A taxonomic revision of Zygodon (Orthotrichaceae)
in southern South America
GRACIELA M. CALABRESE
Real Jardı́n Botánico, Plaza de Murillo 2, E-28014 Madrid, Spain.
e-mail: calabrese@rjb.csic.es
ABSTRACT. The genus Zygodon Hook. & Taylor is revised for southern South America
(Andean-Patagonian and Fuegian regions). Thirteen taxa (twelve species and one variety)
are recognized as occurring in the region: Z. bartramioides Malta, Z. chilensis Calabrese &
F. Lara, Z. hookeri Hampe var. hookeri, Z. hookeri var. leptobolax (Müll. Hal.) Calabrese
comb. nov., Z. intermedius Bruch & Schimp., Z. jaffuelii Thér., Z. magellanicus Dusén ex
Malta, Z. menziesii (Schwägr.) Arnott, Z. papillatus Mont., Z. pentastichus (Mont.) Müll.
Hal., Z. pichinchensis (Taylor) Mitt., Z. reinwardtii (Hornsch.) Bruch & Schimp. and
Z. obtusifolius Hook. All taxa previously considered to grow in southern South America were
studied, and many were typified. Because of the biogeographical connections with other
Southern Hemisphere landmasses, some taxa from other areas (e.g., Australasia or South
Africa) were also considered. The studied taxa are a good representation of the genus as a
whole, thus allowing one to conclude that Zygodon, worldwide, consists of three subgenera:
subg. Zygodon, the species of which have papillose leaf cells that are flat as seen in crosssection, and that can be further divided in two sections, sect. Zygodon defined by leaf basal
cells not in differentiated bands and seta distally dextrorse, and sect. Stenomitrium Mitt. with
leaf basal cells disposed in differentiated alternating bands and seta distally sinistrorse; subg.
Codonoblepharon (Schwägr.) Calabrese stat. nov. comprising species with smooth, flat leaf
cells in cross-section; and subg. Obtusifolium (Malta) Calabrese stat. nov., including species
with papillose, bulging leaf cells. For each of the accepted taxa, keys, descriptions,
illustrations, a complete list of synonyms previously used in the study area (sometimes
expanded with names from abroad), ecology, distribution maps, variability and differentiation from related taxa are provided. Study of the peristome of Zygodon with the SEM
provides new data about its structure and ornamentation.
KEYWORDS. Argentina, Chile, Orthotrichaceae, South America, taxonomy, Zygodon.
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Zygodon Hook. & Taylor belongs to the Orthotrichaceae, a family that includes mosses with small, often
papillose upper leaf cells, mitriform or cucullate
calyptra, terminal perichaetia and sporophytes with
arthrodontous and diplolepidous peristomes consisting of endostomial segments alternating with exo-
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stomial teeth (Goffinet 1998; Vitt 1982). The family
includes four subfamilies according to Vitt (1972):
Zygodontoideae (Schimp.) Broth., Orthotrichoideae
(Schimp.) Broth., Drummondioideae Vitt and Macromitrioideae Broth. The subfamilies are defined by
calyptrae shape, growth of main stem and position of
THE BRYOLOGIST 109(4), pp. 453–509
Copyright Ó2006 by the American Bryological and Lichenological Society, Inc.
0007-2745/06/$5.85/0
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perichaetia: Zygodontoideae and Orthotrichoideae
include acrocarpous mosses while Drummondoideae
and Macromitrioideae comprise cladocarpous mosses.
In addition, the calyptra is cucullate in the Zygodontoideae and Drummondioideae, but mitriform in
the Orthotrichoideae and Macromitrioideae. According to Vitt (1982), the Zygodontoideae includes
Zygodon and Stenomitrium (Mitt.) Broth. Later,
Goffinet and Vitt (1998) considered the Orthotrichaceae to consist of but two subfamilies, each with
two tribes: subfamily Orthotrichoideae (Zygodonteae
and Orthotricheae) and subfamily Macromitrioideae
(Macromitrieae and Schlotheimieae). These authors
presented the first molecular studies on the family to
interpret its phylogeny. According to them, the
Zygodonteae includes Codonoblepharon Schwägr.,
Bryomaltaea Goffinet, Leratia Broth., Zygodon, Leptodontiopsis Broth., Pleurorthotrichum Broth. and
Stenomitrium (Mitt.) Broth., after segregating the
traditional Zygodon in four genera: Zygodon, Stenomitrium, Codonoblepharon and Bryomaltaea.
In the present treatment, I consider Zygodon s.l.
to include acrocarpous mosses with cucullate,
smooth, usually naked calyptrae, exserted capsules
with longitudinal ribs and diplolepidous peristomes.
The main gametophytic characters are the 6
isodiametric, thick-walled, papillose or smooth upper
leaf cells, the costae with ventral guide cells and dorsal
stereids, and the characteristic propagula developed in
the leaf axils.
Zygodon is distributed mainly in tropical and
sub-tropical areas, with most species in America from
Mexico to Tierra del Fuego, especially through the
Andes. Brotherus (1902–1903) recognized 93 species
worldwide (57 in the Americas, 14 in Africa, 13 in
Australia, 10 in Asia and 8 in Europe) and later raised
the number to 112 (Brotherus 1925). Malta (1926), in
the only worldwide revision of the genus, recognized
77 species, whereas Index Muscorum (Wijk et al.
1969) reported 90 taxa, and more recently Vitt (1982)
estimated 52 species worldwide. Currently, 91 species
of Zygodon are recognized according to Crosby et al.
(1999).
After Malta’s monograph (1926), revisions of
Zygodon have been published for Europe and Macaronesia (Düll 1985) and Australasia (Lewinsky 1990). It
has also been treated in regional floras: Moss Flora of
Mexico (Vitt 1994), Prodromus Bryologiae Novo-Granatensis (Churchill & Linares 1995), Flora of Southern
Africa (Magill & Van Rooy 1998), Moss Flora of the
Upper Bureya River (Russian Far East) (Ignatov et al.
2000) and Moss Flora of Central America (Allen 2002).
However, southern South America, despite its richness
in Zygodon, both in terms of taxa number and
abundance (Greene 1986; He 1998; Kühnemann 1938;
Matteri 1985, 2003; Matteri & Schiavone 2002; Seki
1974), lacks an updated revision, here undertaken.
During the course of this revisionary study, one
species was described as new from southern Chile
(Calabrese et al. 2006) and ten were reduced to
synonymy. Twelve species and one variety are
recognized from the Andean-Patagonian and Fuegian
regions: Zygodon bartramioides, Z. chilensis, Z. hookeri
var. hookeri, Z. hookeri var. leptobolax, Z. intermedius,
Z. jaffuelii, Z. magellanicus, Z. menziesii, Z. papillatus,
Z. pentastichus, Z. pichinchensis, Z. reinwardtii and Z.
obtusifolius.
MATERIALS AND METHODS
This revision is based on some 1300 herbarium
specimens from BA, BCRU, BM, CHR, FH, G, H, HBG, HIRO,
JE, MA, MO, NY, PC, PRE, S, TR-VENTURI, UBC, UC, the private
herbaria of Celina M. Matteri (Buenos Aires) and the
Laboratory of Bryology of the Universidad Autónoma
de Madrid (Spain). I also conducted field work during
the austral summers (December–March) for the
period 1995–1998, and April 2001. Specimens collected by the author are in her personal herbarium
with some duplicates in BCRU and MA.
Specimens were studied either in water or
mounted in Hoyer’s (Anderson 1954) or glyceringelatin (Johansen 1940). Illustrations of leaves, cells,
propagula, capsules, peristomes and spores were made
with a camera lucida on an Olympus BX40 light
microscope. Digital microphotographs were taken
with MOTICAM 1300 and Nikon HFX-II cameras.
Peristomes were also studied under the SEM. Mature
intact capsules had their opercula removed and were
dry-mounted on aluminium stubs covered with
adhesive, covered by gold and studied with a Philips
XL30 scanning electron microscope with EDAX Dx4i
analyser at 20 kV and a working distance of 10–14 mm.
In this treatment, all measurements were made
from wet material. Laminal width was measured at
Calabrese: Zygodon in southern South America
the broadest part of the leaf, costal width in the lower
third, cell measurements include the walls, capsule
length excludes the operculum, seta length includes
the vaginula and plant length refers only to the
gametophyte.
The species have been distinguished on the basis
of morphological characters. Descriptions of species
include variability observed in all specimens examined, from the study area and from Australia, New
Zealand, Tasmania, Africa and Central and South
America. Terminology follows Magill (1990).
The regional distribution maps are solely based
on specimens studied by the author. In ‘‘Selected
specimens examined’’ only one specimen per geographical unit is cited, and the total number of
specimens is in parentheses at the head of the section.
A complete list of specimens studied is available upon
request.
TAXONOMIC CHARACTERS
For this study I analyzed and evaluated the
taxonomic characters commonly used in Zygodon. I
also searched for new, stable characters that could be
of use in the taxonomy of the genus. Most Zygodon
species are dioicous and sporophytes are uncommon.
Thus, traditionally, attention has been mainly given to
gametophytic traits. However, in this revision I have
used both gametophytic and sporophytic traits as far
as possible to define taxa.
Gametophyte.
Habit.—Zygodon plants are typically acrocarpic.
Gametophytes are erect, 6 branched, forming loose
(e.g., Z. magellanicus) or dense (e.g., Z. menziesii)
turfs. The exception is the Andean Z. pentastichus
which forms 6 loose mats with plagiotropic stems
and erect branches to 5 cm high. Mitten (1869) and
Brotherus (1902–1903) considered the plagiotropic
habit of great taxonomic value and segregated Z.
pentastichus in an independent section, Zygodon sect.
Stenomitrium Mitt., or at the generic level as
Stenomitrium (Mitt.) Broth.
Size of plants is very variable. Low turfs less than
2 cm high are frequent, and the smallest plants of
some species (e.g., Z. obtusifolius, Z. intermedius, Z.
jaffuelii) reach only 0.2–0.5 cm. On the other hand,
the Andean Z. pichinchensis is always over 4 cm, and
can reach 11 cm tall.
455
Regarding robustness, specimens of some species
are fairly robust (e.g., Z. pichinchensis), while others
are slender (e.g., Z. papillatus). This character varies
according to turf density and height, as it seems
influenced by environmental conditions. However, it
is useful to separate two closely related species, Z.
magellanicus and Z. papillatus (Calabrese et al. 2002).
The color of the upper parts of the plants varies
from yellowish-green (e.g., Zygodon intermedius) to 6
dark olive-green (e.g., Z. obtusifolius), rarely reddishbrown (e.g., Z. pichinchensis).
Rhizoids.—Rhizoids variably cover the stem as a
tomentum, sometimes almost completely, as in Z.
pichinchensis, Z. squarrosus (Taylor) Müll. Hal. and
Z. nivalis Hampe, and occasionally are also developed at older leaf bases (Z. bartramioides and Z.
pichinchensis). Color varies from yellowish-brown to
reddish-brown; the only exception in the genus is Z.
forsteri (With.) Mitt., with hyaline to almost white
rhizoids. The rhizoids are usually smooth to slightly
roughened, although sometimes they can be papillose, a character that separates two otherwise similar
species: Z. magellanicus, with papillose rhizoids, and
Z. papillatus, with smooth or rarely slightly roughened, never papillose, rhizoids (Calabrese et al.
2002).
Stems.—Stems are 6 rounded to pentagonal,
usually with large, uniform, thin-walled central cells
surrounded by 1–3 layers of smaller, thick-walled
cortical cells. This is the most common stem anatomy,
and the only one seen in all the Andean-Patagonian
and Fuegian taxa of Zygodon. However, Z. minutus
Müll. Hal. & Hampe and Z. gracillimus Broth. ex M.
Fleisch., two Australasian members of subg. Codonoblepharon (¼ sect. Bryoides Malta), have different stem
anatomy (Lewinsky 1990): Z. minutus lacks differentiated cortical cells, while in Z. gracillimus all cells
are thick-walled or a few in the center are thin-walled.
Leaf arrangement.—In most Zygodon taxa, the
leaves are spirally arranged, but in species with clearly
pentagonal stems they show a 6 distinct pentastichous disposition, like in the Andean Z. campylophyllus Müll. Hal., Z. longicellularis D. G. Griffin, Z.
pichinchensis, Z. quitensis Mitt., Z. venezuelensis D. G.
Griffin and especially Z. pentastichus.
Leaf orientation.—When dry, leaves are appressed
to erect, straight and keeled in most Zygodon taxa,
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although some species have patent, flexuose to twisted
or rarely homomallous leaves. When moist, leaves
vary from erect to squarrose-recurved, although
spreading leaves are most typical. Zygodon pentastichus differs from any other species in the genus by
its sheathing base and a patent distal part.
Leaf size.—Most taxa of Zygodon have leaves 1–2
mm long. Exceptions are Z. erosus, Z. pentastichus, Z.
runcinatus Müll. Hal. and Z. trichomitrius Hook. &
Wilson, with leaves 2–4 mm long, and Z. obtusifolius
and Z. papillatus with smaller leaves (usually , 1
mm). Leaf width is usually 0.2–0.6 mm.
Leaf shape.—Lanceolate leaves with acute apex
are typical in Zygodon, although three species in the
study area deviate: Z. bartramioides with lanceolaterhomboidal leaves with clearly defined shoulders; Z.
menziesii with 6 lingulate leaves; and Z. obtusifolius
with lingulate leaves and a rounded-obtuse apex.
The rounded-obtuse apex of Zygodon obtusifolius
was traditionally considered diagnostic for subg.
Obtusifolium (Malta) Calabrese (¼ sect. Obtusifolii
Malta). However, the recently described Z. conoideus
var. lingulatus S. R. Edwards (Edwards et al. 2000)
also has rounded-obtuse leaf apices, but belongs to
subg. Zygodon.
Leaf margins.—Margins are usually plane, more
rarely slightly recurved in medial and lower portions
of leaves (e.g., Zygodon menziesii), or lower parts (e.g.,
Z. obtusifolius). When leaves are undulate, this is
especially conspicuous at margins (e.g., Z. hookeri and
Z. reinwardtii; Figs. 20C, 21C).
Most species have margins entire and smooth
toward the leaf base, and entire and papillose in the
rest of the leaf. However, Z. obtusifolius has papillosecrenulate margins for almost their entire length, and
leaf margins in Z. pichinchensis appear spinulose to
spinose by the very long papillae. Species with smooth
leaf cells (subg. Codonoblepharon) have entire and
smooth margins throughout. Finally, some species (Z.
hookeri var. hookeri, Z. reinwardtii and Z. runcinatus)
have denticulate to dentate margins at the leaf apex.
In Z. hookeri var. hookeri and Z. reinwardtii some
totally entire leaves or specimens occasionally occur,
and also some weak denticulations can be found in
species with leaf margins usually entire, such as Z.
hookeri var. leptobolax, Z. jaffuelii and Z. bartramioides.
Although traditionally margin ornamentation has
been considered of great taxonomic value, it shows
wide infraspecific variation and must be used with care.
Leaf costa.—The costa typically ends below the
apex, and seldom reaches the apex or beyond.
Ventrally it is always smooth, and dorsally can be
smooth or papillose, spinose in Z. pichinchensis. In
cross-section it has two ventral guide cells and a band
of dorsal stereids (Zander & Vitt 1979), weakly
differentiated only in Zygodon obtusifolius. Costa
anatomy allows the discrimination of sterile material
of other genera with similar habits. Thus, Leptodontium (Müll Hal.) Hampe ex Lindb. (Pottiaceae) and
Amphidium Schimp. (Rhabdowesiaceae), that in the
past were considered Orthotrichaceae because of their
gametophytic similarity with Zygodon, have central
guide cells and dorsal and ventral stereid bands.
Leaf cross-section.—The leaf lamina in Zygodon is
typically unistratose, except for the bistratose portions
seen only seldom in the margins of Z. menziesii, and
in the recently described Z. bistratus Calabrese & J.
Muñoz, from the Iberian Peninsula, with bistratose
lamina (Calabrese & Muñoz 2006).
Distal laminal cells (Fig. 1).—Cells in the distal
2/3 of leaf are isodiametric (sub-quadrate, hexagonal
or rounded) to oval, 6 thick-walled and usually 8–15
lm in the larger diameter, although the European Z.
forsteri, with cells 15–27 lm, clearly deviates.
Cell surfaces are flat in all species except in Z.
obtusifolius, with distinctly bulging cells (Figs. 1E, 6F).
Most Zygodon taxa have distal cells papillose,
although the group of closely related species considered here as subg. Codonoblepharon, in which I
include Z. forsteri (cf. Matcham & O’Shea 2005), have
completely smooth cells. This is a rare characteristic in
the Orthotrichaceae, where only Muelleriella Dusén,
Stoneobryum D. H. Norris & H. Rob. and few
unrelated Orthotrichum Hedw. species lack papillae.
However, in Zygodon, all species with smooth leaf cells
also share other propagular and sporophytic traits
that support the taxonomic value of this character and
the validity of subg. Codonoblepharon.
Shape, size and number of papillae per cell are
critical for species recognition. Most Zygodon species
have simple papillae (Figs. 1B, D, 8D, 10G, 12E,
14D–E, 17D–E, 19D–E, 20E, 21E–F, 22E–F), with Z.
bartramioides, Z. papillatus and Z. pichinchensis (Figs.
Calabrese: Zygodon in southern South America
457
Figure 1. Cells in the middle portion of leaf. A. Zygodon papillatus (Gay s.n., PC-MONTAGNE). B. Z. magellanicus (Matteri & Schiavone
CM 2449, S). C. Z. pichinchensis (Matteri et al. CM 6144, BA). D. Z. chilensis (Eyerdam 10999-a, UC). E. Z. obtusifolius, the arrow points
to a forked papilla (Cleef 6062, G). F. Z. menziesii (Schwabe 81, S-B1887). Scale: 25 lm.
1A, 15D–E, 16H) also having branched ones intermingled. Finally, Z. obtusifolius has only fused or
branched papillae (Figs. 1E, 6D, F). Papillae can be
acute, sub-acute or blunt, usually less than 5 lm high,
although in Z. pichinchensis they are over 10 lm (Fig.
1C). In Z. pentastichus, Z. chilensis and Z. bartramioides, the only southern South American members
of sect. Stenomitrium, papillae are wider in diameter
than any other species in the genus (Fig. 1D). Among
the studied species, number of papillae per cell varies
from one to three in Z. pichinchensis to six to nine in
Z. hookeri var. leptobolax.
Basal laminal cells (Fig. 2).—Cells in the basal 1/3
of leaf are usually strongly differentiated from distal
cells: smooth or seldom with few papillae, subquadrate to rectangular, hyaline and commonly much
longer. The exception is Z. obtusifolius, with smooth
to weakly papillose basal cells, otherwise identical to
distal ones.
In contrast, Z. bartramioides and Z. pentastichus
have a strong and sudden differentiation between
upper and basal cells. The other species show a 6
gradual transition, although basal cells have a net
morphologic differentiation. The almost undifferentiation of basal cells in Z. obtusifolius has been
considered as a primitive character (Goffinet et al.
1998) and should be of taxonomic importance.
Basal cells are isomorphic except in three species,
Z. bartramioides, Z. chilensis and Z. pentastichus,
where they are arranged in dimorphic longitudinal
bands of 2–9 rows of yellowish, nodulose, thickwalled cells alternating with 2–6 rows of lighter,
hyaline, straight, thin-walled cells. Previously this
character was only noticed for Z. pentastichus, and
considered of no taxonomic value. However, species
exhibiting this basal cell pattern share other leaf
laminal and sporophytic features, which support their
segregation in a distinct group at the infrageneric
rank, as sect. Stenomitrium.
Perichaetial leaves.—In most Zygodon species the
perichaetial leaves are not or only slightly differentiated, and such is the case among the treated
species Z. menziesii, Z. intermedius, Z. jaffuelii and Z.
papillatus. In other species the perichaetial leaves are
differentiated from upper vegetative ones in morphology and areolation, usually having longer,
smooth or slightly papillose cells. Zygodon pentastichus, Z. bartramioides, Z. chilensis, Z. pichinchensis,
Z. reinwardtii, Z. hookeri var. hookeri and Z. hookeri
var. leptobolax clearly show this differentiation, while
in Z. obtusifolius it is less evident.
This character, despite its taxonomic value at
specific level, has not usually been used in keys
because most specimens lack mature perichaetial
leaves.
Propagula.—In the Orthotrichaceae, propagula
are not common but in Zygodon, where most species
have them, only Z. pichinchensis and Z. pentastichus
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Figure 2. Leaf basal cells. A. Zygodon reinwardtii (Skottsberg 435, S). B. Z. hookeri var. hookeri (Matteri & Schiavone CM 4910, BA). C.
Z. obtusifolius (Cleef 6062, G). D. Z. bartramioides (Schwabe 61, JE-HERZOG). E, F. Z. pentastichus (Hosseus 171, JE). Scale A–D, F: 25
lm; E: 0, 25 mm.
among the treated species lack them. Another
distinctive feature is that in other Orthotrichaceae the
propagula develop on leaves or, rarely, on rhizoids,
while in Zygodon they typically originate in leaf axils.
Propagulum length varies between 30–270 lm
and is characteristic for each species. Fig. 3 compares
propagulum size and shape in some of the treated
species. Abundance of propagula is also variable.
Zygodon hookeri var. hookeri, Z. reinwardtii and Z.
intermedius have few propagula, or sometimes lack
them all together, and moreover are difficult to see
because they are hyaline to greenish-hyaline. In
contrast, propagula are frequent and abundant in Z.
bartramioides, Z. jaffuelii and Z. hookeri var. leptobolax, and very abundant and evident in Z. magellanicus,
Z. papillatus and Z. chilensis.
Traditionally, propagula represent an important
source of taxonomic characters, especially when
plants are sterile, because of their morphological
diversification and common appearance. Shape, size,
Calabrese: Zygodon in southern South America
459
Figure 3. Propagula. A. Zygodon reinwardtii (Ule 131, G). B. Z. jaffuelii (RP-6-2285-1b, MO). C. Z. papillatus (lack collector data, FH).
D. Z. magellanicus (Dusén 83, S). E. Z. menziesii (Matteri & Schiavone CM 3768, BA). F. Z. bartramioides (Dusén 362, S). Scale: 30 lm.
color, number per plant and cell organization are of
great use in naming Zygodon species.
Malta (1926) recognized two basic types of
propagula based on their structure: those with only
transverse septa, and those with transverse and
longitudinal septa. However, in this study I have
found a more complex pattern of propagule morphology, with three main types:
a) Fusiform to fusiform-clavate, generally of 3–5 cells,
with transverse septa and seldom some longitudinal ones, usually hyaline, rarely brown (Z.
jaffuelii). It is the most common type in the genus,
seen, among the treated species, in Z. hookeri var.
hookeri, Z. hookeri var. leptobolax, Z. intermedius,
Z. reinwardtii and Z. jafuelii (Fig. 3A–B).
b) Cylindrical to cylindrical-clavate, of 4 or more
cells, with transverse septa or with some longitudinal septa in the apical or basal cells, or both. It
is the most common type in subg. Codonoblepharon. Species from the Andean-Patagonian and
Fuegian regions with this type of propagula are Z.
bartramioides, Z. menziesii and Z. obtusifolius, with
only transverse septa and hyaline walls, and Z.
chilensis, with brownish cells, except often the
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apical and basal hyaline cells occasionally with
some longitudinal septa (Fig. 3E–F).
c) Ovoid to clavate, with a variable number of cells,
with transverse and longitudinal septa, reddishbrown to yellowish-brown. This type is only found
in some local species of sect. Zygodon, Z.
magellanicus (Fig. 3D) and Z. papillatus (Fig. 3C).
Although similar in structure, propagula of Z.
papillatus are smaller, more ovoid and with walls
lighter in color than propagula of Z. magellanicus.
Branched propagula have only seldom been
observed in Z. chilensis (Fig. 12H). Exceptional in the
Orthotrichaceae, branched propagula are only known
in Orthotrichum lyellii Hook. & Taylor, O. obtusifolium Brid. and O. gymnostomum Brid.
Reproductive organs.—Distribution of gametangia is variable in Zygodon and of taxonomic
importance. While in other Orthotrichaceae, such as
Orthotrichum and Ulota D. Mohr, an autoicous
sexual condition is the rule, most Zygodon species
are dioicous, while some are autoicous (e.g., Z.
obtusifolius), and even fewer synoicous (e.g., Z.
reinwardtii).
Malta (1926) discussed the heteroicous sexual
condition, although this is only a funtional character.
He described three types of ‘‘flowers’’ in Zygodon
reinwardtii: ‘‘female flowers’’ (antheridia absent or
reduced and 3–4 archegonia, of which only one is
fertilized), ‘‘male flowers’’ (10–25 antheridia and 1–2
archegonia usually not fertilized), and ‘‘hermaphrodite flowers’’ (2–5 antheridia and 3–4 archegonia, of
which usually only one is fertilized). In fact, all this
cases represent the synoicous sexual condition,
although likely they are functionally unisexual when
archegonia or antheridia are reduced. I have observed
hermaphroditic buds only in specimens of Z.
reinwardtii. Finally, in the usually synoicous Z. hookeri
var. hookeri I once found separated perichaetia and
perigonia in the same plant.
Gametangia are usually terminal in stems.
Nevertheless, Malta (1926) pointed out that in the
dioicous Zygodon schenkei Broth. and Z. perichaetialis
Herzog (not in the study area) perichaetia are 6
lateral on short branches.
Vaginula.—Little is known about the vaginula in
Zygodon, as it has been seldom included in formal
descriptions. Vaginulae can have persistent uniseriate
paraphyses (e.g., Z. hookeri) or be naked (occasionally
with some short paraphyses) as in Z. obtusifolius and
species of subg. Codonoblepharon. Malta (1926) also
indicated that sometimes the upper part of the
vaginula has robust and persistent hairs (e.g., Z.
trichomitrius and Z. pilosulus Müll. Hal.), and
suggested a close relationship between persistence of
these hairs and a pilose calyptra, because the two
species have calyptrae with uniseriate hairs. Ochreae
in Zygodon are typically very short and almost
undifferentiated.
Calyptra.—The calyptra is typically cucullate,
smooth and naked, the latter frequently used to
distinguish Zygodon from other Orthotrichaceae.
However, there are three species with pilose calyptrae:
Z. trichomitrius and Z. erosus Mitt. from South Africa,
and Z. pilosulus from South America. The calyptra of
Z. obtusifolius has been described as having prorate
cells toward the apex (Goffinet & Vitt 1998), and thus
slightly papillose (Lewinsky 1990; Malta 1926). I could
not confirm this because all the studied specimens of
Z. obtusifolius lacked calyptrae because they fall soon
after sporophyte maturation.
Sporophyte.
Seta.—Zygodon always has a relatively long,
straight to slightly curved, usually yellowish to
yellowish-brown, rarely reddish-brown seta.
Torsion of the seta, despite its taxonomic
importance, has not been previously considered in
Zygodon treatments (Düll 1985; Lewinsky 1990; Malta
1926; Vitt 1970, 1984), and only Vitt et al. (1993) used
it to separate Zygodon, Orthotrichum, Ulota and
Groutiella from other Orthotrichaceae. Most Zygodon
species have setae that are distally dextrorse (twisted
to the right, clockwise when seen from the apex) and
proximally sinistrorse (twisted to the left, counterclockwise when seen from apex).
Distally sinistrose setae were considered crucial
by Goffinet and Vitt (1998) to segregate Zygodon
pentastichus at the generic rank as Stenomitrium.
However, I have found distally sinistrorse setae in
three additional species (Table 1). Two of them, Z.
bartramioides and Z. chilensis, which clearly belong in
Zygodon, share with Z. pentastichus ovate distal cells
with 2–4 thick papillae and basal cells in dimorphic
bands, which indicates a taxonomic relationship, and
that they would be the nexus between Z. pentastichus
Calabrese: Zygodon in southern South America
461
Table 1. Setae torsion in Andean-Patagonian and Fuegian Zygodon.
Torsion of the seta
Species
Subg. Codonoblepharon
Z. menziesii
Subg. Zygodon sect. Zygodon
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Subg. Zygodon sect. Stenomitrium
Subg. Obtusifolium
jaffuelii
intermedius
pichinchensis
papillatus
reinwardtii
hookeri var. hookeri
hookeri var. leptobolax
Z. bartramioides
Z. pentastichus
Z. chilensis
Z. obtusifolius
and Zygodon s.str. Finally, Z. obtusifolius also has
distally sinistrorse setae.
The basal part of the seta is sinistrorse in most
Zygodon species, although some specimens of Z.
pentastichus and Z. bartramioides have setae without
torsion proximally, and Z. chilensis always has setae
dextrorse at the base.
Capsule.—Zygodon has exserted capsules, typically erect and eight-ribbed throughout. Despite the
apparent homogeneity, capsules can be 6 cylindrical,
ovoid or ellipsoid to fusiform, and its shape can be
used in species recognition. The exothecial cells are
rectangular, differentiated in bands of 3–6 columns of
thick-walled cells corresponding to the ribs that
alternate with bands of thin-walled cells. Stomata are
superficial and invariably present in the lower part of
the urn and the neck.
Operculum.—In Zygodon the operculum always
has a conic base and usually an oblique beak that
varies from short-rostrate (e.g., Zygodon papillatus) to
long-rostrate (e.g., Z. hookeri). Sometimes it is useful
for species recognition, because the rostrum is
extraordinarily long in some species (e.g., Z. pichinchensis or Z. reinwardtii) and in others very short (e.g.,
Z. papillatus or Z. obtusifolius).
Peristome.—The peristome of Zygodon is arthrodontous and diplolepideous as in all Orthotrichaceae.
It could have been derived from a peristome with an
exostome of 16 free teeth more thickened on the OPL
(outer peristomial layer) than on the PPL (primary
In the distal part
In the basal part
dextrose
sinistrorse
sinistrorse
Sinistrorse or without torsion
dextrorse
sinistrorse
peristomial layer), and an endostome of 16 segments
(Goffinet & Vitt 1998; Vitt 1981).
In Zygodon, the more complex peristome has
sixteen exostomial teeth and sixteen endostomial segments alternating with the teeth, usually united at base
by a low connective membrane. Some species with this
type of peristome also have a preperistome or prostome
(Lewinsky 1990; Malta 1926). One of the most
remarkable tendencies in Zygodon is the gradual loss of
the exostome, from a slight reduction to its complete
disappearance, giving rise to simple peristomes formed
only by endostomial segments. These segments can also
undergo reduction, and thus it is possible to find
endostomial segments well developed, one or two cells
wide, or even rudimentary. As a consequence of
exostome and endostome reductions, the following
peristomial combinations are known in Zygodon:
a) 8 pairs of exostomial teeth and 8 pairs of endostomial segments (e.g., Z. pentastichus, Fig. 9A)
b) 8 pairs of exostomial teeth and an endostome with 8
well-developed segments and to 8 weaker intercalary segments (e.g., Z. obtusifolius, Figs. 6I, 7A)
c) 8 pairs of exostomial teeth and 8 well-developed
endostomial segments (e.g., Z. chilensis, Figs. 12I,
13A)
d) 8 pairs of exostomial teeth diversely reduced and 8
well-developed endostomial segments (e.g., Z.
intermedius, Fig. 18A)
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109(4): 2006
e) peristome of only 8 well-developed segments (e.g.,
Z. reinwardtii, Fig. 20K)
f) peristome of only 8 reduced segments (e.g., Z.
papillatus, Fig. 15J)
g) total absence of peristome (none in the study area,
e.g., the Holarctic Z. viridissimus)
This reductional series can be symbolized by the
following scheme:
8p/16 ! 8p/8þ8 ! 8p/8þn ! 8p/8 ! 8pred./8 ! 0/8 ! 0/8red. ! 0/0
double peristomes
single peristomes
Exostome teeth are reflexed or recurved when
dry, while endostome segments are 6 erect or
incurved. The teeth and the segments have hygroscopic movements that help spore liberation. According to Koponen (1978), the structure of the
peristome is related to hygroscopic movements;
exostomes of equal thickness on the external and
internal surfaces are erect, while exostomes thicker on
the external side are reflexed or recurved. According
to Lewinsky (1990), the type of ornamentation has
influence on exostome movement. For the present
work I used light microscopy and SEM, and confirm
that exostome teeth are ornamented on both the
external (OPL) and the internal (PPL) surfaces, and
that the ornamentation of the two is different. Usually
the OPL has a basal reticulum that varies from
smooth to papillose, while the exostomial PPL is often
ornamented by papillae irregularly arranged or 6
aligned, according to the species. Among the treated
species, only Z. intermedius has the exostome teeth
(that are very reduced) with completely smooth cells.
Also the endostome segments usually differ in surface
ornamentation: while the endostomial PPL is often
smooth or weakly ornamented, the IPL is often
papillose, mostly at base. When the endostomial
segments are reduced (Z. papillatus and all the
synoicous taxa) both PPL and IPL surfaces are
completely smooth. The connective basal membrane
usually has a similar ornamentation to that of the base
of the endostomial segments.
As has been already indicated, in Zygodon
papillatus and the synoicous species of the study area,
peristomes can show great reduction. Spore liberation
in these taxa is thus controlled by the capsule through
the strong longitudinal ribs that reach the mouth.
When dry, the contraction of the ribs closes the
capsule mouth, further blocked by the approximation
of the endostomial segments; when moist, the capsule
mouth opens, separating the endostomial segments,
which makes possible the liberation of the spores. This
mechanism allows for spore liberation only under
moist environmental conditions. As indicated by Lara
et al. (1999), this morpho-functional adaptation to
hygrocastique spore liberation, is also known in other
Orthotrichaceae, such as Macromitrium Brid., Orthomitrium Lewinsky-Haapasaari & Crosby and Orthotrichum. Considering other characters, not all the
species of Zygodon with exostomial reduction are
closely related, which points to morphological convergence that has evolved independently in several
groups of the family.
Spores.—All Zygodon species are isosporous.
Spores are spherical, sometimes slightly flattened, and
usually have an ornamented surface. In the Andean
Patagonian and Fuegian species spores are papillose
(Z. pentastichus, Z. pichinchensis, Z. jaffuelii, Z.
intermedius, Z. papillatus, Z. reiwardtii, Z. hookeri var.
hookeri and Z. hookeri var. leptobolax); foveolate (Z.
bartramioides and Z. chilensis); or smooth (Z.
menziesii and Z. obtusifolius). According to size and
presence of trilete mark, species can be included in
two groups: a) spores 10–15 lm in diameter, usually
with a trilete mark characterize Z. papillatus, Z.
jaffuelii, Z. obtusifolius, Z. menziesii, Z. bartramioides
and Z. chilensis; and b) spores more than 15 lm in
diameter without a trilete mark characterize Z.
pichinchensis, Z. intermedius, Z. pentastichus, Z.
reinwardtii and Z. hookeri.
TAXONOMIC TREATMENT
Zygodon Hook. & Taylor, Muscol. Brit. 70. 1818.
TYPE: Zygodon conoideus (Dicks.) Hook. & Taylor
[basionym: Bryum conoideum Dicks.]
Pleurozygodontopsis Dixon, Ann. Bryol. 12: 51.
1939. TYPE: P. decurrens Dixon
Bryomaltaea Goffinet, Bryology for the Twentyfirst Century 151. 1998. TYPE: B. obtusifolia (Hook.)
Goffinet [basionym: Zygodon obtusifolius Hook.]
Codonoblepharon Schwägr., Spec. Musc. Frond.,
Suppl. 2(2): 142, tab. 137. 1824. TYPE: C. menziesii
Schwägr.
Stenomitrium (Mitt.) Broth., Nat. Pflanzenfam.
Calabrese: Zygodon in southern South America
1(3): 464. 1902 [1903]. TYPE: S. pentastichum (Mont.)
Broth. [basionym: Aulacomnium pentastichum Mont.]
Description. Plants 0.2–15 cm tall, in turfs, rarely
mats, light-green to olive-green, occasionally darkgreen or reddish-brown, growing on bark or rock,
rarely on soil, mortar or cement. Stems erect, rarely
prostrate, simple or branched, rounded to pentagonal
in cross section; cortex of small, thick-walled cells, in
1–3 layers around large, isodiametric, thin-walled,
undifferentiated cells. Rhizoids yellowish-brown to
reddish-brown, exceptionally hyaline, smooth,
roughened or papillose, sometimes covering the stem
as a reddish-brown tomentum. Leaves appressed or 6
twisted when dry, erect-spreading to squarroserecurved when moist, sometimes undulate, typically
keeled, 0.5–3 3 0.15–0.9 mm, ovate-lanceolate to
linear-lanceolate, lanceolate-rhomboidal or lingulate,
sometimes decurrent; lamina unistratose or partially
bistratose; apex acute, rarely rounded-obtuse; margins
undifferentiated, usually plane, rarely recurved (especially in the middle and lower parts of leaf), entire,
papillose or papillose-crenulate, rarely spiculose,
sometimes denticulate to dentate at the apex; upper
laminal cells 6 isodiametric, rounded-hexagonal to
subquadrate or ovate, plane, rarely bulging, papillose
or smooth, thick-walled, 8–15(27) lm in diameter;
papillae usually simple, occasionally branched, blunt,
acute or subacute, 1–9 per cell; basal cells shortrectangular to elongate, rarely isodiametric, smooth
or with scarce papillae, thin- or thick-walled, sometimes nodulose, hyaline or yellowish, isomorphic or
dimorphic; costa simple, usually ending below the
apex, rarely percurrent or excurrent, usually in a deep
furrow, with superficial elongate cells, dorsally
smooth or papillose, usually smooth in the ventral
surface, in section with two ventral guide cells and
dorsal stereids, rarely with 6 homogeneous cells.
Propagula often present in leaves axils, cylindrical,
fusiform, clavate or ovoid, with only transverse septa
or with transverse and longitudinal septa, hyaline or
yellowish-brown to reddish-brown walls. Dioicous,
synoicous or autoicous. Perigonia axillary or terminal;
leaves concave, ovate, apex acute. Perichaetia terminal; leaves differentiated or not from stem leaves,
usually lanceolate to linear-lanceolate with acute,
rarely rounded-obtuse apex. Vaginula glabrous or
pilose. Seta long, erect to slightly curved, distally
463
dextrorse or sinistrorse, usually sinistrorse in the basal
part, exceptionally without torsion or dextrorse at
base, often yellowish-brown; capsule exserted, erect,
ovoid, ellipsoidal or cylindrical, with 8 longitudinal
strong ribs when dry, sometimes slightly contracted
below the mouth, with phaneropore stomata in lower
part of urn and neck; exothecial cells of the ribs
differentiated; operculum conic at base, short to longrostrate, beak straight or oblique; annulus absent;
peristome double, single or absent; exostome teeth
partially or completely united in 8 pairs, ornamented,
reflexed or appressed to the capsule when dry, OPL
papillose, papillose-striate, smooth-reticulate, papillose-reticulate or papillose-lamellate, PPL papillose,
papillose-striate or almost smooth; prostome very
rarely present; endostome of 8 or 16 free segments
alternating with exostome teeth, usually united by a
low basal membrane, rarely fused in the lower 1/3,
inflexed, smooth or ornamented, PPL usually smooth,
occasionally slightly striate or with disperse papillae,
IPL smooth, papillose or papillose-striate. Calyptra
conical, cucullate, glabrous and smooth, rarely pilose
or slightly papillose, covering the upper 2/3 of the
capsule. Spores 10–30 lm in diameter, papillose,
foveolate or smooth, trilete mark distinct or not.
Ecology. Species of Zygodon usually grow on
trees, stumps, also or exclusively on rocks, and rarely
on soil, exposed roots of old trees, sand, mortar or
concrete. Habitat information is scarce for the treated
species, and most data are derived from field work by
the author. In southern South America, they are most
common in the Andean Nothofagus forests, from sea
level to 6 1100 m, always in rather humid places, also
near streams or lakes in riparian N. antarctica forests,
while some species grow in more open and exposed
areas, such as the northern Patagonian NothofagusAustrocedrus mixed forests, that to the east contact
with steppe vegetation. Zygodon species commonly
grow on the native Austrocedrus chilensis, Araucaria
araucana, Fitzroya cupressoides, Nothofagus spp. and
Podocarpus nubigena, and some shrubs, mainly
Berberis. Exotic trees are exceptional, with only one
report of Z. bartramioides growing on Eucalyptus.
In other areas of America, such as Mexico,
Venezuela and Colombia, Zygodon grows at 1000–
4500 m, from the Andean forests to the desert in
Colombia (Churchill & Linares 1995), and Quercus
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109(4): 2006
Table 2. Taxa from the study area and their correspondence with other taxa previously reported.
Taxa previously reported from the study area
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Z.
Z.
menziesii (Schwägr.) Arnott
corralensis Lor.
obtusifolius Hook.
obtusifolius var. antarcticus Malta
pentastichus (Mont.) Müll. Hal.
pentastichus var. crispatulus (Thér.) Malta
pentastichus var. strictifolius Malta
bartramioides Malta
tristis Herzog
jaffuelii Thér.
liebmannii Schimp.
pichinchensis (Taylor) Mitt.
magellanicus Malta
papillatus Mont.
intermedius Bruch & Schimp.
reinwardtii (Hornsch. )A. Braun
reinwardtii var. hyadesii (Besch.) Malta
hookeri Hampe
inermis Malta
theriotii Herzog
Taxa accepted in this work
Taxa previously reported
from other areas
Z. menziesii (Schwägr.) Arnott
Z. obtusifolius Hook.
Z. pentastichus (Mont.) Müll. Hal.
Z. bartramioides Malta
Z. jaffuelii Thér.
excluded
Z. pichinchensis (Taylor) Mitt.
Z. magellanicus Malta
Z. papillatus Mont.
Z. intermedius Bruch & Schimp.
Z. reinwardtii (Hornsch.)
Bruch & Schimp.
Z. hookeri Hampe
Z. hookeri var. leptobolax
(Müll. Hal) Calabrese
Z. chilensis Calabrese & Lara
and Abies forests in Mexico (Vitt 1994). Lewinsky
(1990) provided a thorough description of Australasian Zygodon habitats, while Düll (1985), LewinskyHaapasaari (1998) and Smith (2004) describe the
European and Macaronesian habitats.
Distribution. According to Index Muscorum
(Wijk et al. 1969), Zygodon includes 90 species, of
which more than 50 are distributed in the tropics and
subtropics, while the rest are distributed in the
temperate regions of both hemispheres. The Andean
region, with 49 species reported, can be considered
the main center of diversity for the genus.
For the Andean-Patagonian and Fuegian regions
I recognize 13 taxa (Table 2). Five of them are
endemic (38.5%): Z. bartramioides, Z. chilensis, Z.
jaffuelii, Z. magellanicus and Z. papillatus. Andean
species are Z. pentastichus (from Peru to 478S in
Patagonia and the Juan Fernández Islands) and Z.
pichinchensis (throughout the Andes and Mexico).
Zygodon menziesii and Z. hookeri var. hookeri
(Australia, New Zealand and southern South America), as well as Z. hookeri var. leptobolax (South Africa
and southern South America) show a sub-Antarctic
Z. leptobolax Müll. Hal. (South Africa)
distribution. Finally, widespread in the Southern
Hemisphere are Z. intermedius, Z. obtusifolius and Z.
reinwardtii, the latter also known from Central and
South America, and with isolated reports that need
confirmation for Alaska (very doubtful, but cf. Worley
& Iwatsuki 1970), Madagascar (Crosby et al. 1983),
Philippines (Tan & Iwatsuki 1991), and Tasmania
(Dalton et al. 1991; but see Lewinsky 1990, who did
not mention it for Australasia).
Although it can be argued that the number of
species treated here is low, they are highly representative within the genus and allow one to understand
the relationships between taxa in a wider context.
Indeed, the analysis of taxonomic characters suggests
that the four genera that have been segregated from
Zygodon (Bryomaltaea, Codonoblepharon, Stenomitrium and Zygodon s.str.) have more similarities than
differences from each other, and no clear separation
can be established between them when the variability
of the group is analyzed as a whole. As a consequence,
these four taxa are here considered as part of a
broadly conceived Zygodon, for which a new infrageneric classification is proposed, including three
Calabrese: Zygodon in southern South America
465
Table 3. Historical outline of infrageneric classification of the genus Zygodon and the classification proposed in the present work.
Brotherus (1902–1903) Malta (1923, 1926) Goffinet & Vitt (1998) Goffinet et al. (2004)
Gen.
Gen.
Gen.
Gen.
Zygodon
Stenomitrium
Zygodon
Zygodon
Sect.
Sect.
Sect.
Sect.
Zygodon
Stenomitrium
Obtusifolii
Bryoides
Gen.
Gen.
Gen.
Gen.
Zygodon
Stenomitrium
Bryomaltaea
Codonoblepharon
subgenera, one with two sections. Table 3 shows the
historical outline of the infrageneric classifications as
well as the new classification proposed here.
A Synoptic Key to Infrageneric Taxa.
1. Leaf cells smooth.................. Subg. Codonoblepharon (p. 466)
1’. Upper laminal cells papillose ..................................................2
2. Laminal cells bulging; leaf apex rounded-obtuse..........
.............................................Subg. Obtusifolium (p. 469)
2’. Laminal cells plane; leaf apex acute to acuminate,
exceptionally rounded-obtuse...........(Subg. Zygodon) 3
3. Leaf basal cells dimorphic, porose and thick-walled cells
alternating with bands of straight and thin-walled cells; seta
distally sinistrorse; peristome double .......................................
...................................................... Sect. Stenomitrium (p. 474)
3’. Leaf basal cells never dimorphic, not in differentiated bands;
seta distally dextrorse; peristome double, single or absent
..............................................................Sect. Zygodon (p. 484)
Key to Andean-Patagonian and Fuegian Taxa of
Zygodon.
1. Laminal cells smooth; propagula cylindrical, with only
transverse septa; peristome double......... 1. Zygodon menziesii
1’. Laminal cells papillose; propagula fusiform, cylindrical,
clavate or ovoid, with only transverse septa or with
transverse and longitudinal septa; peristome double or
single ........................................................................................2
2. Laminal cells bulging; basal cells morphologically
undifferentiated from upper cells, except for the
presence of papillae; apex rounded-obtuse; peristome
double; autoicous................................... 2. Z. obtusifolius
2’. Laminal cells plane; basal cells morphologically differentiated from upper cells; apex acute to acuminate;
peristome double or single; dioicous or synoicous ....3
3. Basal leaf cells dimorphic in definite bands; stems erect or
prostrate; leaves clearly 5-ranked; setae distally sinistrorse;
peristome double.......................................................................4
3’. Basal leaf cells not in bands of different colors; stems erect;
leaves not or weakly 5-ranked; setae distally dextrorse;
peristome double or simple ....................................................6
4. Stems prostrate with erect branches; leaves more than
2 mm long; spores large, 17–28 lm in diameter,
papillose .................................................3. Z. pentastichus
Gen. Leratia p.p.
Proposed classification
Subg. Zygodon Sect. Zygodon
Sect. Stenomitrium
Subg. Obtusifolium
Subg. Codonoblepharon
4’. Stems erect; leaves less than 2 mm long; spores small,
10–12 lm, foveolate .......................................................5
5. Leaves lanceolate-rhomboidal; propagula 40–67 3 15–23 lm,
hyaline, usually of 4 cells, with only transverse septa; setae 2–
5 mm long ................................................... 4. Z. bartramioides
5’. Leaves lanceolate to narrow-lanceolate; propagula (75–)87–
120 3 24–30 lm, usually with colored walls, of 4–9 cells,
with transverse septa and sometimes also longitudinal septa;
setae ca. 10 mm long.......................................... 5. Z. chilensis
6. Dioicous; peristome double or single; spores than 18
lm .....................................................................................7
6’. Synoicous; peristome simple, reduced to 8 short
endostomial segments; spores usually more than 18
lm ............................. (Z. reinwardtii-hookeri group) 11
7. Propagula clavate to oblong-clavate, with transverse and
longitudinal septa, colored, abundant ....................................8
7’. Propagula fusiform to clavate, without longitudinal septa,
hyaline or colored, or plants without propagula..................9
8. Propagula large, 50–90 3 30–39 lm, reddish-brown;
rhizoids roughened or papillose; laminal cells with
simple papillae......................................6. Z. magellanicus
8’. Propagula small, 19–32 3 17–25 lm, yellowish-brown;
rhizoids smooth; laminal cells with branched papillae
...................................................................7. Z. papillatus
9. Leaf margins spinulose by large conic papillae; papillae acute
or subacute; plants without propagula; peristome simple
....................................................................... 8. Z. pichinchensis
9’. Leaf margins papillose; papillae low; plants with propagula;
peristome double ..................................................................10
10. Propagula hyaline, fusiform, scarce; peristome double
with reduced exostome of smooth or almost smooth
cells ....................................................... 9. Z. intermedius
10’. Propagula colored when mature, clavate, abundant;
peristome double with exostome of 8 pairs of teeth
distally truncate and ornamented ......... 10. Z. jaffuelii
11. Leaf apex dentate, sometimes denticulate; leaves distinctly
decurrent; spores (15–)18–25 lm; capsules often fusiform,
sometimes asymmetric................................ 11. Z. reinwardtii
11’. Leaf apex entire or denticulate; leaves decurrent, at least
slightly; spores (18–)20–30 lm; capsules cylindrical
............................................................. (12. Z. hookeri s.l.) 12
12. Leaf apex often denticulate; margins undulate; propagula scarce, 36–52 3 12–20 lm, consisting of 3
cells ......................................12a. Z. hookeri var. hookeri
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12’. Leaf apex usually entire; margins usually plane;
propagula abundant, 45–81(90) 3 16–30 lm, usually
consisisting of 3–5 cells12b. Z. hookeri var. leptobolax
Subg. Codonoblepharon (Schwägr.) Calabrese, stat. nov.
Codonoblepharon Schwägr., Spec. Musc. Frond.,
Suppl. 2(2): 142, tab. 137. 1824; Zygodon sect.
Codonoblepharon (Schwägr.) Müll. Hal., Linnaea
18: 669. 1845; Zygodon sect. Bryoides Malta,
Latvijas Univ. Raksti 6: 281. 1923. TYPE: C.
menziesii Schwägr.
Diagnosis. Plants to 2 cm tall, growing on bark
or rock. Stems erect. Leaves lanceolate; apex acute;
cells smooth, not bulging. Propagula cylindrical, with
transverse septa (except Zygodon menziesii var.
angustifolius very rarely with longitudinal septa).
Dioicous or autoicous. Seta dextrorse distally, sinistrorse at base. Peristome double. Spores small, less
than 18 lm in diameter, smooth, trilete mark distinct.
Discussion. This subgenus is defined by the
smooth laminal cells. This difference led Goffinet and
Vitt (1998) to re-establish Codonoblepharon as an
independent genus, a view accepted by Matcham and
O’Shea (2005). However, species of this group share
with other Zygodon species the unique propagula, the
cucullate calyptra, ribbed capsules, as well as the
structure of the double peristome, with 8 pairs of
teeth and 8 endostomial segments keeled and alternate
with exostomial teeth, which in my opinion justify the
maintenance of these species in Zygodon, although at
subgeneric rank. Further support for my taxonomic
arrangement is the position of Z. forsteri, which shows
all previous characters but formerly excluded from
Codonoblepharon by Goffinet and Vitt (1998), later
included on the basis of molecular evidence (Goffinet
et al. 2004), and again excluded by Matcham and
O’Shea (2005) on the basis of its white rhizoids, wider
cells and north temperate distribution.
Previous to this study Zygodon subg. Codonoblepharon was considered to include two species in the
area: Zygodon menziesii (He 1998; Matteri 1989) and
Z. corralensis (He 1998). The latter was considered
endemic to southern South America, but Calabrese
(2003) and Matcham and O’Shea (2005) clearly
demonstrated that it is a synonym of Z. menziesii.
1. Zygodon menziesii (Schwägr.) Arnott, Mém. Soc.
Linn. Paris 5: 233. 1827; Codonoblepharon
menziesii Schwägr., Spec. Musc. Frond., Suppl. 2
(2): 142, tab. 137. 1824. Ind. loc.: ‘‘In nova
Seelandia ad cortices legit cl. Archibald Menzies.’’
TYPE: NEW ZEALAND. STEWART ISLAND: Half Moon
Bay, Jun 1889, Bell 679 (neotype, here designated,
CHR 527696!). NOTE: The type material of Z.
menziesii, kept at G, was lost by the postal service
(P. Geissler, pers. comm., March 1, 2000).
Zygodon drummondii Taylor, London J. Bot. 5: 46.
1846. TYPE: ‘‘Australia, Swan River, Drummond
1843’’ (isotypes, BM!, H, L). Synonymized by
Wilson (1846: 449).
Zygodon corralensis Lorentz, Bot. Zeitung (Berlin) 24:
187. 1866. TYPE: ‘‘Chile, Valdivia, Corral, an den
Kalkmauern der Hafenfestung, Dr. Krause’’
(isotype, NY!). Synonymized by Matcham and
O’Shea (2005: 130).
Zygodon menziesii var. angustifolius Malta, Latvijas
Univ. Raksti 10: 317. 1924. TYPE: ‘‘Neuseeland:
Mauriceville (Gray).’’ (Lectotype, designated
here, Gray 47, BM!; syntypes, Gray 9, Gray 50, all
BM!), syn. nov.
Illustrations. Figs. 1F, 3E, 4–5.
Description. Plants slender, 0.5–2.0 cm tall, in
dense turfs, green above, brown to reddish-brown
below. Stems erect, branched. Rhizoids reddishbrown, roughened. Leaves twisted when dry, erect
when moist, 0.6–1.8(2.1) 3 0.2–0.5 mm, lanceolate
to lingulate, not decurrent, apex acute; margins plane
or recurved in lower 1/2–2/3, rarely bistratose, entire
and completely smooth; laminal cells smooth; upper
leaf cells sub-quadrate to shortly ellipsoidal or
rhomboidal, with 6 thickened walls, 8–12 3 7–12
lm; lower leaf cells sub-quadrate to rectangular,
longer, 15–45 3 10–20 lm, with a group of globose
basal cells; costa ending below the apex, typically
reddish, 30–57 lm wide in the lower 1/3, with
elongate, smooth cells, in section with 2 ventral guide
cells and dorsal stereids. Propagula on reddish-brown
axillary stalks, cylindrical, usually hyaline, greenish or
yellowish-brown, 53–92 3 14–21 lm, (3–)4–7(9)celled, only with transverse septa. Dioicous. Perichaetial leaves undifferentiated. Seta (2.5–)4–6(8)
mm long, dextrorse in the distal part and sinistrorse
to the base; capsule exserted, erect, ellipsoidal to
pyriform, ca. 1.5 mm long, with 8 longitudinal ribs
along the entire capsule length when dry; exothecial
bands differentiated, of 4–5 cell rows each; operculum
conical at base, shortly rostrate and curved; peristome
Calabrese: Zygodon in southern South America
467
Figure 4. Zygodon menziesii. A. Habit when dry. B. Habit when moist. C. Capsule when dry. D. Leaves. E. Leaf apex. F. Leaf basal
cells. G. Leaf cross-sections. H. Propagula. I. Peristome (inner view). J. Spores. A, C from Allison 2263 (CHR); B, dII, F, H from
Schwabe 81 (S-B1887); dI from C. & I. Skottsberg 371 (S); dIII, E, gI from Matteri & Schiavone CM 3768 (BA); dIV from Bell 679 (HII
BROTHERUS); g from Skottsberg 371 (S); I, J from Matteri CM 1534-b (BA).
468
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109(4): 2006
Figure 5. Zygodon menziesii. SEM photographs of peristome. A. Lateral view of the capsule mouth and peristome. B–D. Exostome. B.
OPL, basal. C. PPL, middle portion of teeth. D. PPL, apical. E, F. Endostome. E. PPL. F. Details of IPL ornamentation in the basal
portion of the segments. A–F from Norris 25598 (H).
double; exostome of 8 pairs of teeth, recurved and
appressed to the capsule when dry, dorsal surface
(OPL) with papillae on a reticulum in the basal part
and with papillae aligned or irregularly disposed
above, ventral surface (PPL) papillose, with papillae
longitudinally aligned in the lower 1/2–2/3, and
confluent papillae irregularly disposed at the upper
part; endostome of 8 segments dorsally keeled,
sometimes appendiculate, occasionally with intercalary segments (8þn, 8þ8), PPL smooth, IPL with
papillose lines obliquely oriented at the base, papillae
longitudinally aligned at the central part, and papillae
irregularly disposed (some of them merged) at the
distal part, basal membrane low, continuous, striatepapillose. Spores almost smooth, 15–18 lm in
diameter, trilete mark distinct.
Ecology. Zygodon menziesii usually grows on rocks,
more rarely on bark. In the Andean-Patagonian and
Fuegian region it is found on mortar, concrete and
bricks, and less frequently on bark. Few data are known
Calabrese: Zygodon in southern South America
on the phorophytes, as it has been reported only from
‘‘huahuán’’ (Laureliopsis philippiana (Looser)
Schodde). In Australia and New Zealand it is frequently
found on rock as well; however, it has been also collected
on Eucalyptus, and more rarely, on Nothofagus.
Distribution (Fig. 23A). The distribution of
Zygodon menziesii is typically Subantarctic. It ranges
from Australia, New Zealand and South Africa to
southern South America. In the Andean-Patagonian
and Fuegian region it is found in Argentina from
Tierra del Fuego, and in Chile from Osorno, Valdivia
and Chiloé, as well as in the Juan Fernández Islands.
Differentiation. The subg. Codonoblepharon is
represented in the study area only by Zygodon menziesii.
Confusion of this species with any other in the genus is
unlikely, because it lacks papillae on the leaf cells.
Variability. Zygodon menziesii exhibits considerable gametophytic variability. Leaves are especially
variable in size, width, margin recurvature, apex shape
and costa robustness and color. Regarding leaf size, in
some specimens they hardly exceed 0.6 3 0.2 mm
(e.g., Skottsberg 371, S!) (Fig. 4dI), whereas in others
they reach 1.8 3 0.5 mm (e.g., Streimann 58205, H!).
Leaf margins are usually recurved in the lower 2/3,
althought some specimens have plane margins, especially those extreme expressions with leaves either small
and very narrow, or large and very wide. This variability
was clearly pointed out by Lewinsky (1990: 127–129,
figs. 101, 105, 111), and confirmed by my own
observations. Shape of the apex is also variable; more
often the leaves are wide and suddenly narrowed at
apex, as reported by Malta (1926: 156, fig. 95d), but
there are also leaves more gradually narrowed to the
apex, even in the same plant. The costa is usually very
wide in relation to the leaf width, and although its color
can vary from greenish to reddish, it is typically reddish.
Propagula are also variable. Some of them consist of 3–
4 cells, whereas others have 8–9 cells. Propagula are
usually hyaline-green, but they can be also brownish.
This variation is often present within the same
collection (e.g., Matteri & Schiavone 3768, BA!).
According to Malta (1924) and Lewinsky (1990),
the New Zealand Z. menziesii var. angustifolius differs
from the type variety by the presence of longitudinal
septa in the propagula. However, most propagula in
the type material exclusively have transverse septa, and
only a few in Gray 47 (BM!) and Gray 50 (BM!) also have
469
longitudinal septa; moreover leaves in all the syntypes
are narrowly lanceolate, similar to the leaves in Chilean
materials of the type variety (e.g., Skottsberg 371, S!).
Zygodon corralensis was described to include
plants with narrower leaves gradually ending in an
acute apex, and with smaller propagula than Z.
menziesii. These features are variable in Z. menziesii,
as stated above, and hardly usable to distinguish taxa.
The type material of Z. corralensis and all the
specimens identified as Z. corralensis, including the
only other reported by Malta (Skottsberg 371, see
Malta 1926: 158), fall within the range of variation of
Z. menziesii. The specimen reported as Zygodon
corralensis from South Africa (Arts 1998: 444) was not
available for study, but according to the figure in Arts
(1998) it could belong in Z. menziesii.
Selected specimens examined (76). ARGENTINA.
TIERRA DEL FUEGO: Isla de los Estados, Bahı́a Liberty,
Matteri 1534 b (BA), Isla de Tierra del Fuego, Bahı́a
Buen Suceso, extremo oriental del Morro Sur, 1-II1986, Matteri & Schiavone CM 3768 (BA). CHILE.
REGIÓN V. VALPARAÍSO: Islas Juan Fernández, Masatierra,
Valle Anson, Plazoleta, Skottsberg 371 (S). REGIÓN X.
OSORNO: Yerbas Buenas, centro de recreación Las
Cascadas, Administración, Mahu 22465 (MO 3987813);
VALDIVIA: Corral bei Quitaluto, Hosseus 691 (JEHerzog); CHILOÉ: Chiloé, Lobb 37 (BM 555683a). REGIÓN
XI: Puerto Puyuhuapi, Schwabe 1 (JE -Herzog).
AUSTRALIA. SOUTH AUSTRALIA: Kangaroo Island,
Cape du Couedic road, 1 km S of Rocky River
Settlement, Flinders Chase National Park, 12 km NNE
of Cape du Couedic, Streimann 55009, Musci
Australasiae Exsicc. 468 (CHR 504018). TASMANIA:
Deloraine, Scott 994 (BA). VICTORIA: Ewarts Road,
Barandula Range, 16 km SSE of Wodonga, Streimann
15453 (H). WESTERN AUSTRALIA: along Collie River near
Collie Burn, Norris 25598 (H). NEW ZEALAND. NORTH
ISLAND: Tongariro National Park, Mt. Ruapehu,
Okahune Mtn. Rd., Frahm 29–3 (CHR 492307). SOUTH
ISLAND: Westland, Haast Pass, Frahm 8–23 (CHR
503116). STEWART ISLAND: Horseshoe Point, Deadman’s
Beach, Fife 5977 (CHR 461652). EDERBY ISLAND: NNE of
Sandy Bay, Vitt 8814 (CHR 447876). CHATHAM ISLAND:
Awamata Stream, c. 10 km SW of Waitangi, Fife
11308 (CHR 514636).
SUBG. Obtusifolium (Malta) Calabrese, stat. nov.
Zygodon sect. Obtusifolii Malta, Latvijas Univ. Raksti
470
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6: 282. 1923; Bryomaltaea Goffinet in Goffinet &
Vitt, Bryology for the Twenty-first Century 151.
1998. TYPE: Zygodon obtusifolius Hook.
Diagnosis. Plants small, 0.5–1.5 cm tall, in dense
turfs, often on bark. Stems erect, usually branched.
Leaves lingulate; apex rounded-obtuse; upper laminal
cells isodiametric, bulging, with papillae often forked
or fused, toward the base usually smooth or rarely
papillose, mostly undifferentiated from upper cells.
Propagula cylindrical, with only transverse septa.
Autoicous. Seta sinistrorse throughout. Peristome
double. Spores small, less than 15 lm in diameter,
finely papillose, trilete mark distinct.
Discussion. The diagnostic character for this
subgenus is the bulging nature of the leaf cells. Mostly
undifferentiated basal leaf cells are also rarely seen in
the genus, and help to separate it from other
subgenera. The subgenus only includes Zygodon
obtusifolius, widespread in the Southern Hemisphere,
South-East Asia, Central America and Mexico.
Based on rbcL sequence data, Goffinet et al.
(1998), suggested that Z. obtusifolius would be closely
related to members of subfamily Macromitrioideae,
while sections Zygodon and Bryoides would be
resolved in another clade. Accordingly, the species was
segregated in a new monospecific genus, Bryomaltaea
Goffinet. Goffinet and Vitt (1998: 151) characterized
Bryomaltaea by having obtuse, erect and appressed
leaves. However, these features are also present in
Zygodon conoideus (Dicks.) Hook. & Taylor var.
lingulatus S. R. Edwards, a member of sect. Zygodon.
According to Goffinet et al. (1998), the unexpected
relation of Z. obtusifolius with the Macromitrioideae
could be an artefact or an indication of chloroplast
genome capture from a macromitrioid taxon. Goffinet and Vitt (1998) retained the genus Bryomaltaea
close to Zygodon.
Recently, Goffinet et al. (2004) based on nad5
sequence data, transferred Bryomaltaea obtusifolia to
Leratia as L. obtusifolia (Hook.) Goffinet. These
authors mentioned bulging laminal cells, seta torsion
and prorate cells at the calyptra apex as the
morphological characters shared by Bryomaltaea and
Leratia. However, Leratia has totally smooth capsules
while Zygodon obtusifolius has strongly ribbed ones as
all other species of Zygodon. Torsion of the seta and
calyptra have been previously discussed in this work
(see taxonomic characters), and cannot be considered
to separate Zygodon from Bryomaltea or Leratia. In a
recent paper, Wilbraham and Long (2005) treated Z.
obtusifolius in Bryomaltea, but considered Leratia as a
distinct genus contrary to Goffinet et al. (2004).
Finally, according to Goffinet et al. (2004), trnL-trnF,
rps4, nad5 and LSU data are congruent with their
earlier hypothesis that Zygodon obtusifolius may be
related to the Macromitrioideae, and the clade
including Bryomaltaea and Leratia may indicate a
transition between Macromitrioideae and Orthotrichoideae (Goffinet et al. 2004).
Zygodon obtusifolius shares with all other members of Zygodon the acrocarpic habit, cucullate
calyptrae, all sporophytic features, and especially the
unique propagula which, with the inconclusive
molecular evidence, suggest that it is best treated at
the subgeneric level within Zygodon.
2. Zygodon obtusifolius Hook., Musc. Exot. 2: CLIX.
1819. Codonoblepharon obtusifolium (Hook.) A.
Jaeger, Ber. Thätigk. St. Gallischen Naturwiss.
Ges. 1872–73: 119 (Gen. Sp. Musc. 1: 397). 1874;
Bryomaltaea obtusifolia (Hook.) Goffinet in
Goffinet & Vitt, Bryology for the Twenty-first
Century 151. 1998; Leratia obtusifolia (Hook.)
Goffinet in Goffinet et al., Monogr. Syst. Bot.
Missouri Bot. Gard. 98: 286. 2004. TYPE: ‘‘Hab. In
Nepal. Hon. D. Gardner’’ (lectotype, designated
by Lewinsky (1990), BM!).
Codonoblepharon neglectum A. Jaeger, Ber. Thätigk. St.
Gallischen Naturwiss. Ges. 1872–73:118. 1874;
Zygodon neglectus Hampe ex Müll. Hal., Hedwigia 37:133. 1898. TYPE: ‘‘Nova-Seelandia in fretu
insularum (Colenso).’’ Synonymized by Malta
(1926: 162). NOTE: Codonoblepharon neglectum
has been considered an invalid name (nomen
nudum) by most authors, who only considered
the taxon validly published after Müll. Hal.
described Zygodon neglectus (typified by ‘‘New
Zealand, Knight’’) in 1898. However, A. Jaeger
(1874: 118–119) considered the New Zealand
material collected by Colenso distinct from Z.
obtusifolius Hook., and described them as C.
neglectum with a clear and direct reference to the
description in Hooker (1855), which, according
to the ICBN 32.1, 32.3, and 32.4 (St. Louis Code),
makes the name validly published. Malta correctly considered C. neglectum validly described
Calabrese: Zygodon in southern South America
by A. Jaeger, but his views were apparently
unnoticed by later authors.
Zygodon linguiformis Müll. Hal., Bot. Zeitung (Berlin)
16:163. 1858; Zygodon linguaeformis Hampe ex
Müll. Hal., Bull. Herb. Boissier 6: 95. 1898, nom.
inval. orthogr. err.; Codonoblepharon linguiforme
(Müll. Hal.) A. Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1872–73: 119. (Gen. Sp.
Musc. 1:397). 1874. TYPE: ‘‘Columbia. Ad truncis
arborum: Hampe donavit.’’ Synonymized by
Malta (1926: 162).
Zygodon spathulaefolius Besch., Mém. Soc. Sci. Nat.
Cherbourg 16: 187. 1872. TYPE: ‘‘Mexico. Prope
Mejico, in sylva della Desierta Vieja, sept. 1865,
Bourg. n8 1334.’’ Synonymized by Malta (1926: 162).
Zygodon erythrocarpus Müll. Hal., Linnaea 42: 365.
1879. TYPE: ‘‘Argentinia subtropica, Tucumán
prope Siambon in truncis Alni ferruginei, c. fruct.
maturis et cum Zygodonte ochraceo asociatus.’’
Synonymized by Malta (1926: 162).
Zygodon araucariae Müll. Hal., Bull. Herb. Boissier 6:
95. 1898. TYPE: ‘‘Brasilia, Serra Geral, ad truncos
Araucariae Brasiliensis, Junio 1890: E. Ule, Coll.
N8 665; Minas Geraës, Serra Itatiaia, 1500 m.
altus, ad truncos arborum in sylva, April 1894: E.
Ule, Coll. N8 1918, in Bryotheca Brasiliensi N8
130 sub Zygod. linguaeformi Hpe., qui non
invenitur.’’ (syntypes H-BR! ‘‘Ule 665;’’ BA-MATTERI! ‘‘Ule 665;’’ G-BOISSIER! ‘‘Ule, Bryotheca
Brasiliensi N8 130’’ and ‘‘Ule, Junio 1890’’).
Synonymized by Malta (1926: 162).
Zygodon obtusifolius var. antarcticus Malta, Latvijas
Univ. Raksti 10: 334, 1924. TYPE: CHILE. ‘‘Valparaı́so (Porter), Cordillera de la Costa supra
Angol in truncis arborum (Dusén N8 845); [. . .]
Corral, an Kalkmauern der Hafenfestung
(Krause) [. . .]; o. n. O. (Sainthill) als Z.
obtusifolius Hook.’’ (lectotype, here designated,
‘‘Dusén N8 845’’, S!), syn. nov.
Zygodon obtusifolius var. latifolius Malta, Latvijas
Univ. Raksti 10: 334, 1924. TYPE: ‘‘Mexico:
Federal District, Cima 10,000 ft. (Pringle, Plantae
mexicanae N8 10517 und 10540) als Z. spathulaefolius Besch.’’ (lectotype, here designated,
‘‘Pringle N8 10540,’’ G!), syn. nov.
Zygodon asper Müll. Hal. ex Malta, Latvijas Univ. Bot.
Dārza Darbi 1: 166. 1926, nom. nud.
471
Zygodon rufulus Dusén in Malta, Latvijas Univ. Bot.
Dārza Darbi 1: 166. 1926, nom. nud. in syn.
Illustrations. Figs. 1E, 2C, 6–7.
Description. Plants slender, 0.5–1.5 cm tall, in
dense turfs, olive-green to brownish above, reddishbrown to dark brown below. Stems erect, branched.
Rhizoids yellowish-brown, smooth, well developed at
base and extending up the stems. Leaves erect and
keeled when dry, erect to erect-patent when moist,
0.5–0.9 3 0.15–0.3 mm, lingulate, base not decurrent,
apex rounded-obtuse; margins plane or slightly
recurved in the lower half, papillose-crenulate almost
throughout; upper leaf cells isodiametric, thickwalled, (8–)9–10(13) lm in diameter, bulging,
papillose, 2–3 papillae per cell; papillae low, blunt,
sometimes forked or multiple (several papillae forming a complex single papilla); basal cells isodiametric,
ca 10 lm in diameter, generally smooth, rarely
papillose, especially towards the margins; costa ending
below the apex, 30–48 lm wide in the lower 1/3, in a
deep furrow, with superficial cells elongate becoming
shorter towards the apex, dorsally papillose in the
upper 2/3, ventrally covered by papillose and
isodiametric laminal cells towards the apex, in section
cells 6 homogeneous. Propagula cylindrical, usually
hyaline, rarely yellowish-brown to greenish, 70–85 3
12–15(23) lm, (4–)6–8-celled, with only transverse
septa. Autoicous. Perichaetial leaves slightly differentiated; margins plane; basal cells rectangular, more
elongate than in stem leaves. Seta 5–15 mm long,
sinistrorse throughout; capsule erect, ovoid to ellipsoidal, 1.0–1.5 mm long, with 8 longitudinal ribs;
exothecial bands differentiated, of 4–5 cell rows each,
interrupted by a continuous ring of 3–5 smaller cells
surrounding capsule mouth; operculum conical at
base, shortly rostrate; peristome double; exostome
robust, of 8 pairs of teeth, reflexed or recurved and
appressed to the capsule when dry, dorsal surface
(OPL) reticulate, reticulum almost smooth to densely
papillose, sometimes completely covered in the lower
half with branched or simple papillae, coarsely
papillose in the upper half, ventral surface (PPL)
smooth to roughened-papillose in the lower 2/3,
coarsely papillose at apex; endostome of 8 hyaline,
dorsally keeled segments, occasionally with some extra
segments, dorsal surface (PPL) smooth to slightly
longitudinally striate throughout, ventral surface
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109(4): 2006
Figure 6. Zygodon obtusifolius. A. Habit when dry. B. Habit when moist. C. Leaves. D. Apex. E. Leaf basal cells. F. Leaf cross-section.
G. Propagula. H. Capsule when moist. I. Peristome (inner view). J. Spores. B–G from Dusén s.n. (S); A, H from Ule 665 (HBROTHERUS); I–J from Fendler 25 (G).
(IPL) papillose, papillae in radial rows in basal cells,
aligned or not in the middle and perfectly longitudinally aligned or not towards the apex, basal
membrane low, continuous and striate. Spores finely
papillose or almost smooth, 12–15 lm in diameter,
trilete mark distinct.
Ecology. Zygodon obtusifolius usually grows as
epiphyte, more rarely on rocks, in dense or loose turfs.
It has been found on trunks and roots of old trees
(Araucaria, Quercus, Beilschmidia, Dracrydium and
Weinmania), logs and occasionally on twigs of shrubs.
Distribution (Fig. 23A). In America it ranges
from Chile to Mexico. Also reported from Africa, Asia
(Nepal, Sri Lanka) and Australasia (New Zealand and
Tasmania). In the study area it is known from Región
IX and Región X of Chile.
Differentiation. Zygodon obtusifolius is easy to
recognize by its leaves with bulging cells and roundedobtuse apex, unique features in the genus. The
recently described Z. conoideus var. lingulatus (Edwards et al. 2000) also has rounded-obtuse leaf apices,
but this is a dioicous Northern Hemisphere plant
Calabrese: Zygodon in southern South America
473
Figure 7. Zygodon obtusifolius. SEM photographs of peristome. A. Peristome. B, C. Exostome. B. OPL, middle portion of teeth. C.
PPL, basal and middle portion of teeth. D–F. Endostome. D. Outer surface of the segments, details of the PPL. E. IPL, basal. F. IPL,
apical. A–F from Ule 665 (H-BROTHERUS).
(only known from Surrey, U.K.) with flat laminal cells
bearing simple papillae and marginal cells wider in the
apex than in the medial part of leaves.
Variability. Two varieties of Z. obtusifolius have
been described: var. antarcticus Malta, characterized
by small plants, leaves and capsules, and leaf cells with
strongly thickened walls (Malta, 1924: 334); and var.
latifolius Malta, established to accommodate expressions with wider leaves (Malta, 1924: 334). However,
all these features are found in typical Z. obtusifolius,
which always grows in low turfs, has hyaline to
greenish or yellowish-brown leaves (under light
microscopy), with cells almost isodiametric, bulging
throughout, strongly thick-walled, usually smooth at
leaf base but papillose above. Regarding capsule size, a
specimen reported by Malta as var. antarcticus was
examined, and its capsules are not significantly
smaller than those in typical Z. obtusifolius. It became
clear that the characters indicated by Malta to
segregate the two varieties fall within the variation
range of the typical Z. obtusifolius, and therefore they
are considered synonyms of it.
474
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Zygodon obtusifolius shows variation in some
sporophytic features. It usually has eight endostomial
segments, although there are occasional extra segments. Malta (1926: 164) reported eight segments,
and more rarely 16. A fragmentary prostome, as
reported by Lewinsky (1990: 123), was not seen.
Peristomial ornamentation is also variable. The OPL
is reticulate, and the reticulum walls are either smooth
or papillose. Exostomial PPL varies from strongly
papillose in some specimens to almost smooth in
others. Endostomial PPL has weak longitudinal striae,
although Lewinsky (1990: 123) reported a smooth PPL.
Selected specimens examined (52). Chile. REGIÓN
IX: Chile australis in monte Cordillera de la Costa
supra Angol, Dusén 845 (S).
AUSTRALIA. TASMANIA: Sophia Point, Macquarie
Harbour, Moore 26 (H-BR). BRAZIL. MINAS GERAIS: Serra
do Itatiaia, Ule 130 (G 027205); SANTA CATARINA: Serra
Geral, Ule 22 (G 027204). COLOMBIA. CUNDINAMARCA:
San Cayetano, Hda. Portugal, Cleef 6062 (G 313600).
MEXICO. DISTRITO FEDERAL: Mexique, macizo [?illeg.],
Bescherelle s.n. (G 027238); JALISCO: Slopes of La
Ferrerı́a, above Manantlán, Crum 1053 (G 027239).
NEPAL [locality not stated]. Gardner s.n. (BM 79684a, G
027202). NEW ZEALAND. NORTH ISLAND: North Auckland
District, Leigh Marine Research Station, Fife 7901
(CHR 106607); SOUTH ISLAND: Nelson, Grant NZ 421
(CHR 303269). VENEZUELA. Prope Coloniam Tovar,
Fendler 25 (G 027207, G 027213).
SUBG. ZYGODON
Diagnosis. Plants to (0.2–)0.5–6(15) cm tall,
usually in turfs, on bark or rock. Stems erect or
prostrate, simple or branched. Leaves lanceolate,
rhomboidal-lanceolate or rarely lingulate; apex usually
acute; upper laminal cells 6 isodiametric, roundedhexagonal to ovate, not bulging, always papillose; basal
leaf cells short to long-rectangular, smooth and
differentiated from upper cells, isomorphic or dimorphic. Propagula often present, cylindrical, fusiform,
ovoid or clavate, with only transverse septa or with
tranverse and longitudinal septa. Dioicous or synoicous, occasionally autoicous. Seta distally dextrorse or
sinistrorse, in the basal part sinistrorse, without
torsion or exceptionally dextrorse. Peristome double,
simple or absent. Spores 10–30 lm in diameter,
papillose or foveolate, trilete mark absent or distinct.
Sect. Stenomitrium Mitt., J. Linn. Soc. Bot. 12: 238. 1869.
Diagnosis. Plants in turfs or loose mats. Stems
erect or prostrate. Leaves distinctly 5-ranked, lanceolate to lanceolate-rhomboidal; apex acute to acuminate; basal cells in dimorphic bands of hyaline, thinwalled cells with wide lumina, and yellowish-brown,
strongly thick-walled cells, usually nodulose, with
narrower lumina. Dioicous. Setae distally sinistrorse,
in the basal part without torsion, sinistrorse or
dextrorse. Peristome double.
Endemic to South America, three species grow in
the study area: Zygodon pentastichus, Z. bartramioides
and Z. chilensis.
3. Zygodon pentastichus (Mont.) Müll. Hal., Syn.
Musc. Frond. 1: 675. 1849; Aulacomnium pentastichum Mont., Ann. Sci. Nat. Bot., III, 4: 103.
1845; Stenomitrium pentastichum (Mont.) Broth.,
Nat. Pflanzenfam. 1(3): 464. 1902 [1903];
Pentastichella pentasticha (Mont.) Müll. Hal.,
Oesterr. Bot. Z. 47: 421. 1897. TYPE: ‘‘Chile
australis, ubi primus legit Pöppig, dein C. Gay’’
(PC?, not found).
Macromitrium poeppiggi Duby, Mém. Soc. Phys.
Genève 19: 295, tab. 2, fig. 2. 1868. TYPE: ‘‘In
Andibus Antuccensibus Chili repertam a cl.
Pöppig, in herbario olim Nees von Esenbeck
nunc meo inveni’’. Synonymized by Brotherus
(1902–1903: 494).
Zygodon uncinatus Mitt., J. Linn. Soc. Bot. 12: 234.
1869; Codonoblepharon uncinatum (Mitt.) A.
Jaeger, Ber. Thätigk. St. Gallischen Naturwiss.
Ges. 1872–73: 118 (Gen. Sp. Musc. 1: 396). 1874.
TYPE: [Chile] ‘‘Ins. Chiloé, Lobb’’ (lectotype,
selected here, NY!). Synonymized by Malta (1926:
179).
Pentastichella aurea Dusén, Rep. Princeton Univ.
Exped. Patagonia, Bot. 8(3): 122. 1903, nom. nud.
Pentastichella jaffuelii Thér., Revista Chilena Hist. Nat.
25: 301, tab. 27, fig. 2 a–h. 1921, non Zygodon
jaffuelii Thér.; Zygodon porteri Thér. nom. nov.,
Revista Chilena Hist. Nat. 30: 351. 1926. TYPE:
‘‘Crête des Monts de Las Piedras (Marga Marga),
sur l’écorce des vieux quillayes (Jaffuel, févr.
1917), stérile’’ (isotype, FH!), syn. nov.
Pentastichella pentasticha var. strictifolia Thér., Revista
Chilena Hist. Nat. 28: 131. 1924; Zygodon
pentastichus var. strictifolius (Thér.) Malta, Lat-
Calabrese: Zygodon in southern South America
vijas Univ. Bot. Dārza Darbi 1: 141. 1926. TYPE:
‘‘Temuco (F. Jaffuel)’’ (PC?, not found), syn. nov.
Zygodon porteri Thér. var. crispatulus Thér., Revista
Chilena Hist. Nat. 30: 351. 1926; Zygodon
pentastichus var. crispatulus (Thér.) Malta, Latvijas Univ. Bot. Dārza Darbi 1: 140, fig. 87f. 1926.
TYPE: ‘‘I. Chiloé, Ancud (leg. F. Jaffuel, 1924)’’
(isotype, FH!), syn. nov.
Illustrations. Figs. 2E–F, 8–9.
Description. Plants robust, in lax mats, green
above, brown to dark reddish-brown in the prostrate
stems and lower parts of branches. Stems prostrateascending, profusely covered by a reddish-brown
tomentum, regularly branched, branches erect, to 4
cm high. Rhizoids reddish-brown, smooth. Leaves
clearly in 5 ranks, erect, appressed and keeled when
dry, occasionally flexuose and slightly crisped above,
spreading to recurved from an appressed base when
moist, 2.1–3.0 3 0.6–0.9 mm, lanceolate, apex longacuminate; margins plane, entire and smooth at base,
entire and papillose above; upper and middle leaf cells
isodiametric, hexagonal to rounded, strongly thickwalled, with lumina slightly oval (6–)8–10 lm in
diameter, papillose, strongly differentiated from basal
cells; papillae (1–)3–4 per cell, simple, low and blunt;
basal cells long and narrow, 56–82(90) 3 6–8 lm,
smooth, light bands of thin to slightly thick-walled,
hyaline, larger cells alternating with yellow bands of
strongly thick-walled, porose, yellowish-brown, narrower cells; costa ending below apex, 30–42 lm wide
in the lower 1/3, dorsal superficial cells smooth, in
section with 2 ventral guide cells and dorsal stereids.
Propagula absent. Dioicous. Perichaetial leaves differentiated, erect when moist, 2.3–2.5 3 0.4–0.7 mm,
lanceolate, apex acute to acuminate; margins entire;
cells elongate, smooth or with few papillae, basal cells
not differentiated as in stem leaves. Seta 10–15 mm
long, sinistrorse in the distal part, sinistrorse or
without torsion at base; capsule erect, oblong to
cylindrical, 2.0–2.5 mm long, with 8 longitudinal ribs
throughout; exothecial bands differentiated, of 4–5
cell rows each, interrupted by 3–4 smaller cell-rows
surrounding capsule mouth; operculum conic at base,
with a long beak; peristome double; exostome robust,
of 16 teeth, partially paired (al least joined at base),
typically recurved, dorsal surface (OPL) with coarse,
blunt papillae, densely and uniformly distributed,
covering a basal reticulum hardly visible, ventral
475
surface with papillae in thin vertical lines or irregularly and densely covering the lower 2/3, with coarse
papillae, sometimes joined in the upper 1/3; endostome of 16 very wide segments, shorter than
exostome, fused at base, strongly keeled, dorsally
(PPL) smooth or with short, blunt, coarse, disperse
papillae, ventrally (IPL) strongly papillose, basal
membrane high and continuous. Spores papillose,
(17–)20–28 lm in diameter, trilete mark indistinct.
Ecology. Usually epiphytic, in lax mats on tree
trunks, old stumps and logs, more rarely on rotten
wood; often on old Nothofagus, less common on
Austrocedrus, Araucaria, Aristotelia, Peumus, Eucryphia, Drymis, Gevuina and Pernettya. It has been also
found on rocks, bricks and rarely on soil around
trunk bases.
Distribution (Fig. 23B). It ranges from Peru to
southern Argentina and Chile through the Andes. In
Argentina is known from Neuquén, Rı́o Negro and
Chubut provinces, and from regions VII, VIII, IX, X
and XI in Chile; also reported from the Juan
Fernández Islands.
Differentiation. Zygodon pentastichus shares
some important features with Z. bartramioides and Z.
chilensis, like the ovate upper leaf cells, basal cells
forming distinct bands and the sinistrorse setae to the
apex, suggesting that they are closely related. However, it has leaves loosely clasping the stem at base and
more clearly arranged in five rows, prostrate stems
with erect branches and lacks propagula. Sporophytically, it can be distinguished by operculum shape
(longer than in Z. bartramioides and Z. chilensis), and
by the 16 exostome teeth arched and touching the
capsule wall only at tips, but not appressed to the
capsule wall as in Z. bartramioides and Z. chilensis.
Finally, the spores of Z. pentastichus are larger,
without trilete mark, and papillose (foveolate in Z.
bartramioides and Z. chilensis).
Variability. Zygodon pentastichus is very variable
in branch length, some of which are robust and to 4
cm, while others only extend 1 cm above the
substrate. Leaves vary from very rigid to strongly
flexuose, a condition usually related to water availability; plants from more humid places have flexuose,
longer leaves narrowed in a long-acuminate apex
occasionally crispate when dry, and then not clearly
pentastichous. Although the 5-row disposition of the
leaves can be sometimes difficult to notice when dry,
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Figure 8. Zygodon pentastichus. A. Habit when dry. B. Habit when moist. C. Leaves. D. Apex. E. Leaf basal cells showing
differentiated cell-bands. F. Leaf cross-sections. G. Perichaetial leaf. H. Peristome (inner view). I. Spores. A, B from Calabrese 336
(BCRU); cI from Crosby 12292 (MO); cII from Calabrese 200154 (MA); D–F from Calabrese 4612 (BCRU); G from Calabrese 200159 (MA);
H, I from Thaxter 44 (FH).
Calabrese: Zygodon in southern South America
477
Figure 9. Zygodon pentastichus. SEM photographs of peristome A. Peristome. B–E. Exostome. B. Details of the basal OPL. C.. OPL,
basal and middle portion of teeth. D. PPL, basal. E. PPL, apical. F. Outer surface of endostomial segments (PPL). A, D–F from
Calabrese 169 (BCRU); B, C from Santensson 2210 (FH-BARTRAM).
it is always very distinct when moist. In contrast,
plants from drier places have stiff, erect and appressed
leaves in five regular rows when dry.
The Chilean specimens from Ancud collected by
Jaffuel in 1924 (type of Z. pentastichus var. crispatulus)
and from Guaitecas (distributed by Dusén as Pentastichella aurea) have strongly crispate and flexuose
leaves when dry, but either gametophytically and
sporophytically they are identical to specimens of the
typical variety. Although both are identical, the
Guaitecas material was considered by Malta (1926:
140) to represent a transition towards var. crispatulus.
A similar habit is shown by a collection from MargaMarga (Chile), collected by Jaffuel in 1917 and
distributed as Pentastichella jaffuelii Thér.
The type material of var. strictifolius, collected by
Jaffuel in Temuco, was not available for study. This
variety was described on the basis of its erectappressed leaves and shorter setae and capsules.
However, dry leaves fall into the leaf variability
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previously discussed, as do seta length and capsule
size, which further do not vary significantly in this
species. In this treatment the two described varieties
of Z. pentastichus are included in the range of
variability of the typical expression.
Selected specimens examined (298). ARGENTINA.
CHUBUT: Lago Menéndez, 1-XII-1940, Kühneman s. n.
(BA 5254); NEUQUÉN: Dpto. Los Lagos, picada a
Cascada Los Cántaros, Calabrese 169 (BCRU 1810); RIO
NEGRO: Dpto. Bariloche, Cerro Llao Llao, Calabrese
475 (BCRU 1811). CHILE. REGIÓN V. VALPARAÍSO: Crete des
monts de Las Piedras (Marga Marga), II-1917, Jaffuel
s.n. (FH-BARTRAM 12263), JUAN FERNÁNDEZ: Masafuera,
en cerro Inocentes, Looser 6310 (MO 3987655); REGIÓN
VII. LINARES: Bullileo ca. 50 km east of Parral in the precordillera, near Laguna Amargo, Landrum 1523a (MO
2860320), MAULE: ‘‘Bosque Fisco’’ between Quirihue
and Cobquecura, Landrum 1539 (MO 2856136), TALCA:
Empedrado, Landrum 6341 (MO 3987654); REGIÓN VIII.
Andes Antucenses, Pöppig 33 (G 027208), ARAUCO:
ridge south of Cerro Lanalhue, southwest of Lago
Lanalhue, Fundo Tranquivora, Crosby 13051 (MO
2521905), CONCEPCIÓN: Concepción, IV-1925, Bertho
s.n. (FH); REGIÓN IX. CAUTÍN: Parque Nacional Villarrica,
near Refugio Villarrica, 6.1 km by road south of park
entrance, Crosby 11802 (MO 2521913), Dpto. Villarrica, Pucón, Hosseus 374 (JE-HERZOG), MALLECO: Reserva
Forestal Malleco, Landrum 6340 (MO 3987638),
MALLECO/ARAUCO: Araucaria forest in Parque Nacional
Nahuelbuta, 10 km west of park entrance and 48 km
by road west of Angol, Cordillera Nahuelbuta, Crosby
13069 (MO 2521895); REGIÓN X. CHILOÉ: Isla Chiloé, 5
km by road west of Ancud, Crosby 12335 (MO
2521932), ISLA GUAITECAS: Patagonia occ. in insulis
Guaitecas, Dusén 670 (FH, JE), LLANQUIHUE: Laguna
Verde, Mahu & Tapia 24343 (MO 3987720), OSORNO:
Yerbas Buenas, centro de recreación Las Cascadas,
Mahu 21359 (MO 3987726), VALDIVIA: Valdivia, Hofmann 21 (HBG); REGIÓN XI. AISÉN: Puerto Chacabuco 6IV-1967, Seki s.n. (HIRO 1894-a).
PERU. YUNGAY: Dpto. Ancash, an der Laguna
Llanganuco, 17-X-1973, Hegewald 7576 (MO 3670879).
4. Zygodon bartramioides Dusén ex Malta, Latvijas
Univ. Raksti 10: 320.1924; Pentastichella bartramioides Dusén, nom. nud. in sched.; Zygodon
cylindropyxis Müll. Hal. ex Malta, Latvijas Univ.
Raksti 10: 320. 1924, nom. inval. pro synon.;
Zygodon robustulus Broth. ex Malta, Latvijas
Univ. Raksti 10: 320. 1924, nom. inval. pro synon.
TYPE: ‘‘Chile: o. n. O. [locality not indicated]
(Philippi, Musci frondosi N8 13); o. n. O.
(Pöppig); Angol (Dusén) als Z. robustulus Broth.;
Anden von Villarica (Neger)’’. TYPE: Chile
australis in monte Cordillera de la Costa, supra
Angol oppidam in truncis Araucariae imbricatae,
5-XI-1896, Dusén 362 (lectotype, here designated,
FH-BARTRAM!, isolectotypes, FH-BARTRAM!, S-B6278!,
S-B6279!, S-B6280!; syntype, ‘‘Neger’’, JE-HERZOG!).
Pentastichella robustula Dusén ex Paris, Index Bryol. 4:
320. 1905, nom. nud.
Stenomitrium robustulum Dusén ex Paris, Index Bryol.
4: 320. 1905, nom. nud.
Zygodon robustus Broth. in Malta, Latvijas Univ. Bot.
Dārza Darbi 1: 79. 1926, nom. nud. in syn.
Zygodon tristis Herzog, Rev. Bryol. Lichenól. 29: 196.
1960. TYPE: ‘‘Südchile: Hacienda Freire, an
Eucalyptus, G. H. Schwabe, n8 7’’ (holotype, JEHERZOG!), syn. nov.
Illustrations. Figs. 2D, 3F, 10–11.
Description. Plants robust, 2(4) cm tall, in
dense turfs, green above, brown below. Stems erect,
branched. Rhizoids reddish-brown, usually roughened, extending up the stems and partially covering
them in lower 1/3. Leaves appressed and strongly
keeled when dry, erect when moist, (0.7–)0.9–1.7 3
0.2–0.4 mm at base and (0.3–)0.4–0.8 mm wide at
shoulders, lanceolate-rhomboidal, decurrent, apex
acute; margins plane, entire at base, slightly papillose
above, sometimes lightly denticulate towards apex,
upper leaf cells rounded-hexagonal, isodiametric,
thick-walled, with lumina slightly oval, (6–)8–
12(14) lm in diameter, papillose; papillae (1–)2–4
per cell, usually simple, some branched, to 4 lm high,
blunt; basal cells elongate rectangular to shortly
rectangular, 20–74(90) 3 (3–)6–13(18) lm,
smooth, occasionally with only a blunt papilla, in light
bands of cells with hyaline thin walls alternating with
yellow bands of cells with thickened and colored walls,
sometimes nodulose (especially in older leaves),
suddenly and clearly differentiated from upper cells;
marginal cells shortly rectangular to sub-quadrate;
costa ending below the apex, (25–)29–43(48) lm
wide in lower 1/3, in a deep furrow from base to
Calabrese: Zygodon in southern South America
479
Figure 10. Zygodon bartramioides. A, C. Habit when dry. B. Habit when moist. D. Wet capsule. E. Leaves. F. Perichaetial leaf. G.
Apex. H. Leaf basal cells. I. Leaf cross-sections. J. Propagula. K. Peristome (inner view). L. Spores. A from Schwabe 81 (JE); B, G, iII, jIII
from Schwabe 7 (JE); C, D, K, L from Neger 45 (S); eII, H, iI, jI from Dusén 362 (S); eI, F from Schwabe 61 (JE); iIII from Hosseus 393 (JE).
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Figure 11. Zygodon bartramioides. SEM photographs of peristome. A. Capsule mouth and peristome in lateral view. B–E. Exostome:
B. Details of the basal OPL. C. OPL, apical. D. PPL, middle portion of teeth. E. PPL, apical. F. Outer surface (PPL) of the
endostomial segments. A–F from Dusén 362 (FH).
upper 4/5, flattened with the lamina above, dorsal side
papillose, in section with 2 ventral guide cells and
dorsal stereids. Propagula in brownish-yellow axillary
stalks, cylindrical or clavate, hyaline, 40–67 3 15–23
lm, (3–)4(5)-celled, with only transverse septa.
Dioicous. Perichaetial leaves differentiated, erect when
moist, 1.4–1.7 3 0.28–0.32 mm, narrowly lanceolate,
apex acute; margins entire, partially involute; cells
elongate, smooth or with scattered papillae, undifferentiated at base. Seta 2–5 mm long, sinistrorse in
the distal part, sinistrorse or without torsion at base;
capsule erect, ovoid to ellipsoidal, ca 1.2 mm long,
with 8 longitudinal ribs except at base which is slightly
sulcate to almost smooth; exothecial bands differentiated, of 4 cell rows each, interrupted by a
continuous ring of 2–4 smaller cells surrounding
capsule mouth; operculum conical at base, shortly
rostrate; peristome double; exostome robust, of 8
pairs of teeth, teeth lacunose, partially splitting along
the middle line, recurved and appressed to the capsule
when dry, dorsal surface (OPL) with very coarse,
blunt papillae disposed on the edge of lamellae that
Calabrese: Zygodon in southern South America
form a reticulum in the lower 1/3–2/3, simple papillae
or sometimes coarse lines vertically oriented formed
by the fusion of papillae in the upper 1/3 of the tooth,
ventral surface (PPL) roughened at base, densely
papillose in the upper 2/3; endostome of 8 segments,
dorsal surface (PPL) smooth, ventral surface (IPL)
striate and with papillae vertically and radially
disposed in the lower 2/3, distally with coarse papillae,
basal membrane low, continuous and striate. Spores
foveolate (with numerous depressions or cavities),
10–12 lm in diameter, trilete mark distinct.
Ecology. A typically epiphytic species. Although
data on phorophytes are scarce, it has been found on
Araucaria sp., Nothofagus obliqua (Mirb.) Oerst. and
Eucalyptus sp.
Distribution (Fig. 24A). Zygodon bartramioides is
a Chilean endemic known from Región IX and Región
X (Malleco, Cautı́n, Valdivia and Osorno provinces).
Differentiation. Zygodon bartramioides could be
confused with the closely related Z. chilensis, especially
on the basis of peristome characters. However, there is a
set of features, particularly gametophytic, that separates
both taxa: propagule size, color and morphology, leaf
shape and basal areolation, and seta length. The
propagula are the best distinguishing character,
especially because they are abundant in both species.
Zygodon bartramioides has hyaline, smaller propagula
with only transverse septa, while Z. chilensis has larger,
brownish propagula occasionally branched, often with
longitudinal as well as transverse septa. Leaves shape is
another differentiating character: Z. bartramioides has
lanceolate-rhomboidal leaves whereas they are lanceolate to narrowly lanceolate in Z. chilensis. Basal cells are
usually more shortly rectangular and nodulose in Z.
chilensis than in Z. bartramioides. Finally, setae are 2–5
mm long in Z. bartramioides, but longer, to 10 mm
long, in Z. chilensis.
Variability. The studied collections vary both in
shape and size of the papillae of the upper laminal
cells. Some cells have low and simple papillae, whereas
in other cells of the same leaf they are branched.
Moreover, papillae can be variably tall or short in
different leaves of the same specimen, and also vary in
number, from the common 2–3 to 4. The apices of
older leaves are usually eroded, as are the apical cells,
and thus it is not possible to distinguish papillae.
Zygodon tristis Herzog was described from a sole
481
collection from Hacienda Freire (Schwabe 7), growing
on cultivated Eucalyptus. This is a very poor specimen
that surely represents a young, under-developed
specimen. It has fewer rhizoids, less papillose leaves
with simple papillae (only a few branched), and a
costa slightly papillose dorsally. The examination of
the type of Z. tristis indicates it is a synonym of Z.
bartramioides.
Typification. Of the specimens reported by Malta
(1926: 117), the best collection is Dusén 362, which was
moreover widely distributed to several herbaria. The
specimens kept at S (where the Dusén herbarium is
housed) are sterile, while the material in FH (two
specimens) has sporophytes. The other collection
reported by Malta (Villarrica, Neger, JE!) is large and
fertile, but infected by fungi and there are not duplicates
in the consulted herbaria. Dusén 362 (FH 9678-HERB.
FLEISCHER!) is therefore designated as lectotype.
Selected specimens examined (13). CHILE. REGIÓN
IX. CAUTÍN: Dept. Villarrica: Pucón, Hosseus 393 (JEHERZOG), MALLECO: Chile australis in monte Cordillera
de la Costa, supra Angol oppidam in truncis
Araucariae imbricatae, Dusén 362 (FH, S-B62769, SB6278, S-B6280); REGIÓN X. OSORNO: Termas de
Puyehue, Schwabe 61 (JE-HERZOG), VALDIVIA: Valdivia,
Neger 45 (S).
5. Zygodon chilensis Calabrese & F. Lara, J. Bryol. 28:
97. 2006. TYPE: CHILE. VALDIVIA: Bima District,
Cordillera de Alerce, on bark of Fitzroya
patagonica (alerce), 650 m, 13-II-1958, Eyerdam
11000 (holotype, UC 271784!; isotypes, H!, NY!).
Illustrations. Figs. 1D, 12–13.
Description. Plants robust, 2(5) cm tall, in
dense turfs, green or yellowish-green above and
brown below. Stems erect, frequently branched.
Rhizoids reddish-brown, roughened to papillose,
abundant at base of stems. Leaves appressed when
dry, erect-patent when moist, (1.1–)1.4–1.9 3 0.2–
0.4(0.6) mm, lanceolate to narrowly lanceolate,
keeled, decurrent, with acute apex; margins plane,
entire and smooth at base, papillose above, usually
with 1 or few teeth at the apex; upper leaf cells
rounded-hexagonal, with lumina slightly oval, thickwalled, (9–)11–15 lm in diameter, papillose; papillae
(1–)2–4 per cell, simple, blunt to subacute, rarely
acute; basal cells elongate-rectangular or shortrectangular, (14–)21–40(54) 3 (6–)9–15 lm,
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Figure 12. Zygodon chilensis. A. Habit when dry . B. Habit when moist. C. Leaves. D. Perichaetial leaf. E. Leaf apex. F. Leaf basal cells.
G. Leaf cross-section . H. Propagula. I. Peristome and spores. A, B, D, I from Eyerdam 11000 (UC); C from Eyerdam 10999-a (UC); E
from Seki s.n. (HIRO 750-a); F from Eyerdam 1564-b (UC); G, hI from Eyerdam 11019-a (UC); hII from Seki s.n. (HIRO 545-c).
Calabrese: Zygodon in southern South America
483
Figure 13. Zygodon chilensis. SEM photographs of peristome. A. Lateral view of the capsule mouth and peristome. B–E. Exostome. B.
Details of the apical OPL. C. PPL, inner surface of the tooth and basal outer surface of the endostomial segments. D. PPL, middle
portion of teeth. E. PPL, apical. F, G. Endostome. F. IPL, basal. G. PPL and IPL, apical. H. Spores. A–H from Eyerdam 11000 (H).
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smooth, in light bands of cells with hyaline thin walls
alternating with yellow bands of cells with thickened,
nodulose and colored walls; marginal basal cells subquadrate or rectangular, 9–21 3 9–10 lm wide; costa
ending below the apex, 33–54 lm wide in lower 1/3,
in a deep furrow in the basal 4/5 and in the same
plane with the lamina above, dorsally papillose in the
upper 2/3, in section with 2 ventral guide-cells and
dorsal stereids. Propagula cylindrical or clavate,
usually brown, (75–)87–120 3 (21–)24–30 lm wide,
4–9-celled, sometimes branched and then to 16-celled,
with transverse septa, but frequently also with
longitudinal septa in one or both extremes, conspicuous due to size and color, as well as the usual
abundance, occasionally remaining attached on the
leaf surface. Dioicous. Perichaetial leaves differentiated, erect when moist, 1.90–2.10 3 0.28–0.32 mm,
narrowly lanceolate, apex acute-acuminate; margins
entire, 6 involute; cells elongate, smooth or with
scattered papillae, basal cells undifferentiated. Seta 7–
10 mm long, sinistrorse in the distal part, dextrorse at
base; capsule erect, cylindrical to ellipsoidal, 1.5–2.0
mm long, with 8 longitudinal ribs; exothecial bands
differentiated, of 4 cell rows each, interrupted by a
continuous ring of 3–4 almost isodiametric cells
around capsule mouth; operculum conical at base,
shortly rostrate; peristome double; exostome robust,
of 8 pairs of teeth, each tooth lacunose, partially
splitting along the middle line except at base, recurved
and appressed to the capsule when dry, dorsal surface
(OPL) with coarse papillae on the edge of lamellae
forming a reticulum in the lower tooth half, papillae
and lines (formed by the fusion of papillae) vertically
oriented in the upper tooth part, ventral surface (PPL)
usually almost smooth or roughened at base, sometimes with papillae longitudinally aligned or with
longitudinal smooth lines at the lower part, the upper
1/2–2/3 densely papillose; endostome of 8 hyaline
segments, occasionally with some (1–8) less developed
intercalary segments, dorsal surface (PPL) keeled,
completely smooth or slightly striate at base, ventral
surface (IPL) clearly striate with some scattered
papillae at base, that become gradually aligned in the
upper half, coarsely papillose towards apex, basal
membrane low, continuous and striate. Spores
foveolate, 10–12 lm in diameter, trilete mark distinct.
Ecology. Zygodon chilensis grows on Nothofagus
antarctica (G. Forst.) Oerst., Fitzroya and Podocarpus,
from sea level to 900 m.
Distribution (Fig. 23D). Zygodon chilensis, so far
endemic from Chile, is only known from two
population groups in Valdivia (Región X) and Aisén
(Región XI).
Differentiation. Zygodon chilensis is closely
related to Z. bartramioides. Both taxa have distally
sinistrorse setae, double peristomes, foveolate, small
spores with trilete markings, and basal leaf cells
differentiated in longitudinal bands. However, Z.
bartramioides has lanceolate-rhomboidal leaves with
distinct shoulders, basal leaf cells sharply differentiated from upper cells, blunt and sometimes
branched papillae, smaller propagula and shorter (to 5
mm) setae.
Variability. This species grows in two different
population groups in the study area, which differ in
leaf shape and areolation, and propagule size and
color (Calabrese et al. 2006). Southern specimens have
more narrowly lanceolate leaves with short-rectangular basal cells scarcely differentiated from upper
cells, and propagula usually larger, (75–)105–120 3
(21–)24–30 lm, with 4–9(16) cells, always with
colored walls and more often branched.
Selected specimens examined (7). CHILE. REGIÓN
X. VALDIVIA: Bima District, Eyerdam 10999a (UC
271783, paratype); REGIÓN XI. AISÉN: Estancia Rı́o
Romero, cerro junto al Ventisquero Nef, 19/20-III1967, Seki s.n. (HIRO 1564-b, paratype).
Sect. Zygodon
Zygodon sect. Euzygodon Müll. Hal., Syn. Musc.
Frond. 1: 671. 1849, nom. illeg.
Diagnosis. Plants in turfs. Stems erect. Leaves
distinctly 5-ranked or not, lanceolate or, exceptionally, lingulate; apex acute or rarely rounded-obtuse;
basal leaf cells isomorphic. Dioicous or synoicous,
rarely autoicous. Setae distally dextrorse, sinistrorse in
the lower part. Peristome double, simple or absent.
Widespread in both hemispheres, there are seven
species and one variety in the study area: Zygodon
intermedius, Z. pichinchensis, Z. reinwardtii var.
reinwardtii, Z. hookeri var. hookeri, Z. hookeri var.
leptobolax, Z. jaffuelii, Z. papillatus and Z. magellanicus, the last three endemic to southern South America.
6. Zygodon magellanicus Dusén ex Malta, Latvijas
Univ. Bot. Dārza Darbi 1: 173. 1926. TYPE: ‘‘Hab.
Calabrese: Zygodon in southern South America
485
Figure 14. Zygodon magellanicus. A. Habit when dry. B. Habit when moist. C. Leaves. D. Leaf apex. E. Cells in middle portion of leaf.
F. Leaf basal cells. G. Leaf cross-section. H. Propagula. A, B, cIII, E, hI from Dusén 83 (S); D, F, G, hII from Schiavone & Ochyra CM
6668 (BA); cI, hIII from Matteri & Schiavone CM 2449 (BA); cII from Matteri, Schiavone & Piñeiro CM 5757 (BA).
Chile australis’’. TYPE: ‘‘[Chile:] Patagonia, Punta
Arenas in saxis, P. Dusén N8 83, 27-XI-1895’’
(lectotype, selected by Calabrese et al. 2002, SB6273!; isolectotypes, H!, S-B6274!, S-B6275!).
Zygodon gracillimus Dusén, Rep. Princeton Univ.
Exped. Patagonia, Bot. 8(3): 122. 1903, p. p.,
nom. nud. (Some specimens are Z. magellanicus
while others are Z. papillatus.)
Illustrations. Figs. 1B, 2D, 14.
Description. Plants robust, (0.7–)1.6–2.5 cm tall,
in lax turfs, green above, brown below. Stems erect,
simple or scarcely branched. Rhizoids reddish-brown,
papillose, extending up the stem and partially covering its lower part. Leaves erect and keeled when dry,
patent when moist, (0.7–)0.96–1.75 3 (0.22–)0.32–
0.45 mm, lanceolate, narrower towards base, decurrent, with acute apex; margins plane, entire at base,
papillose above; upper leaf cells rounded-hexagonal,
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thick-walled, 8–13 lm in diameter, slightly smaller to
the margins, papillose; papillae 3–5(6) per cell
(usually 4), simple, blunt to subacute; basal cells
quadrate to short-rectangular towards the margins, 9–
12 lm, rectangular towards the costa, 12–45 3 6–15
lm, smooth; costa ending below the apex, 36–51 lm
wide in lower 1/3, in a deep furrow, superficial cells
elongate on both sides, dorsally smooth, ventrally
covered by papillose laminal cells towards the apex, in
section with 2 ventral guide cells and dorsal stereids.
Propagula in brownish-yellow axillary stalks, clavate,
reddish-brown, 50–90 3 30–39 lm, 5–12-celled, with
transverse and longitudinal septa. Dioicous. Sporophytes unknown.
Ecology. Zygodon magellanicus usually grows on
rock and rock crevices, and only once was found on
bark of Nothofagus pumilio (Poepp. & Endl.) Reiche.
Distribution (Fig. 23B). An Andean-Patagonian
and Fuegian endemic known from Neuquén and
Tierra del Fuego in Argentina and from Región XI
and Región XII of Chile.
Differentiation. This taxon is closely related to Z.
papillatus, which however has smaller (19–32 3 17–25
lm), mostly ovoid and yellowish-brown propagula,
branched papillae and slightly thickened laminal cell
walls, smooth or slightly roughened rhizoids, and
forms slender, more compact turfs. From the
remaining species in the area Z. magellanicus can be
distinguished by having brown propagula with
longitudinal as well as transverse walls.
Variability. Zygodon magellanicus scarcely varies,
with only the leaves ranging from small and broadly
lanceolate to long and narrowly lanceolate. The
variability of this taxon, as well as its differentiation
from Z. papillatus, has been already described in
length by Calabrese et al. (2002).
Selected specimens examined (14). ARGENTINA.
NEUQUÉN: Villa La Angostura, Inacayal, Schiavone &
Matteri CM 4725 (BA, BCRU); TIERRA DEL FUEGO:
Patagonia australis in territorio Rı́o Chico, Hatcher
s.n. (S-B6277); USHUAIA: Arroyo Grande (5 km. NE de
Ushuaia), Siete Cascadas, Matteri & Schiavone CM
2449 (S). CHILE. REGIÓN XI. AISEN: estancia Rı́o Romero,
cerro junto al Ventisquero Nef, 19/20-III-1967, Seki
s.n. (HIRO 1456-c); REGIÓN XII: Penı́nsula Brunswick,
Puerto Isidor, Roivainen 2405 (FH), Fuegia Media,
Lago Blanco, Roivainen 180 (FH-BARTRAM).
7. Zygodon papillatus Mont., Ann. Sci. Nat. Bot., III,
4: 106. 1845; Codonoblepharon papillatum
(Mont.) A. Jaeger, Ber. Thätigk. St. Gallischen
Naturwiss. Ges. 1872–73: 118 (Gen. Sp. Musc. 1:
396). 1874; Z. papillosus Mont. ex Broth. in Engler
& Prantl, Nat. Pflanzenfam. 1(3): 462. 1902
[1903], nom. inval., orthogr. err. TYPE: ‘‘Hab. ad
corticem arborum Chiles australioris invenit hunc
muscum cl. C. Gay. Herb. Mus. Par.’’ (lectotype,
selected by Calabrese et al. (2002), PC-MONTAGNE!).
Zygodon gracillimus Dusén, Rep. Princeton Univ.
Exped. Patagonia, Bot. 8(3): 122. 1903, p. p.,
nom. nud. (Some specimens are Z. magellanicus
while others are Z. papillatus.)
Illustrations. Figs. 1A, 3C, 15.
Description. Plants slender, (0.4–)2.0–4.5 cm tall,
in compact turfs, light green above, brown below.
Stems erect, simple or scarcely branched. Rhizoids
reddish-brown to yellowish, smooth to slightly
roughened, ascending up the stems. Leaves erect when
dry, erect-patent when moist, 0.66–0.85 3 0.14–0.30
mm, lanceolate, narrower towards the base, 6
decurrent, with acute apex; margins plane, entire and
smooth at base, papillose above; upper leaf cells
hexagonal, slightly thick-walled, 9–14 lm in diameter,
coarsely papillose; papillae (3–)4–6(9) per cell,
generally branched, blunt; basal cells short-rectangular, 9–32 3 6–13 lm, smooth; costa ending below the
apex, 28–30 lm wide in the lower 1/3, in a deep
furrow, superficial cells elongate on both surfaces,
dorsally smooth in the lower 2/3, papillose in the
upper 1/3, in section with 2 ventral guide-cells and
dorsal stereids. Propagula ovoid to clavate, yellowish
brown, 19–32 3 17–25 lm, with transverse and
longitudinal septa, although sometimes with only
transverse ones. Dioicous. Perichaetial leaves undifferentiated. Seta 4 mm long, dextrorse in the distal
part, sinistrorse at base; capsule erect, obovate, ovoid
to pyriform, 0.8–1.1 mm long, with 8 longitudinal
ribs; exothecial bands differentiated, of 4–5 cell rows
each; operculum conical at base, shortly rostrate;
peristome rudimentary, of 8 endostome hyaline
segments, dorsal and ventral (PPL and IPL) surfaces
smooth, basal membrane low, continuous and
smooth. Spores minutely papillose, 12–15 lm in
diameter, trilete mark distinct.
Ecology. This species has been only found on
bark. Reported phorophytes are Nothofagus betuloides
(Mirb.) Oerst., N. pumilio and Podocarpus sp.
Calabrese: Zygodon in southern South America
487
Figure 15. Zygodon papillatus. A. Habit when dry. B. Habit when moist. C. Leaves. D. Leaf apex. E. Cells in middle portion of leaf. F.
Leaf basal cells. G. Leaf cross-section. H. Propagula. I. Capsule when dry. J. Peristome and spores. A, B, cI, D–F, H–J from Gay s.n.
(PC); cII, G from Chile (PC); cIII from Dusén II-1897 (S).
Distribution (Fig. 23C). Zygodon papillatus is
known only from Rı́o Negro in Argentina and from
regions VIII, IX, X, XI and XII of Chile.
Differentiation. This taxon is closely related to Z.
magellanicus, which however has larger (50–90 3 30–
39 lm), clavate and reddish-brown propagula, leaf
cells with simple papillae (some branched papillae
may occur very rarely) and very thick walls, papillose
rhizoids, and forms robust, loose turfs. Possible
confusion between Z. papillatus and Z. hookeri var.
leptobolax, both with similar peristomes, is unlikely
considering sexual condition (dioicous in Z. papillatus
vs. synoicous in Z. hookeri var. leptobolax), propagula
(ovoid to clavate, with longitudinal and transverse
septa, yellowish-brown in Z. papillatus vs. fusiform,
only with transverse septa, usually hyaline or green-
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ish-hyaline in Z. hookeri var. leptobolax), and the
simple papillae and spores larger than 20 lm of Z.
hookeri var. leptobolax.
Variability. As for Z. magellanicus, the variability
and differentiation of Z. papillatus was treated in
detail by Calabrese et al. (2002). In any case, Z.
papillatus is morphologically very stable.
Selected specimens examined (30). ARGENTINA.
RÍO NEGRO. BARILOCHE: Reserva Municipal Llao Llao,
Cerro Llao Llao, Calabrese 461 (MA). CHILE. REGIÓN IX:
Chile australis, Cordillera de la Costa supra Angol, 5XI-1896, Dusén s.n. (S); REGIÓN X: Lago Pillaifa,
Schwabe 20 (JE); REGIÓN XI: in valle fluminis Aysén,
Dusén 565 (S, JE, FH-BARTRAM).
8. Zygodon pichinchensis (Taylor) Mitt., Hooker’s J.
Bot. Kew Gard. Misc. 3: 356. 1851; Didymodon
pichinchensis Taylor, London J. Bot. 7: 280. 1848.
TYPE: ‘‘Hab. Andes Quitenses, in montibus
Pichincha et Chimborazo, Jameson’’ (lectotype,
‘‘Pichincha, Jameson,’’ selected here, BM!; isolectotype, G!).
Zygodon setulosus Mitt., J. Linn. Soc. Bot. 12: 237.
1869. TYPE: ‘‘Andes Novo-Granatenses, Weir
334’’ (holotype, NY?, not found). Synonymized by
Churchill et al. (2000: 93).
Zygodon hystrix Dusén ex Paris, Index Bryol., ed. 2, 5:
139, 1906, nom. nud. ORIGINAL MATERIAL: NY!
Leptodontium spinosum R. S. Williams, The Bryologist
33: 73. 1930 [1931]. TYPE: ‘‘Dept. Santander,
Paramo de Santurban near Vetas, Colombia,
3950–4160 meters alt. Killip & Smith 17530, Jan.
17, 1927’’. Synonymized by Zander (1972: 278).
Triquetrella spiculosa Thér., Rev. Bryol. Lichénol. 9:
14. 1936. TYPE: ‘‘Ecuador, rochers du Condorguachana, Benoist 3150’’ (lectotype, PC, not seen).
Synonymized by Churchill et al. (2000: 182).
Illustrations. Figs. 1C, 16.
Description. Plants robust, (2–)5–7(11) cm, in
lax turfs, yellowish-green to brownish-green above,
reddish-brown and tomentose below. Stems erect,
simple or branched. Rhizoids reddish-brown, smooth
or slightly roughened, extending up the stem almost
to the apex, profuse in leaves axils. Leaves clearly in 5
ranks, erect and keeled when dry, squarrose-recurved
when moist, 1.10–1.78 3 0.37–0.67 mm, ovallanceolate, rapidly narrowed in the upper 1/3,
decurrent, apex acute, sometimes ended in a long
hyaline apiculus; margins plane, irregularly spinose by
large papillae; upper leaf cells rounded to subquadrate, isodiametric, thick-walled, 7–11 lm in diameter, papillose; papillae 1–2(3) per cell, simple or
branched, sometimes forked at some height from
base, acute to subacute, to 14 lm tall, occasionally
sharply pointed and hyaline, often fragile and once
broken appearing blunt, frequently longer and spinose
at margins and sometimes also at dorsal portion of
the costa, reaching 30 lm long; basal cells hardly
differentiated from upper cells, short-rectangular, 9–
19 3 5–7 lm, smooth or with 1–2 papillae towards the
margins, gradually longer, rectangular, 15–39 3 7–12
lm, smooth or with 1 small papilla towards the costa;
costa ending below the apex, red-brown, 36–60 lm
wide in lower 1/3, in a deep furrow, dorsally spinosepapillose, in section with two ventral cells and dorsal
stereids towards the base and cells 6 homogeneous
towards the apex. Propagula absent. Dioicous.
Perichaetial leaves differentiated, spreading when
moist, 1.78–2.33 3 0.43–0.45 mm, linear-lanceolate,
apex acute; margins entire towards base and denticulate towards apex; cells in the upper 1/3 with 1–2
small papillae; median cells smooth or papillose; basal
cells smooth, sometimes nodulose, elongate, 30–50 3
6–8 lm. Seta 7–12 mm long, dextrorse distally,
sinistrorse at base; capsule erect, cylindrical to
cylindric-ellipsoidal, ca 2 mm long, with 8 longitudinal ribs throughout; exothecial bands of 5–8 cell
rows each; operculum conical at base, subulate;
peristome simple, exostome absent; endostome of 8
hyaline segments, dorsally keeled in the middle line,
dorsal and ventral (PPL and IPL) surfaces smooth,
basal membrane low, continuous and smooth. Spores
finely papillose, 15–17 lm in diameter, trilete mark
absent.
Ecology. Zygodon pichinchensis usually grows on
crags, cliffs and rock crevices, and also on bark and
soil under shrubs. According to Churchill et al. (2000)
it ranges from 200 to 4800 m. Reported phorophytes
are Nothofagus dombeyi (Mirb.) Oerst. and Diplostephium rhomboidale Cuatrec. In the study area it
grows on wet places in Nothofagus forests.
Distribution (Fig. 24A). In South America, Z.
pichinchensis grows along the Cordillera de los Andes
from Colombia to Tierra del Fuego, avoiding the
more arid areas. It is also known from Central
Calabrese: Zygodon in southern South America
489
Figure 16. Zygodon pichinchensis. A. Habit when dry. B. Habit when moist. C. Leaves. D. Perichaetial leaf. E. Leaf apex. F. Cells in
middle leaf and margins. G. Leaf cross-section. H. Leaf marginal papillae. I. Papillae in dorsal portion of costa. J. Capsule when dry.
K. Peristome and spores. A, B, J from Matteri & Schiavone CM 5133 (BA); cI from Matteri & al. CM 6144 (BA); cII, hII from Jameson
s.n. (BM); cIII from Cleef 5663 (G); D, hIII, iI from Cleef s.n. (G 150899); E from Matteri & Schiavone CM 4778-a (BA); F, G, hI from
Schiavone & Matteri CM 4519 (BA); kI from Cleef 9819 (G); kII from Cleef 8898 (G); hIV from Cleef 8603 (G); hV, iII from Matteri &
Schiavone CM 5103 (BA).
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America and Mexico. In the study area it grows in
Chubut and Santa Cruz provinces of Argentina and
Chilean Región XI and Región XII.
Differentiation. The characteristic leaf cell papillae, as well as the lack of propagula, simple peristome
and dioicous sexual condition make confusion of Z.
pichinchensis with any other taxon unlikely.
Variability. Papillae vary in shape and size (Figs.
1C, 16H–I). In some specimens they are acute and
end in an easily broken hyaline tip, thus giving them a
blunt appearance, especially in the dorsal portion of
the costa; these papillae with hyaline tips are
frequently forked. In others specimens they are very
large and subacute, with the tip neither hyaline nor
fragile. Finally, in the same leaf it is possible to find
branched as well as simple papillae intermingled.
Papillae also vary in size, even in the same leaf,
although they are usually very large (more than 10 lm
long) and spinulose or spinose; only exceptionally
they are smaller and less distinct. Plant size is also
variable, often depending on substrate: plants growing
on soil are larger and more robust, to 11 cm high,
whereas those on rock or bark are the smallest
specimens, just 5–7 cm high. Finally, seta length is
also somewhat variable, from 7 to 12 mm, althought
there is no correlation with other character variation.
Zygodon setulosus was considered a synonym of
Z. pichinchensis by Churchill et al. (2000: 93),
although previously Churchill and Linares (1995: 621)
stated ‘‘according to Griffin III (pers. comm.) these
two species could be synonyms.’’ In neither of these
works nor in the present study could the type (‘‘Andes
Novo-Granatenses,’’ Weir 334) be found. Nevertheless, I studied the other specimen reported by Malta
(1926), who was able to compare it with the actual
type. This specimen (Apollinaire 371, PC!), as well as all
others previously identified as Z. setulosus, belongs to
Z. pichinchensis. According to the original description
(Mitten 1869), Z. setulosus has shorter setae, cylindrical capsules and narrower leaves, while Z. pichinchensis has elongate setae, oblong-elliptical capsules
and wider leaves. As I have already pointed out, seta
length is highly variable and cannot be considered as
diagnostic. Regarding capsule shape, they are always
6 cylindrical or cylindrical-ellipsoidal, and leaf width
varies enough in the same specimen (0.37–0.67 mm)
not to be considered of taxonomic importance. I
concur with Churchill et al. (2000) that Z. setulosus
and Z. pichinchensis are conspecific.
Selected specimens examined (96). ARGENTINA.
CHUBUT: Parque Nacional Los Alerces, arroyo Las
Cascadas, Matteri et al. CM 6144 (BA, BCRU); SANTA
CRUZ: southernmost end of Lago Argentino, Cantino
M-26 (BA, S, H). CHILE. REGIÓN XI. AISÉN: Estancia Lago
Plomo, Ventisquero Soler, 17/20-III-1967, Seki s.n.
(HIRO 1849); REGIÓN XII: Patagonia australis, in monte
Cordillera de los Andes ad Lago Dickson, 20-XII1896, Dusén s.n. (S).
BOLIVIA. LA PAZ. PROV. INQUISIVI: Cumbre
Sayaquira, headwaters of Rio Sayaquira, ca. 2.5 km S of
Estancia Huaña Hkota, Lewis 87–337 (H); PROV.
MURILLO: upper Rı́o Chuquiaguillo, ca. 15–19 km NE of
Plaza San Francisco, Lewis 88–430 (G 322940); PROV. SUR
YUNGAS: ‘‘Taquesi West’’, West side of Taquesi Pass
along Inca Trail to Abra, Lewis 83–2135A-b (MA).
COLOMBIA. ARAUCA: Cabeceras de la Quebrada El Playón,
Patio Bolos, 2 km al ENE del Alto Curisı́, Cleef 8898 (G
313543); BOYACÁ: Macizo alto N de Belén (Mnpio.
Tutasá), Cerro Pan d’Azucar y vecindades, lado SE del
Cerro, Cleef 9819 (G 313542). ECUADOR. PICHINCHA: Bell
383 (G 027219); QUITO: Volcán Cayambe, Balazs 81–09/
A (G 242149). VENEZUELA. MÉRIDA: Rangel, Cordillera de
los Andes, Sierra de Santo Domingo, páramo de
Mucubajı́, just above the Laguna de Los Patos, Griffin,
III et al. 017518 (MA 22496).
9. Zygodon intermedius Bruch & Schimp., Bryol.
Europ. 3: 41 (fasc. 4, monogr. 9). 1838. TYPE:
‘‘Exemplaria Hookeriana, ex insula Novae-Zelandiae missa, cel. Mougeot et Montagne nobiscum benevole communicavere’’ (lectotype, ‘‘New
Zealand, Dusky Sound, A. Menzies,’’ designated
by Lewinsky, 1990, BM!)
Zygodon conoideus (Dicks.) Hook. & Taylor var.
elongatus Hook. & Grev., Edinburgh J. Sci. 1: 132.
1824. TYPE: ‘‘New Zealand; first discovered there
by A. Menzies, Esq. Van Dieman’s Land, Dr.
Spence and Mr. R. Neill’’ (lectotype, ‘‘New
Zealand, leg. Menzies,’’ designated by Lewinsky,
1990, BM!). Synonymized by Lewinsky (1990: 131).
Zygodon brownii Schwägr., Sp. Musc. Frond., Suppl. 4:
317b. 1842. TYPE: ‘‘Adhaerentem ad Leskeam
sciuroidem Hook. in Terra van Diemen a cl.
Brownio lectum vidi’’ (holotype, G, not seen).
Synonymized by Malta (1926: 75).
Calabrese: Zygodon in southern South America
Zygodon angustifolius Hook. & Grev. ex Wilson,
London J. Bot. 5: 449. 1846, nom. nud.
Zygodon brachyodus Müll. Hal. & Hampe, Linnaea 28:
210. 1856. TYPE: ‘‘Australia, Victoria, Sealers
Cove, F. von Müller’’ (isotypes, BM, H, not seen).
Synonymized by Malta (1926: 75).
Zygodon affinis Dozy & Molk. ex Bosch. & Sande Lac.,
Bryol. Jav. 1: 107, t. 85. 1859. TYPE: ‘‘Habitat
insulam Javae; in summo monte Pangerango
Teysmann in herb. Dz. et Mb.’’. Synonymized by
Fleischer (1904: 394).
Zygodon krausei Lorentz, Moosstudien 162. 1864.
TYPE: CHILE. ‘‘Prope Valdiviam, Dr. Krause’’
(lectotype, selected here, S-B6276!; isolectotype,
NY!). Synonymized by Malta (1926: 75).
Zygodon confertus Müll. Hal., Hedwigia 37: 134. 1898.
TYPE: ‘‘Australia, Victoria, F. Campbell’’ (lectotype, selected by Lewinsky (1990), H, not seen).
Synonymized by Malta (1926: 75).
Zygodon gracilicaulis Müll. Hal., Hedwigia 37: 135.
1898. TYPE: ‘‘New Zealand, near Greymouth, R.
Helms’’ (lectotype, selected by Lewinsky (1990),
H, not seen). Synonymized by Malta (1926: 75).
Zygodon hymenodontioides Müll. Hal., Hedwigia 37: 135.
1898, nom. illeg., non Müll. Hal. 1869 [¼ Anoectangium hymenodontoides (Müll. Hal.) A. Jaeger].
TYPE: ‘‘Australia, Gippsland, Moe River, Luehmann’’ (lectotype, selected by Lewinsky (1990), H,
not seen). Synonymized by Malta (1926: 75).
Zygodon nanus Müll. Hal., Hedwigia 37: 135. 1898.
TYPE: ‘‘New Zealand, near Greymouth, R. Helms’’
(lectotype, selected by Lewinsky (1990), H, not
seen). Synonymized by Malta (1926: 75).
Zygodon curvicaulis Dusén, Rep. Princeton Univ.
Exped. Patagonia, Bot. 8(3): 84, fig. 13, pl. IX,
figs. 6–7. 1903. TYPE: ‘‘Patagonia australis’’
(lectotype, here designated, S-B6277!; isolectotype, NY!). Synonymized by Malta (1926: 75).
NOTE: Mixed with Z. magellanicus.
Zygodon pangerangensis M. Fleisch., Musci Fl. Buitenzorg 2: 395, fig. 74. 1904. TYPE: ‘‘M. Fleischer,
Musc. Archip. Ind., N8 280’’ (holotype, FH, not
seen). Synonymized by Malta (1926: 75).
Zygodon coraniensis Herzog, Biblioth. Bot. 87: 63, fig.
21g–i. 1916. TYPE: BOLIVIA. ‘‘In oberen Coranital
an Baumrinde, ca. 2600 m, N8 3413’’ (holotype,
JE, not seen). Synonymized by Malta (1926: 75).
491
Zygodon stresemannii Herzog, Hedwigia 57: 233. 1916.
TYPE: ‘‘ Mittel-Ceram: Epiphyt der Baumfarnregion am Gunung Pinaia, ca. 2530–2750 m, col.
Stresemann N8 311 u. 328’’ (syntypes, JE, not
seen). Synonymized by Malta (1926: 75).
Zygodon transvaaliensis Rehmann ex Sim, Trans. Roy.
Soc. S. Afr. 15: 271. 1926. TYPE: SOUTH AFRICA.
‘‘On trees, Lechlaba, Houtbosch, Transvaal,
Rehm. 500’’ (isotypes, PRE, TR-VENTURI!). Synonymized by Malta (1926: 75).
Zygodon pseudoventricosus Müll. Hal. in Malta,
Latvijas Univ. Bot. Dārza Darbi 1: 77. 1926, nom.
nud. in syn.
Illustrations. Figs. 17, 18A–B.
Description. Plants slender to slightly robust,
0.5–3(5) cm tall, in lax to dense turfs, green to
yellowish-green above, reddish-brown below. Stems
erect, frequently branched. Rhizoids reddish-brown,
roughened, abundant at stem base. Leaves erectflexuose when dry, spreading and flexuose when
moist, (0.63–)0.85–1.45(1.80) 3 0.20–0.32 mm,
lanceolate, narrower towards the base, base not or
slightly decurrent, apex acute; margins plane, entire
and smooth at base, papillose above; upper leaf cells
rounded-hexagonal, isodiametric, strongly thickwalled, 4–10 lm in diameter, papillose; papillae 3–7
per cell, simple, blunt to subacute; basal cells
subquadrate to short-rectangular, thick-walled, 9–40
3 5–15 lm, smooth, occasionally some cells with 1–2
papillae; costa ending below the apex, ca 30 lm wide
in the lower 1/3, in a furrow, dorsally smooth, in
section with 2 ventral guide cells and dorsal stereids.
Propagula rare, fusiform to fusiform-clavate, hyaline,
67–80 3 17–19 lm, 3-celled, with only transverse
septa. Dioicous. Perichaetial leaves undifferentiated.
Seta 3–10 mm long, dextrorse in the distal part,
sinistrorse at base; capsule erect, shortly cylindrical,
pyriform or urceolate, 0.9–1.5 mm long, with 8
longitudinal ribs; exothecial bands differentiated, of 4
cell rows each; operculum conical at base, subulate,
beak curved; peristome double; exostome rudimentary, of few cells, dorsal surface (OPL) almost smooth,
ventral surface (PPL) completely smooth; endostome
of 8 hyaline segments, dorsal (PPL) and ventral (IPL)
surfaces smooth, basal membrane low, continuous
and smooth. Spores papillose, (10–)13–18 lm in
diameter, trilete mark weak.
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Figure 17. Zygodon intermedius. A. Habit when dry. B. Habit when moist. C. Leaves. D. Apices. E. Cells in middle portion of leaf. F.
Leaf basal cells. G. Leaf cross-section. H. Propagula. I. Capsule when dry. J. Peristome and spores. A, cIV, dI, F from Menzies 66 (BM
555688b); cI, dII from TIMB (BM 555688f); cII from Menzies s.n. (BM 555682c); cIII, E from Dusén 413 (S); B, J from C. & I. Skottsberg
M-33 (S); G from Skottsberg 438 (S); H from Streimann HS3047 (H); I from Lara & San Miguel s.n. (HERB. LARA, Universidad
Autónoma de Madrid).
Ecology. Usually growing as an epiphyte on small
branches of shrubs and trees, often on the underside
of the twigs, occasionally at the base of the trunk of
young trees; also rather frequent on rock. In the study
area it has been found on Berberis, mainly on Berberis
ilicifolia Forst. However, in New Zealand and
Australia it is one of the species of Zygodon growing
on a more diverse set of phorophytes, in wet sites as
well as in more open and dry areas.
Distribution (Fig. 24C). Zygodon intermedius is a
widespread species. In the Patagonian and Fuegian
regions, where it is known from Argentina in Rı́o Negro,
Santa Cruz and Tierra del Fuego provinces and from Chile
in regions V (Juan Fernández), VII, IX, X, XI and XII.
Differentiation. Zygodon intermedius has been
very often confused with Z. hookeri, likely because of
their similar habits, the uncertainty of their respective
sexual conditions (resolved by Lewinsky 1990: 136),
and because the peristome of Z. intermedius has been
considered very variable—simple, double or absent
(Malta 1926: 76). According to Scott and Stone (1976:
246), Zygodon intermedius can be distinguished from
Z. hookeri because the latter has wavy leaves with a
denticulate apex, but these characters are variable in
Z. hookeri (cf. discussion under that species), and not
useful to separate them. However, other features
distinguish the two taxa with confidence: Z. intermedius is dioicous, has perichaetial leaves not or very
Calabrese: Zygodon in southern South America
493
Figure 18. SEM photographs of peristomes. A, B. Zygodon intermedius: A. Peristome and capsule mouth. B. Details of a segment and
basal inner surface of the teeth. C–H. Z. jaffuelii. C. Peristome. D–G. Exostome. D. OPL, middle portion of teeth. E. OPL, apical. F.
PPL, apical and in middle portion of teeth. G. PPL, basal. H. Endostome, outer surface of segment (PPL). A, B from Lara & San
Miguel (HERB. LARA, Universidad Autónoma de Madrid); C–H. from Costes s.n. (FH).
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slightly differentiated, a rudimentary exostome,
smaller capsules and spores less than 20 lm in
diameter; whereas Z. hookeri is synoicous (rarely
autoicous), has perichaetial leaves well differentiated,
no exostome, larger capsules and spores typically 20–
26 lm in diameter.
Variability. Three varieties of Zygodon intermedius have been described, one from Java and two from
Africa, none of which has been reported or found in
the Andean-Patagonian and Fuegian regions. The
African var. strictifolius Thér. was described as having
leaves shorter and wider, very slightly (imperceptibly)
acuminate, appressed and not crisped when dry
(Thériot 1930: 38), and has been later reported from
Rwanda and Zaire (Born et al. 1993) and Uganda
(Potier de la Varde 1955). Malta distinguished his new
var. dusenii, from Cameroon, by its leaves relatively
short and wide, erect to patent when moist, and
laminal cells larger, to 13 lm in diameter [not
including the walls] (Malta 1926: 77). Finally,
Fleischer (1904: 395) described the Javanese var.
viridi-gracilior as having leaves shorter and wider than
the type, leaf cells with thinner walls and shorter setae.
The degree of variation of the characters used to
separate these three varieties falls into the range for
typical Z. intermedius, and no taxonomic conclusion
can be reached until more material and, more
importantly, the types, are studied.
No specimen differs significantly from the type.
Nevertheless, a specimen from the Australian Capital
Territory (Streimann 11-VII-1979, BA 32442!) has
abundant propagula, upper leaf cells with thinner walls
than is usual for the species (consequently, the lumina
are wider), with 6–7 papillae each and basal cells often
with 1–2 papillae. Despite these minor differences, all
other characters point to Z. intermedius.
The taxon is somewhat variable in plant size, with
larger plants having larger leaves (to 1.8 mm long,
although they can be only 0.6 mm long, cf. Fig. 17C),
and capsule shape can vary from short cylindrical to
pyriform or urceolate. I think that the observed
variability is driven by environmental conditions, as
this taxon grows in wet as well as in rather dry sites.
Selected specimens examined (209). ARGENTINA.
RÍO NEGRO: Laguna Larga, Isla Victoria, 31-III-1946, De
Barba s.n. (FH-BARTRAM); SANTA CRUZ: Lago San Martı́n,
penı́nsula Cancha Rayada, Skottsberg 438 (S); TIERRA
DEL FUEGO:
Rı́o Chico, Hatcher s.n. (S-B6277). CHILE.
Islas Juan Fernández, Masatierra,
Bahı́a Cumberland, El Paugal, Skottsberg M53 (S);
REGIÓN VII. LINARES: East of Linares along the Rı́o
Atchibuena, along the road through Pejerrey 39.7 km
upstream from the intersection with the road to
Medina and Melado, Taylor & Gereau 11042 (MO
4423010); REGIÓN IX. MALLECO: Reserva Forestal de
Malleco, ca. 70 km east of Collipulli which in turn is
34 km southeast of Angol, Landrum 409 (MO
2556517); REGIÓN X. VALDIVIA: Valdivia, Herzog 5238/a
(JE-HERZOG); REGIÓN XI. AISÉN: Rı́o Exploradores,
Estancia Rı́o Circo, mapa n8 4673 Monte San
Valentı́n, 28-II/1-III-1967, Seki s.n. (HIRO 863-b);
REGIÓN XII. TIERRA DEL FUEGO: Rı́o Cóndor site, Crosby &
Castor 17462 (MO 4442293).
AUSTRALIA. AUSTRALIAN CAPITAL TERRITORY:
Nursery Creek, 42 km SSW of Canberra, Streimann
31393 (CHR 352127); NEW SOUTH WALES: Kosciusko
National Park, Yarrangobilly Caves, Lewinsky 1884
(CHR 350097); TASMANIA: Bothwell Municipality, south
end of Great Lake at intersection of road to Bronte
Park, Norris 33028 (H); VICTORIA: Somer’s Park, 30 km
N of Warburton, Great Dividing Range, Lewinsky 74–
307 (CHR 240266); WESTERN AUSTRALIA: The Cascade
Nature Reserve, 4 km S of Pemberton, Streimann
54247, Musci Australasiae Exsicc. 476 (MA 15263).
BOLIVIA. COCHABAMBA. PROV. CAPARRE: Pampa Tambo,
Hermann 25273 (MO 3670891); LA PAZ. PROV. INQUISIVI:
Pavionani, headwaters of the Rı́o Chimu along the
Milla Milla-Aguilani Trail ca. 2 km SE of Estancia,
Jankho Khalani and ca. 8 km N of Choquetanga, Lewis
89–946 d-3 (MO 3961451). NEW ZEALAND. AUCKLAND
ISLAND: N side of Musgrave Inlet on route to Mt.
Easton, Vitt 9054 (CHR 448821); NORTH ISLAND: Wellington Land District, Mt. Marina, 3-IX-1933, Zotov
s.n. (CHR 6912); SOUTH ISLAND: Marlborough, Kowhai
Bush, Kaikaura, Lewinsky 1122 (CHR 348590). INDONESIA. SUMATRA. WEST-SUMATRA: Urwälder am Südosthang des Singalang, 25-VII-1894, Schiffner s.n. (MA
941). SOUTH AFRICA. TRAANSVAL: in mont. Sechlaba,
Rehmann 500 (TR-VENTURI).
10. Zygodon jaffuelii Thér., Revista Chilena Hist.
Nat. 25: 300, tab. 27, fig. 1a–i. 1921. TYPE: CHILE.
‘‘Los Perales de Marga-Marga, sur troncs de
Bellota, c. fr. (Costes, ann. 1915 et 1916); monts
de Las Piedras (Jaffuel, ann. 1917)’’. (lectotype,
REGIÓN V. VALPARAÍSO:
Calabrese: Zygodon in southern South America
here designated, ‘‘Costes 3-I-1916’’, PC-THÉRIOT!;
isolectotypes, FH!, FH-BARTRAM!).
Illustrations. Figs. 3B, 18C–H, 19.
Description. Plants slender to slightly robust,
(0.2–)0.8–1.7 cm tall, in relatively lax turfs, light green
above, brown below. Stems erect, branched. Rhizoids
reddish-brown, smooth to slightly roughened, profuse
towards stems base. Leaves erect and quite twisted
when dry, erect-patent when moist, (0.6–)0.8–1.5 3
0.2–0.5 mm, lanceolate, base not or slightly decurrent,
apex acute; margins plane, entire and smooth at base,
papillose above, entire or slightly denticulate towards
apex; upper leaf cells rounded-hexagonal, thickwalled, (7–)9–11(13) lm in diameter, papillose;
papillae 4–5 per cell, rarely 6, simple, low, subacute or
blunt; basal cells subquadrate to short-rectangular
towards the margins, (7–)13–25(36) 3 (7–)11–20
lm, larger towards the costa, 16–57 3 13–25 lm,
smooth, with strongly thickened angles; costa ending
below the apex, greenish, reddish in older leaves, 24–
38 lm wide in the lower 1/3, in a deep furrow,
superficial cells elongate, dorsally smooth or with a
few disperse papillae, especially towards apex, ventrally covered by papillose laminal cells towards the
apex, in section with 2 ventral guide cells and dorsal
stereids. Propagula abundant, fusiform-clavate, with
thickened, brown walls when mature, 42–58 3 19–25
lm, (2–)3-celled, only with transverse septa. Dioicous. Perichaetial leaves undifferentiated. Seta 1.5–5
mm long, dextrorse in the distal part, sinistrorse at
base; capsule erect, cylindrical to ellipsoidal, 1.3–1.9
mm long, with 8 longitudinal ribs; exothecial bands
differentiated, of 3–4 cell rows each; operculum
conical at base, rostrate to subulate, beak curved;
peristome double; exostome of 8 pairs of teeth,
distally truncate, patent when dry, dorsal surface
(OPL) reticulate, reticulum usually smooth, rarely
with few papillae, ventral surface (PPL) almost
smooth or slightly roughened towards base, papillose
in the upper half; endostome of 8 short, hyaline
segments, dorsally (PPL) smooth, ventrally (IPL)
smooth to slightly roughened. Spores papillose, (10–)
12–16 lm in diameter.
Ecology. A typically epiphytic species mainly
growing on trunks of ‘‘bellota,’’ Nothofagus obliqua,
but it has also been collected on Populus sp.
Distribution (Fig. 24B). Zygodon jaffuelii seems
495
to be endemic to central and southern Chile, where it
is known from regions V, IX and X.
Differentiation. Zygodon jaffuelii resembles Z.
hookeri in habit. However, it is readily separated from
all other Zygodon species in the study area on
sporophytic grounds: its peristome is double, and the
truncate exostome teeth are a unique feature in the
genus. Furthermore, the basal leaf cells are collenchymatous and oblate to short-rectangular, and the
walls of the mature propagula are brown.
Zygodon jaffuelii has been occasionally confused
with Zygodon hookeri var. leptobolax. However, Z.
jaffuelii is a dioicous species with a double peristome,
undifferentiated perichaetial leaves, brown propagule
walls and smaller spores (always less than 16 lm in
diameter). On the other hand, Z. hookeri var.
leptobolax is a synoicous species lacking an exostome,
with differentiated perichaetial leaves, usually hyaline,
greenish or yellowish propagula and larger spores
(always more than 18 lm in diameter).
Variability. Zygodon jaffuelii hardly varies. The
only character showing some variability is seta length,
ranging from 1.5 to 5 mm. Plants are usually 0.8–1.7
cm tall, but some occasional specimens only reach 0.2
cm. No other structure shows significant variation.
Zygodon jaffuelii has been considered a synonym
of Z. liebmannii Schimp. ex Müll. Hal. (Allen 2002;
Grout 1946). However, Z. liebmannii, a rare species
with few specimens known, most of which have been
studied for this treatment, has lax, longer (35–45 lm),
thin-walled and non-collenchymatous basal leaf cells,
an almost percurrent costa and a commonly decurrent base. On the contrary, Z. jaffuelii has firm,
shorter (13–25(36) lm), and collenchymatous basal
cells, a costa that always ends a short distance from
the apex and the leaf base is not or only slightly
decurrent. Furthermore, Z. jaffuelii commonly has
abundant propagula, while all the studied specimens
of Z. liebmannii, including all known types, lack
propagula. Contrary to my observations, Malta (1926)
described propagula for Z. liebmannii, morphologically similar to those in Z. jaffuelii. I have studied two
out of the three specimens of Z. liebmannii cited by
Malta (1926)—Pico de Orizaba, Liebmann s.n. and
Cima, Pringle 10546—and I could not find the
propagula he reported. Vitt (1994) described Z.
liebmannii as having propagula with 7–8 cells and
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Figure 19. Zygodon jaffuelii. A. Habit when dry. B. Habit when moist. C. Leaves. D. Apices. E. Cells in middle portion of leaf. F. Leaf
basal cells. G. Leaf cross-sections. H. Propagula. I. Peristome (inner view) and spores. A from Jaffuel s.n. (FH); B–I from Costes s.n.
(PC).
Calabrese: Zygodon in southern South America
longitudinal as well as transverse septa. In PC there are
two type specimens of Z. jaffuelii (with duplicates in
FH-BARTRAM, FH-FLEISCHER, MO and NY, all seen)
annotated by D. Vitt as Z. liebmannii. One of them is
in fact Z. jaffuelii, but the other is typical Z. papillatus.
The propagula of Z. papillatus have 7–8 cells and
longitudinal as well as transverse walls, and this
specimen of Z. papillatus seems to be the base of the
confusion in Vitt (1994).
Selected specimens examined (23). CHILE. REGIÓN
V. VALPARAÍSO: Los Perales (Marga Marga) sur troncs
de Bellota, 3-I-1916, Jaffuel s.n. (FH, MO 3987831);
REGIÓN IX. CAUTÍN: Dept. Villarica, Pucon, Hosseus 156
(JE-HERZOG); REGIÓN X. VALDIVIA: Entrance lane to
UACH campus, Isla Teja, Valdivia, Prins RP-6-22851b (MO 3665841).
Specimens examined of Zygodon liebmannii.
MEXICO. VERACRUZ: Orizaba, Liebmann s.n. (BM-SCHIMP.
555687 b, BM-SCHIMP. 555687, both types).
Zygodon reinwardtii-hookeri complex
The Zygodon reinwardtii-Zygodon hookeri complex lacks formal nomenclatural circumscription,
although I have included in it several species of
Zygodon difficult to distinguish that share a unique
combination of characters. They are usually synoicous, have differentiated perichaetial leaves, a simple
peristome of eight endostomial segments, papillose
and large (frequently greater than 18 lm in diameter)
spores and long-rostrate to subulate opercula. This
complex is represented in the study area by Zygodon
reinwardtii, Z. hookeri var. hookeri and Z. hookeri var.
leptobolax. Although the typical expressions of these
three taxa have distinctive features, the morphological
characters show wide variation, which makes it almost
impossible to name specimens with intermediate
characteristics. Specimens from the entire distribution
range of the three taxa have been examined, but no
clear differential morphological characters were
found, consequently more studies are needed to
clarify the validity of the taxa in this complex.
11. Zygodon reinwardtii (Hornsch.) A. Braun ex
Bruch & Schimp., Bryol. Europ. 3: 41 (fasc. 4,
monogr. 9). 1838; Syrrhopodon reinwardtii
Hornsch., Nova Acta Phys.-Med. Acad. Caes.
Leop.-Carol. Nat. Cur. 14(2): 700, tab. 39, fig. a.
1829. TYPE: ‘‘Hab. in Iava [Blume, Reinwardt]’’
497
(type, BM?; there are no original materials at L,
Touw, pers. comm.).
Zygodon moritzianus Müll. Hal., Linnaea 19: 199.
1846. TYPE: ‘‘Columbia, prov. Mérida, in montibus nivosis, Moritz’’. Synonymized by Malta
(1926: 118).
Zygodon denticulatus Taylor, London J. Bot. 6: 329.
1847. TYPE: ECUADOR. ‘‘Andes Quitenses; Monte
Pichincha, Will. Jameson, Nov. 1846’’ (lectotype,
here designated, G-027226!; isolectotype, G027222!). Synonymized by Malta (1926: 118).
Zygodon cylindricarpus Müll. Hal., Bot Zeitung
(Berlin) 11: 59. 1853. TYPE: INDIA. ‘‘Nilghiri’’.
Synonymized by Malta (1926: 118).
Zygodon subdenticulatus Hampe, Ann. Sci. Nat. Bot.,
V, 4: 326. 1865. TYPE: COLOMBIA. ‘‘Bogota, Pacho,
Chiquinquira, Guadalupe, Tequendama, Canoas
et Paramo Choachi, altit. 2090–3600 m, in sylv.
sub n8 2092, 2126, 2129 et 2145, leg. A. Lindig’’
(syntypes, G-027232!, G-027231!). Synonymized
by Malta (1926: 118).
Zygodon andinus Mitt., J. Linn. Soc. Bot. 12: 236.
1869. TYPE: ECUADOR. ‘‘Andes Quitenses, in monte
Pichincha (9000 ped.) in devexu australi, Spruce
117’’ (holotype NY, not seen; isotype, G-027235!).
Synonymized by Malta (1926: 118).
Zygodon circinatus Schimp. ex Besch., Mém. Soc. Sci.
Nat. Cherbourg 16: 187. 1872. TYPE: MEXICO.
‘‘Cordova (F. Müller)’’ (isotype, BM?, not found).
Synonymized by Malta (1926: 118).
Zygodon breviciliatus Thwaites & Mitt., J. Linn. Soc.
Bot. 13: 304. 1873. TYPE: ‘‘In Ceylon, Dr.
Thwaites’’ (holotype, Thwaites 73, NY, not seen).
Synonymized by Malta (1926: 118).
Zygodon glaziovii Hampe, Vidensk. Meddel. Dansk
Naturhist. Foren. Kjøbenhavn, III, 6: 142. 1874.
TYPE: BRAZIL. ‘‘Serra do Itatiaia, Dusén, 1902’’
(isotype, HBG!). Synonymized by Malta (1926: 118).
Zygodon ceratodontoides Müll. Hal., Flora 58: 535.
1875. TYPE: ‘‘Colombia, Ocanna, leg. Wallis.’’
Synonymized by Malta (1926: 118).
Zygodon caldensis Ångstr., Öfvers. Förh. Kongl.
Svenska Vetensk.-Akad. 33(4): 15. 1876. TYPE:
BRAZIL. ‘‘Caldas, Henschen.’’ Synonymized by
Malta (1926: 118).
Zygodon fendleri Müll. Hal., Linnaea 42: 484. 1879.
TYPE: ‘‘Tovar, 1854–55, Fendler Musci Venezue-
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lenses N8 28’’ (lectotype, here designated, G027227!). Synonymized by Malta (1926: 118).
Zygodon gymnus Müll. Hal., Linnaea 42: 484. 1879.
TYPE: ‘‘Tovar, 1854–55, Fendler Musci Venezuelenses N8 27’’ (lectotype, here designated, G027224!; isolectotype, HBG!). Synonymized by
Malta (1926: 119).
Zygodon borbonicus Besch., Ann. Sci. Nat. Bot., VI, 9:
353. 1880. TYPE: ‘‘La Réunion: G. de l’Isle, 1875,
n8 420; P. Lépervanche, 1877.’’ Synonymized by
Malta (1926: 119).
Zygodon hyadesii Besch., Bull. Soc. Bot. France 32: LX.
1885; Zygodon reinwardtii var. hyadesii (Besch.)
Malta, Latvijas Univ. Bot. Dārza Darbi 1: 121.
1926. TYPE: ‘‘Terre de Feu (Dr. Hyades); Patagonia occidentale, ı̂le Wellington: Port-Eden (Dr.
Savatier)’’. (lectotype, here designated, CHILE.
‘‘Terre de Feu. Hyades,’’ NY!), syn. nov.
Zygodon madacassus Schimp. in Wright, J. Botany 26:
264. 1888, nom. nud.
Zygodon paucidens Müll. Hal., Nuovo Giorn. Bot. Ital.
4: 120. 1897. TYPE: BOLIVIA. ‘‘Germain [. . .] 1889
prope Choquecamata provinciae Cochabamba
Bolivianae alto-montosae ex altitudine pedum
10–12,000, ut opinor, collectas’’ (lectotype, here
designated, G-027225!). Synonymized by Malta
(1926: 119).
Zygodon volkensii Broth., Bot. Jahrb. Syst. 24: 241.
1897. TYPE: TANZANIA. ‘‘Kilimanscharo’’. Synonymized by Malta (1926: 119).
Zygodon dives Müll. Hal., Bull. Herb. Boissier 6: 94.
1898. TYPE: BRAZIL. ‘‘Serra do Itatiaia, Dusén,
1902’’ (lectotype, here designated, HBG!). Synonymized by Malta (1926: 119).
Zygodon coarctatus Schimp. ex Paris, Index Bryol.
1372. 1898, nom. nud.
Zygodon argutidens Broth. in Mildbr., Wiss. Erg. Deut.
Zentr.-Afr. Exped., Bot. 2: 148, tab. 13, fig. 8. 1910.
TYPE: UGANDA/ZAIRE. ‘‘Ruwenzori: Butagu-Tal,
Ericaceen-Stämme und Zweige, halbkugelige Polster bilden, ca. 3300 m ü. M. (reichlich fruchtend
im Febr.—1908—n. 2616’’ (holotype H?, not
found). Synonymized by Malta (1926: 119).
Zygodon reinwardtii var. madacassus Cardot in A.
Grandid. & G. Grandid., Hist. Phys. Nat. Polit.
Madagascar 39: 229. 1915, nom. inval., not
accepted by author (ICBN St. Louis, art. 34.1).
Zygodon reinwardtii var. subintegrifolius Malta, Latvijas Univ. Bot. Dārza Darbi 1: 122, fig. 69d. 1926.
TYPE: ‘‘Brasilien, Caldas, Henschen; Columbien,
Bogota, Lindig.’’ Synonymized by Anderson et al.
(1990: 498).
Zygodon lindigii Hampe ex Malta, Latvijas Univ. Bot.
Dārza Darbi 1: 178. 1926, nom. nud.
Pleurozygodontopsis decurrens Dixon, Ann. Bryol. 12:
51. 1939. TYPE: ‘‘Sumatra, Gunong Losir, Atjeh,
Gajolanden, 2940 m, 5-II-1937, van Steenis
10159’’ (lectotype, here designated, BM-DIXON
4044!). Synonymized by Eddy (1996: 16).
Illustrations. Figs. 2A, 3A, 20.
Description. Plants slender to somewhat robust,
1.0–2.8 cm tall, in lax turfs, light-green to yellowishgreen above, brown below. Stems erect, branched.
Rhizoids reddish-brown, smooth, roughened or
slightly papillose. Leaves erect and 6 twisted when dry,
spreading and flexuose when moist, (0.9–)1.1–2.3 3
(0.19–)0.27–0.49 mm, lanceolate, keeled, base decurrent, apex acute; margins usually undulate, less
frequently almost plane, entire at base, papillose above,
usually dentate at apex and in the upper 1/3, rarely
entire or with 1–2 teeth at apex; upper leaf cells
rounded-hexagonal, isodiametric, thick-walled, 8–12
lm in diameter, papillose; papillae (2–)4–6 per cell,
simple, acute to subacute, rarely blunt; basal cells
subquadrate to rectangular, thick-walled, 18–2336–10
lm, shorter towards margins, 9–15 3 6–8 lm, smooth,
ocasionally with 1–2 papillae; costa ending below the
apex, 28–45 lm wide in lower 1/3, in a furrow, dorsally
smooth or with a few dispersed papillae towards the
apex, in section with 2 guide cells and dorsal stereids.
Propagula infrequent and scarce, fusiform, fusiformclavate or clavate, with hyaline or greenish-hyaline,
rarely brown walls, 45–167 3 17–27(36) lm, usually
3–7-celled with only transverse septa, rarely with more
cells and with transverse and longitudinal septa.
Synoicous. Perichaetial leaves differentiated, erect
when moist, 1.5–1.7 3 0.3–0.4 mm, lanceolate, apex
acute-acuminate; margins entire. Seta (5–)9–15 mm
long, dextrorse in the distal part, sinistrorse at base;
capsule erect, rarely oblique, cylindrical or fusiform,
sometimes slightly gibose, narrowed towards capsule
mouth, 1.5–3.2 mm long, with 8 ribs throughout;
exothecial bands slightly differentiated, of 4–6 cell rows
each, interrupted by a continuous ring of 2–6 smaller
Calabrese: Zygodon in southern South America
499
Figure 20. Zygodon reinwardtii. A. Habit when dry. B. Habit when moist. C. Leaves. D. Perichaetial leaf. E. Apices. F. Leaf basal cells.
G. Leaf cross-section. H. Propagula. I. Wet capsule. J. Capsules when dry. K. Peristome and spores. A, B, cII, eII, I, J from P. de la
Bathie (PC-THÉRIOT); cI, hII from Ule 131 (G); cIV, D, hI from Fendler 27 (G); cIII, eI from Skottsberg 435 (S); cV, K from Matteri &
Schiavone TBPA-B 2276 (BA); F, G from Sparre 2044 (S); hIII from Dusén s.n. 29-II-1896 (S); hIV from Germain s.n. (G).
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cells around capsule mouth; operculum conical at base,
subulate, beak curved; peristome simple, exostome
absent; endostome of 8 hyaline segments, dorsal (PPL)
and ventral (IPL) surfaces smooth, sometimes slightly
roughened, basal membrane low, continuous and
smooth. Spores papillose, (15–)18–25 lm in diameter,
trilete marks absent.
Ecology. Zygodon reinwardtii is typically an
epiphyte in Quercus spp., Bonnetia roraimae Oliv. or
Nothofagus spp. forests, usually on small branches and
twigs of shrubs and trees like Drimys winteri J.R.
Forst. & G. Forst., Polylepis sp., Erythrina sp.,
Hypericum sp., diverse Ericaceae or ferns like
Blechnum. It is also occasionally found on soil and
rocky areas with Racomitrium crispulum (Hook.f. &
Wilson) Hook.f. & Wilson. In the Americas it is very
common in the Venezuelan and Colombian paramos.
Distribution (Fig. 24B). This species has a wide
distribution, especially in the tropics. It is known from
Mexico, Central America, South America (from
Venezuela to Tierra del Fuego), Africa, Madagascar,
India, Sri Lanka and SW Asia (China). It has been also
reported from Alaska, Philippines, Java and New
Guinea, but these records are likely confusions with Z.
hookeri var. hookeri, as explained above (p. XX). Malta
(1926) and Lewinsky (1990) discussed the variability of
Zygodon reinwardtii and Z. hookeri, and concluded that
the plants from Australia, Tasmania and New Zealand
belong to Z. hookeri. In the study area it is known from
Santa Cruz and Tierra del Fuego Provinces of
Argentina, and from regions X, XI and XI of Chile.
Differentiation. Zygodon reinwardtii is closely
related to the also variable Z. hookeri, and both can be
difficult to distinguish. Only typical expressions of
both are clearly separated: Zygodon reinwardtii has
decurrent leaves strongly dentate at apex, while Z.
hookeri var. hookeri has slightly decurrent leaves
denticulate at apex. Propagula in the latter are clavatefusiform, smaller (3-celled), and never with longitudinal walls. In addition, Z. hookeri has basal leaf cells
usually larger and capsules are always cylindrical.
Variability. Typical Zygodon reinwardtii has
strongly undulate leaves sharply dentate at the apex,
sometimes in the upper 1/3. Nevertheless, both
features vary considerably. Some leaves have only one
or two well-developed teeth, hardly visible, or totally
lack them, and specimens intermediate in marginal
teeth development are common. Similarly, leaf
undulation is very variable, even in the same specimen. Regarding propagula, although they are a stable
structure with a set of taxonomically important
characters for most Zygodon species, in Z. reinwardtii
they are so variable that they seem to represent
different taxa. Most Z. reinwardtii specimens have
hyaline or greenish-hyaline fusiform to clavate-fusiform propagula, with only transverse septa. Only a
specimen from Choquecamata (Bolivia) has brown to
greenish-brown, clavate propagula with transverse
and longitudinal septa. Despite these differences,
propagula are not frequent or abundant in Z.
reinwardtii, which could hamper correct estimation of
this variability. Sporophytically Z. reinwardtii is a
more stenotypic taxon, with 6 cylindrical to fusiform
and slightly gibbous capsules 1.5–3 mm long. An
anomalous expression, f. cerantodontoiodea Malta
(1926: 121), described from Colombia on the basis of
its strongly curved capsules, was unavailable for study.
On the basis of gametophyte differences as well as
geographical providence, many taxa have been
described around Z. reinwardtii. Many of them were
subsequently synonymized with typical Z. reinwardtii,
mainly by Malta (1926), with only var. hyadesii
(Tierra del Fuego and West Patagonia), var. ruwenzorensis (Thér. & Naveau) Demaret (Ruwenzori) and
var. capillicaulis (Müll. Hal.) Malta (Brazil), being
currently recognized. The type and the many specimens previously identified as Z. reinwardtii var.
hyadesii, that have been examined for this study, have
leaves that fall into the range of variation observed for
the typical expression. There is no correlation between
the paucity of apical teeth, on which basis this variety
was described, and any other feature; in fact most
specimens belong to Z. hookeri var. leptobolax.
Regarding var. ruwenzorensis and var. capillicaulis, I
have reached the same conclusion after studying
specimens from Ruwenzori (Uganda/Zaire) and Serra
do Itatiaia (Brazil), although I delay formal synonymization until types of both are studied. Finally,
Cardot (in Renauld & Cardot 1915: 229) also
mentioned that specimens from Madagascar labelled
Z. madacassus have less dentate leaves, smaller laminal
cells and longer seta. However, he did not accept those
differences enough to consider it different from
typical Z. reinwardtii, even at varietal rank (ICBN St.
Louis, Art. 34.1, Ex. 3), an opinion with which I concur.
Calabrese: Zygodon in southern South America
Selected specimens examined (127). ARGENTINA.
Lago San Martı́n, Brazo NW, Skottsberg
436 (S); TIERRA DEL FUEGO: Lago Fagnano, Skottsberg 776
(S). CHILE. REGIÓN X. LLANQUIHUE: Puerto Varas,
Ensenada, Sparre 2044 (S, FH-BARTRAM); REGIÓN XI.
AISÉN: Rio Aysén, 30-I-1897, Dusén s.n. (S); REGIÓN XII.
ÚLTIMA ESPERANZA: Bahı́a Bellavista, Matteri & Schiavone TBPA-B 2276 (BA).
BOLIVIA. COCHABAMBA. PROV. CHAPARE: sureste
de Laguna Curani, Churchill et al. 19840 (MO
5211664); LA PAZ. PROV. INQUISIVI: Valle de Chimu, 2
km NW of Estancia Chimu and 6 km NE of
Choquetanga, Lewis 38804 d-1 (MO 3960868). BRAZIL.
RIO DE JANEIRO: Parque National de Itatiaia, Pico das
Agulhas Negras, N of Engo Passos and 12.4 km NE of
União, Vitt 21574 (MO 2555716). COLOMBIA. ANTIOQUIA:
Municipio La Ceja, 5,5 km SE de La Ceja hacia La
Unión, Churchill & Sastre-De Jesús 12855 (H); BOYACÁ:
Municipio de Duitama, around Paramo ‘‘La Rusia’’,
Ireland 23568 (CANM 308703, MO 3663203); CAUCA:
Valle del Cauca, Darien, ‘‘Páramo del Duende’’, Cerro
Calima, ca. 60 km N de Cali, Churchill 19556 (MO
5125845); CUNDINAMARCA: Tausa, Cuchilla Laguna
Seca, alrededores Laguna Verde, Churchill et al. 19369
(MO 5125608); HUILA: páramo de Las Papas, cerros y
alrededores de las lagunas La Magdalena y Santiago,
Bischler 872 (MO 5137008); NARIÑO: municipio de
Pasto, Correg. de La Laguna, alrededores Alto
Zapallurco, Ramı´rez & Salas 10835 (MO 4430748).
DEMOCRATIC REPUBLIC OF THE CONGO. Massif du
Ruwenzori, [? illeg.], VII-1929, Humbert s.n. (PC-P. DE
LA VARDE). ECUADOR. Cordillera, Jaeger s.n. (TRVENTURI); NAPO: along road Baeza-Quito, near Cuyuga,
van der Werff et al. 13455A (MO 4420448); PICHINCHA:
vicinity of Rı́o Amalbi, along road leading W from
Nono toward Tandayapa, N slopes of Mt. Pichincha,
Crosby 10540 (MO 2330102); QUITO: Guayrapata,
Spruce 116 (FH-BARTRAM). MADAGASCAR. Mt. Tsaratanana, IV-1924, P. de la Bathie (PC-THÉRIOT). PERU.
CAJAMARCA: Puente Bajo (Chalchuayacu) zw. Encañada
u. Celendin, Hegewald & Hegewald 6615 (MO
3670867); CHAPAPOYAS: Dpto. Amazonas, Chanchillo
zw. Balsas u. Leimebamba, Hegewald & Hegewald
6780 (MO 3670893); OXAPAMPA: Dpto. Pasco, ca. 5 km
von Oxapampa in Richtung Pozuzo, Hegewald &
Hegewald 8441 (MO 3670866); URUBAMBA: Dpto. Cuzco,
Machu Picchu, Hegewald & Hegewald s.n. (MO
SANTA CRUZ:
501
5147610); YUNGAY: Dpto. Ancash, An der Laguna
Llanganuco, Hegewald & Hegewald 7529 (MO
3670869). VENEZUELA. BOLÍVAR: Distrito Piar, Macizo
del Chimantá, sector centro-noreste del Chimantátepui, cabeceras orientales del Caño Chimantá,
Steyermark et al. 128052-A (MO 3061984); MÉRIDA:
Sierra Nevada, teleférico Loma Redonda, camino
hacia los nevados, Cleef & Huber 4787 (MO 3673453).
12. Zygodon hookeri Hampe, Linnaea 30: 632. 1869.
TYPE: AUSTRALIA. ‘‘Hab. ad arbores, mont. Grampian.’’ [1854, Dr. F. Mueller 5] (holotype BM!).
Description. Plants slender to slightly robust,
(0.3–)1.2–3.5 cm tall, in loose turfs, green to yellowish-green above, reddish-brown below. Stems erect,
branched. Rhizoids reddish-brown to yellowish,
smooth or slightly roughened, ascending by stems.
Leaves erect, 6 twisted and keeled when dry,
spreading and flexuose when moist, (0.9–)1.35–2.38 3
(0.25–)0.35–0.55 mm, lanceolate, sometimes narrowed towards the base, base slightly decurrent, apex
acute; margins undulate to plane, entire below,
papillose above, denticulate to entire at apex; upper
leaf cells rounded-hexagonal, 6 isodiametric, thickwalled, (6–)9–14 lm in diameter, papillose; papillae
3–9 per cell, usually simple, subacute, rarely blunt;
basal cells short-rectangular to rectangular, thickwalled, (10–)18–44(63) 3 6–15 lm, smooth or with
1–2 papillae; costa ending below the apex, 30–42(51)
lm wide in lower 1/3, in a deep furrow, superficial
cells elongate on both sides, dorsally smooth in lower
2/3, smooth or papillose towards the apex, in section
with 2 ventral guide-cells and dorsal stereids.
Propagula infrequent or common, scarce to abundant, fusiform to clavate-fusiform, with hyaline or
greenish-hyaline (rarely brown) walls, 36–81(90) 3
12–30 lm, (2–)3–5(6)-celled, with only transverse
septa, ocassionally with transverse and longitudinal
septa and then to 10-celled. Synoicous (very rarely
autoicous in Australia, see Variability). Perichaetial
leaves differentiated, erect when moist, 0.9–1.8 3
0.27–0.39 mm, lanceolate, apex acute-acuminate;
margins entire or with some papillae towards the
apex; cells elongate, smooth at base, smaller and
papillose towards the apex. Seta (4–)10–15 mm long,
dextrorse in the distal part, sinistrorse at base; capsule
erect, cylindrical, 1.3–2.5 mm long, with 8 ribs
througout; exothecial bands differentiated, of 4–5 cell
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rows each, interrupted by a continuous ring of 3–6
smaller cells around capsule mouth; operculum conic
at base, subulate, beak curved; peristome simple,
exostome absent; endostome of 8 hyaline segments,
dorsal (PPL) and ventral (IPL) surfaces smooth,
occasionally slightly roughened, basal membrane low,
continuous and smooth. Spores papillose, (18–)20–26
lm in diameter, trilete mark absent.
12a. Zygodon hookeri Hampe var. hookeri
Zygodon mucronatus Müll. Hal., Hedwigia 37: 136.
1898, nom. illeg., non Broth. 1896; Z. submucronatus Paris, Index Bryol. Suppl. 1: 330. 1900.
TYPE: ‘‘New Zealand’’ (isotype, GOET). Synonymized by Lewinsky (1990: 133).
Illustrations. Figs. 2B, 21.
Diagnosis. Plants slender to slightly robust, (0.4–)
1.2–3.5 cm tall. Leaves 6 twisted when dry, spreading
and flexuose when moist, (0.9–)1.35–2.38 3 0.35–0.55
mm, lanceolate; margins undulate, entire at base,
papillose above, typically denticulate, occasionally
entire at apex; upper leaf cells (6–)9–12 lm in
diameter, 3–6(9) papillae per cell; basal cells
rectangular, 18–44 3 6–13 lm. Propagula infrequent,
scarce, with hyaline or greenish-hyaline walls, 36–52 3
12–20 lm, 3-celled, with only transverse septa.
Selected specimens examined (48). ARGENTINA.
SANTA CRUZ: Dpto. Lago Argentino, Seno Mayo,
pequeña bahı́a al NE del Seno, Matteri & Schiavone
CM 4910 (BA, MA). CHILE. REGIÓN XII. ÚLTIMA ESPERANZA:
glaciar del cerro Balmaceda, margen occidental del
Puerto Toro, Matteri TBPA B 2714 (BA).
AUSTRALIA. TASMANIA: Bothwell Municipality,
south end of Great Lake at intersection of road to
Bronte Park, Norris 33028.(H); VICTORIA: Mt. William,
Grampians, 19 km SSE of Halls Gap, Streimann HS3031 (H). NEW ZEALAND. NORTH ISLAND: South Auckland, near Atiamuri, south of Ratorwa, Allison 486
(CHR 523861); SOUTH ISLAND: Nelson Lakes Nat. Park,
St. Anould, Frahm 23–3 (CHR 503103).
12b. Zygodon hookeri var. leptobolax (Müll. Hal.)
Calabrese, comb. nov.
Zygodon leptobolax Müll. Hal., Hedwigia 38: 113. 1899.
TYPE: SOUTH AFRICA. ‘‘Prom. bonae spei, in fruticetis
prope Rondebosch, aug. 1875, c. fr. vetustis vel valde
juvenilibus: Dr. A. Rehmann, Coll. N8 379,’’ but
really: Cape, near Rondebosch, Rehmann 499
(lectotype, here designated, PRE!). NOTE: Dixon and
Gepp (1923) published a checklist of Rehmann’s
collections from South Africa in which Rehmann
379 belongs to Brachythecium afro-velutinum Müll.
Hal. The only collection of Z. leptobolax is Rehmann
499, published in his exsiccata with the number 147.
Magill (1980) and Magill and Van Rooy (1998) also
considered this collection as the type of Z. leptobolax.
Zygodon inermis Dusén ex Malta, Latvijas Univ. Bot.
Dārza Darbi 1: 174. 1926; Zygodon hyadesii var.
inermis Dusén, nom. nud. in sched. TYPE: ‘‘Chile
australis, ins. Quiriquina in truncis arb., P.
Dusén, 20-IX-1896’’ (lectotype, here designated,
S!; isolectotype, PC!), syn. nov.
Zygodon theriotii Herzog, Arch. Esc. Farm. Fac. Ci.
Méd. Córdoba (Argentina) 7: 47. 1939. TYPE:
‘‘Chile: Prov. Cautin, Dep. Villarrica: Pucón (N8
803, 804 y 822); Prov. Chiloé, Dep. Llanquihué:
Petrohué (N8 497, 521 y 532)’’ (lectotype, here
designated, Hosseus 822, JE-HERZOG!; syntypes,
Hosseus 497, Hosseus 532, Hosseus 803, Hosseus
804, all JE-HERZOG!), syn. nov.
Illustration. Fig. 22.
Diagnosis. Plants slender to slightly robust, (0.3–)
0.8–1.8 cm tall. Leaves twisted when dry, spreading and
flexuose when moist, (1.05–)1.3–2.1 3 (0.25–) 0.31–
0.49 mm, lanceolate; margins plane, sometimes somewhat undulate, entire at base, papillose above, typically
entire at apex, although occasionally with few small
teeth; upper leaf cells 8–14 lm in diameter, with (4–)6–
9 papillae per cell; basal cells short-rectangular to
rectangular, (10–)19–40(63) 3 6–15 lm. Propagula
common, abundant, with hyaline, rarely brown or
greenish-hyaline walls, 45–81(90) 316–30 lm, (2–)
3–5(6)-celled, with only transverse septa., rarely with
also longitudinal septa and consequently to 10-celled.
Selected specimens examined (38). CHILE. REGIÓN
VIII: Chile australis, Ins. Quiriquina, Dusén 226 (H);
REGIÓN IX. CAUTÍN: Dep. Villarrica, Pucón, Hosseus 804
(JE-HERZOG, S-B6271, syntype); REGIÓN X. LLANQUIHUE:
Lac. Llanquihue, Dusén 701 (S); REGIÓN XI. AISÉN: Rı́o
Exploradores, Estancia Rı́o Circo, 2/5-III-1967, Seki
s.n. (HIRO 1133); REGIÓN XII: Otway, Puerto Pomar,
Skottsberg 435 (FH-BARTRAM).
Ecology. Zygodon hookeri usually grows on tree
trunks, shrub branches and fallen logs in forests, more
rarely on rotten wood. In the study area it has been
found on Nothofagus betuloides, Nothofagus pumilio,
Berberis spp. and Austrocedrus chilensis. In New
Calabrese: Zygodon in southern South America
503
Figure 21. Zygodon hookeri var. hookeri. A. Habit when dry. B. Habit when moist. C. Leaves. D. Perichaetial leaf. E. Apex. F. Cells in
middle portion of leaf. G. Leaf basal cells. H. Leaf cross-sections. I. Propagula. J. Capsule when dry. K. Peristome and spores. A, cII, E,
iI, J from Matteri & Schiavone CM 4910 (BA); B, cI, D, F, G, iII from Müller 5 (BM); H, iIII from Linbolt 236 (CHR); kI from Brownlie 824
(CHR); kII from Simpson 118 (CHR).
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Figure 22. Zygodon hookeri var. leptobolax. A. Habit when dry. B. Habit when moist. C. Leaves. D. Perichaetial leaf. E. Apices. F. Cells
in middle portion of leaf. G. Leaf basal cells. H. Leaf cross-section. I. Propagula. J. Peristome and spores. A from Hosseus 497 (JE); B,
cI, iIII, J from Dusén s.n. 20-IX-1896 (PC); cII from Rehmann 499 (PRE); cIII, eI, F, G, H from Hosseus 804 (S-B6271); D from Dusén s.n.
IV-1897 (S); E from Hosseus 822 (JE); eII, iI from Rehmann 499 (PRE); iII from Hosseus 822 (JE); iIV from Mahu 9274 (MO); iV from
Schwabe 45-a (JE).
Calabrese: Zygodon in southern South America
505
Figure 23. Distribution of Zygodon in southern South America: A. Z. menziesii (solid circle) and Z. obtusifolius (open circle). B. Z.
pentastichus (solid circle) and Z. magellanicus (open circle). C. Z. papillatus. D. Z. chilensis.
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Figure 24. Distribution of Zygodon in southern South America: A. Z. pichinchensis (solid circle) and Z. bartramioides (open circle). B.
Z. reinwardtii (solid circle) and Z. jaffuelii (open circle). C. Z. intermedius. D. Z. hookeri var. hookeri (open circle) and Z. hookeri var.
leptobolax (solid circle).
Calabrese: Zygodon in southern South America
Zealand it is also common on Nothofagus spp., and
has been rarely reported from rocks.
Distribution (Fig. 24D). Zygodon hookeri has a
sub-Antarctic distribution. Zygodon hookeri var.
hookeri is known from Australia, New Zealand and
southern South America. In the study area it is known
from Santa Cruz Province of Argentina, and Región
XII of Chile. Zygodon hookeri var. leptobolax is known
from South Africa and southern South America; it has
been found in regions VIII, IX, X, XI and XII of Chile.
Differentiation. Zygodon hookeri var. hookeri is
very similar to Z. reinwardtii, from which can only be
clearly distinguished when both are in their typical
expressions: decurrent leaves strongly dentate at apex
in Z. reiwardtii, and slightly decurrent and denticulate
leaves in Z. hookeri var. hookeri. Other characters such
as propagula size and shape are scarcely useful because
they are rarely observed in both taxa. Capsules are
usually of no use in distinguishing these taxa, except
in the rare occasions when they are 6 fusiform and
slightly gibbose in Z. reinwardtii (always cylindrical in
Z. hookeri var. hookeri). My results show that these
taxa could be synonyms, with Z. reinwardtii being the
oldest name.
Confusion with other species of similar habit,
such as Z. jaffuelii and Z. intermedius is unlikely.
These two species are dioicous, have undifferentiated
perichaetial leaves and double peristomes, while Z.
hookeri is synoicous, has differentiated perichaetial
leaves, and a single peristome.
Variability. Zygodon hookeri varies mainly in leaf
lamina undulation and apex denticulation, as well as
propagule morphology. Leaves are commonly undulate, although plane leaves have also been observed. Leaves apices in the type variety are
denticulate, but vary from denticulate to entire in
var. leptobolax. Finally, propagula are usually fusiform to clavate-fusiform, hyaline and only with
tranverse septa, although two collections from the
northern Andean-Patagonian region (Mahu 9274 and
8629, MO!), have brown, clavate propagula with
longitudinal as well as transverse septa (Fig. 22IIV);
both specimens grew on Austrocedrus chilensis in a
more mesic forest.
Regarding sexual condition, two Australian
collections of Z. hookeri var. hookeri (Streimann 2873
and 3031, H!) are autoicous, although this taxon is
507
almost always synoicous (always in the South
American specimens studied).
Traditionally, Zygodon leptobolax has been considered endemic to South Africa, and Z. inermis and
Z. theriotii endemics from southern South America.
However, all type materials of the three were studied
and no differences among them were found, so I have
concluded that the three names represent the same
taxon. These types are sporophytically identical to the
type of Z. hookeri, which only differs from them by the
more undulate and denticulate leaves, and uncommon, smaller propagula. In the study area there are
specimens exactly matching the type of Zygodon
hookeri, others are identical to South African materials
(i.e., Z. leptobolax), while others are intermediate
between these two morphological extremes. Pending
further taxonomic studies on African Zygodon I prefer
to maintain Z. leptobolax at the varietal rank.
ACKNOWLEDGMENTS
I thank the curators of BA, BCRU, BM, CHR, FH, G, H, HBG, HIRO, JE,
MA, MO, NY, PC, PRE, S, TR-VENTURI, UBC and UC for loan of
specimens. I also thank my Ph.D. dissertation advisors, M. J.
Elı́as (Universidad de Salamanca) and F. Lara (Universidad
Autónoma de Madrid), for their help and support. I am
indebted to Jesús Muñoz (Real Jardı́n Botánico de Madrid) for
his comments on an earlier version of the manuscript, his help
on nomenclatural issues, and providing me with rare literature.
Finally, I am grateful to the Agencia Española de Cooperación
Internacional (AECI) for the fellowship and to the Laboratorio
de Briologı́a (Universidad Autónoma de Madrid) for financial
support for the SEM work.
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