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Revision of "Dieffenbachia" (Araceae) of Mexico, Central America, and the West Indies 

Author(s): Thomas B. Croat 

Reviewed work(s): 

Source: Annals of the Missouri Botanical Garden, Vol. 91, No. 4 (Dec., 2004), pp. 668-772 

Published by: Missouri Botanical Garden Press 

Stable URL: http://www.jstor.org/stable/3298554 . 

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REVISION OF Thomas B. Croat2 

DIEFFENBACHIA (ARACEAE) 

OF MEXICOv CENTRAL 

AMERICAv AND THE WEST 

INDIES1 

ABSTRAC T 

The genus Die,ffienb(lchia Schott has approximately 135 species, most of them occurring in South America. Major 

centers of diversity for the genus include Colombia with 37 species, Ecuador (34), Peru (30), Brazil (27), Panama (20) 

and Costa Rica (13). 'lahere are 26 species in Central America, with 20 species (77Wo) new to science. These are D. 

burgeri Croat & Grayum, D. copensis Croat, D. crebripistillata Croat, D. davidsei Croat 8z Grayum, D. fortunensis Croat 

D. fosteri Croat, D. gr(ll(l(lntesiae Croat, D. horichii Croat 8z Grayum, D. isthrnia Croat, D. killipii Croat, l). lutheri Croat 

D. nitidipetiolat(l Cloate D. obscurinervia Croat, D. panarnensis Croat, and D. standleyi Croat describe(l herein and D. 

beachiana Croat & (,rayum, D. concinna Croat 8z Grayum, D. grayurnii Croat, D. harernelii Croat 8z (Jlayum, and D. 

tonduzii Croat & (rlayLllzl (lescribed elsewhere. Most species range from Nicaraguato Panama. Belize has only 1 species 

of Dieffenbachia; Mexi( o, E1 Salvador, and Guatemala have 2 species, followed by Honduras (3), Nicalarua (6), Costa 

Rica (13), an(l Panalssa (2()). Only a few Central American species could be considered wi(lespread. Altsong the most 

widespread are 1). oe rste *lii Schott and D. wendlandii Schott, both of which range from Mexico to Panarna, as well as 

D. nitidipetiolat(l as(1 1). tonduzii, which range from Honduras to Ecuador. Species enflellsism is hiXrh, especially in 

Costa Rica (t3) an(l lllstllna (9). A total of 9 species are shared hetween Panama and Costa lWie a. Eiglst s)ecies, almost 

31Wo of the total, [-tilltt' into South America. These are D. david ei, D. isthmia, D. killi)ii, 1). Iongis/)(lth(l, D. nitidipetiolata, 

D. obscurirl(rli(l, D. seguine, and D. tonduzii. Most of llsese only extend to Colorzllsia, lout llsêe species, D. 

killipii, D. nitidi/)( tiol(lt(ln and D. tonduzii, range to Ecuador. 0tily D. killipii ranges to tlse ( asteln slo^)e of the Andes. 

Die,ffienbachi(l ()gllirl() Itlnges into Brazil and Bolivia, from the Wt st Indies. 

lVey1l)orfls: Ala(etle, Central America, Dieffenbachia, Mexi(o, South America, West lls({ies. 

The family Ara( eae is worldwide in distribution, Schott ex K. Krause). Im)ortant lo( a1 centers of 

but most species o( ( ur in tropical areas. Its centers generic endemism in the family il( lude Brazil, 

of distribution in(lude both Asia and America whieh has four endemic genera (Bop,tl1era Mayo & 

(Croat, 1979), with 31 genera in the Americas and Ni(olson, Dracontioide.s Ellgl., Ge(lrlllzl N. E. Br., 

28 found in Asia. There are 16 genera in Africa, Zomicarpa Schott), and the Indomalaysian region, 

Madagascar, and the Seychelles. A few genera are with 14 endemic genera. At least 2?550 species, 

restricted to tem)erate regions of the Northern replesenting four-fifths of the totaln o(cur in the 

Hemisphere, inc luding the Mediterranean region Neotropics. The Central American flola comprises 

(Calla L., Lysichiton Schott, Orontium L., Peltandra 19 genera within the Araceae. 

Raf., Sym/)loc(lrl)us Salisb. ex Nutt.), including the With an estimate of more than 955() species in 

Mediterranean region (Arisarum Mill., Ambrosina Latin America, the Araceae are one of the largest 

Bassi, Arum 1,., Dracunculus Mill., Helicodiceros families of flowering plants in the legion. Yet no 

1 This revision could not have been completed without the dedicated and able assistance of my lolllser co-worker, 

Petra S(hmi(lt, who participated in nearly every phase of the work. She continued the research efforts lJnabated while 

I was in the lield. Monica Carlsen and Emily Yates, former and present Research Assistants, respe(tively, were responsible 

lol firlal editing and revision of the manuscript, including final preparation of legend images. Fred Keusenkothen 

was responsible for preparation of scanned images. Special thanks also must go to Cheryl Neurnan and Emily 

Colletti, llesearch Greenhouse managers during the course of this work, who maintained hundreds of Dieffenbachia 

collectiolss in healthy condition. Very special thanks also go to my colleague and fellow aroid specialist Mike Grayum, 

whose kllowledge of the Costa Rican Araceae is unparalleled. Dan Nicolson, from the U.S. National Herbarium, and 

Charlotte Taylor, M0, assisted with nomenclatural issues. John Rawlins, from the Carnegie Museum, Pittsburgh, determined 

most of the beetles collected for this revision of Dieffenbachia. Thanks also go to Helen Young for beetle 

determillations and for information on reproductive biology. Elenor Sauer has proofread or written Latin diagnoses for 

all of the new species. Her expert assistance is greatly appreciated. Finally, I would like to thank my wife, Patricia, 

who has put up with my long absences while doing fieldwork for more than three decades. She was always available 

as an editor and to solve computer and database problems. 

2 p. A. Schulze, Curator of Botany, Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. 

ANN. MISSOURI BOT. GARD. 91: 668-772. 2004.

Volume 91, Number 4 

2004 

Croat 

Revision of Dieffenbachia 

other family is so poorly known taxonomically. with the lowest totals just north of the San Juan 

Plants exhibit considerable morphological plasticity depression (Nicaragua), followed by a marked inat 

all stages of development. Many of the genera, crease approaching the South American continent. 

especially the hemiepiphytic genera Rhodospatha (In this treatment, Central America is defined as all 

Poepp., Monstera Adans., and Syngonium Schott, the area between Mexico and Colombia, whereas 

exhibit complex patterns of heterophylly, with dras- Middle America is all the area between Mexico and 

tically different morphology at different stages of Panama.) Mexico has 2 species, Guatemala (2), Bedevelopment 

(Croat, 1981 [1982]; Ray, 1981, lize (1), Honduras (3), E1 Salvador (2), Nicaragua 

1987). Most members of the family have succulent (6), Costa Rica (13), and Panama (20) (Table 1). 

parts, making them difficult to collect and preserve. Endemism is high, especially in Costa Rica with 3 

Dieffenbachia is one of several medium-sized endemic species and Panama with 9. A total of 7 

genera in the Araceae, with an estimated 135 spe- species are shared between Panama and Costa 

cies. This revision is the first major review of Dief- Rica. Just under one-third (31Wo) of Central Amerfenbachia 

for Central America since Adolf Engler's ican species of Dieffenbachia range into South 

(1915) generic treatment in Das Pf anzenreich. America, these being D. davidsei, D. isthmia, D. 

Dieffenbachia is one of the most important genera killipii, D. Iongispatha, D. nitidipetiolata, D. obscuof 

understory herbs in the family, and it is often a 

dominant component of humid to wet tropical forrinervia, 

D. seguine, and D. tonduzii. 

ests, especially from sea level to 1500 m. It inhabits 

life zones ranging through Premontane moist for- 

MATERIALS AND METHODS 

est (P-mf), Tropical moist forest (T-mf), Premontane This revision is based on more than 29 years of 

wet forest (P-wf), Tropical wet fore.st (T-wf), and Pre- field studies in Central and South America, bemontane 

rain forest (P-rf) (Holdridge, 1967). While tween 1967 and 1996. All but one species were 

most spevies occur in virgin humid forests, the ge- studied live in the field or under cultivation at the 

nus is known from freshwater swamps, stream Missouri lAotani(al Garden. The descriptions have 

tanks, reglowth forest, among lo(k out(rosX aIl(l I)een prepale(l flom }oth living and drie(l spe(io(easionally 

on roa(l l)anks. It often ( onstitutes the mens. The ex(el)tion is D. fo.steri, where the demost 

tonsl)i(uous elelllent of the un(lelstoly vege- s(ril)tioll was lase(l solely on the tyl)e sI)e(inen. 

tation })e(ause of its a}un(la-l(e, fe(luellt (olollial rRle use of ("(llie(l") pre(e(lillg a11 or any lart of 

glowth 11a}it, an(l genelully ]aIge, showy leaves. the des( ril)tioll is an in(lie ation tllat a11 that follows 

The genus t)IOVi4eS a wi(lP valiely of (hoi(e orlla- is lase(l oll let })al iuln nlatel ial ollly. Mol l)hologi( al 

mental plants for horti( ulture, in( lu(ling nlost of the (hala(tes wee *o(le(l dire(tly into a (oml)uterize(l 

ST)et ieS treate,(l he,l e. 

data})ase to ensule parallel an(l sortal)le (lescriy)- 

The genus is (listine t an(l not easily ( onfuse(l tions. 'I'he aloi(l dese liptions (latalease ( ontains 72J4 

with any other aroi(l. It is ( losest to Bog7lera, whie h c halae tel states aplicable to the morphologie al didiffers 

in la(king staminodia sulrounding the pis- versity exlesse(l in DiefWenh(leSli(l (Croat, 1997). 

tillate flowers and in having higher-order venation Dis(ussions and references to illustrations, as well 

reticulate. In contrast, Dieffenb(lchi(l has several as exsi(catae, are stored separately, but tie(l to a 

staminodia surrounding the pistils and higher-order particular spee ies description and to the nomenclavenation 

parallel-pinnate. However, the monotypic tural information by a unique taxon number. Tergenus 

Bognera is endemie to Brazil, and it is not minology and usage in the descriptions in this relikely 

to be a problem with determination of most vision are largely defined by Croat and Bunting 

Dieffenbachia. Alternatively, DiefJenbachia is fre- (1979) and elaborated in Croat (1997). 

quently confused with Philodendron Schott, partic- Ecological zones, though sometimes estimated 

ularly the terrestrial species of the latter with non- from my own experience with Central American 

cordate blades. Philodendron species are mostly vegetation, are largely taken from Holdridge lifehemiepiphytic, 

rarely terrestrial as is Dieffenbachia. zone maps (Holdridge, 1967; Holdridge et al., 

Even when Philodendron species are terrestrial and 1971), where they exist for Central American counhave 

non-cordate blades, they can be distinguished tries, and for Mexico from the "Mapa de tipos de 

by the remotely many-flowered pistils that are not vegetacion de la Republica de Mexico" (Flores et 

surrounded by staminodia, as well as the lack of al., 1971). 

the acrid foul-smelling, irritating sap that is so Herbarium material has been widely distributed 

closely associated with Dieffenbachia. 

and original field vouchers are cited for all herbaria 

Species diversity of Dieffenbachia shows a gen- whose material was seen (see Appendices 1, 2). 

eral diminution from Mexico to Middle America Herbarium material may consist of one of three 

669

670 Annals of the 

Missouri Botanical Garden 

Table 1. Distribution and endemism in Dieffenbachia species from Mexico, Central America and the West Indies. 

X = species present, E = endemic species. 

Country 

Gua- E1 Ni- Col- 

Mexi- te- Salva- Hon- cara- Costa West om- Ecua- Vene- Boliv- 

Species co mala Belize dor duras gua Rica Panama Indies bia dor zuela Brazil ia 

D. aurantiaca X X 

D. beachiana X X 

D. burgeri E 

D. concinna X X 

D. copensis E 

D. crebripistillata E 

D. davidsei X X X 

D. fortunensis E 

D. fosteri E 

D. galdamesiae E 

D. grayumiana X X 

D. hammelii E 

D. horichii E 

D. isthmia X X 

D. killipii X X X X 

D. Iongispatha X X 

D. Iutheri E 

D. nitidipetiolata X X X X X 

D. obscurinervia X X 

D. oerstedii X X X X X X X X 

D. panamensis E 

D. pittieri E 

D. seguine X X X X X X 

D. standleyi X X 

D. tonduzii X X X X X 

D. wendlandii X X X X X X X 

Total number of 2 2 1 2 3 6 13 20 1 8 4 1 1 1 

species (percent 

of endemics) 

(23%) (40%) 

kinds: (1) complete original sets (wild collected); 

(2) sterile original material with an inflorescence 

added from a cultivated plant of the same number; 

and (3) material collected entirely from the cultivated 

plant. Specimens based entirely or in part on 

cultivated material are clearly indicated on the herbarium 

label. 

Herbarium specimens were borrowed from most 

major herbaria including: A, AAU, B, BBS, BM, 

BR, CAS, CAY, CHOCO, CM, COL, CR, CUVC, 

DAV, DS, DUKE, EAP, ECON, ENCB, F, FLAS, 

FSU, FTG, G, GH, GOET, GUAY, HBG, fIUA, INB, 

ISC, K, L, LA, LL, M, MEXU, MICH, MY, NY, P, 

computerized, and the entire collection, including 

many South American species that are being described 

separately, undergoes regular inventories to 

include events such as flowering time and attempted 

hybridizations. This allows electronic access to 

information on status, location, flowering dates, the 

number of vouchers prepared, and the status of the 

description. All specimens cited have been recorded 

in a computerized database for permanent referral. 

See www.mobot.org. 

HISTORY OF THE GENUS DIEFFENBACHIA 

HEINRICH WILHELM SCHOTT 

PMA, PORT, QAP, QCA, QCNE, RSA, S, SCZ, 

SEL, STRI, TEFH, TEX, TULV, U, UC, UMO, US, The genus Dietfienbachia was described in 1829 

USC, USJ, VEN, and WIS. Unless specifically des- by H. W. Schott (Schott, 1829). Schott based the 

ignated as not seen, all specimens are to be pre- genus on a single species, Caladium seguinum 

sumed to have been examined personally. (Jacq.) Vent. (Ventenat, 1800), previously described 

Data on the living collection of Dieffenbachia are as Arum seguine Jacq. (Jacquin, 1763) as well as


2004 

Table 2. Treatment of Dieffienbachia species by H.W. Schott (1860). 

Croat 


Species treated by H. W. Schott (1860) Origin Synonymizations as treated by Engler (1915) 

1. D. cognata Schott Suriname = D. seguine var. Iineata (Mart. ex Schott) Engl. 

2. D. consobrina Schott Brazil = D. seguine var. viridis Engl. 

3. D. costata H. Karst. ex Schott Venezuela 

4. D. gollmeriana Schott Venezuela = D. seguine var. viridis Engl. 

5. D. humilis Poepp. Peru 

6. D. irrorata Mart. ex Schott Brazil = D. seguine var. Iingulata (Mart.) Engl. subvar. 

irrorata (Mart.) Engl. 

7. D. Iineata K. Koch & Bouche New Granada = D. seguine var. Iineata (K. Koch & Bouche) 

Engl. 

8. D. Iingulata Mart. ex Schott Brazil = D. seguine var. Iingulata (Mart. ex Schott) Engl. 

9. D. Iiturata Schott Unknown = D. seguine var. Iiturata (Schott) Engl. 

10. D. macrophylla Poepp. Peru 

11. D. neglecta Schott Jamaica = D. seguine var. viridis Engl. 

12. D. obliqua Poepp. Peru 

13. D. oerstedii Schott Central America 

14. D. picta Schott Unknown 

15. D. plumieri Schott Martinique = D. seguine var. viridis Engl. 

16. D. poeppigii Schott Peru = D. seguine var. viridis Engl. 

17. D. robusta Schott Unknown 

18. D. seguine (Jacq.) Schott Martinique 

19. D. spruceana Schott Brazil = D. humilis Poepp. 

20. D. ventenatiana Schott Suriname = D. seguine var. ventenatiana (Schott) Engl. 

21. D. usendlandii Schott E1 Salvador = D. seguine 

by Linnaeus (1763). The next spe( ies des( ribed marllm (F,ngler, 1879),6 species were treated an(l 

were three from Peru (Poeppig 184tS) D. humilis 2 more c ombinations were made. Between 1879 

Poepp., D. macrophylla Poepp., an(l D. obliqzla and 1899, 14 a(lditional species were describe(l, 

Poepp. 

only 3 of which were recognized by Engler in 1915. 

Between 1852 and 1864 an additional 18 spe- These were D. flaguensis Engl., D. enfleri Engl. 

cies were described, all but 4 of them by H. W and D. olbia L. I,inden 8 Rodigas. 

Schott. Of these only 3 were recognize(l in Engler's In 1899, Engler produced another small work on 

(1915) revision, while the remainder were synony- Dieffenbachia in which he treated 19 species and 

mized or recognized at the subspecific level, mostly made 24 new combinations, but provided no deunder 

Dieffenbachia seguine. Those recognized at scriptions except for one new species, Dieffenbachthe 

specific level were D. picta Schott, D. costata ia aurantiaca Engl. Only 9 species were described 

H. Karst. ex Schott, and D. oerstedii Schott. Be- between this work (1899) and Engler's (1915) retween 

1866 and 1878,12 species were described, vision. Those taxa still recognized by Engler (1915) 

with 6 more of these being recognized as distinct were Dieffenbachia cordata Engl., D. weberbaueri 

in Engler's revision (Table 3). The first major re- Engl., D. gracilis Huber, and D. cannifolia Engl. 

vision of the genus was that of Schott (1860) in his ex Ule. 

Prodromus Systematis Aroidearum, treating 21 spe- The last revision of the genus Dieffenbachia was 

cies, of which Engler later (1915) recognized only by Engler (1915) in Das Pfanzenreich. This revi- 

7 (see Table 2). 

sion contained both keys and descriptions for 27 

species. A total of 11 new combinations were made 

ADOLF ENGLER 

and 6 new species described (see Table 3). The new 

species were D. aglaonematifolia Engl., D. brittonii 

The next revisionary effort made with Dieffen- 

Engl., D. Iongispatha Engl. 8 K. Krause, D. parbachia 

was by Engler (1878) in the treatment of the 

vifolia Engl., and D. pittieri Engl. 8 K. Krause. 

Araceae in Martius's Flora Brasiliensis. In this 

treatment 3 species were included and 16 new 

combinations made (these within D. seguine and D. 

MODERN WORK 

picta). A year later, in his treatment of Dieffen- Very little work was done with Dieffenbachia afbachia 

in DeCandolle's Monographiae Phaneroga- ter Engler (1915). Gleason (1929) described D. pal- 

671

672 

Table 3. Treatment of Dieffienbachia species by A. Engler (1915). 

Annals of the 


Species treated by A. Engler (1915) Origin As treated in this manuscript 

1. D. aglaonematifolia Engl. 

Paraguay 

2. D. antioquensis Linden & Andre 

Colombia 

3. D. aurantiaca Engl. 

Costa Rica D. aurantiaca Engl. 

4. D. bowmannii Carr. 

Colombia & Brazil 

5. D. brittonii Engl. 

Colombia 

6. D. cannifolia Engl. 

Peru 

7. D. cordata Engl. 

Peru 

8. D. costata Klotzsch 

Colombia & Peru 

9. D. daguensis Engl. 

Colombia 

10. D. enderi Engl. 

Colombia 

11. D. gracilis Huber 

Peru 

12. D. humilis Poepp. 

Brazil & Peru 

13. D. imperialis Linden & Andre 

Peru 

14. D. Iatimaculata Linden & Andre 

Colombia 

15. D. Ieopoldii Bull 

Costa Rica 

16. D. Iongispatha Engl. & K. Krause 

Panama 

D. loslgispatha Engl. & K. Krause 

17. D. macrophylla l'oepp. 

Peru 

18. D. obliqua Poepl). 

Peru 

19. D. oerstedii Schott 

Central America D. oe r.stedii Schott 

20. D. olbia J. Linden & Rodigas 

Peru 

21. D. parlatorei Lillde.n & Andre 

Colombia 

D. parlatorei val. m.rmorea Linden & Andre Unknown 

22. D. pflrvifolia Engl. 

Brazil 

23. D. picta (Lodd.) S( hott 

Brazil? 

D. seguine (Jacq.) Schott 

D. picta var. an.gll.sl.ior Engl. 

Unknown 


D. picta var. allgll..sllor subvar. angustifolia 

Engl. 

Unknown 


D. picta var. angll sl ior subvar. jenmanii 

(Veitch) Engl. 

Unknown 


D. picta var. angustior subvar. Iancifolia (Linden 

& Andre) Engl. 

Unknown 


D. picta var. angustior subvar. schuttlesworthi- Unknown 

ana (Hort. Bull.) Engl. 


D. picta var. barraquiniana (Verschaff. & 

Lem.) Engl. 

Unknown 


D. picta var. Iatior Engl. 

Unknown 


D. picta var. Iatior subvar. amoena Hort. Bull. Unknown 


D. picta var. Iatior subvar. carderi Hort. Bull. Unknown 


D. picta var. Iatior subvar. gigantea (Verschaff.) 

Engl. 

Brazil 


D. picta var. Iatior subvar. magniJica (Linden 

& Rodigas) Engl. 

Venezuela 


D. picta var. Iatior subvar. meleagris (Linden 


Ecuador 

D. srleleagris L. Linden & Rodigas 

D. picta var. memoria (Corsi Salviati) Engl. Unknown 


D. picta var. Iatior subvar. mirabilis (Verschaff.) 

Engl. 

Unknown 

D. sequine (Jacq.) Schott 

D. picta var. Iatior subvar. picturata (Linden 


Venezuela 


D. picta var. typica Engl. 

Unknown 


24. D. pittieri Engl. & K. Krause 

Panama 

D. pittieri Engl. & K. Krause 

25. D. seguine (L.) Schott 

= D. seguine (Jacq.) Schott 

Martinique D. seguine (Jacq.) Schott 

D. seguine var. decora (Hort. Vershaff.) Engl. Brazil? 


D. seguine var. Iineata (K. Koch & Bouche) 

Engl. 

Venezuela 

D. seguine (Jacq.) Schott


2004 

Table 3. Continued. 

Species treated by A. Engler (1915) 

D. seguine var. Iingulata (Martius) Engl. 

D. seguine var. Iingulata subvar. irrorata 

(Mart.) Engl. 

D. seguine var. Iiturata (Schott) Engl. 

D. seguine var. Iiturata subvar. wallisii (Linden) 

Engl. 

D. seguine var. nobilis (Hort. Verschaff.) Engl. 

D. seguine var. robusta (K. Koch) Engl. 

D. seguine var. ventenatiana (Schott) Engl. 

D. seguine var. zziridis Engl. 

26 D weberbaueri Engl 

27 D weirii Berkl 

. 

Croat 


Origin 

Suriname & Brazil 

Brazil 

Unknown 

Unknown 

Brazil 

Unknown 

Murlname 

Widespread in West 

Indies, Central and 

South America 

Peru 

Colombia 

As treated in this manuscript 



D. seguine (Jacq,) Schott 

D seguine (Jacq,) Schott 





udicola N. E. Br. ex Gleason from Guayana, and inaccuracy was in part due to a poor understanding 

Jonker and Jonker (1966) described D. elegans A. of the collections that existed at the time, but also 

M. E. Jonker 8 Jonker for Suriname. Three species to the fact that most collections of Araveae have 

were described from Venezuela, D. 1:volivarana G. been made in the last 25 years. Panama, for ex- 

S. Bunting (Bunting, 1963), D. Iiesneri Croat, and ample, has shown vast increases in the number of 

D. Iongil)istil(l Croat (Croat 8 Lambert, 1987). An- spevies eollected and identified sinc e the treatment 

other Venezuelan spee ies, D. duiflae (Steyerm.) G. of Araveae in the Flor(l oJ P(ln(lm(l. Some genera 

S. Bunting was transferle(l (Bunting, 19238) frorr have inereased t)y nearly 544Wo sinee 1944 (Croat, 

S)athis ar/)(l (lui(l(le Steyerm., (les( rit)e(l in 1 951 . L985). I)iefRnl)(l(/li(l now has 2G sl)e( ies re( ogl)iefienl)(lchi(l 

has been so e onfusing that floristi nize(l for Central Ameri( a with 77% of them new 

ae(ounts (lealing with Central Ameri( a have })eell to seien( e sinee work on the revisioll })egan. 'l'his 

of little value. Herllsley (188tS) liste(l only a single re^)resents a 40()z7So inelease from the nlln-l})er lespe(ies, 

D. oer.ste(lii S(hott, fol his Hiologi(l Cell- I)orte(l in Central Amerie a ly )revious workers (Entr(lli-Americ(lrl(l. 

Engler (191tS) rel)orte(l only five gler 1915; Stan(lley, 1937, 1944; Standley d Steyspe(ies 

for (3entral Ameriva, D. oerste(lii, D. (IU- ermark, 1958; Matuda, :1954; Bunting, 19G5). 

r(lntiac(l Ellgl., D. /)ittieri, D. Iongi.s/)ath(ln D. se- Standley's treatment of the Araveae of the Languine, 

an(l D. leo/)ol(lii Bull. The first four were cetilla Va11ey (Stan(11ey, 1931) deait with only 2 

properly eir(ums(ril)e(l largely owing to the faet spee ies, D. oerstedii, which was at least in part corthat 

he was dealing with little beyond the type rectly determined, and D. seguine, 1)robably D. 

speeimens. Dieffenl)achia leopoldii has proven to be standleyi Croat in the present treatment. However, 

a species known only from Colombia. The only in the case of the latter species he did not mention 

plants Engler (1915) called D. seguine were from the almost fully sheathed petiole that is so distinc- 

E1 Salvador the type country of D. wendlandii, tive on D. standleyi. Standley also confused D. sewhich 

he erroneously synonymized with D. seguine. guin.e with other species, mentioning that it ranged 

Dieffenbachia wendlandii differs from D. seguine in to E1 Salvador (probably D. wendlandii) and to the 

having unilocular ovaries, and a long-tapered spa- West Indies (referring to the real D. seguine). 

dix that does not protrude forward at anthesis, as The treatment of the Araceae in the Flora of 

is the case with D. seguine. 

Guatemala (Standley & Steyermark, 1958) is es- 

Matuda (1954) reported only D. seguine for Mex- pecially confusing. In this treatment they dealt with 

ico, whereas Bunting (1965) reported both D. se- four names, D. oerstedii, D. picta (Lodd.) Schott, a 

guine and D. oerstedii for Mexico, the latter being maculate species that I consider synonymous with 

a new report. One of the species that Bunting was D. seguine and not occurring in Central America, 

dealing with, which he named as D. seguine, is rec- and D. pittieri, which I consider to be a narrow 

ognized herein as D. wendlandii. 

endemic in the Canal Zone of Panama. Since they 

These earlier floristic accounts for Central Amer- did not cite specimens, it is difficult to interpret 

ica, in addition to being inaccurate, also grossly their treatment, but their description of D. pittieri 

undercounted the number of existing species. This with a petiole sheathed almost to the apex and 

673

674 

Annals of the 


ranging from Guatemala to Honduras may be what relatively few botanists made many collections of 

I am currently calling D. standleyi. The plant they Araceae. Mike Grayum, also an aroid specialist, 

were calling D. oerstedii probably was, at least in made 68 collections of Diegenbachia in Central 

part, the species named. The plant they called D. America, mostly Costa Rica, out of a total of 73 

seguine was probably D. wendlandii. The mottled Dieffenbachia collections for that country. Barry 

plants they referred to as D. picta may have been Hammel a close associate of Grayum and myself, 

cultivated forms of D. seguine, the mottled forms of made 57 collections of Dieffenbachia in Central 

which are popular with horticulturists and have America. Paul Standley, one of the most ambitious 

been known to persist in areas of abandoned dwell- collectors in the Neotropics made 54 collections. 

lngs. 

Julian Steyermark, another giant in the field of col- 

The Flora of Costa Rica (Standley, 1937) treat- lecting, made 46 collections, but only 13 of these 

ment of DieJ7enbachia was somewhat less confused, were in Central America (Guatemala), whereas the 

probably because it was adapted from Engler's balance were from Venezuela. A1 Gentry also col- 

(1911) treatment. Because several of the species lected a lot of Dieffenbachia, 57 in all, but even 

types came from Costa Rica, fewer names were mis- fewer were from Central America (only 5 collecapplied. 

Here Standley treated D. aurantiaca, an tions). 

endemic to southwestern Costa Rica, D. Ieopoldii The earliest botanist to collect Central American 

(a Colombian species not believed to be in Central species of Dieffenbachia was Friedrich Carl Leh- 

America and perhaps confused with what I am call- mann over a century ago. These were D. killipii 

ing D. killipii in the present treatment), D. pittieri, (Lehmann 5311) and D. tonduzii (I,ehmann 8876), 

and D. seguine. Unfortunately, he cited no speci- the latter as early as 1819. Herman Wendland colmens 

nor gave any characters by which to recognize lected D. wendlandii, in this case l)efore 1858. Oththe 

species. Certainly, no material currently exists er collectors included Adolfe Ton(luz (D. aurantiain 

Costa Rica of D. pittieri or D. seguineX ca) and Henri Pittier, the first to collect D. 

In the Flora of Panama, Standley (1944) did cite Iongispatha (Pittier 2715), D. ob.se llrinervia (Pittier 

specimens for four species. DieJ7enbachia pittieri is 3766), D. pittieri (Pittier 3766), all(l D. killipii (Pitendemic 

to the Isthmus of Panama. Standley (1944) tier 2600), all collected as early (1.s l 911; Paul Stancited 

the type, Pittier 3766, but he also cited ma- dley, who was the first to collee t 1). concinna (Stanterial 

from E1 Valle (Alston & Allen 1839) that rep- dley 36739), D. isthmia (Stan(tle) 29867), and D. 

resents D. crebripistillata. Diegenbachia aurantiaca beechiana (Standley 36840), all in 1924; Paul Allen 

was even more confused, with only one of the cited initially collected D. crebripistill(lt(l (Allen 1839) in 

collections, Woodson & Schery 861, being D. au- 1939; E. P. Killip, D. nitidipetiolal(l (Killip 35113) 

rantiaca. Other collections cited assign to three dif- in 1939; and Louis O. Williams 1). horichii (Wilferent 

species, D. isthmia, D. killipii, and D. ton- liams 28479) in 1965. 

duzii. A total of five species were represented Other Central American species were collected 

among the 12 collections cited as D. oerstedii, only only in the past few decades. DieJ7enbachia burgeri 

one of which actually was that species (Pittier 2836 dates to William Burger and Ronald Liesner (Burin 

western Panama). Other specimens cited under ger & Liesner 7254) in 1970, anel D. davidsei was 

this name belong to D. isthmia, D. killipii, D. ni- not seen until Scott Mori made his collection in 

tidipetiolata, and D. obscurinervia. 

1974 (Mori et al. 4184). Tom Croat encountered 

Dieffenbachia fortunensis in 1976 (Croat 37268), D. 

COLLECTING HISTORY OF DIEFFENBACHIA galdamesiae in 1979 (Croat 49124), and D. panamensis 

in 1974 (Croat 27206). Diegenbachia gra- 

Collecting Araceae has not been particularly yumiana was first noticed by Burger and Matta in 

popular, and this would be especially true for Dief- 1967 (Burger & Matta 4181), D. fosteri by Robin 

fenbachia. Many collectors who have been careless Foster in 1993 (Foster et al. 14649), and D. hamwhen 

collecting Dieffienbachia found themselves melii by David Neill in 1978 (Neill & Vincelli 

badly burned with the often high concentrations of 

oxalic acid in the cut parts. Perhaps this has led 

3484). 

to the generally low numbers of collections of Dief- SUPRAGENERIC CLASSIFICATION OF SUBFAMILY 

fenbachia. A survey of those who collected in Cen- PHILODENDROIDEAE AND RELATIONSHIPS 

tral America, based on the extensive TROPICOS OF DIEFFENBACHIA 

database of collections, shows that aside from myself 

(338 Croat collections for Central America out Engler (1915) placed Dieffenbachia in the subof 

a total of 537 Croat collections in the Neotropics) family Philodendroideae Engl. The same position is


2004 

Croat 


taken by Bogner and Nicolson (1991) in their mod- chieae with two genera (DieJ7enbachia and Bogern 

revision. Grayum (1984, 1990) placed Dieffen- nera) and the tribe Spathicarpeae with eight genera, 

bachia in its own tribe and does not disagree with Mangonia, Taccarum, Asterostigma, Gorgonidium, 

Bogner and Nicolson about its relative placement Synandrospadix, Gearum, Spathantheum, and 

or taxonomic status. However, Grayum substantially Spathicarpa. Their tribe Spathicarpeae differs from 

modified the Philodendroideae by including the ge- the tribe Dieffenbachieae in having tuberous stems 

nus Calla and thus mandated a change in the name and leaves with reticulate higher-order venation. 

of the subfamily to Calloideae Endl. on the basis However, the latter character is also present in 

of priority. This group, as defined by Grayum, is Bognera, one of the two genera in their tribe Diefsubstantially 

larger than that of Bogner and Nic- fenbachieae. In the Dieffenbachia alliance Dieffenolson's, 

with 40 genera arranged in 17 tribes in five bachieae are distinguished from other tribes by 

alliances. This contrasts with 18 genera in 7 tribes having anatropous ovules; the primary lateral veins 

according to Bogner and Nicolson. The latter sys- forming a single marginal vein but also lacking coltem 

closely mirrors Engler's Philodendroideae, ex- lective veins; and, perhaps most importantly, in 

cept for the inclusion of new genera and the syn- having the pistillate portion of the spadix completeonymizing 

of others since the time of Engler. The ly fused to the spathe. Tribe Dieffenbachieae sensu 

size of Grayum's Philodendroideae resulted from Mayo et al. (1997) is distinguished from other tribes 

the inclusion of tribes from the subfamilies Aro- of subfamily Philodendroideae by having stamens 

ideae Adans., Pothoideae Engl., and Lasioideae connate into synandria, and in having several free 

Engl. (Croat, 1990 [1992]). 

staminodia associated with the female flowers. Oth- 

In recent suprageneric classifications by Mayo et er useful distinguishing features include a conal. 

(1997, 1998) all the genera with unisexual flow- stricted spathe partly adnate with the pistillate spaers 

were grouped together in the subfamily Aro- dix, terrestrial caulescent habit, closely parallel 

ideae and further divided into non-ranked groups, primary and minor veins, and seeds with endothe 

perigoniate Aroideae and aperigoniate Aro- sperm. 

ideae. Dieffenbachia, in the latter group, was placed The Dieffenbachia alliance of Mayo et al (1997) 

in the Dieffenbachia alliance, one of four alliances has been supported by molecular studies with chlo- 

(the others called Philodendron, Schismatoglottis, roplast trnI,-F genes (Barabe; Tam et al, 2004) 

and Caladium alliances respectively). These alli- This study shows close relationships between 

ances comprise 33 genera with an additional 37 Spathicarpeae and Dieffenbachieae with Dieffengenera 

being placed in a group labeled "No alli- bachia being paired with Spathicarpa on one 

ance." The classification by Mayo et al. was able branch and with Cerce.sti.s Schott, Rhektophyllum N. 

to make use of the results of the recently completed E. Br., and Culcasia P. Beauv. forming another 

chloroplast DNA studies and anatomical studies of branch of the tree. 

French and colleagues (1995, 1997). DieJ7enbachia, Although both Grayum (1984, 1990) and Bogner 

along with Bognera, was placed in the tribe Dief- and Nicolson (1991) placed Dieffenbachia in its 

fenbachieae, a tribe characterized by having the own tribe, the authors of both systems now agree 

pistillate flowers completely adnate to the spathe. that Bognera, another Neotropical genus, is very 

In the most recent suprageneric classification of close to or within the tribe Dieffenbachieae, sensu 

the family by Keating (2002, 2003a) Dieffienbachia Mayo et al. (1997). Keating (2002, 2003a) simply 

is included in subfamily Philodendroideae, tribe subsumed this tribe (or its genera) within his larger 

Spathicarpeae Schott, along with Bognera, Spa- Spathicarpeae, still in essential agreernent. 

thantheum Schott, Gorgonidium Schott, Synandros- The subfamily Philodendroideae is worldwide in 

padix Engl., Gearum N. E. Br., Spathicarpa Hook., distribution and, sensu Grayum (pers. comm.), it is 

Asterostigma Fisch. & C. A. Mey., Mangonia somewhat equally divided between the Old World 

Schott, and Taccarum Brongn. ex Schott. Keating's and the New World, with 19 genera (17 of them 

revision makes use of a wealth of vegetative ana- endemic) and 761 species in the New World and 

tomical information he has accumulated on the 23 (21 endemic) genera and 354 species in the Old 

family. All members of the Spathicarpeae share in World. Grayum places the Dieffenbachieae in his 

common Type B collenchyma; vascular bundle type Aglaonema alliance with Zantedeschieae Engl. 

II (or occasionally types I or III); and either lack (Zantedeschia Spreng.), Aglaonemateae Engl. 

laticifers or have non-anastomosing type laticifers. (Aglaonema Schott, Aglaodorum Schott), Spathicar- 

The Dieffenbachia alliance of Mayo et al. (1997) peae (Mangonia, Asterostigma, Synandrospadix, 

is distinguished by having simple laticifers and free Taccarum, Gorgonidium, Gearum, Spathanthemum, 

synandria. It contains two tribes, the Dieffenba- Spathicarpa), and Bognereae S. Mayo & D. Nicol- 

675



son (Bognera). Anubiadeae Engl. (1876) (Anubias more foul-scented and caustic than many other spe- 

Schott) and Zantedeschieae are restricted to Africa, cies, and may in part account for less breakage by 

while the Spathicarpeae are restricted to southern deterring animals from the plant. 

South America. 

In some cases DieJ7enbachia may grow in standing 

water. In such cases the stems may be rather 

MORPHOLOGY OF VEGETATIVE STRUCTURES deeply buried in the mud. Some species, such as 

IIABIrF AND GROWTH PATTERNS 

D. grayumiana, are frequent in wet habitats, but 

most species thrive in well-drained soils. The genus 

Dieffenbachia is always terrestrial and caules- is seldom found on road embankments, a habitat 

cent (Croat, 1988 [1990]). In terms of growth be- that is very common for species of Anthurium and 

havior and habit, Dieffenbachia is not as variable Philodendron. This is perhaps because the two latas 

Philodendron, a related genus, but sterile plants ter, principally hemi-epiphytic genera can more 

disassociated with notes about habit can be con- easily become estal)lished on the excessively wellfused 

with those of Philodendron. Stems of Dieffen- drained and poor clay soils of typical road embachia 

are erect, at least on the younger portions, bankments in the Neotropics. 

with older portions of the stem typically becoming Development in Dieffenbachia is never heterodecumbent. 

This creeping portion of the stem is blastic. Instead, changes in internode size and leaf 

sometimes even longer than the erect portion (Fig. size progress without any marked changes in blade 

23A). The overall height of any species is usually shape. However, leaf blades of Dieffenbachia that 

determined by the thickness, strength, and rigidity ultimately become ( ordate at the base are at first 

of the stem. Typically, species such as D. burgeri, simple and acute to obtuse at the base. 

D. hammelii, and most plants of D. oerstedii, with 

small-diameter stems, do not get to more than 1 m 

tall nor do they usually have stems more than 2.5 

STEMS 

cm in diameter. Alternatively, the taller species like Internodes are ty)ically about as long as broad 

D. horichii, D. Ion;,Xisl)atha, and D. standleyi have or even shorter tl(l] broad. Sometimes, as in the 

thick stems and reach heights of 2 to 3.5 m. Cul- case of D. galdan7(.siae, petioles are affixed to the 

tivated plants growing in a pot and unable to be- stem at an oblique tlngle and the internodes are not 

come reclined at the base can grow to indefinite of equal width acloss the diameter of the stem. Inheight. 

A plant of D. standleyi cultivated in the stead, one side is as much as 1 cm wider than the 

greenhouse of the Missouri Botanical Garden grew other. Most commolllyX internodes are glossy to seup 

the side of a wall from a pot to the height of 8 miglossy and smooth though they may be minutely 

m. 

roughened, appearing with a somewhat weak vel- 

The portion of the stem that comes in contact vety sheen, as in tlle case of D. oerstedii. Though 

with the ground becomes rooted at the nodes, but for most species tlle internodes remain moderately 

stems are rarely buried. Instead, the stems usually glossy even in age, when fresh, the internodes may 

creep over the surface of the ground. The growth change promptly. 1 n D. obscurinervia, the stem, 

behavior often leads to vegetative reproduction, though initially semiglossy, promptly becomes 

since on some species the creeping portion of the etched in a deep, areolate pattern to such an extent 

stem tends to produce active branch buds that form that the stem becomes matte and is conspicuously 

new growth. Another feature that tends to induce scurfy. 

additional branching is the often fragile stem ex- Cataphylls are never present on Dieffenbachia. 

tending laterally over the ground, which may be Instead, the new growth on stems of Dieffenbachia 

broken by being walked on by animals. Broken is protected by the sheathing petiole of the precedstems 

invariably produce new branches. These re- ing leaf. 

sults often cause Dieffenbachia to grow in large col- Stem color varies considerably between species 

onies, especially in open, better illuminated areas and even within populations of the same species. 

of the understory of a forest, along stream banks or Species such as D. Iongispatha, which typically 

in open swamps. Examples of colonial growth occur possess stems that are solid green, can sometimes 

in D. crebripistillata, D. isthmia, D. killipii, and D. be variegated with paler colors. Typically, stem col- 

Oerstedii. Some species, such as the taller D. Ion- or variegation is in the form of streaks rather than 

gispatha (to 3.5 m), are usually less colonial. This mottling. The stem may be relatively dark green 

is perhaps because it is a species with a large, stout with even darker streaks, in which case the motstem 

that is less likely to be broken up by the el- tling would not be too apparent, or, as is more freements. 

This species also has a thicker sap that is quently the case, it may be streaked with pale


2004 

%()111P .S[)e('itX,% (1). (/(lVi(lss(i, 1). ,s(^grilitl(') llclVC tilC [)e.t- 

iole sheclll free-en(lillg (K'igS. t

678 

Annals of the 


are solid green on both surfaces. There are consid- with the staminate flowers at the apex and the piserable 

differences in the nllmler of inflorescences tillate flowers at the base. The spadix of Dieffenper 

axiln ranging from solitary on a number of spe- bachia is usually shorter than the spathe by 1-3 

cies at least part of the time or up to eight per axil cm (Fig. 2B). The spadix is contained within the 

as with D. concinna. In any population the number spathe, and is more or less straight or weakly 

of inflorescences per axil may vary. Studies made curved (Fig. 8F) at anthesis, rather than being 

with D. oerstedii in Costa Rica (Valerio, 1984) prominently protruded forward. Some species have 

showed that this species produced primarily one or the spadix protruded forward at anthesis, such as 

two inflorescences per axil. 

D. burgeri (Fig. 4C), D. crebripistillata (Fig. 7C), D. 

horichii (Fig. 13E), D. wendlandii (Fig. 26C), and 

SIATHE 

D. seguine (Fig. 23E). The last species, which ranges 

throughout the West Indies, has a spadix that 

Each spathe in a cluster of inflorescences in 

notably protrudes and is cylindrical, rather than ta- 

Dieffenbachia is subtended by linear-lanceolate 

pered to the apex as in the remaining Central 

bracteoles, each of which are unribbed. The brac- 

American species (Fig. 23F). Diefienbachia seguine 

teoles are typie ally about as long as the peduncles, 

also has a spadix (Fig. 23) that is caught in the 

but can sometimes be shorter or longer. They are 

protruded position when the spathe re-closes. In 

somewhat mar( ese ent. 

contrast to D. seguine, all of the (3entral American 

Spathes of llieffenbachia are uniforrnly green, 

species have spathes that close normally with the 

though the inner surface tends to be somewhat palspadix 

withdrawing inside of the spathe. 

er than the outel surface and frequently much 

Like other members of the Philo(lendroideae, the 

glossier. The OUtUl spathe surface is typically sespadix 

in Dieffenbachia is divide(l up into a fertile 

miglossy to weakly glossy. Sometimes the inner surstarninate 

section at the apex, a sterile section lying 

face of the spathe is somewhat whitish on the inside 

beneath the fertile staminate )oltionX and the pisnear 

the tip. 

tillate portion (Figs. 2B, 7, 18! 2?.5). The lowermost 

The spathe of Illost Diefenbachia is only weakly 

section, as with all Araceae witll llllisexual flowers, 

constricted alsove the spathe tube (Figs. 7A, 16D, 

is the pistillate section. Typie ally lhe pistillate por- 

20F, 22E, 2^-3kJ! 2zlt 25F). When the entire spathe 

tion and the staminate portion (lle of roughly equal 

is fully flattene(l, lhe tube portion is considerably 

lengths, but the length of the stex ile eiegment (when 

wider (Figs. 21S, I SH, 22F, 23F) and the spathe is 

apparent) is highly variable lsolll ill terms of length 

gradually tapere(l from the lower 1/4 to the usually 

and the degree to which stamino(lia and pistillodes 

acuminate apex (Fig. 23F). Sometimes there is a 

are dispersed on it (Figs. 2B, 4(3! .5l), 7, 15H, 18D, 

weak central (onstriction. The point of maximum 

23F, 25G). In most cases the [)istillate region is 

c onstriction c orl esponds to the sterile section of the 

slightly longer than the staminclte. 

spadix betweell the fertile staminate flowers and the 

The fertile staminate portion is somewhat cylinpistillate 

flowers. 

droid to spindle-shaped, broa(lest ill the middle and 

At anthesis tle spathe of Diefenbachia is open 

weakly tapered toward both en(ts, being bluntly 

to about the middle (Figs. 5B, 7A, 13E, 16D), or 

rounded at the apex. The fertile rllale flowers are 

somewhat below the middle, lgut does not open as 

compacted and sub-rounded or with angular marwidely 

as that of Philodendron. At anthesis the 

gins (Figs 2E, 5G). As the spa(lix approaches anspathe 

blade is wider than the tube. Typically only 

thesis the synandria loosen ancl the anthers become 

the staminate portion of the spadix is visible at anvisible 

between the synandria. 'roward the base of 

thesis but sometimes, as in D. crebripistillata, a porthe 

staminate portion of the spa(lix, the flowers are 

tion of the pistillate spadix is also exposed. Closure 

somewhat more irregular in size and shapen but do 

of the spathe after anthesis is sometimes not comnot 

become radically different as in the case of 

plete, with the margins of the spathe meeting irreg- 

Philodendron and Xanthosoma, where the lowerularly. 

At anthesis the blade of the spathe is usually 

most male flowers are sterile and swollen. In conarched 

somewhat forward, hooding the spadix 

trast, Dieftenbachia relies entirely on club-shaped 

slightly. For some species, such as D. crebripistilstaminodia 

surrounding the pistils (Figs. 5E, 14C). 

lata, the spathe may be recurled at anthesis (Fig. 

The lowermost staminate flowers are smaller or very 

7C). 

irregular in shape, sometimes showing signs of two 

SPADIX 

or more flowers fused together. Sometimes the uppermost 

pistils are reduced, apparently sterile, and 

Flowers of Dieffienbachia are unisexual and na- surrounded by sterile male flowers (Fig. 15H). Apked, 

dispersed on the spadix- in the typical manner parently the lowermost staminate flowers are also


2004 

Croat 


more attractive to beetle pollinators, since only the apparently absent, with the fertile staminate and 

lowermost flowers are eaten (Fig. 1B). 

fertile pistillate portions essentially contiguous. The 

length of the sterile portion of the spadix is here 

ANDROECIUM 

defined as that portion which lies between the lowermost 

functional androecium and the uppermost 

Staminate flowers consist of 4 stamens united functional pistil regardless of the presence of pisinto 

a 4- or 5-sulcate truncate synandrium (Figs. tillodes and staminodes along its length. However, 

2E, 5G, 23G, 26E). The apex of the synandrium the sterile portion of the spadix is usually quite 

often has a minute slit or an equidistant series of obvious since a significant proportion of the sterile 

3 minute slits connected at the center. Upon drying section is devoid of flowers. The axis of the sterile 

the apex of the synandrium may become wrinkled portion is typically convex as if the terete axis of 

along the margin and may also have a concave sur- the spadix (now fused) was half sunken into the 

face. The anthers are contiguous or nearly so, af- leafy tissue of the spathe. 

fixed near the upper edge of the synandrium. The 

thecae are obovoid to cylindroid, opening by apical 

PISTII,I,ATE SPADIX 

slits just below the upper edge of the synandrium. 

The pollen is dispersed in slender, subterete filaments 

and typically projects up to 1 cm above the 

surface of the synandrium. When the stamens are 

at anthesis, the shedding pollen can completely fill 

the now closing spathe (Fig. 5F). 

The pistillate portion of the spadix is fused 

throughout its entire length to the spathe. What appears 

to be a stipe is readily apparent at the base. 

The pistillate flowers rarely extend very close to the 

base of the spathe (Fig. 4C). After the presence or 

absence of a sterile segment in the spadix, the 

POLI,EN 

number and disposition of the pistils is perhaps the 

Pollen of Dieffenbachia is released in monads most important taxonomi( character in the infloresthat 

are inaperturate, subisopolar to virtually polar, cence. Unlike many genera in the Philodendroi(leae 

boat-shaped-ellipti( to oblong, or nearly spherie al. that have the pistillate flowers closely aggregate(l 

They are bilaterally symmetrie al or radiosymmetrie on the spa(lix, the pistillate flowers of DiefJenbachia 

(Grayum, 1992). 

are moderately dispersed on the spadix. The degree 

Dieffenbachia pollen is moderately large for Ar- of (lispersal on the spadix is in itself different from 

aveae, ranging from 54 to 99 1lm (as in D. oerstedii) spee ies to spee ies. There are generally only 2 or 3 

and avelaging 79 11m. Exine sculpturing is psilate 

pistils aeross the width of the spadix axis (regardto 

obs(urely verrueulate (as in D. oer.steflii) an(l/or 

less of whether they are lined up or not). Rarely, 

sparingly punctate-foveate to densely foveate with as in the ease of D. killivii, there up to 6 pistils 

scattered compound foveolae (as in D. pittieri and visible across the width of the spadix. At the op- 

D. seguine). Grayum (1992) pointed out that the posite extreme, D. Iongispatha sometimes has a solcompound 

foveolae found on the pollen of D. pittieri itary row of pistils across the spadix axis. The arand 

D. seguine resemble those of Chlorospatha 

rangement of the pistils is generally quite irregular, 

croatiana Grayum. 

lacking any obvious equidistant spacing or alignment 

in rows, but sometimes a series of pistils 

STERILE SPADIX SECTION 

might be arranged in a loose row (as in D. killipii) 

or even in a broad arc across the width of the spa- 

Dieffienbachia typically has a nearly barren sec- dix. 

tion of the spadix lying between the fertile stami- Pistils are 2- or 3-carpellate, sometimes l-carnate 

portion at the apex and the pistillate portion pellate, sessile, depressed-globose to depressedat 

the base (Figs. 5D, 23F). This section occurs in ovoid, pale green, semiglossy, and smooth or 2- or 

the area where the spadix first becomes free from 3-lobate. The stigma is a 2- or 3-lobate cushionthe 

spathe. Only rarely is the transition between like layer that covers the entire apex of the pistil. 

the fertile staminate and the sterile }ortion of the The stigmatic papillae are orange or yellow. The 

spadix clearly defined with the fertile flowers papillae are close, dense, and moderately short, 

abruptly ending on a clear flowerless sterile portion. similar to those of Philodendron, but appear less 

Instead, there is usually an assortment of sterile interspersed with the gelatinous matrix that is so 

male flowers toward the apex of the sterile segment common on the stigmas of that genus. The stylar 

(Fig. 7E). Less frequently there are pistillodes in region is inconspicuous. With the stigma removed, 

the lower half of the sterile segment. In D. beachi- the surface is truncate or broadly sulcate with a 

ana and D. killipii (Fig. 15H) the sterile section is solitary medial or near-medial pore. Ovules are 

679

. . . n . . . 



anatropous, one per locule. Each pistil is surround- were sterile, varying from 1 to 17 per infructesced 

by 4 or 5 claviform, fleshy, white staminodia. ence. 

The staminodia may be fused somewhat at the base, Germination of seeds is usually prompt if the 

often forming a cupuliform structure around the mesocarp is first removed. Valerio (1984) found that 

base of the pistil. Individual staminodia are sub- naked seeds of cleaned berries of D. oerstedii began 

cylindrical, commonly somewhat flattened toward to germlnate wltnln I our days ln a germlnatlon 

the base and usually enlarged, sometimes almost chamber. The percentage of seeds germinating 

subglobular at the apex. Staminodia typically are ranged from 77.3Wo to 100% for different infrucerect-spreading 

or erect, vary considerably in tescences (averaging 91.7Wo). However, berries left 

length, but are almost invariably longer than the intact usually did not germinate, but remainded vipistils, 

usually 2-5 mm long, and held well above able for up to 90 days. 

the pistils. 

Valerio (1984) also reported that although 82Wo 

of the 729 berries studie(l were green, the remain- 

MORPHOLOG Y () F1 FRUITING STRUCTURES der were red or yellowish. All seeds produced only 

eN( l X AND FRUIT 

albino plants. Germinatiorl lates for both the green 

seeds and the abnormal seeds were the same, but 

While the intlorescences are always first erect, the albino plants perishe(l. His studies with seed 

they quickly be(ome reflexed after anthesis. This germination of D. oersteelii, as well as observations 

is probably important to successful development of of other plants in the fiel(l le(l him to conclude that 

the infructescen(e as it prevents the spathe from plants of this species re(uile(l between five and six 

filling with rainwater and thus possible decay. years to reach reproductivf age. 

Fruits develop within the re-closed spathe, which Fruit maturation is nollally a slow process in 

often turns yellow, pale to bright orange, or red as Dieffenbachia, requiring ll) to nine months in the 

fruits begin to nlature. The spathe enlarges some- case of D. nitidipetiolat(l (Young, 1986) (reported 

what during the (ourse of maturation, an(l finally as D. Iongispatha). Valf gio (1984) reported that 

begins to reflex along the middle, especially the fruit maturation in D. *ser.st(elii was approximately 

portion that is fsed to the spadix. In the l)rocess one year. Valerio (1984) sllgt,ested that the seeds 

most of the margins of the spathe slough off, leaving of Dieffenbachia are l)i(l-(lispersed because the 

just the fruiting spadix. The spadix remains arched seeds will not germinate [Xless the fruit mesocarp 

backward with lhe bright red to orange-re(l berries is first removed. At mallll ily the mesocarp of Diefheld 

somewhat apart by the recurving proe ess and fenbachia berries is pasty (l](l somewhat sweet. The 

displayed against the generally pale remains of the fruits would appear to l)e lllo;t suitable to bird disspathe 

(Figs. 1D, 4D). 

persal, being colorful a](l 11aving only a thin layer 

Although fluit develops with self pollination, of edible portion availal)le. 'I'he seeds are soft and 

fruit and seed size are greater when the plants are could easily be destroye(l l)y (hewing, so effective 

out-crossed (Young, 1986). Since Dieffenbachia animal dispersers are plol)al)ly just removing the 

flowers are nake(3 all protection of the flowers must pericarp and thin mesocal ). F ruits of DielJenbachia 

be provided hy the spathe. In the manner of many have been observed bei], eaten by white-faced 

other genera in the Philodendroideae, the spathe monkeys, Cebus capucinll.s (J. Oppenheimer, pers. 

re-closes ovel the female flowers after pollination. comm.), but it is not knowr] whether they are the 

In most cases, protection of the developing fruits primary dispersal agents, sillve cebus monkeys are 

after pollination is provided by the thickness of the known to be generalists ir] lheir eating behavior (J. 

spathe, as is true of other aroid genera such as 

Philodendron, but in Dieffenbachia protection is 

Oppenheimer, pers. comm.). 

also provided by toxins in the sap, which may burn 

the mouth of any animal eating it. However, the 

FLOWERING BEHAVIOR ANI) ItOLLINATION 

pericarp and mesocarp of the berry appear to have Dieffenbachia populations tend to be aggregated, 

no injurious effects when eaten and apparently lack perhaps due as much to the vegetative reproduction 

the sallle toxic effects found elsewhere. 

of the stems as by local germination of seeds. A 

Generally, most pistils of an inflorescence are single clone may have many stems, but relatively 

pollinated and develop into fruits, but the number few stems flower in a given year, and the number 

produce(l per plant is variable. Studies in Costa of inflorescences open on a given day is few (Young, 

Rica on D. oerstedii (Valerio, 1984) showed that the 1988b). Inflorescences borne on a single stem genfruits 

varied from 13 to 43 with an average of 26 erally do not anthesally overlap, thus there is little 

per plant. Valerio also reported that some berries potential for geitonogamy (Young, 1990). For D. ni-


2004 

Croat 


tidipetiolata (reported as D. Iongispatha in Young, to close and the staminate flowers are emitting pol- 

1988b) a reproductive stem may have two to seven len in long filaments (Fig. 5F). In order to leave the 

inflorescences during a single growing season, ma- inflorescence the beetle must literally wade through 

turing at intervals of 3 to 12 days (Young, 1986). pollen, ensuring that it will be covered with pollen 

For D. oerstedii there may be between one and four before flying away to the next receptive infloresinflorescences 

per axil. Valerio (1984) reported that cence. 

plants which produced a solitary inflorescence (34 While some Dieffenbachia species, perhaps 

of the 83 plants studied produced inflorescences) most, have spathes that remain open only 24 hr. 

did not develop fruit. Valerio was unable to ascer- (based on my observations in the field), the pollitain 

whether this was due to a lack of pollinators nation event for D. nitidipetiolata (reported by 

or age of the plants. 

Young as D. Iongispatha) involves three days 

The pollination biology of Dieffenbachia has (Young, 1990). Inflorescences of this species open 

been studied (Croat, 1978, 1983b; Valerio, 1984; in the evening of the first day, but the spadix is not 

Young, 1986, 1988a, 1990). It has long been known initially receptive. On the evening of the second 

to aroid workers that Dieffenbachia and several oth- day the stigmas become receptive (about 17:30) 

er aroid genera, e.g., Philodendron, some Syngon- and the spadix temperature increases to 4°C above 

ium, and Xanthosoma, are visited regularly by large ambient temperature (Young, 1990). It is at this 

scarab beetles (Coleoptera). Young (1986, 1990) time that beetle pollinators arrive, and they enter 

found that D. nitidipetiolata (reported as D. Iongis- the spathe where they generally persist for another 

patha) is pollinated mostly by beetles of the genus full day, departing on the evening of the third day. 

Cyclocephala (Scarabaeideae, Dynastineae) or more Young (1990) found that for most days there were 

infrequently by the genus Erioscelis, also a dynas- fewer than four inflorescences open and in ferrlale 

tine scarab. Other visitors, including Diptera, He- phase at any one time in an area of 700,000 rrl2. In 

miptera, Dermaptera, Thysanoptera, an(l nitidulid her stuLlies at l, Selva in Costa Riea she foun(l 

beetles (Coleol)tera, Nitu(luli(lae), I)rove(l not to that l)eetles flew I)etween 1 an(l 68() m (averaging 

( arry )ol]en and were deeme(l ly Young nol to I)e 

pollinators (Young, 199()). 

8.S m) ))etween (onse( utive visits to 19. nitifli/)etiol- 

t(l (leporte({ as 1). lest7.gis/)(lth(l). 

()ther monoe(ious aloid generae su(h as; l'lliles- Beetle pollination ot l)ie/>nlselehiel is not speler1(1ror1 

an(l X(lethc)somfl, are krown to le }eetle (ies-s,^)e(ifi(. Nine (liffeellt s(arat) teetle sy)e(ies 

[)ollinate(l an(l provide foo(l })y meal-]S; of oil-ri( h of (Sycloee)lzfll(l as wel l as Frioseeli.s ( ollltnl)ie (l Ensterile 

male flowers. ln l)iet/erll)(l(hi(l the foo(1 re- (IIO(I; WUlP fOllrl(l to )OIlitlAtC 1). niti/i/)cJt.iolfltel (rewal(l 

(onsists of the l)rotein-ri(h (lul-shale(l sta- [)orted I)y Young as I). lotZgisisfithfl) at the l,a Selva 

mino(lia surroun(ling ea( h female flower (Young, ltesel ve i] Costa lti( a (Young, ] 986). Still, lliet 

1986, 199()), I)ut the lmeetles have also })een seen fenhach ia sI )ecies l l o(l u( e (lifferent s( ents an(l (liffeeding 

on the lowermost poltion of the fertile sta- ferentially attl ae t pal tie ular leetle spee ies (G. 

minate portion of the sleadix. An indication that the Se hatz, pers. ( omm.). 

staminodial foo(l source is impoltant to the beetles Be(ause of the loose relationship between the 

an(l prefelred over other flolal parts, is that the pollinators ancl any olle species of DiefWenbachia, 

beetles will leave an inflorescence. 

hybridization (loes occasiollally occur (Young, pers. 

The production of scents during flowering is fa- (omm.). It is known that seeds from hybrid plants 

cilitatecl by the thermogenic behavior of the spadix, do germinate and produce viable F1 plants. It has 

which may increase as much as 4°C during anthesis not yet been determined if this F1 generation is 

(Young, 1990). Scent production coincides with the capable of reproducing sexually. Still, this phenomflight 

activity of the scarab beetle pollinators enon of hybridization might explain the many dif- 

(Schatz, 1990). Active movement of pollinators is ferent, seemingly related, but distinct, populations 

usually at dusk, seemingly in direct paths to an of D. nitidipetiolata that occur in some regions, 

open inflorescence. Beetles arrive at the inflores- such as the Rio Guanche region of Colon Province 

cence by first landing on the spadix (Young, 1990), in Panama. Since asexual reproduction is so prevthen 

entering the lower tube portion of the spathe alent in Dieffenbachia, occasional hybridization folwhere 

the pistillate portion of the spathe is at an- lowed by vegetative reproduction might explain 

thesis. Beetles typically remain in the inflorescence these patterns of variation. 

for 24 hr. after their arrival. While in the spathe 

the beetles mate and eat the nutritious staminodia PHENOLOGY 

(Young, 1986) surrounding the pistils. Beetles de- Some species of Dieffenbachia are in flower all 

part the following day as the inflorescence begins year-round. In general, more flowering takes place 

681

682 

Annals of the 


during the rainy season even if the species flowers ama (20), and Costa Rica (13), principally at lower 

at other times of the year as well. This may be tied to middle elevations in the Andes. The genus is 

to the fact that the beetles pollinating Dieff7enbachia also widespread in the Amazon basin as is eviare 

more frequent during wet periods. Although D. denced by the large number of species at low elebeachiana 

and D. burgeri flower primarily in the vations in Brazil with a total of 27 species. Only 8 

dry season in (2osta Rica and PanamaS they occur species occur in the Guianas region in eastern 

in wetter habitats where beetle pollinators are re- South America and the Territorio Amapa of northliably 

available. 

eastern Brazil. The genus is exceptionally abundant 

in the western Amazon basin in the foothills of the 

>Y'rol,o(Jy 

eastern Andes, with 57 species. The wetter forests 

at middle elevations in the foothills of the Andes 

Dieffenbachia has chromosome counts of 2n = 

are particularly rich. For example, there are 5 spe- 

34 and 68 (Petersen, 1989). Grayum argued that 

cies in the vicinity of the Jatun Sacha Reserve 

Dieffenbachia is closely related to Philodendron 

along the Upper Rio Napo near Misahualli in Ec- 

(Grayum, 1984). Petersen (1989) argued that Philuador. 

odendron would not be considered so closely relat- 

As is true for many other genera of Araceae 

ed if its chromosome base number were not 17 as 

(Croat, 1992, 1983a, 1986a, 1986b), the northassumed 

by Grayum. She also argued that the difwestern 

part of South America, especially on the 

ferences in the "size and constitution of the chro- 

Pacific slope of Colombia and adjacent Ecuador, is 

mosomes (small metacentrics of Philodendron versus 

rich in species of Dieffenbachia. About 14 species 

medium to large submetacentrics to subtelocentrics 

occur on the eastern slopes of the Andes in Ecuaof 

Dieffenbachia) also indicate that no close relationdor 

and about 1*S species are known from the Paship 

exists between the two genera" (Petersen, 1989: 

cific slope in Colombia. Relatively [ew species oc- 

128). Petersen ( onsidered the chromosome basic 

cur in southern South America, with only 

number in Philo(lendron to be x = 18 based on the 

Dieffenbachia aS,tl(lonematifolia Engl. occurring in 

fact that all members of Philodendron subg. Mesouthern 

Brazil, I'araguay (Croat & Mount, 1988), 

conostigma (Schott) Engl. have 2n = 18 and suband 

Argentina. (:ollections of only 7 species, mostly 

genus PhilodenXron Schott also has 2n = 18 in 

new species, have been seen for Bolivia. Northpart. 

Grayum (1984) suggested that Daeffenbachia 

central South America is also not particutarly rich 

was close to the tribe Spathicarpeae (subfam. Aroin 

species. Only 2 species occur in the Cordillera 

ideae). Petersen (1989) considered this a reasonde 

Merida in Venezuela, and only 1 of these occurs 

able suggestion since they share the same basic 

in the Cordillera de la Costa (Bunting, 1979; Croat 

number of x = 17 and large chromosomes also with 

& Lambert, l9X7). 

the centromere frequently located distally on the 

A preliminalw key and preliminary descriptions 

chromatids. Petersen (1989) believed that the Aglahave 

been pro(luced for South American lWieffenonemateae 

may be the closest relatives of Dieffenbachia 

species, but many species remain poorly 

bachieae based on the similarity in the constitution 

known. Thus the figures given for South America 

and size of the chromosomes, despite the fact that 

remain tentative. 

the former has a different chromosome base number 

Central American species constitute a group rel- 

(x = 20). Chromosome counts for Bognera reconatively 

isolated from South America, and only seven 

dita (Madison) Mayo & Nicolson, the only other 

species range into South America, especially along 

member of Dieffenhachieae, are also 2n = 32t. Bogthe 

Pacific slope and in the Magdalena River Valley 

nera, a monotypic genus, also shares a karyotype 

in Antioquia Department. The majority of Dieffenof 

medium-sized metacentric to sub-telocentric 

bachia in Central America occur from Nicaragua to 

chromosomes with Dieff7enbachia (Petersen, 1989). 

Panama, and most are relatively widespread in this 

GEOGRAPHICAL DISTHIBUTION AND ENDEMISM 

region. The most widespread species of lWieffenbachia, 

D. oerstedii, ranges from southern Mexico 

The genus lEaegenbachia has approximately 135 to western Panama (lXig. 29). Dieffenbachia wenspecies, 

most of them in South America. It ranges dlandii ranges from Mexico to Costa Rica (Fig. 28) 

from Mexico, throughout Central America, and to and both D. nitidipetiolata and D. tonduzii range 

the West Indies, Trinidad, and most of South Amer- from southeastern Nicaragua to Ecuador (Figs. 30 

ica as far south as Argentina, Paraguay, anel Boliv- & 29, respectively). Irl the latter country, D. nitiia. 

Distribution of species is unequal, with major d ipetiolata crosses the Andes into the Amazon 

centers of diversity including Colombia with 37 drainage in Napo. ReLnaining Dieffenbachia are all 

species, Ecuador (34), Peru (30), Brazil (27), Pan- narrowly isolated. Nine species are shared between


2004 

Croat 

Revision of Dieffenbachia 683 

Costa Rica and Panama, but even these are not independently after the Andes began to arise towidespread 

in the two countries. Those shared are ward the end of the Cretaceous (Raven & Axelrocl, 

D. aurantiaca, D. beachiana, D. davidsei, D. gra- 1974). The fact that there are no truly wide-ranging 

yumiana, D. killipii, D. nitidipetiolata, D. oerstedii, species, i.e., those ranging from Mexico to Brazil, 

D. tonduzii, and D. wendlandii. Dieffenbachia au- attests to this isolation. The high rates of endemism 

rantiaca occurs only in southwestern Costa Rica in in Costa Rica and Panama as well as Mexico perthe 

vicinity of the Osa Peninsula and in adjacent haps reflects the isolation of these areas during pe- 

Panama in the Burica Peninsula area (Fig. 27). riods when the oceans were at much higher levels 

Both Dieffenbachia beachiana and D. grayumiana than they are today, and when the area that is now 

range from northern Costa Rica to western Panama central Panama and Costa Rica was subsequently 

(Bocas del Toro) (Figs. 28, 27), while D. daviclsei clisconnectecl from South America. At the close of 

ranges from northwestern Costa Rica to central the Tertiary period, 18,000 years ago, the sea level 

Panama (San Blas) and Colombia (Fig. 27). was about 100 m higher than today (Holmes, 1969). 

A total of 11 species (42Wo of the total) of Dief- Much earlier the land mass of what is now Central 

fenbachia are endemic to either Costa Rica or Pan- America began to emerge as a series of islands durama. 

Endemism is particularly high in Panama ing the Oligocene epoch with further uplifting durwhere 

8 of the 26 species (31Wo) are endemic. In ing the Middle Miocene. It was not until the Upper 

Panama endemic species are D. copensis, D. cre- Miocene and Pliocene epochs that the final portions 

bripistillata, D. fosteri, D. fortunensis, D. galclame- of the isthmus of Panama emerged above sea level 

siae, D. Iutheri, D. panamensis, and D. pittieri (Figs. (Torre, 1965), and the final connection of Central 

27-30). In Costa Rica 3 of 26 species (12Wo) are and South America was about 5.7 million years ago. 

endemic. Costa Rican endemics are D. burgeri, D. To put these geological events in relation to the 

hammelii, ancl D. horichii (Fig. 27). Neither Mexicc moclern aroicl flora, it should be notecl that even 

nor Miclelle America have enclemic species. cluring this era preculsors to the extant flora proI)- 

DiefJenbflchifl isthmia (Fig. 28), D. Iongisl)eltha ably alreacly existe(l. The angiosperm floras of the 

(Fig. 29), and D. obscllrinervisl (Fig. 29) range Oligo(ene Wele Ivelieve(l to have consiste(! almost 

ae X OSS the l sthmus of Panama from Vel aguas Prov- elltirely of extant genera and with existillg spee ies 

inc e to northel ll Colombia. DieiZenbflc1lia aur(letifl- alllong Oligoe ene atl(l Plio( ene ftol as ('raklllajan, 

a,19. beachiflnfl, and D. grflyllmiflnfl (Figs.27,28) 19G9). 

ale the only s)ecies oc(urring in both Costa Rica Equally important as geo]ogy lo the isolcltion of 

an(l Panama. S(lme spe( ies are fulther isolate({. tlle Celltlal Amelicall aroid flola are ecologi(al fac- 

1)ieffenl)achia allrantiflc f1 an(1 1). burgeri are re- toI s fol Central Amel ic an spee ies of Dieffenl)achi(l. 

stri( te(l to the legion of the ()sa Peninsula in south- Mue h of eastern Panama c onsists of bl oad expanses 

western Costa Ric a (or in the ( ase of D. aurantiac of Tropical moist forest (T-mf) with other, generally 

in Panama on the adjacent Burica Peninsula; Fig. smaller areas of Premontane wet (P-wf) and Tropical 

27) and D. horichii is lestricted to a relatively small wet forest (T-mf) (Holdridge, 1967). In contrast to 

area on the Pacific slope of Costa Rica (Fig. 27). Panama, much of the Choco area of northwestern 

Several other species are also narrowly restricted Colombia consists of much wetter pluvial forest 

or known only from the type specimen. Dieffen- with annual precipitation exceeding 11,700 mm 

bachia fosteri is restricted to northeastern Panama (Gentry, 1982). This broad band of pluvial forest 

(Fig. 27). Dieifenbachia fortunensis is restricted to with its own suite of unique endemic species no 

the northern Chiriqui Province (Fig. 28). Dieffen- doubt acts as a dispersal barrier for species from 

bachia copensis is found only in the Cocle Province regions with less rainfall. It may account for those 

of Panama (Fig. 27), and D. galdamesiae in a small Panamanian and Costa Rican species that skip the 

area south of the Panama Canal (Fig. 28). wettest areas of northwestern South America but re- 

Further collecting in Colombia, especially along occur in the relatively drier areas of mesic western 

the western slope of the Andes, may change the Ecuador. 

distribution of Dieffenbachia7 but the current pat- Middle America has low species diversity with 

terns most likely reflect the realities of life zone Guatemala having only two species and Honduras 

ecology and geologic history of the area rather than three. Only Nicaragua, with six species, four of 

under-collecting. Since relatively few species of Ar- them restricted to the southeastern corner near Cosaceae 

are known to occur at lower elevations on ta Rica, is moderately rich in species. All except 

both the eastern and western side of the Andes, it D. standleyi and D. wendlandii appear to range into 

can be presumed that the evolution of the respec- the country from Costa Rica. The low species ditive 

Amazonian and Pacific coastal floras occurred versity and very low aroid endemism in Dieffen-

684 

bachia in Middle America is perhaps explained by 

its historic remoteness from existing large land 

masses to the north and to the south. The distribution 

of modern aroid species suggests that the 

northwestern part of Middle America may have 

been isolated from Costa Rica in the area of the 

San Juan Depression (Nicaragua). Many of the species 

that occ ur in Costa Rica or Panama enter into 

Nicaragua in only a small area in the southeastern 

Annals of the 


in the United States in Florida, California, and Ha- 

. . 

WAll . 

TAXONOMIC TREATMENT 

Dieffenbachia Schott, Wiener Z. Kunst 3: 820. 

1829. TYPE: Dieffenbachia seguine (Jacq.) 

Schott. 

Seguinurn Raf., F1. Tellur. 3: 66. 121t36 [1837]. TYPE: Separt 

of the country. Though the existing flora of guinurn rnaculaturn Raf. {= Oieffenbachia seguine 

Guatemala (loes not reflect isolation from Mexico to (Jacq.) Schott]. 

Maguirea A. D. Hawkes, Bull. rFOIey Bot. Club 75: 635. 

the same (legree, it is possible that the more ele- 

1948. TYPE: Maguirea spathi((lrpoides A. D. Hawkes 

vated portions of Guatemala, Nicaragua, and Hon- [Diefafenbachia palud icola N . F3. Br.]. 

duras were isolated from major portions of Mexico 

by the Isthlnus of Tehuantepec. With no endemic Terrestrial; caudices thick, often elongate and 

Mexican sy)ecies, the Dieffienbachia flora reveals prostrate, rooting at the lower nodes, the older porless 

endemism than does Anthurium (Croat, 1983a) tions of the stem trailing across the surface of the 

or Philoden(lron (Croat, 1997). Both Mexican Dief- ground, often for considerable (listances; sap often 

fenbachia ale relatively widespread, with D. oerste- milky and frequently with valyillg concentrations of 

dii ranging to Panama (Fig. 29) and D. wendlandii 

oxalic acid, with conspicuous; annular leaf scars. 

ranging to l'ananla (Fig. 28). 

Petioles elongate, amplexicaule slleathed to the mid- 

The reluaining Costa Rican and Panamanian 

dle or sometimes to the apex; s;lleath unequal and 

species not allea(ly discussed above appear not to 

often free-ending at apex; leave o; c lustered in a tight 

have stron> tlffinilies with South American species 

whorl at the stem apex; I)lel e1e .s ovate to oblongand 

clearly (lo not have affinities with othel Cenllal 

ovate, elliptic, oblanceolate. ol ol)ovateX acuminate 

American sy)e( ies. On the other hand, there are 

at apex, acute to obtuse to lous(le(l at base, typically 

subcoriaceous, someli llle S val iegated throughsome, 

reltltively few, Central American species, 

out or in areas along the lzli(ll il) with paler colors; 

among thesl l). killil)ii, D. nitidipetiolata, and D. 

midrib raised on both siulltle ( s; primary lateral 

tonduzZi, tlsal ale likely to be of South Ameli(an 

veins pinnate, usually sunkell (II)oveX raised below, 

origin sin( e they are wide-ranging species in South 

much more prominent than tleA Illinor veins, all lat- 

America. 

eral veins extending to the Illulgills without forming 

a single collective vein lout with several primary 

HORTICU 1 ,'1't J 14 A I SIGNIFICANCE 

lateral veins often forming a se l ies of short, discon- 

Dieffenb(lchi(l, like many other members of the 

Araceae, is one of the world's most important ornamental 

l)lants. Owing to the attractive leaves and 

hardiness tIn(lel the difficult conditions found indoors, 

DieJ/enb(lchia remains one of the most important 

olnamental plants in homes, offices, and 

professional displays. Ornamental aroids play a major 

role in tlle foliage-plant industry in the United 

States, making up about one-third of total foliage 

plant sales (Henny, 1988). Dief7Senbachia is one of 

5 aroid genera routinely among the top 10 foliage 

plants in annual sales volume (Henny, 1988). 

The Araceae hold 7 of the top 13 positions in 

tinuous collective veins; inleAl l)l i mary lateral veins 

sometimes present; minor veils (listinct to obscure, 

usually not markedly raise(le sollletimes connected 

by transverse tertiary veins. I N Iv^LORESCENCES 

shorter than the leaves, 1 to several per axil; bracts 

short and usually inconspicuous; peduncle usually 

somewhat flattened and angulcll with one edge narrowed; 

spathe oblong, persistellle convolute at base, 

often somewhat constricted aI)out midway, opening 

usually only above the mid(lle, usually green on 

outside, somewhat paler within; spadix slightly 

shorter than the spathe, divided into pistillate and 

staminate portions, each with naked unisexual flowers; 

the pistillate portion basal, about as long as the 

terms of overall sales in North America, including staminate portion, fused to the spathe, remotely 

the first two (R. Henney, pers. comm.). An orna- many-flowered (typically 20 to 80 rarely to 160); 

mental cultivar of Dieffenbachia is the second most staminate portion clavate, white, free from the 

important in sales, and the genus consistently has spathe, densely many-flowered, often separated 

one or more taxa on the list of most popular orna- from the pistillate portion by a naked interval. Stamental 

plants in North America. Dieffienbachia minate flowers consisting of 4 stamens united into 

breeding and culture plays a major economic role a 4- to 5-sulcate, truncate synandrium; anthers lat-

Volume 91, Number 4 Croat 685 

2004 Revision of Dieffenbachia 

eral, contiguous, the common connective thick, sessile, depressed globose to depressed ovoid, 2- or 

fleshy, the thecae obovoid to oblong-elliptic, de- 3-lobate; ovules 1 per locule, pale green, semigloshiscing 

by short, apical, pore-like slits; pollen ex- sy, erect, anatropous; placenta axile to basal; stylar 

truded in strands, inaperturate, ellipsoid or oblong region inconspicuous; stigma large, hemispheric or 

or nearly spherical, large, averaging more than 75 2- or 3-lobate, about as broad as the ovary, yellow 

1lm, exine psilate to obscurely verruculate and/or to orange in color. INFRUCTESCENCE with fruitsparingly 

punctate-foveolate to densely foveolate; ing spathe often turning yellow, orange, or red, 

pistillate flowers modelately dispersed but often in breaking up longitudinally in fruit to expose the 

weak rows with 1 to 5(6) pistils per spiral, and colorful berries; fruits baccate, globose or 2- or 3- 

surrounded by 4 or 5 claviform white staminodia, lobed, 1- or 2(3)-seeded; seeds globose or ovoid, 

these longer than the ovary usually 2-5 mm, usu- the testa thick, smooth, green to blackish green; 

ally spreading, sometimes fused briefly at the base; embryo large; endosperm lacking. 

Ovaries 2- or 3-carpellate, sometimes l-carpellate, Chromosomes 2n = 34, 68. 

KFY ro DIEFFENBACHIA OF MEXTCO, C:ENTRAT AMFRICA, AND THE WEST IND1ES 

la. Petiole involute, extending to base of blade and frequently prolonged beyond the base of the blade. 

2a. Larger leaves regularly to more than 45 cm long and 23 cm wide, cuneate at base, coriaceous, usually 

glossy above, not variegated, uniformly green. 

3a. Petiole sheath markedly undulate, at least near the base; Atlantic slope. 

4a. Leaf b]ades matte and somewhat velvety to glisteningly ve]vety above, minutely wrinkled 

on uppel surface; clrying dark blae kish btown to dark ye]lowish brown above; Atlantic slope 

of Panama 21. 1). jvflnslm.entsis Croat 

4b. I,ceaS b1a(1c-s serrlig10ssy to glossy above an(S smoolh on upl)er surface; drying me(lium gray 

to (lark t)rownish gray above, g)a1er ar(1 ye110wish gray to I)alee yee110w-t)rown be10w - --------- 

24. 1). .sl(lr1(ll(loi (roal 

:3b. 1' lio1e shealh rlol rra1ke(11y L1rl(1u1ale; 1'ae ilie s10Z>e 1:3. 1). hori(11ii (r-otll & (,r-lyl1rr 

St>. I ( (IVf s rtlrely rrlo1e tharl 45 ( 1zl 10,-,g 01 2:3 ( rzl wi(1e . 

5(1. 1'1ilzlly 1(1lel1 Vei1lS 6 10 7 )(-1 si(1e; krlow11 olly l1olrl the ty1> 10(llity irl the 1'ar(1rzl (1(1tltl1 

(,, et, . . 22 1). /)i.ll.i.()t-i. 1s,,,r1 

5b. I,i,,,a,y 1t,1e,(,l vei,,s ,ts,,ally 1.5 Ot llO(. 

()t1. 1,tlt illtI(lUS Willl )lillltil\' 1dlPldl V('illS (1())1') lO 17 )('1 si(lf'; 1)ld(lf' t)tlS('OlUlIIS(' lO t011tl(1f'(l; 

1)ld(lP Slil[tl(' m0()SSy 1() tll()(1eld181)' m0()sS), [l()l d1 t11l 1)lilltilt'! Itit('ly Vtili('ttilf'(l; [)('li()lf' IlSIItilIy 

Wilil tI WllitiSil l)t[tl(l 011 IllP Al)tIXiAI Slil[tl('f'; i)tilitililtl (). 1). (r(l)ril)i*slill(ll(l (lo 

6b. Leat 1>1a(1es wi111 )1i1zl1> 1(1l Xa1 vf i1ls 1lslltl11y 1ll01e 11larl It) )( 1 si(1(; 1lslltl11y ( o-(11X1tl1 (l1 

1>(1se, llsllally 1zllle ar(1 sl1>velvely (11o/, 1lslltl11y 1>tl11a1f, ol1erl vtlrif ga1e(1; pf 1io1e- 1ae kit, 

a w1-ilish 1)an(1 o11 the tIl)tIXiclI SIII[tI('E'; Ni(4llAgIItI lo (O0OlllllitI ---------- ---------------- ------ 

- - - - - - - - - - - - - - - - - - - - - - - - -- - - - - - - - - 2 ,5 . I) . l ( ) t I ( l I IZ i i ( j I ( ) ( I I (S J I a y L I I l l 

11). 1)elio1e shetllh er-ee t 10 irlvolule, erl(1ir, sllo1l ol I>la(Je 1>ase (eXe ep1 rare ly orl ll)per 1eaves). 

Tt1. Ulls1leat1le(l porliorl ol petio1e uslltl11y t1it1rlgu1tl1 ir] (10SS se(liorl, s1larply kef1e(1 lo ri(1g(1 (11>axitl11y, 

t11e rrargirls t1-li(kly winge(l 1. 1). (ltlr(lrlli(l((l ,rlg1. 

Ti). 1)etio1e teyo1l(t shenth sul)ter-ete, 1-oun(1e(1 ataxin11y, wil11 rrargirls rour(le(l lo ae ute (rlol wirlge(-1). 

tJd. lvlArllS witll pPliOlUS 01 rrldjol veirls ol 10we- t>1a(1e surlia(e rrlirlutely g1-anular--pu1)eru1erll lo pl1- 

1.>e1 ulerlt. 

9d. 1'rirrary 

1atera1 veins 23 to 36 per si(1ee, (1epa1lint, midril) usua11y dt dbOUt a (3()° angle; 

lili(tl'ih dn(t prililAly IdtElal VE,illS ('0t1Spi('UOUS0y pUt)CIUlUtEl with wllitisll tri(homes ----------- 

2. 1). 6eachiclea Croat & (Jrayum 

b. Priltlaly latelal veins usually less than 18 pairs (rarely lo 22 in D. grayumiana) usually 

departing midril) 40°-6()° at blade mi(lpoinl; midlib and primary lateral veins infonspituously 

glanular-puberulent. 

10a. Blades narrowly oblanceolate, 6.4 times longer than wide, less than 6 cm wide ----- - 

5. D. copensis Croat 

10b. Blades elliptic to obovate-elliptic, oblong-elliptic, to narrowly oblong-lanceolate (rarely 

narrowly oblanceolate in D. fortunensis), 1.7-5.2 times longer than wide, 6.7-39 cm 

wide. 

lla. Petioles with sheaths ending no more than 5 cm from base of blade------------------- 

24. D. standleyi Croat 

llb. Petioles with sheaths ending more than 13 cm from base of blade. 

12a. Primary lateral veins arising at 30°-40° angle; central Panama ---------- 

----- 10. D. galdamesiae Croat 

12b. Primary lateral veins arising at mostly 55°-70° (to 80° in D. grayumiana); 

Costa Rica and western Panama. 

13a. Plants usually stout, 1.5 m tall; blades narrowly ovate, (22) 30-54 

cm long, 10-32 cm wide (averaging 36 x 18 cm), 1.5-2.6 times



longer than wide; Costa Rica to western Panama, 0-1300 m but mostly 

near sea level 11. D. grayumiana Croat 

13b. Plants medium-sized, rarely more than 1 m tall, usually less; blades 

mostly narrowly oblong-lanceolate, 15-33 cm long, 4.2-13 cm wide 

(averaging 21 x 6.8 cm), 2-6.3 times longer than wide; Panama 

(Chiriquf), 900-1600 m 8. D. fortunensis Croat 

8b. Plants with petioles or major veins of the lower blade surface glabrous. 

14a. Petiole sheaths decurrent apically (lacking prominent protruding extensions); Atlantic lowlands. 

15a. Blades drying blackened; Panama (Bocas del Toro), near sea level - ----------- -- 

9. D. fosteri Croat 

15b. Blades drying yellowish brown, greenish to yellowish brown. 

16a. Petioles very glossy, drying as if covered with shellac; Nicaragua to Ecuador 

18. D. nitidipetiolata Croat 

16b. Petioles not glossy, not drying as if covered with shellac. 

17a. Plants less than 1 m tall; blades usually less than 10 cm wide -- ----------- 

12. D. hammelii Croat 8 Grayum 

17b. Stems usually more than 1 m tall; blades more than 10 cm wide. 

18a. Petioles matte; leaf blades usually subcordate at base, usually flecked 

with creamy yellow 11. I). grayumiana Croat 

18b. Petioles semiglossy; leaf blades cuneate to rounde(l or truncate at 

base, usually plain green (rarely mottled with pale gleen). 

l9a. Petioles terete; sheaths decurrent at apex, tighlly inrolled with 

one side completely hidden by the other; Mexie o to Panama but 

but primarily on Pacific slope (except in Oaxa( a and Veracruz 

on Atlantic slope) 26. I). wendlandii Schott 

l9b. Petioles usually obtusely sulcate, sometimes lelete with an obtuse 

medial rib; sheaths open, the margins elect; Nicaragua to 

Cololllbia 16. D. Iongi.s/)(llll(l F:ngl. 8 K. Krause 

1X11). Ietiole sheaths (at least on one side) rounded to auriculate at apex. 

2()a. Blades with more than 18 plimary lateral veins or with primary lalelal veins obscure, 

barely more conspit uous thtln interprimary veins. 

21a. West Indian and Soutll American species; plants with botll sllXly sulcate petioles 

and a thi(kened, protruding spadix; fruits 2- to 3-lol(}(l 

23. I). .s(guine (Jacq.) Schott 

21b. Mainly Central Ameli(an species (D. killipii, D. isthmia, (1X(l 1). obscurinervia 

also in South America). 

22a. Blades with 10 lo 13 pairs of primary lateral veins, these moderately inconspicuous, 

balely more obvious than interprimary ve ills; petioles and 

lower miflribs with pale spots; Central Panama to (ololzll)ia ------ 

----------------- -------------- ---- --- ----------------- -- -- --- 1(). I). obscurinervia Croat 

22b. Blades with 18 lo 28 pairs of primary lateral veins, (Ie( i(ledly more conspicuous 

than tlle interprimary veins; petioles and lowet lzlidrib lacking 

spots; Costa Rica, Golfo Dulce region 3. l). I)llrg(ri Croat 8 Grayum 

20b. Blades variously shaped, matte to glossy above, with 8 to 18 prilllly ltlteral veins per 

side. 

23a. Spadix lacking medial sterile region, the male and femalc- legioIs c.ontiguous or 

nearly so. 

24a. Petioles usually solid green, not densely and conspi( vlotlsly pale-maculate. 

25a. Blades with major veins on the lower surface ganular-puberulent --- 

17. D. Iutheri Croat 

25b. Blades with major veins on the lower surface glabrous ------- 

------------------------------- ---------------- - -------------- -- 15. D. killipii Croat 

24b. Petioles densely and conspicuously pale-maculate. 

26a. Stems more than 1 cm thick; petioles never pale-maculate; primary 

lateral veins usually departing midrib at less than 90°; Costa Rica 

(Osa Peninsula), Panama, and Colombia 15. D. killipEi Croat 

26b. Stems usually less than 1 cm thick; petioles distinctly pale-maculate; 

primary lateral veins usually departing midrib at 90° or more; Costa 

Rica, Sixaola region, Panama, and Colombia -------------------------------- 

------- -- ---------- - - 7. D. davidsei Croat 8 Grayum 

23b. Spadix with an evident medial sterile region, pistillate and staminate portions 

separated by a distinct naked spadix axis. 

27a. Plants usually less than 0.5 m tall 20. D. oerstedii Schott 

27b. Plants to more than 0.5 m tall. 

28a. Blades drying dark brown to blackened on upper surface. 

29a. Blades usually with posterior lobes; lower blade surface drying


2004 

Croat 


yellowish brown to yellowish gray-brown to dark yellow-brown 

or ye]lowish green; stems solid dark green, drying glossy and 

minutely wrinkled (the areoles interspersed with raphide cells); 

Panama, Veraguas Province and the Azuero Peninsula in the 

west (at 700-900 m in Herrera and Los Santos Provinces) to 

Darien Province in the east, and to Colombia 50-800(1000) m 

14. D. isthniia Croat 

29b. Blades lacking posterior lobes; lower blade surface drying grayish 

black; stetns mottled dark green, drying smooth with a dense 

layer of raphide cells (but not minutely wrinkled); Panama, Bocas 

del Toro, sea level 9. D. fosteri Croat 

28b. Blades drying yellowish green to dark gray-green, rarely brownish on 

upper surface. 

30a. Petioles usually white at base; blades usually less than 20 cm 

long 20. D. oerstedii Schott 

30b. Petioles not whitish at base, frequently paler than stem but not 

markedly paler than the remainder of the petiole; blades usually 

more than 20 cm long. 

31a. Petioles usually sharply C-shaped, discolored and whitish 

at the base; blades matte 20. D. oerstedii Schott 

31b. Petioles terete to obtusely C-sllaped or obtusely D-shaped, 

not sharply sulcate, not discolored and whitish at base; 

blades glossy to semiglossy. 

32a. Species of mostly dry habitats in western Central 

America (Mexico to Panama); blades subcoriaceous, 

selrliglossy on upper surface, (15)20-55(65) cm long 

(avelaging 35 x 17 cm); 75 to 900 m in elevation 

26. D. wendlandii St hott 

320-). Spet ies of wet habitals, southeastern Niâragua and 

along the. entile Atlali( slole an(S the ()sa lveellirlsula 

of (4ostcl lli(*a; I)la(les tlso(lelaleZy (*oria(*eotIsX 

gleessy ols 11se tIl)pel .sulfa( e., 1()-.3() ( rts lorig., (avel 

cigint, 2.3 X 12 (n); sea lt'vEl to 2()() 21, nlOStIy tIt 

leess 111as 1()() rll its elevaliors -- - -- - - - - -- 

687 

- -- ------------------------------- ?1. 1). (orl.(itltl(l (31'0dl & (,I'AyLItll 

1. Dieffellbachia aurantiaca 11Jllgl., Anales Tnst. erlding an(l obliquely rouneled to rourl(le(l-aulicu- 

Fis.-6eogt. Nac. Costa Rie a 9: 2()9. 1898. late distally; unsheathe(l ^)ortion 0.tS-2 c m long, 

TYPE: Costa Rica. Puntarenas: "ill silvis pro- angular to + triangular in c ross sec tion, flat to 

pe Santo Domingo atl sinum dule em [Santo bloadly sul(ate a(laxially (sometimes with a medial 

Domingo de Golfo Dulce], fructif. Mart.1896," keel), adaxial margins thin and revolute, acutely 

A. Tonsluz 9961 (holotype, B!; isotypes, CR!, keeled abaxial]y, abaxial margins thickly winged, 

US!). Figures 2, 27A. 

sometimes with a medial keel, thin, revolute, sometimes 

medially keeled at malgins; blades oblong- 

Stout herb, 0.7-2.3 m tall; sap fetid and caustic; 

elliptic to ovate-elliptic, slightly inequilateral, one 

stems erect at apical part, to 2 m long, the older 

side 1-1.5 cm wider than the other side, usually 

portion reclining and up to 2 rll long; internodes 

2.0-4(6) cm long, 5-7(10) cm diam., dark green to 

subcoriaceous, acuminate to abruptly acuminate at 

pale greenX semiglossy to glossy, streaked with paler 

apex, slightly inequilateral and obtuse to rounded 

green; petioles 13-33(48) cm long (averaging 24 cm 

or subcordate (rarely attenuate on one side) at base, 

(26)31-57 x 11-27 cm (averaging 39 x 

long), C-shaped at base in cross sectione moderately 

19 cm), 

spongy, medium green, semiglossy adaxially, matte 1.7-2.6 times longer than wide (averaging 2 times), 

and acutely l-ribbed abaxially, the surface finely ranging from about as long as petioles to 2.8 times 

dark striate, sometimes variegated with paler green longer than petioles (averaging 1.7 times longer); 

markings, sheathing 

margins weakly undulate; upper surface dark 

1/3 to nearly throughout (35%- 

97% their length and averaging 77Wo); sheath 6.5- green, concolorous or sometimes variegated, drying 

25.5(38) cm long (averaging 17.7 cm), medium dark olive-green to medium gray-green, weakly 

green, finely dark green striate, sometimes varie- glossy to semiglossy; lower surface moderately palgated 

with paler green markings, adaxially acute er, matte to weakly glossy, drying pale yellowish 

margins sometimes much paler, with margins in- green; midrib flat to broadly concave above (broadcurled 

but usually not overlapping, the apex free- ly sulcate near base), 5-20 mm diam., sometimes

Volume 91 Number 4 

2004 

Croat 


whitish, faintly stliate above, bluntly acute to V- Panama. It occurs in wet forests and swampy sites 

shaped (a continucltion of the triangular petiole) be- from near sea level to 780 m in Tropical wet forest 

low, drying brown below, fclintly striate; primclry lat- (T-wd) and Prernontane wet forest (P-wfJ life zones 

eral veins (8)12 to 14 per side, departing midrib at (Holdridge, 1967). 

a steep angle, spreading at a 50°-70° angle, grad- Phenology. Dieffenbachia aurantiaca begins to 

ually spreading in a broad curve, slightly to mod- flower in the early rainy season with flower buds 

erately paler than surface in the proximal one-halL having been seen as early as May and as late as 

weakly raised to convex in valleys, flat and darker August, and mature open inflorescences seen bethan 

surface toward margins, drying mostly paler, tween May and December. Fruiting occurs between 

sometimes darker than surface above, weakly con- September and March. 

vex and slightly paler than surface below, drying Discussion. The species is characterized by its 

yellowish and paler than surface or brownish and more or less elliptic, usually unvariegated, greendarker 

than surface below; interprimary veins pre- drying blades, obtuse to subcordate at base, but 

sent or absent, usually with 1 between each pair of especially by the triangular and weakly sheathed 

primary lateral veins in the lower 1/3 of the blade; petioles, decurrent at apex. It is probably most 

minor veins darker than surface, visible but mod- closely related to D. horichii, but differs from that 

erately obscure below. INFLORESCENCES to 3 species in having a longer free portion of the petper 

axil; peduncle (7)10-18 cm long, acutely an- iole, which is more or less triangular. Also vegegular 

on one side, drying striate, 3-6 mm diam.; tatively similar are D. Iongispatha and D. nitidispathe 

17-25(38) cm long, 2-3.5 cm wide, flatten- petiol(lta, but both differ primarily for thfe same 

ing to 5.7-8 cm wide on tube, 2.5-6.'S cm wide at reasons, i.e., that the free portion of the )etiole is 

constriction, 3-4.3 cm longer than the spcldix, glcld- not tliangulal. 

ually long-tcll)eled towald clpex flom the middle, An unusual (olleetion, Croat 677()(), from a reluniformly 

lip>,ht ;,>reen to medium p>,reen thloughout; atively (lry area on the Pa( ifie slove of Costa lti( a 

weakly glossy oulsi(leX glossy insi(le; S0)clthe lXlcl(lP ((rier thatl thP ty[)i('Al SitC,% for this sy)e( ies), hdS a 

2.S-7 ( m wi(le wherl ficittene(l; .s/)ez(lix 16)-2()(3.-3) )Cti()lP tSldt iS (Ies( I ilve(l dS terete th(ugh ttle ( (lrTl 

lorlp>; [tee l)otliorl to l5.rz ( rTl lorlp>,; listillcile Ie(liotl olkletwise rTlal([les 1). (lllr(lrlti(l((l. '1'S1e (ol- 

})olliorl G..r_t.r(l S.5) ( trl lolly l..5-2 ( rTl (licirDl. 1e( 1 iotl looks vaguely l ike 1). niti(li)( tiol(lt(l whi( h 

tSltotlgllolul (tssotilly dtyirl;,r ()-9 tnm (licim.); lerlile is olhetwise lourl(l only on the Atlanti( slo^)e. A 

StcirTlill lle })ollioll 7-12..S ( rTl lorlp>! wSlile.. [cly)ele( 

tOWcltd cl[)CX atl(l WEclkly tapC,[-e(l S{tgrhtly tOWcltd 

liclse; illtetttle(iclles stelile se;,rtllerlt 1..z-4 ( lYl l(,T, 

Willl cl teW S(cllltste(l lliStilS irl lowet half arl(l a [eW 

s(altele( slamitlo(lia in ul)l)er half; listils 51 to 72, 

iltegularly s(attere(l! with .-3 to 4(6 to 7) a(ross the 

wi(lth of the sl)a(ix Se[atate(l frorn one another lly 

1/2 to 4.0 times their wi(lth, (Iey)resse(l-glol)oseX to 2 

tllm long, 1.2-1.6(2.8) nlm ({iam., I)ale gleen; stigmas 

c ushion-sha)e(l to 2 mm high, 2.tS-3 mm 

wide, white; starrlino(liaS to 5 pel pistil, 1.5-2(3.8) 

mm long, flee ol l)liefly united at l)ase, held well 

above the stigmas sometimes united for Illu(h of 

their length; synandria 1-2(3.4) mm diam., margins 

irregularly angled with lounded, linear to 3-sided 

slit medially at apex. INFRUCTESCENCE 19-24 

cm long; spathe orange outside; spadix 8-15 cm 

long; berries red-orange, B & K yellow-red 6/2.5 to 

B & K yellow-red 7/5 to red B & K red 6/10 (Berlin 

& Kay, 1969), subglobose, ovoid to ellipsoid, 7-10 

mm long. 

Distribution and habitat. Dieffenbachia aurantiaca 

is known only from southwestern Costa Rica 

and adjacent Panama, in the region of the Osa Peninsula 

in Costa Rica and the Burica Peninsula in 

rl'on(l|v; ( olle ( tion at lJA origirlally Ial)eale,(l 70tlelllz 

689 

996/7 WdS t-elal)ele(l 7/77 (Iestite tSle [;d('l 1[lAl il 

was also ( ol 1e( 1e(l i rl Mat ( h 1 4;J9(). 11 aI)I)eat S lo [)e. 

i(letili(al to Iwo othet sI)e( imens 1a1)e1e(1 T0slelllz 

9961, tSle ty^)e ( ol1e(>tiorl num[)er. 

Aelellllotl(ll .s1)()( Irn(l1s ('X(1t71.1/1()(l. (j()S I A 141CA. I t

690 Missouri Botanical Garden


2004 

Croat 


Bartolo Limite, 7 mi. W of Puerto Armuelles, Croat 35079 lateral veins 23 to 30(36) per side, departing midrib 

(MO); Burica Peninsula, 8 km W of Puerto Armuelles, 

at a 70°-110° (to 45° toward apex and sometimes 

Croat 21961 (F, MO); Quebrada Manzanillo, 9 km S-SW 

of Puerto Armuelles, Busey 743 (MO); Quebrada Quana- at base) angle, arising acutely then straight to 

banito beyond La Represa, 2 mi. SW of Puerto Armllelles, weakly curved to the margins (then sweeping prom- 

Liesner 114 (MO, US). 

inently toward apex), usually quilted-sunken above, 

convex, puberulent with thick, whitish trichomes, 

2. Dieffenbachia beachiana Croat 8z Grayum, sometimes with adjacent veins alternating ascend- 

Novon 9: 492. 1999. TYPE: Panama. Bocas ing and descending below; interprimary veins usudel 

Toro: Chiriqui Grande-Fortuna, 13.2 mi. W ally present, scarcely less visible than primary latof 

Chiriqui Grande, 8°45'N, 82°lO'W, 310 m, eral veins; minor veins moderately distinct and 

T B. Croat & M. H. Grayum 60130 (holotype, weakly raised below. INFLORESCENCES 1 to 3 

M0-323065!; isotypes B!, K!, PMA!, US!). Fig- per axil; peduncle 9-13 cm x 5-7 mm, drying 3- 

ures 3, 28B. 

4 mm diam.; spathe 10-19 cm long, gradually constricted 

at middle, green throughout, gradually 

Slender herb, 40-100 cm tall; stems briefly 

long-tapered to apex; spathe blade to 3 cm diam. 

creeping at base; internodes 2-6 cm long, 1.5-3.5 

cm diam., medium green to olive-green, sometimes 

when flattened; spathe tube 1-2 cm diam.; spadix 

13-15 cm long; pistillate portion 4.5-6 cm long, 

streaked with cream, semiglossy and obscurely 

roughened. LEAVES clustered near stem apex, 

drying 7 mm diam. throughout; fertile staminate 

portion 5-6.5 cm long, drying 4 mm diam. througherect-arching; 

petioles 17-46 cm long (averaging 

26.5 cm long), sheathed 25Wo-83Wo of petiole (avout; 

intermediate mostly sterile portion usually 2- 

eraging 54%); sheath 10-22 cm long (averaging 

3.5 cm long, 2 mm diam., sometimes with pistillate 

14.2 cm), decurrent at apex; unsheathed portion 

and staminate portions almost contiguous; pistils 46 

(2.5)10-30 c m long, broadly C-shaped in cross secto 

66(100), 2 to 4 situated across the width of the 

tion, dark green to brownish, flattened with ae ute, 

spadix, ovary oblong-ellipsoid, 1.5-2 x 1-1.6 mm; 

erect margins or sharply to bluntly sul( ate a(laxistigmas 

subgloloular, about as broa(l as the ovary; 

stamino(lia narrowly clavate, usually not at all fused 

ally, surface pale green-mottled, matte, usually minutely 

roughened and with scattere(l scales, whitish 

at t)ase, ca. twice as long as pistil; synandria 1.8raphide 

c e11s visible, drying with s(attere(l clusters 

2.6 nlm diam., margins irregularly sut)roun(ledX 

drying smooth an(l light t)rown at apex. INFRUCof 

pustular raised areas with granular-puberulent 

protrusions; I)l(lde.s narrowly elliptie to lan( eolate, 

'I'ESCENCF, with spathe (10)14-16 cm long, somerarely 

narrowly ovate, inequilateral, one side 1-2 

what flattene(l, yellow-green mottled green an(l 

cm wider than the other side, thinly (oria(eous to 

white with darker flec king, maturing to ol ange; spa- 

(lix 6-10 (m long; berrie.s orange, subglot)ose, 6-8 

subcoriaceous, drying papyraceous, weakly bi(olomm 

diam. 

rous, + equilaterally acuminate at apex (the acumen 

to 5 mm long), slightly inequilateral and acute, Distril)ution (ln(X hfll)itflt. Diegenl)lchifl l)eflchirounded 

or truncate (rarely subcordate in Panama) *lnfl ranges from northeastern Costa Rica to western 

at base, 16-41 cm long, 6.5-15 cm wide (averaging Panama (Bocas del Toro, Chiriqui, and Veraguas) 

28 x 10.7 cm), 1.8-5.3 times longer than wide at elevations of 40 to 800 m. In Costa Rica it occ urs 

(averaging 2.6), 0.7-1.9 times longer than petioles on the Atlantic slope of the Central Cordillera and 

(averaging 1.12 times longer than petioles); margins the Cordillera Talamanca, ranging from the Saracrisped-undulate; 

upper surface dark green (in piqui region to Tortuguero and Siquirres. 

Panama sometimes mottled with white or cream), Phenology. DieJ:fenbflchifl beflchiflnfl initiates 

semiglossy (rarely matte), lower surface semiglossy inflorescences in the late rainy season with plants 

to weakly glossy or matte, slightly paler; midrib flat- reaching anthesis in the dry season (January 

raised above, often striate, usually concolorous through April). Immature fruits have been seen 

above, convex to thicker than broad below, puber- from April to September and mature fruits have 

ulent with thick, whitish trichomes below; prim(lry been seen from April to December. 

Figure 2. Dieffenbachia aurantiaca (Croat 35292). A. Habit, showing open inflorescence.-B. Close-up of spathe 

and spadix (spathe held in a fully splayed position). C. Close-up of leaf showing fully sheathed petiole.-D. Leaf 

cluster with open spathe, showing side view. E. Close-up of portion of spadix with pistils showing stigmas and the 

overtopping club-shaped staminodia. F. Close-up of staminate portion of spadix showing strands of pollen emerging 

from between androecia. 

691

692 

:: A: 

Annals of the 


Figure 3. Diefenbachia beachiana. A. A series of leaves showing adaxial surface (Croat 44283). B. Close-up 

of leaf cluster showing variegated leaves and a cluster of inflorescences (Sims 8). C. Adaxial surfaces of leaves and 

open inflorescences. D. Close-up of inflorescence showing emergent portion of the spadix and adaxial surface of 

petiole. C, D. (Croat 68444).


2004 

Croat 


Discussion. The species is characterized by its Cultivated specinzens. Costa Rica. Henny 5 (M0). Hemoderately 

thin, quilted blades with many broadly 

redia: Finca La Selva, O.T.S. Field Station, oliginally collectecl 

by Jim Beach, cultivated at Missouri Botanical Garspreading 

primary lateral veins and crisped-undu- den (MBG), vouchered as Croat 59152 (M0). 

late margins with the lower midrib and primary lateral 

veins puberulent. DieJJ:7enbachia beachiana is 3. 

similar to D. galdarnesiae7 a species from central 

Panama that also has somewhat puberulent midribs 

on the lower blade sudaces. That species differs in 

having the pubescence much shorter, merely granular-puberulent, 

and has fewer than 22 veins per 

side (vs. 23 to 36 for D. beachiana) that arise from 

the midrib at a 30°-40° angle (vs. 70°-110°). In 

addition, Panamanian populations of D. beachiana 

differ in having their blades usually white- to 

Dieffenbachia burgeri Croat & Grayum, sp. 

nov. TYPE: Costa Rica. Puntarenas: Palmar 

Norte-Panamanian border7 3 km N of turn-off 

to Rincon, 8°48'39"N, 83°16'18"W, 110 m, 10 

Sep. 1996, E B. Croat & D. Hannon 79211 

(holotype, MO-5170493!; isotypes, AAU!, B!, 

CAS!, CR!, COL!, DUKE!, EAP!, F!, GH!, 

HUA!, INB!, K!, M!, MEXU!, NY!, PMA!, 

RSA!, US!, VEN!, UB!, WU!). Figures 4, 27B. 

cream-mottled, whereas those of D. galdarnesiae Planta terrestris, 0.5-1 m; internodia 0.5-2.0 cm longa, 

are always solid green in color. Dieffenbachia be- 1.2-2.5(3.5) cm diam.; petiolus 13-48 cm longus, vaginatus 

1/3_2/3 longitudinis; vagina 8.5-20 cm longa, inequiachiana 

is also somewhat similar to D. grayurni- lateralis acute ad apicem apex; pars libera 5.3-29.5 cm 

ana7 but that species differs in having the veins longa; lamina oblongo-lanceolata vel anguste elliptica, 

merely minutely granular in slender rows on drying, 25-40 cm longa, 5-16 cm lata, nervis primariis lateraliand 

in having somewhat more ovate and wider bus (13)18-28 utroque, inflorescentia l -7 in quoque axblades. 

illa; peduneulus 7.5-12 cm longus; spatha 1].5-15(21) 

cm longa; spadix 9-15.6 cm Songus; piKstilla 21-37. 

A single collection (Cro(t 6695X) from Verclguas 

Province in Panama (lifferx in having the primary Herlg, (). S- 1 m tall, sa, milky, sharply foetid; 

latercll veins curving more prominently towar(l the interno(les glossy to semiglossy, (lrying fre(luently 

apex than rllatericll frolll eIsewhele ill westeln l'cln- lllatte an(l millutely warty, ()..S-2 ( m long, 1.2anla 

and (3oStcl Ri( a. In cld(lilion, the pul)es;( en( e 2..5(.-3..5) ( m (lianl., (lark green to }v1a( kis;h green, 

is nlerely granulclr-l)ul)el ulerlt l ather than pubet u- 

t. H(lltl/tl(l 966()7 (-olle( 

te(l clt l,(l Selvcl in Heledica 

Province in Costcl ltica, ploba})ly repreF;ents 

a hybrid between D. beachiana an(l 1). (on(illn(l. 

The specilllen has tfile sliglltly s( abrous petioles 

an(l the impressed Inajol veins of D. b()el(hieln(l, lxut 

the blades are mole or less elliptic and dry the 

colol of D. concinn(l. 

A*lflltiotl(ll s/)ecilrze/ls S()(M (E()S'I'A 141(CA. Heresflia: 

llfo leje-llfo Sardinalito, Atl. slo^)e of Vol( tirl 13atva, (,r(l- 

rllm bX99 (M()); Sat1 Jose-lvlo. Viejo, vi(. of Chilarrlarlle, 

l 1.6 mi. N of (2aril)larle o, Cro(lt be'.X.58 (}3, Cll, M()); Iq'i1( a 

La Selva, rlear Rfo l'uetlo Viejo, BurW,rer & Stolze 5754 

(CK, 1'^); Zona Plote(tora, N slolles of Vol(an l1arl)a, Rfo 

Peje-Rfo Gua(imo, Grelyum & KSchltz 3220 (DUK1,). Li- 

mon: Rio Pa( uare-Quebra(la l)iablo, ea. 2.5 km 1E ol Si- 

quirres, Grayum 7698 (M0); la. rl'oltugueton Est. Agua 

Fria, R. Robles 1234 (CR, M0); I, Colombiana Farm, 

United FIU jt CO, Standley 36840 (US). Puntarenas: Gol- 

fito, Par. Nac. Esquirlas, Quebrada Gamba, Sanchez 539 

(M0). San Jose: Vazquez de Coronado, Braulio Carrillo 

NP, along hwy. San Jose to Siquilres, trail to Rio Sucio, 

Croat 7X77l (INB, M0, WU). PANAMA. Bocas del 

Toro: Chiriqui Lagoon, Water Valley, von Wedel 1438 

(M0); lO km SW of Chiriqui Grande, Thompson 4937 

(CM); Fortuna Dam-Chiriqui Grande, 7.3 mi. N of bridge 

over Fortuna Dam, 3.2 mi. N of Continental Divide, Croat 

& Grayum 60255 (M0); Chiriqui Grande-Fortuna, 7.7 mi. 

W of Chiriqui Grande, 1.5 mi. W of Punta Pena, Croat & 

Grayum 60094 (K, M0, SC75); near hwy. to Chiriqui 

Grande, McPherson 11816 (M0). Veraguas: vic. of Santa 

Fe, 8 mi. N of Escuela Circlo Alto de Piedia, Croat 66953 

(M0, PMA, SCZ, US). 

693 

Iryirlg (Iark })rown to })Iae k Or yellow-l)rown Or yel- 

low-greerl. I,lq,AVF,% s( attere(l along stenl with areas 

exose(l ex( e)t neal a)eX; /seliol().s 1.-3-48 ( nl long 

(averagi ng 26) ( m long), (Iark green, s;u rf:d( e some- 

W}at rll()ttle(lX weakly r)ale-streake(l neal lease, 

glossy to s;enliglossy, sheathirlg 1/3 tO 2/3 the length 

of the yetiole (().S their average }ength); sheath 8.5- 

2() ( nl long (averaging 12.6 ( m), sometinles soli(l 

(reamy white, with margins often drying thin an(l 

light l)rown, the tiy) inequilaterally ae ute; un- 

sheathe(l portion 5.3-29.5 c m long (averaging 12.9 

cm), variable in cross section, from subterete to C- 

shape(l or U-shape(l, sharply sulcate to somewhat 

flattene(l near apex a(laxially, the margins obtuse to 

acute; I)lade.s oblong-lane volate to narrowly elliptic, 

25-40 cm long, 5-16 cm wide (averaging 30.5 x 

10.5 cm) 1.9-3.7 times longer than wide (averag- 

ing 2.8 times longer than wide), 0 83-1.6 times lon- 

ger than petiole (averaging 12 1 times longer), 

slightly inequilateral, one side 0.4-1.4 cm wider 

than the other side, thinly coriaceous to subcoria- 

ceous, slightly to moderately bicolorous, gradually 

long acuminate at apex, narrowly acute to rounded 

and equilateral or slightly inequilateral at base (one 

sometimes up to 6 mm higher on the midrib than 

the other); upper surface solid dark green, matte to 

weakly glossy (sometimes variegated with pale mot- 

tling), drying gray-green to dark olive-green; lower 

surface slightly paler, semiglossy to weakly glossy



Figure 4. Dieffienbachia burgeri. A, C, D. (Croat & D. Hannon 79211). A. Side view of whole plant showing 

roots, stem, leaves, and inflorescences. B. (Croat & Grayunz 59814). Cultivated collection with open inflorescence. 

C. Close-up of inflorescence with spathe cut open. D. Mature infructescences, one open with berries exposed.


2004 

Croat 


(sometimes matte or weakly glossy on both surfac- life zones (Holdridge, 1967). It occurs in wet forests 

es), drying medium yellow-green to yellow-brown, and swampy sites. 

midrib above flat to sharply sulcate, sometimes flat- Phenology. Flowering in D. burgeri occurs in 

raised, 3-5 mm wide, semiglossy, concolorous, dry- the late wet season to early dry season from Noing 

slightly darker than surface, convex to thicker vember to March, and mature fruits have been seen 

than broad below, convex toward apex, slightly pal- from early April to early October. 

er than surface, drying dark brown to yellow-brown, Discussion. The species is characterized by the 

often acutely l-ribbed, primary lateral veins (13)18 petiole sheathed 1/3z/3 of its length, with the sheath 

to 28 per side, departing midrib at an acute angle, acute at the apex, and the usually matte, greenishthen 

spreading at (40°)45°-70° angle, sweeping drying leaf blades with numerous primary lateral 

along margin, not forming collective veins, sunken veins. The blades are quite variable, ranging from 

to quilted above, convex beneath, drying darker or narrowly oblong-elliptic to ovate-elliptic, oblonglighter 

than surface, the interprimary veins slightly lanceolate or narrowly elliptic. Dieffenbachia burless 

prominent than primary lateral veins, minor geri is most easily confused with D. oerstedii and 

veins moderately close and darker than surface differs from that species in having the petiole 

(drying weakly raised), surface minutely speckled sheath acute at the apex, rather than free-ending 

on drying. INFLORESCENCES 1 to 7 per axil, pe- and rounded to auriculate as in D. oerstedii. In adduncle 

7.5-12 cm x 4 x 7 mm, medium ;reen, dition, D. burgeri has more primary lateral veins, 

moderately glossy (sap with a faint, unusual scent), numbering (13)18 to 28 per side (vs. (4)6 to 9(11) 

spathe 11.5-15(21) cm long at anthesis, 1.1-1.9 cm per side in D. oerstedii). 

longer than the spadix, glossy to semiglossy on both Burger & Liesner 7254 describes the spathe at 

surfaces, flattening 2.7-3 cm diam. at constriction, anthesis as having a strong unpleasant aroma. 

dark green outside throughout, slightly paler inside, Etymology. The species is named in honor of 

spathe blade < 2 cm diam. when furled, spathe William Burger of the Field Museum of Natural 

tube to 10 cm long, 1.5-2.0 cm diam., flattening History, long-time expert on Costa Rican plants, 

3.7-7 cm wide, spathe blade 2.7-3.7 cm diam. at who ma(3e the first collection of the spec ies as well 

anthesis, .a(lix 9-15.6 cm long at anthesis, free as most of the earliest (olle(tions during his exportion 

44.7 ( m long; pistillate portion 5-6 t m peditions t() Costa Ri(a. 

long, the fertile staminate portion 4-6.5 cm long, 

5-6 mm diam., directed slightly forward out of the 

spathe, sterile intermediate segment 1.5-3 ( m long, 

Par(lty/)es. (:()S'I'A R1(:A. Puntarenas: Palmar, Sur- 

(:ha(arita krll 287, (a. 3 km NX of Cha(arita, (rrayllnt et. 

l. 7547 (INf3, MO); Refugio Na(. Gc)lfito, Fila Gamba, 

5 mm diam., with many staminodia near base and 

few just below fertile staminate portion, pistils 21 

to 37, moderately closely spaced, almost (ontiguous, 

to 3 dispersed across spadix width, weakly 

Rio (Carlaza (trainage, Grayllnt & G. Herrera 9236 (CR, 

MO); Fila Barriganes, ca. I km %, 3 km W of Canasas, 

Croat & Grflyunt 59814 (COL, CR, INB, MO, PMA, 

QCNE, SCZ, TEFH, US); Piedras Blancas Rinc6n 3.7 mi. 

W of Pan-Am Hwy., Croat 67692 (MO); Chacarita-Rincon 

constricted below middle, green to pale yellow, de Osa, ca. 6 km W Chacarita, Croat & Grayunz 59731 

stigmas pale yellow, 1-1.5 mm on drying, stami- (CR, MO); Rincon de Osas ridge letw. Quebrada Aparicio- 

Quebrada Aguabuena, Grayunz et al. 4014 (CR, MO); W 

nodia about as long as or slightly longer than pis- of Rincon de Osa, Croat & Grayunz 59851 (MO); Par. Nac. 

tils, somewhat flattened and thickly swollen, sub- Sector Esquinas, YiC. of Fila Gamba hills behind Esquinas 

globular and somewhat truncate at apex synandria Rain Forest Lodge, along Quebrada Negra, at end of side 

drying irregularly somewhat rounded, the margins rd. off of Villa Bricena to Golfito Rd., Croat & D. Hannon 

79286 (CM, INB, MO); San Jose, along rd. to San Isidro 

usually irregularly turned upward, 1-1.4 mm diam. 

del General, Burger et al. 10671 (F, MO); Golfito, Queat 

apex, orange to yellow-orange, green, narrowly brada Negra, Marten 848 (F); Osa, ca. 5 km W of Rincon 

an;led on one side. INFRUCTESCENCE with de Osa, Burger & Gentry Jr. 8898 (CR, F); near the airfield 

spathe usually orange, fruiting spadix 5-7 cm long, ca. 4 mi. W of Rincon de Osa, Burger & Stolze 5489 (F); 

ca. 3 cm wide, berries orange-red to red-orange dry- 

W of airstrip, Utley & Utley 1100 (CR, F). San Jose: 

Perez Zeledon Cant6n, San Isidro General-Dominical, Fila 

ing pale yellow-brown, ovoid, 7-8 mm lon;, seeds Tinamastes, Croat & D. Hannon 79112 (INB, MO, QCNE, 

drying blackened, minutely warty. 

USM); San Isidro del General-Dominical, above Alfombra, 

Burger & Baker 10121 (F, MO). 

Distribution and habitat. Dieffenbachia burgeri 

is endemic to southwestern Costa Rica on the Pa- 4. Dieffenbachia concinna Croat & Grayum, Nocific 

slope from Dominical to the Osa Peninsula, von 9: 492. 1999. TYPE: Costa Rica. Puntarsea 

level to 500 m, rarely 1(300 m elevation, in enas: Palmar Norte-Panamanian border, 3 km 

Tropical wet forest (T-wf) Premontane wet forest (P- N of jet. to Rinc6n, 8°48'39"N, 83°16'18"W, 

wf), and sometimes in Premontane rain forest (P-rf) 110 m, 10 Sep. 1996, 1: B. Croat & D. Hannon 

695



79191 (holotype, MO-5170498!; isotypes, peduncle (2.5)8-17.5 cm long (averaging 10.5 cm 

AAU!, B!, CAS!, COL!, CR!, DUKE!, F!, GH!, long), somewhat compressed dorsiventrally in cross 

HUA!, INB!, K!, M!, MEXU!, NY!, PMA!, US!). section, 3-4 mm wide on drying; spathe 11-25.7 

Figures 5, 28A. 

cm long, 1.5-2.0 cm wide when furled (as long as 

or up to 2 times longer than peduncle and aver- 

Small, erect herb, 0.6-1.3 m tall, sap milky; in- aging 1.6 times longer), constricted 4.5 cm above 

ternodes glossy, 1.5-4(7.5) cm long, 1-4 cm diam., base, the constricted area 1.8-3 cm wide when flatmedium 

green to dark green, obscurely marbled tened, medium to pale green outside, somewhat 

light to medium green. LEAVES erect-arching, pet- darker on tube except white on open face, uniformioles 

7.5-25(35) cm long (averaging 17 cm long), ly paler within, drying dark brown to blackened 

terete and obtusely sulcate near apex or subterete throughout; spathe blade 3.3 cm wide when flatto 

C-shaped or D-shaped in cross section, flat to tened; spathe tube 4.5-8.5 c m wide when flattened, 

shallowly sulcate adaxially (the margins obtuse to paler inside; spadix protruding forward at anthesis, 

acute), rounde( or sometimes acute abaxially, mod- 12-14 cm long; free portion 6.5-9.5 cm long; piserately 

erec t, semiglossy or almost matte, dark tillate portion tapered towal(l apex, 4.5-9 cm x 5- 

green to olive-green, blotched striate or streaked 12 mm, stipe and axis pale green, fertile staminate 

white or silver-green, drying green to sometimes portion white, tapered slighlly toward both ends, 8- 

light yellow-br(wn, sheathing from lower 2/5 to near- 9 cm long, 7-10 mm diam. midway; sterile stamily 

throughout (averaging 0.63 of length); sheath nate portion 2.5-4 cm long; mostly sterile inter- 

6.7-13(20) cm long (averaging 9.5 cm), to 1.5 cm mediate segment 5-7 mm diam., mostly bare but 

diam., margins erect to involute, the tip asylnmet- with a scattering of staminodia on both ends (esrically 

auriculate to rounded and free-ending, with pecially at base); pistils 42 lo 65, closely spaced; 

one side obtuse to rounded, with the other side ovary globose, 1-2.5 mm diam., pale green; stigma 

acute to obtuse; unsheathed portion 1-13 cm long, pale yellow; staminodia 1-5 mm long, up to 3 times 

to 1.5 cm (liall.; blades usually + elliptic, ovate- longer than pistil, somewhal llattened toward base 

elliptic, rarely ovate or broadly lanceolatee 16-36 and free at base, tapering and somewhat globular, 

cm long (averagirlg 23 cm long), (7.8)10-] 5.5(20.5) 1-2 mm diam. at apex; svllandria with flowers ircm 

wide (averaging 12 cm wide),1.5-2.9 times lon- regularly rounded, 0.t


2004 

Croat 


D G 

Figure 5. Dieffenbachia concinna. A-E, G. (Croat 78318). F. (Croat 67594). A. Close-up of adaxial surface of 

blade. B. Crown of plant with inflorescences, one open. C. Close-up of stem and petiole bases showing variegations. 

D. Close-up of opened inflorescence exposing entire spadix. E. Close-up of inflorescence with portion of spadix 

showing elongated staminodia overtopping pistils. F. Close-up of spathe showing emerging threads of pollen filling 

spathe. G. Close-up of male portion of spadix showing androecia with anthers exposed. 

697



size (to 1.3 m vs. 0.75-1 m in D. oerstedii), and Raven 21532 (F), Rinc6n de Osa, Quebrada Apariciomore 

coriaceous blades typically broadest at the 

Quebrada Aguabuena, Gray7lm et al 3982 (CR, MO), vic 

of Boscosa, at Quebrada Aguabuena, Croat & D. Hannon 

middle (vs. broadest below the middle in D. oerste- 79238 (COL, INB, MO, NY, PMA, TEFH, TEX, UB), Secdii) 

with more numerous primary lateral veins (6 to tor Esquinas, vic. of Fila Gamba hills behind Esquinas 

12(14) vs. mostly 6 to 9 for D. oerstedii). In addi- Rain Forest Lodge, along Quebrada Negra, at end of side 

tion, the blades of D. concinna are glossy above rd. off of Villa Bricena to Golfito Rd., Croat & D. Hannon 

79291 (CM, DUKE, INB, MO, P), Refugio Nac. Golfito, S 

and almost rounded at the base versus weakly tributary of Rio Canaza, Grayum et al. 9250 (MO), W side 

glossy to matte and subcordate in D. oerseedti. Fila Gamba, ca. 6 km from Golfito airport, Croat & Gra- 

Grayum et al. 10588, from the headwaters of the yalm 59930 (CM, K, MO), Rfo Claro, 2.5 mi. SE of Golfito, 

Rio Piedras Blancas in Puntarenas Province, is un- Croat 67594 (MO), Orotina-Jaco, valley of Rio Grande 

usual not only in occurring much higher (to 900 m) 

Tarcoles, 1 km S of Quebrada Ganado, vic. Hotel Pink 

Paradise, Croat 79076 (INB, MO), Par. Nac. Corcovado 

than most collections of this species but in having Alonkey Woods, Kernan & Phillips 831 (CR, MO), Lower 

a broader leaf (to 19 cm wide) with a longer petiole Lookout Trail, Kernan 748 (CR, MO); Est. Sirena, Que- 

(to 30 cm). In other respects it agrees with other sada 51 (INB, MO, US, CR), Penfnsula de Osa, Estacion 

collections of D. concinna. Hammel 9660, collected Biologica Los Patos, 8o34'N, 83o31'W, 200 m, 2 Sep. 

1993, R. Aguilar 2195 (CR, INB, MO); 0-1 km W of park 

at La Selva in Heredia Province in Cesta Rica, headquarters at Sirena, Liesner 2871 (CR, MO), Pan-Am 

probably represents a hybrid between D. beachiana E4wy.! Rinc6n, Croat & D. Hannon 79167 (MO), Rinc6nand 

D. concinna. The specimen has the slightly Rancho Quemado, just S of Rincon near Rio Rinc6n, 

scabrous petioles and the impressed major veins of Croat & D. Hannon 79170 (HUA, INB, MEXU, MO, 

D. beachiana, but the blades are more or less el- 

QCA, RSA, WU), vic Palmar Norte, Allen 5669A (F, UC), 

Punta Banco, M. Cha1a(lrrfa et al 257 (CR); Coto Brus, 

liptic and dry more the color of D. concinna. Croat Guaymi Reserve, river trail along Rio Lim6ncito near jct. 

& Hannon 79199, collected south of Palmar Norte, Of Villa Palacios school trail, Koshear 59 (CR), Golfito, 

appears to be a hybrid between D aurantiaca and Refugio de Vida Silvestre, Marten 789 (CR), Carr. Inter- 

D. concinna. Its blade shape and maculations, and am.-Sinai, Fila Huac.as! (a. 4 km NE (3Dy road) at Las 

Huacas (;'Venecia"), (;relum & Evans 10156 (CR, MO); 

numerous inflorescences suggest l). concinna, while Palmar Norte-3alisco, (Juebrada Benjamin, Graymm et al. 

the broadly sulcate petiole favors D. aurantiaca. 9962 (CR, MO). San Jose: above Palmar Norte, Croat 

35708 (MO). NICARA(tUA. Rio San Juan: Res. in{lio- 

Additional specimens examined. COSTA RICA. He- Mafz, Cano el Tambor t)ranch of Rio San Juan, Rueda et 

redia: near Puerto Viejo along rd. near Rfo Sucio, Croat al. 4070 (MO). PA N A MA. Panama: rd. to Puerto 

35688 (MO), Croat 35702 (MO); Finca La Selva, OTS 3imenez, Osa, 40 km W of Panam. Hwy., rt. 2, L. Gomez 

Field Station on the Rfo Puerto Viejo just E of its junction 19531 (MO). 

with the Rfo Sarapiquf, Hammel 9688 (DUKE), Croat Cultivated collections. Costa Rica. Puntarenas: Esqui- 

78732 (INB, MO, W). Limon: La Colombiana Farm, Unit- nas, 25 km SE of Palmar Sur, along Pan-Am Hwy., origed 

Fruit Co., Standley 36739 (US); headwaters of Quebra- inally collected by Bru( e. McAlpin (Selby 85-33), 25 m, 

da Mata de Limon, Finca Anai (Sixaola region), Grayum 2 Jan. 1994, cultivate(l at MBG, vouchered as Croal 

et al. 4447 (MO); Cerro Tortuguero junto a la Barra de 77281 (MO); 15 May 1996, Croal 78318 (MO). 

Tortuguero, R. Robles 2090 (CR, INB, MO), Barringer et 

al. 1973 (F); Par. Nac. Tortuguero, Go'mez-Laurito 7856 5. Dieffenbachia copensis Croat, sp. nov. TYPE: 

(CR); Boca de las Lagunas de Tortuguero, Burger & An- 

Panama. Cocle: E1 Cope, forest on Continental 

tonio 11249 (F, MO); Pococi, Refugio de Vida Silvestre 

Barra del Colorado, SE base Of Cerro del Tortuguero, Gra- Divide above El Cope, 8°38'NX 80°38'W, 700yum 

et al. 11139 (CM, CR, INX, MO, USJ), Rfo Chirri- 900 m, 27-29 Apr. 1985 B. Hantmel 13636 

p6cito-Rfo Sardina, Refugio Nac. Barra del Colorado, Gra- (holotype, M0-3284876!). Figures 6, 27B. 

yum 9777 (CR, MO); Rio Reventaz6n, Finca Montecristo 

below Cairo, Standley & Valerio 48960 (US). Puntaren- Herba 45 cm alta; internodia 7 mm longa, 7 mm diam., 

as: McAlpin 85-33 (SEL); 2 km NW of Chacarita, 30 km petiolus (17)21-29.5 cm longus, vagina S12.5 cm longa; 

S of Palmar Sur, Grayum & Fleming 8119 (CR, INB, MO); pars libera 10.5-15.5 cm longa, lamina anguste oblanceoridge 

betw. Rio Riyito (valley of Laguna Chocuaco) & lata, 25.5-33.3 cm longa, 5.2 cm lata, nervis primariis 

Quebrada Banegas, S of Cerro Rancho Quemado, Grayum lateralibus 9-12 utroque, inflorescenlia 1 per asillam, peet 

al. 7567 (MO); Rinc6n de Osa Rancho Quemado, ca. dunculus 5 cm longus, spatha 15.7 cm longa, 1.2 cm lala; 

10 km W of main Rincdn-Pto. Jimenez Rd., Croat & Gra- spadix 12.3 cm longus. 

yum 59785 (CR, MO); Osa, 6 km W of Rinc6n, Grant & 

Rundell 92-01928 (CR, MO, US); near the airfield ca. 4 

Small herb, to 45 cm tall; internodes about as 

mi. W of Rinc6n de Osa, Burger & Stolze 5461 (CR, F), long as broad, 7 mm diam., drying matte, dark 

Osa Penfnsula, NW of airfield, ca. 5 km W Of Rinc6n de brown, finely and irregularly folded; petioles 

Osa, Burger & Liesner 7196 (CR, F, MO, PMA); Osa Pen- (17)21-29.5 cm long, sheathed 41Wo-56Wo their 

insula, Nicolson 3393 (BM, US); Costena-Cruces, Rfo Pie- length, sheath 8-12.5 cm long, narrowly acute at 

dras Blancas, Cerro Anguciana, Grayum et al. 10588 

(INB, MO, CR); Rio Sandalo, Dodge 10198 (F, MO), apex, drying dark brown to medium yellow-brown; 

Dodge 10020 (F, MO); Palmar Norle de Osa, Allen 5669 free portion 10.5-15.5 cm long, subterete, obtusely 

(EAP, UC, US); Osa Peninsula, 4 mi. W of Rincon de Osa, 

sulcate, drying medium yellow-brown to dark


2004 

f i 

Figure 6. Dieffienbachia copensis (Hammel 13636). Type specimen. 

J 

Croat 


l 

E 

j t. F 

, * 

-/ 

Ar 3C e3 e 

D i ef f snbach 1 a copense CfOtt 

MISSOURI 

699 

BOTANICAL GARDEN 

HERBARIUM 

N° 3284876 

PANAMA *ts2Fi.:-: @s :; 

t'S? s':k-'sX 

Prov . COCL E: 

El Copa. Forest 

above town. 

on continental dvd 

8v-38tNJ 80--38tW. 

700-9OO rn 

Terrestr 1 al 

On slope In frstZ Lvs wavy con mrSn drk 

green aboveJ pale secondery velns 

X mpre s se d abov e. 

hpr/ 27-29/ t 985 

B. Hammel t 3636 

M I SSOUR | BOTAN I CAL GARDEN FIERBAR I UM ( MO }

700 

Annals of the 


brown, minutely granular to somewhat scabridu- free section to 1.1 cm long in D. copensis). Dieffenlous; 

blades narrowly oblanceolate, 25.5-33.3 cm bachia galdamesiae differs in having blades prolong, 

5.2 cm wide, 6.4 times longer than wide, nar- portionately broader, 2.7-4.2 times longer than 

rowly long-acuminate at apex, narrowly attenuate at wide, versus 6.4 times longer than wide in D. cobase, 

dark green and matte above, moderately paler 

and semiglossy below, drying dark gray-green and 

penszs. 

minutely granular above, light yellowish gray be- 6. Dieffenbachia crebripistillata Croat, sp. nov. 

low; midrib concolorous and narrowly raised above, TYPE: Panama. Cocle: Alto Calvario, above E1 

eonvex and granular-puberulent to hispidulous and Cope, ca. 6 km N of E1 Cope, Atlantic slope 

slightly darker below; primary lateral veins 9 to 12 along trail which leads down to Las Ricas, Lipairs, 

arising at an acute angle then spreading at mon 8 San Juan, 8°39'S, 80°3C'GT, 710-800 

45°-50° angle, plominently curved upward along m, 22 June 1988, YS B. Croat 68746 (holotype, 

the margins ancl extending to above the origin of MO-3610884!; isotypes, B!, (2AS!, COL!, 

the next higher veins, paler and weakly sunken DUIkE!, F!, GH!, HUA!, INB!, K!, M!, MEXU!, 

above, convex an(l slightly darker below, drying SY!, PMA!, US!). Figures 7, 27A. 

narrowly raise(l and granular; minor lateral veins 

weakly visible; c ross-veins equally visible on lower Planta 30-100(120) cm alta; interno(libl 2.5-4 cm longa, 

1.5-3(5) cm diam.; petiolus 7.5-24 ( lls longus; lamina 

surface; lower surt:ace moderately granular. INFL0ovata 

vel raro elliptica vel oblongo-elliptica, (22)27- 

RESCENCE sol i lary, longer than the petioles; pe- 46(55) cm longa, 10-25 (m lata, nelvis primariis lateralduncles 

5 cm long (Irying 2.5 mm diam., pale yel- ilous (10)15-17 utroque; inllorescentis 1-2 per axillam; 

low-brown; sp(lth() 1 5.T cm long, 1.2 cm diam., pale peedunculus (8.5)10-14 cm longus; sl)alls{l 13-28 cm longreen, 

clrying me(liurrl yellow-brown, narrowly long- ga, 2.0-3.2 cm lata; spadix 13-26.5 ( lls losgus; parte pistillata 

6-10(12) cm longa, 8-12 mm lll. 

acuminate at tlle tal)ex (acumen 1.7 cm long); spadix 

12.3 cm long; fle e portion 8.5 cm long; pistillate Short, thick-stemme(l herb, 3()-1()()(120) cm tall; 

portion S cm lollg. 4-S mm diam.; staminate portion sap foul-smelling, eventually tulnilltg white; inter- 

5.5 cm long, z1 lllrll cliam., drying yellow-brown; nodes clark green, glossy, 2.5-4 ( lll long, 1.5-3(5) 

mostly nake(1; illte.rmediate sterile portion 1.1 cm (m diam.; petio1Kes 7..S-24 cm long (averaging 16 

long, 2.S mm (lialll. on drying, with only a few sta- e m long), somewhat s)ongy, soli(l (ltark green or 

minodia scattere(t throughout its length; pistils 38, -with a white band on the abaxial surface, conspic- 

2 to 3 situated across the width of the spadix; ovary uously sheathed ranging from 0.7 to fully through- 

0.5-6.0 mm (liarll.; stigmas about as broad as the out the length (but rarely ending mole than 1 cm 

ovary; stamino(lia 2 to 4 per pistil, 1-1.4 mm long, lrelow the blade and often extending beyond the 

slightly thickene(t toward the apex, not at all broad- l)ase of the blade); sheath to 1.5 cm high, 7.5-19.5 

ened toward the l)ase; the sterile male flowers in 2 em long, inequilaterally rounde(l to auriculate at 

rows, somewhal separated from the fertile flowers; apex; unsheathed portion usually not apparent but 

synandria 2.2-2.tA mm long, 1.6-2.0 mm wide, ir- when present 1-7 cm long (averagmg 1.6 cm long), 

regularly rounded to rhombic at apex. INFRUC- t)roadly sulcate, the margins shalp; blades ovate to 

TESCENCE not seen. 

larely elliptic or oblong-elliptic, (22)27-46(55) cm 

long, 10-25 cm wide (averaging 37 x 18 cm), 1.5- 

Distribution (I nd habitat. Dieffenbachia copen- 3.3 times longer than wide (avelaging 1.9 times 

sis is known only from the type specimen in the longer), 1.9-3.7 times longer than petioles (aver- 

Cocle Province of Panama at 700 to 900 m in the aging 2.3 times longer than petiole)* somewhat in- 

Lower montane rain forest (LM-rf) life zone (Hold- equilateral, one side up to 0.5-2.7 crll wider, modridge, 

1967). 

erately coriaceous, abruptly ac u minate and 

Discussion. The species is characterized by its downturned at apex, obtuse to rounde(l at base, the 

narrowly oblanceolate, narrowly long-acuminate basal portion of the blades held solllewhat erect; 

blades with densely granular primary lateral veins upper surface glossy to moderately glossy, usually 

and a gl anular-hispidulous midrib on the lower sur- solid dark green sometimes mottled with pale 

face. The species is somewhat similar to both D. green, slightly bicolorous; lower surface glossy, palgaldamesiae 

and D. fortunensis, differing from both er; midrib flat-raised to flat-rounded and paler esin 

having proportionately narrower blades that are pecially near base, 12-20 mm wide, becoming obbroader 

well above the middle. In contrast, the tusely convex toward apex and diminishing before 

blades of D. fortunensis are broadest well below the reaching the apex above, broadly flat-rounded to 

middle and have the pistillate and staminate por- narrowly rounded and pale green to white below 

tions of the spadix contiguous or nearly so (vs. a (especially near the base); primary lateral veins


2004 

Croat 


Figure T. Dieffenbachia crebripistillata (Croat 75172). A. Potted plant with open inflorescence (Croat 75172). 

B. Close-up of crown of plant with open inflorescence. C. Close-up of inflorescence with pollen emerging from 

androecia (Croat 49243). D. Close-up of inflorescence with tube flared out and spathe blade recurved. E. Closeup 

of inflorescence showing pistillate portion, intermediate sterile section, and lower section of staminate portion. 

701

702 

Annals of the 


(10)15 to 17 per side, sunken to weakly sunken been made during the late rainy season (Augustabove, 

raised below; interprimary veins obscure December). 

above, obscurely visible below; minor veins mod- Discussion. This species is related to D. Ionerately 

distinct on lower surface. INFLORES- gispatha, which differs in being a much more ro- 

CENCES 1 to 2 per axil; peduncle (8.5)10-14 cm bust and taller plant (to 3.5 m), having petioles that 

long, flattened laterally, pale green; spathe 13-28 dry green and with a long unsheathed portion, and 

cm long, 1.3-1.5 times longer than peduncle, pale pistils that are much larger (to 7 mm diam.), many 

to medium green outside (sometimes white at an- fewer in number (only to 26), and widely spaced. 

thesis), paler within, unmarked, + oblong, acumi- See that species for additional discussion of the 

nate at apex, only weakly constricted midway (con- differences. Though D. crebripistillata usually has 

stricted area to 3.2 cm wide when flattened); spathe solid green blades, some plants from Cocle (at La 

tube 1.5-3 cm wide when furled, 8-8.5 cm wide Mesa near E1 Valle de Ant6n and along the Conwhen 

flattened; spathe blade to 2.0-3.2 cm wide tinental Divide north of Llano Grande) have palewhen 

furled; spadix 13-26.5 cm long (averaging 18 maculate leaves. Typically the petiole of Dieffencm 

long), the free portion 8.5-11 cm long; pistillate bachia crebripistillata is fully sheathed; however, 

portion 6-10(12) cm long, 8-12 mm wide; fertile some plants from the Cerro Jefe region (Croat 

staminate portion 5-9 cm long, 6-10 mm diam.; 35915, Croat & Zhu 76644, and Dwyer 9497) or 

the mostly sterile intermediate portion (1)1.5-2.5 the Rio Guanche region (D¢Arcy 9678 and Croat 

cm long, with some sterile male flowers in the up- 11420) rarely have a significallt free portion of the 

per 1/3; pistils (57)80 to 100, scattered, frequently petiole 3-5(7) cm long. 

adjacent, up to 5 in a row across the width of the Collections from lowland (:ol6n (Croat 75172, 

spadix, never more than 5 mm apart; stigmas pale D'Arcy 9678) are somewhat illtermediate between 

orange, densely pul)escent; staminodia white, + ob- D. Iongispatha and the mole tv,nical D. crebripistillong, 

ca. 2-3 mnl long, rounded at apex; synandria lata from the highlands of l'tlllama. The petioles 

irregularly rounde(l at apex, smooth or weakly dim- are not as fully sheathed, witll the free portion of 

pled medially, to 2.6-2.8 mm diam.; pollen exsert- the petiole ranging from 4 to 5 cm long in these 

ed in slender c realll-colored threads to 3 mlll long. specimens. 

INFRUCTESCEN(LIdS with berries ovoid to sul)el- Most collections from E1 (ol)e in Cocle Province 

lipsoid, orange to })right red, 8-11 mm long. have fully sheathed petioles all(l leaf blades proportionately 

smaller and nallower, averaging only 

Distribution sled habitat. Dieffenbachia crebri- 25 cm long (vs. an average of .-37 cm for all other 

pistillata is endemie to central and eastern Panama populations of the species) all(l 2.5 times longer 

at (100)250-800(975) m, in Tropical wet forest (T- than wide (vs. an average of 1.9 times longer) with 

wf), Preniontane wet forest (P-wf), and Premontane petioles averaging only 11..5 ( lll longer (vs. an avrain 

forest (P-li) life zones (Holdridge, 1967) in Co- erage of 16 cm for other po^)ulations). These colcle, 

Col6n, Panama, and San Blas Provinces. While lections (Antonio 3039; (vroat 44558, 44682, 

the species mostly occurs in a few areas of mod- 49155, 67576; Croat & Zhll 76757; FolsonI 1265, 

erately high elevations (including Santa Rita Ridge, 3211, 6218; and Sytsrna & An(lersson 4573) differ 

Cerro Bruja, Cerro Jefe, Cerro Campana, and La in having smaller elliptic bla(les. The E1 Cope re- 

Mesa), it dips down to as little as 100 m along the gion is well known for having tllany ende-mics, and 

Colon Province coast in the area of Rio Guanche these plants may represent differences that may uland 

Portobelo. Most of these collections differ in timately prove to be important, but for now are behaving 

petioles that dry less orange and have the ing considered as D. crebripistillata. One collection 

petiole sheath ending somewhat below the blade, from E1 Cope in Cocle Province (Croat 74861) difnever 

overtopping the blades. As mentioned else- fers from the above only in having the petiole free 

where under D. nitidipetiolata (and below) these above the sheath for a distance of 4.5 cm. 

collections may represent hybrids with either D. ni- Etyrnology. The specific epithet is derived from 

tidipetiolata or D. Iongispatha. 

"creber" (meaning crowded together) and "pistilla" 

Phenology. Dieffenbachia crebripistillata flow- (pistils) and refers to the very closely aggregated 

ers principally at the end of the dry season (April) pistils on this species in contrast to the widely sepand 

in the early part of the rainy season from May arated pistils of D. Iongispatha with which the speto 

July, less commonly during the balance of the cies had long been confused. 

dry season (January-March) and in the latter part 

of the rainy season (August-October). Fruiting time 

is poorly known, but most fruiting collections have 

Paratypes. PANAMA. Cocle: N of E1 Valle, Cerro 

Gaital, Knapp 5758 (MO); E1 Valle de Anton Region, at 

La Mesa, 3.2-6 mi. above E1 Valle, Luteyn & Kennedy


2004 

Croat 


1612 (DUKE), FLORPAN, Guerra et al. 5252 (PMA), Lgewis 

et al. 1753 (M0), Allen & Alston 1839 (M0); La Mesa, 

Finca Mandarinas, along rd. to Finca Furlong, Croat 

67115 (AAU, B, CAS, JAUM, M, M0, P, PMA, TEFH, 

19 Nov. 1984, M. H. Grayu7n G. Herrera, A. 

Matute & E Chavarr^'a 4483 (holotype, M0- 

3229671!; isotype, INS!). Figures 8, 278. 

VEN), 1 km E of Finca Macarenita, Croat 7478 (M0); 

Divide SW of La Mesa, Stein & Hamilton 990 (M0); La 

Mesa, above E1 Valle de Antbn, ca. 2 km W of Cerro 

Pilon, Croat 37361 (F, M0); above E1 Valle on rd. to Cerro 

Pilon, Croat 25352 (M0); 2 km W of Cerro Pilon, Sullivan 

537 (M0); 3 km N of E1 Valle de Anton, Wilbur et al. 

15664 (DUKE); above E1 Valle, Croat 13403 (M0); La 

Pintada-Coclecito, 5.3 mi. N of stream at Llano Grande, 

Atlantic slope, Croat 49243 (HUA, MEXU, M0, PMA, 

SEL, VEN); rd. to Coclesito, 12 mi. from Llano Grande, 

Herba 30-100 cm alta; internodia (1.0)1.5-5 cm longa, 

1.5-1.9 cm diam.; petiolus 3-11.5 cm longus, 5-6 mm 

diam., vaginatus 0.4-0.97 longitudinis, pars libera subteres 

vel D-formata, (1.5)2-5.5 cm longa; lamina oblongoelliptica, 

9-26.8 cm longa, 2.3-9 cm lata, nervis primariis 

lateralibus (6)9-15 utroque; inflorescentia 3 per axillam; 

pedunculus 1.5 I cm longus; spatha W16 cm longa; spadix 

(7.5)10-12.5 cm longus; baccae 8-11 diam.; semina 

2-3. 

Churchill et al. 3993 (MO); past Llano Grande on rd. to 

Cascajal, Sytsma et al. 4398 (M0); Llano Grande-Cas- Slender herb, 30-100 cm tall; sap clear; intercajal, 

Hammel 7212 (M0); vic. Alto Calvario, 7 km N of nodes (1.0)1.5-5 cm long, 1.5-1.9 cm diam., weak- 

E1 Cope, Folsom 3211 (MO), Folsom 6218 (M0), Folsom ly glossy to semiglossy, minutely roughened, usu- 

1265 (M0), Ant6nio 3039 (M0); above E1 Potroso sawmill, 

Cont. Divide, N of E1 Cope, Sytsma & Andersson 4573 

ally with an even mixture of medium yellow-green 

(M0); 9.4 km above E1 Cope, Croat 44682 (MO), Croat and black-green streaks with no obvious back- 

44558 (M0); Alto Calvario, ca. 4.6 mi. above E1 Cope, ground color, sometimes medium yellow-green to- 

Croat 74861 (M0); Alto Calvario, 5.5 mi. N of E1 Cope, ward base with black-green streaks, reversing to 

3.5 mi. N of Escuela Barrigon, Croat 67576 (MO, PMA), 

dark black-green with yellow-green streaks toward 

Croat & Zhu 76757 (HUA, INB, K, M0, PMA, TEX), 

Croat 49155 (M0, RSA, SCZ); Rio Indio, on rd. from Por- the apex, drying obviously minutely wrinkled, stritobelo 

to Nombre de Dios, Croat 33568 (M0). Colon: ca. ate and matte. LEAVES spreading-pendent to pen- 

5 mi. SMt of Portobelo, Croat 14175 (M0, PMA); 2 mi. S dent; petioles 3-11.5 cm long (averaging 6.5 cm 

of Portobelo, Croat 11420 (M0); S approach of Cerro Bru- long), 5-6 mm diam., arched-spreading from stem, 

ja from Rio Escandaloso, Hammel 3140 (M0); 9-12 mi. 

E of Trans-Isthmian Hwy. on Santa Rita ridge, Thompson 

sheathed from 0.4 to 0.97 (averaging 0.73) the 

4874 (CM, K, M0); Santa Rita Ridge, 26 km from Trans- length of the petiole, paler than stem, white to yel- 

Isthmian Hwy., Knapp et 1ll. 1717 (M0); Rfo Gatun, Por- lowish green speckled, especially near the base, the 

tobelo, Antonic) 3820 (M0); Rio Guanche, (a. 3-5 mi. base often white, tinged with pink and dark-striate; 

above bridge inland, Croat 26193 (M0), D'Arcy 9678 sheaths auriculate-prolonged at apex (often appear- 

(M0); (a. 3-5 km above briclge, Croat 36958 (M0); ta. 

3-5 nli. inland, Croat 26239 (M0); tSabanitas-Portobelo, ing + acute on dried specimens), the margins in- 

Rfo Piedras Lumber Rd, 6.T mi. E of Sabanitas, 3.9 mi. rolled on both sides throughout their length, the 

up logging rd., Croat 75172 (M0). Panama: Par. Nac. opposite sides usually in contact and sometimes in 

Altos de Campana, Buena Vista, Chame, FLORPAN, Es- part overlapping; unsheathed portion (1.5)2-5.5 cm 

pinosa et al. 4723 (M0, STRI), Espinosa & E. Martz'nez 

3308 (PMA); Sendero del Tigre, Correa & Montenegro 

long, subterete to D-shaped or broadly and obtusely 

10419 (PMA); trail to summit, Correa & Montenegro sulcate, the margins moderately blunt; blades nar- 

10647 (PMA); Cerro Campana, ca. 1 mi. from Interamer- rowly to broadly oblong-elliptic, 9-26.8 cm long, 

ican Hwy., Croat 35951 (F, M0, MY, PMA, QCA), Croat 

14707 (M0); along trail to summit, Croat 17210 (M0, 

PMA, UB, VEN), Croat 12123 (M0), Croat 12086 (M0), 

Croat 14789 (M0), C. E. Smith Jr. & H. M. Smith 3389 

(F), Luteyn & Kennedy 1837 (DUKE), Hammel 3781 

(MO), Luteyn 3188 (DUKE), Croat 25213 (MO); Cerro 

2.3-9 cm wide (averaging 20.4 x 5.4 cm), broadest 

near the middle, (2)2.74.8 times longer than wide 

(averaging 36 times longer than wide), 1.9-6.2 

times longer than petiole (averaging 3.1 times longer 

than petiole), slightly inequilateral, one side 3- 

Campana, 8.6 mi. SW of Capira, Luteyn 1010 (DUKE); 

6.1 mi. above Pan-American Hwy., Croat 74760 (M0); 5- 

10 km NE of Altos de Pacora, Mori & Kallunki 6026 

(M0); rd. past Altos de Pacora, 3-3.5 mi. NE of Altos de 

Pacora, T.8-8.2 mi. above Pan-Am Hwy., Croat 68699 

(M0); 0.8 mi. beyond turn-off to Altos de Pacora, Croat 

& Zhu 76644 (MO); 4.6 km beyond peak on rd. to Altos 

de Pacora, 26.3 km from Interamerican Hwy., Croat 35915 

(M0); Cerro Jefe, Dwyer & Gentry 9479 (DUKE, M0). 

9 mm wider than the other, sometimes weakly falcate, 

thinly coriaceous, gradually to abruptly longacuminate 

at apex (acumen down-turned and apiculate), 

acute to rounded or cordulate and nearly 

equilateral or markedly inequilateral at base (one 

side sometimes also ending lower on the midrib); 

margins sometimes undulate; upper surface plain 

dark green in Costa RicaX sometimes variegated 

with irregular light green blotches in Panama pop- 

7. Dieffenbachia davidsei Croat & Grayum, sp. ulations, glossy to semiglossy; lower surface slightly 

nov. TYPE: Costa Rica. Limon: headwaters of paler, matte to weakly glossy; drying dark brown to 

Quebrada Mata de Limon, central fork & hills dark gray-brown above, yellow-brown to yellowbetw. 

central & westernmost forks, Finca Anai green below; rnidrib flat at base, becoming weakly 

(Sixaola region), 9°35'N, 82°39'W, 25-40 m, convex toward the apex, concolorous or paler 

703

704 

r 

Annals of the 


Figure 8. Dieffenbachia davidsei. A. Potted plant showing habit with several inflorescences. B. Blade adaxial 

surface with quilted primary lateral veins. C. Close-up of stem showing mottled petiole and stems. D. Crown of 

plant with cluster of inflorescences, and petiole showing free-ending sheath apex. E. Abaxial surface of leaf blade 

and immature inflorescence. F. Close-up of open inflorescence.


2004 

Croat 


clbove, convex and paler than surface below, fre- The species frequently occurs in mature forest 

quently pale-mottled; primary lateral veins (6)9 to along streams. 

15 per side, flat to sunken7 weakly quilted above, Phenology. lliefienb(zchia daridsei flowers in 

raised and darker than surface below, drying usu- the rainy season sometime after May (one plant 

ally darker than surface below, sometimes paler seen with both flower buds and seemingly mature 

than surface, usually departing midrib at 65°-90° infRorescences in lclte October). Mature fruits have 

angle throughout most of its length (rarely with the been seen during the dry season, in the late rainy 

veins in the distal half of the blade arising at 30°- season? and early dry season from November 

50°), prominently curved toward the apex in the through April. Immature fruits have been seen in 

outer 1/3, those near the base to as much as 120° late April indicating that flowering may begin even 

clngle, sometimes forming a weak sigmoid curve, earlier than May. 

the primary lateral veins in the upper 2/3 to 1/2 some- Discussion. The species is distinguished by its 

times loop-connected for some distance, forming a slender, dark, semiglossy, mottled stemsS mottled 

weak collective vein 2-5 mm from margin, but this petioles with a prolonged, usually acute sheath 

usually not extending all the way to the apex; in- apex, and more or less oblong blades with rather 

terprimary veins almost as prominent as primary prominent primary lateral veins that spread at nearlateral 

veins, flat to sunken above, weakly raised ly 90°. 

and darker than surface below. INFLORESCEN- Dieffenbachia davidsei is probably most closely 

CES usually to 3 per axil, sometimes with more related to D. obscurinervia. That species differs in 

than one axil producing inflorescen(es; /eduncles usucllly being cl larger plant (to 1.5 m) with plom- 

1.5-4 em long, 4-5 mm (tiam., ovoi(l to D-shape(l inently scurfy brown internodes, petiole sheclths 

in c.ross se( tion; spathe 8-16 cm long, 3.1.-4.8 which clre tyl)i(cllly only rounde(l clt the clpeX (not 

times longer than pedunc le, e oria( eous! semig]ossy free-en(ling), cln(l [)lclflex which hclve weclkly (leve,lon 

hoth sulfates, me(lium lo light gleen outsi(le, ol)e(l I)rimclly lcltelcll veins thcltcllisecltcln angle, 

slightly 1)a1e1 within; tuT)e olulol, lo someewhcll el- tlstlcllly cll)otlt 4.5° lo lle llli(lril). 

lil.)soi(l, G-7 cm long, ol)lo]g-lfll( volclle wherl All.[o|g[ l[]e [)IbOa(lly sI)Xea(linn l)l'illlal'y lcllPldl 

ol)ele(l, a( llle at apex (a(+lillell tIl)i( lllcilP ['01' 2 rilill); VEillE t11P 011P 0t lllP IllOSl (liSlill('liVP ('llcil('lPliSli(bE 

.Nvelelit (7.5)1()-12.5 e m loly,* uI) lo 1.7 em shollet 0t' 1). el(llpi(l..sei (willl llle VEill.S [lELlcllly S[)l'vtl(lillS 

lhan the s^)clthe; pistillate ^)oltioll (4.()).5..5-() ( m fr0nl lilP nli(lril) cit a l)lOcl(l AnglC PVEI1 to nEcil tilC 

long, (())tS-I() rnm wi(le (the clXiS ('a. 5 t11l11 (licllll.)t A})UX 01 tilP 1)ltl(lU), cl fEw ('OllU('tiOllE [tOtil thP ,1 

alrrlosl ( olltig^.lous with the stalrlill.lle l)oltioll (a I ltino-Carli ltocl(l in l'cinatila (est)e( icilly lhe I'MA 

wi(le stelile illtermeeticlte se(liorl lclekirg); the l)is- XllUPt 01 I(lre(le.s 94()) hcive t)rinicily lciteral veinx 

tillclle [IOWEI S more wi(tely St)ct( C(l llUcil tilC cl[)eXe lhat xl)tel(J cil atlgles of u^) lo 4tS°. 'lnhe lvelreele.s 94() 

4-6 rrlnl cll)cllt; l)istils 2()-.-J.S, ovaly glol)ose, ^)ale olle( tiol] alKso 11as t1101'e (001StVi( uouSly [)unt lEllC 

gleen; stiglna l)right yellow with 10llr l)l ushy l)al)il- ^)etiolex tharl (Jovs lle M() sheet of ll]C SalnC nUtIIlae, 

2-2.4 Illnl diam.; stamino(lia whiteX usuall.y bel. It iXs l)ossil)le that l'(lre(le.s 94() is a hyl)li(l bet)roa(lly 

unite(l at base an(l tolming an often nearly tween l). delviel ei al(l I). ol).scllrinel-vi(l. IXoth sI)ec 

onlplete bowl around the l)istil, somewhat flattene(l ('ies OC('UI in the Nusigandf area along the FJ1 

throughout, barely thickene(l at the a,oex, 3.4-5 mm 1,lano-Calti Road. Dieffent)(lchia ol)sezlrinervia also 

long; stam inate portion oblong 4-6 c m long, has c onspic uously mottled petioles, but its veins 

creamy white, bluntly pointed at apex; synandria arise at a much more ae ute angle. The stems of D. 

irregularly 4- to 6-lobed, drying 1.5-2.3 mm diam., obsezlrinervia differ markedly in being matte and 

deeply depressed with the margins turned up. IN- scurfy-brown rathel than semiglossy, dark black- 

FRUCTESCENCES with spadix 5.5-7 cm lc)ng, green with yellow-green mottling as in D. davidsei. 

berries orange, 8-ll mm diam., subglobose, 2- to Another unusual vollection, possibly also a hybrid, 

3-seeded. 

is Croat & Zhu 76666 from La Mesa in Cocle Province 

near E1 Valle de Anton. It has the glossy in- 

Distribution and habitat. DieJ%enbachia david- ternodes of D. davidsei but the veins of D. obscusei 

ranges from northeastern Costa Rica to Panama rinervia. It is perhaps just an unusual collection of 

(Darien Province) and Colombia (Choco) in Pre- D. obscurinervia. 

montane wet fnorest (P-wf), Tropical wet forest (T-wf), Etymology. The species was first collected in 

and Premontane rain fnorest (P-rf) life zones (Hold- Panama by Scott Mori in 1974 along the E1 Llanoridge, 

1967). In Costa Rica it is known only from Carti road (Mori et al. 4184), and later in Costa 

the Sixaola region at 20-40 m elevation. In Panama Rica by Gerrit Davidse and C. H. Hamilton (Davit 

ranges from 100 to 900 m in Panama Province. idse & Hamilton 23617) in 1983. It is named in 

705



honor of Gerrit Davidse of the Missouri Botanical base, minutely granular-roughened throughout with 

Garden, co-editor of the Flora Mesoamericana se- white raphide cells; unsheathed portion 3-15.5 cm 

ries. 

long, sharply C-shaped, flat-sulcate, with margins 

Paratypes. COSTA RICA. Limon: Quebrada Mata de 

Limon, Finca Anai (Sixaola region), Grayum fst al. 4440 

(CR, MO). PANAMA. Bocas del Toro: Fortuna Lake 

Area, Fortuna-Chiriquf Grande, 1.6 mi. N of Cont. Divide, 

prominently raised; blades narrowly oblong-lanceolate, 

rarely sometimes narrowly ovate, or narrowly 

oblanceolate, 15-33 cm long, (3.5)4.5-9.5(13) cm 

wide (averaging 21 x 6.8 cm), 2-6.3 times longer 

Croat & Zhu 76533 (MO). Cocle: Cano Blanco del Norte- 

Cano Sucio, Davidse & Hamilton 23617 (MO, PMA); E1 

Cope, Cont. Divide, Hammel 13685 (MO); E1 (:ope, ca. 1/2 

mi. N of Cont. Divide at Alto Calvario, Croat 75100 (MO); 

Atl. slope, Hammel 5607 (MO). Darien: Par. Nac. del 

Darien, of Rfo Pucuro, vic. Tacarcuna, ca. 18 km E of 

Pucuro, Hammel et al. 16397 (MO). Panama: E1 Llano- 

Cartf Rd., 9.8 km from Interam. Hwy., Mori et al. 4184 

(MO). San Blas: trail to Cerro Camucanala from Rfo Titamibe, 

de Nevers et al. 4709 (MO, PAN); (:omarca de 

Kunayala, Nusigandl, E1 Llano-Cartf Rd., 10.1 mi. N of 

than wide, 0.38-1.48 times longer than petiole (averaging 

1.1 times longer), + equilateral or weakly 

inequilateral, one side 5-15 mm wider than the 

other side, thinly coriaceous to sulgcoriaceous, drying 

papyraceous, usually quilted, weakly lgicolorous, 

+ equilaterally or sometimes inequilaterally 

acuminate or sometimes acute at apex (acumen 

sometimes lgriefly apiculate), equilateral to slightly 

inequilateral an(l attenuate to acute, rarely rounded 

Interam. Hwy., Paseo Mariska, Croat & Zhu 77017 (INB, at lgase; margins usually conspicuously undulate, 

MO); E1 Llano-Cartf Rd., de Nevers & H. Herrera 3975 drying crisped; upper surface glossy to semiglossy 

(MO); km 18, Galdames et al. 1330 (STRI); Nusigandi, E1 

or weakly glossy, medium to dark green, sometimes 

Llano-Cartf Rd., betw. Nusigandf and 1 mi. N of Nusigandi, 

Croat & Zhu 76597 (HUA, MO); Sede de Campo mottled white or cream; lower surface matte to 

de PEMASKY, ca. 2() km on E1 Llano-Cartf Rd., Paredes weakly glossy, mo(lerately paler; m.idril) flat or flatet 

al. 940 (MO, STRI); Isla Nargana, tributary of Rfo Dia- raised and concolorous above, thicker than broad 

blo, Galdames et al. 1570 (PMA, STRI). Veraguas: Cerro 

Tute, above Alto de l'iedra, McPherson 10725 

to convex below drying lgrownish matte and darker 

(MO). CO- 

LOMBIA. Choco: Nuquf, Corr. Arusf, Est. Biol. E1 Amar- than surface to slightly paler than surface, granulargal, 

along trail to Arusl, Croat & Mor(l 83685 (MO). puberulent to conspicuously granular below; primary 

lateral veins 1() to 17 per side, departing mid- 

8. Dieffenbachia fortunensis Croat, sp. nov. rib at an actlle angle, then moderately curved 

TYPE: Panama. Chiriquf: along road between toward the margin at 55°-90° (mostly 55° in the 

Fortuna Lake and Chiriquf Grande, 4.5-5 km middle of the }lade but at more acute angles toward 

N of dam of Fortuna Lake, 8°43'N, 82°17'W, the apex and more olgtuse angles toward the lgase), 

1100-1135 m, 8 Mar. 1985, E B. Croat & M. turning upwar(l along the margins and forming a 

H. Grayum 60002 (holotype, M0-349012!; iso- series of fine collective veins along the margins, 

type, PMA!). Figures 9, 28A. 

some of which extend to the apex, matte, prominently 

sunken to narrowly or weakly sunken and 

Herba 4s100 cm alta; internodia 1-5 cm longa, 0 8- concolorous at)ove, thicker than broad or convex 

2.0 cm diam., petiolus 12-28 cm longus, vaginatus 0.37- 

0.83 longitudinis; vagina 6.5-14.5 cm longa, decurrens ad 

apicem; lamina oblongo-lanceolata, raro interdum anguste 

ovata vel anguste oblanceolata, 15-33 cm longa, 4.5-9.5 

and darker than surface, granular-puberulent to conspicuously 

granular below, drying darker than surface, 

interprimary veins usually about as conspicucm 

lata, nervis primariis lateralibus 10-17 lltroque; inflo- ous as primary lateral veins, minor veins moderately 

rescentia solitaria; 

12-18 cm longa; 

pedunculus 

spadix 9-11.5 

4.7-7 em 

cm longus. 

longus; spatha 

obscure to distinct; "cross-veins" loose-connected, 

weakly oblique, drying minutely dark-punctate. IN- 

Slender herb, 40-100 cm tall, sap lacking dis- FLORESCENCFJS 1 per axil, peduncle 4.5-7 cm 

tinctly foul aroma, stem erect except at base, con- long, spathe 12-18 cm long, 2-3.7 longer than spataining 

inconspicuous, mThite raphide cells, inter- dix, 1-1.5 cm diam. on tule when furled, gradually 

nodes 1-5 cm long, 0.8-2 cm diam., unmarked, contracted midway, gradually long-tapered to apex, 

semiglossy, dark green to medium green, sometimes uniformly pale green on both surfaces, spadix 9- 

minutely mTrinkled and glistening, drying black- 11.5 cm long, drying 3-3.2 mm diam., free portion 

ened, moderately smooth (lzut minutely granular on 1.5-2 cm long, drying 2 mm diam., pistillate pormagnification). 

LEAVES erect-spreading, + clus- tion 4.24 cm long, drying 7 mm diam. throughout, 

tered at/near stem apex, petioles 12-28 cm long fertile staminate portion 54.5 cm long, drying 4 

(averaging 19.6 em long) sheathing for 0.37-0.83 mm diam. throughout, mostly naked intermediate 

their length (averaging 0.58 their length), sheath portion of spadix 1.5-2.0 em long, drying 2 mm 

6.5-14.5 cm long (averagin; 10.1 cm), with mar- diam., with a few sterile pistillate flowers in the 

gins striate, decurrent distally, dark green, drying lower half, pistils 35 to 38, well-spaced, rarely 

blackened to dark bromTn, surface matte, striate at more than 2 dispersed across spadix width, de-


2004 

Croat 


Figure 9. DiefJenbachia fortmnensis. A, B. (Croat 600021. A. Habit of sterile plant. B. Leaf blade adaxial 

surface. C-F. (Croat 76340). C. Leaf blade abaxial surface. D. Stem showing petiole bases and free-ending sheath. 

E. Close-up of stem showing acutely and broadly sulcate petiole. F. Stem showing internodes and petiole bases. 

707

708 

Annals of the 


pressed-globose, to 1.6 mm long, 1 mm diam., pale mi.l.l, Croat & Grayunz 60346 (MO).Chiriqui: Gualagreen; 

stigma yellow, about as broad as the pistil; ca-Chiriqui Grande, 1 km S of Cont. Divide & Bocas del 

Toro boundary, Croat 66866 (MO);7.2 mi. beyond Los 

staminodia white, 1-1.4 mm long, conspicuously Planes de Hornito, Croat 67837 (MO);8.3 mi. NW of Los 

thickened, drying pale orange-brown; the synan- Planes de Hornito, Croat 49971 (MO); rd. to Fortuna Dam 

dium 2-2.6 mm long, 1.6-2.0 mm wide, white, dry- site, N of Gualaca, 22.7 mi. beyond Rio Esti bridge, Croat 

ing yellow-brown, the margins irregularly angular- 48672 (MO,PMA);4.8 mi. beyond IRHE facilities at dam, 

Croat 68012 (INB, MO,PMA), Croat & Zhu 76340 (MO), 

subrounded. INFRUCTESCENCE with spadix to 6 Thonipson 4951 (CM,MO); Fortuna Dam area, Quebrada 

cm long; berries not seen. 

Los Chorros-Quebrada Hondo, to N of reservoir, Churchill 

& Churchill 6158 (MO), Churchill & Churchill 6159 (MO). 

Distribution (nd habitat. Dieffenbachia fortunensis 

is apparently endemic to Panama, known 

only from the fi ortuna Dam region of Chiriqui Prov- 

9. Dieffenbachia fosteri C l oal, sp. nov. TYPE: 

ince (hence the epithet) at 900 to 1600 m in Pre- 

Panama. Bocas del Toro: l jaguna de Chiriqui, 

montane rain Jorest (P-rf) and Lower montane rain 

Nuri, 15 km E of Punta (Llicamola, Ensenada 

forest (LM-rf) life zones (Holdridge, 1967). 

de Catavela to Quebra(la Nuri, 8°55'N, 

Phenology. Flowering occurs during the late 81°49'W, elevation neal sea level7 20 Mar. 

dry season an(l early wet season from March to June 

1993, R. Foster, A. Herre, F. Kalko & C. Hanwith 

fruits maluling in the late rainy season after 

dley 14649 (holotype, PMA!). Figure 27A. 

September. 

Planta (0.4)1-2 m alta; interno(litl 1. ) ( m longa, 1.5 cm 

Discussion. It is distinguished by its usually diam. in sicco; petio]us 10-29 ( rll lorlrllsX vaginatus 1/3-1/2 

small stature, lzloderately thin, prominently veiny, longitudinis, pars ].it)era subteles; Illlila ovato-elliptica 

undulate, narl owly oblong-lanceolate to narrowly vel lanceolata vel anguste ovaltl, (1 1)17-28 cm longa, 

(6)7-14 cm lata, nervis primblriis ll( alibus 8-13 utovate 

blades willl a flat-raised midrib, and petioles 

roque, inflorescentia I per axilltlll: I)( (lunculus 9.5 cm 

with the sheath (le(urrent at its apex and with a longus; spatha 20 ( n] longa, 2 ( l] (litll.; spadix 18 cm 

long, free, shaluly flattened-sulcate distal portion. longus; pars pisti]].ala ().5 em lollrtl (il] Illl( tu); baccae 6- 

Also charactel ist ic are the granular to granular-pll- 7 mm diam in si(co, I-lo(ulares. 

berulent majot veins on the lower blade surface. 

Slender herb, (().4)1-2 nl t t11; sap smelling of 

The species is c losest to D. beachiana, the latte 

oxalic acid; interno(les medilltzl >XXn, dark greendiffering 

in having the major veins on the lowel 

mottled, 1.5 cm long, drying l.5) ( z1 (liam. LEAVES 

surface cons,oie uously whitish puberulent an(l 

clustered at/near stem apex; )(ltioles 10-29 cm 

blades with ptirllary lateral veins more numerous 

long, drying 2-3 mm diam., 91le itlling 1/3 to 1/2 their 

and spreading c onsistently at broader angles (most 

length; sheath margins dl y i n> + hyaline and 

to ca. 90° throughout most of the blade rather thats 

brown, sheath apex with one si(lej ol)scure to roundonly 

a few VeillS near the base of the blade). 1Xed, 

with the other side acute; unsleathed portion 

terms of pubese ence D. fortunensis is closer to 1). 

of petiole subterete, oval in ( toss section, 12.5galdamesiae, 

whi(h also shares granular puberul- 

18.5 cm from base of blade; I)leldel.s ovate-elliptic to 

ence rather tllan the loose puberulence of D. belanceolate 

or narrowly ovate, (1/1+)17-28 x (6)7-14 

achiana, but I). g(ldamUesiae differs from D. fortucm, 

2-2.4 times longer than wi(le, somewhat inenensis 

in having the primary lateral veins 15 to 17 

quilateral, one side 1-1.5 c l wi(let than the other 

per side and arising at a 30°-40° angle (vs. 10 to 

side, acuminate at apex, the (1( llnen apiculate, 1 

17 per side, an(l at a 55°-90° angle). Dieffenbachi 

mm long, inequilateral an(l oltllse or rounded at 

galdamesiae is also restricted to areas east of the 

base; midrib convex to flat-ltlis( (l (Ind concolorous 

Panama Cana], while D. fortunensis is restricted to 

above, drying somewhat flattene({ alld concolorous 

far western Panama. 

above, convex and densely hisi(llllous, darkerthan 

The species has been confused with small plants 

surface below, drying striate witll white inclusions 

of D. Iutheri, a similar species from Cerro Colorado 

below; primary lateral veins 8 to 1.-3 per side, weakin 

Chiriqui Province. That species usually has widly 

sunken above, raised beneath, (leparting midrib 

er leaf blades (to 15 cm) that are ovate and dry less 

at an acute angle then spreading at 65°-80° in the 

blackened. In addition, that species has the pistillower 

1/2 of the blade, spreading at 45°-55° toward 

late and staminate portions of the spadix contiguapex, 

extending almost straight toward the margins 

ous, and lacks the mostly sterile section between 

then broadly sweeping toward the apex; upper surthe 

pistillate and staminate portion as is present on 

face dark green, matte, drying dark brown to some- 

D. fortunensis. 

what blackened; lower surface slightly paler than 

Paratypes. PANAMA. Bocas del Toro: along Cont. above, weakly glossy, drying slightly less grayish 

Divide rd. N off main Fortuna-Chiriqui Grande Hwy., on black than upper surface, densely grayish-speckled


2004 

Croat 


below. INFLORESCENCES 1 per axil; peduncle 9.5 Small herb, to 50-80 em tall; sap lacking foul 

cm long, 5 mm diam.; spathe 20 cm long, 2 cm odor but weakly caustic to skin; stem supported at 

diam., gradually long-tapered, green throughout, base by adventitious roots; internodes 6-20 x 0.5the 

tube flattening to 3.5 cm wide; spadix 18 cm 2.0(3.0) mm diam., dark green, weakly to moderlong, 

with pistillate portion to 6.5 cm long in fruit; ately glossy; petiole scar oblique, one side 3-4 mm 

berries 6-7 mm diam., dried, l-locular. 

wider than the other; petioles 18-43 cm long (averaging 

29 cm long), densely granular-puberulent, 

Distribution and habitat. DiefJ7enbachia J7osteri 

sheathed 0.25-0.66 their length (averaging 0.43 

is only known from the type in western Panama 

their length); sheath erect to involute, 6.5-20 cm 

near sea level in Bocas del Toro Province in the 

long (averaging 12.3 cm long), inequilaterally 

Tropical moist forest (T-mf) life zone (Holdridge, 

rounded to acute at apex; unsheathed portion 10- 

1967). 

25 cm long (averaging 16.5 cm long), sharply C- 

Phenology. Little is known about the phenology 

shaped, broadly concave adaxially; blades oblongof 

this species, but the type collection was in early 

elliptic,27-60 cm long, 6.7-18 cm wide (averaging 

fruit in March. Flowering probably occurred some- 

40.8 x 11.7 cm), 2.7-4.2 times longer than wide 

time during the rainy season. 

(averaging 3.6 times longer), 1.1-1.8 times longer 

Discussion. Dieffenbachia fosteri is characterthan 

petioles (averaging 1.2 times longer than petized 

by its slender, rather weakly sheathed petioles 

ioles), acuminate at apex, acute at base, inequilaand 

especially by its ovate-elliptic, somewhat 

teral, one side to 1.5 cm wider than the other, thinly 

blackish-drying blades. It is apparently not closely 

coriaceous, drying moderately thin; upper surface 

related to any other Central Americcln species, 

dark green and semiglossy, drying dark brown to 

though in the shape of blades and llle de(urrent 

olive-green; lower surfae e slightly pcaler, dlmost 

sheath clpCX it is most similar to D. killi/)ii. That 

matte, clrying weakly glossy, grayish to yellowish 

spee ies (liffers in having semiglossy l)la(les thclt (I r y 

brown; mi(lri/) broatlly ( onvex-flattene(l clnd ( onmore 

Or less green. It (ould possiluly 1)e confuse(l 

( olorous at)ove ndr r owly r oun(ledX niatte (larker 

wilh 19. grr(lyltmi(lll(l, cl spe(ies ranging alollg the 

cin(l slcirsely grcinulcil-t)ul)el ulent telow (Irying 

(2cllit)i)eclll (oclst of (:ostcl Ri(cl lo westelll l'tlllcllz 

darkel ll-lall SUI'fcl('C! [ltltte,,(l to ,i(lge(l with 0ne or 

all( larotcltely o( ( urling in the alecl wl-lere 1). /o.sleri 

tjolh tnargins tarrowly laise(s; /)rim,(lry l(lt,er(ll Xleill.s 

WclE (olle(te(l; tlowevel; the latler (liffers frolls /). 

1.S Io 17 I)er si(se cirising at .-3()°-4()° angle .sweel)- 

/os/,el-i ill llclvillS llcll10wly ovclte-sut)(ol(late tElcl(sPS 

ing l)rolilinently tow Ir(l tlse ll)ex with as Inally d,% 

ttlclt ale tyl)i( cllly ( olsl)ie uously lclle-X-llcl( ultlte a^s(l 

4 to 5 of thenl silnultaneously (oursilig along the 

also in (ryillg olive-green to yellowisll l)lown rcltte 

rnaigill within 1 ( In of the nsaigirl? somelilnes forinhclrs 

tElclf kete(s. 

ing weak ( olle( tive veins weakly (luilte(l-sunken 

A/ylzloloA,ry. 1'11e sI)ee ies is ncll-lse(l ill [oôl of 

at)ove, ( onvex cln(l .slclrsely glclnulclr-T)ut)elulenl 

one of its (olle(torsX llobin Foster, from the liel(l 

dn(t s()mewhat se u I fy tel(w, (1yillg (Iarkel thcln sur- 

Museuln of Naturcll History in (:hicago, an exl)ert 

f:ae e with the ( entel ( ollal)Se(t an(l the margins ofon 

Neotroi( al plant etology and the vegetalion of 

ten thin and upturned; Illinor veins ohse urely to 

the N votropie s. 

noderalely visible and (larkel than surface above, 

(llying moderately (tistincl; (ross-veins sometimes 

10. Dieffenbachia galdarnesiae Croat, sy). nov. drying moderately distine t below. INFLORES- 

TYPE: Panama. Comarca de San Blas: Pe- CENCFJS to 3 per axil; peduncle 5.5-15 cm long, 

masky, Sen({ero Nergan Igar, km 15 on E1 L1a- 5-8 mm diam., drying 2.5-5 mm diam.; spathe 14tlO-CaI-ti 

Road, 9°20' N, 78°58 ' W, 350 m, 2 25 cm long, 1.6-2.5 times longer than peduncle, 2 

July 1994, C. Galdames, T B. Croat & M. Alba cm diam. at tube, green on both surfaces; spadix 

1222 (holotype, MO-5548504!; isotypes, 15.7-18 cm long; free portion 9.5-12 cm long; pis- 

AAU!, B!, COL!, F!, GH!, INB!, K!, MEXU!, tillate portion of spadix 7.3-10 cm long, 8 mm 

NY!, PMA!, RSA!, S!, SCZ!, UB!, US!, VEN!). diam.; staminate portion of spadix 7-8.5 cm long, 

Figures 10, 28A. 

6 mm diam.; the mostly sterile intermediate portion 

Planta 50-80 cm alta; internodia 6-20 mm longa, 0.5- 1.3-2.5 cm long with a few pistillodes extending in 

2.0(3.0) mm diam.; petiolus 18-43 cm longus, C-formatus, the lower 1/2; pistils 59 to 68, moderately closely 

vaginatus 0.25-0.66 longitudinis; lamina oblongo-ellipti- spaced, 2 to 4 situated across the width of the pisca, 

27-60 cm longa, 6.7-18 cm lata, nervis primariis la- tils; ovaries 1.2-2.0 mm diam.; stigmas 1.6-2.2 

teralibus 15-17 utroque; inflorescentia 3 per axillam; pedunculus 

5.5-15 cm longus; spatha 14-25 cm longa, 2 diam., depressed-globose; staminodia club-shaped, 

cm diam.; spadix 15.7-18 cm longa; pars pistillata 7.3- 1.8-2.8 mm long, 0.4-1.0 mm wide, flattened, free 

10 cm longa, 8 mm diam. 

to base, not expanded toward the base, thickened 

709

710 

Annals of the 


Figure 10. Dieffienbachia galdamesiae (Croat 76560). A. Habit. B. Leaf blade adaxial surface. C. Close-up 

of stem showing roots and petiole bases. D. Petioles and clusters of inflorescences. 

and somewhat granular-puberulent at apex, syn- Phenology. The species has been seen in flowandria 

2.4-2.8 mm diam., irregularly rounded, er in June as well as in October. Mature fruits have 

broadly sulcate to truncate with slightly overlapping been seen in December. 

edges, the margins + crenulate. INFRUCTES- Discussion. The species is characterized by its 

CENCES not seen. 

moderately small habit, only to 80 cm tall, the gran- 

Distribution and habitat. Dieffenbachia galda- ular-puberulent petioles and major veins on the 

mesiae is endemic to Panama, known only from lower blade surface. It is most closely related to D. 

central Panama on both sides of the isthmus in the beachiana and D. fortunensisn differing from both 

Tropical wet forest (T-wf) life zone (Holdridge, 1967) in having the primary lateral veins ascending at 

at 350 to 500 m elevation. It occurs in swampy less than a 50° angle. It may prove to be only subconditions 

in creek beds in virgin forest in nearly specifically distinct from D. fortunensis. 

full shade. 

Etymology. The species is named in honor of


2004 

Croat 


Chilean botanist Carmen Galdames, a long-time broadly flat-raised and striate toward apex) slightly 

resident of Panama, who has collected in Panama paler and dark green-spotted to concolorous above, 

for the SCZ herbarium at the Smithsonian Tropical convex to narrowly rounded and white or narrowly 

Research Institute. Carmen was the first to bring acute and paler below; primary lateral veins 13 to 

the species to my attention and also collected the 18(to 22) per side, gradually arising at a steep angle 

type specimen. 

from the midrib, then spreading in a broad curve 

at 55°-80° angle, (those near the apex to 25° angle, 

Paratypes. PANAMA. Panama: E1 Llano-Carti Rd., 

6.8 mi. from hwy., Croat 49124 (MO). San Blas: Comarca those near the base sometimes 90°-110° angle and 

de Kunayala, Nusigandi, E1 Llano-Carti Rd., 9 mi. N of sometimes forming a sigmoid curve), deeply sunken 

main hwy., Nergan Igar (Nergan Trail), Croat & Zhu above, convex below, forming a series of weakly 

76560 (MO, PMA). Veraguas: Dos Bocas del Rio Calov- developed collective veins that eventually merge 

eborita, 16-17 km NW of Santa Fe, Dressler 5316 (MO). 

with the margin; at least the midrib and primary 

lateral veins sometimes minutely farinose-granular; 

11. Dieffenbachia grayumiana Croat, Novon 9: 

minor veins moderately obscure below. INFL0- 

494. 1999. TYPE: Costa Rica. Limon: Refugio 

RESCENCE 1 to 3 per axil, sometimes subtended 

Nacional Barra del Colorado, forests and pasby 

a reduced leaf with a fully sheathed petiole (the 

tures between Rfo Chirripocito and Rio Sarsheath 

emarginate at apex) and a reduced leaf 

dina l Sardinal], 10°8 ' N, 83°5 ' W, 12 m, M. H. 

blade, 12-15 x 3.5-6.5 cm; peduncle 8-12 cm 

Grayum 9773 (holotype, M0-4370212!; isolong, 

drying 2-3 mm diam.; spathe 16.5-23.5 cm 

type, INB!). Figures 11, 27A. 

long, 3-4 cm longer than the spadix, 4.0-5.0 cm 

Stout herb, 1-1.5 m tall; stems erect at apical wide at base, to 2.5-3.tS cm wide at constriction, 

part, the o]der portion reclining for up to 1.5 m; 2.5-3 cm wide on blade (to 7 em wide when flatinternodes 

2.5-8.tS cm long, 2-3.5(-10) em diam., tened), uniformly light green to medium green on 

(lark green, glossy, variegated with eream-yellow or hoth surfac es, weakly glossy throughout outsi(le, 

pale green (sometimes medium green with (lark somewhat glossier within; )ez(lix 15-27 enl long; 

green lines as in Cro(lt & Grayum 60]4f9); /)etioles free portion 7-1t3 ( rn long; isti]]ate portion 8.0- 

(24-)t3()-59 ( m long (averaging



Figure 11. Dieffenbachia grayumiana. A. Habit (Croat 60149). B-D. (Croat 74953). B. Leaf blade adaxial 

surface. C. Blade abaxial surface. D. Apex of stem showing heavily sheathed petioles with sharply sulcate free 

portion, and speckled petiole bases.


2004 

Croat 


from January to June, rarely as eclrly as November. rante-13ocas del Toro, near Milla 5, Croat & Porter 16499 

Fruiting collections have been seen only from May 

(MO); Chiriqui Grande-Fortuna, 3 mi. W of Chiriquf 

Grande, Croat & Grayum 60149 (K, MO, PMA, US)7 Gualto 

September. 

aca-Chiriqui Grande, 8.1 mi. S of Punta Pena, Croat 74953 

Discussion. The species is characterized by its (MO, PMA). Chiriqui: Gualaca-Chiriqui Grande, 1.4 mi. S 

narrowly ovate, typically subcordate, mottled 

blades; weakly sheathed, decurrent petioles; and 

of Punta Pena, Croat 74945 (MO), M. Akers 78A (MO). 

variegated stems and petioles. Also characteristic 12. Dieffenbachia halnlnelii Croat & Grayum, 

are blades that are frequently glossy on the upper Novon 9: 496. 1999. TYPE: Costa Rica Hesurface 

and matte or nearly so on the lower surface. redia: Finca La Selva, O.T.S. Field Station on 

The major veins are sometimes minutely farinose- the Rio Sarapiqui, 50-80 m, M. H. Grayum 

granular on the lower sudace. In this regard it is 7670 (holotype, M0-3491533!; isotypes, B!, 

similar to D. beachiana: a species with puberulent CR!, K!). Figures 12, 27B. 

major veins on the lower blade surface. Aside from 

pubescence type, D. beachiana also differs in hav- Small herb, 25-40(70) cm tall; sap not fouling 

much narrower blades (1.8-5.3 times longer smelling; stem becoming decumbent and subrhithan 

wide) with more numerous pairs of primary zomatous at base; internodes 1-1.5(2.7) cm long, 

lateral veins (23 to 36 pairs). 

on lower portions, 3.5-7.5 cm long toward the apex, 

Dieffenbachia grayumiana is superficially simi- 0.5-2 cm diam., glossy, dark blackish green, drying 

lar to D. seguine from the West Indies in the shape yellowish brown to gray-green, weakly striate. 

and coloration of its dried blades but lacks the LEAVES erect-arching; petioles 7-25(35) cm long 

sharply sulcate petioles, the bicclrpellclte ovaries, (averaging 15 em long), erect, green or mottled with 

and the protruding stubby spadix with cl reflexed dull yellow-gl efen, (lrying greenish to sometimes 

spclthe b]cl(le seen in the Lltter. 

yCll()WiSh br()Wn, XUltd('8 (11 ying mdttC, XhUdthing 

A ( olle( tion from BO( clS (Je1 r1ôro Provine e in f()r (().



< - s - 

Figure 12. Diegenbachia hammelii. A, E. (Croat 78758). A. Habit of plant in the wild. B. Potted flowering 

plant showing habit (Duke 81-157). C. Potted plant with inflorescence at anthesis (Croat 78731). D, F. (Croat 78731). 

D. Leaf bases and cluster of inflorescences, the one on the right at anthesis. E. Crown of plant with an open 

inflorescence. F. Crown of plant with a cluster of inflorescences (one cut away to expose full-sized berries).


2004 

Croat 


darker than surface or slightly paler than surface erately numerous primary lateral veins. One blade 

below; the interprimary veins almost as conspicu- margin is usually plane and one margin usually mious 

as primary lateral veins; minor veins darker nutely undulate. 

than surface below, drying moderately faint; "cross- In Costa Rica it is most easily confused with D. 

veins" darker than surface below, drying moderate- Oerstedii or small plants of D. grayumiana, both of 

ly faint. INFLORESCENCES 1 to 3 per axil; pe- which can be distinguished in having the petiolar 

duncle 4.5-11 cm long, subterete in cross section; sheath auriculate at the apex rather than decurrent 

spathe (8)10-16 cm long, 1.3-2.7 times longer than as in D. hammelii. At La Selva, where the species 

peduncle, medium green throughout, and weakly was first studied, D. hammelii is uncommon. One 

glossy outside, medium green and glossy inside, collection (Hammel 8784) reported that the sap was 

cuspidate to acuminate at apex; spathe tube 6.5- 

12.5 cm long, 1.2-2.3 cm diam., free portion 3-6 

not foul smelling, as is the case with many species. 

Additional 

cm wide when flattened; spadix 6.5-15.5 cm long; Finca La 

free portion 4.7-7.5 cm long; pistillate portion 5- jo, Grayum 

8.5 cm long, 6-7 mm diam. throughout, fused with Boca de 

specimens seen. COSTA 

Selva, OTS Field Station on 

2772 (DUKE, MO). Limon: 

las Lagunas de Tortuguero, 

RICA. Heredia: 

the Rfo Puerto Vie- 

Tortuguero, near 

Burger & Antonio 

spathe for up to 4 cm; fertile staminate portion ta- 11224 (CR, F), Cerro Coronel, E of Rio Zapote, 1 km from 

pered to both ends, moderately acute at apex, 

banks of Rfo Colorado, Stevens 24257 (CR, MO), 2 air km 

(1.7)3-4.5 cm long, 6-7 mm diam. throughout; sterile 

intermediate segment (0.8)2-3 cm long, with a 

SSE of Islas Buena Vista 

Herrera 31077 (CR, MO); 

Vista in the Rfo Colorado, 

in the 

3.5 

Davidse 

Rfo Colorado, Davidse & 

air km S of Islas Buena 

& Herrera 31154 (MO), 

few scattered staminodia throughout; pistils 26 to Ref. Barra del Colorado [Sardinal], Grayum et al. 9744 

43, ovoid, sparse to moderately dense, loosely scat- (CR, MO); Pococi, Tortuguero, N end of Lomas de Sierpe, 

tered in clusters of 2 to 4 with up to 3 across the 

S from Rfo Tortuguero, Grayum et al. 11169 (CR, INB, 

width of spadix but often with spaces between the 

groups of pistils up to twice the width of the spadix, 

MO). San 

NP, along 

Sucio, Olcl 

Jose: Vazquez de 

San Jose to Siquirres 

Carillo Station site, 

Coronado, Braulio 

Hwy., along trail 

Crot 78758 (COL, 

Carrillo 

to Rfo 

GOET, 

sometimes in a spiral with up to 5 to 6 pistils; ova- INB, MEXU, MO, PMA, TEFH, UB, WU). NICARA(>UA. 

Rio 

ries l-locular, 2.4 mm long, 2.4 mm diam.; stigmas 

San Juall: near Cano Chontaleno, 20 km NE of El 

1.6-1.8 mm diam.; stamino(lia 2.8-3 mm long, up 

to twice as long as pistils, free fronl one another at 

Castillo, 

Sabalos, 

Neill & Vitlcelli W484 

KS8(zlisk 8()§JU (M()). 

(M()); "Marcelo" near Rio 

base, thi(kened at both ends, white at apex, but 13. Dieffenbachia horichii Croat & Grayum, sp. 

often translucent midway and dryillg flattene(l an(l nov. TYPE: Costa Rica. San Jose: Canton Perez 

very thin; synandrium bluntly 4- to S-side(l, mar- Zele(lon, along road betw. San Isi(lro Generalgins 

irregularly shaped towar(l the hase, + rounde(l Dominical, Fila Tinamastes, 9°18'24"N, 

at apex, 2-2.5 mm diam. INFRISCTESCENCE with 83°46'11"W, 900-1100 m, 7: B. Croat & D. 

spathe pale orange outside; spa(lix 22 ( m long; {)er- Hannon 79115 (holotype, MO-05095465!; isories 

orange to bright red, obovoid-ellipsoid, 1 cm types, AAU!, B!, CAS!, COL!, CR!, DUKE!, F!, 

long, 8 mm diam. 

GH!, HUA!, INB!, K!, MEXU!, NY!, P!, PMA!, 

Distribution and habitat. DieJ%enbachia hammelii 

occurs in southeastern Nicaragua (Dpto. Rio 

QCNE!, RSA!, S!, SCZ!, TEFH!, TEX!, UB!, 

US!, VEN!, MIU!). Figures 13, 27A. 

San Juan) and northern Costa Rica from sea level Planta terrestris, 1.0-1.5(2) m; internodia 1-3 cm lonto 

100 m in the Tropical wet forest (T-wf) life zone ga, 4-6 cm diam.; petiolus 8-33 cm longus, vaginatus fere 

omnino; vagina 

(Holdridge, 1967), in wet forests and swampy areas ovato-elliptica, 

on the Atlantic slope. 

mariis lateralibus 

Phenology. Flowering plants of D. hammelii quoque axilla; 

libera 1.5 

2Sb0 em 

1v21 

pedunculus 

cm longa; lamina elliptica vel 

longa, 9-30 cm lata, nervis priutroque, 

inflorescentia 3-6 in 

8.5-19 cm longus; spatha 

have been seen from March through May and also 14.5-32.5 cm longa; spadix 13-17 cm longus; pistilla 43- 

69. 

July, while mature fruits have been seen in August 

and September. Cultivated plants at the Missouri Stout herb, 1-1.5(2) m tall; sap white, copious, 

Botanical Garden flowered in mid July and mid Oc- foetid, caustic; stem erect on younger parts, to 1.2 

tober. 

m long and reclining on older parts, internodes 1- 

Discussion. The species is characterized by its 3 cm long, 4-6 cm diam., semiglossy to glossy, dark 

small stature, typically 25-40 cm; its glossy, de- green to medium green; petioles 8-33 cm long (avcumbent, 

subrhizomatous stems; weakly sheathed, eraging 18.7 cm long), weakly glossy, sheathing 

matte-drying petioles (sheath decurrent at apex); nearly or completely throughout; sheath medium 

and moderately small, more or less oblong-elliptic, green streaked with yellow-green, margins involute, 

weakly inequilateral green leaf blades with mod- the tip free-ending and irlequilaterally rounded-au- 

715

716 

Annals of the 


Figure 13. Dieffenbachia horichii. AX B. (Croat & Hannon 79115). A. Habit of flowering plant. B. Crown of 

plant with cluster of inflorescences. CX DX E. (Croat 79073). C. Plant with inflorescence at anthesis. D. Close-up 

of stem with petiole bases. E. Close-up of lamina base and close-up of inflorescence. 

riculate (auricle sometimes extending up to 1.5 cm nate at apex, + equilateral and obtuse to rounded 

beyond blade); unsheathed portion lacking or to 1.2 (rarely acute or narrowly rounded) at base, margins 

cm long (rarely to 6 cm long), obtusely somewhat weakly undulate; upper surface dark green, semiflattened 

in cross section; blades narrowly to broad- glossy to highly glossy, drying dark gray-green to 

ly elliptic to ovate-elliptic, 26-60 x 9-30 cm (av- dark yellow-brown; lower surface weakly glossy to 

eraging 45 x 19 cm), 1.9-3.4 times longer than matte, moderately paler, drying yellow-brown to 

wide (averaging 2.5 times longer), 1.7-4 times lon- yellowish green; midrib broadly and shallowly 

ger than petiole, slightly inequilateral, one side sunken to flat-sunken above, 5-20 mm diam., con- 

0.5-1.2 cm wider than the other side, subcoria- vex and bluntly low-triangular below, drying light 

ceous to coriaceous, somewhat bicolorous, acumi- brown to dark brown paler than surface below; pri-


2004 

Croat 


mary lateral veins 14 to 21 per side, departing mid- but flowering collections have also been seen in 

rib at a mostly 30°-40° angle above middle, often September. Immature inflorescences have been 

70°-90°, often arising at an acute angle, spreading seen from November to May, and mature fruits have 

to the margins, broadly curved toward apex, even- been seen from the late rainy season (November) 

tually merging at margins, weakly sunken above, to early rainy season (late June). 

weakly convex below, drying brownish and darker Discussion. The species is characterized by its 

than surface below; the interprimary veins usually fully sheathed petioles, involute petiolar sheath, 

darker than surface, 1 between each pair of primary and by its thick and more or less elliptic, semilateral 

veins; minor veins visible, slightly darker glossy to glossy, mostly unvariegated blades that 

than surface, drying moderately obscure on lower dry somewhat greenish to yellow-brown. 

surface. INFLORESCENCES 3 to 6 per axil; pe- Dieffenbachia horichii is closely related to two 

duncle 8.5-19 cm long (averaging 13.5 cm), 3-6 other species with petioles fully sheathed or nearly 

mm diam., drying striate; spathe 14.5-32.5 cm so. These are D. panamensis and D. standleyi. In 

long, 4-5.5 cm diam. (averaging 20.8 cm long), comparison with D. horichii: D. panamensis has 

2.5-4 cm diam. at constriction, 0.9-2.3 times lon- similarly shaped blades and petioles fully sheathed, 

ger than the peduncle (averaging 1.5 times longer but that species differs in having the petiole sheath 

than the peduncle), gradually long-tapered toward usually flaring and recurled rather than involute as 

apex from midway, light green to medium green in D. horichii. It also has leaf blades with the upper 

throughout; spathe tube 3 x 2.3 cm diam. when surface matte and subvelvety that dry blackened. 

furled (flattening 4.0-5.5 cm wide), the constricted Another difference is that D. panamensis occurs 

area flattening 2.5-4.0 cm wide; spathe blaele 2.5- principally on the Atlantic: slope of central Panama, 

4 cm wide at anthesis; spadix 13-27 cm long (av- whereas D. horichii occurs on the Pacific slope of 

eraging 17.4 c m long); free portion 10-13.5 ( m Costa Rica. 

long; pistillate portion to (6.5)8-10.8 cm long (av- DieJ/en/)lchi(l horichii is likely to I)e (onfuse(l 

eraging 8.3 c m long), 8-17 mm wide, drying 8 mln with D. .st(ln(lleyi: another sr)ee ies with fully or 

diam.; feltile staminate )ortion 8. 3-12 cm long, mostly winpr,(l Xetioles. That sI)ee ies oe ( urs; on the 

c rearm-( olore(le mo(lerately tapere(l towar(l aI)ex an(l Atlantie slole of Hon(luras an(l Nie arapr,ua, an(l (lifweakly 

talxere(l towar(l baxe, 7-12 mln (liam. mi(l- fers lxy 11avinpr, the [)etiole slleatll eree t an(l re( urle(l 

way; mostly xtel ile interlne(liclte segnlerlt 1 .7--3.7 outwar(l alont, the margins an(l a(ute at the ay)ex 

cm long, witll a few xcattele(l xtamino(lia through- with the sheath margins marke(lly un(lulate (vs. tlle 

out; pixtilx 4.-3 to 69, irregulal Iy scattere(l, neal Iy sheath malgins illvolute an(l smooth). 'I'he letioles 

contiguouse .-3 to 4(6 to 7) (lisl)erse(l acloxs xla(lix of l). .ft(lMdleyi are also longer on average, frequentwidth? 

xelalate(l from one another by ul) to 4 tinlex ly more than 25 c m long, an( average 3() (m long 

their width; ovary depresse(l-globoxeX 2 mm long, (vs. fre(uently 1ess than 25 c rll long, averaging less 

yellow-green; xtigma cushion-S;hape(l, 2.8-3.4 mm than 2() ( m long for l). horichii). Tn a(i(itionX the 

diam., about twice as wide ax thick and usually upper midrib on the l)lades of D. stclol(lleyi is broa(lbroader 

than the pistil at anthesix, yellowish; xta- ly concave, whereas on D. horichii the midril) is 

minodia white, 3 to 5 per pistil, 3-4 mm long, free l)roadly (onvex with a medial sulcus lout with the 

or briefly united at base; synan(lria 1.6-3.0 mm entire midrib sunken in a valley. 

diam., subrounde(l, depressed medially at apex, Most of the typical material of D. stan(lleyi has 

drying orange-brown. INFRUCTESCENCE 19-24 been collected in the Lancetilla Valley and its vicm 

long; spathe orange outsi(le; spadix 8-15 cm cinity in HoncAllras, and has blades considerably 

long; berries red, subglobose, ovoid to ellipsoid, 7- longer on average than those of D. horichii. Some 

10 mm long. 

collections in Nicaragua and in western Costa Rica 

are unusual. Stevens 7457 from Zelaya and Moreno 

Distribution and habitat. Dieffenbachia horichii 17142 from Matagalpa dry a darker yellow-brown. 

is known only from the Pacific slope of Costa Rica The only other species in Central America that has 

from the Carara reserve and Puriscal region to the a fully sheathed petiole is D. tonduzii. Dieffenbach- 

San Isidro region and Dominical. It occurs in Pre- ia tonduzii is distinguished by having smaller and 

montane rain forest (P-rf) and transition forests be- thinner blades (rarely to 45 cm long and 20 cm 

tween Tropical wet forest (T-wf) and Premontane wide) that are usually matte to weakly glossy above 

rain forest (P-rf) life zones (Holdridge, 1967), from (vs. usually glossy in D. horichii) with more primary 

sea level to 900 m. 

lateral veins (18 to 25 vs. usually fewer than 20 in 

Phenology. Dieffenbachia horichii begins to D. horichii). Grayum 4757: from the Carara Reflower 

in the early rainy season from May to July, serve in Puntarenas Province, is perhaps a hybrid 

717



between D. horichii (Grayum 4756) and D. oerstedii sheath 8.5-21 cm long (averaging 14 cm), with 

(Grayum 4765). It differs in having petioles that are margins drying thin, light brown and minutely unnarrower 

and less fully sheathed. 

dulate, the tip inequilaterally acute to emarginate 

Etymology. The species is named in honor of and free-ending; unsheathed portion flattened or 

horticulturist Clarence Horich, who made the first rounded and becoming weakly sulcate toward apex 

collection of the species. 

in cross section (never sharply sulcate), blunt to 

Paratypes. COSTA RICA. Puntarenas: San Jose 

Province, Playa Dominical-San Isidro del General Baru, 

'rinamastes, Burger et al. 10669 (F, MO), Burger & Baker 

10137 (CR, F), along Quebrada Bonita, Carara Res., Grayum 

et al. 5723 (INB, MO), Finca E1 Eden, km 183, R. 

2, ca. 400 m E of Santa Marta, L. G6mez 22951 (B, CM, 

CR, MO); Quebrada Bonita, Carara Res., Grayun 4756 

(CR, MO), Croat 79073 (EAP, INB, MO, PMA), hills at 

SW part of Montanas Jamaica, ca. 2.5 km NE of Bijagual 

moderately acute, rarely broadly and bluntly sulcate; 

blades oblong-ovate to narrowly ovate, (12)15- 

30(39) cm long, (6)10-26 cm wide (averaging 26 

x 16 cm), 0.78-1.77 times longer than petiole (averaging 

1.1 times longer), inequilateral, one side 

0.5-2.6 cm wider than the other side thinly coriaceous, 

abruptly to gradually acuminate at apex, 

acute to cordate at base, the sides often + unequal, 

de Turrubares, Carara Res., Grayu1m et al. 5467 (MO). usually subcordate with at least one side subcor- 

San Jose: Par. Nac. Braulio Carrillo, Quebrada Sanguijuela, 

Chavarrfa & Untana 157 (CR, MO), Zona Prot. La 

Cangreja, vic. Quebrada Grande, ca. 2 km NNE of Mastatal 

de Puriscal, Grayun 8638 (CR, MO), ZP La Cangreja, 

ca. 1.5 km E of Santa Rosa de Puriscal, Grayun et al. 

8336 (CR, MO), San Jose rd. from Parrita to Santiago de 

Purriscal, Barringer 1790B (CR), Res. Biol. Carara-Est. 

Quebrada Bonita, Chac6n 1406 (CR), Cordillera Talamanca, 

Rfo Hermoso, Finca E1 Quizarra, L. Willianes et al. 

28479 (F, NY, US); Acosta, along Rfo Parritilla, ca. 1 km 

date at base; upper surface matte to weakly glossy, 

dark green, drying dark gray-green to sometimes 

blackened; lower surface matte, paler, drying yellowish 

gray-brown to dark yellow-brown; midrib 5- 

8 mm diam., flat to broadly and obscurely sunken 

at base, weakly raised toward apex above, concolorous 

to slightly paler than surface, frequently 

much paler than surface or even white toward apex, 

E of Zoncuano, Grflyum el al. 11174 (INB, MO); Perez 

Zeled6n, Fila Tinanlaste, Valverde 741 (CR, MO); Puriscal, 

Fila Tufares, Salitrclles de Puriscal, Goncez-Laurito 7792 

(CR); Z.P. La Cangreja, Cerros de Puriscal, San Martfn de 

Puriscal, La Fila Vala Blanca, J. Morales 2035 ((21t, INB, 

MO). 

Cultivated specimens. Costa Rica. 800-900 z1, cultivated 

at Munich as 1203/74, Horich s.n. (M). 

often with a light green streak distally! drying paler 

than surface or darker than surface al)ove, convex 

to bluntly round-raised, drying somewhat flattened 

with acute ribs below, drying darkel than surface 

below; primary lateral veins 7 to 12(15) per side, 

departing midrib at a 45°-55° angle (to 30°-50° at 

apex, 50°-90° at base), arising a( utely, then 

14. Dieffenbachia isthlnia Croat, sp. nov. 

TYPE: Panama: along trail betw. Rfo Maje & 

Quebrada Brava, 60 m, 4 May 1976, 1: B. 

Croat 34656 (holotype, M0-240198!; isotypes, 

B!, K!, PMA!, US!). Figures 14, 28A. 

spreading weakly or not at all sunkell often greenish 

toward apex, sometimes raise(l 1leal midrib, and 

diminishing toward margins above; lllinor veins obscure 

above, obscurely visible to not visible below. 

INFLORESCENCES straight to sliglltly curved, 3 

to 5 per axil, bracteoles 9-20 cm long; peduncle 

Planta 

2-3(4.5) 

plerumque 

cm diam., 

ad 1 m, 

atroviridia; 

internodia 

petiolus 

0.5-3 

11-34(40) 

cm longa, 

cm lon- 

(2.5)4-12.5 cm long, 6-10 mm diall., sometimes 

flattened on one side in cross sec.ti(-)n, green; spathe 

gus, vaginatus 0.4-0.77 longitudinis, vagina 8.5-21 cm 15-17.5(23) cm long, medium green outside, slightlonga, 

lamina oblongo-ovata vel anguste ovata, (12)15- 

30(39) cm longa, (6)10-26 cm lata, plerumque subcordata, 

nervis primariis lateralibus 7-12(15) utroque, inflorescentia 

3-5 per quoque axillam; pedunculus (2.5)4ly 

paler green inside throughout, except the apical 

portion white at anthesis, to 1.7 c m lollger than the 

spadix, elongating somewhat after closing; spathe 

12.5 cm longus, spatha 15-17.5(23) cm longa, spadix tube less than 2 cm diam. when furled, spathe 

10.5-15.8 cm longus; pars feminea 6.5-9.5 cm longa, 1.4 blade 2.5-3 cm diam. at anthesis; s)


2004 

Croat 


Figure 14. Dieffenbachia isthmia (Croat & Zhu 77115). A. Leaf blade adaxial surface. B. Close-up of petioles 

showing inflorescence. C. Close-up of female portion of spadix showing pistils and staminodes. D. Infructescences, 

one cut off to show the fruits. 

719

720 

Annals of the 


larly somewhat rounded, margins often irregularly & Croat 906 (DUKE), Croat 15173 (MO), Croat 5819 

turned upward. INFRUCTESCENCE with spathe to 

(MO), Croat 6308 (MO), Croat 6502 (MO, PMA), Croat 

4576 (MO), Killip 39979 (US), Kenoyer 188 (US); 12 mi. 

23.5 cm long, usually orange; spadix, 9-12 cm S of Col6n on Rio Providencia, Tyson &: Blurn 3998 

long, ca. 3 cm wide, berries orange to red or orange- (PMA); Frijoles vic., Pittier 3754 (US), Bailey 335 (F); 

red, drying pale yellow-brown, ellipsoid, T-8 mm Gatun Lake, Standley 41107 (US), Standley 40960 (US), 

long, 6 mm diam. (the stigmatic area ca. 2 mm AIaxon et al. 6812 (US), Maxon et al. 6820 (US), Standley 

31266 (US); Rio Paraiso, above East Paraiso, Standley 

wide); seeds drying dark brown, 6 mm long, 5 mm 29867 (US). Colon: Rio Guanche, ca. 5 km upstream 

diam., to 3 mm thick, moderately smooth, caved from rd. to Portobelo, Harernel &: Trainer 14767 (MO). 

outward on funicular side. 

Darien: PN Cerro Pirre, vic. Rio Perisenico, Croat &: Zhu 

77115 (INB, MO); 112 mi. E of Bayano Dam Bridge, vic. 

Distril)utio71 (ln(l habitat. Dieffienbachia isthmia Cangl6n, Antonio 4546 (MO, PMA); trail from Cana to 

ranges from Panallla to Colombia (Antioquia and Colombian border alollg Rio Setgandi, Gentry et al. 28574 

Choco). The spee ies is highly variable ecologically, (COL, MO); E1 Real-l'inogana, Duke 5014 (MO); 3 mi. N 

occurring )rin( i)ally in Tropical moist J7orest (T-mf) 

of Santa Fe, Tyson et (ll. 4631 (SCZ), 4632 (MO, SCZ); 2 

mi. E of Santa Fe, Tyson et al. 4834 (STRI); PN del Daand 

drier parts of Premontane wet J7orest (P-wf) life rien, between Rio Tol)alisa 8 Rio Pucuro, ca. 17 km E of 

zones at 50-8()()(1 ()()()) m elevation, but also in Pre- Pucuro, La Laguna alecl, Harernel et al. 16262 (CAS, COL, 

montane moist /or()st (P-mf), Premontane wet forest MO, PMA); airstrip at Cana gold mine area, Croat 38010 

(P-wf), and Tro/)ie (ll wet forest (T-wf) life zones (MO), 38057 (MO); S of E1 Real, Alturas de Nique, near 

Cana mine, rd. to Boen de Cupe, McPherson 11591 (MO); 

(Holdri(lge, 1967) in Colombia. In Panama, the Cana region, ca. 1.5 klrs from Cana trail to Boca de Cupe, 

species ranges fX om Veraguas Province to the McPherson 15037 (M()). Herrera: NZZ of Las Minas, near 

Azuero Peninsultl ill the west (700-900 m in Her- Chepo, on Montosa (Je Chepo, McPherson 10958 (MO); 

rera an(l I os Salltos I'rovinces). 

Dist. Las Minas, Ch(\|o, loma E1 Montuoso, Galdarnes et 

al. 1626 

Phenology. It'lowering for D. isthmia oecurs in 

(MO, PMA, US); Las Minas, base of E1 Higo, 

Galdarnes et al. 248f) (I'MA, US). Los Santos: Loma Priethe 

early rainy se.lsion from May to September (rare- ta, Cerro Grande, I(l(i.s et al. 2195 (COL, DUKE, MO, 

ly in Novellll) l )^ with fruits maturing during the UC). Panama: Disll ilo (:hepo, Puerto Coquira, Zarnbrano 

dry season arl(} etltly rainy season of the following & Delgado 1336 (PM \); San Jose Island, I. Johnston 1165 

year, mostly ill Al)lil and May, but with many col- (GH); vic. Bayano llk( dam near Canita, Gentry & Tyson 

1653 (MO, PMA); (Jllilllcin Lewis et al. 3251 (MO); 3.8 

lections ma(le i ll I ru i t from August to Oe tol)er. mi. E of Rio Ipeti, louel slopes of Serrania de Maje, HuJ2t 

Discus.siosl. '1'11e species is characterize(l by its & Jacobs 1997 (M()); Itfo Majo-Quebrada Brava, ca. 2 mi. 

moderately small (usually less than 1 m tall and upstream from watell'tllls near edge of Bayano Lake, Croat 

with stems usually less less than 2.5 cm diam.) 34745 (MO); Isla 13tlstlno, Garibaldi 68 (MO); near Chihabit 

and frequently subcordate blackish-drying 

man, ca. 2 mi. up lwlo La Maestra, Kennedy 1193 (F). 

Veraguas: near plol)osed route of rd. from E1 Cortezo to 

blades which often have a white streak on the distil Arenas, Harernel 5.X72] (MO); Azuero Peninsula, trail behalf 

of the flattened (not flat-raised) midrib. In Pan- tween Jobero and lleadwaters of Rio Pedregal, Croat 

ama, it is probal)ly most easily confused with D. 34475 (COL, F, M(), I'MA); 18 km NZZ of Las Minas, Cerro 

killipii, which differs in having proportionately nar- Alto Higo, Harernel 12(98 (MO); "Los Girasoles," Escuela 

Agricola Alto Piedltl ( a. 5 km NW of Santa Fe, Dressler 

rower (typically only to 16 cm), usually subcordate 4716 (DUKE, F, M(), I'MA); 18 km W of Las Minas, Cerro 

greenish-drying blades with the uppel midrib Alto Higo, Harernel jU98 (MO); Cerro Delgadito just NW 

weakly flat-raised rather than merely flattened of Cerro Tute, S of Fiarlta Fe, Luteyn 4043 (DUIkE); Dist. 

above. 

Montijo, Isla Coiba, (erro de La Torre, Galdarnes et al. 

2286 

Though its range does not overlap with D. isth- 

(MO); Isla Coila, Rio Escondido vic., Galdarnes et 

al. 2252 (MO). COI ()MBIA. Antioquia: Mutata, Bajiramia, 

D. oerstedii may be confused with this species. Nuevo Oriente RoafJ, Brand & Ascanio 277 (COL). Cho- 

The latter, ranging from Mexico to central Panama, co: Riosucio, PN Natural Los Catios, Campamento de Tildiffers 

in having a usually sharply sulcate petiole upo, Forero et al. 1723 (COL, MO). 

and blades that dry greenish to yellowish green 

rather than blackened as is usually the case in D. 15. Dieffenbachia killipii Croat, sp. nov. TYPE: 

. I - 

IsthUmla. 

Panama. Cocle: vic. of E1 Valle de Anton, La 

Etymolog;y. The species epithet ;;isthmia" re- Mesa, forested flat area near Finca Macarenita, 

fers to its dominance in the area of the Isthmus of 8°36'N, 80°0T'M1, 800 m, 6 July 1994, YS B. 

Panama. 

Croat & G. Zhu 76666 (holotype, MO- 

04612287!, isotypes, AAU!, B!, CAS!, COL!, 

Paratypes. PANAMA. Canal Area: Barro Colorado 

Island, Aviles 24 (MO), Shattuck 397 (MO); Lutz Trail, CR!, DUKE!, F!, GH!, HUA!, INB!, K!, M!, 

Croat 5378 (F, MO, RSA, SCZ, STRI), Croat 10133 (MO), MEXU!, NY!, PMA!, SCZ!, US!, VEN!, WU!). 

Croat 5317 (MO), Croat 5896 (MO), Croat 7712 (MO), Figure 15, 28B. 

Croat 11291 (F, NY, SCZ), Croat 5183 (MO), Croat 10982 

(MO, SCZ), Croat 5709 (MO), Foster 865 (DUKE), Luteyn Planta terrestris, 40-100 cm.; internodia 1.5-5.5 cm


2004 

Croat 


Figure 15. Dieffenbachia killipii. A. Habit showing plant with quilted primary lateral veins (Croat 56902). B. 

Leaf blade adaxial surface (Croat 78247). C. Close-up of adaxial surface of blade (Croat 74759). D. Habit of 

flowering plant (Croat 76666). E, G. (Croat 75154). E. Close-up of stems. F. Habit of flowering plant (Croat 76666). 

G. Close-up of two inflorescences, one at anthesis. H. Close-up of pistillate and sterile staminate portions of spadix 

(Croat 76259). 

721



longa, (0.8)1.5-3(4) cm diam; petiolus (4)6-20124.5) cm vex, slightly paler to concolorous above, thicker 

longus, vaginatus 2/5 longitudinis vel fere omnino; lamina 

than broad and narrowly rounded to almost roundoblongo-elliptica 

vel oblongo-ovato, (13)19-30(33.5) cm 

longa, (4.5)7-16(21.3) cm lata, nervis primariis laterali3ous raised, matte, paler than surface to almost conco- 

8-12 utroque; imSorescentia 1-4 in quoque axilla; pedun- lorous below, drying brown with ridges, darker than 

culus 4.5-9 cm longus; spatha 12-19 cm longa; spadix surface below; primary lateral veins 8 to 12 per 

12-15 cm longus; pistilla 20-37. 

side, arising at an acute angle, then spreading at 

mostly (40°)45°-70° angle (rarely to 30°, sometimes 

Medium-sized herb, 40-100 cm tall, stem creepto 

80° toward base, rarely to 110° at base), freing 

over surface of ground at base, then erect; sap 

quently forming collective veins that merge with the 

milky, unscented; internodes initially weakly glossy, 

margin higher up on the blade, narrowly sunken to 

often faintly dark green and medium yellowish 

weakly quilted-sunken above, thicker than broad to 

green-marbled at lower nodes, becoming glossier in 

convex and weakly pleated-raised, darker than surage, 

1.5-5.5 cm long, (0.8)1.5-3(4) cm diam., meface 

to almost concolorous below, usually drying 

dium to dark green or olive-green or black-green, 

darker than surface; minor veins few, obscure 

drying dark yellow-brown to orange-brown, rarely 

above, obscurely visible and darker than surface to 

dark brown, epidermis sometimes fissured in a 

moderately distinct below. INFLORESCENCES 1 

cracked network so as to appear corky in some arto 

4 per axil; bracts 9-20 cm long; peduncle 4.5-9 

eas. LEAVES scattered along stem, denser near 

cm long, 6-7 mm diam., medium green, white at 

apex; petioles (4)S20(24.5) cm long (averaging base; .s/)athe 12-19 cm long at anthesis, 1-2 cm 

12.2 cm long), firm to spongy, slightly paler or dark- longel than the spadix, medium green throughout, 

er than stem, medium green to dark green, matte sometimes faintly dark green-lineate on faded areas 

to weakly glossy, faintly striate toward the base, outsi(le, slightly paler and glossy inside; spathe 

drying greenish to grayish yellow to brown, sheath- tube 2-3 cm diam. when closecl, 5.5-7.5 cm wide 

ing for 2/, to fully throughout (0.4-1 the petiole when [lattened, 1.5-2 cm (liam. at constriction (flatlength 

and averaging 0.69); sheath 3-18 cm long, tening 3.5-4 cm wide); spathe bla(3e 2.5-3 em 

(averaging 8 ( m), with the sheath margins not dry- dianl.; spadix bluntly pointe(lX weakly protruding 

ing markedly different than the remainder of folwal(l at anthesis, 12-1tS (m long; free portion 

sheath; sheath apex with the tip free-ending and 5.5-() ( m long (sometimes with a few l)istillate flowinequilaterally 

acute to emarginate, sometimes dry- ers itl the basal portion); pistillate portion 5-8.5 cm 

ing acute; unsheathed portion C-shaped and sharp- long. 1 0 mm diam. throughout; fert i le staminate 

ly sulcate to narrowly and sharply sulcate to sub- portiotl 4.5-6.8 cm long, 7-8 mm diam. midway, 

terete and weakly sulcate or obtusely and narrowly sometimes bluntly pointed (frequently with the 

sulcate in cross section; blades oblong-elliptic to witheled portion weakly protruding out of the front 

oblong-ovate, rarely narrowly ovate, (13)19- of the spathe after anthesis); sterile intermediate 

30(33.5) x (4.5)7-16(21.3) cm (averaging 25 x 11 seglllent to ca. 5 mm diam., but usua]ly absent with 

em), 1.5-4.2 times longer than wide, as long as or the )istillate and staminate portions alrnost conti;up 

to 4.8 times longer than petiole (averaging 2.3 uous; pistils 20 to 37, well-spaced, sometimes agtimes 

longer than petiole), inequilateral, one side gregated into weak rows, frequent Iy irregularly 

0.5-1.5 cm wider than the other side, sometimes gapy)ed, 2 or 3(6) dispersed across s^)adix width, 

falcate, subcoriaceous to weakly coriaceous, mod- widely spaced at base and at apex, )ale yellowerately 

bicolorous, acuminate to gradually acumi- green, 2.4 mm diam., 1.4 mm high; stigma 1.8-2.2 

nate at apex inequilateral, sometimes inequilater- mm diam.; staminodia very thickene(l and mostly 

ally rounded to subcordate, rarely acute at base; joined at base, tapered gradually toward apex and 

margins moderately straight on one side, frequently not markedly thickened, sometimes broadened latmarkedly 

undulate on other side; upper surface se- erally and apparently consisting of a union of 2 

miglossy, dark green, frequently white, pale green staminodia, sometimes with 2 pistils contiguous 

or yellowish green-spotted or white-streaked, dry- and apparently sharing staminodia; synandria 1.2ing 

dark brown to olive-brown or gray-green; lower 1.4 mm diam., ca. 4 per spiral, irregularly rounded 

surface paler, matte to weakly glossy, drying yellow- to 4- to 6-sided, drying widely spaced, the margins 

brown; sinus less than 1 cm deep, rarely to 2.5 cm of apex markedly turned upward. INFRUCTESdeep; 

rnidrib flat-raised, 3-5 mm wide, sometimes CENCE to 23 em long; spathe orange outside; spasulcate 

toward base, usually in moderately deep dinc 9-10.5 cm long, 2.S em wide; berries red to 

valleys, usually concolorous, sometimes paler than reddish orange or orange-red, drying pale orangesurface 

aboveX sometimes weakly 3- to a-grooved brown, ellipsoid, 2- to 3-lobed, 7-8 mm long, S 

on upper surface, drying flat-raised to broadly con- 10 mm diam.; seeds 1 to 2 per berryS drying dark


2004 

Croat 


, . . 

brown, flattened on funicular side, to 4.9 mm long, 50480 and Plowman & Davis 4094, both collected 

4.4 mm diam., drying smooth, 3 mm thick. along the Baeza to Tena Road, at 1410 and ca. 

1743 m, respectively, and Croat 72573 south of 

Distribution and habitat. Dieffenbachia killipii Coca at 300 m. An unvouchered collection made 

ranges from southwestern Costa Rica (Puntarenas) by Tan, Halton, and Besse at the Auca Oil Field at 

to Panama and the western slopes of the Andes in 240 m is also the same species. It has been vouch- 

Colombia (Antioquia, Choco, Tolima, and Valle) ered from the Marie Selby Botanical Gardens' livand 

Ecuador (Esmeraldas, Guayas, E1 Oro, Pichin- ing collection (SEL 79-0090) by Plowman 14121 

cha, Los Rios, and Manabi on the Pacific slope and and Ingram 1124. 

in Napo on the Atlantic slope) at 0 to 900 m ele- Gentry et al. 65306, from Quindio Department, 

vation. In Panama it occurs in Tropical wet forest Colombia, at 1400 m, may also be this species, but 

(T-wf) and Premontane wet forest (P-wf) life zones it is from higher elevation than other Colombian 

(Holdridge, 1967) and ranges from Cocle Province collections. A collection made in Venezuela at Maito 

Darien Province in the east. In Colombia it is quetia at 30 m near the Caribbean (Andre' 457A) 

known from Tropical wet forest (T-wf) and Tropical appears to also be D. killipii. However, this may be 

rain forest transition to Premontane wet forest (T-rf/ questioned because Andre 457 was labeled as hav- 

P-wf) life zones. 

ing been collected in Colombia at Angostura de 

Phenology. F8lowering occurs in D. kill ipii Honda (Tolima). Both collections were dated Dethroughout 

the year, with most flowering specimens cember 1875. 

having been made in the late rainy season from In South America the species may be confused 

September to November. Fruiting collections were with D. enderi, described from the valley of the Rfo 

made mostly during the late dry season and early Cauca in Colombia. That species is similar in havrainy 

season of the following year from May to July. ing the petiole sheath auriculate and free-ending at 

Few fruiting collections were Kseen between AuguKst the apex, and in having the staminate and pistillate 

and May. 

spadives (ontiguous. Tt diffels in having a thicker 

Di.scu.s.sion. The specieKs is chara(terized by its stem (to 8-9 cm diam.), longer blades (to 50 cm 

medium stature (to 1 m), slen(ler sulcate petioles long) with ae ute hases and more numerous primary 

with the Ksheath inequilaterally auli(ulate at apex lateral veins (( a. 24 lel side, spae e(-l 1.5-2 ( m 

(an(l uxually ending well helow the l)ase of the apalt). Tn (ontraKst, L). killi/)ii has hla(lex that are 

l)la(le) an(l espe(ially ly the flequelltly Ksul)cor(late roundeel or sul)( or(3ate at the hase, ;an(l fewer than 

bladeKs (whie h dry yellowish green) with a flat- 15 primary lateral veins. It is also similar to D. 

raised mi(lrib. Also unuKsual for the sl)e(ies is the elelguellsi.s from the valley of the Rio Dagua in Valle 

near absenfe of a Ksterile portioll between the pis- Department, whie h similarly has staminate and l)istillate 

and staminate spadie es. 

tillate zones ( ontiguous on the spa(lix. That spee ies 

Dieffenb(lchiel killipii is similar to L). Ieo)oldii, eliffers in having larger hlades (to 40 c m long) with 

whieh waKs descIibed from a cultivate(l plant of un- many elose primary lateral veins (up to 25), an(l a 

known specific origin, believed to have been ob- petiole only 5 cm long. 

tained in Colombia. Engler saw living material of Dieffenbachia killipii is probably most easily 

this species and illustrated it in his 1915 revision. confused in Central America with D. isthmia, which 

While it has very similar leaves, D. Ieopoldii differs differs in having stems that dry usually blackened; 

from D. killipii in having the pistillate and stami- somewhat more ovate, blackened blades; and a spanate 

portion of the spadix separated by a sparsely dix with a more or less sterile portion between the 

flowered, almost sterile section. In addition, while staminate and pistillate portions. The species may 

the upper surface of D. killipii is semiglossy, D. also be confused with D. oerstedii from Costa Rica 

Ieopoldii is described as having a blade "velvety and western Panama. That species shares leaves of 

green" on the upper surface in the notes of N. E. 

Brown (on specimen of Lehmann 1052), at least 

similar shape and size, but it has blades that are 

matte, rather than semiglossy, on the upper surface 

suggesting that the upper surface was matte. Fi- and dry greenish or yellowish green. In addition, 

nally, the midrib of D. Ieopoldii is described as D. oerstedii has petioles that are typically more 

white, whereas this is not the case with D. killipii. sharply sulcate than those of D. killipii. The petiole 

Collections from the eastern slopes of the Andes base is green in D. killipii and whitish in D. oerin 

Napo Province, Ecuador, appear also to be this stedii. Furthermore, the spadix has staminate and 

species but more studies are needed to confirm this, pistillate portions nearly contiguous in D. killipii, 

as they would be the only collections of D. killipii but separated by a distinct sterile portion in D. oerin 

the Amazon basin. The collections include Croat stedll. 

723

724 

Annals of the 


In Panama the ranges of D. oerstedii and D. Dil- at Rio Agua Salud, Croat 12353 (MO); Rio Frijol on Pipelipii 

do not actually overlap, but come close in Chi- line Rd., 6 m N of Gamboa, Tyson 1443 (FSU, MO, SCZ); 

Frijoles-Monte Lirio, Killip 12154 (US); N of Frijoles, 

riqui on the western slopes of the country. All Standley 27413 (US); W of the Canal, near Gatun, Stanknown 

collections there occur in the mountains at dley 27224 US); rd. S-10 N of Escobal near junction with 

900 to 1300 m. Dieffenbachia oerstedii occurs in Rd. S-1, Croat 12489 (MO); 1.5 mi. N of Escobal, Croat 

Panama in western Chiriquf Province at highland 12491 (MO, SCZ); near Limbo Gun Club camp, 10 mi. W 

of Gamboa on Pipeline Rd., Lazor & Tyson 3492 (FSU); 

sites and with an outlying population at E1 Cope in 

lake shore along Gatun River valley, Pittier 6845 (US); 

Cocle Province, while D. killipii ranges in general vic. Gamboa, Pittier 2600 (US); Pipeline Rd. N of Gamno 

further west than the Azuero Peninsula and Ver- boa, Kennedy 455 (F). Chiriqui: 8.8 km past Gualaca on 

aguas Province in Panama. There is an unusual rd. to Chiriquf Grande, Hoover 1324 (MO); Gualaca-Foroutlying 

population of D. killipii in the area of the tuna Dam site, 2.8 mi. beyond Los Planes, Croat 48816 

(MO); Cerro Colorado, 15.6 mi. above Ri) San Felix, Croat 

Osa Peninsula in Costa Rica (Kennedy 1594). A 48437 (MO). Cocle: E1 Limon, Merl(li(ta 1-10 (PMA), 

collection from Cerro Colorado (Croat 48437) in Mendieta 1-101 (PMA), Mendieta 1-121 (I'MA); vic. of E1 

Chiriqui Provin(e has blades slightly larger than Cope, PN E1 Cope, 5-6 mi. N of 191 (3ope, below Old 

most specimens of the species. 

Rivera saw works area, Croat & Zhu 76746 (CAS, DUKE, 

MO); 5 hours' walk N from Alto Calvalio to Rio Blanco, 

Croat 74785) from La Mesa in Cocle Province, 

Sytsma et al. 2414 (MO); La Mesa, al)ove E1 Valle de 

Panama, is unusual in having a series of rather Anton, Croat 14388 (MO), Croat 14X()(9 (MO); trail beyond 

prominent collee tive veins and in having the minor La Mesa towards Los Llanos and the l)ol(ler between Cocle 

veins distine t when fresh (typically they are rather and Panama Provinces, N of E1 Valle (It Anton, Luteyn 

obscure). It WilEe in fact, so unusual that it was ini- 

3175 (DUKE), Croat 37383 (MO); I ,(l Me sa above E1 Valle, 

Gentry 7423 (MO, NY); 5 mi. N ol El Valle de Anton 

tially mistakell for a Xanthosoma. It is, however, Luteyn 1203 (DUKE); La Mesa trail towtlt(ls Cerro Cara 

within the (legree of variation for D. killipii. coral, NE of E1 Valle de Anton, Lut(yl1 S/80 (DU1EtE); vic. 

Kress 77-8.X() and 77-831 from the vicinity of E1 Valle, Bartlett & Lasser 16678 (MlX,XlJ, MO); Penono- 

Santa Fe in Velaguas, Panama, are unusual collec- me-Coclecito, 5.6 mi. N of Llano Clallde, along Rio Cascajal 

5.6 mi. N of Llano Grande, 1.4 Illi. 1N of Cont. Ditions 

in havi1g the staminate and pistillate portions 

vide, Croat 67480 (MO, SCZ); at L.l Mesa! 3.2 mi. above 

of the spa(lix sie)arated by as much as 1 cm. In E1 Valle, 0.1 km E of Finca Macalelsita, Croat 74789 

this regard tlley are similar to D. Iutheri, but differ (MO), Croat 74792 (INB, MO, I'MA)* Ctl. I km W of rd. 

from that sl)e( ies in lacking the granular puberu- betw. Finca Mandarinas and Fin(l l7tlosr, Croat 67197 

lent major veills on the lower blade surface. (MO); area between Cano Blan(o (lel Norte, Cano Sucio 

and Chorro del Rio Tife, Davidse & llolll illon 23599 (MO); 

Some colle( tions of the species from Darien rd. to Coclesito, logging camp 12 1lsi. 1'l0lil Llano Grande, 

Province, Panama (e.g., Antonio & Hah77 4405, Churchill et al. 4032 (MO), Churallill (1 (tl. 4125 (MO), 7 

Duke 15591, lvolar7>co 1485, and Schmalzel 1212), km N of E1 Cope, near Rivera SawXlille /olsom & Collins 

are unusual in having petioles more fully sheathed, 6436 (MO); La Mesa, above E1 Valle (Je Ant6n, ca. 2 km 

W of Cerro Pil6n, Croat 37479 (M(), I'MA, WU); base of 

sometimes to less than 1 cm from the base of the 

Cerro Pil6n above E1 Valle, Gentry & I)ll1^ ( r 3643 (DUKE, 

blade or even with the petiole sheathed throughout. F, MO). Colon: PN Chagres, See( iols lJo(luer6n, Rio San 

The Costa ltican population of D. killipii is note- Juan de Pequeni, Espinosa et al. S()l l (I'MA), Espinosa 

worthy in that the single collection made there in & Guerra 3762 (PMA), Espinosa (t *X.l. 4478 (PMA); Dist. 

the Osa Peninsula is quite disjunct from the nearest Donoso, Campamento Botija, J. I' )le.leo et al. 1905 

(PMA); Rio Providencia, 12 mi. S ol (olorl, Tyson & Blum 

population in Panama, where the species has not 3954 (FSU, SCZ); Rio Guanche, betlwe X ls I'uerto Pil6n and 

been collecte(l west of Cocle Province. The species Portobelo, ca. 1.5 mi. S of rd., Cro(lt & %hu 76253 (MO, 

is to be expet ted at other sites in Veraguas and in PMA), Croat & Zhu 76259 (MO, XlX l,, S(2Z); lower Rio 

Chiriqui Prov i nce, Panama. 

Guanche, Dressler 4688 (PMA); N ol l{fo (Juanche, Dav- 

Etymology. The species is named in honor of 

idse & D7Arcy 10098 (MO); Rfo (,tlull( he, 3-5 km above 

bridge on Col6n-Portobelo Rd., Cro(lt 7(9329 (MO), Croat 

the late E. P. Killip, botanist at the U.S. National 79359 (MO), Sytsma 1658 (MO); rlear Peluca, km 25.6 

Herbarium and one of the more prodigious plant from Transisthmian Hwy. on rd. to Norrll)re de Dios, Kencollectors 

in the Neotropics, who was one of the nedy 2661 (MO); ca. 8 km E of Pina, Thompson 4816 

earliest collectors of this new species. 

(CM, MO); Portobelo-Nombre de Dios, 1.2 mi. beyond the 

jet. of rd. to Isla Grande, Croat 49805 (INB, MO); Nuevo 

Paratype.s. COSTA RICA. Puntarenas: Osa Penin- Tonosi-Rio Indio, Portobelo-Nombre de Dios, Croat 33551 

sula, 2.5 mi. SW of Rinc6n, Kennedy 1594 (MO). PAN- 

AMA. Bocas del Toro: Cerro de Bocatorito, Peterson & 

Annable 6768 (MO). Canal Area: 12 mi. S of Col6n, Tyson 

et al. 4486 (SCZ); 12 mi. S of Col6n, near Rio Providencia, 

Tyson & Blum 3997 (MO, SCZ); Pipeline Rd. 10 

mi. from Gamboa gate, Croat 15082 (DUKE, MO), ca. 7- 

8 km N of Gamboa, XKnapp 2275 (MO); Pipeline Rd. near 

(AAU, MO, QCNE); Achiote, McPherson 9176 (MO); Santa 

Rita ridge rd., ca. 22 km from transisthmian hwy., Hammel 

et al. 14472 (MO). Darien: Quebrada Biboto (Peccary) 

off Rio Areti, Duke 13601 (MO); 1-3 mi. N of Paya, 

Duke & XKirkbride 14019 (MO); Rio Cocalito, Whitefoord 

& Eddy 136 (BM, MEXU, MO); PN Cerro Pirre, vic. headquarters 

on Rio Perisenico, Croat & Zhu 77102 (COL, 

Gamboa, Clewell & Tyson 3306 (MO, SCZ); Pipeline Rd. HUA, INB, L, MO, NY, QCA, SCZ, TEFH, US, VEN);


2004 

Croat 


Paya-Pucro, Stern et al. 433 (GH, MO, US); Punta Guayabo 

Grando to Rio Jague, Antonio & Hahn 4405 (MO); 5 

of E1 ISlano-Carti Rd., Schatz 1079 (B, K, MO, US). Veraguas: 

rd. beyond Escuela Agricola Alto Piedra, NW of 

mi. W of Yaviza, Schmalzel 1212 (MO), Schmalzel & Al- Santa Fe, Pac. slope, 0.6 mi. beyond fork in rd., Croat 

verson 1199 (MO, PMA); vic. of gold mine at Cana, Croat 

37605 (HUA, INB, K, MO); S of E1 Real, headwaters of 

Rio Pirre at fork known as Dos Bocas, Kennedy & Foster 

395 (DUKE); Pinogana-Yaviza, lDuke 5169 (MO); 10 mi. 

49056 (MO); Escuela Agricultura above Santa Fe, Kress 

77-830 (DUKE), 77-831 (DUKE). COLOMBIA. Antioquia: 

Rio Anori valley, Quebrada La Tirana-Providencia, 

28 km SW of Zaragoza, Alverson et al. 376 (WIS). Choco: 

S of E1 Real on Rio Pirre, Duke 5437 (MO); trail from Rio Quibdo-Istmina, 6.6 km S of Quibdo, Croat & Cogollo 

Pucro to Quebrada Maskia, Duke 13082 (MO); Rio Tucuti 52174 (COL, MO); ca. 10-15 km S of Quibdo, and 8-10 

upstream ca. 2 hrs. (piragua) vic. Tucuti, Duke 5260 (MO); km E, Grayum et al. 7657 (MO); Sautata, PN de Los Ka- 

Punta Guayabo Grando-Rio Jaque, Antonio & Hahn 4406 tios, Renterza 10680 (HUA); Rio Atrato, Tagachi, Forero 

(MO); Cerro Canlpamento, S of Cerro Pirre, Duke 15591 et al. 9007 (COL); Bahia Solano, Corr. E1 Valle, Quebrada 

(US); Rancho Frio, halfway up slope of Cerro Pirre from 

Piji Vasal, Folsom 6247 (MO); gold mine at Cana, Sullivan 

745 (MO); Bajo Lepe, 7 km al SE of Boca de Cupe, Po- 

Tundo, tributary of Rio Valle, Espina et al. 2902 (MO); 

San Jose del Palmar, Rio Torito, Finca "Los Guaduales," 

Forero et al. 6272 (COL), 6489 (COL). Tolima: Angostura 

lanco 1485 (PMA); base camp Cerro Pirre NP, Croat de Hondo (Magdalena), Lehmann s. n. (K). Valle: Bajo 

68961 (COL, MO, PMA); PN del Darien, RENARE sta- Calima Region, Buenaventura-Malaga, km 11, Croat 

tion, McDonagh et al. 433 (BM), 439 (BM, MO); Est. Biol. 69343 (CUVC, MO); Pulpapel Headquarters, Bay 278 

Rancho Frio at N base of Cerro Pirre, ca. 9 km S of E1 (MO); 5 km N of main Cali-Buenaventura Hwy., Croat & 

Real, Quebrada Perisenico, de Nevers et al. 8267 (CAS, Bay 75707 (MO); Cali, Vereda Pico de Aguila, Gamboa 

MO); Rio Coasi, Cana-Coasi trail, R. Hartman 12488 (K, et al. 71 (MO); Rio Dagua, Buenaventura, Lehmann 5311 

MO, PMA); Rio Pavarando, 10 km NE of Jaque, D7Arcy (K). ECUADOR. Cotopaxi: Guayacan-Montenegro, N of 

& Sytsma 14498 (MO); Rio Pirre, 2.5 nli. above E1 Real, Pucayacu, Croat 73792 (QCNE). Guayas: Cordillera 

Duke 5099 (MO); Rio Torti, 38.6 mi. E of Bayano Dam Chong6n-Colonehe, Cerro La Torre, Cornejo & Bonifaz 

Bridge, neal Torti, Antonio 4638 (MO); Rio Tuquesa, Cle- 3107 (GUAY). E1 Oro: ca. 11 km W of Pirlas on r(l. to 

zio 221 (MO), 226 (MO); Rlo Cllito, frorrl Yaviza at junc- Arenil las, Thompson 16() (MO); 7.6 km f l om Tahuin on 

tion with Rfo (2hucunaque to (a. 1 hl. I-vy outboar(l Irom 

jet., Burcll e/ *11. 1123 (DAV, M(), NY); (a. 1() krrl upstream 

Irorzl Na%areJtll, H(lArl /42? (1X, M()); Rfo Ja(luee Valley, 

K/l(l)/) & M(lll(l W()89 (M()); l{fo 13t11sas, Kllr.s(lr & 

ld. to 1'ie(3ras, 7'hom/).son 1E?8 (MO); Machala-l,oja, 25 kr 

AF: of jur( tiorl irl rd. to Pirlas, (,zro(lt T()72.T (M(), QCNIq,). 

Eslll+ralflas: (Leerro Mutilees, (,zornejo & Boni/(lz T1(31 

((,UAY); Saralo l)orlirgo (lee los (3olora(los-Esrzleeral(3as, t3() 

CSol(^^ 4 (S(iC), /.T (S(jXJ). Pallallla: I'irin-(2arasas trail kls NW ol Sarllo l)orlsirlgo, tS.tS krls NW ol ()uirlirs(lee ( ro(l.l. 

neear l'il ia, Olzke 14X4f) (M()); I1JI VAllE (Je Ma(Jrorlo-lJa T.ST4(J (M()); () krls beeyorlcJ l)i(lge oveer l{fo Estreral(las 

Savla, 2.S Illi. N 01 I1JI Valle (JeX Mtl(JIorlo ;3.S rlli. N 01 (11e11 Sarl Maleoe rcJ. to I1JSI}lPIAI(IAS ;II)OrI) (t1. ().() kGlL 

turrloll lo St1ll Jose, 1 1.6 ti. N ol 1 as Mal>arilas, vic. 

(2helasX valle y ol ltfo Marsolf, ('roo/ & %/Ill 77()48 ((JA%9 

IJI)I3 MI1JXlJ M()^ I)MA, NY SI1JI,, 'I I1JXE US); N ol'(Jatlil2e 

(Iro(l/ 14.S()4 (M()); (Jel'l'0 (Jaltll)til, 1'.5 kzl SW OF 1'1lllza 

Cily 0tL Ile-tXt. Hwy., Morz &^ lXol/esz 7698 (M()); (t1. I() 

lseyol(l Ulliv. Ieclll-l. I tliS VAlg(1S 'I'OIr(S-I1JXt. I1JXI). Mutile, 

l{fo Mlilee ('roo/ .SSeSL?(3 (IIUA M(), (v)(A, US); 11)ag1a- 

Sa,, I,oe >,o ,(0., Meleli.sot. ./. (ll. .S()()e (k', SkJIJ); Ha( if ,,(0tt 

(,lJaytls ( (1. 2?() kzl S Ol EJSI}lE lAI(LlS7 or/(r (/ ell. 4282 (M()); .ry & I,(orl(.s 7¢X1()8 ((,UAY, M()); lJilsn (,)lJillil(l-lJilSa, 

6.1 li. alxove lar-AIz. Hwy., 3.2 TL1i. I)eyor(J [)ark er- (,. 1 kzl W of filo. I)ol-zlin;,ro-lXJslzelul(0as Hwy., (0e)allil(r 

tIar(e & (,ua(Ja Ike 14346 (US); 

Cerro Jefe Region, vic. Finca Vega, 2.3 mi. above Lago 

Cerro Azul, 4.1 mi. above old Pan-Am. Hwy., Croat 75154 

tricia Pilar-24 de Mayo, Dodson et al. 8415 (MO); Rio 

Blanco, Santo Domingo-Esmeraldas havy., 3 km S of km 

24, Croat 50684 (MO); Santo Domingo de Los Colorados, 

(MO), Croat 11570 (F, MO, SCZ); Rio Maje, above first 

waterfall, Croat 34442 (F, MO, PMA); Rio Torti, S of Pan- 

Rancho Brahman, ca. 10 km NNZvT of Santo Domingo, Sparre 

14122 (S); Hacienda Zaracay, Sparre 15182 (S), Acosta 

am. Hwy. near village of Upper Torti, Folsom & Mauseth Solis 10896 (F, S); Tsachila Chiguilpe, Cer6n et al. 29159 

7844 (MO). San Blas: SW of Puerto Obaldia, Croat (QAP); Santo Domingo-Quevedo, km 11.5, Est. Gustavo 

16803 (MO); Ailigandi, Hammel & D'Arcy 5018 (MO), A. Orces, Quishpe & Davila 82 (QAP, QCNE); km 41, Zak 

Jones & Tejada 275 (PMA); Kuna Yala Nusigandi FS, NW et al. 5726 (QCA); vic. of Montalvo, 40 km E of Babahoyo, 

725



Holm-Nielsen et al. 2769 (AAU, NY, Q(:NE, S)7 Hacienda wide), 1-2 times longer than petioles, coriaceous to 

Monica, 12 km E of San Carlos, Sparre 19397 (S); Haci- subcoriaceous, semiglossy, bicolorous, weakly inenda 

Clementina, Harling 313 (S); Babahoyo-Montalve, 

Sparre 14556 (S), Cornejo & Bonifaz 4828 (GUAY, MO); equilateral, one side 0.6-3.5 cm wider than the oth- 

Centro Cientifico Rio Palenque, Croat 38669 (MO, er, usually short-acuminate at apex, sometimes 

QCNE), Dodso7l & Tan 5338 (SEL), Croat 73826 (MO, acute to rounded with a short acumen, acute to 

QCNE), Croat 50655 (MO), Fallen & Ray 860 (SEL); rounded at base, with the edges turned up near the 

Quevedo, Asplund 15574 (S)7 Dodson 6188 (RPSC, SEL); 

Vinces, Mocachi-Palenque, Jauneche forest, km 70 Quevbase; 

upper surface dark green, drying gray-green 

edo-Palenque, via Mocachi, Dodson et al 10594 (GUAY, to dark olive-green, rarely yellowish brown; lower 

MO, SFL). Manabi: Portoviejo-Pichincha rd., ca. 20 km surface slightly paler, drying yellowish green, rarely 

E of San Placido, Harling & Andersson 24778 (GB, MO); yellowish brown7 midrib flat, 1-2 cm wide at base, 

Cerro Montecristo, S of Manta, Sparre 19488 (S); Mach- concolorous or slightly paler than surface above, 

alilla NP, zona de San Sebastian, Gentry et al. 72496 (MO, 

QCNE); Chone-Santo Domingo Rd., ca. 20 km NNE of drying slightly paler than surface and weakly raised 

Flavio Alfaro, Montanas de Convento, Harling & Anders- above, concave to prominently raised on lower surson 

18898 (GB); Hacienda Don Juan, 10 km NE of Jama, 

N of Rio Don Juan, Neill et al 11683 (QCNE); 223 km S 

of Pedernales, 3.5 km SW of Camarones, I)elinlss 452 

(AAU, NY, MO). Napo: Auca, Plowman 14121 (F). Pastaza: 

Coca-Rio Tiguino, t35.23 km S of Coca, Croat 72573 

(MO). Pichincha: vic. Hotel Tinalandia, 9.6 km E of Santo 

Domingo de los Colorados, Croat 55666 (GUAY, MO, 

QCA); Machachi-Santo Domingo, 19.3 km E of Alluriquin, 

Thompson & Rawlins 1104 (MO); Centinela, Montanas 

de Ila, 12 km E of Patricia Pilar, L0jtnant & Molau 

15839 (AAU); Santo Domingo-Quevedo, Patricia Pilar, 

face, drying brownish; primary lateral veins 15 to 

26 pairs, sunken above, convex below, arisin; at an 

acute angle to the midrib then spreading at an angle 

of 45°-60°(70°), sometimes to 90° near the base 

of the blade, sometimes drying moderately wrinkled; 

interprimary veins lacking or 1 between each 

pair of primary lateral veins, sometimes almost as 

prominent as the primary lateral veinsS minor veins 

indistinct Juvenile telades with acute base and sol- 

Dodson et al. 14638 (M0, QCNE), Gentry et al. 26709 id green midrib. INFI,ORESCENCES 1 to 3 per 

(MO); vic. of Santo Domingo de Los Colorados, Peripa, axil; peduncles (5.5)7-25 x 1.5-2 cm; spathe 

SW of Santo Domingo, Croat & Nunez 82064 (MO, 

QCNE); Nanegalito-Pto. Quito Rd. km 113, ENDESA, 5 

medium to dark greerl broadly curved, long-acukm 

W of San Vincente Andoas, Croat 82829 (MO), Croat minate, 2748 cm long, 2.5-8.2 times longer than 

et al. 83795 (MO); Rfo Silanche, Quito-Pto. Quito, km peduncle, to 4 cm wi({e at anthesis, tube flattenin; 

113, Rodriguez 262 (NY, QCA); Santo Domingo-Quininde 5-12.5 cm wide, oorlstlicted area 2.5-3 cm diam., 

rd., km 41, Zak et al. 5414 (QCA); vic. Maquipuquna 

flattening 3.2-5 ( m wide, spathe blade 3-6 cm 

Res., rd. to Maquipucuna Lodge, LSeimbeck R. 306 (AAU). 

Sucumbios: Lago Agrio-Baeza, ca. km 107, Croat 50480 wide at anthesis, flattening to 6-12 cm wide mid- 

(MO). VENEZUELA. Maiquetia, Andre 457 (K). way, the distal inner surface sometimes white when 

Cultivated specimens. Ecuador. Napo: Auca Oil Field, open; spadix (21)35-38 cm long; free portion 12- 

23 Sep. 1991, Ingram 1124 (MO). 

19 cm long; pistillate portion of spadix (except 

sometimes the uppermost part) fused to spathe 13- 

16. Dieffenbachia longispatha Engl. & K. 

15 cm long; fertile s;taminate portion (8)11-14 cm 

Krause, Pflanzenr. IV, 23 Dc(Heft 64): 44. 

x (9)12-14 mm ((3rying 6-9 mm diam.); mostly 

1915. TYPE: Panama. Colon: Fato (Nombre de 

sterile intermediate portion (2)3-4.3 cm long with 

Dios), July 1911, H. F: Pittier 3838 (holotype, 

a few scattered staminodia in the upper half (some- 

US!; isotypes, B!, F!, M0!). Figures 16, 29B. 

times to throughout ite; length); pistillate flowers 10 

Terrestrial, (1)1.5-3.5 m tall; sap very foul and to 26, round or barely bilobed, widely spaced 5- 

pungent; stem prostrate at base, then erect; inter- 10(20) mm apart, forming in a single irregular row 

nodes 4-12 cm diam., with leaf scars prominent, or scattered but usually no more than 2 flowers 

dark green, semiglossy, drying dark brown to or- across the width of the spadix (rarely 3); ovaries 

ange-brown; petioles thick and succulent, semiglos- pale green, 4-7 mm diam.; stigmas 4-6 mm diam., 

sy, usually solid dark green, rarely streaked with yellow to orange, somewhat broadly bowl-shaped, 

pale green, 23-55 cm long (averaging 36 cm long), 5-7 mm thick on the edge, medially with 1 to 2 

sheathed to about middle (0.58-0.85 their length, somewhat elongate lobes, the lobes 1-1.5 mm 

averaging 0.72); sheath 25X1 cm long (averaging diam., somewhat longer than broad; staminodia 5 

25 cm long), inequilaterally rounded at apex, some- to 6 per pistil, white, irregular, 2-6 x 2-3 mm, 

times weakly free-ending; unsheathed portion 4.0- much flattened at base, less so toward the apex, 

30.5 cm long (averaging about 11 cm), C-shaped often somewhat puckered at the apex; synandria in 

and obtusely sulcate or + terete with a faint flat spirals of 4 to 7 each, 3-4 mm wide, subrounded, 

rib adaxially; blades oblong-elliptic, 41-72 x 17- drying light yellow-brown and concave at apex. IN- 

38 cm (averaging 53 x 24 cm), 1.7-2.9 times lon- F1lUCTESCENCES 17-24 cm long; berries 1.5-2 

ger than wide (averaging 2.3 times longer than cm diam., often deeply emarginate at both ends and


2004 

Croat 


Figure 16. Diedfenbachia longispatha. A. Habit with open inflorescence. B. Plant habit with cluster of inflorescences. 

C. Crown of plant with close-up of leaf base with open inflorescence. D. Inflorescence at anthesis with 

staminate spadix exposed. A, C. (Croat & Zhu 76257); B, D. (Croat & Zhu 76203). 

727

728 

Annals of the 


appearing to be a double fruit, bright yellow to or- comm.). Dieffenbachia longispatha and D. nitidiange, 

mesocarp ca. 2 mm thick, soft, sweet and petiolata do not occur together in Costa Rica since 

tasty at maturity; seeds oblong, 7-8 mm diam., D. Iongispatha barely crosses west of the Panama 

brown to black, smooth. 

Canal. In eastern Panama D. Iongispatha is found 

principally in areas of Tropical moist forest (T-mf) 

Distribution and habitat. Dieffenbachia longis- near E1 Real. 

patha ranges from central Panama to northern Co- Smaller plants of D. Iongispatha may also be 

lombia, mostly from sea level to 180 m, but perhaps confused with D. crebripistillata. In addition to beto 

250 m (owing to a collection at Rio Tuquesa with ing a plant of smaller stature (rarely to 1.3 m tall), 

no elevation reported), occurring in Tropical moist it has more fully sheathed petioles (often to the 

(T-mf), Prernontane wet (P-wf), and Tropical wet for- base of the blade and sometimes overlapping the 

est (T-wf) life zones (Holdridge, 1967). In Panama blade) that usually have a whitish band abaxially 

it occurs on both coasts, but it is relatively rare on and dry conspicuously yellow or orange-brown (vs. 

the Pacific slope. 

green for D. Iongispath(l). In addition, D. crebripis- 

Phenology. While collections of D. Iongispatha tillata differs in having smaller spathes (to 28 cm) 

have been ma(le in both flower and fruit year-round, with the pistils closely aggregated on the spadix, 

flowering OcCUIs y)rincipally during the first half of with (57)80 to 100 pistils versus 10 to 26 for D. 

the rainy season between June and August, while Iongispatha. Finally, though there may be a certain 

fruits mature throughout much of the dry season overlap in their ranges in areas along the Atlantic 

and the first liall' of the rainy season, especially coast in Colon Provine e! D. crebripistillata generally 

from Februaly to September. 

occurs at higher elevatiollst mostly 250 to 975 m. 

Discussion. I)iegenbachia longispatha grows 

most frequently .llong streams, in deposits of sedi- Additional specimens e.w-(l lr1itled. PANAMA. Bocas 

ment along lak s in standing water, but also in del Toro: 1.5 km N ol Ll Cluta, Peterson 6405 (US). 

Canal Area: Barro Co l ox tIdo I s l and, Armour Trail 2000, 

deep soil in tl( lorest understory. Frequerltly it is 

Croat 11321 (F, MO, US): Btlll)our Trail 225, Croat 6471 

found solitary ol irl small clusters. While the spe- (MO); Barbour Trail 2()()* C/-eelt 7/36 (MO); Drayton Trail, 

cies may colollize it is typically much less ( olonial near end, Croat 5767 (M()); Dlayton Trail 1610, Croat 

than other sl)e( ies in Panama and Coloml)ia, such 5761 (MO); Lake Trail, Cro(lt 6276 (MO, SCZ); Oppenas 

D. isthmia, 1). oerstedii7 or D. killipii. 

heirner 67-1-3-1244 (M()); Miller Trail 300, Croat 4095 

(MO); Miller trail 1400, 

The species is; characterized by its usually tall 

(Xlro(/t 17394A (MO); Miller Trail 

1375, Croat 7455 (MO); I'(llson Trail 400, Croat 4134 

((1)1.5-3.5 nl lligh) and robust stem (usually 6-12 (MO); Snyder-Molino Tltlil, (XOat 6192 (MO); Van Tyne 

cm diam.), an(l ly the petiole being inconspicu- Trail, Stimson 5277 (DUKI,* STRI); Wheeler Trail 300, 

ously sheathe(l and decurrent at the apex with a Croat 4125 (MO); Zetek 'I'ltlil 500, Fuertes Cove, Croat 

long subterete Iree portion beneath the bla(le, this 5259 (MO), Croat 6409 (M()); ().6 mi. N of Gamboa near 

Rio Frijol, Tyson 1438 (M()); vi(. Curundu Housing area 

drying charae teristically olive-green or dark larown. of Albrook Air Force bas^, I'alque Metropolitano, Croat 

Perhaps most ( haracteristic of D. Iongispatha are & Zhu 76203 (AAU, (E l{, M ( ), US); Gatun Locks-Fort 

the widely scattered, moderately sparse and very Sherman, 1 mi. E of Foll fils^lman, Croat & Zhu 76266 

(MO, US); Rio Grande-Pe (llo Vidal, rd. to Arraijan, large pistils all(l the usually long spathe. Dieffen- 

Pittier 

bachia longi.s/)atha is typically one of the tallest 

2715 (US); near Fort l


2004 

0.9-1.7 cm wider than the other, weakly to narrow- 

ly acuminate to acute and apiculate at apex, round- 

ed and usually inequilateral, sometimes acute and 

decurrent on the petiole at base, moderately un- 

dulate along the margins, moderately coriaceous to 

subcoriaceousX very dark green and semiglossy to 

weakly glossy above, sometimes mottled in a band 

along the midrib (especially heavy on one side be- 

low the middle) with yellow-green to cream above, 

drying dark gray-brown to dark brown and matte to 

weakly glossy above, slightly to moderately paler 

and weakly glossy to matte below, drying yellow- 

Croat 


Llano-Cartl Rd., near border with San Blas Province, 

Croat 67399A (MO); Rfo TapiaX Standley 26156 (US), 

Standley 28238 (US). San Blas: E of Puerto Obaldia, 

Croat 16923 (MO). COLOMBIA. Antioquia: San Luis, 

Guillernio & Cardenas L. 863 (HUA, JAUM). Choco: 

Acandi, Rio Cuti, Corr. Unguia, Roniero-Castaneda 6436 

(COL); Nuqui, Corr. Arusi, vic. Arusi, Rio Arusi, Croat & 

M. Mora 83777 (= Mora 394) (MO). Meta: PN Natural 

Tinigua, Serrania Chamusa, Centro de Investigaciones Primatologicas 

La Macarena, Camp Colombia Stevenson 379 

(COL). 

green, weakly glossy; rnidrib slightly flat-raised, 

concolorous and 4-5 mm wide, weakly ribbed and 

rmedially sulcate above, narrowly rounded and 

slightly paler below, drying flat-raised and slightly 

darker yellow-brown above; primary lateral veins 14 

to 16 per side, arising at an acute angle, then 

spreading at 25°-70° angle (25°-45° angle toward 

the apex, 65°-70° toward the base), weakly quiltedsunken 

to narrowly sunken and concolorous, sometimes 

in weak valleys above, darker and convex to 

17. Dieffenbachia lutheri Croat, sp. nov. TYPE: round-raised, minutely granular-puberulent to sca- 

Panama. Chiriqui: along border of Bocas del bridulous below, drying weakly and narrowly 

Toro Province, Cerro Colorado, above mine, raised, concolorous above, irregularly angular and 

1600 m, originally collected by Luther et al., yellow-brown below; minor veins on upper surface 

cultivated at Marie Selby Botanical Gardens in part weakly etched, concolorous, those on the 

(acc. #86-0873), 4 Oct. 1991, S. 1E Ingram lower surface moderately obscure to easily visible, 

1146 (holotypeX MO-4224350!; isotypes, B!, darker than surface; lower surface densely pale- 

COL!, K!, MEXU!, PMA!, SEL-065912!, US!). spotted, the spots regularly rounded and evenly 

Figures 17, 27B. 

spaced. INFLORESCENCE 1 per axil; peduncle 4- 

Planta 0.6-1.5 m alta; internodia brevia, 2.0-2.4 cm 

diam.; petiolus 23 cm longus, vaginatus ().7 lorlgitudinis; 

lamina anguste ovata, (24)27-34 ( m longa, 14-1 5 em latcl, 

6 cm long, 1 cm diam., drying dark yellow-brown, 

3.5-8 mm wide; spathe pale green on botll surfaces, 

white near tip, acuminate at apex, 16.5-20.S cm 

nervis primclliis lateralibus 14-16 utroque; inflores(enti 

1 per axillatn; pedunculus 4-6 (m longus, 1 (rn (ticlm.; 

spclthcl pcllli(le viridis, 16.t5-2().tS ( ltl lorlgcl. 

long, flattenecl to 3.5 ( m wide on tube, drying yellowish 

l)rown; sl)(l(li-x 14-20 ( m long (ine lu(ling 1- 

1.tS (m long sti^)e); istillate ortion of sI)a(}ix 7.()- 

HerT), ().6-1.5 m tall; say) not srrlelling of oxali( 7.t3 ( m x 8-9 rnrrl, ( om)letely ( ontiguous with staa(id; 

i71tUrl7.0(les short, to 2 (nl long, 2.()-2.4 (m minate )ortion or nearly so; starninate lsor tion of 

liam., (lark green an(l semiglossy, (Irying Illatte an(l syxa(lix 5. 3-8 ( rn x 5-9 rnm on (Irying, with the 

t)laekene(l; /)etioles ea. 2tS ( m long, (lark greell an(l renlnants of lhi(ke-le(l the(ae visilvle along u^)y)er 

weakly glossy, smooth or with fine, of)S( ure lines on margills; îslils 55 to 6(, 2 to 4 or tllem in a sy)iral 

fresh matelial, sheathe(l ().7 their lengthe (Irying ae loss the wi(lth of the slea(lix, soliletililes mo(lel- 

(lark olive-gray towal d l)ase, somewllat yel lowish ately s)arsely sI)ae e(Se (Iryilly sonletimes ^)romibrown 

toward apex; sheath ae ute on one si(le at nently ('OllVeX an(l the I)istils thus visilvle fronl lsoth 

apex, the other side narrowly loun(Se(l at al)ex, with to) an(l si(le views; pistils 1.4-2.3 mm wide; stigt)oth 

si(les c onfluent, the sheath mal gins (Irying mas pale yellow, 1.f3-l.7 mm diam., drying with 

thin, pale brown, the innel surface of the sheath short, mostly ae ute, mostly ereet strigose tric homes; 

pale yellow-green and very glossy, drying light yel- staminodia 2-3 mlll long, 1.0-1.6 mm wide, marklow-brown, 

much paler than the outer surface; free edly thiekened at the apex, sometimes broader than 

portion of the petiole oval to sharply C-shaped in long, white, drying yellow-brown; synan(lria creamy 

cross section, 7 cm long, sharply sulcate to flat- white, drying irregularly rounded to angular, 0.5tened 

with sharply ridged margins; blades narrowly 3.0 mm wide, yellow-brown, the apex sunken on 

ovate, (24)27-34 x 14-15 cm, 1.9-2.2(2.7) times drying, mediuln yellow-browll with pale, rounded, 

longer than wide, slightly inequilateral, one side cellular inclusions. Berries not seen. 

729 

Distribution and habitat. DieJ2fenbachia lutheri 

is apparently endemic to the region of Cerro Colorado 

in Chiriqui and Bocas del Toro Provinces at 

1390 to 1600 m in the Lower rnontane rain forest 

(LM-rf) life zone (Holdridge, 1967). 

Phenology. A flowering collection was made in 

Panama, and the species has flowered in cultivation 

in October at the Marie Selby Botanical Gardens. 

Discussion. The species is most similar to D. 

Iongispatha in having a considerable portion of the 

petiole unsheatlled and obtusely sulcate; the sheatl



Figure 17. Dieffenbachia 

lutheri. A. Habit of potted plant. B. Plant habit with inflorescence. C. Close-up 

of stem showing petiole sheath and sulcate petioles. D. Close-up of inflorescence.


2004 

Croat 


acute at the apex; and a blade of generally similar matte, moderately smooth; petiole scars 4-5 mm 

shape and dried color. It differs from D. Iongispatha thick. LEAVES erect-spreading, clustered at apex 

in occurring at a much higher elevation (1390- of stem; petioles 1344 cm long (averaging 28 4 cm 

1600 m vs. 180 m), in having the staminate and long), 1-1.5 cm diam. midway, sheathed 0.3-0.76 

pistillate portions of the spadix contiguous (sepa- their length (averaging 0.48), very glossy as if varrated 

by a conspicuous, mostly sterile portion in D. nished, rarely only semiglossy, dark green, drying 

Iongispatha), and in having a spathe that is much orange-brown to blackened; sheath 14-34 cm long 

shorter (to 20.5 cm vs. 27-48 cm). 

(averaging 14.3 cm long), erect to involute, acute 

Diefenbachia lutheri is also similar to D. killipii and decurrent (but sometimes weakly elevated) at 

and even shares with that species the contiguous apex; the unsheathed portion 5-22 cm long (averpistillate 

and staminate portions of the spadix. It aging 13.4 cm long), usually terete to C-shaped, 

differs from D. killipii, however, in having the major sometimes obtusely and narrowly to broadly sulcate 

veins on the lower surface granular-puberulent or V-sulcate, rarely D-shaped; blades oblong-elliprather 

than glabrous. 

tic to oblong-lanceolate, or ovate-elliptic, gradually 

Dief7nenbachia lutheri might be confused with an- acuminate to acute, equilateral to inequilateral at 

other high-elevation species, D. crebripistillata, but apex, usually only weakly inequilateral with one 

that species occurs only up to 950 m, lower than side 0.3-1.5 cm wider (but can be up to 3.5 cm 

D. Iutheri, and differs in having a fully sheathed wider) than the other, acute to obtuse or rounded 

petiole that dries yellow-brown. Diefenbachia luth- at base, usually equilateral, sometimes inequilatereri 

has a petiole that dries dark gray and is un- al with one side acute and the other rounded, 19sheathed 

for more than 5 cm at apex. 

59 x 7-27.5 cm (averaging 38 x 12.6 cm), 2.1- 

Etymology. The species is named in honor of 4.8 times longer than wide, broadest usually at 

botanist Harry Luther from the Marie Selby Botan- middle, 0.7-2.4 times longer than petiole (averagical 

Gardens, Sarasota, Florida, who collected living 1.38 times longer than petiole), subcoriaceous 

ing material of the type species. 

to thinly coriaceous, moderately to weakly bicolo- 

Paratypes. PANAMA. Bocas del Toro: Cerro Colorado, 

9.2 mi. W of Chame, Croat 69070 (MO). Chiriqui: 

Cerro Colora(So, 18.6 mi. N of Rio San Felix, 6.6 mi. beyond 

Chame, Croat 7ti007 MO); 34.1 km of Rio San Felix, 

Croat 37268 (M0). 

rous; margin sometimes weakly undulate; upper 

surface dark green and glossy to semiglossy, plain 

green or rarely weakly variegated with pale green 

in some areas or throughout much of the blade, 

sometimes plain green with just the midrib whitened, 

drying gray-green; lower surface moderately 

18. Dieffenbachia nitidipetiolata Croat & Gra- paler, matte to weakly glossy, drying yellow-green; 

yum, sp. nov. TYPE: Panama. Bocas del Toro: midrib + flattened (slightly sunken toward base), 

Valle del Silencio, along Rfo Changuinola, ca. concolorous or slightly paler, sometimes white 

1 km above mouth of Rio Teribe, vic. Teribe above, convex or bluntly acute below, drying pale 

Indian population, disturbed forest among co- yellow-brown often with a medial ridge on drying; 

coa plantations, 9°21'40"N, 82°31'40"W, 100 primary lateral veins (10)12 to 20(22) per side, dem, 

25 June 1994, 7: B. Croat &: G. Zhu 76449 parting midrib at 20°40°(55°) angle on narrower 

(holotype, MO-04614032-33!; isotypes, AAU!, leaves (especially in Panama), often to 70°-85° an- 

B!, CAS!, COL!, CR!, F!, GH!, INB!, K!, gle on broader blades (mostly in Costa Rica), weak- 

MEXU!, NY!, PMA!, US!). Figures 18, 30A. ly arcuate-ascending, weakly sunken above, weakly 

Planta 50-75(120) cm alta; internodia 0.7-1.5 cm lon- convex and slightly paler below; minor veins visiga, 

1.5-2.5(4.0) cm diam.; petiolus 13-44 cm longus, va- ble, not prominent below. INFLORESCENCES 

ginatus 0.3-0. 76 longitudinis; vagina 14-34 cm longa, mostly 1 to 5 per axil; bracts 2-ribbed throughout; 

acuta ad apicem; pars libera 5-22 cm longa; lamina obpeduncles 

11-18(24) cm long, 0.9-1.3 cm diam., 

longo-elliptica vel oblongo-lanceolata vel ovato-elliptica, 

19-59 cm longa, 7-27.5 cm lata, nervis primariis later- somewhat flattened, medium green, semiglossy; 

alibus (10)12-20(22) utroque; inflorescentia 1-5 in spathe 15-36 cm long, 1.3-2 times longer than pequoque 

axilla; pedunculus 11-18(24) cm longus; spatha 

15-36 cm longa; spadix 13-29 cm longus; pistilla 48- 

60(79). 

duncle, uniformly pale green to greenish white or 

greenish cream, becoming pale yellow post-anthesis, 

semiglossy to matte outside, slightly paler and 

Herb, 50-75(120) cm tall; sap moderately foul- glossy within, spathe tube 2-2.9 cm diam., flattensmelling; 

stems erect or partially reclining; inter- ing to 8-9 cm wide, weakly constricted above the 

nodes 0.7-1.5 cm long, 1.5-2.5(4.0) cm diam., tube to 1.7 cm diam., the constricted area to 5 cm 

glossy to semiglossy, dark green to medium green, wide when flattened, narrowly acuminate at apex; 

smooth, drying dark brown to light yellow-brown, spathe blade 2.5-3.5 cm wide at anthesis, flatten- 

731

732 

Annals of the 


Figure 18. Dieffenbachia nitidipetiolata. A. Habit. B. Close-up of leaves, adaxial surface. C. Close-up of 

crown of plant showing newly opened leaf and inflorescence. D. Opened inflorescence exposing spadix. AX B. (Grayun 

3218); CX D. (Croat 76942).


2004 

Croat 


ing to (3.7)4.5-5.7 cm wide midway; spadix 13-29 cies differs in having a minutely granular upper 

cm long, ca. 1-9 cm shorter than the spathe; the surface, a matte-drying lower midrib with promifree 

portion 11-16 cm long; fertile staminate por- nent raphide cells (vs. drying glossy without raphtion 

5.7-10.5 cm x 6-11 mm, slightly broader at ide cells), and matte-drying petioles (vs. drying 

middle, gradually tapering toward both ends, blunt- glossy). 

ly rounded at apex; male flowers 4 to 6 per spiral, In the Rio Guanche area of Colon Province, Pansubrounded 

to hexagonal, sometimes depressed ama, the species may be hybridizing with D. cremedially, 

the margins irregularly and smoothly in- bripistillata since both species occur along the Rio 

dented; pistillate portion 5.5-14 cm long, 11-15 Guanche and in the Santa Rita area further inland 

mm diam., extending down to the very base of the and at higher elevations. Some plants have the free 

spathe; pistils 48 to 60(79) globose, green, to 2.5- portion of the petiole over half its length on aver- 

3 mm diam., moderately closely spaced except age, which is characteristic of D. nitidipetiolata 

sometimes in uppermost 1 cm (the distance be- (but not of D. crebripistillata), yet the free portion 

tween them generally equaling or up to twice their is not notably glossy as is usually the case for D. 

width), usually to 3 per spiral or 4 to 6 in an arch nitidipetiolata. Examples of these collections are 

across the width of the spadix; stigma yellow, de- McPherson & Merello 8235 from Santa Rita Ridge 

pressed-globose, simple; staminodia 3 to 5 per flow- and Thompson 4874 from Rio Guanche. The Mcer, 

clavate, white, 2-6 mm long, up to 2 times lon- Pherson and Merello collection also describes the 

ger than pistil, somewhat flattened and mostly petiole as having a white streak on the abaxial surcontiguous 

at base (sometimes united in pairs), face, a feature otherwise known only in D. crebrisometimes 

slightly thickened and somewhat puck- pistillata. 

ered at apex, drying orange; staminate portion interrupted 

from piKstillate portion by a + naked por- P(lrfityl)es. C()S'rA RICA. Alajuela: -34 km beyond 

tion, 2.tS-4.S c m x 5-9 mm with a few Ksterile San l4aril6n 1)eyond l jos Angeles, I,llteyn W@8.T (l)UKIS); 

llo(lt. 4.X36)8 (M()); (,lJcililo-(,ual)i 

lees 14(l., A(lrritlMevr 2416) (t''). Hereflia: 1, Xeelneal 

sea level to 12()() m in wettel palts of Trol)ic 

va at Xalul)i(lili, llT7l()ll.()Z 1()3 (M()); 1, X,011 Plole(lola 

moi.st foresl ( 1 -mf), Premont(lrle wet /orest (P-wf), Rio (,rU('ililO, (,r(zl-lllzl & ks(&h(ltz .X218 (I)U Ktg); La Selva, 

Tro/)ie(ll wet forest (T-wf), an(1 Premontarle rairl /or- ()'1'S tX'ieel(l Aluliorl orl ttlee Rfo Puello Viejo, E of its jcnl. 

est (P-rf) 1ife zones (Holdli(lge, 1967). 

wiltl ttle>. ttfo SaIal)iqui, Ctro(lt 442.S4 (M()), H(lltlmel 8123 

Phenology. Dief/erl b(lchi(l niti(lil)etiol(lt(l flow- 

(I)UKt, tX', M()), Burger & kSXtolze .57.53 (tX', (,,t), Fol.som 

92()() (I)U Ktg), H(ltnmel 1()()82 (I)UKIS), Beflch 14tSb 

ers principally at the beginning of the lainy season (l)UKIS), Beflch 1439 (DUKIi, MO), M(lcL)ougal 1()27 

(June-September), but also as early as March and (DUKIL), Jitzzezzez 1()3 (MO); Rio Salapi(lui, 2 km S of San 

as late as October, with the heaviest flowering ap- Miguel, Lent 37 (tX'); Sarapiqui, Croat 44307 (MO), Grapalently 

in August. Fruitillg collectiolls 11ave been yum. 2225 (DUKE, MO); San Jose and Pto. Viejo, vie. of 

Chilamante, 11.6 mi. N of Cariblanco, Croat 68379 (MO). 

made from May through January. 

Limon: Hitoy Cerere reserve, SW of Valle lga Estrella, 

Discussion. The species is characterized by its along Rfo Cerere to ca. 1 km upstream from Quebrada 

long, glossy petioles with the unsheathed portion Barrera, Grayum & Hammel 5785 (MO); Res. Biol. Hitoy 

being moderately elongate, 0.7-0.24 cm of their Cerere, Valle de la Estrella, trail to Cerro Bob6cara, G. 

length, and which dry glossy as though varnished, Herrera 4116 (CR, MO); N of Quebrada E1 Molinete, Rfo 

Chirrip6-Rfo Corinto, Grayum & Jacobs 3524 (MO); Ref. 

and by the moderately closely spaced pistils. It is Nac. Barra del Colorado, Rfo Chirrip6cito-Rfo Sardina, 

most easily confused with D. Iongispatha, differing Grayum 9844 (CR, MO); Rfo Catarata, Burger et al. 1()323 

from that species in having the pistils much smaller (F, MO, SEL); BriBri-Rfo Sixaola, Rfo Catarata, Burger & 

and aggregated and from D. crebripistillata in hav- Antonio 1()9()5 (F, MO, PMA); BriBri-Caribbean coast, 

Croat 43217 (MO). Puntarenas: E of Monteverde on the 

ing the petiole almost unsheathed (vs. sheathed 

Pacific watershed, Burger et al. 1()699 (MO); Palmar Normostly 

to the apex in D. crebripistillata). The spe- te-Panamerican border, 3 km N of turn-off to Rinc6n, 

cies may be confused with D. hammelii. That spe- Croat & O. Hannon 79199 (INB, MO, US). San Jose: 

733



Vazquez de Coronado, Braulio Carrillo NP, along Hwy. San km SW of Pato, Croat 56070 (CHOCO, MO); Rio Condoto- 

Jose to Siquirres, trail to Rio Sucio, site of the Old Carillo Rio San Juan, Killip 35113 (COL, US); Alto de Buey, Gen- 

Station, Croat 78788 (IBE, INB, MO, WU). NICARA- try & Forero 7342 (COL, MO); Bahia Solano, Puerto Mu- 

GUA. Zelaya: Cano Montecristo, E of Campamento Ger- tis, Gentry & Forero 7217 (COL, MO); Bahia Solano, Croat 

man Pomares, P. Moreno & J. Sandino 15160 (MO); Rio 57462 (CHOCO, COL, MO); N of Bahia Solano, Cerro 

Punta Gorda, Atlanta, "La Richard," 200 m SE, Moreno Mecana, Juncosa 1795 (MO); Mecana, Quebrada Resa- 

& Sandino 12976 (MO). PANAMA. Bocas del Toro: Rio quita, Juncosa 1898 (MO), 1912 (MO); Rio San Juan, 

San Pedro7 Gordon 55C (MO); Santa Catalina7 4 Dec. Noanama, Forero et al. 4571 (COL); Rio Bicordo7 tributary 

19677 Blackwell et al. 2704 (COL7 MO7 UC); Chiriqui La- of Rio San Juan7 Noanama7 Forero et al. 4659 (COL) 

goon area, Cocoa Cay, von Wedel 2892 (COL, F, MO, UC); Acandi, Quebrada Sardi, E1 Paramo, Roldan et al. 1199 

above RR stop at Milla 7.5, Croat & Porter 16303 (MO) (HUA, MO); Corr. San Francisco, Vereda Coquital, sitio 

Croat & Porter 16412 (MO), Croat 38120 (HUA, INB, E1 Paramo, Quebrada Zardi, Betancur et al. 1177 (HUA); 

MO, UB, VEN), Croat & Porter 16375 (MO); Chiriqui Vereda Coquital, Fonnegra et al. 2899 (HUA, NY), 2907 

Grande-Fortuna, 7.7 mi. S of Chiriqui Grande, 1.5 m W (HUA, NY); Nuqui, Corregimiento Arusi, vic. of Arusi, 

of Punta Pena, Croat & Grayam 6()116 (MO), Akers 78 Est. Biol. E1 Amargal, Croat & Mora 83652 (= Mora 27()) 

(MO); IRHE vic., Carrasquilla & Mendoza 1239 (MO, (MO); Corr. Arusi, Est. Biol.. E1 Amargal, Mora 170 

PMA); Almirante-Ojo de Agua, 3-6 km W of Almirante, (COL). ECUADOR. Esmeraldas: Zapallo Grande, Rio 

Croat 38229 (F, MO, PMA); about 10 mi. W of Veraguas Cayapa, Barfod et al. 48348 (AAU); Lita-San Lorenzo Rd., 

border, McPherson 11401 (CM, MO); Valle del Silencio, 0.9 km E of E1 Durango, 19.8 km W of Alto Tambo, Croat 

along Rio Changuinola, ca. 1 km above mouth of Rio Ter- et al. 82520 (MO); Playa de Oro7 1 km from Rio Santiago7 

ibe, vic. Teribe Indian population, Croat & Zhu 76422 Ceron & J. Corozo 33858 (QAP), 33947 (QAP), Ceron & 

(INB, MO, WU); Cocoa Cay, vic. of Chiriqui Lagoon, von J. Corozo 34097 (QAP); San Lorenzo-Mataje, departing 

Wedel 2892 (F); Rio Teribe, vic Teribe II, IRHE, Carras- main Lita-San Lorenzo Hwy., 11.6 km N of Gasolinera 

quilla 2005 (PMA); Changuinola-Tuibe Rivers, Zigla Rd., San Lorenzo, 2.9 km W of main Lita-San Lorenzo Hwy., 

Lazor et al. 2578 (MO); limit trail of Par. Intl. La Amistad, Croat et al. 84041 (AAU, B, CAS, COL, DUKE, G, K, M, 

from Quebrada Boca Chica to Quebrada Bonyic, Polanco MO, NY, P, UB, US, WU); Cotacachi Cayapas, San Mi- 

1591 (PMA). Chiriqui: Fortuna Lake Area, 3.4 km N of guel, Rio Cayapas, Sector Loma Linda, Tipflz et al. 2280 

Quebrada Chorro, 1.6 mi. N of center of bridge over lake, (MO, QCNE); Rio Cayapa, Zapallo Grande, Barfod et al. 

Croat 74958 (MO); vic. Fortuna Dam, McPherson 9829 48154 (AAU, MO); Zapallo Grande, Kvist et al. 48348 

(MO); Quebrada Los Chorros-Quebrada Hondo, N of For- (AAU, QCA); Rfo Cayapa, Zapallo Grande, Kvist & Asanza 

tuna Lake, Churchill & Churchill 6105 (MO); Fortuna- 40756 (AAU); Rio Grande, at Zapallo Grande, Barfod & 

Chiriqui Grande, 0.7 mi. NW of center of dam, Croat & Skov 60101 (AAU); Rio San Miguel, upstream from Pueb- 

Zhu 76482 (INB, MO, PMA); trail from rd. near Forestry lo Cayapas, Holm-Nielsen et al. 25355 (AAU); San Lor- 

Nursery to Rio Hornito, Thompson 5028 (CM). Cocle: enzo, Sparre 18326 (S). Morona-Santiago: Santiago-Mo- 

Cont. Divide, N of Penonome on rd. to Coclesito, Hammel rona, Rio Morona, 23.4 km E of Santiago, Croat 87448 

4049 (MO). Colon: Santa Rita Ridge rd., ca. 8 mi. E of (MO, QCNE). 

Transisthmian Hwy., McPherson & Merello 8235 (MO); 

Santa Rita Ridge Rd., ca. 1 hr. walk from end of rd., 

Antonio 4488 (MO); Santa Rita Ridge Rd. 6.5 mi. E of 1 9. Dieffenbachia obscurinervia Croat, sp. 

Boyd-Roosevelt Hwy., Croat & Zhu 76942 (CR, GOET nov. TYPE: Panama. San Blas: trail from dock 

GUAT, HUA, INPA, ISC, LE, MO, PMA TEFH, UB, to airport to village of Cangandi, 9°24'N, 

WISC); Puerto Pilon-Portobelo, Rfo Guanche, ca. 1.5 mi. 79°24'W, 3-30 m, 26 Mar. 1986, G. de Nevers, 

S of rd., Croat & Zhu 76254 (MO, PMA); Rio Guanche, 

3-5 km above bridge, Croat 79348 (B, INB, MEXU, MO, H. Herrera & S. Charnley 7409 (holotype, M0- 

US), Sytsma 1695 (MO); 3-7 km above bridge, Hammel 3475792!; isotypes, B!, US!). Figures 19, 29A. 

et al. 4882 (MO); ca. 3 km above bridge, Croat 49761 

(MO); Santa Rita-Serra Llorona, Galdames et al. 1138 Planta 0.8-1.5 m; internodia brevia, (1)3 1(5) cm lon- 

(STRI); Rio Iguanita, Croat 49768 (MO). Darien: W side ga, 1.5-2.5(3) cm diam.; petiolus 8-12 ( m longus, vaginof 

Cerro Pirre, Croat 68854 (MO); Rio Tuquesa, ca. 2 km atus ad medium vel ad apicem; lamina anguste elliptica, 

air distance from Cont. Divide, near upper gold mining 20-30 cm longa, 5-8 cm lata; inflorescentia 1-3 per axcamp 

of Tyler Kittredge, Croat 27161 (MO); Serrania de illam; pedunculus (1.7)3-5 cm longus; spatha 11.5-20 cm 

Pirre, Rio Cana-Rio Escucha Ruido, above Cana Gold longa; spadix 10-18 cm longa, cum parte F)istillata 5-5.5 

Mine, Croat 37746 (MO). Panama: E1 Llano-Carti Rd., cm longa. 

near border with Province of San Blas, Croat 67379A 

(MO). San Blas: Comarca de Kunayala, Nusigandi, E1 Slender herb, 0.8-1.5 m tall, stem decumbent on 

Llano-Carti Rd., 10.1 mi. N of Inter-am. Hwy., then ca. older parts, weakly rooted, leaf scars inconspicu- 

0.5 mi. N Paseo Mariska, Croat & Zhu 77023 (MO); SE ous; internodes glossy, (1)3-4(5) cm long, 1.5of 

Puerto Obaldia, Croat 16769A (MO); trail from dock to 

airport to village of Cangandi, de Nevers et al. 7407 (MO, 

2.5(3) cm diam., dark green, epidermis fissured mi- 

PMA); E1 Llano-Carti Rd. km 16.7, trail W to waterfall 5 nutely into a netlike reticulum becoming uniformly 

km from rd., de Nevers & S. Charnley 5898 (MO, PMA); fissured and brown-scurfy with age; petioles 8-12 

Canagangi, de Nevers et al. 5727 (MO). Veraguas: Boca cm long, averaging 10.6 cm long, sheathing 1/2 to 

de Concepcion, in Golfo de los Mosquitos, McPherson throughout (average 0.8 their length), light green or 

11381 (MO). COLOMBIA. Antioquia: Mutata, Corr. Longani, 

Callejas et al. 5601 (HUA). Choco: trail from Rio yellow-green with the surface dark green-splotched 

Mecana to Alto de Mecana, Gentry & Juncosa 41037 or dark green with white or yellow spots (sometimes 

(MO); Serrania de Baudo, Las Animas-Pato, Rio Pato, 10 with spots forming a repeated series of irregular


2004 

Croat 


0d -- 

Figure 19. Diegenbachia obscurinervia. A7 B. (Croat 12660). A. Cluster of several potted plants with plant on 

left with open inflorescence. B. Close-up of crown of plant showing adaxial and abaxial surfaces and inflorescence 

at anthesis. C. Close-up of stem showing mottled epidermis and speckled petioles (Croat 61234). 

inflorescence showing both pistillate and staminate portions of spadix (Croat 13849). 

D. Close-up of 

735



bands); sheath 6.7-10 cm long, the tip usually Cocle) to northern Colombia (Antioquia and Choco) 

rounded to inequilaterally emarginate, rarely de- from 30 to 800 m in Tropical rnoistforest (T-mf) and 

current, frequently with one side emarginate and Tropical wet forest (T-wf) life zones (Holdridge, 

with the other side acute; unsheathed portion lack- 1967). 

ing or to 5.2 cm long, sharply to bluntly sulcate; Phenology. The species has been seen in flowblades 

narrowly elliptic, 20-30 cm long (averaging er from May to December and fruiting collections 

24 cm long), 5-8 cm wide (averaging 7.5 cm wide), have been reported from from September to May. 

2.4-5.2 times longer than wide, 1.6-3.1 times lon- Discussion. Dieffenbachia obscurinervia is charger 

than petiole, slightly inequilateral, one side acterized by its scurfy brown internodes, conspic- 

0.7-1 cm wi(ler than the other side, subcoriaceous, uously splotched petioles, and narrow blades with 

weakly bie olol ous, gradually acuminate, briefly apic- weakly developed primary lateral veins. It is not 

ulate at al)ex, {lequently somewhat falcate, cuneate easily confused with any other species but was into 

obtuse ol X ounded, often weakly inequilateral, correctly called D. pittieri, a species considered rerarely 

weakly sube ordate at base, upper surface dark stricted to the Isthmus of Panama, for many years. 

green, semit,lossy, drying dark yellow-green to dark See that species for a discussion of the differences. 

olive-greels; lowel surface slightly to moderately paler, 

drying yellow-green, midrib slightly raised to flat 

above, con( ololous, drying concolorous above, convex 

to narlowly rounded below, dark green, densely 

Paratypes. PANAMA. Canal Area: Pipeline Rd.7 mi. 

1-37 N of GarYlboa7 S. Knapp 1042 (MO); Pipeline Rd. at 

Rio Agua Salud7 Croat 12352A (MO); Barro Colorado Island7 

Armour 'I'rail 12807 Croat 8623 (MO); Drayton Trail 

pale yellowish green-spotted, drying paler than sur- 10007 Croat /2660 (MO); Drayton 19107 Croat 6775 (MO7 

STRI); Draytoll 1490, Croat 5759 (M()). Cocle: vic. E1 

face or datket than surface below; primary lateral 

Valle de AntI)ll, La Mesa7 Finca Maarenita7 Croat & Zhu 

veins 10 to 13 I)er side, weakly etched above, weak- 76666A (M ( )* I'MA). Colon: Santa Rita Ridge7 Croat 

ly raised or llot at all raised below; the interprimary 13849 (MO); Nombre de Dios7 Cro(lt t57.X29 (MO); Cocle 

veins alnsost as conspicuous as primary lateral del Norte7 ZI(lrnmel 4484 (MO); I'ollofelo-Nombre de 

Dios7 1.2 lzli. I)eyond jet. of rd. lo Isla Grande7 Croat 

veins, milsor veins moderately distinct below. IN- 

49791 (M()7 I'MA); Rio Guanche7 (,. .S kln upstream from 

FLORES(E , N (E IiS 1 to 3 per axil; peduncle (1.7)3- rd. to Portobeloe Hammel & Train(r /4786 (MO); Loma 

5 cm longe somewhat flattened in cross section; de la GloliaF near Fato (Nombre (le l)ios)7 Pittier 3847 

spathe 11.5-2() c m long, 3.2-9.5 times longer than (US). Panalll.'l: Balboa West7 Za/l(lt(l (t (ll. 289 (PMA); 

peduncle, (lark green or pale green throughout, along trail ol'l' I,larlo-Carti Rd.7 4.6 rrli. from jet. with Pancaudate-ae 

lllninate, sometimes sharply reflexed below 

apex, .V)adix 10-18 cm long; free portion 5-9 

cm long; pistillate portion 5-5.5 cm long; sterile 

Am. Hwy.7 M(l'l1(rson & Merello 8/4.S (MO). San Blas: 

Rio Cangandl (2angandi7 H. Herrer(l 245 (MO7 PMA); Nusigandi7 

alotlg ll Llano-Carti Rd.7 ().7 km beyond Cuna 

headquarterst 11.6 km from Pan-Am. Hwy.7 Croat 75149 

segment less than 1 cm long, 6 mm diam., with (KS MO). C()l,()MBIA. Antioquia: Slo Anorl valley7 near 

scattered staminodia throughout; pistils 20 to 40, 

moderately ( losely spaced, scattered, 2-2.6 mm 

diam.; stigma depressed-globose, 2 mm diam., staminodia 

white, usually 3 per pistil, 1.5-2(3.4) mm 

Planta Provi(lell( ia7 Shepherd 438 (M(), WIS); Anori7 Corr. 

Prov.7 Soejart(l & Renteria 3556 (HUA); %aragoza7 Rio Anori7 

Quebra(ltl l, 'rirana-Rio Anori7 2 km N of Quebrada 

La Tirana7 3 k lll ll priver from Planta Providencia7 Alverson 

et al. 283 (C()l,); Carepa7 Tulenapa Reserva (ICA)7 Turbolong, 

flattene(l, oblong to clavate, grooved medially, Mutata7 40 kzl S of Turbo7 Callejas et al. 4873 (MO7 NY). 

Choco: Nu(ltlt, Quebrada Chaqui7 Galeano et al. 4825 

drying flattened, thickened at apex; synandria ir- 

(MO); Quibddx, l, Concepcion7 15 km E of Quibdo7 Archer 

regularly rounded with margins undulate, minutely 2215 (US); Cor(loba7 Tierralta7 Rio Esmeraldas-Rio Simi7 

warty, widely sunken at apex, 1.8-2.6 mm diam., 2 km above ( orlfluence7 Bernal et al. 1158 (COL). 

dark yellow-brown, matte on drying. INFRUC- 

TESCENCE broadly arching; spathe mostly decid- 20. Dieffenbachia oerstedii Schott, Oesterr. 

uous; spadix to 8 cm long; berries red, broadly el- Bot. Z. 8: 179. 1858. TYPE: Guatemala. Agualipsoid, 

to 1.5 cm long, less than 1 cm diam. cate, A. S. Oersted s.n. (holotype, C!). Figures 

Common name. Abior (Kuna) (de Nevers et al. 

20, 29A. 

7409). 

Herb, 30-75(100) cm tall (usually less than 50 

Distribution and habitat. Dieffenbachia obscu- cm); stem erect or partially reclining, often conrinervia 

ranges from central Panama (as far west as spicuously clustered with numerous plants; sap 

Figure 20. Dieffenbachia oerstedii. A. Potted plant with inflorescences and leaves with mottled blades (Croat 

68449). B. Potted flowering plant with pale midribs (Croat & Zhu 76794). C. Leaves and post-anthesis inflorescence 

showing ovate-lanceolate blades with scarcely sunken primary lateral veins (Croat 63208A).-D. Close-up of


2004 

Croat 


Figure 20. (Continued) stems showing the free-ending petiole sheath (Croat 36297). EX F. (Croat 68449). E. 

Flowering plant with open inflorescences. F. Close-up of flowering plant showing open spathe blade and exposed 

stamlnate portlon of inflorescences. 

737

738 

Annals of the 


moderately foul; leaf scars conspicuous, transverse veins" sometimes visible. INFLORESCENCES 1 to 

or oblique (when oblique, up to 2x as wide on one 2 sometimes 3, rarely 4 per axil; peduncle (2.2)3.5side 

as the other), forming a T or forming a W on 12(22) cm long (average 7.6 cm long), l/lo as long 

open side of sheath; internodes semiglossy to glossy, as spathe to fully as long as the spathe, averaging 

weakly warty on magnification, 1-5.T cm long, 0.8- 0.43 as long, 6-13 mm diam., 2-4 mm diam. on 

2(3) cm diam., dark olive-green to blackish green, drying, somewhat flattened in cross section; spathe 

medium green or dark green, sometimes variegated (7.5)10-17(25.5) cm long, weakly constricted above 

with streaks of paler green, drying dark brown to tube, medium green on both surfaces, sometimes 

dark yellow-brown, glossy, smooth; petioles 10- tinged whitish on back throughout (the median rib 

20(.30) cm lollg, (averaging 13.7 cm long), 3-4 mm green), weakly glossy to semiglossy outside, glossy 

diam., sheathing 1/3 to 3/4 (rarely nearly throughout) on inside, narrowly acuminate at apex; tube to 2- 

their length (avelaging 0.7 their length), medium 2.8 cm wide, 1.5-2.3 cm thick when furled, to 8 

green, matte, sometimes finely streaked throughout cm wide when flattened; spathe blade to 2.5 cm 

with darker green, usually weakly glossy and white diam. when furled, 3.0-8 cm wide at anthesis; spanear 

base; sheath 2.5-21.5 cm long (averaging 9.3 dix (7)10-17(21) cm long (averaging 14.6 cm), 1- 

cm long), pale green to white on lowermost clasping 5 cm shortel than spathe; pistillate portion 4.3portion 

(contl asting sharply with much darker 7.5(10) cm long (averaging 6.4 cm long), 5-10 mm 

stem), with margins involute, the tip with one side diam.; fertile staminate portion 3-8.5 cm long (averect, 

free-ending and rounded to auriculate, with eraging .5..3 ( m long), 5-7 mm diam., broadest at 

the other side rounded to acute (sometimes not free- middle, tapeling toward both ends, especially at 

ending in Coe le Province, Panama); unsheathed 

bluntly y)oi llted apex; mostly steri le intermediate 

portion 5-1.S (m long, C-shaped to U-shaped in 

segment (1)2-3.2 cm long, to 2.5 mm diam. (dried), 

cross section, ( onvex adaxially, acute to bluntly anwith 

a few u i(lely scattered stel ile staminate flowers 

gled on margills; I)leldes ovate to narrowly ovate or 

in uppel 3/4 clnd a few pistillate flowers in lower 1/4, 

ovate-lanceolate or rarely oblanceolate, (5.5)14sometime.s 

with only sterile stanlillate flowers scat- 

22(35) x (1.7)4-14(21.5) cm, (averaging 20.1 x 

tered thlotlt,llout, rarely almost l)ale; pistils (26)33- 

8.5 cm), 1.4-5.9 tilileS longer than wide (averaging 

46(54), lo(lerately closely spat e(l except in the 

2.5 times longer than wide), 0.8-2.7 times longer 

lower 1..S em and the upper ] (m (the distance 

than petiole (avel aging 1.5), often inequilateral, 

between theln generally equaling or twice their 

somewhat thinly ( oriaceous to subcoriaceous, moderately 

bicolorous, gradually acuminate and apicwidth), 

uS to 4 in a row across tlle width of the 

ulate at apex (the acumen 5 mm long), acute and 

spadix, subt,lobose, 1.5 x 1.4-2.6 mm; stigma deweakly 

decurrent to obtuse or more often rounded, 

pressed-globoseX yellow, 1 mm wide, puberulent; 

sometimes subcordate at base; upper surface matte 

style cavitolln with a weak central dome; staminoand 

subvelvety to weakly glossy, dark green, fre- dia clavate, white, drying orange, 2-3(4.5) mm 

quently splot(hed light or medium green or with long, 1-2 nlm wide at apex, 2-3 times longer than 

white areas especially near midrib (all variations pistil, usually weakly fused at base, sometimes well 

frequently found in a single population), drying separate(l? weakly flattened, thickened toward apex; 

dark brown to gray-green, concolorous, sometimes synandria 3 to 5 per spiral, 1.4-2.0 mm diam., the 

faintly dark-stliate; lower surface matte to weakly margins slllooth to irregularly and smoothly inglossy 

(epidermal cells raised and sometimes trans- dented, smooth and subrounded at apex. INFRUClucent), 

drying yellow-green to yellow-brown, some- TESCENCE to 22 cm long; spathe orange outside; 

times faintly dark-striate, slightly paler; rnidrib flat- berries bright red, globose, with 13 to 43 per spadix, 

raised to obtusely flat-raised above, concolorous or 4-6 mm diam. 

sometimes white above (sometimes mottled on 

plants with mottled leaves, sometimes faintly dark- Distribution and habitat. Dieffenbachia oerstestriate 

on both surfaces), slightly paler and convex dii ranges from Mexico (Veracruz, Oaxaca, Chiapas, 

to broadly convex to bluntly acute below; prirnary and Tabasco) mostly along the Caribbean slope in 

lateral veins (4)6 to 9(11) per side, departing midrib Central America in Guatemala, Honduras, Nicaraat 

a 40°-45° angle, broadly arcuate-ascending, gua, and Costa Rica (where it occurs on both 

sunken to quilted-sunken above, slightly darker slopes) and west-central Panama. The species also 

than surface, weakly convex-pleated below; inter- occurs on the Pacific slope of E1 Salvador (Depart- 

1 

primary velns weaK or as consplcuous as prlmary ment of Ahuachapan). The species ranges from sea 

lateral veins; minor veins indistinct above, moder- level to 1260 m in Tropical dry (T-df), Tropical rnoist 

ately distinct to moderately obscure below; "cross- (T-mf), Tropical wet (T-wf), Prernontane wet (P-wf),


2004 

Croat 


and Prernontane rain forest (P-rf) life zones (Hold- averaging 14 cm long (sheathed 0.48-0.9 the petridge, 

1967). 

iole length). In contrast, plants from Heredia Prov- 

Phenology. Flowering of D. oerstedii occurs ince in Costa Rica are smaller than average with 

throughout the year, but with the heaviest flowering blades averaging 13.9 cm long and 5 cm wide (2.9 

at the end of the dry season and the first part of times longer than wide). 

the rainy season, April to September. Fruit matu- The distribution of Dieffenbachia oerstedii in Panration 

is more regularly scattered throughout the ama is apparently disjunct with some collections 

year. 

known from far western Panama, but the species also 

Discussion. This species is characterized by its occurs in the vicinity of E1 Cope and La Mesa in 

small stature (generally less than 1 m tall); fre- Cocle Province. The populations in E1 Cope have 

quently clustered stems; and sharply C-shaped pet- somewhat smaller leaves (9.5-15 cm long), but differ 

ioles that are whitish at the base and sheathed only slightly from those in Costa Rica. The E1 Cope 

mostly 1/3_3/4 their length, with the sheath obscurely material is otherwise morphologically identical to 

free-ending and inequilateral with at least one side populations of DP oerstedii such as those found at La 

usually merely rounded or acute rather than auric- Selva in northern Costa Rica. 

ulate. Also characteristic are the moderately thin, The petiole sheath of the Cocle populations is 

usually small, + ovate-lanceolate blades, which are merely rounded on one side and acute on the other 

generally acute to rounded or truncate basally and (vs. auriculate on one side and rounded on the othmatte 

or only weakly glossy adaxially. 

er side for typical material of D. oerstedii). In ad- 

Dief77enbachia oerstedii is the most widespread dition, the midrib is not faintly striate on both surand 

ecologically variable species in Central Amer- faces as is usually the case with typical D. oerstedii. 

ica. It is extremely malleable in terms of leaf size, The petiole scars are much more oblique than usual 

leaf blade shape, and markings, with different with the internodes longer on one side of the stem 

blade coloration types (cf. Fig. 20A, B) all found than on the other (vs. more nearly perpendicular to 

within the same population. For example, in pop- the stem with the internodes of equal width on both 

ulations at Rio Grande de Taracoles at Carara in sides of the stem for typical D. oerstedii). Despite 

Costa Rica (0-200 m), the plants of this species the strong morphological similarities, plants from 

are robust. The populations of D. oerstedii at 1400 Cocle Province, Panama, differ markedly in their 

m in Braulio Carillo National Park (also Costa Rica) ecological requirements. Cultivated plants, for exhave 

small anel delivate leaves. In Costa Rica, the ample, will survive well only in an area of nearly 

leaf blades tend to be matte or only weakly glossy 100% humidity. Alternatively, the Costa Rican maon 

the upper surface. However, elsewhere in Cen- terial from Heredia does very well in the drier parts 

tral America the blades are often more glossy. A of the greenhouse. 

collection from E1 Salvador (Selby 77-3()78) has In Mexico, Dieffenbachia oerstedii may be conblades 

that are initially rather glossy on the upper fused with D. wendlandii since they have similar 

surface and weakly glossy on the lower surface (al- leaf blades. Living material of the latter is easily 

beit soon turning only weakly glossy above). Plants separated in having petioles with the free portion 

in Panama and Costa Rica have blades 2-11 cm terete or obtusely and weakly sulcate (vs. sharply 

wide and acute to rounded at the base, whereas in C-shaped and sulcate in D. oerstedii), the base solid 

other parts of Central America from Nicaragua to green, and in having the sheath decurrent at the 

Mexico the blades are often 14-18 cm wide with apex with one margin overlapping the other (vs. 

subcordate leaf bases. 

rolled in from both sides in D. oerstedii, with both 

Standley (1937) considered Dieffienbachia leopol- sides visible and often protruded at apex and the 

dii (Bull. Catal. 1878) similar to or synonymous with sides inequilaterally rounded to auriculate) Com- 

D. oerstedii, but the type specimen indicates that it pared with D. oerstedii, D. wendlandii is typically 

was actually collected in Colombia, where D. oerste- a larger plant with longer stems, thicker internodes, 

dii does not range. 'The species is actually closer to and larger leaves up to 33 cm long and 9.5-15 cm 

D. killipii (see discussion under D. killipii). wide, and with the midrib 1.2-1.4 cm wide (vs. 5- 

Collections from Mexico and Belize are larger on 6 mm wide for D. oerstedii). In addition, D. wenaverage 

than those in Panama, Costa Rica, and dlandii has the stigma bowl-shaped at anthesis with 

Nicaragua, with blades 16.5-35 cm long and 6- a protruded central dome. In contrast, the stigma 

16.5 cm wide (averaging 23 x 11 cm). Petioles for of D. oerstedii is cushion-shaped. 

the same area range from 8.5 to 30 cm long, about Diegenbachia oerstedii is similar to D. killipii in 

as long as the blades to 1.7 times longer than the general blade shape and size as well as in often 

blade with the sheath from 7.5 to 21.5 cm long, 

having the 66flat-raised" upper midrib, but D. killipii 

739

740 

Annals of the 


often differs in having petioles not at all whitish at Additional specimens seen. BELIZE. E1 Dorado, 

the base, blades which are typically semiglossy (vs. 

Schipp 386 (F, GH, K, MO, NY); Caves Branch, halfway 

up St. Herman's Hill, Whitefoord 1176 (BM, MO); Cayo, 

matte and subvelvety to weakly glossy in D. oerste- Stann Creek Districts, Hummingbird Hwy., betw. mi. 25dii), 

and especially in having the spadix with the 34, Dwyer & Diecknian 13008 (MO); Vaca Plateau, Vaca 

pistillate and staminate portions nearly contiguous Falls, Balick et al. 2080 (MO); Stann Creek, Cockscomb 

rather than well separated. 

Basin, Jaguar Pres., 10 km W of Maya Center, off S Hwy., 

Balick et al. 2716 (MO); Toledo, vic. of trail to Esperanza 

Herbarium material of Dieffenbachia oerstedii beginning 1 mi. N of Columbia Forest Station, Vanderveen 

may be confuse(l with D. seguine from the West 590 (MO); Columbia For. Stat., Dwyer 9920 (MO); San 

Indies. Both x,oee ies sometimes have blades of sim- Jose trail to Esperanza 1 mi. N of Columbia Forest Station, 

ilar shape, an(l lenth have petioles that are com- Croat 24250 (MO); San Jose, 6.T rlli. N of Columbia Forest 

Station, Dwyer 11184 (F); Bladen, ttichardson Creek, afparably 

sheatlle(l and sharply C-shaped on the free fluent of Bladen Branch, lower ptlt-t of Maya Mountains, 

portion. The lwo sI)ecies differ in stature, with D. Davidse & Brant 31879 (MO); alorlg Bladen Branch from 

Oerstedii beirlg IsUe h smaller and with blades matte, Richardson Creek to Quebrada (le Oro, Davidse & Brant 

often subvelvely, whereas those of D. seguine are at 32366 (MEXU, MO); Bladen, Solornon Camp, vic. of the 

jet. of Richardson Creek and 131a(len Branch, foothills of 

a minimum weakly glossy on the upper surface. The the Maya Mountains, Davidse & lXrant 32410 (MEXU, 

auricles on tle fIee-ending sheath of D. seguine are MO); Bladen, Bladen Nature Rexelve, ca. 2 air km N of 

both rounde(l al tlle apex, whereas in D. oerstedii upper Bladen Branch, Davidse 3.S$S02 (BRH, MO, SEL); 

one of the si({es of the sheath is almost rounded at Bladen Res., Ek Xux archaeolor,i( bll site, Davidse & Holst 

36041 (MO, MY); upper Bladels 13ltlnch basin, along main 

apex and the otle! is usually acute at apex. How- Bladen Canyon, Davidse & Hollelll(l 36505 (BRH, MO); 

ever, the sheall, llsat subtend inflorescences in D. Columbia Res. near Crique N( rtot Whitefoord 3284 (BR, 

Oerstedii may 1sas e the apex auriculate on both MO). COSTA RICA. Alajuela: I (lr,tlna Hule, NE of Cerro 

Congo, Luteyn 3342 (DUIkE). > sides. 

(IIlz.l)ert & Rogerson 619 

(A); canyon of Rio Cariblan(o (XX(l W slope and summit 

The tyl)e loctllily of Dieffenbachia oerstedii col- of ridge between Rio Cariblars( o (XX(l Quebrada Quicuyal, 

lected by ()e lst (l lemains in doubt but it may be SW of Cariblanco, GrayunI et (ll. f)/S)4 (MO); 3 km N of 

from Gualerzlll. 'I'he type specimen contains only La Luisa and 15 km N of Gteeia. !llfllr/)hy & Jacobs 1289 

(DUKE, MO); San Carlos, Hal(l^ (t the name ''ScXsgtlisillo'' on the label. Also penciled 

Hel 1799 (MO); 

Cordillera de Tilaran, San Ralssols-Bajo Rodriguez, vic. La 

on the label ix llse name "Aguacato," perhaps add- Balsa, 8.9 mi. NW of centel ol Sals l4am6n, Croat 68069 

ed at a latel (|lle. Engler (1915: 15) cited a col- (CR, K, MO); 10 km N of }3ija¢zla, Croat 36473 (DUKE, 

lection ma(le ly ()ersted from "Schottige Berwalder F, INB, MO); Upala, Brasilia l.r) kzl S of town, Finca de 

des Berges Aguae ate" (perhaps Cerro Aguacate in Mario Jir6n, Herrera 1636 (lNI3. M()); 22 km NE of Quesada 

by air, 4 km W of Muellx Stlls Carlos, Liesner 14102 

Sierra de las Minas). He also listed "sanquinello" (MO); 22 km NE of Quesad;l l)y ait; 4 km W of Muelle 

(Engler, 191.5: 1 S) as a common name for D. oer- San Carlos, Liesner 14155 (13, K, M()); 2 km N of Santa 

stedii (note the (lifference in spelling from that orig- Rosa, Liesner et al 15035 (13. Ml'aXU, MO); Cant6n de 

inal label). 

Upala, Colonia Blanca, Finx (s lS io Negro, Rivera 1560 

(INB, MO); rd. to Los Angelees. NW of church in San Ra- 

It is possiT)le that Dieffenbachia oerstedii occurs 

m6n, Stone 3317 (DUKE); Sars lltltssd)n-Bajo Rodriguez, 12 

in the Lesser Anti]les and in French Guiana. How- km NW of Los Angelos, 16 kzl NW of San Ram6n, Croat 

ard 11744, pulpoltedly collected along the road be- 78830 (MO); vic. km markels I 1-12, ca. 7 km NW of Los 

tween Rousseau and Sulfur Spring on Dominica, Angelos, 11-12 km NW ol Stlls llam6n, Croat 78859 

(INB, MO); Rio Grande de 'I'til( ol(s, vic. Capulin, Stanappears 

to l)e tllis species. Another specimen coldley 

40223 (US); Rio Segundo, ( (l. I km N of Intl. Airport, 

lected in Frelle h Guiana (Croat 74319) along the Croat 36855 (MO); Rio Zapote. I, islopes of Volcan Mira- 

Cayenne-Regina highway, as well as cultivated ma- valles, W of Bijagua, Burger e t (ll. / /695 (F, MEXU, MO) 

terial going by the name "D. oerstedii variegata" San Carlos (Pital), 1 km N de lSa l jegua, Haniniel 20231 

(Lyon 67-1086), appears to also represent this spe- (INB); Rio Samen hacia Upala, Aguilar 5204 (INB, MO); 

San Ram6n, ca. 13 km NW ol Surl Ram6n, Luteyn 3263 

cies. The same taxon is in cultivation in Port-au- (DUKE); along rd. from San Ram6n, N of Balsa, ca. 13.8 

Prince, Martinique (perhaps originating from km N of bridge over Quebrada Volio, Stevens 13761 (MO); 

French Guiana). Since the species is not apparently 3.5-4 mi. W of San Ram6n, Croflt. 467$S5 (INB, MO); Rio 

widespread in the areas mentioned, the field-col- Grande, San Isidro, Carvajal 273 (K, MO); Upala, 13.8 

km N of Bijagua, Croat 36444 (MO); Upala Cant6n, Est. 

lected plants may in fact be plants escaped from Biol. San Ram6n, Dos Rios, Chinchilla 137 (INB, MO); 

cultivation. Living specimens studied of the mate- Upala, Rio Zapote, 4 km NNE of Bijagua, Croat 36297 

rial from French Guiana and the cultivated material (MO); 3.5 km W of Fortuna, 2.5 km NW of New Volcan 

from the I,yon Arboretum are characterized by hav- Arenal, Taylor & Taylor 11660 (MO). Cartago: slopes of 

Miravalles above Bijagua, L. Goniez 19038 (MO); Cant6n 

ing the midrib creamy white from the lower 1/4 to 

de Turrialba, 6 km W of La Suiza on the rd. to Pacayitas, 

the upper 1/4 Of the blade. A similar feature is found Kennedy & Solonion 4629 (F, INB, MO). Guanacaste: 

in some populations of D. oerstedii in Costa Rica. SW slopes of Volcan Rinc6n de la Vieja and Volcan Santa


2004 

Croat 


Maria, Burger & Pohl 7825 (F); La Tejona, N of Tilaran, tel Pink Paradise, near sea level, 5 km S of bridge over 

Standley & Valerzo 46000 (US); PN Guanacaste Est. Mengo, 

Volcan Cacao, INBio 186 (INB, MO); Tilaran, San Pedro 

de Rio Chiquito, Monteverde, Haber & Bello 7191 

(CR, MO); La Cruz-Santa Cecilia, Finca La Pazmompa, 

Rzos et al 109 (CR); PN Rinc6n, Rivera 623 (K, MO); Las 

Rio Agujas on rd. to Jaco, Croat 79075 (INB, MO); Sector 

Esquinas, vic. Fila Gamba hills behind Esquinas Rain 

Forest Lodge, along Quebrada Negra, at end of side rd. 

off of Villa Bricena to Golfito Rd., Croat & D. Hannon 

79284 (CAS, CM, COL, F, MEXU, MO, NY, SEL, TEX, 

Nubes de Rio Chiquito l km NW of village on Cont. Divide, 

Haber & Atwood 9163 (INB, MO); PN Guanacaste, 

VEN, WU); Nicoya Peninsula, Curu, Sanders et al. 19322 

(MO); Canton de Buenos Aires, Ujarras, headwaters of Rio 

Est. Cacao, Carballo et al. 47 (MO); Cord. de Guanacaste, 

Rinc6n de la Vieja, near refugee camp, along rd. NW of 

Kuiye, trail to Olan, Chacon 364 (F, MO); Coto Brus, Osa 

Peninsula, Sabanillas de Limoncito, L. G

742 

Annals of the 


38647 (F). Jutiapa: vic. Jutiapa, Standley 7S669 (F) be- of Yaxchilan, Breedlove 33839 (DS); vic. Palenque artween 

Mita and Asuncion Mita, Steyermark 31763 (F). Pe- chaeological site, Davidse et al. 20326 (MEXU7 MO), 6- 

ten: between Finca Yalpemech on Rio San Diego and San 12 km S of Palenque on rd. to Ocosingo7 Breedlove 34977 

Diego on Rio Cancuen, Steyermark 45410 (F). Quezal- (DS); Pichucalco, cerca Campo Aviacion, F: Miranda 7S46 

tenango: Retalhuleu, near Chivolandia along rd. to San (MEXU); Solusuchiapa, 24 km below Ixhuatan along 

Felipe, Standley 87180 (F). Retalhuleu: Pueblo Nuevo road to Pichucalco, Breedlove 24166 (DS). Oaxaca: Choa- 

Stricker 342 (US), Rio Samala, vic. San Felipe, Steyermark pan, Mpio., San Juan Lalana, trail Jalahui-Arroyo San Pe- 

34555 (F). Sacatepequez: below Barranco Hondo, Stan- dro, Noriega & H. Vasquez G. 1353a (MEXU). Tabasco: 

dley 88988 (F). San Marcos: 3 km SW of San Rafael Pie Cerro las Campanas, 3 km E of Teapa, ca. 50 km S of 

de Cuesta, Harmon & Fuentes 4740 (MO); Rio Chopal, Villahermosa, Conrad et al. 2863 (MO);vic. of Teapa, Tea- 

Finca E1 Porvenir, S-facing slopes of Volcan Tajumulco, pa-Tacotalpa, 3.1 m E of Teapa, Croat & D. Hannon 

Steyermark 37456 (F); Volcan Tajumulco, Steyermark 65370 (MEXU, MO); 3 km E of Teapa along rd. to Jalapa, 

37153 (F); near El Molino, Standley 78523 (F); Rio de Croat 40107 (MO); Grutas de C)cona near Teapa, Davidse 

los Esclavos valleye near E1 Molina, Standley 60715 (F) et al. 29516 (MEXU, MO); raeotalpa, 3 km E of Lazaro 

Volcan Tecuamburro, Heyde & Lux 4654 (GH, K, NY, US) Cardenas, Cowan 2063 (MEXU, MO); Teapa, Cerro las 

Solola: S-facing slopes of Volcan Atitlan, above Finca Campanas, 3 km E of Teapa, c a. 50 km S of Villahermosa, 

Moca, Steyermark 47908 (F). Suchitepequez: near Pueb- Conrad & Conrad 2882 (MO). Veracruz: Cordova, Bourlo 

Nuevo, Standley 66844 (F), near Las Lajas, Standley geau s.n. (P); Las Palmas-Catemat o, km 1S, Leija & Garza 

58291 (F). HONDUItAS. Coyol, Carleton 508 (US), Stan- 3341 (MEXU); Las Palmas-(3cltemaco, km 18, C-12-A 

dley 26305 (F, (H, US). Atlantida: along rd. for munic- Gonzalez 3341 (MEXU);LasCru(es, Las Choapas, G6mezipal 

water supply of Tela, Lancetilla Botanical Gardens, Pompa 1505 (F); Ebiotrolottl, A/laya 1 (MEXIJ); 6.5 km 

on rd. ca. 2 mi. WXW of Tela and S of main hwy., Croat from Santiago Tuxtla and 3.6 km on trail to Cerro del 

& D. Hannon 6462L? (MO, TEFH, US); Campamento Que- Vigfa, Gonzalez 5597(MEXU);(atemaco, 10 km N of Sonbrada 

Grande ca. l() km SW of La Ceiba, base of N slope tecomapan, vic. Hotel Playa E.S( ondida, Nee 23733(B, F 

of Pico Bonito, Lie.s/ler & Mefia 26236 (MO); Lancetilla, GH, MEXU, MO, NY); E sidev of entrance of Laguna de 

4 km S of Tela, Me/l(l 190 (TEFH); sendero a la Pica, E1 Sontecomapan into the Gulf of Mexico, 7 km NE of Son- 

Dorado, Cruz 354 ('I IEFH); ca. 3 mi. S of Tela, We/)ster et tecomapan, Nee 22595 (E M()); (Cerro Cochinitos, near 

al. 12624 (US), W(6.ster et aI. 12625 (F, MO, US); Tela, Catemaco, R. Hernandez .S12 (F MEXU, US); Playa Esvalley 

above Exp. Stat., MacDougal et al. 3193 (ENCB, condida, 12 mi. airline NN\X' of Sontecomapan, Holstein 

F, MO, NY), Macl)ellg(ll et al. 3299 (MO); near Tela, Pfei- & Arnibruster 20425(U(3);(;t.EAl,,i.S of Catemaco near 

fer 2163 (US), KSltKl/dley 52702 (F, US); 10 mi. Sld of Tela Zapoapan on rd. to Acayu(.ll vlJoor() Jr. & Bunting 8928 

along Rfo Lane etilla, Croat 42645 (INB, MEXlJ, MO (BH); 5.7-6 mi. from CateAlllul( o orl rd. to Santecomapan 

PMA, TEFH); llSo San Alejo, S of San Alejo, KSt(lndley Moore Jr. & Blleting 893X(B[f); Catemaco-Montepio, 12.1 

7700 (F), La Cei))a area, 35 km S of La Ceiba oll rd. to km beyond end of asphalt llw. poltion, 22.2 km N of 

Olanchito, Madi.so/Z. 712 (GH); La Ceiba, Mt. Cangrejal, Catemaco, along border trail to tllee l os Tuxtlas Res., Croat 

Mt. Cangrejal, l a (3eeiba area, Y7lncker et al. 8395 (1z, GH, 78687 (MO); along rd. betweetl (gateemaco and Montepio, 

K, MO, US). Colllayagua: ca. 2 km S of Lake Yojoa 2.6 km S of Los Tuxtlas F iel(l Sttltion, Croat 78695 (MO 

Balick et aI. 1738 (MO). Copan: 24 km E of Santa Rita, WU); 3 mi. SW of Sontecollltly)alle I os Tuxtlas, Barlow s.n. 

Harmon & Dwyer 4()35 (MO, MEXU); Nueva Al( adia, 6 (US); Chinameca, Pajapan, 7( lle (t *1l. 4466 (MEXU); Comi. 

S, Harmon & l+5uentes 6419 (MO). Gracias a Dios: atzacoalcos, 6 mi. E of Cont%l(oul(osX Hwy. 180, Croat 

Ahuas Bila, 200 km SW of Puerto Lempira, Nelson & Cruz 40062 (CM, MO, QCA, TEF H); Hidalgotitlan, 3 km SW 

9215 (MO, TEFH, UMO). Yoro: Cordillera Nolliere de of Campamento La Laguna? N e)2W'993(F, M, NY, XAL); 

Dios, hills S of San Jose de Texiguat, Davidse et al. 34478 Hidalgotitlan, 1 km SE of Agustin Melgar, Nee 29752 

(MO). MEXICO. Campeche: E1 Maculisal, 40 knl al sur (MO); Rio Soloxuchil, entre Hnos. (3edillo y la Escuadra, 

de la carretera, Escarciga, Chetumal, Bravo s.n. (MEXU M. Vasquez et al. V-907(XAIU);lcls Choapas, Las Cruces, 

30199) (MEXU). Chiapas: Chiapas, Calbrera et al. 1937 Nerling & Gomez-Pampa /.5()T (1q); Misantla, 8 km S of 

(MO); Izabal, Mpio. Livingston, E. Martinez et al. 23175 Misantla on rd. to Xalapa, Ma(li. esl 589 (GH, SEL); San 

(MEXU); 2 mi. S of Chiapas border along Hwy. 19S, 8 mi. Andres Tuxtla, Est. de Biol. Iro^)i(al Los Tuxtlas, N of 

N of Pichucalco, Croat 40087 (MO), Croat 4008(? (MO); San Andres Tuxtla betweell Xollte^( omapan and Montepio, 

60 mi. SE of Palenque, Croat 40163 (CHIP, MO); near Croat & D. Hannon 63129 (MEXU, MO), 19 Aug. 1972, 

ruins at Palenque, KSpellman et al. 164 (F, MO, NY); 2.5 Madison 627 (GH); San Andres Tuxtla, N and E sides of 

mi. N of Isthuatan, Croat 47868 (MO); Bochil-Pichucalco, Laguna Encanatada, 3 km NE of San Andres, Nee et al. 

17.1 km SW of Pichucalco, Croat 78678 (MO); 2() km S 24759 (F, K, MO, NY, SP, XAI,), Ibarra 455 (MEXU), 

of Palenque on rd. to OcosingoS Mayo & Madison 301 (K); Ibarra & Cedillo 1804 (MEX, MO, NY), barra 645 

Mpio. Ixhuatan, Clarke 60 (DS); Ococingo, 19 km NW of (MEXU, MO), Dillon et al. 1837 (F, MO, NY), Cedillo E 

Crucero Corozal, E. I#artinez 13445 (MEXU, MO); Tene- 3645 (MEXU, MO), Calzada 338 (F, MEXU), Chazaro 416 

japa, along Rio Tanate at Habenal, paraje of Mahben- (F, MEXU); Monte Pio, 15 km al W de Catemaco, L. Gonchauk, 

Breedlove 12752 (F); Palenque Archaeological zalez 1473 (ENCB, MEXU); Santiago Tuxtla, 6.5 km de 

Site, 3 mi. S of Palenque, Thorne & Lathrop 40559 (RSA); Santiago Tuxtla y 3.6 km camino al cerro del vigia, Leija 

Rio Cuxtepeques, near Finca Gadow, Breedlove 40155 & Garza 5597 (MEXU); Santiago Tuxtla, cerca Madero, 

(DS); Las Margaritas, Breedlove & McClintock 34192 (DS); Ramamoorthy et al. 3763 (MEXU); Tezonapa, 5 km SW 

Mapastepec, Rfo Testecapa, 10 km SE of Mapastepec of Motzorongo, vic. Felipe E. Martinez, R. Robles G. 815 

Breedlove & Thorne 30681 (DS); Ocosingo, 5 km SW of (MEXU, XAL); Tlapacoyan, along Rio Tablazos, upstream 

Santo Domingo, 120 km SE of Palenque on rd. to Bonam- from Puente de Tomata, 6 km SSW of Tlapacoyan, Nee et 

pak, Davidse et al. 20440 (CM, MEXU, MO); Ocosingo, 3 al. 26103 (F, NY, XAL). PANAMA. Chiriqui: vic. David, 

km NW of Vertice del Chisoy camino a Boca Lacantumf Pittier 2836 (MO); 13-20 km W of Rio Chiriqui Viejo, 

E. Martfynez S. 13630 (MEXU); Rlo Usumacinta, at ruins D¢Arey 10766 (MO); rd to Rfo Serrano, Folsom et al. 2111


2004 

Croat 


Figure 21. Dieffenbachia panamensis. A. Habit of flowering plant (Croat 67526). B. Potted plant showing ovate 

blades (Cirino s.n.). C. Habit showing ovate-elliptic blades and inflorescences (Croat 74856). D. Crown of plant 

with blade showing quilted blades and close-up of open inflorescence (Croat 67526). 

(MO); W of Tole, Rfo Cuvibora, Hammel 6264 (MO); Burica 

Peninsula, vic. of Puerto Armuelles, San Bartolo Limite, 

10-12 km W of Puerto Armuelles, Croat 22173 (MO). 

Cocle: 4-6 km N of E1 Cope, Montenegro & Chung 1462 

(PMA, STRI), Aranda et al. 2860 (PMA); La Mesa, N of 

680-770 m, 25 Mar. 1993, T B. Croat 74856 

(holotype, MO-4342614!; isotypes, B!, CAS!, 

COL!, DUKE!, F!, GH!, INB!, Lt, K!, MEXU!, 

NY!, PMA!, UB!, US!, VEN!). Figures 21, 

E1 Valle de Ant6n, 2 km W of Cerro Pil6n, Croat 37351 

(MO). Veraguas: vic. Santa Fe, 1.7 km past Escuela Alto 

30A. 

Piedra, Croat & Zhu 76859 (MO); 6-7 km past school, 

Nee 99121 (MO), Luteyn & Wilbur 4569 (DUKE). 

Cultivated plants. Mexico. Veracruz: Catemaco, cultivated, 

Bunting 1692, Lankester Gardens, Cultivated, Kew 

70-76-494 (K); Finca La Selva, originally collected by Helen 

Young, 17 June 1971, Croat 68449 (CM, CR, MO). 

Planta 0.75-2.5 m; internodia brevia, 34 cm diam.; 

petiolus 10-14 cm longus, vaginatus ad apicem; vagina 

effusa; lamina ovata ad elliptica, (26345-75 cm longa, 

(13.5)1740 cm lata; inflorescentia 3 per axillam; pedunculus 

10-18 cm longus; spatha 13-20 cm longa; spadix 

12-18 cm longa, cum parte pistillata 5-9 cm longa. 

21. Dieffenbachia panarnensis Croat, sp. nov. 

TYPE: Panama. Cocle: vic. of E1 Cope, 4.1 mi. 

N of village, along old logging road which 

leads down to the lowlands, 8°39'N, 80°36'W, 

743 

Stout erect herb, 0.75-2.5 m tall; internodes 

short, typically tnore than twice as broad as long, 

1-3 cm long (typically somewhat longer on one side 

than on the other), 3-4(5) cm diam., dark green,



moderately glossy. LEAVES rosulate, erect-spread- this species to be certain of its flowering and fruiting 

to spreading; petioles (6)10-14 cm long (aver- ing phenology. However, flowering collections have 

aging 10.6 cm long), moderately erect, dark green been made in January, March, July, and September, 

(white at base), sheathed throughout, the margins and fruiting collections during September and Noflaring 

to recurled, sometimes incurled, markedly vember. 

crisped-undulate nearly throughout; blades ovate to Discussion. The species is recognized by its 

elliptic, (26)45-75 cm long, (13.5)17-40 cm wide fully sheathed petioles that are usually flaring to 

(averaging 4.5.9 x 24.4 cm), 1-2.4 times longer recurled, by the moderately coriaceous, somewhat 

than wide (averaging 1.6 times longer than wide), velvety, weakly quilted blades, and usually broadly 

markedly ine(3uilateral (one side 2.5-5 cm wider convex midribs. It has up to three large infloresthan 

the otlle) ol)tuse to rounded at apex (the tip cences (mostly more than 30 cm long) per axil. 

inequilatelul all(l weakly acute to acuminate), acute The species may be most closely related to D. 

to roun(le(l at l)ase, moderately coriaceous, mod- standleyi, which ranges from Nicaragua to Honduerately 

l)icololous, dark green, matte and somewhat ras along the Atlantic slope. Both share petioles 

velvety to glistellirlgly velvety above, weakly quilt- mostly fully sheathed, and have the sheath margins 

ed and minlltely wrinkled on upper surface, much rolled outwatd, but D. pananiensis differs in having 

paler to mo(lel ately paler and slightly glossy to blades more coriaceous, matte, and somewhat velmatte 

below (llying dark blackish brown to dark vety to glisteningly velvety above, weakly quilted 

yellowish l)own above, yellowish brown to grayish and minutely wrinkled on the u)per surface. 

brown below; midrib broadly convex or sometimes Though tlle species does not o(cur in the same 

flattene(l, 1..5-2 c m wide, slightly paler above, area, it colll(l be confused with O. horichii, a spesomewhat 

nlore (onvex and slightly paler below; cies with a similarly fully sheatGle(l petiole, but the 

primary lat.(ral 1peins modelately quilted, (7)10 to margins ol which are incurle(l lather than flaring. 

20 per side. al ising at an acute angle, then spread- The blades of O. horichii dry nlostly greenish rather 

ing at 4tS°-()()° angle, often at a somewhat wider than somewlsat blackened. Die/j(lll)(lchia horichii is 

angle Otl ose si(le, obtusely sunken above, convex restricted to the Pacific slope of (4osta Rica, ranging 

to obscllrels X aise(l and dal ker below; interprimary from sea le veL to 900 m, whi le D. panamensis is 

veins usucllly 1 I)er pair of primary lateral veins, primarily ksown from the Atlalltie slope of Panama 

about equal to I)rimaries; minor veins not visible. (also know s [l om the Pacific S lo,oe at higher ele- 

INFLORES(411,NCES 3 per axil; peduncles 10-18 vations). 

cm long (avetaging 12.5 cm long), somewhat flattened, 

me(lium gleen, matte, weakly striate drying, 

Paratylv(.s. IANAMA. Cocle: Allo Calvario, 7 km N 

of Cope, FolxeZzl et al. 8264 (M0); Alto Calvario, old lumto 

6-10 mlll (lianl.; spathe 13-20 cm long (aver- ber trail to Itls Ricas, Lim6n & Sclrl Juan, Croat 68820 

aging 18.4 ( m long), medium to pale green, weakly (M0); Alto (>llvario Region, 4.5 mi. 1N of E1 Cope, 2.5 mi. 

and finely striate on back surface, with darker, N of Escuelbl 13a1rigon, Croat 675L?6 (M0); 8 km above E1 

oblique lines extending out to the margins, inner Cope, Harrll)7(l 778 (M0), Folsom & Collins 6486 (M0). 

Darien: Rfo Iuquesa, ca. 2 air kzl flom Cont. Div., vic. 

surface sliglltly paler; spadix 12-18 cm long; free of Kittredge hold mine, Croat 272?()6 (M0). Panama: 

part 8.5 crn long; pistillate portion 5-9 x 1.8-2.2 above E1 (2ope, 28 km NW of Penorlome, Read & Watson 

cm, narrovve(l slightly toward the apex; sterile sta- 84-75 (US). 

minate portion 3-4.2 x 1-1.2 cm; mostly sterile 

intermediate see tion 1.2 cm long, with a few scat- 22. Dieff¢nbachia pittieri Engl. 8 I(. I(rause, 

tered stamino(lia in the lower half; fertile staminate Pflanze-lr. IV. 23 Dc(Heft 64): 42. 1915. 

portion 5-7 c m long, the flowers irregularly 5-sid- TYPE: Panama. Canal Area [Colon1: Gamboaed, 

2.5-3 nlrn diam.; pistils 3.5-4 mm diam., ir- Las (3 l uces [currently Madden Forest Reregularly 

rounded at apex, closely packed, almost serve], 50-80 m, 2 July 191 1, H. F: Pittier 

contiguous; staminodia club-shaped, 4-5 mm long, 3766 (holotype, US!; isotype, B not seen). Figgenerally 

extending above the pistils, broadest at ures 22, 29B. 

base and apex, the tips 1.5-2 mm diam. Fruits orange. 

Herb, to probably less than 1 m tall; internodes 

1-1.5 cm long, 1.5-2 cm diam., smooth, drying 

Distribution and habitat. Dieffenbachia pana- dark yellow-brown with conspicuous pale petiole 

mensis ranges principally along the Atlantic coast scars; petioles 10-11 cm long, fully sheathed, freeof 

Panama from 50 to 850 m, on both slopes along ending and weakly auriculate on both sides at apex; 

the Continental Divide east of the Canal Area. blades obliquely oblong, 17.8-20.5 cm long, 6-8.0 

Phenology. There are not enough collections of cm wide, 2.9-3.1 times longer than wide, 1.8 times


2004 

Figure 22. Dieffenbachia pittieri. Herbarium type specimen. 

Croat 


g7 

a d

746 

Annals of the 


longer than petioles, shortly acute and markedly a broad sterile segment rather than having them 

inequilateral above, acute to obtuse at base, pri- contiguous or nearly so. 

mary lateral veins 6 to 7 per side, concentrating 

mostly in lower half of the blade, extending parallel 23. Dieffenbachia seguine (Jacq.) Schott, Wieto 

the midrib, then spreading at 25°-30° angle, ner /. Kunst 1829(3): 803. 1829. Arum seguinearly 

straight to weakly curved to the margin and ne Jacq., Enum. Syst. P1. 30. 1760. Arum sesweeping 

up along the margin, drying moderately guinum, orth. var. Caladium seguinum (Jacq.) 

obseure above, convex and darker than surface be- Vent., Mag. Encycl. 30. 1801. Diegenbachia 

low; minor veins parallel with obscure and irregular plumieri Schott, Oesterr. Bot. Wochenbl. 69. 

ross-veining on drying; surface drying matte, mod- 1852. TYPE: Plumier, Descr. P1. Amer. tab. 

el-ately dark yellow-brown above, light brown and 61 (lectotype, designated here). Figures 23, 

finely pale-speckled below. INFLORESCENCES 2 30B. 

pel axil; peduncles 3.5-5 cm long, drying light CaladiunI niaculatunI Lodd., Bot. Cab. tal. 608. 1822. 

brown, 2 mm diam.; spathe 17 cm long, to 3.5 cm 

wide when flattened; spadix 14.5 cm long; pistillate 

portion of spadix to 8.5 cm long, 4 mm wide; staminate 

portion of spadix 5 cm long; mostly sterile 

Dieffenbachia niaculata (Lodd.) G. l)on, in Sweet, 

Hort. Brit. ed. 3, 632. 1839. Dieffenbflchia niaculata 

(Lodd.) Bunting, Baileya 10: 145. l 963, nom. superfl. 

TYPE: Lodd., Bot. Cab. tab. 6()8. 1822 (lectotype, 

designated here). 

segment 2.2 cm long, drying 2 mm diam., with a 

few scattered cup-like pistillodes throughout its 

length; pistils 59, borne in a cup-like disk; staminodia 

mostly missing. 

Dic)Jfenbachia literata Schott, Oesterr. Bol. Wochenbl. 68. 

1852. Dieffenbachia seguine fo. litllrata (Schott) 

Engl., F1. Bras. 3(2): 175. 1878. I)i(.ffenbachia seguine 

subvar. Iiturata (Schott) lErlr,l., Bot. Jahrb. 

Syst. 26: 568. 1899. Dieffenbachi(l .s(guine var. Iiturata 

(Schott) Engl., Pflanzenr. 1V, 2.< Dc(Heft 64): 

Distribution and habitat. Dieffenbachifl pittieri 

is known only from the type in the Isthmux of Panama 

in an area of Tropical moist forest (T-mf) life 

zone (Holdridge, 1967) at less than 200 m elevation. 

It is most unusual to encounter an endemic 

species in this life zone, but the species is distinct 

from other species in Dieffenbachia. There is a possibility 

that this could be a hybrid between D. kil- 

47. 1915. TYPE: based on a cullivtlted collection 

fide Schott (not seen). Schott Pailllillr 1874 serves 

as the type [microfiche 64. c T] (I( ( lolype, designated 

here). 

I)ietfienbachia lineata K. Koch 8 BoLlel>, Index Sem. 

(Berlin) 14. 1853. Dieffenbachifl seglline fo. Iineata 

(K. Koch & Bouche) Engl., F1. Blas. :s(2): 175. 1878. 

Dieffenbachia seguine subvar. Iill(ata (Schott) Engl., 

Bot. Jahrb. Syst. 26: 569. 1899. Dicwffenbachia seguine 

var. Iiturata (Schott) Engl., Pflanzenr. IV, 23 

lipii and D. isthmia because hybridization does oc- Dc(Heft 64): 47. 1915. TYPE: New Cranada, based 

c ur in the wild in Dieffenbachia. Its greatest on a cultivated collection at the Bf rlin Botanical 

Garden, not sczen. Schott illustratioll 1867 serves as 

esemblance is with D. killipiiS which also occurs the type [microfiche 64 d T] (le(lol!|)e, designated 

in this area and dries a similar color. Dieffnenbachia here). 

pittieri differs from D. killipii in having a dark Di().ffenbachia robusta Schott, Oesterr. I3OI. Wochenbl. 65. 

brown, smooth-drying stem (light yellow-brown, 1854. Dieffenbachia seguine var. robllsta (K. Koch) 

Engl., Bot. Jahrb. Syst. 26: 568. 1Xf39. TYPE: Lousually 

conspicuously ridged stems for D. killipii), cality unknown, cultivatczd colle( tioll of unknown orfully 

sheathed petioles (usually with the petiole igin at Berlin Botanical Garden. S( hott illustrations 

sheath ending well below the base of the blade for 1907-1908 serve as the type [mi( I-ofi( he 66 b2, b3] 

D. killipii), blades with the midrib flat on the upper (lectotype, designated here). 

Dicvffenbachia consobrina Schott, Syn. Aroid. 131. 1856. 

surface ("square-raised" for D. killipSi), and a spa- TYPE: Brazil. Rio Negro, Martius s. rl . (holotype, M) . 

dix with the pistillate and staminate portions widely Dieffenbachia poeppigii Schott, Syn. A Z oi(l. 130. 1856. 

separated by a long, mostly sterile portion (vs. with TYPE: Peru. Without locality, Poep/)ig s.n. (holotype, 

the pistillate and staminate portions more or less 

W destroyed during World War II); S( hott illustration 

1903 serves as the type (microfiche 64. b 6) (lectocontiguous 

in D. killipii). 

type, designated herc). 

Dieffenbachia pittieri has been confused with D. Dieffenbachia cognata Schott, Syn. Aroid. 130. 1856. 

Obscurinervia (Standley, 1944; Croat, 1978), but it TYPE Suriname. Paramaibo, I}Iartius s.n. (holotype, 

proved to share little in common with that species 

M). [See also Schott paintings 1845 & 1846.] 

Di().ffenb(lchi(l gollnterian(l Sf hott, Oesterl. Bot. /. 8: 387. 

once the type specimen was examined. It differs l858. rl=YPE: Venezuela. Without lo( ality, Collmer 

from that species in having smooth, not scurfy s.n. (holotype, B apparently lost); Sf hott drawing 

stems, unspotted, fully sheathed petioles, blades 1857 (microfiche 64 b. T) serves as the type (lectowith 

distinct primary lateral veins and with a midtype, 

designated here). 

Die.JJenbachia neglecta Schott, Bonplandia T: 30. 1859. 

rib flat, not "square-raised" above, and a spadix TYPFi: Jamaica. Without localityS Distin (holotype, 

with pistillate and staminate portions separated by K!).

Volume 91 Number 4 

2004 

Croat 


Figure 23. Dieffenbachia seguine. A. Habit of plant showing the long creeping caudex and erect portion of stem 

apex (Croat 60613). B. Potted plant showing habit and leaves with variegated blades and inflorescences at anthesis 

(Croat 69695). C. Close-up of crown of plant with abaxial surfaces of blades and side view of the spathe (Croat 

53921). D. Petiole showing slightly free-ending, inequilateral sheath and acutely sulcate free portion above sheath 

(Croat 68494). E. Crown of plant showing speckled petioles and inflorescence at anthesis (Croat 77298). F. Spathe 

at anthesis, cut open to show inner surface of tube. G. Close-up of inflorescence showing apical half with spathe 

blade somewhat spreading and staminate portion of spadix protruding somewhat forward. H. Portion of spadix showing 

bicarpellate pistils. F, G, H. (Croat 71828). 

747

748 

Annals of the 


DiefJenbachia ventenatiana Schott, Bonplandia 7: 30. green and glossy soon becoming dark green, se- 

1859. Dieffenbachia seguine subvar. ventenatiana miglossy to almost matte, smooth. LEAVES arching; 

(Schott) Engl., Bot. Jahrb. Syst. 26: 568. 1899. Dieffenbachia 

seguine var. ventenatiana (Schott) Engl. petioles 10-34.5 cm long, averaging 29.7 cm long, 

Pflanzenr. IV, 23 Dc(Heft 64): 48. 1915. TYPE: Su- medium green and weakly glossy, rarely white, 

riname. Without locality, Hostrnann s.n. (holotype, sometimes densely pale green maculate, sheathed 

K!). 

2/3_85 or more their length, free part of petiole 5-7 

DiefJenbachia lingulata Mart. ex Schott, Prod. Syst. Aroid. 

334. 1860. Dieffenbachia seguine var. Iingulata cm long, acutely sulcate; sheath 20-28 long, pale 

(Mart. ex Schott) Engl., F1. Bras. 3(2): 175. 1878. green and matte on inside, usually acute on one 

Diefferl6achia seguine subvar. Iingulata (Schott) side, rounded on the other side, sometimes inequi- 

Engl., lSot. Jahrb. Syst. 26: 568. 1899. TYPE: Brazil. laterally acute on both sides, curved inward along 

Para: (J. M(lrtius 3307 (holotype, M!). 

the margins but the sides not contacting, faintly 

Dieffienb(lchi(l irror(lta Schott, Prod. Syst. Aroid. 335. 

186(). Die.lRn6achia seguine fo. irrorata (Schott) striate throughout but especially noticeable toward 

Engl., Fl. IAras. 3(2): 175. 1878. Deffenbachia se- the base, the basal portion of the sheath often perguine 

sul)var. irrorata (Schott) Engl., Bot. Jahrb. sisting after much of the petiole falls free; blades 

Syst. 26): 5623. l899. Dieffenbachia seguine subvar. ovate-lanc eolate, 17-38.5 cm long, 10-20 cm wide 

irrorflt(l (S( holt) Engl., Pflanzenr. IV, 23 Dc(Heft 64): 

48. 1()15. 'I'YI'Id: Brazil. Para: C. Martius 2640 (ho- 

(averaging 32.1 x 17.3 cm), 2.7-3.3 times longer 

lotypt?, M!). 

than wide, 1.2-1.6 times longer than petioles, ine- 

Die,0enbachi(l ( ons/)lircata Schott, J. Bot. 2: 52. 1864. quilateral, one side 1.5 cm broader, inequilaterally 

Die.,ffierl b(l ( hi(l seguine fo. conspurcata (Schott) Engl., rounded at base or with one side acute the other 

F1. Bras. -3(2): 175. 1878. TYPE: Brazil. Para: Schott rounded, subcoriaceous, semiglossy, medium-dark 

painti rlgs 184(3 & 1850 (microfiche 65 b5 & b6) 

selve ax thee ty)e (lectotype, designated here). green, sornetimes mottled with pale green or white, 

Dieffenb(x(11i(l b(lrr(l.(lliiniana Verschaff. & Lem., Ill. Hort. especially along midrib, moderately paler below; 

ll: 1. 3297. 1X64. Dieffenbachia seguine fo. barra- midrib fklttened-convex and slightly paler, ca. 6 

quini(111(l (Vels(haff. & I,em.) Engl., F1. Bras. 3(2): mm wi(le above, narrowly rounded and slightly pal- 

]74. 129729. 1)i()@fenbachicl seguine subvar. barraquiniarla 

(Veels(htllf. & Lem.) Engl., Bot. Jahrb. Syst. 

er below, sometimes darker green-maculate in low- 

26: 5()tA. 189(). Dieffenb(lchia picta var. barraquini- er half; )r-imary lateral vein.s 13 to 15(19) per side, 

ana (Vels( hall. & Lem.) Engl., Pflanzenr. IV, 23 arising tlt 40°-50° angle, quilted-sunken and con- 

D((Hell 64): S(). 1915. TYPE: not designated; the colorous tlbove, concolorous and concave below; inilluslltiorl 

ill lll. Hort. 11: t. 387. 1864, serves as terprimclly veins usually present and nearly as conthet 

ty^)e (le( lolype, designated here). 

Dieffenba(11.i.(l ;,yiA,r(ll1tea Verschaff., Ill. Hort. 13: t. 470, spicuous; as primary lateral veins; minor veins 

471. 1866. 1)ieffenbachia picta subvar. gigantea moderately indistinct, arising from the midrib and 

(Vers( llaff.) Etlgl., Bot. Jahrb. 26. 569. 1899. TYPE: parallelillg the primary lateral veins. I1NFLORESnot 

(leesignatee(l; the illustration in Ill. Hort. 13 470- CENCES 1 to 4 per axil, usually solid green but 

471. 1866, setves as the type (lectotype, designated 

here). 

sometimes pale greenish yellow maculate through- 

Dieffenba(l1ifl wellli. ii Linden, Ill. Hort. 17: t. 11. 1870. out (ma( ulations sometimes appearing in irregular 

DieJkIlb(lc 11i(l seguine subvar. wallisii (Linden) transverse rows on spathe); peduncles 2.5-14 cm 

Engl., Iflarl>enr. IV 23 Dc(Heft 64): 47. 1915. long, 7-X x 8-12 mm diam., medium green, weak- 

TYItId: Brazil. Rio Negro, 1866, Wallis s.n. (holotype, ly glossy, faintly dark green-striate; spathe 11-24 

K!). 

Dieffenba(hia bra.siliensis Veitch, Cat. 12. 1875. DiefJen- cm long, abruptly acuminate at apex, gradually 

bachi(l picta fo. brasiliensis (Veitch) Engl., F1. Bras. constricted above tube, in the upper 2/3, medium- 

3(2): 176. 1878. TYPE: Brazil. Not designated; dark green and semiglossy to matte outside, slightly 

drawing on p. 12 of Veitch Cat. serves as the type paler and glossy within; spathe blade at anthesis 

(lectotype, designated here). 

DiefJenbachia picturata L. Linden & Rodigas, Ill. Hort. 

stiffly erect then recurving near apex; spathe tube 

39: l()l. t. 163. [CLXII] 1892. Dieffenbachia picta 7-10 cm long, 1.2-1.8 x 2.0-2.5 cm diam.; spadix 

Schott subvar. picturata Engl., Bot. Jahrb. Syst. 26: 10-19 cm long; the naked portion at base 1.5-2.5 

570. 1899. TYPE: Venezuela. Illustration in Ill. cm long; pistillate portion of spadix 4-8.5 cm long; 

Hort. vol. 39: t. 163. 1892, serves as the type (lec- pistils 20 to 25, closely aggregated except in upper 

totype, designated here). 

DiefJenbachia seguine (Jacq.) Schott subvar. ventenatiana 10-12 mm, with up to 3 of them in a loose spiral 

(Schott) Engl., Bot. Jahrb. Syst. 26: 568. 1899. Dief- across the spadix but usually with 1 or 2 at any 

fenbachia seguine (Jacq.) Schott, var. ventenatiana level on the spadix; ovary bicarpellate, markedly 

(Schott) Engl., Pflanzenr. IV. 23 Dc(Heft 64): 48. bilobate, rarely 3- or 4-locular (and 3- to 4-lobate, 

1915. Diefenbachia ventenatiana Schott, Bonplandia 

7: 30. 1859. TYPE: Suriname. Hostnian (K!). 

respectively), pale green, semiglossy, 3.2-3.6 x 

2.1-2.4 mm; stigmas pale orange, doubled (one for 

Herb, to 1.5 m tall; stems reclining at base then each locule) but usually fused along the adjoining 

erect; internodes 1.7-5 x 1.5-4.0 cm, medium margins, 2.3-4.3 x 2.0-2.4 mm diam.; staminodia


2004 

Croat 


4 per pistil, 2.8-3.1 mm long, 0.8-1.0 mm wide at wendlandii. The latter species was even erroneousapex, 

slightly thickened toward apex, flattened and ly synonymized with D. seguine by Engler (1915). 

free at base, the base equally as wide or up to twice Both species sometimes have blades of similar 

as wide as the apex; fertile staminate portion of shape, and both have petioles that are comparably 

spadix 5-7.5 cm long, 5-15 mm diam., slightly sheathed and sometimes sharply C-shaped on the 

broader midway, tapering slightly toward both ends, free portion. However, D. wendlandii differs in havbluntly 

acute to rounded at apex, at anthesis pro- ing pistils with a single ovule. The staminate spadix 

truding forward out of spathe and usually being of D. wendlandii is also more or less tapered toward 

trapped there by the closing spathe; staminate flow- the apex and not protruding strongly forward at aners 

5 to 6(8?) per spiral, (1.6)2.2-3.5(4) mm diam., thesis, whereas in D. seguine the spadix is stubbysquarely 

rounded to rounded, sometimes broadest ellipsoid. 

perpendicular to the axis, smoothly rounded at I agree with Bunting (1979), who synonymized 

apex, sometimes with a transverse linear slit me- both D. picta Schott and D. rrzaculata (Lodd.) G. 

dially; anthers 5 to 6 per synandrium, shedding Don under D. seguine. This species has been the 

their pollen well below the rim of the synandrium; most confusing of all Araceae in the number of epthe 

mostly naked portion of spadix 2.0-4.0 cm long, ithets (subspecies, varieties, subvarieties, and 

6-8 mm diam., medium green with 1 pistil in lower forms) recognized, over 60 in all. Some are varieties 

1/3 and with 2 to 3 staminodes in upper 1/2, some- of D. picta, others varieties of D. seguine or comtimes 

with only a few staminodia scattered in upper binations between the same two taxa. It is not clear 

3/4. Berries bright red or orange. 

why this should be so. Throughout its range it is 

Distribution and habitat. Dieffenbachia seglline 

no more variable than any other species of Dieffenranges 

throughout much of the West Indies from bachia, but it is quite widespread and this brings 

Cuba (Ileana Arias, Havana, pers. comm.), Jamai- it into contac t with more workers. Most of the epc 

a, Hispaniola, and Puerto Ric o, through the LeKsKser ithets were l)ased on Schott names, but Engler 

AntilleKs to 'I'rinidad and South Amerie a, there rang- treate(l most of these as varieties or sul)varieties of 

ing throughout much of Venezuela (Ama%onaKs, Ar- D. .seguine or D. /)icta. All of the taxa involved 

agua, A)ure, Bolivar, Caral)obo, I)elta Amae UIo, share a suite of chara(ters that make it unique, 

namely the shar^)]y sulcate l)etioles the )rotru(ling 

Distrito Fe(Setal, Fal((')n, 1,ta, Miran(0a, Monagaks, 

Nueva Esparta, Portuguesa, Su(re, Ta(hila, 'lruji- an(l thickene(l staminate poltion of the sl)a(lix, an(l 

llo, Yarae uy, %ulia), eksl)e( ially in the Cordillera de bit arpellate ovaries. 

1a CoKsta, to (,uyana and Suriname (Boggen et al., A e ultivate(l e ollectioll from Gl enada (Croat 

1992), Frell( h (uiana, eakstern Brazil (Amal)a, 77298) ha(l sap that was; c elely-s( ente(l, not at all 

Amazonaks, (Joials, Maranhao, Para, RondoniaS Sao smelling of oxalic aci(l, tyl)ical of so rnany DiefWen- 

Paulo), and west to the lowlands of Colombia (Meta, b(lchi(l species. Croat 78X2S, frorll Venezuela, also 

Vaupes, Vie hada), eastern Ecuador (lfapo), and Bo- has a sweet- rather than a foul-se ented sap. 

livia (Pando, Santa Cruz). 

Engler (1915) placed Dieffenbachi(l lanciJolia 

DiefWenlv(lchifl seguine flowers and develops ma- Linden & Andre into synonymy with D. picta, I)ut 

ture fruit to some extent all year-round, but with the former is a distinct species from Antioquia Demore 

flowering occurring in the late dry season and partment, Colombia. Also excluded are combinathroughout 

the early part of the dry season between tions of the name, DieJfienbachia picta Schott forma 

March and September. 

lancifolia (Linden & Andre) Engl. and D. picta 

The species is characterized by having petioles Schott subvar. Iancifzolia (Lind. & Andre) Engl. 

that are shorter than the blade and sharply sulcate Also erroneously placed in synonymy with D. 

on the free portion, with the sheath margins acute picta was Dieffenbachia nieleagris Linden & Rodito 

rounded at the apex and with ovate-lanceolate gas and the combination subvariety nieleagris Engl. 

blades. The best character for separation of the spe- That species is from Ecuador (see illustration 159 

cies, however, is the unusual inflorescence with in Ill. Hort.) and looks like it might be synonymous 

large, bicarpellate ovaries and a spathe that is with D. spruceana Schott. 

somewhat spread backward while the staminate Dieffenbachia shuttleworthiana Engl. was synportion 

of the spadix protrudes prominently forward onymized with D. picta but should be excluded 

and persists after the spathe has closed. along with the following combinations: D. picta for- 

The species was long considered to be a common ma schuttleworthiana (Regel) Engl. and D. shuttlespecies 

in Central America but that material proved worthiana Hort. Bull. This Colombian species is 

to be mostly D. oerstedii, and in some cases, D. not closely related to D. seguine. 

749

750 

Annals of the 


Additional specintens examined. Andre 1202 (K)7 Ca- niark 91896 (NY7 US); Los Caracas-Higuerote Rd.7 bedet 

6030 (K). BOLIVIA. Pando: Nicolas Suarez7 a few tween km 5 and Osma7 Bunting & Fernandez 3295 (NY); 

km N of Cobija7 Beck 17103 (LPB7 MO). Santa Cruz: Pico Naiguata7 Fila Las Delicias Naiguata7 vic. Las Deli- 

Velasco7 10 km SE of buildings of Est. Flor de Oro7 Nee cias7 Bunting & Manara 2126 (NY)7 Bunting & Manara 

41364 (MO7 NY). BRAZIL. Engler 226 (K, P), Engler 227 2119 (NY); Rio San Julian7 just above Caraballeda7 Bun- 

(K7 US). Mato Grosso Sul: Catharino et al. 1909 (SP). ting 2044 (NY). Falcon: PN Quebrada de la Cueva E1 

Para: Mpio. Capanema7 Rfo Quatipuru7 near Miraselvas7 Toro7 Liesner et al. 7681 (MO). Lara: Dtto. Jimenez7 PN 

ca. 30 km by rd. W of Braganc,a7 Davidse et al. 18102 Yacambu7 Quebrada Negra7 Davidse & Gonzale: 21012 

(MO); Rfo Anajas7 Anajas-Vista Alegre7 Cuanta do Anajas7 (MO). Miranda: Cerros del Bachiller7 5.5 km S of village 

Beck et al. 295 (NY)7 Rfo Anajas7 Ilha do Marajo7 Prance Santa Cruz on Hwy. 9 between Guatire and Cupira7 Croat 

et al. 30247 (MO). Rondonia: Alta Floresta do Oeste7 53956 (MO); Cerros del Bachiller7 near E end7 10 air km 

Rodovia P-507 Gonfalves et al. 224 (MO). Sao Paulo: Sao W of Cupira7 Steyerniark & Davidse 116647 (MO)7 PN 

Paulo7 Instituto de Botanica de Sao Paulo7 Secao de Fi- Guatopo7 Fila la Guzmanera7 Santa Teresa-Altagracia de 

toecologia7 Lotto s.n. (K7 MO). COLOMBIA. Vaupes: Rfo Orituco7 ca. 8 km S of jet. of rd. to Caucagua7 Bunting et 

Kuduyari7 tributary of Rfo Vaupes7 Schultes & Cabrera al. 2072 (NY)7 2074 (NY); Sta. Teresa-Altagracia de Or- 

17890 (NY). Vichada: Rfo Meta7 Manati7 Cuatrecasas ituco7 Aristeguieta 1764 (NY)7 Croat 21741 (MO); Cau- 

4225 (US). ECUADOR. Napo: Auca Oil Field7 Ingram cagua7 600 m7 Bunting et al. 2073 (NY). Monagas: Mor- 

1169 (MO7 SEL). t'RENCH GUIANA. Vic. of Saul7 route ichal Las Tetas7 tributary of Rfo Tonoro7 near Aguasay7 

to Bellizon7 100-300 m past Eaux Claires7 Croat 74137 Montes s.n. (MO); Caripe-Santa Ines Rd.7 ca. 3 km E of 

(MO7 US7 VEN); Piste de Belizon7 pk 21.87 Billiet et al. Teresen7 Colorado-Los Cigarrones7 Rio Colorado7 Bunting 

6257 (MO)7 Crique (Jabaret7 bassin de 170yapock7 Creniers 2729 (NY); Jusepin-Barcelona Rd.7 6 km SW of Jusepfn7 

9896 (CAY7 K). Cayenne: Colline de Montravel7 Ile de Bunting 2738 (NY); Caripe-Maturin Rd.7 Bunting 2678 

Cayenne 7 Le Goff A. 95 (CAY7 MO) 7 Piste de Saint-Elie7 (NY); Morichal E1 Esfrerzo7 Jusepin7 M. He redia 60 (MO). 

15.7 km S of ORS'l'OM 

camp7 Prevost 3258 (CAY7 MO)7 Nueva Esparta: Margarita Island7 Juan Griego trail7 J. 

Prevost 3580 (CAY7 MO); Centre ORSTOM de Cayenne A. Johnston 305 (NY7 US). Portuguesa: Quebrada A1- 

Pre'vost 3382 (CAY, MO); Saint-Laurent-du-Maroni7 Saul7 gatrobo7 tributary of Rfo Morador7 7 krn NE of Boca de 

trail to Mont (Jalhao, Mori & Gracie 18717 (MO7 NY). Morlte7 22 km NE of Rio Suruguapo7 kSt(yermark et al. 

PERU. Loreto: Allo Amazonas7 Dtto. Manseriche7 Pongo 127204 (MO); Guanare-Biscucuy Rd.7 1 l luente7 21 km 

de Manseriche7 t?. IVoj(1ts et al. 607 (B7 MO7 US7 WU). abovee jet. of Guanare-Barinas rd.7 BZ111.1.is1.g 3321 (NY); 

VENEZUEI jA. Alllzllas: Rio Guainia7 S of Maroa7 Ma- Dtto. Araure7 Rio Auro (1JA IJUCia)7 NW of Sta. Lucia7 

guire et al. 3646/ (NY); Cano Mosquito7 Cano Marieta7 Aysz1(lrd & Ortega 3090 (M()); Dtto. (^^l lrsare7 Guanare- 

Lister & Colche.ster 272 (K)7 Tencua7 Colchest(r 2108 (K)7 Bat ias7 Mpio. Mesa de Cava( as7 Sterp,tio.s ( I al. 7940 (MO7 

Dpto. Atabapo7 Rfo Clll-ucunuma7 Raudal Picure7 J. Pe'rez P()lWl). Sucre: E1 Pilar-(ffvIaliquen7 El Pilar and Guari- 

& M. Sosa 671 (M()); Cano Majagua7 E. & S. Zent 2189 que.Z! 4-10 km S of E1 Pi]al7 (Croat 5438() (MO). Tachira: 

(MO); Casiquiare7 Rfo (ûainia7 Bunting et al. 4108 (NY); 8 klll S of E1 Pinal7 Steyermark et al. 1()2()36 (MO7 MY); 

Dtto. Atures7 Rio (2ataniapo7 45 km SE of Puerto Ayacu- Sarl Cristobal-Chorro del lndio7 Cano Xe(to-La Florida 

cho7 Steyerniark (t *ll. 122412 (MO). Apure: Dtto. San krn 14-20 al E de San Cristobal7 Buntill;,r 11687 (MO)7 

Fernando7 Rio AIau( a, 5 km SW of E1 Faro, Davidse & vi. I as Minas7 N of La Laguna7 16 krrl Sl of Santa Ana7 

Gonzalez 13410 (M()). Aragua: betw. Maracay and Ocu- Ste-(rnlark & Liesner 118891 (MO). Trujillo: Sabanamare 

de la Costae Henry Pittier NP7 1.6 km N of summit7 Mel(loza7 ca. 5 km below Betijoque7 Bllr7ting 2829 (NY). 

Croat 60567 (CM7 1R MO); Pittier Park7 Paraiso trail to Yaracuy: Las Trincheras-El Cambur7 N ol Salom (NE of 

Pico Periquito7 Cro(lt 21412 (MO); Ocumare-Portachuelo7 Ni>ua7 W of Valencia) on rd. to Candelal i l7 Croat 54649 

Steyerniark & Kogers 119364 (MO7 USS); Portachuelo- (CM MO); Marin and Aroa7 Sierra de Atoa7 Dtto. Felipe 

Paraiso Trail7 neal Itclncho Grande7 Bunting et al. 1937A between Albarico and Tesorero7 9.8 krn 1N of jet. of hwy. 

(NY); Rancho Gtan(le to the Toma7 Bunting et al. 1949A 1 at Marin7 Croat 60613 (MO); Sierra de Aloa7 PN Yurubi7 

(NY); 12 km NW of lXancho Grande7 Bunting 2035 (NY) 0-2 km N of San Felipe7 Rio Yurubi7 I,i(.wner & A. Gon- 

Bunting 2036 (NY), Bunting 2622 (NY); Maracay7 campos zale.z 10126 (MO7 NY); Sierra de Aroa7 vic. Aracal7 7 km 

de la Facultad de Agionomia y Centro de Investigaciones above San Felipe7 Bunting et al. 1997 (NY); Cerro La 

Agricolas7 Univ. Cential de Venezuela7 near Pozo del Dia- Chapa7 N of Nirgua7 Steyerniark & Buntltlg 97728 (NY); 

blo7 Bunting 2026A (NY); vic. Maracay7 Bunting 2027 Dtto. Nirgua-Dtto. San Felipe7 Cerro l a (hapa7 7 km N 

(NY); Rio Tuy7 NE of Maya7 7 km SE of Colonia Tovar7 of 1\ irgua7 Davidse et al. 20914 (MO); (e rro La Chapa7 

Steyerniark & Liesner 121834 (MO). Bol*ar: La Gran Steyerniark et al. 100213 (NY7 US); Cerio I a Chapa7 Stey- 

Sabana7 E1 Dorado-Santa Elena7 km 2027 S of Salto Kama7 erniark et al. 100318 (US)7 Steyerniark et *ll. 100239 (US); 

Davidse et al. 4844 (MO); base of Altiplanicie de Nuria7 Dtto. Nirgua7 Salom-La Candelaria Rd.7 (4umbre Game- 

Bunting & Holniquist 4319 (NY); W of Hato de Nuria7 lotal7 Meier et al. 5164 (MO7 VEN); I)tto. Bruzual7 

Steyerniark 88864 (NY7 US); Rio Karuai7 base of Sororo- Montana de Maria Lionza7 W of Sorte7 Sleyerniark et al. 

pan-tepui, W of La Laja7 Steyerniark 60796 (NY); near E1 124955 (MO); Dtto. San Felipe7 PN Yurul)i Ayniard et al. 

Paito7 5 km S of Cano Paso Ancho7 Bunting & Trujillo 2714 (MO); Dtto. Urachiche7 Quebrada Higueronal7 W of 

2236 (NY); Kamarata7 Bogner 965 (M). Carabobo: PN Urachiche7 vic. Sabana de Mendez7 Steyerniark et al. 

San Esteban7 Rio San Esteban7 Benftez de Roias et al. 124629 (MO); Rio Guayabito7 15 km N of Marin7 Steyer- 

4661 (MO7 MY); Rio San Gian7 5-6 km S of Borburata7 niark & Bunting 105293 (MO). GREATER ANTILLES: 

Steyerniark 94331 (MO7 US). Delta Amacuro: Rio Ama- HAITI. Nash 611 (NY); massif de la Lelle7 Port au Prince7 

curo between "vuelta larga77 and Cerro Wuausa7 Dept. An- Mantflurry7 Eknian 1983 (K). HISPANIOLA. SANTO DOtonio7 

Trujillo et al. 17397 (MY). Distrito Federal: Fila MINGO. San Cristobal: Cordillera Central7 San Cristo- 

Las Delicias above Naiguata7 Bunting & Manara 2124V bal7 Hato Damas-Mano Matuey7 Croat 68582 (MO). Bar- 

(NY)7 Drake 7 (P7 US); Cerro Naiguata7 6 km SW de los ahona: Fuertes 564 (GH); Santo Domingo7 Zanoni et al. 

tanques de la Electricidad de Caracas (Cocuizal)7 Steyer- 27170 (GH7 MO7 NY); E1 Aguacate7 La Leonor7 Moncion7


2004 

Croat 


Liogier 13307 (GH, NY); Carib Territory, Bataka, J. Higgins 

119 (MO, NY); Sierra de Baoruco, Arroyo La Travesia 

near the La Travesia sugar mill, Zanoni et al. 25159 (MO, 

NY). E1 Seibo: E1 Guaraguao-Los Hurados, Miches-Higuey, 

km 15, Mefia & Raniirez 9929 (MO, NY). Monsenor 

Nouel (Peravia): Cordillera Central, Rio Yuboa, Zanoni 

et al. 23113 (MO, NY); Cordillera Oriental, Arroyo de 

Agua, 11.7 km W of E1 Valle, Croat 68494 (MO); 9.2 km 

SE of Miches, along Rio Yeguada, Arroyo Santiago, Croat 

68547A (JBSD, K, MO). Samana: 2.3 km S of Playa E1 

Valle, Meiia & Zanoni 6614 (MO, NY); Sanchez, Las Canitas 

Mts., N. Tclylor 48 (NY); Sanchez Ramirez, Rose et cll. 

4375 (NY); Rio Cevicos, W of Cevicos, 22 km E of Cotui, 

Zanoni & Pinientel 23420 (MO, NY). San Cristobal: San 

Cristobal, a Jina (de Yamasa), 16 km del Parque Central 

de Yamasa, Zanoni & Pinientel 23456 (MO, NY); near 

Rio Nigua, Lavastre 1845 (NY); near Nigua River, La 

Toma, Augo 1693 (NY); E1 Dajao, Bayaguana, Liogier & 

Liogier 20185 (NY). JAMAICA. Port Antonio: Hitchcock 

s.n. (MO); Fury River, Harris 8361 (NY); John Crow Mts., 

Britton 4126 (NY). Portland: foothills of John Crow Mts., 

W of Ecclesdown, GrayunI et al. 9973 (K, MO). St. Elizabeth: 

Frenchman, Proctor 38601 (NY). PUERTO RICO. 

Cultivated speciniens. Society Islands. Leeward. Raiatea 

Island, cultivated in Uturoa, 17 Dec. 1926, J. t 

Moore 451 (MO). U.S.A. Hawaii: Honolulu, Kalihi Valley, 

15 Mar. 1956, Harris & Neal s.n. (BISH); Foster Gardens, 

3 Mar. 1962, Miyashiro s. n. (BISH); Lyon Arboretum, 10 

Sep. 1975, 13. Herbst & S. Ishikawa 5459 (BISH); Oahu, 

5 May 1954, Won s.n. (BISH); Honolulu, Kaimuki, 5 May 

1954, R. Won s.n. (BISH); Kapalama Heights, Kamehameha 

Girls School, 6 June 1932, Judd et al. s.n. (BISH). 

Missouri: Missouri Botanical Garden, originally obtained 

from J. Henny, Apopka, Henny 7 (MO), Croat 78287 (MO); 

13 Sep. 1990, Miller & Schniidt 5551 (MO). Puerto Rico. 

Rio Abajo (originally collected by Thonipson 3238), 19 

June 1997, Croat 78345 (MO). Hispaniola. Border of PN 

Los Haitises, 27 Apr. 2000, Luther s.n. (Holst 6237) (MO). 

Grenada. Originally collected by John Criswick, Grenville, 

Grenada, 1993, Croat 77298 (MO, VEN). Belize. Cayo: 

Nabatunich, near Sukkotz, 17°06'N, 89°W, 24 Jan. 1990, 

Balick et al. 2359 (MO). Venezuela. Carabobo: originally 

collected by Bunting from a Garden in Valencia, 300-500 

m, (Bunting 13515) Croat 78323 (MO, VEN); Jardin Botanico, 

Caracas, 19 Aug. 1976, Croat 38344 (MO). 

Without exact locality: Engler 2793 (K), Sintenis R 2793 

(K), Britton & Britton 9664 (NY), Britton & Britton 7897 

(NY); Mariccao, Sargent 643 (MO); Scln Sebastian, Sargent 

340 (M()); L>lguna Tortuguero, Howard & Nerling 16978 

(A); Quebrcl(la T'rieta, 1Ll Ver(le Field Stcltion in I,uqui]lo 

24. Dieffenbachia standleyi Croat, sp. nov. 

TYPE: F4ontluras. Laneetilla Botanical Gardens, 

ea. 2 mi. WSW of Tela and S of main 

hwy., 15°44'N, 87°27' W, 70-90 m, 9 Feb. 

Experimental Forest, W si(le of Lu(luillo Mts., W/lilehill 8 

(M()); Luquillo Mts., Palrrleer-FIoli(la, klll 28.1, K. W(lA,trler 

T58 (A); Rfo At)cljo Stclte Foest, Hwy. ()21, CSlro(ll. 6()867 

(M(), INY, I'MAe REA); ktll 21.S orl Rle. 2, W of Cclr(lelclticl, 

Solomosl .S761 (M()); Mclytlguf z vi( ., (louell .S.S() 

1987, YE B. Croat & D. Hannon 64638 (holotyl)e, 

M()-344288t3!; isotypes, B!, CAS!, COI,!, 

EAP!, '!, CH!, INB!, K!, MEXU!, PMA!, NY!, 

Tl



Figure 24. Dieffienbachia standleyi (Croat 42676). A. Habit of cultivated plant. B. Stem and base of leaves 

C. Upper part of stem showing abaxial surface of leaf blade and cluster of inflorescences. D. Close-up of petiole 

bases showing the markedly undulate petiole sheaths.


2004 

Croat 


brown below; rnidrib slightly paler and flat to sul- Specimens from Nicaragua differ somewhat from 

cate and finely paler striped above (especially near those in Honduras, having leaf blades smaller on 

the base), sometimes white in distal 2/3, convex to average, ranging from 37 to 69 cm long and 13 to 

narrowly rounded, thicker than broad and paler to 31 cm wide (averaging 51 x 22.3 cm), but the 

slightly paler below, drying 8-12 mm wide, medium petioles in both areas average 30 cm long. The 

yellow-brown, finely ribbed with the ribs minutely Honduran populations have petioles consistently 

and obscurely scabridulous on magnification; pri- fully sheathed, whereas plants in Nicaragua often 

rnary lateral veins 12 to 24 per side, arising at an have a free portion of the petiole above the sheath 

acute angle, then spreading at 40°-65°(70°) angle, ranging from 1 to 9 cm (averaging 4.4 cm long) with 

obtusely sunken and weakly quilted, paler toward the petioles only rarely fully sheathed. 

midrib above, bluntly angular and + concolorous Blade shape is quite variable. Blades can be 

below, drying pale yellow-brown; lower surface with from 1.9 to 2.8 times longer than wide on the same 

minor veins moderately obscure, arising mostly plant (e.g., Croat & Hannon 64638). Plants from 

from the midrib but also from the primary lateral the Lancetilla Valley in Honduras have proportionveins. 

INFLORESCENCES 2 to 4 per axil; peduncle ately longer blades, with some plants having blades 

to (7.5)15-29 cm long (averaging 16 cm long), up to 3.2 times longer than wide (Croat 42676). 

somewhat flattened; spathe matte, 20-50 cm long Specimens collected at Lancetilla and its surround- 

(averaging 28.3 cm long), 1.2-3.1 times longer than ings have blades that range from 1.9 to 3.5 times 

the peduncle (averaging 1.6 times longer), 2.5-3.0 longer than wide (averaging 2.5 times longer than 

cm diam.; spadix 20-30 cm long (averaging 25 cm wide). Specimens from areas other than Lancetilla, 

long); pistillate portion of spadix 12-13 cm long; and outside Honduras, have blades 1.6 to 2.7 times 

staminate portion 15-16 cm long. Fruits to 1 cm longer than wide (averaging 2.1 times longer than 

long, closely packed, reddish orange to scarlet at wide). 

maturity with the spathe light yellow. 

A c ollection from Zulia, Venezuela, between l, 

Frfa and San Juan de Cold)n (Croat 78289) differs 

Ditstriblltion (lnd h(ll)it(lt. Die,t,tenbflchi(l .st(ln- in having the petiole less fully sheathe(l an(l in havdleyi 

ranges from Honduras (Atlanti(la, Comayagua, ing narrower bla(les up to 2.6 times longer than 

Cortes, El Paraiso, (Jrae ias a l)ios, ()lane ho, Yoro) broa(l an(l drying somewhat less hlackene(l. It may 

to Nicaragua (Matagalpa an(l %elaya) at 3()-1()0() [)rove to t)e the same species. If so, it wouI( [)e the 

m elevation. Most c ollee tions in Hon(luras were only (:entral American sI)eeies (with the [)ossit)le 

made in the lian(etil]a Va11ey. 

exce,Dtion of D. killi)ii) that ranges from Central 

Phenology. F]owering s)ee imens of D. .st(lndle- Americ a to Venezuela. 

yi have been seen from May through Septeml)er, Etynology. Diefjfenl)lchi(l .standleyi was first 

although a few flowering c o11ee tions were also made collectecl in 1928 by Paul Standley at Lan(etilla, 

as early as February and March. Fruits start to de- Hon(luras. Standley was not only the author of most 

velop in February and reac h full size during March c)f the aroid treatments for most existing Central 

or April (in Nicaraguan collections) or July-August American floras, but he also described a number 

(in Honduras). 

of new species of Araceae, as well as collecting 

Discussion. The species is characterized by its several paratypes of D. standleyi. The species is 

stout stem, fully winged petioles, yellow-brown-dry- named in his honor. 

ing ovate-elliptic blades with 12 to 24 pairs of primary 

lateral veins, and a long spathe (20-50 cm). Paratypes. HONDURAS. Atlantifla: 5 km S of La 

It is most easily confused with D. horichii and may Ceiba, Madison 705 ((;H); near Lancetilla, Yuncker 4961 

(F, MO, NY); Lancetilla valley, near Tela, Pfeifer 2124 

ultimately prove to be inseparable from that spe- 

(US); near Puente Alto stop on S.F. Co. R.R., E of Ceiba, 

cies. Dieffenbachia horichii differs in having shorter Yuncker et al. 8551 (F, GH, MO, NY, US); ca. 3 mi. S of 

petioles with the sheath involute and moderately Tela, Webster et al. 12581 (DAV); vic. Tela, near Rio Lansmooth, 

in contrast to the petiole sheaths erect and cetilla, above Exp. St., MacDougal et al. 3303 (MO, NY, 

even curled outward as well as being undulate 

US); Tela, ca. 10 mi. SE of Tela along Rio Lancetilla, 

Croat 42625 (MO), Croat 42676 (MO), Standley 53146 (F, 

along the margins in D. standleyi. 

US). Comayagua: Siguatepeque, Standley & Chacon 

A collection by Stevens et al. 20998 from Mata- 6701 (F). Cortes: 2-3 mi. SW of Omoa on rd. from Puerto 

galpa is unusual in being described as 3 m tall with Cortes to Guatemalan border, Croat 42556 (MO); E1 Para 

trunk 1() cm in diameter. Another Stevens col- aiso, Yuscaran, Rio de los Aguacates, Standley 25700 

(EAP). Gracias a Dios: Ahuas Bila, 200 km SW of 

lection (7457) from Zelaya Department is notewor- 

Puerto Lempira, Nelson & Cruz 9291 (TEFH, UNAH, US). 

thy in drying darker yellow-brown than other col- Olancho: Catacamas, Standley 18786 (EAP); along Rio 

lections. 

Olancho, W of main Tegucigalpa-Catacamas Hwy., ca. 1 

753

754 

Annals of the 


km upstream from and NW of Puente Boquer6n, 8.6 mi. throughout (this white coloration continuing onto 

SW of Catacamas, 6 mi. SW of Sta. Maria del Real, Croat the midrib), sheathed virtually throughout; sheath 

& D. Hannon 64109 (INB, MEXU, MO, TEFH, US); along 

Rio Olancho, on rd. Gualaco-San Esteban at Rio San erect to involute (rolled inward throughout in age), 

Martin, 19.3 mi. E from Gualaco, 8.7 mi. SW of San Es- free-ending and unequally rounded at apex, proteban, 

Croat & D. Hannon 64317 (B, MO, US); Rio Olan- longed to 2 cm beyond the base of blade; uncho, 

San Esteban-Bonito Oriental, 19 mi. NE of San Es- sheathed part obsolete or rarely to 1 cm long (when 

teban, Croat & D. Hannon 64476 (MO); Refugio de Vida 

Silvestre "La Muralla," Nelson & Andino 16247 (TEFH); 

evident obtusely flattened); blades ovate to ovate- 

Mpio. Jano, 16 mi. NE of La Union along the rd. to Olan- elliptic or oblanceolate-elliptic, (15-)25-48(-63) 

hito, Davidse et. al. 35484 (MO). Yoro: near Puente cm long, (8-)15-32 cm wide (averaging 34 x 16 

Grande, on a tributary of the Rio Agua (Rio Puente cm), broadest near the middle, sometimes below, 

Glande), Blackmore & Chorley4073 (MO). NICARAGUA. frequently above the middle, 1.4-2.5(-3.5) times 

Matagalpa: El Tollsa Rd., Neill 1571 (MO); ridge along 

td. betw. fJa Dalsta arld La Luna, E of Esquipulas, Stevens longer than wide (averaging 2.1 times longer than 

11786 (CAS, M() TEX); Macizos de Penas Blancas, SE wide), 1.9-3 times longer than petioles, spreading 

side, drainage ol (ssuebrada E1 (ssuebradon, Hda. San Mar- to erect-spreading, inequilateral, one side 1-3 cm 

tin, horder with l)epartmento de Jinotega, Stevens et al. wider than the other, gra(lually to abruptly acumi- 

2()998 (MO); Matagalpa-Siuna, 1.5 km al NE de Los Angeles, 

Moreno 17/42 (MO); E1 Trebol, 7 km S of Penas 

nate, sometimes acute at apex, inequilaterally cor- 

T3lancas, rd. to fi l 'luma, Moreno & Robleto 20526 (MO). dulate at base, one side sometimes broadly rounded 

Zelaya: Finca Waylawas, Pipoly 4479 (MO, US); Siuna- to obtuse, the other side cordulate, sometimes in- 

Matagalpa, ca. 31.4 km beyond Rio Uli (near Wani), ca. equilaterally acute, subcoriaceous, often conspicu- 

8.9 km beyond llosa Grande La Balsama, Stevens 7457 

ously quilted, moderately l)icolorous; upper surface 

(MEXU! MO); Cerro l ivico, 7 km NE of Siuna, Neill 3629 

(MO); Rio Sucio, Ed of Bonanza, Stevens 12311 (MO), Ste- usually solid dark to medium green, sometimes 

vens 8044 (MO); Res. Bosawas, Bonanza, Cerro Cola Blan- conspicuously to sparsely variegated with pale 

ea, vic. Cacerio (le Vitinia, Rueda & Coronado 6557 (MO). green or pale yellow thloughout much of the surface, 

the mottling large ol small, but somewhat re- 

25. Dieffenbachia tonduzii Croat & Grayum, stricted to the area midwcly t)etween the midrib and 

Novon 9: 497. 1999. TYPE: Panama: Bocas del margin, matte to weak]y glossy, sometimes appear- 

Toro: Valle del Silencio, along Rio Changuinola, ing weakly velvety, drying glay-green to olive-green 

ca. 1 km above mouth of Rio Teribe, vic. Teribe or dark brown; lower surfcl( e much paler and matte 

Indian population, disturbed forest among co- to weakly glossy, silvely-gleen, drying yellowish 

coa plantations, 9°21 '40"N, 82°31 '40"W, less green to yellow-brown helow; niidrib flat to broadly 

than 100 m, 25 June 1994, 1: B. Croat & G. rounded and moderately to strongly paler, pale 

Zhu 76452 (holotype, MO-04611212!; isotypes, green or sometimes creanly white above (sometimes 

AAU!, B!, BM!, BR!, CAS!, CM!, COL!, CR!, only toward the apex), blutly acute to obtusely an- 

DUKE!, F!, GH!, HUA!, INB!, ITIC!, JAUM!, gular and paler, sometillles white or creamy white 

K!, L!, LE!, M!, MEXU!, NY!, P!, PMA!, below, (0.6-)1-1.7 cm wi(Se; priniary lateral veins 

QCA!, QCNE!, R!, RSA!, S!, SCZ!, SEL!, (14 to)l8 to 25(to 30) per side, arising at an acute 

TEX!, UB!, US!, VEN!, WU!). Figures 25, angle and spreading at 45°-90°, sometimes reflexed 

29B. 

toward the base, promillently to weakly and obtusely 

sunken above, convex to weakly raised and 

Terrestrial herb, 0.5-1.5 m tall, usually to less darker than surface or concolorous below, some of 

than 1 m tall; internodes 1.5-4.5(-6) cm long, 1.5- the lowermost with a weak fold near the base (Croat 

3(-4.5) cm diam., usually solid dark to medium & GrayunI 60112), somelillles convex-pleated begreen, 

sometimes faintly marbled with gray-green low; interprimary veins almost as conspicuous as 

or yellowish gray throughout (on plants that also the primaries; minor veins moderately to distinctly 

have streaked petioles), initially weakly glossy, be- visible, darker than surface below. INFLOREScoming 

semiglossy to glossy, often with a subvelvety CENCES 1 to 2(to 4) per axil, often with two orisheen; 

petiole scars manila to white, curved down- ented in opposite directions; peduncle (3-)6-17 cm 

ward on the opposite side of the stem and ending long (averaging 10.3 cm long), 7-8 mm diam., 

unevenly; petioles 10-24 cm long (averaging 17.6 weakly glossy, dark to medium green, sometimes 

cm long), held + erect, medium green (except with pale yellow-green streaks; spathe (12-)15-28 

sometimes white to pale green at base), almost cm long (averaging 20 cm long), 2-4 times as long 

matte to weakly glossy, weakly striate (especially as peduncle, acuminate at apex, convolute to about 

near the base), narrowly rounded to obtusely an- the middle in lower part, matte to weakly glossy 

gular on abaxial surface and often white medially, outside, glossy within, solid medium green on both 

sometimes streaked in a variegated pattern surfaces, gradually and weakly constricted some-


2004 

Croat 


Figure 25. Diegenbachia tonduzii. A. Close-up of stem, showing both creeping and erect portions. B. Habit 

with inflorescences and variegated blades. C. Crown of plant with open inflorescence. D. Close-up of stem showing 

petioles streaked with creamy white. E. Close-up of stem showing solid green petioles. F. Close-up of spathe with 

the blade portion open (Croat & Zhu 76452¢ type plant). G. Inflorescence open to show upper male flowers and 

lower portion of spadix with pistils and staminodia (Croat 66533). A, B, C. (Croat & Zhu 76452¢ type plant); D, E. 

(Croat 76450). 

755

756 

Annals of the 

Missou ri Botanical Garden 

what above the middle; spathe tube 1.5-3 cm diam. Because of the fully sheathed petioles it can be 

when closed, 6.8-9 cm wide when flattened; con- confused only with D. horichii and D. crebripistilstricted 

area 4.3-4.5 cm wide flattened; spadix (9- lata. Both of the latter typically have much larger 

)16.5-25.5 cm long, about as long as or up to 3.0 leaves that are not at all cordulate at the base 

cm shorter than the spathe; free portion 7.8-8.5 cm (though they are rarely broadly and weakly subcorlong; 

pistillate portion (4.8-)6-11.5 cm long; mostly date). Dieffenbachia tonduzii is polymorphic with 

sterile portion sparsely flowered to naked (rarely regard to leaf markings in the same way as D. oerlacking, 

as in Croat 70768), 1.8-4.0(-8.5) cm long stedii, and is quite variable in all respects. It apwith 

0.6-1.5 cm totally bare, the uppermost portion pears, in Panama, to hybridize with D. oerstedii. 

with a few staminodia, sometimes with a few scat- Croat & Zhu 76857C from the vicinity of Santa Fe 

tered staminodia throughout, the lower half some- in Veraguas Province is apparently a hybrid with 

times with an occasional pistil and much reduced characteristics intermediate between D. tonduzii 

staminodia, rarely with the female flowers + equi- and D. oerstedii. 

distant and nearly contiguous with staminate part The species is similar to D. daguensis, a Col- 

(Croat 70768); fertile staminate portion (4-)5.5-10 ombian species described from less than 200 m 

cm long, 7-10 mm diam., slightly broader midway, elevation on the Rio Dagua in Valle Department. 

weakly tapering to apex and base, bluntly pointed That sl)ecies also has many rather close primary 

at apex; staminate flowers 5 to 6 per spiral, + lateral veins and a fully sheathed petiole but differs 

rounded in outline, crenulate along margins, trun- in having the staminate and pistillate sections of 

cate at apex; sterile male flowers irregularly the spa(lix contiguous or nearly so. In addition, it 

shaped, 1.8-2.5 mm diam.; pistillate portion of spa- differs in having much shorter petioles (described 

dix to 11 cm long, 9-10 mm diam.; female flowers as being up to 5 cm long). 

(15 to)48 to 62, closely aggregated except in the A South American species, D. parlatorei Linden 

upper 1.5 cm of spadix, 4 to 5 across the width of & An(l re, also sometimes has petioles fully 

the spadix (upI)ermost pistil borne on an almost sheathe(l, but differs from D. tonduzii in having leaf 

bare segment of the spadix); pistils pale cream-yel- blades usually broadest al)ove the middle, semilow 

to pale yellow-green, smooth, 2.3-3.5 mm glossy on the lower surface, and the midrib often 

diam.; style (after stigma has fallen) sharply cupu- broadly l ounded and spongy. It also has the primary 

liform, 1.5-1.7 mm diam. with a single central lateral veins arising at a 40°-60° angle from the 

pore; stigmas yellow; staminodia clavate, white, 2- midrib (often at more than 60° and sometimes up 

3 mm long, mostly contiguous and sometimes fused to 90° in D. tonduzii). 

at base. INFRUCTESCENCES with spathe pale Cro(ll 70900, from 250 m in Choco Department 

yellow; berries red to red-orange, 5-8 mm diam. of Cololllbia appears to be D. tonduzii but differs 

in sevelal ways. It has leaves with the midrib flat- 

Distribution and habitat. Dieffenbachia tondu- raised aI)ove with the margins undercut. It also has 

zii ranges from southeast Nicaragua throughout stems that appear scurfy (though weakly glossy if 

Central America to the Pacific slope of Colombia rubbed clean). Another difference is that the petiole 

(Antioquia, Choc6, Cauca, Valle) and Ecuador (Es- sheath is more prominently free-ending and submeraldas, 

Loja, and Los Rios), from sea level to acute at the apex. In addition, the free portion of 

1400 m, in Tropical wet forest (T-wf) and Premon- the petiole is broadly and sharply sulcate. 

tane rain forest (P-rf) in Central America and in 

Tropical wet forest (T-wf) and Premontane wet forest Additio/lal specimens examined. COSTA RICA. Alajue- 

(P-wf) and Tropical wet forest transition to Premon- la: rd. to Colonia Virgen del Socorro, Rfo Sarapiquf, Stevens 

13564 (MO), Croat 68336 (CR, MEXU, MO, TEFH); Cantane 

(T-wf/P) in Colombia. 

t6n de Alajuela, Grayum & Murakami 9939 (CR, MO); Fin- 

Phenology. Flowering in D. tonduzii occurs ca Los Ensayos, ca. 11 mi. NW of Zarcero, Croat 43629 

throughout most of the year with flowering collec- (CM, MO); Canas-Upala, 10 km N of Bijagua, Croat 36472 

tions seen from February through November. Most (MO); Rfo Zapote, Canas-Upala, Rfo Zapote, 4 km NNE of 

Bijagua, 

collections have been made from April through Au- 

Croat 36260 (MO); Cordillera de Tilaranv San Ram6n-Bajo 

Rodrfguezv Rfo Cataratitas, Croat 68097 (INBv 

gust. According to the collections, fruits mature MO); Cordillera de Tilaran, San Ram6n-Bajo Rodrfguez, 

throughout the year but with the greatest concen- vic. of km 19.5 NW of San Ram6n, Croat 78838 (MO); 17tration 

from October to January. 

20 km NNW of San Ram6n by rd. on way to San LorenzoS 

The species is characterized by its fully sheathed 4-7 km N of Balsa, Ljiesner & Judziewicz 14797 (CRv MO); 

San Ram6n-Bajo Rodrfguez, 3S37 km NW of San Ram6n, 

petioles, usually matte to weakly glossy, sometimes 

Croat 68196 (CMS MO, W); Naranjo-Aguas Zarcas, along 

weakly velvety blades with cordulate bases and nu- Hwy. 15, 8 km NE of Quesada, Croat 46945 (K, MO, PMA); 

merous, broadly spreading primary lateral veins. Arenal Volcano, Funk et al. 10626 (CR, US), Funk et al.


2004 

Croat 


10718 (CR); San Ram6n-Fortuna, ca. km 25, D. Smith et (CR, INB, MO) Puntarenas: Cordillera de Tilaran, Bosque 

al. 1059 (DUKE, MO); Cordillera de Guanacaste, Montev- Eterno De Los Ninos, Laguna Poco Sol, 18 km ENE of 

erde, San Gerardo Biol. Sta., 1.5 km NE of Station, D. Monteverde, Haber et al. 10824 (CR, INB). San Jose: Can- 

Penneys 633 (CR, INB, MO); Cordillera de Tilaran, Bosque 

Eterno De Los Ninos, Rio Penas Blancas, Laguna Poco Sol 

on del Rio Grande de Orosi, Chacon et al. 1488 (CR, MO); 

La Hondura, Standley 36314 (US); Vazquez de Coronado, 

Haber & Zuchowski 11175 (CR, MO). Cartago: Tucur- Braulio Carrillo NP, along Hwy. San Jose to Siquirres, along 

rique, Tonduz 12874 (G, US); Rio Naranjo, Finca E1 Cedral, 

Orosi, Lent 4042 (F, MO, NY, SEL); La Vuelta, Tucurrique, 

trail to Rio Sucio, site of the Old Carillo Station, Croat 

78787 (MO). NICARAGUA. RSo San Juan: "Los Filos" 

Tonduz 503 (US); 1.5 mi. E of Cachi, 10.2 mi. NE of jet. near Loma Los Filos, Rio Santa Cruz, Salick 8153 (MO). 

at Paraiso, Croat 47088 (MO); along Camino Raiz de Hule, Zelaya: ca. 6.3 km S of bridge at Colonia Yolaina and ca. 

SE of Platanillo, Croat 36727 (MO), Croat 36747 (MO), 0.8 km S of ridge of Serranias de Yolaina on rd. to Colonia 

Croat 36821 (MO); Cant6n de Turrialba, Rio Reventazon, Manantiales, Colonia Somoza, Stevens 6420 (MO); Rio Pun- 

CATIE, Turrialba, Grayum et al. 9469 (CR, MO, NY), 3 

km E of Turrialba "Los Espaveles" nature trail, Rio Reventa 

Gorda, Atlanta, mouth of Cano el Guineo, Moreno & 

Sandino 12855 (MO), Moreno & Sandino 12891 (MO); Rio 

taz6n, Liesner et al. 15330 (MO); Turrialba, Instituto Inter- Punta Gorda, Atlanta, Cano Negro mouth of Rio Chiquito, 

am., Lent 639 (F), Lent 694 (F); 12 km S of Turrialba by 

air, 4 km SE of Pejibaye along Rio Gato, Liesner 14394 

Moreno & Sandino 12917 (MO); Rio Punta Gorda, Atlanta, 

mouth of Cano del Oro at Rio Chiquito, Moreno & Sandino 

(MO); Tres Equis, 1.5 km E of Turrialba-Lim6n Hwy., Lies- 12955 (MO); Rio Punta Gorda, Cano E1 Guineo, Tellez et 

ner et al. 15365 (MO); PN Tapanti, Oropendola trail, Nilsson al. 4875 (MEXU, MO). PANAMA. Bocas del Toro: Limit 

et al. 632 (CR); Oropendola trail, Nilsson et al. 377 (CR); trail, Parque Intl. La Amistad, from Quebrada Boca Chica 

Monumento Nacional Guayabo, Perez 1 (CR); Monumento to Quebrada Bonyic, Polanco 1615 (PMA); 7.7 mi. W of 

Nacional Guayabo, Santa Teresita, Rivera 1718 (CR, K). 

Guanacaste: E1 Arenal, Standley & Valerio 45262 (US); 

Chiriqui Grande, 1.5 mi. W of Punta Pena, Croat & Grayum 

60112 (CR, MEXU, MO, PMA); Chiriqui Grande-Fortuna, 

E1 Real, Standley & Valerio 45206 (US); Cordillera de 13.2 mi. W of Chiriqui Grande, Croat & Grayum 60139 (B, 

Guanacaste, Rinc6n de la Vieja, near refugee camp, along MO, PMA); Fortuna-Chiriqui Grande Hwy. near Cont. Div., 

rd. NW of Quebrada Grande, Barringer et al. 4039 (F). 1.1 mi. from main hwy., Croat & Grayum 60355 (MO); 

Heredia: E of San Ram6n, Loiselle 106 (MO), Porto Viejo- Chiriqui Grande-Fortuna, above waterfal1, 1.6 mi. N of 

Rio Sucio, Croat 35753 (F, MO, PMA); La Selva, Puerto Cont. Div., Croat & Zhu 76450 (CR, MO, PMA, SCZ, US); 

Viejo de Salapiqui, Croat 44317 (MO); I, Selva, OTS Field 

Station on the Rso Puerto Viejo just F, of its j(t. with the 

Changuinola, near IJuzon, Kenz1edy.X253 (MO); Changuinola-Altnirante, 

Mile 7.5, Cro(lt & I'orter 16249 (MO); (7ual- 

Rso Sarapiquf, H(lmmel 8874 (l)UKk, 11, M()), (Jr(lyalrrl a( a-(4hiriqui (7ran(1e, 6.6 trsi. N of bri(lge over Fortuna 

278() (I)UK1 , M()); 1, Selva, 6 kns l)y r(l. frorls Rlo l)PjE I,ake, Cro(lt 667XS (M()); (/uala( a-(3hiriquf (7rande, 1.6 mi. 

crossing S krs StSk, of Masasay, 5C'/I(I/.Z & (Jr(lyllm 7()6 N of Corst. I)iv., Cr()(lt 749X() (M()); (Juala( a-Chiri(ui 

(DUKId); lvuello Viejo just 1d of j( l. with Rfo SaIay)i(Uf, 

Folsom 1()116 (I)UK19); ()( ( i(lerstal tail, Kr(^.sx 84-16X() 

(lran({ee, X.1 rrsi. S of l'unta l'eena (Jro(lx 749H (MO); N of 

Forluna l)arls, M(lther.s()sl 11 129 (M()); r(s. to (Chiri(uf 

(SEl,); Rfo S[J(*io rseeal lvolto viPj0, (>ro(lt .XT682 (M()); Zona (^ran(le, M(l'hers0s1 7X71 (M()); 5.t-3 rlsi. N of 11orturla l)arrs, 

Protee(tola N slo^)es of Vol( airs 13ar1)a, })etw. Itio l)PjE All(l 

Rio (lUa( illso (,r(lyaltll & KSlhel.z W174 (I)U K11,). Lilllon: 

the.n 1.4 nsi. W alorlg gravel r(l. lo (Lorst. I)iv. trail, (ro(lt & 

X,hll 76X28 (M(), I'MA); along CoIsl. I)iv. trail, M(l'llersos 

along hwy. :32 {lorls Turriall)a to l,ills(5rl ( a. 11 rlsi. S of 

Siquillees (>ro(lt 4X.X@S (IS(E, M(), l'MA, WlA(2); (ûapiles 

6W65 (M()); tortuna l)ans-(Chili(uf (7lan(1ee, I mi. fronl Corst. 

I)iv., Chllrf hill & Cllllrcl1ill 62tH (M()); ltio (Cri( ansola, Firs- 

I,eosl 72?() (11'); (Ûa)ile.s, 'loro Aurarillo, lJeos1 26573 ((LIt), a St. l,ouis-KorlkirltoeX W)o(lsosl Jr. (t (ll. 79()9 (t, M()). 

Fin( a Arlai, at he.a(1waters of (Juebra(la Mata (lee l,irls6n, W 

of Mata (se T,inson (Sixaola legion), (Jr(lyun1 & Schfltzt5279 

Chiriqui: (7ua1a( a-11'orturla l)ans, 1 () rlsi. NW of Los Planes 

de Hornit(, Crc)(lt 50()49 ((2R, M()); vi(. 11ortuna l)am Of 

(CR, M()); 11irs(a Castilla, Do(Ige & Coerger (9489 (M()); 7 ltio Chiriqui, Croat 66533 (B, CM, lEl\CT3, l)UKIL, KY(), 

km SW of BriT31i, 1,. Gomez et al. 2()405 (B, MO); (Iraitlage I., MO, NY, OOMOT(), (2CA, SAIt, TkzX, US); 4.5-5 km 

of Rio Parisnlirla and Ri() Reventazon, Sht1k & Molina 

4288 (l)UKkJ); 1() mi. S of Punta Cahuita, ca. 3 mi. S of 

N of Fortuna l,ake, Croat & GrKlyum 60070 (INB, MO); 7.9 

mi. I)eyond (NW of) I,os l'lanes de Hornito, Croat 499.W 

turnoff to BrilXri, Croat 43201 (MO); Ref. IJarra del Colo- (MO); (ualaca-Bocas del Toro bolder, km 111, Gordon 3369 

rad(, Rfo Chirrip6cito-Rio Sardina, Grayum 9804 (CR, (PMA); lE ortuna Dam site N of Gualaca, 7.7 mi. beyond Los 

MO); Par. Tortuguelo, Est. Biol. Agua Frfa, Sendero Las Planes de Hornito, Croat 48778 (MO). Cocle: vic. E1 Valle 

Lomas, 5 km from station, Robles 1142 (CR, MO); Sendero de Anton, La Mesa, Finca Macarenita, Croat & Zhu 76653 

Los Ralldales, 8 km .SF, from station, Robles 1158 (CR (BM, C, FSU, GOET, INPA, ITIC, L, LE, MO, QCA, PMA 

MO); PN Tortuguero, Lomas de Sierpe, 4 km NE of station R, TEFH, UB). Darien: Mamey, Whitefoord & Eddy 372 

along Rfo Sierpe, Robles et al. 2050 (CR, G, MO); Tortu- (BM, MO, PMA). San Blas: Rio Armila, 10 km WSW of 

guero Cant6n, BriBri-Suretka, Barringer 3525 (CR, F); Res. Puerto Obaldia, Mori et al. 6814 (MO). Veraguas: valley 

Indfgena Talamanca, Sukut, mouth of Rfo Sukut at Rfo of Rio Dos Bocas, 11 km from Escuela Agricola Alto Piedra 

Uren, Hammel et al. 17548 (CR, MO); Rfo Reventaz6n, (above Santa Fe) on rd. to Calovebora, Croat 27490 (MO); 

Finca Montecristo below Cairo, Standley & Valerio 48997 NW of Santa Fe, 11 km from Escuela Agricola Alto de 

(US); Rfo Segundo, Asunci6n, L. Go'mez & Herrera 23477 Piedra, in valley of Rio Dos Bocas, Mori et al. 3817 (F, 

(MO); Rfo Sixaola, BriBri-Caribbean coast, Baker & Burger MO); 0.6 mi. beyond Escuela Agricola Alto Piedra, Croat 

90 (F, MO); Lim6n-Shiroles, Rfo Sixaola, 0.9 mi. SW of & Folsom 33989 (MO); 1.7 mi. past Alto Piedra School, 

Bambu, 6.5 mi. SW of BriBri, Croat 43298 (MO); Rfo Six- Croat & Zhu 76858 (MO, PMA); beyond Escuela Agricola 

aola, ca. 0.5 mi. SW of Bambu, ca. 3 mi. NE of Bratsi, Alto Piedra, Croat 49070 (MO); 3-5 mi. N of Santa Fe, 

Croat 43266 (CR, MO); Tortuguero Cant6n, BriBri-Sixaola, Gentry 3035 (MO); vic. Escuela Agricultura Alto Piedra 

NW of Paraiso, Barringer et al. 3479 (CR, F); Cordillera near, Antonio 2994 (MO); 0.6 mi. beyond Escuela Agricola 

de Talamanca, headwaters of Quebrada Kakebeta below di- Alto Piedra, Croat & Folsom 34042 (MO); Cerro Tute revide 

between Rio Xikiari and Rio Boyei, Grayum 10858 serve, along ridge to summit, Croat 66993 (HUA, MO, 

757

758 

Annals of the 


PMA); trail to top of Cerro Tute, Croat 48903 (MO, PMA); de Bimbe, Croat 57000 (CM, MO); Centinela, 12 km E of 

Cerro Tute, Sytsma & Antonio 3006 (MO). COLOMBIA. Patricia Pilar on border with Los Rios, Gentry 26705 (MO); 

Antioquia: Murri, La Blanquita, Rfo Murrf, Transect 7, Centinela, Montanas de Ila, 13 km E of Patricia Pilar, ca. 

Gentry et al. 75903 (MO); Villa Arteaga, Gutierrez, G. & 

Barkley 17115 (COL); PN Natural "Las Orquideas," Ven- 

54 km S of Santo Domingo, Hammel & Trainer 15836 (MO). 

ados arriba, Rfo Venados, A. Cogollo et al. 3462 (JAUM, 

MO); Mutata, Rfo Chontadural, Hacienda E1 Darien, Fonrl.egra 

1344 (HUA); Carepa, Est. Exp. de Tulenapa (CA), 

(>(llle.j(l.s et al. 9704 (NY). Choco: Serranfa de Baudo, Las 

Anilllas-Pato, Rfo Pato, ca. 4 km SW of Pato, Croat 56112 

(Cl-f()CC), JAUM, K, MO); Medellfn-Quibdo, km 208.5, 9 

kls W of Tutunedo, ca. 9 km E of Quibd6, Croat 56205 

26. Dieffenbachia wendlandii Schott, Oesterr. 

Bot. Z. 8: 179. 1858. TYPE: E1 Salvador. Santa 

Ana, H. Wendland s.n. (holotype, GOET!). 

Figures 26, 28B. 

(CH()CO, COL, CUVC, HUA, MO); Quibd6-Medellfnv km 

Stout herb, to (0.8)1.2-2(3) m tall; sap strong and 

103z). 14 km E of Tutunendo, Croat 56282 (CHOCO, COL 

JAUM, MO, PMA); San Jose del Palmar-Novita, vic. Santa foul-scented (only weakly foul-scented in some At- 

Rosa, (>roat 56625 (COL, HUA, MO); Quibdo-Medellfn, 25 lantic slope populations); stems erect, decumbent 

,l,i. E of Quib(l6, (>roat 52300 (F, MO, PMA); ca. 2 km E at base; internodes dark green to blackish green or 

of Playa de Oro, Croat 57427 (CHOCO, MO); Pueblo Rico medium green, glossy to semiglossy, 1-5 cm long, 

(Risalalda)-Istmina (Choco), Quebrada Anton, 15 km W of 

Santa Cecilia, 6 km SV of Choco-Risaralda border, Croat 

2-5.2 cm diam.; petioles (11.5)16-32(65) cm long 

7()S*()0 (MO); Me(lellfn-(Juibdo, 85 km W of Bolfvar, Croat (averaging 25 cm long), matte, medium dark green, 

46).X1() (MO); Ae andi-Serranfa del Darien, Juncosa 619 sometimes finely darker green-striped throughout 

(M()); Pueblo Ri(o (Risaralda)-Istmina (Choco), 1 km W of (or at least near the base), often weakly glossy to- 

(,tlulato and Rio Guarato at Risaralda and Choc6 border, 

ward the apex, moderately spongy, obtusely and 

(vro(lx 70868 (CM, MO); Nuquf, Quebrada Chaquf, Galeano 

46()() (MO); Arusf, E1 Amargal, trail to Arusf, Mora 51 shallowly sulcate (sometimes more acutely sulcate 

((j()I,), Croat & Mor(l 83696 (MO); Quibdo, Tutunendo-Alto near the apex) to D-shaped, sometimes with a slen- 

({e1 Viente R(l., 25 km N of Quibdo, Callejas & Jangoux der erect margin or often terete in populations in 

26t (HUA). Narillo: Rfo Timbiquf, Lehmann 8876 (K); easteln Mexico, sheathed (0.3)0.5-0.9 their length; 

vtllley of Rfo Im})i, l'asto-Tumaco, vic. "Palmar" 3 km NW 

ol Rie aulte, ca. 1 knl E of Texas Gulf Pipeline Maintenance 

sheath (6)12-30(45) cm long (averaging 18 cm 

Station, Rfo Inll)i, C8roat 71461 (MO). Risarallla: Pueblo long); sheath sometimes markedly undulate on that 

Ri(o, Santa Ce(ilia, Quebrada La Calera, Betancur et al. portion clasping stem, inc urled throughout its 

3()54 (MO). Valle del Cauca: Cali-Buenaventura, Lobot,tlerrero-Cisneros, 

Quebrada la Guinea at 1.2 kln E of Ciseros, 

Croat 62831 (COL, HUA, MO, UB); vic. Queremal, 

llfo Cava, Croat & Gaskin 80391 (CUVC, MO); Cordillera 

()(cidental, Rfo Digua, Cuatrecasas 15053 (US); Bajo Callength 

with the margins incurled to erect and 

touching or well-spaced, initially decurrent or nearly 

so at the apex and with one side completely hiding 

the other from above the middle, sometimes 

ima, Buenaventura-Malaga, km 51.3, Croat 71017 (MO); with the margins somewhat erect and eventually + 

lAuenaventura-Malaga, Pulpapel facilities at km 9, Croat emarginate with one side rounded, the other side 

70099 (MO). ECUADOR. Esmeraldas: San Lorenzo, Rfo 

Palavi, vic. AWA encampment, Hoover et al. 

rounded and somewhat free-ending, rarely with the 

3161 (MO); 

Awa camp to Rfo Palavf, Hoover et al. 3968 (MO); Lita-San apex })roadly rounded and free-ending; unsheathed 

l,orenzo rd., 14.2 km W of Rfo Lita Bridge (below Lita), portion of petiole (1)3.5-12(24) cm long (averaging 

Croat et al. 82305 (MO); 17.3 km E of Rio TululbfX Croat 8 cm long), terete or thicker than broad, 12-14 mm 

83126 (MO, WU); 33.0 km E of Gasolinera San Lorenzo, 

diam., 13-15 mm thick, usually obtusely sulcate 

Rfo San Jose, 1.1 km N Lita-San Lorenzo rd, Croat 83889 

(AAU, F, GB, MO, NY, QCA, S); Lita-San Lorenzo Rd., 1.2 except terete in some areas on the Atlantic slope 

km W of E1 Durango, 21.1 km W of Alto Tambo, Croat et of Mexico (the sulcus sometimes broader toward the 

*11. 82441 (MO, QCNE); Lita-Carondelet Rd., km 16, apex); blades narrowly ovate to ovate-elliptic, 

Schwerdtfeger 21422 (MO); Bilsa Biol. Res., Montanas de (15)20-55(65) cm long, (9)10-22(28) cm wide (av- 

Mache, 35 km W of Quininde, 5 km W of Santa Isabela, 

eraging 35 x 17 cm), 1.3-2.4 times longer than 

Pitrnan & Bass 995 (MO, QCNE); Fila de Bilsa, 7 km E 

of San Jose de Bilsa, ca. 80 km due SW of Esmeraldas, 12 broad (averaging 2 times longer than broad), subkm 

SE of E1 Salto on Atacames-Muisne Rd., Gentry et al. coriaceous to moderately coriaceous, acute and 

72955 (MO); Eloy Alfaro, comuna de Corriente Grande (Rfo 

Chimbagal, tributary del Cayapas), Yanez et al. 1387 (MO); 

Res. Ecol. Cotacachi-Cayapas, Charco VicenteS Rfo San Miguel, 

Palacios & Tirado 11287 (MO, QCNE); Quininde, 

Herrera-Los Monos, headwaters of Rfo Aguacatal, Palacios 

13626 (CM, MO, QCNE, US); NE of Las Golondrinas, Sitio 

apiculate to abruptly or gradually acuminate at 

apex (sometimes rounded and apiculate), acute to 

obtuse or rounded and attenuate at base, sometimes 

rounded to subcordate, slightly inequilateral, one 

side 0.8-3.0(4.5) cm narrower than the other; upper 

La Bella Jungla, Cooperativa Unidos Venceremos, Palacios 

11452 (MO, QCNE). Loja: Rfo Pichima, Forero 719 (COL, 

MO). Pichincha: Rfo Palenque Science Center, halfway 

between Quevedo and Santo Dominga de los Colorados, 

Gentry et al. 24700 (MO); rd. E of Santo Domingo-Quevedo 

rd. (beginning 10.S km N of Patricia Pilar), Caserfo Palmar 

surface dark green, semiglossy to weakly glossy, 

drying gray-green to yellow-brown; lower surface 

paler, matte or nearly so, drying yellow-green to 

yellow-brown (both surfaces dark yellow-brown on 

very old specimens); midrib flat to broadly sunken


2004 

Croat 


Figure 26. Dieffenbachia wendlandii. A. Habit, plant with open inflorescence. B. Adaxial blade surface. C. 

Potted flowering plant with open inflorescence (Croat 39749). D. Open inflorescence at anthesis (Croat 47219). 

E. Staminate portion of spadix and spathe blade. F. Close-up of spathe showing part of pistillate portion, the mostly 

sterile portion and the base of the staminate portion A, B. (Croat 47219); D, E, F. (Croat 47219). 

759

760 

Ez D. fosten (0, -8 ' ; < * ( \ 

Annals of the 


86° 85° 84° 83° 82° 81° 80° 79O -78° 

,$ ''''-- / - X A 

11°- X * 'd'§i, 11° 

4' ,, tT 

10°- 8 \ 10° 

< A 0 \+X < , 

9O * D. aurantiaca X X L,> 'X,> < 9O 

8° A D. horichii

15°- * D. galdamesiae ^ (, < 15 


2004 

Croat 

Revision 

of Dieffenbachia 

- 

85° 84° 83° 82° 81° 80° 79° 7 18° 77° 76° 

11° i e A 11° 

10°-< \ -10° 

B + ' ' FS, s 8 

* D. fortunensis f S 

A 100 o 100 Kilometers < 

_ j d 

85° 84° 83° 82° 81° 80° 79° 78° 77° 76° . 

1010° 85° 80° 75° _ 70° 65° 

10°- 9 _ X -10° 

S D. beachiana 2 * ¢ 

* D. killipii f 4 X 

A D. standleyi n 

N 

l l 

70° 65° 

Figure 28. Distribution map. A. Dieffenbachia concinna, D. fortunensis, D. galdamesiae, and D. isthmia. B. 

Dzeffenbachia beachiana, D. killipii, D. standleyi, and D. wendlandii. 

to flattened and concolorous to weakly paler above 

with fine, close, slightly paler striations, convex to 

round-raised or narrowly rounded and slightly to 

moderately paler beneath, usually faintly striate, 

drying somewhat orange-tinged and paler than the 

;. 

761 

surface, 5-10(18) mm wide; primary lateral veins 

(6)7 to 11(13) per side, weakly and obtusely quilted-sunken 

above, convex, weakly pleated-raised 

beneath, arising at an acute angle then spreading 

at (40°)55°-70° angle, slightly paler than surface 

.

762 

, C72 

89° 86° 83° 80° 77° 74° 

l l l l l l 

10o- }aN tra Ft 

7°- < A *- 

* D. Iongispatha < * 

* D. piffieri t t 

* D. tonduzii t t 

4°- >l 

H 

200 0 200 Kilometers t 

Annals of the 


'0 B .?'. 

oo 

n z r t 

X 

z | 

AX 

> 

89° 86° 83° 80° 77° 74° 

Figure 29. Distribution map.-A. Dieffenbachia obscurinervia and D. oerstedii. B. Dieffenbachia longispatha, D. 

pittieri, and D. tondazii. 

when fresh, usually drying darker than surface, closely parallel marginal veins that do not form a 

sometimes lighter than surface7 moderately straight collective, minor veins on lower surface moderately 

or weakly curved to near the margin then gradually obscure on fresh material7 more prominent and 

curved upward along the margin to form a series of darker than surface on drying. INFLORESCENCE 

. 

71 ° 

. 

-10°


2004 

a- 

Croat 


75° 7p° 6 15° 6p° 5 15° 59° 4 15° 

15°-< t i 

; f _ V * D. seguine 

B ), R . fi. 

80° 75° 70° 65° 60° 55° 50° 45° 

Figure 30. Distribution map. A. Dieffenbachia nitidipetiolata and D. panamensis.-B. Dieffenbachia seguine. 

1 to 4 per axil, bracteoles to 25 cm long, peduncle 

(7)12-22 cm long, 1-1.5 cm x 0.8-1.3 cm diam., 

subterete, pale medium green, weakly glossy, faintly 

and finely striate-streaked, spathe (16)25-32 cm 

long, narrowly acuminate to cuspidate-acuminate at 

apex (the tip turned back), medium-green, weakly 

glossy to semiglossy outside, equally colored and 

glossy within with weak, darker, short oblique lines 

running between the parallel vertical veins 

throughout the length of the spathe, the spathe tube 

rs 

763 

-20° 

-15° 

-10° 

S12 cm long, (2.7)3.5 1.5 cm diam. when furled, 

6.5-12.5 cm wide when flattened, with dense minute 

depressions scattered throughout the tube, 

constricted portion of spathe (2)3.3-4.0 cm wide, 

3.5-8 cm wide when flattened; spathe blade 3.7- 

4.5 cm wide, flattening to 3.3-6.3 cm wide, spadix 

(12)1>29.3 cm long, 2.0 1.7 cm shorter than the 

spathe, scarcely protruded forward, its stipe 1-2 cm 

long, 1.3 cm diam., the free portion of the spadix 

9-11 cm long, pistillate portion (5.5)8.5-10 cm 

-5o

764 

Annals of the 


long, 1.2-1.7 cm diam., 0.9-1.1 cm on drying (ra- til August or rarely September. What appears to be 

chis 0.9-1.2 cm diam.), the upper 1 cm sometimes a secondary flowering period may occur in the early 

with as few as two apparently fertile flowers, sta- dry season because flowering collections have been 

minate portion of spadix (7)9-16 cm long, the fer- seen in December and February. Most fruiting 

tile staminate portion (4.5)7-12.5 cm long, gradu- specimens have been made in the dry season and 

ally tapered toward both ends, (0.7)1-1.3 cm diam. early wet season from January to May. 

midway, 7 mm diam. 1 cm below the apex; mostly Discussion. The species is characterized by its 

sterile intermediate portion of spadix 2.0 l.0 cm robust, but moderately short stature, dark green, 

long, 7-9 mm diam., with a few aborted pistillate semiglossy stems; narrowly ovate to ovate-elliptic 

flowers in the lower half and a few sterile male yellowish brown, to yellow-green-drying leaf 

flowers in the apical half, often with a totally barren blades; but especially by the partially sheathed petsegment 

of up to 1-2.8 cm long; pistils (33)45 to ioles with the sheath margins decurrent or ending 

55, depressed-globose, weakly pale yellow-green, abruptly and rounded at apex with a free, unmoderately 

glossy, (2)3 to 5 across the width of the sheathed sulcate portion 1-9 cm long at the apex. 

spadix, 3.0-3.7 mm diam., 1.5-1.7 mm high; stig- In addition, the species has an unusually large 

ma yellow to pale orange, 0.6-1 mm high, 2.0-2.7 spathe for the size of its leaves, frequently exceedmm 

diam., sometimes broadly sunken medially, ing 30 cm long. 

sometimes with a prominent, protruding dome held Populations of 1). wendlandii on the Atlantic 

slightly above the outer ring; staminodia 3 to 4, free slope have petioles completely terete, rather than 

to the base, broadened toward the base, 0.8-1.2 sulcate adaxially, })ut this varies even on the same 

mm wide at base and sometimes partly fused, usu- individual. In ad(lition, these populations are more 

ally slightly thicker near apex, (2.5)3.5-5 mm long, likely to have the ^)etiole sheath even more decidwhite, 

slightly flattened, 1.5-3.5 mm wide, about as edly decurrent at the apex. At least the populations 

long as the pistils; synandria 4 to 7 visible per spi- of plants in Oaxaccl at middle elevation above Valle 

ral, 3.5 l.0 mm diam., widely spaced at base, trun- Nacional have sal) that is only mildly odorous, 

cate and smooth at apex, irregularly rounded, pale whereas elsewhere the sap is malodorous, smelling 

tan, becoming bowl-shaped and brown except for somewhat like sk ll 1l k or peccary. 

white, erect margins, margins sometimes crenate, DieJ%enbachia 1lperldlandii is easily confused with 

with thecae 6 to 8 per synandrium, these subglo- D. oerstedii on the eastern slopes of Chiapas and 

bose, ca. 1 mm long, held just below the apex of Veracruz, but D. Ilnendlandii is much more robust 

the synandrium. INFRUCTESCENCE with spathe than plants of D. oerstediip which are usually less 

green at maturity; berries red to bright orange, 6-8 than 1 m tall and have internodes usually less than 

mm diam. 

2.5 cm in diameter: In contrast, the stems of D. 

wendlandii are rarely less than 2.5 cm in diameter 

Distribution and habitat. Dieffenbachia wen- and are usually 4-5 cm in diameter. Other differdlandii 

occurs principally in seasonally dry habi- ences in D. oerstedii are the sharply sulcate (rather 

tats on the Pacific slope of Central America, rang- than terete to obtusely sulcate) petioles that are 

ing from central Mexico (Oaxaca and Chiapas) to white (rather than green) at the base, and the 

Guatemala (Escuintla, Huehuetenango, San Mar- sheath apex of which at least one margin is rounded 

cos, Suchitepequez), E1 Salvador, Honduras (Mor- and prominently free-ending. In contrast, D. wenazan), 

Nicaragua (Esteli, Granada, Matagalpa, Nue- dlandii on the Atlantic slope is more than 1.5 m 

va Segovia, Zelaya), Costa Rica (Puntarenas tall at maturity, has internodes more than 3 cm in 

Province), and Panama (Veraguas) at elevations of diameter, and petiole sheath margins acute to only 

75 to 900 m. In Mexico the species occurs on the weakly protruded at apex with the free portion of 

Atlantic slope only in the State of Oaxaca in the the petiole at most obtusely sulcate. Another way 

Serrania de Juarez at 250-705 m and in Veracruz D. wendlandii differs from D. oerstedii is by its typin 

or near the Estacion Biologica de Los Tuxtlas ically larger spathe, over 25 cm long versus less 

near the Caribbean coast at 5-165 m elevation. The than 20 cm long for D. oerstedii. 

only collection of the species in Panama, from Ba- Previously most material of D. wendlandii from 

hia Honda in Veraguas Province, is unusual in hav- Mexico and Guatemala has been mistakenly called 

ing blades more ovate and in drying pale green. It D. oerstedii, and indeed herbarium material without 

may prove to be a new species. 

good field notes is difficult to separate. Several of 

Phenology. DiefNfenbachia wendlandii flowers the collections of D. oerstedii made in lowland Veprincipally 

in the rainy season, beginning in May racruz (Holstein & Armbruster 20425; Nee 23 773^ 

and especially in June and July, but continuing un- 29752 and 29993) appear to have the petiole


2004 

Croat 


sheath somewhat decurrent without a conspicuous America. The material previously determined as D. 

free-ending apex. They are described as plants no seguine in Central America has proven to be either 

more than 1 m tall. It is possible that these rep- D. oerstedii with blades larger than normal, or D. 

resent hybrid plants since both D. wendlandii and wendlandii. 

D. oerstedii occur in lowland Veracruz. 

Additional speciniens exaniined. COSTA RICA. Ala- 

Dieffenbachia wendlandii has been confused juela: Rfo Cacao, near Atenas, L. Goniez 19572 (MO). 

with D. standleyi in eastern Nicaragua. However, Puntarenas: rd. to Monteverde, Wilbur et al. 15861 

that species occurs only on the Atlantic slope and (DUKE). EL SALVADOR. Wendland 410 (GOET); San 

Salvador, Calderon 914 (US), Renson 266 (NY, US). GUAhas 

petioles sheathed completely to the apex and a TEMALA. Escuintla: Engler 2389 (GH); Escuintla-Aloproportionately 

longer blade, ranging from 1.9 to tenango, mi. 6, Croat 42050 (DUKE, MO); Cucunya at 

3.5 times longer than wide, and averaging 2.5 times San Andre Osuna, Seler 2389 (G), Seler 2398 (GH). Huelonger 

than wide. In contrast, D. wendlandii has huetenango: Cerro Victoria, Finca San Rafael, Sierra de 

los Cuchumatanes, Steyerniark 49637 (F). San Marcos: 

blades ranging from 1.3 to 2.4 times longer than 

Finca Armenia near La Trinidad above San Rafael, Croat 

wide and averages only 2 times longer than wide. 40790 (ENCB, MO); Rio Ixpal, below Rodeo, Standley 

Throughout its range in western Mexico and Central 68724 (F); Santa Rosa, Chiquimulilla-El Ahumado, N of 

America, D. wendlandii is variable in the blade Los Cerritos, Standley 79561 (F); E of Cuilapa, Standley 

shape with the proportionately broader blades oc- 78161 (F). Suchitepequez: Mazatenango, Morales 1047 

(F, CR); vic. Tiquisate, Steyerniark 47684 (F, MO). HONcurring 

in Mexico and Guatemala, where they av- DURAS. Francisco Morazan: San Antonio de Oriente, 

erage 1.8 times longer than wide. The collections above E1 Jicarito, Standley 21077 (F). MEXICO. Chiafrom 

E1 Salvador and Nicaragua, on the other hand, pas: Escuintla-Monte Ovando, 2.8 km NW of Turquiz, 

have leaf blades averaging 2.3 times longer than Croat 47510 (MO); Escuintla-El Triunfo, ca. 1 mi. N of 

Escuintla, Croat 43813 (MO); Tapachula-Nueva Aleman, 

wide. The populations in Mexico and Guatemala 

mi. 4, Croat 43791 (CHIP, CM, MO); Mapastepec, Sierra 

have shorter petioles (averaging 19.8 cm long and de Soconusco, Croat & D. Hannon 63340 (MO); Tapachu- 

22.6 cm long, respectively) than those in E1 Sal- la-Union Juarez, at km 13.5, 1.3 mi. N of Trinidad, Croat 

vador and Nie aragua (averaging almost 30 e m 47219 (CHIP, MEXU, MO); Acacoyagua, Eji(la Eas Golong). 

lan(lrinas, Cerro Mt. ()van(lo Trail, Croflt 78480 (MO); 

A(apetagua, K. Hern(in(lez 47.S (MEXU, M(), NY); Es- 

The single Costa Rican colle( tion made in Pun- ( uintla, Esperan>a, Mfltllfl(l 167e55 (F, MEXU); Esperanza, 

tarenas Provine e, along the dry roa(l ay)l)roaching Mfltll(l(l 16X66) (MEXU). Oaxaca: Tuxtepec, 6 nli. W of 

the Montever(le reserve, is noteworthy in })eing so Valle Na(. on Hwy. 17tS, Croflt. .X9749 (AAU, MEXU, MO, 

far out of the range of the sl)ecies. I)iei/enb(lchi(l IASA, SKIJX TE



ever, this is a distinct species from Antioquia De- Dieffenbachia picta subvar. angustifolia Engl., Bot. 

partment, Colombia. 

Jahrb. Syst. 26: 569. 1899. TYPE: based on a 

Dieifenbachia picta Schott forma lancifolia (Lin- cultivated plant of unknown origin, not seen. 

den & Andre) Engl. and Dieffenbachia picta Schott No type listed by Engler. 

subvar. Iancifolia (Linden & Andre) Engl. are com- Dieffenbachia seguine var. minor Engl., Bot. Jahrb. 

binations of Dieffenbachia lancifolia Linden & An- Syst. 26: 567. 1899. TYPE: No type listed by 

dre that also must be excluded from consideration Engler. 

in this revision of Central American Dieifenbachia. Dieffenbachia picta var. Iatior Engl., Bot. Jahrb. 

Dieffenbachia meleagris Linden & Rodigas, Syst. 26: 569. 1899. TYPE: No type listed by 

,olaced in synonymy of D. seguine by Engler (1915); Engler. 

however, this is a distinct species from Ecuador, Dieffenbachia picta var. Iatior Engl., Bot. Jahrb. 26. 

possibly one synonymous with D. spruceana Schott. 569. 1899. TYPE: Not seen. No type listed by 

The recombine(l Dieffenbachia meleagris Linden & Engler. 

Ro(ligas subvar. meleagris is also to be excluded. Dieffenbachia picta subvar. memoria Engl., Bot. 

Dieffenbachia shuttleworthiana Engl. was syn- Jahrb. Syst. 26: 570. 1899. TYPE: based on 

onymized with D. picta by Engler (1915), but it living material at Berlin Botanical Garden 

must be excluded along with the combination (also Buitenzorg), not seen. No type listed by 

D. picta Schott forma schuttleworthiana (Regel) Engler. 

Engl. and D. shllttleworthiana Hort. Bull. This Col- Dieffenbachia /)icta subvar. mirabilis Engl., Bot. 

ombian species is not closely related to D. seguine. Jahrb. Syst. 26: 570. 1899. DieJ%enbachia mirabilis 

Verseh. ex Engl., in DC., Monogr. Phan. 

SPEC1ES INCERTAE SEDIS 

2: 448. 12J78. TYPE: based on a cultivated 

plant of unknown origin by Verschaffelt, not 

The following species names are believed to be seen. No type listed by Engler. 

synonyms of Dief/enbachia seguine based on the re- Dieffenbachi(l l)icta var. angustior Engl., Bot. Jahrb. 

vision of Dieffenb(l chia by Engler (1915); however, Syst. 26. 569. 1899. TYPE: No type or locality 

owing to the (lestruction of many herbarium spec- listed, not seen. No type listed by Engler. 

imens during World War II, lectotypification was Dieffenbachia seguine fo. viridis Engl., F1. Bras. 

impossible. The specimens were no doubt available 3(2): 174. l 878. Dieffenbachia seguine subvar. 

during Engler's time, and it is assumed that he saw viridis Engl., Bot. Jahrb. Syst. 26: 567. 1899. 

the material. 

Dieffen6(lcXlia seguine var. viridis Engl., Pflanzenr. 

IV. 2


2004 

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manta Massif, Gran Sabana, Venezuela. Bot. Soc. Venez. publica Mexico. 1:200,000. Secretaria de Recursos 

Ci. Nat. 25(106): 29-33. 

Hidraulicos, Mexico. 

. 1965. Commentary on Mexican Araceae. Gentes French, J. C. 1997. Vegetative anatomy. Pp. 9-24 in S. J. 

Herb. 9: 289-382. 

Mayo, J. Bogner & P. C. Boyce, The Genera of Araceae. 

. 1979. Una sinopsis de las Araceae de Venezuela. Royal Botanic Gardens, Kew. 

Revista Fac. Agric. (Maracay) 10: 139-290. 

, M. Chung & Y. Hur. 1995. Chloroplast DNA phy- 

. 1988. Notes on a Guayana Dieffenbachia (Ara- logeny of Ariflorae Pp. 255-275 in P. J. Rudall, P. J. 

ceae). Phytologia 65: 390. 

Cribb, D. F. Cutler & C. J. Humphries (editors), Mono- 

Croat, T. B. 1978. Flora of Barro Colorado Island (Panama cotyledons: Systematics and Evolution. Royal Botanic 

Canal Zone). Stanford Univ. Press, Stanford. 

Gardens, Kew. 

. 1979. The distribution of Araceae. Pp. 291-308 Gentry, A. H. 1982. Evidence for phytogeographic patin 

K. Larsen & L. B. Holm-Nielsen (editors), Tropical terns as evidence for a Choco refuge. Pp. 112-136 in 

Botany. Academic Press, London. 

G. T. Prance (editor), Biological Diversification in the 

. 1981 [1982]. A revision of Syngonium (Araceae). Tropics. Columbia Univ. Press, New York. 

Ann. Missouri Bot. Gard. 68: 565-651. 

Gleason, H. A. 1929. Studies on the flora of northern 

. 1983a. A revision of Anthurium (Araceae) for South America XI. New and noteworthy monocotyledons 

Mexico and Central America. Part 1. Mexico and Mid- for British Guiana. Bull. Torrey Bot. Club 56: 8-14. 

dle America. Ann. Missouri Bot. Gard. 70: 211-240. Grayum, M. H. 1984. Palynology and Phenology of the 

. 1983b. Dieffenbachia. Pp. 234-236 in D. N. Jan- Araceae. Ph.D. Dissertation, University of Massachuzen 

(editor), Costa Rican Natural History. Univ. Chicago setts, Amherst. 

Press, Chicago. 

. 1990. Evolution and phylogeny of Araceae. Ann. 

. 1985. The large monocots of Panama. In W. G. Missouri Bot. Gard. 77: 628-677. 

D'Arcy & M. D. Correa A. (editors), The Botany and . 1991. Systematic embryology of Araceae. Bot. 

Natural History of Panama: La Botanica e Historia Nat- Rev. (Lancaster) 57: 167-203. 

ural de Panama. Monogr. Syst Bot. Missouri Bot. Gard. . 1992. Comparative External Pollen Ultrastruc- 

20: 5-12. 

ture of the Araceae and Putatively Related Taxa. Mon- 

. 1986a. A Revision of Anthurium (Araceae) of ogr. Syst. Bot. Missouri Bot. Gard. 43. 

Mexico and Central America. Part 2. Panama. Monogr. Hemsley, W. B. 1885. Biologia Centrali-Americana 3: Part 

Syst. Bot. Missouri Bot. Gard. 14: 1-205. 

18. Plates 97-100. R. H. Porter and Dulan, I,ondon. 

. 1986b. The distribution of Anthurium (Araceae) 

Henny, R. J. 1988. Ornamental aroids: (3ulture ancl breedin 

Mexico, Middle America and Panama. Selbyana 9: 

ing. Pp. l-33 in J. Janick (editor), Horticultural Re- 

94-99. 

views, Vol. X. Timber Presi,, Portland. 

. l988 1l99()]. Fgcology and lifeforms of Araceae. 

Holelridge, L. R. l967. IJife Zc)ne licology. l'rol)i(al Sci- 

Aloideana ll(3): 4-55. 

. 199() 1 l 992l. A ( omparison of aroid ( lassifi( ation 

ence (2enter, San Jose, (3osta ltie a. 

systems. Aroideana 13: 44-64. 

, W. C. (Jrenke, W. H. Hatheway, '1'. lziang & J. A. 

. l992. Species divelsity of Araceae in Coloml)ia: 

rFosi Jr. 1971. 'orest linvironmenti, in Tropical Zorlees. 

A preliminary survey. Ann. Missouri Bot. Gard. 79: 17- 

Pergatnon Ivress, Oxford. 

28. 

Holmes, J. At. l969. On the absolute fall of sea-leveel dur- 

1997. A revision of Philodetldron sulvgenus Phil- ing the (Juaternary. Palaeogeogr. Paleoc limatol. Paodendron 

(Araceae) for Mexicc) and Central Amerita. laeoecol. 6: 237-239. 

Ann. Missouri Bot. Gard. 84: 311-704. 

Jacquin, N. J. 1763. Selettarum Stirpium Americanarum 

& G. S. Bunting. 1979. Standardization of An- Historia. Ex officina Kransiana, Vienna. 

thurium dese riptions . Aroideana 2: 15-25. 

Jonker-Verhoef, A. M. E. & F. P. Jonker. 1966. Notes on 

& N. Lambert. 1987. The Araceae of Venezuela. the Araceae of Suriname III. Acta Bot. Neerl. 15: 130- 

Aroideana 9: 3-214. 

146. 

& D. Mount. 1988. Araceae. Pp. 7-46 in R. Spi- Keating, R. C. 2002. Vol. 10. Araceae. In D. F. Cutler & 

chiger & J. M. Mascherpa (editors), Flora del Paraguay. M. Gregory (editors), Anatomy of the Monocotyledons. 

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tionship to a revised classification of the genera of Ar- 

Cullen, J. 1978. A preliminary survey of ptyxis (vernation) ac.eae. Ann. Missouri Bot. Gard. 9]: 485-494. 

in the angiosperms. Notes Roy. Bot. Gard. Edinburgh . 2004b. Systematic occurrence of raphide crystals 

37: 161-214. 

in Araceae. Ann. Missouri Bot. Gard. 91: 495-504. 

Engler, A. 1878. Araceae. Pp. 105-107 in Martius (edi- Linnaeus, C. 1763. Araceae. Pp. 1367-1374 in Species 

tor), Flora Brasiliensis 3(2). Vienna. 

Plantarum, 2nd ed. Stockholm. 

. 1879. Araceae. Pp. 2-681 in A. & C. DeCandolle Matuda, E. 1954. Las Araceas mexicanas. Inst. Biol. Mex- 

(editors), Monographie Phanerogamarum 2. Paris. ico 25: 176. 

. 1899. Beitrage zur Kenntnis der Araceae. 17. Mayo, S. J., J. Bogner & P. C. Boyce. 1997. The Genera 

Revision der Gattung Dieffenbachia. Bot. Jahrb. Syst. of Araceae. Royal Botanic Gardens, Kew. 

26: 564-572. 

, & . 1998. Pp. 26-73 in K. Ku- 

. 1915. Anubiadae, Aglaonemeteae, Dieffenba- bitzki (editor), The Families and Genera of Vascular 

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Pp. 1-78 in A. Engler (editor), Das Pflanzenreich IV. (except Orchidaceae). Springer, Berlin. 

23 Dc(Heft 64). 

Petersen, G. 1989. Cytology, systematics and chromosome 

Flores, G., L. Jimenez, X. Madrigal, R. Moncayo & F. numbers of Araceae. Nordic J. Bot. 9: 119-166. 

Takaaki. 1971. Mapa de Tipos de Vegetacion de la Re- Poeppig, E. F. 1845. Araceae. Pp. 83-91 in E. F. Poeppig 

767



. 

& S. E. Endlicher, Nov. Gen. & Spec. F. Hoffmeister, 6. D. crebripistillata Croat 

L. 

elpzlg. 

7. D. davidsei Croat & Grayum 

Raven, P. H. & D. I. Axelrod. 1974. Angiosperm bioge- 8. D. fortunensis Croat 

ography and past continental movements. Ann. Missouri 9. D. fosteri Croat 

Bot. Gard. 61: 539-673. 

10. D. galdamesiae Croat 

Ray, T. S. 1981. Growth and Heterophylly in an Herba- 11. D. grayumiana Croat 

ceous Vine, Syngonium (Araceae). Ph.D. Thesis, Har- 12. D. hammelii Croat & Grayum 

vard University, Cambridge. 

13. D. horichii Croat & Grayum 

. l 987. Cyelic heterophylly in an herbaceous vine, 14. D. isthmia Croat 

Syngonium (Araceae). Amer. J. Bot. 74: 16-26. 15. D. killipii Croat 

Schatz, G. 199(). Chapter 7. Some aspects of pollination 16. D. Iongispatha Engl. & K. Krause 

biology in Cerltral American forests. Pp. 69-84 in K. 17. D. Iutheri Croat 

S. Bawa & M. Hadley (editors), Reproductive Ecology 18. D. nitidipetiolata Croat 

of Troy)i(.al 11oreest Plants. Parthenon, Park Ridge, New 19. D. obscurinervia Croat 

Jersey. 

20. D. oerstedii Schott 

Schott, H. NV. 12929. Fur Liebhaber der Botanik. Wiener 21. D. panamensis Croat 

%. Kunst 1829(3): 803. 

22. D. pittieri Engl. 

. 186(). Ilodromus Systematis Aroidearum. Typis 23. D. seguine (Jacq.) Schott 

Congregationis Mechitharisticae, Vienna. 

24. D. standleyi Croat 

Standley, P. C. 19.-s1. Araceae. Arum Family. In Flora of 25. D. tonduzii Croat & Grayum 

the Lanceti]la Valley, Honduras. Publ. Field Mus. Nat. 26. D. wendlandii Schott 

Hist., 130t. Seel . 1(): 118-124. 

. 1937. Flora of Costa Rica, Part 1. Publ. Field 

Mus. Nat. Hist., Bot. Ser. 18: 131-146. 

1944. A l ae eae. In R. E. Woodson, Flora of Pan- 

APPENDIX 2 

tNDEX TO EXstcCATAE. TYI'F, sICtEs tN BoLDFACE 

ama. Ann. Missouri Bot. Gard. 31: 1-60. 

Acevedo-Rodriguez 47()5 (2.-3); Acosta Solis, M. 10896 

& J. Ste.ye.rmark. 1958. Araceae. Pp. 304-363 in (15); Aguilar, R. 256 (20), 21'35 (4), 2380 (20), 2412 (20), 

Flora of Gu at e llsal a. Fieldiana, Bot. 304. 

5204 (20); Aguilar et al 43()2 (20); Akers, M. 78 (18), 

Takhtajan, A. lt3()9. Araceae. Pp. 8, 108, 199, 239, 247, 78A (11); Allen 5669 (4), .5()()9A (4), 5972 (1); Allen 8z 

251 in Flowelillg lvlants, Origin and Dispersal. [Trans- Alston 1839 (6); Alvarez, A. & P. Herrera 699 (15); A1lated 

flom lAtlssiarl text by C. Jeffrey, Kew.] Smithsonian verson et al. 283 (19), 376) (15); Anaya 1 (20); Andre 1202 

Institution, Wasllington, D.C. 

(23), 457 (15); Antonio 2t3()4 (25), 3039 (6), 3820 (6), 

Tam, S.-M., P. (:. 13oyce, T. M. Upson, D. Barabie, A. Brun- 4488 (18), 4546 (14), 46).sEJ (1

(15) 1 266() ( l 9) 1 34() 3 (6), l 32J49 ( 1 ()) l l . 5 (6) 1g1t3()() 


2004 

Croat 


Bunting & Trujillo 2236 (23); Bunting et al. 1937A (23), 

1949A (23), 1997 (23), 2072 (23), 2074 (23), 2073 (23), 

(10),49155 (6),49243 (6),49310 (25),49761 (18),49768 

(18), 49791 (19), 49805 (15), 49932 (25), 49971 (8), 

4108 (23); Burch et al. 1123 (15); Burger & Antonio 50049 (25), 50480 (15), 50655 (15), 50684 (15), 50725 

10905 (18), 11198 (18), 11224 (12), 11249 (4); Burger & (15), 52300 (25), 5378 (14), 53921 (23), 53956 (23), 

Baker 9968 (11), 10121 (3), 10137 (13); Burger & Gentry 54380 (23), 54649 (23), 55549 (15), 55629 (15), 55666 

Jr. 8898 (3); Burger & I iesner 7111 (20), 7196 (4), 7254 (15), 56070 (18), 56112 (25), 56205 (25), 56282 (25), 

(3); Burger & Matta 4181 (11); Burger & Pohl 7825 (20); 56625 (25) ? 57000 (25), 57427 (25), 57462 (18), 59152 

Burger & Stolze 5024 (18), 5461 (4), 5489 (3), 5753 (18), (2), 60567 (23), 60613 (23), 60867 (23), 62831 (25), 

5754 (2); Burger et al. 1323 (20), 10323 (18), 10669 (13), 66533 (25),66732 (25),66866 (8),66953 (2),66993 (25), 

10671 (3), 10699 (18), 10734 (11), 11695 (20), 11926 67115 (6),67197 (15),67329 (19),67379A (18),67399A 

(20); Busey 743 (1). 

(16), 67480 (15), 67526 (21), 67533 (20), 67576 (6), 

Cadet 6030 (23); Calderoll 914 (26); Callejas & Jan- 67594 (4), 67692 (3), 67700 (1), 67837 (8), 68012 (8), 

goux 2692 (25); Callejas et al. 4873 (19), 5661 (18), 9704 68069 (20), 68()97 (25), 68195 (18), 68196 (25), 68336 

(25); Calzada 338 (20); Camp E-3599 (15), 3653 (15); (25), 68358 (2), 68359 (11), 68379 (18), 68449 (20), 

Carballo et al. 47 (20); Carleton 508 (20); Carlson 153 68494 (23),68547A (23),68582 (23),68679 (15),68699 

(20); Carrasquilla 2005 (18); Carrasquilla & Mendoza (6), 68746 (6), 68820 (21), 68854 (18), 68961 (15), 

1239 (18); Carrasquilla & Rincon 304 (15); Carvajal 273 68971 (16), 69070 (17), 69343 (15), 70099 (25), 70868 

(20); Catharino et al. 1909 (23); Cedillo, T. 3645 (20); (25), 70900 (25), 71017 (25), 71119 (15), 71461 (25), 

Ceron & J. Corozo 33858 (18), 33947 (18), 34097 (18); 72573 (15), 73792 (15), 73826 (15), 74137 (23), 74759 

Ceron et al. 29159 (15); Chacon 364 (20), 507 (11), 553 (15), 74760 (6), 74789 (15), 74792 (15), 74853 (20), 

(12), 1406 (13); Chacon et al. 1488 (25); Chavarria & 74861 (6), 74930 (25), 74945 (11), 74952 (25), 74953 

Umana 157 (13); Chavarrfa et al. 257 (4); Chazaro 416 (11), 74958 (18), 75007 (17), 75099 (20), 75100 (7), 

(20); Chinchilla 137 (20); Churchill & Churchill 6105 75149 (19), 75154 (15), 75172 (6), 75195 (16), 77281 

(18), 6158 (8), 6159 (8), 6252 (25); Churchill & de Nevers (4),77283 (20),77298 (23),78287 (23),78318 (4),78323 

4333 (15); Churchill et al. 3993 (6), 4032 (15), 4125 (15); (23), 78345 (23), 78480 (26), 78678 (20), 78687 (20), 

Clarke 60 (20); Clewell & Tyson 3306 (15); Clezio 221 78695 (20), 78711 (26), 78720 (26), 78731 (12), 78732 

(15); Cogollo et al. 3462 (25); Colehester 21()8 (23); Con- (4),78733 (11),78758 (12),78771 (2),78784 (20),78787 

rad sT Conrad 2882 (20); Conrad et al. 2863 (20); Cornejo (25), 78788 (18), 788



5727 (18), 7407 (18), 7409 (19), 8267 (15); Delinks 452 

(15); Dillon et al. 1837 (20); Dodge 10020 (4), 10198 (4); 

Dodge & Goerger 9489 (25), 10157 (20); Dodson 6188 

(15); Dodson & Dodson 11130 (15); Dodson & Tan 5338 

(15); Dodson et al. 8415 (15), 10594 (15), 14638 (15); 

Hahn 142 (15), 945 (23); Hammel 122 (2)! TT8 (21), 3140 

(6), 3586 (20), 3781 (6), 4049 (18), 4101 (20), 4298 (14), 

4484 (19), 5378 (14), 5607 (7), 6264 (20), 7212 (6), 8123 

(18), 8167 (11), 8212 (12), 8270 (20), 8273 (12), 8415 

(20), 8606 (20), 8617 (11), 8748 (12), 8784 (4), 8846 (2), 

Donnell Smith 7813 (20); Drake 7 (23); Dressler 4688 8873 (11), 8874 (25), 9688 (4), 9772 (4), 9922 (20), 

(15), 4716 (14), 5316 (10); Dryer 1681 (11); Duke 5014 

(14), 5099 (15), 5169 (15), 5260 (15), 5437 (15), 13082 

(15), 13601 (15), 14346 (15), 15591 (15); Duke & Kirk- 

10081 (4), 10082 (18), 10449 (20), 12320 (12), 13636 

(5), 13685 (7), 20231 (20); Hammel & D'Arcy 5018 (15); 

Hammel & Grayum 14169 (1); Hammel & Kernan 16661 

bride 14019 (15); Duss, P. 3790 (23), 21496 (23); Dwyer 

2077 (15), 4565 (15), 9920 (20), 11184 (20); Dwyer & 

Dieckman 13008 (20); Dwyer & Gentry 9479 (6). 

Eggers 14183 (15), 15095 (15); Ekman 1983 (23); Engler 

226 (23), 227 (23), 2389 (26), 2793 (23); Espina et 

(4); Hammel & Trainer 13962 (20), 14211 (20), 14767 

(14), 14786 (19), 15836 (25); Hammel et al. 4882 (18), 

6871 (20), 14472 (15), 16262 (14), 16397 (7), 17548 (25), 

20114 (20); Harling 313 (15); Harling & Andersson 

16727 (15), 18898 (15), 19371 (15), 24778 (15); Harmon 

al. 2902 (15); Espinosa & Guerra 3762 (15), 3939 (15); 

Espinosa & E. Martinez 3308 (6); Espinosa et al. 3138 

& Dwyer 4035 (20); Harmon & Fuentes 4740 (20), 6419 

(20); Harris 8361 (23); Harris & Neal s.n. (23); Hartman 

(l5), 3611 (15), 4478 (15), 4723 (6), 10035 (20). 

Faden et al. 76/82 (20); Fallen & Ray 860 (15); Fishlo( 

k 326 (23); Folsom 1265 (6), 1444 (15), 3211 (6), 4054 

(20), 6218 (6), 6247 (15), 8712 (20), 9200 (18), 9329 (12), 

9724 (11), 10116 (25); Folsom & Collins 436 (15), 6486 

(21); Folsom & Mauseth 7844 (15); Folsom et al. 2111 

(20), 6848 (15), 6852 (15), 8264 (21); Fonnegra 1344 

(25); Fonnegra et al. 2899 (18), 2907 (18); Forero 719 

(25); Forero et al. 1723 (14), 4571 (18), 4659 (18), 6272 

(15), 6489 (15), 9007 (15), 9047 (15); Forther 10213 (20); 

Foster, R. B. 865 (14); Foster et al. 14649 (9); Fuentes 

R. 12488 (15); Henny 5 (2), 7 (23); Herbst & S. Ishikawa 

5459 (23); Heredia, M. 60 (23); Helnandez 473 (26), 542 

(20), 605 (20), 1318 (26), 2586 (26); Hernandez & Vazquez 

551A (20); Herrera 1636 (20)! 4116 (18); Herrera, 

G. et al. 2928 (20); Herrera, H. 245 (19); Heyde & Lux 

4654 (20); Higgins, J. 119 (23); Hill, S. 24812 (23); 

Hitchcock s.n. (23); Hodge 2929 (23); Hodge & Hodge 

3174 (23); Holm-Nielsen et al. 2769 (15), 25355 (18); 

Holst 62370 (23); Holstein & Allbruster 20425 (20); 

Hoover 1324 (15); Hoover et al. :3161 (25), 3968 (25); 

Horich s.n. (11), s.n. (13); Howar(l 11564 (23), 11744 

367 (20); Fuertes 564 (23); Funk et al. 10626 (25), 10718 

(25). 

Galdames et al. 1138 (18), 1330 (7), 1570 (7), 1626 

(23); Howard & Nevling 16978 (2:3); Huft & Jacobs 1997 

(14). 

Ibarra 167 (20), 455 (20), 645 (2()); Ibanez et al. 1829 

(14), 2252 (14), 2286 (14), 2486 (14); Galeano, G. 4600 

(25); Galeano, G. et al. 4825 (19); Gamboa et al. 71 (15), 

91 (15); Garibaldi 68 (14); Garnier 772 (26); Gentry 3035 

(26); Ibarra & Cedillo 1804 (2()); lltis et al. 30338 (20); 

INBio 186 (20); Ingram 1124 (15)! l 146 (17), 1169 (23), 

76452 (25). 

(25), 26705 (25); Gentry & Dwyer 3643 (15); Gentry & 

Forero 7217 (18), 7342 (18); Gentry & Juncosa 41037 

Jacobs 2159 (20); Jimenez 10:3 (18). VI (18); Johnston, 

I. 1165 (14); Johnston, J. R. 305 (23). Judd et al. s.n. 

(18); Gentry & Lajones 73108 (15); Gentry & Tyson 1653 

(14); Gentry et al. 3406 (15), 24700 (25), 26709 (15), 

28574 (14), 72496 (15), 72955 (25), 75903 (25); Gomez 

19038 (20), 19531 (4), 19532 (20), 19572 (26), 20562A 

(20), 20565 (20), 22016 (20), 22951 (13); G6mez & Herrera 

23477 (25); Gomez et al. 20405 (25), 21108 (20); 

G6mez-Laurito 7792 (13), 7856 (4); G6mez-Pompa 1505 

(20); GonSalves et al. 224 (23); Gonzalez 1473 (20), 671I 

(23); Juncosa 619 (25), 797 (15). 1795 (18), 1898 (18), 

1912 (18); Jones, A. & Tejada 275 (15). 

Kennedy 455 (15), 1193 (14)! 1.594 (15), 2661 (15), 

3253 (25); Kennedy & Foster 395 (15); Kennedy & Solomon 

4629 (20); Kenoyer 188 (14); Kernan 381 (20), 748 

(4); Kernan & Phillips 831 (4); KeBw 70-76-494 (20); Killip 

12154 (15), 35113 (18), 39979 (l4); Knapp 1042 (19), 

2165 (4), 2275 (15), 5758 (6); Kllap^) & Mallet 3089 (15); 

(20), 3341 (20), 5597 (20); Gordon 55C (18), 339 (25); Knapp et al. 1717 (6); Koshear 59 (4); Kress T7-830 (15), 

Grant & Rundell 92-01928 (4); Grayum 2225 (18), 2288 

(11), 2771 (20), 2772 (12), 2780 (25), 2840 (4), 4756 (13), 

T7-831 (15), 84-630 (25), 84-1622 (20), 84-1632 (11); 

Kufer, J. 394 (20); Kursar & Coley 4 (15); Kvist & Asanza 

4765 (20), 6194 (20), 6887 (11), 6899 (2), 6918 (20), 40756 (18); Kvist et al. 48348 (18). 

6925 (20), 6936 (2), 7620 (12), 7698 (2), 8638 (13), Lasser 16678 (15); Lavastre 1845 (93); Lazor & Tyson 

9251 (20), 9277 (20), 9773 (11), 9777 (4), 9804 (25), 3492 (15); Lazor et al. 2578 (18); l eClezio 135a (16); Le 

9830 (11), 9844 (18), 10588 (4), 10858 (25); Grayum & Goff A. 95 (23); Lehmann s.n. (15), 5311 (15), 8876 (25); 

Evans 10156 (4); Grayum & Fleming 8119 (4), Grayum Leija & Garza 3341 (20), 5597 (20); Leimbeck, R. 306 

& Hammel 5785 (18); Grayum & G. Herrera 7829 (20), (15); Lent 37 (18), 161 (1), 639 (25) 694 (25), 2789 (20), 

9139 (1), 9236 (3); Grayum & Jacobs 3524 (18), Grayum 4042 (25); Lent et al. 3374 (18), Leon 720 (25), 26573 

& Murakami 9939 (25); Grayum & Schatz 3174 (25), (25); Lewis et al. 1753 (6), 2195 (14), 3251 (14); Liesner 

3206 (11), 3218 (18), 3220 (2), 5279 (25); Grayum & 114 (1), 1736 (3), 2871 (4), 14102 (20), 14394 (25); Lies- 

Sleeper 6100 (20); Grayum & Warner 5710 (20); Grayum ner & A. Gonzalez 10126 (23); Liesner & Judziewicz 

et al. 3982 (4), 3983 (4), 4014 (3), 4440 (7), 4447 (4), 14797 (25); Liesner & Mejia 26236 (20); Liesner et al. 

4483 (7), 4961 (20), 5467 (13), 5719 (20), 5723 (13), 7681 (23), 15035 (20), 15122 (20), 15144 (18), 15330 

5862 (20), 7547 (3), 7549 (1), 7567 (4), 7657 (15), 8038 (25), 15365 (25); Liogier 13307 (23); Liogier & Liogier 

(11), 8336 (13), 8345 (20), 9250 (4), 9469 (25), 9744 (12), 20185 (23); Lister & Colchester 272 (23); Lloyd 237 (23); 

9962 (4), 9973 (23), 10578 (20), 11116 (11), 11139 (4), Loiselle 106 (25); (20); L0jtnant & Molau 15839 (15); 

11163 (11), 11169 (12), 11174 (13); Grove 01 (20); Guer- Lopez Garcia & Martin 122 (26); Lotto s.n. (23); Luteyn 

ra & Liesner 2871 (4); Guillermo, J. & D. Cardenas L. 1010 (6), 1203 (15), 3175 (15), 3180 (15), 3188 (6), 3263 

863 (16); Gutierrez, G. & Barkley 17115 (25). 

(20), 3342 (20), 3385 (18), 4043 (14); Luteyn & Croat 

Haber & Atwood 9163 (20); Haber & Bello 7191 (20); 

Haber & Hammel 1799 (20); Haber & Zuchowski 9251 

906 (14); Luteyn & Kennedy 1612 (6), 1837 (6); Luteyn 

& Wilbur 4569 (20); Luther s.n. (23). 

(20), 11175 (25); Haber et al. 4979 (20), 10824 (25); Maas et al. 7834 (1); MacDougal 1027 (18), 1090 (20),


2004 

Croat 


3193 (20), 3299 (20), 3303 (24); Madison 589 (20), 627 

(20), 705 (24), 712 (20); Madison et al. 5008 (15); Machilla 

352 (20); Sargent 340 (23), 643 (23); Schatz 1079 

(15); Schatz & Grayum 700 (20), 701 (20), 706 (25); 

guire et al. 36461 (23); Marten 789 (4), 831 (20), 848 (3); 

Martinez 3016 (20); Martinez et al. 22713 (20), 23175 

Schipp 386 (20); Schmalzel 1212 (15); Schmalzel & A1verson 

1199 (15); Schubert & Rogerson 619 (20); Schultes 

(20), 23474 (20), 23693 (20); Martinez, E. 2287 (26), 

13445 (20), 13630 (20), 16131 (20); E. Martinez S. & 

& Cabrera 17890 (23); Schwerdtfeger 21422 (25); Seler 

2389 (26), 2398 (26); Shank & Molina 4288 (25); Shat- 

Aguilar 12435 (20); Matuda 16369 (26), 16765 (26); Max- tuck 397 (14); Shepherd 438 (19); Simmonds s.n. (23); 

on et al. 6812 (14), 6820 (14); Mayo & Madison 301 (20); Sinaca 830 (20); Sintenis, P. 2793 (23); Skutch 5328 (1); 

McAlpin 85-33 (4); McDade s.n. (16); McDonagh et al. 

433 (15), 439 (15); McDowell 148 (20), 769 (11), 1012 

Slane 634 (23); Smith 2239 (20); Smith, C. E., Jr. & H. 

M. Smith 3389 (6); Smith, D. A. 134 (20); Smith, D. et 

(20); McPherson 7371 (25), 9176 (15), 9829 (18), 9865 al. 1059 (25); Smith, H. H. & G. W. Smith 1411 (23) 

(25), 10725 (7), 10958 (14), 11129 (25), 11381 (18), Soejarto & Renteria 3556 (19); Solomon 5761 (23); Sparre 

11401 (18), 11591 (14), 11816 (2), 15037 (14); McPher- 14122 (15), 14556 (15), 15182 (15), 15499 (15), 18326 

son & J. Aranda 10095 (2); McPherson & Merello 8143 (18), 19397 (15), 19488 (15); Sparrow & Brewster 108 

(19), 8235 (18); Meier et al. 5164 (23); Mejia & Ramirez (20); Spellman et al. 164 (20); Sperry 517 (20), 525 (20), 

9929 (23); Mejia & Zanoni 6614 (23); Mena 190 (20); 567 (20), 570 (20); Standley TT00 (20), 7935 (20), 18786 

Mendieta 1-10 (15), 1-101 (15), 1-121 (15); Miller & (24), 20085 (20), 21077 (26), 21414 (20), 25700 (24), 

Schmidt 5551 (23); Miller & Taylor 5937 (23); Miller et 26156 (16), 26305 (20), 27224 (15), 27413 (15), 28238 

al. 753 (15); Miranda, F. 7546 (20); Miyashiro s.n. (23); (16), 28732 (16), 29867 (14), 31266 (14), 32242 (20), 

Montenegro, D. & Chung 1462 (20); Montes s.n. (23); 32794 (20), 36314 (25), 36739 (4), 36840 (2), 38942 (20), 

Moore, J. W. 451 (23); Moore Jr. & Bunting 8928 (20), 40223 (20), 40960 (14), 41107 (14), 44754 (20), 52702 

8933 (20); Mora 51 (25), 70 (18); Moraga 173 (20); Mo- (20), 52924 (20), 53146 (24), 53990 (20), 55444 (20), 

rales 1047 (26), 2035 (13); Morales et al. 5413 (20); Mo- 58291 (20), 60715 (20), 63529 (20), 63620 (20), 65041 

reno 15290 (26), 16421 (26), 16464 (26), 17142 (24); (20), 66844 (20), 68724 (26), 72447 (20), 75669 (20), 

Moreno & Henrich 8427 (26); Moreno & Robleto 20526 78161(26), 78523 (20), 79561 (26), 87180 (20), 88988 

(24); Moreno & Sandino 12855 (25), 12891 (25), 12917 (20); Standley & Chac6n 6701 (24); Standley & Valerio 

(25), 12955 (25), 12976 (18), 15160 (18); Mori & Bolten 45008 (20), 45206 (25), 45262 (25), 45592 (20), 46000 

7698 (15); Mori & Gracie 18717 (23); Mori & Kallunki (20), 48960 (4), 48997 (25); Stein & Hamilton 990 (6); 

3591 (15), 6026 (6); Mori et al. 3817 (25), 4184 (7), 6814 

(25); Murphy & Jacobs 1289 (20); Moreno, P. & J. Sandino 

15160 (18). 

Nash, G. 611 (23); Nee 9023 (16), 22595 (20), 23733 

(20), 29752 (2()), 29993 (20)! 41364 (23), 99121 (20); Nee 

et al. 24759 2())! 26103 (20); Neill 1571 (24), 1657 (26), 

3629 (24); Neill & Vin(elli 3484 (12); Neill et a1. 11683 

(15); Nelson & Andino 16247 (24); Nelson & Cruz 9215 

Stergios et al. 7940 (23); Stern et al. 433 (15); Stevens 

6027 (26), 6420 (25), 7457 (24), 8044 (24), 11786 (24), 

12311 (24), 13564 (25), 13761 (20). 23642 (11)! 24257 

(12); Steverls et al. 17591 (26)! 20998 (24), 24699 (12); 

Stevensone P. 379 (16); Steyerlnark 31763 (20)! 34555 

(20). 37153 (20), 37456 (20), 38647 (20), 38775 (20), 

41699 (2()), 44754 (20), 4541() (20)! 45869 (20), 47684 

(26), 47908 (20), 49321 (20), 49637 (26), 52070 (20), 

(20), 9291 (24); Nevling & G6mez-Pampa 1505 (20); Ni(- 60796 (23), 88864 (23), 91896 (23), 94331 (23); Steyerolson 

2()6() (23), 3393 (4); Nilsson & Manfredi 505 (20); mark & Bunting 97728 (23), 105293 (23); Steyermark & 

Nilsson et a1. 377 (25)! 632 (25); Noriega & H. Vasquez Davidse 116647 (23); Fiteyermark & Liesner 118891 (23), 

G. 1353a (20). 

121834 (23); Steyermark & Rogers 119364 (23); Steyer- 

Ocampo 3411 (20); Oersted s.n. (20); Opler 246 (20); mark et al. 100213 (23), 100239 (23), 100318 (23), 

Oppenheimer 67-1-3-1244 (16). 

102036 (23), 122412 (23), 124629 (23), 124955 (23), 

Palacios 11452 (25), 13626 (25); Palacios & Tirado 127204 (23); Stimson 5277 (16); Stone 3317 (20); Stricker 

1287 (25), 11327 (15); Paredes et al. 940 (7); Peneis, D. 342 (20); Sturrock 352 (23); Sullivan 59 (15), 537 (6), 

633 (25); Perez 1 (25); Perez, J. & M. Sosa 671 (23); 

Peterson 6405 (16); Peterson & Annable 6768 (15); Pfeifer 

2124 (24), 2163 (20); Philcox 8231 (23); Picado & 

745 (15); Sytsma 1658 (15), 1695 (18); Sytsma e.t al. 2414 

(15), 4398 (6); Sytsma & Andersson 4573 (6); Sytsma & 

Antonio 3006 (25). 

Gamboa 134 (20), 138 (1); Pipoly 4479 (24); Pitman & 

Bass 995 (25); Pittier 2600 (15), 2715 (16), 2836 (20), 

Taylor 216 (20); Taylor, N. 48 (23); Taylor & Taylor 

11660 (18); Tellez et al. 4466 (20), 4875 (25); Thompson 

3754 (14), 3766 (22), 3838 (16), 3847 (19), 6845 128 (15), 160 (15), 441 (26), 3238 (23), 4593 (15), 4816 

(15); Pittier & Durand s.n. (1); Plowman 14121 (15); Po- (15), 4874 (6), 4937 (2), 4951 (8), 5028 (18); Thorne & 

lanco 1485 (15), 1591 (18); Polanco et al. 1905 (15); Por- Lathrop 40559 (20); Tipaz et al. 2280 (18); Tonduz 503 

ter et al. 4282 (15); Prance et al. 30247 (23); Prevost 3258 

(23), 3382 (23), 3580 (23); Proctor 20987 (23), 32251 

(25), 9961 (1), 12874 (25); Trujillo et al. 17397 (23); 

Tyson 1438 (16), 1443 (15), 4632 (14), 6700 (16); Tyson 

(20), 38601 (23). 

& Blum 3954 (15), 3997 (15), 3998 (14), 4099 (15); Tyson 

Quesada 51 (4), 175(20); Quishpe & Davila 82 (15). et al. 4486 (15), 4631 (14), 4701 (16), 4834 (14). 

Ramamoorthy et al. .3763 (20); Raven 21532 (4); Read 

& Watson 84-75 (21); Renson 266 (26); Renteria 10680 

Utley & Utley 1100 (3). 

Valerio s.n. (20), 246 (20), 462 (1), 1355 (20), 1356 

(15); Rios et al. 109 (20); Rivera 623 (20), 758 (20), 1560 (20); Valverde 741 (13); Vanderveen 590 (20); Vasquez, 

(20), 1718 (25); Robles, R. 815 (20), 1142 (25), 1158 M. et al. V-907 (20); Villacorta et al. 314 (20), 408 (20); 

(25), 1234 (2), 2090 (4); Robles et al. 2050 (25); Rodri- von Wedel 1438 (2), 2892 (18). 

guez, A. & Estrada 142 (20); Rodriguez, G. 32 (20); Rodriguez, 

J. 262 (15); Rojas et al. 607 (23); Roldan et al. 

Wagner, R. 558 (23); Webster et al. 12581 (24), 12624 

(20), 12625 (20); Wendland s.n. (26), 410 (26); White- 

1199 (18); Romero-Castaneda 6436 (16); Rose et al. 4375 foord 1176 (20), 3284 (20); Whitefoord & Eddy 136 (15), 

(23); Rueda et al. 4070 (4); Rueda & Coronado 6557 (24). 372 (25); Whitehill 8 (23); Wilbur 28249 (20), 37243 (20) 

Salick 8092 (12), 8153 (25); Sanchez 539 (2); Sanders 37337 (20); Wilbur & Jacobs 34819 (20), 34962 (20), 

et al. 19322 (20); Sandino 1248 (26); Sandoval & Chin- Wilbur et al. 15664 (6), 15861 (26); Williams, L. et al. 

771



28479 (13); Witherspoon & Witherspoon 8401 (15), Won 

s.n. (23); Woodbury et al. s.n. (23); Woodson Jr. & Schery 

861 (1); Woodson Jr. et al. 1909 (25). 

Yanez et al. 1387 (25); Yuncker 4961 (24); Yuncker et 

al. 8395 (20), 8551 (24)? 8826 (20). 

Zak et al. 5383 (15), 5414 (15), 5726 (15); Zambrano 

& Delgado 1336 (14); Zanoni & Pimentel 23420 (23) 

23456 (23); Zanoni et al. 23113 (23), 25159 (23), 27170 

(23); Zapata et al. 289 (19); Zent, E. & S. 2189 (23) 

Zuniga 216 (20).

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