 Shrub suberect, 5-6 ft. Stems yellow, sulcate. Internodes 1-26 cm. Spines stout,

concolorous, sulcate, 1.5-2-5 cm. Leaves with oblong-elliptic blade, 24 x 1.2-3 cm., con- tracted to petiole (0.3-) 1-25 cm.; apex rounded; margins 20-30-spinose (0.5-1 mm.)- serrate (0.5 mm.), distant 0.5-2 mm.; both sides closely finely reticulate; above dull light green; below epruinose, paler green. Racemes 1&15-fld., 45-56 cm., including peduncle (04)-) 1-1.5 (-2) cm. Pedicels 8-13 mm. ; bracts 1 mm. Flowers 12 mm. diameter, H.C.C. 603 externally. Prophylls concolorous, lanceolate, 2 x 0.75 mm. Outer sepals 3-!i x 2 mm., elliptic-obovate. Median sepals 5-5 x 4.75 mm., broadly obovate. Inner sepals 7 x 5.5 mm., obovate. Petals (H.C.C. 2/1), 6.5 x 3-5 mm., elliptic, obtuse, entire; base clawed, with oblong, separate, non-marginal, discolorous glands, 1 x 0.5 mm., situated above the claw. Stanzens 4mm., not produced, truncate. Ovules 2. Berries oblong, epruinose, 9-10 x 5 mm.

Honan: Lu shan hsien, seed only.

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186 L. W. A. AERENDT Cultivated: L.A. 395, one of the plants raised from this seed (and inaccurately dis-

tributed as B. virgetomrn); fl. 19 May 1943; fr. 24 Sept. 1943, 25 Sept. 1944 (Types, 0, BM).

The leaves of B. h o n a n e n s i S with their contracted base and conspicuous petiole associate it with this section, while the larger flowers are more characteristic of the following section, Vulgares. This species may be regarded a a link between the two sections.

Sect. Vulgares Schneid., Bull. Herb. Boissier, (2), 5, 660, 1905; Sargent, PI. Wils. 3,440, 1917

Stem glabrous, mostly yellow when mature. I&tmnks (1.S) 2-3 (-4) cm. Leaves deciduous ( 1 . 5 ) 3-6 (-10) cm. long, including decurrent petiole 1-10 (-15) mm.; base cuneate; margins mostly closely spinose; always more or less reticulate. Racenzes open, formal (lo-) 15-30-fld., (2-5-) P 7 (-10) cm. Pedk4.s (2-) 5-9 (-12) mm. Flowers 8-12 mm. diameter. StamRns not, or scarcely, produced, truncate or rounded. 04yUles 1-2. Berries mostly oblong, 8-12 mm.

North, Central, and South East Europe, Lazistan, Armenia, N.W. Persia, Kashmir, Punjab Himalaya, Garhwal, Yunnan, W. Hupeh, Shansi, Chihli, Mongolia, Manchuria, Amurland, Korea, Japan.

KEY 47, to Sect. Vulgares (Maps 2 and 33) Spines foliaceous; fruit lustrous, firm, globose. 250. B. koreana Spines not foliaceous; fruit dull, soft, oblong.

Leaves entire, or with few inconspicuous spinules. Stems pale yellow; leaves concolorous green below.

Fruit bright red, estylose; stems sulmte; leaves 3-6 cm. Fruit dark red to purple, stylose; stems subterete; leaves 1-2 cm.

248. B.&ntalia 249. B. concolor

Stems dark red. Leaves 1-26 cm., scarcely reticulate; below grey, pruinose. Leaves 2-4 cm., closely reticulate; below green, epruinose.

~ V W with Closely (16) 20-60-~pinOSn. Stems dark red, or red-brown, subterete.

Leaves below grey, pruinose. Petals aa long as inner sepals; style nil. Petals shorter than inner sepals; style short.

Leaves 1-4 cm. ; racemes 3-4 cm. ; petals entire. Leaves 5-10 om.; racemes 5-10 cm.; petals emarginate.

Leaves below green, epruinose.

Stems pale yellow, sulmte. Petals longer than inner sepals.

Leaves green, epruinose below. Leaves grey, pruinose below.

Petals shorter than inner sepals. Leaves green, epruinose below.

Leaves above lustrous; pedicels 8-13 mm.; petals entire. Leaves dull; pedicels 5-8 mm.; petals emarginate.

Spines 10-20 mm. ; leaves openly or slightly reticulate. Spines 3-6 mm. ; leaves closely reticulate.

Leaves grey, pruinose below.

257. B. agricoka Cf. 256. B. farreri

253. B. bretschmideri 255. B.hryana

256. B. farreri 254. B. @yamnth

251. B. vulgarie 252. B.xlaxi$ora

258. B.drgpmsana

259. B. amur& 260. B. hraii

261. B.regeliana

(248) Berberb orientalis Schneid., BuU. Herb. Boissier, (2), 5,666,1905; B. Fedtschenko, in Komamv, Fl. U.R.S.S., 7,557, 1937

Shrub 6-7 ft. Stem sulcate, bnght yellow. Spines 1 (-3)-fld., 4 4 mm., rarely to 2 cm. Leaves 3-6 x 13-24 (-3) cm.; ovate-elliptic; decurrent petiole 2-4 mm.; margins entire

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or iIiconspicuously spinulose-subsermlate, with the minute spinules distant 3 4 mm., by the leaf apex, but 0-75 mm. long and 1-2 mm. distant towards the base; both sides dull, concolorous, closely reticulate. Bacem~s ( 1 6 ) 20-25 (-30)-fld., 4-7 cm. long, with peduncle 1-2 (-5) mm. Pedicels (5, by apex-) 7-8 (-10, below) mm. Berries bright red, sblong, 10-11 x 5-6 mm. Seeds purple.

N.W. Persia: Mt. Ghilan, fr. Aucher-Eloy 4039 (Type, K). Tid ish Armenia : Koosoodagh, Sintensis 3183 ; Gumuschkane between Sorda and

Russian Armenia : according to Schneider. L:tzistan: according to Schneider. According to Fedtschenko, the following varieties may be distinguished.

‘Toke, Sintenis 3437 (both, K).

KEY 47A Racomes P 7 em.

Leaves EM above described. 248. B. o r i e n t d i s var. oriental is Leaves narrower. 248a. B. o r i e n t d i a var. litoralis Rupr. Leaves coriaceous, acute. 2483. B. orientalis var. emarginata (Boiss.) Grossh.

Racemes 2-3 em. 248c. B. 0rientdi.q var. brachybotrys Boiss.

This species is not in cultivation. It hm some relation botanically and geographically m t h the Sect. Integerrimae, but is a t once distinguished by its bright yellow sulcate stems.

(249) Berberis concolor W. W. Sm., Not. Bot. Card. Edinb. 11, 199, 1920 Shmb 5-8ft. Stems shoots slightly sulcate; mature subterete, pale yellow, not or

scarcely, in parts and very sparsely verruculose. Spines concolorous, 1 (-3)-fld., 6-12 (-15) mm. Leaves 10-20 (-27) x 3-5 (-9) mm.; narrowly obovate; cuneate, subsessile; obtuse ; entire, rarely with 1-3 spinules ; branched venation distinctly elevated, both sides finely reticulate; below concolorous green, sparsely finely papillose. Racemes sometimes compound below, 12-20-fld., 26-45 (-7) cm. Pedicels 2-3 mm., in the com- pound parts of the inflorescence, otherwise 5-7 (-10) mm. ; bracts 1-2 (-2.5) mm. F h e r s 6-7 mm. diameter. Outer sepak 2.5-3 x 2-26 mm., broadly ovate. Inner sepals 4-4*!j x 3-3.25 mm., obovate. Petals 3.5 x 2.5 mm. ; apex shortly incised; base shortly clawed. Stamens 3 mm., slightly produced, truncate. Ovules 1 (-2). Berries dark red, to red-purple, with short style.

N.W. Yunnan: Tung-chuling, fl. 31 May 1913, Kingdon-Ward 315 (Type, E); East Tibet border, Doker la, 13,000 ft., July 1913, Kingdon-Ward 707 (E) (this specimen has broader leaves, to 18 x 9 mm.); Atuntze, 9250 ft., 14 Sept. 1937, Yu 10207 (E); Doker la, Mekong-Salween divide, 12,000 ft., Sept. 1921, Forrest 20292 (K), ‘Shrub, 443 ft.; fruit purplish red’ (this specimen also has broader leaves, to 23 x 10 mm.). Not in cultivation.

(250) Berberis koreana Palib., Acta H&i Petropl., 17, i, 22, t. i, 1899; Nakai,

Shrub compact, very stoloniferous, erect, %4 ft. Stems with young shoots green; mature dark red, sulcate, not verruculose. Spines mostly conspicuously foliaceoua, encircling the utems, otherwise 3-7-fld., and palmate. Leaves to 6.5 x 3.2 cm., including petiole 1-8 Inm. ; oblong-elliptic to oblong-obovate; base contracted, apex rounded; margins 1%2o-spinose (1 mm.)-serrate (0.5 mm.), distant 2-5 mm.; both sides closely reticulate; above with the young leaves conspicuously mottled, or with the veins marked in, red, but finally a dull light green; below grey, pruinose. RaGemes 12-20-fld., 4-6 em. Pedicek 6 8 I-. F h e r s H.C.C. 3-311. Prophylls ovate, acute, 2-5 x 1-25 mm. Outer sepals 2.5 x 1-25 mm., ovate, subacute. Inner sepals 5 x 3.5 mm., obovate. Petah 4.5 x 3 mm., obovate, entire ; base cuneate with very conspicuous, oblong, separate, non-marginal

Flora Kmeana, J . Colt. Sci., Tokyo, (Art. 1, 26), 41, t. v, B, 1909. (Fig. 47)

Page 188: Berberis and Mahonia : A taxonomic revision

188 L. W. A. AERENDT glands, 0.7 x 0-3 mm. Stamm 4 mm., produced, rounded. oeyules 1-2. Berries red, lustrous, firm, globose, 7-8 mm.

Korea: Prov. Kyong-hui, 1899, Palibin, 8.n. (Type, K); in montibus Ouen-san, near Seoul, July 1906, Faurie 539 (BM); Aug., 1901, same loc., in ledgea, Farges 219.

Cultivated: L.A. 226, fl. 4 June 1942; fk. 30 Sept. 1942. A plant highly to be recom- mended for English gasdens, where its dark stems and unusual spines (of the tspe found generally in the S. American Congestifioraa) give distinction, in addition to the great beauty of the brightly marked spring leaves, the vividly red autumn foliage, the abundant flowers (late enough to escape spring frosts) and the brilliant berries.

(251) Berberis vulgaris L., 8p. PZ. 1: 330,1753

sepals. Oprules 2. Berries red, oblong, eatylose, dull.

account in J. Arn. Arb. 4,223-30, 1923.

Stems yellow, angled or sulcate. InJlo7escence racemose. Petals longer than inner

For detailed synonymy and a list of cultivated forms of B. vdgurk, see Schneider’s

KEY 47B Petals entire.

Leaves green above. Shrub 6-7 ft., with stems slightly angled or sulate.

Leaves with apex rounded. 251a. B. &aria var. vulgaris Leaves with apex acute. 251 b. B. vulgaris var. &ifolia

251~. B. vdgaria var. mluzta Leaves purple above. 251d. B. vulgar& var. puqnmfol ia

Pet& subentire, mulate. 251 e. B. x crendata

(251 a) Berberis vulgaris var. vulgaris Spiw 1-3-fld., 1-2 cm., sulcate. h v e a 2.5-6 x 0.7-3.5 cm., oblong, obovate, or

elliptic; decurrent petiole 1-8 mm.; mmgins 18-36-spinose (1 mm.)-sedate, distant 1.5-2.5 mm., opedy or scarcely reticulate; both sides dull. h n m 15-25-fld., 4-6 cm., including peduncle 1 cm. Pedicels 6-12 mm.; bracts 1-2 mm. Fbwers 1 cm. diameter. ProphyUs small. Outer se@ 3 x 2 mm., ovate. Inner sepccls 5 x 3 mm., obovate. Petals 5.5 x 3 mm., obovate, entire. ovules 2. Berries 9-10 x 4-5-5 mm.

Europe: Switzerland, fl. 5500 ft., June 1887, Gamble 18868 (K); May 1897, Gamble 26082 (K). According to Fedtschenko, generally in central Europe to the Balkans, and in European Russia from Ladoga to h e n , Upper and Middle Dnieper, Volga to Don, Black Sea border, Crimea and Lower Don, and northern Caucasus. According to Willkomm and Lange, Prodr., Fl. Hisp. 2,901,1880, it occum in wooded regions in the mountains of East, Central and Northern Spain, and the Pyrenees valleys a t 3000 ft.

Shrub 3-4 ft., with stems very sulcate.

Cultivated: very common, but might always be replaced by a better species.

(251 b) Berberis vulgaris var. acutifolia (Prmtl) Schneid., syn. B. acutifoliu Prantl., Id. Sern. Wireburg., 7 , 1873

(251 c) Berberis vulgaris var. sulcata Ahrendt, var.nov., ram& valde sulcatis; syn. B. Bulcata Hort.

Spiw stout (1 (-3)-fld., 1-3 cm. Leaves 4-5-7.5 x 1.6-3 cm., including very short petiole; obovate-elliptic; base contracted to petiole, 5-10 mm. ; margins 25-35-spinose (1.5-2 mm.)-serrate (1 mm.), distant 1.5-3 mm. ; apex obtuse to subacute; distinctly and fairly closely reticulate; below slightly paler green. Racenzes (&) lO-ZO-fld., 3 - 6 5 cm., racemose to subumbellate, including peduncle 1-15 mm. Pedicels 5-9 mm. ; bracts 1.2 mm., broadly ovate, acute. Inner se& 46-4-75 mm. Petals 4-75 x 2.75 mm., entire. Berries (immature), 8 x 3.5 mm., red, ellipsoid, estylose.

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Berberis and Mahonia 189

Albania: Tomor range, Abbas Ali peak, limestone cliffs, 6500ft., fr. 12 Aug. 1935,

Cultivated: fl. 24 Apr. 1948; fr. 20 Sept. 1947, at Maidwell Hall, Northampton. Alston & Sandwith 2460 (Type, K). Low growing shrub, up to 4 ft.

(251 d ) Berberis vulgaris var. purpurifolia Ahrendt, nom.nov., foliis rubro-pur- pureis; syn. B. vulgaris var. purpurea Hort., non Lodd. ex Loud., Hort. Britt.

Cultivated: fl. May 1933; fr. Oct. 1933. A garden form with dark reddish purple leaves; the usual garden name seems preoccupied since 1850 for a purple-berried form. There have been recorded references to plants of B. vulgaris with white, yellow, or black berries, but I have not seen any to correspond to these. Schneider refers to a var. subintegrifolia Giraudias, in Bull. Ass. pyr. (1896), (syn. var. rnicrophylla Coste in Bull. SOC. Bot. Fr., 1897) with lower habit, leaves 1.5-2 cm., more distinctly reticulate and more firm in texture, with margins ‘serrata vel subintegerrima’, short (scarcely to 3 cm.) 8-1 2-fld., racemes.

References for the other varieties or forms of B. vulgaris, whose nature seems in- definite, or which can only be traced in historical record are as follows.

(i) Var. serotinu Schrad., in Linnaea, 12, 364, 1838-late flowering; fruit pale red. (ii) Var. sanguinea Schrad., 1.c.-fruit deep blood red. (iii) Var. JEava Schrad., 1.c.-fruit golden yellow. (iv) Var. apyrena (asperma) Schrad., 1.c.-fruit seedless. (v) B. lucida Schrad., 1.c. 363, which appears to be a form of B. vulgaris with ‘leaves

somewhat lustrous, racemes aa long a~ the leaves, petals entire, fruit deep red, flowering 10-12 days before B. wdgaris, and with habitat Iberia’.

(vj) Var. viohcea Lodd. ex Loud., Hort. Britt. pp. 138-9, 1850-with violet. berries. (vii) Var. alba Lodd. ex Loud., 1.c.-white berried. (viji) Var. lutea Lodd. ex Loud., 1.c.-with pale yellow berries. (ix) Var. purpurea Lodd. ex Loud., 1.c.-with purple berries. (x) Var. nigra Lodd. ex Loud., 1.c.-with black berries.

Miller, in Card. Dict., ed. 8, 1768, refers to: (xi) ‘Berberis vulgaris without stones.’ He adds that this quality seems accidental,

that transplanted suckers produce stoned fruit, and deduces that it is the age which occasions the variation.

pp. 138-9, 1859

(xii) A form of B. vulgaris with ‘sweet black fruit’.

(251e) Berberis x crenulata Schrad., Linnaea, 12, 362, 1838 According to Schrader, this differs from B. vulgaris in having the flowering stems slender

and nodding, and slender green shoots; the leaves oblong-obovate, and the leaves of the sterile stems oblong-ovate ; the petals yellow, and not flame coloured, with the apex more or less crenulate (rarely denticulate) ; the leaves being somewhat greyish below. Accord- ing to F&. Vilm., p. 18, this is derived from B. vulgaris x B. canadensis.

(252) Berberis x laxiflora Schrad., I.c., 367, 1838; Schneid. J. Am. Arb. 4,213, 1923

KEY 47c Fruit ultimately dark red.

Stems pale yellow-grey. Leaves on flowering stems narrowly oblong-obovate, subobtuse, subentire ; racemes

Leaves on flowering stems oblanceolate, acute, closely spinose ; racemes peduncled. subsessile. 252a. B. x h x i j o r a var. laxiflora

252 b. B. x luxijora var. oblanceolata Stems purple-brown. 252c. B. x 1axJEora var. langeana

Fruit ultimately bright red. 252d. B. x ‘Brilliant’

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190 L. W. A. AERENDT

(252 a) Berberis x laxiflora var. laxiflora Shrub 5-7 ft. Stems very sulcate ; young shoots pale red, mature yellow. Spines 1-3-fld.,

7-15 mm., sulcate, with longer lateral pa&. Leaves 2-4.5 (-6.5) x 0.9-2 (-3.2) cm., obovate; subobtuse to subacute; attenuate to petiole 7-10 (-18) mm.; entire or indis- tinctly spinom; openly reticulate, below greyish pruinose. Racemes 4-9 cm., pendulous, sessile. Pediceb 6-10 (-20, below) mm. ; bracts 1.5-3 mm. Flowers 7-9 mm. diameter. ProphyuS small, broad, ovate. Outer sepda 2.5 mm., broadly ovate. Inner sepals 4.5- 5mm., elliptic-obovate. Petals M-5mm., obovate, subemarginate. O W 2, sub- sessile. Be& oblong ellipsoid, 10 x 5 mm., eatylose, at &t pale red, finally deep blood red.

Cultivated: by origin, reputed fo be B. vdgaris x B. &inemis. Not cultivated in England now.

(252b) Berberis x laxiflora var. oblanceolata Schneid., 1.c. 4, 214, 1923 Spines to 8 mm. Leaves 20-40 x 6-13 mm.; margins 8-15-spinose (0.75 mm.); reti-

culate. Racemes 3-7 cm. ; peduncle 1-2 cm. Pedicels 4-7 (-10) mm. Outer sepds 2 mm, Inner sepda 3-54 mm. Pet& 4-45 mm., emarginate. Be& 11 x 6 5 mm., a t first pale red, becoming dark red. Habit of B. chinensis. Not cultivated now in England. The former variety haa the broad leaves of B. vulgaris combined with the entire margin of B. chinensis; while this variety has the narrow leaves of B. chinensis with some serration derived fkom B. vulgaris. In both these varieties the pale yellow stems of B. wuZgariS, and the dark red berries of B. chinensis appear. In the following variety, the dark red stems of B. chinensis are manifest.

(252c) Berberis l d o r a var. langeana Schneid., 1.c. 4, 215, 1923 Spines discolorow, yellow, 1-3-5 cm., sulcate. Leaves 2-6 x 0.&2-5 cm. ; obovate-

elliptic or with few spinules. Racenzerr 2-5-4 cm. Pedkda 5-7 (-12) mm. Outer sepals 3-4mm. Inner se@ 4-5-6-8mm., slightly emarginate. Petals 5-6.5mm. Ovules sessile. Berries 10x5 mm.

Cultivated: origin in U.S.A., from seed of B. chinensis labelled B. serotina Lange. Not cultivated in England.

(252 d ) Berberis x ‘Brilliant’ Shrub 6-7 ft. Stem with young shoofs red; mature slightly red-brown, soon pale yellow

brown, sparsely distinctly verruculose. Leaves 3.5-6 x 3.5cm., ovate, contracted to petiole 3-5 mm. ; obtuse, entire, rarely with a few spinules; openly reticulate; above dull, below grey, pruinose. Racemes 16-26-fld., 7-10 cm., including peduncle 1-2 cm. Pedicds red (4-) 6-8 (-10) mm.; bracts 1-2 mm. Fbwer8 H.C.C. 3/1. ProphyZls lanceokate, 1-5 x 1 mm. Outer sepals 3 x 1-75 mm. Inner sepals 5 x 3.75 mm. Petals 5-5x 3-75 mm., obovate, entire ; glands conspicuous, orange, separat>e non-marginal, 0.9 x 0.45 mm. Stamens 4.5 mm., not produced. Berries bright red, H.C.C. 20.

Cultivated: L.A. 255, 0. 12 May 1942; fr. 30 Sept. 1942. This plant is the best of several known under this name. Its main characteristics are those of B.xhxi$ora. Since, however, its origin is unknown, it cannot be referred o by a varietal name attached to a binomial which belongs to hybrids between two known species. It may, equally well, be derived from, or have connexion with B. regeliana. It is recorded because of its outstanding ornamental garden value in berry, on account of which it might well replace B. vulgaris in cultivation.

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Berberis and Mahonia 191

(252e) Berberis x cerasina Schrad., Linnma, 12, 379, 1838 Shrub 5-6 ft. Stems terete; young shoots purple; mature red-brown. Spines 5-7-fld.

Leaves 2 4 x 1-1.2 cm., oblong-obovate; attenuate to distinct petiole; obtuse; margins spinose-serrate. Racemes 3-5 cm., erect to subpendulous. Pedicels 5-6 mm. Outer sepals ovatie-suborbicular. Inner sepals obovate-suborbicular. Petals emarginate. Berries ellipsoid, cherry red.

Cultivated: origin uncertain, but recorded that the leaves present no autumn colour. Not cultivated now in England.

(252f) Berberis x alksuthiensis Ahrendt, aff. B. vulgari sed ramulis rubro-brunneis, foliis subcoriaceis, petalis sepalis interioribus brevioribus distinguenda

Stem glabrous, mature sulcate, red-brown. Spines 1-2 cm. Leaves deciduous, to 6 x 2.8 cm., obovate, rounded; base contracted to decurrent petiole 1-3 mm. ; reticulate; fairly closely spinose-serrate; below, at first grey, pruinose, finally green. Racemes 10-2O-fld., 4-6 cm. Pedicels glabrous, 7-10 mm. Flowers 9-10 mm. diameter. Outer sepals yellow, 3 x 2 mm., ovate, acute. Median sepals 4-5 x 3-3.5 mm., oblong-ovate, obtuse. Inner sepah 6 x 4 mm., elliptic-obovate. P&.h 5 5 - 2 5 x 3 mm., obovate, acute, incised with rounded lobes. Ovules 2. Stumm 3.25 mm., not produced, truncate. Berries red, epruinose, 12-13 x 5-6 mm., oblong with very short style.

Cultivated (Kew 196125) : fl. 23 May 1942; fr. 13 Oct. 1939, a t Kew. The texture of the leaves is between those of its reputed parents B. vulgaris and B. chitria.

(253) Berberis bretschneideri Rehder in Sargent, Trees and Shrubs, 2 , 21, t. 110, 1907

Stem red-brown, terete. Spines solitary, to 1 cm., paler yellow-brown. Leaves 4-6 :r( 1-8-3 cm. ; oblong-obovate, obtuse ; attenuate; margins densely spinose-serrate; above bright yellow-green, reticulate ; below pruinose, grey, less veined. Racemes 10-1 5-fld., 2 - 5 4 cm., including peduncle 5-10 mm. Flowers 6-8 mm. diameter. Prophylls ovat'e, 2 x 1.5 mm. Outer sepals reddish, 2.5 x 2 mm., elliptic-obovate. Inner sepals 4-5 :r( 2-5-2.75 mm., obovate. Petals 4-5 mm., marginate. Ovules 2. Berries dark red to red-purple oblong, estylose, 10-12 x 5.5-6.5 mm.

Chihli: near Pekin, seed only, 1880, Bretschneider, 8.n. Cultivated: in V.S.A., from seed collection cited, Type. I have not seen a plant of

this species cultivated in England.

(254) Berberis pachyacantha Koehne, Deutsche Den&. 170, 1893; Parker, A Forest Flora for th.e Punjab, with Hazara and Delhi, 12, 1918

Shrub 6-10 ft. Stem dark red, stout, subterete. Intern& 2-4 cm. Spines often absent, otherwise 1 (-3)-fld., 2-6 (-15) mm., plane to subterete. h c e s from 3-5-55 cm. x 1-1-7 cm. to 7-10 x 3-4 cm. ; oblong-obovate; base contracted to petiole 7-17 mm. ; apex rounded or obtuse ; margins 1630-spinose (0-25-0-5 mm.)-subsemlate, distant 1-3 mm. ; closely reticulate; below concolorous green, epruinose, epapillose. Racemes 15-25-fld., 4-10 cm. long, including peduncle 1-1.5 cm. Pedicels 6-10 mm.; bracts 1 mm. Outer sepals 3-3-75 x 2-2.25 mm., ovate, obtuse. Inner sepah 5-6 x 3.5-4 mm., obovate. Petals p4.5 x 2.5-3 mm., acutely emarginate with acute lobes. Ovules 1-2. Berries red, oblong-ovoid, or obovoid, estylose, epruinose, 8-10 x 4-5 mm. Xeeds black, 6 x 2.5 mm.

Ksshmir: Kishtwar, 8000-10,000 ft., and Sind valley, 9OOO ft., Thomson, s.n. Kala- pani, Kamrinala, Astor valley, Inayat, 10 July 1901, Duthie 25468; Pass from Logan to Oolso, fl. 17 June 1847, Hooker 1241 B; Naboog High Pass, fl. 16 May 1847, Hooker 1241

Page 192: Berberis and Mahonia : A taxonomic revision

192 L. W. A. AERENDT (all, K); Gurez, Kishenganga, 8000 ft., fr. 23 Sept. 1940, Ludlow & S h e d 8229 (BM); Mantnar Nullah, Bringhi valley, 11,0oOft., fr. 18 Oct. 1940, Ludlow & SherrifF 8272 (BM); Sinthan Paas, head of Bringhi vdey, 11,OOOft., fr. 24 Oct. 1940, Ludlow & S h e d 8181 (BM).

Punjab Himalaya: Simla, Nagkanda, 9500 ft., 30 May 1878, Gamble 60721) (K). Garhwal: 10,OOO ft., Schlich 25 (l), (0). According to Parker, common at 8ooo-

12,OOO ft., in moist shady places. Cultivated: from Ludlow & Sherrif€ 8229. I had a plant from 1942 to 1950 when it

died; it is to be hoped that other specimens from this number survived. This species has been confused with the Nepal B. petwhris which has entire thick leaves, and con- spicuously long style. B. bhmirana differs from B. p a c h y w n t b conspicuously in its yellow stems and subacumimte leaves.

(255) Berberis henryana Schneid., BuU. Herb. B&,qier, (2), 5, 664, 1905; (2), 8,261,

Shrub 6-8 ft. Stem lustrous red-brown, subterete to subsulcate. Spines sometimes absent, mostly weak, 1-3-fld., occasionally on the shoots to 3 cm. Leaves 1-5-3 (-6) x 0.8-1-8 (-3) cm.; elliptic-obovate; obtuse; cuneate to short decurrent petiole; margins 12-20-spinose (1-1.5 mm.)-serrulate; distant 1-3 mm.; openly, or scarcely, reticulate; above dull light green; below grey, pruinose. Racernee 10-20-fld. (2-) 3-6 cm. long, including peduncle 1-2 om. Pediwh 5-9 mm.; bracts 1-16 mm. Prophylki lanoeolate, scuminate, 1-16 mm. Outez se+ 2-5-3.5 x 1.5-2 mm., oblong-obovate. Inner sepals 56-66 x 4-44 mm., obovate. P& 5-6 x 4-5 mm., oblong-obovate, incised; glands elliptic, separate. Stumm 36-4.5 mm., not produced, truncate. Ovuka 2. Berries red, scarcely pruinose, ellipsoid with short style, 9 x 6 mm.

W. Hupeh: near Patung, Henry 5470 B (Type, K); Changyang hsien, mountains, May 1900, W h n (Veitch) 645 (K); shrub 5 ft.

E. Szechuan: Tschen-kaxi-tiu, Farges 550, cited by author, as also: Shensi: Tai-pai-shan, Sept. 1899, Giraldi 2310; Kian &an, Lao-y-shan, 4 Aug. 1897,

1908; Sargent, PI. Wils. 3,440, 1917

Giraldi, 2303.

(256) Berberis farreri Ahrendt, spmov., aff. B. hnryanue a qua foliis multo angus- tioribus saepe integris, crebre reticulatis, subtus viridibus, pettllis subintegris, distinguenda

Stem subterete to subsulcate; mature red-brown. InternodRs 2-3 cm. Spines often absent, sometimes weak, 1-3-fld., 3-8 mm. Leaeres 13-30 (4) x 3-6 (-15) mm.; base contracted to decurrent petiole 3-6 (-15) mm. ; margins often subentire, yet often 10-20- spinose (0.35-0.5) mm.-subserrulate; finely and fairly closely reticulate both sides; below ultimately epruinose, green. Ru.c.eme.9 15-22-fld., 3-4 cm. Pedida glabrous, slender (5 mm., apex-) 7-11 mm. Flowers c. 8 mm. diameter. Outer sepals 3 x 2.5 mm., broadly obovate, rounded. Inner se+ 5.25 x 2.25 mm., narrowly obovate, rounded. Petuh 4 x 1.75 mm., narrowly obovate, obtuse, subentire; slightly clawed, with oblong, separate, non-marginal glands, 0-8 x 0.3 mm. Stamens not produced, obtuse. Ovules 2.

West Kansu: Lo sin, Lang chan, 9 July 1918, Licent 4138 (K); seed only, Farrer 318. Cultivated: from Farrer 318, fl. 5 June 1939 (Type, 0), ex H. G. Hawker, Stode,

Devon, comm. A. B. Jackson.

(257) Berberis aQricola Ahrendt, sp.nov., ramulis atrorubria sulcatis, foliis integris,

Stem sulcate, mature dark red. Spines yellow, sulcate, stout, 1-2 cm. I&& 1.5-3 cm. Leaves obovate to 28 x 13 mm., entire; venation branched, scarcely sub- reticulate; below grey, pruinose. Racenzes 16-24-fld., often verticillate, rarely compound

infloreacentiis racemosis, fructibus stylosis distinguenda

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Berberis an& Mahonia 193

below, 4-7.5 cm. long. Pedhls 5-7 mm., slender, with bracts 1.5 mm. F k r s c. 10 mm. diameter. Outer sepals 3-3.5 x 1-5-2 mm. Inner s e p b 5.5 x 4 mm., obovate. Petals 4.5 x 2-25 mm., obovate, entire. Stamens 3 mm., scarcely produced, rounded-truncate.

S.E. Tibet: Prov. Kongbo, Dzala, Pasum Chu, 12,5OOft., fl. 3 July 1947 Ludlow, S h e H & Elliot 14071 (Type, BM), 'shrub W f t . , in hedges bordering cultivation'; Kyonka, Pasum Chu, 12,000 ft., fl. 19 June 1947, Ludlow, Sherrif€ & Elliot 13937 (BM), 'shrub 4-5 ft., growing on the edges of fields'. The latter number has slightly shorter racemes, and slightly larger flowers. This would appear to be a very attractive plant, but it is not in cultivation.

ovules 1-2. style distinct.

(258) Berberis atroprasina Ahrendt, Card. Chron. (3), 112, 155, Oct. 1942 Shrub 5-6 ft. Stems with young shoots green; mature yellow-brown, subangled to sub-

sulaate. Internodes 2-4 cm. Spines concoloroua, sulcate 1 (-3)-fld., 2-5 (-8) mm. Leav,esto 5 x 1-7 cm., though on the shoots to 8.5 x 2.5 cm., including decurrent petiole 5-12 mm.; elliptic-obovate; attenuate, cuneate ; obtuse; margins 15-25-spinose (0-5-1 mm.)-sub- serrulate, 1-2 (-3) mm. distant ; both sides fairly closely and distinctly reticulate ; above lustrous dark green; below a t first slightly pruinose grey, finally somewhat paler, but still deep green; on the shoots dull and brighter green above, the margins entire or with 2-6 spinules. Racemes 12-20-fld., 3-5 (-8) cm., including peduncle 3-10 mm Pedicel8 8-12 mm.; bracts 1-1.5 mm. Prophylls 2.25 x 1 mm., red, subacuminate. Outer sepal8 4 - 5 5 x 2-25-2.5 mm., oblong-obovate. Inner sepals 6 x 3.5 mm., rhombic-ovate. Petals 5 x 2.5 mm., obovate-elliptic, entire; base shortly clawed, with separate glands 0.9 x 0.7 mm. Stamens 3.5 mm., not produced, truncate. O v u h 2-3. Berries deep carmine red, oblong, 8-9 x 4 4 . 5 mm., with style absent or not distinct.

N.W. Yunnan: 1932, seed only, Forrest, s.n. (inaccurately distributed as no. 29012). Cultivated: from the Forrest seed cited, fl. May 1940; fr. 17 Oct. 1940, Sept. 1942

( T Y P , 0).

(259) Berberis amurensis Rupr., BuU. A d . Sci. St-Pdtersb. 15, 260, 1857; B. Fedt-

Shrub 8-11 ft. Stem more or less angled or sulcate, mature pale yellow to grey. Intemzodes 2.5-4 (-7) cm. Spines 3-fld., 1-2 cm. Leaves oblong, ovate or elliptic; con- tracted to a decurrent petiole; margins 40-60-spindose (0.5 mm.) subserrulate, distant 1-2 (-3) mm.; apex acute; both sides openly, or more or less indistinctly, reticulate, and conoolorous green. Pedicel8 5-7 (-10) mm. Outer sepals 3 x 2 mm. Inner sepah 5.5- 6 x 33-325 mm., all obovate. Petals 4.5-5 x 2.5-3 mm., slightly emarginate. O v u h 2. Berries 10 x 6 mm., oblong, estylose, red, slightly pruinose blue at the base.

schenko in Komarov, 3'1. U.R.S.S., 7,557, 1937

K E Y - 47 D

Recomes (S) 4-10 cm. long, 10--25-fld. Leaves 5-10 x 2.5-5 cm. 259a. B. anturenab var. amuremia Leaves 5-7 x 3-5 em. 259 b. B . amuremia var. latqolia

2590. B. amuremia vm. licentii Racemes 16-3.5 cm. long, (&) 8-15 (-20)-fld.

(2590) Berberis amurensis var. amurensis Rcmmes 4-10 cm. long. S.:E. Manchuria: fr. 1860, Maximowicz, s.n., Iter secundum (K). Amurland : Amur river valley (Type, Leningrad). N. Chihli: Purdom 34. N. Shansi: Ma-kia-pou, 30 July 1914, Licent 669 (K).

13 JOURN. LI". SOC. BOTANY-VOL. LvII

Page 194: Berberis and Mahonia : A taxonomic revision

194 L. W. A. AHRENDT Korea: Prov. S. Kanlego, Eiko, bush of 15 ft., only one seen, fr. 22 Sept. 1917, Wilson

9225 (BM, K); Prov. Kogan, Kongo-am, 4-8 ft., in thickets, not common, fr. 12 Oct. 1917, fiwit scarlet, Wilson 9249 (K).

According to Fedtschenko, a t edges of woods, and banks of mountain streams on stony mil in the Far East: Ueeurak valley.

Cultivated: from Wilson seed at Maidwell Hall, Northampton, fr. 20 Sept. 1947.

(2593) Berberis amurensis vax. latifolia Nakai, PI. Coreanna (J. Cdl. Sci., Tokyo, 26, Art 1) 41, 1909

Racenzes 3-5 cm. long. Korea: above Do-do, 1 June 1917, bush 3-6ft., local, Wilson 8566 (Type, K). Not

cultivated.

(259~) Berberis amurensis var. licentii Ahrendt, vamov., foliis minoribus, racemis abbreviatis

Leaves to 4.5-6 x 1.6-3 cm., or 5-5-6x 1-7-2 cm.; decurrent petiole 1 cm.; apex acute; margins 12-20-spinose (0.5 mm.)-sedate (0.5 mm.). Racemes 1-5-3.5 cm. Pediuh &9mm.;bractslmm. Berries9x6mm.

Central Shansi: King-hsien, fl. 5 May 1916, Licent 1770 (Type, K). South Shansi: Nan-keou, limy country, immature fruit, 21 June 1915, Licent 1223 (K). Chihli: Tientsin, Suan Yang Tong, Licent 2866 (K). Central Mongolia: fr. 11 Sept. 1917, Licent 3542 (K). Not cultivated.

(260) Berberis hersii Ahrendt, Card. IUu8.t. 64, 426, 1944. Stem yellow, finely sulcate. Spines mostly absent, occasionally weak, solitary,

3-6 mm. Leaves 2.5-5 x 1-14 cm., obovate; contracted to decurrent petiole, 6-15 mm.; margins 15-3o-spindose (0.5 mm.)-subsedate, distant 0.5-1.5 mm. ; both sides closely reticulate; below at first grey, pruinose, finally green. R a c e w 15-30-fld., 3-5 cm. Pedicel% slender, dark red (2-) 4-6 (-8) mm.; bracts 1-16 mm. F’lowers 8 mm. diameter. P r q h y l h red, acute, 2 x 1 mm. Outer se@ 3-3.5x2-2-5mm., ovate, obtuse. Inner selpals 4.5-5 x 3-3-5 mm., obovate. Petals 4-46 x 2.5-3 mm., elliptic; shortly emar- ginate; shghtly clawed, with oblong, separate, non-marginal glands, 0.6 x 0-4 mm. Stixmn-s 2.5 mm., not produced, truncate, or retuse. Ow&s 2. Berries ellipsoid, estylose 9 x 5.5 mm., red, epruinose.

ShitaFli: Nan yang shan, 6500 ft., fr. 22 Sept. 1923, Hers 2730, 2783. Cultivated: from Hers 2783, a t Kew, fl. 12 May 1939; fr. 21 Oct. 1939 (Type, 0 ) ; from

Hers 2730, a t Kew, fl. 21 Apr. 1939.

(261) Berberis regeliana Koehne ex Schneid., Mitt. Dtsch. Dendr. Ges. 1905, 119, 1905; syn. B. vulgaris var. japonicu Regel., Acta H&i Petropol. 2 , 415, 1873; Ranch. et Sav., Enum. PI. Jap. 2,273; Nakai, PI. Coreana (J. Coll. Sci., Tokyo, 26, Art. 1,) 41,1909; syn. B. japonicu (Regel.) Schneid., Bull. Herb. Boissieer, (2), 5, 667, 1905; syn. B . amurensis var. japonica (Regel) Rehder in Bailey, Cycl. Amer. Hort. 1,154,1900

Shrub 5-6ft. Stenzs pale yellow-grey, angled, sulcate. Spines 3-fld., slender, con- colorous 1-2-2.2 cm., sulcate. Leaves 3 x 1.5-2 to 6 x 3-4 cm., excluding decurrent petiole 7-15 mm.; base contracted; apex rounded; obovate; margins 25-40 spinose (1 mm.)- s e d a t e , distant 1.P2.5 mm.; very closely reticulate; above dull green; below whitish grey, pruinose, papillose. Injlorescenw (5-) 10-15 (-18)-fld., openly racemose (2-) 3-6 cm. long. Pedicels 4-9 mm. Flowers 8-10 mm. diameter. outer s e w 2.75-3 x 1.8 mm. Inner sepals 5-6 x 3 4 mm., all obovate. Petals 46-65 x 2-5-3 mm. O& 2. Berries red, slightly bluish pruinose at the base, oblong, estylose, 10-11 x 4-5 mm.

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Berberis and Mahonia 195

Japan: Prov. Nambu, 1865, Tschonoski, 8.n. (Type, K); Kuroshi 26 Sept. 1889, Faxlrie 1287; d'Akita, 5 May 1888, Faurie 2119; Cote de Fukuyama, 2 June 1889, F a d e 3806; Yeso near Hakodate, Small, s.n. Hokkaido, Fukuyama, Prov. Oshima, 18 July 1890, Mirjaba & Tokomuithu, s.n.; Hondo, 26 May 1914, Wilson 6763, 6 July 1914, Wilson 7131, 18 Oct. 1914, Wilson 7686; also Wilson 6802,6904 (all, K).

Cultivated: L.A. 28, 0. 6 June 1942; fr. Nov. 1942. This species is often distributed inaccurately as B. hrmna or B. sieboldii.

The following hybrids, which are insufficiently known, are placed a t the end of this section to which they apparently belong.

(262a) Berberis x meehanii Schneid. ex Rehd., Manual of Cultivated Trees and Shrubs,

Sterns angled, grey-brown. Spinea solitary, 1-2cm. Leaves elliptic to obovate, 2 . 5 4 cm. ; densely spinose-serrate; below reticulate. Racenzee 4-9 cm. Berries ellipsoid, red, slightly pruinose.

Chltivated: in U.S.A.; origin unknown. Distributed by Meehan as B. concinnu. Not cul1;ivated in England.

247, 1927

(262b) Berberis x bella Schneid., J. Am. Arb. 4, 205, 1923 Stems with young shoots purple; mature yellow-grey ; subangled. Internodes 1.5-3 cm.

Spines 1-3-fld., stout, yellow, 1 4 cm. ; subsulcate. Leaves 2 4 x 0-8-2 em., oblong-obovate to elliptic; cuneate to petiole 1 cm.; margins entire or with 1-6 inconspicuous spinules; ope.nly reticulate; below grey, pruinose (but on shoots more ovate, acute, and spinose, but with shorter petiole). Racemes dense, 4-6 em., including peduncle 1-1.5 cm. Pedicels 4-7mm.; bracts 1.5-2mm. Flowers 7-8mm. diameter. ProphyuS small. Stumns apioulate. Ovules 2, subsessile. Berries ovoid to ellipsoid, 10-11 x 5-6 mm., pale red, with style absent or inconspicuous. Seeds purple.

Cultivated: (Type, Herb. Am. Arb.). Not cultivated in England.

Sect. Brachypodae Schneid., Sargent, PI. Wils. 3, 442, 1917 Stems mostly pubescent or puberulous, mature yellow. Internodes (1-) 2-5 (-6) 6 cm.

Xpiiles fairly stout, 1-3-Ad. (1-) 1.5-3 (4) cm. Leaves deciduous (2-) 4-8 (-12) cm. long, including petiole 6 2 0 mm., mostly closely spinose, always pubescent or puberulous above or below. Racenzes 20-50-fld., mostly very narrow and spikelike, (2-) 6-12 cm. long. Pedicels 2-5 (-8) mm. long; bracts mostly as long as or longer than the pedicels. Flowers 8-10 mm. diameter. Prophylls sometimes in two series. Sepals in three series. Petals emarginate. Sturnens generally scarcely produced, truncate. Berries red, generally estylose (6) 9-11 mm. long, but black in one hybrid.

Kansu, Hupeh, Shensi, Shansi.

KEY 48, to Sect. BRACHYPODAE (Maps 2, 28, 34) Berries red.

Stom always glabrous; petals longer than inner sepals. .Leaves P 1 0 om.; subacuminate; fruit 10 x 9 mm., lustrous. Leaves 2-4 cm., subacute; fruit 10 x 5 mm., dull.

Pedicels (I-) 2-3 (-4) mm. ; bracts longer than, subequal to, or at least half as long

263. B. giraldia 272b. B. x bombycinu

Stoms pubescent, at any rate at k t .

as, the pedicels. Petals longer than inner sepals.

Mature stems pale yellow-grey ; leaves closely spinose. Leaves 3-6 cm., acute; prophylls 4, in 2 series; sepals 9, in 3 series, pedicels

13-2 pubescent. 265. B. brachypoda

Page 196: Berberis and Mahonia : A taxonomic revision

196 L. W. A. AERENDT

KEY 48 (continued) Leaves 6-12cm., subacumimte; prophylls 2, sepalls 3, both in one Series; pedicels glabrous. 266. B.8alicaria

Mature stems dark red-brown; leaves subentire. 267. B.

S t e w with shoots pubescent, and mature stems glabrous ; leaves very pubescent ; Petals shorter than inner sepals.

pedicels 1-2 mm.; bracts much longer, 2-3 mm.; fruit subglobose 4-5 mm. 268. B. stenostachya

Stems with shoots and mature stems puberulous; leaves only pubendous; pedicels 2 - 5 4 mm., with bracts (1.6-2-5 mm.), shorter or subequd.

269. B. mit~ollia

270. B.&escens 271. B.mqnifolia

Berries blue-black. 272a. B. x gibbsii

Pedicels 5-10 mm.; bracts 1 mm., very much shorter. Leaves 2-4 cm., entire or subentire; racemes 2-4 cm. Leaves 6-10 cm., closely spindose; racemes 6-10 cm.

(263) Berberis giraldii Hesse, Mitt. Dts&. Dendr. Ges. 1913, 272,1913. (Fig. 50) Shrub 6-8 ft., with sparse, little branched, loosely arching and spreading stems. Stems

slender, sulcate, yellow. Internodes 3-6 cm. S p i w solitary, 1-2 cm. Leuves 4-10 x 1-5- 4 cm., including petiole 1-2 om.; rhombic-ovate; base contracted; margins 30-50- spinose (1 mm.)-serrate (1 mm.), distant 1-3 mm.; scuminate; openly reticulate; above dull green; below slightly paler lustrous green, with pubescence on veins and veinlets. Racemes 25-50-fld., spikelike, 6-1Ocm. long, including peduncle 2-3cm. Pedicels glabrous, 2-3 mm. ; bracts subequal 1-5-2-5 mm. Pr0phyU.s in three series. OuterpophylLs at mid-point of pedicel, red, lancedate, 2 x 0-5 mm. Hdian and iraner propJayZLs in two series of three, a t apex of pedicel appressed to the flower, lanoeolate, truncate, acumi- nate, 1.5-2 x 0-5-0.75 mm. F’lowers pale, H.C.C. 2. Outer 8 e p . b 2-2.25~ 1-16 mm., partly red, oblong-ovate, obtuse. Median se@ 3 x 2.5 mm., oblong-elliptic. Inner 8epa.k 5 x 4 mm., obovate. Petals distinctly longer than inner sepals, 5.5-6 x 2 - 6 3 mm., oblong-obovate; acutely em-ate, with mute lobes; conspicuously clawed, with glands subconcolorous, oblong, separate, marginal, 1 x 0.5 mm., situated above the claw. Skcmens 4-46 mm., not produced, truncate. Ovules 2, sessile. Berries firm, very lustrous, estylose, ellipsoid-globose, 10 x 9 mm.

Shensi (2 ) : the author refers merely, with a brief description in German, to plants growing from seed (from N. China,), presumably collected by Giraldi. As some of this seed produced the plant cited below, the specimens from it may be regarded aa a sub- stitute for the type.

Cultivated: L.A. 120, raised by the late W. J. W c h a n t from seed sent by Hesse, fl. 24 June 1942; fi. 22 Oct. 1941 (BM). A most remarkable species in having so many prophylls aa well aa sepals. In cultivation, a plant conspicuous in its unusually shaped leaves, which are bright red when young, and magnificent in berry. Its unique garden value is somewhat spoiled by its ungainly habit. My own plant died some few years after observations were made, and, on inquiry, I found that Mr Marchant’s plants at Wimborne were also dead. It is uncertain whether the species is still in cultivation. Plants which paas under this name are often B. geraldii Hort., which is B. aggregata Schneid., or a form of this.

With a sparse description, but apparently very closely related to B. giraldii is:

(264) Berberie subsessillflora Pamp., N m Ciona., Ital., N.S., 22,293, 1915

N. Hupeh : Zan-lan-scian, U-kia-pi, Oct. 1913, Silvestri 4064; Lao-sciu-ze-ze, Nov. 1912, Silvestri 4099. Not in cultivation. For another rare species &om this locality compare B. zanlanscianen& (Sect. Wallichianae, no. 62).

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Berberis and Mahonia 197

(265) Berberis brachypoda Maxim., BUU. A d . Sci. St-P&rsb. 23, 308, 1877 Shrub compact, 3 4 ft. Stern sulcate; young shoots green, pubescent; mature yellow-

grey, glabrous or subglabrous, not verruculose. Internodes 2-3 cm. Spines sulcate con- colorous, 3-fld. (I-) 2-3 (4) cm. long. Leaves 3-7-5 x 1-2-3.3 cm. ; oblong-elliptic; acute; shortly cuneate to decurrent petiole 5-10 mm.; margins 25-40-spinose (06-1 mm.)- subserrulate, distant 1-2 mm. ; reticulate; above dull light green, shortly pilose; below concolorous, elongated pilose. Rmmes 20-30-fld., spikelike 7-12 cm., including peduncle 1.5-4 cm. Pedicels pubescent, 2 4 mm.; bracts 2-4 mm. Plowers vividly red without, 8 mm. diameter. Prophylls red, lanceolate, acuminate, in two series, each of two. Outer prophylls situated near mid-point of pedicel. Inner pophylls appressed, aa usual, to the flower as epicalyx. Outer sepah 2 x 1.5 mm., red, prophyll-like, ovate, acute. Median sepals 3 x 2.5 mm., oblong-obovate. Inner sepals 4.5 x 3 mm., obovate. Petals longer, 5 x 2.75 mm.; apex acutely emarginate, with acute lobes; glands oblongorbicular, separate, non-marginal. Stamens 3 mm., not produced, truncate. Ovules 1-2. Berries hn, lustrous, blood red, oblong, 9 x 5 mm., excluding short, thick, distinct style.

Kansu: 1875, Piasezki, s.n., Type (W); Sin long chan, 9 July 1918, Licent 4087 (K). Shansi: Yeo-teou, 6 June 1916, Licent 2008 (K); (N.) between Kachen and Ponshien,

11 July 1915, Licent 1448 (K). Cultivated: L.A. 46,0. 6 June 1942; fr. 3 Oct. 1939. This species is rare in cultivation,

but might well be more extensively planted, aa it provides, with a small and compact habit, ornamental racemes of flower and fruit. Its flower structure is unusual and some- what similar to that of B. giraldii.

(266) Berberis salicaria Fedde, Bot. Jb. 36, Beibl. 82, 42, 1905; syn. B. brachypoda var. salicuria (Fedde) Schneid, Bull. Herb. Boissier, (2), 8, 262, 1908.

Shmb 6 ft. Stem pubescent, mature yellow-grey. Spines 1-3-fld., 2-4 cm. Leaves 3-12 x 1 4 cm. ; lanceolate, subacuminate; margins 15-40-spinose (1 mm.)-subsedate, distant 1-2 mm.; narrowly reticulate; both sides concolorous green; above dull; below slightly lustrous. Racemes spikelike, 7-12 cm. Pedicels glabrous, 2-3 mm. Flowers (Z immature) 2.5 mm. diameter. Prophylls ovate, acute. Outer sepals shorter than, and petals longer than, inner sepals. Ovules 2.

Shensi: Si ku tzui shan, and Monte tu lian pin, Giraldi 7019 (Type, K). Not in culti- vaticin.

(267) Berberis gilgiana Fedde, Bot. Jb. 36, Beibl. 82,43, 1905 Sturns puberulous; mature red-brown to purple. S p i w solitary, 0-8-2.7 cm. Leaves

1 4 r: 0.3-1-5 cm., including short petiole; base attenuate; apex acute; margins entire, or 2-9-spinose (0.25-0.5 mm.)-subserrulate, distant 2-5 mm., towards apex; clearly narrowly reticulate; both sides dull pale green, finely puberulous. Racewas spikelike, 4 6 em., including pubescent peduncle 1-3 om. Prophylls 2, ovate, acute. Outer sepals small. Inner sepab obovate. Petals slightly longer than inner sepals.

Shensi: Lu kiu piu, Giraldi 2307; Kan y shan in Lao y shan, fl. 2 May, Giraldi 2308; Quan tou shan, Giraldi 2309 (Type, K); Tai-pei-shan, Purdom, 6, 8 (K).

(268) Berberis stenostachya Ahrendt, sp.nov., ram& foliisque valde pubescentibus ; bracteis pedicellis multo longioribus ; petalis sepalis interioribus brevioribus ; fructibus tantum 5 mm. longis, distinguenda

Stem angled-sulcate ; young shoots very pubescent; mature glabrous, pale yellow. InternodaP 16-26 cm. Spines 3-fld., 1-2 cm. Leaves to 6.5 x 2.5 cm., including petiole P I 0 mm. ; contracted to decurrent petiole; apex acute or subacute; margins very sinuate 10-20-spinose (0.5-1 mm.)-serrate (0.5-1 mm.), distant (I*&) 2-4mm. ; reticulate; pilose

Page 198: Berberis and Mahonia : A taxonomic revision

198 L. W. A. AHRENDT both sides; above dull green; below slightly paler green. Racepnes 2!%0-fld., spikelike, 2-5-45 cm. long. PedktA pubescent, 1 (-2) mm. long; bracts pubescent, much longer, 2-3 mm. F h r 8 c. 5 mm. diameter. Prophyb 1 x 0.5 mm., red, ovate, acute. outer sepal4 2 x 1 mm., ovate. Inner 8epd.s 3.5 x 1-5 mm., obovate-elliptic. Petale 3 x 1.25 mm., emargiwte with acute lobes; base cuneate with widely separate marginal elliptic glands, 0.3 x 0.2 mm. Stmaem 2 mm., not produced, truncate. oerules 2. Berries red, epruinose, estylose, ellipsoid, 5 x 3.5 mm. S& brown.

Kansu: seed, Fhck 15829. Cultivi4.d: from Rock 1-5829, fl. May 1939; fr. 13 Oct. 1939 (both at K) (Type, 0).

(269) Berberia mitifolia Stapf, Bot. Mag., 154 t. 9326, 1931; syn. B. brachylpoda

Shrub 6-8 ft. Stems with young shoots finely sulcate, green or slightly reddish, puberu- lous; mature, subsulcate, yellow, remaining puberulous, black-ve~~uculose. I&& 2-4 (-5) cm. Spine8 1-3-fld., stout, 1 - 5 4 cm., sulcate, concolorous. Leaw 3-5-6.5 x 1- 2.2 cm., including petiole 1-16 cm.; oblong-obovate; base cuneate; apex generally acute; margins 15-25- (-#)-spinom (1 mm.)-serrate (1 mm.), distant 1-2 (-3) mm.; above slightly lustrous yellow green, subglabrous; below subconcolorous green, with pubescence on the veins and veinlets; openly reticulate both sides. Rcceenzes 20-30-fld., 5-9 cm. Pedicels puberulous, 3-4 (sometimes in fruit, to 6) mm. ; bracts 16-26 (-3) mm. F k r s H.C.C. 2. Pr0phyU.s red, 1-8 x 1.2 mm. outer sepal.8 3 x 2-2-25 mm., ovate, obtuse. Median 8epd.a 4.55 x 3-3.5 mm., oblong-elliptic. Inner s e p h 5-6 x 44.5 mm., obovate. Petals 4-5-5.5 x 3-3-5 mm., oblong-elliptic; apex emargjnate, with two acute lobes; base scarcely clawed, with obovate, separate non-marginal glands, 0.8 x 0-6 mm. Stumm 36-3-75 mm., slightly produced with apex truncate a t perimeter, but centrally apiculate or subapiculate. ovoc2ea (1-) 2. Be& red, epruinose, estylose, oblong, 10-11 x 6-7 mm.

W. Hupeh: Fang Hsien, uplands thickets, 10,40&10,70Oft., fr. Oct. 1910, Wilson 4416; Hsing shan h e n , woodlands, 42003200ft., fl. June; fr. Nov. 1907, Wilson 554 (K).

Cultivated: at Kew, from Wilson 4416, for plate of Bot. Mag. t. 9326 (Type, K); also from same origin, L.A. 72, fl. June 1938, 18 June 1941,20 May 1942,26 Apr. 1945; fr. 31 Oct. 1939.

var. g.ibbsii Hort., in part

(270) Berberis pubescens Pamp., Nuovo ciorn., Ital., s.n., 17, 273, 1910; syn. 'B.

Stems with young shoots reddish, pubescent ; mature yellow-grey, glabrous. I&& 1-2 cm. S p i w mostly solitary, 1-2-5 cm., sulcate. Leaves 2-3 (-4) x 0.5-0.7 (-1) cm., obovate, mucronate; attenuate to decurrent petiole, 7-10 (-15) mm., included; margins mostly entire, rarely with a few spinules; reticulate and pubescent both sides, but more markedly veined below. Bwew 2-4 cm., including peduncle 1-1-5 cm. Pedicele 5-8 mm.; bracts 1 mm. Be* p6 mm.

gilgiuna' Schneid. in Sargent P1. Wils. 3,440, 1917, in part, non Fedde

N. Hupeh: Monte Niang-niang, Silvestri 718, Type. Not cultivated.

(271) Berberis magnifolia Ahrendt, sp.nov., aff. B. lyubescenti sed foliis majoribus, 5-10 om. longis, crebre spinoso-serrulatis; racemis elongatis, distinguenda

Shrub 6-7 ft. Stem8 with young shoots reddish, puberulous ; mature slender, yellow, gla- brous, sulcate. Internodes 2-5 em. Spin.e.8 1-3-fld., 1-5-3 cm. Leaves 5-10-5 x 1-6-36 cm.; elliptic or rhombo-elliptic; petiole 1-1.5 cm. ; margins 20-40-spinulose (0.5 mm.)-sub- s e d a t e , distant 0-5-2 mm. ; apex acute or subacute; openly reticulate both sides; above dull light yellow-green; below paler, greyish pruinose; both sides puberulous on the veins and veinleta. Racemes sometimes compound below, 25-35-fld., 6-10 cm. long, in- cluding peduncle 2 4 cm. Pedic4.8 subglabrous, or very slightly puberulous, 5-10 mm.;

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Berberis and Mahonia 199

bracts 1-14mm. Prophylls in two series of three each, appressed to the flower as epicalyx. Outer and inner prophylls both reddish, lancedate, acuminate, 2.5-3 x 0.4- 0.8 mm. Flowers varying in colour from H.C.C. 6412 to 112. Outer sepals lanceolate, acute, 2.5-3 x 1-16 mm. Hedian sepals 3.75 x 2-5 mm., oblong-elliptic, rounded. Inner sepals 4.5 x 2-75 mm., obovate. Peta2s 3 x 1.75 mm., elliptic, acute, subentire or shortly to minutely emarginate with acute lobes. Stamens 2.5 mm., slightly produced, truncate. Ovules 2. Berries red, epruinose, 9 x 5 mm., with short thick style.

Cultivated: L.A. 737, fl. May 1948; fr. 2 June 1952 (Type). An interesting species of this small section. It has been inaccurately distributed as B. francisci-ferdimndii, but probably comes, as do the other species, from seed received from the northern Chinese provinces.

The following two hybrids, owing to their hairy leaves, must be placed in this section.

(272 a) Berberis x gibbsii Ahrendt, hybr.nov., ramulis foliisque pubescentibus

Stem sulcate; young shoots puberulous; mature yellow, puberulous. Leavm to 30 x 13 mm., subentire or with 1-6marginalspinules; belowgreen, sparsely pilose. Racenaerr 12-200-fld., 2-5-5 cm.,includingpeduncle1-5-2.5cm. Pedi&glabrous,4-6 (-8, below) mm. ; bracts 1-5-2 mm. Inner sepals 4.5 mm. Petale 3 mm. Stuntens not produced, truncate. Ovules 2. Berries black, pruinose, blue, 8 x 5 111111. Style absent or not distinct. Seeds brown.

Cultivated (Kew 7-951-31, Gibbs): fl. 23 May 1940; fr. 18 Oct. 1939 (at K) (Type, 0). An interesting hybrid, presumably between a species of this section, and a black-fruited one.

fructibus atrocaeruleis

(272 b) Berberis x bombycina Ahrendt, hybr.nov., ramulis glabris, foliis pubeacenti- bus, petalis sepalis interioribus longionbus, fructibus rubris

Stems glabrous, sulcate, yellow, verruculose. Internodes 2-36 cm. S p i w weak, 10-1 5 mm., 1-3-fld. Leaves 2.54.5 x 0.7-1.3 cm. ; elliptic, subacute; attenuate to de- current petiole 4-10 mm. ; margins 20-40-spinose (0.5-1 mm.)-subsemulate, distant 0-5-1 mm. ; reticulate; below paler green sublustrous, sparsely pilose. Racemes 7-14-fld., 1.543 cm. Pedicels glabrous, 3-5 mm. ; bracts much shorter, ovate, acuminate. Prophylk ovate, subacuminate, 2.3 x 0.9 mm. Outer sepals 24'5-3 x 1.5 mm., oblong. Inner sepal8 5 x 3.5 mm., obovate. Petals 5 x 2 mm., oblong, subacute to obtuse, entire; scarcely clawed, with separate, submarginal, ovate, oblong, or obovate glands 1 x 0.7 mm. Shrv~ns 2-75 mm. Ovules 2, shortly stipitate. Berries bright red, epruinose, ellipsoid, 9-10 x 4.5-5 mm. Style absent or indistinct.

Cultivated (Wisley 59/36): fl., fr. 1938, a t Wisley (Type, 0). This plant was o r i ea l ly labelled B. h t e i , a plant of which may be one of the parents, the other belonging to Sect. Brachypodae.

KEY 49. to Sects. Polyanthae and Sherriffianae (Maps 4 and 38) OVulm3 1-2.

Inflorescence paniculate. A Inflorescence not paniculate B

Inflorescence paniculate. C Ovules 3-5 (-7, in two hybrids).

Inflorescence not paniculate. KEY 53 A. Style 1-1.5mm. 273. 23. polyantha

Style absent or to 0.5 (-0.75) mm. Leaves green, epruinose below, scarcely or openly reticulate.

Leaves entire; fruit stylose; pedicels 4-7 mm. ; bracts x pedicel. Ovules 2-3, subsessile. 279. B. xanthophlaea

Page 200: Berberis and Mahonia : A taxonomic revision

200 L. W. A. AH~ICNDT

KEY 49 (continu&) Ovule 1, stipitate. 280. B. 8oht$ora

Stems yellow, glabrous. 286. B . e d g e d b m stems red, puberulous. 287. 33. kunawurensis

Leaves spmose; f i t estylose; pediceh 1-4 mm.; bracts &l xpedicel.

Leeves grey, pruinose below; closely reticulate. stems puberulous.

Pediceh 1-4 mm. ; bracts (1-3 mm.), +2 x pedicel. Stems red; fruit Mtyl-; bracts 1-2 x peaid. 296. B. tuylorii Stems yellow; fruit stylose; bracts +l x pedicel.

Panicles (6) 9-15 (-25) om.; fruit ovoid, pink. 274. B. pdtii Panicles 1-2.5 om. ; fruit globose, red. 275. B.a$rregatcc

Pedicels 5-10 mm.; bracts 1 mm. 276. B. rockii

Pedicels 2-5 mm.; panicles 10-24-fld.; leaves 1 : 2.54 . Stems glabrous.

Fruit estylose, black, pruinose blue. 298. B. sherrisi Fruit stylose, red, epruinose.

Leavea narrow, 1 : 4, always entire. 278. B. oblanceobta Leaves broader, 1 : 2-5-3, sometimes spinom. 277. B. brdpnkuhta

Pedicels 5-18 mm.; panicles 3040-fld.; leaves narrower, 1 : (3-) 4-7. 281 b. B. x leachiana

B. Leaves green, epruinose below; stems glabrous, dark red; fruit 8-10 mm., estylose. Leaves entire; infiorescence 1-5-fld.; pedicels 7-16 mm. 291. B. hmkyana Leaves with 3-12-marginal spinules; inflorescence 5-12-fld.; pedicels 4-7 mm.

Cf. 287. B. kzmawu*eneie Leaves grey, pruinose below; sterne pale yellow; fruit P 5 mm., stylose. Stems puberulous ; leaves spinose ; pedicels 1-2 mm. ; bracts subequal.

Stems glabrous; lesves entire; pedicels 4-9 mm.; bracts much shorter. 289. B. arido-calida;

290. B.he&ropia C . Ovules 3-5.

Leaves below green, epruinose; bracts longer than pedicels.

Leaves below grey, pruinose; bracts subequal to or shorter than pedicels. 288. B. grandibracteata

F h i t pruino~, eatylose. Stems pale yellow; leaves narrow, 1 : 3 4 ; pedicels 6-15 mm.; bracta 1 mm.,

Stems dark red; leavee broad, 1 : 1.6-2 (-2-5); pedicels ( 1 . 6 ) 2-4 (-5) mm.; much shorter; fruit red, pruinose blue. 281. B. b e a n h a

bmta longer, or subequal; fruit black, pruinose blue. 297. B. syalalbica

Panidea 6 1 7 cm. ; stems subterete. 285. B. k d w w a m Pmkles 1-5 cm.; stems very sulate.

Fruit epruinoae, stylose.

Sterne dark red, puberulous; bracts subequal to pedicels.

Stems glabrous, dark red, or pale yeUow; bracta much shorter than 282. 33. metapolyantha

pediceh. 283. B. x m m i w ovulee 5-7. 284. Hybrids

Sect. Polyanthae Schneid., BuU. Herb. M & r , (2), 5, 814, 1905 Stems mostly sulcate. Internodes 1.5-3-5 cm. Spines (1-) 3-fld., (1-) 16-26 (-3) em.

Leaves mostly deciduous, rarely semi-evergreen; margins entire or with 3-11 (-25) spinules; nearly always closely reticulate, and mostly pruinose-grey below. InJibrtmew (2) 12-100-fld., panicdate (except in the Subsect. Pseudopolyanthae where the in- florescence is much reduced and fascicled or subfaacicled in form, and only 6-12 mm.

bracts generally shorter, but sometimes longer. Flowers M m m . diameter. Petals IOW), e l ~ ~ t h d , 2-10 (-26) ~ m . low. Pedicels 1-7 IIM. (to 15 111111., only in B. beank~);

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Berberis and Mahonia 201

nearly always emarginate, and shorter than inner sepals. Stamens mostly not, or scarcely, produced and obtuse or truncate. Ovules 1.5. Berries red, generally epruinose, often soft, 6-8 (-12) mm., mostly with short style 0%-0.5 (-0.75) mm.

Hazara, Kaahmir, Punjab, Jaunsar, Garhwal, Kumaon, Nepal, Bhutan, Burma, Yunnan, Kweichou, Szechuan, Hupeh, Kansu.

Subsect. Eupolyanthae Ahrendt, Kew BUZZ. 1939,273, 1939 Stem angled or sulcate; mature yellow. h u e s deciduous (except in B. polyantha);

margins entire, or with 3-8 (-25)-spinules. Inflmescence paniculate, 15-40 (-100)-fld., (2) 6-25 cm. long. Pedicd.9 1-5 (-10) mm. Flowers 5-9 mm. diameter. Ovules (1-) 2 (-3:). Berries shortly stylose (except only in B. rockii).

Iiansu, W. Szechuan, Hupeh, Yunnan, S.E. Tibet (Charme district).

KEY 50, to Subsect. Eupolyanthae (Map 35) Style 1-1.6 mm. ; berries narrow, dark red. Style short, 0.2-0.5 (-0.75) mm., or absent; berries brighter red.

273. B. polyantha

Leaves with spinose margin; below pruinose grey. Stems puberulous ; leaves broad (1 : 2) ; stamens produced.

Style distinct; pedicels stout, 1-4mm., shorter than bracts, or not more than

Panicles narrow, (6-) 9-25 cm. long; pedicels 2.5-4 mm., with shorter (1-2 mm.)

Panicles broad, 1-2.5 cm. long; pedicels 1-2 mm., with longer (1.5-2.5 mm.)

Style absent ; pedicels slender, 5-10 mm., 5-7 times as long as the bracts ; stamens

twice as long; stamens obtuse.

bracts; petals shorter than inner sepals. 274. B.pra&?ii

bracts; petals as long as inner sepals. 275. B. aggregata

apiculate. 276. B. rockii

277. B. brevipaniculata Stems glabrous; leaves narrower (1 : 2 . 5 3 ) ; stamens not produced.

Leaves with entire margin. Leaves below pruinose, grey. Stems glabrous.

Leaves narrow (1 : 2.5-3), sometimes spinose. 277. B. brevipaniculata Leaves narrower, 1 : 4, always entire. 278. B. oblanceolata

Stems puberulous. 275 b. B. agg~egata var. integrifolia

Panicles P 9 cm. ; spines 3-fld. ; ovules 2-3, subsegsile; stamens not produced.

Panicles 3-4 cm. ; spines solitary; ovules solitary, stipitate; stamens produced.

Leaves below epruinose, green.

279. B. xanthqvlduea

280. B. aolutiflora

(273) Berberis polyantha Hemsley, J. Linn. SOC. (Bot.), 29, 302, 1892; Schneid. in Sargent, PI. Wik . 1, 376, 1913; Ahrendt, J. R. Hort. SOC. 79,190, fig. 3, Apr. 1954

Shrub 10-14 ft. Stem glabrous or subglabrous; mature yellow-brown, sulcate. Inter- d s 1.5-3 cm. Spines mostly solitary, concolorous, 1-3 cm. Leaves persistent or semi- evergreen, 1.2-4.5 x 0.5-1-7 cm. ; oblong-obovate; base contracted to decurrent petiole, 2-7 mm.; apex rounded; margins 5-11-spinose (1 mm.)-serrate (1 mm.), distant 2.5- 5 mm. ; closely reticulate both sides; above sublustrous dark green; below grey, pruinose, papillose. Panides (30-) 50-100-fld., broad and much branched, 6-12 cm. long, 3-5 mm. broad. Pedicels 2 4 mm.; bracts 1-5-2.5 mm. Prophylk ovate, acute, 3 x 1.25 mm. Outer sepals 3 - 5 4 x 1.75-2 mm. ovate, acute. Inner sepals 5-6.5 x 2-5-3 mm., obovate. Petals 3.5-4 x 2 mm., obovate; apex incised with acute lobes; base clawed, with oblong glands, 0.7 x 0.5 mm. Sbmn-s 3 mm., produced, obtuse. O w l a 2, subsessile. Berries dark red or dark red-brown, very narrowly ovoid, 7-8 x 3 4 mm., excluding a conspicuous style, 1-14 mm. long. Seeds p q l e , 5.5 x 2.5 mm.

Page 202: Berberis and Mahonia : A taxonomic revision

202 L. W. A. AHRENDT W. Szechuan: near Tachien lu, 9000-13,000ft., fl. , fr. Pratt 206, 704 ( T y p , K).

Tachien lu 8500-10,700ft., July, Sept. 1908, Wilson 1048 (K); Tachien lu, Kangtin Ksien, a t road side, 8500-9500 ft., Fang 3519 (K).

Cultivated: from Wilson seed, at Maidwell Hall, Northampton, fl. 14 May 1948; fr. 20 Sept. 1947. This is the only cultivated plant of the species in England which I know still to be alive. There waa one other at Kew (no. 352), of which there are dried specimens in the herbarium there. Attempts to root cuttings all f d e d in 1948, 1949 and 1950. B. pJyantha appears to be a species (or possibly a natural hybrid) with a very small region for its habitat, where, possibly, it fails to propagate itmlf, as it defies attempts a t propagation in this country.

(274) Berberis prattii Schneid., Sargent, PI. Wils. 1, 376, 1913; syn. B. aggregatu var. pdii Schneid., 1.0. 3, 443, 1917

Shrub fairly erect, 6-10 ft. Stem with young shoots puberulous and slightly reddish; mature pale yellow, subpubemlous or subglabrous, finely verruculose, sulate. I n t e d 1 - 6 3 om. Spines terete, 13-fld., 5-10 (-15) mm. Learn deciduous, 1-3 (4) x 0-5-1.4 (-1.8) cm., including petiole 1-3 mm.; base contracted; apex rounded; mmgins 3-8- spinose (1-1.5 mm.)-dentate (1-2 mm.), distant 5-10 mm. ; slightly lustrous fairly light yellowish green above; below grey, p ~ i n o ~ e , finely closely papillose; both sides closely reticulate. Pedicels 2 . 5 4 mm.; bracts 1-1-5 (-2-5) mm. F h r a H.C.C. 3. Prophylls absent. Outer 8epa.h lanceolate, subacuminate, 4 4 x 1 - 5 2 mm. Median se@ 4-5 x 1.5-2 mm., ovate-lanceolate, subacuminate. Inner ae@ 5.5 x 4 mm., obovate. Petals 4.5 x 2.25 mm., narrowly obovate; acutely emarginate with acute lobes; scarcely clawed, with separate oblong non-marginal glands, 1 x 0-5 mm. Stamna 2.5 mm., produced, obtuse. Ovules 2, sessile. Berries firm, ovoid, bright pink, 6 x 4.5 mm. Style 0.3-0-5 mm.

KEY 5 0 ~ Leaves with spinose margins.

Paniclea erect, with VerticiUate paniculate portions brsnching at intervals from the rachis; the inflomcence being stiff and 15-80-fld., and (6) 10-15 (-26) om. long.

Pedicels of fruit straight. 274a. B. pmttii var. pmii Pedicels of fruit recurved. 274b. B. pmttii var. recurvata

274c. B. prdtii var. Zuxipenddu Leaves with entire margins. 274d. B. x 'Stonefield Glow'

Paniclea subpendulous, more open, 4-6 (-10) cm. long.

( 2 7 4 ~ ) Berberis prattii var. prattii

Wilson 1261 (Type, K); Wilson 1300; Pratt 80 (K).

(Veitch) 3153, fl. 18 July 1939. A magnificent plant foE cultivation.

W. Szechuan: Tachien lu, thickets to south-eat, 8400-9000 ft., fl. June; fr. Oct. 1908,

Cultivated: L.A. 93, fl. 3 June 1942,l July 1943; fr. 28 Oct. 1944; also at Kew, Wilson

(274b) Berberis prattii var. recurvata Schneid., Sargent, PZ. Wikr. 1, 377, 1913; syn. B. aggregata vm. recurzlata Schneid., 1.c. 3,443, 1917

W. Szechuan: Mupin, thickets, 6500-8000ft., fl. June; fr. Oct. 1908, Wilson 1073

Cultivated: L.A. 194, fl. 24 June 1942; fr. 12 Sept. 1940. For cultural purposes scarcely (TYP, K).

distinguishable from the previous variety.

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Berberis and Mahonia 203

(274~) Berberis prattii var. laxipendula Ahrendt, J. R. Hort. SOC. 79, 192, Mar. 1954

This differs from the preceding two varieties in its subpendulous panicles, which are shorter, 4-6 (-10) cm. long, and proportionately broader, and of a looser nature, with somewhat larger berries, 6-7 x 4-46 mm., and slightly longer style (0.7 mm.).

W. Szechuan: no locality, Wilson (Veitch) 3152. Cultivated: from the seed cited &s collected by Wilson, a t Kew (nos. 351437/05)

(Type, K). A photograph appears in Gard. Chron. 4 Nov. 1942 (inaccurately labelled B. polyantha). In the autumn of 1953, this variety was shown a t the R.H.S., by Mr E. H. M. Cox, Glendoick, Perthshire. Compare Bot. Mag. t. 8549, Apr. 1914.

(274d) Berberis x ‘Stonefield Glow’, Ahrendt, J. R. Hort. SOC. 67, 131, Apr. 1942 Outer sepals ovate, acute, 3 x 2 mm. Median sepals 5 x 3 mm., obovate, acute. Petals

4.5 x 3 mm., incised, acutely emarginate; glands 0.9 x 0.6 mm. Stamens not produced, truncate. Berries soft, red, depressed-globose, 7-8 x 7-8 mm. Style 0.25-0-5 mm.

Cultivated: origin L.A. 92, 0. 31 July 1939; fr. Nov. 1939. Most likely B. prattiix B. subcaul&tu. Hybrids which fall under this heading are commonly distributed as B. plyantha or B. prattii.

(275) Berberis aggregata Gchneid., BuU. Herb. Boissier, (2), 8, 203, 1908; Sprague, Bot. Mag. 143 t. 8722, 1917; Ahrendt, J. R. Hort. Soc. 79, 191, 193, 1954

Shrubdensely branched, 3-5 ft. Stemverysulcate ;young shoots puberulous ; mature pale yellow. Intermodes 18-27 mm. Spines 3-fld., slender, 8-15 mm. Leaves 8-25 x 6 1 1 mm.; oblong-obovate ; base contracted, subsessile ; apex rounded ; closely reticulate ; above dull yellow-green; below grey, pruinose. Panicles dense, sessile, 10-30-fld., 1-26 cm. long. Pedicels 1-2 (-2.5) mm. ; bracts slightly longer, 1-5-2.5 mm. ProphyUs ovate, acute, 1 x 0.6 mm. Outer sepah 2.5 x 1.75 mm. Inner sepah 3.5 x 2.5 mm., all elliptic. Petals 3.5 x 2 mm., obovate; apex narrowly emarginate; base clawed, with oblong glands, 0.7 x 0.5 mm. Stamens 2-25 mm., produced, obtuse. ovules 2, subsessile. Berries aoft, eubglobose, pale red, 6-7 mm. Style 0.5 mm.

KEY 5 0 ~ Shins pale yellow. Bracts slightly longer than pedicels.

Leaves spinose. Leaves entire.

Bracts much shorter than pedicels. Shms red-brown.

275a. B. aggregakz vm. aggregata 275b. B. aggregata var. integrifolia

275c. B. x ‘Stonefield Dawn’ 275d. B. x ‘Ruby Watson’

(27Sa) Berberis aggregata var. aggregata

aa broad as long. Leaves with margins 3-8-spinose (1-16 mm.)-sedate , distant 2-3 mm. Panicles

Kansu (E): Potanin, s.n. (Type). Szechuan (N.W.): Aug. 1903, Wilson (Veitch) 3155 (K). Cultivated: L.A. 24, fl. June 1937; fr. Nov. 1937.

(275 b) Berberis aggregata var. integrifolla Ahrendt, var.nov., foliis integris Leaves to 22-30 x 7-12 mm. Panicks narrow, 2-4 cm. Pedicels puberulous in flower,

glabrous in fruit, 1-2 (-2.5) mm.; brads 1-5-2.5 (-3) mm. Berries 7 x 4-5 mm. Styk 0.4 mm.

Page 204: Berberis and Mahonia : A taxonomic revision

204 L. W. A. AERENDT Szechuan: (W.) Pao-hsing-hsien, 6500 ft., fr. 17 Sept. 1936, 8000 ft., Chu 3863, and

fl. 25 June 1936, Chu 2939 (Type, E); Sept. 1904, Wilson (Veitch) 3155a (K). 'Shrub 6 ft.'

Yunnrtn: Yunnan-sen, fl. Maire 1997 (K). Not cultivated.

(275c) Berberis x 'Stonefield Dawn' Ahrendt, J. R. Hort. Sw. 67,131, Apr. 1942 Leaves pale yellowish green. Berries globose, 4-5 mm., H.C.C. 21. Cultivsted: fl. May 1940; fr. Dec. 1940.

(275d) Berberis x 'Ruby Watson', Ahrendt, h., 132, 1942

H.C.C. 2211.

(276) Berberis rockii Ahrendt, sp.nov., ram& puberulis, foliis marghibus spinoso- serratis, pedicellis longioribus, !%lorn., gradimis cum bracteis minutis; in- floreclcentiis panicdath, fructibus estylosis, distinguenda

Stems eulcate; young shoots puberulous; mature pale yellow brown, pilose-vermculose. I&& 2.54 cm. Spines weak, 13-fld., 2-7 mm. Leaves deciduous, to 3.7 x 1.8 cm.; obovate-elliptic; base contracted to decurrent petiole, 1-2 mm.; apex rounded; margins 6-20-spinose (1 mm.)-serrate (0-5-1 mm.), distant 2-3 mm.; closely reticulate; above slightly lustrous bright yellow-green; below grey, pruinose, not, or scarcely, papillose. Panicles loose, 25-40-fld., 6-9 cm. long, including peduncle 1-3 cm., with an ocwional extra b a d flower with long pedicel 1-2 cm. Pedicek very slender, puberulous, 5-10 mm. ; bracts 1 mm. (or at base of subpedunclea to 1-5 mm.). E"l0Wers H.C.C. 3. ProphyuS red, 1-2 x 0-5-0-8 mm. Outer sepals 2-5-3 x 1-1.25 mm., oblong-ovate, acute. Inner s e p h P 5 x 3-4 mm., obovate-spathulate. Petals 4-25-4-5 x 2-2-25 mm., elliptic; apex acute, incised; base slightly clawed with elliptic, separate, submarginal glands, 0.8 x 0.5 mm. Stamens 2.5-2-75mm., produced, shortly apiculate. O& l (-2). Bewies dull red, oblong, estylose, 8 x 3-54 mm. Seed-s pale yellow, 5-5-5 x 2 mm.

Stems dark red-brown, puberulous. Lecves persistent. Berries 5.5-6.5 x 4.5-5-5 mm.,

Yunnan: Moting shan, 13,000 ft., Sept. 1932, seed, Rock 23339. Cultivated: at Kew, fr. 13 Oct. 1939; also at Watlington, fl. 5 May 1943 (Type, 0) .

(277) Berberis brevipaniculata Schneid., BUZZ. Herb. Boissier, (Z), 8, 263, 1908 Stems glabrous, sulcate; mature pale, brown to red-brown. Internodes 1.5-2-5 cm.

Spines 1-3-fld., 10-17 mm., below broadly sulcate. Leavee 10-30 x 4-9 mm.; narrowly obovate; baae contracted, subsessile; apex rounded, spathulate or retuse, and mucronate; margins entire, occasionally 1-2-spinulose (0.25 mm.)-subserrulate ; above openly re- ticulate; below grey, pruinose, indistinctly veined. Panicla loose, 14-24-fld., 2-3 cm. Pedicels (2-1 3-4 mm.; bracts 1 mm. Flowers 5-6 mm. diameter. Inner sepals about 4 mm. long. Petals subequal to inner sepals, narrowly ovate, subacute. Stamens not produced. ovules sessile.

Hupeh: Ichang, fl. 1888, Henry 4675 (Type, K). Not cultivated. Hybrids of B. aggre- gata are sometimes distributed under this name.

There fall under this heading in the key (though unconnected with this species), two hybrids which may be distinguished as follows.

KEY 50c Petals and inner sepals subequal; leaves rounded or emarginate at apex.

Leaves indistinctly veined below; margins sometimes spinose. Leaves closely reticulate below ; always entire.

277. B. brevipaniculala 277a. B. x Sibbertoft Coral

277 b. B. x Crimson Bead Petals shorter than inner sew; laves acute at apex.

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Berberis and Mrchonia 205

(277a) Berberis x ‘Sibbertoft Coral’ Stanley; ex Ahrendt, J. R. Hort. Soc. 67, 131, Apr. 1942

Berries ovoid, 6-7 x 56-5 mm., H.C.C. 21.

(2773) Berberis x ‘Crimson Bead’ A. R. Ahrendt ex L. W. A. Ahrendt, J. R. Hort. Soc. 67, 131, Apr. 1942

Berries oblong, 4.5-5 x 3.54 mm., crimson (H.C.C. 22, a darker shade). Named by my mother, this is an ornamental hybrid with its prolific drooping panicles of deep coloured berries.

(278) Berberis oblanceolata (Schneid.) Ahrendt, Kew Bull. 1939, 275, 1939; syn.

.Leaves to 24 x 6 nun., acute, entire, sessile, closely reticulate, pruinose grey below. Panicles narrow, 10-15-fld., 2 - 5 5 cm. long. Pedicels 2-3 (4) mm.; bracts shorter. OV& 2.

W. Szechuan: valley of Hsao-chin-ho, near Monkong-ting, 7500-9750 ft., fl. June 1908, Wilson 2868 (Type, K). ‘Decumbent bush, 3-5ft.; flowers deep yellow.’ Not cnl tivated.

(279) Berberis xanthophlaea Ahrendt, J. Bot., Lond. 79 (Suppl.), 73, 1942 stems angled-sulcate, mature very pale yellow, not verruculose. Inikmodes 1-2 cm.

S p i w concolorous, 3-fld., stout, terete, 1.2-2.3 cm. Leaves 1-2.6 x 0.8-1.3 cm. ; base cuneate, sessile; apex obtuse or rounded; margins entire; both sides with distinct branched venation and indistinct fine close reticulation; below papillose, a t first grey, pruinose, ultimately slightly paler lustrous green. Panicles 15-30-fld., (2 .6 ) 3.5-6 (-9) cm. long, including peduncle 1-3 om., sometimes with a few extra flowers at the baive. Pedicels 4-7 mm. ; bracts 1.5-2 mm. ProphyUs 1-3-16 x 0.9-1.25 mm., ovate, acute. Outer sepals 3-3-5 x 2-3 mm., elliptic. lnnev sepd.3 5-6 x 2.5-3.5 mm., elliptic-obovate. Petakr 3-5-4.5 x 2.5-3 mm., obovate; apex deeply incised; base slightly clawed, with oblong, widely separated, submarginal glands, 0-7-1 x 04-0.5 mm., situated above the claw. Stamens 2.25-3 mm., not produced, truncate. Ovules 2-3, subsessile, or with stipe up to one-sixth the length of the ovule. Berries red, pruinose blue, oblong-obovoid, 7-9 x 5-6 mm. Style 0.5 mm.

S.E. Tibet: Charme district, Chayul Chu, 12,000 ft., 25 Oct. 1938, with one old berry, Ludlow, S h e d t Taylor 6406 (Type, BM); ‘to 5ft., on dry gravel terraces’; Char Chu, 10,500 ft., 1 Nov. 1938, fl. Ludlow, SherriE & Taylor 6651 (BM), ‘shrub 4 4 ft., on dry stony river banks’. Not in cultivation.

This is remarkable for the period of the year when it was collected in flower; it is the westerly species of this subsection, and is a link with the still more westerly Subsect. Su1)polyanthae.

(280) Berberis solutiflora Ahrendt, sp.nov., ramulis glabris, foliis integris subtus

Intemzoh 1.5-2 cm. Spines solitary 1-2 cm. Leaves 10-25 x 4-10 mm., obovate, ses- sile, entire, obtuse ; both sides with openly elevated reticulation ; below epruinose, con- colorous green. Panicles loose, 12-24-fld., 3-44 x 2-3 cm., including peduncle 1-14 cm., and with subpeduncle of the branched portions 7-10mm. long. Pediceb glabrous, 4-6 mm.; bracts 1-1.5 mm. Prophylh lanceolate, acuminate, 1-75 x 0.5 mm. Outer sepal8 2.5 x 1 mm., ovate, acute. Inner se@ 5 x 2-25 mm., oblong-obovate. Petah 3-75 x 2.25 mm., obovate; base shortly clawed, with elliptic, widely separated, marginal glands, 0.6 x 0.4 mm. Stamns 3-25 mm., apex produced, rounded-truncate. O v u h sofih‘y, stipitate, the stalk being half as long aa the ovule. 8tyk short.

B. p l y a n t h var. oblanceoluta Schneid., Sargent, PI. Wils. 1, 376, 1913

viridibus, paniculis laxis, ovulis solitariis stipitatis distinguenda

Page 206: Berberis and Mahonia : A taxonomic revision

206 L. W. A. ~ E N D T

Yunnan Mekong-Yangtle divide, Bei-ma-shan, S.E. of Atuntze, 12,OOO ft., fl. May- June 1932, Rock 22835 (Type, K). ‘Shrub 3 &., in alpine meadows.’ Not in cultivation.

Subsect. Metapolyanthae Ahrendt, J. Bot. Lond. 79 (Suppl.), 75, 1942 Panicles 10-20-fld., 2-5 cm. long. Bracts shorter than pediwls. Ovules (3-) 4-5 (in

Yunnan, Szechuan. hybrids, to 7).

KEY 51, to Subsect. Metapolyanthae (Map 35) ovules 3-5.

Leaves openly reticulate; margins spinose; pedicels 5-18 mm., with bracts very much shorter ; fruit dark red, conspicuously pruinose.

Leaves fairly narrow, 1 : 3-4; panicles 10-20-fld., 2-4cm.; ovules 3-5; fruit 10 x 6 mm., pruinose mauve, estylose. 281. B. beaniana

Leaves narrower, 1 : 4-7; panicles 3MO-fld., 6 6 cm.; ovules (&) 3; fruit 8 x 4 mm., pruinose blue, stylose. 281 b. B. x h?u.chiana

Leaves closely reticulate; margins entire (except in hybrids); pedicels 1-5 mm., with slightly shorter bracts, 1.- mm. ; fruit bright red, epruinose, stylose.

Stems dark red, puberulous. Leaves narrow, 1 : 2.5-3, apex rounded; bracts subequal to pedicels.

Leavea broad, 1 : 1-2, apex retuse; bracb much shorter than pedicels. 282. B. metqmlyantha

282b. B. m y i Stems glabrous. 283. B. x carrnim and CARMINEA hybrids

Ovules 5-7. 284. POLYANTIUE x ANQULOSAE hybrids

(281) Berberis b e a n i a ~ Schneid., Sargent, PI. Wils. 3, 439, 1917; Stapf in Bot. Hug. 144, t. 8781, 1918

Shrub fairly densely branched, compact, 6-8 ft. Stems glabrous; young shoots green; mature pale yellow-grey, angled-sulcate, sparsely finely verruculose. Internodes (1.5) 2-3 (4) cm. Spines 3-fld., 1-2-5 em., plane. Leaves deciduous, 2-5 x 0.6-1.3 cm.; narrowly elliptic to ovate-elliptic; apex acute to subacuminate; base attenuate to decurrent petiole, 1-3 mm.; margins !j-lZ-spinose (1-16 mm.)-serrate (0.5 mm.), distant 2-5 mm.; above scarcely lustrous deep green, openly reticulate; below grey, pruinose, finely papillose. Panidea loose and broad, lO-ZO-fld., 2 4 cm. long. Pedicels (6-) 9-15 mm.; bracts 1.5 mm. Flowers H.C.C. 2/1-3/1. Outer sepal8 2-3 x 1-1-25 mm., oblong-ovate, yellow without discoloration. Median sepal8 4-5 x 2.25-3.25 mm., oblong-ovate, sub- obtuse. Inner sepal8 6-66 x 3.544 mm., obovate. Petals 5 x 3-3-25 mm.; apex sub- entire or minutely emarginate; base cuneate with oblong, separate, non-marginal, sub- concolorous, glands, 1-14 x 0.5-0.6 mm. Stamens 3-34 mm., slightly produced, rounded. Ovules 3 4 , sessile. Berries ovoid, estylose, 9-10 x 5-6 mm., dark red, heavily pruinose mauve.

W. Szechuan: 1904, Wilson (Veitch) 1930, seed only. Cultivated: from above seed (Type plant at Kew); LA. 26, derived from cuttings of

the type plant; fl. 24 June 1942, 20 June 1945; fr. 12 Nov. 1940. The leaves persist long before falling.

A hybrid may be distinguished aa follows.

KEY 51a Leaves with mar- 5-12-sphose; inner sepals 6-6-5 mm.; berries 10 x 6 mm., quite

Leaves with margins 2-4-spinose; inner sepals 4-5 mm.; berries 8 x 6 mm., sometimes estylose. 281. B. beaniana

with extremely short style, not exceeding 0-25 mm., otherwise estylose. 281 a. B. x Stonefield Mauve

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Berberis and Mahonia 207

(281a) Berberis x ‘Stonefield Mauve’ Ahrendt, J . R. Hort. Soc. 67, 134, Apr. 1942

deciduous. The leaves are broader than in B. beaniana, of which it is a seedling, and quickly

Cultivated: fl. 30 June 1938; fl. 1 Nov. 1938.

(281 b ) Berberis x leachiana Ahrendt, J . Bot., Lond. 80 (Suppl.), 81, 1942 Stem terete, verruculose. Spines solitary, slender, 10-18 mm. k w e s 30-60 x 4-

9 mm., or to 40 x 13 mm. ; narrowly elliptic, acute, mucronate; petiole 3-5 mm.; margins 4-10-spinose (1 mm.)-serrulate, distant 2-4 mm. ; slightly lustrous deep green; below openly veined, pruinose, grey. Panieles broad, 4 4 x 3 4 cm. Pediceh glabrous, 4-18 mm. ; bracts 1.5-2.5 mm. Prophyth 2 x 1.5 mm., yellow, subacute. Outer sepals 4 x 3 mm., elliptic. Inner sepah 4.5 x 3.75 mm., obovate. Petals 4.25 x 3 mm., obovate, entire; cuneate with separate, ovate, acute glands, 1 x 0-6 mm. Stamens 3.25 mm., not pro- duced. Berries 7-8 x 4 mm., red, pruinose blue. Styk 0.25 mm.

Cultivated: at Wisley, 0.29 June 1939; fr. 17 Nov. 1939. This may also be a hybrid of B. beanianu.

(282) Berberis metapolyantha Ahrendt, J . Bot., Lond. 79 (Suppl.), 75, 1942 Stems sulcate. S p i m 3-fld., 1-1.5 cm. Leaves semi-evergreen to 25 x 10 mm., obovate,

cuneate, sessile, entire, rounded, closely reticulate, dull green. Panicles 10-14-fld., 3-5 cm., reduced to a raceme, compound below, a t apex of stem. Pedicels puberulous, 3-5 mm. ; bracts 2-3 mm. Outer sepals 2 x 1 mm., oblong-obovate, rounded. Inner sepals 4 x 2.25 mm., obovate. Petals 3 x 1-5 mm., obovate, oblong-elliptic. Stamens 2.5 mm., slightly produced, shortly apiculate. Berries oblong-globose, 6.5 x 6 mm. Styk 0.5- 0.75 mm. W. Szechuan: seed, Forrest 27773. Cultivated: a t Kew, from above seed, fl. June 1939; fr. 25 Oct. 1939 (Type, 0). Some

of the seed from this number has produced plants with a more reduced inflorescence. B. awyi, too, includes plants with a variable inflorescence. Accordingly, i t seems possible that both B. metapolyantha and B. coryi are natural hybrids between B. aggregata and B. u4lsonue.

KEY 5 1 ~ Leaves narrow, 1 : 2-5-3.5.

Leaves 1 : 2.5; bracts 2-3 mm., subequal to pdicels; style 0-5-0-75 mm.

Leaves 1 : 3.5; bracts 1 mm., much shorter than pedicels; style 0-25 mm. 282. B. metapolyantha

282a. B. x colburyensis Leaves broad, 1 : 1-2. 282b. B. coryi

(282 a) Berberis x colburyensis Marchant, Catabgm of Trees and Shrubs, 21, 1937, angli ce

Cultivated: in the nursery of the late W. J. Marchant, Wimborne, Dorset.

(282,3) Berberis coryi Veitch, G a d Chron. (3), 52, 321, 1912; New and Hardy Plants, W . China, 7, 1913

Leaves semi-evergreen, 10-25 x 8-12 mm. ; spathulate-cuneate; entire apex; retuse. Panicles 3-5 cm., sometimes reduced to short racemes or subfascicles, 7-20-fld. Pedieels 4-5 mm. Berries oblong-globose, 5-6 x 4-5-6 mm. Style 0.5 mm.

Yiinnan: rocks, cliffs, dry rocky slopes on the N’Marka-Salween divide, 9ooO-10,OOO ft., 1917-19, Forrest 18516 (Type, K). ‘Shrub 1-1) ft., flowers orange; fruit purplish red.’

Page 208: Berberis and Mahonia : A taxonomic revision

208 L. W. A. AJXRENDT Cultivated: by Veitch originally; also L.A. 521; fr. Dec. 194-4; fl. 18 July 1945,3 July

1952. The berries are pinkish red; possibly Forrest’s remark refers to an old discoloured h i t on his flowering stem. As has been remarked above this is poasibly a hybrid be- tween B. aggregata with panicles of flowep and B. wilsonae with semi-evergreen leaves and fascicled flowers. On account of its showing sometimes a purely fascicled inflores- cence, it has often been referred to as B. subcauj- PEW. coryo‘, which would have to be corrected to B. wilsonae var. wryi, since B. .subcad- is now regaxded as a variety within B. wileonae, and in any caae on account of the pubescent stems. In view of its character, it seems desirable to leave it as originally named.

(283) Berberis x carminea Chittenden ex Ahrendt, J. R. H o d SOC. 67,132, Apr. 1942 Leaves 15-35 x 4-8 mm. ; margins 2-3-spinose, distant 2-5-4 mm. ; reticulate; grey-

green; below pruinose, whitish grey. Panicles 10-16-fld., including peduncle 7-10 mm., or sometimes 3-5 cm. long, with a peduncle 14-26 cm. Pedkls slender, 2-5 mm.; bracts 1.25 mm. Outer se& yellow, 3 x 2 mm., ovate, acute. Median sepds 5 x 3 mm., oblong-ovate. Inner se& 6.5 x 4 mm., ovate, rounded. Petals 4-25 x 1.75 mm., elliptic; apex retuse with acute lobes; glands suborbicular, 0.6 x 0.5 mm. Stamens 3 mm., scarcely produced, truncate. Berries H.C.C. 21, ovoid, 8-9 x 6-5-7 mm. Style 0.5 mm.

Cultivated: at Wisley, fl. 12 July 1938; fr. 16 Nov. 1938 (Type, BM).

KEY 510 Carminea hybrids, Stem glabrous. InJlorescence panicdate. Ovules 3-5.

Panicles with peduncle 1-3 cm. ; leaves deciduous. stems pde, yellow. Leavas ~pin~ee; flower diameter 9-11 mm.; fruit 8-9 mm. Leaves entire ; flower diameter 7-9 mm. ; fruit 9-1 1 mm.

Stems dark red; leaves entire. Leaves broad, 1 : 2.5. Leaves narrow, 1 : 3-54.

Fruit 6.5 mm., carmine. Fruit 8.5 mm., vermilion.

Panicles sessile or subsessile. Leaves deciduous.

Stems yellow; fruit 9 x 9 mm. Stems red; fruit 6 x 6 mm.

Leaves spinose; flower diameter 7-9 mm.; fruit 7 mm. Leaves entire; flower diameter 10-12 mm.; fruit 8.5 mm.

Leaves semi-evergreen.

283. Canninea 283a. Sparkler

283 6. Autumn Cheer

283c. Fireflame 283d. Aurora

283e. Buccaneer 283f. PirateKing

283g. Barbaroma 283L Bountiful

(283a) Berberis x ‘Sparkler’ Chittenden ex Ahrendt, J. R. Hort. Soc. 67,132, Apr. 1942 h u e s broad, 1 : 2.5. Berries ovoid-subconical, mandarin red, H.C.C. 17/1. Cultivated: a t Wisley, fl. 12 July 1938; fr. 16 Nov. 1938.

(2833) Berberis x‘Autumn Cheer’ Chittenden ex Ahrendt, I.c., 1942 Berries oblong-ovoid, 6-7 x 5-6 mm., H.C.C. 19. Cultivated: a t Wisley, fl. 12 July 1938; fr. 16 Nov. 1938.

(2836) Berberis x ‘Fireflame’ Chittenden ex Ahrendt, l.c., 1942 h v e a nmow, 1 : 3-5. B e r h ovoid, 6-5 x 5-5 mm., H.C.C. 21. Cultivated: at Wisley.

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Berberis and Mahonh 209

(283d) Berberis x ‘Aurora’ Chittenden ex Ahrendt, I.c., 1942 Berria globose-ovoid, 8.5 x 8 mm., H.C.C. 18. Cultivated: a t Wisley.

(283e) Berberis x ‘Buccaneer’ ex Watson; Ahrendt, l.c., 1942 Berries globose, 9 mm., almost estylose, partly whitish, ultimately Geranium Lake,

H.C.C. 20, possibly showing influence of B. janaeSaana as well aa of B. aggregatu and B. wihnae var. subcaulialata.

Cultivated : Watson’s nursery, Killiney.

(283f) Berberis x ‘Pirate King’ Waterer ex Ahrendt, l.c,, 1942 Bem‘es H.C.C. 19/1. Cultivated: Waterer’s nursery, Bagshot, Surrey, fl. 4 May 1937; b. 9 Nov. 1937.

(283g) Berberis x ‘Barbarossa’ Watson ex Ahrendt, l.c., 1942 Berries ovoid-globose, H.C.C. 19. Cultivated: Watson’s nursery, Killiney.

(283h) Berberis x ‘Bountiful’ ex A. R. Ahrendt ex Ahrendt, l.c., 1942 Berries subglobose, H.C.C. 19. Cultivated: Stonefield, Watlington, Oxfordshire, fl. 23 July 1938; fr. 11 Oct. 1938.

Polyanthae x Angulosae hybrids. Sterns glabrous. Inflbrescence paniculate. Ovules 5-7. Berries H.C.C. 19.

KEY 511, Leaves deciduoue, broad, 1 : 2-2.5; apex rounded; flowers 8-9 mm. diameter; fruit

Leavea semi-evergreen, narrow, 1 : 4.5-6; apex subacute; flowers 10-12 mm. diameter; ovoid-conical, 9 x 8 mm. 284. Autumn Beauty

fruit ovoid-globose, 7 mm. 284 a. Knockvale Scarlet

(284) Berberis x ‘Autumn Beauty’ Chittenden ex Ahrendt, l.c., 134, 1942 Cultivated: a t Wisley, fl. 12 July 1938; fr. 16 Nov. 1938.

(284a) Berberis x ‘Knockvale Scarlet’ Mulligan ex Ahrendt, l.c., 1942 Cultivated : Knockvale House, Belfast.

Subsect. Subpolyanthae Ahrendt, Kew Bull. 1939, 274, 1939 Leaves deciduous. Injlorescence paniculate, 10-40-fld., (1 .6) 2-43 (-17) cm. long. Hazara, Kashmir, Punjab, Jaunsar, Tehri, Garhwal, Nepal, Bhutan.

KEY 52, to Subsect. Subpolyanthae (Map 36) Leaves below pruinose, grey; panicles (5-) 8-17 cm.; pedicels 4-15 mm., (2-) 3-10 times

as long ae their brects; ovules 3-4. 285. B. k o e h w n a Leaves below epruinose, and more or less concolorous green; panicles 1-5-4 cm.;

pedicels 1-2 (-4) mm., shorter than, or at most 1-2 times as long as, their bracts. 14 JOTJRN. LINN. SOC. BOTANY-VOL. L M

Page 210: Berberis and Mahonia : A taxonomic revision

210 L. W. A. AERENDT

=Y 52 (continued)

Leaves with spinose margin; ovules 2. Stems glabrous, yellow, subterete; leaves mostly broad, 1 : 2-2-6; bracts subequal

to, or longer than, pedioels; petals shorter than inner sepals. 286. B. edgeumthiaraa

287. B. kunawrenais h v e a entire; ovules 3-6. 288. B.grandi6racteata

Stems puberulous, red-brown, sulcate; leaves narrow, 1 : 3-4; bracts shorter than pedicsls; petals subequal to inner sepals.

(285) Berberis koehneam Schneid., Bd. Herb. Boissier, (2), 5, 814, 1905. (Fig. 48) Shrub &loft. Stems glabrous; mature subterete or subangled. Internodes 1.5-

2.5 cm. Spinerr yellow-brown, terete, 1-3-fld., 1-2 cm. Leaves deciduous, 1 - 5 4 x 0.7- 1.6 cm.; oblong-obovate; base cuneate, subsessile; apex rounded; margins mostly 2-4- spinose (0.5 mm.)-serrate (1-5 mm.), distant 4-7 mm., sometimes subentire; both sides with distinct branched venation; above bright green, scarcely reticulate; below grey, pruinose, papillose, not reticulate. Pan* loose, ( 1 6 ) 20-40 (-60)-fld., (6-) 8-17 cm. long, including peduncle (1-) 2-3 (-5) cm., and with subpeduncles of the branched portions, 1-2 cm. Pedi& (2-) 4-7 (-15) mm.; bracts 1.5 mm. Outer sep& 2-5-3.5 x 16-2-25 mm., ovate, acute. Median sepals 4 - 5 3 x 3 . 5 4 mm., oblong-ovate, acute. Inner sepuk 6.5-7.5 x 4-5 mm., obovate. Petah 5-6 x 3-4 mm. ; apex rounded, incised; base cuneate, with obovate, separate, non-marginal glands, 0-8-1 x 0.5-0.6 mm. Stuwn.s 36-3.75 mm., not produced, truncate. ovules 3-4, shortly stipitate. Bewies bright red epruinose, oblong, 8 x 4 mm. Style absent or very short.

I ~ Y 52a Mature stems red-brown to dark red. Mature stems yellow.

285a. B. lwehmna var. koehneana 2853. B. koehneana; var. aurallaea

( 2 8 5 ~ ) Berberis koehneana var. koehneana Kumaon: Duthie 5309 (Type) ; Almora, Budhi, Byans, 9500 ft., 1923, Parker 2074. Nepal (W.): "jnkar, 1923, Parker 2083 (all, K). (W.): Burchula Lekh, near Jumla,

9oooft., open places in mixed forest, ahrub 6-8ft., 10 July 1952, Polunin, Sykes & Williams 4514 (BM); Kunrigaon, south-east of Jumla, 900 ft., shrub, 5-6 ft., at edge of mixed forest on steep slopes, 22 July 1952, Polunin, Sykes & Williams 4862 (BM); Rara Daha, 9800 ft., 13 Aug. 1952, Polunin, Sykes & Williams 5151 (BM) ; Dozam, near Simikot, 8500 ft., shrub 6-8 ft., 4 June 1952, Polunin, Sykes & Williams 4272 (BM).

Cultivated: at Wisley, fl. 18 July 1939; fr. 11 Nov. 1938, and at Kew, fr. 2 Nov. 1939. A magnificent fruiting shrub, which could to advantage be aa widely used as B. prattii in gardens.

(285b) Berberis koehneana var. auramea Ahrendt, var.nov., ramulis aureis S t m yellow, hrete, glabrous, scarcely verruculose. Internodes 1.5-3 cm. Spines

2-10 mm., 1-3-fld. Leaves to 35 x 13 mm., or to 45 x 12 mm., obovate; margins 2 4 - spinose (1-5 mm.)-serrate (1 mm.) ; venation branched and slightly elevated; below grey, pruinose. Panicles 5-lox 3-4 cm., 20-30-fld., with peduncle nil to 5 mm. Pedicels 4-15 mm.; bracts 1-5 mm. Outer sepah 3.5 x 2-5 mm., ovate, acute. Inner sep& 5.5 x 2.75 mm., oblong-obovate. Petals 4-5 x 3 mm., entire. Stamens scarcely produced, truncate.

Central Nepal: Langtang, 8-10 ft., on stony slopes, 11,500 ft., 22 June 1949, Polunin 506 (Type, BM); Marsiandi, 11,500 ft., stony hillsides, 27 June 1950, Lowndes L 1064 (BM). Not in cultivation.

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Berberis and Mahonia 21 1

(286) Berberis edgeworthiana Schneid., BUU. Herb. Boissier, (Z), 8,263, 1908; Parker, Flora .for the Punjab with Hazara and Delhi, 14, 1918; syn. B. brachybotrys Edgeworth, Trans. Linn. Soc. Lond. 46, non C. Gay, 1845

Shrub 2 4 ft. Stems subangled, pale yellow, black-verruculose. Internodes &20 mm. Spines 1-3-fld., 5-18 mm., plane. Leaves 11-32 x 6-15 mm., obovate-elliptic; base cuneate, subsessile; apex acute to subacute; margins 3-8-spinose (1 mm.)-serrulate (0.5 mm.), distant 2-5 mm.; openly reticulate; both sides epruinose, epapillose, sub- lustrous concolorous green. Panicla 10-25-fld., 1-54 cm., subsessile. Pedicels 1-2 mm. ; bracts subequal. Flouters H.C.C. 211. ProphyUs c. 2 x 1 mm., lanceolate, acute. Outer s e p k 3-4 x 2-2.5 mm., oblong, obtuse. Inner sepals 5-55 x 3.5-4 mm., obovate. Petals H.C.C. 2, elliptic-obovate, 4.5-5 x 3 mm. ; apex deeply and narrowly incised; base slightly clawed, with concolorous, oblong, separate, non-marginal glands, 0.7 x 0.4 mm. Sta7nens 3 mm., produced, obtuse. Ovules 2. Berries bright orange-red, firm, oblong, estylose, 7-8 x 4 4 . 5 mm.

N.W. Himalaya: locality not further specified, fl. 8OOO-11,OOO ft., 1844, Edgeworth 65 (Type, K); fl. May 1898, 10,500 ft., Gamble 26752 (K).

Punjab: Chamba State, Bhandal valley, Chadbiant, 8500 ft., fr., 20 Oct. 1919, Parker, s.n. (K); Simla, 9500 ft., June 1885, Collett 723 (K).

Jrmnsar: 10,OOO ft., Karama peak, fl. May 1892, Gamble 23746 (K); Mundali, 8500 ft., fl. June 1895, Gamble 25542 (K).

Garhwal: 10,OOO ft., Schlich 27 (2) (0). According to Parker, from the Punjab Himalay& (Chamba, Kunawur and Simla hills)

to Garhwal, 8OOO-12,OOO ft. Cultivated: L.A. 215, fl. 24 June 1942; fr. 22 Oct. 1943. The cultivated specimens have

somewhat shorter bracts, as is the case with Parker’s fruiting specimen of 1919, from which they may well be derived.

There are two specimens, in the Kew herbarium, which have narrow leaves as in the following species, B. kunawurensis, but otherwise conform to, and seem to belong to, B. edgeworthiam, namely, Jaunsar: Karama peak, 9OOO ft., fl. May 1892, Gamble 23601 ; and Punjab: Tutra Tach, 10,000 ft., 27 May 1898, Duthie 21083. It seems undesirable to place these as a variety within B. edgeworthiana since, together with the existence of Parker’s specimen with the shorter bracts, they rather suggest the existence of a series of natural hybrids connecting the populations of the formal species. This is the more likely as their habitats overlap.

(287) Berberis kunawurensis Royle, Ill. Bot. H i d . 64, 1834; Parker, 1.c. Shrub 3 4 ft. Stems with young shoots puberulous, red ; mature glabrous, red-brown, to

yellowish red. Internodes 1-2 cm. Spines concolorous, 3-fld., slender, subsulcate, 1-2-5 cm. L e a v u 16-45 x 5-15 mm. ; narrowly obovate; base slenderly cuneate, subsesaile; apex acute to subacuminate; margins 4-7-spinose (0.5-1 mm.)-serrate (0.5-1 mm.), distant 2 4 rum. ; openly veined, scarcely reticulate ; below subconcolorous green, epruinose, epapillose. Panicles lMO-fld., 1-54.5 cm., including peduncle 5-15 mm. ; subpeduncle 5-7 Inm. Pedicek 1.54 mm.; bracts 1-2 mm. Inner sepals and petccls 4 4 . 5 mm., subequal. Ovules 2. Berries 6.5-8 x 44.5 mm., oblong, with short style.

Hazara: Thandiani, 8500 ft., fr. June 1910, Parker, s.n., plant from fairly deep soil, an isolated specimen, fl. 1896, Duthie 19131.

Keshmir: Poshiana, 8OOOft., fl. 26 June 1902, Drummond 13893 (this is the ‘B . vulgaris var. aetnensis from Kashmir’, of Hooker & Thomson).

Punjab: Kunawur, 1834, Royle, s.n., Type; Rogi, 9OOO ft., fl. 1885, Drummond herb. no. 1223; Simla, Jarhorha, fl. June 1891, l0,000ft., Gamble 23690. Chmba, NWW, 7000-8000 ft., fl. 29 May 1896, Duthie 18320.

Tehri: Rikshin, 10,OOO ft., fl. May 1898, Gamble 26855 (all, K). 14-2

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212 L. W. A. ~ E N D T

According to Parker, common along the Miragan ridge in Hazara, and in Kagan, but not, as k y l e claims, in Kmawur, or even in Chamba. Usually a dense p t c h in exposed situations.

an extreme N.W. species in the Himalayas (con- bed, in fact, to Hazara and @an on the Hazara4.W. Hashrmr * hntier) , in contrast to the slightly more easterly B. e d g e W k m (lying from Punjab to Gahrwal). My deter- mination of herbarium sheets doea not conwpond with this, a fact which may be further evidence in favour of the existence of a seriea of linking forms; ultimately we may regard this whole population &B the variable contents of one larger and more widespread species. From h h m i r , Tilptra forest, near Rampur, Jhelum valley, come two specimens from lO,OOOft., on 8 July 1940, Ludlow & S h e d 7765, 7766 (BM), with reduced 4-12-fld., subracemoae, to subfawicled inflorescences.

Parker would seem to regard this

(288) Berberis grandibracteata Ahrendt, sp.nov., atT. B. edgewmthianae et B. kuw- wuren.si sed foliis integris crebre reticulatis, bracteis pedicellis distincte longio- ribus, ovulia 3-5, stylo distincto, distinguenda

Stemsglabrous,sulcate; youngshootsreddish; matureyellow-brown. Internodes 1-2 cm. Spinm 3-fld.. 8-13 mm. Leaves 10-20 x 4-7 mm., obovate, cuneate, sessile, entire, acute, closely reticulate both sides ; below subconcolornus green, sublustrous. Panicles 10- 15-fld., seasile, 1-1.5 cm. long. Pedkds 1-2 mm., in flower, 1.5-2-5 mm., in fruit; bracts longer, 1.25-3 mm. 0vule.s 3-5. Berries oblong-globose, 6 x 5-6 mm. Xtyk 0.5 mm.

Bhutan: seed. Cultivated: a t Headfort; dried specimens inscribed ‘believed to have come from

Bhutan’ fl. (Kew sheet 1323); fr. (Kew sheet 1324) (Type).

Subsect. Pseudopolyanthae Ahrendt, Kew Bd. 1939, 274, 1939 Stems mostly dark red, sulcate, sometimes pale yellow. Leaverr mostly entire semi-

evergreen, grey, pruinose, below; always closely reticulate both sides. Injbreswnce 2-8 (-12)-fld., mostly fascicled, sometimes racemose-subfascicled, 5-12 (-25) mm., never paniculate. Pedicel% (1.S) 4-8 (-12) mm.; bracts often much, rarely slightly, always distinctly, shorter. OvuEes (2-) 3-5. Be& pink to red, 4 4 x 3-6 mm. Style 0.25- 0.5 mm., distinct.

Kansu, Szechuan, Yunnan, Kweichou, Burma, S.E. Tibet (Prov. U). This interesting subsection, was at first placed by Schneider in the Sect. Sinenses, which then also contained the present Sect. Franchetianae. With the latter, this subsection shares the mostly entire leaves, the few-flowered inflorescence, but M e r s in its mostly smaller, always closely reticulate leaves, and always small berries with distinct style. Later this subsection was transferred to the Sect. Polyanthae, with which there are obvious relationships; but it lacks the chief mark of the Polyanthae, the ram paniculate in- florescence. But it will be seen that B. arkb-ealida is closely related to B. aggregata, and that the latter and B. pattii hybridize so freely with the B. w i & m varieties as to suggest some kinship. Again, although there has been no experience of cultivation in this case, there would appear to be relationship between B. qrandibracata and B. beesiana.

-Y 53, to Subsect. PseudopolyantJue (Map 37) Ovules 1-2 (-3).

Stems pale yellow; leaves below grey, pruinose; pedicels 1-9 mm.; fruit 4-5 mm. Stems puberulous ; leaves spinose ; pedicels 1-2 mm. ; bracts subequal.

Stems glabrous; leaves entire; pedicels PQ mm.; bracts much shorter. 289. B.arido-calidcr

290. B. heteropsie

291. B. hemeleyana Stems dark red; leaves below green, epruinose; pedicels 8-16 mm.; fruit 10 mm.

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Berberis a d Mahonia 213

Ovules (3-) 4-5. Stems puberulous. Stems pale yellow.

Leaves green, epruinose below. Leaves grey-white, pruinose below.

292. B. beekana 293d. B. x Tom Thumb

Stems dark red ; leaves grey-white, pruinose, below, and semi-evergreen. Leaves broad, 1 : 1.25-2; shrub erect, 6 ft. Leaves narrow, 1 : 3-5-4; shrub semi-prostrate, to 3 ft.

Cf. 282 b. B. coryi

293. B. &m var. w i l s m (Key 53c) Stems glabrous.

Leaves blue-green; apex obtuse, rounded or spathulate; fruit globose.

Leaves yellow-green; apex acute; fruit ovoid, or oblong-ovoid. 294. B. wilsonae var. aubcadialatcG (Key 53d)

295. B. w i k m var. stap$ana (Key 53e)

(289) Berberis arido-calida Ahrendt, Kew BuU. 1939, 273, 1939 Shrub suberect, 4-5 ft. Stems sulcate; young shoots slightly reddish, puberulous;

mature pale yellow, puberulous, black-verruculose. I&rnodes 1-2 cm. Spines 3-fld., slender, finely sulcate, 8-13 mm. Leaves deciduous 10-20 x 5-10 mm.; elliptic-obovate; base cuneate, subsessile; apes subacute; margins 2-6-spinose (1 mm.)-sedate , but often entire on new shoots; both sides closely reticulate; above dull yellow-green; below grey- pruinose. InJEorescence 6-10 (-14)-fld., racemose-subfascicled, not paniculate, 8-15 mm. Pedicels 1.5-2 mm.; bracts 1-1-5 mm. Ppophylls lanceolate, 2 x 0.8 mm. Outer sepals 3 x 1 5 mm., ovate, acute. Inner sepals 4 x 2-5 mm., obovate. Petals 3.5 x 2 mm., ovate- elliptic; apex scarcely emarginate; base truncate with suborbicular glands, 0.7 x 0.6 mm. Stamens 2.25 mm., slightly produced, obtusely subapiculate. O v u h 2. Berries 6 5 x 3- 4 mm., oblong-ellipsoid. Style 0.5-0.75 mm.

Kansu: Siku-Satanea ranges, 1914, seed, Farrer 355, ‘Small, growing in open upper alpine turf, with tightly bunched blood-scarlet berries, a native of hot, dry, regions’; 1914, fl. Mayer 2206 (K) ; West Kansu, Chone and Touchow district, 8OOO-9OOO ft., 25 Apr. 1912, seed, Purdom 826.

Cultivated: introduced by Farrer, 355, fl. 1 July 1938, 6 June 1942, 18 July 1945; fr. 2B Oct. 1939 (Type). This species has also been raised from Forrest 12355, and is related to B. aggregata.

(290) Berberis heteropsis Ahrendt sp.nov., aff. B. wilsonae, sed ramulis glabris

8te.m glabrous, sulcate, mature pale yellow. Intemzodea 1-2-5cm. S p i w 3-fld., 5-15 mm. Leaves 15-25 x 4-8 mm.; narrowly obovate; cuneate, sessile; entire; obtuse or rounded; closely reticulate; below grey, pruinose. Inflorescence 4-8-fld., fascicled or race- mose-subfascicled, 5-15 mm. Pedicels glabrous, 4-9 (-12) mm. ; bracts much shorter. Ovukp (1-) 2. Berries 4-5 x 3 4 mm., oblong-ovoid. Style 0*5-0.75 mm.

Kweichou: Long-ly, fr. 15 Nov. 1907, Cavalerie 3042 (Type, K). ‘Fruits roses, fleurs jaunes; rare.’

Szechuan: Lu-shan-hsien, 3250ft., 16 Oct. 1936, Chu 4006 (E). ‘Along roadsides, 4 4 ft., common.’ Not cultivated.

pallide flavis, ovulis tantum 1-2 distinguenda

(291) Berberis hemsleyana Ahrendt, sp.nov., aff. B. Urilsonae, sed foliis subtus viri- dibus, pediceh longioribus, ovulis tantum 2, fructibus multo majoribus et estylosis, distinguenda

Stem glabrous, sulcate, dark red, fairly or markedly stout. Internodes 6-15mm. Spina 8-25 mm., stout, orange, sulcate. Leaves to 22-28 x 7 mm., entire; closely reti- culate, oblanceolate; apex acute; below at first slightly grey, pminose, finally very pale

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214 L. W. A. AIERENDT lustrous green. Injbreswnce subfascicled or racemose-subumbellate, 1-26 cm. long. Pedi& 7-16mm., glabrous, stout. Outer sepals 3.5x2-5mm., ovate. Inner se& 5.5 x 3-5 mm., obovate. Petals 3-5 x 2 mm., obovate, clawed. Stamens 3 mm., truncate. ovules 2 (-3). Berries 10 x 5 mm. oblong, eatylose.

S.E. Tibet: h v . U, Ganden, 25 d e a east of Lhasa, 13,5OOft., fr. 22 Sept. 1943, Ludlow & S h e d 9948 (Type, BM). ‘Shrub 5-6 ft., on hillsides among roses and Cotawasters’; Lhasa, 13,OOOft., 17 May 1943, Ludlow & S h e d 9504 (BM). ‘Open rocky hillsides.’ Not cultivated.

(292) Berberis beesiana Ahrendt, sp.nov., ramulis flavia, foliis subtus viridibus,

Stem mature pale yellow-brown. Spines 3-fld., 1&15 mm. Leaves obovate, sessile, entire, acute, closely reticulate; slightly lustrous deep green; below dull, epruinose, green. O W 4-5. Berries globose, 6 mm. Style 0.2 mm.

inflomcentiis faaciculatis vel suburnbellatis, o& 4 4 , distinguenda

KEY 53B Stem puberulous; leaves to 12 x 5 mm., below paler yellow green; inflorescence 6-10-fld.,

Stems glabrous; leaves 10-25 x 3-7 mm., below concolorous; inflorescence 2-4-fld., fascicled; pedicels 2-3 mm.; bracts 0.6-1 mm.

subumbellato; peduncle 1-10 mm. ; pedicels slender, 5-8 mm. ; bracts 1-1.6 rnm.

2920. B. b&nu var. b&na

292 b. B. beeaiana var. glabra

(292a) Berberis beesiana var. beesiana Bhutan: seed, 10,OOO ft., from a 10 ft. high bush, Bees 3491. Cultivated: at Glasnevin, fr. 12 Oct. 1920 (Type, at Kew, H/1913/20).

(292b) Berberis beesiana var. glabra Ahrendt, var.nov., ramulis glabris, floribus

N.E. Bhutan: Me La, Cho La, valley, 12,500 ft., fl. 30 June 1949, Ludlow, S h e d & Hicks 20432 (Type, BM). ‘Bush 5-7 ft., among patches of shrub in otherwise open valley.’ Not cultivated. This should be compared with B. &eumthianu.

paucis, pedicellis longionbus, inter &a, distinguenda

(293) Berberis wilsonae Hemsley, Kew Bull. 1906,151, 1906; Rain, Bot. Mag. 138,

Shrub with arching branches, spreadmg 4-6 ft. acroas, but only 1-3 ft. hlgh. Stems dark red, sulcate. Internodes 1-2cm. S p i w 3-fld., slender, 1-2cm. Leaves semi- evergreen, 10-25 x 2 - 5 4 mm. ; obovate-spathdate; cuneate, d e ; rounded; closely reticulate both sides; dull above; below grey, pruinose. Injbresscence PT-fld., fascicled, 7-10 mm. long. Pedi& P 7 mm. ; bracts 1 mm. ouler s e p b ovate, acute. Inner sepals obovate. Petals obovate; narrowly emarginate with subacute lobes. Btamens 3 mm., slightly produced, obtuse. ovules 3-5. Bewies soft, pinkish red. Style 0.5 mm.

t. 8414, 1912.

KEY 53c Shrub 2-3 ft.

Leaves 10-25 x 3-6 mm. Leaves 4-8 x 1-2 nun.

Shrub scarcely 1 ft. high.

293a. B. d o n a e var. evilsonas 293b. B. wilsonae var. pannfolh

2930. B. wileonae var. fauoea

(293 a) Berberis wilsonae var. wilsonae

P& 4 x 2 mm. Berries 6 mm., globose. Leave.9 dull grey-green above. Outer se@ 3-4 x 2-3 mm. Inner sep& 5.5 x 3.5 mm.

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Berberis and Mahonia 215

W. Szechuan: roadside, 7800 ft., Aug. 1903, Wilson (Veitch) 3154 (Type, K). Kungtin hsien, Tachien lu, 2-3 ft., 9OOO ft., 22 Sept. 1928, Fang 3522 (K).

Yunnan: ledges of cliffs, and rock slopes, 2-3ft., l0,000ft., July 1923, Forrest 27044 (K).

Cultivated: L.A. 83, fl. 24 Aug. 1939, June 1942; fr. 12 Oct. 1938. One of the most widely cultivated Barberries, though probably out-numbered by the many hybrids (named and casual) which its varieties have produced in English gardens.

(293 b) Berberis wilsonae var. parvifolia (Sprague) Ahrendt, stat. nov.; syn. B. parvi- folia Sprague, Kew BuU. 1908,445, 1908 non Lindley, 1847

Outer sepals 2 mm. Inner sepals 3.5 mm. Petals 2-75 mm. Berries globose, 4 mm. Szechuan (N.W.) : Min valley, 1904, Wilson (Veitch) 3154A (Type, K). Cultivated: a t Kew, from seed of Forrest, 12979.

(293~) Berberis wilsonae var. favosa (W. W. Sm.) Ahrendt, stat.nov.; syn. B. favoscr W. W. Sm., Not. Bot. Card. Edinb. 11, 200, 8, 1919

TJpper Burma: Hpimaw, 7000 ft., Aug. 1914, Kingdon-Ward 1852 (Type, E). Prostrate dwarf shrub, rising less than 1 ft. from the limestone rock over which its long branches trail. Not cultivated.

(298d) Berberis x ‘Tom Thumb’ Morris ex Ahrendt, J . R. Hwt . SOC. 67,133, Apr. 1942

Differs from B. wilsonae mainly in its yellow stems. Cultivated : by Mr Sidney Morris, a t Earlham Hall, Norwich, being raised from seed of

Wisley hybrids. See J. R. Hort. 8oc. 49, 1923; Oarden, 87, 555, 1923.

(294) Berberis wilsonae var. subcaulialata (Schneid). Schneid, &t. Bot. 2. 57,298,

Stem9 glabrous. Leaves conspicuously blue-green. Fihwers fascicled. Berries 6 mm., globose (but of variable shape in hybrids).

Szechuan: between Tachien lu, Tseku, and Mekong, 1894, seed, Soulie, s.n. ; Muph, 7000 ft., Wilson 1267.

Yunnan: Weihsi, west of Tung-chuling, 9750 ft., 14 Nov. 1937, Yu 10708 (E), shrub, 1-2 ft., common. Also Forrest 11500, 15316 (K).

Cultivated: from Soulie’s seed, Type. Also, in England, from Yu’s seed, 10708; 0. 6 June 19-12, 21 Aug. 1943, 18 July 1945; fr. Oct. 1942. Further distinction must be made here.

1918; syn. B. submulialata Schneid., Fedde Rep. Sp. Nov. 6, 267, 1909

K E Y 53D Inflorescence 3-7-fld., always fascicled. 294. B. wilsonae var. subcauliaEata Inflorescence 7-15-fld., mostly subracemose-fascicled.

Fiuit globose-ovoid to globose, 5-6.5 x 4-6.5 mm. ; leaves markedly semi-evergreen. Fruit globose. Fruit not globose.

294a. B. wilsonae var. guhtzunica

294 b. B. x Comet 294c. B. x Coral

Fruit globose -ovoid. Fruit oblong-ovoid.

Fruit ovoid-conical, 7-9 x 6-7 mm. ; leaves subdeciduous, or less conspicuously persis- tent.

Flowers small; sepals in two series, inner 4-25 mm. Flowers larger; sepals in three series, inner 5.5-6 1~1111.

294d. B. x Firefly 294e. B. x Fireball

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216 L. W. A. ~ N D T

(294a) Berberis wilsonae var. guhtzunica Ahrendt, comb.nov.; syn. B. eubcQulialatcG

Differs from 294 only in its racemose-subfascicled, instead of p d y fascicled, in-

S.W. Szechuan: Muli, Guhtzun, 10,OOO ft., 5 Dec. 1937, Yu 1 4 W ( T y p , E). ‘Com-

Cultivated: from Yu 14840, fl. 4 Bug. 1943, 18 July 1945; fi. 9 Dec. 1943. Also, at

vm. sui;tzu7c;ea; Ahrendt, J. Bot., Limd., 79 (Suppl.), 76, 1942

Boreaceme.

mon.’

Liss, Gamble 30879, from F o m t seed (dried apcimen, K).

(2943) Berberis x (Comet’ Chittenden ex Ahrendt, J. R. Hort. Soc. 67,133, Apr. 1942 Be&q H.C.C. 19. Cultivated: a t Wisley.

(294c) Berberis x ‘Coral’ Chittenden ex Ahrendt, I.c., 1942 B e w h H.C.C. 21. Cultivated: a t Wisley, fl. 12 July 1938; fr. 16 Nov. 1938.

(294d) Berberis x (Firefly’ Chittenden ex Ahrendt, I.c., 1942 Berries H.C.C. 18. Cultivated: a t Wisley, fl. 12 July 1938; fi. 16 Nov. 1938.

(294e) Berberis x ‘Fireball’ Chittenden ex Ahrendt, I.c., 1942

BerriM H.C.C. 18. Cultivated: at Wisley, fl. 12 July 1938; fr. 16 Nov. 1938.

(295) Berberis wilsonae var. stapfiana (Schneid) Schneid., ost. Bot. 2.57,298, 1918; syn. B. sta/@na Schneid., Kew BuU. 1912, 35, 1912; Hutchinson, Bot. Nag. 43, t. 8701, 1917

Leaves 10-20x24 mm., acute. P e d M 2-4 mm.; bracts scarcely 1 mm. Outer Be& 2.5 x 2 mm., broadly obovate, rounded. Inner se& 4 x 3 mm., spathuhte- obovate, 4 x 3 mm. Petals 3-5 x 1.75 mm., obovate, emarginate with rounded lobes. Stamens 3 mm., slightly produced, truncate. Berries ellipsoid or ovoid, 4-5 x 3-4 mm. Style 0-5 111111.

W. China: seed sent to Messrx Vilmorin. Cultivated: f h m seed cited, Vilmorin 4039 (Type, K).

(295a) Berberis wilsonae var. latior Ahrendt, var.nov., foliis latioribus h u e s 5-17 x 2-5 mm., acute, yellow-green. InJlorescence S-g-fld., racemose-sub-

fascicled. Pedieels 2-3 (-4, in fnrit) mm.; bracts 1 mm. Prophylla 2 x 1 mm. Outer s e p h 4 x 3 mm., obtuse. Inner sepals 5 x 3 nun. P& 4-5 x 2-5 mm., acutely emar- ginate, with acute lobea. 8tmnen.s 3mm., shortly apiculate. Berries oblong-ovoid, 7X5mm.

Yunnan: seed only, Forrest 30459. Cultivated: from Forrest 30459, fl. 3 June 1942,7 July 1945; fr. 27 Oct. 1944 (Type, 0). Distinguish its foIlowing further varieties and hybrids.

KEY 53E Idloreaceme 3-7-fld., always fmcicled. 295. B. w i l s o m var. stcvpfiana InfIoreacence 7-12-fld., racemose-subfasoicld.

haves broad, 1 : 2.5-3. 295a. B. udsonae var. latior

Page 217: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 217 Leaves narrow, 1 : 5.

Fruit oblong-ovoid, 5-6 x 3-3-6 mm., H.C.C. 19. Fruit globose-ovoid, 6 x 6 mm.

295 b. B. x Ferax 295c. B. x smithianu

(295 b) Berberis x ‘Ferax’ Chittenden ex Ahrendt, J. R. H w t . Soc. 67, 133, 1942 Cultivated: a t Wisley.

(29Bc) Berberis x smithiana Sprague ex Ahrendt, I.c., 1942 Cultivated: Guernsey, Caledonian nursery of Measrs C. Smith (Type, K).

Sect. Sherriffianae Ahrendt, J. Bot., Lond. 79 (Suppl.), 77, 1942 Stems dark red, or red-brown, when mature. Internodes 1-3cm. Spines 1-3-fld.,

P l i m m . , discolorous, yellow. Leaves deciduous; above dull green; both sides with branched venation, sometimes reticulate ; below grey, pruinose. Panicles 10-70-fld., 4-16 cm. long. Pedicels 1-5 mm. ; bracts 1-5-3 mm., from 4 to 14 times as long as their pedicels. Petals shorter than inner sepals. Fruit black or purple, pruinose blue, 6.5- 10 mm. long, always estylose.

S.E. Tibet: Provinces Pome, Kongbo, and Tsari district.

KEY 54, to Sect. Sherriffianae (Map 38) Bracts subequal to, or much longer than, their pedicels; stamens not produced, rounded-

truncate. Leaves closely reticulate; ovule ; solitary. 296. B. taylorii Leaves openly reticulate, or with only branched venation; ovules 3-5. 297. B. gyalaica

298. B. sherrifii Bracts only half as long as pedicels; stamens produced, apiculate.

(296) Berberis taylorii Ahrendt, J. Bot., Lond. 79 (Suppl.), 71, 1942. (Fig. 52) Stems puberulous, subterete; mature light red-brown. Internodes (0.7-) 1-1.5 (-2) cm.

Spi.nes solitary, 5-11 mm., yellow-brown. h u e s obovate, 1.5-3 x 0-7-1-3 em., with de- current petiole 1-3 mm. ; margins entire (rarely 1-3 spinules) ; obtuse or rounded ; above yellow green. Panicles narrow, (20-) 30-50 (-70)-fld., 4 - 6 9 cm. long, including peduncle (0.6-) 1-14 cm. Pedicels puberulous, 1-2 mm.; bracts longer, puberulous, 2-3 x 1- 1.25 mm. Outer sepals 3 x 1.5 mm., ovate, acute. Inner sepals 4 x 3.75-4 mm., obovate, orbicular. Petals 3-3-5 x 2-25-3 mm., obovate; apex acutely emarginate with acute lobes; base clawed, with oblong, separate, rounded glands, 0.6 x 0-3 mm. Stamens 3 mm., not produced, rounded. O w b solitary. Berries narrowly ovoid 8 x 2-5 mm., estylose dark red or purple with blue bloom. Seeds pale yellow-brown.

S.E. Tibet: Prov. Kongbo, Lilung Chu, 9800ft., fr. 3 Oct. 1938, Ludlow, Sherriff & Taylor 7163 (Type, BM); ‘the fruiting specimen of 5180, with fruit reddish-purple’; Peru La along path in Quercus ilex thicket, 0 . rather greenish yellow, up to 5 ft., fl. 8 July 1938, Ludlow, Sherriff & Taylor 5180 (BM); Lamdo, 13 July 1938, Ludlow, S h e d & Taylor, 5821 (BM) ; ‘common on the banks of the Tsangpo river’; Lilung Chu, near Molo, 29 June 1938, Ludlow, S h e d & Taylor 5686 (BM); ‘little seen; buds greenish yellow’; Chab, Gyamdu Chu, left bank, 9800ft., fl. 29 July 1947, Ludlow, Sherriff & Elliot 14204 (BM); ‘open hill slopes and in lanes bordering cultivation, 3-5 ft.; flowers greenish yellow’.

Prov. Pome: Nambu La, 10,5OOft., fl. 9 July 1947, Ludlow, Sherriff & Elliot 15351 a (BM).

Cultivated: L.A. 264, from type collection 7163-in the University Botanic Garden, Oxford.

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218 L. W. A. AHRENDT

(297) Berberis gyalaica Ahrendt, Gurd. Chrm. (3), 109, 101, Mar. 1941; Sealy, Bot. Mag., 145, N.S., t. 22, 1948. (Fig. 53)

Shrub 6-9ft. Stems sulcate, not or scarcely verruculose. Leaves elliptic, sessile. Pan* 1530-fld.; peduncles 1-2 (4) cm.: subpeduncles of branched parts 4 7 mm. Flowers H.C.C. 2/1. ProphyUs concolorous, 2-3 x 1-1-5 mm., oblong, acute. Outer sepaLs 4-5 x 34.5 mm., oblong-obovate. Inner sepals 6 x (4) 5-6 mm., oblong-suborbicular. Petals 4 x 3 4 mm., broadly obovate; apex rounded, narrowly incised with rounded lobes; base clawed, with oblong-elliptic, widely separated, submarginal glands, 0-7 x 0.5 mm., situated above the claw. Stamens 3 mm., not produced, truncate. Ovdea 3-5. Bewim oblong-ovoid, 9-10 x 4-5 mm., black pruinose blue, estylose. S& pale yellow.

KEY MA

Stems and pedicels puberulous. Leavens spinose, 6-15 x P 9 mm.; panicles 3-7 cm. Leavens entire, 15-30 x a 1 4 mm.; paniclea 8-10 cm.

297a. B. gycclaiccl var. g y & h 297 b. B. gyahiw var. maxirni$ora

297c. 23. gg/ala& var. minuaka Stems and pediceh glabrous; lerrvea entire, 15-30 x 8-12 mm.; paniclens 3-5 cm.

(297a) Berberis gyalaica va. gyalaica Pedicels (1-5-) 2-3 mm.; bracts 1.75 x 2.75 mm. Leaves with margins 3-5-spinose

(1 mm.)-serrulate, but entire on shoots. S.E. Tibet: Prov. Pome, Gyda, 9oo0-10,OOO ft., 19 July 1924, bush 5-8 ft., in thickets

on exposed side, seed, Kingdon-Ward 5962; Temo la, 10,500 ft., fr. 4 Oct. 1947,34 ft., on dry ground, Ludlow, SherriiT & Elliot 15825 (BM).

Cultivated: fl., fr., 1939 (d of KW 5962), a t Exbury; fl. 8 July 1943; fr. 28 Oct. 1943

The cultivated plants were, a t fmt, incorrectly numbered Kingdon-Ward 5936, which (Tgpe, BM).

is the type of B. johannis.

(297b) Berberis gyalaica var. maximiflora Ahrendt, varmov., paniculis multo

Panicles 8-16 cm. long. Pedicels 2-5 mm.; bracta 2 4 mm. Be& 13 x 7 111112. S& red-brown.

S.E. Tibet: Prov. Pome, Temo La, 11,OOO ft., 3 4 ft., dry ground, 4 Oct. 1947, Ludlow, S h e d & Elliot 15826 (Type, BM).

Cultivated: from Type collection.

elongatis

(297c) Berberis gyalaica var. minuata Ahrendt, var.nov., ram& et idorewentiis glabris, foliis integris, paniculis abbreviatis

Pedicds 1-2 mm. ; bracts 2-3 nun. Seeds yellow-brown. S.E. Tibet: Prov. Kongbo, Tsangpo valley, Tse, 95OOft., 10 Oct. 1947, Ludlow,

S h e d & Elliot 13304 (Type, BM). Cultivated: from Type collection.

(298) Berberis sherrifEii Ahrendt, J. Bot., Lond. 79 (Suppl.), 77, 1942 &,%nu glabrous; young shoots green, slightly sulcate; mature red-brown, subterete to

subsulcate, slightly verruculose. Leaves 12-24 x 5-13 mm. ; obovate; attenuate to petiole 1-5 mm., entire. Panicles 10-20-fld., 3-5 om. Pedicek glabrous, 2-4 (-5) mm.; bracts

3.5 x 2.5 mm., oblong, acute. Median se@ 4.25 x 3.5 mm., oblong-elliptic. Inner sep& 6 x 5 mm., obovate. Pe&h 5 x 3-75 mm., obovate; apex minutely acutely emmghate,

1.5-2-5mm. F ~ T S H.C.C. 1. P~OphylLS 2 ~ 1 * 4 m m . , Oblong, acute. Oarter sepals

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Berberis and Mahonia 219

with acute lobes ; base scarcely clawed, with widely separated, non-marginal glands, 0.8 x 0.4 mm. Stamens 3 mm., slightly produced and shortly apiculate. Ovules 1-2 (-3). Berriu 6.5-7 x 3 4 mm., oblong-ovoid, black pruinose blue, estylose.

H.E. Tibet: Tsari district, Migyitun, Tsari Chu, 1O,ooO-11,OOO ft., on edge of forest on steep hillside, 24 Oct. 1938, Ludlow, S h e d & Taylor 6629 (Type, BM); ‘bush 3-6 ft.’

Cultivated: from type collection, fl. 29 June 1945; 11 July 1952; fr. 20 Nov. 1944.

I ~ Y 55, to Sects. Heteropodae, Crataeginae and Ulicinae stems puberulous. 299. B. loudonii Stems glabrous.

Mature stems yellow; fruit estylose. Leaves with 8-25 marginal spinules.

Leaves green below; fruit 9 x 5 mm. Leaves grey below; fruit 15 x 10 mm.

Leaves below green, epruinose. Leaves below grey, pruinose.

302. B. mtnenais 317. B. x notubilis

304. B. garcim 312. B. stolonifera

Leaves entire.

Mature stems dark red, or purple; leaves entire; pedicels 4-10 (-20) mm. Fruit with distinct style, (0.3-) 0.75-1-5 mm.

Stems terete; leaves 3-6 cm., below pruinose, grey. Stems sulcate; leaves 1-3 cm., below epruinose, green.

307. B. oblonga

Style short, scarcely 0.3 111111. ; fruit scarcely pruinose. Ovules 3-4; leaves narrow, 1 : 4-5; petals as long as inner sepals.

300. B. cratuegina Ovules 2; leaves broad, 1 : 2; petals shorter than inner sepals. 301. B. hispanica

Leaves broad, 1 : 2.5 ; pedicels 2-5 mm. ; petals longer than or equal to inner

Leaves often narrower, 1 : (2-) 3-5; pdieels 5-12 mm.; petals shorter than

Style long, 0.5-1.5 mm.; fruit conspicuously pruinose; ovules 2.

sepals; style 0.5-0.75 mm. 308. B. cretica

inner sepals; style 1-1.5 mm. Leaves fairly narrow, 1 : 2-3. 309. B. calliobotrys Leaves narrower, 1 : 4-5.

Leaves reticulate; inflorescence racemose, 2-3 cm. 310. B. wazuristanica Leaves not reticulate ; inflorescence subfascicled, 1-1.5 cm.

311. B . gambleana Fruit estylose.

Leaves grey, pruinose below ; petals longer than inner sepals ; fruit conspicuously

Leaves 10-30 x 6-15 mm.; inflorescence subumbellate; fruit 9-10 x 9 mm.,

Leaves 7-15 x 6-12 mm. ; inflorescence subfascicled; fruit 8 x 4 mm., oblong.

pruinose.

obovoid. 313. B. psewlumbellata

314. B. royleana Leaves epruinose, green, below ; petals shorter than inner sepals.

Fruit conspicuously pruinose. Fruit 12 x 8 mm. ; leaves 2-5-55 cm. Fruit 7 x 4 mm. ; leaves 1-2.5 cm.

Fruit epruinose, or scarcely pruinose. Leaves broad, 1 : 1.5-2.5.

305. B. nzaderenais 315. B. aitchisonii

Ovules 4-6 ; stamens apiculate. 316. B. heteropoda Ovules 2-3; stamens truncate.

301. B. hispanica

Cf. 305. B. nzaderensis

Ovules solitary. 306. B. libanotica Ovules 3-5. Sect. ULICINAE

Leaves 1-2.5 cm. ; pedicels 5-8 mm. ; fruit 9 mm. Leaves 2.5-5.5 cm.; pedicels 8-20 mm.; fruit 12 mm.

Leaves narrow, 1 : 4-7; fruit epruinose.

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220 L. W. A. AHRENDT

Sect. Crataeghae Ahrendt, sectnov., ramulis plenunque atrorubris, f o h deciduis subtua plenunque viridibus, lase nervatk, inilomcentiis fasciculatis vel subrace- mosis, fructibus atris, epruinosis vel vix pruinosis, estylosis vel stylis indistincte visibilibua, vix 0-3 mm. longis.

Stenzs dark red, glabrous (except only in B. loudonii, B. gar&); always sulcate. Spines (1-) 1-5-2.5 (-3) cm., yellow or yellow-brown. h u e s deciduous; venation mostly openly branched, occasionally reticulate. In.escence 3-20-fld., fascicled or subracemose, 1-2.5 (4) cm. long. Pedkds glabrous 4-11 (-20) mm.; bracts 1-2 (-26mm.), from one-quarter to one-twelfth the length of the pedicel. Plowers 6-8 (-10) mm. diameter. ovules 1-2 (4). Stamens not, or slightly produced, rounded, or truncate. Fruit black, generally epruinose or very slightly pruinose, occasionally more pruinose. Style absent, or scarcely 0-3 mm. long.

Madeira, Spain, Morocco, Algeria, Sardinia, Corsica, Sicily, S. Italy, Caucaaus, Turkey, Armenia, Syria.

Fruit with short style, not or scarcely pruinme.

KEY 56, to Sect. Crataeginae (Map 3 and 39)

Stems yellow, pubcent. 299. B. loudonii Stems red, glabrous.

300. B. crataegina 301. B. hiapanicu

Fruit estylose; leaves green, ep&ose below. Leaves with 10-2O-marginel epinulea; inflorescence (6-) 10-20-fld. 302. B. aetnenais Leaves entire; inflorescence 3-10 (-15)-fld.

Ovules 3-4; leaves narrow, 1 : 4-5; petals as long as inner sepals. Ovules 2; leavea broad, 1 : 2; petals shorter than inner sepals.

Stems yellow. 304. B. garciae Stems dark red.

Leaves 25-55 x 8-20 mm. ; pedicel8 8-20 mm. ; fruit 12 rnm., slightly p&ose.

Leaves 10-20 mm. long; pedicels 4-8 mm.; fruit 9 rnm. 305. B. d r e n s i s

Leaves narrow, 1 : 5. 306. B. libanotica Leaves broader, 1 : 2-2.5. 301a. B. hiqtmnica var. hackeliixrua

(299) Berberis loudodi Ahrendt, sp.nov., ramulis flaveacentibus pubeacentibus, foliis deciduis subtus pruinosis, fructibus atrocaedeis vix pruinosis stylo brevis- simo.

Shrub fairly dense, with stiff, arching branches, 5-6ft. Stem s u l ~ t e ; young shoots green puberulous; mature yellow-brown, puberulous. I n t e e 1 . 5 3 em. Spines 1-2 cm. Leaves to 27xlOmm. obovate, subsessile, entire, acute; both sides with slightly branched venation; above dull light grey-green; below grey, pruinose. Inflorescence 6-lO-fld., umbellate-racemose, 1.5-3 cm. Pedicels glabrous, 4-8 (-11) mm. ; bracts 1 mm. Flowers H.C.C. 1/1-1/2. ProphyUs red, oblong, acute, 1-.5-2*75 x 1-1-5 mm. Outer sepals 2.25-2.75 x 1-5-2 mm., broadly ovate, subacute. Inner sepal8 5-5-6 x 44 .25 mm., obovate. Petals 5x4mm., obovate, rounded; entire; cuneate with orange coloured glands. Stamens 4mm., not produced, truncate. O v d ~ 2. Berries black, almost epruinose or with slight blue bloom, 7 x 4 nun. Style very short. Seeds pale yellow- brown.

1 Caucasus: without locality; distributed aa B. chinenads. Cultivated: L.A. 123, fl. 29 Apr. 1943,26 Apr. 1945; fr. 28 Oct. 1944 (Type, 0).

(300) Berberis crataegina DC., Syst. 2, 9, 1821 Shrub 6-7 ft. Stem glabrous, sulcate, mature lustrous dark red. Internodes 1-25 cm.

Spines yellow, stout, sulcate, 1-2-5 cm. h u e s 15-3Ox 3-7 mm., or to 48x 8 mm., oblanceolate, attenuate, sessile, entire (rarely 1-2 spinules), acute; both sides con-

Page 221: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 22 1

colorous sublustrous deep green, with openly branched venation. Racemes (6-) 8-14 (-18)-fld., (1-) 3-5 om. long. Pedicels slender, 5-9 mm.; bracts 1 mm. Flowers 7 mm. dkmeter. Inner se& 4-5 x 3 mm. Petals 4-5 x 2.5 mm. Stamns 3.5 mm., produced, truncate. Oprules 3 4 . Fruit black, epruinose, or scarcely pruinose, oblong-ovoid, 6-7 x 3.5-4 mm. Style short, distinct.

Under this name Fedtschenko describes a plant with terete stems, solitary ovulm, and dark red fruit, and may be referring to B. orientalis. B. crataegina is not in cultivation in any variety. B. crataegina waa described by De Candolle from plants collected in Asia Minor without spec& locality. According to Loudon in Arb. et Frut. Brit. 1, 305, 1838, 'young plants of what are said to be this species are in the garden, but have not yet flowered'.

KEY 56n Spines mostly solitary; racemes mostly 3-5 cm.

Leaves bright green, 1 : 3-4; racemes 1Ck15-fld. Ideaves paler grey-green, narrower (1 : 5-6) ; racemes 8-12-fld.

m a . B. cr&gina var. crataegina

300b. B. crataegina var. armenha 300c. B. cratuegina var. lyeiea Spines mostly 3-fld. ; racemes 1-3 cm.

(300 a) Berberis crataegina var. crataegina Asiatic Turkey: Anatolia, Selefke, Ala.ya Mts., July 1930, Byles H. 1793; below Ruster

Estref's house, Chankaya, near Ankara, one bush 6f t . high, 3250ft., 6 May 1926, Lindsay 19.

Clappadocia, 4200-6500 ft., 29 May 1898, Siehe 36 (all, K).

(30C)b) Berberis crataegina vm. armeniaca Schneid., BuU. Herb. Boissier, (2), 5 , 657, 1905

Turkish Armenia: Erzinghan, prope Albushik-chan, fl. 6 May; fr. 17 July 1890, Sintenis 3002 (K).

(300~) Berberis crataegina var. lycica Schneid., 1.c., 1905 Asiatic Turkey: Lycia, Elmaly on hills, fl. 13 May 1860, Bourgeau 12 (Type, K).

(301) Berberis hispanica Boiss. et Reut., Pugill. P1. Nov. 3, 1852; Willk. et Lange, Prodr. 3'1. Hisp. 3, 901, 1880; syn. B . australis Moris ex Schneid., Bull. Herb. Boissier, (2) , 5 , 685, 1905; syn. B. vulgaris var. australis Boiss., Voy. Bot. Esp. 2 , 2 p.p. 1839

Shrub s t a y suberect, to 5 ft. Stems glabrous, sulcate; young shootsgreen; mature dark red. Internodes 1-2 cm. Spines yellow-brown, 1-3-fld., 1-2 cm., with shorter lateral parts, slightly sulcate. Leaves 10-25 x 6 1 0 mm. ; elliptic-obovate; base cuneate to petiole 1 4 mm. ; entire; acute, mucronulate (0.7 mm.) ; both sides with open branched venation, scarcely subreticulate ; epapillose ; concolorous slightly lustrous green. Inflmescence 6-15-fld., subracemose, subfascicled, or subumbellate, 1-2 (-4) cm. long, including peduncle 3-5 mm. Pedicels slender, glabrous, 5-8 mm.; bracts 1-2 mm. Fbwers c. 6 mm. diameter. Outer sepals H.C.C. 211, oblong-ovate, acute, 2 x 1.5 mm. Inner sepals concolorous, 44 .25 x 2.3-2.6 mm., oblong-obovate. Petals H.C.C. 3 4 , obovate, 3 x 2 mm. ; entire ; base cuneate with ovate, acute, separate, submarginal glands, 0-7 x 0.3 mm. Stamens 2-2.5 mm., slightly produced, rounded. Ovules 2. Berries bla.ck, scarcely or slightly pruinose blue, ovoid, 9 x 5 mm.

Page 222: Berberis and Mahonia : A taxonomic revision

222 L. W. A. AHBENDT

KEY 5 6 ~ Fruit with very short style, C. 0.3 m. Fruit estylose.

301a. B. h+nb var. lhpanicu 301 b. B. hbpanb var. hackdi~na

(301 a) Berberis hispanica var. hispanica South Spain: Prov. Almeria, C ~ R O Almirez, 5 July 1908, St. Lager am.; Sierra de

Maria, N. slopes above Maria, fl. 30 Apr. 1928, Ellman & Sandwith 990. Prov. Granada, on Mt. Cerro Calm, 10 milea S.E. of Granada, 5000-6000 ft., 24 May

1924, Ellman & Hubbard 838; Sierra Nevada, 30 June, 21 July 1876, Winkler, 8.n. (a, K) ; seed only, Envers Todd, s.n.

North Africa, Algeria: Col de Tizougarine, 3 July 1879, Reboud 2352 (K); Pente sud du Djebel-Mahmel, fl. 6 June 1853, Bal- 946 (K) (this specimen has the longer in- florescence, 2-4 cm.).

S. Morocco: Greater Atlas, Mt. Teza, fl. May 1871, Hooker, s.n. (K). ‘Fruit dark blue.’ Cultivated: L.A. 431, from seed collection by Envers Todd, fl. 5 May 1943; fi. 22 Sept.

1943, 21 Sept. 1944.

(301 b) Berberis hispanica var. hackeliana (Schneid.) Ahrendt, comb.nov.; syn. B. australis var. h k l & a n a Schneid., Is., 659, 1905

South Spain: Prov. Granada, Sierra Nevada, ad. flum. Genil., Hackel, 8.n. 1876 (Type). Not in cultivation.

(302) Berberis aetnensis Real., Flora sicula, 1, 28,1826; Roem & Schult., Syst. Veg. 7 , i, 2, 1829; syn. B. vulgaris var. aetnensis (Presl) Fiori et Paoletti, Fl. Anal. Ital. 1, 529, 1898

Shrub compact, globose, 1-2 ft. Stem glabrous; young shoots slightly reddish; mature pale yellow, sulcate. Spines 3-fld., 1-2 cm. h u e s (1-) 2-36 cm. x (5-18 mm. ; oblong- obovate; margins 8-10-spinose towards apex; above grey green; below green epruinose. Racema (6-) 10-20-fld., ( 1 . 5 ) 2.5-4 (-7.5) cm. Pedicd.3 6-10 mm.; bracts 1-5-2 mm. Flowers 6-7 mm. diameter. Outer sepah 3.5 x 2 mm., elliptic. Inner sepah 4.5 x 3 mm., obovate. P e e 3-5 x 2 mm., obovate; glands separate, submarginal, 0.7 x 0-5 mm., elliptic. Stanzens 2.75 mm., not produced, truncate. O& 2. Fruit, while unripe red, finally when ripe, black, epruinose, or scarcely pruinoae, oblong, estyloae, 8-9 x 4-5 mm. Seeds dark brown.

KEY 56c Leaves 2S35 mm. long; idloreaceme 2-5-4 (7.5) cm., 10-20-fld.

Leaves obtuse. 302a. B. aetnenda vm. aetnenda Leaves acute. 302 b. B. aetmd var. cakdwku

303. B. boia&ri Leaves 10-20 nun. long; inflorescence 16-25 mm., 6-12-fld.

(302a) Berberis aetnensis var. aetnensis

in cultivation.

(302 b) Berberis aetnensis var. calabrica Schneid., Bull. Herb. Boksier, (2), 5,660,1905

Sicily: sterile regions around Mt. Etna, 6000-7500 ft., 1871-2, Gandoger 36 (K). Not

South Italy: Cahbria, Mt. Pollini, Rig0 356 (Type). Not cultivated.

(303) Berberis boissieri Schneid., 1.c. 1905 Internodes 7-15 mm. Spines concolorous, stout, 3-fld., sulcate, 1-5-26 cm. h u e s

obovate, 10-20 X 5-13 mm. ; margins 10-15-spinose (0.5-1 mm.)-serrate (0.5-1 mm.),

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Berberis and Mahonia 223

distant 2-2.5 mm.; apex rounded; below green. Pedicels 4-8 (-10) mm.; bracts 1-S 2.5 mm. Berries black, epruinose, estylose, oblong-ellipsoid, 7-8 x 4-5 mm.

Corsica: Mt. Renoso, Pozzi. 29 June 1917, Forsyth-Major H292-23; Tortello, fr. 26 Aug. 1916, Forsyth-Major, H292-33 (both, K).

Sardinia: March 1828, Thomas (K, sheet 1641). Not in cultivation.

(304) Berberis garciae Pau, Not. Bot., P1. Esp. 2 , 6, 1889; Willk. Suppl. Prodr. F1.

Stem yellow. haves entire, rarely with a few spinules. Pedicels 6-12mm. Fruit

East Spain: Aragon, Prov. Teruel, Sierra de Javalambre, Reverchon 405, 3190. A

Hisp. 314. 1893, Schneid., 1.c. 659, 1905

6-7 x 4-5 mm., estylose.

very little known species, not in cultivation.

(305) Berberis maderensis Lowe, Hook., J. Rot., Hew Card. Misc. 8, 289, 1856; Lowe, Man. Fl. Madeira, 1, 8 , 1857; syn. 3. lycioides Lowe l.c., 1856, pro syn., non Stapf.

Stem glabrous; mature dark red-purple. Internodes 2-4 cm. Spines absent, or 3-fld., 1-2 am. Leaves 2-5-5 x 1-2 cm., including petiole 3-8 mm. ; obovate; base attenuate; margins entire, rarely 14-spinules ; reticulate; below green, epruinose, epapillose. In- f i r e m m e 5-10 (-12)-fld., racemose-subfascicled, 2-2.5 cm., or partly racemose, partly umbellate, with an extra basal flower, and 3-4.5 cm. long. P e d k k (6-) 8-15 (-20) mm. Outer sep& 3 x 1.5mm., ovate, acute. Inner sepuls 6x3-5mm., obovate. Petals 4.5 x 2-5 mm., obovate. Stamens 4 mm., not produced, truncate. Berries black, pruinose blue, 11-12 x 8 mm., estylose.

Madeira: Agoa de Bica, Rib. da Metade, 13 Sept. 1838 (Type); Canhas, June 1855 Moniz in herb. Lowe; without precise locality, 1856, Mason s.n.; on rocks, Jardin da Serra, 4000-4500 ft., fl. May, fr. Nov. 1865, Mandon 3 (all, K). Not in cultivation.

(306) Berberis libanotica Ehrenberg ex Schneid., Ill. H a d . Laubh. 1, 310, 1905 Xtem glabrous, mature dull dark red. Spines 1-3-fld.) yellow-brown, sulcate, 1-3 cm.

Leaves 10-25 x 2-5-5 mm. ; oblanceolate; base attenuate, sessile; entire, acute ; venation both sides branched, scarcely openly reticulate ; below green, concolorous, epruinose. InJorescence Sg-fld., racemose-subfascicled, umbellate, or racemose-subumbellate, 8-12 (-20) mm., including peduncle, 1 4 mm. Pedicels slender, 4-7 mm. Outer sepals 2.5 x 1.5 mm., ovate. Inner sepals 5 x 3-25 mm., elliptic. Petals 4 x 3 mm., obovate, entire. Stawns 3-5 mm., not, or scarcely produced, truncate. Ovules solitary. Berries black, epruinose, or scarcely pruinose, oblong-obovoid, 9 x 5-6 mm.

Lebanon, Sanin, subalpine region, 5,20Of., 17 June, 1897, Bornmiiller 35; Aucher- Eloy 391 ; Zebdain, near Damascus, top of Mt. Garbus, 5000 ft., 4 June 1855, Kotschy 24 (all, K). Not in cultivation.

Sect. Heteropodae Schneid., BUU. Herb. Boissier, (2)) 5, 457, 1905; Ost. Bot. 2. 67, 216, 1918

Laroes deciduous, mostly entire. InJorescence 2-10-fld. Pknuers 4-8 mm. diameter. Ovules 1-6. Fruit black, sometimes pruinose, blue or white

Crete, Cyprus, Greece, Turkestan, Sinkiang, Afghanistan, Baluchistan, Wazaristan, Kashmir, Punjab.

KEY 57, to Sect. Heteropodae (Maps 3, 12, 40, 41) Style conspicuous (0.5-) 0.75-13 mm. long. Subsect. CRETICAE

Stems terete; leaves 3-6 cm., below grey, pruinose; inflorescence 10-20-fld., PI cm. long; ovules 3; petals longer than sepals; fruit 11 x 7 mm. 307. B. oblonga

Page 224: Berberis and Mahonia : A taxonomic revision

224 L. W. A. AHRENDT

KEY 57 (CmtiTaUed) Stem sulcate; laves 1-3.5cm., below green, epruinose; infloreacence 3-8-fld., 6-

Leaves to 17 x 7 mm. ; intlomnce 6-12 mm. ; pedicels 2-5 mm. ; petals longer than

Leaves to 36 x 12 mm.; intlomnce 15-30 mm. ; pedicels 5-10 mm. ; petals shorter

30 mm. long; ovules 2; fruit 7-8 x 6-6 mm.

inner sepals; fruit epruinose; style 0-5-0.75 mm.

than inner sepals; fruit conspicuously pruinom; style 1-16 mm.

308. B. Oreticcr

h v e s broad, 1 : 2-3. 309. B. d W b O e 8

Leaves n8lTow, 1 : 4-5. Leaves reticulate; intloscence racemose, 2-3 cm. long. Leaves not reticulate; intlorcsnce subfescicled, 1-1.5 cm. long.

310. B. w a z w k w n k

311. B. gambltwu Style absent. (Subsect. PSEWDUMBEILATU.)

Stem yellow; inflorescence 10-20-fld., 6-9 cm. long; pedimls 1-2 om.

Stems dark red, or purple; inflorescence 2-9-fld., 1-5-3 (-5) cm. long; pedicels 4-10 312. B. atolonqera

(-15) Leaves grey, pruinose below.

Fruit 8-10 x 4-9 mm. Leaves 10-30 x 6-16 mm.; inflorescence subumbellate; fruit 9-10 x 9 mm.,

Leaves 7-15 x 8-12 mm.; inflorescence subfascicled; fruit 8 x 4 mm., oblong. obovoid. 313. B. peudu?nbd&a

314. B . r o y h w Fruit 15 x 10-11 mm. 317. B.xnotabiEis

Leaves 1-2.5cm. long; pedicels 4-8mm.; ovules 2-3; fruit 7x4-5mm., con-

Leaves 3-6 cm.; pedicels 9-17 mm.; ovules 4-6; fruit 12 x 6-12 mm., epruinose

Leaves green, epruinose below.

spicuously pruinose. 315. B. aitchkmii

or m l y pruinose. 316. B.h&wopdu

Subsect. Creticae Schneid., BUU. Herb. Boksier, (2), 5, 657, 1905 S t e m glabrous; mature always dark red or purple. Internodes (7-) 10-20 mm. Spinerr

1-2 cm. Leaves deciduous, 14.5 cm., entire. In.ore.scena 3-8 (-20)-fld., fascicled, race- mose-subfascicled, umbellate or pseudumbellate, ( 0 . 6 ) 1-2 (-3) cm. Pedials 2-8 (-12) mm. Fbwers 4-8 mm. diameter. ovulee 2 (-3). Fruit black, sometimes pruinose, 56-8 mm. long, including style 0-5-1.5 mm.

(307) Berberis oblonga (Regel) Schneid., B d . Herb. B&sier, (2), 5, 458, 1905; B. Fedtachenko in Komarov, Fl. U.R.S.S., 7, 556, 1937; syn. B. heteropoda var. obknzga Regel, Acta Horti PetropoE. 5 , (i), 227, 1877

Shrub to 7 ft . , loosely branched, spreading. Stem terete to subterete ; young shoots green ; not or scarcely verruculose. Spines yellow-brown, 3-fld., stout, plane, sometimes with a basal stalk to the triple set. Leaves 3-6 x 1.3-2-3 cm., including petiole (6) 10-15 mm.; oblong-obovate; base contracted; entire, rarely a few marginal spinules; apex rounded; openly veined; dull pale grey-green; below pruinose, grey. Race- loose, 10-20-fld., 4-7 cm. Pedicels glabrous, 5-10 mm.; bracts 1-2 mm. Flowers H.C.C. 3. Prophylk absent. LYepal.8 in one series of three, 4.5 x 4 mm., broadly obovate. Petaks 5 x 3.5 mm., obovate, rounded, entire; gland8 orange, oblong, separate, non-marginal, 1.5 x 0-8 mm. 0vu.k-y 3, subsessile. Berries oblong, pruinose blue, 10-11 x 6-8 mm. Styk 1 mm.

!Curkestan: Tschatkal (Chotkola), 8000-10,000 ft., fr. 1876, Regel, 8.n. (Type, K). According to Fedtschenko, on the stony slopes of the mountains of Tian-shan and Pa&-Altai floristic regions of U.S.S.R.

Cultivated: L.A. 391, fl. aday 1940,3 June 1940; fr. Sept. 1942,7 Oct. 1942. The fruit does not properly ripen in cultivation.

Crete, Cyprus, Greece, Turkestan, Afghanistan, Baluchistan, Wazaristan.

Page 225: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 225

(308) Berberis cretica L., Sp. PI. 1, 331, 1753; Flura Gmeca, 14, 35, t. 342. (Fig. 51) Shrub 3-5 ft. Stem glabrous, stout, sulcate; mature lustrous. Spines discolorous

yellow to orange. Leaves 10-17 x 4-7 mm., obovate; base contracted, cuneate, sessile; entire; acute to subacute; both sides openly reticulate, with lateral veins distinct and veinlets more or leas indistinct; above deep green; below paler green. Infirescence 3-6-fld., fascicled, 6-8 (-12) mm. Pedicels 2-5 mm., in flower, P 7 mm. in fruit. Outer sepals 4 x 2.5 mm., elliptic. Inner s e e 4-4.5 x 3 mm., obovate. Pet& 44.75 x 3 mm., obovate, entire ; glands discolorous orange, oblanceolate, separate, submarginal. O d e d 2, with stipes subequal in length fo the ovule. Stumna 3-5-4.5 mm. B e r r h 6-7 x 4- 5 mm., ellipsoid, at first red, ultimately black, epruinose, or but very slightly pruinose. Style 0.5-0-75 mm.

Crete: Laasithi, 5000 ft., 19 May 1914, Gandoger 2340; White Mts., above 3000 ft., Omalo, 11 June 1938, Grant 29, and 20 June 1937, Lemperg 524 (all, K).

Cyprus: Troodos, 39005600 ft., 4 May, 24 June, 22 July and 6 Sept. 1937, Kennedy 450,451,452, 453 (K).

Greece: in Monte Olymp Thessalie 3,000 ft., 29 July 1927, Handel-Mazzetti s.n. (K). ‘Mature fruit dark blue.’ According to Loudon, in Arb. et F&. Brit. 1, 304, 1838, B. cretica is stoloniferous, has black fruit, and waa in cultivation in 1759. I have seen no cultivated specimen, and presume that it is no longer to be found in English gardens. B. cretica var. serratifolia Poir, ex Loud., 1.c. p. 305, 1838, is of doubtful identity.

(309) Berberis calliobotrys Aitch., J. Linn. Soc. (Bot.), 19,151,1882, nomen; Koehne, Deukche Dendrologie, 168, 1898; Schneid., Bull. Herb. Boissier (2), 5, 137, 661, 1905. (Fig. 49)

#terns sulcate. Spines 3-0d., plane. Leaves 8-35 x 4-12 mm.; oblong-obovate; attenuate to decurrent petiole, 1-6 mm.; entire, rarely 14-spinose (1 mm.)-serrulate; acute ; both sides distinctly reticulate, concolorous green, epapillose. Inflorescence 5-8-0d., racemose-subfascicled, 1-5-3 cm., including peduncle 1-10 mm. Pedicels 5-8 (-10) mm. Outer sepals 3-54x2-2-5 mm. Inner sepalzi 6x4.5 mm Petals 5 0 5 x 3 mm., obovate. Stamens 4 mm., not produced, truncate. Ovules 2. Berries black, conspicuously pruinose white, globose-obovoid, 7-8 x 5-6 mm., including style 1-1.5 mm.

Afghanistan: Kuram valley, between Sergal and Sikaram, 8000-10,000 ft., 4 June 1879, Aitchison 490, (Type, K); Kuram valley, Drekalla, a large shrub with peculiar swollen fruit, 17 Aug. 1879, Aitchison 176/4; Kuram valley, Habibkalla, 4 May 18i9, Aitchison 176; Aitchison Shalizan, fr. June 1879 s.n. (all, K).

Baluchistan (British): Gustoi, fr. 25 May 1897, Harsukh 20451; Zarghun, Aug. 1885, Lace, s.n. (all, K).

Kashmir: Batkoot, Pahlgam, 6500 ft., 7 May 1940, Ludlow t S h e d 7601 (BM). Oultivated: plants have been raised from no. 7601.

(310) Berberis wazaristanica Ahrendt, J. Asiat. SOC. Beng. (Sci.), (3), 11, 3, 1945 atems sulcate. Spines 3-fld., subconcolorous with the dark red stems. Leaves 10-

25 x 2-5 (-6) mm., oblanceolate; base attenuate; margins very entire; apex subacute or subobtuse. InJEorescence 5-8-fld., subfascicled, 1-1-5 cm. long. Style c . 1 mm.

Wazaristan: Pirghal, 8000-11,500ft., in immature fruit, 15 May 1895, Harsukh 15620 (Type, K). Not in cultivation.

(311) Berberis gambleana Ahrendt, l.c., 1945 &ems sulcate. Leaves 15-35 x 3-7 (-8) mm., oblanceolate; base attenuate, sessile;

margins entire; apex subacute; both sides openly veined with only veins and branched veinlets visible; both sides concolorous green. Inflorescence Pg-fld., racemose, 2-3 cm.

15 JOURN. LINN. SCC. BOTANY-VOL. LVII.

Page 226: Berberis and Mahonia : A taxonomic revision

226 L. W. A. ~ E N D T

long. P&i& 6-12mm. long. Berries black, with blue bloom, globose, 6mm., ex- cluding style c. 1 mm. long.

Baluchistan: Zarghun, fr. Aug. 1885, Gamble 1 (Types K).

Subsect. Pseudumbellatae Ahrendt, J. Bot., Lond. 80, 104, (Suppl.), 1943 Stem glabrous, generally dark red. Internodes (5 mm.-) 1-26 (4) cm. Spines mostly

1-2-5cm., occasionally w& and 1-9mm. Leaves deciduous, entire or with few in- oonspicuous spinules. Inflorescence 3-8 (-20)-fld., fascicled, racemose-subfaacicled, um- bellate or subumbellate, 1-26 cm. long. Ped& P 1 6 mm., rarely to 20 mm. Fknuers 6-8 mm. diameter. Ovules 1-2 (-6). Fmit, black, pruinose, estylose.

!I’urkeatan, Afghanistan, h h m i r , Punjab.

(312) Berberis stolonifera Koehne et Wolf, F a Rep. Xp. Nov. 12, 129, May 1913 Shrub stolonifemus. Stem mature, sublustrous pale yellow-brown. Spines 1 (-3)-fld.,

very slender, 7-13 mm. Leaves with lamina 2-56 x 1-2.3 cm., excluding petiole 8 mm. to 4.5 cm. long; oblong-obovate; base cunectte; entire or with few inconspicuous marginal spinulea; only lateral veins visible, but these prominent both sides; above bright green; below grey, pruinose, very closely, broadly papillose. Racemes open, 10-20-fld., 6-9 cm. long, including peduncle Pedkek 10-15 mm., occasionally to 2 cm., a t base of the inflorescence; bracts 1-5-2mm. Inner 8epa-h 5mm. Petals 4mm. Stamens 3mm. Ovules 3, stipitate with stalk as long aa the o d e . Berries 9 x 5-6 mm., ovoid, pruinose Pulplea

Turkestan: the authors cite no locality and no specimens. Not in cultivation.

(313) Berberis pseudumbellata Parker, Kew BUU. 1921,118,1921 ; syn. ‘B . unzbellata’ Parker, Fwat Fl. Punjabwith Hwraand Delhi, ed 2,15,1924; syn. ‘B. unzbeua2Q’ Parker, Fore& F1. Punjab with Hamra and Delhi, 15, 1918, non Wall. ex Don.

Stem subterete, angled or finely sulcate, slender; mature dark red to red-brown. Spines mostly solitary, 1-8 (-15) mm. Leaves 10-25 (-30) x 6-15 mm.; oblong-obovate; base contracted to petiole 5-1Omm.; entire, rarely with 1-5 inconspicuous spinules; apex rounded; both sides with lateral veins and a few branching veinlets visible, indis- tinctly above, elevated below; above dull light grey-green; below grey, pruinose, finely greenish grey. Inflorescence 3-5 (-7)-fld., umbellate to subumbellate, 1-5-26 cm., occasionally to 5-6 cm., including peduncle 8-10 mm. P&i& stout, 6-9 (-15) mm.; bracts 1 mm. ProphyUs yellow, oblong-ovate, 1.5 x 1 mm. Outer sepals 2.75 x 1.75 mm., ovate. Inner sepals 5 x 3 mm., obovate. Petals longer, 5-6 x 3 mm.; clawed, with approximate submarginal glands within the claw. Stamens 4 mm., not produced, trun- cate. Ovules 3. Berries hard, pithy, black, conspicuously pruinose blue or purple, estylose, obovoid-subglobose, 8-10 x 6-9 mm.

Punjab: Chamba State, Ravi valley, Ulansa, fr. 7300 ft.; 17 Bug. 1920, Parker, s.n. (Type); fl. 3 May 1920, 7100 ft., Parker, 8.n.

Kashmir: 19 Apr. 1890, Hunter-Weston 10158 (K); Nilang Suss, 9500-10,OOoft., Schlich 27 (5) (0); Upper Bringhi valley, 6500 ft., fr. 17 Oct. 1940, Ludlow & S h e d 8271 (BM), ‘shrub 5-7 ft.; fruit dark purple, covered with bloom, on open stony hill- sides’; Kagan, Sind valley, 6000ft., fr. 8 Nov. 1940, Ludlow & SherriE 8291 (BM), ‘leaves turning dark red; open stony ground’; Gauhan, Bringhi valley, 26 May 1940, Ludlow & S h e d 7616 (BM), ‘river bank’; Kufwara, Lolab valley, 5500 ft., fr. 15 Oct. 1940, Ludlow & Sherriff 8269 (BM).

According to Parker (1s. 1915, as umbellata), in the inner dry valleys (4000-9000 ft.) of Kagan (Jar6d to Nahan), and K u n a m (common in the Baspa valley).

(I%. 54.)

Page 227: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 227 Cultivated: from Ludlow & SherrX 8291, fl. 31 May 1951, at the University Botanic

Garden, Oxford; in cultivation also from numbers 8271, 8269.

(314) Berberis royleana Ahrendt, J. Asiat. Soc. Beng. (Sci.), (3), 11, 5, 1945 rs’tenzs more or less sulcate. Spines 3-fld., more or less concolorous red with the stems,

stout. Leaves 7-15 x 6-12 mm., including petiole 1-2 mm. ; broadly oblong-obovate: base contracted; margin entire; apex rounded; both sides with branched venation. In,oresmnce 3-8-fl., fascicled, or umbellate-subfascicled, 1-14 cm. long, subsessile, or with peduncle 1-3 mm. Pedicels 5-10 mm. Berries pruinose grey, immature, 8 x 3.5 mm., estylose.

Kashmir: Srinagar, Tahkt-i-Suliman, 5000-8000ft., 7 May 1892, Duthie 10816

(315) Berberis aitchisonii Ahrendt, 1.c. (3), 11, 4, 1945; syn. B. crutaegim var. uzbulim Schneid., Bull. Herb. Boissier, (Z), 5, 657, 1905; syn. B. densi$ora Wen- delbo in Nytt. Bug. Bot. 1, 80, 1952

~ Y t e w u dark red, sulcate. Leaves 10-25x4-10mm., excluding petiole 1 4 m m . ; obovate ; attenuate; entire ; acute, mucronulate; both sides concolorous green with open elevated reticulation. Inflorescence (2-) 4-43 (-8)-fld., fascicled or racemose-subfascicled, 1.2-2 cm. long, or racemose and 2-3 cm. Pedicels (4) 5-8 mm.; bracts 1.5-2 mm. Ovules (I-) 2-3. Berries black conspicuously pruinose grey-white, estylose, or very short and indistinct style, 6-7 x p4.5 mm.

Afghanistan: Kuram valley, near Sergal, 900&1l,OOOft., 10 Aug. 1879, Aitchison 176/5 (Type, K); hills north of Kaiwas, 1O,o00-11,OOO ft., 3 July 1879, Aitchison 176/3 (K); Kabul, Konigsberger, s.n. (Type of Schneider’s variety cabulica, in Herb. Vienna). Hirtdu Kush Mts., Tirich Mir, Shokor Shal, C. 11,400 ft., 6 July 1950, Wendebo.

(316) Berberis heteropoda Schrenk., Enurn. PI. Nm. 1,102,1841 ; B. Fedtschenko in

Shrub 6-7ft. Stems finely sulcate, ultimately subterete, dark red. Spines 3-fld., slightly paler, 5-10 mm., plane to subterete. Leaves 2-4 (-6) x 1-2.7 (4) om., excluding petiole 3-10 mm. ; obovate-elliptic ; base contracted; maxgins entire, or indistinctly spinose ; both sides reticulate, subconcolorous, or below scarcely paler and lustrous. InJorescence 4-9-fld., fascicled or umbellate, 1-5-3 (-5) cm. Pedicels 9-17 mm.; bracts 1-5-3 mm. Outer sepals 5 x 4 mm., elliptic, rounded. Inner sepals 7 x 5 mm., obovate. Petals 6 x 4 mm., obovate-spathulate, entire, cuneate. Stamens 4-5 mm., produced, apioulate. Ovules 4-5(-6), stipitate. Berries black, slightly pruinose blue, estylose.

According to Fedtschenko, B. heteropoda is to be found in Dzungaria, Tarbagatai, and is generally distributed from Mongolia to the Southern Altai, and Kashgaria (Kuldja).

(Twe , K.)

Komarov, PI. U.R.X.S. 7,555, 1937

KEY 57A Flvj t oblong. Fruit globose.

316a. B. heteropoda var. heteropoda 316 b. B. heteropoda v&r. sphaerocarpa

( 3 1 6 ~ ) Berberis heteropoda var. heteropoda Sxnkiang: Dzungaria, Schrenk, 8.11. (Type, K); fl., Krassnov., s.n. (K). As far as I am

aware this is not in cultivation; the few plants I have seen bearing this name are B. obloqa, a Turkestan species with exceptional flower structure and a long style.

(316 b) Berberis heteropoda var. sphaerocarpa (Kar. et Kir.) Ahrendt, stat.nov. ; syn. B. sphaerocarpa Kar. et Kir., Bdl. Soc. Nat. MOSC. 15, 376, 1842

Sinkiang: Alatau Mts., Karelin & Kiriloff 1176 (Type, K); Schrenk, 8.n. (0). 15-2

Page 228: Berberis and Mahonia : A taxonomic revision

228 L. W. A. AHRENDT

(317) Berberis x notabilis Schneid. in Silva Tarouca et Schneid., Umere Freiland-

8 h d fairly upright, 6-8 ft. Stems glabrous, subterete to h e l y sdmte; young shoots green, tinged red; mature yellow, or grey-brown. Internodes 1 - 5 4 cm. S p i w 1-3-fld., sulcate, 5-15mm. Learn 2 - 5 5 x 1.3-2-2 cm., obovatetoelliptic; basecontractedh petiole 6-15 mm. ; margins indistinctly 15-24-spinose (0.5 mm.)-aerrulate; apex rounded, mucro- nulate; dull grey-green and openly veined above; grey, pruinose and subreticulate below. Injeorescence 14-2O-fld., 4-6 cm., racemose, sessile. P e d W 10-15 mm.; bracts 2 mm. Flowers golden yellow. ProphyU.8 oblong-ovate. Outer sepals similar, 3.25 mm. Inner sepal8 6 mm., oblong-obovate. P& 6 mm., obovate. Stanzens 4-4.5 mm., scaxcely produced. oerules 3 4 , stipitate, with stalks as long aa the ovule. Berries ovoid to ellipsoid, dark red, pruinose, estylose, 15 x 10-11 mm. S d purple.

Cultivated: origin in U.S.A., from seed labelled B. ~ r o p o d a , about 1895 (Type in Herb. Am. Arb.). Not cultivated in England.

Laubg., ed. II, 16,1922; J. Am. Arb. 4,203,1923

Sect. IJUcinae Schneid., Bull. Herb. Boissier, (2), 5, 4-00, 1905 Shrub8 1-3 ft. L3tew.w glabrous, mature dark purple. Internodes 5-10 (-12) mm.

Spines 1-2cm., discolorous, yellow, plane or subsulcate. Leaves deciduous, narrow, 1 : 4-7, subenervate; below concolorous green, epruinose. InJloreseence 2-7-fld., 4-15 mm. long. Pedkeh 2-5 mm. Be& black, epruinose, or scarcely pruinose, estylose, 4-8 mm.

N.E. h h m i r , W. Tibet, Sinkiang.

KEY 58, to Sect. Ulicinae (Maps 3,40 and 41) Leaves m o w , 1 : 6-7; inflorescence 6 9 mm. long; pedicek 3-6 nun.; fruit 4-6 mm.

haves less -ow, 1 : 4-6; inflorescence 10-15mm. long; pediceh 2-3mm.; fruit 318. B.ulicim

8 mm. 319. B. kaahgarka

(318) Berberis ulicina Hook. f. et Thorns., FZ. Id. 227, 1855. (Fig. 55) Internodes 5-8 mm. Leaves 7-15 x 1-2.5 mm., linear-lanceolate; margins with 1-2

spinules. I n . e s c e w 3-6-fld., fascicled, or fascicled-subumbellate. Inner sepals 5 x 3 mm. P& 4 x 2 mm., emaxginate. Xtanzens 3 mm. O h 3-5. Berries globose.

W. Tibet: adjacent to Nubra, 14,000-16,000 ft., dry rocks, 1849, Hooker, s.n. (Type, K). Kashmir: Ladakh, Sapur, 13,000-16,000 ft., Thornson, s.n.; Kardong, fl. 13,500 ft.,

23 June 1929, Ludlow 506 (BM); Durgu, foot of Chang La, 12,700 ft., 18 Aug. 1941, Ludlow & Sherriff 8563 (BM), ‘shrub 2-3 ft.; habitat, edges of cultivation; fruit indigo blue, with very slight blue bloom ’; the only Berberis seen in Ladakh. I do not know if this species is in cultivation. I raised one tiny plant from no. 8563, but this died about 1947.

(319) Berberis kaschgarica Rupr., Xert. Tianschan, 38, 1869; Maxim., PI. Tangut. 31, t. 23, figs. 1-11 1889; B. Fedtschenko in Komarov, Fl. U.R.S.X. 7,555,1937

Intemzodes 8-12 mm. hve.s 10-20 (-30) x 2 4 (-7) mm. ; oblanceolate; cuneate, ses- sile; entire or with 1-3 marginal spinules (1-2mm.), distant 3-6mm. Inflorescence 2-7-fld., racemose-subfascicled. Inner sepals 4-5 mm. Petah 4.5 mm., emarginate. Stamens 2.5 mm. Ovules c. 4. Berries ovoid, 8x 6 mm.

Sinkiang: Tian-shan, Ili Fluss, Krassnov, s.n., 1866; Yarkand, 8 Aug. 1870, Hen- derson 327, and 30 Sept. 1870, Henderson 742 (all, K).

According to Fedtachenko, generally found in the district Junghu, Kashgaria, in particular in the valley of the Sungtu, 50-60 milea north of Kashgar. Not in cultivation.

Page 229: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 229

Group Australes Schneid., Bull. Herb. Boissier, (2), 8, 264, 1908; sectionibus (Actin- amnthae, Microphylkce, Corymbosae, Buxijoliae, Montancce, Ilicijoliae, Laurinnze, Latifoliae, Virgatae, Conjertae, Agapaknses, Quindiuewes, Goudotiae, Truxillenses, Panicuhtue) ut in clavibus 59-91, et in descriptionibus 320497, circumscriptis ; a Septentrionalibus ut in clavi 1 distinguenda

~!tems variable. Spines often absent, manifold, palmate, foliaceous, otherwise weak, rarely strong. Leaves nearly always evergreen, chartaceous, or coriaceous ; rarely de- ciduous. InJlorescence solitary, fascicled, umbellate, pseudracemose-subumbellate, race- mose or paniculate. Pedicek 1 mm. to 5 cm. Bracts almost invariably shorter than pedicels sometimes foliaceous. Flowers often golden yellow to reddish orange. ProphyuS present or absent. Sepals in from (1-) 2-3 (4) series. Petals shorter or longer than inner sepals; apex entire or emarginate; base cuneate or clawed, with glands indistinct or conspicuous. Stamens shorter than petals ; below the anthers sometimes dentate, more often edentate; apex not produced or produced, and then truncate, obtuse or apiculate. Ovules 1-12. Berries as far as is known, always black; mostly, but not always, pruinose. Style often insignificant, i.e. absent, or short, up to 0.5 mm.; often significant, (1-) 1.5- 5 mm. long.

Habitat: South America, and Central America, includmg Falkland Islands and Juan Feimindez.

Subgroup Euaustrales Ahrendt, foliis sempervirentibus et rigide chartaceis vel plus- minusve crasse coriaceis, inflorescentiis paucifloratis et pistillis estylosis, vel inflorescentiis multifloratis et pistillis conspicue stylosis vel foliis deciduis, inflores- centiis paucifloratis et pistillis conspicue stylais.

South and Central Chile, South and Central Argentine, Juan Fernhdez Islands, Falkland Islands, Uruguay, Brazil. ( Z O O - ) 30-56" S. latitude.

KEY 59, to Euaustrales Style absent, or short and insignificant (scarcely 0.4 mm. long).

Leaves chartaceous; spines foliaceous, or manifold, occasionally 3-fld., but then almost

Leaves not chartaceous, but thin, or coriaceous; spines solitary or 3-fld., or absent. invariably palmate. Sect. ACTINACANTHAE

Ovules 2. Sect. MICROPHYLLAE Ovules 3-12.

O v u l s 3-5 ; leaves mostly deciduous ; inflorescence peduncled, subumbellate.

Ovules 612; leaves more or less evergreen; idorescence mostly solitary or Sect. CORYBTBOSAE

fascicled, rarely elongated. Sect. BUXIFOLIAE Style significant, conspicuous, 1-5 mm. long.

Leaves thin, papyraceous, more or less deciduous; flowers mostly solitary or fascicled, inflorescence rarely elongated. Sect. MONTANAE

Leaves thick, coriaceous, evergreen ; inflorescence mostly umbellate to racemose, sometimes fascicled.

Leaves stiff or rigid; stems mostly terete, often dark or pubescent, sometimes yellow, rarely sulcate ; inflorescence nearly always umbellate to racemose.

Leaves more flexible; stems very sulcate, glabrous, pale yellow ; inflorescence often Sect. ILICIFOLIAE

fascicled. Sect. TRIGONAE

Sect. Actinacanthae Schneid., Bull. Eerb. Boissier, (2), 5 , 143, 1905 &'pines 3-11-fld., 5-10 (-20) mm. long, mostly palmate or foliaceous. Leaves evergreen,

nearly always s t a y chartaceous, rarely more thinly papyraceous, mostly more or less coarsely dentate, with 1-7 (-22) teeth to each margin. In$oresceme only occasionally

Page 230: Berberis and Mahonia : A taxonomic revision

230 L. W. A. ~ E N D T

solitary, mostly 2-20-fld. Stanaens often dentate. ovules (2-) 3-5 (-10). Fruit black, mostly pruinw blue, estylose or with insigniscant style scarcely 0.4 mm. long.

Chile, Argentine, 30°-66” S. latitude.

KEY 60, to Subsects. of Sect. Acdnacanthae Spines (3-) manifold, mostly palmate, not foliaceous. Subseot. EUACTRTACANTHAE spines foliac0oue. Subsect. CONOEBTIFLORAE

Subsect. Euactinacanthae Schneid., I.c., 1905 KEY 61, to Subsect. EUACTINAOANTFUE

Leaves entire, or with 1-7 m8rgina.l teeth; stiffly or thickly C-UE. Infforeecence 6-8 (-20)-fld.; pi nee 3-fld., palmate, 3-8 mm.; stamem dentate; fruit

ovoid, estylose. Leaves dentate, elliptic. 320. B. brachyantha Leaves entire, elliptic to suborbicular. 321. B.fEorida

Inflorescenoe 2-6-fld. Inflorescence (I-) 2-6-d. ; spines 3-5 (-7)-fld.

Leaves margined white, paler below. Leaves uniformly green above.

322. B. nZarginata

Stamens dentate; stems yellow; leaves green below; flowem fascicled; pedicels 7-10 IIM.

h v e s dentate. Leavea spathulate-cuneate; margins with 1 spinule; spines 3-fld.

323. B. brevifoliac Leaves oblong, orbicular, or elliptic; maxgins with 3-4 qhules; Style absent.

Spines 1-2 cm. long, 3-fld., slightly palmate; leaves suborbicular, 8888ile,

Spinea 7-11 mm. long, 3-7-fld.. very palmate; laves oblong, with petiole articulate below the blade. 324. B. g r e w d h ~

16-2-5 cm., articulate above the blade. 325. B. actinacanthu Leaves entire; style short. 326. B. phiJippii*

Leaves green be1ow.t Stamem dentate.

Leaves of two kin&, concolorous below; qines absentor palmate.

Leaves uniform, paler below; spines subterete, scarcely palmate. 327. B.p lymnpha

328. B. gknwrata 329. B. bidepltata Leaves white or grey below, pruinom.

Flowers solitary. Stamens dentate; petah shorter than inner sepals.

Flowers 7-8 mm. diameter. haves EpklOSe; p”9Wl” C. 5 m. 330. B. antwoam Leaves entire; pe&cel” 1&20 mm. 331. B. x gwmeyensia

Flowers 12 mm. diameter. Cf. 497. B. b~loche& Stamens dentate; petals subeqd to inner sepal”. 332. B. heterophylla

Leaves with 18-22 epinoSe teeth, thinly papymceous. 333. B.coquirnbe&$ Imperfectly known; not able to be placed in key. 334. B. lnorenonis

* B. ph-i haa sulcate yellowish grey atems and short style; if the stems are red and temte, the style absent, and the spinea somewhat palmate, refer to B. rotund$oZia.

t Compare B. ilicifolia which has long 3-5-fld. spines, but large flowere with long pdceh. large leaves and many ovules, all of which charecters distinguish it from B. gbmerafu. Compare also B. trigmu with somewhat palmate spinea but which is distinguished by ite long narrow, entire leaves. In addition, B. rotum&f&, B. ilicifolia and B. t r i g m are all distinct in their coriaOeoua leaf character. Clearly they properly belong to the sections in which they have bwn p M , but are connected to the Actinacanthee by their tendency towards palmete spines.

3 B. pear& should also be compared with B. coqwknbeneis. The former haa coriaceous leaves and conspicuous style (1-6mm.); while the letter has thinly papyrsceons leaves and style absent or ineignificsnt.

Page 231: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 231

(320) Berberis brachyacantha Phil. ex Reiche, Fl. Chile, 1,42, 1895 Chile: Prov. Talca, Constituci6n.

(321) Berberis florida Phil., Linnaea, 33, 5, 1864; Reiche, PI. Chile, 1, 40, 1895 Imves green below. Pedicels 4-5 mm. Chile: Prov. Colchagua, according to Reiche, Cord. de Colchagua, 6700 ft., flowering

in December.

(322) Berberis marginata C. Gay, Fl. Chil., 1, 88, 1845; Reiche, Fl. Chile, 1,42, 1895 Spines 3-7-fld., palmate. Stems terete, glabrous. h v e a with 3-7 marginal spinules. Chile: Prov. Valdivia, according to Lechler, Berb. Amer. Austr. 37, 1857, environs of

Lake Llanquihub (Yanquigue).

(323) Berberis brevifolia Phil. ex Reiche, Fl. Chile, 1, 42, 1895 & e m sulcate. Spines 3-fld., palmate. Pediceh 7-10 mm. Chile: Prov. Colchagua, according to Reiche, Cahuil. The preceding four species are but little known, from slender descriptions without

material. However, sufficient information is given to place them in the key, which contains most of the known facts about them. This being so, it seems proper to preserve and place the names so that, when further material is forthcoming, they may not be renamed accidentally.

(324) Berberis grevilleana Gillies ex Hook., Bot. Misc. 3, 136, 1833; syn. B. actin-

&ems sulcate. Spines 3-fld., with the lateral parts shorter than the middle, only slightly palmate and thickened at the base. Leaves rhombo-elliptic, 10-20 x 5-7 mm. (according to Philippi), or 8-20 x 5-13 mm. (according to Schneider), reticulate, sessile. Berries ovoid, estylose.

Chile: Prov. Santiago, according to Reiche and Lechler. Prov. ValparaiSo, according to Schneider. Prov. Colchagua, according to Reiche, who records that the species flowers in November.

(3261) Berberis actinacantha Mart. in Roem. et Schult., Syst. Veg. 7, i, 12, 1829;

Shrub M f t . , with spreading arching branches. Stem slightly sulcate, glabrous; mature yellow. Spines 3-7-fld., green, palmate, 5-12 mm. h u e s oblong, 10-25 x 5- 10 nim., contracted to petiole 15-25 mm.; margins 3-4-spinose (1-2 mm.)-dentate (2-3 mm.); above dull grey-green; below dull, greenish; both sides openly reticulate. Flowers fascicled 2-6. Pedicels glabrous, 7-10 mm.; bracts 3 x 1 mm. ProphyyuS 3 x I -25 mm., ovate, subacute. Outer sepals 3.25 x 1.5 mm., ovate, acute. Median sep& 4 x 3.25 mm. Inner sepals 5-6 x 4-4.5 mm., obovate. Petals 4-45 x 2.5 mm., oblong-ob- 0vate;subacute to obtuse,entire;basecuneatewithin&stinct glands, 1 x 0-5mm. Stamen8 dentate, truncate. Ovules 5-7. Berries black, pruinose blue, globose, estylose, 6-7 mm.

Chile: baths of Collina, Macrae (Type) ; Prov. Santiago, Cord. de 10s h a n o s , 1853-61, Philippi, s.n. Prov. Valparaiso, according to Edwards, common near Valparaiso, not a plant of the coast, but inhabits the first range of the Cordilleras. According to Hooker in Bot. Misc., Valparaiso, Bridges, s.n. ; Cuming 434. According to Lechler, common in the Central Provinces, flowering November.

Cultivated: L.A. 53, fl. 5 May 1944; fr. 3 Aug. 1944. A good garden plant, with its restricted habit, distinct leaves, and fine display of orange coloured flowers.

Reiche, PI. Chile, 1,42,1895, refers to a variety mollis, from Rancagua, San Fernando, with leaves softer, and more closely veined below.

mntha var. grevilhnu (Gill.) Schneid., BUU. Herb. Boissier, (2), 5, 145, 1905

Argentine: Prov. Mendoza, valleys of the Andes, according to Hooker.

Edwards, Bot. Reg. 31 (N.S. 17), t. 55, 1845. (Fq. 56)

Page 232: Berberis and Mahonia : A taxonomic revision

232 L. W. A. AERENDT

(326) Berberis philippii Ahrendt, nom.nov.; syn. B. u- Phil. Anal. Univ. Santiago, 1872, 663, fide Reiche, Fl. Chile, 1, 34, 1895, non Wall. ex Don; syn. B. lwterophy1l.u var. plwa@ra Reiche, I.c. 43, 1895.

Stems grey, slightly sulcate. S p * w more or less sulcate, 5-7-fld., very broadly pal- mate. Leaves enthe. In$me-smnce c. 4-fld., umbellate. Stumem dentate. Bemies globose, with short style.

Chile: Prov. Valdivia, Val del Rio Bueno; Cord. de Cauquenes. Not cultivated.

(327) Berberis polymorpha Phil,, Linnaea, 28: 664, 1856; Reiche, F1. Chile, 1, 35,

Stems some with 3-5-fld., palmate spines, and membranaceous leaves, quite entire; others with no spines, and spinose coriaceous leaves; all the leaves concolorous below. Inflorapcence 44-fld., subracemose, with fairly long peduncle. F k r s c. 6 mm. diameter. Shmm edentate. Style absent.

Chile: Prov. Ruble, Chilltin, Dec. 1855, Germain, s.n. (Type). According to Philippi, CordiIlera near town of Chillhn, flowering December. Not in cultivation.

(328) Berberis glomerata Hook. et Am., Bot. of Beech. Voy. 5, 5 , 1830; Reiche,

Stems puberulous; mature dark red. Internodes 4-8mm. S p ' w 5-fld., often sub- terete, less often palmate, 6-10 mm. long, red-brown. h u m rhombic-oblanceolate (or spathulate-obovate), 5-10 x 2-4 mm. (or to 15 x 5 mm.); cuneate, sessile; apex mucro- nate; margins 1-2-spinose (1 mm.)-dentate (1.5 mm.); subenervate; above greygreen; below slightly Instroue paler green, epapillose.

1895

Fl. Chile, 1,34, 1895

GY 61a I m v m with 1 spinde to each margin Leaves with 2 spinule to each margin

(328a) Berberis glomerata var. glomerata InfEorescence 3-5-fld., umbellate or subumbellate, 8-12 mm., including peduncle

4-6 mm. Pedicel% glabrous, 1-24 mm.; bracts subequal 1-2 mm. Flowers 3-4 mm. diameter. Outer 8epd.s 2 x 1.5 mm., ovate, shortly acuminate. Inner sepals 2-75-2 mm., broadly obovate. Petals 2 x 1.25 mm., obovate. Shmm 1.5 mm.; below the anthers conspicuously lanceolate-acuminate-dentate ; apex scarcely produced, rounded-truncate. Owles 2. Berrks with pedicels 2-4 mm.; c. 6.5 x 4.5 mm., oblong, pruinose blue. Style 0-25-0.4 mm.

Chile: Prov. Coquimbo, IUapel, Dec. 1862, ex Herb. Reed, fr., and fl. ex Herb. Hook., 1867 (both K). According to Reiche in Prov. Coquimbo (La Serena, Arquero, Anda- collo), Rancagua, flowering Oct. According to Hooker, Prov. Coquimbo. Not in cultivation.

328a. B. g b n w ' a t a var. glomerata 328b. B. glomerata var. &bknsriarrcl

(328b) Berberis glomerata var. zahlbruckneriana (Schneid.) Ahrendt, statmov. ;

Shrub 3-4ft. Stenas scarcely angled, slightly puberulous, red-brown. Spines dense, 1-3-5-fld., brown, sulcate. Leaves spathulate-obovate, to 10-15 x 5 mm. Infore-smme 3-5 (-7)-fld., racemose-subfascicled. Pedkds 4 mm.; bracts 2 mm. Berries 5-6 x 4- 5 mm., shortly stylose.

Chile: Prov. Coquimbo, Illapel, Philippi, 8.n. (Type); Oct. 1929, Clarence Elliott 526 (K). Not in cultivation.

syn. B. zuh&ruck&m Schneid., BuU. Herb. Boiseier, 8er. (2), 5: 145 (1905)

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Berberis and Mahonia 233

(329) Berberis bidentata Lechler, Berb. Amer. Azcstr. 11, 1857; Reiche, Fl. Chile, 1, 36,1895

Chile: Prov. Valdivia, Cord. de Ranco, according to Lechler. Not cultivated.

(330) Berberis antucoana Schneid., Bull. Herb. Boissier, (2), 5, 144, 1905 Stems glabrous, subangled; mature yellow. Internodes c. 1 cm. Spines 5-fld., yellow,

sulcate, 5-12 mm. Leaves 8-20 x 6-13 mm. ; oblong-rhombic; base rounded or truncate; margins with 1-3-spinose teeth. Flowers solitary, c. 7 mm. diameter. Pedicels c . 6 mm. Petals shorter than inner sepals. Stamens edentate, produced.

Chile: Prov. Antuco, Cord. de Antuco, 1828 (Type) ; Sierra Velluda, 1829, Philippi, s.n. Not in cultivation.

(331) Berberis x guernseyensis Ahrendt, hybr.nov. (B. actinacantha x ? B. buxi- folia), spinis palmatis, f o b chartaceis, floribus parvis u t in B. actinacantha; ramulis puberulis, foliis coriaceis integris, floribus solitariis, staminibus edentatis ut in B. buxifolia

Stems sulcate, puberulous, red-brown. Spines yellow, palmate, sulcate, to 15 mm. long, 2.5 mm. broad at base. Leaves with hypoderm, to 20-25 x 9 mm., subsessile, entire, acute, mucronate (1-2 mm.) ; above slightly lustrous; below dull, subconcolorous; both sides with but few veins visible, and slightly elevated. Flowers nearly always solitary. Pedicels 1-2 cm. Outer sepals 4 x 2.75 mm. Median and inner sepals equal, 5 x 4 mm., obovate. Petals 3.5 x 2-25 mm., obovate; cuneate, truncate at base; entire. S t a ~ n s 3 mm., edentate; not produced, truncate.

Cultivated: a t Wisley, fl. 1938. The plant had, it seems, been received from the Caledonia Nursery of C. Smith, a t Guernsey, under the name B. actinacantha, but I have very little doubt that it is a hybrid of the origin stated. (497) B. barilochensis may also be a natural hybrid between Sects. Actinacanthae and Buxifoliae.

(332) Berberis heterophylla Juss. ex Poir., Encycl. Mdth. Bot. 8, 622, 1809; Hook., Exotic F b a , 1, t. 14, 1823; Job, Rev. Mus. La Plata, N.S., Sect. Bot., 5, 46, fig. 10, 1942; syn. B. tricuspidata Smith ex DC. Syet. Veg. 2 , 17,1821, pro syn.

A‘hrub 3-4 ft. Stems angled, dark brown. Spines 3-fld., stout, 10-15 mm. Internodes 1-2 cm. Leaves recorded as of two kinds: (a) young, pale green, entire, soft; ( b ) older, dark, lustrous green, rigid, aristate, tridentate, the margins having each one spinose tooth towards the apex about one-third of the way down from the apex; all the leaves obovate, cuneate, slightly veined, to 25 x 10 mm. F k r s solitary. Pedicels 7-15 mm. Ouier sepals 3 x 2.5 mm., ovate, obtuse. Inner sepals 5-6 x 3.54.5 mm., ovate, obtuse. Petals 4.5-6 x 3-5 mm., orange, obovate; apex rounded, but slightly crenulate, other- wise entire; base truncate with separate non-marginal glands, oblong, 0.8-1.2 x 0.2- 0.4 mm. Stamens c . 4 mm., below the anthers apiculate-dentate (1 mm.); apex not produced, retuse-truncate. Ovules 8-10. Berries pruinose blue, 6 mm., globose, estylose.

Chile: Prov. Magellan, according to Reiche, Cord. de Aculco. Argentine: according to Job, from 41” lat. to Magellan Straits. Cultivated: from seed collected (Magellan) and raised by Mr Collingwood Ingram a t

Berienden Grange, Kent ; my plant, from this distribution, has not flowered. Hooker queries the identity of the plant he figures, which is described above, with

‘the true heterophylla’, which ‘is stated to be 1-seeded’. However, his plant seems to be the same as that figured by Job in ‘Los Berberis de la regi6n de Nahuel Huapi’, cited above, in which fifteen Argentinian species are provided with most attractive and clear illustrative figures. As suggested in the introduction, it is the number of ovules which

Page 234: Berberis and Mahonia : A taxonomic revision

234 L. W. A. AHRENDT the plant produces which is relevant, not the proportion of these which happen to mature into seeds under varying circumstances. Job gives '4-7 seeds', which is not inconsistent with the o d e s being as stated. Loudon in Arb. et Frut. Brit. 1,305,1838, says that this species is 'the ilicifolia of English gadens', and, according to Lindley, it 'has nothing but its raxity to recommend it '. With regard to the two kinds of leaves, it may be remarked that, generally, it is usual for Berberk with stiff, coriaceous or char- taceous leaves, to have the new leaves soft and paler green (or red). I have quoted the old description because this seem to have inspired the specific name; but I doubt whether this relates to a specific chaxacter. The plant in cultivation is semi-evergreen, or in cold spells almost deciduous. Numbers 330-332 are intereating because, with their solitary flowers, they tend t o w d the Sect. Buxifoliae. B. antucoana is a first link in this direction. B. x gwmey& is a gaxden hybrid between the two sections; and B. hetero- phyUa, with its non-pahate (albeit stout) 3-fld. spines, and many o d e s lies still farther towards the Sect. Buxifoliae. But the texture of the leama suggests that it should be retained within the Actinacanthae. Still further in this direction, and possibly best placed within Sect. Buxifoh, lies B. bariloche&.

(333) Berberis coquimbensis P. Muiioz, A@. Tecn., Ch&, 8, 79, 1948; BoZ. SOC.

Shrub some 3-4 ft. high. S p k e . ~ 611-fld., rigid, up to 1 cm. long, revolute, mostly 5-8 x 2-6 mm. Leaves ch-us, ovate, glabrous, papillose, 10-25 x 7-11 mm. ; base attenuate to a petiole 1-5 mm.; margins with 18-22 spinules. InjIormcem ClO-fld., fascicled, or subumbellate with a peduncle 0-5-2 cm. long, and 0.4 mm. thick. Pedi& pubescent; bmts 1 mm. Outer se@ small. Inner sep& 4 - 5 4 x 3-4 mm. Petals 3-4.5 x 2.5-3 mm., with basal glands, 1 . 5 ~ 0 - 5 mm. Stamem 2 mm., dentate; apex truncate. Berries subglobose, 5 x 4 mm.

Chile: h v . Coquimbo, Department de Ovalle, Bosque de Fray Jorge, 27 Sept. 1942 (Type-S.G.O.57575); same locrtlity, 1350-1700 ft., 26 Sept. 1935, Muiioz 1371.

Argent. Bot., 3, 117, 1950

(334) Berberis morenonis Kuntze, Rev. CJen. 3, (ii), 3, 1898 * Stem glabrous. Spins 3-5-fld. Leaves oblong-ovate; at base of stems remotely

spinose-dentate, with a 2-4cm. long petiole; towards the apex, thinner, scarcely coriaceous; margins 2-3-spinose.

Argentine: Provinces Santa Cruz, Patagonia, 50'-53" S. latitude, Moreno 768 (Type).

Subsect. Congestitlorae Schneid., BUU. Herb. Boissier, (2), 5, 146, 1905 Chile: Provinw Vdparaiso, Colchagua, Made, Malleco, Cautfn, Valdivia, 33'-40" S.

latitude. KEY 62, to Subsect. Congestitlorae

h v e s below densely pilose-pubescent ; flowers solitaxy ; fruit oblong.

Leaves below glabrous; intloreacence 620-fld. ; fruit globose. 335. B. pilosifolia

Leaves enervate or subenervate; ovules 1-2. 336. B. varii&wa Leaves reticulate; ovules 3-5.

Stems pubescent; leaves thinly chastaceous, without hypoderm; pedicels 1.6-3 mm. Leaves 4 4 mm., papillose below; inflorescence 6-10 mm.; peduncles 3-6 mm.;

Leaves 1630 mm., epapillose below; inflorescence 3-4 cm.; peduncles 2-3 cm.; flower diameter 3 mm. 337. B. cr&pa

flower diameter 6 mm. 338. B. congeetiflora

Leaves hely , closely reticulate, orbicular. 339. B. hakeoidee Leaves coarsely, openly reticulate, elliptic. 340. B.ho?rido

Stems glabrous; leaves tbickly chartaceous, with hypoderm; pedicels P 6 mm.

Page 235: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 235

(335) Berberis pilosifolia Ahrendt, sp.nov., spinis foliaceis, foliis subtus pubes-

Shrub 3 4 f t . Stems glabrous; young shoots red; mature. pale yellow-brown. Internodes 5-16 mm. Spines foliaceous, the basal portion 5-10 mm. broad, surrounding the stems, margined with 5-7 spinose ( 3 4 mm.) teeth (3-5 mm.). Leaves thick, 2-5 x 0.7-1-8 cm., excluding petiole 6-15 mm.; oblong, rigid; apex acute; margins 5-8-spinose (3-5 mm.)- spreading-dentate (3-5 mm.), distant 5-8 mm. ; above glabrous, sub-bullate, indis- tinctly veined, dull, pale grey-green; below dull, coarsely elevated openly reticulate and compicuously pilose-pubescent. Flowers solitary. Pedid in fruit, glabrous, 7-10 mm. long. Berries black, almost epruinose, oblong, estylose, or with style scarcely 0.2 mm. long. Ovules 3-5.

Chile: Prov. Maule, Cauquenes, 3000-4000ft., Dec. 1927, Clarence Elliott 114 (Type, K). Not in cultivation.

centibus, floribus solitariis, valde distincta

(336) Berberis variiflora Schneid., BUU. Herb. Boissier, (2), 5, 147, 1905 S t e m puberulous subangled, pale. Internodes 10-15 mm. Leaves 16-20 x 7-18 mm.,

oblong-elliptic to suborbicular, with petiole c. 1 cm. ; margins spinose-dentate; enervate. Injlorescence 4-6-fld., part fascicled, part subumbellate, with peduncle 1-5-2.5 cm., with an extra flower at its base. Pedid 2-5 mm., but of the basal flower to 10 mm. Flowers 4 mm. diameter. Petals subemarginate and subequal to inner sepals; stamens dentate. Ovules (1-) 2. Berries ellipsoid to subglobose, estylose, pruinose blue, 5 x 4 mm.

Chile: locality cited by author asvilla Rica, 1897, Neger, s.n. (Type, Herb. Munich; K).

(337) Berberis crispa C. Gay, FZ. Chil., 1, 86, 1845; syn. 3. actinacanthu var. crispa (C. Gay) Reiche, FZ. Chile, 1, 41, 1895

Stems slender; mature dark red-brown, sulcate, pubescent. I n t e d 5-9 (-12) mm. S p i m foliaceous, the basal part encircling the stem being 2-5 mm. broad, margined with 5-7 spinose (2-3 mm.) teeth (2-4 mm.). Leaves thinly coriaceous, oblong to oblong- orbicular, 5-9x245-6mm.; base contracted to petiole c. 1 mm. long; margins with 3-4 spinose (1-2 mm.) teeth (1-2 mm.) ; both sides reticulate; above dull; below pruinose grey. InJlorescence 4-8-fld., fascicled or umbellate-subfascicled, 6-10 mm. long, in- cluding peduncle 3-6 mm. Pedicels glabrous, 16-24 mm. ; bracts puberulous, subequal 1-25-2425 mm. Prophylls 1.25 x 0-4 mm., lanceolate. Outer sepals 1.5 x 1 mm., elliptic. Inner sepals 2 x 1.6 mm., obovate. Petals 2 x 1.5 mm., obovate. Stamens 1.5 mm., below the anthers lanceolate-dentate. Bendes globose, pruinose blue, estylose, 3 4 mm.

Chile: Prov. Valparaiso, in convall. amoena Quebrda de Chlorillis prope Quillota, fl. July, Poeppig., 8.n. (0) ; seed, 1928, Robinson. According to Reiche, lugares esteriles de lrts provincias centrales.

Cultivated: from seed collected by Mr G. W. Robinson (Kew 188-28)-fl. at Watling ton, 5 May 1941, 18 Apr. 1944. Shrub 2-3 ft.

(338) Berberis congestiflora C. Gay, FZ. Chil., 1, 75, 1845. (Fig. 57) Shrub dense with slender, arching branches. S t e m slightly sulcate; young shoots

puberulous; mature glabrous, or subglabrous, pale yellow. I n t e d 2.5-6 cm. Spines foliaceous, the basal part encircling the stems, 5-10 mm. broad, and margined with 10-20- spinose (16-3 mm.) teeth (0.5-1 mm.). Leaves thinly coriaceous, ovate-orbicular, 1-2.5 cm. ; base subcordate; petiole 2-5 cm., articulate towards the apex; margins 6-20-spinose ( 1 . 5 3 mm.)-serrate (0.5-1 mm.), distant 1-2 mm. ; apex rounded; both sides with clearly elevated open reticulation, and indistinct fine reticulation ; above dull grey-green ; below paler, grey, pruinose. In@escence 8-10-fld., corymbose-umbellate, 3 4 cm. long, in- cluding peduncle 2-3 cm. Pedicek 2-3 mm.; bracts 1-1.5 mm. Flowers 5-7 mm.

Page 236: Berberis and Mahonia : A taxonomic revision

236 L. W. A. AERENDT diameter. Outer sepals ovate, acute, 2 x 1-5 mm., H.C.C. 711. Median aepals concolorous, 3 x 2 mm., ovate-elliptic. Inner sepals 5 x 3 mm., oblong-obovate. Pet& [H.C.C. 8/1), 4 x 3 mm., obovate ; entire; cuneate with elliptic, separate, non-marginal, discolorous, orange-red glands, 0.7 x 0.3 mm. Stamem 2.5 mm. ; below the anthers acute- to acumi- nate-dentate; apex not produced, truncate. ovules 3-5. Berries M m m . , globose, estylose, pruinose blue.

Chile: Prov. Cautin, Temuco, 500ft., fr. Jan. 1928, Clarence Elliott 332; Dec. 1930, Clarence Elliott 586; 400 ft., Dec. 1939, Sandeman 366 (all, I(); seed Comber 913.

Prov. Valdivia, according to Philippi. Prov. Malleco, according to Philippi, who records that the species flowers from Sept.

to Nov., the fruit being ripe in December. Cultivated: from Comber 913, fl. 9 June 1942; fr. 16 Aug. 1941. Cf. (372). B. knig&ii.

(339) Berberis hakeoides (Hook f.) Schneid., BUZZ. Herb. BoiSSier, (2), 5, 146, 1905;

Shrub with few branches, 8-12 ft. Stem glabrous, yellow. Spines foliaceous, the basal part surrounding the stems being 5-12mm. broad, and margined with 7-14 spinose (2-3 mm.) teeth (2-3 mm.). Leaves thick, rigid, oblong-orbicular, 2 4 x 1.75-3.5 cm., excluding petiole 1-3 cm.; margins 7-15-spinose (2-3 mm.)-dentate (2-3 mm.), distant c. 3 mm.; apex rounded; above pale grey-green with indistinct venation; below grey, papillose, h e l y reticulate. Inflot.eceence 8-20-fld., sessile, fascicled, or racemose-sub- fascicled, 5-15 mm. long. Pedicek 4-7 mm. Outer sep& ovate, acute, 2 x 1 mm. Median a e f i 3-5 x 2.5 mm., oblong-ovate, rounded. Inner sepals 5 x 3.5 mm., obovate. Petals H.C.C. 8, spathulate, 3-75 x 2 mm., apex rounded truncate, slightly emarginate. Stamens 2 mm., below the anthers conspicuously lanceolate-dentate; apex not produced, trun- cate. Ovules 3-5. Berries 5 mm., globose pruinose blue, estylose.

syn. B. umgesti@ra var. h u h & Hook. f., Bot. Mag., t. 6770,1884

Chile: seed, 1861, near Arguilhie, Pearce, 8.n. Cultivated: Plant fig. in Bot. Mag. (Type) ; fl. c. 1884, at Veitch’s Coombe Wood nursery;

fl. 17 Apr. 1943,28 Apr. 1944; fr. 3 Aug. 1944.

(340) Berberis horrida C. Gay, Fl. Chil., 1,84,1845; syn. B. actinacantha var. kids

Stem glabrous, yellow. Spines at base of branches 5-10-fld., or foliaceous, but at apex 3-fld. Leaves 2 - 5 6 x 1-2 cm., excluding petiole 4 4 mm. ; rigidly chartaceous; margins 5-8-spinose-dentate ; below papillose. F’lowers fascicled 6-10. Stamm dentate. Berries oblong, estylom, 6 x 3 mm., black, scarcely pruinose.

Chile: Prov. Colchagua, Philippi, s.n. According to Lechler, in the Central Provinces. Not cultivated.

(C. Gay) Reiche, Fl. Chik, 1,42, 1895

Sect. Microphyllae Ahrendt, sectmov., foliis plusminusque sempervirentibus ; ramulis pallide flavia; ovulis tantum 2; fructibus pruinosis caeruleis; staminibus edentatis

Chile: Prov. Magellan. Argentine: Provinces Santa Cruz, Tiem del Fuego. Falkland Islands.

GY 63, to Sect. Microphyllae Leaves mostly 1-2 cm. long; spines 1-2 cm. long; inflorescence 2-3-fld., subfascicled.

Leaves mostly 3-4 cm. long; spines weak; intloreacence 8-14-fld., subracemose, P 6 cm. 341. B. nzicrophylh

long. 342. B.lilloana

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Berberis and Mahonia 237

(341) Berberis microphylla Forst., Comm. Qoett. 9,29, 1789; Dippel, Handb. Laubh.

Internodes 8-15 mm. Stem yellow. Spines 3-fld., 1-2 cm. long. Leaves (lo-) 15-25 (-30) x (6) 7-12 (-15) mm.; elliptic-obovate; apex acute; entire; base cuneate with petiole 4-8 mm. ; above dark green, scarcely lustrous; below grey, pruinose. Inflorescence 2-3-fld. Pedkls glabrous, 3-5 mm., but in fruit to 9 mm. long. Flowers 1 cm. diameter, pale yellow. Outer sepals reddish, 2.5 x 1.5 mm., ovate. Inner s e w 6 x 4 mm., obovate. Pet& 4 x 3 mm., obovate-spathulate, entire. Stamena 2.5 mm., edentate; not produced, tnuicate. Ovules 2. Berries globose, estylose, pruinose blue, 6 mm.

Argentine: Tierra del Fuego, Forster, s.n., Type; seed, Collingwood Ingram, s.n. Prov. Santa Cruz, Patagonia.

Chile: Prov. Magellan, collector unknown, s.n. Falkland Islands: 1839-45, Hooker, s.n. (0). Cultivated: L.A. 737, from Tierra del Fuego seed, collected by Mr Collingwood Ingram.

3, 134, 135, fig. 71, 1893; Hook. f., F1. Antarctica, 2, 231, t. 87, 1847

(342) Berberis lilloana Job in LiUoa, 17, 82, 1949 A small tree. Spines 3-%d., weak. Leuves membranaceous, 3-4 x 1.5 cm., margins

completely entire; petiole c. 1 cm.; venation slight. Racemes 8-14-fld., longer than the leaves. Pedicels 5-7 mm. Stccmens dentate. Ovlcles 2. Berries estylose.

The author provides only the facts cited, and cites no l o d t y or specimens.

Sect. Corymbosae Schneid., Bull. Herb. Boissier, (2), 5, 812, 1905 Leaves deciduous. Injiorescence 4-12-fld., 3-5 cm., more or less umbellate, with

peduncle 2-3.5 cm., rarely fascicled. Ovules 3-5 (-6). Berries estylose, or with style very short and insignificant.

Juan F e r h d e z Islands. Chile : Provinces Antuco, Concepci6n, muble. Range 34"-38" S. latitude.

KEY 64, to Sect. Corymbosae Stenls pubescent; flowers fascicled; leaves always reticulate and spinose; ovules 0.

Stems glabrous; inflorescence more or less umbellate and peduncled; leaves entire; ovules 3-5.

Stamens edentate, truncate ; spines mostly absent, occasionally 3-fld., but weak ; leaves acute; peduncles 1-2 em. ; fruit 8 x 6 mm. , ellipsoid, pruinose, shortly stylose.

Stems pale yellow-grey, lustrous; leaves thin, entire, with petiole 1-2 cm., and broad (4 x 2.5 om.) closely reticulate lamina. 344. B. corymbosa

Stems dark red-purple, dull; leaves thick, spinose, with petiole 2-6mm., and narrower (3.5 x 1-3 cm.) lamina, a t most openly reticulate. 345. B. masafuerana

Strtmens dentate; spines 3-5-fld., palmate; berries 5 x 4 mm., subglobose, black, epruinose, estylose. 346. B. rotundayolia

343. B. setigrifolia

(343) Berberis setigrifolia Ahrendt, sp.nov., inter sectionem ramulis pubescentibus; foliis reticulatis marginibus spinoso-serratis ; dorescentiis fasciculatis ; ovulis 6 ; distinguenda

8tt:m-s pubescent, terete; youngshootsred; mature pale yellow-brown. Internodes 3-5 cm. Spines absent. Leaves mostly ovate, and acute, but sometimes oblong-elliptic and sub- obtuse, 2-33 x 14-2 cm.; base rounded with petiole 5-12 mm.; margins 6-12-spinose (2-2.5 mm.)-dentate (1 mm.), distant 3-6 mm.; both sides closely reticulate with elevated veinlets; above dull green, or almost so; below grey, pruinose, papillose.

Page 238: Berberis and Mahonia : A taxonomic revision

238 L. W. A. ~ E N D T

Flowers fascicled 3-5. Pedkds glabrous, 2 - 5 4 cm. Fbwers 1-5 cm. diameter. Outer and inner sepah both 9 x 7 nun., obovate. Petals 6 x 3 mm. Sbmw 4 mm., edentate; apex produced, obtuse. Ovala 6. Styk absent.

Chile: without l o d t y , lo00 ft., Dec. 1865, Pearoe 8.n. (Type, K). ‘A deciduous little bush with erect branches and orsnge-red flowers.’ Not cultivated.

(344) Berberis corymbosa Hook. et Am., Bot. Mbc. 3,135,1833; Reiche, Fl. Chile,

Stem glabrous, pale yellow-grey, lustrous, subterete. Spines often absent, sometimes 1-3-fld., 1-5 mm. h w thin, elliptic, acute, entire, with cuneate base, 2-4 x 1-2-5 cm., excluding petiole 8-20 mm. ; both sides finely reticulate; green, papdlose, below. P e d i d slender, glabrous, 1-2 cm. Stamens dentate, trwmte. Berries ellipsoid, pruino~e blue, 8 x 6 mm., excluding style 0-5 m.

1,34,1895

KEY MA Inflorescence 4,7-fld., corymbose-umbellate, 2-5-36 cm. long, including peduncle 1-2 cm.

Inflorescence &12-fld., corymbose-paniculate. 344a. B. wr.yTnbosa var. cory~nbosa 344 b. B. eosl/mbosa vm. pankulata

(3444 Berberis corymbosa vm. corymbosa Juan Femhdez : without locality, Philippi, s.n. Type.

(3446) Berhris corymbosa var. paniculata Phil., A d . de La Univemidad, Santiago, 1872,664,1872; Reiche, Fl. Chile, 1, 35, 1895.

Juan Femhdez: without locality, Philippi, s.n. Type. According to Philippi, both varieties flower September to October. Neither is in cultivation.

(345) Berberis masafuerana Skottsberg, Phanerogams of Juan Fernh&z, 125, no.

Shrub 3-5 ft. Stenzs slender, slightly sulcate; mature dull, dark, red-purple. Spines 1 (-3)-fld., 3-5 mm. Leaves 2-3.5 (4-8) x 0.8-1.3 (-2-8) cm., including petiole 2 4 (-8) mm. ; obovate-elliptic; entire, reticulate; slightly paler and pruinose below, but with the new leaves suborbicular and minutely spinulose. InJlorescence c. 11-fld., racemose, c. 5 om., including peduncle 2-3 cm. Pedicela 6 7 mm.; bracts 2-3-5 mm. Otherwise unrecorded.

Juan Fern4ndez: Masafuera, Q. de la Cam, one seedling by the side of a waterfall, a t 700 ft. ; larger shrubs seen on the wall of a canyon in an utterly inaccessible place, but some branches were obtained by means of a lass00 (no. 503) ; Masahem, Q. de la Loberia, one shrub on steep rock in the stream (Skottsberg) ; Maaafuera, Douglas, 8.n. (0). Not cultivated.

57, 1921

(346) Berberis rotundifolia Poeppig et End., Nova #en. w Sp. PI. 2, 63,1839 Stem terete, glabrous; mature dark red. Spines 35-fld., palmate, 4 4 3 mm. Leaves

thin, entire with rounded apex; base contracted to a petiole 1-2-5 cm., which is articu- lated below the broadly elliptic lamina, the latter measuring 2-4-3.6 x 1-6-2.4 cm. ; dull green above; below grey, pruinose; both sides not, or scarcely, reticulate. Inflorescence umbellate, 5-7-fld., 3-5-5 cm. long, including peduncle, 2-3-5 cm. Pedicels glabrous, 1-1.4 cm. ; bracts 1-1.5 mm. Flower diameter 10-12 mm. Sepals sulphur yellow. Petals orange. Stamens dentate. Berries black, epruinose, globose-ellipsoid, 5 x 4 mm.

Chile: Prov. Antuco, Andes, close to Pic0 de Pilque, Poeppig 747 (Type).

Page 239: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 239

I'rov. Ruble, 1856-7, Germain, s.n., according to Schneider. Prov. Concepci6n, acwrding to Gay. The palmate spines and dentate stamens relate this species with the Actinacanthae

amongst which Schneider Erst placed it. B. rotundifoiia may well be. regarded aa linking the Actinacanthae and Corymbosae.

Sect. Buxifoliae Schneid., BUU. Berb. Boi8&r, (2), 5, 139, 1905 Leaves mostly evergreen, sometimes deciduous. Plowers solitary, or fascicled 2-5,

rarely elongated umbellate-racemose. 0vuk.s 6-12. Bern& mostly estylose, otherwise with style very short, insignificant (to 0.4 mm.).

Chile; Provinces Colchagua, Antuco, Valdivia, Made, Ruble, Llanquihu6, Magellan. Argentine: Provinces Catamarca, mrdoba, Neuqudn, Rio Negro, Santa Cruz. Uruguay: Monte Video. Brazil; Prov. Minas Germ.

KEY 65, to Sect. Buxifoliae Leaves broad, 1 : 1.5-2.6 (-4). (Subsect. EWWIFOLLAE.)

Corolla with 6 petals, as usual. Spines present.

Flowers solitary, occasionally fascicled 2-3 (-4).

Flowers fascicled 2-4; stems glabrous. Spines 3-7 mm. ; pedicels 4-8 mm.

Fruit stylose ; leaves spinom. Fruit estylose ; leaves entire.

Leaves without hypoderm, thin; stamem edentate.

347. B. hieronymi 3486. B. buxifolia var. nudu

349. B. spinosisaima Spines 2-5-5 em.; pedicels 2-3 cm.; leaves entire; fruit stylose.

Flowers solitary; stems pubescent, at any rate at Gret. Stems terete.

Spines absent, or weak, generally up to 7 (-20) mm., 1 (-3-fld.). Leaves coriaceous, acute; stems dark red-brown. 348. B. buxifololia Leaves thin, rounded or spathulate ; sterns pale yellow-grey.

350. B. parodii Cf. 497. B. bardochen&8 Spines stronger, 1.5-2 cm., sometimes 3-6-fld.

Leaves broad, 1 : 1-5, acute; petals much shorter than inner sepals;

Leaves narrow, 1 : 3, obtuse; petah slightly shorter than inner sepals;

stems sulcate.

stamens not produced. 351. B. montevevidensis

stamens produced. 352. B. campos-pwtoi Leaves with hypoderm, thick; stamens dentate. 353. B. cuneatu

Racemes 3-5-fld. ; stamens dentate. 354. B.michay Racemes 6-12-fld.; stamens edentate. Cf. 496. B. andeana

348c. B. buxifolia var. i n e m i s 355. B. serrato-den-

356. B. polypetdu

Inflorescence racemose-submbdak.

Spines absent. Leaves entire; stems red-brown. Leaves spinose ; stems yellow-grey.

Corolla recorded as with 12 petals.

Inflorescence 1-4-fld. ; leaves with hypoderm. Leaves narrow, 1 : (5-) 7-20. (Subsect. EM-PETRTFOW.)

Leaves linear, 1 : 12-20, mostly entire; flowers solitclry ; pedicels 5-8 mm.

Laves broader, 1 : (5-) 7-12, sometimes serrate; inflorescence 2-4-fld.; pedicels 357. B. empetrifolia

8-12 IIM. 358. B. wawTa7Mc

Infkwescence 5-10-fld. ; leaves without hypodem. 359. B. mutabilia

Page 240: Berberis and Mahonia : A taxonomic revision

240 L. W. A. AHRENDT

Subsed. Eubdoliae Schneid., 1.0. 141, 1905 (347) Berberis hieronymi Schneid., BUU. Herb. Boiesie*, (2), 5; 394, 1905

Stem with young shootsred, sometimesslightly pubemlous ; matureghbrous. Internodes 6-12 mm. Spina 3-fld., 3-7 mm. Law obovate, to 13 x 5 mm.; baae cuneate, sessile; margins entire, or with tridentate apex produced by one tooth each side towards apex (spinulw 1.5 mm. long, serrations 1-1-5 mm. long) ; mucromte; with only a few elevated lateral veins; both sides dull, below paler, epapillose. Infloe8cew 3-5-fld., fascicled or subumbellate. Pedkds glabrous, 4-6 mm. F k r diameter c. 6 mm. ProphyUe 2-2.5 x 0.5-1 mm., lanceolate. outer 8epa.h 3 x 1-75 mm., ovate, obtuse. Inner sep& 4 x 3-5 mm. elliptic. Petah 4 x 3 mm. ; base with separate, ovate, acute glands, 0.8 x 0.5 mm. Stamm 3.5 mm., not produced, truncate. O v d a c. 6. Sty&? short.

Argentine: Prov. mrdoba, Sierra Achah de C%rdoba, 15 Nov. 1878, Hieronymus, 8.n. (Isotgpe, K); 1876,1877,1883, Hiemnymus s.n., according to Schneider. Not cultivated.

(348) Berberis buxifolia Lam., IU. Gen., t. 253, fig. 3, 1792; Poir., in Encyd. M&?wd., Bohnique, 8,619, 1809; Schneid., B d . Herb. BoiSSier, (2), 5, 142, 1905; Job in Rev. Mw. La Plata (N.S., sect. Bot), 5,34, fig. 5,1942; Hook. f., Bot. Mag., 106, t. 6505, 1880. (Fig. 58)

Stem terete; mature red-brown. Internotla 1-2 cm. Lxzm co+ceous, elliptic, entire; above dull da k green; below grey, pruhose. Stclmens dentate. Owdes 6-12. Berrim globose, pruinose blue, estylose.

KEY 65a Leaves epapillose below; pedicels3-16 mm.; fruit 6-8 mm.

Pedicels 10-18 mm. ; stems pubescent; flowers SOL-. Spines present, weak, P 7 mm. long. Shrub spreading, 6 ft. tall and through. Shrub compact, 2 fi. high and through.

348a. B. M f o l i a var. M f o l i a 3486. B. M f o l i a var. mm

Spines absent. 348c. B. bux@olia var. inermis

Stems pubescent. 348d. B. M f o l i a var. antarctica Stems glabrous. 348e. B. M f o l i a var. nud.ci

348f. B. M f o l i a var. papillosa

Pedicels 3-7 mm.; spines 4-7 mm.

Leaves papillose below; pediceh 15-30 mm.; fruit 12-16 mm. long.

(348a) Berberis buxifolia var. burdfolia; syn. B. d&i% Sweet, The British Flower Garden, (2), 1, 100, 1831; Maund, The Bohnht, t. 42, 1837; non Paxton.

Shrub to 7 ft., with arching branches. Stem mature pubescent. Leave8 to 17 x 9 mm. Flowers solitary, 1446 mm. diameter. Pedicels puberulous 14-18 mm. Outer sepals, H.C.C. 7; narrowly oblong-obovate, 6 x 3 mm. Inner sepal8 concolorous, 7-8 x 4-6 mm., oblong-obovate. Petals H.C.C. 9/1-10; elliptic-obovate, 4.5-6 x 2-3.5 mm., entire, cuneate ; glands lanceolate separate, non-marginal, concolorous, 2 x 0-5 mm. Stamens 4-46 mm., edentate, but slightly swollen below the anthers; apex produced, retuse with acute margins. Be& 6 mm. Chile: Prov. Antuco, Poeppig 107,667. Prov. Magellan, Commerson, 8.n.; Punta Are=. wastes and slopes, very common,

Argentine : Prov. Neuquhn, Cerro h h a r i , Comber 415 (K) ; San Martin de 10s Andes ;

Prov. Rio Negro, San Carlos de Bariloche.

fl. 26 Nov. 1950, Brooke 7052 (BM).

Lago Nahuel Huapi.

Page 241: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 241 According to Job (who gives the latter citations) the general distribution is 37'45" S.

Chltivated: L.A. 443, fl. 7 May 1942; fr. Aug. 1942. Photograph in &rd. Illust. latitude.

p. 617, 28 Nov. 1941.

(3483) Berberis buxifolia var. nana A. Usteri, Mitt. Dtsch. Dendr. Ges. 8 , 90, 1899 Outer sepals H.C.C. 712, narrowly obovate, 7 x 2 mm. Inner sepals concolorous,

10 x 6.5 mm. Petals H.C.C. 911, oblong-obovate, 6 x 4 mm., entire, truncate. Stamens 5 mm., edentate, not produced, truncate.

Cultivated: L.A. 3, fl. 17 Apr. 1941, 12 May 1942, 13 Apr. 1943,26 Apr. 1944.

(348c) Berberis buxifolia var. inermis (Persoon) Schneid., Ill. Handb. Laubh. 1, 302, 1904; syn. B. inemis Persoon, Syn. P1. 1,387, 1805

(348d) Berberis buxifolia var. antarctica Schneid., B d . Herb, Boissier, (2), 5, 142, 1905

Leaves sub-evergreen or sub-deciduous, 18-30 x 7-12 mm. Pedi& 3-5 mm. Chile: Cape Horn, 56" S. latitude, Hooker 1098 (Type).

(348e) Berberis buxifolia var. nuda Schneid., I.c., 1905

Berries 8 mm. Stems glabrous; mature dark red. h v e a 13-22 x 6-12 mm. Flowers fascicled 2-4.

Chile: Prov. Valdivia, Villa Rica, 1897, Neger, s.n. (Type).

(348f) Berberis buxifolia var. papillosa Schneid., IW. H a d . Laubh. 1, 302, 1904; syn. B. dulcis Paxton, Mug. Bot. 10, 171, 1843, non Sweet nec Maund

Shrub habit rigid, 3-5 ft., branches a little inclined to turn downwards. Spines slender, fairly long. Leaves obovate; apex subacute to subobtuse, to 15 x 7 mm., entire; below pruinose, papillose. Flowers solitary, 15-17 mm. diameter. Pet%& 1-5-3 cm. Outer sepals 8 x 5 mm. Inner sepals 9 x 6 mm., all obovate. Petals smaller. Bemh globose, 12-15 mm.

Chile: Prov. Valdivia, Villa Rica, Neger, s.n. (Type). According to Lindley B. buxifolia comes from Magellan, and may be distinguished

from B. dulcis of Valdivia, the latter with larger, more spathulab leaves which are more deciduous, and larger fruit. The earliest B. dulcis, that of Sweet, seems identical with B. buxijolia var. buxijolia. Lindley's Valdivian plant, with larger fruit, appears to correspond with B. dulcis of Paxton, which has somewhat obtuse leaves, and, according to Paxton's figure, very large fruit. Schneider's variety seems to be the same it9 this, having also the very long pedicels; but he does not describe the fruit as of such large size as I'axton does. It may be that Paxton's figure exaggerates this feature; or that Schneider's B. buxifolia var. pa/pillosa and Paxton's B. dulcis are two close varieties, or two forms of one variety.

(349) Berberis spinosissima (Reiche) Ahrendt, stat.nov. ; syn. B. buxi,folia var. spinosissima Reiche, PI. Chile, 1, 39, 1895; syn. B. bwxifolia var. mcracanthu Phil. ex A. Usteri, Mitt. Dtsck. Dendr. Ges. 8, 90, 1899

Stems glabrous; mature dark red, sulcate. I & d 1 3 c m . Spines very stout (2-5--) 3-5 cm. long, noticeably discolorous, paler yellow-red, 1-3-fld., with the lateral parts shorter. Leaves deciduous, 10-20 x 3-7-5 mm., obovate, entire, slightly undulate; venation branched; above dull grey green; below grey, pruinose, epapillose or scarcely papillose. In,,oreswnce (1-) 2-3 (4)-fld., fascicled. Ped& glabrous, slender, green,

16 JOURN. LlNN. S0C.-BOTANY POL. LVII

Page 242: Berberis and Mahonia : A taxonomic revision

242 L. W. A. AERENDT 2-3 cm. Ozcter 8epal.s H.C.C. 7, oblong-elliptic, 4 x 3 mm. Inner sep& concolorous, 7 x 5.5 mm., obovate. PetaEa H.C.C. 9,6 x 4-5 mm., oblong-obovate; entire; base cuneate with concolorous, separate, non-marginal, lanceolate glands, c. 2.25 x 0.5 mm. S t a ~ m 3.5 mm., edentate; apex apiculate. Ovules 9-12. Berrdes globose, pruinoSe blue, 6 mm., excluding distinct style, 0.4 mm.

Argentine: Prov. Neuqukn, Pulmari, J d n de 10s Andes, 3500 ft., fr. 16 Jan. 1926, Comber 417 (K).

Chile: Prov. Ruble, Cord. de C W n , according to Philippi. Prov. Valdivia, Hohenacker 524, according to Schneider. Cultivated: from Comber's collection, no. 417, fl. 5 May 1941; fr. Aug. 1940. This species is very distinct from B. buxifolia in its quite deciduous leaves of thin

texture, its markedly strong spines, its apiculate stamens, and the very distinct, though short style. Comber givea the following field notes-'a large bush, 12 ft. high by 20 ft. wide. Fruit round, black, with bloom, edible; hardy'.

(360) Berberis parodii Job, Darwinians, 5, 184, fig. 1, 1941; Rev. M w . La P h

Tree 15-20 ft. high. Stem with young shoots reddish; mature pale yellow or grey. Internodes 1-4-2 cm. Leavee fascicled 6-10; dull grey-green; below pruinose, grey; entire; obovate-spathulate ; apex rounded or subtruncate ; base contracted to , and articulate with, petiole 3-9 mm. Spine% 3-fld., sulcate, 1-2 cm. E'lowers 8-10 mm. diameter, with all components obovate. Outer s e p h 4 x 2.5 mm. Inner se& 5 x 3.5 mm. P W 3-5 x 2.5 mm., entire. Stamem 3 mm., edentate; apex produced, shortly apiculate or obtuse. Ovules 8-12. Berries globose, 7-9 mm., estylose, pruinose blue.

Argentine: Prov. Neuquth, Lago "raful, 17 Jan. 1935, Cabrera & Job 396,410 (Type, La Plata) ; also herto &yer, Lago Espejo, Rio k p a n r t .

Prov. Rio Negro, San Carlos de Bariloche. Localities according to the author, who gives the general distribution aa 40'42 S. latitude.

(N.S., sect. Bot.), 5, 38, fig. 7, 1942

(361) Berberis montevidensis Schneid., BuU. Herb. BoiS.&er, (2), 5, 143, 1905

Stem thick, pubemlous, sulcate; mature red-brown. I n t e e 6-1Omm. S p i w stout, 5-12 mm., yellow-brown, sulcate. Leaves 10-15 x 7-10 mm., broadIy obovate, or obovate, entire, acute; below papillose. FZinm-8 solitary. Pdi& glabrous, 10-15 mm. Outer sepal.s 5 x 3 mm. Inner se& 8 x 5 mm., obovate. Pekrls 4-6 x 2-25 mm., obovate, entire. Stamens 4.5 mm., dentate; not produced, truncate. ovules c. 9.

Uruguay: Monte Video, ex Herb. Brown (Type, Vienna). Cultivated: at Wisley, 00wering May 1939.

(362) Berberis campos-portoi Brade, Rodriqueaia, 9, no. 20,43, 1946 Shrub to 6ft. Stem more or leas flexible; young shoots pubescent ; mature yellow brown,

sulcate, puberulous, but becoming finally glabrous and grey. Spines la-fld., to 15 mm. long, sulcate below. Leaurn subcoriaceous, glabrous ; obovate-spathulate ; completely entire; base subsessile; apex obtuse, mucronate; above dull; fascicled 5-12, to 23 x 8 mm. Flowers 8olitrtry, golden yellow, 8-12 mm. diameter. Peokd.~ 10-12 mm. Outer sepab ovate-lanceolate, 6x3mm. Inner sepab broadly ovate, obtuse, 7-8x5mm. Pek& obovate-suborbicular, slightly shorter than the inner sepals, 6 x 5-6 mm. Stamens dentate; apex produced, obtuse. Ovary with style absent or very short and thick.

Brazil: Prov. Minas Geraes, Serra do Capara6, 30 Oct. 1922, 7700ft., P. Campos- P6to 1148; Serra do Caparab, Caaa Queimada, 8000ft., 25 Sept. 1941, Brade 17020 (Type, in Herb. do Jardim Bot. de Rio de Janeiro, 45941). This species is not in culti- vation, but is intereating in its marked extension of the range of the section. Mainly,

Page 243: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 243 the Sect. Buxifoliae is recorded from 30" S. lat., extending southwards, in the west, that is in Chile and adjacent Argentine. An exception is B. mtevidensis which, still at 30" S. lat., extends the range to the east coast. B. campos-portoi, while still in the east, presents the section with a species some 10" farther north.

(353) Berberis cuneata DC., Syst. 2, 16, 1821; Job, Rev. Mus. La Phta (N.S., sect.

Ashrub 5-7ft. Stem glabrous, sulcate, red-brown. Spines 3-fld., M c m . , sulcate. h z v e s thick, rigid, subsessile, 15-20 x 4-6 mm. ; margins by apex 1 (-2)-spinose (0.5 mm.)- serrate (2-3 mm.), being thus generally tridentate, rarely entire; obovate-cuneate ; above bright green; below papillose, dull green. Flowers solitary. Pedicels 1-2 cm. Outer sepals 3 x 1 mm., oblong-ovate, obtuse. Inner sepals 6 x 3.75 mm., oblong-obovate, rounded. Petals 5 x 3 mm., oblong-obovate; apex entire; glands c. 0.8 x 0.4 mm., separate, non-marginal, close to the truncate base. Stamens below the anthers, con- spicuously acuminate-dentate ; apex not produced, truncate. Berries obovoid-globose, estylose, pruinose blue, c. 7 mm.

Bot), 5, 48, fig. 11, 1942

Argentine: Prov. Santa Cruz, Patagonia, east coast, Rio Santa Cruz, Berg s.n. I?rov. Neuqudn, Voldn Copahu6, 6 Mar. 1888, Kurtz, s.n. (La Plata 25100). Yrov. Rio Negro, Bahia Mpez, 15 Jan. 1935, Cabrera s.n. (La Plata 1362). The last

two citations are according to Job who gives the general dktribution as Patagonia, 35'42 S. lat. Not cultivated.

(354) Berberis michay Job, I.c., 5, 55, fig. 14, 1942 Stem pale brown. Spines 1.5-2 cm., with shorter lateral parts, sulcate below. Leaves

15-30 x P 8 mm. ; petiole 3-5 mm. ; narrowly obovate ; entire, sometimes tridentate ; above dull green; below pruinose, greyish mauve; venation slight. InJorescence 3-5-fld., very loosely racemose, 610 om. long. Pedicds 1-2 om.; bracts 3 4 mm. Flowers 5-8 mm. diameter. Sepals recorded aa 4, measuring 6-5 x 3.5 mm. Petah in one series of 3, oblong-triangular, obtuse, 4.5 x 2 mm. StamRns conspicuously lanceolate-acuminate- dentate. Berries globose, black, pruinose blue, 8-10 mm., estylose.

Argentine: Prov. Santa Cruz, Arroyo del Bote, 'Iter patagonicwn' no. 354, Mar. 1914 (Inst. Darwinion) (Type).

I'rov. Rio Negro, Lago Nahuel Huapi, 1 Nov. 1914, Meunier 8.n. The author gives the general distribution aa Cordilleras, 41"-51" S. lat.

(353) Berberis serrato-dentata LechIer, Berberides Americae Austr., 16, 1857 ; Skotts- berg, Vet. -AM., Handl. (N.S.), 56, 228, 1914; non Reiche.

Stem green to grey. Spines absent. Leaves narrowly obovate, to 12-15 x 5 mm.; base attenuate; margins spinose-serrate; above lustrous; below paler green; texture coria- ceous. InJloresCence fascicled or subracemose. Pedicels long. Berries ovoid, estylose.

Chile : Prov. Valdivia, Lechler 2985 (Type) ; Cord. de Valdivia, Pilad&, flowering Nov. and Dec., 'with the aspect of an Escallonia'. Not cultivated.

(356) Berberis polypetala Phil., A d . Uniu. Santiago 1872, 666, 1872 Leave membranaceous, ovate-suborbicular ; base attenuate to petiole ; paler below.

InJrmescence with peduncle two to three times aa long as the leaves. Petals recorded as 12 in number.

Chile: Prov. LlanquihuB, Cdbuco, laa faldaa del Voldn. Not cultivated.

16-2

Page 244: Berberis and Mahonia : A taxonomic revision

244 L. W. A. ABRENDT

Subsect. Empetrifoliae Schneid., BuU. Herb. m r , (2), 5, 139, 1905 (357) Berberis empetrifolia Lam., IU. Cen., t. 253, fig. 4, 1792; Poir., in. Encycl.

M&!?wd. Bot. 8,621, 1809; Schneid., BUU. Herb. BoisSier, (2), 5,140,1905; Lindley, Bot. Reg. 26, t. 27, 1840; Job, Rev. Mw. La Plata (N.S., sect. Bot.), 5, 32, fig. 4, 1942; syn. B. revoluta Sm. ex DC., Syst. 2,16,1821, pro sp. (fig. 59)

Shrub subprostrate, 6-12 in. high. Stem terete; young shoots puberulous; mature sub- glabrous, dark red. Leaves linear oblong, 10-20 x 1-1-5 mm.; margins entire, revolute; enervate; slightly lustrous dark green; below grey, pruinose. Flowers H.C.C. 6-7. ProphyuS 2.5 x 1.5 mm., ovate, acute. Ozltersepale 3.5 x 1-25 mm., oblong-obovate, acute. Median sepals 4-25 x 1-75 mm., narrowly ovate, acute. Inner sepals 7-25 x 3 mm., nar- rowly obovate, acute, or subacute. Petals 4.5 x 2 mm., oblong-obovate, emarginate with indistinct, oblong, separate, non-marginal basal glands, 1.2 x 0.5 IIM. Stamens 4.5 mm., shortly dentate; apex not produced, truncate. Ovules 8-12. Bern& 6-7 mm., globose, eatylose, pruinose blue.

General distribution of species (i) in Chile, Provinces Santiago to Magellan, 33'45" S. lat., and (ii) according to Job, in Argentine, Provinces Cahnmcat o Tierra del Fuego, 28O-55" S. lat.

KEY 6 5 ~ to Subsection Empetrjfoliae Spinea 6-12 mm., shorter than the leaves (1-2 cm.) which are oonvolute.

Spines 15-26 mm., longer than the lavea (1-2 cm.) which are scarcely convolute. 357a. B. tmpetr$oliu var. empetkfolia

357 b. B. empeh+folia vm. mzg&nica

(357 a) Berberis empetrifolia var. empetrifolia Chile: Prov. Magellan, Punta Arenas, waste near sea, Brooke 5993 (BM), fl. 24 Nov.

1950; Prov. Santiago, Philippi; Prov. Ruble, chiusn, 1855, Germain, s.n.; Prov. Valdivia, Villa Rim, 1897, Neger, s.n. ; Prov. Antuco.

Cultivated: fl- Apr. 1939; fr. June 1939.

(357b) Berberis empetrifolia var. magellanica Schneid., Bull. Herb. Boissier, (2), 5, 140, 1905

Chile: Prov. Magellan, Commerson, s.n.; Sandy Point, Hohemcker 1065

(358) Berberis wawrana Schneid., I.c., 140, 1905 S t e m glabrous, scarcely sulcate, red-brown, sometimes slightly pruinose. Internodes

c. 1 cm. Spines 3-4-fld., brown, below broadly sulcate, 3-14 mm. Leaves 10-30 x 1.5- 2.5 mm. ; lanceolate; entire or with 1-2 spinules near the middlea of the margins; at first plane, kally broadly revolute; below papillose. In.e.sct?nce 24-fld., faacicled or sub- umbellate, glabrous. Pedicds 8-12 mm. long. Flowers 8-10 mm. diameter. Petals sub- equal to inner sepals. Stamem dentate. Ovules 8, sessile.

Chile: Prov. Colchagua, Philippi, s.n. ; without locaJity, Seibold-Wawm 3039.

(359) Berberis mutabilis Phil., A d . Univ. Suntiago, 1872, 665, 1872; Schneid., 1.c. 141, 1905

Stems subsulcate, dark brown, often pruhose. Internodes 12-18 mm. Spines weak, 8-12 mm. Leaves 10-20 x 1 - 5 4 mm., oblanceolate, entire, subplane. Flowers fascicled P10. Pedicels c. 1 cm. Flowers c. 8 mm. diameter. Petals shorter than inner sepals.

Chile: Prov. Maule, Hacienda de Cauquenes, Caj6n del Arriem, Dessauer, 8.n. 1875. Prov. Valdivia, Villa %cay 1897, Neger, 8.n. Not in cultivation.

Page 245: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 245

Sect. Montanae Ahrendt, sect.nov., foliis plerumque deciduis ; inflorescentiis 1-3 (-15)-

Chile : Provinces Santiago, Colchagua, Maule, Ruble, Cautin, Valdivia, LlanquihuB. Argentine: Provinces Neuquh, Rio Negro. Range 33"42" S. lat.

fld.; ovulis (5-) 6-9; stylo valde elongato, 2-3 mm. long0

KEY 66, to Sect. Montanae Inflorescence P15-fld., subumbellate or racemose, 3-6 cm. long.

Leaves obovate to spathulate; apex rounded; inflorescence subumbellate, 4-6-fld. ;

Leaves lanceolate ; inflorescence 10-15-fld., subracemose ; stamens edentate. stamens dentate. 360. B. cabrerae

361. B. mikuna Inflorescence 1 (-3)-fld., 1-1.5 (-2) cm. long.

Mature stems dark red. Stems glabrous, very stout; flowers fascicled; stamens much shorter than petals.

Stems puberulous, less stout; flowers solitary; stamens subequal to petals. 362. B. mntana

363. B. chillunensis Mature stems pale yellow-brown, puberulous. 364. B. coletioides

(360) Berberis cabrerae Job, Rev. Mus. La Plata (N.S., sect. Bot), 5,60, fig. 16, 1942 Stems glabrous, slender. Spines 1-3-fld., 8-10 mm. long, deep brown, sulcate. Leaves

mernbranaceous, obovate to spathulate ; apex rounded; venation notably branched, finely reticulate; 15-30 x 8-18 mm. Injbrescence 4-6-fld., subumbellate, 3-6 cm. long, including peduncle 5-15 mm. Pedicels 1-2 cm. Prophylls 2, obtuse, 4 x 2.5 mm. Outer sepuls 7 x 3 mm., elliptic. Median sepals 10 x 6 mm., obovate. Inner sepals 15 x 8 mm. Pet& (in two series of three), 8 x 5 mm., with basal glands. Stamem shortly dentate; apex shortly apiculate.

Argentine: Prov. Rio Negro, Bariloche, Cerro Lopez, 5000 ft., Cabrera 325, 15 Jan. 1935 (Type, La Plata no. 26065).

Prov. Neuqu6n, Comber 798 (1925-27) (K). General distribution, according to the author, 38"41" S. lat.

(361) Berberis mikuna Job, Lalba, 17, 81, 1949 Shrub to 20 ft. high. Spines 3-fld. Leaves lanceolate, membranaceous, entire, often

sinuate, subsessile, with apex mucronulate. Racemes 10-15-fld. Flowers yellow. Pro- phylls 3, triangular. Sepals in two series each of three, obovate. Pet& in two series of three. Stamens apiculate. Berries black. The author provides only the above facts and citeai no specimens or localities.

(362) Berberis montana C. Gay, Fl. Chil. 1, 90, 1845; Job, Rev. Mw. La Plata

Shrub very erect in habit, stiffly branched, 7-12 ft. high, much less through. Stems very stout, always glabrous ; young shoots reddish; mature dark, red or red-brown, angled- sulcate. Internodes 7-25 mm. Spines concolorous, stout, 3-fld., 7-12 mm. Leaves 10- 20 x 6-11 mm., deciduous, thin, obovate, entire; both sides green; papiuose below; venation slight, with only very few veins visible. Flowers fascicled 2-3. Pedicels ( 1 6 ) 15-20 mm. Flowers 1.5-2 cm. diameter. Outer sepals 5-6 x 3-4 mm., oblong. Median sepals 5-8.5 x 4,4-5 mm., elliptic. Inner sepals 95-10 x 6-7 mm., obovate. Petals 7-8 x 5-6 mm., obovate, entire, or slightly truncate-emarginate; base cuneate with lanceolate, acuminate glands, 1-2 x 0.2 mm. Stamens 4-45 mm., dentate, shortly apiculate. Ovules (5-) 6-9. Berries pruinose blue, globose, 64fx 6-v.5 mm., excluding stvle 2-3 mm.

(N.S., sect. Bot.), 5, 58, fig. 15, 1942. (Fig. 60)

Page 246: Berberis and Mahonia : A taxonomic revision

246 L. W. A. AHRENDT Chile: h v . Cautin, Volch Lla,ima, 3600 ft., fl. Feb. 1927, Werderm- 1277 (K). Prov. Vddivh, Cord. de h c o , Lechler 1990 (Type). Argentine: Prov. Neuqudn, 1925-7, Comber, seed number 280; Cerro Belvedere, 12 Jan.

Prov. Rio Negro, Cerro Lopez, 30 Jan. 1934, Pard 6128 (La Plata). According to Philippi, also in Chile, in the Provinces Santiago, Made (Cauquenea),

Colchagua, Ruble (Chdh) , fl. Jan. According to Job, generally, S.W. from Argentine, Prov. Rio Negro to Chile, Prov. Llanqujhu6.

Cultivated (Comber's seed no. 280): fl. 11 May 1939, 18 May 1940; fr. 30 June 1943.

(363) Berberis chillanensis (Schneid.) Sprague ex Bean, New Flora and Silva, 5,50, fig. 16 (1932), anglice; Kew Bus. 1932,455, 1932; Job, 1.c. 44, fig. 9,1942; s p . B. snontana vm. chiuanensis Schneid., BUU. Herb. BoiSSier, (2), 5, 395, 1905

Sh& erect, to 6ft. S m puberulous, moderately stout. S p i m 4-8mm. Leavea deciduous, thin, P 1 5 x2-7 mm., entire, acute. Flowers solitary. Pedicels glabrous, 5-10 mm. oute* sepal8 3-5 x 14-24 mm., oblong-ovate. Median se@ 7-8 x 3.54 mm., oblong-elliptic. Inner se@ 8-10 x 5-6 mm., obovate. Petals 5-5.5 x 4 mm., obovate; apex subtruncate to retuse. Stumm 4.5-5mm., subequal to petals dentate. Ovules (6) 6-12. Otherwise as in B. wwntanu.

1933, Cab- 200 (La Plata).

KEY 66A Pedioels glabrous; stamens apiculate. 363a. B. chiuanensds var. chillanenais Pedicels puberulous; stamene t8rUn-h. 363 b. B. var. hirsutiw

(363 a) Berberis chillanensis vm. chillanensis Chile: Prov. Ruble, Cord. de ChUn, 1856-7, Germain s.n. (K; Type, Vienna). Argentine: Prov. Neuqudn, Pino Hachado, Casa Blana, 38" 35' S., 5 Feb. 1920, Parodi

2072, according to Job; BGo de Palo Mahuida, 6000 ft., 16 Jan. 1926, Comber 415a (KO.

(3633) Berberis chillanensis var. hirsutipes Sprague lx., 1932; Bot. Mag., 160

ProphylLs absent. Outer sep& H.C.C. 5, ovate, obtuse, 3.5 x 2 mm. Median sepals 6 x 3 nun., ovate, obtuse. Inner sepals 9 x 5 mm., rhombo-elliptic, obtuse. Petals H.C.C. 8, obovate, 6.25 x 4 mm., entire, rounded or subtruncate. Starnem 5-56 mm., shortly dentate; apex not produced, truncate.

Argentine: Prov. Neuquh, Suangulo, 6500 ft., fl. 16 Jan. 1926, Comber 479 (Type, K) ; fr. 16 Jan. 1926, Pulmazi (Queseria), 3500 ft., Comber 415c (K).

Cultivated: fl. 22 May 1941, 15 Apr. 1943, 26 Apr. 1944; fr. 30 June 1943.

t. 9503, 1938.

(364) Berberis coletioides Lechler, Berb. Amer. Austr. 38, 1857 ; syn. B. numtana var. coletdoides (Lechl.) Schneid., Bull. Herb. Boissier, (Z), 5, 395, 1905

Stew puberulous; mature yellow-brown. Leaves entire. Pedicek 5-10 mm.

KEY 6 6 ~ S t e m stout; spines stout, 3-5-fld., 1 6 1 5 nun. ; leaves to 15 x 6 nun. ; flowera fascicled 2-3.

Stems slender; spines weak, 6 8 mm. ; leaves to 12 x 6 nun. ; flowers solitary.

(364 a) Berberis coletioides var. coletioides

364a. B. wletioides var. coletwides

364b. B. wletioides var. gracilis

Chile: h o v . Santiago, Philippi, 8.n. (Type). Not cultivated.

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Berberis and Mahonia 247

(364 b) Berberis coletioides var. gracilis (Schneid.) Ahrendt, stat.nov. ; syn. B. m- t a m var. gracilis Schneid., l.c., 1905.

Chile: Prov. Valdivia, Villa Rica, Neger, s.n., 1897 (Type, Munich).

There lie here hybrids between the Sects. Buxifoliae and Ilicifoliae. With the former they share the possessim of 6-12 ovules; and with the latterthe conspicuously long style. Discrimination is as follows.

KEY 67, to hybrids between Sects. Buxifoliae and Ilicifoliae Ovules 6-12

Flowers solitary. Flowers not solitary.

366. B. x antimianor

Stamens not, or slightly, produced; apex truncate, rounded or obtuse.

Stamens much produced, conspicuously and slenderly apiculate. 367. B. x stenophyllu

368. B. x irwinii

under 369. B. darwinii Ovules 3-5. . . compare related hybrids falling within section Ilicifoliae and placed

(366) Berberisxantoniana Ahrendt, Card. Chron., 3rd ser., 108, 25, 20 July 1940, Seamus, Card. Chron., (3), 127, 3, Jan. 1950; syn. B. darurinii var. macrophylla Hort. (ex Catalogue, Smith, Newry, Northern Ireland)

Shrub stoloniferous, spreading, and of ungainly habit. Stems terete; mature red- brown, pubescent. In te rnoh 7-17. Spinas c. 1 cm. h u e s thick, rigid, 9-15 x 6-10 mm., elliptic, mostly entire, rarely with an occasional spinule; above lustrous dark green; below much paler yellow-green, slightly papillose. Flowers solitary. Pedicels 1-2-2.7 cm. ; bracts 3 4 mm. Plowers 14 mm. diameter. Prophylk absent. Outer sepals H.C.C. 7, oblong-obovate, subobtuse, 5 x 2.6 mm. Median sepals 6 x 3 . 5 4 mm., oblong-elliptic. Inner sepals 8-10 x 6-7 mm., broadly elliptic. Petids H.C.C. 8, obovate-elliptic, 6 x 3.5- 4 mrn.; entire; glands separate, non-marginal. Sta7nens also 6 mm. long, but notably protruding beyond the convolute petals; shortly acutely dentate; apex not produced, truncate. Ovules 8-10. Berries globose, pruinose blue, 7.5 mm., excluding style 2.5 mm.

Cultivated: in Smith’s Daisy Hill Nursery, Newry, Northern Ireland, as B. darwinii x B. buxifolia.

L.A. 103, from the above original source, 0. May 1942 and Apr. 1943; fr. no record

Thisis amost distinct and fairlyrecent hybrid. Thewell-known B. x stenophylhhasleaves of textureintermediate between its parental sections (Sect. Buxifoliae, B. ernpetrifolia, and Sect. Ilicifoliae, B. darwinii); it has the many ovules of the former combined with the subracemose inflorescence and long style of the latter. The later hybrid, B. x antoniana, between the same sections (Sect. Buxifoliae, B. buxifolia, and Sect. Ilicifoliae, B. dar- winii) has leaves of the same rigid texture and very lustrous surface as the latter, but entire as in the former ; contrary to B. x stenophyllu, the inflorescence takes the solitary form of B. buxifolia ; the multiovulate and long-stylose characters are, however, similarly assooiated, thus bringing both hybrids together in their position between the Sections.

(Type, 0).

(367) Berberis x stenophylla Lindl., Gard. Chron. 1864, 460, 1864; Schneid., Ill.

Shrub 6-10 ft., with arching branches. Stems terete; mature dark red-brown, puberu- lous. Internodes 7-15 mm. Spines solitary, 2-5 mm. Leaves fascicled 5-6 ; narrowly elliptic, to linear-elliptic, 8-17 x 2-3 mm. ; entire, very revolute, subenervate; above

Handb. Laubh. 1, 298, 1904

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248 L. W. A. ~ E N D T

lustrous dark green; below much paler yellow-green. I n , 7-14-3d., racemose, 2-3 cm. Pedi& glabrous, 3-6 mm.; bracts greenish, ovate, acute, 1-75 x 0.75 mm. Flowers 7-9 mm. diameter, H.C.C. 5. ProphyUs concolorous, oblong-ovate, subacute, 3 x 1-5 mm. Outer 8epd.s 3.5 x 2 mm., oblong-obovate, obtuse. Inner seg& 6-66 x 3.25- 4 mm., oblong-obovate. Petals 56-6-25 x 3-3-25 mm., obovate, entire; cuneate with glands narrowly obovate, subacuminate, 1.3 x 0.4 mm. Stam.en.s 4 mm., shortly dentate; apex not produced, truncate. OwuZa 7-10. Berries globose, 6-7mm., pruinose blue, excluding style 1-1-5 mm.

Origin: in the nursery of Mesm Fisher and Holmes, Handsworth, near Sheffield, about 1860.

Cultivated: L.A. 21, fl. Apr. 1942; fr. June 1942. Many further hybrids have been raised from 23. xstenophyUa (vide Ahrendt, ‘The

Berber& stenophyllu hybrids’, J . R. Hort. SOC. 74, 36-40, Jan. 1949); these may be dis- tinguished as follows.

KEY 67a Leaves all narrow, 1 : 4-8, and revolute.

Leaves entire. Flowers not red in bud, or externally when open. Flowers golden yellow. H.C.C. 5-6.

Petals entire and shorter than inner sepals. Leaves not exoeeding 20 x 3-5 mm. Leaves to 40 x 7-8 III~TI.

367. B. x stenophyk 367a. Var. btifolia;

Petals emarginate and longer than inner sepals. 367 b. Vm. autumdia Flowers orange, H.C.C. 7-8.

Leaves dark green, lustrous; flowers H.C.C. 8. Racemes 4-7-fld.; 10-15 mm. long; petah shorter than inner sepals.

Racemes 7-14-fld., 15-25 mm. long; petals longer than inner sepals. 367c. Var. rig&

367d. Var.compacta

Leaves dull blue-green; flowers H.C.C. 8. 367e. Var. g h u c a Leaves of shoots coloured; flowera H.C.C. 7. 367 f. Var. pkturata

Petals shorter than inner sepals. 367g. Var. wrallina Petals longer than inner sepals. 367h. Var. corallina c0m;pacta

Leaves tridentate. 367j. Var. reflexa

Leaves partly m o w , 1 : 5-6, entire and revolute and partly broad, plane, tridentate,

Flowers not red in bud, or externally when open, golden yellow, H.C.C. 6; petals

Leaves otherwise, diverse; petals shorter than inner sepals.

Flowers conspicuously red in bud, and to some extent externally when open.

Leaves not all narrow.

or spinose.

longer than or equal to inner sepals. The spinose leaves with (1-) 2-3 marginal spinules. The spinose leaves tridentate, with only 1 marginal spinule each side.

367k. Var. diver,@folia;

Shrub, habit loose, to 3-4 ft. 3671. Var. gracilis Shrub, habit compact, to 1 ft. 367m. Var. gracilia nam

petals shorter than inner sepals. 367n. Var. semper$orens Flowers red in bud, and more or less so externally when open, orange, H.C.C. 7;

Leaves all broad, 1 : 2-5-3-5, tridentate, plane. Racemes 68-fld. ; petals shorter than inner sepals.

Flowers not red without; inner sepals 4 mm. Flowers conspicuously red without; inner sepals 6 mm.

367p. Var. Brilliant 367q. Var. coccinea

367r. Var. Crawley Gem Racemes 7-14-fld.; flowers red without; petals longer than inner sepals.

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Berberis and Mahonia 249

There can be seen in the above key a gradual transition in leaf shape and margin from B. empetrifolia towards B. &&nii. In both these species the petals are shorter than the inner sepals. There is no record of the source of the longer petals in some of the hybrids.

(368) Berberisxirwinii Byhouwer, Mitt. Dtsch. Dendr. Ces. 1923, 253, 1923; syn.

Shrub compact, dense, 3-5ft. Stems terete, dark brown; young shoots puberulous; mature glabrous. Internodes 7-13 mm. Spines 1-3-fld., weak, 3-6 mm. Leaves 12-20 x 3-5 mm. ; spathulate-cuneate; tridentate; above lustrous dark green; below, sometimes a t first grey, pruinose, Gnally dull, paler yellow green. Racemes 44-fld. Pedicels 4 4 mm.; bracts 3 4 mm. F1?4nuers H.C.C. 7, not red without. Prophylls con- colorous with sepals, ovate, acuminate, 3 x 1.5 mm. Outer sepals 5 x 2.75 mm., narrowly oblong. Inner sepab 7 x 3 mm., oblong. Petals oblong-obovate, 5-5 nun. long, the outer series 3-36 mm. broad; the inner series 2-26 mm. broad; entire or subretuse; glands indistinct. Sturnens 4 4 mm., edentate; apex with conspicuously slender apiculate connective, 0.75-1-75 mm. long. ovuh 6-9. Berries globose, pruinose, 6-7 mm., ex- clutling style 1-14 mm. long.

B. x stenopkylla var. irwinii Hort

Cultivated: fl. 23 Apr. 1939, 5 May 1944; fi. Aug. 1944. Byhouwer cites the synonymy with B. x stenophyUu var. irwinii for his hybrid, which

is printed ' B. inwinii' ; a brief description is given but no specimens are cited.

Sect. Ilicifoliae Schneid., Bull. Herb. Boissier, (2), 5, 802, 1905 Leaves evergreen, almost always with hypoderm; margins dentate. Ovules 2-5 (only

in .B. wmberi to 10). Fruit black, mostly pruinose blue. Style always conspicuous, 1 6 - 5 mm. long.

C!hile : Provinces Valparaiso, Santiago, Curico, Talca, Ruble, Concepci6n, Bio-Bio, Antuco, Malleco, Cautin, Valdivia, Chiloe, Magellan.

Argentine: Provinces Cbrdoba, Buenos Aires, NeuquBn, Rio Negro, Chubut, Santa Crux.

TJruguay : Prov. Paysandu. Brazil. General range, 32"-56" S. lat.

J ~ Y 68, to Subsects. of Sect. Ilicifoliae Spines weak or absent, at most 1-9 mm. long; stems either glabrous and pale yellow, or

pubescent and dark red or dark brown. Stems pubescent, dark, red or brown; inflorescence (7-) 1@30-fld.

Stoms glabrous, pale yellow or grey; inflorescence (1-) 6--8-fld. Subsect. DARWINIANAE Subsect. EUILICIFOLIAE

Subsect. CHILENSES Spines strong, 1-3 em.; stems mostly dark red and glabrous.

Subsect. Darwinianae Ahrendt, subsect. nov., inter sectionem ramulis pubescentibus,

Stems pubescent, dark, red or brown. Spines weak, 3 (-5)-fld., 1-7 mm. InJEOrescem

Chile : Provinces Ruble, Concepcih, Bio-Bio, Antuco, Malleco, Cautin, Valdivia,

Argentine: Provinces NeuquBn, Rio Negro, Chubut, Santa CNZ. Range 37'42" S. lat.

atrorubris vel atrobrunneis distinguenda

( 7 4 10-30-fld., racemose. Berries pruinose blue. Style 3-5 mm. Ovules 3-5.

Chiloe.

Page 250: Berberis and Mahonia : A taxonomic revision

250 L. W. A. AHEENDT

KEY 69, to Subsect. Darwinianae Sbms terete.

Leaves not orbicular, entire or with 1-3 sphules to each margin; stems dark brown. Flower diameter 10-12 mm.; stamens not, or scarcely, produced; leaves 5-16 (-20) x

Flower diameter 16-18mm.; stamens conspicuoudy produced; leaves 15-30~ 3-8 (-10) mm., broad, 1 : 1.5-2.

5-10 mm., narrow, 1 : 3.

369. B. darwinii

310. B . l o l o g d 371. B. coatulata

stems sulate. 372. B. lcni&ii Leaves orbiculax, many toothed; stems dark red.

(369) Berberis darwinii Hook., Icon. PI. 7, t. 672,1844; Bot. Hag. 77, t. 4590, 1851;

Shrub densely branched, 5-8 ft. Stem all very pubascent ; young shoots green ; mature dark brown. Internodes 7-25mm. S p i w 5-1Omm. Leaves 10-20 x 5-12mm., thick, rigid; mostly hexagod, with apex acute and muoronate, and cxmh margin with two spinose teeth; base cunate; enervate; above luetrous dark green; below very much paler, scarcely lustrous, yellow-green. Racemes (5) 10-30-fld. Pedkh red, 6-10 mm.; bracts 3-5 mm. F k r s H.C.C. 8. Outer se@ 3-75 x 2 mm., reddish, ovate, acute. Median se@ 4 4 x 2.5-3 mm., elliptic-ovate, obtuse. Inner sepak 6.5-7.5 x 4-5 mm., oblong- obovate. Petals 5-5-6 x 3-3.5 mm., elliptic-obovate, rounded; entire; basalglanda oblong- orbicular, 0-5x0.4mm. Stanzene 5-5mm., not, or scarcely, p d u d and rounded truncate. 04yUles 4. Berries globose, 6-7 mm., pruinose blue. Style 3 mm.

Job, Rew. Mus. La Plata (N.S., sect. Bot.), 5, 63, 1942. (Eg. 61)

KEY 69a

Racemes formal, 12-30-fld., 4-6 cm. long. Racemes subfascicled, or subumbellate, 5-15-fld., only 1-3 cm. long.

379a. B. darwinii var. &m’nG

Leavea lustrous above; shrubs 5-8 ft. high. Inner sepals 6-7 mm.

Flowers orange throughout. 369 b. B. darwinii var. rnug&niCa Flowers notably red without, partly suffused red.

Leaves to 18 x s 9 mm. 369c. B. darwinii var. pendula Leaves to 18 x 14 mm. 3693. B. darwinii ‘Flame’

369e. B. danvinii ‘Fireflame’ Cf. 370. B. x lologe.rzsiS

Leaves spinose-serrate. 369f. B. &mint% vm. prostrata Leaves entire. 3698. B. darwinii var. naw

Inner sepals 8-10 mm. Leaves to 27 x 6 mm.; stamens truncate. Laves to 27 x 9 mm.; stamens produced.

Leaves dull above; shrubs 1-4 ft. high.

For the hybrids 369, c-g, vide Ahrendt, J. R. H w t . Soc. 74,3640, Jan. 1949.

(369a) Berberis darwinii var. darwinii Chile: Prov. Valdivia, Bridges, s.n. (0). Lindley refers to this collection. Chiloe Island, Darwin, 8.n. (Type) ; seed only, Lobb, s.n. Province Antuco, Poeppig ( ?), 1828, according to Schneider. Prov. Concepci6n, 1893-6, Neger, sm., according to Schneider. Provinces Ruble, Bio-Bio, Malleco, Cautin, to Prov. Valdivia, Rio Palane, according

Argentine: Prov. Rio Negro, Bariloche, Cabrera (La Plata). Prov. Chubut, Patagonia, seed only, Lobb, s.n. Cultivated: according to Lindley, introduced by Lobb’s seed collections cited above,

to Philippi.

and now one of the most widely cultivated Berberis in England.

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Berberis and Mahonia 251

(369b) Berberis darwinii var. magellanica Ahrendt, J. R. Hort. SOC. 74, 40, Jan. 1949, nom.nud.; a B. darwinii var. darwinii inflorescentiis tantum 5-10 (-15)- floratis, 1-3 cm. longis, subfasciculatis vel subumbellatis distinguenda

Argentine: Prov. Santa Cruz, near Straits of Magellan, King, 8.n. (Type, 0). Yrov. NeuquBn, Lago Lolog, Comber 803 (K). Cultivated: L.A. 5, fl. Apr. 1944; fr. June 1944.

(370) Berberis x lologensis Sandwith, Kew Bull. 1928,108,1928; Comber, Card. Chron. (3), 89, 334, May 1931; Job, l.c., 1942

#terns very pubescent, slightly angled, dark brown. Spines W-fld., 3-7 mm. Leaves iu m B. darwinii, but to 3 x 1 cm., mostly spathulate-obovate, tridentate by apex (but on the shoots larger and with 3-4 marginal teeth each side). Inflorescence umbellate- racemose, 3-7-fld. Pedicels 5-15 mm.; bracts 2.5-3.5 mm. ProphyuS suffused H.C.C. 713, ovate, acute, 3 x 1-5 mm. Outer sepals H.C.C. 8, elliptic, 6.5-7.5 x 3 4 mm. Inner sepals concolorous, 10 x 6 mm., oblong-obovate. Petals H.C.C. 9, obovate, 65-76 x 3-5-5 mm. Stamens 5.5 mm., much produced (1 mm.), truncate. Owules 34. Berries pruinose blue, globose, f3-7 mm. Style 3 mm.

Argentine: Prov. NeuquBn, Lago Lolog, 3400 ft., 20 Feb. 1927, Comber 1061 (Type, K). 'A possible hybrid between Comber 797, B. linearifolia, and Comber 803, B. dam-nii. Three plants found. Habit intermediate; growing with the other two species.'

Chltivated: L.A. 432, fl. 26 Apr. 1944; fr. Aug. 1944.

(371) Berberis costulata Gandoger, BUZZ. SOC. Bot. Fr. 59 (1912), 705, 1913 Sterns red, tomentose. Leaves thick, orbicular; base cuneate; apex obtuse; margins

with many spinose teeth; midrib elevated. In@wescenCes racemose. Pedicels 10-12 mm. Berries ovoid. Style 4 mm.

C!hile: Chiloe Island, by Quell6n, Skottsberg 310 (Type). Not in cultivation.

(372) Berberis knightii (Lindl.) K. Koch, Dendr. 1,413, 1869; syn. Mahonia knightii hod. ex Lindl., J. Hort. SOC. Lond. 5 , 20, 1850

Stems red-brown, sulcate. Leaves suborbicular, petiolate. Chile: Locality unknown. L'indley, J. Hort. SOC. Lond., 5, 20, 21, 1850, describes this species as a 'tufted, erect,

unattractive shrub, scarcely exceeding 1 ft. in height, with slender downy shoots, and orbicular, somewhat cordate flat leaves, fringed with slender spines, and seated upon downy stalks, more than twice as long as themselves; they are pale-green, and only a little darker on the upper than on the under side.. . .The spines. . .are broad and deeply divided in a palmate manner.' It was, he says, found in Knight and Perry's nursery, and waa possibly introduced to cultivation there by Anderson in King's expedition to the Magellan Straits.

This description reminds one, as regards the leaves, of B. hakeoides, which, however, is a tall shrub ; has glabrous, yellow, stems ; leaves grey below, and petioles only about as long as the leaves; and foliaceous spines. It may also be compared with B. congestiJEora.

Subsect. Euilicifoliae Ahrendt, subsect.nov., inter sectionem, ramulis glabris pallide flavis ; inflorescentiis 2-6 (-9)-floratis, fasciculatis vel subumbellatis distinguenda

Spines absent, or weak, 1-5-fld., 1-7 (-9) mm. long. Stamens edentate. Ovules

Chile : Provinces Concepcih, Valdivia, Magellan. Argentine : Provinces Cbrdoba, Buenos Aires, NeuquBn, Santa Cruz. Uruguay: Provinces Paysandu. Brazil. Range 30"-53" S. lat.

3-5 (-10). Style 1-5-3 (-5) mm.

Page 252: Berberis and Mahonia : A taxonomic revision

252 L. W. A. AHRENDT

KEY 70, to Subsect. Euilicifoliae Leaf margins with 1-3 (-0) coarse spinose teeth; lamina elliptic, 1-3 cm. long; leaves

Spines absent; inflorescence 2-3-fld., fascicled; pedicels 2.5mm.; ovules 10; style

Spines present; inflorescence 3-8-fld., umbellate or subracemoee; pediceh longer,

fascicled.

3-5 mm.

glabrous; ovules 2-6; style 1.5-3 mm.

373. B.deri

Pedicels 5-9 mm. ; stems subterete. Pedicels 1-2 cm. ; stems sulmte.

374. €3. ru8dfolia

Infioreacenm umbellate; bracts one-third to one-quarter as long as pedicels;

Leaves with mar& spinules and teeth each 2-3 mm. long; bracts 4-7 mm. ;

Leaves with marginal spinules and teeth each 1 mm. long; bracts 2-5 mm.;

Inflorescence subracernose; bracts half as long as pedicels; leaves almost dull

LBaf margins with 10-20 6ne spinules; lamins hoeolate, 4-10 cm.; leavea mostly

leaves very lustrous above, with attenuate base.

375. B. ilicgolia flower diameter 18-22 mm.

flower diameter 12-16 mm. 376. B. pseudo-ilic$ollia

above, with base contracted. 377. B. aubantarctica

solitary. Spin= absent; leaves without hypoderm. 378. B. mgeriana Spines pment; leaves with hypoderm. 379. B. peurcei

(373) Berberis comberi Sprague et Sandwith, Xew BUU. 1927, 175, 1927; Job, Rev.

Shrub 1 4 ft. Intemzodes 2-4 cm. Xtem terete. Spines absent. L a v e s very thick and rigid, 1.7-3.5 x 1-2.5 cm. ; oblong-elliptic; apex mucronafe; base contracted to a petiole 2-5 mm. ; margins 1-3 (4)-spinose (1-3 mm.)-denhte (1-3 mm.), distant 5-10 mm. ; above almost dull pale green, below greyish pruinose, subenervate. FZmuem 14-18 mm. diameter, fascicled 2-3. P e d M 2.5 mm.; bracts scarcely 1 mm. Sep& 5 in number; outer 2 elliptic-orbicular, 4-5-43 x 4 4 . 5 mm. ; inner 3 elliptic-obovate, 7-10 x 4 - 5 7 mm. Petals 5 in number, similar and subequal to inner sepals. Stanzens produced, apiculate ovules 8-10.

Argentine: Prov. Neuquhn, Cerro Lotena, 3000 ft., 24 Sept. 1925, Comber 49 (Type, la). General distribution 37O-41" S. lat.

Cultivated: L.A. 430 (derived from seed of Comber 49). It has not yet flowered.

M m . La Plata (N.S., sect. Bot.), 5, 53, fig. 13, 1942. (Fig. 62)

(374) Berberis ruscifolia Lam., Ill. Qen., t. 253, f. 2; Poir. in Encycl. M&h. Bot. 8,

Stems terete. Leaves thick, rigid, 2-4 (-6) x 0-7-1.3 (-2) cm., oblong- or spathulate- obovate ; cuneate, subsemile; acute, mucronate; margins 1-2-spinose (1-2 mm.)-dentate ( 2 4 mm.); above lustrous pale green, subenervate; below lustrous, scarcely paler green, veined, sometimes openly reticulate. InJorescence M-fld., umbellate-subfascicled, 1.5- 2-5 cm. long. Pedicels glabrous, 5-9 mm. ; bracts 1 - 5 2 mm. Stanzens not produced, truncate.

619, 1809; Eichler, Mart. Fl. Bras. 13 (i), 232, 18fX

Brazil. According to Eichler, in agro Bonariensi, Commerson, 8.n. Uruguay: Prov. Paysandu, Concepcicin on the Argentine border, Lorentz 1065 (0). Argentine: Prov. Buenos Aires, Tweedie 183 (0). Prov. Cbrdoba, Hieronymus 34,94; Lorentz 158,430; Chacra de la Merced, Hierony-

mus 588; Rio Zebrtllos, Sierra Chica de C&rdoba, Galander, s.n., according to Schneider. Prov. San Luis, Cerco de Rio Grande, Galander, s.n.; Partido h u l , Boca de Las

Sierras, Oeten 213, according to Schneider. Prov. Chubut, Rio Chico, Lorentz, s.n., according to Schneider.

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Berberia and Mahonia 253

Cultivated: 1950, 1954, seen in the University Botanic Garden, Oxford. Not yet flowered. The leaves are thinner, and though stiff, a little more flexible than those of B. unnberi and B. ilicifolia.

(375) Berberis ilicifolia Forst., C m m . Goett 9,28, 1789; Hook., Bot. Mag., 73 t. 4308, 1847; Hook. f. Fl. Antarctica, 2, t. 86, 230, 1845

Il’hrub loosely branched. Stem very sulcate. Spines 3-5-fld., fairly stout, 10-15 (-20) mm. Leaves thick, rigid, 1.5-4-5 (-6) x 0.6-1-8 (-2.5) cm., oblong-elliptic; cuneate; aristate; margins (2-) 3-5 (-6)-spinose (1-2 mm.)-dentate (2-3 mm.), distant 6-12 mm.; both sides openly veined, with veins and branching veinlets clearly visible, sometimes also with less clear impressed subreticulation ; above grey-green, very lustrous ; below lustrous paler yellow-green. Inflorescence 4-7-fld., subumbellate or subfascicled. Pedicels slender, red, 15-22 mm.; bracts 4-7 mm. F k r s 17-22 mm. diameter. Prophylls absent. Outer sepals 4-5 x 2-2-5 mm., ovate, acute. Median sepals 7-9 x 4-5 mm., elliptic. Inner sepah 10-12 x 8-9 mm., obovate. Petals 9-10 x 6-8 mm., obovate, entire. Stamens 5-8 mm.,eproduced, apiculate. ovules (3-) 4 4 . Bewies pruinose blue, obovoid, 8 x 6-5 mm., excluding style 2-3 mm. long.

Argentine: Prov. Tierra del Fuego, Forster, 8.n. ( T y p ) ; Cuming, 8.n. (0). Chile: Prov. Magellan, Cape Horn, Hermit Island, 1839-43, Hooker, s.n. (0). Philippi specifies Punta Arenas in Tierra del Fuego, and gives November to March as

the period offlowering. Lindley, J . Hort. SOC. Lond. 5,9,1850, relates that a plant of this species flowered a t Kew, but died and ‘the species needs to be re-introduced’. A century later it still seems not to be in cultivation. Plants passing under this name are often the bigeneric hybrid x Mahoberberk neubertii.

(376) Berberis pseudo-ilicifolia Skottsb., Svensk. V e t . - A M . H a d . , N.S., 56, no. 5, 226, 1914, in obs.; B. sew&-dentatu Reiche, An. Univ. Chile, 88, 89, 1894, non LeChl.

Leaves 4 x 1.8 cm., elliptic; petiole 5 mm.; margins, for the apical half, 4-6-spinose (1 mm.)-serrate (1 mm.), distant 3-6 mm., with open venation, and part of the marginal vein visible. Inflorescence few-fld., subumbellate. Pedicels 10-15 mm. ; bracts 2.5 mm. Outer sepals 5-6 x 3 mm., elliptic, rounded. Inner sepals 8 x 6 mm., obovate, rounded. Petals 7 x 6 mm., broadly obovate, entire, base clawed, with separate non-marginal glands, 1 x 0.5 mm., situated above the claw. S t a ~ ~ edentate; apex produced, obtuse or rounded. Style 2-3 mm.

Chile: Prov. Valdivia, Lechler 3232 (Type). Not in cultivation.

(377) Berberis subantarctica Gandoger, Bull. SOC. Bot. Fr. 59, 705, 1913 According to the author, this differs from B. ilicifolia, in the less attenuate bases to the

leaves which are duller and shorter; in the inflorescence which is subracemose and not umbellate; in the bracts which in fruit are +g instead of the length of the pedicels; and :in its shorter style.

Argentine: Prov. Santa Cruz, Patagonia, or Tierra del Fuego, Skottsberg 293 (Type). A little known species not cultivated.

(378) Berberis negeriana Tischler in Engler, Bot. Jb. 31,640, 1902; reimpr. in Tischler, Berberid. und Podophyll. 49 1902

X t e m sulcate. Leaves 5-10 x 2 4 cm., narrowly ovate; apex acute or acuminate, margins with 10-20 spinose serrations; below scarcely paler green. Inflorescence 5-8-fld., umbellate-subracemose. Flowers c. 8 mm. diameter. Outer sepals 2-2.5 x 0-6-1 mm., ovate-hnceolate. Inner se@ 4-45 x 2-3 mm., ovate, subacute. Petals entire, as long aa

Page 254: Berberis and Mahonia : A taxonomic revision

254 L. W. A. AERENDT inner sepals. Stamens subequal in length to petals. ovules 2 (-3). Be& e p d o s e . Style 3-5 mm.

Chile: Prov. Concepci6n, Neger, 8.n. (Munich). Prov. Valdivia, Villa Rim, Neger, 8.n.

(379) Berberis pearcei Phil., Linnarea;, 33, 4, 1864; Reiche, Fl. Chile, 1, 35, 1895;

Shrub 3-4ft. Stem slender, 3-5mm. diameter. 8p.w absent, rarely present. Leaves ovate-lanceolate, lustrous above, dull below, 3-9 x 1-2-5 cm.; petiole 3-5 mm. long; margins with 15-18 spindose teeth. InJEorescence umbehte or subumbellate, SM-fld., 3 4 cm. long. Pedicels slender, 1 - 6 2 cm. ; bracts 2-3 mm. Prophyfi 3 x 1.5 mm. 0ut-w sepuh 6 x 2.5 mm. Inner eepuh 8 x 2 mm. Petals 6 x 4 mm. 8tamn.s edentate, produced, obtuse. ovules 3. Be- ovoid-globose, 7 mm., excluding style 2.5 mm.

Job, Rev. Mw. Lcc Pzata (N.S., sect. Bot.), 5, 63, fig. 17, 1942

Chile: Prov. Vddivia, Roquete de Rtmco, Pwce, s.n. (Type). Argentine: Prov. Neuqudn, Cemo Colorado, Lolog, 1925-7, Comber 859 (K); Nor-

quinco, 1925-7, Comber 604 (K); Cerro Belvedere, 10 Jan. 1935, Cabrera 199; Puerto Correntoso, P d 11683. Prov. Rio Negro, Cerro Otto, Bariloche, 7 Jan. 1935, Cabrera 132, according to Job.

Subsect. Chilenses Schneid., Bull. Herb, Bokwkr, (2), 5, 801, 1905 Shm dark red, &brow. S p i ~ stout, 1 - 6 3 ~ m . long. Chile: Provinces Valparak, Santiago, Curico, Talca. Range 32"-36" S. lat. This subsection waa 0rigir.d~ included by Schneider as a subsection of the succeeding

Sect. Laurinm, but it appears more properly to belong here. The few species which it contains are known only from the very brief descriptions, all given by their authors more than a century ago. !l!heiw are just sdicient for the provision of a key which should be useful for identification when fresh materid is available.

I(EY 71, to Subsect. Chilenses Leaves narrow, 1 : 4, each margin having, mostly, only one spinose tooth; racemes

Leaves broad, 1 : 2, each margin having 3-5 spinose teeth; racemes 3 cm. or more in subfascicled, 1-5-2 om., shorter than the leaves. 380. B. brachybotria

length, longer than the leaves.

h sves obovate, broad-t by 8px. Leaves thick, with hypoderm.

Leaves below grey, pruinose; fruit globose. 381. B. chile& Leaves below yellow-green, epruinose; fruit ovoid. 382. B. ferox

383. B. fmgmna Leaves thin. 384. B.diffusa

(380) Berberls brachybotria C. Gay, FZ. Chil., 1,81 ,1W; syn. B. gayi I(. Koch, Hort.

Leaves lanceolate, to 20 x 5 mm., margins mostly each with 1 tooth. Rmnzes sub-

Chile: Prov. Vdparaiso, environs of Vdpmaim, according to Philippi.

(381) Berberis chilensis Gillies ex Hook., Bot. Misc. 3, 135, 1833

Leaves elliptic, or rhombo-elliptic, broadest at their mid-points.

Dendr. 19,66, 1853, non Citerne

fascicled, 1 - 6 2 cm. long.

Chile: Prov. Tdca, Gillies, s.n., according to Hooker. Prov. Vdparaiso, Bridges, s.n., acwrding to Hooker. According to Philippi, to be found in dry plains in the central provinces of Chile. Seed

waa collected about 1926 by Clarence Elliott who raised from it two plants which subsequently died.

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Berberis and Mahonia 255

(382) Berberis ferox C. Gay, Fl. Chil., 1, 80, 1845; syn. B. chilensis var. ferox (C. Gay) Reiche, FL Chile, 1, 38, 1895

Chile: Prov. Santiago, according to Philippi.

(383) Berberis fragrans Phil. ex Reiche, FZ. Chil., 1 , 38, 1895. Chile: Prov. Curico, Vichuqubn, according to author.

(384) Berberis diffusa C. Gay, 1.c. 80, 1845; syn. B. chilensis var. diffwa (C. Gay) Reiche, 1.c. 38, 1895. 1.c.

Chile: Prov. Santiago, 'Fue observada una vez en la misma ciudad de Santiago'.

Sect$. Trigonae (Schneid.) Ahrendt, stat.nov. ; syn. Sect. L A ~ A E , subsect. Trigonae Schneid., Bull. Herb. Boksier, (2), 5,669,1905 ;ramulisvalde sulcatis, glabris, annotinis pallide flavis ; foliis sempervirentibus, oblanceolatis, integris, subenervatis, supra nitidis, subtus cinereis pruinosis papillosis ; idorescentiis 4-15-floratis ; pedicellis 1.54 cm. longis; staminibus edentatis, apice non productis, obtusis vel truncatis; fructibus caeruleo-pruinosis; stylis 2-5 mm. longis.

Chile : Provinces Talca, Ruble, Concepcih, Antuco, Cautin, Valdivia. Argentine: Prov. Neuqu6n. Range 35'40" S. lat.

&Y 72, to Sect. Trigonae Lea\ es narrow, 1 : 4; inflorescence racemose-subfascicled, or subumbellate, 8-15-fld.,

4-5 cm. long; pedicels 18-25 mm. long; ovules 3; fruit ovoid, 10 x 4 mm.; style 1.5- 2 Inm. 385. B. trigona

Leaves narrower, 1 : 5-6; inflorescence fascicled, 44-fld.; pedicels 2-4 cm.; ovules 5-7; fruit ellipsoid, 8 x &-7 mm. ; style 3-5 mm. 386. B. l inemi fo lk

(385) Berberis trigona Kunze ex Poeppig et Endlicher, Nova Gen. 2, 63, t. 187, 1839 Shrub 4-6 ft. Spines 3-fld., palmate. Leavas acuminate, plane, 3-5 x 04-1-25 cm.

Pedicel8 16-26 (-3.5, below) cm.; bracts 2-3 mm. Fhuer8 orange. Prophylls 2 x 1 mm., whitish, ovate, acuminate. Inner sepals 6 x 2.25 mm., narrowly obovate. Petals 5 x 2.5 mm., obovate, entire.

Chile: Prov. Antuco, Poeppig 108, 860, 861 (Type). Prov. Talca, according to Reiche. Prov. Concepci6n, according to Reiche. Not in cultivation.

(386) Berberis linearifoh Phil., Linnaea, 28, 663, 1856; Reiche, h., 39, 1895; Comber, Card. Chron. p. 4,2 May 1931; Job, Rev. H w . La Plata (N.S., sect. Bot.), 5, 41, fig. 8, 1942; h l y , Bot. Hug., 161, t. 9526, 1938. (Fig. 63)

Shrub 4-8 ft., sparsely branched, habit loose. Intem&!e.s 2-3 cm. h v a oblong- obovate to oblanceolate; apex obtuse, mucronate (1-1.5 mm.) ; margins slightly revolute. Pedicels red, slender. ProphyUs H.C.C. 14, ovate, acute, 2-2-5 x 0.8-1-5 mm. Outer sepah concolorous, 3-3-25 x 1-1-25 mm. Median sepah 7 x 2 mm., lanceolate. Inner sepah 10 x 44.5 mm., obovate-elliptic. Petals H.C.C. 11, obovate-elliptic, 7-8.5 x 3-4 mm., apex rounded, entire; base cuneate with indistinct glands. Stamens 5.5mm. Berries ellipsoid, 8-10 x 6-7 mm.

KEY 72A Spin= 3-fld., subpalmate, 3-8 nun. ; leaves 2 5 (-6) x 04-1 cm.

Spines absent; leaves P 7 em. long. 386a. B. linmrifolia vm. l i w r i f o l i a

3863. B. linearifolia vm. h g v o l i a

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256 L. W. A. AERENDT

(386a) Berberie linearifolia var. linearifolia

Neuqudn, 5000-6000 ft., Clarence Elliott 601 (K). Chile: Prov. Valdivia, summit of the Puerto Blest Pass fkom Chile to Argentine, Prov.

Prov. Cautin, Vol& Llaima, 3600 ft., Werdermann 1277 (E). Argentine: h v . Neuqudn, Lago Lolog, 3400 ft., 25 Nov. 1926, Comber 797, loosely

erect, in moist shady woodland (K). According to Lechler, in the Cord. Nahuelbuta of h v . Valdivia, and Straits of Magellan. Photograph in Gar& IUust. p. 618, 28 Nov. 1941.

Cultivated: LA. 63 (Comber), fi. Apr. 1937; fr. June 1937; LA. 406 (Clarence Elliott), fl. Apr. 1944; fr. June 1944. A seedling from the latter introduction has been named (by Mr W. J. Marchant, who raised it) (386c) B. linecGri,folia var. Jewel. It is much more vividly and persistently red in bud, and has smaller leaves, and ovoid fruit, 9-10 x 6- 6-6 mm., excluding style 2-5 mm. Examined (L.A. 437) in fl. Apr. 1945; in fr. 20 June 1945.

(386b) Berberis linearifolia vax. longifolia (Reiche) Ahrendt; syn. B. trigom var. limg~oliu Reiche, Fl. Chile, 1, 36, 1895

Chile: Prov. Ruble, Valle del estero del chillsn.

KEY 73, distinguishing between Sects. Ilicifoliae, Trigonae and Laurinae Stems pubescent. Spines weak, 1-7 mm. long. Sect. ILICIFOLIAE (Damuinianae) Spines strong, 1-2 cm. long. Stems yellow ; leaves reticulate ; racemes 2.5-3-5 cm. 3%. B. weberbaueri Stems purple; leaves scarcely veined; racemes 5-8 cm. 404. B.peruviana

Stems glabrous. Stems dark, red to purple.

Leaves entire. Leaves scarcely reticulate; flowers 1.5 cm. diameter; ovules 3-5. 394. B. p e m s a Leaves very reticulate; flowers 4-6 mm. diameter; ovulea 1-2. 395. 23. f w w m ~ 1 a a :

Leaves with 3-5 teeth to each margin. Sect. ~ C I F O L I A E (Chilemes) Leaves with many teeth to each margin. 403. B. trollii

Leaves dentate.

Stem pale yellow. Leaves entire.

Stems terete or subterete. Sect. LAWRINAE stems very sulcate. Sect. T~ICONAE

Spines weak, 1-9 mm. Sect. ~ C I F O L I A E (Euilicqolke) Inflorescence 1-6 (-8)-fld. Inflorescence 8-15-fld. Sect . L A W R ~ A E (E&u&nae)

Leaves spinose-serrate.

Spines strong, 1-5 cm. SeCt. LAURINAE (3?'leXUOSCM)

The following section, Laurinae, is largely, both botrtnically and geographically, between the subgroups Euaustrales and Aequinoctiales; it appeam to upset the balance of the classification to attach it to either in preference to the other. Accordingly I leave it as an Intermediate Section, separating Euaustrdes and Aequinoctiales, and provide the above additional key.

The Intermediate Sect. Laurinae Schneid., Bull. Herb. Boissier, (2), 5,669,1905 Schneider included as subsections the Trigonae and the Chilemes, which, in this

classification are immediately preceding, but he omitted the Flexuosae which he had included as a subsection of the C o r y m W . This section links and slightly overlaps the

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Berberis and Mahonia 287

two great subgroups Euaustrales and Aequinoctiales, particularly geographicdy ; botanically it leans towards the former through the Ilicifoliae.

Leave8 evergreen, mostly entire, sometimes dentate. InJlorescence (&) 8-25-fld., racemose. OzruJe.8 1-5. Stamens nearly always dentate. Fruit with conepicuous style, 16-3 mm. long. Brazil: Provinces Minas Gem, St Paul.

N. and central Chile: Provinces Antofagaata, Ca.uth, Vddivia. Bolivia: Provinces Cochabamba, La Paz. Peru: Provinces Lima, Junin, Ancachs, Puno, Apurimm, Hdnuw, Cuzco. Ecuador : Provinces Pichincha, Le6n. N. Argentine: Provinces Salta, T u c m h , Catammca, La Rioja, C6rdoba. The distribution range is: (i) in the east, 19-32" S. lat., 4&58" W. long., in Brazil and

Uruguay; and (ii), in the west, from the Equator to 30 (-40") S. lat. 65-80" W. long, in Ecuador, Peru, Bolivia and north (rarely central) Chile.

Uruguay.

KTCY 74, to Subsects. of Sect. Laurinae Spines weak, P-9 (-15) mm. long., 1-3-fld. Spines strong, (1-) 1-54 cm. long, 3 (-5)-fld.

Subsect. EULAURKNAE Subsect. FLEXUOSAE

Subsect. Eulaurinae Schneid., BuU. Herb. BAder, (2), 5, 670, 1905 Characters a.s in sectional diagnosis and key. 8km mature pale yellow, glabrous.

S. Brazil: Provinces Minas Geraes, St Paul. TJruguay. N. Chile: Prov. Antofagaata. S . Peru: Prov. Cuzco.

Pe.dic& 4-10 mm. ; bracts 2-4 mm.

KEY 75, to Subsect. Eulaurinae (Map 43) Leaves sessile, or subsede, narrow, (1 : 3-4), 6-7 cm. long, entire or with 1 tooth to each

margin. Leaves quite entire; stems terete; stamens not produced, truncate.

Leaves very lustrous above; racemes 3-6 mm. long, 6-15-fld.; fruit ovoid, pruinose blue.

Leaves ovate, obtuse, below lustrous green; petals shorter than inner sepals.

Leaves obovate, acute, at first pruinose grey below; petals longer than inner

Leaves almost dull above, apex aoute; racemea 5-10 em. long, 15-25-fld.; fruit

387. B. laurina

sepalF3. 389. B. glaucescena

globose, black, epruinose. Ovules 3-5; spine5 present. 390. B.coriacea Ovules 2; spines absent. 391. B. andreana

392. B. l i tordh

393. B. citernei

(387) Berberis laurina Billbg., Thunb., PI. Bras. Dec. 1, 8 (1817), t. 2, f. 2; E i d e r ,

Leaves with hypoderm, 3-7 x 1-2 cm., subenervate both sides. Pedicels 5-7 mm.; bracts 2-3 mm. Racemes with peduncle 5-15 mm. Outer sepals 3 x 1-75 mm. Inner sepals 5 x 4 mm. Petals 4 x 3.5 mm. Ovules 3-5.

Leaves with 1 tooth towards apex of each mar@ (tridentate) ; stems subsulcate.

Leaves petiolate (4-8mm.), with lamina orbicular, 2-4cm. long, lustrous above; margins each with 10-16 spinose teeth.

Mart., Fl. Bras., 13 (i), 230, t. 52, 1864

17 J0UIt;N. IS". S0C.-BOTANY VOL. LVII

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258 L. W. A. AERENDT

woody shrubberies, from Wdaa in this province, to Monte Video. Not cultivated. S. Brazil: Prov. Mbas Germ, Wawra 464 (K); wcording to Eichler in plains and

(388) Berberis claussenfi Citeme, Berber. et E r y t h r q . p. 152, 1892. This is described by the author as differing from B. cmiacea in its broader leaves, and

from B. Eaurina by its leaves being not papillose, and having venation impressed above. S. Brazil : Prov. Minaa Geraes, Claussen, s.n. (Type).

(389) Berberis glaucescens St Hil., FZ. Brm., p. 46, t. 10,1825 Internodes 2.5-6 cm. 8pks 3-fld., 7-15 m. Leaves 3.5-6-5 x 1-4-2-4 cm., oblong-

obovate; above grey-green; below, a t first prujnose grey, W y dull, yellow-green, epapillose; both aid- with only a few lateral veins, and the marginal vein, elevated. Racem~s 10-18-fld. Pediceb glabrous, slender, 6 7 mm.; bracts 2-3.5 mm. Outer sepals 2.5 x 1 mm., Itmceolate. 3-5 x 3 m., obovate. Stamens 2 mm., shortly dentate, truncate.

S. Brazil: Prov. Cisplathe, in syl.vulis a fbibus antiquh Lusitanicis usque ad urbem Maldonado. St Hilaire, 8.n. (Isotype, K). Not cultivated.

IWMV 8ep& 3.25 x 2.5 mm., obovak.

(390) Berberis coriacea St Hil., 1.c.; syn. B. seuowiana vax. t2tambotrys Schneid.,

I&& 3-5 cm. Leavas with hypodem, obovate, subenervate below slightly paler. BuU. Herb. Boissier, (2), 5, 801, 1905

Pedicds 7-10 mm.; bracts 3 4 mm. Racemes with peduncle 1-54 cm.

KEY 75A Leaves broad (1 : 2-53), 3-7 x 1-2-5 cm. Leaves narrow (1 : 5), to 8.5 x 1.6 om.

3Wa. B.coriaeeavax.coriaeea 390 6. B. coldacea VW. oblanceqololia

(390a) Berberis coriacea vax. coriacea

vated. S. Brazil: h v . St Paul, 1841, Sellow, s.n. (K); Curitiba. St Hilaire, s.n. Not culti-

(390b) Berberis coriacea vm. oblanceifolia Ahrendt, vas.nov., foliis angustioribus, oblanceolatis

Leaves oblanceolate, baae slenderly cuneate to petiole 1-6mm.; above dull; below pder ; both sides with hterd and margind veins and veinlets visible. See& dark purple.

S . Brazil: Prov. St Paul, Rio, fr., 1861-2, Weir 415 (Type, K). Not cultivated.

(391) Berberis andreana Naudin, &m HortkcoEe, Paris, 1899, 9, 1899 I&& c. 3 cm. Spines abwnt. Leaves fascicled c. 4; to 4-7 x 1.6 cm., or to 5 x 2 om. ;

obovate to elliptic; venation branched to subreticulate; below green. Racemes loose, including peduncle 1.5-2.5 cm. PediA 5-9 mm. ; bracts 3-5 mm. Prophylls 1.5 x 0.5 mm., lanceolate. Outer sepals 2-25 x 1.5 mm., ovate. Inner sepals 3 x 2-25 mm., elliptic. Petals 5 x 3 mm., obovate, obtuse, entire; glands c. 1 x 0-3 mm., separate, non-marginal. Stamens 3 mm. ovules 2, stipitate, with stalk about one-third as long as the o d e . Uruguay: seed collected by Edouard Andd. Cultivated (Type) and recorded as

suitable for growing in the Mediterranean region (1899). A shrub 5 ft. high after 4-5 years in cultivation. Not cultivated in England.

Page 259: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 259

(39%) Berberis litoralis Phil., Florulu atacarnensis, no. 4, 7, 1860 Stems subsulcate. Leaves to 1.6 em.; margins each with one tooth towa~ds apex;

above very lustrous; below green, epapillose. 6-15-fld. Stamens edentate. Berries ovoid. Style 1 - 6 2 mm. K. Chile: Prov. Antofagasta, Department fitoral, ad Aquam Miguel Dim dictam in

litore deserti atacamensis, 900 ft. Not cultivated.

(393) Berberis citernei Ahrendt, nom.nov.; syn. B. gayi Citerne, Berber. et Erythrosp. p . 152, 1892; Macbride, Fl. Peru, in Field Mus. Publ. Bot., 13, pt. 2, p. 673,1938, non Koch, Hort. Dendr., p. 19, 1853

Leaves with blades orbicular, 2 4 em., contracted into petiole 4-8 mm. ; margins with 10-l6 spinose serrations ; above lustrous, subenervate ; below almost dull, epapdose, helly reticulate. R a c e w c. lz-fld., 2-4 cm. long. Pedicels glabrous, c. 4 mm.; bracts puberulous.

Peru: Prov. Cuzco, Cuenca, Gay 1164, Type. Not cultivated.

Subsect. Flexuosae (Schneid.) Ahrendt, comb.nov. ; syn. Sect. C O R Y ~ ~ ~ B O S U , Subsect. Flexuosae Schneid., BUU. Herb. Boissier, (2), 5, 812, 1905

Stems mostly terete (except where occasionally recoded aa subsulcate or submgld) .

Ecuador : Provinces Pichincha, Le6n. Peru: Provinces Lima, J d n , Ancachs, Puno, Apurimac, Hdnuco, ~ C O . Bolivia: Provinces Cochabamba, La Paz. S. Brazil: without locality. Central Chile: Provinces Cautin, Valdivia. Uruguay. N. Argentine: Provinces Salta, Tucumh, Catamarea, La Rioja, C6rdoba.

Spi?tes 3 (-5)-fld., stout (1-) 1-65 cm. long. Stantens often apiculate.

KEY 76, to Subsect. Flexuosae (Map 43) Leaves quite entire.

Mature stems purple. Leaves scarcely reticulate; flower diameter 1.5 cm.; ovules 3-5. Leaves closely reticulate; flower diameter 4 mm.; ovuleg 1 (-2).

Stems pubescent; leaves with suborbicular blade; spines 1-1.5 em.

Stems glabrous; leaves with blade narrower (1 : 2 - M ) ; spines 1.5-3-5 em.

394. B. jlexwsu 395. B. m o n o v e m

Mature stems pale yellow, or whitish.

396. B. weberbuueri

Pedicels (3-) &7 mm. ; bracts 2-3 mm. ; petals longer than, or aa long as, inner

Leaves lustrous above, green below, apex acute; pedicels glabrous 5-7 nun.;

Leaves slightly lustrous below, 1-75-3 om. broad (1 : 1*5-2.5), subenervate

sepals.

petals longer than inner sepals.

or with few lateral veins; internodes 3-5 cm. ; fruit 5-6 x 6 5 mm. 397. B. valdiviunu

398. B. seUo&w

399. B.haUii

Leaves dull below, 8-15 mm. broad (1 : 3) reticulate ; internodes 1-3 cm. ;

Leaves dull above, pruinose grey below; pedicels pubescent, 3-5 mm.; petals

Pedicels 1-2 em. ; bracts P 7 nun. ; leaves with apex rounded; petals shorter than

fruit 7 x 6 mm.

as long as inner sepals.

inner sepals. Leaves thin, without hypoderm; below pruinose, grey. 400. B. cmmututa

17-2

Page 260: Berberis and Mahonia : A taxonomic revision

260 L. W. A. AHBENDT

KEY 76 (mat.) Leaves thick, with hypodezm; below epruinw, p n . InfloFescence (6) 8-14-fld., fascicled-recemose, or umbelhte-raoemose, 2-3 cm.

long; pediceh 6-16 mm.; petals much shorter than inner seph; O v u l e s 46; fruit 10-13 x 6-7 mm. 401. B. diva+

Inflorescence 12-20-fld., racemose, P 7 cm. long; pedioele 1-2 cm.; petals scarcely shorter than inner sepals; ovules 3; fruit 6-9 x 3-6 mm.

402. B. 9.mdddlii Leaves with margins spinofJe-serr&te.

Leeves lustrous above; spines only 1-2 cm. long. Leavea broad, 1 : 1-6-2.

Leavea with many mq$ntd spinulm. Leeves with few marginal spinules.

Pet& longer than inner sepale. Petals shorter than inner sepals.

Leaves dull above; spines 3-6 cm. long. Leaves IlmTOw, 1 : 6-0.

403. B. trollii

404. B.pmviana 405. B. peudo-~.nubsa

406. B.spimdosa 407. B.armata

(394) Berberis flexuosa R. et P., FZ. Peruv. et Chil., 3,62, t. 281a, 1802; Mubride,

h u m without hypodem, obovate, fG3 x 0.9-1-6 cm., including petiole, 3-5 mm.; both sides with loosely branched venation, scarcely subreticulate; below dull, paler green, papillose. Race- 8-14-fld., 3-4 cm. long. Ped+ pubescent, 4-8 mm.; bracts 2-3 mm. F k r s pale yellow. Inner sep& 7 x 2.25 mm. Petds 10 x 3 mm., entire, all the components obovate. Si?umm 7 mm., apiculate. Be& oblong-ellipsoid, 9 x 5 mm.

Peru: Prov. Lima, Cheuchin, Ruiz & Pav6n (Type). Prov. Junin, Taxma, Dombey, Weberbauer 1726, rtccording to Macbride. Not in

Fl. Peru, in Fkld .Mw. Publ. Bat., 13, pt. 2, p. 672, 1938

cultivation.

(395) Berberis monosperma R. et P., 1.c.; Macbride, 1.c. p. 677, 1938

puberulous, 8-12 mm. F h e r s small (4-6 mm. diameter, according to Macbride). h v e a dull, reticulate, papillose below, 3-4x 0.8-1 cm. Racemes 8-14-fld. Pedicel8

Peru: Prov. Lima (?), Huamatanga, Ruiz & Pav6n (Type). Prov. Junin, Tarma, Ruiz & Pav6n. Prov. Ancachs, Tdenga, 11,OOO ft., Weberbauer 2851. Prov. Puno, Agapata, 10,OOO ft., Lechler 2642. Not in cultivation.

(396) Berberis weberbaueri Schneid., Bot. Jb. 42,83,1908; ilhcbride, I.c., 680,1938 Stems angled. Spinea 3-fld., 1-2 cm. Leaves 1-5-3 (4) x 1-2-3 cm., reticulate. Racemes

GlO-fld., 2-5-36 cm. long. Pedicek 4-8 mm., puberulous; bracts 1 - 5 3 mm. Flowers 10-12mm. diameter. Petu.Zs slightly longer than inner sepals. Stantens apiculate. ovules 2.

Peru: Prov. Ancachs, Cajatambo, Ocros, 11,300 ft., Weberbauer 2680 (Type). Not cultivated.

(397) Berberis valdiviana Phil., Linnuea, 28, p. 609, 1856; and ex Reiche, 3'1. Chile, 1, 37, 18%; Sealy, Bot. Mug., N.S., t. 139, 1951

Stems with young shoots purplish at first, soon greenish yellow; mature pale yellow- brown. Spines sulcate. Leaves stiff, thick, with hypoderm, 2-5-5 x 1-3 cm., excluding petiole 1-5 (-8) mm. ; elliptic, oblong-elliptic, or oblong-ovate; entire (but partly coarsely dentate on young plants), subenervate, or with few lateral veins and some openly branched venation slightly impressed; above slightly lustrous deep green; below much paler

Page 261: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 261 yellow-green, Gnely papillose, subenervate, or with few lateral veins and veinlets slightly elevated. Racemes 10-18 (-25)-fld., 3-4-5 cm., long. Prophylk usually solitary, ovate, acute, 1 . 2 5 ~ 0.75 mm. Outer sepak (H.C.C. 6), 2 x 2 mm., broadly obovate, sub- acute. Inner sepals concolorous, 3.5-4 x 2-2.5 mm., obovate. Petals (H.C.C. s), 4A.5 x 2.5-3 mm., obovate, rounded, entire; base cuneate with separate lanceolate glands, 1 x 0.3 mm. Stanzens 3 mm., edentate; apex produced, shortly apiculate. Ovules 3-5. Berries pruinose blue.

GY 76a Leaves to 6 5 x 2-3 om. ; racemes with peduncle 1-2 cm. ; fruit ovoid.

Leaves to 5-5 x 2 cm. ; racemes sessile, or with peduncle 2-8 mm. ; fruit obovoid. 397a. B. valdiviana var. valdiwiam

397 b. B. vaI.diviana var. gmaXfolia

(397 a) Berberis valdiviana var. valdiviana Chile : Prov. Valdivia, Philippi 554 (Isotype, K). BOP. Cautin, Temuco district, Bulnes Road, 700-1500ft., fr. Dec. 1930, Clarence

Cultivated: L.A. 268,0. May 1940; fr. Aug. 1952. Photograph in Qard. Illust., p. 617, Elliott 598 (K).

26 Nov. 1941.

(397 b) Berberis valdiviana var. gracilifolia Ahrendt, var.nov., foliis agustioribus racemis subsessilibm, fmctibus obovoideis, distinguenda ; syn. B. valdiviana ‘varia con las hojas angostas’ Phil., ex Reiche, 1.c.

Chile: Pror. Valdivia, NalalhuB, 650ft., fl. Sept. 1924; fr. Jan. 1925, Hollermayer (Werdermann) 682 (Type, K). Not in cultivation.

(398) Berberis sellowiana Schneid., BUZZ. Herb. Boissier, (2), 5, 801, 1905 Stem arching. S p i m 3-fld., terete, 2-4 cm. Leaves chartaceous, to subcoriaceous,

with hypoderm, 264.5 x 0.8-1.6 cm., narrowly obovate; petiole short; both sides with lateral veins elevated, more finely openly reticulate. Racernes 12-15-fld., 3 4 cm. long, including peduncle, 1-2 cm. Petals obovate, slightly longer than inner sepals, and with base clawed. O& 3-4. Berries ovoid-globose.

Brazil: Sellow, s.n., without locality (Type, Herb. Berlin). Lhguay: Mt. St Lucia, Arechavaleta 3018 (Berlin).

(390) Berberis hallii Hieron., Bot. Jb., 20, 49, 12, 1895 Stem subangled, grey-brown. Spines 1-2 cm. Leaves to 3.5 x 2 em.; petiole 4-6 mm.

finely reticulate. Racernes sometimes compound below. Prophylb acuminate, 2-3 mm. Inner sepah 4 x 3 mm., elliptic. Petals 4 x 2 mm., oblong.

&I? 7 6 ~ Racemes subpendulous, 3-5 cm. Racemes erect, 7-10 cm.

399a. B. laallii vm. Wlii 3996. B. laallii var. wagneriana

(399a) Berberis hallii var. hallii; syn. B. M i i var. typica Schneid., BUZZ. Herb. Boissier (ser. 2), 5, 813, 1905

Ecuador: Prov. Pichincha, Quito, Jameson 11; Hall 17 (Type).

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262 L. W. A. AERENDT

(399b) Berberis hallii var. wagneriana Schneid., B d . Herb. Boissier, (2), 5,813; 1905 Ecuador: Prov. W n , Hochebene la Tmunga, 1858, Wagner, 8.n. (Type).

(400) Berberis coxnmutata Eichler, Mart. Fl. Bras., 13, pt. 1, 231, 1864; Macbride, Fl. Peru, in Field Hua. Publ. Bot. 13, pt. 2,672, 1938, excluding synonymy with B. divaricata

Inteepnodes 2-5-5 om. Spines stout, 3-fld., 1-2 cm. Xtem red-brown, soon becoming yellow-brown. h v w 2-5.5 x 0.8-143 cm., or to 6.6 x 3 cm., elliptic, mucronate; petiole 5-10 mm. ; both sides with lateral veins, and branched veinleh, both elevated, and with fine reticulation; above dull grey-green; below grey, pruinose, papiuoSe. Racemes 10-20-fld., 3-5-6 (-8) cm., including a peduncle (1-) 1-5-2 (-3) cm. long. Ped& glabrous, 6-10 mm., slender; brade at base of stems folimeous, 10-20 x 2 - 5 5 mm., but a t the apex of the stems reduced, simple, 3-5 mm. long. ProphyUs ovate, acute, 2 x 1.5 mm. Outer sep& 3.5 x 2.5 mm., ovate, acute. Inner se& 5-6 x P 5 mm., obovate. Petals 3 - 5 4 x 2 - 5 4 mm., obovate, entire; glands sepaxate, non-maxginal, 0.5 x 0.2 mm. Stamem 3 mm., dentate; apex produced, shortly apiculate. 0vuk.s 3-5.

Brazil: according to Eichler. S. Bolivia: Prov. (?), Calderillo, 10,400ft., fl. Jan. 1904, Fiebrig 3216 (K); Rosa1

Peru: Prov. Apurimac, Weberbauer 5911, according to Macbride. Prov. Hdnuoo, Sawada 78. Prov. Cuzco, Gay 2191.

6700 ft., roadside, fl. 4 Oct. 1949, Brooke 5763 (BM).

(401) Berberis divaricata Rusby, Mern. Torrey Bd. C1.4,203, 1895; and Bull. N.Y. Bot. 6%. 4,321,5 Sept. 1907

Stem lustrous. Intermodes 2-4 cm. Spines plane. Learn to 4 x 1.2 cm., or to 3.6 x 1.4 cm., obovate or elliptic; both sides openly veined; above grey-green almost dull; below dull, much paler yellow-green, finely papillose, indistinctly reticulate. Race- with peduncles 5-2Omm. P e d a stout, slightly thickened at apex. Outer s e p h 2.5 x 1.5 mm. Inner e e p h 7 x 4 mm. P e a 5 x 2 mm., spathulate, entire. Omib (3-) 4.5. Be& ovoid, pruinose blue.

Bolivia: Prov. Cochabmba, Pocona, 8200 ft., fl., 11 Nov. 1928, Steinbach 8692 (E) ; Calomi, 1 Jan. 1949, Brooke 5058 (BM); Songo, Nov. 1890, fk., Bang 863 (Isotype, K). Not cultivated.

(402) Berberis weddellii Lechl., Berb. Am. Austr. p. 21, 1857 Stem shoots h t reddish; mature yellow. I&& 3-4 cm. Spines stout, 3-fld.,

1-2 cm. Leaves to 5 x 1-8 cm., obovate; entire, subrevolute; above dull grey-green, subenervate or with few distinct lateral veins; below, dull, paler, yellow-green, papillose with lateral veins distinct, and veinlets indistinct, but both elevated. Race- with peduncle 1-2 cm. long. Outer sepds 3 x 2 mm., oblong-ovate, muminate. Inner q v a h 6-6.5 x 5 mm., obovate. Petah 6 x 4 mm., obovate. Stamena 4 mm., shortly apicdate, dentate. O& 3. Be& oblong. Bolivia: Prov. La Paz (Larecaja), near Sorata, Canamulloni, in thicketa, 9800-

10,400 ft., fr., fl. 9 Jan. 1860, Mandon 864 (K). Not cultivated.

(403) Berberis trollii Diels, Notizbl., Bot. Gart., Berl., 11, 781, 1933 Stem glabrous, terete, lustrous red-brown. Leavea rigid, 3-6 x 2-3 cm. ; petioles

1-5 mm.; margins with many spinose teeth; above very lustrous, scarcely or slightly veined, below epapillose. Racemap dense, 8-15-fld., 1 - 5 2 cm. Pedicds glabrous, 5-6 mm.

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Berberis and Mahonia 263

Outer sepals 2.5 x 2.5 mm., ovate. Inner sepals 6 x 4 mm., obovate. Petals 5-6 x 3 4 mm. Stamens 4.5 mm., apiculate. O v u h 6-8.

Bolivia: Prov. (?), Padeava, Cuevaa, in evergreen shrubberim, 11,300 ft., fl. 24 Sept. 1927, Troll 219 (Type, Berlin). Not cultivated.

(404) Berberis peruviana Schellenberg, Bot. Jb., 50 (Beibl. iii), 6, 1913; Macbride,

.Stem; mature purple, pubescent, ultimately subglabrous. Leaves to 4 x 2.5 cm ., obovate; petiole 1-7 mm.; margins with few spinose teeth; above lustrous, smooth, scarcely veined or subenervate; below dull, papillose, reticulate. Racemes many-fld., 5-8 cm. Pedicels 8-10 mm. ; bracts 4-5 mm. Inner sepals 5 mm. Petals 7 mm.

Peru: Prov. Apurimac, Convencion, near Cuzco Border, and mouth of Rio Pampas, 9100 ft., Weberbauer 5987 (Type). Not cultivated.

Flora of Peru, l.c., 13, 678, 1938

(405) Berberis pseudo-spinulosa Job, Rev. Mm. La Plata (Botcinicu, no. 41), 8, 129, 1943; syn. B. spinulosa Grisebach., in Abh., Kmig. Ces. aOtt., 19, 70, 1874, non St Hil.

iPwub 3-5 ft. high. Stem mature, yellow. Spines 3 (-raxely, 5)-fld., curved, sulcate below, 1-2 cm. long, the lateral parts being somewhat shorter than the centre. Leaves 3-5 x 1.5-2 cm., ovate to lanceolate; petiole 3-6 mm.; margins sinuate, 34-spinose (1 mm.)-serrate (1 mm.), distant 7-12 mm.; below paler green. R m w 10-15-fld., with the flowers 8-10 mm. diameter. Pedicels 8-10 mm.; bracts 1-2 mm. ProphyUs 2-5 x 2 mm., triangular. Outer sepah 3 x 2 mm. Inner sepds 5.5 x 3 mm. Petals 4 x 3 mm., with discolorous glands. Stamens 3.5 mm.; below the anthers edentate, or slightly swollen. 04uUle-s 4-7. Berries subglobose, 5-8 mm. Style 1.5 mm., in flower 2-3 mm. long in fruit.

Argentine: Prov. Salta, Department Guachipas, Alemania, Laa Pavas, 6700ft., 10 Jan. 1937, Job, s.n. (La Plata 043971, Type).

I'rov. TucumAn, Luna, 10,OOO ft., 10 Nov. 1912, Rodriguez 591. Yrov. Catamarca, Sierra de Belkn, Jan. 1872, Lorentz, s.n.; Valle de las Granadillaq

Prov. La Rioja, Cuesta de la Puerta de Pedra, Sierra Velasco, 8 Jan. 1879, Hieronymus

Prov. Cbrdoba, Sept. 1875, Berg, s.n. Generally from North of Argentine to 32" S. lat.

cerca de Yacutula, Feb. 1872, Lorentz, 8.n.

& Niederlein 72.

(408) Berberis spinulosa St Ha., Fl. Bras. Mer. 1, 45, 1825; Eichler, Mart. Fl. Bras.

Stem lustrous pale yellow. Spines 1-2 cm. Leaves 5-76 x 1-26 cm. ; petiole 4-9 mm. ; margins by apex with 5-6 spinose teeth ( 2 4 mm.); above lustrous; below green; with only a few lateral veins either side. Berries globose, 10 mm.

S. Brazil: Prov. St Paul, in the south, woods near Curitiba, St Hilaire, s.n. (Type).

13 (i), 232, 1864

(407) Berberis armata Citerne, Berberid. et Erythrosp., p. 152, 1892; Macbride, F1. Peru, Lc., 13, 668, 1938

Stems pale whitish yellow. Spines 3-5-fld., 3-5 cm., stout. Leaves 3-4 x 2-5-3.5 cm., without hypoderm; margins 5-8-spinose-serrate; veins indistinct; above dull; below papillose or subpapillose. Racemes 6-12-fld., sometimes compound below, 2.5-4 cm. Fkwet-8 6 mm. diameter.

Peru: Prov. Cuzco (?), without locality, 1839-40, Gay, s.n. (Type). Not cultivated.

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264 L. W. A. AHB,ENDT

Subgroup Aequincdales Ahrendt, foliis sempervirentibus coriaceis ; inflorescentiis paucitloratis et pistillis cormpicue stylosh, vel pistillis estylosis et inflorescentiis plenunque multifloratis

S p i m nearly always weak or absent, rarely stout. Leaves evergreen, mostly, but not always, without hypoderm. In$lorescnce. mostly many-flowered, generally paniculate, and then with fruit eatylose, or very short; otherwise few-flowered, and then with fruit often, though not always, distinctly stylose. O v d a 2-5 (-8). Berries black, with or without blue bloom.

(? Mexico), Nicaragua, Costa Rim, Venezuela, Colombia, Ecuador, Peru, Bolivia, North Chile (Provinces Tacna, Tarapaca, Antofagaata, Atamma). (20'-) 10" N. lat. to 10" (-22") s. lat. ( G O - ) 70-85" w. long. No species of this subgroup is in cultivation in England, now.

KEY 77, to Sects. of the Aequinoctiales (Maps 44,45) Inflorescence not paniculrrte.

Style conspicuously long (1-) 1 - 6 2 mm. Leavea 3-8 cm. long, broad, petiolate. Sect. LATIFOLIAE Leaves 6-20 (-30) mm. long, narrower, 1 : 2-4, subsessile.

Inflorescence 1-6 (-8)-fld., fascicled or subumbellate.

a c t , VWATAE Style absent, occasionally short and up to 0-4 mm. long, (rarely 0%5-0*75 mm.)

Ovulea (I-) 2-5; leavea somewhat narrow, 1 : ( l e g ) 2-3. Ovules 8-12; leaves broad, 1 : 1-3.

Moregcence (8-) 10-30-fld., more or less formally racemose. Flowers 1 4 2 cm. diameter with the longest component 7-11 mm.

Flowers 4-10 (-12) mm. diameter, with the longeat component 2-6 (-6) mm.

Sect. CONFERTAE Sect. AUAPA~NSES

Sect. QUIND~UENSES

Sect. GOWDOTIAE Sect. !cRUxrLLENSES

k v e a with baae abruptly contracted to an articulate petiole. h v e a with baae ouneate to decurrent petiole.

Inflorescence p&culate. k t . PANICULATAE

Sect. Latifoliae Schneid., BuU. Herb. BoisSier, (2), 5, 449, 1905 Leaves evergreen, below papillose. Style c. 2 mm. Peru, Costa Rim.

KEY 78, to Sect. Latifoliae Leaves entire ; petioles long; intlorescence fascicled. 408. B. lalifolia Leaves spinose; petiolea short; inflorescence subumbebte. 409. B. nigricam

(408) Berberis htifolia R. et P., FZ. p m v . et chi.?. 3,52, t. 282a, 1802 Stem glabrous. S p i w (1-) 3-fld., 3-8 mm. Leavrn 3-5-7-5 x 1-8-3 cm. ; oblong-

obovate; base contracted in petiole 3-6 cm.; entire; reticulate; very lustrous above. Fk~wers fascicled 3-6. Pedicels 10-18 111111. ovules 3-4. Fmit black, eprujnose, 7 x 3 mm., excluding style 2 mm.

Peru: Palo-Amarillo, Huajampe, Pav6n, 8.n. Prov. Hu&nuco, in cold grovea, Pillao, Ruiz and Pav6n (Type).

(409) Berberis nigricans 0. Kuntze, Rev. Gen. 1, 10, 1891 Leaves lustrous above; papillose below. In$oremmx subumbellate. Costa Rica: Llanuras Prov. (on boundary of C h g o Province), Volc&n Iraz6,8500-

10,OOO ft., Kuntze, s.n.

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Berberis and Mahonia 265

Sect. Virgatae Schneid., Bull. Herb. Boissier, (2), p. 148, 1905 &ems mostly dark, red or purple. Spines absent or weak, 1-9 mm., rarely to 2 cm.

Laves 6-36 x 2-12 mm., subsessile, entire, or with 1-5 marginal spinules. Flowers mostly solitary, sometimes fascicled or subumbellate, 2-6 (-8). Ovules 2-5 (-6). Xtyle conspicuous, (1-) 1.5-3 mm., s i m c a n t .

Peru : Provinces Junin, Puno, Cuzco, Huhnuco. Ecuador: Prov. Pichincha. Bolivia: Provinces Cochabamba, La Paz.

KEY 79, to Subsects. of Sect. Virgatae Petals shorter than inner sepals; flowers mostly solitary; stamens, where known, not

Petals longer than inner sepals; inflorescence 2-8-fld. ; stamens where, known, produced, produced, truncate; leaves to 12-25 mm.

apiculate; leaves to ( 1 6 ) 25-30 mm.

Subsect. RECTINERVIAE

Subsect. EWIRGATAE

Subsect. Euvirgatae Schneid. Peru: Provinces Junin, Puno, Cuzco, Huhnuco.

KEY 80, to Subsect. Euvirgatae Leaves below tomentose.

Spines absent. Spines, 1-2 cm.

410. B. tonzentosa 411. B. huenkeana

Leaves below glabrous. Stems, when mature, dark red.

Leaves to 25-30 mm., grey pruinose below, with 3-8 marginal s p h u h . Stems pubescent ; pedimls 3-5 111111. Stems glabrous; pedicels 1.5-3 cm.

Leaves to 14 mm., green below, entire.

Leaves with spinose margin; flower diameter 6-8 mm. Leaves entire; flower diameter 15-17 mm.

412. B. carinata 413. B . h u a n w M

414. B. virgata

415. B. parwzfira 416. B.lutea

Stems, when mature, pale yellow.

(410) Berberis tomentosa R. et P., Fl. Peruv. et Chil. 3, 52, t. 282b, 1802 Btenzs glabrous, angled; mature lustrous dark red. Spines absent. Leaves without

hypoderm, to 5 x 3.6 cm., ovate; petiole short; margins 34-spinose (1.5-2 mm.) distant 6-12 mm. ; below whitish tomentose. InJloreseenee 24-fld., fascicled. Pedicels 2-3 cm. Flowers 1.4 cm. diameter. Inner sepah 6 x 3.5 mm., elliptic. Petals 8 x 4.5 mm., obovate. Stamens apiculate. Berries ovoid, pruinose blue, 3-4 x 2.5-3 mm., excluding style c. 1.5 mm.

Peru : Prov. Junin, Concepci6n, Pavh, s.n.

(411) Berberis haenkeana Presl., ex Schult., Syst. 7 (ii), 1616, 1830.

ways. A very slightly known species. Said to differ from B. tomentosa, in having stout spines, 1-2 cm., and possibly in other

Peru : locality unknown.

(412) Berberis carinata Lechler, Berb. Am., Austr., p. 41, 1857: Macbride, Flura of

stems terete, pubescent, dark red. Spines absent. Leaves with hypoderm, 15-25 x 7-12 mm.; margins with 5-8 spinose serrations; above lustrous; below grey, pruinose,

Peru, in Field. Nus. Publ., Bot. 13, pt 2, p. 671, 1938

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266 L. W. A. AHRENDT papillose. Plowers fascicled 2-6. Pedicels pubescent 3-5mm. Se& (5 in number, according to Macbride), shorter than the acuminate petals. ~~~ edentate.

I(EY 80a Leaves with marginal spjnulea 1-14 mm. long. Leavea with marginal sphulea 2-2.5 mm. long.

412a. B. c4zrinat.a var. eari.rtClta; 412b. B. c4winat.a var. echinata:

(412a) Berberis carinata vm. carinata Peru: Prov. Puno, Agapata, Lechler 2644.

(412b) Berberis carinata vm. ecbinata Diels, Notdzbl., Bot. &rt. Bed. 11, 782, 1933 Peru: Prov. Cuzco, Hills of Saxaihuaman, 11,700 ft., Herrera, s.n. (Type).

(413) Berberis huanucensis (Schneid.) Macbride, FZ. Peru, 1.c. p. 674; syn. B. virgata

Stem glabrous, terete, purple. Spines absent or 3-fld. and 5-9mm. long. L e u v ~ 12-30 x 5-9 mm. ; margins 3-5-spinose-serrate towmds apex; below pruinose, papillose. Pediwh (1-) 1-5-3 cm.

var. h u m d Schneid., B d . Herb. Boissier, (Z), 5, 391, 1905

Peru: Prov. Hdnuco, Pav6n, s.n. ( T p , in Herb. Boias.).

(414) Berberis virgata R. et P., FZ. Peruv. et Chil. 3, 51, t. 281, 6, 1802. (Fig. 64) Sterns slightly angled, glabrous; mature dark red-purple. Spines solitmy, or 3-fld. and

stipitate, the portions being 2-3 mm. long. Leaves with hypoderm, 6-14 x 3-6 mm.; ovate-elliptic; attenuate, cuneate; entire; above lustrous with loosely branched, or at most openly subreticulate, venation; below green, papillose, more h e l y reticulate. Flowers fascicled 2-4. Pediwh glabrous, 1-14 cm. Flowers c. 1 cm. diameter. Sepals, five in number, obovate-orbicular, 4 x 2-5-3 mm. Petals, five in number, obovate, 7 x 3-35 mm., entire. Sturnem 5 mm., edentate, produced, apiculate. Ovules 2. Berries obovoid-ellipsoid, 6 x 4 mm., excluding style 16-2 mm.

Peru: Prov. Hu4nuc0, woods, H u m - h w i , Mathews 859; Huarica, Ruiz and Pav6n

Prov. Cuzco, above Yammanche, 13,300 ft., Weberbauer 4946. (Type).

(415) Berberis parviflora Lindl., J . R. H&. Soc. 2,243,1847; 5,6, 1850 Stems yellow. Spines P7mm. , 3-fld. h v u to 3.7x1-2cm., spathulate; base

cuneate; margins with 1 spinose (Zmm.)-serration (3mm.) by apex leaving a lobe (7.5 mm.), mucronate (3 mm.); below green. F h r s fascicled 6-6. Pediwh 2-3 om. Kbwers very small.

? Peru: type cultivated fiom a plant ‘presented to the R.H.S. when small, about 1844, by Messrs Lee of Hammemmith who were not a w m whence it came. Its resem- blance to B. virgatiz raises a suspicion that it is mme South American species; a true, slender, evergreen, perfectly hardy and rather handsome.’ Lindley’s figure does not show a very significant style.

(416) Berberis lutea R. et P., Pl. Peruv. et Chil. 3,51, t. 280; Micbride, l.~., 676,1938 Stem slightly angled, puberulous; mature pale yellow-brown. Spines absent. Leaves

with hypoderm, 12-25 x P 9 mm.; oblong-obovate; apex mucronate; base with short petiole; entire; above very lustrous, below scarcely paler green; venation both sides openly branched, not reticulate; the leaves arising in fascicles at the apex of scaly shoots, 1-1-5 cm. long, &rising from the nodes of the stems. P k r 8 fascicled 3-6 (-8). P e d i d

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Berberis and Mahonia 267

puberulous, 8-16 mm. Flowers 15-17 mm. diameter. Outer sepals reddish. Inner sepals yellow, obovate, 4.5 x 2.5 mm. Pe.tu.h 9 x 4.75 mm., obovate, entire; glands oblong, prominent. Stamens 8 mm., edentate; produced, apiculate. Ovules 6. Berries almost epruinose, 9 x 5.5 mm., excluding style (1-) 14-2 mm.

Peru: Prov. Amazonas, according to Schneider, Chachapoyas, Mathews, 8.n. Prov. HuAnuco, according to Ruiz and P a v h , near Panao, Pillao, Chaclla and Muiia.

Subsect. Rectinerviae Ahrendt, subsect.nov., petalis sepalis interioribus brevioribus ; staminibus plerumque non productis sed truncatis; floribus plerumque solitariis ; foliis 12-25 mm. longis.

Ecuador: Prov. Pichincha. Peru: Prov. Cuzco. Bolivia: Provinces Cochabamba, La Paz.

KEY 81, to Subsect. Rectinerviae Stems glabrous.

Pedicels glabrous, 1-5 mm. ; ovules 2; flowers 80litcrry. Leaves narrow, 1 : 3-5, lustrous, entire; pedicels 3-5 mm.; flowers 8 mm. diameter;

Leaves broad, 1 : 2.5, dull, margins with 3-5 spinules; pedicels 1-2 mm.; flowers ovules stipitate. 417. B. rectinervia

12 mm. diameter; ovulea sessile. 418. B. ruabyana

Leaves entire ; flowers solitary. 419. B. cliflcwtw&8 Leaves spinose; flowers fmcicled. 420. B.pavoniana

Pedicels pubescent, 6-16 mm.

Stems pubescent. Pedicels 5-7 mm., pubescent; flowers solitary, 9-10 mm. diameter; style 1-5-2 mm.

Pedicels 10-15 mm., glabrous; flowers fascicled, 14 mm. &meter; style 0.5-0.75 mm. 421. B. phyllamntha

Cf. 422. B. qwweanu

(417) Berberis rectinervia Rusby, Mem. Torrey Bot. C1. 3, no. 3, 5, 1893 Stem subterete, or subangled, purple, verruculose. Internodes 5-10 mm. Spines

absent, or very weak, 2-4 mm. Leaves with hypoderm, 7-12 x 2-3.5 mm.; narrowly obovate, sessile, entire, lustrous; below green, papillose; both sides subenervate. Flowers solitary. Pedimh glabrous, (1-) 3-5 mm. Outer s e p k 4 x 1.5 mm., oblong-obovate, subacute. Inner sepals 5 x 3 mm., oblong-elliptic, obtuse. Petals 4-25 x 3 mm. ; apex rounded, entire; base cuneate with separate lanceolate glands, 1.5 x 0.4 mm. Stamens 3.5mm., edentate; apex not produced, truncate. Ovules 2, stipitate, the stipes being about as long as the ovule.

Bolivia: Songo, Nov. 1890, Bang 857 (K).

(418) Berberis rusbyana Ahrendt, sp.nov., aff. B. rectinerviae, sed foliis latioribus, marginibus spinosis, opacis ; pedicellis brevioribus, floribus majoribus, ovulis semilibus, distinguenda

Shrub to 10 ft. Stems terete, purple, verruculose. IntemLodes 8-15 mm. Spines absent or 1 3 m m . , often subpalmate and 1-5mm. broad at base. Leaves very rigid, with hypoderm, 15-25 x 7-10 mm. ; oblong-obovate, sessile, obtuse; margins 3-5-spinose (1-5-2 mm.)-serrate (1 mm.), distant 2-6 mm.; almost dull both sides; above grey-green with lateral veins indistinctly elevated ; below much paler yellow-green, epapuose, with elevated branched venation. Flowers solitary. Pedicels glabrous, 1-2 mm. Outer sepal8 3.5 x 1 mm., lanceolate, acute. Median s e p k oblong-ovate, or oblong-elliptic, 7.5 x 2-5 mm. Inner s e e similar, 7.5 x 3-25 mm. Petals 6 x 4 mm., obovate, entire, cuneate.

Page 268: Berberis and Mahonia : A taxonomic revision

268 L. W. A. ~ N D T

Stumna 4 mm., dentate; apex not or soarcely produced, rounded-truncate. 0vu.h 2, subseasile. Berries unknown, but with pedicels 2-4 mm.

Bolivia: Prov. Cochabamba, Department Chapare, near Colomi, Cerro La Abra, 11,000 ft., about 17" S. lat., fl. 27 June 1929, Steinbach 9874 (Type, E).

(419) Berberis cliffortioides Diels, in Notizbl. Bot. &rt. Bert., 11, 782, 1933;

Stems glabrous, purple, sulmte. Lec;cves 7-12 x 2 - 4 5 mm., thick, with hypoderm, oblanceolate; attenuate, sessile; entire, with visible lateral veins, but not reticdate; below concolorous, papillose. Flowers solitary. Pediixh 6-10 mm., pubescent. Inwr sepals 4 mm. Petals 3.5 mm. Style 1 - 6 2 mm.

Peru: Prov. Cuzco, in collibus Saxaihuaman, 11,400 ft., fl. Deo. 1928, Herrera 2205a. Prov. Junin, in the Cordillera of Punto on the route from Huancayo to Collorbamba,

Macbride, F1. Peru, in Fi& Nus. Publ. B d . 13, pt. 2, 671, 871,1938

fl. Nov. 1866, Raimondi 2562.

(420) Berberis pavoniana Ahrendt, sp.nov., aff. B. diffortwidi sed foliis marginibus

Stems glabrous. I n t e A 8-15 mm. Spines absent. Leaves 11-22 x 3-6 m., thick, with hypoderm, obovate; sessile; entire; dull above, and openly veined; below reticdate, slightly paler green, epapjliose. Flowers fascicled 2-3. P e d W pubescent, 9-16 mm. Outer se@ small. Inner se@ 7 x 3.5 111111. Petals 6 x 5 mm., obovate, entire; basal glands lanceolate, acuminate, separate, non-marginal, 1.5 x 0.5 mm. Stumna 5 mm., dentate; apex not produced, rounded-truncate. O v u h 3.

Ecuador: Prov. Pichincha, PAramo de Tiopullo (0-1"s. lat.), ll,OOOft., 1825, fl. Jameson 567 (Type, E).

spinosis, floribua fascicdatis distjnguenda

(421) Berberis phyllacantha Rusby, Hem. T m e y Bd. Cl. 6,4,1893 Stems pubescent, subterete or subangled, pale yellow. Intermah 10-20 mm. Spines

mostly absent or weak, rarely to 1-2 cm. Leaves thick, with hypoderm, 10-25 x 4- 11 mm., obovate; cuneate, sessile; margins 2-3-spinose (1-2 =.)-serrate (1-1.5 mm.), situated towards apex of lwf; above lustrous, with lateral veins indistinct ; below dull, yellow-green, subenemate, papillose. Fkmer8 solitary. Pd& pubescent, 5-7 mm.; bracts 2 mm. Outer se@ 3 x 1.25 mm., ovate, acute. Median se@ 5 x 2 mm. Inner sepakr 6 x 3.5 mm., obovate. PetQls 5 x 3 mm., obovate; glands ovate, 0.6 x 0.5 mm. Stumna 44.5 mm., dentate; apex not or scarcely produced, rounded-truncate. Ovules 4. Bemkw ovoid, pruinose blue, 8 x 5 mm., excluding style 1 - 6 2 mm.

Bolivia: Prov. La Paz, Sorata, Nov. 1892, Bang 1614; Mandon 862 (K).

Sect. Confertae Ahrendt, sect.nov., aff. sectioni Virgatat. sed stylo null0 vel brevissimo

S p i w 2-10 mm., mostly weak or absent, h u e s evergreen; cuneate; sessile or sub- sessile, nearly always with hypoderm. F h e r s solitary, or inflorescence 2-6 (-8)-fld., 1-3-5 cm. long. Style absent or insigniscant (up to 0.3 mm.). (In two species only, is the style 0-5-0-75 mm., and these clearly form a link with the section Virgatae.)

Colombia: Prov. Cundimmarca. Ecuador : Provinces Pichincha, Cuenca, Chimborazo. Peru: Provinces Cajamarca, Puno, Cuzco, Lima, Ancachs. Bolivia: Provinces La Paz, Potosi, Tarija. Range 4' N., to 15' S. lat.

Page 269: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonirt 269

KEY 82, to Sect. Confertae Style distinct, 0*60*75 mm. ; stems terete; dark, red or purple; leaves lustrous above dull

Stems pubescent; flowers fascicled 2-3; pedicels 10-15 mm., glabrous; flower diameter

Stems glabrous; flowers solitary; pedicels 3-5mm., pubescent; flower diameter

below.

15 mm.; petals shorter than inner sepals; leaves entire.

10 mm. ; petals longer than inner sepals; leaves with spinose margins.

422. B. sprmceana

423. B. paueidentata Style absent or insignificant, scarcely 0.3 mm. long.

Leaves below pruinose, white or grey. Stems pubescent ; style absent. 424. B. rari&wa Stems glabrous.

Style short ; leaves spinose ; stamens truncate. 425. B.aaxicola Style absent; leaves entire; stamens subapiculate. 426. B. humbertiana

Leaves below always, or finally, @;reen epruinose. Stems pubescent.

Mature stems pale yellow ; pedicels pubescent. Inflorescence 1-4-fld., solitary or fascicled, sometimes subumbellate ; pedicels

2-8 mm. Spines absent, occasionally present and weak, 2-4mm. long, possibly sub-

palmate; pedicels 4-8 mm. ; leaves subenervate to subreticulate; inflores- cence 6 1 0 mm. long. 427. B. conferta

Spines conspicuous, 3-fld., 1-1.5 cm. long; pedicels 2-3 mm.; leaves more distinctly reticulate; inflorescence 1-24 cm. long. 428. B. bolivbnu

429. B. lobbiana 430. B.psiloclarla

427c. B. conferta var. hypopyrrantha

Inflorescence subracemose, 3-1O-fld., 2-4 cm. long; pedicels 1-2 cm.

Mature stems dark, red, purple, or black; pedicels glabrous.

Mature stems pale yellow; pedicels pubescent. Mature stems dark, red, purple, or black; pedicels glabrous.

Stems glabrous.

Leaves fairly narrow, 1 : 2-63, entire. Pedicels 2-3 111111. ; ovules solitary; petals as long as inner sepals; spines

Pedicels 1-2 cm.; ovules 3-5; petals shorter than inner sepals; spines absent strong, conspicuously discolorous. 431. B. chryaacantha

or weak. 432. B. globosa Leaves broad, 1 : 1.61.75; margins with 2-0 spinules.

Leaves reticulate with 2-3 marginal spinules ; inflorescence subracemose,

Leaves not reticulate, with 3-6 marginal spinules; inflorescence fascicled, 3-6-fld. ; pedicels 6-10 n ~ n . 433. B. rig&

1-3-fld. ; pedicels 2-4 mm. Cf. 430. B. psiloclada

(422) Berberis spruceana (Schneid.) Ahrendt, stat.nov. ; syn. B. conferta var. spruceana Scheid., Bull. Herb. Boissier, (2), 5, 393, 1905; Macbride, Fl. Peru, in Field Mus. Publ. Bot. 13, pt. 2, p. 676, 1938.

Internodes 5-10 mm. Spines absent, or solitary and 1-2 mm. long. Leaves 12-20 x P7 mm., narrowly obovate. Pedicels 10-15 mm. ; bracts 1-1.5 mm. Outer sepals 5.5 x 3.5 mm. Inner sepals 7.5 x 4-5 mm. Petals 6 x 5 mm., broadly obovate; glands ovate, 1.2 x 0-6 mm. Stamens 4 mm., produced, truncate. Ovules 3. Style 0.5-0.75 mm.

Ecuador : Prov. Chimborazo, Valle de Pangor, secus limitem sylvarum, 10,000- 12,000 ft., fl. Oct. 1858, Spruce 5549 (Isotype, K). ‘Avery rare shrub, 12 ft. high; flowers orange.’

Peru: Prov. Cajamarca, Chota, west of Huambos, 9750 ft., Weberbauer 4157, 260, according to Macbride.

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270 L. W. A. AKBENDT B. spm~ceana and the following species B. pamkkntata, on account of their longer

styles fall, properly, in between the Sects. Virgatae and Codertae. However, their styles are not of the really signiiicant length which characterizes certain South American sections (amongst them the Virgatae); accordingly, it seems best to include them in the Confertae, placing them here, where they will be adjacent to the Virgatae. Spruce’s two volumea confain several detailed maps of the areaa of his travels in tropica.1 South America.

(423) Berberis paucidentata Rusby, BUZZ. N.P. Bot. Gdn. 4, 321, 5 Sept. 1907

h u e s thick, 9-16 x P 9 mm.; margins 1 (-2)-spinose (1-1.5 mm.)-senate (1-16 mm.), subenervate; below slightly veined. P e d W 3-5 mm.; bracts 1-2 mm. Inner sepals dark red, ovate, acute, rigid, 4-5 mm. Petal;p 5-6 mm. BemS pruinose blue, 8-9 mm. Style 06-0.75 111111.

Bolivia: without locality, Bang 1828 (Isotypes, E, K). Rusby describes the venation as coarse and prominent above, which is not the case in the specimens cited.

(424) Berberis d o r a Lechl., Berb. Am. Awrtr., p. 33, 1857 Stem puberulous. Leavea below grey, pruinose, papillose. Style absent. Bolivia: Prov. La Paz, Andes around La Paz, according to Lechler. Prov. Potosi, Yungas, Weddell 4275.

(425) Berberis saxicola Lechl., 1.c. p. 42, 1857; Macbride, FZ. Peru, 1.c. 13, pt. 2,

Stem terete, glabrous, purple. Spines absent. Leaves 3-6 cm. long, below pruinose white, reticulate; margins with 3-10 spinose serrations. Flowers solitary, with short pubescent pedicels. O w u k 5-7. Sturnem truncate. Style very short, but distinct.

Peru: Prov. Puno, rocks. Sachapata and Tabina, 15,300 ft., Lechler 2096 (Type). Prov. Cuzw, Pam de Trea Cruces, according to Macbride, Pennell 13911. ‘Petals

p. 679, 1938

scarlet.’

(426) Berberis humbertiana Macbride, FZ. Peru, 1.c. 13, pt. 2, p. 674, 1938 Stem glabrous, terete. Spines 3-0d., 35 mm. Leaves rigid, 8-10 x 4-5 mm. ; mostly

entire, rarely with one tooth each side; obscurely openly reticulate; above sublustrous; below dull, papillose. Inflorescence 4-6-fld., 6-15 mm. long. P e d W 2-2.5 mm. Inner s e f i 3 mm., orbicular. Petals 3 mm. long. Stamens subapiculate. Berries globose, estylose.

Peru: Prov. Cuzco, near Cuzco, Gay, s.n. (Type, Paris).

(427) Berberis conferta Kunth in Humb. et Bonpl., Nov. Gem. et Sp. PI., 5, 69, tab. 430, 1821; Hieron., Bot. Jb. 20,49, 11, 1895. (Fig. 65)

Stem yellow, pubescent except in variety hymrrantha. Spines mostly absent, otherwise very short, -nun., and sometimes subpalmate. Inflorescence solitary, or fascicled 2-4, only 4-10 mm. long. Pedicds 4 4 (-10) mm., pubescent.

Ecuador : Prov. Pichincha. Peru: Provinces Cajamarca, Puno.

KEY 82A Stems pubescent.

Flowers often faecicled; leaves spinose. 427a. B. confertu var. conferta Flowers solitary; leaves entire. 427 6. B. conferta vm. karsteniccncc

4270. B. wnferta var. hypopgm-antha Stems glabrous; leaves entire; flowers solitary.

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Berberis and Mahonia 27 1

(427a) Berberis conferta var. conferta Stm..~ terete; young shoots puberulous, pale red-brown; mature glabrous, pale yellow-

brown. Intemzodes 5-13 mm. Spines3-fld., palmate,24mm. long, base 1 mm. broad. Leaves with hypoderm, (lo-) 1P16 (-18) x (5) 6-7 (-9) mm.; obovate; margins 2-4-spinose (1 mm.)-dentate (1.5-2 mm.), distant 3-6 mm., situated by apex; base cuneate, entire; above subenervate, lustrous ; below reticulate, concolorous green, epruinose, epapillose. Flowers fascicled 2-4. Pedicels puberulous, 4-8 (-10) mm. ProphyuS 1-1-5 mm., ovate, acute. Petals 5.5 x 4 mm., obovate, entire. Stamens 3 4 mm.; below the anthers very shortly dentate, or sub- dentate; apex not produced, rounded. Ovules 4-5. Style absent, or scarcely 0.2 mm.

Peru : Prov. Cajamarca, between Cajamarca and Magdalena, Humboldt, s.n. (Type, according to Kunth).

Prov. Lima, Rio Blanco, 15,000 ft., according to Macbride. Ecuador: 9,000-11,000 ft., fl. Oct. Lehmann 4693 (K). Prov. Cuenca, woods about Contrayerba, according to Hieronymus.

Outer sepals 5 x 2.75 mm. Inner sepak 5-25 x 3.5 mm.

(427 b) Berberis conferta var. karsteniana Schneid., Bull. Herb. Boissier, (2), 5, p. 393,

Stems terete. Internodes 5-9 mm. Spines mostly absent. Leaves 8-12 x 3-7 mm., or to 18 x 6 mm., entire, reticulate below, epapillose. Pedi& 4-7 mm.

Ecuador: Prov. Pichincha, Quito, 12,000 ft., Karsten, s.n. (Type).

1905

(427c) Berberis conferta var. hypopyrrantha Schneid., Bot. Jb. 42, 82, 1908 Shrub 1-2 ft. Leaves 10-30 mm. long; margins with 2-10 spinules from mid-point to

apex; reticulate; below paler green. Flowers solitary. Pedicels red, 5-10 mm. Petals subequal to the inner sepals. Stamens not produced. Ovules 5. Style very short.

Peru: Prov. Puno, Cuyocuyo, 10,400 ft., Weberbauer 853 (Type).

(428) Berberis boliviana Lechl., Berb. Am. Austr. 21, 303, 1857 ; Macbride, Fl. Peru. 1.c. p. 670, 1938; syn. B. conferta var. boliviana (Lechl.) Schneid., Bull. Herb, Boissier, (2), 5, 393, 1905

Stem pubescent, yellow. Spines 1-3-fld., 1-1.5cm. Leaves 1Ox3mm. to 12-17x 5 mrn., entire ; pale above; below concolorous, epapdlose; closely reticulate. In$ores- cencct fascicled or subumbellate, 2-4-fld., 1-1-5 (-2.5) cm. long. Pedicels pubescent, 2-3 mm. Petuk and inner sepals c . 3 mm. Style absent.

Bolivia: Prov. Potosi, Agopaya, 11,700 ft., Weddell 4139. Prov. JA Paz, near Peru boundary, Pentland s.n. (Type). Peru: Prov. Cuzco, Pachar, 10,000 ft., Pennell 13690.

(429) Berberis lobbiana Schneid., Bot. Jb. 42, 83, 1908; Macbride, Fl. Peru, 1.c. p. 675, 1938; syn. B. conferta var. lobbiana Schneid., Bull. Herb. Boissier, (2), 5 , 394, 1905

Stem terete, puberulous; mature pale yellow-brown or pale red-brown. Internodes 1-1.5 (-2) cm. Spines absent, very rarely present and then weak. Leaves 15-28x6- 10 mm. ; elliptic-obovate; base cuneate, sessile; entire, very rarely with one spinose tooth each side ; apex conspicuously acute, mucronate (1-5-2 mm.) ; above lustrous, light green, openly reticulate ; below sublustrous, concolorous, more closely reticulate. In- Jlorescence (3-) 6 8 (-10)-fld., partly fascicled, partly umbellate-racemose, 2-3.5 cm. long, with the peduncle of the latter part 1 cm. long. Pedicels puberulous, 1-2 cm. long; bracts 2-4 mm. Flowers 10-12 mm. diameter. Outer sep& 5-5.5 x 3 mm. Inner sepah 7 x 6 mm., broadly obovate. Petals 6.25 x 5 mm., broadly obovate; base with a distinct claw. Stamens 5-5.5mm. long, edentate; apex not produced, truncate. O& 4-5. Berries estylose, or with style not distinct.

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272 L. W. A. ~ E N D T

Ecuador (East Central): Lobb 341 (Isotype, K). Peru: Prov. Huhuco, Huamaliee, south-west of Monz6n, 11,700 ft., Weberbauer

3305.

(430) Berberis psiloclada (Schneid.) Ahrendt, statmov., syn. B. wnferta, var. pilo-

Stems; mature dark purple-brown to black. Spine8 short, palmate, 1 (-3)-fld. Leuvu with hypoderm, 8-25x5-16mm.; margins with 2-6 teeth from middle to apex; not reticulate; light green above; warmly paler green below. Flowers fascicled 1-3. Pedkds glabrous, 2-4 mm. Flowers yellow, red without. Berries ovoid-oblong, 8 x 6 mm., with distinct short style.

Peru: Prov. Ancachs, above Hwaz , 10,40Oft., fl., 6 Jan. 1903, Weberbauer 3225 12,OOO ft., Weberbauer (Type) 3278.

duds Schneid., Bot. Jb., 42, 82, 1908

(431) Berberis chrysacantha Schneid., Bot. Jb. 42, 81, 1908 Stems glabrous; mature dark purple, subsulate. Spins conspicuous, orange-yellow,

1.5-2-5 cm. Leave% without hypoderm, 10-18 x 3-54 nun., narrowly obovate; acute, entire; below concolorous; reticulation elevated. I n , e % m m 3-5-fld., racemose-sub- fascicled. Pediceh 2-3 mm., glabrous; bracts 1-1.25 mm. Petals obovate, emaxginate, as long aa the inner sepals. Stitmem apiculate. O d e a solitary. Style very short. Bolivia: Prov. Tarija, Condorhusssu, west of Tarija, 9750 ft., fl. 7 Feb. 1904, Rebrig

3052 (Isotype, BM).

(432) Berberis globosa Benth., P1. Hurtweg., p. 158, Bug. 1845 Stem terete. Intenzoda9 6-14 mm. Spina absent or 3-fld., 2-6 mm. Leuvu without

hypoderm, 15-30 x 6-10 nun., obovate-elliptic; base cuneate, sessile; entire; above lustrous green with close elevated reticulation; below, at first grey and pruinose, hally dull paler green with open reticulation, papillose. InJlorescence 3-6 (-8)-fld., fascicled or racemose-subfascicled, 1-2.5 cm. long. P e d W glabrous, 1-2 cm. ; bracts 3-4 mm. ProphyUs lanceolate, acuminate, 1.5 x 0.5 mm. Outer sepds 2 mm. Inner sepaJs 4 x 2.5 mm. Petals 2.5 x 1-75 mm., obovate. Stumn8 2-2.5 mm., edentate, apichte. Ovules 3-5. Berries globose.

Colombia: Prov. Cundinamarca, Bogot&, lO,OOOft., shrub, Wft., Hartweg 879 (Type, K).

(433) Berberis rigida Hieron., Bot. Jb. 20, 49,11, 1895 Stem glabrous, subsdcate; mature purple-black. 8pine.s 3-fld., 4-1Omm. Leaves

without hypoderm, 10-30 x 7-18 mm.; margins 2-3-spinw (1.5-2 mm.)-serrate (2- 3 mm.), distant 4-7 mm.; above lustrous, with loose, coarse, elevated, reticulate vena- tion; below dull, paler, papillose, similarly veined. [email protected]' M-fld., subracemose, 2-4 cm. long, including peduncle 1-2 cm. Pedicels glabrous 7-10 mm.; bracts 3 mm. Prophylh lanceolate, 2-5 x 0.8 mm. Outer sepah 4 x 2 mm., oblong-elliptic. Inner sep& 5-6 x 3 - 5 4 mm. P d 5-6 x 3-36 mm. ovules 4 4 . Stamem c. 3 mm., apex apiculate. Ecuador: Prov. Cuenca, Warszewicz 3 (26) (Type); Punfa. de Caja, no. 806 (K).

Sect. Agapatenses (Schneid.) Ahrendt ; s p . Sect. Actinacanthae Subsect. AQAPATENSES Schneid., BuU. Herb. Boissier, (2), 5, 148, 1905

kve . s coriaceous, evergreen, 2-4 cm. long, with hypoderm, suborbicular, or broadly elliptic, below papillose. Spine8 not palmate. Inglorwceence 2-6 (-8)-fld. Ovules 8-12. Fruit estylose.

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Berberis and Mahonia 273

Peru: Prov. Huiinuco. Bolivia: Prov. Cochabamba. Range 10-17" S. let.

KEY 83, to Sect. Agapatenses, Stems pubescent; leaves 2-3.5 x 2-3-3 cm., suborbicular; base cordate, or Cruncate ;

Skms glabrous; leaves narrower, to 4 x 3 cm., with rounded base; slightly veined, not closely reticulate; petiole 1-2 cm. 434. B. agap&n&

reticulate. 435. B. benoistiana

(434) Berberis agapatensis Lechl., Berb. Am. Austr. p. 44,1857; Macbride, Fl. Peru,

A!II%W terete; mature lustrous red-brown. Spines absent, or weak, 1-3-fld., 6-8 mm. Leaves with margins 8-12-spinose (1-1-5 mm.)-serrate (0.5-1 mm.), distant P 7 mm. (though on young leaves more entire and cuneate); almost dull above; below pruinose. Flowers fascicled 2-6 (-8). Pedi& 12-20 mm. Stanzens ( 2 ) dentate. Berries 8 x 4 mm., immature.

in Field Mus. Publ. Bot. 13, pt. 2, 669, 1938

Peru: Prov. Puno, Agapata to Sachapata, Lechler 2646 (Type). Bolivia: Prov. Cochabamba, fr. 1891, Bang 1091 (K).

(435) Berberis benoistiana Macbride, FZ. Peru, 1.c. 669, 1938 Inner sepals 6 mm. Petals 4 mm. Berries epruinose. Peru: Prov. Huitnuco, Tambo de Vaca, on a fog-drenched summit in open mossy

places, 13,00Oft., Macbride 4455 (Type, Herb. Field. Mus. Chicago). 'Slender shrub, 3 ft. high, with bright yellowish red flowers, and black fruit.' Macbride notea that the papillae below the leaves are 80 abundant as to seem to form a tissue.

Sect. Quindiuenses Schneid., Bull. Herb. B&eSir, (2), 5, 809, 1905 &ems terete, mature pale yellow or yellow-grey. Spines absent, or weak, P 8 mm.

Lecaves evergreen, mostly without hypoderm, and nearly always closely reticulate. Racemes l0-25-fld., 5-20 cm. long. Pedieels (8-) 12-20 (-25) mm. Flowers 14-2 cm. diameter. Outer sepals 2-4 mm. Medkn sepals 4-6 mm. Inner sepals 8-11 mm. Petals 5-7 mm. 0vuk.s 2-8. Style absent, or indistinct (except in B. odifoZkz).

Colombia : Provinces Nado, Magdalena or Boyad. Ecuador: Prov. Pichincha. Bolivia: Prov. Potosi, and North Chile. Range 5" N. to 10"s. lat.

KEY 84, to Sect. Quindiuenses Leaves with 6 6 0 marginal spindes, below mostly epruinose and green.

Leaves 7-16 cm. long, with 20-60 spinules to each margin; below concolorous green;

Ideaves 2-6cm. long, with 6-9 spinules to each margin; below much paler green;

Leaves entire (rarely with a few occasional inconspicuous sphulw); below pruinose,

ovules 6-9; style absent. 436. B. ciliariS

ovules 2-3; style 0.5-1 mm. 437. B.ovali,folia

white or grey; style absent or indistinct. Leaves closely reticulate.

Stamens apiculate; pedicels 8-12 mm. Stamens truncate; pedicels 12-25 mm.

438. B. quinduerrsis 439. B. grandifiora

440. B. aurahumen& Leaves subenervate, or with a few lateral veins. 18 JOURN. LZNN. SOC. BOTANY-VOL. LVII

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274 L. W. A. AERENDT

(436) Berberis ciliaris Lindl., J . R. Hort. Soc. 5,7,1850; syn. B. km$ora Citerne non

Sterns glabrous ; young shoots slightly purplish ; mature pale yellow-grey. Spines absent, or 1-3-fld., 4-9 mm. Leaves above dark green, slightly lustrous; closely reticulate both sides. Sbmm scarcely or slightly produced, truncate or obtuse. O v u h 6-9.

schrad.

KEY 84 A Leaves with apex mute, below epruinose, subconcolorous green.

h v e a with apex rounded, below plvinose white. &a. B. ciliarie vm. dkrk

436 b. B. ciliaris var. obtzLsata

(436~) Berberis ciliaris vm. cillaris Leaves 7-15 x 2-54? cm., or to 14 x 4 4 cm. ; margins 20-6O-spinose (1.5-2.5 mm.)-

sedate, distant 1-54 mm.; below subconoolorous sublustrous green, epapiuose; apex acute. In$oreswnce 10-15-fld., racemose, 5-12 cm. long, including peduncle 3-5 cm. Pedkth 1-2 cm.; bracts 3-5 mm. Outer sepals 6-5 x 3 mm. ovate, subobtuse. Median and inner sepals both 10 x 4-5 mm., elliptic. Pet& 10 x 5-6 mm. Stanzena 7 mm.

Bolivia: Prov. Potoai, Yungaa, Weddell 4226 (K). North Chile: Choro, 10,OOO Et., 7 Feb. 1950, Brooke (BM); Bridges, s.n. (aa Zaxiea)

(K). Brooke remarks that the flowera have a sweet scent.

(4363) Berberis ciliaris var. obtusata Ahrendt, var.nov., foliis apice rotundatis, subtus dbo-pruinosis, inflorescentiia longioribus distinguenda

Leaves below white, pruinose, papillose, to 11 x 4.5 cm., oblong-ovate, apex obtuse to rounded; margins 30-spinose (spreading, 1.54mm.)-mrruIate. R- 9-14-fld., loose, 1P21 cm. long, including peduncle 6-12 cm. PediceZa 2-4 cm. ; bracts 4-6 mm. Sep& in four series. Outer sep& 5 x 2 mm. Outer median sepals 7-5 x 5 mm. Inner median sepals 9-10 x 6-7 mm. Inner sepals 10 x 7 mm. Petals 9-10 x 6-5 mm. &mem 6-5 mm.

Bolivia: Unduavi, 10,OOO ft., 3 July 1950, Brooke 6608 (Type, BM).

(437) Berberis ovalifoh Rusby, Desm$tk of Three Hudred New Speck? of S&h

Sterns terete to very finely sulate. Spinea 4-8 mm., stout. Leaves (2-) 3-4.5 (-6) x 1-1-8 (-3) om.; elliptic to oblong-obovate; thick; petiole 1 cm.; margins 6-9-spinom (1-2 mm.)-serrate (0-5 mm.), distant 2.54 mm. ; sublustrous grey-green above, with branched venation; below very much paler green and very ihely reticulate. Racemes (7-) 10-25-fld., 4-10 cm. long, including peduncle 1-5 cm. Pedicels 6-20 mm.; bracts 2-54 mm. Outer se& 5-5 x 3 mm. Inner se& 8 x 4 mm., all elliptic. Petals 6 x 3 mm., obovate. Stamem 4 mm., not produced, truncate. O v d a 2 (-3). Style 0-5-1 mm.

Amriuzn Plank?, p. 16, Dec. 1920

Bolivia: Brooke 6108 (BM).

(438) Berberis quinduensis Kunth in Humb. et Bonpl., Nm. Gkn. 5,70, t. 432,1821 Interrwuh 1-63 cm. Spines absent, or 1-3-fld., to 9 mm. Leaves to 4 x 1.6 or 6 x

2.2 cm.; obovate; base cuneate, seasfie; apex obtuse; margins entire, rarely with 2-5 inconspicuous spinules near apex; above dull green; below grey, pruinose, epapillose; both sides closely reticulate. Race= 12-25-fld., loose, 5-1Ocm. Pedkds glabrous, slender, 8-12 nun. ; bracta 2-3 mm. ProphyyuS truncate, acumhate, 1-5 mm. Outer sepals 3 x 1.5 mm., ovate, acute. Median sepals 4 x 2.5 mm., ovate-elliptic, obtuse. Inner

Page 275: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 275 sep& 8-9 x 5 - 5 4 mm., elliptic. Petals 5 x 3-5 mm., obovate, cuneate. Stamem 4 mm., produced, apiculate. O v u h (3-) 4-5.

Colombia: Prov. Nariiio, Paato, Tuquerres, 9800 ft., 1851-7, fl. Triana, s.n. (K).

(439) Berberis grandiflora Turcz., BuU. SOC. Nut. Mosc. 27 (ii), 287, 1855; syn.

& e m terete to subterete. Internodes 4-7 cm. Spines 3-fld., 3-6 mm. Leaves 3-6 x 2 . 5 4 4 om. ; base contracted to petiole 1-1-5 cm. ; margins entire, rarely with two to five inconspicuous spinules by apex; above slightly lustrous, with close h e elevated reticu- lation ; below pruinose, grey-white, papillose, with elevated reticulation more open and coarse. Inflorescence racemose or umbellate-racemose, 10-20-fld., 5-14 CIL. long, in- cluding peduncle 2 4 cm. Pediwls glabrous, 1-26 cm.; bracts c . 3 mm. Outer sepals 3 x 1 mm. Median sepals 6 x 2 mm. Inner sepals 9 x 6 mm. Petals 7 x 4 mm. Stamens truncate. Berries ellipsoid, or obovoid. Style absent or indistinct ; slightly pruinose blue, 8 x 5 mm. Seeds black, 5 x 3 mm.

Ibuador: Prov. Pichincha, Mt. Pichincha, west side, 10,OOO ft., Jameson 78 (E), and 12,000ft., Jameson, s.n. (E), 1857-9, Spruce 5548 (0). In 1850, at 13,OOOft., Jameson, 8.n. (Isotype, K); Jameaon 281 (Isdype of B. jameS0n;i Turcz., non Lindl.).

(440) Berberis aurahuacensis Lemaire in Van Houtte, Fl. des Serres, 4, t. 334, Apr. 1848

Ntem terete; young shoots green; mature yellow. Spines absent, or 5-fld., 3-5 mm. long. Leaves of two kinds-(a) the lower with cordate base, and petiole 1-3 cm., almost aa long aa the blade, (b) the upper, obovate-elliptic, attenuate at the baae to a very short flat petiole; blades 2.5-5 x 1-2 cm., or to 4.2 x 2 cm. ; margins subundulate; mostly entire (rarely with 1-5 inconspicuous spinules); with a few lateral veins (and part of the marginal vein a t most) showing on either side, sometimes subenervate, never reticulate ; below pruinose, white. InJorescence (8-) 10-20 (-30)-fld., racemose, subverticillate, 5-12 cm. long including peduncle 1-2-5 cm. Pedieels glabrous, stout, 6-10 mm. ; bracts 2-3 mm. Outer sepals golden yellow, 3-4 x 1.75-2-5 mm. Median sep& 5-6 x 3 4 mm. Inner sepals 10 x 7-8 mm., oblong-obovate. Petals orange, 7-8 x 4-5 mm., obovate; apex acutely emarginate with rounded lobes. Style absent or indistinct.

Colombia: Prov. Magdalena, or Prov. Boyad (?), Rio Hacha, near the village of Aurahuacu-Taquina of the Sierra Nevada, 9OOOft., near the snow line, Linden s.n., according to Lindley, an elegant shrub, with stiff, erect branches.

B. jampmmii Turcz., l.c., 286, non Lindl., 1850

Sect. Goudotiae Schneid., Bull. Herb. Boissier, (2), 5 , 811, 1905 Stem terete, glabrous. Spines 3-fld., 5-12 mm. Leaves evergreen; base distinctly

petiolate, with petiole (0.5-) 1-2 cm., articulate below the lamina, without hypoderm. In$oreswnce racemose or racemose-subfascicled (6-) 8-2O-fld., 2-5-1 1 cm. long. Berries with very short style, or estylose.

Venezuela: Provinces Zamora, M6rida. Colombia: Provinces Antioquia, Huila, Cundinamarca. Ecuador: Prov. Pichincha. Peru : without locality. Range from the Equator to 8" N. lat.

KEY 85, to the Sect. Goudotiae Leaves entire; stems (where known) pale, yellow or grey.

Lmves with revolute margin, papillose below; racemm 12-20-fld., 8-11 em. long, often compound below.

18-2

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276 L. W. A. ~ E N D T

BY 85 (cont.) 441 B. discolor 442. B. m r i t z i i

Pedimk pubescent; leaves subenervate, pruinw grey below. Pedi~ls glabrous; leaves closely reticdate, epruinose &reen below.

Leaves with margin not revolute; racemes 6-16-fld., not compound below; pedicels glabrous.

Idorewence sessile, racemose-subfascicled, 2.5-3-5 cm. long; leaves epapillose below; petals as long as inner sepals.

lif7orescence peduncled, 8cBmoB8, 44cm. long; leaves papillose below; p e ~ . shorter than inner sepals.

443. B. guilache

444. B.papJlosa Leaves with spinose margins.

Petiole (2-) 5-10 mm.; leaves with blade not orbicular, 1 : (1*75-) 2-3. Leaves white below; stems yellow. Leaves green or grey-green below; stems dark, purple or black.

444. B.ppdloea

Leaves reticulate above, below pruinose grey, papillose; w g i n s with 5-8 spinules; racemes 0-12-fld., 0-8 cm. long, very loose; pedicels 12-20 mm.

445. B. stuebelii Laves enervate above, openly reticulate and epruinose green, and epapillose

below; m a r e with 3-5 spinules; racemes 10-20-fld., 4-6 cm., d e w ; pedicels 7-12 IIM. 446. B. gowlotii

447. B. bumronis

(441) Berberis discolor Turcz., Bud. Soc. Nat. Mosc. 27 (ii), 287, 1855; Pittier, J. Wmh. A d . Sci. 21,136,131; syn. B. nutam Lindenet Planch., PI. Columb. 8 (1863); Spraee, Kew Bull. 1926, p. 35,1926

InJloresCence 12-20-fld., racemose, 8-11 cm. long, often compound below. Pedicels pubescent, 8-13 mm.; bra& 2-3 (-4) mm. O& 2.

Venezuela: h v . Zamora, 10,OOO ft., Sierra Nevada de Mdrida, Funck and Schlim 1124 (Type); Phramos de Santo Doming0 y Chachopo, 10,7OOft., fl. 14 Sept. 1922, Jahn 1120.

(442) Berberis moritzii Hieron., Bot. Jb. 20, Beibl. 49, p. 16, 1895; Pittier, 1.c. Spines 3-fld., stout, 4 4 3 mm. Leaves 36-76 x 1 - 3 3 cm.; petiole P 1 5 (-20) mm.

Racemes 10-20-fld., rarely compound below, 8-11 (-15) cm. long. Pedicels glabrous, 7-12 mm.; bracts c. 2 mm. Inner se@ 3 x 1.5 mm., ovate, acute. Petals 5 x 3 mm., much longer than inner sepals, obovate, obtuse. Stamem edentate, apiculate. ovules 2.

Venezuela: Prov. Zamora, east side of mountains of cord. de Mdrida, Moritz 1309 (K). Prov. Mhrida, Alpine belt of the mountains above Mdrida city, fl. Feb., according to

Petiole 1-26 cm. ; leaves papillose below, with orbicular blade.

Pittier.

(443) Berberis guilache Triana et Planch., Ann. Sci. Nat. Bd., ser. 4, 17,454, 1862 Stems with young shoots subsulcate; mature terete, pale yellow. Internodes 4 4 cm.

Spines 3-fld., 4-9 nun. h e r e s 2-5-5.3 x 13-24 cm., excluding petiole, 5-10 (-20) mm. ; oblong-obovate ; entire, sometimes very slightly revolute ; above slightly lustrous deep green, with close coarse elevated reticulation; below slightly pruinose, dull, greyish with fine closely elevated reticulation. Racemes (6-) &15-fld., 2-5-3.5 cm. long. Pedkds glabrous, 7-10 (-15) mm. long. Prophylla 2, lanceolate, acuminate, 3.5 x 1.5 mm. Outer sepals 5 x 2-5 mm., oblong. Inner sepds 6 x 4 mm., obovate. P&ls 6 x 3.5 mm., obovate; apex slightly retuse; base with elliptic glands, 0.8 x 0.5 mm. S t u m 4 mm., produced, shortIy apiculate. O d e s 4-5.

(444) Berberis papillosa Benoist, Pubt. Mw. H k t . Nat. Paris, 3 (iii), 76,1933

Colombia: Prov. Antioquia, 1851-7, fl. 7800 ft., Triana s.n. (K).

Leaves with articulate petiole; white or whitish below.

Page 277: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 277

KEY 85a Leaves with spindose margin, slightly veined. Leaves with entire margin, reticulate.

M a . B. papillosa var. papillosa

Leaves to 6 x 3 cm., very lustrous above; inflorescence racemose-subumbellate,

Ideaves to 6 x 4 cm., dull above; inflorescence racemose, 6-8 cm. long; style absent. 4-6 cm. long; style short, scarcely 0.4 mm. 444 b. B. papillosa var. aequatorialis

444c. B. papillosa var. opacifolia

(444a) Berberis papillosa var. papillosa Stem yellow. Leaves without hypoderm, papillose and whitish below; margins with

3 4 fine spinules; 6-9 lateral veins visible; petiole 5-12mm., articulate a t the apex; blade 2-5 x 1-2 cm.; hypoderm absent. Racemes 4-5 cm. long. Pedicels 8-10 mm.; bracts 3 mm. Flowers 6-7 mm. diameter. Stamens produced, subapiculate. Berries ellipsoid with short distinct style.

Ecuador: Prov. Pichincha, cool forest below San Jose de Minas, 3 Mar.1931, Benoist 3950 (Type).

(444 b) Berberis papillosa var. aequatorialis Ahrendt, var.nov., foliis reticulatis

Leaves above very lustrous; below pruinose white, finely papillose. Racemes 6-12-fld., subumbellate, 4-6 cm. including peduncle, 2-3 em. Pedicels glabrous, 10-13 mm. ; bracts 3 mm. Outer sepals small. Inner sepals 7 x 4 mm. Petals 6 x 3 mm., obovate; apex entire, base with lanceolate, separate, marginal glands, 1-5 x 0.5 mm. Stamens 5 mm., edentate, apex produced and subapiculate. Ovules 2-3. Style very short.

Ecuador: Prov. Pichincha, Andium Quitensium, Jameson 186 (Type, E).

integis supra nitidis

(444c) Berberis papillosa var. opacifolia Ahrendt, var.nov,, foliia integris reticu- latis supra opacis ; fructibus estylosis

Ltlaves with rounded apex, above dull green; below pruinose white, a t any rate a t first, but tending to become epruinose but much paler than above. Pedicels 5-12 mm.; bracts 2-4 m n . Prophyh lanceolate, 3 x 1 mm. Outer sepals oblong, 3 x 1-5mm. Median sepak 4 x 2 mm. Inner sepals 6 x 3 mm., all oblong. Petals 4 x 3 mm., obovate, entire; glands oblong, 1.5 x 0.5 mm., situated at the truncate base of the petal. Stamens 3 mm., produced, apex apiculate. Ovules 2. Style absent.

Ecuador: Prov. Pichincha, Andium Quitensium, west slope of Mt. Pichincha, 12,000 ft., Jameson 10 (Type, E).

(445) Berberis stuebelii Hieron., Bot. Jb. 21, 308, 1895 Stems lustrous, purple, terete. Leaves 2 4 x 1-2.5 cm.; margins with 4-6 spinules;

both sides with coarse elevated reticulation; below papillose, pruinose. Racemes 6-10-fld., 6-8 cm. Pedicels 1-2 cm.; bracts 3-5 mm.

Colombia: Prov. Huila, Pkamo de Huila, Stuebel 282a (Type); Ptiramo de Moras, Stuebel253 b.

(446) Berberis goudotii Triana et Planch., Ann. Sci. Nut. Bot., ser. 4, 17, 52, 1862 Shrub 5-7 ft. Stems; mature black. Internodes 1-2.5 cm. Spines 3-fld., 4-8 mm.

Leaves 2-5 x 1-1.6 cm. ; elliptic; petiole 1-3 mm.; margins 3-5-spinose (1-2 mm.)-serrate; above lustrous, enervate ; below, a t first slightly pruinose, Snally paler yellow-green, not papillose or scarcely distinctly so, with open elevated reticulation. Racemes 10-25-fld., 4-6 cm. Pedicels glabrous, 7-12 mm. ; bracts 3-4mm. Prophy& 2 x 0*75mm., lanceolate.

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278 L. W. A. AEIRENDT

Outer sepal-s 2.5 x 1.5 mm., ovate. Medbnsepds 4 x 26mm., ovate. Inner sepals 6 x 3mm., obovate. Petah 6 x 3 mm., obovate. 8 k . m ~ ~ ~ 4.5-5 mm., edenhte; apex much produced, conspicuously slenderly apiculate. Otndes 3.

Colombia: ROV. CunWmca, Cord. of the Andes, near Bogot&, lO,OOOft . , fl. Hartweg 878 ( T w ) ; Mb. east of Bogoa, 2 Feb. 1899, Sprague 180 (K).

(447) Berberis buceronis Macbride, CandoUea, 6,3, 1934; Fl. Peru, 2, in Field Mw.

Stenzs arching. Spines 2-3-fld., stout, 7 mm. long, 1.5 mm. thick, distinctly curved. Leaves with blade 2-4 x 2-4 cm., orbicular, base deeply cordaSie, with a slender puberu- lous petiole, 1-24 cm. long; margins undulate, spinom-dentate; above slightly lustrous, subenemate; below dull, papillose prominently reticulate. InJloreSCence corymbose- racemose, partially compound. Ped,ic& to 10 mm.

Peru: without locality, Weberbauer 6994 (Type).

Publ. Bot. 13, pt. 2, p. 671, 1938

Sect. Truxillenses Schneid., B d . Herb. BoisSier, (2), 5, p. 804, 1905 Spines weak or absent. Stem when mature, mostly black or dark, red or purple,

often terete. Leaves evergreen, sessile or with short petiole, mostly without hypoderm. 1n.lorescenee (S-) 10-30-fld., racemose, sometimes compound below. Fruit with style absent, indistinct, or extremely short and insignikmt.

Mexico (2). Nicaragua. Venezuela: Prov. Truxillo. Peru: Provincea Cajamarca, Cuzco, Amazonas, Anmhs. Colombia: Frovincea Nariiio, Mmiquita, Smtander. Ecuador: Frovinces Loja, Pichincha, Azuay, Chimborazo. Bolivia: Prov. 1 Larecaja. Range, 15" S. lat. to 20" N. lat.

KEY 86, to Sect. Truxillensea Leaves 6-10 cm. 472. B. &&foliu Leaves 2-5 (-6) om. bPedicela 1-5 mm.

Leaves orbicultlr, spinose; stems terete; petals shorter than inner sepals. Leaves with 2-3 mar@ spindes, white below; stems dark red. Leaves with 8-14 marginal spinules, grey below; stems pale yellow.

449. B. W b a t a

467. B. podophylla Leavw not orbicuh, but broad, 1 : 14-2.5; entire.

Mature stems glabrous; leaves 2-5 x 1-2-2.5 cm. Leaves 3-5 x 1.62.5 cm. ; petals shorter than inner sepals.

Mature stems pale yellow-brown; leaves acute. 466. B. chinaboeka Mature stems black; leaves rounded. 454. B. edentata

471. B. wettsteiniuna h v e s 15-25 x 8-15 mm.; petals 84 long as inner sepals. Mature stems pubescent; leaves 6-15 x 3-8 mm.; petals longer than or equal to

inner sepals. Mature stems pale yellow-brown, terete; leaves naxrow, 1 : 3-4.

Mature s tem dark red, sulcate; leaves broad, 1 : 2. 469. B. ke.eissEe9+r~a

470. B. rechingeri Pedicels (6-) 8-15 111111.

Leaves below persistently pruinose, white or grey. Leaves entire; ovules 2.

Stamens truncate ; leaves reticulate. 453. B. jelakiana Stamens apiculate; leaves slightly veined. 468. B. brumalis

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Berberis and Mahonia 279

Leaves spinose; ovules (3-) 4-5 (-7). Pedicels glabrous ; leaves reticulate.

Bracts 2-3 mm., one-quarter to one-fifth 88 long aa their pedicels. Stems dark red-purple. 459. B.denaa Stems pale yellow-brown. 458. B. bergeriana

452. B. prolijka Bracts 5-7 mm., three-quarters as long as their pedicels. Pedicels pubescent; leaves indistinctly, scarcely, or not, reticulate; petals

longer than or equal to inner sepals. Petals longer than inner sepals; leaves ( 1 : 2-26) dull above with 6-13

marginal spinules; bracts subeqd to pedicals. 461. B. pichiwhen& Petals equal in length to inner sepals; leaves broader (1 : 16-1.8), lustrous

above with 2-3 marginal sphulm; bracts only one-qumter to one- fifth ps long aa their pedicels. 462. B.engk?~iana

Leaves below always, or finally, epnrinose, green. Leaves entire; lustrous above.

Pedicels puberulous ; leaves closely reticulate both sides. Leaves 1-2 em. long; petals as long aa inner sepals; ovules 4-5; stamens

Leaves 2 4 c m . long; petals shorter than inner sepals; ovules 2; stamens apiculate. 465. B. hirtdipea

truncate. 451. B. TUX&%& 450. B. r i g i d i f o h Pedicels glabrous; leaves subenervab or indistinctly veined.

Mature stems and pedicels glabrous. Leaves spinose.

Leaves above subenervate or indi6tinctly veined; mature stems black; ovules 2.

Petals shorter than inner sepals. Petals FS long as inner sepals.

Stamens scaxcely produced, truncate; stems pale yellow. Stamens produced, apiculate.

Mature stems black; petals as long 88 inner sepals. Mature stems yellow; petals shorter than inner sepals.

450. B. rigidifolicc 455. B. dryand&phy&

464. B. reicheano

460. B. lechlerkzna 457. B. p & b w

Leaves closely reticulate both sides; ovules 3-5.

Mature stems and pedicels pubescent; ovules 5-6. Stamens apiculate; petals shorter than inner sepals; leaves lustrous, 2.5-

Stamens truncate; petals as long as inner sepals; leaves dull, 12-24 x 6- 5.5 x 1.3-243 cm. 456. B. r e t i n a d

463. B. aaxorum 13 mm.

Subsect. Eutruxillenses Schneid., I.c., p. 807, 1905 Stamens mostly truncate, or obtuse, rarely apiculate. Ovules mostly 2, occasiondy

(Mexico ? ) . Nicaragua. Venezuela : Prov. Truxillo. Peru: Prov. Cajamarca, CUZCO, Amazonas. Range 10" S. lat. to 20" N. lat.

3-5 (-7).

KEY 87, to Subsect. Eutruxillenses Spines 5-fld. 448. B. fwinoaa Spines not 5-fld., but absent, 3-fld. or palmate.

Leaves not, or scarcely, revolute; pedicels (3-) 6-15 111111. Leaves with spinose margins.

Petals shorter than inner sepals. Leaves below white; pedicels 3-6 mm. Leaves below, at first slightly pruinose, finally green; pedicels 6-10 mm.

449. B. dealbata

450. B. rigidifolia

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280 L. W. A. AHBENDT

KF,Y 87 (cont.) Petals as long re, or longer than, inner sepals.

Stems terete; pedimls 6-9 m.; bracts 6-7 m., subequel to their pdicels;

Stems sulcate; pediwls much longer than brmte; ovules 2. ovule43 6-7. 452. B. proliJicQ

455. B. dryan&phylla Leaves with entire margins. Pedicels glabrous. Pedicela pubescent.

Leaves below green, epruinose. Leave43 below grey, pruinose.

Leave43 revolute; pediwk 1-2 m.

Cf. 450. B. rigid$ololia

451. B. truxillensiS 453. l3. jelskiclna 454. B. edentatcl

(448) Berberis farinoaa Benoist, Publ. H w . Hi&. Nat., Park, 3 (iii), 75,1933 Spines 5-fld. to 14mm. long. Leaves without hypodem; below papillose, pruinose

whitish; 2.5-7 x 14-2.2 cm. InfEoresCence glabrous, racemose with long peduncle, 12-15 cm. long in all, sometimes compound below. PerEicecs 6-11 mm.; bracts 3-5 mm. Flowers c. 8 mm. diameter. Otherwise unrecorded.

Ecuador: Prov. Pichincha, Phramos de San Juan near Quito, 26 Feb. 1931, Benoist 3939 (Type).

(449) Berberis dealbata Lindl., Bot. Reg. 21 (N.S., viii), t. 1750, 1836; Loud., Arb.

Shrub 4-5 ft. Stems terete, red-purple. Spinexi absent. Leaves sparse, mostly solitary, to 2.5 x 2.5 cm.; ovate-orbicular, to subrhombic; coriaceous; convex; base with purple petiole 2-5 mm.; margins undulate, 2-3-spinose-dentate by apex, the teeth and sur- mounting spinules each measuring 1-2 mm.; venation openly branched, not reticulate; above blue-green; below white. Racema drooping, 20-30-fld., 2-3 cm. long, including peduncle 1-2 cm. Pedi& 3-5 (-6) mm. ; bracts minute. Pr0phyll.s 2 x 1 mm., ovate, acute. Outer 8epd.s 3.5 x 2 mm., elliptic. Inner sepals 4.5-5 x 3-3.25 mm., obovate. Petals 445x2.75-3mm., obovate, entire; base cuna te with glands, 1.3-1.5~0.3- 0.4 mm., contiguous at their bases.

( 2 ) Mexico : according to Lindley. According to Loudon, I.c., the berries are at first red, then purplish, the shrub is hardy and was introduced to cultivation about 1830. It was apparently described from cultivated material. Lindley remarks that, although white below, the leaves have not a pruinose, or pubescent undersurface, the white being due to discoloration of the cuticle. The flowers are sweetly scented. There seems to be a little doubt as to the exact habitat of this species; if, as Lindley stated, it does come from Mexico, it is the only Berberis recorded from there, and provides a marked northerly extension of the range of the genus, into the region of M a b k .

et Frut. Brit. 1,307 and 4,2537, 1838

(450) Berberis rigidifoliaK unth in Humb. et Bonpl., Nov. Gem. 5,70, t. 431, 1821

S t e m terete, glabrous; mature black, subvermculose. Spines 3-fld., 2-6 mm. Leaves 3-5 x 1-14 cm., obovate-elliptic; base cuneate, subswile; apex obtuse, mucronate; margins entire, or towards apex 3-7-spinose (2-2.5 mm.)-serrate (1 mm.), distant 3-6mm. above lustrous bright green, subenervate, or very indistinctly and h e l y openly reticu- late; below with open elevated reticulation, at first slightly pruinose and grey, &ally paler dull green. Racemes 10-15-fld., 4 4 cm. long. PediCeEs glabrous, 6-10 mm. ; bracts 2-3mm. ProphylEs small. Inner Bepala 6x4.5mm. Petals 5x4.5mm. Otherwise unknown.

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Berberis and Mahonia 281

C!olombia : Pbntae Expeditiones Botanicae. Mutisii Vice-Regni Novae Granadae, 1783-1808, no. 108 (K).

(451) Berberis truxillensis Turcz., Bull. SOC. Nat. Mosc. 27(ii), 285, 1855; Pittier, J., Wash. A d . Sci., 21, 136, 1931; syn. B. Zycioides Linden et Planch. ex Triana et Planch., Prod., p. 52, 1862, non Stapf. (Fig. 66)

Stem glabrous, terete, yellow. Internooh 3 4 cm. Spines absent. Leaves 2 0 4 0 x 5- 11 mm. ; obovate; cuneate, sessile; entire, rounded; both sides with close elevated reticu- lation; above slightly lustrous green; below dull yellow green, papillose. Racernee (6-;I 8-12 (-15)-fld., 2-5-3 cm. long. Pedids pubescent, 6-12 mm., sometimes quickly becoming glabrous, but with more persistently pubescent rhachis ; bracts 1-2 mm. Prophyk lanceolate, acuminate, 2 x 0.6 mm. Outer sepals 2-25 x 1-25 mm., ovate. Inner septl 4 x 2.75 mm., obovate. Petals 3.5 x 2.25 mm., obovate. Stamns truncate. 0vzr.les 2. Berries estylose, epruinose, obovoid, 7-5 x 4 mm.

Venezuela: Prov. Truxillo, Agua de Obispo, 9750ft., fr., Funck and Schlim 754 (Type); same locality, Linden 306 (K).

Prov. Zamora, Pasum del Morro, 8400 ft., Jahn 1062.

(452) Berberis prolifica Pittier, J . Wash. A d . Sci. 21, 135, 1931 Shrub 3-6ft. Stems terete, glabrous. Spines palmate, 1-2mm. long. Leaves with

lamina 2 4 . 5 x 1-2 em., excluding decurrent petiole 7-11 mm., the upper 6 5 mm., arti- culate above to the lower 3-6mm.; margins h e l y spinose; venation with 7-8 clear lateral veins, and reticulation faint; below pruinose grey. Racemes many-fld., 4-10 cm. Pedicels glabrous, 8-9 mm.; bracts 5-7 mm. Sepals (five in number), 4-6 x 2 4 mm., ovate; acute or acuminate. Petals (three in number), 8 x 6-6-5 mm., ovate, obtuse; glands 1 x 1.5 x 0.7 mm. Ovules 4-7.

Venezuela: Prov. Mdrida, Quebrada de Saivay, 13,000 ft., fl., fr. 17 Apr. 1930, Gehriger 41 (Type); Canada de Chachapoto, 12,00Oft., fr. 22 Jan. 1922, Pittier 13258. Both localities near San Rafael de Mucuchies.

(453) Berberis jelskiana Schneid., Bull. Herb. Boissier, (2), 5, 808, 1905; Macbride, PI. Peru, in Field Mus. Publ. Bot., 13, 2, p. 675, 1938

Internodes 1-2 em. Leaves 20-30 x 8-12 mm., narrowly obovate, entire; below pruinose grey, papillose ; with finely elevated reticulation. Raeemes lO-15-fld., sometimes compound below, 6 6 em. long. Pedicels puberulous, stout (7-) 10-15 mm. ; bracts 2-3 mm. Flowers 11 mm. diameter. Stamens truncate. Ovules 2. Style absent.

North Peru: Prov. Cajamarca, Cutervo, Jelski 232 (Type).

(454) Berberis edentata Rusby, Mem. N . Y . Bot. Gdn, 7,239, 1927 Leaves thick, rigid, 4-5 x 1.5-2.5 cm.; oblong-ovate; petiole

5 mm.; margins entire, revolute; apex rounded; veins prominent both sides. Race- many-flowered, compound below, with stout rachis. Pedicels 1-2 mm. Petals orbicular 5-6 x 5-6 mm. Stamens shorter than petals. Berries estylose, 5-6 x 4-5 mm.

Bolivia: sunny banks at Pongo de Quine, 11,500 ft., 12 July 1921, Mulford Expedition to the Amazon valley, White 169 (Type). ‘Shrub, 4-6 ft.’

Mature stems black.

(455) Berberis dryandriphylla Diels., Notizbl. Bot. art. Berl. 11,781,1933; Macbride,

Mature stems black, slightly sulcate. Spines 3-fld. Leaves 3-65 x 1.5-2-5 cm.; oblong- obovate; petiole 2-8 mm. ; margins coarsely dentate; openly reticulate. Raeemes many-

Fl. Peru, lx., p. 672, 1938

Page 282: Berberis and Mahonia : A taxonomic revision

282 L. W. A. AEQZENDT flowered, 4 4 cm., including peduncle, c. 1 cm. PerEieels purple, c. 1 om. long. Inner s e p h 5-6 x 3.5 mm., elliptic. Petacls subequal to inner sepals. Stamm 3-5 mm., pro- duced, obtuse to truncate. chrules 2. Styk absent, or indistinct.

Peru: Prov. Cuzco, V d e del Forachtay, 7500 ft., Lucmachayoc, Nov. 1927, Herrera 1742 (Type).

Subsect. Pichinchenaes Schneid., Bull. Herb. Boksier, (2), 5, 805, 1905 Leaves mostly spinose-serrate. Stamem produced, mostly apioulate. oencles 3-7.

Colombia : provinces N d o , Mariquita, btander. Ecuador: Provhces Loja, Pichincha, Bzuay, Guayas.

Be& estyloee.

KEY 88, to Subsect. Pichinchenses Stamem truncate; leaves reticulate, and green below; ovules 4-5.

Stems and pedicels pubegcent; petals as long as inner sepals. Stems and pedicels glabrous; petals shorter than inner sepals.

Leaves with sphose margin. 463. B. aaxcwum

464. B. reicheana Leaves entire. Cf. 466. B. &+mboen&

Leaves entire; bracts 3-5 mm. Stamens apiculate.

Mature stems yellow; pedicels 3-5 mm., equal to their bracts; petals shorter than

Mature stems red; pedicels 5-10 mm., twice as long as their bracts; petals as long inner sepals.

as inner sepals.

Cf. 466. B. chimboensia

465. B. hhllipea Leaves spinose.

Bracts 6 7 mm., subequal to their pedicels; petals longer than inner sepals.

Bracts 2-3 (-7) mm., half to one-sixth as long as their pedicels; petals not longer 461. B.&himhm&

than inner sepals.

reticulate. Leaves with 4-9 marginal spinules; below green, epruinose; bractg P 7 mm.,

half as long as pedicels; petals shorter than inner sepals. Leaves with 2-3 marginal spinules; below white pruinose; bracts 2mm.,

one-quarter to one-sixth as long as their pedicele; petals as long as inner Bp&. 462. B . e n g k r k ~

Pedicels pukcent , 8-15 mm.; leaves with lateral veins visible, but not

456. B. retinerrcia

Pedicels glabrous; leaves reticulate. Stems pale yellow; petals shorter than inner sepals.

Leaves below green, epruinose. 457. B.pdopoda Leaves below grey, pruinose. 458. B. bergerkna

Inflorescence racemose-subfascicled, sometimes compound below. 459. B. &ma Inflorescence racemose. 460. B. lechJerkmz

Stems dark, red to black.

(456) Berberis retinervia Triana et P h c h . apud Wedd., Chkw. And. 2, 296, Nov.

Stems terete; mature dark red, very pubescent. I&& 15-25 mm. Spines absent. L e a v a to 55 x 28 mm., obovate; margins 69-spinose (2-3 mm.)-subserrulate, distant 4-8 mm.; above lustrous deep green, with fine close elevated reticulation; below much paler, dull yellow-green, papillose, with venation slightly branched but not reticulate. Racenzes 10-18-fld., 5-7 cm., including pubescent peduncle, 1.5-2.5 cm. Pedicels pubescent, 8-15 mm.; bracts 4-7 mm. Prophyllls hceolate, 2.5-0-6 mm., acuminate. Outer s e p h 3 x 2 mm., ovate, acute. Outer d i a n sepds 5 x 3 mm., ovate, obtuse.

1861; Ann. Sci. Nat. Boi?., ser. 4, 17, 53, 1862

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Berberis and Mahonia 283

Inner median sepals 5.5 x 4 mm., elliptic. Inner se@ 6.5 x 5 mm., elliptic. Petals 5.5 x 3.3 mm., rhombic-ovate, obtuse, entire, base clawed, with lanceolate glands, 1 x 0.3 mm., contiguous and situated within the claw. Stamens 3.5 mm. Ovules 5-6.

Cblombia: Prov. Nariiio, Volcano of Pasto, 11,700-12,700 ft., Jameson s.n. (Type).

(457) Berberis psilopoda Turcz., Bull. SOC. Nut. Mosc. 27 (ii), 285, 1855; syn. B. toli- mensis Linden et Planch., ex Triana et Planch., Ann. Sci. Nat. Bot., ser. 4, 17,52, 1862

&ems terete; mature glabrous, yellow, venuculose. Spines stout, 3-5-fld., to 6 mm. Leaves 3 4 x 1.5-2 cm., obovate: elliptic; subsessile; obtuse; margins 10-16-spinose (1-1.5 mm.)-subsedate, distant 1 - 6 2 (-3) mm. ; both sides closely reticulate ; above lustrous; below paler green, papillose. Racem~s 12-25-fld., 6 5 cm. long, including peduncle, 1-2 cm., sometimes compound below. Pedicels 6-12 mm.; bracts 2 mm. Prophylls ovate, acute, 2.5 x 1.5 mm. Outer sepals 3.5 x 2 mm., oblong-ovate. Median sepals 5 x 2-75 mm., oblong. Inner sepa.?.s 5.5 x 3.5 mm. obovate. Petals 4-5 x 2.5 mm. Stattttns 3 mm., produced apiculate. Ovules 4. Berries obovoid, 8-9 x 5-6 mm., estylose. pruinose blue.

Colombia: Prov. Mariquita, Tolima, Linden 912 (Type of both psilop& and toli- w,miS).

(458) Berberis bergeriana Schneid., Bull. Herb. Boissier, (2), 5, 805, 1905 Stems terete; mature glabrous, yellow. Internodes c. 2.5 cm. Spines 3-5-fld., terete,

5-10 mm., brown, minutely black-verruculose. Leaves 2 4 x 1-1-14 cm., oblong-ovate, subsessile; margins 5-8-spinose (1.5-2 mm.)-serrate (0.5 mm.)-distant (2-5)- 3-5 mm. ; above lustrous, finely but indefinitely reticdate; below dull, at first pruinose grey, often ultimately slightly paler green, with elevated reticulation. Racemes c. 8 cm., often compound below. P e d i d 8-12 mm. ; bracts 2-3 mm. Flower diameter c. 8 mm. Petals much shorter than inner sepals. Ovules 4-5.

Colombia: without locality, Linden 902 (Type in Herb. Bobs., and 0).

(459) Berberis densa Triana et Planch., Ann. Sci. Nut. Bot., ser. 4, 17, 53, 1862; syn. B. globosa var. &ma Schneid., BUU. Herb. Boissier, (2), 5, 805, 1905

fitems terete; glabrous, mature dark red-purple. Leaves to 30 x 13 mm.; mostly with each margin with 1-2 spinules, occasionally entire; with fine elevated reticulation both sides ; below papillose. Injlorescence racemose-subfascicled or subumbellate, 8-15-fld., 2.5-4 cm. long. Pedieels 10-15 mm.; bracts c. 3 mm. Ovules 3-5.

Colombia: Prov. Santander, Pamplona, Funck and Schlim 1280 (Type).

(460) Berberis lechleriana Schneid., l.c., p. 806, 1905 A'tems terete, glabrous; mature black. Intemzodes 1.5-2 cm. Spines dark brown,

1-3-fld., 3-7 mm. Leaves 17-25 x 6-12 mm., obovate; with 2-3 marginal spinules; reticulate both sides, below yellow-green. Racemes erect, 2-5-5 cm. Pedicels 8 mm.; bracts 3 mm. Flowers 12 mm. diameter. Petals as long as inner sepals. Ovules 3.

8. Ecuador: Prov. Loja (?), Pearce, s.n. (Type, Vienna).

(461) Berberis pichinchensis Turcz., BuU. SOC. Nut. Mow. 27 (ii), 286, 1855 &ems with young shoots puberulous ; mature glabrous. Internodes 2-3 cm. Spines absent

or &7-fld., 2 4 mm. Leaves 2 . 5 4 x 1-2-1.7 cm., ovate-elliptic; petiole 2-5 mm.; margins 6-13-spinose-(1-5-25 mm.)-serrate (1 mm.), distant 3-8 mm. ; above dull, indistinctly reticulate ; below paler, slightly pruinose, papillose, with close elevated reticulation. Racemes sometimes compound below, 10-25-fld., P 5 cm. long, including peduncle

Page 284: Berberis and Mahonia : A taxonomic revision

284 L. W. A. AHRENDT c. 1 cm. Pedkh stout pubescent, 5-8mm.; bracts 6 7 mm. Prophylls lanceolate, 3 x 0.5 mm. outer 8epal.s 2-5 x 1.75 mm. Inner sep& 4 x 2.5 mm., obovate-elliptic. Petals obovate, 6 x 4 nun., entire; glands indistinct. Stamens 4 mm., produced, apicu- late. ovules 4.

Ecuador: Prov. Pichincha, Mt. Pichincha, fl. May and June, 13,00Oft., Jameson 585 (E); Jameson 9 (E).

(462) Berberis engleriana Schneid., Bull. Herb. Boksier, (2), 5, p. 807, 1905 Stem glabrous when old. Internodes 10-15 mm. Spines absent or 2-5 mm. Leaves

thin, 2.5-3-5 x 1 - 6 2 cm.; obovate-elliptic; petiole 6-10 mm.; margins with 2-3 small spinules ; above lustrous, scarcely reticulate ; below pruinose white, Snely reticulate. Racenzes, sometimes compound below, subpendulous, 5-7 cm. Pediceh puberulous, 8-10 mm.; bracts 2 111111. F k r 8 8-10 mm. diameter. Petah aa long as inner sepals. ovules 4.

Ecuador: 13,000 ft., without locality or collector’s name (Type, Vienna).

(463) Berberis saxorum Ahrendt, sp.nov., inter sectionem ramulis pedicellisque pubes- centibus ; foliis reticulatis, spinoso-serratis, subtus viridibus ; sepalis interioribus petalisque aequilongis; staminibus truncatis; ovulis 4 4 , distinguenda

Internodes 1-14 cm. Spines 1-3 (-!j)-fld., 3-6 mm. Leaves to 23 x 13 mm.; elliptic; margins 44-spinose (1.5 mm.)-serrate (0.5 mm.), distant 3-6 mm.; dull both sides; fairly closely reticulate ; below paler green, epapillose. In$ormcence 6-lO-fld., umbellate- facemose, 1-5-2 cm. Prqhylh 3 x 2 mm., ovate, acute. Outer sepals 3.5 x 2.5 mm. Mediun and inner se@ 5-5-5x3.5-4mm., all elliptic. Petals 5-5.5x3*54mm., obovate, entire. Stumm 5 mm., not produced, truncate. Ovules 5. Style absent.

Ecuador: Prov. Pichincha, Mt. Pichincha, on rocks, 14,000 ft., Jameson, s.n. (Type, E).

(464) Berberis reicheana Schneid., I.c., p. 806, 1905 Stem terete. Intemzodes 10-15 mm. Spines stout (1-) 3 (-5)-fld., 6-12 mm. Leaves

20-35 x 8-14 mm.; narrowly obovate; cuneate, subsessile; margins mostly 2-4-spinose; both sides with elevated reticulation; above slightly lustrous; below at h t slightly pruinose grey but ultimately epruinose dull green, papillose. Racemes 3-5-5 cm. Pedicels 5-8 mm.; bracts 2.5-3-5 mm. Outer sep& 3 x 3 mm., ovate. Medkn sepals 6 x 3 mm., elliptic. Inner se@ 7 x 4 mm., obovate. Petals 6 x 2.5 mm., oblong-obovate, cuneate, entire. S&mm 3 mm., not produced, rounded-truncate. Ovules 4-5.

Ecuador: Prop. Azuay, west side of Mt. Azuay, fl. Aug. 1859, Spruce 6033 (Type, Vienna, K). ‘Dense shrub; 8 ft. high; flowers orange.’

(465) Berberis hirtellipes Ahrendt, sp.nov., inter sectionem ram& glabris; pedicellis puberulis; foliis integris tantum 1-2cm. longis, subtus epruinosis viridibus, reticulatis ; sepalis inkrioribus petalisque aequilongis; staminibus apiculatis, distinguenda

Stem glabrous; mature red-brown, verruculose. Internodes 5-15 mm. Spines absent. k v e s to 20 x 7 mm.; petiole 2 mm. ; margins entire; with close elevated reticulation; above lustrous; below paler green, epapillose. InJEoresCence 1.5-2-5 cm., racemose or subumbellate. Pedicel8 pubescent (5, below)-8-10 mm.; bracts 3-5 mm. Inner sepals 5.5 x 8.5 mm. Petals 5.5 x 2-5 mm., both oblong; glands oblong, 1.5 x 0.5 mm., marginal. kVtamns produced, apiculate. Ovules (3-) P 5 . Style absent.

Ecuador: Jameson, s.n., fl. 1825 (Type, E).

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Berberis and Mahonia 285

(466) Berberis chimboensis Schneid., BuU. Herb. Boissier, (2), 5, 811, 1905 Stems terete, glabrous; mature yellow-brown. Intermah 3-4 cm. Spines 3-5-fld.,

stout, 8-15 mm. Leaves 3-54 x 1.5-2.5 cm., or to 6.5 x 2.7 cm. ; obovate; petiole 1 4 mm. ; entire, rarely with 1-2 marginal spinules; apex acute or subacute; both sides with elevated reticulation ; above scarcely lustrous ; below with very fine closer reticulation, papillose, at first slightly pruinose finally dull paler yellow-green. Racemes 3-5-5 cm. Pedicels 3-5 mm.; bracts 3 4 mm. Outer sepals 2.5 x 1.5 mm., ovate, acute. Inner sepak 5 x 2.5 mm., oblong-elliptic. Petals 3 x 2 mm., obovate.

Ecuador: Prov. Chimborazo, Chapacoto, Chimbo, fl. Dec., 8500-9750 ft., Lehmann 5493 (Isotype, K).

Subsect. Keisslerianae Schneid., I.c., 808, 1905 Leaves evergreen. Pedicels 2-5 (-10) mm. ; bracts 1-2 mm. Stamens produced apicu-

late. Plower diameter, 4-6 mm. 0wle.s 1-3. Peru: Prov. Ancachs. Bolivia: Prov. (? ) Larecaja.

KEY 89, to Subsect. Keisslerianae Leaves 8-25 mm.

Leaves with spinose margin; stems terete yellow. Leaf margins with 8-14 spinules, reticulate; pedicels 4-5mm.; pet& shorter than

Leaf margins with 1-5-spindes (rarely entire), only slightly veined; pedicels inner sepals. 467. B. podophyh

6-10 mm.; petals as long as inner sepals. 468. B. brunzalis

469. B. keisaleriana

Stems sulcate. 470. B. rechingerd 471. B. wettateiniarna Stems terete.

Leaves 6-10 cm. 472. B. denaifolia

Leaves entire; pedicels pubescent. Petals longer than inner sepals. Petals only as long as inner sepals.

(467) Berberis podophylla Schneid., Bot .Jb. 42,84,1908; Macbride, Fl. Peru, in Field

Stem glabrous. Spines 3-fld., 10-18mm. Leaves orbicular with a truncate base; margins with 8-14 spinules ; reticulation elevated; below epapillose, pruinose, grey. Racemes 36-5 cm. Pedicek glabrous, P 5 mm.; bracts 1-1.5 mm. Petah slightly shorter than inner sepals. Stamens produced, scarcely apiculate. Ovules 2 (-3). Style absent.

Peru : Prov. Ancachs, Cajabamba between Samanco and Huaraz, 9750 ft., Weberbauer 3120, 3137.

Mus. Publ. Bot., 13, pt. 2, 679, 1938

(468) Berberis brumalis Macbride, CundoUeu, 6,4, 1931; FZ. Peru, I.c., 2,670, 1938 Stems glabrous. Spines absent. Leaves t o 4 x 2 cm.; petiole 7 mm.; margins revolute,

mostly entire, occasionally with 2-5 spinules ; above lustrous and slightly veined; below dull, pruinose, papillose, subenervate. InJoracem unrecorded. Pedicels 6-10 mm. Petals as long as inner sepals and each scarcely 3.5 mm. long. Stamns much produced, apiculate.

Peru: without locality, Weberbauer 6799 (Type).

(469) Berberis keissleriana Schneid., BuU. Herb. Boissier, (2 ) , 5, 808, 1905 Stems terete, or subterete, puberulous; mature yellow-brown. Intemzodes 8-12 mm.

Spines orange, subterete, stout, 3-0d., 15-25 mm. Leaves 8-12 x 2-5 mm., oblanceolate

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286 L. W. A. ARRENDT with short petiole; above grey-green; below papillose; openly reticulate. Racence~ 8-12-fld., 1-5-2-5 cm. P e d W puberulous, 4-5 mm,; bracts 1 - 6 2 mm. Outer sepals 1.5 x 0.5 mm., ovate, a c e a t e . Inner sepak 2 x 1.25 mm., obovate. Petals 2-2-25 mm. S t a m w apicdate. ovules 1-2.

Bolivia: without locality, Cuming 192 (Typ, Vienna); 1850, Bridges, 8.n. (Herb. Boies.).

(470) Berberis rechingeri Schneid., BUU. Herb. Boissier, (2), 5, 808, 1905 Stem sulcata or subsulcate, puberulous; mature dark red or red-brown. Intemwoks

8-12 mm. Spines absent or solitary, 3-8 mm. Leaves 10-16 x 5-8 mm.; obovate, sub- sessile; entire, subacute; both sides with open elevated reticulation; below not, or scarcely, paler green, but with the veins more distinct. Racemes 8-12-fld., 1-15 cm. Pedicds puberulous, 2-3 mm. ; bracts 1-16 mm. Outer selpals 2-7 x 0.8 mm., ovate. Inmr sep& 3 x 2.5 mm., broadly obovate. Petals aa long &B inner sepals. Stamm apiculate. ovules 2.

Bolivia: Prov. ( 2 ) Larecaja, near Combaya, in thickets, Mandon 863 (Type, Vienna; K).

(471) Berberis wettsteiniana Schneid., I.c., 809, 1905 Stems glabrous, scarcely sulcate, m.inutely verrucdose. Internodes 1-5-2 cm. h'pines

1-3-fld., terete, yellow-brown, 8-14 mm. Leaves with hypoderm, 15-25 x 8-15 mm., obovate, cuneate; entire, rarely with 1 spinule to each margin; above openly veined; below, scarcely papillose and more distinctly veined. Racernee many-flowered, compound below, to 4cm. long, including peduncle. Pedicels puberulous, h l l y subglabrous, 3-5 mm.; bracts 1 - 5 4 mm. Flowers c. 6 mm. diameter. Petals scarcely emargimte, but aa long as inner sepals. Stamens shortly apiculate. ovules 3.

Bolivia: without locality, Cuming 195 (Type, Vienna); 1850, Bridges, 8.11. (Herb. Boiss.).

(472) Berberis densifolia Rusby, Descriptdans of Three Hundred New Species of

Stem terete; mature dark, red or purple. Leaves thick, with hypoderm, 6-10 x 2.5- 4 cm. ; margins coarsely dentate. Racemes 5-7 cm., including peduncle 1-26 cm. Petals and inner 8epah of equal length, 6 x 3.5 mm.

Bolivia: Unduavi, 10,OOO ft., Odober 1885, Rnsby 508 (Type); November 1910, Buchtien 2854.

South American Plants, pp. 16, 17, Dec. 1920

Sect. PanicUlatae Schneid., BUU. Herb. BOiSeier, (2), 5, 815, 1905 Sp'nes 3-7-fldd., w d , 2-10 mm., sometimes absent. Stmu mostly terete, occasionally

sulcate, mostly glabrous. Leaves evergreen, more or leas closely reticulate. InJEorescence panimhte, 25-260-fld. Flowers mostly 4-8 (-10) mm. diameter, though in three species &B large aa 1&15 mm. diameter. Pedicds short or long, but always much longer than their brads. Be+ black, mostly pruinose blue, eatylose, or occasionally with style 0.1-0.5 (-1) mm.

Venezuela: Prov. M6rida.. Colombia: Provinces Bolivar 2, Cauca, Mariquita, Cundinamarca. Ecuador: Provinces Cuenca, Loja, Chimborazo, Tunguragua, Pichincha, Azuay. Peru: Provinces Amazonas, Cajamarca, Huhuco, Cuzco. Bolivia: lod t iea uncertain. Range 10' N. to 13" S. lat.

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Berberis and Mahonia 287

KEY 90, to Sect. Paniculatae

Leaves with spinose margins. Stems pubescent; flower diameter 10-15 mm.

Leaves with 10-20marginal spinules, below slightly pruinose grey; panicles 20-40-fld. ;

Leaves with 20-40 marginal spinules, below epruinose green; panicles 100-140-fld. ; outer sepals shorter than median and inner sepals.

outer sepals aa long as median and inner sepals.

473. B. colombiana

474. B. verticillata Sl;ems glabrous; flower diameter &8 (-10) mm.

Leaves epruinose, green, below; pedicels 5-10 mm. Leaves revolute, subenervate above ; pedicels pubescent ; flower diameter

Leaves not, or scarcely, revolute reticulate above; pedicels glabrous; flowers 8-10 mm.; o d e s s 4 . 475. B. pindilicensis

6-8 mm. diameter; ovules, where known, 1-2. Mature stem dark red, terete. 476. B. jamsonii* Mature stems pale yellow, more or leas sulcate.

Leaves with 3-7 marginal spinules; petals shorter than inner sepals.

Leaves with 10-30 marginal spinules; petals equal to or longer than inner 477. B. warezewiczii

sepals. Stems slightly sulcate; panicles 14-22 cm. long; stamens apiculate.

Stems very sulcate; panicles 6-12 cm. long; stamens truncate. 478. B. pectinata

479. B. beauverdiana

Leaves elliptic; mature stems purple. 480. B. schwerinii Leaves orbicular ; mature stems pale yellow. 481. B. jkbrigii

Leaves white, pruinose, below; pedicels 2-5 mm.

Leaves entire. Leaves below pruinose, white or grey.

Pedicels pubendous; leaves revolute, petals shorter than inner sepals. :Pedicels glabrous; leaves not or scarcely revolute.

482. B. nutuw

Flowers 10-14 111121. diameter. 483. B. hyperythra Flowers 7-8 mm. diameter; petals as long as inner sepals.

484. B. qhum

Cf. 485. B. barbeyana

Leaves a t &st pruinose white below. 485. B. barbeyanu Leaves always green below.

Leaves 6-11 cm. long; panicles 20-30-fld.; pedicels 10-25 mm. Leaves 2-6 cm. long; panicles 5C9O-fld.; pedicel8 6-12 mm.

Leaves always, or &ally, green below.

Leaves with margin very revolute. O d e s 4; panicles 25-50-fld. 486. B. eimonSii Ovules 2; panicles 60-250-fld. Cf. 488. B. m d t i f i r a

Leaves not, or scarcely, revolute. Stems distinctly aulcate; leaves above dull, or scarcely lustrous; pedicels

pubescent. Stems dark red; leaves indistinctly veined 8bOV0; panicles 30-40-fld.; and

7-loom. long; flower diameter 7-8mm.; ovules 3-4; stamens apiculate. 487. B. paniculata

Stems pale yellow; leaves closely reticulate above; panicles 50-250-fld., 12-30cm. long; flower diameter 4-5 (-7)mm.; ovules 2; 6tamenS truncate. 488. B. rndifiora

Stems terete or subterete.

Pedicels glabrous. Stem pale yellow.

Leaves to 3.5 cm. long, broad, 1 : 1.6-1-7; pedicel8 5-7 mm. 489. B. bumliaefolio

* Cf. 495. B. a k i e n h u d .

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288 L. W. A. AERENDT

KEY 90 (cont.) Leaves 46.6 cm. long, narrower, 1 : 2.5; pedicele 8-17 nun.

490. B.witeUina Pedicels pubescent.

491. B. loxensis apiculate.

truncab. 492. B. hochreutinerana 493. B. lehmnnii

494. B. w8chqffeltelta

Leaves below concolorous green; petals as long as inner sepals ; stamens

Leaves below paler green; petals shorter than inner sepals; stamens

Stems dark red-purple. Not completely determinate, but seemingly distinguished by its dentate stamens.

(473) Berberis colombiana Ahrendt, spmov., inter sectionem foliis marginibus spinoso- serratis, floribus ma@, sepalis exterioribus parvia, ram& pubescentibus, distinguenda

Stem pubeacent. h a v e s to 7 x 2.2 cm., oblong-elliptic, subsessile; margins 12-18- spinose (1 mm.)-subsemdate; above dull or scarcely lustrous, with coarsely but fairly closely elevated reticulation; below scarcely or slightly pruinose, grey, with distinct branched venation, but less conspicuous reticulation. Panicles 2W-fld. Pedicels very pubescent (elongated pilose, the hairs being about 0.15 mm. long), stout, 8-11 mm. long; bracts 3-5 mm. Flowers 13-15 mm. diameter. ProphyUs 2.5-3 x 1-1.3 mm., triangular. Outer s e p h 4 4 x 2-5-3 mm., oblong-ovate. Median se+ 7-8 x 2-5-3 mm., narrowly oblong-elliptic. Inner sepah 7-84 x 44-55 mm., obovate. Petals 6-7 x 4.55.5 mm., obovate, clawed. Sh- 5 mm., scarcely produced, truncate. Ovules 2.

Colombia: Phramo de Herveo, Fresno, Tolima, light woods and ravines in PAramo, 12,000 ft., fl. 14 Aug. 1939, Hanbury-Tracy 610 (Type, K). ‘A shrub, 4 4 ft. high, with orange flowers.’

(474) Berberis verticillata Turcz., BuU. Xoc. Nat. M a c . 27 (ii), 288, 1855; syn. B. quindiW Tr. et Pl., non Kunth.

Stems puberulous; young shoots subsulcate; mature terete, red-brown. Spines absent, or M-fld., 3-10 (-15) mm. Leaves thick, with hypoderm, to 8.5 x 3 cm. ;narrowly obovate, subeessile, obtuse; margins 2040-spinose (1.5-2 mm.)-subsedate, distant 1-2 mm. ; both sides closely reticulate; above slightly lustrous; below paler green, almost dull, papillose. Pan& 100-140-fld., 20-27 cm. long, with stout rachia, including peduncle 6-10 cm., the subpeduncles of the branched portions being 2-4 cm. Pediuh pubescent. 5-9mm.; bracta usually N m m . long, sometimes foliaceous. Flowers c. lOmm, diameter. ProphyUs 2, lanceolate, acuminate, 5 x 1-5 mm. Outer, median and inner sepah all 7 mm. long, but the outer sepals 2.5 mm. broad, while the median and inner sepals are both 3.5 mm. broad. Petals obovate, 4 x 2 mm.

Colombia: Prov. Cauca, Linden 1122 (K, 0). Prov. Mariquita, near Quindiu, !Criana, s.n., according to Schneider, this being the

source of the synonym.

(475) Berberis pindilicensis Hieron., Bot. Jb. 20, Beibl. 49, p. 12, 1895. (Pig. 67) 8 k m sulcate; mature yellow, glabrous. Spines 3-5-fld., palmate, 4 9 mm. Leaves

with hypoderm, to 6-7 x 2-3-26 cm.; narrowly obovate; petiole 2-6 mm.; apex obtuse or rounded; margins revolute (6) 10-20 (-30)-spinose (1.5-2 mm.)-serrate (06 mm.), (1 .6) 2-2-5 (4) mm. apart; above sublustrous, slightly and indistinctly veined or sub- enervate; below epapillose, scarcely paler yellow-green. Pan& (60-) 100-140-fld.,

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Berberis and Mahonia 289

10--20 cm. long, 3-5 cm. broad, with a stout rachis, including a peduncle 6 9 cm. Pedicels stout, pubescent, 4-6 (-7) mm. ; bracts (2-) 2.5-3-6 ( 4 . 5 ) mm., lanceolate, acuminate. Plowers deep yellow. Prophylls c . 1.5 x 1 mm., broadly ovate, acuminate. Outer sepals 2-5-3 (-3.5) x 2-2.5 mm., ovate-triangular, obtuse. Inner sepals 5-6 x 2.5- 3 mm., elliptic to obovate. Petals 6 5 x 2.75-3.75 mm., obovate. Stamns 3-3-5 mm., produced, apiculate. Ovules 3 4 . Berries globose, 6-7 mm., estylose.

Ecuador: Prov. Azuay, Cuenca, East Pindilic Andes, near Pindilic, about 10,000 ft., Lehmann 4654 (K).

Prov. Loja, without locality, Lehmann 6504 (K), 1857-9, Spruce 5449 (E). f iov. Chimborazo, according to Schneider, south of Chimborazo 1858, Wagner, s.n. I?rov. Tunguragua: Mocha (Ambato), 10,500 ft., fl. July 1939, Sandeman 56 (BM, 0)

‘The only place I saw it, and very restricted here.’

(476) Berberis jamesonii Lindl., J. R. Hort. Soc. 5, 8, 1850; non Lemaire, non Turcz; syn. B. glauca Benth., PI. Hartweg. p. 125, 1843, non Kunth.

Sterns terete, glabrous; mature dark red or red-brown. Internodes 2 - 5 4 cm. Spines 3-fld., 2-5 mm. Leaves without hypoderm, 3-5.5 (-7) x 1.2-2.2 (-2.8 em.); petiole 1-7 mm.; margins by apex with 2-6 spinules; above lustrous, closely finely reticulate; below, a t f i s t slightly pruinose, soon epruinose, and finally much paler dull green, papillose, closely coarsely reticulate. Panicles 30-40 (-50)-fld., 6-8 cm. long. Pedicels glabrous, (8-) 10-15 mm.; bracts 3 mm. ProphyUs lanceolate, acuminate, 2.5 x 0.5 mm. Outer sepals 2.5 x 2 mm., ovate, acute. Median sepals 4 x 3 mm., oblong-ovate, obtuse. Inn.er sepals 5.5 x 4 mm., broadly elliptic. Petals 5 x 2.75 mm., oblong-ovate; apex entire; glands separate, lanceolate, 1 x 0-3 mm. Stamens 3.5 mm., produced, apiculate. Ovules solitary. Berries estylose, pruinose blue. Seeds black, obovoid, 5 x 3 mm.

Ecuador: Hartweg 710 (K). Prov. Pichincha, according to Lindley, seed from Quito was sent to Messrs Veitch, and from St Martha to Glendinnings nursery; the type was derived from cultivated plants from these origins. Lindley remarks: ‘ I t is a very beautiful bush with leaves of the deepest green and very lustrous, nearly three inches long, entire or with a few spinules. Veitch of Exeter says it stood several winters there, but, in very severe weather, the shoots and young wood have been injured.’

(477) Berberis warszewiczii Hieron., Bot. Jb. 20, Beibl. 49, p. 13, 1895 Stems glabrous, yellow-brown. Spines 3-5-fld., 5-7 mm. Leaves with hypoderm, to

5 x 3 cm.; oblong; petiole 2-5 mm., articulate above the blade; apex obtuse; margins 3-7-spinose (1-1.5 mm.) ; both sides lustrous green, reticulate. Panicles glabrous. Pedicels 6-9 mm. Prophylls 1 mm. Inner sepals ovate. Petals oblong-obovate, shorter than inner sepals.

Ecuador: Prov. Azuay, Cuenca, Warszewicz 4 (25).

(478) Berberis pectinata Hieron., l.c., 14, 1895 Sferns glabrous, subsulcate, pale yellow-grey. Leaves to 5-6 x 3-34 cm., with hypo-

derm, oblong-elliptic; petiole 4-7 mm., articulate above the blade; margins 15-30- spinose (1-1.5 mm.)-serrate (1-1.5 mm.) ; both sides somewhat lustrous green, reticulate. Panicles glabrous, 14-23 em. long. Pedicels p5 mm. PrqhyUs 1.8 mm. Inner sepals 4 x 3 mm., obovate-elliptic. Stamem apiculate. Ovules 2.

Petals 4.5 x 2 mm., oblong-obovate.

Ecuador: without locality, Warszewicz 2 (27).

19 JOURN. LINN. SOC. BOTANY-VOL. LVIT

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290 L. W. A, ~ E N D T

(479) Berberis beauverdiana Schneid., BUU. Herb. Boissier, (2), 5, 817, 1905 Stems glabrous, very sulcate; mature yellow. Internocia 2-5-4 cm. Spines 5-7-fld.,

3-9 mm. Leaeces without hypoderm, to 7 x 3 cm., oblong-elliptic; petiole to 2 mm. long; margins 14-20-spinoae (1-1-5 mm.)-subserrulate, distant 2 - 5 4 mm. ; above slightly lustrous, reticulate, though often somewhat in&tinctly ; below c l d , concolorous, epapjlloae, finely reticulate. Pan& 40-70-fld., 6-12cm. long, including peduncle (2-) 5-7 cm. Pedic.i& glabrous, 5-8 mm.; bracts 2-5-3 mm. ProphyUs triangular, 1-1-25 mm. Outer sepal8 3 x 1.5 mm., ovate-elliptic. Inner sepals 4.5 x 2.75 mm., obovate. Petals 4.5 x 3 mm. Stamem 3.5 mm., apex slightly produced, or rounded- truncate. Ovules 2.

Peru: Prov. Amazonas, Mathews s.n. (K). Prov. Cajamarca, Chota, Huambos, Weberbauer 4199, according to Macbride, who

cites a MS. note of Schneider that this is possibly a variety with entire leaves and longer pedicels.

(480) Berberis schwerinii Schneider, BUU. Herb. B&sier, (2), 5, 818, 1905 Stems glabrous, subsulcate; mature dark, purple. Internodes c. 2.5 cm. Spines brown,

3-5-fld., 4 4 mm. Leaves with hypoderm, 2-3.5 x 1.5-2 cm., ovate-elliptic; subseasile; margin 8-15-spinose; apex rounded; sca,rcely reticulate either side; above lustrous; below pruinose white. Panicka 7-9 cm. long. Pediah 4 5 mm. long; bracts 2-3-5 mm. Flowers c. 7 mm. diameter. Petals slightly shorter than inner sepals. Stamens truncate. ovules 2.

Ecuador: Prov. Azuay, around Cuenca (2 ) (Type, Vienna).

(481) Berberis fiebrigii Schneid., Bot. Jb. 42, 85, 1908 k v e - s 3.24 x 3-36 cm., slenderly spinose all round the margins, below white and

papillose. Panicles 4.5 cm. long. P e d a 2-3 mm.; bracts 1-2 mm. Flowers 5 mm. diameter.

Bolivia: Huayavilla, 6OOO ft., fl. 6 Dec. 1903, Fiebrig 2508 (Type).

(482) Berberis nutans Linden et Planch., PI. Columb., p. 8,1863; Sprague, Kew Bull.

Stem glabrous, terete, yellow. Intemzodes P 7 cm. Spines 3-fld., yellow, terete, 5-10 mm. h u e s 4-43 x 1.5-3.5 cm., elliptic or oblong-elliptic; subsessile; margins sub- revolute, entire, or subentire with very few occasional spinulea; apex acute; above lustrous, closely finely reticulate ; below pruinose white, with coarser elevated reticu- lation. Panicles (1%) 20-30 (-40)-fld., 7-18 cm., including subglabrous peduncle, 3-5 cm. ; sometimes not completely paniculate but part raceme and part panicle; rachis slightly puberulous. Pedicek glabrous, 1-2 cm.; bracts 4-6 mm. Prophylls 3 x 1 mm., lanceolate. Outer sepals 4 x 2 mm., oblong. Hedian sepals 5 . M x 3 mm., elliptic. Inner sepals 7 x 4.5-5 mm., oblong-elliptic. Petals 4.5 x 3.54 mm., broadly ovate; base clawed. Stumem 3-5 mm., produced, apiculate. Ovules 4-5. Berries conspicuously pruinose, grey or white, broadly ellipsoid, 9 x 7.5 mm., with style absent or indistinct. Seeds purple.

Venezuela: Prov. Mhrida, Sierra Nevada de Mkrida, 11,600 ft., Schlim, s.n. (Type); Linden 306; Phramo de Conejos, sheltered gullies and terraces iq open phramo, 13,000 ft., fl., fr., 24 Sept. 1938, Hanbury-Tracy 157 (K). ‘Shrub, 7-8 ft., with rich yellow flowers.’ Berberis diswlop and B. moritzii both differ in having a racemose inflorescence and petiolate leaves; B. discolor has, also, pubescent pedicels and subenervate leaves; B. moritzii differs, also, in the epruinose green undersurface of the leaves, and in the petals being as long as the inner sepals.

1926, p. 35, 1926

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Berberis and Mahonia 29 1

(483) Berberis hyperythra Diels, Biblioth. Bot. 29 (Heft l l6 ) , 86, 1937 Stem grey. Spines 1 cm. Leaves chartaceous, lustrous, conspicuously reticulate, with-

out hypoderm ; below paler, greyish, pruinose, less reticulate, papillose, elliptic, or ovate or subobovate-elliptic ; base slenderly cuneate; apex obtuse or shortly acute ; margins thickened, entire or with few spinules, 3-6 x 1-5-3 cm. InJlorescence glabrous, paniculate; peduncles to 5 cm. long. Pedicels slender, 1-1.5 cm. Plowers orange, red without, 1-1.4 cm. diameter. Ovules 6.

Central Ecuador: Prov. Chimborazo, Tipococha, 10,700 ft., fl. 15 Aug. 1933, Diels 544 (Type) . Not cultivated.

(484) Berberis glauca Kunth in Humb. et Bonpl., Nov. Gen. 5 , 71, t,. 333, 1821 Leaves without hypoderm, 6-11 x 2.5-5.5 cm., oblong-elliptic, or oblong-ovate; petiole

1.3-2.3 cm., articulate below the blade; margins undulate, entire, or subentire; both sides with fairly close reticulation; above scarcely lustrous bright green ; below pruinose white, epapillose. Panich 25-30-fld., 10-16 cm., including peduncle, 5-8 cm. Pedicels glabrous (0-8-) 1-2.5 cm. ; bracts 3 mm. Outer sepals 2.5 x 0.6 mm., lanceolate. Median sepals 3 x 1.8 mm., ovate, obtuse. Inner sepals 4-5 x 2.25 mm., oblong-elliptic. Petals 4-5 x 2.75 mm., obovate, entire. Stamens 2.75 mm., produced, rounded. Ovules 2-3. Berries ovoid.

Colombia: Prov. Cundinamarca, before 1867, Holton 676 (K).

(485) Berberis barbeyana Schneid., Bull. Herb. Boissier, (2), 5, 816, 1905 Stem glabrous; young shoots somewhat purplish; mature a t first red-brown, soon

becoming yellow-brown, black-verruculose. Internodes 2-5 cm. Spines mostly solitary, stout, 5-9 mm., sometimes absent or 3-fld. Leaves fairly thick, but without hypoderm, 2-5.8 x 1-2.7 cm., obovate; petiole 3-5 mm. ; apex rounded; margins entire; both sides closely, finely reticulate ; below epapillose, a t first pruinose white, finally almost green and scarcely pruinose. Panicles (6) 50-70 (-90)-fld., (8-) 10-14 cm. long, including peduncle 3-5-5 cm. Pedieels glabrous, 6-12 mm. ; bracts 2.5-3.5 mm. Outer sepals 2.5 x 2 mm., ovate, subacute. Inner sepals and petals both 4.5 x 3 mm., oblong-obovate. Stamens 3-5-3.75 mm., slightly produced, rounded-truncate. Ovules 3 4 . Style very short.

Peru: Prov. Amazonas, Chachapoyas, Mathews, s.n. (Isotype K, 0).

(486) Berberis simonsii Ahrendt, sp.nov., inter sectione foliis integris subtus eprui- nosis opace viridibus, marginibus distincte revolutis ; paniculis floribus 25-50, pedicellis glabris; petalis sepalis interioribus brevioribus ; ovulis 4 ; stylo brevis- simo distincto, distinguenda.

Internodes 2 4 cm. Spines absent, or 3-fld., and 4-8 mm. Leaves to 5.5 x 2.7 cm. ; petiole 3-7 mm. ; margins entire distinctly revolute ; both sides closely reticulate ; below epapillose, dull, but scarcely paler, green. Panicles 7-10 cm., including peduncle, 1 4 . 5 cm. Pedicels glabrous, 3-8 mm.; bracts 1.5-2 mm. ProphyUs triangular, 1.8 x 0.9 mm. Outer sepals 2 - 5 3 x 16-1-75 mm., ovate, obtuse. Median sepals 3.5-4.5 x 1.5- 2-5 mm., elliptic. Inner sepals 5 x 3.5 mm., oblong-elliptic. Petals obovate, 4.5 x 2.5 mm. Stamens 3 . 5 4 mm., produced, obtuse. Ovules 4. Style 0.25-0.5 mm.

Ecuador: without locality, Simons, 8.n. (Type, K); Jameson, s.n. (E).

(487) Berberis paniculata Juss. ex DC., Syst. 2 , 12, 1821 Sfems glabrous, sulcate; mature red-purple. Spines 3 (4)-fld., 5-9 (-11) mm. Leaves

with hypoderm, to 5 x 1-6 cm., subsessile; entire, rarely with 1-2 marginal spinules; above slightly lustrous and indistinctly veined ; below also slightly lustrous concolorous

19-2

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292 L. W. A. AHRENDT green, but finely, closely reticulate. Panicles 3W-fld., narrow, 7-10 cm. Pedicels pubescent, 5-9 mm.; bracts 3-4 mm.; peduncle (2-5-5 em.) and rachis often glabrous. Outersepaki2-5-3 x 2-2-5 mm. Innersepals5 x 3.5 mm., oblong-elliptic. Petals 4 x 3 mm., obovate. S t a m w 4 mm., produced, apiculate. ovules 3-4.

Ecuador: Prov. Pichincha, Quito, Jameson 634 (K, 0); Lehmann 4654 (K). Peru : without locality, Jusaieu s.n., according to Macbride.

(488) Berberis multiflora Benth., PI. Hartweg., p. 124, Dec. 1843 Shrub 6-10 ft. Stem glabrous. Internodes 5-7 cm. Spines 3-fld., 3-7 mm. Leaves

without hypoderm, to 9 x 3-5 cm., obovate, or oblong-obovate, sessile or with petiole 1-2mm.; margins entire, subrevolute, subundulate; both sides with elevated reticu- lation, scarcely lustrous concolorous green and epapillose.

K E Y 9oA Panicles 200-250-fld., 16-30 cm.; pedicels 3-4 mm.; flowers 4.5 mm. diameter.

Panicles 50-1O0-fldd., 6-12 em.; pedicels 6-12 mm.; flowers 10 mm. diameter. 488a. B. mdtijzma vm. na&$?wa

488 b. B. mdtijzma vm. dveclcene

(488a) Berberis multitlora var. multiflora Panicles 20&250-fld., very compound, including peduncle (5-) 8-12 (-15) cm. long,

this and the subpeduhcles of the branched portions (1-2 cm. long) both pubescent; bracts 2-4 mm. Pedicek pubescent, 3-4 mm., with their bracts only 1-2 mm. B”lOwer8 44 .5 mm. diameter. Prophg& lanceolate acuminate, 1-5 mm. Outer sepals 2 x 1 mm., ovate. Inner sepals 3 x 1.5 mm., elliptic. Petah 3 x 2.25 mm., broadly obovate. Stanzens 2 mm., not produced, truncate.

Ecuador: F’rov. h j a , near Saraguro, Hartweg 708 (Type, K).

(488b) Berberis multiflora var. calvescens Schneid., BUU. Herb. Boissier, (2), 5, 816,1905

Stems very sulcate;youngshootsreddish;maturepaleyellow. Leavesthin, to8 x 3*7cm., obovate-elliptic ; petiole 1-10 mm. ; both sides slightly lustrous, concolorous green, with conspicuously branched venation, but finer, closer, reticulation. Panicles 50-100-fld., 6-12cm. long and 6-7cm. broad, including peduncle 3-4cm. Pediwk glabrous, 6-12 mm. ; bracts 3 4 mm., but sometimes foliaceous. Prophylh triangular, 1 x 0.5 mm. Outer sepals 3-3-5 x 2-2.5 mm., ovate, acute. Inner sepah 6 x 3.5 mm., oblong-obovate. Petals 5-25 x 3.5 mm., obovate; glands lanceolate, 1.5 x 0.4 mm., separate, marginal. Stamens 3.5 mm., slightly produced, rounded-truncate. Ovules 2.

Ecuador: 1857-9, Spruce 6037 (Isotype, K).

(489) Berberis bumeliaefolia Schneid., BdI. Herb. Boksier, (2), 5, 818, 1905 Stems glabrous, slender, subterete or finely subsulcate; mature pale yellow. Inte9xxdes

1.5-2.5 cm. Spines absent or 1-3-fld., 2-5 mm. Leaves without hypoderm, 2.5-3.5 x 1.7-2 cm. ; petiole 5-10 mm.; broadly obovate; margins entire; apex rounded; both aides finely reticulate; below subconcolorous green, papillose. Panicles ZMO-fld., 8-12 cm., including peduncle 3-6 cm. Pedi& glabrous, slender, 5-7 mm.; bracts 1-5-2 mm. Outer sepals 2.25 x 1 mm., ovate, acute. Inner sepals 3 - 5 4 x 2.5-3 mm., broadly elliptic or obovate. Petds 4 x 2 mm., obovate. Stamna truncate. Ovules 2.

Bolivk: without locality, Bridges, 8.n. (Type, Herb. Bob., K); Cuming 191 (Vienna).

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Berberis and Mahonia 293

(490) Berberis vitellina Hieron., Bot. Jb. 20, Beibl. 49, p. 15, 1895 Znternodes 4-6 cm. Stem terete, yellow. Spines absent, or 3-fld., 3-6 mm. Leaves

without hypoderm, 4-65 x 1.6-2.4 cm.; petiole very short; margins entire; both sides narrowly reticulate ; below sometimes a t Grst slightly pruinose, finally, or always, pale green; papillose. Panicles 25-35-fld., (8-) 12-16 (-24) cm., including peduncle to 6 cm. Pedicels slender, glabrous, 8-17 mm. ; bracts 1-1.5 (-2) mm. Ovules 2. Berries pruinose bliie, obovoid, 7-5 x 4.5 mm., excluding style c. 1 mm.

Venezuela : Prov. M&rida, Tovar, 1854-5, Fender 16 (K) ; by El Pergual de la Lagunita, not far from Colonia Tovar, Moritz 795. According to Pittier, Aragua, El Pesjuali de la La,gunita, on the road from Caracas to Colonia Tovar, fl. July, Moritz 795 (Type): Colonia Tovar, fl. Dec. 1924, Allart 473; upper belt of Naiguata, forming isolated bushes, fl. 24 May 1913, Pittier 6269; La Cienega, Silla de Caracas, 7600 ft., fl. 21-29 Dec. 1918, Pittier 8303.

Colombia : without locality, Karsten, s.n. (leaves more pruinose below).

(491) Berberis loxensis Benth., PI. Hartw., p. 125, Dec. 1843; Macbride, PI. Peru, in Field Mus. Publ. Bot., 13, pt, 2, 675, 1938

Stem glabrous, terete ; mature pale yellow, verruculose. Internodes 1.5-3.5 cm. Spines ubsent or solitary and 2-5mm. long, occasionally 3-5-fld. Leaves with hypoderm, to 4.5 x 1.7 cm., obovate-elliptic ; subsessile, or petiole 1-3 mm. ; apex mostly obtuse, some- times acute and mucronate ; entire not revolute ; both sides closely reticulate ; lustrous green, epapillose. Panicles 5-9 cm., including peduncle 1-3 cm., 2540-fld. Pedicels puberulous, 3-6 mm. ; bracts 1-1.5 mm. Outer sepals 2 x 1 mm., ovate, acute. Inner sepals 3 x 2 mm., obovate. Petals 3 x 2 mm., obovate. Stamens shortly apiculate. Ovules 2.

Ecuador: Prov. Loja, mountains near Loja (Loxa), Hartweg 709 (Type, K, 0). Peru: Prov. HuBnuco, near Huhuco, 5800 ft., along trail, according to Macbride:

I’rov. Cajamarca, Huambos, 8000 ft., Weberbauer 4199. l3. loxensis is recorded as having been cultivated by Veitch from seed introduced by

Lobb; the cultivated plants were described as having leaves blunt and bright green, flowers very small in erect peduncled panicles standing clear of the leaves, and as having stood (with Veitch) unsheltered through the winter of 1849. Like the few other species of the Aequinoctiales which were cultivated in England in the 1840’5, this has probably been lost to cultivation for a century.

and 6000 ft., Yanano.

(492) Berberis hochreutinerana Macbride, Candollea, 6, 2, 1934; FZ. Peru, 1.c. 673 8tem glabrous, subterete. Spines 3-fld., to 7 mm., fairly slender, being scarcely

1 mm. thick a t their base. Leaves to 7-8 x 3 cm. ; oblong-obovate; apex rounded; entire; above lustrous and finely reticulate; below paler and dull with rather prominent veins. Panicles broad, 10-15 cm. long. Pedicels puberulous 5-7 mm. ; bracts 3 4 mm. Inner sepals 5-53 mm. Petals shorter than inner sepals. Stamens 3.5 mm., with apex scarcely produced.

Peru: Prov. Cuzco, south-east of Cuzco, Mt. Acomayo, 10,000 ft., Weberbauer 6831 (Isotype, K).

(493) Berberis lehmannii Hieron., Bot. Jb. 20, Beibl. 49, p. 15, 1895 Stem terete, glabrous; mature red-purple. Spines 3-fld., 5-9 mm. Leaves without

hypoderm, 6-12 x 4-7 cm. ; petiole 7-15 (-20) mm. ; elliptic, entire; both sides with closely elevated reticulation ; above lustrous green ; below epapillose, dull, slightly paler green. Panicles 50-100-fld., 10-30 cm. long, including peduncle 3-6 cm., the sub- peduncles of the k .anched parts being only 1-2 cm. PerEicels glabrous, 4-8 mm. ; bracts

Page 294: Berberis and Mahonia : A taxonomic revision

294 L. W. A. AERENDT (2-) 3-4 mm. Flowers yellow. Prophylls triangular, 1.5-2 x 1-25 mm. Outer sepal8 2.75 x 1.75 mm., oblong-ovate. Median se@ 4 x 3 mm., oblong-ovate. Inner sepak 4 x 3.25 mm., obovate. Petals 4 4 . 5 x 2-2.5 mm., obovate; apex slightly retuse; base cuneate with oblanceolate glands, 1 x 0.25 mm. Stamens 3 mm., not produced, truncate, or slightly retuse. Ovules 2.

Ecuador: Prov. Azuay, Andes, west of Cuenca, Yerba-Buena, and Molleturo, 8700- 9800 ft., Lehmann 4603 (I.sotype, K).

(494) Berberis verschaffeltischneid., Bull. Herb.BoisSier, (2),5,821,1905; syn. B. jame-

Leava ovate to ovate-oblong, coriaceous; base attenuate ; very glabrous; lustrous above ; below paler ; margins undulate-sinuate-spi-~rrate. Flowers large, chrome yellow, in much branched, many-flowered panicles. Ped&eels puberulous. Stamem dentate.

The type is 6.om a plant cultivated about 1859, flowering in England in May and June, but of unknown origin; apparently, a small shrub.

soni Lemaire, Ill. Hort. 6,201, 1859, non Lindl., non Turcz.

APPENDIX 1, to the genus Berberis

(495) Berberis aldenhamensis Ahrendt, &rd. IUm. 21 Dec. 1946 Stems ; young shoots glabrous; mature deep red, lustrous, somewhat angled and suIcate.

Internodes 2-5-3.5 cm. Spines orange, weak, 3-fld., 5-10 mm., scarcely sulcate below. Leaves deciduous, 3-4-5 x 1-2-2 cm., elliptic; base contracted to decurrent petiole 3-8 mm.; apex acute; margins 8-15-spinose (0 -61 mm.)-subsemdate, distant 3-5 (-7) mm.; texture subcoriaceous; above dull deep green with distinctly impressed open reticulation ; below lustrous, and only slightly paler yellow-green, with less conspicuous venation. Punidea (80-) 100-140 (-2OO)-fld., 13-16 cm. long, very much branched and 7-10 cm. broad, with a thick green glabrous rachis, and peduncle only 1-2 cm. long, the subpedimcles of the branched portions being 1-5 (-10) mm. P e d i d glabrous, green, 5-10 (-12) mm. Prophylls 3, triangular, greenish, 3 x 1 mm. outer sep& similar and sub- equal to these. Inner se@ 3-54 x 3 mm., broadly elliptic. Petals oblong, 3-5 x 2 mm. ; apex entire; basal glands approximate, non-marginal. Stamem 2.5 mm. ; edentate; apex not produced, rounded-truncate. O& 2. Berries 9 x 5 mm., oblong-obovoid, estylose, epruinose, at h t green, red when immature, finally black.

Cultivated: L.A. 504, fl. 19 June 1946; fr. 12 Oct. 1946 (Type, 0). The genus Berberis is so complex that, when presented with unusual cultivated material, it is sometimes desirable to know its history. This plant was grown by the late W. J. Marchant, at Wimborne, Dorset ; he obtained it from a Somerset garden : here, it had arrived when bought by auction at the dispersal of plants from the great garden at Aldenham, Hert- fordshire. Most unfortunately its history goes no further back. The great paniculate inflorescence presented by this plant is shown in (a) the Chinese sections Polyanthae (B. prattii, B. plyantha, B. koehneana), and Sherriiiianae; from these B. aldenharnensis differs in its leaf texture; from the Polyanthae in its lack of style also; from the Sher- riffianae as well in its much greater inflorescence. (b) The Himalayan Asiaticae and Tinctoriae, which have the leaf texture of B. aldenhanzensis, but differ in having much fewer flowered inflorescences of a corymbose-paniculate nature (B. chitria), or in the form of a raceme compound below. (c) The tropical South American Paniculatae, with which this specimen closely corresponds.

Some internal evidence suggests that it may belong to the Paniculatae. Against this conclusion, there are only two, inconclusive, facts: (i) That the leaves are deciduous. (ii) That I can find no record of the introduction of seed and raking of plants of the Berberis of the Aequinoctiales for nearly a century, although a few dried specimens have been collected.

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Berberis and Mahonia 295

Concerning the deciduous nature of the leaves: the Himalayan Tinctoriae in many winters will remain completely, or partly, evergreen (even in Oxfordshire) ; in severe winters they will lose all their leaves in late January or February, appearing thus as defoliated (but otherwise unhurt) evergreens, rather than as deciduous plants. On the contrary, B. aldenhmensis, in my experience, loses its leaves in the autumn (even in a cold greenhouse), and appears genuinely deciduous. It is possible that one of the Aequinoctiales may be deciduous, or, even more likely that it may be able to flourish in England as a deciduous plant.

If B. aldenhamensis is a member of the Aequinoctiales, it appears closest to B. jame- sonii, from which it may be distinguished as follows:

Kay 91 Stems terete; leaves lustrous with 2-6 spinules; panicles 30-50-fld., 6-8 cm. long;

476. B. jamesonii pedicels 10-15 mm. ; petals 5 mm., longer than inner sepals; ovules solitary.

Stems sulcate; leaves dull, with 8-15 spinules; panicles 80-200-Ad., 13-16 cm. long; pedicels 5-10 mm.; petals 3.5 mm., as long as inner sepals; ovules 2.

495. B. aldenharnensis

R. aldenhamensis may also be compared with certain entire leaved species: B. leh- mannii, with terete stems and lustrous leaves; B. multijlora var. multijlora, with ex- tremely sulcate pale yellow stems, and pubescent inflorescence; B. barbeyanu with finally yellow stems, more ovules, and style.

Otherwise, it may be regarded as a possible natural or garden hybrid between the Sects. Polyanthae (e.g. B. koehneanu), and Asiaticae (e.g. B. lycioides). Again it may well be an unrecorded species of Sect. Sherriffianae.

APPENDIX 2. Species of Berberis, group Australes, subgroup Euaustrales, not further classified

(496) Berberis andeana Job, Lilloa, 17, 81, 1949 h'hrub 12-26 ft. high. Stems grey. Spines 1-3-fld. Leaves fascicled 5-10) coriaceous,

ovate-lanceolate, petiolate ; margins entire, sinuate ; apex mucronate ; venation slight. Racenzes 6-12-fld. F h e r 8 golden yellow. Prophylls 4, ovate. Sepals in one series of three. Petals in two series of three. Stamens edentate. Fruit black, pruinose blue.

Only the above facts are furniahed by the author, and no localities or specimens are cited.

(497) Berberis barilochensis Job, Rev. Hm. La P h t a (N.S., sect. Bot.), 5,50, fig, 12, 1942

Ashrub to 5 ft. Internodes 1-1.5 cm. Spines 1-5-2 cm., 3-fld. (but in young plants and shoots, and sometimes a t the base of stems, 3-5-fld.). Leaves coriaceous, ovate-lanceo- late, subsessile; above bright green; below dull green, 10-20 x 6 7 mm.; apex acute, mucronate; margins sinuate, either entire or 1-3-spinose (1-2 mm.)-serrate (0.5 mm.), the teeth spreading. Plowers solitary. Pedicels 1-5-2-5 cm. Outer sepals 4 x 2.5 mm. Inner sepals 6.5 x 4.5 mm. Petals 5 x 4 mm., cuneate, entire; basal glands 1 x 0-5 mm., separate, non-marginal. Stamens edentate, produced, obtuse. Ovules 6-9. Berries black, pruinose blue, subglobose, estylose, 10 mm.

Argentine: Prov. Rio Negro, San Carlos de Bariloche, Cabrera 33,5 Jan. 1935 (Type, La Plata 26118). Not cultivated. This species has some connexion with the few solitary- flowered, many-ovuled species of the Sect. Actinacanthae, cf. B. antucoana, B. hetero- phylla. It is also very close to the Sect. Buxifoliae, and to B. buxifolia itself. It may be a natural hybrid between the two sections.

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296 L. W. A. AHRENDT

PART I11

The Genus Mahonia Nutt.

NOTE ON THE DISTINCTNESS OF THE GENUS, AND THE USE OF THE GENERIC NAMX MAHONIA

T h generic name. The earliest generic name is Odostenwn R a h . , in A m r . Monthly Mag. 2,265, (1817), 4, 192, (1819) and in J . Phys. 89, 259, (1819). The use of the name Odostemon in 1817 was closely followed by the name Mahonia Nuttall, Gen. A m r . 211, n. 307, (1818). During the nineteenth century, however, neither name was much used, and the species were generally referred to the genus Berberis.

In 1901 Fedde revealed and recorded the main structure of the genus. He proposed sectional divisions which expressed the structure so well that, despite the many species since separated or fieshly discovered, his original scheme (covering the thirty-seven species to which he gave a key) must remain unaltered. Fedde marked the beginning of the twentieth century by treating this genus as distinct from Berberis, and for it he used the name Mahonia. This procedure involved the providmg of fresh combinations for many species which had been described as Berberis, the spec& epithets being transferred accordingly. In 1913 Schneider described a number of new species, gave a key to eighteen Asiatic ones, and, using the name Mahonia, continued to regard the genus as distinct. In January 1917 Takeda presented a scholarly paper on the Asiatic species; in this he separated a large number of Himalayan species, many of which had previously been referred to Mahonia napaulensis (a custom parallel to the one which had referred so many Asiatic evergreen Berberis to B. wallichiana). Takeda set a high standard of detailed description, was the h t to investigate very deeply the flower structure, and made important research regarding the most reliable diagnostic characters to be found. However, he made no attempt to claasify the genus, and gave no key; he continued to regard the genus as distinct, and used the name M a h i a .

In 1922 Standley described certain of the Mahonias of North America, giving a key to twenty-two of them. He continued to regard the genus as separate, but revived the earlier generic name Odostenwn. However, the International Rules specify the name Mahonia amongst those to be conserved against the strict application of the rules; consequently the revival of the name Odostemon was soon discontinued.

In 1934 Abrams dealt with the species of the PacSc States, giving a key to ten of these, and using the name Mahonia which has now become thoroughly established (except in a few horticultural circles where the name Berberis still prevails).

The distinctness of the genera. It appears to have been the general custom in the nine- teenth century to mention only two distinctions between Berberis and Mahonia, and then to dismiss these as insdEcient for generic distinctness. The characters cited were : (1) Dentate stamens present in Mahonia, but not in Berberis. (2) Glands to the petals present in Berberis but not in Mahonia. But this is not so. Some (South American) Berberis possess dentate stamens, while probably only between one-quarter and one-half of the recorded species of Mahonia possess them. Again, the petals of Mahonia do possess glands; sometimes these are indistinct and may have been overlooked by the earlier writers; on the other hand, they are, sometimes, quite as distinct as in the genus Berberis.

The distinction between the genera rests on other grounds. The leaves of Berberis are simple, of Mahonia pinnate. This is the one universal distinction to which there is no exception. To this must be added the fact that three-quarters of the species of Mahonia possess a form of inflorescence never found in the simple-leaved Berberis, a fascicle of several dense spikelike racemes. Only the minority share a Berberis-like form of in- florescence. Finally, there are no spines on the stems of Mahonia; but these are nearly

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Berberis and Mahonia 297

always present in Berberis, the exceptions forming a tiny minority. There is thus a con- currence of characters which can be held to justify the separation between the genera on the basis of simple and pinnate leaf.

KEY 92, to the two groups of the genus Mahonia Bracts of the inflorescence persistent and long, (1-) 1.5-4 em. Bracts of the inflorescence deciduous and short, 2-5 (-8) mm.

Group ORIENTALES Group OCCIDENTALES

Group Orientales Ahrendt, bracteis inflorescentiarum persistentibus et (1-) 1 . 5 4 em.

Lea$ets in 1-25 pairs. InJlorescence always a cluster of fascicled racemes, with long basal persistent bracts. Pedicels with floral bracts shorter than, equal to, or often several times longer than themselves. Flowers yellow, almost invariably without pro- phylls. Berries black, generally pruinose blue or grey.

Distribution: Himalayas from Kashmir and Punjab to Bhutan, S.E. Tibet, S.W. China (Szechuan, Yunnan, Hupeh, Kweichou, Kwangsi, Kwangtung, Chekiang), South India, Assam, Manipur, Burma, Annam, Siam, Philippine Islands, Formosa, Sumatra. There is one species from the U.S.A. which is a remarkable exception from the otherwise rigid demarcation between the nature and habitat of the groups.

This group needs no further analysis into sections being coterminous wit'h the Sect. Longibracteatae Fedde.

longis. Folia plerumque crasse rigida hypodermate praedita. (Maps 1 and 46)

Sect. Longibracteatae Fedde, Bot. Jb. 31, 30, Aug. 1901

KEY 93, to species of the Sect. Longibracteatae Leaves below pruinose white, fairly thin, with closely elevated reticulation above.

Leaves below nearly always green, rarely pruinose or white, and then subenervate above. 562. M . hypoleuca

FLoral bracts (4) 6-10 (-15) mm. long, two to six times as long as their pedicels.

Floral bracts (1-) 2-5 (-6) mm., (0.2%) 0 . P 1 (-1.5) times as long as their pedicels. Subsect. LONGIBRACTEATAE

Leaves petiolate, that is with the bare portion of the rachis, below the basal pair of leaflets, (2-) 5.5-15 em. long, occupying from one-third to one-half of the length of the rachis.

Leaves subsessile, that is with the lowest pair of leaflets 1.5-2.5 (-4) cm. above the base of the rachis, the short petiole being only one- twentieth or less of the length of the rachis.

Flowers bracteolate. A Flowers ebracteolate.

Leaflets entire. Subsect. INTEGRIFOLIOLATAE* Leaflets with margins spinose-dentate.

Subsect. NERVOSAE

Leaflets with their margins closely toothed, i.e. with ( 1 s ) 12-26 teeth,

Leaflets with their margins sparsely toothed, i.e. with 2-10 (-12) teeth, 2.5-9 mm. apart. B

1-2 (-2.5) em. apart. Stamens dentate. Subsect. DENTATAE Stamens edentate.

Leaflets distinctly petiolulate. 531. M . bodinieri Leaflets sessile or subsessile.

Leaves with mostly ( 1 s ) 12-25 pairs of leaflets. Leaves with 3-8 (-12) pairs of leaflets.

Stamens apiculate.

* Cf. 526b. M.fEavida var. integrijoliola.

C

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298 L. W. A. AHRENDT

KEY 93 (coat.)

Stamens not apiculate. Racemes loose, with pedicels (0*8-) 1-2 cm. long.

Racemes dense, with pedicels 2-5 (-7) mm. Subsect. DOLICHOPOD~E

Stamens not produced, truncate; style insignificant, i.e.

Stamens produced, rounded, obtuse, or truncate; style absent, or up to 0-7 (-1) mm. long.

significant, i.e. (1-) 1.5-3 (-5) mm. long.

E

F

A. Leaflets contiguous or overlapping, dull above, racemes 5-9 cm. long; pedicels 7-10 mm. ; ovules 6-8; Style 1 mm. 510. M. bracteolata

Lsaflets very remote; racemes 10-20 cm. long; pedicels 4 6 mm. ; ovules 4-5; style very short. 538. M. m i r e i

B. Leaflets with 12-26 marginal teeth. Bracts only 4 x pedicels. Bracts subequal to pedicels.

517. M . wmgnifica

Leaflets contiguous or separate, with 12-16 marginal teeth,with narrow (1 : 4-5) apical cusps measuring 15-20 x 3-4mm.; above subenervate; racemes 7-13 cm. long, fascicled 3-4; pedicels and bracts sub- equal, 5-9 mm. ; glands indistinct.

Leafletscontiguous oroverlapping,with 1&26marginalteeth,withbroad 1 : 2-5) inconspicuous apical cusps measuring 10 x 4 mm.; above re- ticulate; racemes 4-5 cm. long, fascicled 5-8; pedicels and bracts subequal, 3 4 mm.; glands distinct. 508. M. p lyodon ta

Leaflets with 30-50 very fine marginal teeth. 542. M. leptodonta:

C . Racemes compound below; leaflets broad, 1 : 2-2.5. 543. M. klossii

532. M. pachkshirem'8

Racemes not compound below. Style short, 0-5-1 mm. ; leaflets remote, rarely contiguous. Leaflets very narrow, 1 : 5-6; margins with 1-3 teeth. Leaflets broader, or with more teeth. Bracts slightly longer than, or subequal to, pedicels.

548. M. oiwkensis

Leaflets with 6-12 marginal teeth; stamens apiculate; ovules 4-5; glands

Leaflets with 3-6 marginal teeth; stamens truncate; ovules 1-3. distinct. 521. M. lomaPiifolfolia

Ledets slightly variable in shape (1 : 2-54) , mostly 7-10 x 2-2.5 om., very remote being 4-6 cm. apart; glands distinct; ovules 3.

546. M.morrisonensis Leaflets very variable in shape (1 : 1-3-5), to 6 x 14-2 cm.; leas remote,

being 1 4 cm. apart; glands indistinct; ovules 1-2. 515. M . diqarifomnata

Bracts only half as long as pedicels. Leaflets very variable from ovate-orbicular to lanceolate, (1 : 1-4); mar-

ginal spino~e h t h very variable k depth (1-10 -.) and number. Cf. 531. M. bodinicri

Leaflets more constant from oblong-lanceolate to lanceolate (1 : 3-4) ; marginal spinose teeth 3-6 in number, 2-4 mm. deep.

520. M . aristulata 511. M . confertu Style very long, 2-3 mm. ; leaflets conspicuously overlapping.

D. Racemes 5-10 (-15) mm. long, fascicled 5-8 (-9); pedimls longer than, or subequal to, bracts; leaves to 28-32 cm. long.

Bracts only half as long as pedicels. Style 0.2-0.3 mm.; ovules 2 (-3). 525. M . pycnqhylla Style 0.7-0.8 IIUII. Cf. 519. M. jaunsareks

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Berberis and Mahonia 299

Bracts subequal to pedicels. Style 0.2-0.3 mm. ; ovules 1-2 ; leaflets with 2-5 marginal teeth.

Style 1-2 mm.; ovules S 5 ; leaflets with 5-10 marginal teeth. Leaflets thick, overlapping, broad (1 : 1-2) ; petals entire. Leaflets thin, mostly separate, narrower (1 : 1-5-3.5) ; petals emarginate.

523. M . manipurensis

509. M . hicksii

514. M . borealis Racemes 16-30 cm. long, fascicled (6-) 8-15; outer sepals small, 2-2.5 mm.

Leaves up to 28 cm. long; leaflets broad (1 : 1-2.5), with 3-4 marginal spinules; ovules 2-3 (-4).

Style absent. 528. M . salweenensis Style present.

524. M . napadensis 518. M . amnthifolia

Style extremely short; leaflets lustrous above. Style 1 mm. long; leaflets dull above.

Leaves 30-60 cm. long; leaflets narrower (1 : 2-5-3.75); ovules 4-6; style distinct. Leaves 30-45 cm. long; leaflets fairly broad (1 : 2 .5 ) ; racemes fascicled 6 1 0 ;

pedicels 5-10 mm., with much shorter bracts, 3-6 mm.; style 2 mm. 544. M . siamnsis

Leaves 50-60 em. long; leaflets narrow (1 : 3.25-3.75); racemes fascicled 12-15; pedicels 4-5 mm., with slightly longer bracts, 5-6 m.; style 1 mm. 526. M.Jlavida

E. Style absent; fruit 15 x 10 mm. ; leaflets pruinose, whitish below.

Style present (except in M . tikushiensis); fruit 6-9 mm.; leaves green below. 529. M . sheridaniana

Leaflets in more than 3 pairs; pedicels 3-7 mm. Petals as long as, or longer than, inner sepals.

Petals shorter than inner sepals. Bracts longer than pedicels. Bracts shorter than pedicels.

536. M . schochii or 537 M . nitens (Key 98)

563. M . huiliensis

533. M . roxburghii

530. M. confusa

Outer sepals fairly large, 4 mm. long. Outer sepals small, mostly 1.5-2-5 111111. long.

Leaflets very narrow (1 : 6), base cuneate. Leaflets broader (1 : I-4), base truncate or rounded.

Bracts 4-7 mm. long, 0-7-0.9 times as long as their pedicels. Leaflets more or less narrow, 1 : 2-5.

Racemes 1625 cm. long; bracts about two-thirds as long &S their

Racemes 3-6 em. long; bracts subequal to their pedicels. pedicels. 541. M . japonica

549. M . discolorzfolia

539. M . fargesii

Ovules 6; fruit estylose. 547. M . tikushielzais Ovules (2-) 4-5; fruit stylose.

Leaflets broad, 1 : 1-2; bracts subequal to their pedicels.

Bracts 1.5-3 (-4) mm. long, 04-0.6 times as long as their pedicels.

Leaflets mostly, or all, contiguous or overlapping. Leaflets finely spinose-serrate. Leaflets coarsely aristate-dentate.

522. M . simonsii

545. M . sumatremiis 540b. M . bealei var. planifolia

Fruit globose, 5-6 mm. Fruit ellipsoid, 10 x 6 mm.

Leaflets mostly remote or separate. Leaflets with margins finely spinulose-serrate.

Leaflets in 4 4 pairs, lowest 2-3 cm. above base.

Leaflets in 6 8 pairs, lowest 1-1.5 cm. above base. 527. M . honcockhna

535. M.fordii

Page 300: Berberis and Mahonia : A taxonomic revision

300 L. W. A. AHRENDT

KEY 93 (cont.) Leaflets with margins comely aristate-dentate.

Leaflets subenervate; racemes 5-10 cm. ; ovules 3-4.

Leaflets h e l y subreticulate; racemes 2-3 cm. ; ovules 2. 540. M . bealei

534. M.ponzensk Cf. 557b. M . fortunei var. szechuanica Leaflets in 1-3 pairs; pedicels 1-2 mm.

F. Racemes fascicled more than 12; pedicela 2-3 mm.; bracts 2 mm.

Racemes fascicled 5-8; pedicels 5-7 111111. ; bracts 3-6 mm. Cf. 516. M . grifithii

Ovules 4; leaflets openly reticulate; margins with 3-7 coarse teeth.

Ovules 5 ; leaflets closely reticulate ; margins with 8-12 finely appressed spinules. 512. M . dolichostylis

513. M . dwlowiana

Subsect. Eufongibracteatae, subsect.nov., bracteis floratis quam pedicellis 2-4-plo longioribus

Leaves with 1-8 pairs of leaflets, the latter with 3-9 (-15) pairs of marginal teeth. Pedieels 2-5 mm.; bracts (6) 6-10 (-15) mm., two to four times as long as their pedicels. Prophylls absent. Stamens edentate, with apex not, or scarcely, produced and rounded or truncate. Ovules 1 4 .

S.E. Tibet (Monyd, Tsari, Zayul), Manipur, S.W. Szechuan, W. Yunnan, Kwangsi.

KEY 94, to Subsect. Eulongibracteatae Leaves with only 1-2 pairs of leaflets, and petiole 2-4-5 cm. long. Leaves with more leaflets, and petiole scarcely 1.5 cm. long.

499. M . bijuga

500. M . subimbricata Leaves with 11-13 pairs of leaflets. Leaves with (3-) 6 7 (-9) pairs of leaflets.

Style absent or very short. Flower bracts very broad, .encircling the pedicel. 501. M . eurybracteata Flower bracts of usual narrower proportions, merely supporting the pedicel.

Leaflets with a conspicuously narrow (1 : 4-6) cusp at the apex; margins with

Leaflets narrow (1 : 4); ovules 4; bracts 4-5 mm., twice as long as their

Leafleets less narrow (1 : 3); ovules 1 (-2); bracts 6-12 mm., three to four

Leaflets with inconspicuous broad cusp (1 : 1-5-2.5) at the apex; mar@ with

(5) 7-18 teeth.

pedicels. 502. M . taronensis

times as long as their pedicels. 503. M . veitchiorurn

3-9 teeth. Petals as long as inner sepals; ovules 4. 504. M . calumicaulis Petals shorter than inner sepals; ovules 2. 505. M . longibracteatu

Style 1-1-5 mm. long. Racemes 11-18 cm. long; bracts 3-6 times as long as their pedicek.

Racemes 6-11 em. long; bracts twice as long as their pedicels. 506. M . monyulensis

507. M . feddei

(499) Mahonia bijuga Handel-Mazzetti, Symbolm Sinicae, 7, 331, fig. 6, 1931 Leajets with margins spinose (1.5 mm.)-serrate (4-5 mm.), distant 7-14 mm., remote;

apex with inconspicuous cusp, c. 23 x 10 mm. Pedicels 2-26 mm.; bracts 4-5 mm. Otherwise unknown.

Szechuan: Ningyuen, 10,700 ft., 16 Apr. 1914, Handel-Mazzetti 1430 (Type).

Page 301: Berberis and Mahonia : A taxonomic revision

30 1 Berberis and Mahonia

(500) Mahonia subimbricata W. Y. Chun & F. Chun, J . A m . Arb. 29, 420, 1948 Leaves 12-22 cm. long, very close and overlapping with 11-13 pairs of leaflets, the

lowcr pairs always conspicuously overlapping, the upper overlapping or a t least con- tiguous. Leajets, the lateral pairs slightly unequal, slightly decreasing in size from the middle (where they are 2.5-3.5 x 1.5 cm.) to the ends where they are 1-5-2.5 (-3.5) cm. long, varying from ovate to narrowly ovate; apex with long, acute cusp; margins with 3-5 teeth above, and 6-8 teeth below; stiffly coriaceous to rigid; above olive green, below at first slightly greyish pruinose, finally sublustrous yellow-green ; with three primary veins slightly impressed above, and no veinlets. Racemes 7-9 cm. long, fascicled, c. 13, the flowers set c. 4 mm. apart, and the bracts of the inflorescence measuring 12 x 5 mm. Pedicels slender, 3 mm.; flwal bracts 6 mm. Outer sepals 2 x 1.5 mm., broadly ovate. Median sepals (3 x 2 mm.) and inner sepals (3 x 1-75 mm.) both oblong-obovate. Petab as long as inner sepals or slightly shorter; apex obtuse, entire; glands linear-elliptic, spreading. Stamens produced, obtuse. Ovules solitary.

Kwangsi : Ching-Hsi hsien, An-teh district, Tung-kong village, shrub on forested slopes, flowers yellow, 21 Sept. 1935, S. P. KO 55791 (Type, National University of Kwangsi). Not cultivated.

(501) Mahonia eurybracteata Fedde, Bot. Jb. 31, 127, 1901 Leaves to 30-35 cm. long, with 5-8 pairs of leaflets, the lowest half the size of the

others and very close to the base of the rachis. Leajets lustrous above, below dull paler yellow green; oblong; base cuneate; margins revolute, with 2-5 teeth towards the apex; with some lateral veins slightly branched into veinlets, 7-12 x 2.5-3.5 cm. Racemes 6-15 cm., with their bracts 2 x 1-25-1075 cm. Pedicels 2 mm.; floral bracts c . 3 mm. Outer sepals small. Petals shorter than the inner sepals. Style absent.

Szechuan: by Chiachu-pa, Nanchnan, 1891, Bock & Rosthorn 1251 (Type). Not cultivated.

(502) Mahonia taronensis Handel-Mazzetti, Anz. Akud. Wiss. Wien, Math.-2Vat. 60

Shrub 3 4 ft. Leaves 1 8 4 2 cm. long, with 5-7 pairs of leaflets, the lowest pair small and close to the base. Leajets 6-13 x 1.5-3 cm., lanceolate, sessile; margins 12-20- spinose-serrate (1-3 mm.) ; cusp conspicuously narrow, 1 :4; texture thin; above dull olive green; below yellow-green; with deeply impressed narrow midrib, 5-8 very oblique lateral veins, and some more branched venation. Racemes 5-8 cm., fascicled 3-5. Pedzcels 2-2.5 mm.; bracts 4-5 mm. Berries globose, 6 mm., estylose, pruinose blue.

S.E. Tibet : Prov. Zayul, near Burma border, Taron valley, Djion-djiang, 27" 55', 9400-9700 ft., 9 July 1916, Handel-Mazzetti 9447 (Type).

(503) Mahonia veitchiorum (Hemsl. et Wils.) Schneid. in Sargent, PI. Wils. 1, 383, 1913; Takeda, Notes R. Bot. Card. Edinb. 6, 232, 1917; syn. Berberis veitchiorum Hemsl. et Wils., Kew Bull. 1906, 152, 1906

.Leaves oblong, with the lowest pair of leaflets small and c. 5 mm. above the base. Leajets fairly thin, mostly oblong-elliptic, sometimes oblong-ovate, all almost equal, or all slightly increasing in size from base to apex of the rachis, 6-9 (-12) x 2-3 (4) cm., the basal pair being only 2-3 x 1-2 cm. ; the terminal leaflet 10-11 x 4 4 . 5 cm., elliptic; base of leaflets cuneate or rounded; margins spinose-serrate; apex with conspicuous elongated narrowly acute cusp ; above slightly lustrous with venation composed of distinct, and sometimes impressed lateral veins and veinlets, not reticulate ; below paler yellow green with similarly branched, but elevated, venation, sometimes slightly reticu- late. Racem~s 6-12 cm., fascicled 4 4 ; bracts of the idorescence 20 x 3-5 mm. Pedicels

(1923), 181, 1924; Symbolae Sinicae, 7, 328, 1931

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302 L. W. A. AHRENDT 2-3 (4) mm.; bracts 6-12 mm. Outer sepals 5.75 x 2-25 mm., oblong-ovate, acute. Median se+ 6 x 2.75 mm., oblong-elliptic. Inner sepah 7 x 3 mm., obovate. P e a 5.5 x 2-25 mm., obovate; acutely emarginate; with rounded lobes; basal glands distinct. Ovules 2. Fruit with very short or indistinct style.

KEY 94A Leaflets with 10-18 mrrrginal teeth, and narrow apical cusps (1 : 4) ; floral bracts 8-12 mm.

Leaflets with 7-9 marginal teeth, and narrower apical cusps (1 : 5-6) ; floral bracts 6-8 m. 503a. M . veitchiorunz vm. weitchiorurn

503 b. M . veit&aOrUm var. k ingdm-mdhna

(503a) Mahonia veitchiorum var. veitchiorum Leaves to 32-40 x 10-12 cm., with 4 4 (-7) pairs of very oblique contiguous leaflets

with decreasing internodes, 5 6 - 4 6 cm. Leajfets with margins 10-18-spinose (1.5- 2-5 mm.)-serrate (1-1-5 mm.), distant 4-43 mm.; apex with cusp 20125 x 4-6 mm. Fruit subglobose, 5 mm.

Szechuan: cliffs, 2000-4000 ft., fl. July 1903, Wilson (Veitch) 3142 (Type, K), shrub 2 4 ft.; Mt. Omi, fl. June 1904, Wilson (Veitch) 4725 (K), 4 ft.; Henry 8993 (K).

Yunnan: amongst scrub, open situations in side valleys, fr. Forrest 7890 (K), a shrub, 3-5 ft., in the hills to the eaat of Tengyueh, at 6000-7000 ft. ; Machang-Kai valley, north of Tengyueh, a t 6000-7000 ft., a shrub 4-8 ft., with bright yellow fragrant flowers, in open situations on forest margins, Forrest 9756 (K).

S.E. Tibet: Prov. Zayul, near Rima, by Assam border, 6500-7500 ft., in broad-leafed forest with smttered conifers; undershrub, stems erect, 3 ft. flowers bright cobalt yellow, 29 Mar. 1950, Kingdon-Ward 19266 (BM). This also has a very narrow cusp, 17-23x 3-4 mm., and is an example of the Chine-Tibetan flora of the Upper Lohit (Zayul Chu) valley on which Kingdon-Ward has remarked. Its serrations are deeper (2-4 mm.).

Not cultivated, but apparently worthy of introduction.

(503 b) Mahonia veitchiorum var. kingdon-wardiana Ahrendt, var.nov., jugis foliolorum pluribus; foliolis marginibus parce dentatis, apice angustiore cus- pidatis, magis remotis, distinguenda.

h u e s 14-27 x 10-12 cm. Leaflets in 5-8 pairs separate or remote, with their inter- nodes slightly decreasing from 3.5 to 3 cm.; margins 7-9-spinose (2-3 mm.)-serrate (2-3 mm.), distant 6-10 mm. ; set very obliquely on the rachis. Fruit 6-7 x 4 4 . 5 mm., ellipsoid.

S.E. Tibet: Tsari district, Migyitun, 9000-10,0o0 ft., shrub 2-4 ft. high in fruit in the forest, usually in colonies, 10 July 1935, Kingdon-Ward 11956 (Type, BM).

(504) Mahonia calamicaulis Spare et E'isch., Rew BUZZ. 1929, p. 249,1929; Chatterjee Rec. Bot. Surv., India, 16, No. 2, p. 38 (1953)

Shrub 2-3 ft. Leaves 15-30 cm., with 6 7 p& of sessile leaflets. h$h% with the small lowest pair generally only 5mm. from the base; but occasionally this small, (3 x 11.3 cm.) basal pair are absent leaving a pseudo-petiole 5-10 cm. long; the remaining leaflets increase in size from the base of the rachis either to the middle or the apex, 76-11.5 x 2-5-56 cm., ovate to lanceolate; margins with 5-9 spinose teeth; apex cusp broad and inconspicuous ; base cuneate ; venation slightly branched, not prominent. Racemes 6-12 cm. long, fascicled, with peduncle to 2 cm., and bracts of the inflorescence c. 1.7 cm. long. Pedicel8 3 4 mm., with floral bracts 6-10 mm. long. Outer sepals small,

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Berberis and Mahonia 303

ovate-suborbicular. Median and inner sepals equal, oblong, 5 mm. long. Petals about 5 mm. long, narrowly obovate, slightly emarginate with obtuse lobes. Ovules 4.

S.E. Tibet: Prov. Zayul, Delei valley, 7000-8000 ft.; dwarf erect undershrub, of the temperate rain forest, in shade under trees on ridges; flowers bright yellow, fragrant; leaves very hard and leathery; Kingdon-Ward 8058 (Type, K). Not in cultivation.

(505) Mahonia longibracteata Takeda, Notes R. Bot. Card. Edinb, 6 , 236, 1917 Shrub 6-10 ft. Leaves to 25 x 11 cm., oblong-lanceolate, with 5-6 pairs of contiguous

or overlapping leaflets, whose internodes decrease, 4-5-3 cm., the basal pair being only 2.5 x 1.5 cm., c. 5 mm. above the base of the rachis. Leaflets fairly thick, ovate, sub- equal, but slightly elongating from base to apex, 5-7 x 3-3-5 em., the terminal leaflet being larger and with a petiole 1 cm. long; margins 3-5-spinose (2-3 mm.)-serrate (2-4 mm.) distant 1-2 cm. ; apical cusp conspicuous, 15-17 x 5-8 mm. ; above lustrous with fairly close h e l y impressed reticulation ; below paler yellow green, much less veined. Outer sepals 6 6 x 1-2 mm., ovate. Median sepals 5-6 x 3 mm., oblong-elliptic. Inner sep ls 5-6 x 3 mm., obovate. Petals 4.5 x 3 mm., oblong-elliptic with indistinct glands. ovules 2.

W. Yunnan: East flank of Tali range, open situations amongst scrub, 8000-10,000 ft., July-Aug. 1908, Forrest 4345 (Type, K). Not cultivated.

(506) Mahonia monyulensis Ahrendt, sp.nov., a speciebus precedentibus subsectionis

Leaves 2448 x 11-16 cm., narrowly elliptic, with 5-9 (-10) pairs of leaflets with inter- nodes decreasing from 5 to 1.5 cm., and their size decreasing from the middle to the ends, the basal pair being 10-15 mm. above the base. Stirpules linear-oblong, 10 x 1 mm. Leaflets 5-5-9 x 3 4 . 5 cm., mostly contiguous or overlapping, rigid, thick; margins 2 4 (-5)-spinose (2.5 mm.)-dentate (5-6 mm.), distant 1-7-35 cm.; apical cusp 15-20 x 10 mm., acuminate, fairly distinct; both sides slightly lustrous, with open venation, not reticulate ; above with venation indistinct ; below paler yellow green with venation elevated. Racemes 11-18 cm. long, fascicled 5-11 ; bracts of the inflorescence 4 cm. x 5-6 mm. Pedicels slender, 1.5-24 mm.; floral bracts lanceolate, acuminate, 5-9 (-15) x 1-5 (-3) mm. Outer sepals 2.5 x 1.5 mm., oblong. Median sepals 4 x 2.75 mm., elliptic- obovate. Inner sepals 6 x 3 mm., elliptic. Petals 4.5 x 2.25 mm., elliptic-obovate; apex shortly acutely emarginate, with two acute lobes; basal glands distinct. Stamens 4 mm. Ovules solitary. Style 1 mm. or longer.

S.E. Tibet: Prov. Monyul, on the East Bhutan frontier, between Pang Chen and Le, Nyam Jang Chu, 7000 ft., a few shrubs only, seen along the river bank, flowers golden yellow, 28 Mar. 1936, Ludlow & Sherriff 1247 (Type, BM). Not in cultivation.

stylo longo distinguenda; a M . feddei ut in clavi 94 distincta.

(507) Mahonia feddei Ahrendt, sp.nov., a M . monyulensi racemis brevioribus, bracteis

Leaves with 4-5 (-6) pairs of leaflets, the basal pair, measuring only 3-5 x 2-3.5 em., being situated 10-15 mm. above the base of the rachis. Leaflets, apart from the basal pair, increasing in size from base to apex, from 7-5 x 5 to 9 x 6 cm. ; terminal one 11 >: 8 cm.; oblong-elliptic; margins 3-6-sphose (2-3 mm.)-dentate (4-6 mm.), distant 12-25 mm.; apex with indistinct cusp; openly and loosely veined; much paler yellow green below. Racemes 6-11 cm. Pedicels 1-2 mm.; bracts 3-4 mm. Flowers similar to the preceding species. Stamens produced, truncate. Style c. 1 mm.

Manipur: Sirhoi, on the edge of forest, on lee side of ridge, 7500 ft., shrub 4-6 ft., flowers golden yellow, 7 Apr. 1948, Kingdon-Ward 17222 (Type, BM). Not cultivated.

florum pedicellis eorum tantum duplo longioribus distincta.

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304 L. W. A. AEIRENDT

Subsect. Dolichostyles Ahrendt, subsect. nov., f o b jugis foliolorum plerumque 4-9; jugis infimis multo minoribus supra b a s h rhachiais tantum P10mm. insertis; foliolis subtus epruinosis, flavo-viridibus, marginibus eorum plerumque dentibus 3-8; pedicellis 3-10 mm. longis, bracteis florum plerumque multo brevioribus, raro aequilongis; stylis 1-3 mm. longis

E. Bhutan, Yunnan, Szechuan. Leaflets in (3-) 4-9 (-20) pairs; margins with 3-8 (-20) teeth; below yellow-green;

lowest leaflets small, close to base. Racemes 4-20 cm. Pedicels 3-10 mm.; bracts 2-5 mm., occasionally equal to the pedicels, generally much shorter. Sturnns edentate. Ovule8 4-8, as far as is recorded. Style 1-3 mm.

KEY 95, to Subsect. Dolichostyles Bracts as long BB pedicels.

Leaflets with 12-26 marginal teeth. Leaflets with 3-8 (-10) marginal teeth.

508. M . polyodonta 509. M . hicksii

Bracts only one-third to two-thirds as long as pedicels; leaflets with 3-9 (-12) marghal teeth.

Prophylls present; ovulea 6-8. Prophylls absent.

510. M.bracleolaSa

511. M . cmferta Leafteb in 10-20 markedly overlapping pairs. Leaflets in 5-9 mostly separate or remote pairs.

P e a as long as inner sepals. 512. M . dOl'khoStyli8 Petals shorter than inner sepals. 513. M. duclouxhm

(508) Mahonia polyodonta Fedde, Bot. Jb. 31, 126, 16 Aug. 1901; Takeda, Notes

Leaves 20-30 cm. long, with 5-6 pairs of leaflets, the basal pair small, 2-3 x 1-1.5 cm., very close to the base of the rschis. Leu$& oblong-ovate, the second pair measuring 3 4 x 2-3 cm., and the others 7-10 x 2-3 (4) cm. ; terminal leaflet with petiole c. 1.5 cm. ; base truncate or subcordate; margins with 3-12 teeth on the lowest three pairs, and 20-26 teeth on the upper pairs; dull above; lustrous below. Racenzes 4-8 cm. Pediceh 3 4 mm.; bracts 3-4 mm. Berries ovoid, 4-6 mm., excluding style 1 mm., pruinose blue.

R. Bot. Gard. Edinb., 6 , 236, 1917

Szechuan: Bock & Rosthorn 2043 (Type). Not cultivated.

(509) Mahonia hicksii Ahrendt, sp.nov., a M. p l y h t a dentibus pauculis distincta,

Leaves c. 36 x 14 cm., oblong-oblanceolate, with the internodes of the leaflets de- creasing from 5 to 1-5 cm., on ascending the rhachis, the small (2.5 x 2.5 cm.) lowest pair of leaflets being only 1 cm. above the base of the rachis. Leafets in 7-9' pairs; apart from the basal pair, their length increases from 6 cm. at the base of the rachis to 8 cm. at the mid-point, thence decreasing to 6.5 cm. at the apex; their breadth decreases con- tinuously from 6 cm. at the base to 3 cm. at the apex; their shape thus changes from ovate-orbicular to narrowly ovate; all the leaflets are set obliquely, and the terminal leaflet measures 7 x 4 cm. ; bases truncate; margins (3-) 5-8 (-10)-spinose (1-2 mm.)- dentate (5 mm., at base to 2 mm. at apical), distant 8-12 mm. ; apical cusp 18 x 8 mm., inconspicuous ; the pairs are contiguous below, but mostly overlapping, especially above ; venation indistinct above, branched below. Racem~s 7-13 cm. long, fascicled, c. 8. Pedicd.~ 2 - 5 4 (-5) mm. ; bracts 3-4 (-5) mm., subequal to them or very slightly longer. Outersepah 45x205 mm. Mediansepals 545x3 mm. Inner sepals 6.5x3.5 mm., all the sepals being ovate. Petals 5 x 2-5 mm., narrowly obovate, entire. Stamens edentate, with apiculate apex. Ovules 4 4 . Style 1 mm. or more.

E. Bhutan: !l'raahiyangse Dzong, scrub land on dry hillside, 5500 ft., flowers yellow, 27 Apr. 1949, Ludlow, S h e d & Hicks 20200 (Type, BM). Not in cultivation.

a speciebus ceteris subsectionis bracteis pedicellisque aequilongis distinguenda.

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Berberis and Mahonia 305

(510) Mahonia bracteolata Takeda, Notes R . Bot. Gard, Edinb, 6 ,228, Jan. 1917 Leaves to 30 x 12 cm., oblong, with (3-) 4-5 (-7) oblique, contiguous or overlapping

pairs of leaflets, their internodes decreasing (3.5-) 3-1.8 cm., from base to apex of the rachis, the much smaller basal pair being c. 1 cm. from the base. Leajleb thick, rigid, ovate; apart from the basal pair fairly equal, 7-8 x 2.5-3.5 cm., the terminal leaflet being scarcely larger but with a petiole 1-1.5 cm. long; bases broadly cuneate; margins 5-8 (-12)-spinose (1.5-3 mm.)-serrate (1.5-3 mm.), distant 12-18 mm.; apex with a very inconspicuous cusp, 8 x 6 mm. ; both sides dull; above grey-green, indistinctly veined ; below paler, subenervate. Racemes 5-9 (-10) cm. long, fascicled 3-4; bracts of the in- florescence 10-15 x 7-10 mm. Pedicels (5-) 7-10 mm. ; floral bracts 2-3 (4) mm. Pro- phylls absent a t apex ofracemes, but present (1-2) a t the base, similar to the bracts but smaller. Outer sepals 3 x 2.5 mm., ovate, acute. Median sepals 5 x 3.5 mm., ovate, obtuse. Inner sepals 5 x 4 mm., oblong-ovate. Petals 7 x 3 mm., oblong-elliptic; apex slightly retuse ; glands distinct. Stamens 4.5 mm., produced, obtusely subapiculate. Ovules 6-8. Style c . 1 mm.

Yunnan : dry and open situations in the margins of pine forests on the divide between the Hoching and Lichiang valleys, 7000-9000 ft., fl. 10 Nov., Forrest 7364 (Type, E, K). Shrub 3-7 ft. with flowers bright yellow tinged purplish in the bud, and fragrant. Not cultivated.

(511) Mahonia conferta Takeda, I.c., p. 230, Jan. 1917 Leaves to 35 x 7 cm., oblong-obbnceolate to oblong-subspathulate, with 10-20 pairs of

obliquely pointing, overlapping leaflets, their internodes decreasing along the rachis from 3 to 1.3 cm., the much smaller basal pair situated c. 5 mm. above the base of the rachis. LeafEets thick, rigid, oblong-ovate, with the length increasing from 3 to 5 cm. from base to apex, and the breadth slightly decreasing from 2.5 to 1.8 cm., so that the shape changes from broadly to narrowly ovate ; bases truncate ; margins 34-spinose- (2-2.5 mm.)-serrate (3-5 mm.), distant 1-2 cm.; apex with a fairly conspicuous but bent cusp, 25 x 8 mm. ; terminal leaflet c. 6 x 3 cm. ; above lustrous, subenervate, below lustrous, paler green, loosely veined. Racenzes 12-18 cm. long, fascicled 4-6; bracts of the inflorescence 10-15 mm. long. Pedicels slender, c. 6 mm.; floral bracts 3 4 (-5) mm. Outer sepals 2.75 x 2.25 mm., oblong-ovate. Median sepals 5 x 3 mm., elliptic. Inner sepals 9 x 4 mm., oblong-ovate. Petals 7.75 x 3-5 mm., oblong-ovate; glands distinct. Stamens 5 mm., slightly thickened below the anthers; apex produced, shortly apiculate. Ovules 3. Style c . 1 mm.

Yunnan: Feng chen Lin, mountain forests, 6500ft., fl., shrub 4ft., Henry 10180 A (Type, K). Not in cultivation.

(512) Mahonia dolichostylis Takeda, I.c., p. 229, Jan. 1917

Leaves to 40 x 15 cm., oblong, with 6-8 pairs of slightly oblique, remote leaflets, with internodes decreasing from 6 to 3.5 cm., the much smaller lowest pair set c. 5 mm. above the base of the rachis. Leafrets rigid, narrowly oblong-ovate, their length increasing (4459.5 cm.) from the base to the apex of the rachis, but their breadth remaining fairly constant, 2.8-3.2 cm., the terminal leaflet being scarcely larger; bases rounded-sub- truncate; margins 4-8-spinose (1-2 mm.)-serrate (1-2 mm.), distant 8-18 mm. : apical cusp inconspicuous, c. 8 x 6 mm. ; above lustrous yellow-green, with venation scarcely even subreticulate and indktinct ; below slightly lustrous, paler, with only some lateral veins visible. Racemes 10-18 em., fascicled c. 6; bracts of t,he inflorescence 10-15 mm. Pedicels 5-7 mm.; floral bracts 3 mm. Outer sepals 4 x 3 mm., ovate, acuminate. Median sepals 5 x 3 mm., ovate, obtuse. Inner sepals 9 x 4 mm., oblong-obovate. Petals 9 x 3 mm.,

20 JOURN. LINN. SOC. BOTANY-VOL LVlI

Page 306: Berberis and Mahonia : A taxonomic revision

306 L. W. A. AHBENDT narrowly oblong; apex incised with rounded lobee; glands distinct. Stamens produced, retuse. Ovules 4. Style 3 mm.

Yunnan: vicinity of Yunnan-sen, Maire 2003 (Type, K). Not cultivated.

(513) Mahonia duclouxiana Gagnepain, Bull. Soc. Bot. Fr. 55 (4th series, 8), 87, 1908 Leaves with stout rachis, and the lowest pair of leaflets close to the base. LeaJrets

in 5-9 pairs. R m m m 15-27 cm. Pedi& 5-7 mm.; bracts 2-5-4 mm. Style c. 2 mm.

KEY 95A Leaflets broad, 1 : 1.5-2.5, conspicuously reticulate.

Leafleta narrower, 1 : 2-54 , indistinctly veined. 5130. M. duclouxiana var. ducbuxiana

513b. M. d- ’ na var. hiluiw

(513a) Mahonia duclouxiana var. duclouxiana Leaves to 40 cm. long, with 7-9 pairs of leaflets. h . e h ovate, 3-12 x 24.5 cm. ; base

truncate or subcordate ; margins 10-12-subappressed-spinulose-serrulate ; above h e l y but conspicuously whitish reticulate; below paler and very openly reticulate. Racemes 15-20 cm. long. Outer sepals %3 mm., ovate, obtuse. Median s e p k 6 mm., acuminate. Inner sepals 8 mm., lanceolate. Petuls c. 7.5 mm., apex retuse, glands distinct. Stamens slightly produced, subtruncate. Ovules 5. Berries globose, 6mm., excluding style c. 2 mm.

Yunnan: near roads to Lou-kiou-en, 10 Feb. 1905, Ducloux 3055; on the road from Yunnan-sen to Kiu-tsin fou, not far from Ma-long, 20 Mar. 1902, Ducloux 2323 (Type); Gnon-kay ( H o b ) , 24 Mar. 1885, Delavay 2353.

(513b) Mahonia duclouxiana par. hilaica Ahrendt, var.nov., foliis angustioribus, minus nervatis, distinguenda.

Leaves to 40 x 14 om., narrowly obovate, with 5-8 (-9) pairs of separate t.0 remote leaflets, their internodes decreasing &om 7 to 3.5 cm., along the rachis, c. 5 mm. above the base of which is set the small baaal pair measuring 3 x 1.75 cm. Lea$& fairly thin, but rigid; oblique; their length increasing (3-10 cm.) constantly from base to apex; but their breadth increasing (1.5-3.5 cm.) from the base only to the middle, and then decreasing (3-5-2-4 cm.) h m the middle to the apex; thus, towards the base they are ovate, acute, 3.5-5.5 x 1.5-3 cm. ; towards the middle they are narrowly ovate and sub- acuminate, c. 8-5 x 3.5 cm. ; towards the apex they are lanceolate and acuminate, 9-5 x 3 cm., or 10 x 2.4 cm. ; bases rounded; margins 7-9 (-10)-spinose (1-2 rnm.)-serrate (1-2 mm.), distant 7-15 mm.; apex without cusp; above scarcely lustrous with indistinct elevated reticulation ; below scarcely paler sublustrous, with venation loosely elevated. Racemes 20-27 cm. long, fascicled c. 8; bracts of the inflorescence c. 12 x 6 mm. Ovules 4-5. Berries pruinose blue, ellipsoid, 8-9 x 56-65 mm., excluding style c. 2 mm.

W. Yunnan: Shunning, Hih, 8000ft., a common tree, 20-30ft., among forest, fr. 26 June, 1938, Yu 16471 (Type, E). Although many of Yu’s collections are in cultivation, I have not met this among them.

Subsect. Acanthifoliae Ahrendt, subsect.nov., foliolis in6mis ad baaim rhachidis ap- proximatis foliolis ceteris margine plerumque dentibus 3-9 ; apice inconspicue cuspi- datis; pedicellis 1-7 mm., bracteis florum 13.5 mm., duplo brevioribus ad 1.5-plo longioribus; petalis sepalis interioribus semper brevioribus; staminibus plerumque apiculatis; stylo semper conspicuo, plerumque 1-2 mm. longo.

Leaves with basal leaflet close to the base of the rachis. h . e t s in 5-20 pairs; margins with 3-9 (-14) teeth; apex with inconspicuous cusp. Racemes 7-20 cm. long, fascicled

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Berberis and Mahonia 307

2-12. Pedicels 1-7 mm., with their bracts (1-3.5 mm.) from one-half to 1.5 times at3

long as themselves. Petals shorter than inner sepals. Stamens mostly produced, apicu- late; rarely not produced, truncate. Ovules 1 4 . Style (0.G) 1-2 mm.

Kashmir, Punjab, Jaunsar, Tehri, Kumaon, Nepal, Sikkim, Bhutan, S.E. Tibet, Assam, Manipur, Burma.

KEY 96, to Subsect. Acanthifoliae Bracts longer than pedicels.

Leaflets in 5-9 pairs; pedicels 5-4 mm.; bracts 3-5 mm.; outer sepals large, 5 mm.;

Leaflets in 12-20 pairs; pedicels 1-2mm.; bracts 1-5-2 mm.; outer sepals small, stamens produced, apiculate ; ovules 3-4. 514. M . borealis

1.5 mm.; stamens not produced, truncate; ovules 1-2. 515. M . new species Bracts shorter than pedicels.

Pedicels 1-3 mm. ; bracts subequal. Leaflets in 12-20 pairs, narrow (1 : 1-4); racemes 7-13 cm. long.

Leaflets in 8-12 pairs, broad (1 : 1-2.5); racemes 13-20 cm.

Leaflets very narrow (1 : 3--6), with margins with 1 6 1 4 teeth; stamens not pro- duced, truncate. 517. M . magni,fica

Leaflets broader, 1 : 1-2.8 (-4), with margins with 2-6 teeth; stamens apiculate, where known.

Cf. 515. M . new species 516. M . griffithik

Pedicels 3-7 mm. ; bracts only one-half as long as pedicels.

Leaflets broad, 1 : 1-2.8; pedicels 4-7 mm.; bracts 2-3.5 mm. Leaflets at apex overlapping; racemes 14-24 em.; style 1-1-5 mm.

Leaflets at apex separate, or subcontiguous; racemes 7-14 em. ; style 0.75-1 mm. 518. M . acunthifoliu

519. M . jaunsarek 520. M . aristulata Leaflets narrower, 1 : 3-4; pedicels 3-4 mm.; bracts 1.5-2 mm.

(514) Mahonia borealis, Takeda, Notes R. Bot. Gard. Edinb. 6 , 221, Jan. 1917: Chatterjee, Rec. Bot. Surv. India, 16 (no. 2), 41, 1953

Leaves to 40 x 15 cm., oblong-oblanceolate, with (6) 6-9 pairs of usually separate leaflets, though they are contiguous or slightly overlapping by the apex; their inter- nodes decreasing along the rachis from 7 to 3 cm. Leaflets thin, ranging from narrowly oblong-ovate to oblong-oblanceolate ; their size, both length and breadth, increasing continuously from base to apex (6) 7-10 x ( 1 . 6 ) 1 . 7 3 (4) cm. ; the smaller ovate basal pair measure 3 x 1.7 cm., and are set 5-10 mm. up the rachis; bases broadly rounded or truncate; margins 4-8 (-9)-spinose (1-2 mm.)-serrate (2-4 mm.), distant (5-) 7-12 (-20) mm.; apical cusp 13-20 x 2-6 mm., not very conspicuous; above dull grey-green, with indistinct open subelevated reticulation; below with elevated branched venation, and fine close reticulation. Racemes (5-) 8-12 (-20) cm. long, fascicled 4-9, with thick rachis and bracts of the inflorescence measuring 10 x 7 mm. Pedicels thick, 3-4 mm. ; floral bracts 3-5 mm. Outer sepals 5 x 2.5 mm., ovate, acute. Median sepals 7 x 2.5- 3 mm., oblong-ovate, subacute. Inner sepals &8.5 x 3-3-5 mm., oblong-obovate. Petals 5-6 x 3-3-5 mm., oblong; apex acutely emarginate with obtuse lobes; glands distinct. Stamens slightly swollen below the anthers, but scarcely subdentate ; apex produced, apiculate. Ovules 34. Berries 5-8 x 5 mm., globose, to ovoid, pruinose blue, excluding style 1-2 mm.

KEY 96 A

Racemes 5-12 em. long; fruit ovoid, 8 x 5 mm. Racemes 12-20 om. long; fruib globose, 5 mm.

514a. M . borealis var. borealis 514b. M . borealis var. parryi

20-2

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308 L. W. A. AHRENDT

(614~) Mahonia borealis var. borealis Kashmir: Royle, 8.n. (Type, 9). Tehri: 7000 ft., Mussooree, 1892, Stewart 8.n. (K). Kumaon: 7000 ft., Strachey & Winterbottom 1 (K). Punjab: Chamba state, Kainthli reserve, 7000 ft., 6 June 1919, Parker, 8.n. (K):

‘ common near Kagiar, in Kainthli reserve, Kalatop and Dalhousie range, in moist forest undergrowth’; 1844,5000-7000 ft., Edgeworth, s.n. (K).

(514b) Mahonia borealis var. parryi Ahrendt, var.nov., rammis elongatis, fructibus

Assam: Lushai hills, north of Vanlaiphai, 6000 ft., fl. Feb. 1927, 5000 ft., Mar. 1928, Parry 149,149a (Type, 9). ‘Lushai name Pualleng; a yellow dye is made from the sap.’

Chatterjee, I.c., p. 81, dealing with the alkaloid contents of this species and &f. nucni- purensis, remarks on their close chemical similarity which does not support their morpho- logical differences, suggesting that the latter may be a further variety of M. borealis. However, in his taxonomic discrimination he retains them as separate (I.c. p. 41), as I had already done in these pages (originally written in 1942-3, and revised in 1952). Against the weight of chemical a f i t y , there has to be set the considerable weight of such a constant character as the significant style both in Berberis and Mahonk.

(515) Mahonia new species. Leaves to 40 x 10 cm., oblong, with (1%) 15-20 pairs of leaflets, nearly always separate,

but sometimes subcontiguous towards the apex, towards which their internodes decrease from 4 to 1 cm., the small basal pair measuring 1-26 x 1-25, being orbicular and set only 2-5 mm. a bovethe base of the rachis. LeaJlets thick, rigid; apart from the basal pair, their length increases from (2 -5 ) 4-6 cm., passing from the base to the middle, and then decreases from 6 to 4 cm., passing on to the apex; but their breadth constantly decreases from 4 to 1 cm., passing from the base to the apex; thus their shape is suborbicular at the base, lanceolate t o w a d the middle, and ovate towards the apex of the leaves; margins 3-5-spinose (1-1.5 mm.)-dentate (3-8 mm.) &hnt 1.5-2 cm.; apex cusp in- conspicuous; leaflets placed only slightly obliquely, the terminal one measuring 6 x 1-8 cm. ; both sides sublustrous with loose elevated venation, branched, but scarcely subreticulate. Racenzes 7-14 cm. long, fascicled c. 7; bracts of the inflorescence 20 x 9mm. Pedieels 1-2mm.; floral bracts 1.5-2mm. Outer se@ 14xlmm. , ovate. Median sepals 2.5 x 1-5 mm., oblong-elliptic. Inner se@ 46 x 1.5-2 mm., oblong- elliptic. Petals 3.25 x 1.5 mm.; apex alightly emarginate; glands indiatinct. Stamerrp 2-25 mm., scarcely produced, rounded-truncate. Oeyules 1-2. Style 1 mm. or more.

S.E. Tibet: Prov. Monyul, Bhutan frontier, Nyam Jang Chu, between Le and Pangchen, 7500 feet, fl., 2 Nov. 1938, a shrub up to 8 ft., Ludlow, Sherriff & Taylor 7011

This species is remarkable for the variation in size and shape of the leaflets on the same leaf. It differs from M. boredis in the more numerous thickly leathery leaflets. Unfortunately the specimens in the British Museum (Natural History) on which the description is based seem to have been destroyed in an &-raid; hence it is left unnamed.

Garhwal, Landour, 1831, Vicary 42 (K).

globosis.

(516) Mahonia grmthii Takeda, l a , p. 217, 1917; Chatterjee, I.c., p. 37, 1953 Leaves oblong-elliptic with 8-12 (-13) pairs of leaflets, overlapping towards the apex,

contiguous near the middle, subcontiguous at the base; their internodes decreasing from 6 cm. (below) to 4 cm. (middle) to 1.5 cm. (apex); the small basal pair, c. 2 x 1-5 cm., being only 5-10 mm., above the base of the rachia. LeaJlets from broadly to narrowly, ovate, their size (both length and breadth together) increasing from 3-7-5 x 2 - 6 3 cm.,

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Berberis and Mahonia 309

from the base to the middle, then decreasing from 7-5-45 x 3-2 cm., from the middle to the apex ; bases rounded-cuneate or subtruncltte; margins 3-5-spinose (1-1.5 mm.)-ser- rate (1.5-2.5 mm.), distant 1-2 cm.; apical cusp inconspicuous, c. 2 x 1-5 om. ; above sublustrous with c. 5 oblique lateral veins impressed; below dull, scarcely paler with few veins and veinlets. Racemes 12-20cm., fascicled c. 11; bracts of the inflorescence c. 1 cm. Pedicels thick, 2-3 mm.; floral bracts 16-24 mm. Be& ellipsoid, slightly pruinose blue, 9 x 6 mm., excluding style 1.5-2 mm.

Bhutan: fr. Grifith 1739 (Type, K). Not cultivated.

(517) Mahonia magnifica Ahrendt, sp.nov., bracteis florum pedicelh eorum multo brevioribus ; stylis aigdcantibus ; foliolis elongatis, angustis ; staminibus apice truncatis, distinguenda.

Leaves to 60 x 25 cm., with 10-11 pairs of leaflets, the basal pair, 1.5 x 2-1-1.5 cm., set 5-20mm. above the base of the rachis; the internodes of the remainder decreasing from 6 to 1.8 cm. in passing from baae to apex of the leaf. LeaJets lanceolate; their length increasing from 10 to 14 cm., passing from the base to the middle, and then decreasing from 14 to 11 cm., in passing on to the apex; their breadth steadily decreasing from 3.5-3-1-8 cm., in passing from base to middle to apex; bases broadly cuneate to sub- rounded; margins 10-18-spinose (1-1-5 mm.)-serrate (1-3 mm.), distant 4-12 mm.; above dull, indistinctly openly reticulate; below paler, lustrous, with venation more distinct, finely reticulate. Raames 10-20 cm., erect, fascicled c. 12; bracts of the inflorescence 12-15 x 6-8 mm. Pedicels 2 4 (-5) mm., slender; floral bracts 1-2 mm. Outer sepals 2-3 x 2 mm., oblong-ovate. Median sepals 5 x 2.5-3 mm., elliptic. Inner sepals 6-6.5 x 3 mm., oblong-elliptic. Petals 4.5 x 2-26 mm. Stamens 3 . 5 4 mm., scarcely produced, truncate. Style 1-1.5 mm. or longer.

Manipur: Tusom, Khulen, 6500 ft., a small tree, branching repeatedly with brilliant yellow flowers, 19 Nov. 1948, Kingdon-Ward 18325, (Type, BM). Not cultivated.

(518) Mahonia acanthifolia G. Don, Cen. Syst. 1, 118, 1831; Takeda, 1.c. p. 219, 1917; Chatterjee, 1.c. p. 39, 1953

Leaves to 48 x 12 cm., oblong-lanceolate, with 8-11 pairs of obliquely set leaflets, over- lapping for the apical half, contiguous by the middle, separate below, the lowest pair (1-2 x 1-2 cm.) set 15 mm. above the base. Leajlets fairly thick, rigid; above dull light green, becoming darker, slightly lustrous, with age, slightly veined ; broadly to narrowly oblong-ovate; their length increasing from 4.5 to 6.5 cm., passing from base to middle, then slightly decreasing from 6.5 to 5.5 cm., on continuing to the apex; their breadth continuously decreasing from 3.5 to 2 cm.; the terminal leaflet being larger, 6.5-7.5 x 3 - 5 4 cm., with a petiole c. 1.5 cm.; bases of lateral leaflets truncate; margins 2 4 - spinose (1-2 mm.)-serrate (4-6 mm.), distant 1.5-3 cm. Racemes 15-23 cm. long, fascicled 3 4 ; rachis 2.5-4 mm. thick; bracts of the inflorescence 2-5-3 x 1.2-1.5 cm. Pedicels 4-6 mm. ; floral bracts 2-3.5 mm. Outer sepals 1.5 x 1.5 mm., broadly ovate. Median sepals 4-5 x 3 mm., ovate, obtuse. Inner sepals 7 x 3 mm., oblong-elliptic. Petals 6 x 3 mm., oblong-elliptic, narrowly incised; glands distinct. Stamens 3.5 mm., apex distinctly conical-apiculate. Ovules 3 4 . Berries oblong, pruinose blue, 10-12 x 5-7 mm., excluding style 1-1.5 mm.

Nepal: 1821, Wallich 1480 (Type, K); Pundoah, 1820, Wallich 1480C (K). Sikkim: Darjeeling, Thio gapsek kum, 7000-8000 ft., fl. Hooker 41 (K), ‘fruit ripe in

May’; Oct. 1904, 6000 ft., Drummond 14814 (K), a specimen with leaflets thinner and more dentate; Darjeeling, fl. Oct. 1874, Gamble 1032 (K), and fr. Mar. 1877, Gamble 403SB (K); fl. 1880, Gamble 8561 (K).

Assam: Naga Hills, Japvo, 8000 ft., on precipitous densely forested slopes, not rare, 24 Nov. 1949, Kingdon-Ward 19088 (BM).

Page 310: Berberis and Mahonia : A taxonomic revision

310 L. W. A. AEIRENDT Kumaon: Blinkworth, 8.n. Cultivated: L.A. 717, without flowers or fruit, 1948; also seen, a t Bishopsteignton,

South Devon, Aug. 1951, a tree some 20 ft. high and 25 ft. through.

(519) Mahonia jaunsarensis Ahrendt, sp.nov., aff. H. acanth&,folh a qua folio&, a b a s h versus apicem, longitudine semper increecente, wd latitudine satis con- stante; inter se versus apicem foliorum non imbricatis sed plusminusve separatk, recedit

Leaves to 28 x 13 cm., obovate, with 5-7 pairs of leafleta mostly separate, but some- times subcontiguous towards the apex of the leaf; their internodes gradually decreasing from 5 to 3.5 cm. on passing from base to apex of rachis which is 2 mm. thick; the small basal pair measuring 12 x 8 mm., and set only 5-7 mm. above the base. Leaflets broadly ovate to broadly lanceolate, their length increasing from 3.5 to 7 cm., from base to apex, while their breadth remains constant at 3.5 cm.; bases broadly cuneate, rounded, or subtruncate; margins 3-5-spinose (15-3 mm.)-dentate (3-6 mm.), distant 1-2 cm. ; apical cusp inconspicuous; terminal leaflet 9-2 x 4 cm., with petiole 1-5-2 cm.; above slightly lustrous; both sides subelevated subreticulate. Racema 7-14 cm. long, fascicled 5; bracts of the inflomcence c. 1-5 cm. long. P e d S 5-7 mm.; floral bracts 2-3.5 mm. Pmit (immature), pruinose blue, ovoid, 8 x 4 mm., excluding style 0.7-1 mm. (possibly longer a t maturity).

Jaunsar: Chakrata, 7000 ft., fl. 9 Apr. 1897, Rich 3 (B 33) (Type, K); same locality, 6OOO ft., Jadi forest, Nand 73 (0), ‘Jaunsar name Khoru, a large shrub wmmon in moist places, and under oak forests about Chakrata; Chakrata, Marshall, s.n. (0). Not cultivated.

(520) Mahonia aristulata Ahrendt, sp.nov., foliolis in ambitu, et apice conspicue

Leaves with 11 pairs of ledleh, their internodes decreasing from 5.5 to 1.2 cm., from base to apex, the bawl pair measuring 12-20 x 6-7 mm., and set c. 1 cm. above base. Leaflets oblong-lanceolate or lanceolate; from the base to the middle of leaf, their length increases from 5.5 to 7 cm., while their breadth remains constant, 1.5 cm.; from the middle to the apex their sue (both length and breadth together) decreases, from 7 x 1-5 to 4 x 0-9 cm.; bases broadly rounded or subtruncate; margins 3-6-spinose (2 mm.)-serrate (2 mm.) ; fascicled c. 2; bracts of the inflorescence 10-12 mm. Pedi& slender, 3-4 mm. ; floral bracts 1.5-2 mm. Otherwise 11nknown.

Upper Burma: Myitkyina, 6 milea from Kangrang, Naw Mu Pa, fl. 15 Dec. 1938, KingdomWard 17489 (Type, K): ‘shrub 5 ft. high with red flowers.’ Not cultivated.

aristatis, inter alia distincta

Subsect. Napaulenses Ahrendt, subsect.nov., foliis plerumque jugis 3-9 foliolorum ; bracteis florum pedicellis e o m brevioribus ad paululum longioribus; staminibus productis, apice apiculatis, interdum obtuse subapiculatis ; stylo brevissimo vel nullo

Leajleta in 3-9 (-20) pairs; margins 3-11-dentate. Pedicth 2-7 (-10) mm.; floral bracts 2-6 mm. Pmphylls absent. Xtapnens mostly apiculate, occasionally obtusely subapiculate. Ovules 14. Style insigdicant, absent or extremely short.

Nepal, Assam, Manipur, N.W. and W. Yunnaq.

&Y 97, to Subsect. Napaulenses Leaflets mostly narrow, 1 : 5-6, in 12-20 pairs. Leaflets broader, 1 : 1-3 (-4-5), in 3-9 pairs.

521. M . l m r i i f o l i a

522. M . simneii Ledets fairly narrow, 1 : (2 .5 ) 3-4; petals 8~ long SB inner sepals.

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Berberis and Mahonia 311

Leaflets 1 : 1-2 (-3), much broader; petals shorter than inner sepals. Outer sepals large, 3.5-4 mm.; inner sepals less than, or at most equal to, twice

Leaflets broad, 1 : 1.2-2; racemes 4-8 cm.; pedicels 3-4 mm.; bracts 2-3 mm.;

Leaflets narrower, 1 : 2-3; racemes 1 6 2 0 cm.; pedicels 5-8 (-10) mm.; bracts

Outer sepals small, 2-2.5 mm.; inner sepals 2.5-3 times as long as the outer sepals.

the length of the outer sepals.

ovules 1-2. 523. M . nzani.pzlrenais

4-6 mm.; ovules 4-5. 524. M . napaulensia

Style very short. Ovules 2 ; leaflets contiguous to overlapping; pcxhmls 3.4 x bracts.

Ovules 3-5; leaflets remote to separate; pedicels subequal to bracts. 525. M . pycnophylla

Bracts 5-6 mm.; median sepals 3 mm.; inner sepals 6 mm.; stamens apiculate. 526. M.$a&

Bracts 2-3 mm.; median sepals 5.5 mm.; inner sepals 8 mm.; stamens obtusely subapiculate . 527. M . hancockiana;

Style absent. 528. M . salweenenk

(521) Mahonia lomariifolia Takeda, Notes R. B d . Gard. Edinb. 6 , 231, Jan. 1917; M . alexandri Schneid., Bot. Gaz. 43, 519, June 1917

Leaves to 60 x 15 cm., oblong-elliptic, with (1%) 1 6 2 0 pairs of leaflets which are only slightly oblique, contiguous above and separate towards the lower part of the rachis; their internodes decreasing from 6 to 1-2 cm., in passing up the rachis, the small sub- orbicular basal pair measuring c. 1 x 1 cm., and set 5-10 mm. above the base. LeaJlets rigid, not very thick, varying along the rachis from oblong-ovate to oblong-lanceolate ; lengths decreasing from 7 cm. a t the middle of the leaf towards both ends, being 4 cm. at the apex and 2 cm. below (excluding the basal pair) ; but their breadth decreasing con- tinuously in passing up the rachis from 15 to 7 mm. ; terminal leaflet 5 x 1 cm. ; margins 3-7-spinose (1-16 mm.)-serrate (1.5-3 mm.); apical cusp to 15 x 8 mm., below, or 25 x M m m . , above where it is fairly conspicuous; above dull light green; very slightly veined or enervate. Racemes 9-15 cm., fascicled, c . 7; bracts c . 1 mm. Pedicels slender 3-5 mm. ; bracts 2 4 mm. Outer sepals 3 . 5 4 x 1.5 mm., ovate, obtuse. Median sepals 8 x 3 mm., elliptic-obovate. Inner sepals 9 x 3-25 mm., elliptic. Petals 6.5 x 3 mm., oblong-obovate, narrowly incised. Stamens c . 3 mm., slightly produced, subapiculate. Ovules 3-5. Berries ovoid, 6-7 x 4-5 mm., pruinose blue. Style very short.

Yunnan: Milo district, in mountain woods, fl. Henry 10309 (K); Mekong, and side valleys, 6 ft. high a t 8000-12,000 ft., Sept. 1904, Forrest 141 (K); a shrub 6 1 2 ft. high in shady, moist situations, and gullies on hills, to the north and east of Tengyueh, with bright yellow non-fragrant flowers, Nov. 1912, Forrest 9244 (K) ; Atuntze, rare, 13 ft. high among thickets, 9500ft., 14 Sept. 1937, Yu 10212 (E); Mekong valley, Aiwa, common, 1-2 ft. high, evergreen shrub under mixed forests, a t 8300 ft., 7 Dec. 1938, Yu 22881 (E) (the last is a specimen with longer style).

S. Szechuan: between the towns Yen Yan Hsien and Yung-ming between villages Wo-lo-ho and Cho so, mountain side 8700 ft., 15 June 1914, Schneid. 1588 (Type of alemndri, Am. Arb.).

Cultivated : by Major Johnstone, Chipping Campden, Gloucestershire, from seed col- lected on Forrest’s last journey, in Yunnan (see Bot. Mag. t. 9364). This desirable, but tender, Mahonia is beginning to be fairly widely cultivated, particularly on account of its very graceful foliage. I saw a bush about 10 ft, high, in Aug. 1951, a t Lindridge, near Bishopsteignton, South Devon.

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312 L. W. A. AHBENDT

(522) Mahonia shonsii Takeda, Notes R. Bot. aard, Edinb. 6,222,1917 ; Chatterjee,

Leaves with 6-9 pairs of contiguous to overlapping leaflets, their internodes decreasing from 5 to 3 cm., up the rachis, and the baa1 pair measuring 2-2-5 x 1-5-1-75 cm. set c. 5 mm. above the base. k f l e t s narrowly ovate to lanwlate; their length being 11 cm. at the middle of the leaf, but decreasing to 7 om. at either end; their breadth below the middle being fairly wmtant, 2.5-3 cm., but above the middle slightly decreasing to 2-2-5 cm.; the terminal leaflet being equal to the others by the apex; bases broadly rounded; margins 7-ll-spinoae (1 mm.)-serrate (1-2 mm.), distant 5-12 mm.; apex cusp inconspicuous, 10-20 x 5-10 mm. ; both sides yellow-green, sublustrous with con- spicuous openly reticulate venation. Racemes 10-15 cm., fascicled, c. 8, with rather thick rachis. Pedicels 2-3 mm.; floral bracts 1-5-2.5 mm. Outer sepak 2 x 1-5 mm., ovate, acute. Median sepals 5 x 2 mm., oblong-elliptic, obtuse. Inner sepals 7 x 2 mm., oblong- obovate. Petals 5 x 2 mm.; glands minute. ovules 3. Styk very short.

Rec. Bot. Sum. India, 16, No. 2, 39, 1953

Assam: Khasia Hills, Nov. 1850, Simons, s.n. (Type, K). Not cultivated.

(523) Mahonia manipurensis Takeda, 1.c. p. 222, 1917; Chatterjee, 1.c. p. 41, 1953 Leaves to 30-40 x 8-10 cm., oblong; with 7-9 pairs of contiguous, overlapping leaflets,

their internodes decreasing from (5) 36-16 cm., in passing up the rachis; the basal pair measuring 1-5-2 x 1-1.3 cm., and situated c. 6-12 mm. above the base. Lea$ets thick, rigid, ovate; all about the same length, 4-5-56 cm., but with their breadth slightly decreasing from 3.5 to 2-5 cm. ; bases truncate-subcordate; margins 2-4-spinose (2 mm.)- dentate (3-5 mm.), distant 1.5-2-5 cm. ; apical cusp inconspicuous, 10-14 x 8-10 mm. ; terminal leaflet similar to the others near the apex, but larger 5-5.5 x 26-3 cm. ; dull above with indistinct subreticulate venation ; below paler with branched elevated venation. Racemes 4-8 cm., fascicled 3-5; bracts of the inflorescence 10-15 mm. Pedicek slender, 3-4 mm. ; floral bracts 2.5-3 mm. Outer sepak 4 x 2 mm., ovate, acute. Median sepals 7 x 4 mm., oblong-ovate, obtuse. Inner sepals 8-9 x 4-5 mm., oblong-obovate. Petals 7 x 3.5 mm. Stumm produced, obtusely subapiculate. Ovules 1-2. Xtyle very short.

Manipur: KaRsoorie range, on the frontier of Burma, 8000ft., fl. Jan. 1882, Watt 5956; Sirvhifurar ranges, north-east, 8000 ft., fl. Apr. 1882, Watt 6472 (Type, K); Saura pass, range on the frontier of Burma, 40 miles nearly due north of Ukhrul, above Tusom, Khulem, 7000 ft., fl. 20 Nov. 1948, where Kingdon-Ward saw this, in tree form, with the trunk 30 in. in girth. Not cultivated.

(524) Mahonia napadensis DC, Syst. 2,21,1821; Takeda, 1.c. p. 216,1917; Chatterjee,

Leaves to 40 x 12 cm., oblong-oblanceolate, with (3-)4-7 oblique pairs of contiguous or slightly overlapping leaflets; their basal pair measuring only 2 x 1-5 cm., and being situated (0 .5) 1-2 cm. above the base; the internodes of the remainder decreasing up the rachis from 5.5 to 3.5 cm. Leaflets thick, rigid, ovate; their length increasing from 5.5 to 9.5 cm., fkom the base to the middle of the rachip, but then decreasing slightly from 9.5 to 8 cm., in passing to the apex; their bredth being fairly constant, 3-4 cm.; the terminal leaflet being hceolate, 9.5 x 4.5 cm. ; bases broadly rounded or subtruncate; margins 4-10-spinose (1-2 mm.)--dentate (3-8 mm.), distant 7-18 (-25) mm.; apex acuminate, or with sometimes inconspicuous cusp, 15 (-25) x 10 mm. ; above slightly lustrous; below dull ; venation clearly branched above, subreticulate below. raceme^ 14-20 cm. long, pendulous, fascicled 6-7; bracts of the inflorescence c. 2 cm. Pedhds slender, 5-7 (-10) mm. ; floral bracts 4-6 mm. Outer sepak 3-4 x 2 mm. Median sepals 5-7 x 3-3.5 mm., both these series being oblong-ovate, to oblong-elliptic. Inner sepals 7-9 x 3-54 mm. Petds 4-54 x 3.5-5-5 mm. ; apex emarginate with rounded lobes; glands

1.c. p. 36, 1953

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Berberis and Mahonia 313

distinct. Sturnens with apex distinctly conical apiculate. Ovules 4-5. Berries 9 x 6 mm., excluding style c. 0.4 mm., heavily pruinose blue.

Nepal: Naranihetty, fl. 15 Nov. 1802, Hamilton, s.n. (Isotype, K); Jan. 1821, Wallich 1480.

W. Nepal: Lekh Rangchi, in dense wet oak and rhododendron forest, shrub 3 4 ft. tall a t 7500 ft., 3 Apr. 1952, Polunin, Sykes & Williams 3832 (BM).

Cultivated: L.A. 719; fl. 25 Mar. 1949. Seen also, Aug. 1951, 6 ft. tall, a t Lindridge, South Devon.

(525) Mahonia pycnophylla (Fedde) Takeda, Notes R. Bot. Gard. Edinb. 6,218, Jan. 1917; Chatterjee, 1.c. p. 40, 1953; syn. M . nepaknsis var. pycnophylla Fedde, Bot. Jb. 31, 124, 1901

Leaves to 31 x 16 cm., oblong, with 5-6 pairs of mostly contiguous leaflets; the basal pair measuring 3.5-4 x 3 cm., mt 5-10 mm. above the base; the internodes of the re- mainder decreasing from 8 to 3-7 cm. passing up the rachis. Leaflets thick, rigid; apart from the basal pair, ovate and fairly equal in size (9-11 x 4-5-5 cm.) ; bases truncate- subcordate; margins 3-5-spinose (2 mm.)-serrate (1-5-3 mm.) ; apical cusp inconspicuous, 2-5 x 1.5 cm. ; above sublustrous with indistinct subelevated reticulation; below scarcely paler, dull, with venation branched and scarcely even subreticulate. Racemes 12-22 cm. fascicled c. 10; bracts of the idorescence c. 12 x 7 mm. Pedicels slender, 5-7 mm. ; bracts 1-5-2 mm. Outer sepals 2-2.5 x 2 mm., ovate-obtuse. Median sepals 5-6 x 3 mm., ovate. Inner sepals 7 x 3-5 mm., elliptic. Petals 5-5.5 x 2-5-3 mm., narrowly obovate, with distinct glands. Stamens 3.5 mm., conical apiculate. Ovules 2. Style very short.

Assam: Khasia Hills, Tserra, 400M500 ft., Grifilth 450 (Type, K). Not cultivated.

(526) Mahonia flavida Schneid., in Sargent, PI. Wils. 1, 382,1913; Takeda, 1.c. p. 227 Shrub 6-8 ft. Leaves to 70 x 25 cm., oblong-elliptic; with 7-8 pairs of obliquely placed,

separate to remote leaflets, the much smaller basal pair being set c. 5 mm. above the base; the internodes between the others being fairly constant along the rachis a t 5-6.5 cm. Leaflets fairly thick, their size (6-12 x 3 . 5 4 cm.) gradually increasing from base to apex; the terminal leaflet being much larger, to 17 x 4.5 cm., with a petiole 3 cm. long; bases rounded; margins 6-9-spinoae (1-2 mm.)-serrate (1-2 mm.), distant 8-15 mm. (varietally with 0-2 teeth); apex with a broad inconspicuous cusp, 2.5 x 1 cm. ; both sides yellow green, slightly lustrous, with indistinct elevated reticulation ; ovate to elliptic-lanceolate. Race- 11-22 cm. long, fascicled c. 20; bracts of the inflorescence 3 4 cm. long. Pedicels 4-5 mm.; floral bracts 3-6 mm. Outer sepals 2 x 2 mm., ovate, acute. Median sepals 3 x 2 mm., oblong-ovate. Inner sepals 6 x 3 mm., oblong-obovate. Petals 5.5 x 2.25 mm., emarginate with rounded lobes. Stamens 3.75 mm., dilated below the anthers; apex apiculate. Ovules (3-) 4-5. Style very short.

K E Y 97A Leaves with 6-9 teeth t,o each margin. Loaves entire or with 1-2 teeth to each margin.

(526a) Mahonia flavida var. flavida

Not cultivated.

526a. M . fEavida vm. jta& 526 b. M . &vida var. integrifoliola

Yunnan: mountains to the south-east of Mengtze, 5000 ft., Henry 10180 (Type, K).

(526 b) Mahonia flavida var. integrifoliola Handel-Mazzetti, Symbolae S i n i w , 7, 332, 1931

Yunnan: Hsiao-Magai, near Yunnan-fu, 1914, Schneider 311 (Type).

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314 L. W. A. ~ E N D T

(527) Mahonia hancockiana Takeda, Notes R. Bot. Gbd. Edinb. 6, 231, Jan. 1917

Leaves to 30 x 12 cm., oblong-oblanceolate, with 4-6 pairs of obliquely placed remote leaflets, the much smaller b a d pair being 2-3 cm. above the base of the rachis; remaining internodes decwing upwards, c. 7 ,4 , 3, 2.5, 2 cm. Leaflets fairly thin and flexible, the small basal pair measuring 26-3 x 1 - 5 2 cm.; the others, from base to apex, decreasing in size (both length and breadth together) from 6-5 x 2-2-1-7 cm. ; terminal leaflet larger, oblong-ovate; bases broadly cuneate or rounded; margins 5-6-spinose (1-1.5 mm.)-serrate (1-1-5 mm.), distant 4-11 mm.; apex cuspidate-acuminate, the in- conspicuous cusp measuring 18 x 8 mm. ; above dull grey-green, subenervate; below much paler with branched veins and veinlets subelevated. Racemes 12-16 cm., fascicled c. 4. Pedicels 4-5 mm. ; bracts 2-3 mm. Outer sepah 2.5 x 2 mm., ovate, obtuse. Median sepak 5.5 x 3 mm., ovate-elliptic. Inner sepals 7.5 x 3.5 mm., obovate. Petals 6 x 2.75 mm., elliptic-obovate; apex acutely emarginate with rounded lobes; glands dis- tinct. Starnens 4.5 mm., apex produced, obtusely subapiculate. Ovules 3-5. Style short, thick.

Yunnan: Mengtze, in forest, 9000-9500ft., fl. Feb. 1894, Hancock 151 (Type, K). ‘Flowers rich yellow.’ Not cultivated.

(528) Mahonia salweenensis Ahrendt, sp-nov., foliolis contiguis vel imbricatis, in ambitu versus basim rhachidum ovatis sed vemw apicem lanceolatis, semper longitudine increscentibus sed latitudine decrescentibus ; bracteis florum pedi- cellis eorum subaequilongis; petalis sepalis interioribus brevioribus; staminibus apiculatis; stylo nullo, distinguenda.

Leaves to 28 x 12 cm., narrowly obovate with 6-7 pairs of continguous or overlapping leaflets, of which the much smaller baml pair are set only c. 5 mm. above the base, the remaining internodes deoreaaing upwards from 5 to 2.5 cm. LeaJEets ovate at base to lanceolate a t apex; their length always increasing upwards from 4.5 to 8 cm.; their breadth always slightly decreasing upwards from 4 to 3-5 cm. ; bases truncate ; margins 3-4-spinose (1-2 mm.)-dentate (3-6 mm.), distant 1 - 5 3 cm.; both sides sublustrous; venation indistinct above, open below. Racema 17-21 om. long, fascicled c. 8; bracts of the inflorescence c. 3 x 1 cm. P e d s 3-5 mm.; floral bracte 2 - 5 4 mm. Outer sepals 2.5 x 1 mm. Median sepds 3-5 x 2-25 mm. Inner s e p k 6 x 3 mm., all oblongelliptic. P e W similar, 5 x 2.5 mm. ; apex slightly emarginate; glands distinct, oblong, separate, non-marginal, 0.7 x 0.3 mm. S h w 3.5 mm., apex distinctly apiculate. Ovules 2-3. Style absent.

N.W. Yunnan: Salween valley, Sekai, 5500 ft., fl. 17 Nov. 1938, Yu 22999 (Type, E). ‘Casual, 2-4 ft. high, by margin of forest.’ Not cultivated.

Subsect. Japonicae Ahrendt, subsect.nov., foliia jugis 3-9 foliolorum; foliolis subtus pallidius flavo-viridibus ; bracteis florum pedicellis eorum fere semper brevioribus, raro vix longioribus ; staminibus plerumque haud productis, truncatis, interdum leviter productis apice obtusis, non apiculatis; infra antheras edentatis ; prophyllis nullis (M. m i r e i solum excepto); stylo nullo vel brevissimo, ad 0.3 mm. longo.

h j l e t s in 3-9 (-15) pairs, paler yellow green below. Pedi& 2-6 (-10) mm. long; floral bracts 1-6 (-9) mm. long, mostly 0-5-0-9 times as long as their pedicels, occasionally slightly longer. Stanzens mostly not produced, truncate; occasionally produced, obtuse; always dentate. Prophylls absent, except in M. ntairei. Ovules 1-7. Style absent or very short, up to 0-3 mm. long.

S.E. Tibet (Prov. Pome, and Pachakshiri district), Manipur, Yunnan, Szechuan, Hupeh, Kweichou, Kwangtung, Chekiang.

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Berberis and Mahonia 315

-Y 98, to Subsect. Japonicae Ledets with 2-16 more or less coarse, teeth to each margin.

Leaflets in %5 pairs; fruit 13-15 x 10-12 mm., estylose. Leaflets in 5-13 pairs; fruit, where known, 6-10 x 4-6 mm., mostly very shortly stylose.

529. M . aheridaniam

Ovules 1-2. Ledets narrow, 1 : 5-8. 530. M . confuaa Leaflets broader, 1 : 1-2-3 (-4.5).

Ledets petiolulate, in 10-13 pairs. 531. M . bodinieri Leaflets sessile, in 5-9 pairs.

Leaflets with 3-16 marginal teeth, mostly dull, sometimes lustrous above. Leaflets with 12-16 marginal teeth; bracts as long as, or slightly longer

than, pedicels; outer sepals lanceolate, 4 x 1 nun. 532. M . pachakahirensis

Leaflets with 3-6 (-9) marginal teeth; bracts shorter than pedicels; outer sepals ovate, broad (1 : 1.3-1.5).

Ovules 2. Outer sepals large, 4 x 3 mm.; leaflets in 4-5 pairs, slightly lustrous.

Outer sepals small, 2 x 1.3 mm.; leaflets in 7-8 pairs, dull. 533. M . roxburghii

Racemes 1-3 cm. ; inner sepals 6 mm. ; stamens produced, obtuse. 534. M . pornensis

Racemes 8-12 cm.; inner sepals, 4.5 mm.; stamens not produced, truncate. 535. M . fordii

Ovules solit,ary. 536. M . schochii Leaflets with only 2 coarse teeth to each margin; very lustrous. 537. M . nitem

Ovules (%) 4-7. Leaves with 5-9 pairs of leaflets.

Prophylls present ; leaflets very lustrous, reticulate. 538. M . mairei Prophylls absent ; leaflets dull, indistinctly veined.

Ovules (5-) 6-7. 539. M . fargeaii Ovules (3-) 4-5.

Leaflets 1 : 1-2.5; terminal leaflet broad; racemes 5-10 cm., erect; pedicels 4-6 mm. ; bracts 2-3 mm. ; median sepals 6 mm. long.

Leaflets 1 : 2-3.5; terminal leaflet narrow; racemes 1CL25 cm. pendulous; pedicels 6-10 mm. ; bracts 4-7 mm. ; media sepals 3.5 mm. long.

540. M . bealei

541. M . japonica Leaves with 12-20 pairs of leaflets. Cf. 521. M . lomariifolia

542. M . kpkdonta Leaflets with 30-50 h e teeth to each margin.

(529) Mahonia sheridaniana Schneid. in Sargent, PI. Wih, 1, 384, 1913; Takeda,

Leaves to 30 x 10 cm., oblong-oblanceolate, with 3-5 pairs of obliquely placed separate leaflets; the small basal pair, 1-5-2 x 1-15 cm., set about 5 mm. above the base; the remaining internodes decreasing from 5 to 3 cm. up the rachis. Leajets thick, rigid, ovate; length increasing up the rachis from 4 to 7-5 cm., while the breadth remains fairly constant a t 2-3 cm. ; terminal leaflet much larger, 10 x 3-35 cm., with petiole 1-5-2-5 cm. ; bases broadly rounded, or subtruncate; margins 3-4-spinose-(l-2 mm.) serrate ( 3 4 mm.), distant 8-20 mm.; apical cusp inconspicuous, 14-20 x 6-9 mm. ; above dull, indistinctly veined ; below with venation little branched and slightly elevated. Racemes 5-6 cm. long, fascicled c . 3; bracts of the inflorescence c. 1 cm. Pedicels c . 4 mm. ; bracts c. 2 mm. Outer sepals 1.5 x 1 mm., oblong-ovate. Median sepak 3.5 x 2.5 mm., ovate. Inner sepak 6 x 4 mm., elliptic. Petals 6 x 3 mm. Stamm 3.5 mm., not pro- duced, truncate. Berries ellipsoid, pruinose blue, estylose, 13-15 x 10-12 mm.

Hupeh: Changyung, woods, 8000 ft., fl., fr. 1SApr. 1900,Wilson (Veitch) 426 (Type, K). ‘Shrub 4 ft. high.’ Not cultivated.

Notes R. Bot. Card. Edinb. 6, 237, 1917

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316 L. W. A. AERENDT

(530) Mahonia confusa Sprague, Kew BUU. 1912, p. 339; 1914, p. 232; Takeda, N d e s R. Bot. Gard. Edinb. 6, 234, 1917; syn. bI. zenutnii Schneid., in Sargent, P1. Wils. 1,378,1913

Shrub 3 4 ft. Leaves to 45 x 12 cm., oblong-oblanceohte, with (6) 6-8 pairs of very obliquely placed leaflets, mostly separate, sometimes subcontiguous ; the basal pair similar to the others, 2.5-5 x 04-1-2 cm., and set 5-15 (-20) 111111. above the base of the rachis; the remaining internodes decreasing from 5 to 3 cm. Leaflets fairly thin, nar- rowly ovate to elliptic-lanceohte; bases cuneate; cusp inconspicuous, 25 x 5 mm.; above dull green. Outer se@ 2.5 x 1.5 mm., ovate, acute. Median se@ 3-9 x 2.5 mm., elliptic. Inner sepals 4.1 x 2.5 mm., elliptic. Petals 3.75 x 1.6 mm., apex very acutely emarginate, with rounded lobes. Sfarnena 2*75mm., produced, truncate. OvUEes 2. Berries obovoid, pruinose blue. Style indistinct, or absent.

KEY 988, Terminal l d e t petiolste, with petiole 2-2-5 om. long. 530a. M . confusa vsr. confusa Terminal l d e t sessile. 530b. M. w n f w vsr. boumei

( 5 3 0 ~ ) Mahonia confusa var. confusa Leaflets with length increasing from b w to apex from 7 to 10 cm., but with their

breadths fairly constant a t 1-5-1.8 (-2) cm. ; terminal leaflet scarcelylarger, 9-11 x 2-3 cm., long petiohte; margins 3-5-spinose (1-2 mm.)-serrate (1-2 mm.), distant 9-18 mm. ; with inconspicuous lateral veins ; below dull, with venation very open and elevated. Racenzee 7-12 cm., fascicled 6 7 ; bracts of the inflorescence 1-15 cm. Pedicel-s 26-36 mm. ; floral bracts 2-2.5 mm. Fruit 7 x 5 mm.

Hupeh: Ichang and immediate neighbourhood, Feb. 1887, fl. Henry 3117 (Type, K); fr. Oct. 1887, Henry 3351A (K); Apr. 1901, fl. Wilson (Veitch) 268 (K).

Szechuan: Mt. Omei, fr. Nov. 1904, Wilson (Veitch) 3143 (K).

(530b) Mahonia confusa var. bournei Ahrendt, var.nov., foliolo terminali sessili,

Lea$& from the base to the apex increasing from 25 x 5 mm. (the smaller basal pair) to 7 om. x 9-12 mm.; margins 3-5-spinose (06-2 mm.)-serrate (0.51 mm.), distant 7-14 mm. ; venation slightly branched, and slightly elevated, both sides. Racemes 5-8 cm., fascicled 3-5. Pedicels 3 mm. ; bracts 2 mm. Fruit 6 x 4 mm.

Hupeh : Chung-chung, Mao-chua-tbih, Jan. 1886, Bourne, 8.n. (Type, K) ; Ichang, Nanto and mountains to northward, Feb. 1887, Henry 2689 (K). Neither variety is in cultivation.

There probably also belongs near M . eonfusa, Berberis ganpinensis GveilI6, Bull. SOC., Agric., Sci. et Arts de la Sarthe, 59,317,1904; Fedde Repert. Sp. Nov. 6 , 372, 1909. This is also referred to by IRveill6 in a footnote as (530c) &z?mnia ganpinensis (Uvl.) Fedde, and described as having slightly veined narrowly lanceolate leaflets, short crowded racemes, elongated fruit with subnil style. Kweichou, by Gan-pin, rare, Oct. 1897, Martin & Bodinier 1929 (Type).

distinct&

(531) Mahonia bodinieri Gagnepain, B d . SOC. Bot. Fr. 55 (4th ser., 8), 85, 1908 Leaves with stout r a c k and 10-13 pairs of leaflets, the lowest very close to the base.

Leufrets with basal pair suborbicular, and shape varying to lanceolate at the apex; in- cluding the basal pair, along the r a c k the length increases from 3 to 11 cm., but the breadth slightly decreases from 4 to 3 cm.; margins on the broad lower leaflets with coarse teeth (5-10 mm. deep), on the ovate middle leaflets with smaller teeth (3-5 mm.

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Berberis and Mahonia 317

deep), and on the leaflets by the apex with fewer fine teeth (1-2 mm.) deep ; bases petio- lulate, truncate-cuneate ; venation openly reticulate. Racemes 12-25 cm. Pedicels very slender, with floral bracts about half as long as themselves. Outer sepals small, ovate, acute. Median and inner sepals about twice the size of the outer. Petals subequal to the inner sepals, emarginate and with indistinct glands. Stamens with apex truncate but tridenticulate. Ovules 2.

Kweichou: Gan-pin et Mount du Colloge a Kouy-yang, fl. 19 July 1898, Bodinier 2465 (Type). Not in cultivation.

(532) Mahonia pachakshirensis Ahrendt, sp.nov., foliis foliolorum jugis 5-6, satis tenuiter coriaceis ; bracteis florum pedicellis eorum equilongis ; sepalis exterioribus a anguste lanceolatis ; staminibus truncatis ; ovulis 2 ; stylo brevissimo, distinguenda.

Leaves to 36 x 9 em., narrowly oblong, or oblong-elliptic; with 5-6 pairs of very obliquely placed separate leaflets; the very small basal pair inserted 10-15 mm. up the rachis; the remaining internodes decreasing from 5 to 3 cm. Leaflets fairly thin, apart from the basal pair, all fairly equal c. 9 x 3.4 cm.; bases symmetrical, broadly cuneate; margins 12-16 spinose (2 mm.)-serrate (1.5 mm.), distant 2.5-5 mm.; above lustrous, subenervate, or with venation indistinct ; below dull with openly branched elevated venation, and fine indistinct reticulation. Racemes 7-13 cm., fascicled 3-4; bracts of the inflorescence ovate, 12 x 6 mm. Pedicels slender (P) 5-9 mm. ; floral bracts 6-9 mm. Outer sepah 4 x 1 mm., lanceolate, acuminate. Median sepals 46-4-75 x 2-26 mm., narrowly obovate. Inner sepals 4.5-5 (-5.5) x 3-3-25 mm., elliptic. Peta2s 4.5 x 2 mm., narrowly elliptic ; apex acutely emarginate, with acute lobes ; glands indistinct. Stamene slightly produced, rounded-truncate. Ovules 2. Style short.

S.E. Tibet: Pachakshiri district, Lalung, 8000-9000ft., fl. 10 May 1938, Ludlow, Sherriff & Taylor 3727 (Type, BM). ‘Shrub 3-4 ft., fairly common on banks in dense wet mixed forest ; leaves shiny; flowers primrose yellow, slightly fragrant.’ Not cultivated.

(533) Mahonia roxburghii (DC.) Takeda, Notes R. Bot. Card. Edinb. 6,218,1917; Chatterjee, 1.c. p. 37, 1953; syn. M . napaulemis var. roxburghii DC, Prodr. 1, 109, 1824; syn. Berberis pin& Roxburgh, PI. Indim, 2, 184, 1832, non Lag.

Leu$& in 4-5 pairs, the small lowest pair, 6-8 x 3 4 mm., set 1-2 cm. above the base; remainder fairly equal, 6-8 x 3 4 cm., but with the terminal leaflet larger; bases broadly cuneate to rounded; margins with (2-) 3-5 (-6) teeth; above sublustrous. Racemes 8-12 om., with slender rachis. Pediceb slender, P 5 mm.; floral bracts 2-2-5 mm. Outer sepals 4 x 3 mm., ovate. Median sepals 7 x 4 mm., oblong-ovate. Inner sepals 8 x 3 mm., elliptic. Petals 7 x 3 mm., oblong-elliptic; apex deeply obtusely incised, with obtuse lobes. Stamns not produced, truncate. Ovules 2. Style short.

Manipur: Hamilton 342 (BM); Wallich 1480 D; seen by Kingdon-U’ard, flowering in 1948, ‘a slim shrub 6-8 ft. high, with leaflets almost as broad as long’; possibly this is a variety with broad leaflets.

(534) Mahonia pomensis Ahrendt, sp.nov., foliis foliolorum jugis c. 7 remotis; foliolis parce spinoso-dentatis, apice inconspicue cuspidatis ; racemis tantum 1-3 cm. longis ; bracteis pedicellis gracilibus eorum brevioribus ; staminibus truncatis ; ovulis 2 ; stylo distinct0 sed brevissimo, distinguenda.

Leaves with c . 7 pairs of leaflets; the basal pair, 13 x 12 mm., set only 0.3 mm. above the base; the remaining internodes decreasing upwards from 8-3 cm. Leafets with sizes grading from base to apex, 3.5 x 2.5 cm., 4.5 x 3.7 cm., 4.5-7 x 2 . 8 4 cm., 6-8 x 2.5- 3.7 cm., the narrowly ovate terminal leaflet measuring 8.5 x 2.7 cm. ; margins 4-6- spinose (1.5-2 mm.)-dentate (3-5 mm.), distant 8-18 mm. ; apical cusp inconspicuous,

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318 L. W. A. AHBENDT 10-16 x 8-12 mm.; above dull, with venation much branched but rather indistinct; below much paler, with venation elevated, and more branched to subreticulate. Ra~ernerr 1-3 cm., fascicled c. 4; bracts of the inflorescence 1-5 cm. P d i A slender (2-) 3-4 (-5) mm.; floral bracts (I-) 15-24 (-3) mm. Outer se@ 2 x 1 mm. Median sepab 3-5 x 2 mm. Inner se@ 6 x 3-25 nun. Pe&& 5 x 3 mm., obovate, entire. Stanzens pro- duced, truncate. Ovules 2. Style distinct, but very short.

S.E. Tibet: Prov. Pome, near Trulung, Tongyuk Chu, 7500 ft., 9 Jan. 1947, Ludlow, Sherriff t Elliott 12092 (Type, BM). ‘Shrub 4-8 ft., flowers yellow, tinged red on the outside; in dense mixed forest.’ Not cultivated.

(535) Mahonia fordii Schneid. in Sargent, PI. WA. 1,383,1913; Takeda, Notes R. Bot.

Leaves to 40 x 10 cm. oblong, with 7-8 pairs of obliquely placed, distinctly remote leaflets; the basal pair, 3 x 1.4 cm., set 1.5 cm. above the baae; the remaining internodes decreasing from 9 to 3 cm. .Lea,& thin; apart from the basal pair, all fairly narrowly ovate, and fairly equal in size, or very slightly decreasin@; towards the apex, 8-6 x 2.5- 2 cm. ; terminal leaflet oblong-ovate, similar to others near the apex but larger; bases broadly rounded; margins 5-9-apinose (1-1-5 mm.)-serrate (1-1-5 mm.), distant 1- 1.8 cm. ; apical cusp fairly conspicuous, 15-20 x 3-6 mm., somewhat bent; above dull, slightly paler below; both sides enervate or very slightly veined. Racem 8-12cm., fascicled 5-7 ; bracts of the inflorescence 1-2 cm. PerEicele 3-5 mm.; floral bracta2-3 mm. Outer sepals 2 x 1.6 mm., ovate. Hdkn se@ 3 - 5 4 x 2 - 5 4 mm., ovate, obtuse. Inner se@ 4.5 x 3 mm., elliptic. Petals 4 x 2-28 mm., elliptic; apex emmgiuate with rounded lobes. Stamens 2.6 m., not produced, truncate. 0vuJe-s 2. Berries immature, 7 x 5 mm., obovoid. Style 0.25 mm.

Kwangtung: North River, Nov.-Dec. 1888, fl., fr. Ford. 17 (Type, K). Not cultivated.

Card. Edinb. 6,227,1917

(536) Mahonia schochii Schneid. ex Handel-Mazzetti, Symbo2ae Sinicae, 7 , 329, 10 Aug. 1931

Leaflets in c. 6 oblique pairs, contiguous above, separate below; apart from the small basal pair, narrowly ovate with their internodes deeming upwards from 9-4 (-3) cm. ; their length decreasing from the middle to the ends, while the breadth remains fairly constant, (7-) 9-12 (-14) x 3 cm. ; the terminal leaflet longer but much narrower; bases cuneate; margins 3-5-spinose (2-3 mm.)-serrate (2-3 mm.), distant 8-18 mm. ; dull yellow-green both sides, much paler below; venation indistinct above; veins and veinlets elevated below. Petals as long as, but narrower than inner sepals. Stanzens produced, truncate. O& solitary.

Kweichou: in silva mixta, prope Lang tai ting, h t e x nanus, 6500 ft., 14 Oct. 1916, Schoch 410 (Type, K). Not cultivated.

(537) Mahonia nitens Schneid. in Sargent, PI. Wils. 1, 379, 1913 Shrub 4-6 ft. Leaves to 25 cm. long, with 5-7 pairs of leaflets with internodes c. 3 cm.

Leaflets lustrous both sides, more or leas concolorous, with slightly elevated open reticu- lation below. Leaflets, both lateral and terminal, more or less equal, ovate, 6-8x2.5- 3.2 cm., oblique, base cuneate, margins entire for basal half but with two coarse sinuate spinose teeth each side from the middle to the apex, the apex having a conspicuous cusp. Racemes to 14 om., with small bracts. Pedicels 2-3 mm., with slightly longer bracts. F h e r s red without. Outer sepals oblong-ovate. Inner sepals similar, larger. Petah broadly oblong, as long as the inner sepals. Stamens edentate, truncate. Ovules 2, sessile.

W. Szechuan: Hungya Hsien, red sandstone cliffs, 3500 ft., Sept. 1908, Wilson 2881 (Type). Not cultivated.

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Berberis and Mahonia 319

(538) Mahonia mairei Takeda, Notes R . Bot. Card. Edinb. 6 , Y28, 1917 Leaves with 6-8 pairs of very remote leaflets, the lowest pair close to the base. Leajets,

apart from the small basal pair, oblong, 7-10 x 2 - 5 4 cm., oblique; margins with 5-12 teeth ; apex cuspidate-acuminate ; above very lustrous with conspicuously elevated reti- culation; below dull. Racemes 12-20 cm., with stout rachis, fascicled 6-7. Pedicels 4-5 mm. ; floral bracts 3 4 mm. Prophylls linear, situated on the basal half of the pedicel. Outer sepals 2 x 2 mm., ovate, obtuse. Median sepals 4 x 3.5 mm., ovate-orbicular. Inner sepals 7 x 4 mm., oblong-ovate. Petals 6 x 3 mm., oblong-elliptic; acutely emarginate with rounded lobes. Stamens 4-5 mm., produced, rounded. Ovules 4-5. Style short.

Yunnan: near Yunnan-sen, Maire 101 (Type, E). Not cultivated.

(539) Mahonia fargesii Takeda, 1.c. p. 235, Jan. 1917 Leaves to 30 x 9 cm., oblong-oblanceolate, with 6-7 fairly oblique pairs of leaflets,

remote towards the base, separate to subcontiguous towards the apex; the two lowest small pairs set 5-8 mm. and 2-3 cm. respectively above the base of the rachis, the remaining internodes decreasing upwards from 4.5 to 2.5 cm. Leaflets thick, rigid, ovate ; the basal pair 2 x 1.5 cm., the remainder increasing in size from the base to the middle (3.5-6 x 2-5-3-5 cm.), then decreasing in size towards the apex, 6-4.5 x 3.5-2.5 ; the terminal leaflet larger, 56-7 x 1-8-3 em., with petiole 1-2 cm.; bases subtruncate; margins 3-5-spinose (2-3 mm.)-dentate (4-6 mm.), distant 1-2 cm.; above dull, with venation openly reticulate, but indistinctly elevated ; below dull, slightly paler, sub- enervate. Racemes 11-18 om., fascicled c. 11 ; bracts of the inflorescence 10-20 x 6-8 mm. Pedicels 5-7 mm.; bracts 4-6 x 1-5-2 mm. Outer sepals 2.5 x 1.5 mm., ovate, acute. Median sepals 5 x 2.5 mm., oblong-ovate. Inner sepals 5.5 x 2.5 mm., oblong-elliptic. Petals 5 x 2 mm., elliptic; glands distinct. Stamens 2-5 mm., not produced, truncate. Ovules 5-7.

Szechuan : Tchen-keou, Farges, s.n. (Type, K). Not cultivated.

(540) Mahonia bealei (Fort.) Carr. FZ. de Serres, 10, 166, 1854-5; Takeda, 1.c. p. 225, 1917; s-p. Berberis bealei Fortune, Guard. Chron. 1850, p. 212; Hook. Bot. Mag. 81, t. 4852, 1855; syn. M . japonicu Hort., non (Thunb.) DC.

Shrub 4-7 ft. Lea,ves to 45 x 20 cm., narrowly obovate, with 5-8 distinctly separate (or varietally contiguous) leaflets; the basal pair, 4 x 3 cm., set 10-15 mm. above base; the remaining internodes fairly constant, 6-7 cm. LeaJEets thick, rigid, ovate; from the base to the apex their length slightly increasing and their breadth slightly decreasing, SO

that they change from broadly to more narrowly ovate, 7-10 x 6 4 cm. ; terminal leaflet much larger, 14 x 20-12 cm., conspicuously broad; base broadly cuneate to subrounded; margins (2-) 3-6 (-8)-spinose (3-4 mm.)-serrate (3-8 mm.), distant 12-20 mm. ; cusp distinct, but not conspicuous, 2.5 x 1 cm. ; above dull, sometimes becoming very slightly lustrous, grey-green ; below dull, paler yellow green ; venation indistinct both sides. Racemes erect, 5-10 cm., fascicled 6-9 (-12) ; bracts of the inflorescence 10-15 mm. Pedicels 4-6 mm.; floral bracts 2-3 mm. Outer sepals 2.5 x 2.5 mm., ovate, subobtuse. Median sepals 6x4mm. , ovate. Inner sepals 7-8x34mm., oblong. Petals 6 - 7 ~ 3-3.5 mm., obovate; apex emarginate with rounded lobes; glands distinct. Stamens 3-75-44 mm.; not produced, rounded-truncate. Berries ovoid, 10 x 6 mm., pruinose grey. Style short, scarcely 0.1 mm., or absent.

KEY 9 8 ~ Leaflots remote or separate ; apical leaflet very broad and conspicuous ; flowers large ;

Leaflets contiguous or overlapping; apical leaflet less conspicuous; flowers small; ovules 2. ovules 3-4. 540a. M . be&i var. bealei

540b. M . bealei var. p k n ~ o l i a

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320 L. W. A. AHRENDT

(540a) Mahonia bealei var. bealei Hupeh: Ichang, in flower, Henry 3283, 3250; Patung, Henry 1450 (all, K). Cultivated: in 1855, figured in Bot. Mag. t. 4852 (Type). L.A. 720, fl. 11 Feb. 1944

(W); fr. 23 June 1954 (B). This and M. jwpcmka are both very Widely cultivated, but, unfortunately, generally with the specific names reversed. Even without flower or fruit, they can easily be distinguished in cultivation by the very broad terminal leaflets of M . beaki and the narrow ones of M . japmica. Uncertainty may a&, however, because many plant8 raised from seed are hybrids between these two species.

(540b) Mahonia bealei var. planifolia Hook. f. et Thorns. Bot. Mag. 81, t. 4846, 1855

Outer se@ 2 x 1-5 mm. Mediun sepals 4 x 2.5 mm. Inner sepals 5-6 x 2.5-3 mm. Cultivated, and figured in Bot. Hag. t. 4846 (Type). According to Hooker, the smaller

flowers of this variety are only ‘6 mm. long’, while those of the preceding variety are ‘1 cm. long’; further, vhile the racemes of the latter are erect, those of the former are pendulous. I have doubts about the association of var. phnifolia with M. bealei. Com- pare here the M a h i a referred to aa Berberis cumanqainea Fortune, in J . R. Hort. SOC. 8, 226, 1852.

In J . R. H&. Soc. 7,225,1852, Lindley remarks that M. bealei waa found in a garden at Hwuychow by Fortune, who gave it the name bealei in the Gardener’s Chronicle in 1849.

(541) Mahonia japonica (Thunb.) DC., Syst. 2, 22, 1821; Takeda, I.c. p. 241, 1917; syn. M . japmka vm. gracillima Fedde, Bot. Jb. 31, 118, 120 (1901); syn. I h japonica Thunb., FZ. Jap. 79,1784; Ic. PI. Jap. 4, t. 2,1802

Leaves to 45 x 14 cm., oblong-oblanceolate, with 6-9 pairs of slightly oblique, dis- tinctly separate leaflets; the basal pair, c. 2 x 2 em., set c. 5-10 mm. above the base; the remaining internodes decreasing upwards from 6 to 3-5 cm. Leajlets thick, rigid; except for the basal two pairs, their length increasing and breadth slightly decreasing from base to apex, their shape correspondingly changing from narrowly oblong-ovate to lanceolate, 5-9 x 3-25-2-25 cm. ; terminal leaflet narrow, c. 9 x 3-5 cm. ; bases rounded to subtrun- cate; margins (1-) 2-5 (-6)-spinose (3-5 mm.)-dentate (5-10 mm.), distant 2-3.5 em.; both sides indistinctly openly veined; above slightly lustrous deep grey-green; below dull paler, yellow-green. Racemes pendulous, 10-25 cm. long, fascicled c. 10; bracts of the inflorescence c. 2 cm. Outer sepals 2.5 x2.5 mm., ovate, acute. Nedian sepals 3-5 x 2.5 mm., ovate. Inner sepals 7 x 4 mm., ovate. Petah 6.25 x 3 mm., oblong- elliptic; apex incised With rounded lobes; glands indistinct. 8tamens 4.5 mm., not pro- duced, truncate. O& 4-5. Berries 8-9 x 4-5 mm., ovoid, pruinose grey. Style very short.

K E Y 980 Leaflet margins with 2-6 teeth; terminel leaflet petiolate. %la. M . japonica vm. j a p m b M e t mergina with 1-2 teeth; terminal l ed& segsile or s u k i l e .

541 b. M. japonica var. thfwca

(541a) Mahonia japonica var. japonica

reputed to be cultivated introductions.

Tsang 20223 (K): ‘fairly common in dry silt or sandy soil, as scattered shrubs’.

Japan: Tokyo, May 1906, Teraaaki, 8.n. This and all other Japanese specimens are

Kwangtung: Loh, Chuang district, Ching Uen shan, near Kau Fung, 2 Nov. 1930,

Chekiang: according to Lindley. Cultivated: L.A. 721; fl. 27 Jan. 1944; fr. 30 June 1944.

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Berberis and Mahonia 321

(641b) Mahonia japonica var. trifurca (Loud.) Ahrendt, comb.nov.; syn. Berber& trifurcu Loud., Encycl. Fl., Suppl. 11, 1346, 1842; Lav., Arb. Segrez, p. 16, fig. 4B, 1877; Fort. in Lindl. and Paxt., Fl. Qard. 3, fig. 24, 1852-3; Lindl. J. R. Hort. Soc. 7 , 226, 1852

Chekiang: according to Lindley, in a cottage garden near the coast.

(542) Mahonia leptodonta Gagnepain, BUU. Soc. Bot. Fr. 85, 166, 1938 Leaves with up to 10 pairs of leaflets, the basal pair not very close to the base. h $ e t a

fairly equal, ovate-acuminate, base rounded-subcordate, margins with up to 50 very fine spinulose seda t ions , apex cuspidate, veins few, 4-6 cm. x 10-15 mm. Racemes 6-7 cm., fascicled 5-6. Pedkk 5-8 mm.; bracts 2 mm. Outer sepals 2 mm., ovate, acute. Inner s e p k 6-7 mm., oblong. Petuk 3 mm. (without glands according to the author). Skcmns below anthers dilated; apex not apiculate. St$e absent.

Yunnan: Mt. Tchen-fong-chan, 2200 ft., Maim, s.n. (Type). Not cultivated.

Subsect. Siamenses Ahrendt ; foliis plerumque foliolorum jugis 4-10; foliolis plerumque supra nitidis; bracteis florum pedicellis eorum fere semper brevioribus, raro paulu- lum longioribus ; petalis sepalis interioribus semper brevioribus ; staminibus plenun- que truncatis, raro apiculatis; ovulis plerumque 3-6; stylis nullis vel brevimimis, interdum significantibus. h$& in 6 1 0 (-14) pairs; nearly always lustrous above. Racemes 3-35 cm. long,

sometimes compound below, or subpaniculate. Pedicels 2-9 (-12) mm.; floral bracts 2-5-6 mm., mostly shorter than the pedicels. Stamens mostly truncate, rarely apiculah. Berries, where known, 5-9 x 4-6 mm., obovoid, ovoid, or globose. Style rarely absent, often very short and insigniscant (up to 0.4 mm.), sometimes significant, 0.7-2 mm.

Siam, Annam, Sumatra, Formosa.

KEY 99, to Subsect. Siamenses Ovules 2 ; racemes branched or subpaniculate. 543. M.kb88ii Ovules (3-) 4-6; racemes not much branched, if at all.

Sttyle (0*5-) 0.75-2 mm. Racemes 2&35 om.

Stamens apiculate; style 1.5-2 mm. 544. M . siamensis Stamens obtuse; style distinct. Cf. 553. M. philippinen.&

Racemes 5 2 0 cm. ; stamens truncate; style 0.5-1 mm. Leaflets in 5-7 pairs, contiguous or overlapping; raoemes 10-20 cm.; pedicek

Leaflets in 7-15 pairs, remote; racemes 5-8 em.; pedicels 2-4 mm.; bracts 5-9 mm. ; bracts one-half as long as pedicels. 545. M. sumatrensie

longer than pedicels. 546. M. morrisonensis Style absent or very short, not exceeding 0.4 mm.

Leaflets broad, 1 : 1.75. Ledeta narrow, 1 : (%) P 6 .

Leaflets in 12-14 pairs, very lustrous above. Leaflets in 4-6 pairs, dull above.

547. M . tikushiensis

548. M . ViWakEn868

549. M . discvlorl,foliu

(543) Mahonia klossii Baker f., J. Nut. Hist. Soc., Siam, 4, 127, 15 Nov. 1921 Leajets in 8-12 pairs; lowest pair small and close to base, others subequal, 56-10 x

2-54cm., from broadly to narrowly ovate; margins with 3 4 teeth; above lustrous. InJlorescence according to the author ‘paniculate’, and shorter than the leaves. Berries ovoid-globose, shortly stylose.

Annam: shrub 10-15 ft., a t 6500 ft., Langbian Peak (Type). Not cultivated. 21 JOURN. LINN. SOC. BOTANY-VOL. LVII

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322 L. W. A. AHRENDT

(644) Mahonia siamensis Takeda, Kew BUU. 1915 p. 422, 1915 Leaves with 6-10 pairs of contiguous or overlapping leaflets (though slightly separate

below); the b a d pair, 3 x 1.8 cm., set c. 1 cm. above the base; the remaining internodes decreasing upwards from 6 to 3 cm. k $ e i % from ovate to lanceolate; their length increasing from 4-7 to 8-10 cm., from the base to the middle, thence decreasing from 8-10 to 6-9 cm. towarda the apex; their breadth fairly constant at 2 - 5 4 cm., below the middle, but decreasing to 2-3 cm. upwards above the middle; terminal leaflet equal to the others at the apex; bases rounded-subtruncate or subcordate; margins 5-9-spinose (1 mm.)-serrate (1-1-5 mm.), distant 1-2 om.; cusp inconspicuous, 5-7 x 3-5 mm.; above very lustrous and indistinctly veined; below much paler dull yellow-green, with few elevated veins. Racemes (lo-) 20-30 (-35) cm. long, fascicled 6-11, somewhat loose flowered, with rachis, 3-5 mm. thick. Pedicek (51) 6-10 (-12) mm.; floral bracts (2-) 3-4 (-6) mm. Outer s e p k 2 x 2 mm., ovate, acute. Median se& 4 x 3 mm., ovate. Inner sepals 8 x 4.5 mm., oblong-elliptic. PetaEs 7 x 4 mm., elliptic, much incised, with rounded lobes. Stamens apiculate. O& 4-6. Berries globose, 7-9mm., excluding style, 1.5-2 mm., conspicuously pruinose blue.

Siam: Chiengmai, Doi Sutep, 50003500 ft., a shrub to 10 ft. high in evergreen jungle, fl. 13 Apr. 1910, Kerr 1107 (K); a shrub, 12 ft. high, fl. 11 Apr. 1914, Ken 1107A (Type, BM, K); Doi chien duo, a shrub 6 ft. high on open rocky ground, at 6800 ft., fl. 6 Nov. 1922, Kerr 6616 (K). Not cultivated.

(545) Mahonia sumatrensis Merrill, Contr. Am. Arb. 8,55, 1934 S h d 6ft . high. Leaves with 5-6 (-8) pairs of leaflets, mostly contiguous or over-

lapping (though separate below); basal pair 2 x 1 cm., scarcely 5 mm. above the base of the rachis; the remaining internodes decreasing u p w d from 8 to 3 cm. Leufits apart from the b a d pair, oblong-ovate, all subequal; by the base 8 x 3 cm.; towards middle 10 x 3-5 cm. ; by apex 9 x 3 cm. ; terminal leaflet also 9 x 3 cm. ; bas- rounded-subtrun- cate; margins (3-) p5 (-6)-spinose (2-3 mm.)-dentate (3-5 mm.), distant 1-2 cm.; cusp fairly conspicuous, 15-20 x 5-7 mm. ; above lustrous yellow-green, fairly distinctly veined ; below sublustrous, subconcolorous green. Raceww.~ 10-22 cm., sometimes slightly branched below, over a short part of the rachis, fascicled 8-12. PerEicels 5-9 mm., very slender; floral bracts 3 4 mm. Outer selpals 2-3 x 16-26 mm., ovate. Median sepals 445 x 2-25 mm., elliptic. Inner selpals 6-7 x 3 mm., elliptic. Pek& 5 x 2 mm., oblong- elliptic; apex emarginate with obtuse lobes; glands distinct. Stumem 3.5 mm., scarcely produced, truncate. Ovules 4. Berries globose, pruinose blue, 5-6 mm., excluding style 0-5-0-75 mm.

Sumatra: east coast, Berastagi, 4500 ft., second growth forest, fl. 24 Apr. 1925, Yates 1560 (Type, New York Bot. Gard.); same loo., 18 Feb. 1932, Baugham 11111, and fr. 8 Oct. 1929, van Leeuwen 12904 (K); in forest, fl., fr. Feb. 1921, Ridley, s.n. (K); Atjeh, Boerni, Geredong, 750043500 ft., 3 Sept. 1934, van Steenis 650 (K). Not cultivated.

(546) Mahonia morrisonensis Takeda, Notes R. Bot. Gard. Edinb. 6,239, 1917 Leaves to 50 x 10 cm., oblong-oblanceolate, with (7-) 10-15 pairs of distinctly remote

leaflets; brtsal pair 1-1-75 x 0.7-1-5 mm., set c. 5 mm. above the base of the rachis; the remaining internodes decreasing from 9 to 2.5 cm. Leafits ovate to lanceolate ; their size (both length and breadth) increasing from base to middle from 4.5 x 1.5 to 7.5 x 2.2 cm. (-10 x 2.5 cm.), thence decreasing to the apex to 5 x 1.5 cm. ; bases rounded or subtrun- cab; margins 3-7 (-lO)-spinoae (1-2 mm.)-serrate (1-3 mm.), distant 1-2 cm. ; apical cusp inconspicuous, c. 20 x 5 mm. ; above lustrous with elevated venation ; below paler green with distinct branched venation. Racemes to c. 7 cm., fascicled c. 4. Pedicels 2-3 mm.; floral bracts 2 - 5 3 mm. Outer se& 2.5 x 1 nun., ovate, acuminate. Median se&

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Berberis and Mahonia 323

3.5 x 1.5 mm., obovate. Inner sepals 5 x 2 mm., oblong-obovate. Petals 4 x 1.75 mm., oblanceolate; apex acutely emarginate with rounded lobes; glands distinct. Stamem not produced, truncate. Ovules 3. Berries 6 x 4 mm., ovoid, excluding style 0.75-1 mm. ; pedicels in fruit 3 4 mm. ; bracts 5-6 mm.

Formosa: Mt. Morrison, fl., fr. 18 Nov. 1906,7500 ft., Kawakami & Mori, s.n. (Type); Tayataka, dense Picea forest, 7500 ft., 21 Oct. 1912, Wilson 960 (K). Xot cultivated.

(547) Mahonia tikushiensis Hayata, Icones Plantarum Formosanarum, 5, 5, 1915

Leaves c. 25 x 10 cm., oblong, with 9-11 pairs of leaflets; the basal pair very small and close to the base; the remaining internodes constant, c. 3.5 cm. Leaflets decreasing insize from the middle to the ends, to 7 x 4 cm., overlapping; terminal leaflet about equal to the middle ones, with a petiole 2-2.5 em.; lustrous both sides; pale yellow-green below; elevated reticulation more clear above than below; margins 3-5-spinose (2 mm.)-dentate. Racenzes c. 10 cm. long. Outer sepals 2.5 x 2 mm., ovate, obtuse. Medkn sepals 5 x 3.5 mm., similar. Inner sepals 7 x 4 mm. Petals 5-5 x 3 mm. ; apex rounded in outline, with incision 1 mm. deep, flanked by two contiguous rounded lobes; glands 1.5 mm. Stamens truncate. Ovules 6. Style absent.

Formosa: Tikushizan, Shimada, s.n. Mar. 1915 (Type).

(548) Mahonia oiwakensis Hayata, 1.c. 6, 1, 1916 Leaves 2 5 4 0 x 10-20 cm., with 12-14 pairs of leaflets; basal pair 1.5-2.5 x 0.7-1-5 em.,

with 1-2 teeth. Leaflets with their length increasing and breadth decreasing, along the rachis, 8-10 x 2-1.4 cm., narrowly ovate to lanceolate; terminal leaflet longer, linear- lanceolate; bases rounded to cordate; margins with 2-3 h e spinulose teeth; very lustrous above; paler below; midrib slightly impressed above and raised below; lateral veins indhtinct and scarcely elevated above, slightly elevated below. Racemes 9 cm. ; bracts of the inflorescence 3.5 x 2 cm. Berries conical-ovoid, stylose, 9 x 6 mm., pruinose blue.

Formosa: Musha Oiwake, Hayata, s.n. May 1916 (Type); Gokwanzan, Mori s.n. Feb. 1910. Not cultivated.

.

(549) Mahonia discolorifolia Ahrendt, sp.nov. ; foliis foliolorum jugis 4-6 ; foliolis ovatis ad lanceolatis, margine dentibus 1-6, apice conspicue cuspidatis; bracteis florum pedicellis eorum subaequilongis; staminibus truncatis, distinguenda.

Leaves to 43 x 9 em., oblong with 4 4 pairs of very oblique leaflets, separate by the apex, remote below; the basal pair 4-5 x 15-14 cm., set 6-8 mm. above the base; their internodes decreasing from 7 to 3.5 cm. up the rachis. Leaflets varying up the rachis from ovate to lanceolate, their length continuously increasing and breadth decreasing, 76-10 x 2.5-1.8 cm.; bases cuneate; margins 1-6-spinose (1.5-2 mm.)-serrate (2-3 mm.), distant 7-20 mm. ; apex with a conspicuous bent cusp, 30 x 7 mm. ; terminal leaflet 10-12 x 2-24 em., with petiole 2-3 cm. ; both sides with indistinct branched venation; above dull grey-green; below much paler yellow-green, subreticulate. Racemes 3-6 em., fascicled c. 3. Pedicels subequal to their bracts. Outer sepals 2.5 x 1.5 mm., ovate, acute. Median sepals 3.25 x 2 mm., ovate, obtuse. Inner sepals 4 x 2 mm., elliptic. Petals 3.25 x 1.5 mm., oblong-ovate; apex retuse, with rounded lobes. Stamens 2-25 mm., not procluced, truncate.

Formosa: Prov. Kagi, Arisan, Matsugama, 8700ft., fl. 10 Feb. 1918, Wilson 9816 (Type, K), ‘rare shrub, in forests, 5-8 ft. high.’ Not cultivated.

21 -2

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324 L. W. A. AERENDT

Subsect. Dentatae Ahrendt, submct.nov., staminibus infra anthers dentatis dis-

Leaflets in 6-12 pairs, often conspicuously overlapping; below yellow-green; margins with 3-8 teeth. Racenaes 5-33 cm. long. Pedkeh 2-10 mm., floral bracts much shortar or wbequal. Pekrls shorter than inner sepals. Skzmew below the anthers acutely dentate ; apex truncate or apicubte. ovolles 2-6. 8tyl.e short, or up to 1 mm. long.

tinguenda.

Sikkim, South India, Kweichou.

f i ~ 100, to Subsect. Dentatae Ovules 2; outer sepals c. 1.6 mm. long. Ovules 4-6; outer sepals 3-4 mm. long.

550. M . leveiUeana

Racemea 15-33cm. long; pedicels 7-1Omm.; bracta 2-3mm.; stamens truncate;

Racemes 6-9 em. long ; pedicels 3-4 mm. ; bracts subequal to pedicels ; stamem apicu- ovules 5-6; style 1 mm. 551. M . h c h & i

late; ovules 4; style short. 552. M . e % k k i d

(550) Mahonia leveilleana Schneid., in Sargent, PI. Wik. 1,385,1913; syn. M. &am (Uvejlld) Rehder, J . A m . Arb. 17, 322, 1936; syn. Berberis cham Uveilld, Bad. Soc. Bot. Fr. 51,289,1904, non Petzold & Kirchner, Arboretum bluSca&me, p. 132, no. 101, 1864

Leaves to 45 cm., with 8-9 pairs of leaflets; the lowest pair much smaller, and close to the base. Leaflets very variable; the lowest two paire rhombic-ovate, 3-5-55 x 2.5- 3 cm. ; the middle pairs obliquely ovate-elliptic, up to 10 x 3.2 cm. ; those towards the apex ovate-lanceolate, 7-9 x 2-5-3 cm. ; bases rounded or truncate; apex acuminate; terminal leaflet oblong-ovate, 11 x 4.3 cm.; margins unddte , with 4-8 teeth; openly reticulate with lateral nerves distinct. Raeenzes to 18 cm. long. Pedkds 2-3 mm.; floral bracts 1-16 mm. outer se* 1.5 mm. long. Petals slightly shorter than inner sepals. O v u h 2, sessile.

Kweichou: around Kouy-yang, 7 July 1898, Bodinier 2469 (Type). Not cultivated. A note is necessary concerning the correct name of this species. In 1904 Gveill6

gave to it the name Berber& +am. In 1905 Schneider published a B. +am (a true Berberis) by raising to specific rank B. Senensis var. +am Franchet. In 1913 Schneider remarked that LBvejll6’s B. &am waa only a manuscript name, and consequently it wa8 not necessary to transfer this invalid specific name when referring the plant to the right genus; further, he did not consider such a procedure desirable on account of his own use of the specific epithet +am for a true Berber&; Schneider therefore gave to Leveille’s plant the name Mahoniu leveilleana.

Now Rehder has remarked that the name B e r b e ~ elegans Uveill6 was validly published for this Mahoniu, and from this fact he draws two conclusions: (1) that for the true Berberis h a m of Schneider, a fresh name must be provided, since elegans must now be regarded a.s a later homonym already preoccupied by Uveill6 for the Mahonia; consequently, in a footnote, and with bare synonymy, he has proposed BerberisschneQderi. (But this requisite specifk name had already been supplied-it is B. amoena Dunn and has been used for the Berberis species in question in these pages) ; (2) that, since the name B. elegam Uveill6 haa been validly published for the M h i a , the correct name for the latter must be M . ekgans (IRiveill6) Rehder. [See under (193).]

The reawning of this shtement is, of course, quite correct. But Leveille’s Berberis &am itself, of 1904, is a hter, and invalid, homonym. In 1864, in Petzold t Kirchner, Arboretum Mmmvieme, p. 132, no. 101, there is described a B. ekgans Hod. Lips. This has no Latin diagnosis, but such is not demanded for validity a t this early date. The long

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Berberis and Mahonia 326

description given in German suggests that their plant is B. lycium Royle. The fact that the earlier homonym of Kirchner is also a submerged synonym for B. lycium does not affect the matter; it has quite obviously been based on a plant very difFerent from that of Uveilld’s later homonym; in fact on a Berberis and not on a MaJwnia.

Since Gveillt?s name is, therefore, itself invalid, the transfer of the epithet elegans, to Mahonia is rendered illegitimate by the existence of the name M. leveilleana. Consequently, and fortunately (for I agree with Schneider’s view as to the desirability of letting the name elegans lapse), the correct name for this species is Mahonk leveilleana Schneider.

(551) Mahonia leschenaultii (Wall.) Takeda, Notes R. Bot. Card. Edinb. 6, 223, Jan. 1917; Chatterjee, Rec. Bot. Sum., Indk, 16 (no. 2), 38, 1953; syn. Berberis Eeschenuultii Wall. ex Wight et Am., Prodr. Fl. Pen. Ind. Or. 1, 16, 1834; Wight, Icon. P1. Or., t. 940 (1845) ; syn. B. nepalensis var. leschenuultii Hook. f. et Thorns. Fl. Brit. I d . 1, 109, 1872

Leaves to 48 x 13 cm., oblong, or spathulate-oblanceolate, with 8-10 pairs of slightly oblique and conspicuously overlapping leaflets; the lowest pair, 1.5-3 x 1.3-2-2 om., set c. 5 mm. above the base; the remaining internodes decreasing from 6 to 2.5 cm. Leaflets thick, rigid ; from base to apex their breadth continuously but slightly decreasing from 3 to 2 om. ; but their length increasing from the base to the middle from 2.5 to 5.5 cm., thence towards the apex slightly decreasing from 5.5 to 4.5 cm.; the terminal leaflet subequal to the adjacent ones; the lateral leaflets sometimes as large as 8-5 x 4.2 cm. ; brtses subcordate-truncate ; margins 3-7-spinose (2-3 mm.)-dentate (3-7 mm.), distant 1-2 cm. ; cusp inconspicuous, 15 x 12 mm. ; above very lustrous with venation and some elevated reticulation indistinct ; below scarcely paler, subenervate. Racema 15-33 cm. long, fascicled 7-13; bracts of the inflorescence c. 15 mm. long; rachis 2-3 mm. thick. Pedicels slender, 7-10 mm.; floral bracts 2-3 mm. Outer sepals 3 x 3 mm., broadly ovate, obtuse. Median sepals 4-5 x 3-4 mm., oblong-ovate. Inner sepals 6-7 x 4 mm., oblong- ovate. Petals 5-6 x 3 mm. ; basal glands distinct ; apex acutely emarginate with rounded lobea. Stamns scarcely produced, truncate. Ovules 5-6. Berrk pruinose blue, sub- globose, c. 7-8 mm., excluding style c . 1 mm.

South India: Nilghiri Hills, fl. WalIich 1479 (Type, K); fr. Wight 49,50 (K); Falconer Shala, fr. 6 June 1897, Bourne 309 (K); Gunwattu Shola, fl., and immature fr. 25 Dec. 1898, Bourne, s.n. (K); Hohenacker 1125 (K); Viscount Gough 53 (K); Ootamacund, 7500 ft., fl. Sept. 1883, Gamble 12443 (K). Not cultivated.

(552) Mahonia sikkimensis Takeda, 1.c. p. 220, 1917; Chatterjee, 1.c. p. 40, 1953 Leaves with 8-12 pairs of leaflets, mostly overlapping or contiguous, though separate

near the base; the lowest pair, 2.5 x 1.5 om., set 8-15 mm. above the base. Leaflets varying from broadly to narrowly ovate; from base to apex their internodes decrease from 4 to 1.5 cm., and their sue (both length and breadth together) increases from P 8 x 2-3-5 cm. ; the terminal leaflet is similar to, but slightly larger than, the adjacent ones; base truncate; margins (3-) 4 4 (-6)-spinose ( 3 4 mm.)-dentate (4-6 mm.), distant 1-2 cm. ; apical cusp inconspicuous, 15-20 x 7-9 mm. ; venation indistinct both sides; paler below. Racema 5-9 cm. long, fascicled c. 5. Pedicels c . 4 mm.; bracts subequal. Outer sepals 36-4 x 1.5-2 mm., ovate, acute. Median sepals 6 x 3 mm., oblong-ovate, acute. Inner sepals similar, 7 x 3-5 mm. Petals 6 x 3 mm., oblong-elliptic; basal glands conspicuous ; apex acutely emarginate, with rounded lobes. Stamens apiculate. Ovules 4. Style short.

Sikkim: Latong, 7000ft., 1885, fl. King’s collectors (Type, Herb. Calcutta, fig. in Herb. Kew). Not cultivated.

Page 326: Berberis and Mahonia : A taxonomic revision

326 L. W. A. AERENDT

Subsect. Dolichopodae Ahrendt, subsect.nov., racemis conspicue laxis; pedicellis longis (0-7-) 1-2cm. longis; bracteis multo brevioribus, tantcm 2-3mm. longis dis- tinguenda.

Lmve-s subeeseile, with 2-10 pairs of l d e t a , the lowest pair small and close to the base. Stanzens dentate, not produced, truncate.

Philippine Islande, Yunnan, Szechuan.

~ E Y 101, to Subsect. Dolichopodae Leafiete in 5-10 pairs, below epruinose yellow-green; racemes 9-20 cm. long; pedicels

Leaflete in 2-3 pairs, below pruinose white, at my rate at first; racemes 25-55 cm. long; 7-12 mm. ; petals subequal to inner sepals.

pedicels 16-20 mm. ; petals shorter than inner sepals.

553. M . philippinensis

Leavea reticulate. 554. M. gracilipea Leavea with 3-5 parallel longitudinal veins. 555. M. subtr ipl ined

(553) Mahonia philippinensis Takedrt, Notes R. Bd. Card. Edinb. 6, p. 244, Jan. 1917;

L4ave.s 15 x 5-30 x 8-10 cm., with 5-9 (-10) pairs of leaflets which are approximate towards the apex, more widely separate towards the middle, and remote towards the base; the lowest pair c. 2 x 1 cm., set 10-15 mm. above the base; the remaining internodes decreasing upwards from 4 to 2 cm. Leu. shape varying up the rachis from ovate to lanceolate; from the base to the middle, the length increasing but the breadth constant, 3 x 2-7.5 x 2 cm. ; from the middle towards the apex their size (both length and breadth together) slightly decreasing from 7-5 x 2-6 x 1.5 cm.; terminal leaflet similar to, but larger than, the adjacent ones; base truncate; margins 3-6-spinose (2-3 mm.)-dentate (2-3 mm.), distant 1-2 mm.; apical cusp fairly conspicuous, c. 20 x 8 mm. ; above dull, subenervate; below much paler with branched elevated venation. Racenzes 9-17 (-25) cm. long, loose, fascicled c. 7; rachis 1-5-2 mm. thick. PediceLs 7-10 (-12) mm.; floral bracts (2-) 2 - 5 3 mm. Outer sepah 2 x 1.5 mm., ovate, obtuse. Median sepals 3-36 x 3 mm., ovate. Inner sepah 6 x 3 mm., elliptic. Petals 6 x 3 mm., elliptic; basal glands distinct; apex acutely emarginate with rounded lobes. StamRns not produced, truncate. ovules 2. Berries pruhose blue, globose, c. 8 mm., excluding style, 0.25 mm.

Philippine Islands: Luzon, Prov. Benguet: fl., fr. Loher 60 (Type); Baguio, fl., fr. 12 Nov. 1904, Williams 1460 (K); Mar. 1904, Elmer 5929 (K). Not cultivated. Takeda and Schneid. described this independently, but gave it the same specific name.

Schneid., Bd. Gbz. 63, 520, June 1917

(554) Mahonia gracilipes (Oliver) Fedde, Bot. J b . 31, 128, 16 Aug. 1901; Takeda, 1.c. p. 237, 1917; syn. Berber6 gracilipes Oliver, in Hook, Ic. Pl., ser. iii, 8, t. 1754, 1887

Leaves to 64 x 12 cm., oblong-oblanceolate, with 2-3 pairs of very oblique, remote laflets; the lowest pair similar to the others, only slightly smaller, and set 3-7 cm. above the base of the rachis; the remaining internodes decreasing upwards from 15 to 9 cm. LeaJets fairly thick, all subequal, 8-11 x 2.5-3.5 cm., or 4-12-5 x 3-5 cm., or to 13-5 x 4.5 cm. ; terminal leaflet 12-5-15.5 x 3-5.5 cm., with petiole 4 - 6 7 cm. ; base cuneate ; margins entire below, but towards apex 2-5-spinose (1-2 mm.)-serrate (2-3 mm.), dis- tant 1-2 cm. ; cusp fairly conspicuous, 20-25 x 7-8 mm. ; above dull with somewhat elevated openly reticulate venation; below, at first pruinow white, jinally dull paler green, with elevated branched venation, scarcely reticulate. Racenzes very loose, 25- 55 cm. long, faacicled, c. 3; bracts of the inflorescence 2-2.5 cm. PediceLs very slender, 1.5-2 cm. ; floral bracts 2 mm. Outer sepah 2.5 x 1.9 mm., ovate, acute. Median sepah 4 x 2.75 mm., ovate, acute. Inner sepals 4.5 x 2-75 mm., elliptic. Petals 3.5 x 2-5 mm.,

Page 327: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 327 obovate; apex slightly retuse with rounded lobes. Stamens 2.25 mm., apex not produced, truncate.

Szechuan: Mt. Omei, 4000ft., Faber 85 (Type, K); Mt. Omei, Nov. 1902, Wilson (Veitch) 3144 (K), ‘on cliffs, 2 ft. high’; Aug. 1904, shrub, 3 ft. high, Wilson (Veitch) 4727 (K); Omei shan, fl. 16 July 1930, Fang 7670 (K). Not cultivated. This species shows a tendency towards Subsect. Nervosae.

(555) Mahonia subtriplinervis (Pranch.) Fedde, Bot. Jb. 31, 129, 16 Aug. 1901 ; syn.

Leaflets in 3 4 pairs, very pruinose below; base cuneate ; apex acute ; margins, below the mid-point entire, above spinulose; terminal leaflet much larger; conspicuous in their venation with 3-5 parallel longitudinal nerves, the outer ones being almost as clear as the midrib. Racemes loose flowered with slender pedicels three times as long as the flowers, which are pale yellow.

Yunnan: in woods of the region around Tchen-fong-chan, Delavay 5024 (Type). Not cultivated.

Berberis subtriplinervis Franch. Bull. Mw. Hist. Nut. Pa&, 1, 63, 1895

Subsect. Integrifoliolatae Ahrendt, subsect.nov., foliolis marginibus integris distinguenda.

Only one species has, so far, been recorded under this heading:

(556) Mahonia shenii Chun, J . Arn. Arb. 9, 127, 1928 Bhrub 3-7 ft. Leaves to 40 cm., with 1-2 pairs of leaflets; internodes 6-4 cm. Leaflets

elliptic to ovate-lanceolate, the basal pair scarcely smaller, 8-13 x 3-5 cm,; terminal leaflet 13-15 x 6 cm., oblong-elliptic ; base cuneate ; margins entire ; venation subreticu- late above; below much paler yellow-green, with indistinct venation. Racemes c. 10 cm., fascicled 6-8; bracts of the inflorescence lanceolate, 1-2 cm. Pedicels 3-4 mm.; floral bracts 1 mm. Berries depressed globose, 6-7 mm., estylose, pruinose blue.

Kwangtung : north river region, about 20 miles north-west of Chengkou, on the road to Pan lung tsze, west slope of shaded ravine, fr. 24 Dec. 1927, Chun 5850 (Type, Am. Arb.). Not cultivated. This species, again, with its lack of the small basal pair of leaflets tends towards the following Subsect. Nervosae.

Subsect. Nervosae Ahrendt subsect.nov., foliis petiolatis; foliolis inferioribus non versus basim rhachidis, et non multo minioribus, distinguenda.

Hupeh, Szechuan, Yunnan, h a m , and one species (the only one from this subgroup) in 1J.S.A.

KEY 102, to Subsect. Nervosae LeaAets very narrow, 1 : (6) 6-10; racemes only 3-5 cm. long; stamens apiculate.

Leaflets in 1-4 pairs, narrow (1 : 67), with 6-10 marginal teeth. Leaflets in 8 pairs, narrower, 1 : 7-10, with 15-20 marginal teeth.

Leaflets subconcolorous below; racemes 5-6 cm. ; bracts subequal to pedicels; stamens

Leaflets paler below; racemes 10-20 cm. long; bracts one-half as long as pedicels;

557. M . fortuwi 558. M . eetosa

Leaflets broad, 1 : 1.3-2.5; racemes (5) 10-20 cm. long.

truncate; ovules 2; style short. 559. M . decipiens

ovules 5-8; fruit globose, 6-8 mm. Leaflets in 2-3 pairs; stamens apiculate; style 2 mm. Leaflets in 4-10 pairs; style short or absent.

560. M . annamim 561. M . nermosa

Page 328: Berberis and Mahonia : A taxonomic revision

328 L. W. A. AHXENDT

(557) Mahonia fortunei (L5dl.) Fedde, Bot. Jb. 31,130, 16 Aug. 1910; syn. Berberis

S A d to 6 ft. h u m 8-23 x f5-18 cm., spathdate obovate with 1-4 p a h of distinctly remote oblique leaflets; small basal pair absent, the lowest pair being similar to the others and set (2-) 4 7 cm. above the baae of the rachis; remaining internodes de- creasing from 3.5 to 2.5 cm. L e u . fairly thin, oblong-lanceolate to elliptic-lancedate; all subequal, or, from baae upwards, with their size very slightly increasing; terminal leaflet larger, sessile; mmgins with 6-10 teeth; apex acute to subacuminate, without cusp; above dull deep green, with few veins generally indistinct, some veinlets openly elevated ; below paler sublustrous yellow-green, with branched elevated venation. Racema 3-5 cm. long, fascicled 4-8; bracts of the inflorescence, 7-12 mm.

fo*ounei Lindl. J. R. Hort. Sm. 1,231,1846

KEY 102A Leaves 16-23 x 10-18 cm., with 3-4 pairs of leaflets, the lowest P 7 cm. above the base;

ledeb 10-12 x 1.5-1-9'cm.; terminal leaflet 12-16 x 1.7-2 cm.; base cuneate mute; margins 6-10 (-12)-spinose (1-2 mm.)-serrate, distant 1-2 cm.; pedicels 2-4 nun., with bracts 2-3 mm., shorter or subequal.

Leaves 8-12 x 6-9 om., with 1-2 (-3) pairs of leaflets, the lowest 2-3 cm. above the base;

dentate (2.5-6 mm.), distant 6-12 mm.; pedicel~ only 1-2 mm., with longer bracts 1-5-2.5 mm. 557b. M. fortune$ var. Bzechmnicu

557a. M. fortunei var. fortunei

leaflets 6-8 x 1.2-1.3 cm.; base cuneate, WU~B; -gins 6-9-spino~e (1-1.5 -.)-

(557a) Mahonia fortunei var. fortunei Outer se+ 2-25 x 2 mm., ovate, acute. Mdkn s e e 4-5 x 2.5 mm., oblong. Inner

se@ 5 x 2.5 mm., oblong-elliptic. Pet& 3.5 x 1.6 mm., entire. Xtamm 2 4 mm., dis- tinctly conical-apiculate.

Hupeh: 1843, Fortune, am. (K). Cultivated: in garden at Shanghai, fl. 1846, Fortune 32 (Type, K); L.A. 724, fl. Oct.

1938. Them is, apparently, a hybrid, M. xhe.terophyUa Schneid., in IU. Handb., Laubh. 1, 318,1906, which, according to Zabel, is M. aquifdium x M. fortunei.

(557b) Mahonia fortunei var. s z e c h d c a Ahmndt, vm.nov., foliis foliolorum jugis tantum 1-3; foliis petiolisque eorum et fofiolis multo minoribus; foliolis margini- bus c r w dentatis; pedicellis multo minoribus, sed bracteis eorum distincte longioribus, distinguenda.

Ovulee 2. Berries globose or oblong-globose, 4 4 mm., estylose, pruinose blue. Szechuan: Lu-shan hsien, 4000 fi., among bushes, a shrub 1-2 ft. high; leaves dark

green, beneath light green, smooth; flowers yellow; fr. 19 Oct. 1936, Chu 4057 (Type, E).

(558) Mahonia setosa Gagnepain, BuU. Xoc., Bot. Fr. 55 (4th ser., 8), 86, 1908 Leaves with 8 pairs of leaflets, and with very slender rachis; lowest pair of Ieaflets

very remote from the base. LeaJlete linear-lanceolate, G. 7 cm. x 7-12 mm. ; base cuneate ; margins 15-20-spin0se (3-4 mm.)-serrulate; apex acuminate; reticulate. Racenzes 3-4 cm. Outer sepals ovate, acute. Median and inner sepals subequal, lanceolate. Petals scarcely shorter than inner sepals; baae clawed, with distinct glands. Stamtxw with apex produced, obtusely subconical. ovules 2, stipitate.

Yunnan: near Tiou-pa-teou, fl. Oct. 1894, Delavay 6830 (Type). Not cultivated.

Page 329: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 329

(559) Mahonia decipiens Schneid., in Sargent, PI. Wils. 1, 379, 1913 Shrub to 6 ft. Leaves to 20 x 11 cm., narrowly obovate with 2-3 pairs of more or less

contiguous, oblique, leaflets; small basal pair absent, the lowest pair set 3-6 cm. up the rachis; the remaining internodes decreasing upwards from 4 to 3 cm. Leajets rigid, all similar, but slightly increasing in size upwards, 4 4 x 34-5 cm.; terminal leaflet 8-9 x 5-55 cm., petiole 2.5 cm.; base of lateral leaflets truncate or slightly cordate; margins 4-6-spinose (1-5-2 mm.)-dentate ( 2 4 mm.), distant 1-2 cm.; apex subacuminate with inconspicuous cusp, c. 2 x 1.7 cm. ; both sides dull, subconcolorous, with indistinct open elevated reticulation. Racemes 5 6 cm., fascicled c. 2; bracts of the inflorescence G. 1.5 cm. Pedicels 4 mm.; floral bracts 4 x 2 mm. Outer sepals 2 x 1 mm., ovate, obtuse. Median sepals 3 x 2 mm., ovate. Inner sepals 5 x 3 mm., oblong-elliptic. Petals 4.5 x 3 mm., obovate, emarginate. Stamns not produced, truncate. Ovules 2. Style short.

W. Hupeh: Chang-yang-hsien, woodlands, 5000ft., fl. Apr. 1907, Wilson 2884 (Type, K). Not cultivated.

(560) Mahonia annamica Gagnepain in Lecomte, FZ. Gen. Indo-Chine, 1, 157, 1907;

Shrub 15-20 ft. Leaves with 2-3 pairs of leaflets, and slender rachis; small basal pair of leaflets absent, the lowest pair being situated about the mid-point of the rachis. ZeafEets ovate, 8-10 x 2.5-4 cm., all similar and subequal; margins 7-8-spinose (1-2 mm.)- serrate ( 2 4 mm.); openly reticulate; above grey-green; below paler. Racemes c . 20 cm. Pedicels with bracts half as long as themselves. Outer and d i a n sepals ovate. Inner 5epal.s lanceolate, at least twice as long as the median sepals, sometimes even longer. Petids lanceolate, obtusely acuminate; glands prominent. StamRns with apex apiculate with long centre tooth flanked by two shorter ones. Ovules 5. Berries pruinose blue, globose, 6-7 mm., excluding style c. 2 mm.

Annam: Haut Donnai, plateau de Lang-bian, 5-6 ft., at 8400 ft., Jacquet 629 (Type). Not cultivated.

I.c. p. 84, 1908

(561) Mahonia nervosa (Pursh) Nutt., Gen. A m r . Pl. 1, 218; 2, n. 307 (1818); syn. Berberis glumaeea Spreng, Syst. 2, 120, 1825; Lindley, Bot. Reg. t. 1426, 1831; Lindley, J. R. Hort. Soc. 5, 18, 1850; Mahonia glumacea DC., Syst. 2, 1821; syn. Berberis nervosa, Bot. Mag. 68, t. 3949, 1842

Leaves with 3-10 pairs of leaflets, separate to remote, the leaves being 40-60 cm. long; the lowest pair of leaflets set (3-) 6-10 cm. above the base; the remaining internodes decreasing upwards from 6 to 4 cm. LeaJlets thick; their length from the base to the middle increasing from 4 to 7 cm., thence to the apex decreasing from 7 to 5 cm. ; their b d t h continuously but slightly decreasing from base to apex, 35-26 (-2) cm. ; their shape changing upwards from ovate to lanceolate ; terminal leaflet with petiole 2-3 cm. ; margins 8-15-spinose (1-5 mm.)-serrate (1.5 mm.), distant 8-12 mm.; both sides sub- concolorous, loosely veined, dull, somewhat more grey above and more yellow below. Racemes (7-) 10-20 cm. long, fascicled 2 4 ; bracts of the inflorescence 2-3 cm. Pedicels (6) 6-12 (-15) mm. ; floral bracts ( 2 . 6 ) 4-6 (-10) mm. Outer sepals 3 x 1-75 mm. Median sepals 5-5-5 x 2-75-3 mm. Inner sepals 7 x 4 mm. Petals 56-45 mm., emarginate. Sta7nens 4 mm., edentate. Berries with style absent or very short, slightly pruinose blue, globose, 6-8 (-9) mm. Ovdm 6-8. Seeds black.

British Columbia: Howell, s.n.; Yale, Macorun 66, fl. 6 May 1875. Idaho: Latah Co., woods 4 miles east of Farmington, fr. 26 June 1892, Sandberg 507. California: Lobb 177. Washington: Cascade mountains, dry woods in the upper valley of the Nesqually,

fl. 6 May 1895; fr. 3 Sept. 1895; Allen 136.

Page 330: Berberis and Mahonia : A taxonomic revision

330 L. W. A. AHRENDT Vancouver Island: on rocky ledges, Renfrew district, Edinburgh, 2000ft., 1 Aug.

(all, K) 1912, Rosendhal643. In Allen 136, in flower the pedicels are 6-8 mm., and their bracts 3-4 mm., and there

is a short style to the ovary; but in his fruiting specimen, the pedicels are only 4 4 mm., and their bmta 4-5 mm. long.

The emliest specific name of this plant is mmosa, when it was called Berberis nervosa. The earliest name, as a M a h i a , is M . glutnuma DC. The transfer of specsc names from one genus to another has demanded that the name M . w o s a be used. However, Lindley, in J. R. Hort. SOC. 18,1850, wing the generic name Berberis persists in using the title B. glumacea; he rejects the earlier specific name, n e m a , in the following words: ‘Pursh conferred the name mrvosa upon a plant with the leaves of glurnmea and the flowers of aqui;folium. As there is no such plant as a Barberry with the leaves of one species and the flowers of another, it was clear that the name was given to a nonentity, and must be cancelled.’

It is certainly true that Pursh did make this error, which is at once perceived on account of the dentate stamens which he has conferred on his B . nervosa.

Nuttall did not correct this point but included M . m m a under the headmg of the genus Mahonia which he describes as possessing dentate stamens as a generic character ; as we have seen, only a minority of species of M a h i a have dentate stamens.

This subsection is related in its petiolate leaves to the North American species which comprise the following, and h a l , part of this work. In M . m m a a , the only species of the ORIENTALES and LONQIBRACTEATAE to come from the western hemisphere, this relation- ship is supported geographically. As its chief botanical characters demand its inclusion in the present section, it comes appropriately at the end ofi t ; thus the Subsect. N e m a e , and the species M. nervosa, p m n t a link with the OCCIDENTALES (and the three sections which it comprises).

APPENDIX 3 The following species of M a h i a are not su5ciently fully recorded to be classified:

(562) Mahonia hypoleuca Takeda, Note5 R. Bot. Qard. Edinb. 6, 238, Jan. 1917; syn.

Leaves to 50 x 12 cm., oblong-oblanceolate, with 8-9 pairs of slightly oblique, separate to remote, leaflets; the small basal pair, 23 x 1.6-2.5 cm., ovate, coarsely toothed, set 1-2 cm. above the base of the rachis; the remaining ipternodes decreasing from 6 to 4.5 cm. Lea$& fairly thin, all ovate; their size increasing from the base to the middle from 4.5-8 x 3 4 . 5 cm. ; but from the mid-point to the apex, their size slightly decreasing to 5-5-6-5 x 3 . 2 4 cm. ; terminal leaflet similar and scaxcely larger; base broadly rounded, subtruncate; margins subentire, or with 3-7 inconspicuous teeth; apical cusp incon- spicuous, 5 x 3 mm. ; above sublustrous yellow-green, with few lateral veins impressed, and fine reticulation indistinctly elevated ; below conspicuously pruinose white, with open reticulation distinctly elevated, and closer reticulation more finely marked. Plowers and fruit unknown.

Yunnan: Mengtze, south-west mountains, 6000 ft., Henry 9863 (Type, K). A shrub of 3 ft., not in cultivation, but distinct in the pruinose under-surface of the leaves.

M . nivea Schneid., Bot. Qaz. 63, 521, June 1917

(563) Mahonia huiliensis Handel-Mazzetti, Symbdcce Sinicae, 7 , 329, 10 Aug. 1931 Leaves 18-25 cm. long with 7-9 pairs of leaflets, mostly contiguous; rachis slender

with internodes (3-) 2-1-3 cm. ; lowest pair small, close to baae. LeaJlets ovate-lanceolate, slightly larger a t middle, to 8 x 2.5 cm. ; terminal leaflet larger; base broadly rounded, slightly oblique; margins (3-) 5-8-spinose (c. 1 mm.)-serrate (16-3 mm.), distant

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Berberis and Mahonia 331

7-17 mm. ; apical cusp 17 x 7.5 mm. ; lustrous above. Racemes to 18 cm. Pedicels c . 2.5 mm., with slightly longer bracts. Outer sepals 1-16 mm., narrow. Inner sepals broad, 2.5 mm. Petuls c. 3 mm. long, emarginate. Stamens edentate, not produced, truncate. Ovules 3. Style scarcely 0.5 mm., indistinct.

Szechuan: by Huili, 7000-7600 ft., 22 Oct. 1914, Handel-Mazzetti 5632 (Type). Not cultivated; probably of the Subsect. Japonicae.

(564) Mahonia caesia Schneid., Bot. Guz. 63, p. 519, June 1917 Shrub 4-10 ft. Leaves with 6-9 pairs of leaflets, the leaves being c. 25 cm. long, the

basal pair of leaflets very close to the base, and the remaining internodes 2.5-1.5 cm. Leaflets narrowly lanceolate, slightly larger in the middle than a t the ends, thickly coriaceous, smooth, above yellow green, below somewhat pruinose, base rounded-truncate ; margins undulate, with 5-9 coarse spinose (1.5-2.5 mm.) teeth. Otherwise unknown.

N.W. Yunnan: mountain sides between Lichiang fu, and River Yangtze, by Yung- peh-ting, 3 July 1914, Schneider 1723 (Type, Am. Arb.). Not cultivated.

APPENDIX 4 Berberis trqaeanthoides DC. Syst. 2 , 18, 1821, and Berberis caraganuefolia DC. l .~ . ,

belong not to the genus Berberis or Mahonia but to the genus Caraganu.

Mahonia. Group Occidentales Ahrendt, inflorescentiis paucifloratis et fasciculatis vel subumbellatis, vel racemosis laxis solitariis, vel laxe paniculatis ; vel racemis fasciculatis cum foliolis distincte reticulatis vel nervatis hypodemate null0 ; bracteis inflorescentiis deciduis tantum 2-5 (-8) mm. longis.

North West, and Central, America.

KEY 103, to Sects. of the Group Occidentales (Maps 1 and 47) Inflorescence many-flowered, in raceme8 (fascicled or solitary), or in panicles; leaves

reticulate or veined, without hypoderm, fairly thin in texture. Inflorescence a fascicle of (2-) 3-5 more or less dense and erect racemes.

Inflorescence a solitary loose pendulous raceme or panicle.

faintly veined, more or less stiff, often thick and rigid.

Sect. AQU~FOLIATAE Sect. PANICULATAE

Sect. HORRIDAE Inflorescence few-flowered, fascicled, umbellate, or subumbellate ; leaves enervate or

Sect. Aquifoliatae Fedde, Bot. Jb. 31, 69, 75, 16 Aug. 1901 Leaves with 2-6 pairs of leaflets. Racemes 1-9cm. long, fascicled 2-7. Pedicels

5-10 mm. ; floral bracts 1-5 mm., from one-quarter to one-half a,a long as their pedicels. Fknner diameter (7-) 10-12 mm. Inner se@ ( 4 - g ) 6-8 mm. long. Ovules (3-) P 9 . Stanzens mostly dentate below the anthers; apex generally not produced, truncate. Berries mostly estylose, rarely with style 1-2 mm. long.

British Columbia, Washington, Arizona, Wyoming, Dakota, New Mexico, Oregon, Idaho, California, Mexico, Guatemala.

KEY 104, to Sect. Aquifoliatae (Maps 48 and 49) Leaflets suborbicular. 565. M . amplectens Leaflets not suborbicular.

Leaves with only one pair of leaflets. Leaves with 2-4 (-6) pairs of leafleh.

Fruit 15 mm. long, ovoid-attenuate.

581. M . eutriphylla

577. M . incerta

Page 332: Berberis and Mahonia : A taxonomic revision

332 L. W. A. ~ E N D T

KEY 104 (con$.) Fruit 6-10 mm. long, broadly ovoid to subglobom, not attenuate.

Stamens edentate. 580. M. anpattfolia Stamens dentate.

Style 1-2 mm.; prophylls towards base of pedicel. Ledeta broad, 1 : 2-2.7; margins with 10-12 h e teeth. Ledeta narrow, 1 : 3-4; margins subentire.

584. M . grad& 583. M . 8ubintegrifolia

Style absent ; prophylls either absent, or situated at mid-point of pedicel, or tLt the apex of the pedicel a p p d to the flower.

Ledeta lustrous above. Leaves with 2 pairs of leafleta; pedicels 2-6 mm., with prophylls at their

Leaves with (2-) 3-6 pairs of ledeta; pedicels 6-10 mm.; prophylle absent, 582. M . quinqwfolh mid-point.

or at mid-point of pedicel. Ovulea 3-6.

M e t e rigidly c o r i m u s ; prophylls at mid-point of pedicel.

Leaflets thinly coriaceous; prophylls absent. 578. M . vohnia

573. M . Iyiz"Tiana (Key 105)

566. M . t o l - d 567. M . dictyota

571. M . aquifolium; 572. M . 8092nei;

Ovules 6-7; petals, 4 mm., shorter than inner sepals. Ledeta narrow, 1 : 4; margins with 6-12 teeth. Ledeta broad, 1 : 1-75; margins with 3-4 teeth.

Ledeta dull above. Ledeta very broad, 1 : 1-16, with rounded apex; prophylls a b n t .

Ledeta narrower, 1 : 1.7-3, with acute apex. 568. M . repens

Ovulea 6-8. Prophylls present, a t mid-point of pedicel; ledeta h e l y serrate.

Prophylls absent ; ledeta oarsely dentate.

Shrub &6 in. high; prophylls absent. Shrub 6-12 ft. high; prophylls present.

of pedicel; stamens apiculate.

pedicel; stamens truncate.

569. M.purnila 570. M . califmica

574. M.nuna Ovules 3-6.

Ledeta broad, 1 : 1.6-2, in 1-2 pairs; prophylls from middle to base

Ledeta narrower, 1 : 2-3, in 3-5 pairs; prophylls a t mid-point of 579. M . schiedeana

576. M. pinndat

Subaect. Euaquifoliatae Ahrendt, subsect.nov., foliis jugis (1-) 2-4 (-6) foliolorum, jug0 infirno aliis simili supra basim rhachidis (1-) 2-5-6 cm. inserto; racemis densi- 0oratis fasciculatis 2-7; prophyllis nullis (M. pmila , M. pin&, prophyllis v e r s ~ medium pedicellorum, excepta) ; staminibus dentatis (M. amplectente excepta); fructibus glauco-pruinosis, estylosis.

British Columbia, Washington, Arizona, Wyoming, Dakota, New Mexico, Oregon, Idaho, California.

KEY 105, to Subsect. Euaquifoliatae (Map 47) Leafleta orbicular, or suborbicular, 3-6 cm. broad. Ledeta not orbicular, 1-3 (-6) cm. broad.

565. M . amplecten.3

ovules (6) 6-9. Ledeta narrow (1 : 4) with 6-12 teeth to each margin. Ledeta broader (1 : 1-2-2) with 3-5 (-8) teeth to each margin.

566. M . toluuce&8

Ledet margins with coarse spinose (3-4 mm.)-teeth (3-10 mm.) Margins with 3-4 teeth, upper surface very lustrous with open, indistinct re-

ticulation; racemes 4-7 cm. 567. M. dictyota

t See Key 1050.

Page 333: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 333 &* with 8-12 teeth, upper surface dull with conspicuous reticulation; 570. M . ca l i fo t -nk~

568. M.repens

569. M.purnila

racemes 2 . 5 5 cm. Leaflet margins with finer spinose (1-3 mm.)-teeth (1-3 mm.), or entire.

W e t s with apex rounded; prophylls absent. Leaflets with apex acute; prophylls present at mid-point. of pedicel.

Ovules 3-5. Pedioels without prophylls.

Ledeta distinctly lustrous above.

longer than inner sepals. M e t s , below, epapillose (or almost so), and subconcolorous green; petals

M e t s , below, papillose and pruinose grey; petals shorter than inner sepals. 571. M . aguifolium

572. M . mnnei 573. M . piperiana

Leaflets greewh grey below, with 12-16 teeth to each margin. Leaflets whitish grey below, with 7-10 teeth to each margin.

Leaflets dull, or scarcely lustrous, above; petals shorter than inner sepals. Leaflets distinctly spinose-serrate, not convolute.

Leaflets always green. Shrub 4-6 in. high. 574. M. nana Shrub 2-5 ft. high. 575a. M. x wugneri

Ledets, at first creamy and pink in colour, Snally red. 575c. M. x m s e r i

Leaflets plane, entire. 575d. M. x herveyi Leaflets conspicuously convolute. 575e. M. x convoluta

575 f . M . x undulata

576. M . pinnutut

(565) Mahonia amplectens Eastwood, Proc. Cd$. A d . Sci. (ser. 4), 20, 145, 18 Dec.

Shrub 1-2 ft. Leaves with 3 pairs of leaflets overlapping at their bases, the lowest pair 4 cni. above the base. Leaflets suborbicular, 3-5 cm. wide, except for the terminal leaflet ; marginal serrations distant 5-10 mm. ; veins glaucous. Otherwise unrecorded.

California (S.E.): San Bernardino County, Santa Rosa Mts., Nov. 1904, no flowers or fruit, Trask 66 (Type, Herb. Calif. Acad. Sci., no. 81281). Not in cultivation.

(566) Mahonia toluacensis (Bean) Ahrendt, comb.nov.; syn. B. toluacemis Hort.,

Shrub 4-6 ft. Leaves to 16 x 9 cm., with 3 pairs of remote leaflets; their internodes decreasing from 4 to 2.5 cm. ; the lowest pair subequal to the others, situated 2.5-3.5 cm. above the base. Leajlets thin, lanceolate or narrowly ovate, all subequal, 5-66 x 1.2- 1-7 cm., or to 7 x 2 cm.; terminal leaflet similar with a petiole 1.3-2-5 cm.; the lowest two pairs with a petiole 5-15 mm., but the upper pair sessile or with a petiole only 1-3 (-5) mm.; margins 7-10-spinose (1.5-2 mm.)-serrate (2-5 mm.), distant 5-12 mm., undulate ; apex acuminate ; both sides closely reticulate, above impressed, below elevated ; above very lustrous; below paler yellow-green, lustrous. Racemes 1-2.5 cm., fascicled c. 4; bracts ovate, acute, 5 x 3 mm. Pedicels 7-10 mm.; floral bracts pale green, ovate, acute, 3.5 x 1-5-2.5 mm. Flowers H.C.C. 2. Prophylls nil. Outer se@ 2-2.5 x 1-1.5 mm., ovate, acute. Median sepals 4 x 2 mm., oblong. Inner sepals 6.5 x 3 mm., obovate-sub- spathulate. Petals 5 x 2.5 mm., oblong-ovate. Stamens 3 mm., conspicuously lanceolate- acuminate-dentate; apex produced but very truncate. Ovules c. 8. Berries pruinose blue, estylose, ovoid, 5-6 x 3 4 mm.

Leaflets convolute or entire.

Pedicels with prophylls at mid-point. Leaflets above very lustrous with venation scarcely visible. W e t s above dull, or scarcely lustrous, with venation openly reticulate.

1931

Gard. Chron. p. 435, 1868; Bean, Trees and Shrubs, 3, 52, 1933; 2, 294, 1950

California : Los Angeles County, Toluca. Cultivated: L.A. 737; fl. 21 May 1942; fr. Aug. 1942 (BM).

t See Key 1 0 5 ~ .

Page 334: Berberis and Mahonia : A taxonomic revision

334 L. W. A. AHRENDT

(567) Mahonia dictyota (Jepson) Fedde, Bot. Jb. 31, 89, 16 Aug. 1901; syn. Berberb

Leaves with (2-) 3 pairs of contiguous or overlapping leaflets, the lowest pair being 1.5-5 cm., above the base of the rachis. Leaflets ovate, subequal, c. 4 x 2.5 cm.; margins 3-4-spinose (3-4 mm.)-dentate (3-6 mm.); above very lustrous, openly or in- distinctly reticulate ; below dull with conspicuous closely elevated reticulation. Racemes 4-7 cm., fascicled c. 5; bracts of the inflorescence c. 5 mm. long. Pedicels glabrous, 5-7 mm.; floral bracts 1-2 mm. long. Plowers c. 7 mm. diameter. Prwhylls absent. Outer sepals 2.2 x 1-25 mm., ovate. Inner sepals 4.5 x2-5 mm., obovate. Petals 4 x 2.25 mm., emarghate. oencles (4-) 5-7. Berries ellipsoid, or ovoid-ellipsoid, estylose, pruinose blue, 9-11 x 5-7 mm.

Arizona: Pima County (N.E.), Santa Catalina Mts., fr. 4 Aug. 1914, Rehder 266 (K); vicinity of Tucson, fl. 15 Apr. 1908, Rose 11794 (K).

California: Yuba County, Marysville, Jepson, s.n.; Los Angeles County, San Diego, Palmer, s.n. Not cultivated.

dictyota Jepmn, BwU. Torrey CI. 18, 319

(568) Mahonia repens G. Don, Gen. Syst. 1, 118, 1831 Shrub stoloniferous, to 2-3 ft. Leaves 15-27 cm. long with (1-) 2-3 pairs of leaflets,

the lowest pair being (1-) 2.54 (-6) cm. above the base. Leuflets ovate, similar, sub- equal; base rounded or subcordate; apex rounded or obtuse, rarely subacute; above dull; below papillose, at first slightly pruinose, ultimately epruinose or subpruinose, paler green; above fairly closely but not very distinctly veined; below less distinctly veined. Racema 5-8 cm., fascicled 5-7. Pedicels 5-8 (-11) mm.; floral bracts 2.5 x 1.5 mm. Prophylls absent. Outer sepals 2 x 1 mm., ovate. Median s e p h c. 4 x 2 mm., oblong. Inner se@ 7x4mm., obovate. Petals c. 6x3mm., oblong-obovate; apex deeply incised, with two acute lobes. Ovules 6-9. Stamem dentate; apex not produced, trun- cate. Berries globose, 8-9 x 6-9 mm., estylose, pruinose blue.

IiEy 105a Leaflets (&) 5-6 (-7) x ( 2 . 5 ) 3-4 (-5.5) em., ovate; margins undulate, distinctly 5-9-

Leaflets 5-8 x 4-6 em., broadly ovate; margins not undulate, entire or with few incon- spino~e (2-3 emate ate (2-4) -. d k h t 8-15 -. 568a. bf. repma vm. repem

spicuous spinulea. 568 b. M. repem var. rotundifolics

(568a) Mahonia repens var. repens Colorado: Custer County, 8000ft., fr. 1891, Cusack 501 (0); 4800ft., a rare plant,

19 Aug. 1903, Copeland 3810 (K); according to Jepson, in the Rocky Mountains. Jepson cites the following much wider distribution : British Columbia, Washington,

Oregon, Nevada, Utah ; and North California (Del Norte, Siskiyou, and Modoc Counties), stretching southwards (into Lassen, Plumes, Sierra Nevada, Placer, Alpine, Mono, and Inyo Counties). Fedde cites, also, Wyoming, Dakota and New Mexico.

Cultivated: L.A. 38A, fl. May 1933, May 1935; fr. Aug. 1935.

(568b) Mahonia repens var. rotundifolia Fedde, Bot. Jb. 31, 82, 16 Aug. 1901 Leaves 12-22 x 5-13 cm., with leaflets in (I-) 2 (-3) pairs, approximate, contiguous or

overlapping; the lowest pair 3-7 cm. above the base; internodes decreasing from 4.5 to 3.5 cm. Leaflets similar, subequal; margins entire, or inconspicuously 6-12-spinose (0.5-1 mm.)-serrulate (0.5-1 mm.); above dull; below paler, papillose, pruinose, greyish, finally greenish; terminal leaflet 8 x 6 cm., or 7.5 x 7 cm. Pedicek slender, 7-11 mm.

Cultivated: a t Wisley, fl. May 1938. This variety is of cultivated origin.

Page 335: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 335

(569) Mahonia pumila (Greene) Fedde, Bot. Jb. 31, 82, 1901 ; syn. B. pumila Greene,

&hrub scarcely up to 1 ft. high. Leaves 8-14 cm. long, with 2-3 pairs of separate leaflets; basal pair similar to others (1-) 3-5 cm. above the base; internodes decreasing from 2-5 to 1-5 cm. Leajtets at first red, then purplish, finally deep grey green, all similar; from base to apex slightly increasing, 2.5-5 x 1.5-243 (-3) cm.; terminal leaflet larger, with petiole 1-1.7 cm.; base broadly cuneate to subtruncate; margins (4-) 5-8 (-10)- spinose (1-1-5 mm.)-serrate (1 mm.), distant 3-8 mm.; apex acute; above dull, with indistinct branched venation ; below grey-green, scarcely pruinose, sparsely papillose, openly reticulate. Racemes 2 4 cm., fascicled c. 5; bracts of the inflorescence 3-6 mm. Pedicels 5-10 mm.; floral bracts 1-2 mm. Prophylk a t mid-point of pedicels, ovate, acute, 2.5 x 2 mm. Outer sepals 3 x 1-75 mm., ovate, acute. Median sepals 5 x 3.25 mm., obovate, acute. Inner sepah 8 x 4.5 mm., oblong-obovate with base clawed. Petals 6-5 x 3 mm., oblong elliptic; apex rounded but deeply incised (1.5 mm.), with obtuse lobes. Stamens 4-5 mm., dentate; apex not produced, rounded-truncate. Ovules (5-) 6-43. Berries estylose, globose-obovoid, 9-10 x 8-9 mm.

Oregon: Josephine County, east base of coast Mts., near Waldo, fl. 27 Apr. 1892, Howell 1405 (K); dry ground, summit of Grant’s Pass, Crescent City Road, 3 July 1918, Peck 8095 (K).

Jackson County, Siskiyou Mts., 15 Aug. 1925, Thompson 441 (K). According to Jepson along the inner north coast range and higher Sierra Nevada foothills, from north California to south of Oregon, 3000-5000 ft.

Pdtonia, 2, 161, 1891

Lake County (Type). California : Siskiyou County, Siskiyou Mts., adjacent to Oregon (Josephine and Jackson

Cultivated: L.A. 739, fl. 29 Apr. 1942, 4 Apr. 1943; fr. 28 July 1954. Stoloniferous. Counties).

(570) Mahonia californica (Jepson) Ahrendt, comb.nov.; syn. B. californicu Jepson, Fl. Calif. p. 549, 1922

Shrub 3-6 (-10) ft. Stems rigidly erect, little branched. Leaves 4-10 cm. long, with 2 4 pairs of leaflets, subequal. Leaflets 2.5-5 cm. long, very pale or greyish, and con- spicuously reticulate; margins with 8-12 very coarse spreading aristate teeth, whose depth is from one-third to two-thirds the breadth of the body of the blade which is very undulate. Racemes densely many-flowered, 2.5-5 cm. long. Pedicels 6-10 mm.

California (west central): Solano County, Pellego Hills, in the north-west of the county, Jepson, s.n. (Type). According to Jepson, on dry rocky interior foothills, 300-1500 ft., or up to 6000 ft., on desert ridges, of the inner coast range south to Cis- montane, south California, and apparently also in the south Sierra Nevada foothills (east of Fresno county). Not in cultivation. The coarsely-toothed leaves are like some species of Sect. Horridae which stretches into California, but the inflorescence is distinct (see Key 110).

(571) Mahonia aquifolium (Pursh) Nutt., Gen. A m r . PI. 1, 212, 1818; syn. Berberis aquifolium Pursh, PI. Amer. Sept. p. 219, 1814.

Shrub 3-6 ft. Leaves with (2-) 3 4 (-6) pairs of leaflets; the lowest pair 2 - 5 4 cm. above the rachis. Leajets ovate, subequal, 4-8 x 2 4 cm.; margins with 6-12 teeth, distant 5-12 mm. ; above very lustrous with conspicuous close impressed reticulation ; below dull, a t first slightly greyish becoming yellow-green and sublustrous, epapaose. Racemes 5-8 cm., fascicled 3 4 . Pedicels 5-8 (-10) mm. ; floral bracts %4 mm. Prophylls absent. Outer sepals 1.5-2-5 x 0.75-1.5 mm., ovate, acute. Median sepals 4-5.5 x 3 4 - 5 mm.,

Page 336: Berberis and Mahonia : A taxonomic revision

336 L. W. A. AHRENDT obovate. Inner sepah 6-8 x 2 6 - 3 5 mm., oblong-obovate. Pet& 7-9 x 3 4 mm., nar- rowly oblong-obovate, apex acutely emarghata and biapiculate. Stamens 5.5 mm., dentate, apex not produced and truncate. Ovules (3-) 4-5. Be& globose, estylose, conspicuously pruinose blue.

~ E Y 1 0 5 ~ Leaflete coarsely toothed, spinose (2-3 mm.)-dentate (3-4 mm.).

M e t e finely toothed, spinom (0-26-0.6 I n 1 n . ) - W d 6 b (0.25-0-6 mm.).

(571a) Mahonia aquifolium var. aquifolium British Columbia: Great Rapids, rich soil, among rocks, 11 Apr. 1806, Lewis, 8.11.

(Type, Herb. Philadelphia A d . Sci.; photograph in Herb. Hew). Selkirk and Rocky mountains, half a mile above Carbonate, 10,OOO ft., 10 June 1906, Shaw 208 (K). Washington: San Juan Islands, Brown's Island, 25 J m e 1917, Zeller 846 (K). Lewis County (west): Caacade mountains, upper valley of the Nesqudy (i.e.

c. 50' 51" N., 118" W.), 8 May 1895, Allen 135 (K). Oregon: Wasco County, H d river, Whited 1068, fl. 7 Apr. 1924 (K); Tolmie, ex

Herb. Hook. (K). Cultivated: p h t s under this name are some of the most commonly grown shrubs in

England. They vary in leaf from very lustroy to quite dull, and from coarsely to finely toothed; and the leaflets vary considerably in breadth. It seems apparent that this cultivated population, grown from garden seed, consists largely of w u a l hybrids, probably mainly between M . aquifoium and M . repem. Certain distinct forms have received informal garden namm as varieties of M. aquifolium; as, however, these all differ in having petals shorter than inner sepals, they have been treated as distinct hybrids under number 575. Also in cultivation ie M . pinnatu hom California. The dis- tinctions between this and M. aquifolium are at h t sight obscured in cultivation owing to the hybrid variation mentioned. From the true M . aquifolium, it is at h t sight distinct in its dull more finely serrate leaves; also, it has petals shorter than inner sepals, but is most M y distinguished by the prophylls at the mid-point of the pedicels; see under (576), M . pinnatu.

571a. M . aquifolium var. wpifolium

571 b. M . aquifolium var. lyallii

(571b) Mahonia aquifolium var. lyallii Ahrendt, var.nov., foliolis marginibus spinulis dentibusque gracilibus.

Oregon: Casca.de mountains, boundary commission, 1859, Lyall, 8.n. (Type, K). Seeunder (557),M. f0TtUWiforhf. x heterophyMaScbneid. =M.aquifolium x M . fortunei.

(572) Mahonia sonnei Abrams in Phytologia, 1, ii, 92, July 1934 Leu$& lustrous bright green above, below greenish and papillose, margins with 12-16

teeth. Racemes with bracts 2-5 mm. Sta- dentate. Be& oblong. Otherwise un- recorded, but presumably as in M. aquifolium.

California: Nevada County, rocky banks of the Truckee river, 11 Aug. 1884, Apr. 1885, Sonne 11 (Type, Sheet 95828, Dudley Herb.). Not cultivated. See under (576), M. pinnata.

(573) Mahonia piperiana Abrams, 1.c.

Otherwise as in M. 8Wnei.

10176 (Type); same locality, fr. 1927, Abrams 12136.

Leu$& lustrous bright green above; below grey, papillose; margins with 7-10 teeth.

Oregon: Jackson County, near the Siskiyou summit, 9 Mar. 1924, Abrams & Benson

Page 337: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 337

General distribution, usually on thinly wooded slopes, arid transition zone. Oregon : Jackson and Josephine Counties to California : Mendocino County and north

of Lake County. Not cultivated. See under (576), M. pinnuta.

(574) Mahonia nana (Greene) Fedde, Bot. J b . 31, 83, 16 Aug. 1901; syn. B. nana

Shrub to 1 ft. Leaves with c. 3 pairs of leaflets, the lowest about 5 em. above the base. Leaflets ovate, fairly equal, 6-5 x 3 em.; base rounded; margins 4-6-spinose (1-2 mm.)- serrate (1-2 mm.); apex acute; above dull or scarcely lustrous; below grey-green. Racemes 6 5 em. long, with stout rachis (1-1-5 mm. thick), fascicled 2-3. Pedicels fairly stout, 6-10mm.; floral bracts 3mm. Ovules 3-5. Berries globose, c. 9mm., estylose, pruinose blue. Seeds purple.

Colorado (west central) : Gunnison watershed, 7720 ft., fr. 10 Aug. 1901, Baker 764 (K). (Specimen determined by Dr Greene.)

East Washington: Spokane County, Hangman Creek, 1510 ft., 25 May 1893, Sandberg & Leiberg 73 (K). The leaflets of this more western specimen, in 5 pairs, have up to 13 teeth to each margin, and it has slender pedicels 10-15 mm. long; it may be compared with M. aquifolium.

Generally in the Rocky Mountains from Idaho and Montana in the north, southwards to Arizona and New Mexico, according to Pedde.

The hybrid forms which have been named in gardens are as follows.

Greene, Pittonia, 3, 98, 1891

( 5 7 5 ~ ) Mahoniaxwagneri (Hort.) Ahrendt, nommud., R.H.S. Dict. Card. 3, p. 1235,

Leafrets in 3-5 pairs, fairly narrow, 1 : 2-3; margins with 4-10 teeth. Ovules 4-5. Stamens dentate. Berries globose, 6-8 mm., eatylose, pruinose blue (all as in both parent species). h$et.s dull and petals shorter than inner sepals (as in M. pinnatu). Pr0iphyU-s absent (as in M. aquifolium). Distributed as M . aquifoliumxkt. pinnata, which this plant appears to be.

Nov. 1951, foliolis opacis ut in M. pinnutu, prophyllis nullis ut in M. aquifolio

Cultivated: L.A. 741, fl. May 1942; fr. Aug. 1942.

KEY 105c Leafleta dull, reticulate Leaflets sublustrous, scarcely reticulate

575a M . x wagnerri 575 b M . x aldenhamemia

(575 b) Mahonia x aldenhamensis (Hort.) Ahrendt; syn. 1El. quifolium var. dden-

Cultivated: fl. a t Kew, 24 Apr. 1944. This is very close to M. wagneri and may well be hamensis Hort.

another hybrid between the same species.

(575c) Mahonia x moseri (Hort.) Ahrendt, nom.nud., R.H.S. Dict. Card. 3, p. 1234, Nov. 1951; foliolis primo pallide luteis et pallide rubris, ultimo rubris; pe t ah sepalis interioribus brevioribus

Cultivated: L.A. 55, fl. May 1935, May 1944; G. Aug. 1944. A highly ornamental hybrid.

(575d) Mahonia x herveyi (Hort.) Ahrendt; syn. M. aquifolium var. rotundifolk herveyi Hort. ; foliolis opacis, latis, apice obtusis; pe t ah sepalis interioribus brevioribus

Leaflets dull, and broader and with apex more rounded than in M. aqugolium. Pro- phylls absent. Outer sepals prophyll-like, 2 x 1.5 mm., ovate, acute. Median sepals

22 JOURN. LINN. SOC. BOTANY-VOL. LVII

Page 338: Berberis and Mahonia : A taxonomic revision

338 L. W. A. AERENDT 3 x 1-5 mm., oblong, obtuse. Petals 5.25 x 2.5 mm., narrowly obovate, mutely emarginate with rounded lobes. Ovules 4-5. Stamem 3-75 mm., dentate; apex truncate.

Inner s e p k 8 x 3-5 mm., narrowly obovate.

Cultivated: 0. Kew, 2 May 1947. Fedde, 1.c. p. 82, records Berberis repens var. rotundifollia krveyi (Hort.) Desf., which

he identifies with M a h i a repem var. rotundifollia.

(575e) Mahonia x convoluta (Hort.) Ahrendt ; syn. M. aquifolium var. convoluta Hort. ;

Leaflets dull, margins revolute and h e l y sparsely spinulose or entire, conspicuously

Cultivated: fl. Kew, 24 Apr. 1944.

foliolis opacis, marginibus revolutis, longitudine convolutis

convolute.

(575 f ) Mahonia x undulata (Hort.) Ahrendt ; syn. M. aquifolium var. undulata Hort. ; foliolis valde nitidis, parce nervatis, pedicellis prophyllis versus medium eorum

Leujets very lustrous, and very slightly veined. Prophylk present at mid-point of pedicels.

In the lustrous character of the leaflets and the presence of the prophylls this is inter- mediate between M. quifolium and M . pinnata (which possess respectively the former and the latter of these characters) in a way reverse to M. x wagneri which combines the opposite features (namely, respectively, the lack of prophylls and the dull leaves of these species). However, the scarcely veined leaflets differ from the reticulate ones of both species.

(576) Mahonia pinnata (Lag.) Fedde, Bot. Jb. 31,86,1901; syn. B. pinnuta Lag., Eklech. Hort.Ma&t.,6,1803; 14,1816; nonKunth; syn.Y. fmcic&ri.sDC.,Xyst. 2,19,1821

Shrub 8-10 ft. high. Leaves with 3-5 pairs of approximate or contiguous leafiets, the lower leaflets being 1-3 cm. above the base of the rachis, then with their internodes decreasing up the rachis from 4 to 1.5 cm. Leujets subequal; (3-) 4-5 (-6) x 1.2-1.4 em.; base broadly cuneate; margins undulate, 4-6-spinose (1-2 mm.)-serrate (2-3 mm.), dis- tant 3-7 mm.; apex acute; above dull; below paler, greyish, pruinose; both sides reti- culate. Racemes 4 4 cm. long, fascicled c. 5. Pedkls 3-7 (-10) mm. long, with floral bracts 1.5-2-5 mm. ProphyUs situated towards the mid-point of pedicel, ovate, 1.5-2 mm. long. Outer sepals 3-3-5 x 2-25-2.5 mm., broadly ovate. Median sepah 5 x 2.75 mm., obovate. Inner sepak 6 x 3 mm., oblong. Petals 5-5 x 2.75 mm., oblong-obovate; apex rounded and acutely incised with rounded lobes. Stamens dentate; apex not produced, truncate. Ovules 4 4 . Berries subglobose, 6x6mm., estylose, pruinose blue. Seeds purple-brown.

California: Monterey County, Monterey, type locality; 1828, Hartweg 1643 (K).* Much confusion has surrounded the plant populations and names associated with this

specific epithet pinmta. (1) In 1803, Lagasca set up the binomial B. pinnata, with further reference to it in

1816. Details are meagre, but it is clear that he refers to a ‘Berberis’ with pinnate leaves, i.e. a M a h i a , that its leaflets are spinose-toothed, that it was growing in the Madrid garden, from which it was typified, from seed gathered by D. L. Nde, with the record that its habitat is around Monterey and Nutka. There is a Monterey in the Nuevo Leon State of Mexico, and a Monterey County (and town) in California.

* Monterey: J. D. Hooker and A. Gray, s.n., 1877; 1853-4, near the 35th parallel of latitude, Bigelow, s.n.,; 1833. Douglas, 8.n. The leaves have become green below in some of these specimens.

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Berberis and Mahonia 330

(2) In 1821, Kunth used the same binomial, B.pin&, for a plant figured and briefly described, and clearly stated to come from Moran in Mexico; this is in the Hidalgo State. Among other characters, the figure shows the leaflets to have 8-10 spinose- teeth to each margin, the inflorescence to consist of loose, long, racemes, either solitary, or occasionally fascicled 2-3; and the fruit to be ellipsoid.

(3) Also in 1821, De Candolle used the binomial M . fascicularis as a synonym for 3. pin& Lag.; in doing so, he cites as habitat ‘N. America, around Monterey and Nutka (N6e ex Lag.), and in New Spain near Moran’. His description states that the leaflet margins have 6 6 spreading teeth, and that the racemes are dense and crowded, their pedicels bearing prophylls. But he suggests that ‘the Mexican plant from a Spanish garden differs in its fewer (2-3) racemes which are pendulous and not erect, and may be a quite different species’. Thus he throws doubt on the identitkation of B. pinnatu Lag., with B. pinnata Kunth. But the phrase ‘the Mexican plant from a Spanish garden’ is confusing; it may imply that B. pinnata Kunth was thought to have been based on a different plant of Lagasca’s; it could imply that Lagasca’s plants included also the Mexican one-but against this is the clear statement that De Candolle means the N. American Monterey in connexion with the plants of Lagasca.

(4) In 1823, Sims, in Bot. Mag. t. 2396, figures a B. fasciculuris which he regards as synonymous with the three names already cited, i.e. with B. pin& Lag., B. pinnata Kunth, and M . fasciculuris DC. Sims’s figure is based on a plant which flowered in Feb. 1823 a t Boyton, where it was grown by A. B. Lambert, who had raised it from seed received from the Spanish botanist Lagasca. Sims deduces that ‘ obviously this plant is his [i.e. Lagasca’s] species, and consequently De Candolle’s M . fasciculuris and a native of New Spain’. Sims repeats De Candolle’s query concerning the identity of Lagasca’s species with that of Humb. et Bonpl., ex Kunth, from Mexico with ‘the fewer racemes described and figured’. Sims then suggests that the plant he has figured was ‘thought possibly to be the same m aquifoliurn Pursh ’, but this is ‘under one foot high ’, whereas the plant ‘brought to the Madrid garden by M. N6e from New Spain is erect and P 5 ft. high’. Sims figure shows a plant with very coarsely dentate leaves and (with one ex- ception) without prophylls on the pedicels. The figure is somewhat indeterminate, and may well represent a plant of M . aquijolium, which is somewhat erect, and does grow 3-5 ft. high. In this matter, Sims may have been confusing M . aquifoliurn with M . nana or M . repens. It is quite possible, therefore, that Sims’s figure was drawn from a specimen of M . qui fol ium grown from that part of N6e’s collection which was gathered from Nutka-in British Columbia, the home of this species; on this hypothesis, the Madrid garden contained more than one species grown from N6e’s introductions. However, since Lambert’s Boyton plant was grown from garden seed from Madrid, it may well have been a hybrid, as are so many plants called M . aquifolium in present-day English gardens. Having seen no herbarium specimen of B. pinnata Lag., as interpreted by M . fascicularis DC., from British Columbia, it seems best to associate the Nutka citation with a collection of M . quifolium distinct from the Californian species from Monterey. Sims’s discussion and figure of B. fascicuhris is best ignored; in any case, it is not in the main line of development of this investigation, as it is intended to be merely a later and figured representation of M . famicularis DC., with a deliberate reversion to the generic name Berberis.

At this stage, out of the confusion it becomes clear that, apart from the likely collection of M . aquifolium from Nutka, we are dealing with: (a) Lagasca’s Californian species of 1803, from Monterey, and ( b ) Kunth’s Mexican species of 1821, from Moran, and that they were recognized as possibly distinct by De Candolle, in 1821.

(5 ) I n 1829, this distinction was explicitly accepted by the use of the binomial B. moranelwis Hebenstr. et Ludw., ex Schult., Syst. Veg. 7, 17, as a synonym for B. pinncLtu Kunth from Moran in Mexico, as distinct from B. pinnatu Lag.

22-2

Page 340: Berberis and Mahonia : A taxonomic revision

340 L. W. A. AERENDT (6) In 1901 Fedde used the generic name M a h i a for the Californian plant which

became M . pinnata (Lag.) Fedde. Fortunately the identity of the plant was made more certain by Hartweg's no. 1643, collected in Monterey, in 1828. Fedde, however, includes the Mexican localities in his citations.

(7) In 1922 Standley referred to a species from Guatemala which, he says, 'has hitherto been confused with M . fmcicdaris DC. which ranges from South California to Vera Cruz '. He thus, again, cites the Californian and Mexican localities as producing the same species, while separating the Guatemala plant as M . volcaniu.

In 1950, Johnston used the generic name M a h i a for the Mexican plant as M . nzoranensiS (Hebenstr. et Ludw., ex Schult.) Johnston, as Fedde had previously done for the Californian M. pinnda. But Johnston makes quite clear the geographical dis- tinction, stating that M . manens i s 'is the most widely distributed M a h i u of Central Mexico. Fedde and Standley make this identical with the plants of the Pacific coast and the U.S.A., which are B. pinnata Lag., 1803, and M. jaseicularis DC, 1821 ; but the name B. n w r a d is eamntially a renaming of B. pin& HBK, which is a (later) homonym of B. pin& La@;. It comes from Mexico only'.

In conclusion it may be well, h t , to record the geographical distinctions involved between these, and some closely related species, by considering them in order from north to south:

(i) British Columbia, Washington, San Juan Islan&, extreme north of Oregon (Wasco county), i.e. c. 45-51" N. lat., 118-123" W. long.-M. apifolium. (Part of NCe's col- lection and some of Lagasca's resulting Madrid plants, and Sims's figure of Mr Lambert's Boyton plant.)

[ii) Extreme south of Oregon (Jackson County), extending into the northern part of California on the western, sea float, side (Mendocino county, and north of Lake county), i.e. 3943" N. lat., 123-124" W. long.-M. pipwiana, closely related to M . aquqolium.

(iii) Northern California on the eastern, inland, side, (Nevada county) c. 39" 30' N.: lat., 120-121" W. lat.-M. sonnei, a h closely related to M . apijolium.

(iv) C o d Southern California (Monterey county), i.e. c. 35"-36" N. lat., 121" W. long.-M. p*nna&z, a speciea much more distinct from H. aguifdurn.

(v) Inland cantral Mexico, c. 21" N., 100" W.-M. mmawn-si.8. (vi) S.W. Guatemala, c. 1P16" N. lat., 91-92" W. long.-M. wlcania. Finally, to indicate the botaniical distinctions involved :

KEY 1 0 5 ~

571. M. aquifoliunt British Columbia to

573. M.piperianu i Key 105 572. M . ~onnei

Leaflets not very rigid, with spreading and fairly coarse spinose-serration, with spinules 1-5-2 mm. long, and teeth 1 - 5 3 mm. deep; racemes 4-10 cm. long.

Ledete grey below, dull above, the lowest being 1-3 cm. above base of rachis; racemes fascicled c. 5, erect, dense-flowered, P6cm. long, with short pedicels

North California. Pedicels without prophylls.

Pedicels with prophylls.

3-7 (-10) DXXL long; fnrit s~bglobose C. 6 x 6 IIM. p.Uino~e blue. 576. M . (South California)

Leaflets green below, lustrous above, with lowest one only 5-10 mm. above base of rachis; racemes solitary or fascicled 2-3, pendulous, loosely flowered, 7-10 om. long, with pedicels (0-17-) 1-2 cm. long; fruit ellipsoid, c. 8 x 6 nun. epruinose, black. 585. M . manens is (Mexico)

Leaflets very rigid, green below, lustrous above, margins spinulose-subsedate, the minute spinulea appressed to the margin; racemes c. 3 cm. long.

578. M. volcania (Guatemala)

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Berberis and Mahonia 341

Subsect. Schiedeanae Ahrendt, subsect.nov., foliis jugis 1-6 foliolorum, jug0 h h o aliis simili, supra basim rhachidis (1-) 2-7 em. inserto; pedicellis prophyllis prae- ditis, plerumque versus medium vel versus basim positis, sed interdum apice, flori quasi epicalyce appressis ; staminibus dentatis (M. angustijolia excepta) ; fructibus saepe distincte stylosis, interdum estylosis.

Mexico : Coahuila, Nuevo Le6n, San Luis Potosi, Querhtaro, Puebla, Hidalgo and

Guatemala : Esquintla and Huehuetenango Provinces. Oaxaca States.

KEY 106, to Subsect. Schiedeanae (Map 49) Fruit 15 111111. long, ovoid-attenuate. Fruit 6-10 mm. long, not attenuate.

577. M . incerta

Pedicels 1-7 (-10) mm.

the base. Ledets dull above; pedicels 5-11 mm., with prophylls situated from the middle to

Leaflets in 3-5 pairs; stamens truncate. Leaflets thin with fairly comm marginal serration; dull; apex acute.

Leaflets rigid with h e marginal serration; lustrous, apex obtuse. Cf. 576. M . pinnutat

578. M. volcania Leaflets thick, in 1-2 pairs; stamens apiculate. 579. M. schiedeana

Stamens edentate. 580. M. angustifolia Stamens dentate.

Leaflets lustrous above; pedicels 1-5 mm.

Leaves with only one pair of leaflets; prophylls appressed to the flowers; ovules 7-11. 581. M. eutriphylla

Leaves with 2-6 pairs of leaflets; prophylls from mid-point to base of pedicel; ovules (where known) 4-5.

Leaflets broad (1 : 1.3-2), in 2 remote pairs, margins with 7-10 teeth; racemes 7-11 cm. long; prophylls at mid-point of pedicel.

Leaflets narrower (1 : 2-4), in 2 subentire pairs or 3-6 overlapping pairs with serrate margins ; racemes 2-6 cm. long ; prophylls towards base of pedicel.

582. M . quinquefolia

Leafletu in 2 subentire pairs. 583. M. subintegrifolia Leaflets in (%) 3-6 pairs, margins with 10-12 teeth. 584. M. gracilis

Pedicels (0.7-) 1-2 cm. 585. M. muranenaiat

(577) Mahonia incerta Fedde, Bot. Jb. 31, 93, 16 Aug. 1901

Leaves with 3 4 pairs of leaflets, the lowest pair subequal to the others and situated 3-5 cm. above the base. St ipuh c. 5 mm. Leaflets ovate to lanceolate, 3 4 x 1-2 cm. ; base cuneate or subrounded, and subpetiolulate; margins with 5-10 spinose teeth; above deep lustrous green, or at leaat sublustrous; below dull, paler green. Racemes long; bracts of the inflorescence 2-3 mm. long. Pedicels 8-10 mm.; floral bracts 1-15 mm. Pr0phyJl.s 1 mm. long, situated a t mid-point of pedicel. Berries ovoid, with attenuate apex, 1.5 cm. long.

Mexico : Hidalgo State, between Real del Monte and Atotonilco et Chico, Ehrenberg 1288 (Type). Not cultivated.

(578) Mahonia volcania Standl. et Steyerm., ‘Studies of Central American Plants’, in

Shrub 3-6 ft., with few stems. Leaves with short petiole and 2-43 approximate pairs of leaflets. Leajlets rigidly coriaceous, sessile, ovate or oblong-ovate, 2 4 x 1-1.7 cm. ; margins finely appressed spinuloae-serrulate ; apex rounded or obtuse ; above lustrous

t See Key 1050.

Publ. Bot. Ser. Field Mu8. Nut. Hist, Chicago, 23, pt 1, 6, 14 Jan. 1943

Page 342: Berberis and Mahonia : A taxonomic revision

342 L. W. A. AHRENDT with pale prominent subreticulation ; below paler green with prominent reticulation. Racemes 3 cm. long. Pedicele slender, rigid, to 10 mm., with prophylls near mid-point. Outer sepah broadly ovate 2 mm. long. Petals 6-7 mm.

Guatemala: Esquintlii, Prov., pine forest, s l o p of Voldn de Agua, along the trail above Santa &ria de Jesbs, about 10,OOO ft., Feb. 1939; Standley 65221 (Type); same locality, Jan. 1937, Johnston 580.

Huehuetenango Prov., region of Chemal, ll,OOOft., Dec. 1940, Johnston 1690. Not in cultivation. See under (576), M. pinnatu.

(579) Mahonia schiedeana (Schldl.) Fedde, Bot. Jb. 31, 90, 16 Aug. 1901; syn. B.

h u e s with 1-2 pairs of leaflets, the lowest pair set (1-) 2-7 cm. above the base. Leaflets thick, rigid, broadly ovate, 2-3 x 1.5-2.5 cm.; base broadly cuneate; margins undulate, 3-6-spinose (2-3 mm.)-dentate (2-3 mm.); apex shortly acute; above dull grey- green; below dull, paler green. Racema fascicled 1-3, 1-2 (-3) cm. long; bracts of the inflorescence 3-5mm. long. Pedieels slender, 7-11 mm. long; floral bracts 1-1.5 (-2) mm. Prophylh lanceohte, acuminate, situated between the middle and the base of the pedicel. Outer s e p h 2 x 1 mm., ovate. Inner s e p h 4 x 2.25 mm., obovate. Petals 3-25 x 1.75 mm. Stamens c. 3 mm., apex produced. ovules 3-6. Berries 8-86 x 56-33 mm., ovoid, estylose, pruinose blue. Seeds purple brown.

Mexico: Mexico State, Sierra de las Cruces, a dense shrub, flattening against cliffs, 12,000 ft., fr. 24 Oct. 1895, Pringle 6218 (K).

Hidalgo State, Barranca above Santa Fey fl. 22 Mar. 1899, Pringle 6975 (K), shrub 8-12 ft.

Oaxaca State, between Guantotalapa and Tlachichucha, 1828, Schiede (Type). Nuevo Le6n State, Sierra Madre Mts., 40 miles south of Saltillo. Not cultivated.

schiedeuna Schldl., Bot. Ztg. 12, 654, 1854, non Dippel

(580) Mahonia angustifolia (Hartw.) Fedde, 1.c. p. 91,16 Aug. 1901 ; syn. B. anquati-

Shrub 10-12 ft. tall. Leaves with 2-4 pairs of leaflets, the slightly smaller lowest pair 1-2 cm. above the base. Lea$& oblong-oblanceolate, 10-20 x 7-5-10 mm. ; base swile, rounded; margins 5-7-spinose (1 mm.)-serrate (0.5 mm.), distant 3-6 mm.; apex acute; above sublustrous; below dull, slightly paler. Racemes 3-4 cm.; bracts 2 mm. Pedkls c. 2.5 mm. ; floral bracts 1-5 mm. ProphyUs lanceolate, situated towards the middle of the pedicels. Outer s e p h 2.5 mm. Inner sepals 5 x 3 mm. Petals 4 mm., emarginate with obtuse lobes. Stamens edentate. Berries subglobose.

Mexico: Hidalgo State, between Actopan and Pachuw, Haxtweg 270 (Type, K). Not cultivated.

f o l k Hartw., ex Benth., PI. Hartw., 34, 1840

681) Mahonia eutriphylla Fedde, I.c. p. 91,16 Aug. 1901; syn. M . trlifolia Cham. et Schlecht. MSS., in Herb.; syn. B. eutriphylh (Fedde) C . H. Muller, Arner. Midl. Nat. 27, 484, 1942

Leaves 6-11 x 4-5 cm., with only one pair of leaflets set (2-) 3-7 cm. above the base. Leafeta oblong-ovate, 3 x 1.2-1-8 cm., to 4 x 2.2 cm. ; base broadly cuneate; margins 5-6-spinose (2-3 mm.)-serrate (2 mm.), distant 6-10 mm.; apex acute; apex leaflet with a petiole 8-12 mm. ; above lustrous, closely finely veined and reticulate; below dull. with reticulation more open and more distinct. Racenzes 1-2 cm. long; bracts of the in- florescence 5 x 3 mm. PediceLs 1-14 (-3) mm. long; floral bracts 2 x 1 mm. ProphylLs 1-25 x 1-25 mm., at apex of pedicel appressed to the calyx. Outer s e e 1.75-2 x 1.25- 1.75 mm., ovate, acute. Median sepals 3-36 x 2-26 mm., elliptic. Inner sepals 6-7 x 2.75-3.5 mm., obovate-elliptic. Petals 5-6-5 x 3-3-5 mm., obovate-elliptic; apex emarginate with acute lobes; glands distinct. Sturnens 5 mm., dentate; apex slightly

Page 343: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 343

produced, truncate. Ovules 7-9. Berries black, epruinose, ovoid, 10 x 7.5 mm., excluding style 0.25-0.6 mm. long.

Mexico: Mexico State, Crucero to Agua Blanca, among rocks in pine woods, 15 ft. high, fr. 1 Dee. 1936, and fl. 5 May 1936, Hinton 8826 (K) ; fr. 2 Aug. 1936, Hinton 8902 (K).

Coahuiln State, Mun. de Cuatro Cienegas, Sierra de la Cailon del Agua, 9 Sept. 1939, in moist shady arroyos, specimen with only leaves and dried peduncles, Muller 3244 (U.S.N.A.).

( 2 ) Hidalgo State, ( 1 ) Kaukando bei La Encarnacibn, 1840, Ehrenberg 1109 (Type). Muller remarks that the 'citation of the type is not very illuminating'. Not in culti-

vation.

(582) Mahonia quinquefolia (Standley) Standley, Publ. Bot. Ser. Field Mus. Nut. Hist. Chicago 4, 205, 1929; syn. Odostemon quinquefolia Standley, Proc. Biol. SOC. Wash. 31, 133, 29 Nov. 1918

Leaves 9-15 cm., with stout rachis, and 2 pairs of sessile, subequal, remote leaflets, the lowest pair 3 4 . 5 cm. above the base. LeaJlets ovate to oblong-ovate, 3-5.3 x 1.5- 4 cm.; base rounded or broadly cuneate; margins with 7-10 teeth; apex varying from subacute to subobtuse; both sides lustrous green with prominent venation and fine reticulation. Racemes with stout rachis and peduncle, 7-11 cm. long. Pedicels 2-5 mm.; floral bracts small. Prophylls near middle of pedicel. Outer sepals small, acute. Inner sepals ovate, obtuse. Stamens dentate.

Mexico: Puebla State, on the Cerro Matzize, vicinity of San Luis Tultitlanapa, July 1907, Purpus 2775 (Type, U.S. Nat. Herb. 841065); Cerro de Paxtle, Purpus 2774 (with larger leaflets).

(583) Mahonia subintegrifolia Fedde, Bot. J b . 94, 16 Aug. 1901 Leaves clustered a t the apex of the stems, with 2 pairs of leaflets, the lowest pair being

0-5-1.5 cm. above the base of the rhachis. Lea$ets ovate-lanceolate, 20-30 x 5-10 mm. ; base cuneate; margins mostly entire, rarely with 1-5 minute spinules; apex acute or acuminate ; above lustrous green; below slightly paler, sublustrous. Racemes suberect, 2-3 cm. Pedicels 2-5 mm. ; floral bracts 1.5-2 mm. Prophylls 1 mm., similar t o the bracts, situated near tho base of the pedicel. Outer sepals very small. Median and inner sepals larger. Petals shorter than inner sepals and entire. Stamens dentate. Berries pruinose blue, with style 1-2 mm. long.

Mexico: Hidalgo State, Atotonilco, 20'15' N, 98"50' W, Ehrenberg 978 (Type).

(584) Mahonia gracilis (Hartw.) Fedde, 1.c. p. 95, 16 Aiig. 1901 : syn. B. gracilis Hartw.

Leaves with (2-) 3-5 (-6) pairs of contiguous or overlapping leaflets, the lowest being (2-) 2-5-36 (4) cm. above the base. Leajets ovate, 3 . 5 4 x 1.5 cm. to 5 x 2-3 cm.; margins 10-12-spinose (0.5 mm.)-serrate (0.6 mm.), distant 2 4 mm. ; apex acute; both sides finely reticulate ; above bright green very lustrous ; below sublustrous, subcon- colorous or slightly paler green; petiole of the terminal leaflet 1-1.5 cm. Racemes fascicled 1-3, dense, suberect (3-) 4-6 (-8) cm. long; bracts of the inflorescence 2-3 (-5) mm. Pedicels (1-) 2-3 (4, in fruit) mm. long; floral bracts 1 mm. Prophylls situated towards base of pedicel. Outer sepals 1.25 x 1 mm., ovate, acuminate. Median sepals 3.25 x 2 mm., oblong-obovate. Inner sepals 6 x 3.5 mm., oblong-obovate. Petals 4.5 x 2.6 mm., obovate, entire. Stamens below the anthers dentate ; apex conical-apiculate. Ovules (3-) 4-5. Berries pruinose blue, ovoid, 10-12 x 5.5-7 mm., excluding style 1- 1.5 mm.

Mexico: Nuevo Le6n State, Sierra Madre near Monterey, fr. 1 June 1899, Pringle 2530 (K).

ex Benth., P1. Hartw. 34, 1840

Page 344: Berberis and Mahonia : A taxonomic revision

344 L. W. A. AERENDT

Palmer 8. San Luis Pot& State, San Migue'lito, SchaEner 711 (K); without locality, Parry L%

Hidalgo State, Zimaph, Hartweg 271 (Type, K). Oaxaca State, 7700 ft., 1840, Galeotti 4641. South Texas: acwrding to Fedde. Not in cultivation. '

(585) Mahonia moranensis (Hebenstr. et Ludw.) I. M. Johnston, J . A m . Arb. 31, 190, 1950; syn. B. m a n e m i s Hebenstr. et Ludw., ex Schult. f., Syst. vq. 7,17, 1829; syn. B.pkataKunth in Humb. et Bonpl., Now. Gen., 5,71, t. 434,1821,non Lag. 1803; non M. pinnata (Lag.) Fedde

Leaves with 4-5 pairs of leaflets, the lowest. 5-10 (-40) mm. above the base of the rachis; subequal, including the basal pair. h$h% with base rounded; margins 8-10- spinose (0.5-1 mm.)-serrate (1-5-2.5 mm.), distant 5-10 mm.; apex subacuminate; re- ticulate; above lustrous light green; below paler green. Racenzes loose, pendulous, 7-10 cm. long, solitary, or fascicled 2-3. Pedieels slender, (0-7-) 1-2 cm. long ; floral bracts small. Prophylls situate towards mid-point of pedicel. Petals obovate, entire, scarcely shorter than the inner sepals. Bem'es broadly ellipsoid, c. 8 x 6 mm., with a broad sessile stigma; black, epruinose. (See Key 105 D.)

KEY 106a Leaflets fairly broad, 1 : 2.5, measuring c. 6.5 x 2-5 cm. pedicle~ 7-12 mm.

Leaflets narrower, 1 : 4 4 , measuring 5-6 x 1-1-5 cm. pedicel8 1-2 cm.

(585 a) Mahonia moranensis var. moranensis

585a. M . rnura& var. w a n e 4

M . mranensiS var. cachira 585 b.

Leaves with lowest pair of leaflets 5-10 mm. above base of rachis. Mexico: Hidalgo State, Moran (Type). ALSO, Barranca de Santa Fe, Schmitz 4814B. Querbtaro State, Querbtaro, 1829, Uhde 1048. Vera Cruz State, Orizaba, Linden 993. Apart from the type, the above citations are according to Fedde under M. pin&

which he confused with this species. Mexico State, Temascaltepec district, Las Cruces, fl., fr. 10,500ft. 8 Feb. 1933,

Hinton 3262 (K), a tree 30ft. high with ripe fruit black. Cnrcero, 84OOft., 17 Feb. 1934, Hinton 5591 (K). Hornos, in pine forest, 10 Feb. 1936, Hinton 8915 (K).

There are also specimens from M e s h Viejo, 6 Apr. 1932, Hinton 488 (K); and Com- unidad, 8000ft., fr. 9 May 1933, Hinton 3850 (K). Further, without locality, fl. 1848-9, Gregg 640 (K).

(585b) Mahonia moranensis var. cachira (Fedde) Ahrendt, comb.nov.; syn. Jf. pin- nub var. cachira Fedde, Bot. Jb. 31, 88, 16 Aug. 1901

Leaves with lowest pair of leaflets 3-4 cm. above base of rachis. Mexico: Mexico State, Toluca, Andrieux 542 (Isotype, K).

Sect. Paniculatae. Fedde, Bot. J b . 31, 71, 77, 16 Aug. 1901 Inflorescence simply, and generally loosely, racemose ; or paniculate. Mexico, Cuba, Guatemala, Costa Rica.

KEY 107, to Subsects. of Sect. Paniculatae. (Map 49) Inflorescence racemose. Inflorescence paniculate.

Subsect. LAXIRACEMOSAE Subsect. EUPANICULATAE

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Berberis and Mahonia 345

Subsect. Laxiracemosae Ahrendt, subsect.nov., inflorescentiis racemosis semper soli-

LeaJetsin 2-6 pairs; lowest pair (2-) 3-7 cm. above the base of rachis; margins entire or with up to 20 spinose teeth; reticulate; below epruinose, green. Ovules 2 4 . Style absent, short, or up to 1 mm.

tariis, plerumque laxis, non paniculatis.

KEY 108, to Subsect. Laxiracemosae Pedicels 2-4 nun. long; leaflets in 2-4 pairs.

Leaflets very thin, with spinulose margin; prophylls present; style short.

Leaflets thick, with entire margin; prophylls absent; style long.

Lowest pair of leaflets only 5-10 mm., above the base of the rachis; leaflets spinose;

Lowest pair of leaflets (2-) 3-7 cm. above the base of the rachis; leaflets entire; style

586. M . andrieuxii 587. M . chochoco

Pedicels 1-2 cm. long; leaflets entire, in %6 pairs.

style absent. Cf. 585. M . mranensis

distinct. Leaflets with apex rounded, or obtuse, rarely subacute. 588. M . longipes Leaflets with apex acute to subacuminate. 589, M . tenuifolia

The racemes of this subsection are generally very loose on account of their long pedi- cels. In the first two species, although the pedicels are short, the racemes are not dense, since the flowers are scattered somewhat sparsely along the elongated rachis.

(586) Mahonia andrieuxii (Hook. et Am.) Fedde, Bot. Jb. 31, 103, 16 Aug. 1901;

Leaves 8-15 cm. long, with 3 pairs of leaflets, the lowest pair set 2-3 cm. above the base. Lea$ets very thin, 3-5 x 1-1.5 cm.; margins 15-20-spinose (0.5 mm.)-subserrulate, distant 2.5-6 mm.; above slightly lustrous dark green; below paler; both sides openly reticulate. Racemes 4-10 cm. Pedicels 2 4 mm. ; bracts 0.5 mm. Prophylls set towards middle of pedicel. Outer sepals c . 3 mm. Inner sepals c. 4.5 mm. Petals c . 3.75 x 1-75 mm., entire. Stamens dentate. Ovules 2-3. Style short.

syn. B. andrieuxii Hook. et Am., Bot. Beech. Voy. p. 318, 1841

Mexico: without locality, 1883, Andrieux 469 (Type, BM). Not cultivated.

(587) Mahonia chochoco (Schldl.) Fedde, 1.c. p. 103, 16 Aug. 1901 ; syn. B . chochoco Schldl., Bot. Ztg. 12, 652, 1854

Shrub 20-35 ft. Leaves 7-15 cm., with 2 4 pairs of leaflets; oblong-elliptic lowest pair 3-5 cm. above the base. Stipules nil. Leaflets thick, 4-6 x 1-24 cm.; base cuneate; margins undulate, completely entire ; apex obtuse, rounded, or slightly retuse ; both sides closely reticulate ; above very lustrous dark green ; below slightly paler, sublustrous. Racemes loose, 6-8 cm. long; bracts of the inflorescence 4-5 mm. long. Pedicels 3 4 mm. ; floral bracts 1-15 mm. ProphyUs absent. Outer sepals broadly ovate, subacurninate. Median sepals broadly ovate. Inner sepals elliptic. Petuls obovate, entire, slightly shorter than inner sepals. Ovules 4. Berries ovoid, 6-9 mm., excluding style 0.5-1 mm.

Mexico : Hidalgo State, by Minoschtla, Tschatschakala, Chochooala, 1840, Ehrenberg 1034 (Type, BM), Ehrenberg 1250; Palo, mountains, Dec. 1831, Berlandier 927.

(588) Mahonia longipes (Standley) Standley, Publ. Bot. Ser. Pield Mus. Nut. Hist. Chicago, 4, 205, 1929; Gentry, Brittonia, 6, 309, 1948; syn. Ohstemon kmgipes Standley, Proc. Biol. SOC. Wash. 31, 133-, 29 Nov. 1918

Leaves 20-30 cm. long, with slender rachis and 5-6 pairs of remote leaflets, the lowest pair being 4-55 cm. above the base of the rachis, the remaining internodes decreasing upwards from 4 to 2 cm. Leafrets thin, oblong-elliptic, their size decreasing from base to

Page 346: Berberis and Mahonia : A taxonomic revision

346 L. W. A. AHRENDT

apex, 5-25 x 1.6 cm.; baae cuneate, sessile; margins completely entire; apex obtuse, rarely subacute, mucronate; above dull pale green; below concolorous green, epruinose; both sides finely, fairly closely reticulate, though somewhat more coarsely below. R m w loose, 15-22cm. long, including peduncle 5-7cm. long. Pediceh slender, 1.5-2 cm. long. 0mZe.s 2. Berries obovoid, or subpyrifom, verypruinom, 10 x 6 - 6 5 mm. excluding a thick style varying from 0.3 to 1 mm. long. 8eed.s dark purple.

Mexico: Durango State, San Rambn, 21 Apr.-18 May 1906, Palmer 103 (Isotype, K). Not cultivated.

(589) Mahonia tenuifolia (Lindl.) Loud. ex Fedde, Bot. J b . 31, 105, 1901; syn. B. tenuifolia Lindl., Bot. Reg. 24, Nisc. 64, 1838; and Bot. Reg. 30, t. 26, 1844.

Shrub to 10 ft. Leaves with 3-6 pairs of leaflets, the lowest pair 5-7 cm. above the baae of the rhachia, the remaining internodes decreaaing upwards from (6-) 4-26 cm. LeccJEets thin, 6-10 x 1-5-2-5 cm. ; baae aubsesaile; margins undulate, completely entire; apex acute to subacuminate; above lustrous dark green; below paler yellow green; both sides closely reticulate. Rmnzes 10-30cm., very loose; bracts of the inflorescence c. 4 mm. Pedicels slender, 1-2 cm. long; floral bracts 1-16 mm. Prophylk at apex of pedicels appressed to calyx. oute7 sepals very small. Median s@ larger. Inner sepah 4-5-5 x 2-2-5 mm. Petds c. 3 mm., shorter than median and inner sepals. Stamens dentate. h b 2. Berries obovoid, 7-8 x 4 4 mm., excluding style 0.4 mm. long.

Mexico: Vera Cruz State, below Mt. Orizaba, near Zaquapan, 3000 ft., 1836, Hartweg, 262 (Type, BM); banks of creeks near Zaquaph, in moist localities, Sept. 1929, Purpus 13079 (K).

Cuba: fl. 1860-4, Wright 1855 (K). This species is probably the same as B. ( M a h i a ) frazinijoliu Hook., Ic. P1.4, t. 329-

330, 1841, which is briefly described but well figured, and is recorded as coming from Xalapa, collected by Galeotti.

Subsect. Eupaniculatae Ahrendt, subsect.nov., inflorescentiis paniculatia Lea$et.s in (2-) 3-8 (-13) pairs, reticulate, often petiolulate (14mm.). Prophylls

mostly present, and situated from the middle to the apex of the e c e l s . Stanzens mostly dentate, occasionally dentate. ovules, where known, 5-7. Fruit mostly estylose. Style, if present, short.

Mexico, Guatemala, Costa Rica.

KEY 109, to Subsect. Eupaniculatae Leaflets entire.

Leaflets in (3-) 4-7 pairs with apex acute or acuminate. Leavea 16-25 cm. long, with petiole 3-4 cm.; 1eaflet.e acute, subsessile; inflorescence

Leaves 40-80 cm. long, with petiole 9-15 cm. long; l d e t s acuminate, petioldate; formally panicdate with pedicels 6-10 mm. long. 590. M. ehrenbergii

inflorescence cymose-panicdate with pedicela 1618 mm. long. 591. M. chiape&

Leaflets in 2-3 pairs with apex rounded. 592. M. johnstonii

Leaflets broad (1 : 1-1-3), in 3-7 pairs; margins with (P) 6-20 spinulose teeth pedicels Leafleta with margins with (2-) 6 2 0 spinose teeth.

5-20 mm., from 2.5 to 6 times as long 88 their bracts. Leaflets with the loweat pair (1-) 1-6-23 cm. above the base of the leaf rachis. Stamens dentate; leaflets above dull, or very slightly lustrous.

Ledeta with apex acute to subacuminate. Leaflets fairly -OW (1 : 3), in 5-8 pairs; margins 12-20 finely spinulo~e-

serrulate; 6-13 cm. long, subconcolorous green below. Style present. Cf. 593. M. pan-

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Berberis and Mahonia 347

Style absent. 594. M . lanceolata

2-6 cm. long; paler green below. 595. M . dicin0 Leaflets with apex rounded. 596. M . pall&

Leaves with petiole 4 4 cm., dull above; petals shorter than inner sepals; style

Leaves with petiole 5-10 mm., very lustrous above; petals as long as inner

Leaflets broader (1 : 1-2), in 3-4 pairs; margins 6 6 coarsely spinose-dentate;

Stamens edentate.

short. Cf. 593. M . paniculata

sepals; style absent. 597. M . hartwegii Leaflets with the lowest pair only 2-5 (-10) mm. above the base of the leaf rachis.

Leaflets very lustrous, oblong-ovate (1 : 2-2.5), with apex acute; stamens edentate.

Leaflets dull, or scarcely lustrous, broader, (1 : 1-1-2), with apex rounded; Cf. 597. M . hartwegii

stamens dentate. Leaflets 5-11 x 3-9 cm., in 3-5 pairs; pedicels 15-20 mm. Leaflets P 5 x 2-3 cm., in 5-6 pairs; pedicels 5-10 nun.

598. M . pmi i 599. M . zimapana

2-4 mm. subequal to their bracts. 600. M . argutu Lcaflets narrow (1 : P 5 ) , in (5) 7-13 pairs; margins with 2-3 spinose teeth; pedicels

(590) Mahonia ehrenbergii (Kunze) Fedde, Bot. Jb. 31, 106, 16 Aug. 1901; syn. B.

Shrub 3 4 ft. Leaves 15-25 cm., with 3-7 pairs of leaflets, the lowest pair 3 4 cm. above the base of the rachis. Leaflets 3 4 - 5 x 1-2 cm.; base subsessile; margins entire; apex acute; both sides lustrous and closely reticulate. P a n i c h 25-35 cm. long, including peduncle 10-15 cm. long; bracts of the inflorescence 4-6 mm. long. Pediceb 5-10 mm.; floral bracts 1-3 mm. Prophylb situated towards middle of pedicels. Petals emarginate. Stamens dentate.

ehrenbergii Kunze, Linnaea, 20, 45, 1847

Mexico (south) : seed only, Ehrenberg, s.n. Cultivated: Halle Botanic Garden, fl. May 1844 (Type) .

(591) Mahonia chiapensis Lundell, Lloydia, 3,209,1940 Shrub, glabrous, 10 ft. high. Leaves 40-60 cm. long, including petiole 9-15 cm. long

the lowest of the 6-7 pairs of leaflets being set well up the rachis. LeaJEets entire, chartaceous, ovate to lanceolate, 8.5-13.5 x 2.7-6 cm.; bases rounded (on the lower part of rachis) or abruptly acute (on the upper part of rachis); apex acuminate, apiculate ; midrib impressed above, elevated below ; reticulate, prominently so below ; petiolules 3-7 mm. long. Inflorescence cymose-paniculate to 50 cm. long, including peduncle up to 12 cm., with slender wiry branchlets drying black; bracts of the inflorescence 8 mm. long. Pedicels to 1.8 cm.; floral bracts a t base. Prophylls situated on pedicel. Sepals in three series, outer 2 mm., inner 7 mm. long. Petals to 5.5 mm., emarginate. Stamwi 3-3-5 mm: filaments thick; a t their apex below the anthers, dentate. Ovary ovoid.

Mexico (south): Chiapas State, Mt. Ovando, in shade, 6700 ft., 14-18 Nov. 1939, Eizi Matuda 3996 (Type, University of Michigan Herb.). Not cultivated.

(592) Mahonia johnstonii (Standley et Steyerm.) Standley et Steyerm., Publ. Bot. Ser. Field Mus. Nut. Hist. Chicago, 23, 6, 1943; Standl. et Steyerm., ‘Flora of Guatemala’, in Fieldiana, Botany, 24 (part I V ) , 277, 1946 (Chicago); eyn. B. johnstonii Standl. et Steyerm., Publ. Bot. Xer. Field Mus. Nat. Hist. Chicago, 22 (no. 3, publication 478), 140, 10 Sept. 1940

Shrub 6-8 ft., glabrous, with stems densely leaved, the internodes between the latter being short. Leaves 2-6 cm. long, with slender rachis and long petiole, and (2-) 3 (-4)

Page 348: Berberis and Mahonia : A taxonomic revision

348 L. W. A. AERENDT pairs of leaflets. Leaflets 3-5 x 1-5-2-5 cm., oblong-ovate; base sessile; margins entire, revolute; apex rounded or very obtuse, sometimes retuse; above lustrous with very prominent venation ; below paler with prominent venation and close reticulation. Other- wise unrecorded.

Guatemala: Sacatephquez Prov., Parramos, 14 Dec. 1938, Johnston 1525 (Type, Herb. Field Mus. Chicago); damp thicket near Pastom, 5200 ft., 14 Dec. 1938, Standley 59895.

Santa Rosa Prov. (north), Baja Verapaz, in pine oak forest, 30 Mar. 1939, Standley 69801.

Zacapa Prov., Sierra de las I6na.s. According to the authors, endemic to Guatemala, where it is found on dry, brushy, often rocky, hillsides at 4300-5500 ft. Not cultivated.

(593) Mahonia paniculata Oerst., Kjoeben?uvn Vidensk., M a . , 36, 1856; Fedde, 1.c. p. 107 ; Stand. et Steyerm. Fkldiana, 24 (part IV), 277,1946; syn. B. paniculata Hemsl., Biol. Centr. Amer. 1, 24, 1879; B. hemskyi Donn. Smith in Pittier, $7. Costa Rica, 2,47, 1898, non B. hemskyam Ahrendt

Leaves with 7-8 pairs of leaflets, lowest pair 4-8 cm. above the base, the remaining internodes decreasing from base to apex of rachis, 4 - 5 4 cm. LeaJletS the lowest pair 5 x 1.7 cm., the others subequal, 7-5 x 2.5 cm. to 9-5-12 x 3 4 cm. ; margins 12-20-spinu- lose (0.5 mm.)-serrulate (0.5 mm.), distant 2 4 mm.; apex shortly acuminate with mucro 3 4 mm. long; above dull; below scarcely paler yellow-green; both sides with h e l y elevated reticulation, below with coarse conspicuous lateral venation. Panicles 18-25 om. long, including peduncle 6-14 cm. ; subpeduncles of branched parts 1 - 6 3 cm. Pedicels 7-12 (-20) mm.; floral bracts 2-3 (-5) mm. ProphyuS 1 mm., situated towards middle of the pedicels. Outer sepal-s 2 x 1.5 mm., ovate, acute. Inner s e p h 5 x 3 mm. Petals 4*25mm., emarginate, with rounded lobes. Stamens very slightly dentate or subdentate; apex apiculate. Ovules 5-6. lctyle short.

Guatemala: Volcsln de Fuego, 10,500 ft., 10 Jan. 1874, Salvin, s.n. (K). Costa Rica: Llanuras Prov. (on border of Cartago Prov.), Vol& Irazb, by Palo de

San Juan, 8000-9000ft., 1857, Oerstedt, 8.n. (Isotype, K); alao, according to Fedde, Phramo, dans les clairihres du versant S. du Barba et de l’Iraz6, Pittier 868.

San Jose Prov. Hofmann, s.n., according to Fedde. Not cultivated. Hemsley keeps Salvin’s specimen from Guatemala distinct aa Beiberk sp. 20.

(594) Mahonia lanceolata (Benth.) Fedde, Bot. Jb . 31, 92, 16 Aug. 1901; syn.

Leaves with 5-8 pairs of remote leaflets; the lowest pair similar to the adjacent ones and 1 - 5 4 cm. above the base of the rachis; the remaining internodes decreasing from base to apex of rachis from 6 to 4 cm. Leujlets thin, lanceolate; size slightly increasing from base to apex (6-) 9 x (2-) 3 cm. to 13 x 3.5 cm., or 12 x 4 cm. ; basal pair sometimes slightly more ovate, 4 4 x 1 - 5 2 cm. ; terminal leaflet similar and subequal to adjacent ones, subsessile or with petiole 1-2 cm. ; base cuneate, petiolulate (1-3 mm.); margins 12-16-spinose (1-1.5 mm.)-serrate (0-5-2 mm.), distant 4-8 mm.; apex acuminate; above scarcely lustrous ; both sides hely, closely reticulate, impressed above and elevated below. Panicles 50-100-fld., with thick rachis, 25-3Ocm. long, including peduncle. S u b e d u w of the branched portions slender, 1-6 cm. ; bracts 3 4 mm. Pedicels slender 8-12 mm.; floral bracts 2-3 mm. Prophylk 1 mm., situated between middle and apex of pedicel. Outer sepah 2.5 x 1.5 mm., ovate, acute. Median sepals 6 x 3.5 mm., elliptic or oblong-elliptic. Inner sepals similar, 8 x 3.5 mm. Petals 7 x 4 mm., obovate-elliptic; base distinctly clawed, with small glands; apex shortly retuse with rounded lobes. Stamens 5 mm., dentate; apex not produced, truncate. Ovules 5. Berries at first slightly pruinose, soon and finally eprujnose and black, estylose, ovoid, 11-12 x 7-8 mm.

B. laneeolata Benth., P1. Hartw. p. 34, 1840

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Berberis and Mahonia 349

Mexico: Mexico State, mountains near Apulco (Aculco), Hartweg 269 (Type) ; Temn- scaltepec district, Nanchititla, fr. 6 Oct. 1933, Hinton 4103, and fl. 21 Mar. 1935, Hinton 7547 (BM, K).

Hidalgo State, Barranca, below Trinidad ironworks, 5500 ft., 4-6 ft., 9 Sept. 1906, Pringle 10301 (BM).

Not in cultivation.

(595) Mahonia ilicina Schdl., Linnaea, 10, 236, 1835; Fedde, Bot. Jb. 31, 112; syn.

Shrub 4-9 ft. Leaves 7-15 cm., with (3-) 4 (-5) pairs of leaflets, the lowest pair 1- 2.5 cm. above the base of the rachis. Leajets 2-6 x 1-3 cm. ovate; base subpetiolulate; margins 4-6-spinose (2-3 mm.)-serrate (1-5-2-5 mm.), distant 6-10 mm.; scarcely or slightly lustrous both sides ; below paler green, with distinct branched venation, coarsely prominent reticulation fine and somewhat open. Panicles 10-15 cm. long; bracts of the inflorescence 4-5 mm. long. Pedicels 5-10 mm. long; floral bracts 2-3 mm. long, but those of the subpeduncles 5-7 mm. long. Prophylls situated near the middle of the pedicels. Outer sepals small, broadly acuminate. Petals shorter than inner sepals, and emarginate. Stamns shortly dentate. Ovules 5-7. Berries globose, 5 mm., estylose, pruinose blue.

B. ilicina (Schdl.) Hemsl. Biol. Centr. Amer. 1, 23, 1879

Mexico : between Guantolapa and Tlachichuca, Schiede 300 (Type). Hidalgo State, Atotonilco, 1838-9, Ehrenberg 977 ; near La Encarnacih, 1830, Schiede,

s.n., according to Fedde. Not in cultivation.

(596) Mahoniapallida (Hartw.) Fedde, 1.c. p. 109,16 Aug. 1901 ; syn. B.&Zida Hartw.

Shrub 5-8 ft. tall. leave.^ 25-30 cm. long, with 5-6 pairs of overlapping leaflets, the lowest pair being 1-15 em. above the base. Leaflets 6-9 x 3 4 cm.; petiolule 2-3 mm. ; margins very undulate, 9-11-spinose (1.5 mm.)-serrate (0.5 mm.), distant 5-9 mm.; apex rounded, mucronulate ; above dull, or scarcely lustrous ; finely closely reticulate ; below paler, yellow-green, openly reticulate. Panicles 20-40 em. long, including peduncle, 5-15 cm. long; bracts of the inflorescence, 6-9 mm. Pediceb 5-10 mm. Prophylls situated at mid-point of pedicels. Petals incised. Stamens dentate. Ovules 6-7. Berries globose, 7 mm., pruinose blue. Seeds purple-black, 4-5 mm.

Mexico (south): Hidalgo State, San Jose del Oro, Zacualtipan, Hartweg 268 (Type, K, BM); hills below Cardonal and ZimapBn, and beyond to Atotonilco, then by La Majada, San Jose del Oro.

in Benth., PI. Hartw. p. 34, 1840; Hemsley, 1.c. p. 23, 1879

Oaxaca State, 6700 ft., 1894, Pringle 4812.

(597) Mahonia hartwegii (Benth.) Fedde, Bot. Jb. 31, 109, 16 Aug. 1901; syn.

Leaves 25-50 cm. long, with 5-7 pairs of leaflets, the lowest pair being much smaller, 2-4 x 1-2 cm., and situated 5-10 mm. above the base ofthe rachis; the remaining inter- nodes decreasing from base to apex of rachis from 8 to 4 cm. k $ e i % thin, oblong- ovate, subequal (apart from the basal pair), (6-) 7-9 (-10) x (2.5-) 3-3-5 ( 4 . 5 ) cm., or to S x 5 cm.; petiolule 1-3 mm.; margins 10-20-spinose (1 mm.) serrulate (scarcely 0.5 mm.), distant 3 4 mm.; apex acute; above very lustrous, subenervate or indis- tinctly veined ; below slightly lustrous paler yellow-green, with branched venation clearly elevated and indistinct open reticulation; terminal leaflet with petiole 2-3 em. long. Panicles 20-35 cm. long, loosely (7-) 10-18-fld., with a thick peduncle 3-9 om. long, the subpeduncles of the branched parts measuring 1 4 cm. Pedicels 5-10 (-12) mm.,

B. hartwegii Benth., PI. Hartw. p. 34, 1840

Page 350: Berberis and Mahonia : A taxonomic revision

350 L. W. A. AERENDT glabrous; floral bracts 2-26 mm. ProphyuS 1 mm. long, situated on the pedicels 3 4 mm. below the calyx. Outer se& 2-26 x 2 mm., broadly ovate, obtuse to rounded. Inner sepals 5-5-6 x 2-26 mm., narrowly oblong-elliptic. Pela2s 5.5-6 x 3-36 mm., obovate, emarginate. Stamens c. 4.5 mm., edentate. OvuEap c. 7. Style absent.

Mexico: Hidalgo State, by Contadero, between Tulo and St Barbara, 1839, Hartweg 272 (BM, K, 0). Not in cultivation.

(598) Mahonia paxii Fedde, Bot. Jb. 31, 113, 16 Aug. 1901

Leaves 15-30 cm. long, with 3-5 pairs of leaflets, the lowest pair situated only 3-4 mm. above the base of the rachis. Lea$& 5-11 x 3-9 mm. ; petiolule 3-4 mm. ; margins 7-17-spinose (1 mm.)-serrate (1-1-5 mm.), distant 6-15 mm.; apex rounded; both sides scarcely lustrous, openly reticulate. Panicles 15-25 cm. long; bracts of the idoreacence 5-10 mm. long. Pedicds 1-5-2 cm. long; floral bracts 2-5 mm. long. ProphyUs situated towards the middle of the pedicel. Petads emarginate. Berries subglobose, 5 mm.

Mexico: Hidalgo State, between Zimaphn and La Enmrnaci6ny 1M, Ehrenberg 1108, 1110 (Type).

(590) Mahonia zimapana Fedde, 1.c. p. 111, 16 Aug. 1901 Leaves 10-15 cm. long, with 5-6 pairs of leaflets, the lowest pair situated only 5 mm.

above the base of the rachis. L e a , 4-5 x 2.3 cm.; petiolule 3-4 mm.; margins 6-10- spinose (2 mm.)-serrate (1-5-2 mm.), distant 5-10 mm.; above slightly lustrous; below dull, paler; both sides openly reticulate. Panicles 20-25 cm. long, with bracts of the inflorescence 4-5 mm. long. Pedicels 5-10 mm. ; floral bracts 2 mm. ProphyUs situated towards the middle of the pedicels. Petals emarginate. StamRns dentate.

Mexico: Hidalgo State, Las Verdoaas, by Zimapfin, Ehrenberg 1078 (Isotype, BM). Not cultivated.

(600) Mahonia arguta Hutch., Card. Chron. 43, 82, 8 Feb. 1908; and Bot. Mag. 135, t. 8266 (1909); syn. B. arguta (Hutch.) Bean, Trees and Shrubs, 3, 42, 1933, non Franch.

Shrub 5-7 ft. Leaves to 30 om. long, with (5-) 7-11 (-13) remote pairs of leaflets, the lowest pair only 5-10 mm. above the base of the rachis, and the remaining internodes decreasing from base to apex of the rachis from 4 to 2 cm. LeaJlets lanceolate, or oblong- lanceolate; the basal pair only 25 x 6 mm., the others 5-9 x 1-2 cm.; base cuneate, sub- sessile; margins (1-) 2-3-spinose (1-1.5 mm.)-serrate (1-3 mm.), distant 7-20 mm., but sometimes subentire ; terminal leaflet subsessile ; both sides lustrous and conspicuously reticulate. Panicles 50-60-flowered7 3040 cm. long, including peduncle 10-14 cm., the branched portions being 24-flowered. Pedieels 2 4 mm. ; bracts subequal. Flowers pale yellow. Outer sepal5 3 x 1.75 mm., elliptic. Median and inner se+ 6 x 3 mm., elliptic. Petals 4x3mm., oblong; emarginate with obtuse lobes. Stamens 6mm., edentate; apex not produced, truncate. Fruit globose, c. 8 mm., estylose.

Mexico or Central America. The origin of this p h t is uncertain; it was received at Kew in 1907, from Glasnevin,

where it had been cultivated for 28 years previously; the plant from which the type specimen WM taken flowered at Kew in May 1907 and May 1908, being kept in a dold house for the winter, but outside in the summer.

Sect. Horridae Fedde, Bot. J b . 31, 70, 77, 1901 Shrubs 3-12 ft. high. Leaves with 1-3 (-9) pairs of leaflets, and with petiole 2 mm.-

5 cm., this being the distance of the lowest leaflets above the of the leaf rachis. Lea,$& chartaceous, thin or thick, but always stiff or rigid. Inforeswnce 3-8-flowered7 mostly fmcicled, umbellate or subumbellate, occasionally in the form of a short few-

Page 351: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 351 flowered raceme. Pedicels 4 2 0 mm. ; floral bracts mostly 1-3 mm., occasionally folia- ceous, 3-9 mm. ProphyUs present, in varying positions according to the species. Flowers 5-12 mm. diameter. Sepals in 1-3 series. Inner sepals 5.5-8 mm. Petals always shorter than inner sepals. Ovules (4) 5-9 (-18). Berries mostly black, pruinose blue, but occasionally red, reddish white or reddish yellow.

U.S.A. : Texas, New Mexico, Arizona, Colorado, Utah, California. Mexico: Coahuila, Sonora, Nuevo Leon and San Luis Potosi States.

KEY 110, to Sect. Horridae (Maps 49 and 51) Leaflets very broad, 1 : 14-1.6, only 6-20 mm. long, with apex obtuse or broadiy acute,

Leaflets narrower, 1 : 2-6, and (15 mm.-) 2-6 cm. long, with apex acute to acuminate, in 5-9 pairs. 601. M . muelleri

in 1-4 pairs. Leaflets fairly narrow, 1 : 2-3 (-4), not or scarcely revolute, shortly petiolulate.

[Inflorescence densely, many flowered, racemose. Cf. 570. M . californica] Inflorescence sparsely few-flowered, mostly fascicled, umbellate or subumbellate.

Leaves with only one pair of leaflets; stamens edentate; fruit blue-black; style

Leaves with 2-4 pairs of leaflets; stamens dentate (where known) ; style absent. distinct. 602. M . trifoliolatu

Ovules 4-8. Leaflets thick, in 2-4 pairs; lowest pair only 1-2 m m . above base; margins

Leaves slightly veined or subenervate; bracts not foliaceous, 1-3 m m . with 3-6 teeth; fruit 10-15 mm. long.

long; fruit black, pruinose blue, 10 x 15 mm. diam. Seeds almost black, 4-5 mm. long. 603. M . fremontii

608. M . higginsae

red, epruinose, l(r12 mm. long. 604. M . swaseyi

margins with 10-18 teeth; fruit red, epruinose, 6-7 mm. long.

Fruit yellowish red, 6-8 mm. diam. ; seeds red-brown, 3 mm. long

Leaves closely reticulate ; bracts foliaceous, 3-9 mm. long; fruit yellowish

Leaflets thin, in 2 pairs; lowest pair (5mm.-) 1.5-2.5 cm. above base;

605. M . nevinii Ovules 8-18; fruit red, epruinose, 5-8 mm. long. 606. M.haematocarpa

607. M . pinifolia Leaflets very narrow, 1 : 5-6, revolute, almost linear, sessile.

(601) Mahonia muelleri I. M. Johnston, J . Am. Arb. 31, 189, 1950 Leaves with 5-9 pairs of leaflets, the lowest pair set only 2-3 mm. above the base of the

rachis, the remaining internodes being 8-10 mm. Leaflets elliptic or ovate-elliptic, the basal pair 6-10 mm. long, the remaining ones subequal, 10-20 x 7-12 mm. ; base obtuse or subtruncate; margins finely spinose (1-1-5 mm.)-serrate (1 mrn.); apex obtuse or broadly acute; terminal leaflet scarcely larger, with petiole 3-8 mm. ; chartaceous, thin, stiff, scarcely rigid; above dull grey-green; below paler. InJlorescence very loosely sub- umbellate or subracemose, 3-5-flowered, with rachis 1 - 6 2 cm. long; bracts of the inflorescence 6 x 3 mm. Pedicels 5-10 mm. long; floral bracts 1-23 mm. long. Plowers unknown. Berries subglobose, 6-7 mm., inflated. Ovules 6-10.

Mexico: Nuevo Leon State, road from Pablillo to Cieneguillas, about 15 miles south- west of Galeana, very abundant in the lower pinyon belt, 15 June 1934, C. H. & M. T. Mueller 803 (Type, Am. Arb.). The author remarks that this is a plant ofthe lower pinyon belt in the east slope of the Sierra Madre Oriental of N.E. Mexico; and that it is most closely related to M . f r e m t i i , which belongs to the west of U.S.A. and N.W. Mexico. M . fremntii has pallid, thick, very rigid, narrow, coarsely dentate, subacuminate, lobulate and very undulate leaflets. M . muelleri differs in its much thinner, less pallid, very much less rigid leaflets, which are not undulate, but have nearly flat surfaces, and only minutely toothed margins.

Page 352: Berberis and Mahonia : A taxonomic revision

352 L. W. A. AHRENDT Cultivated: L.A. 747. A plant which I received in 1950, which did not flower, and

which was labelled (obviously inaccurately) as M. frenzontid, is probably this species.

(602) Mahonia trifoliolata (Moric.) Fdde, Bot. Jb . 31, 96, 16 Aug. 1901; Johnston, 1.c. p. 190, 1950; syn. B. trifoliolatu Moric., P1. N w v . Am’r. 113, t. 69, 1841; Hemsl., Bid. Centr. A m r . 1, 24, 1879; syn. B. trvfoliata, Lemaire, Fl. Sew@, 1, 130, t. 56, 1845; B. ilicifolia Scheele, Linnaea, 21,591, 1848, non Forst.; B. roe- m r k n a Scheele, L i n w , 22, 154, 352, 1849

Shrub 3-7 ft. Leaves trifoliolate, having but one pair of leaflets, set 1.5 cm. above the base. Leaflets fairly thick, 2-6x1-1.5 cm.; margins (1-) 2 (-3)-spinose (1-2mm.)- dentate (2-5-7 mm.), distant 1.5-2 cm.; terminal leaflet sessile; both sides subenervate. Inflorescence 3-6-fld., 1-2-5 (-4) cm. long, subfascicled, semile. Pedicels 5-8 mm.; floral bracts 1-5-2 mm. Prophylls appressed to calyx. Outer sepals small, ovate, obtuse. P& entire, shorter than the suborbicular inner sepals. Stamens edentate. Berries black, pruinose blue, subglobose, 6-7 x 4.5-5 mm., excluding style 0-25-0-5 mm. long.

Ledeta above slightly lustrous green, and epapillose. M e t s above conspicuously grey-green, and very minutely and closely papillose.

602a. M . t ~ f o l w h var. t k f o l i o h

602b. M. trfoliohta var. g k w x

(602a) Mahonia trifoliolata var. trifonolata

9131 (K); and, according to Johnston: Texas: Cmliad County, Goliad, thickets on dry calcareous soil, fl. 10 Mar. 1916, Palmer

Nueces County, Corpus Christi, Heller 1384 (K). Travis County, Austin, 1872, Hall 10. Coma1 County, New Braunfels, Lindheimer 322,575,661. Bexar County, San Antonio, 1853, Thurber, s.n. Frio County, near Rio Frio, on the old road between Laredo and San Antonio, Mar.

Bee County, north-east of Beeville, Wiegand 599. Kleberg County, Sinclair, 8.n. Duval County, neax San Diego, Mar. 1887, Sargent, s.n. Jim Hogg County, Hebronville, Hanson 341. Johnston summarizes the distribution thus: it is conlined to Texas, south of Austin

to Corpus Christi. Although to be expected in Mexico, north of Tamaulipas State, no Mexican collection has been seen.

1828, Berlandier 1437 (Type).

(602b) Mahonia trifoliolata var. glauca I. M. Johnston, J. A m . Arb. 31, 190, 1950;

Texas: Hood County, Comanche Peak, near Granbury, dry limestone ledges, 15 Sept.

Armstrong County, Gamble Ranch, Palmer 13910. Brewster County, Chisos mountains, Moore & Steyermark 3121. Taylor County, near Camp Berkeley, Tolatead 6880. Medina County (north), south of Tarpley, McVaugh 7671. Terrell County, Fedora, Palmer 33576. Presidio County, Davis mountains, Moore & Steyermark, 8.n. Culberson County, 11 miles north of Van Horn, Waterfall 4613. Arizona: Pima County (north-east), Santa Catalina mountains, 1881, Vasey, 8.n. (K). Mexico: Coahda State, without further locality, immature fr., 1898, Palmer 139 (K). Chihuahua State, vicinity of Santa Eulalia, 4300 ft., 28-29 Apr. 1908, Palmer 135 (K),

syn. B. trvfoliata Hartw. ex Lindl., Bot. Reg. 27, Misc. 68, 1841

1914, Palmer 6544 (K); 1846, fl., fr. Lindheiner 322 (0). Also, according to Johnston:

and 5-10 June 1908, Palmer 341 (K); near Chihuahua, fr. 1885, Pringle 261 (K).

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Berberis and Mahonia 353

San Luis Potosi State, seed, Hartweg, at Hacienda del Espiritu Santo, on the road between Zacatecas and San Luis Potosi. Johnston describes this as 'the most widely distributed variety, covering large areas in west Texas, and southwards into Mexico' (north central). It appears to stretch even farther west into Arizona. Johnston remarks that the varieties have botanical and geographical distinction.

It would appear that the former, green, variety lies in south-east Texas from 27" 50'- 30" 20' N. lat., and 97-99" W. long., while the second, grey, variety, lies in west Texas to Arizona (presumably including New Mexico), from 30-32" 50' N. lat., and 101-110" W. long., and in Mexico from 22-32" N. lat., and 101-107" W. long.

Cultivated: it is the latter, grey, variety which is in cultivation in England. This introduction may be based on the plants grown from Hartweg's Mexican seed (collected a t the southernmost part of the area of distribution), on which plants the name B. tri- foliata was conferred.

Lindley remarks of his B. trifoliata, which is M . trifoliolutu var. gluucu, 'Hartweg found it on the road from Zacatecas to San Luis de Potosi, where it covered large tracts of country, of an immense plain, otherwise occupied chiefly by Opuntias, Yuccas, and stunted plants of Prosopis dulcis '.

(603) Mahonia fremontii (Torr.) Fedde, Bot. Jb . 31, 98, 16 Aug. 1901; syn. B. trifoliatu Ton, in Whipple's Rep. Expl. Survey, 4, 63, no. 4 (part), 1856, non Hartw. ex Lindl., 1841, non Lemaire; B.fremontii Torr., Bot. Mex. Bound., p. 30, 1858; Johnston, J . A m . Arb. 31, 190, in obs. 1950

Leaves 3-10 cm. long, with (1-) 2-3 (4) pairs of leaflets, the lowest pair only 1-2 mm. above the base. Leaflets thick, rigid, oblong-lanceolate; 2-6 x 1-1.5 cm.; margins 3 4 (-6)-spinose (1-2 mm.)-dentate (2-4 mm.), distant 3-7 mm.; dull grey-green, sub- enervate or indistinctly veined. Injlorescence 3-5-fld., fascicled or racemose-sub- umbellate, 2-5 cm. long; peduncle nil, or 1-1.5 cm. long. Pedicels (4) 8-12 (-15) mm. ; floral bracts 1-3 mm. Prophylls situated, one towards the middle of the pedicels, one towards their apex appressed to the calyx, ovate, acute, reddish, 2.5 x 1.5 mm. Outer sepals whitish yellow, 3.5-3.75 x 15-1-75 mm., ovate, subacute. Median sepals 6.5- 7 mm., obovate, rounded, slenderly clawed. Petals 5 x 3.5 nim., apex entire; base scarcely clawed, or cuneate with oblong, acute, marginal glands, 0-6 x 0.2 mm. Stamens 4mm., dentate; apex not produced, subtruncate to slightly rounded. Ovules 5-8. Berries black, pruinose blue, estylose, ovoid, 1-5 x 1 cm.

Colorado (west): Hinsdale County, valley of the Rio Grande, below Douana, Passy, s.n. (K).

Utah (south, 38" N. lat.); fr. 1874, Parry 5 (K); 3500 ft., in gravel, fr. 5 Aug. 1894, Jons 3196 (BM); 4500 ft., in gravel, fl. 28 Apr. 1894, Jons 5122 (BM).

Arizona (south): Pima County, Tanque Verde mountains, Tucson, 4 July, 1935, Nelson 1459 (K).

California : according to Johnston. Mexico : Sonora State ; Lower California, according to Johnston who records that

' M . fremontii ranges in the deserts of western U.S.A. from west Colorado, and north-west Mexico westward into California and south to Lower California'.

Cultivated: fl. Apr. 1937; the idorescence in this instance was entirely fascicled. Closely related to M . fremontii in its leaflets is (570), M . cal(fornicu, which has,

however, the dense racemes of the Aquifoliatae, and seems to lie between the sections, botanically and geographically and (608) M . higgimae with yellowish red fruits.

(604) Mahonia swaseyi (Buckley) Fedde, 1.c. p. 102, 16 Aug. 1901; syn. B. swmeyi, Buckley in Young, Fl. Texm, p. 152, 1873

Shrub 4-8 ft. Leaves 5-9 x 4-6 cm., with (2-) 3 (4) pairs of remote leaflets, the lowest pair shorter and broader, c. 3 x 1-75 cm., set only 1-2 mm. above the base of the rachis,

23 JOURN. LINN. SOC. BOTANY-VOL. LVII

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364 L. W. A. AHRENDT

the remaining internode8 decreasing upwards from 15-20 to 10-12 mm. Leaflets moder- ately thick, 2-5-3.2 x 0-6-0.8 cm. ; base sessile or with petiolule 1-2 mm. long; margins 4-6-spinose (1-1-5 mm.)-serrate (1-1-5 mm.), distant 3-6 mm.; above grey-green; below pruinose grey; both sides fairly closely reticulate. In@eswnce 44-fld., 4-6 cm. long, loosely racemose. Pediceh 7-14mm.; bracts foliaceous, ovate, acute, 3-9x3-5mm. Prophy& set towards the middle of the pedicels, 2.5 x 1-5 mm., ovate, acute. OpruEes 5-7. Berries globose, 10-12 mm., estylose, white or yellow, slightly reddish.

Texae (south, c. 30" N. lat., 98" W., long.). Hays County, Buckley, s.n. (Type); Plank, s.n. Comal County, dong rocky streams by Devil's Backbone, 16 June 1917, Palmer

Cultivated: 1937 ; this p h t did not flower. This and the following are apparently rare, 12179 (K).

restricted, endemic species.

(605) Mahonia nevlnii (Gray) Fedde, Bot. Jb. 31, 102, 16 Aug. 1901; syn. B. nevinii

S h d sparsely branched, 3-5 ft. Leaves to 10 x 6 cm., with 2 pairs of remote leaflets, the lowest pair (0.5) 1-5-2-5 cm. above the base of the rachis; the second pair 1-2 cm. above the latter. h f l e t s thin, stiff, lanceolate, similar, subequal, 2-3.5 x 0.7-1.3 cm. ; margins 10-18-spinose (1-2 mm.)-serrate (0.5-1 mm.), distant 2 - 5 4 mm.; apex acumi- nate; terminal leaflet slightly larger, with petiole 5-9 mm.; above blue-green to grey- green, with characteristic indistinct white venation; below pruinose, grey, scarcely or not papillose. Inflorescence 4-8-fld., racemose-subumbellate. Pedicels 5-10 (-12) mm.; floral bracts 2-3 mm. Flowers 5-6 mm. diameter, H.C.C. 5, but greenish without in the region close to the apex of the pedicel. ProphylLs situated at the apex of the pedicels, appressed to the calyx but with their tips reflexed, but sometimes 1-3 mm. below the apex; two outer prophylls triangular, 1-75 x 1 mm.; one inner prophyll, triangular, acuminate, 3 x 1 mm. S e p h in only one series, 5.5 x 3-25 mm., oblong-obovate. P e a 4.5 x 3 mm., elliptic-obovate, subacutely emarginate with subacute lobes. Shwns 3-3-25 mm. ; below the anthers conspicuously lanceolate-acuminate-dentate ; apex not produced, truncate. Berries globose, red, epruinose, estylose, 6-7 mm. Seeds purple, 3 mm. long.

S. California: Los Angeles County, by Los Angeles, Nevin, s.n. (Type); Darland, s.n. (K, sheet 1395).

Cultivated: fl. Apr. 1937; also flowering at the University Botanic Garden, Oxford, Apr. 1952.

Gray, Sympt. Fl. N. Amer. 1 (i) 69, 1895

(606) Mahonia haematocarpa (Wooton) Fedde, 1.c. p. 100, 16 Aug. 1901; syn.

Leavm with (1-) 2-3 pairs of leaflets, the lowest pair only 2-5 mm. above the base. S t i w 0.5-2 mm. h f l e t s thick, rigid, narrowly oblong-ovate, or oblong-lanceolate, 1.5-4 x 0.5-1 cm.; margins 2-4-spinose (1-2 mm.)-serrate (3-5 mm.), distant 6-12 mm.; above dull grey-green, subenervate. Inflorescence 5-7-fld., openly umbellate-racemose. Pediceb 1-2 cm.; floral bracts 1-3 mm. Prophylls 2, one situated at the middle point of the pedicels, the other situated at the apex of the pedicels appressed to the calyx. outer se& broadly ovate. Mediun sepah obovate. Inner se& obovate. Petals slightly shorter than inner sepals, and with apex emarginate. Stamens dentate. Ovules (9-) 12-18. Berries red, ovoid-globose, estylose, 7-8 mm. Seeds purple, 3 mm. long.

B. haemat0Ca;rlmc Wooton, Bull. Torrey Bot. CJ. 25,304, 1898

Texas (eaat): 1847, Hall 161 and Lindheimer 175 (K). Colorado (west): Sen Miguel County, between Dry Creek and E. Gunnison, 4500-

4900 ft., Purpus 100.

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Berberis and Mahonia 355

New Mexico (south) : Otero County, Sacramento mountains, fk. 18 Aug. 1914, Rehder

Lincoln County, Mescalero Agency, in the white mountains, 58(H)ft., Wooton 376

Dona Ana County, Organ mountains, 5800 ft., 1899, Wooton, s.n. (Type). Arizona: among rocks on Apache Trail, above Fish Creek canyon, fi. 5 Apr. 1935,

Nelson 1785 (K). Mexico (north-west) : Sonora State, Sasali, 1892, Pringle, s.n. According to Johnston,

'the large area in New Mexico, Texas, and adjacent Mexico, separating the range of H. fremontii and M . mwlleri, is occupied by M . haematocarpa. This is closely related to M . fremontii and M . muelleri and, in N. &om and New Mexico, even intergrades with it. In the east portion of its range, where it occurs closest to the area of M . muelleri, M . haemtocurpa becomes relatively stable and a readily delimited species. Its rigid, undulate, pallid, coarsely toothed, lanceolate leailets with the terminal one frequently sessile and larger than the others, separate it readily from M . muelleri.'

M . muelleri seems quite distinct amongst this section; while M . fremontii and M . haematocarpa are very close and supedicially very similar. Their leaves and leaflets are very much alike, and they share the twofold distribution of their prophylls. The basic difference between them lies in the blue-black fruit and 5-8 ovules of M . fremontii, and the red fruit and 12-18 ovules of M . ?mematocarp.

Cultivated: July 1942; this plant did not flower.

312 (K).

(1897).

(607) Mahonia pinifolia Lundell, in 'Studies of Mexican and Central American plants', Publ. Carneg. Imt . , no. 478,209, 1937; syn. B. pinifolia (Lundell) Muller, Amer. Mid.!. Nat. 27, 484, 1942

Shrub glabrous, to 10 ft. Stems short, thick. Leaves mostly gathered together at the apex of the stems, coriaceous, with 1-2 pairs of leaflets, and short petiole which is pale red and 1-5 mm. long, and somewhat spreading and sheathlike at the base. LeaJlets sessile, rigid, aculiform, linear in appearance, being narrowly lanceolate, 15-30 x 3-5 mm., with a revolute margin which is generally entire but sometimes with a few teeth; apex mucronate. InJtorescenw few-flowered in short racemes. Pediwls slender, 3-5 mm. long, pale red. Flowers unknown. Ovules 5-9. Berries red, globose, estylose, 5-8 mm. S& 3-5-4 x 1-2-2 mm., red-brown.

KEY 11OB Leaflets in 1-2 pairs, very revolute and needle-like. Leaflets always in 1 pair only, more palmate.

607a. M . pin$oZia var. pinifoZcolia 607b. M . pinifolia var. coahuilensis

(607 a) Mahonia pinifolia var. pinifolia Mexico: San Luis Potosi State, collected in a shallow, alkaline basin on the road

between Charcas and Santo Domingo, 7 Aug. 1934, Lundell 5608 (Type, Herb. Univ. Michigan).

(607 b) Mahonia pinifolia var. coahuilensis (Muller) Ahrendt, comb.nov. ; syn. B. pini-

Mexico: Coahuila State, Cafion del Pajarito. Sierra de la Madre, Mun. de Cuatro Cienegas, 6 Sept. 1939, Muller 3171 (Type, U.S.N.A.). "his interesting species is not in cultivation.

Muller remarks : ' This species presents an interesting illustration of evolutionary trend. The narrow revolute leaflets are clearly derived from the broader leaflets of the Aquifofiatae. Although the leaves are in some measure suggestive of B. trifoliohta, the

folia var. coahuilemis Muller, 1.c. p. 485, 1942

23-2

Page 356: Berberis and Mahonia : A taxonomic revision

356 L. W. A. AERENDT presence of dense cylindrical papillae on the undersurface clearly denies that relationship. Lundell failed to mention this character in an isotype in the herbarium at Tucson.’

In this comexion, comparison may be made with (601 b), kl. triifoliokcta var. g k c w ; (569), M. purnila; (572), M . sonnei; (573), M . p i p r k ~ w h i c h have papillose leaflets- and (571), H. apuifdaum, where the leaflets are epapillose or scarcely papillose.

(608) Mahonia higginsae (Mum) Ahrendt, combmov.; syn. Berberis higginsae Mw, Aliso 4, 91,1958

‘With much the same habit as B. fremrmtii Tom., stiff, erect, 1-2-5 m. tall, glaucous ; leaves rigid, strongly crisped, spine-toothed with spines 2-34 mm. long; terminal leaflet squarish-ovate to lance-ovate, 15-36 cm. long, 1 - 6 2 cm. wide; lateral leaflets more or less squarish-ovate, 1-2-5(-3) cm. long, 1-2 cm. wide; racemes about 3 - 5 6 cm. long; flowers yellow, about 5 mm. long; berries yellowish red, about 6-7 mm. long; seeds dark red-brown, plump, 2.7-3.2 mm. long. ( h t e x 1-26 m. altus, rigidus, glaucus; foliis rigidis, valde crispatis, spinoso-dentatis, spinis 2-36 mm. longis; foliolis 1-7, terminal- ibua quadmto-ovatis aut Imam-ovatis, 1-5-35 cm. longis, laterdibua 1-2-5(-3) cm. longis; racemis 3 - 5 6 cm. longis; floribus luteis, 5 mm. longis; baccis luteo-coccineis, 6-7 mm. longis; seminibus fusco-rubiginosis, 2.7-3-2 mm. longis.)

Type, dry rocky point at Boulevard, eastern Sen Diego County, California, 12 July 1929, growing at edge of chaparral and near Quercus agrifolia, Johnson & Stark 1351 (RSA sheet 1428). Other specimens seen: Dubber, north-east of Jacumba, San Diego Co., Ethel Higgim 15479; Walkers Ranch near Jacumba, Abranzs 3693. Fifty miles south-east of Tecate on Laguna Hansen Road, Lower California, F. W. Peirson 5866 and Munz 9534; Cieneguita, Sierra Juarez, Lower California, Wiggins 9178; between Neji Rancho and Alaska, Lower California, Wiggins & G k p i e 4165; and Los Aguajitos, 18 milea north-north-west of Rosarito, Lower Californiia, Wiggins 10,017. This proposed species grows in dry rocky places and on slopes at about 3000 to 4500 ft. It resembles B. fremntii, to which it has been referred, in foliage, but has juicy berries ca. 6-8 mm. in diameter instead of dry sated ones 1CL15mm. in diameter; has red-brown seeds ca. 3 mm. long instead of almost black ones 4 5 mm. long. From B. haewadomrpa Wooton, of somewhat similar appearance it differs in its yellowish red rather than plum- coloured berries and in its terminal leaflets being mostly not more than twice as long as wide, instead of 2 to 5 times as long as wide.’

It is a pleasure to dedicate this species to Ethel Bailey Higgins, This species being unknown to me, the original description is reproduced above.

APPENDIX 5 On hybridization between the genera Berberis and Mahonia

It is noticeable in gardens where plants of the two genera are freely grown that self- sown seedlings do not appear as freely of Mahonia as of Berberk. In fact such seedlings are generally confined to those of the (generally hybrid) garden forms of M. aquifolium which produce similar forms. M. japonim and M . bealei, though common in gardens, seem not to produce casual seedlings; but true plants can be raised from carefully sown seeds. The two genera rarely hybridize in gardens. Nevertheless one intergeneric hybrid, on which C. K. Schneider based the hybrid genus x Mahobepberis in his Ill. H a d . Laubh. 1, 806 (1905), has been known for over a hundred years, and three others have arisen during the past twenty years, two in Sweden, one in the United States. They are discussed by G. Kriissmann in Deutsche Baumschuk, 2,300 (1950) and by D. Wyman in Armoldk, 18, 9-12 (1958), who notes their immunity to rust.

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Berberis and Mahonia 357

(609) x Mahoberberis neubertii (Baum.) Schneid., I.c. ; syn. B. neubertii (Hort.) Baum., apud Lemaire, Ill. Hort. 1, Misc., p. 111, 1854, published by Lemaire and Ver- schaffelt; Schneid., 1.c. p. 316, 1906

This hybrid between ‘ B. atropurpurea ’ (according to Baumann), interpreted, doubt- less correctly, as B. vulgaris by Schneider, and M . quifolium appeared in 1850, in the nursery of Baumann, as the result ‘evidently of an artificial cross’ between the species. It forms a fairly evergreen bush 3 4 ft. high, with some finely toothed large obovate leaves like those of B. vulgaris, and some coarsely toothed ovate acute leaves like those of M . aquifolium. The plant is sterile, and was named after Prof. Neubert of Tubingen University (Wurtemberg) ; the type specimen, seemingly, came from Verschaffelt’s nursery. Schneider (1.c. p. 316) remarks that it was then often distributed in gardens as ‘ B. ilicifolia ’. This is still the case.

(610) x M. aquicandidula Kriissmann, 1.c. (1950) Raised in Sweden and reputed to be B. cundidulu x M . aquifolium.

(611) x M. aquisargentii Kriissmann, 1.c. (1950) Raised in Sweden and reputed to be B. sargentii ( Z B. sargentianu) x M . apuqolium.

(612) x M. miethkeana Melander & Eade in Nut. Hort. Mag. 3, 257 (1954)

(1958) to be very similar to x M . apuisargentii imported from Sweden.

(613)

Raised in the United States (near Tacoma, Washington) and stated by Wyman

Berberis glazioviana Brade, in Arquivos do Jardim Botdnico do RW de Juneiro,

Shrub &loft. high. Stems erect, slightly sulcate, glabrous, brown when young. Spines threefold, 7-10 mm. long, broadened towards base. Leaves glabrous, subsessile, fascicled 5-12 ; coriaceous; obovate with cuneate base; apex obtuse, mucronulate; margins on new shoots 4-10-aristate-serrate, but on the mature stems quite entire; venation subreticulate or very openly reticulate ; above dull green ; below paler, slightly greyish; (2.5-)3-5 cm. x 1-2(-3) cm.; almost plane. InflOTescence 3-5-fld., 4-5 cm. long, umbellate or subumbellate with peduncle, 3-44 cm. long. Pedicels 6 1 0 mm. with basal acuminate bracts, 1-2 mm. Flowers 8-10 mm. diam. Prophylls c. 3 mm. long, narrowly ovate, acuminate. Sepals 4.5-5 mm., suborbicular, rounded. Petals 5-55 mm. long, orbicular, rounded, entire ; basal glands c. 0.5 x 0.3 mm., separate, non-marginal, situated above the scarcely clawed base of the petal. &‘tamens edentate, apex scarcely produced, obtuse. Style apparently 3 mm. or more.

Brazil: State of Rio de Janeiro, Serra do Itatiaia, planalto 13 km. da Estrada Nova, 7800 ft., 22 Nov. 1954, A. C. Brade 21293 (Type, Herb. Brade; cotype Herb. do Parque Nacional do Itatiaia).

Among the sect. Laurinae, subsect. Eulaurinae, this species seems to be distinct in its few -flowered umbellate inflorescence, apart from which it seems to approach most nearly to B. glaucescens which has leaves a t first greyish pruinose below, and petals longer than inner sepals. Comparison may also be made with the similar inflorescence of sect. Ilicifoliae, subsect. Euilicifoliae, particularly B. rwcifolia which differs in its dentate leaves and subsessile inflorescence, and B. ilicifolia, also with dentate leaves, and with larger flowers with apiculate stamens. The description given is based on that of the author and his figures. He refers also to the indeterminate Berberis itutiniae Glaz. from this region.

14, 276, 1956; fig. 1-10.

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368 L. W. A. AE~RENDT

ADDBNDA

Sect. Walllchianae, subsect. Subleves (41e) Berberis hayatana Mizushima, in Qzcestione to Fornwsan Barberries, I, Contri-

butbwfmm the Research Institute for Naturad Reeources, no. 670; no. 35, p. 31, 1964. IX; ? B. formosana Li, in J . Wash. Ad., Sci., 42,41 (1952), non Ahrendt

Stem very glabrous; mature yellow; sulcate, stout, c. 1-5mm. thick. Intermodes 1-1.5(-2.5) cm. Spine.9 3-fld., a t right angles, slender, mostly h e l y sulcate below, the centre one 1-17 mm. long. Leuves fascicled 2-3(-5); thinly coriawus, without hypo- derm; oblong, to elliptic- or ovate-oblong; above dull, with midrib impressed; below slightly lustrous green with veins distinctly elevated; margins 3-7-spinose (1 mm.)- serrate, distant 3-4 mm. ; 1.5-2.5-3.5 x 06-1-1.2 cm., broadest a t their midpoints. Flowers fascicled (1-)2, diam. 8 mm. Pedicel.~ very slender, quite glabrous, 8-10 mm. ProphyUs 3, triangular-ovate, acute, c. 1.5 mm. Outer sepals broadly ovate- or obovate- elliptic, apex rounded, 5 mm. long. Inner sepah obovate, rounded, 5 mm. Petals 4 nun., obovate; baaal glands ovate, rounded; apex distinctly emarginate. Starnem c. 2-5 nun., apex slightly produced, subtruncate. Ovule solitary, with stipe as long as itself. Berries ellipsoid, c. 6 mm. long, black, epruinose, or possibly very slightly pruinose; estylose or with stigma subswile.

Formosa: Province Taihoku-Mt. Tai-heizan, Rato-gun, 17 May 1917, Hayyata s.n. (Type, in Herb. Univ. Tokyoensis); mountains near Mmroahu, Taihoku-ayu, 17 July 1952, T. Suzuki 7258 (Type of B. fornaosana Li, in Herb. Univ. National Taiwanensis).

The above admirable description, provided by the author, is so full and clear that the species can, at once, be placed in the subsection Subkves, characterized by yellow, sulcate stems; thin leaves without hypoderm; solitary ovules and black epruinose berriea with short style. B. hayatmm is the only Formosan member of the Subkves, and difFers generally from those previously known through its very few-flowered fascicles and lack of style. B. subk& dif€ers in its conspicuously long ovule stipe, and petals longer than inner sepals. B. dolk?wskvnm is distinct in its exceptionally long stamens, and B. ernilii in its lanceolate, acuminate prophylls and subsessile ovulea. In the general indicative Key 3 for the sect. Wallichhae, B. hayatam would, according to the author’s description of its venation, be associated with 52, B. silvieolu.

Mizwhima makea two important statements : (1) The mountain flora of Formosa is now known phytogeographially to have striking

a%hities with that of south-west China and the Himalayas. (2) He hoped to settle the relationship between Formosan Barberries and the Sino-

Himalayan ones, but was unable to come to any clear conclusion on the subject. Within $he framework of this aim, he investigated the arguments for regarding as

conspecific the Yunnan B. subacuminata, the Kweichou B. b6color and the Formosan B. mingetsensis. Thia is an effort parallel to the suggested merging of the Nepal uniovulate B. wallichiana and the Javan 4-odate B. xanthoxyh. Clearly Mizushima could not make out a caae which satisfied him, his decision being ‘I hope to refrain from uniting the Formosan B. minget-sensk fully to B. bicolor of China.’ He says ‘the number of ovules, a t least in the WaUichianae group, deservea no importance in separating two plants akin to each other in other characteristics’. This view enables him to think conspecifically about the 2-oded B. biwlor and the 6 5 - o d e d B. mingetsenak; but it is at variance with the conclusions which I have recorded. On the other hand, Mizushima has entered carefully and fully into the anatomical characters of the leaves, a basis for discrimination which Schneider investigated, began to use, but soon abandoned-a course which I have followed.

Clearly one’s conclusions wil l depend upon the diagnostic weight given to the char- acters. If one is pre-determined to identify Pacific species with related Sino-Himalayan

Page 359: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 359 ones, it will be necessary to assess the weight of diagnostic character in accord with this pre-conception. My aim haa been to try to assess this weight independently, on its own merits. This has resulted in revealing Himalayan, Chinese, and Pacific counterparts linked within their own particular sections, thus :

Sect. Wallichianae Replicatae. Formosa-16. B. nuntoensis.

Manipuranae. Formosa-27. B. mingetsensis. Yunnan, Szechuan-14. B. taliensis; 15. B. grodtnzannkna.

Acuminatae.

Subleves .

Barandanae .

Leves.

Manipur-21. B. mmipuranu; Burma-26. B. coxii. Formosa-48. B. aristato-serrulata. Yunnan49 . B. acurninata; 50. B. dehvayi; 54. B. subacuminuta;

52. B. silvicola. Kweichou-53. B. bicolor; 56. B. xanthoclada. Tibet-55. B. leptopoda. Fo rmosa41 A. B. hyatuna. Burma-Yunnan41. B. subbvis. Kweichou-43. B. ernilii. Formosa-59. B. kawakumii; 60. B. alpicola. Java-57. B. xanthoxylina. Philippines-58. B. barandam. Formosa-72. B. brevisepala. Nepal-73. B. poluninii. Bhutan-74. B. subpterocluda. Punnan-78. B. levis.

Formosa-B. morrisonensis. Kansu-B. diaphnu. Szechuan-B. aemulaw, B. tischleri. Shensi-B. circumserrata.

Sect. Angulosae, Subsect. Diaphanae

In this way, the striking afities between the Formosan and Sino-Himalayan Berberis populations are clearly displayed. Each Formosan species has one or more Sino- Himalayan counterparts which are quite distinct. These atKnities are shown by a natural loose co-sectional, rather than a forced tight conspecific, presentation. In particular, my analysis would associate, not B. mingetsensis, but B. aristato-sew-, with B. sub- muminuta and B . bicolor. Whereas each Formosan species has its Sino-Himalayan counterpart, the converse is not the case. For nine out of the fourteen subsections of the Wallichianae, no Formosan counterpart is known; and, outside this section, we know only the one Angulosae species given above. There is some internal evidence for re- cognizing an a f i i t y between the Wallichianae and the Angulosae ; doubtless it was this a fk i ty which led Schneider originally to place B. pruinosa with the Angdosae; the presence of B. morrisonensis in Formosa may be regarded as further evidence of this afbity. This note, written on 15 October 1958, soon after I iirst read Mikushima's paper, could not be inserted in position in the text which had then been printed.

ACKNOWLEDOEB~ENTS I am grateful to Dr W. T. Stearn for drawing my attention to certain descriptions

which have been incorporated in the addenda, and for his help in checking a number of references.-Author.

At the time when this taxonomic revision was undertaken Article 69 (3) of the International Code of Botanical Nomenclature was not in force.-Editor.

Page 360: Berberis and Mahonia : A taxonomic revision

360 3c

O O O Q

4 a 4b

013 5 a 5 b

36 2 a 2 b 2 c 3 a I A

4c 4d Q; 5 c 5d , 5 e

6c 6

Fig. 1. Stem section: a, terete (B. insignia); b, subterete (B. wallichiana); c, angled (B. manipurana); d , sulcate (B. sublewis).

Fig. 2. Leaf margin: a, closely spinulose subseda te (B. sieboldii); b, remotely spinose serrate (B. julknae); c, coarsely aristate dentate (B. veitchii).

Fig. 3. Laaf petiole: a, sessile (B. wikonae); b, decurrent to distinct petiole (B. virgetorum); c, con- tracted to articulate petiole (B. actinacantha).

Fig. 4. Leaf shape: a, ovate (B. umbellafa); b, oblong-elliptic (B. dziantha); c, obovate (B. chitria); (1, elliptic (B. manipuramz); e, Ianceolate (B. szlblevis); f, oblanceolate (B. wilsonae); g, linear- lanceolate (B. Zepidifolia); h, linear or linear-oblong (B. imolkta).

Fig. 5. Apex of Berberis leaves: a, mute (B. dielsiccluc); b, obtuse (B. diaphana); c, rounded (B. cir- cwnserrata); d, acuminate (B. gagnepaini var. lanceolata). Apex of Mahonia leaflets: e, M . japoniCa vm. trifurca; f , M . bealei; g, M . veitchimum.

Fig. 6. Forms of inflorescence: a, solitary (B. a n g u h a ) ; b, fascicled (B. hoookeri); c, umbellate (B. amoena); d, subumbellate (B. sieboldii); e, racemose-subumbellate (B. fmest i i ) ; f, pedunculata raceme (B. UuZgariS); 9. subsessile raoeme (B. &n&Zia); h, spike-like raceme (B. brachypda); i, raceme compound below (B. dielsiana).

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Berberia and Mahonia 361

Fig. 6 (cont.). j , panicle (B. prattii); k, short congested panicle (B. aggregata); I , corymbose panicle (B. chitria); m, lax raceme (Mahonia bngipes).

Fig. 7. Apex of petals: a, rounded, entire (B. dasyclada); b, obtusely emarginate with obtuse lobes (B. Zeptopoda); c, acutely emarginate with obtuse lobes (B. verruculosa); d, acutely emarginate with acute lobes (B . i w i g n k ) ; e, obtusely emarginate with acute lobes (B. Zudlowii var. saxi- cl~&la); j, acute, narrowly incised (B. k a n s u e k ) ; g, rounded, narrowly incised (B. veitchii). All x4.

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362 L. W. A. ABR.EX?DT 8 a 86

10a 10b 1oc

1% 12h

126

121

r 12j

11 a

9 6 9 c

11 b

I

12d 12e

j 12k 121 12m

9 d

8 11 c

12n 120

13b Fig. 8. Base of petals: a, truncate or cuneate, with separate submarginal glands (B. uertuculoaa);

b, clawed, with spproximate non-msrginal glands (B. &i). All x 4. Fig. 9. Apex of stamens: a, conspicuously apiculate (B. inOinii); b, shortly spiculate (B. o s m m h i i ) ;

c, truncate, produced (B. 53ewwxh8a); d, truncate, not produced (B. ta&&). All x 8. Fig. 10. Apex of filsmenta: a, slightly swollen, not dentate (B. h k e o k h ) ; b, shortly dentate (B.

c - r s i s vsr. hirszctapes); c, conspicuously lanceolate-dentate (B. chitria). In the very great majority of species there is neither tooth nor welling, the filaments are edentate. All x 8.

Fig. 11. O v a q snd ovulea : a, four ovules some sessile some subseasile or shortly stipitate (B. t a l i ~ ~ b ) ; b, four ovules distinctly stipitate with st& as long as the ovule (B. dasyduda); c, solitary ovule with long stalk (B. dlevia) . All x 8.

Fig. 12. Fruit and style: a, ellipsoid, estylose (B. thunbergi6); b, oblong, estylose, (B. orthbotrys); c, oblong, with ahort distinct style (B.julianae); d,globose, with long significant style (B. damkii); e, ovoid, with attenuate apex snd thick bent style (B. d iaphna) ; f, globose with thick bent style

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Berberis and Mahonia 363

(B. dictyopAyUa); g , narrowly oblong with distinct short style (B. chitria); A, i, j , narrowlyoblong and bent estylose (B. johmmk); k, small, oblong estylose (B. Zecomtei); 1, small, ovoid with short,

narrowly oblong (B. hookeri); 0, large ellipsoidal, estylm (B. Z a i ) . All x 1). Fig. 13. Pedicels and bracts: a, flower with medium-sized pedicel and short basal bract (B. mani-

p ~ a n a ) ; b, flower with short pedicel cmd bract subequal in length (B. aggregata); c, flower with short pedicel and conspicuously longer enveloping bract ( M a h i a veatchiorum); d, pedicels conspicuously slender at b e , thickening remarkably towards apex (B. imrm8&); e, bract situated a t middle of pedicel ( M a h i a pinnata). All x 14.

Fig. 14. Detail and cross-section of leaf: a (B. replicata), underside of leaf showing strongly revolute, spinose margin, x 8; b (B. replieata), cross-section of leaf, x 4.

Fig. 15. B. chitria, x .$ (no. 113a). Fig. 16. B. cm’aria, x 8 (no. 119). Fig. 17. B. umbellata, x .$ (no. 124). Fig. 18. B. kongboensis, x 8 (no. 131).

distinct style (B. ZepZifoZia); m, small, globose, with short distinct style (B. wilsonae 1; n. large,

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364 L. W. A. AHRENDT

Fig. 19. B. padam&, x ) (no. 37). Fig. 20. B. dhdha, x ) (no. 6). Fig. 21. B. &nsoZitcr, x 8 (no. 36). Fig. 22. B. indqni8 var. tongbe&, x 8 (no. 2d). Fig. 23. B. VE99YUdQ8a, X ) (no. 18). Fig. 24. B. deimzwdha, x ) (no. 44). Fig. 25. B. Zycioides: a, fruiting branch, x 8; b, stamen, x 4; 0, ovary and ovules, x 4 (no. 98). Fig. 26. B. glaucoeap , x Q (no. 99).

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Berberis and Mahonia 29 30

365

Fig. 21. B. ludlowii, x 3 (no. 136). Fig. 28. B. everestiana, x 8 (no. 138a). Fig. 29. B . tsanqpoem's, x ) (no. 141). Fig. 30. B. coneinna, x 8 (no. 140). Fig. 31. B. kumaonensis, x ) (no. 142). Fig. 32. B. ciroumserrata, x 3 (no. 144). Fig. 33. B. dictyophylla vw. campylogyna, x 8 (no. 153c). Fig. 34. B. john&, x 8 (no. 177). Fig. 35. B. cooperi, x 8 (no. 178). Fig. 36. B. minutiflora, x 8 (no. 190). Fig. 37. B. graminea, x 8 (no. 191). Fig. 38. B. amoena, x 8 (no. 193). Fig. 39. B. y u n n a n e d , x ) (no. 206).

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366 L. W. A. AE~RENDT

Fig. 40. B. forratii, x 8 (no. 226). Fig. 41. B. kphifolkz, x 8 (no. 224). Fig. 42. B. jameaiana, x 8 (no. 238a). Fig. 43. B. vernae, x 8 (no. 229). Fig. 44. B. &boldii, x + (no. 210). Fig. 45. B. didyoneura, x 8 (no. 207). Fig. 46. B. Prirgetorum, x 8 (no. 216). Fig. 47. B. koreana: a,fruiting branch, x 8; 6 , sterile shoot showingfolisceousspi, x 8 (no. 250).

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Berberis and Mahonis 367

Fig. 48. B. koehneam, x 8 (no. 285). Fig. 49. B. callwbotrys, x 8 (no. 309). Fig. 50. €3. giraldii, x 8 (no. 263). Fig. 51. B. creticu, x 4 (no. 308). Fig. 52. B. taylorii, x 8 (no. 296). Fig. 53. B. gyalaica (no. 297). Fig. 54. B. pseudumbellata, x 8 (no. 313). Fig. 55. B. ul ic im, x 8 (no. 318).

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368 L. W. A. ABRENDT 56 57a 57c 58

64

59

66 Fig. 56. B. actimmntha, x Q (no. 325). Fig. 57. B. cung~tijzOra: a, fruiting branch, x j; b, leaf from sterile stem, x Q; c, stamen, x 4 (no. 338). Fig. 58. B. Mfol ia , x Q (no. 348). Fig. 59. B. empetrifolia, x 8 (no. 357). Fig. 60. B. -am, x j (no. 362). Fig. 61. B. dam.nii, x j (no. 369). Fig. 62. B. mberi , x Q (no. 373). Fig. 63. B. Zinearifolia, x Q (no. 386). Fig. 64. B. virgaia, x 8 (no. 414). Fig. 65. B. confer&, x f (no. 427). Fig. 66. B. t d l l e n s d s , x Q (no. 451). Fig. 67. B. pindilicensds, x Q (no. 475).

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Berberis and Mahonia 369

Map 1. Berberis groups Septentrionales and Australes. Mahonia, groups Orientales and Occidentales.

I I I 1 I l l I 1 1 I I I I I 1 160 140 120 I00 80 60 40 20 0 20 40 60 80 100 120 140 160

Map 2. Berber&, sects. WALLICHIANAE, TINCTORIAE, INTEGERRIMAE, DASYSTACHYAE, VuLamm, CANADENSES and the localized BRACHYPODAE (B).

24 JOURN. LINN. SOC. BOTANY-VOL. LVII

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370 L. W. A. AHBENDT

I W

I I I I I I I I I I

Map 3. B e r b h , se~te. CBATAEQINAE, H~TEBOPODAE, ASIATICAE, the localized ULICINAE (U), FRANOHETIANAE and TSCHONOSKYANAE.

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Berberis and Mahonia 371

50

k5

I0

5

0

5

:0

24-2

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372 L. W. A. ATZRENDT

Map 5. I. Berbmis, subsects. INSIONES and HOOEERIANAE, the species indicated by numbers in circles. INSIONES: (1) daaycZuda, (2) iwignk, and vctrietiea a-e, (3) inerassata. and variety, a, (4) bypokedna. HOOK~IANAE: (5) hookeri, (6) eaU&&ha, (7) chrysosphaera, (8) osntastonii. 11. Weatarn species of other subsections of Berberis with their numbem in squares. REPUCATAE: (9) eollettii, (1O)p*, (ll)gri@h&na,(l2)amub&, (13)replieato. M~NIPIJRANAE: ( 2 1 ) m n i - puraw, (22) hchamnii, and variety, 5, (26) cozii. TRIACANTHOPHO-: (29)venwta. SWLEVES: (41) mbZwia. and varieties, a-c. ACCIMINATAE: (55) Zeptopoda. EUWALLICXKIANAE: (68) d i - &&na. (71) bhutanemia. LEVES: (73) poluninii, (74) mApe+oclada. SOULIEANAE: (87) tarone&.

Map 6. Berberis, subsects. REPLICATAE and VEI~EUCULOSAF,. Sect. REPLICATAE: (9) collettii, (10) p a e c i p u a , (11) griBthbna, (12) amdilis, (13) replica&, (14) t c c l i e k , (15) grodtmannb, (16) nantoemia, Subsect. VER~WCULOSAE: (17) candi~&, (18) verrucuZom, (19) aamyana.

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Berberis and Mahonia 373

Map 7 . Berberis, sects. MANJPURANAE and TRIACANTHOPHORAE: subsect. MANIPURANAE: (20) para- specta, (21) mnniprana, (22) hchananii, (22b) buchnanii var. tauangemis, (23) fallax, (25) wardii, (26) coxii, (27) mingetsenais, (28) impedita. Subsect. TRIACANTHOPHORAE : (29) venwta, (30) triacanthophora, (31) weitchii, (32) gagnepainii, (32b) gagnepainii var. suboosa+n, (3.1) phanera, (35) lubrh .

Map 8. Berberis, subsects. SANGUINEAE and SUBLEVES. Subsect. SANQUINEAE: (36) iwolita, (37) pan- lanewis, (38) dawidii, (39) sanguinea, (40) petrogena. Subsect. SWLEVES: ( 4 1 ) sublevis, (42) dolichostemon, (43) emilii.

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374 L. W. A. AHRENDT

I I 25

20

15

10

5

C

Map 9. Berber&, subsects. AG-ATAE and BBANDANAE. Subsect. Ac-ATAE : (44) dehmcadh, (45) valida, (46) arguh, (47) fdl.cio.a, (48) a&tut~-eenu2ata, (49) aczkrnhzfa, (50) d t h ~ ~ , (51) 8 i r n d . u ~ ~ . (52) silvicob, (53) bdcolw, (54) aubacUmina;to, (55) leptopoda, (56) xa&w&du. Subsect. BAIZANDANAE: (57) x a d w x y h , (58) baradam, (59) h&mG, (60) c d p k d ~ .

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Berberis and Mahonia 375

Map 10. Berberis, subsects. EUWALLICHIANAE and LEVES. Subsect. Euw-c-~ : (61) micro- petala (62) zunlamcianerrsis, (63) bergmunniae, (64) j d i a a e , (65) willeana, (66) dumicokz, (67) ferdinandicoburgii, (68) wallichiana, (69) sargentkm, (70) recumata, (71) bhutanensis. Subeect. LEVES: (72) brevisepakc, (73) poluninii, (74) mibpteroclada, (75) lwphylla, (76) wangii, (77) hob- eraepedan, (78) l e v k , (79) SubcOriaCeu.

Map 11. Berber&, subsect. SOULIEANAE: (80) muirei, (81) schneideriana, (82) atrocarpa, (83) s d h m , (84) lempergiana, (85) nemosa , (86) chingii, (87) tarorrenais, (88) hibberdiaw. Subsect. PRUINOSAE : (89) p i n o s a , (90) centi;fEora, (91) cawuleriei.

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376 L. W. A. AHRENDT

Map 12. Berber& distribution of the sect. ASIATICAX in detail, compared with that of sect. HETERO- PODAE in general: (92) asiatica, (93) heteracantku, (94) baluchktanicu, (95) Zycium, (96) khasaana, (97) parkeriana, (98) lycwides, (99) ghucocarp, (100) hain&i.

Map 13. Berber& group Septentrionalea: sect. TINCTOEIAE, and its subsects. POTILNINIANAE, E m - c - TORIAE, C ~ T R I A E , UMBELLATAE. B. hoktii and B . granatii (of subsects. CHITRIAE) and B . xanthoxyylon (of subsect. BARANDANAE are its only members of the Septentriondes south of the equator.

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Berberis and Mahonia 377

Map 14. Berber&, subsect. EUTINCTORIAE: (105) petwlaris, (106) huegeliana, (107) ceylanica, (108) nil- ghiriensis, (109) zvightiana, ( 110) timtoria, (1 11) a f s k n i ~ .

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378 L. W. A. ARRT~NDT

Mep 15. BerberiS, subaect. EUTINCTORIAE: (111) a f g k n h , subsect. CEITELAE: (112) naioranlk, (113) chit7i0, (114) sikkirnmreis, (115) ceratqphylla, (116) amk, (117)jlorjbunda, (118) a w , (119) co*iaria, (120) h o W , (121) gmntii, (122) petitiana, (123) amw.

Map 16. Berber&?, subsects. UMBEZLATD and L I N D L J z Y ~ A E : (124) umbdldcr, (125) brandisigno, (126) waltmanu, (127) dtt$h&mtau, (128) tiurmsOniana, (129) kashmiranu, (131) k o n g b o d . (132) lindhyatau, (133) forskdiancl.

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Berberis and U o n i a 379

Map 17. Berberis, sect. ANQULOSAE and its subsecta. CONSIMILES, DI~PHILNAE, DICTYOPHYLLAE, EUANOULOSAE, JAESCIXKEANAE, SIBIRICAE and SWANQULOSAE.

90 95

Map 18. Berberb, subsect. EUANQULOSAE: (134) anguloaa, (135)parbep&, (136) Zudlowii, (137) macro- s e m , (138) everestiana, (139) erythroclada, (140) wncinna, (141) tsangpoensaS, (142) k u - 6 , (143) temolaica.

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380 L. W. A. AERENDT

Map 19. Berber&, subseota. DIAPHILNAE, DICTYOPHYLLAE and SIBIRICAE. Subsect. DIAPHANAE: (144) Circunzserrata, (145) dkphana, (146) aernuhm, (147) wri-8omd8, (148) t k c h h % (149) virea- cem, (150) elliotii. Subsect. DICTYOPHYLLAE: (152) approxha% (153) d&%yophyhYurr, (154) ambro- zyano, (155) s t W z b n o , (166) md&?&, (157) d d k e p a h , (158) &*QhWrn. Subsect. SIBIRICAE : (159) boreali-sdnensis, (160) &&a, (162) &ark.

Map 20. Berbek, subsect. JAESCHEEANAE: (163) burwnica, (164) lasaockma, (165) apkulata, (166) zayuhmz, (167) hmbertii, (168) mtmifolia, (169) hobsonii, (170) kartanica, (171) h i d i m , (172) hrnilhmiana, (173) jaeachkeana.

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Berberis and Mahonia 381

40

30

20

70 80 90 100 Map 21. Berber&, subsect. SUBANQULOSAE: (176) trdChohaem&o&s, (177) johnnk, (178) cooperf,

(179) orthobotrya, (180) arnbdgua, (181) @nor&, (182) platyphylh, (183) 8uberwt.a.

Map 22. Berberis, subsect. CONSIMILES: (186) consim%s, (187) fmon&w, (188) puru?%rescena, (189) Shf?W&9lU.

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382 L. W. A. ~ D T

Map 23. Berber&, sect& FBANCHETIANAE, TSCI~ONOSKYANAE, and SIEBOLDXAE.

Map 24. Berberb, subsecte. M ~ ~ ~ G O R A E and EU"CE~TANAE. Subsect. MINWJ!IFLOEAE: (190) m+n4Ma, (191) g r a m h . Subsect. EURRANCHETIANAE: (192)fracnchetb, (193) m e w , (194) thibetiea, (196) teeonztei, (197) tsarongerrsis, (198) bodmni+,

Page 383: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 383

Map 25. Berberis, subsect. YUNNANWSES: (200) t o m e & h a (201) pap'Uqera (202) microtricha, (203) rnehmgen&, (204) humido-umbrosa, (205) dawoensks, (206) yunnanensis, (207) dictyomura, (209) bracteata.

Map 26. Berberk, group Septentrionales: subsect. SIEBOLDIAE: (210) Sieboldi, (211) re tkdfh , (212) dubia, (213) oritrepha, (214) quelpaer ted .

Map 27. Berberis, sect. TSCHONOSKYANAE: (215) mou&zcana, (216) virgetorum, (217) thunbergii, (220) 8ilva-taroucclna. (221) tsclumoslcyama.

Page 384: Berberis and Mahonia : A taxonomic revision

384 L. W. A. AEEGENDT

5'

4

3

I Map 28. Berber&, sect. CANADENSEB.

Map 29. Berberis, sect. SINENSES: (224) kpidtfolb, (226) p a l h 8 , (226) fmmti i , (227) o h i w k , (228) poire+%.

Page 385: Berberis and Mahonia : A taxonomic revision

386 Berberia and Mahonia

Map 30. Berber&, sect. INTEQERBIMAE: (229) vermoe, (230) purdornii, (231) caroli, (232) &n&bra. (233) twunnannica, (234) integerrim, (235) ilienaiS, (236) nZMnntula740, (237) francisci-ferdinandi, (238) j a m d a m .

Map 31. Berberis, derwi&n-a and turunnannk: (232a) densi,fEora vaz. &ma&n-a, (232b) dem&wa vw. mamacantha, (232c) &&&n-a var. macrobotrys, (232d) &nsi,lEora var. SeWatifOlda, (234) in- tegerrima, (236) nummuhria.

25 JOURN. LINN. SOC. BOTANY-VOL. LVII

Page 386: Berberis and Mahonia : A taxonomic revision

386 L. W. A. ~ D T

Map 32. B e r b b , sect. DASYSTACEYAE: (239) kamue&, (240) &el&-, (241) dolkhobotrys, (242) fed- deana, (243) daeystachya, (244) zabdiana, (245) chhng&, (246) anhweknsis, (247) honanedn.

Msp 33. B e r b b , sect. V a a a a ~ s : (248) oriental&, (249) corrcolor. (260) kwmm, (251) vulgaris, (253) bretschnekderi, (254) pmhyaeantha, (255) hnryana, (256) farre?& (257) &la, (258) atro- pranina, (259) m u r d , (261) regdkm.

Page 387: Berberis and Mahonia : A taxonomic revision

Berberis a d Mahonia 387

25-2

Page 388: Berberis and Mahonia : A taxonomic revision

388 L. W. A. ABRENDT

Bbp 36. Berberb, subsects. E U P O L Y ~ A E , METAPOLYA”EAE: (273) gdyantha, (274) pm-i, (275) aqgrqata, (277) brewipataidda, (278) oblOmwAtu, (279) mdwphkaea. (280) SO~octijEO*o, (281) beankna, (282) mefapolyan8hn.

34

32

30

Map 36. Berber48, subsect. SWPOLYANTHAE: (285) h e h w m , (286) &emhiancl., (287) hmuwr&.

Page 389: Berberis and Mahonia : A taxonomic revision

389 Berberis and Mahonia

Map 37. Berber&, subsect. PSEUDOPOLYANTHAE: (289) ariok-ealida, (290) heteropk, (291) hemleyanu, (292) b&m, (293a) wilaonae var. wi€.sonae, (293b) wilsong var. pamrifolia, (294) & m e var. .subdiahta, (294 a) wilsoni var. guhtzuniea, (294 b) Berber& x ‘Comet ’.

30 30

29 29

28 28

27 27

Map 38 Berberis, seot. SHERRIFFIANAE: (296) taylorii, (297) q y h i m , (298) sherrifli.

Page 390: Berberis and Mahonia : A taxonomic revision

Map 40. Berberis, sects. HETEROPODU and ULICINAE. HETEROPODAE (CRETICAE); (307) oblonga (308) wetdcea, see Map 39, (309) mllwbotrys, (310) wam&fanim, (311) ga?nbhm. HETEROPODAE (PSEUDUMBELLATAE): (312) stolont7era, (313) peududeuat0, (314) royleana, (315) aitchisonii, (316) heteropoda. ULICINAE: (318) uZiOina, (319) kocrc7tgark-z.

Page 391: Berberis and Mahonia : A taxonomic revision

Berberis and Mahonia 391

Map 41. An extension of map 40 showing the full range of B. heteropodu and B. oblonga. The remaining spp. of the sects. HETEROPODAE, and ULICINAE ere shown by asterisks.

Page 392: Berberis and Mahonia : A taxonomic revision

392

0

5

LO

i5

40

45

50

55

Page 393: Berberis and Mahonia : A taxonomic revision

393 Berberis and Mahonia

Map 43. Berberis, sect. LA-NAE: subsects. EULAURINAE and FLEXUOSAE. EULATJRINAE: (387) hu- rim, (388) claussenii, (389) ghwxacem, (390) Coriacea, (391) adreuna, (392) ZdtoraZis, (393) ci temi. FLEWOSAE: (394) $emmsa, (395) ynmpemm, (397) valdkviana, (398) seUowio?ta, (400) corn- mu&&, (401) divarimfa, (402) w e d d d h a , (404) pewvhna, (405) pseudo-spkdosa, (406) spkulosa, (407) amnata.

Page 394: Berberis and Mahonia : A taxonomic revision

t

09

I

D

!i

OC

I 1 OL SL

&a~5x~jT "fr 'M '7 V6€

Page 395: Berberis and Mahonia : A taxonomic revision

Berber& a d Mahonia 395

Page 396: Berberis and Mahonia : A taxonomic revision

396 L. W. A. ~ D T

45

40

35

3c

f5

35

30

Map 48. Mahunk, eect. AQUIFOLIATM: subsect. EUAQUIWOLIATAE.

Page 397: Berberis and Mahonia : A taxonomic revision
Page 398: Berberis and Mahonia : A taxonomic revision

398 L. W. A. AHRENDT

Nap 51. Mahonkz, sect. HOIWDAE.

Page 399: Berberis and Mahonia : A taxonomic revision

Index

Genus Berberis L. 26-295, 357-9

Illustrations are con6ned to pp. 360-8. Appendices 1 (p. 294) and 2 (p. 295) comprise alphabetical lists of species

Serial numbers of species are given in parentheses, for example (325)

bracts and pedicels, 363 (Fig. 13) diagnostic characters of, 2 filaments, apex of, 362 (Fig. 10) fruit and style, 362 (Fig. 12) geographical distribution of, 21, 369-98 (maps) inflorescence, forms of, 360 (Fig. 6) keys, use and nature of, 24 leaves: apex of, 360 (Fig. 5); cross-section of, 363

(Fig. 14); margins, 360 (Fig. 2); petioles, 360 (Fig. 3); shapes, 360 (Fig. 4)

ovary and ovules, 362 (Fig. 11) petals, apex of, 361 (Fig. 7); base of, 362 (Fig. 8) stamens, apex of, 362 (Fig. 9) stem, sections of, 360 (Fig. 1)

Groups, sections and subsections Actinacanthae Schneid. (sect.), 229 Acuminatae Ahrendt (subsect.), 59 Aequinoctiales Ahrendt (subgroup), 264 Agapatenses (Schneid.) Ahrendt (sect.), 272 Agap&ense-s Schneid. (subsect.), 272 h g d o s a e Schneid. (sect.), 108 Asiaticae Schneid. (sect.), 85 Australes Schneid. (group), 229 Barandanae Schneid. (subsect.), 63 Brachypodae Schneid. (sect.), 195 Buxifoliae Schneid. (sect.), 239 Canadenses Ahrendt (sect.), 170 Candid.rclae Schneid. (subsect.), 45 Chilenses Schneid. (subsect.), 254 Chitriae Schneid. (subsect.), 95 Confertae Ahrendt (sect.), 268 Congestiflorae Schneid. (subsect.), 234 Consimiles Ahrendt (subsect.), 147 Corymbosae Schneid. (sect.), 237 Crataeginae Ahrendt (sect.), 220 Creticae Schneid. (subsect.), 224 Darwinianae Ahrendt (subsect.), 249 Dasystachyae Schneid. (sect.), 182 Diaphanae Ahrendt (subsect.), 121 Dictyophyllae Ahrendt (subsect.), 126 Empetrifoliae Schneid. (subsect.), 244 Euactinacanthae Schneid. (subsect.), 230 Euangulosae Schneid. (subsect.), 112 Euaustrales Ahrendt (subgroup), 229 Eubuxifoliaa Schneid. (subsect.), 240 Eufranchetianae Ahrendt (subsect.), 152 Euilicifoliae Ahrendt (subsect.), 251 Eulaurinae Schneid. (subsect.), 257 Eupolyanthae Ahrendt (subsect.), 201 Eusilaenses Schneid. (subsect.), 170 Eutinctoriae Schneid. (subsect.), 93 Eutruxillenses Schneid. (subsect.), 279 Euvirgatae Schneid. (subsect.), 265

Groups, sections and subsections (CON.)

Eutndgarea Schneid. (subsect.), 182 Euwallichianaa Schneid. (subsect.), 66 Fhxwsae Schneid. (subsect.), 259 Flexuosae (Schneid.) Ahrendt (subsect.), 259 Franchetianae (Ahrendt) Ahrendt (sect.), 151 Franchetianae Ahrendt (subsect.), 151 Goudotiae Schneid. (sect.), 275 Heteropodae Schneid. (sect.), 223 Hookerianae Ahrendt (subsect.), 39 Ilicifoliae Schneid. (sect.), 249 Insignes Schneid. (subsect.), 36 Integerrimae Schneid. (sect.), 174 Jmschkanae Ahrendt (subsect.), 133 Keisslerianae Schneid. (subsect.), 285 Latifoliae Schneid. (sect.), 264 Laurinae Schneid. (sect.), 256 Leves Ahrendt (subsect.), 72 Lindleyanae Ahrendt (subsect.), 106 Manipuranae Ahrendt (subsect.), 46 Metapolyanthae Ahrendt (subsect.), 206 Microphyllae Ahrendt (sect.), 236 Minutifloras Ahrendt (subsect.), 151 Montanae Ahrendt (sect.), 245 Panicdatae Schneid. (sect.), 286 Pichinchenses Schneid. (subsect.), 282 Polyanthae Schneid. (sect.), 200 Potaninianae Ahrendt (subsect.), 92 Pruinosaa Schneid. (subsect.), 80 Pseudopolyanthm Ahrendt (subsect.), 212 Pseudumbellatae Ahrendt (subsect.), 226 Quindiuenses Schneid. (sect.), 273 Rectinerviae Ahrendt (subsect.), 267 Replicatae Ahrendt (subsect.), 41 Sanguineae Schneid. (subsect.), 55 Septentrionales Schneid. (group), 28 Sherrfianae Ahrendt (sect.), 217 Sibiricae Ahrendt (subsect.), 131 Sieboldiae Ahrendt (subsect.), 163 Sinenses Schneid. (sect.), 171 Soulieanae Ahrendt (subsect.), 76 Subangulosae Ahrendt (subsect.), 140 Subleves Ahrendt (subsect.), 57 Subpolyanthae Ahrendt (subsect.), 209 Tinctoriae Schneid. (sect.), 91 Triacanthophorae Schneid. (subsect.), 50 Trigonae (Schneid.) Ahrendt (sect.), 255 Trigonae Schneid. (subsect.), 255 Truxillenses Schneid. (sect.), 278 Tschonoskyanae Schneid. (sect.), 165 Tschonoskyunae Schneid. (subsect.), 165 Ulicinae Schneid. (sect.), 228 Umbellatae Schneid. (subsect.), 104 Verruculosae Ahrendt (subsect.), 45

Page 400: Berberis and Mahonia : A taxonomic revision

400 IndeX

Genus Berberis L. 26-295, 357-9 (cont.)

Groups, sections and subsections (wnt.) Virgatae Schneid. (sect.), 265 Vulgares Schneid. (aect.), 186 Wallichianae Schneid. (aect.), 30 Yunnsnenses Ahrendt (sub-sect.), 157 Yunmnensea Ahrendt (subsect.), 140

Spedes and varieties actinacantha Mart. (325), 231, 368 (Fig. 56)

var. c&pa (C. Gay) Reiche, 235 var. gr&nu (Gill.) Schneid., 231 var. horrida (C. Gay) Reiche, 236

acuminata Franch. (as), 61

czcuminda Veitch, 51 acertifolio Prantl, 188 aemulans Schneid. (146), 124 aetnemis presl(302). 222

var. aetnensis (302a). 222 var. oslebrioa Schneid. (302b), 222

c z c u m i e Stepf, 53

affinis G. Don ( l l s ) , 100 afghanica Schneid. ( l l l ) , 95 africana Hebemtr. et Ludw. ex Schultea f. (123),

agapatensis Lechl. (434), 273 aggregate Schneid. (275). 203 var. aggregate (275u), 203 var. integrifolia Ahrendt (275b), 203 var. pmttii Schneid., 202 var. rmmiata Schneid., 202

agricola Ahrendt (257), 192 sitChis0nii Ahrendt (315), 227 aldenhamenais Ahrendt (495), 294 x alksuthienais Ahrendt (252f). 191 alpicola Schneid. (60), 66 amabilis Schneid. (12). 43 ambigua Ahrendt (180), 144 ambrozyana Schneid. (154), 128 amoena Dum (193), 154, 365 (Fig. 38)

var. amoena (193a), 154 var. moloenaia Ahrendt (193c), 154 var. umbeMors Ahrendt (193b). 154

var. amuremis (259a), 193 var. j u m k (Regel) Rehd., 194 var. latifolia Nakai (259b), 194 var. licentii Ahrendt (259~) . 194

104

amuremia Rupr. (259), 193

andeana Job (496). 295 andreana Naudin (391), 258 undrkuxii Hook. et Am., 345 angdosa Wall. ex Hook. f. et Thoms. (134), 113

var. angulosa (134u), 113 var. hv6pe.a Franch., 151 var. faaciculata Ahrendt (134b). 114 var. Hook. f. et Thoms., 114

ungwtifdiu Hartw. ex Benth., 342 anhweienwia Ahrendt (246), 185 anniae Ahrendt (104), 93 x antoniana Ahrendt (366), 247 antucoana Schneid. (330), 233 apiculata (Ahrendt) Ahrendt (l65), 135 approximata Sprsgue (152), 127

var. mrnpyzogym Ahrendt, 128

Spedes and varieties (cod.) uqu+jo&m Purah, 336 mp&a (Hutch.) Bean, 350 arguta (-oh.) Schneid. (46), 60 arido-dida W d t (289), 213 wistota collett, 97 aristete DC. (118), 101 wistota Parker, 90

var. rniorontha Hook. f. et Thorns., 96, 99 var. pmv$mva Hook. f. et Thorns. MS., 99

aristato-Berrulste Hayats (48). 61 armete Citeme (407), 263 d u t k G S t h , 43 &tiice Roxb. ex DC. (92), 86

var. asiatica (920). 86 var. clarkema Schneid. (92b), 87

asmyana Schneid. (19). 46 atrocerpe Schneid. (82), 77

VW. 8 t r O C q ( 8 2 ~ ~ ) . 77 var. longiptw Abrendt (82c), 77 var. mbintegra Ahrendt (82b), 77

atropraeina Ahrendt (258), 193 aurahutwed L e e (440), 275 x ‘Aurora’ Chittenden ex Ahrendt (283d), 209 uzurtralis Moris ex Schneid., 221

var. haekdiuna Schneid., 222 x ‘AutumnBeeuty’ Chittendenex Ahrendt (284),

x ‘Autumn Cheer’ Chittenden ex Ahrendt (283b).

baluchistanh Ahrendt (94), 87 barandana Vidal(58), 65 x ‘Barbaroese’ Watson ex Ahrendt (283g), 209 barbey- Schneid. (485), 291 berilochemis Job (497). 295 bedei Fort., 319 beaniana Schneid. (281). 206 beauverdiena Schneid. (479), 290 beesisns Ahrendt (292), 214

var. blxsuma . (292a), 214 var. glabrs Ahrendt (292b). 214

xbella Sohneid. (262b). 195 benoistiens Macbride (435), 273 bergeriana Schneid. (458), 283 bergmanniae Schneid. (63), 67

var. acanthophyb Schneid. (63b), 68 var. bergmanube (63a), 67

bhutanensie Ahrendt (71), 72 bicolor L6veU6 (53), 62 bidentata Lechl. (329), 233 boisaieri Schneid. (303), 222 boliviana Lecbl. (428), 271 x bornbycine Ahrendt (272b), 199 boreali-sinemis Nakai (159), 131 boschanii Schneid. (198), 157 x ‘Bountiful’ A. R. Ahrendt ex Ahrendt (2831),

brechyamntha Phil. ex Reiche (320), 231 brechybotria C. Gay (380), 254 bruchybotrys Edgeworth. 211 brachypoda Maxim. (265). 197

209

208

209

var. gihaeii Hort., 198 var. 8alieOria; (Fedde) Schneid., 197

Page 401: Berberis and Mahonia : A taxonomic revision

IdCX

Genus Berberia L. 26-295, 357-9 (wnt.)

40 1

Species and varieties (wnt.) bracteata (Ahrendt) Ahrendt (209), 163 brandisiana Ahrendt (125), 106 bretschneideri Rehd. (253), 191 brevifolia Phil. ex Reiche (323), 231 brevipaniculata ScIineid. (277), 204 brevipeS (Franch.) Schneid., 151 x breviscapa (Ahrendt) Ahrendt (208e), 163 breviseqala Hayata (72), 72 x ‘Brilliant’ (252d), 190 X bristolensis Ahrendt (6a), 40 brumalis Macbride (468), 285 x ‘Buccaneer’ Watson ex Ahrendt (283e), 209 buceronis Macbride (447), 278 buchananii Schneid. (22), 48

var. buchananii (22a), 48 var. tawangensis Ahrendt (22b), 48

bullata Ahrendt (24), 49 bumeliaefolia Schneid. (489), 292 burmanica Ahrendt (163), 134 buxifolia Lam. (348), 240, 368 (Fig. 58)

var. antarctica Schneid. (348d), 241 var. buxifolia (348a), 240 var. inermis (Persoon) Schneid. (348c), 241 var. m.mracantha Phil. ex A. Usteri, 241 var. nana A. Usteri (348b), 241 var. nuda Schneid. (348e), 241 var. papillosa Schneid. (348f), 241 var. 8 p * k & m u Reiche, 241

cabrerae Job (360), 245 caespdtosa Madden MS., 120 calcipratorum Ahrendt (158), 130 califmica Jepson, 335 calliantha Mulligan (6), 40, 364 (Fig. 20) calliobotrys Aitch. (309), 225, 367 (Fig. 49) csmpbellii Ahrendt (124c), 105 campos-portoi Brade (352), 242 canadensis Mill. (222), 170 candidula Schneid. (17), 45 ca-pilhrk Cox ex Ahrendt, 115 caraganaefolicl DC., 331 carinata Lechl. (412), 265

var. carinata (412a), 266 var. echinata Diels (412b), 266

x carminea Chittenden ex Ahrendt (283), 208 caroli Schneid. (231), 176

var. hoanghnaiS Schneid., 175 cavaleriei LBveill6 (91), 83

var. p i r w s a Byhouwer, 79 centiflora Diels (go), 82 x cerasina Schrad. (252e), 191

ceratophylla G. Don (115), 100 ceylanica Schneid. (107), 94 chekiangensis Ahrendt (246), 185 x chenaultii (Hort. ex Cat. L. Chenault) Ahrendt,

x ‘Cherry Ripe’ A. R. Ahrendt (185), 147 chilensis Gill. ex Hook. (381), 254

var. diflusa (C. Gay) Reiche, 255 var. feros (C. Gay) Reiche, 255

var. chillanenais (363a), 246 var. hirsutipes Spregue (363b), 246

n.n. (32h), 54

chillanensis (Schneid.) Sprague ex Bean (363), 246

26

Species and varieties (COW.)

x chilternensis Ahrendt (31a), 52 chimboensis Schneid. (466), 285 chinensis Poir. (227), 173

var. chinensis (227a), 173 var. paphlagonica (Schneid.) Ahrendt (227b),

174 chingii Cheng (86), 79 chitria Lind. (113), 97, 363 (Fig. 15)

var. chitria (113a), 97 var. occidentalis Ahrendt (113b), 97 var. sikkimnais Schneid., 99

chrysacantha Schneid. (431), 272 chrysosphaera Mulligan (7), 41 ciliaris Lindl. (436), 274

chochoco schldl., 345

var. ciliaris (436a), 274 var. obtusata Ahrendt (436b), 274

var. circumserrata (1444, 122 var. occidentalior Ahrendt (144b), 122 var. subarmata Ahrendt (144c), 123

circumserrata Schneid. (144), 122, 365 (Fig. 32)

citernei Ahrendt (393), 259 claussenii Citerne (388), 258 cliffortioides Diels (419), 268 x colburyensis Marchant (282a), 207 coletioides Lechl. (364), 246

var. coletioides (364a), 246 var. gracilis (Schneid.) Ahrendt (364b), 247

collettii Schneid. (9), 42 colombiana Ahrendt (473), 288 comberi Sprague et Sandwith (373), 252, 368

x ‘Comet’ Chittenden ex Ahrendt (294b), 216 commutata Eichler (400), 262 conciqnaHook.f.etThoms. (I40),118,365(Fig. 30)

(Fig. 62)

var. brevior Ahrendt (140b), 119 var. concinna (140a), 119 var. extensiflore Ahrendt (14Oc), 119

concolor W. W. Sm. (249), 187 conferta Kunth (427), 270, 368 (Fig. 65)

var. boli&na (Lechl.) Schneid., 271 var. conferta (427a), 271 var. hypopyrrantha Schneid. (427c), 271 var. karsteniana Schneid. (427b), 271 var. lobbkna Schneid., 271 var. pdocluda Schneid., 272 var. qwrweana Schneid., 269

congestiflora C. Gay (338), 235, 368 (Fig. 57) var. hukeoades Hook. f., 236

cmsanguim Fort., 320 consimilis Schneid. (186), 148 cooperi Ahrendt (178), 142, 365 (Fig. 35) coquimbensis P. Muiioz (333), 234 x ‘Coral’ Chittenden ex Ahrendt (294c), 216 Coriaceu Brandis, 90 coriacea St. Hil. (390), 258

var. coriacea (390a), 258 var. oblanceifolia Ahrendt (390b), 258

&riu Collett, 90 coriaria Royle ex Lindl. (119), 102, 363 (Fig. 16)

var. coriaria (119a), 102 var. patula Ahrendt (1 19b), 102

JOURN. LINN. SOC. BOTANY-VOL. LVII

Page 402: Berberis and Mahonia : A taxonomic revision

402 I*

Genus Berberis L. 26-295, 357-9 (cont.)

Species and varieties (cod.) coryi Veitch (282 b), 207 corymbosa Hook. et Am. (344). 238

var. corymbom ( 3 4 4 ~ ) . 238 var. panicdata Phil. (344b), 238

costdata Candoger (371), 251 coxii Schneid. (26). 49 cMIteegina DC. (300), 220

var. armeniaca Schneid. (300b), 221 var. uabulh Schneid., 227 var. crataegba (300a), 221 var. lycica Schneid. (300~) . 221

x crenulata Schcad. (251e). 189 cretica L. (308). 225, 367 (Fig. 51) crimson Bead’ A. R. Ahrendt ex Ahrendt

criSpa C. Gay (337), 235 cuneata DC. (353). 243 darwinii Hook. (369), 250, 368 (Fig. 61)

(277b), 205

var. darwinii (3690), 250 var. macroplbyUa Hort., 247 var. magellanice Ahrendt (3693). 251

daayclada Ahrendt (l) , 36 dwystechya Maxim. (243). 184 davidii Ahrendt (38). 56 dawoensis K. Meyer (205), 161 dowsonii Bsan, 167 dedbata Lindl. (449), 280 decandolleana Ahrendt (130). 107 x declineta Schrad. (2080). 162 var. oxyphylla Schneid. (208d). 163 dehacantha Schneid. (44), 60, 364 (Fig. 24)

var. oolido Schneid.. 60 dehvayi Schneid. (50), 61

var. dehvayi (500). 61 var. wechinensis Ahrendt (50b), 61

&ma Schneid., 56 dens& Tr. et P1. (459), 283 densiflora Bob. et Buhee (232), 176 de&*a Wendelbo, 227

var. bungeana Ahrendt (232e), 177 var. densiftore (232a), 176 var. mamacantha Boiaa. (2326). 176 var. mscrobotrys Ahrendt (2320). 177 var. serratifolia Boiss. (232d), 177

densifolia Byhouwer, 44 densifolia Rusby (472). 286 diaphana Ildaxim. (145), 123 diaphatna Rehd., 123

var. Circurn8ewatu Schneid., 122 var. ditbphana (145a), 123 VBr. teChiensk Ahrendt (145b), 123 var. d o a Ahrendt (145~) . 124

dictyoneura Schneid. (207), 161, 366 (Fig. 45) var. bra&& Ahrendt, 163

dictyophylla Franch. (153), 127 dictyophyk Hook. f., 127

128, 365 (Fig. 33) var. campylogyna (Ahrendt) Ahrendt (153e),

var. dictyophylla (153a), 127 var. epruinosa Schneid. (153b), 128

dictyotu Jepson, 334 dielsiana Fedde 1240). 183

Spedes and varieties (cont.) diffusa C. atry (384), 255 discolor Turcz. (441), 276 divaricata Rusby (40l), 262 dolichobotrya Fed& (241), 183 dolichcatemon Ahrendt (42), 59 dryandriphylla Diels (455), 281 dubis Schneid. (212). 164 duloie Pexton, 241 dulcis Sweet, 240 dumicola Schneid. (66), 69 x durobrivensis Schneid. (228a), 174 duthitwm Schneid. (127), 106 edentata Rusby (454), 281 edgeworthiam Schueid. (286), 211 ehrenber~i Kunze, 341 eIegan.9 LBveill6, 324 eikgam (Franch.) Schneid., 154 elliotii Ahrendt (NO), 126 x emarginata Willdenow (161), 132

emilii Schneid. (43), 59 x ‘Emperor’ Chittenden, n.n. (183a), 146 empetrifolia L8m. (357), 244, 368 (Fig. 59)

var. empetrifolia (357a), 244 vm. magellanica Schneid. (357 b), 244

vm. britmx@ Schneid. (161b), 132

engleriana Schneid. (462). 284 erythrocleda Ahrendt (139). 118 var. erythroclada (139u), 118 var. trulungensis Ahrendt (I39b), 118

&+hyk (Fedde) C. H. Muller, 342 everestiana Ahrendt (138). 116, 365 (Fig. 28)

var. evematism ( 138 a;), 1 17 var. nambuensis Ahrendt (138b). 117 var. ventosa Ahrendt (138c), 117

faberi S h e i d . (151), 126 fallmiom Schneid. (47), 60 fallax Schueid. (23). 49 farinosa Benoist (448), 280 farreri Ahrendt (256), 192 f u w a W. W. Sm., 215 faxonitma Schneid. (187), 148 feddeans Schneid. (242), 184 fendleri Gray (223), 171 x ‘Ferax’ Chittenden ex Ahrendt (295b), 217 ferdinandi-coburgii Schneid. (67). 70 var. ferdinandi-coburgii (67a). 70 var. vernalis Schneid. (67b), 70

ferox C. Gay (382), 255 fiebrigii Schneid. (481). 290 fi.eti0‘ Schneid., 158 x ‘Fireball’ Chittenden ex Ahrendt (294e), 216 x ‘Fireflame’ Chittenden ex Ahrendt (283c), 208 x ‘Firefly’ Chittenden ex Ahrendt (2943), 216 fbxuosa R. et P. (394), 260 floribunda Wall. ex G. Don (117), 100 var. afinia Ahrendt, 100

florida Phil. (321). 231 formosana Ahrendt, 65 fornw8ana Li, 358 forrestii Ahrendt (226), 172, 366 (Fig. 40) forskaliene Schneid. (1331, 108 . ,. . I _ fortzcnei Lindl., 328

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Genus Berberis L. 26-295, 357-9 (eont.)

403

Species and varieties (cant.) fragrans Phil. ex Reiche (383), 255 franchetiana Schneid. (192), 153

var. franchetiana (l92a), 153 var. glabripes Ahrendt (192b), 153 var. mecrobotrys Ahrendt (192c), 153

francisci-ferdinandi Schneid. (237), 179 fremntii Torr., 353 gagnepainii Schneid. (32), 52

vm. Nipes Ahrendt (32f), 53 var. gagnepainii (32a), 52 var. lanceifolia Ahrendt (32d), 53 var. lanceifolia f. pluriflora Ahrendt (32e),

vm. omeiensk Schneid. (32c), 53 var. praastans Ahrendt (32g), 54 var. subovata Schneid. (323), 52 Tottenham var., 45

53

gambleana Ahrendt (311), 225 ganp’mmk Lbvei116, 3 16 garciae Pau (304), 223 gayi Citerne, 259 gayi K. Koch, 254 x gibbsii Ahrendt (272a), 199 gilgiane Fedde (267), 197 gilgiana Schneid., 198 giraldii Hesse (263), 196, 367 (Fig. 50) ghum Benth., 289 glauca Kunth (484), 291 glaucescens St. Hil. (389), 258 glaucocarpa Stapf (99), 90, 364 (Fig. 26) ghioviana Brade (613), 357 globosa Benth. (432), 272

var. &ma Schneid., 283 glomerata Hook. et Am. (328), 232

var. glomerata (328a), 232 vm. zahlbruckneriana, (Schneid.) Ahrendt

(32883, 232 g l u m Spreng., 329 goudotii Tr. et PI. (446), 277 gracilipm Oliver, 326 grmdk Hartw. ex Benth., 343 graminea Ahrendt (191), 152, 365 (Fig. 37) grandibracteata Ahrendt (288), 212 grandiflora Turcz. (439), 275 grantii Ahrendt (121), 103 grevilleana Gill. ex Hook. (324), 231 griffithiana Schneid. (1 l) , 43 grodtmannia Schneid. (15), 44

vsr. flavoramea Schneid. (153), 44 var. grodtmannia (15a), 44

xguernseyensis Ahrendt (331), 233 guilsche Tr. et PI. (443), 276 gyalaica Ahrendt (297), 218, 367 (Fig. 53)

var. gyalaica (297a), 218 var. maximiflora Ahrendt (297b), 218 var. minuat8 Ahrendt (297c), 218

haenmtocarpa Wooton, 354 haenkeana Presl ex Schult. (411), 265 hainesii Ahrendt (loo), 90

va,r. brevifilipes Ahrendt (100b), 91 var. hainesii (100a), 91

hakeoides (Hook. f.) Schneid. (339), 236

Species and varieties (wnt.) hallii Hieron. (399), 261

var. hallii (399a), 261 var. typka Schneid., 261 var. wagneriana, Schneid. (3993). 262

h d t o n i a n a Ahrendt (172), 137 hartwe& Benth., 349 x hauniensis Schneid. (113d), 98 havatana Mizushima (41 e), 358 helenae Ahrendt (34a), 55 hemsleyana Ahrendt (291), 213 he7nsleyi Donn. Smith, 348 henryana Schneid. (255), 192 hersii Ahrendt (260), 194 heteracantha Ahrendt (93), 87 heterophylla Juss. ex Poir. (332), 233

var. pZur$ora Reiche, 232 heteropoda Schrenk (316), 227

var. heteropoda (316a), 227 var. o b h g u Regel, 224 var. sphaerocarpa (Kar. et Kir.) Ahrendt

(316b), 227. heteropsis Ahrendt (290), 213 hibberdiana Ahrendt (88), 79 hieronymi Schneid. (347), 240 higgimm Munz, 356 himslaica Ahrendt (171), 137 hirbllipes Ahrendt (465), 284 hispanica Bob. et Reut. (301), 221

var. hackeliana (Schneid.) Ahrendt (301 b), 222 var. hispanica (301a), 222

hobsonii Ahrendt (169), 137 hochreutinerana Macbride (492), 293 holocraspedon Ahrendt (77), 74 holstii Engler (120), 102 honanensis Ahrendt (247), 185 hookeri Lemaire (5), 39

var. candidula Schneid., 45 var. hookeri (5a). 39 var. kztijolia Bean, 40 var. microcarpa Ahrendt (5c), 40 var. platyphylla Ahrendt (54 , 40 var. viridis Schneid. (5b), 40

horrida C. Gay (340), 236 horrida Junghuhn, 64 huanucensis (Schneid.) Macbride (413), 266 huegeliana Schneid. (106), 94 humbertiana Macbride (426), 270 humido-umbrosa Ahrendt (204), 159

var. dispersa Ahrendt (204c), 160 var. humido-umbrosa (204a), 160 var. inornata Ahrendt (204b), 160

x hybrido-gagnepainii Suringar (17a), 45 h p r y t h r a Diels (483), 291 hypokerina Airy-Shaw (a), 38 hypoleuca Lindl., 87 iberica Stev. et Fisch. ex DC., 173

var. paphhgoniea Schneid., 174 ignorata Schneid. (LSL), 145 ilicifolia Forst. (375), 253 iZic$olia Scheele, 352 iliCina (Schldl.) Hemsl., 349 iliensis Popov (235), 178

26-2

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404 ITWkX

Genus Berberis L. 26-295, 357-9 (cozrt.)

Species and varieties (COW) impdita Schneid. (28), 50 incrassata Ahrendt (3). 38 var. bucahwangensk Ahrendt (36), 38 var. inc-ta (3a), 38

inemG Persoon. 241 insignis Hook. f. et Thorns. (2), 36

var. elegentifolia Ahrendt (2b), 37 var. gouldii Ahrendt (2f). 38 var. insignis (2a), 37 var. shergmnensis Ahrendt (2e), 37 vex. tongloensis Schneid. ( 2 4 , 37,364 (Fig. 22) var. zelaica Ahrendt (2c), 37

insolita Schneid. (36), 56, 364 (Fig. 21) integerrima Bunge (234), 178 inikgewimcr Franch., 179 var. bzrhseana Schneid., 177 var. denSi&wa Schneid., 176 var. eriwaned-s Schneid., 177 var. pyrocarpa Regel, 179 var. stenophylla Msxim., 176 var. turC0mcmnnk.a Schneid., 177

x interposita Ahrendt (5e). 40 x irwinii Byhouwer (368). 249 jaeschkeana Schneid. (173), 138

var. bimbilaica Ahrendt (173~) . 139 var. jaeschkeana (173a), 138 var. usteriena Schneid. (1736), 139

jrtmesiana Forrest et W. W. Sm. (238). 179, 366 (Fig. 42)

var. jamesiana (238a), 179 var. leucocarpa (W. W. Sm.) Ahrendt (238~) .

var. saepium Ahrendt (2386). 180 180

jantesoni Lemaire, 294 jamesonii Lindl. (476). 289 jantesonii Turcz., 275 japoniCa (Regel) Schneid., 194 javanica Miq. MS., 64 jelskiana Schneid. (453), 281 johannis Ahrendt (177). 141, 365 (Fig. 34) johnstonii Stendl. et Steyerm., 347 julianae Schneid. (64). 68 var. julianae (64a), 68 var. oblongifolia Ahrendt (646), 69 var. patungensis Ahrendt (sac), 69

vm. kansuensis (239a), 183 var. procera Ahrendt (2396), 183

kartanica Ahrendt (170), 137 kaachgarica Rupr. (319), 228 kaehmirana Ahrendt (129), 107 kawakamii Hayata (59), 65 kawakamii Seay, 65

kansuensis Schneid. (239), 182

var. formosans (Ahrendt) Ahrendt (59b), 65 var. kawakamii (59a), 65

keissleriana Schneid. (469), 285 kerriana Ahrendt (101). 91 x kewensis Schneid. (95e), 89 khaaiana Ahrendt (96), 89 knightii (Lindl.) K. Koch (372), 251 x'Knockvde Scarlet' Mulligan ex Ahrendt

(284a), 209

Species and varieties (eont.) koehneana Schneid. (285), 210, 367 (Fig. 48) var. aurama Ahrendt (2856), 210 var. koehn- (285a), 210

kongboensia Ahrendt (131), 107, 363 (Fig. 18) koreana Pelib. (260), 187, 366 (Fig. 47) kumeoneneis & h i d . (142), 120, 365 (Fig. 31) kunswurensk Royle (287), 211 lambertii Parker (167), 135 lanceold0 Ebnth., 348 Iasioclema Ahrendt (164), 134 latifolia R. et P. (408), 264 laurina Billbg. (387), 257 laxi&wa Citarne, 274 x laxiflora Sobrad. (252), 189 var. langeana Schneid. (252c), 190 var. laxiflora (252a), 190 var. oblanoeolata Schneid. (2526). 190

x leachitma Ahrendt (281 a), 207 lechleritma Schneid. (460), 283 lecomtei Schneid. (196), 155 lehmannii Hieron. (493), 293 kichtlinii Hort., 167 xlemoinei Ahrendt (161a), 132 lempergiana Ahrendt (84), 78 lepidifolia Ahrendt (224), 171, 366 (Fig. 41) kptmlada Diels, 154 leptopods Ahrendt (55). 63 leschnaultii Wall. ex Wight et Am., 325 kucocorpa W. W. Sm., 180 levis Franch. (78), 75

var. brachyphylla Ahrendt (786), 75 var. levis (78a). 75

libanotics Ehrenberg ex Schneid. (306). 223 liechtensteinii Schneid. (103), 92 lilloana Job (342), 237 lindleyana Ahrendt (132), 108 linearifolia Phil. (386), 255, 368 (Fig. 63)

var. linedolia (386a), 256 var. longifolia (Reiche) Ahrendt (3866), 256

liophylla Schneid. (75), 74 var. conglobeta Ahrendt (756), 74 var. liophylla (75a), 74

litorelis Phil. (392), 259 lobbiana Schneid. (429), 271 x lologensis Sandwith (370), 251 loudonii Ahrendt (299), 220 loxensis Benth. (491), 293 lubrica Schneid. (35), 55

ludlowii Ahrendt (136), 114, 365 (Fig. 27) lucida schrad., 189

var. capillmi8 (CoxexAhrendt) Ahrendt (1366),

var. deleica (Ahrendt) Ahrendt (136c), 115 var. ludlowii (136a), 115 var. sakdened-s Ahrendt, 116 var. saxiclivicola Ahrendt ( 136 d), 1 15

lutea R. et P. (416), 266 lycioides Linden et P1. ex Tr. et Pl., 281 lycioides Lowe, 223 lycioides Stepf (98), 89, 364 (Fig. 25) lycium Royle (95), 87

115

var. lycium (95a), 88

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Genus Berberis L. 26295, 357-9 (ma. )

405

Species and varieties (&.) lycium Royle, var. simlensis Ahrendt (95c), 88

var. subfascicularis Ahrendt (95b), 88 var. subvirescens Ahrendt (954, 88

x macracantha Schrad. (118a), 101 var. pulchra Schneid. (118b), 102

macrosepda Hook. f. et Thoms. (137), 115 var. dekk Ahrendt, 115 var. macrosepala (137a), 116 var. sakdenensis (Ahrendt) Ahrendt (137 b), 116 var. setifolia Ahrendt (137c), 116

msderensis Lowe (305), 223 magnifolia Ahrendt (271), 198 mairei Ahrendt (80), 76 manipurana Ahrendt (21), 47 marginata C. Gay (322), 231 masafuerana Skottsb. (345), 238 mximowiczii Regel, 167 x meehanii Schneid. ex Rehd. (262e), 195

mekongensis W. W. Sm. (203), 159 metapolyantha Ahrendt (282), 207 michay Job (354), 243 micrantha Ahrendt (1 12), 96 micropetala Schneid. (61), 67 microphylla Forst. (341), 237 microtricha Schneid. (202), 159 mikuna Job (361), 245 mingetsensis Hayata (27), 50 minutiflora Schneid. (190), 151, 365 (Fig. 36)

var. glabramea Ahrendt (190b), 152 var. minutitlora (190a), 151

miqueliana Ahrendt (219), 168 mitifolia Stapf (269), 198 monosperma R. et P. (395), 260 montana C. Gay (362), 245, 368 (Fig. 60)

VW. ChiUunenSis Schneid., 246 var. coletiaides (Lechl.) Schneid., 246 var. qrucilia Schneid., 247

montevidensis Schneid. (351), 242 naoranemb Hebenstr. et Ludw. ex Schult. f., 344 morenonis Kuntze (334), 234 moritzii Hieron. (442). 276 morrisonensis Hayata (147), 124 mouilhana Schneid. (215), 166 mucrifolia Ahrendt (168), 135 muliensis Ahrendt (156), 129

var. atuntzeana Ahrendt (156b), 130 var. beimanica Ahrendt (156c), 130 var. muliensis (156a), 129

multiflora Benth. (488), 292 var. cdvescens Schneid. (488b), 292 var. mdtiflora (4880), 292

mutabilis Phil. (359), 244 nana Greene, 337 nantoensis Schneid. (16), 44 negeriana Tischler (378), 253 nemorosa Schneid. (85), 78 nepalemb var. hchenaultii Hook. f. et Thoms.,

nerwosu Pursh, 329 neubertii (Hort.) Baum., 357 nevinii Gray, 354 +icans 0. Kuntze (409), 264

325

Species and varieties (COW.) nilghiriensis Ahrendt (108), 94 xnotabilis Schneid. (317), 228 nummularia Bunge (236), 178

var. nummularia (236a), 178 var. pyrocarpa Schneid. (236b), 179 var. schrenkiana Schneid., 178 var. s i n k Schneid., 179 var. szovi tzbnu Schneid., 176

nutans Linden et P1. (482), 290 oblanceolata (Schneid.) Ahrendt (278), 205 oblonga (Regel) Schneid. (307), 224 orientalis Schneid. (248), 186 oritrepha Schneid. (213), 165 orthobotrys Bienert ex Aitch. (179), 142

var. canescens Ahrendt (179f), 144 var. conwayi Ahrendt (179b), 143 var. orthobotrys (179a), 143 var. rubicunda Ahrendt (179e), 144 var. rupestris Ahrendt (179d), 143 var. sinthanensis Ahrendt (179c), 143

osmastonii Dunn (8), 41 x ottawensis Schneid. (218), 168

ovalifolia Rusby (437), 274 x oxoniensis Ahrendt (63d), 68 pachyacantha Koehne (254), 191 pallens Franch. (225), 172 pdlida Hartw., 349 pniculata Hemsl., 348 paniculata Juss. ex DC. (487), 291 padanensis Ahrendt (37), 56, 364 (Fig. 19) papillifera (Franch.) Koehne (201), 158 papillosa Benoist (444), 276

var. purpurea Schneid. ex Rehd. (218a), 168

var. aequatorialis Ahrendt (444b), 277 var. opacifolia Ahrendt (444c), 277 var. papill- (444a), 277

paraspect8 Ahrendt (20), 47 paravirescens Ahrendt (188), 148 parisepala Ahrendt (135), 114 parkeriana Schneid. (97), 89 parodii Job (350), 242 xparsonsii Schneid. (113c), 98 parviflora Lindl. (415), 266 pawifolda Sprague, 215 paucidentata Rusby (423), 270 pavoniana Ahrendt (420), 268 pearcei Phil. (379), 254 pectinata Hieron. (478), 289 peruviana Schellenb. (404), 263 petiol& Wall. ex G. Don (105), 93

var. garhwdana Ahrendt (105b), 94 var. petiolaris (105a), 94

pe t i t im Schneid. (122), 104 petrogena Schneid. (40), 57 phanera Schneid. (33), 54 philippii Ahrendt (326), 232 phyhan tha Rusby (421), 268 pichinchensis Turcz. (461), 283 pilosifolia Ahrendt (335), 235 pindilicensis Hieron. (475), 288, 368 (Fig. 67) pinifolia (Lundell) Muller, 355 var. eouhuilensis Muller, 355

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406 I&

Oenue Berberis L. 26-295, 367-9 (oont.)

Species and varieties (mat.) pin- Kunth, 344 pin- Lag., 338 pinnuta Roxburgh, 317 x ‘F’irate King’ Waterer ex Ahrendt (283f), 209 platyphyb (Ahrendt) Ahrendt (182). 145 podophylla Schneid. (467). 285 pohtii Schneid. (228), 174 poluninii Ahrendt (73). 73 polyantha H e d . (273), 201

polymorpha Phil. (327), 232 polypetala Phil. (356), 243 potaninii Biaxim. (102). 92 praecipua Sohneid. (lo), 42

var. major Ahrendt (lob), 43 var. praecipua (lOa), 42

prainkna Schneid. ex Stapf, 58 prattii Schneid. (274). 202 var. laxipendule Ahrendt (274~) . 203 var. prattii (274a), 202 var. recurvata Schneid. (274b), 202

prolifice Pittier (452), 281 xprovincielis Audibert ex Schrad. (16lc), 132

pruinosa Franch. (89), 80

var. obka- Schneid., 205

var. 8m& (Koehne) Scheid., 157

var. barresiana Ahrendt (89e), 81 var. brevifolia Ahrendt (89g), 82 var. brevipea Ahrendt (tigo), 81 var. longifolia Ahrendt (89b), 81 var. pruincaa (89a), 80 var. punctah Ahrendt (89j), 82 var. serratifolia Ahrendt (89h), 82 var. tenuipea Ahrendt (89f), 81 var. viridifolie Schneid. (89d). 81

pseudo-ilicifolia Skottsb. (376), 253 pseudo-spinulosa Job (405). 263 pseudumbellata Parker (313). 226. 367 (Fig. 54) psiloclada (Schneid.) Ahrendt (430), 272 psilopoda Turcz. (467). 283 pubescens Pamp. (270). 198 p z c m i l c c Greene. 335 purdomii Schneid. (230), 175 pyonophyUa Bienert ex Schneid., 176 quelpaertensis Nakai (214), 165 quindiuensis Kmth (438). 274 guindiw& Tr. et Pl., 288 rariftora Lechl. (424), 270 rechingeri Schneid. (470), 286 rectinervia Rusby (417). 267 recurvata Ahrendt (70). 71 regeliana K w h e ex Schneid. (261), 194 x regleriana Notcutt (176), 140 x rehderiana Schneid. (222a). 170 reicheana Schneid. (464). 284 replicata W. W. Sm. (13), 43 var. d k p r Ahrendt, 43

reticdata Byhouwer (211), 164 retinervia Tr. et P1. (456), 282 TIWQ~W~Z Sm. ex DC., 244 rigid8 Hieron. (433), 272 rigidifolia Kunth (450). 280 rockii Ahrendt (216). 204

Speciee and varieties (cont.) rmme&ana Saheele, 352 rotundifolia Poepp. et Endl. (346), 238 royleana Ahrendt (314). 227 (? x ) rubroetiUe Chittendem (184). 146

var. chealii C h d ( IMb) , 147 var. crawleyensis Ahrendt (184a), 146

x ‘Ruby Wetsan’ Ahrendt (275d), 204 rufescena Ahrendt (199), 157 mbyana Ahrendt (418), 267 ruscifolia Lam. (374). 252 salicaria Fedde (266), 197 sanguinea Franch. (39), 57

sargentiana Schneid. (69), 71 saxicola Lechl. (425), 270 aaxonnn Ahrendt (463), 284 mhidmna Schldl., 342 8-2hn&& Rehd., 164 schneideriam Ahrendt (81). 76 schweriuii Schneid. (480), 290 sellowkma Schneid. (398), 261

var. miomphyllo Hort. apud Creasy, 56

var. tetonobotrys Schneid., 258 8 W & k hll@, 173 xaerrataKoehne (198a), 157 aerrato-dentata k h l . (355), 243 serroto-dentda Reiche, 253 setigrifolia Ahrendt (343), 237 shemkua ahrendt (189), 148 shemilfii Ahmndt (298), 218 x ‘Sibbertoft Coral’ Stanley ex Ahrendt (277a).

sibirica Pall. (160), 131 sieboldi Miq. (210). 164, 366 (Fig. 44) 8- ’ (Schneid.) Ahrendt (114), 99

205

var. baileyi Ahrendt (114b). 99 var. glabramee Ahrendt (114c), 90 var. sikkimensie (114a), 99

~ k O h Y8hbe,169 silva-teroucena Schneid. (220), 169 silvicola Schneid. (52), 62

var. angustata Ahrendt (52b), 62 var. silvicola (52a), 62

aimonsii Ahrendt (486), 291 simulam Schneid. (51), 62 8inensie DC., 173

var. c b ~ ~ & Regel, 174 var. elegotle Franch., 154 var. 2WPUccgonica Schneid., 174 var. @/pica Franch., 155

xsmithian8 Sprague ex Ahrendt (296c), 217 solutiflora Ahrendt (280), 205 soulieana Schneid. (83). 77

var. paucinervata Ahrendt (83b), 78 var. soulieana (83a), 78

xspaethii Schneid. (113e). 98 x ‘Sparkler’ Chittenden ex Ahrendt (283a). 208 sp. Rehd. (1927). 79 & ~ ~ O U Z T ~ Her. et Er., 227 q W r a Fedde, 92 apinmissima (Reiche) Ahrendt (349), 241 Spinurosa Griseb., 263 aphdosa St. Hil. (406), 263

Page 407: Berberis and Mahonia : A taxonomic revision

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Genus Berberis L. 26-296, 357-9 (cont.)

407

Species and varieties (cont.) spraguei Ahrendt (208), 162

var. pedunculata Ahrendt (208b), 162 var. spraguei (208a), 162

sprucema (Schneid.) Ahrendt (422). 269 8krpJiana. Schneid., 216 stearnii Ahrendt (195), 155 atenophyk Hance, 77 x stenophylla Lindl. (367), 247

stenostachya Ahrendt (268), 197 stiebritziana Schneid. (155), 129 stolonifera Koehne et Wolf (312), 226 x ‘Stonefield Dawn’ Ahrendt (275c), 204 x ‘Stonefield Glow’ Ahrendt (274d), 203 x ‘Stonefield Mauve’ Ahrendt (281a), 207

stuebelii Hieron. (445), 277 subacuminata Schneid. (54), 62 subantsrctica Gandoger (377), 253 szcbcaul4alrua Schneid., 215

var. guhtzunku Ahrendt, 216 subcoriaxrea Ahrendt (79), 75 suberecta Ahrendt (183), 146 sublevia W. W. Sm. (41), 58

var. inVindi Hort., 249

var. exqukita Ahrendt (ale), 58 var. grandifolia Schneid. (ald), 58 var. microcarpa (Hook. f. et Thoms.) Ahrendt

var. sublevis (41 a), 58 subpteroclda Ahrendt (74), 73

var. impar Ahrendt (74b), 73 var. minoripes Ahrendt (74c), 73 var. subpteroclda (74a), 73

(alb), 58

subsessiliflora Pamp. (264), 196 subtripher& Franch., 327 8ulcata Hort., 188 awaseyi Buckley, 353 taliensis Schneid. (14), 44 taronensis Ahrendt (87), 79

var. taronensis (87a), 79 var. trimensis Ahrendt (87 b), 79

taylorii Ahrendt (296), 217, 367 (Fig. 52) temolaica Ahrendt (143), 120

var. artisepala Ahrendt (143b), 121 var. temolaica (143a), 121

tenuifolicc Lindl., 346 thibetica Schneid. (194), 154 thomsoniana Schneid. (128), 106 thunbergii DC. (217), 166

var. argenteo-marginata Schneid. (217f ), 167 var. atropurpurea Chenault (217g), 168 var. erecta (Rehd.) Ahrendt (217j), 168 erecta Rehd., 168 var. globra Franch., 155 var. maximowiczii (Regel) Regel (217e). 167 var. minor Rehd. (217d), 167 var. papillifera Franch., 158 var. pluriflora Koehne (217b), 167 var. plurijEora erecta Anon., 168 var. rubrifolia Ahrendt (217h), 168 var. thunbergii (217a), 167 vm. uniflora Koehne (217c), 167

tinetoria Lesch. (1 I O ) , 95

Species and varieties ( c o d . ) tischleri Schneid. (148), 124

var. abbreviata Ahrendt (148b), 125 var. tkchleri (148a), 125

tolimensis Linden et P1. ex Tr. et Pl., 283 toluacensi.9 Hort., 331 tomentoss. R. et P. (410), 265 tomentulosa Ahrendt (200), 158 x ‘Tom Thumb’ Morris ex Ahrendt (293d), 215 tragaeanthoidas DC., 331 triacanthophora Fedde (30), 51 trichohaematoides Ahrendt (176), 141 t r k q i d d u Smith ex DC., 233 trifoliata Hartw. ex Lindl., 352 trifoliuta Lemaire, 352 trzfoliata Torr., 353 trifolwlata Moric., 352 trtfurea Loud., 321 trigona Kunze ex Poepp. et Endl. (385), 255

trollii Diels (403), 262 truxillensis Turcz. (451), 281, 368 (Fig. 66) tsangpoensis Ahrendt (lal), 119. 365 (Fig. 29) tsarica Ahrendt (162), 133

var. ritangensis Ahrendt (162 b), 133 var. tsarica (162a), 133

taarongensis Stapf (197), 156 var. bre&cupa Ahrendt, 163 var. megacarps Ahrendt (197b), 156 var. tsarongensis (197a), 156

tschonoskyans. Regel (221), 169 turcomannica Kar. ex Ledeb. (233), 177 t u r m n n i c a Rehd., 177

var. m f o l i a Reiche, 256

var. buhseana (Schneid.) Ahrendt (233b), 177 var. den9il;fEora Rehd., 176 var. turcomannica (233a), 177

ulicina Hook. f. et Thoms. (318), 228,367 (Fig. 55) umbellata Lindl., 108 umbellata Parker, 226 umbeUata Phil., 232 umbellata Wall. ex G. Don (124), 104, 363

var. brianii Ahrendt (124b), 105 var. umbellata (124a), 105

wterionu (Schneid.) Parker, 139 var. apiculata Ahrendt, 135

valdiviana Phil. (397), 260 var. gracilifolia Ahrendt (397b), 261 var. Phil. ex Reiche, 261 var. valdiviana (397a), 261

valida Schneid. (45), 60 validimpala Ahrendt (157), 130

var. primoglauca Ahrendt (157b), 130 var. va1idiseps.h (157a). 130

x vanfleatii Schneid. (174a), 140 v d o r a Schneid. (336), 235 veitchii Schneid. (31), 51 veitchiorurn Hemsl. et Wils., 301 venusta Schneid. (29), 51 vernae Schneid. (229), 175, 366 (Fig. 43) verruculosa Hemsl. 0t Wils. (IS), 46, 364

(Fig. 17)

(Fig. 23) f. viridescena Schneid., 46

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408 Ind&X

Genus Berberls L. 26-295, 357-9 (cod.)

Species and varietKes (cont.) versohaffelti Schneid. (494), 294 verticillata Turcz. (474). 288 x vhorh i i Schneid. (174), 140 u6rescena Hook. f., 148 virescenS Hook. f. et Thoms. (149), 125

var. igraoratu (Schneid.) Ahrendt, 145 var. macrocarpa Bean, 148

virgata R. et P. (414), 266, 368 (Fig. 64) var. huanwxmb Schneid., 266

virgetorum Schneid. (216), 166, 366 (Fig. 46) vitellina Hieron. (490), 293 vulgaria L. (251), 188

var. acutifolia (Prantl) Scheid. (2516), 188 var. aetnensis (Presl) Fiori et Paoletti, 222 var. auatTdk Boiss., 221 var. emorginata (Willd.) Gordon, 132 var. japonie0 Regel, 194 var. p e d b Audibert, 132 var. purpurm Hort., 189 var. purpurifolia Ahrendt (251d), 189 var. S d C r t t 8 Ahrendt (251~), 188 var. vulgaris (251a), 188 V W . . 189

wallichiana DC. (68), 71 walliehiana Hook. f., 39

var. Collett et Hemsl., 42 var. qrdipea Ahrendt, 58 var. Wfdk Hook. f. et Thorns., 40 var. microcarpa Hook. f. et Thoms., 58 var. pa.ll&ia &is, 46 var. pall& Hook. f. et Thorns., 72 pamifolk Franch., 56 f. ar- Franch., 60

walhrana Ahrendt (126), 106 wangii Schneid. (76), 74 wardii Schneid. (25), 49

Species and varieties (cont.) warszewiczii Hieron. (477), 289 x watlingtonensis Ahrendt (21a), 48 wawram Schneid. (358), 244 wazaristtmica Ahrendt (310), 225 weberbaueri Schneid. (396), 260 weddellii Lechl. (402), 262 wettsbiniana Schneid. (471), 286 wightiane Schneid. (log), 95 willeana Schneid. (65), 69

var. serrulata Schneid. (656), 69 var. willeane (65a), 69

wilsontle Hemsl. (293), 214 var. f a v w (W. W. Sm.) Ahrendt (293c), 215 vm. guhtzunica (Ahrendt) Ahrendt (294a),

var. Iatior Ahrendt (295a), 216 var. parvifolia (Sprague) Ahrendt (293b), 215 var. stapfictna (Schneid.) Schneid. (295), 216 var. subcaulialata (Schneid.) Schneid. (294).

var. wilsonae (293a). 214

216

215

x Wintonensis Ahrendt, n.n. (63c). 68 wisleyensia Ahrendt (34), 54 x wokitzgensis Ahrendt, n.n.. 45 xanthoclada Schneid. (56). 63 xenthophlaes Ahrendt (279). 205 xanthoxylon Hesskarl ex Schneid. (57), 64

vm. junghuhniana Ahrendt (578). 64 vm. aumatranica Ahrendt (57c), 64 var. xmthoxylon (57a), 64

var. pldyphyllia Ahrendt, 146 yunnanenSie E’ranch. (206), 161, 365 (Fig. 39)

zabeliana Schneid. (244), 184 m W M m Schneid., 232 &cienensis Pamp. (62), 67 zayulens Ahrendt (166). 135

XMahoberberis Schneid., 357

aquicandidula Kriissmann (610). 357 aquisargentii Rriissmann (611). 357

miethkeana Melander et Eade (612), 357 neubertii (Baum.) Schneid. (609), 357

Genus Mahonia Nutt. 296-356

end 5 (p. 356) comprise alphabetical lists of species

Serial numbers of species are given in parentheses, for example (597)

Illustrations are confined to pp. 360-3. Appendices 3 (p. 330), 4 (p. 331)

bracts and pedicels, 363 (Fig. 13) diagzlostic characters of, 2 lilamente. apx of, 362 (Fig. 10) fruit and style, 362 (Fig. 12) geogrephicel distribution of, 21, 369-98 (maps) infloremence, forms of, 360 (Fig. 6) keys, use and nsture of, 24

leaves: apex of, 360 (Fig. 6); cross-section of, 363 (Fig. 14); margins of, 360 (Fig. 2); petioles of, 360 (Fig. 3); shapes, 360 (Fig. 4)

ovaq and ovules, 362 (Fig. 11) petels: apex of, 361 (Fig. 7); Inwe of, 362 (Fig. 8) stamens, apex of, 362 (Fig. 9) stem, sections of, 360 (Fig. 1)

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ITUZeX

Genus Mahonia Nutt. 296356 (cont.)

409

Groups, sections and subsections Acanthifoliae Ahrendt (subsect.), 306 Aquifoliatae Fedde (sect.), 331 Dentatae Ahrendt (subsect.), 324 Dolichopodae Ahrendt (subsect.), 326 Dolichostyles Ahrendt (subsect.), 304 Euaquifoliatae Ahrendt (subsect.), 332 Eulongibracteatae Ahrendt (subsect.), 300 Eupaniculatae Ahrendt (subsect.), 346 Horridae Fedde (sect.), 350 Integrifoliolatae Ahrendt (subsect.), 327 Japonicae Ahrendt (subsect.), 314 Laxiracemosae Ahrendt (subsect.), 345 Longibracteatae Fedde (sect.), 297 Napaulenses Ahrendt (subsect.), 310 Nervosae Ahrendt (subsect.), 327 Occidentales Ahrendt (group), 331 Orientales Ahrendt (group), 297 Paniculataa Fedde (sect.), 344 Schiedeanae Ahrendt (subsect.), 341 Siamenses Ahrendt (subsect.), 321

Species and varieties acanthifolia G. Don (518), 309 x aldenhamensis (Hort.) Ahrendt (575b), 337 akzalzdri Schneid., 311 amplectens Eastwood, (565), 333 andrieuxii (Hook. et Am.) Fedde (586), 345 angustifolia (Hartw.) Fedde (580), 342 m a m i c a Gagnep. (560), 329 aquifolium (Pursh) Nutt. (571), 335

var. aldenhamensis Hort., 337 var. aquifolium (571a), 336 var. convoluta Hort., 338 var. lyallii Ahrendt (571b), 336 vm. rotundifolia herveyi Hort., 337 var. undulata Hort., 338

arguta Hutch. (600), 350 aristulata Ahrendt (520), 310 bealei (Fort.) Carr. (540), 319

var. bealei (540a), 320 var. planifolia Hook. f. et Thorns. (540b),

320 bijuga Handel-Mazzetti (496), 300 bodinieri Gagnep. (531), 316 borealis Takeda (514), 307

var. borealis (514a), 308 var. parryi Ahrendt (514b), 308

bracteolata Takeda (510), 305 caesia Schneid. (564), 331 calamicaulis Spare et Fisch. (504), 302 californica (Jepson) Ahrendt (570), 335 chiapensis Lundell (59l), 347 chochoco (Schldl.) Fedde (587) , 345 conferta Takeda (511), 305 confusa Sprague (530), 316

var. bournei Ahrendt (530b), 316 var. confusa (530a), 316

x convoluta (Hort.) Ahrendt (575e), 338 decipiens Schneid. (559), 329 dictyota (Jepson) Fedde (567), 334 disoolorifolia Ahrendt (549), 323 dolichostylis Takeda (512), 305

Species and varieties (cont.)

duclouxiana Gagnep. (513), 306 var. duclouxiana (513a), 306 var. hilaica Ahrendt (513b), 306

ehrenbergii (Kunze) Fedde (590), 347 ekgans (LBveillB) Rehd., 324 eurybracteata Fedde (501), 301 eutriphylla Fedde (581), 342 fargesii Takeda (539), 319 faa*ri.s DC., 338 feddei Ahrendt (507), 303 flavida Schneid. (526), 313

var. flavida (526a), 313 var. integrifoliola Handel-Mazzetti (526b), 313

fordii Schneid. (535), 318 fortunei (Lindl.) Fedde (557), 328

var. fortunei (557a), 328 var. szechuanica Ahrendt (557b), 328

fremontii (Torr.) Fedde (603), 353 ganpinensis (L6veille) Fedde (530c), 316 glunzacea DC., 329 gracilipes (Oliver) Fedde (554), 326 gracilis (Hartw.) Fedde (584), 343 griffithii Takeda (516), 308 haematocarpa (Wooton) Fedde (606), 354 hmcockiana Takeda (527), 314 hartwegii (Benth.) Fedde (597), 349 x herveyi (Hort.) Ahrendt (5754, 337 hicksii Ahrendt (509), 304 higginsae (Munz) Ahrendt (608), 356 huiliensis Handel-Mazzetti (563), 330 hypoleuca Takeda (562), 330 ilicina Schldl. (595), 349 incerta Fedde (577), 341 japonica (Thunb.) DC. (541), 320 japonica Hort., 319

var. gracillinza Fedde, 320 var. japonica (541a), 320 var. trifurca (Loud.) Ahrendt (541b), 321

jaunsarensis Ahrendt (519), 310 johnstonii (Standl. et Steyerm.) Standl. et

Steyerm. (592), 347 klossii Baker f. (543), 321 knightii Hort. ex Lindl., 251 lanceolata (Benth.) Fedde (594), 348 leptodonta Ga,gnep. (542), 321 leschenaultii (Wall.) Takeda (551), 325 leveilleana Schneid. (550), 324 lomariifolia Takeda (521), 31 1 longibracteata Takeda (505), 303 longipes (Standl.) Standl. (588), 345 magnifwa Ahrendt (517), 309 mairei Takeda (538), 319 manipurensis Takeda (523), 312 monyulensis Ahrendt (506), 303 moranensis (Hebenstr. et Ludw.) I. M. Johnston

(585), 344 var. cachira (Fedde) Ahrendt (585b), 344 var. moranensis (585a), 344

morrisonensis Takeda (546), 322 xmoseri (Hort.) Ahrendt (575c), 337 muelleri I. M. Johnston (601), 351 nana (Greene) Fedde (574), 337

Page 410: Berberis and Mahonia : A taxonomic revision

410 I*

Genus Mahonia Nutt. 296-366 (d.)

Species and varieties (cont.)

napadensis DC. (524), 312 vm. rmbzcrghii DC., 317

ne+& vm. pymwphyu0 Fedde, 313 nervosa (Pursh) Nutt. (561), 329 nevinii (Gray) Fedde (605), 354 new sp, (515). 308 nitem Schneid. (5357, 318 niveu Schneid., 330 oiwakensk Hayate (548), 323 pachaksbnsis Ahrendt (532), 317 pallida (Hsrtw.) Fedde (596), 349 peniculata Oerst. (593), 348 paxii Fedde (598), 350 philippinensis Takeda (553), 326 pinifolia Lundell (607), 356

vm. coahuilensis (Muller) Ahrendt (607b), 355 vsr. pinifolicz (607a). 365

pinnate (Lag.) Fedde (576), 338 vm. cuehim Fedde. 344

piperha Abrema (573). 336 polyodonta Fedde (508), 304 pomensiS Ahrendt (534), 317 pumila (Gmne) Fedde (569), 335 pycnophylla (Fedde) Taked8 (525), 313 quinquefolia (Stand.) Standl. (582), 343 repens G. Don (568), 334

var. repens (568a). 334 vm. rotundifolia Fedde (5686). 334

Species and varieties (m.) schiede- (Schldl.) Fedde (579), 342 schochii Schneid. ex Hendel-Maezetti (536), 318 setosa Gagnep. (558), 328 s h e d Chun (556), 327 sherideniana Schneid. (529), 315 siamensis Takeda (544), 322 slklumsnsis Tekeda (552), 325 shonsii Takeds (522), 312 sonnei Abrams (572). 336 subimbricata W. Y. Chun et F. Chun (500), 301 subintegrifolia Fedde (583), 343 subtriplinervis (Fmnch.) Fedde (555), 327 s-trensis Merrill(545). 322 swaseyi (Buckley) Fedde (604). 353 taronensis Hendel-M[azzetti (502). 301 tenuifolia (Lindl.) Loud. ex Steud. (689), 346 tikushiensis Hayata (547). 323 toluecensis (Been) Ahrendt (566). 333 t~foZlda Cham. et Schlecht. MSS., 342 trifoliohta (Moric.) Fed& (602). 352

. .

vsr. glauca I. M. Johnston (6026), 352 vm. trifoliolata (602a), 352

x undulate (Hort.) Ahrendt (575f). 338 veitchiorum (Hemsl. et W&.) Schneid. (603), 301

vm. kingdon-wardiana Ahrendt (503b). 302 vm. veitchiorum (503a), 302

volcania Stend. et Steyerm. (578), 341 x wagneri (Hort.) Ahrendt (575a), 337 zemanii Schneid., 316 zimapana Fedde (599), 350

roxburghii (DC.) Takeda (533), 317 salwwnensis Ahrendt (528), 314

Cmagene, 331 I l a japonica Thunb., 320

Odostemon hgipea Standl., 345 odostemon q?4irUpfolia stiuldl., 343