(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Children's Library | Biodiversity Heritage Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "Contributions from the University of Michigan Herbarium."

CONTRIBUTIONS FROM THE 

UNIVERSITY OF MICHIGAN 

HERBARIUM 



UNIVERSITY OF MICHIGAN HERBARIUM 

ANN ARBOR, MICHIGAN 

20 OCTOBER 1997 



CONTRIBUTIONS FROM THE 

UNIVERSITY OF MICHIGAN 

HERBARIUM 



VOLUME 21 



UNIVERSITY OF MICHIGAN HERBARIUM 

ANN ARBOR, MICHIGAN 

20 OCTOBER 1997 



Copyright 1997 
5ity of Michigan Herbari 
All rights reserved 



Printed in the United Sta 



about the availability and prices of previous volume 
iddress the Director, University of Michigan Herbari 
lilding, Ann Arbor, Michigan 48109-1057, U.S.A. 

20 October 1997 



Revision of Pterandra (Malpighiaceae) Chi 

Excentradenia, a new genus of Malpighiaceae from South An 



neotropical Malpighic 
lit flora of Mt. Murud 



Mexic 
Miscellanc 


O ISMA 


EL Calzada J., Gabriel Flores F. & Oswaldo Tellez V. 1 
1 Sphagnum— iO Howard Crum 1 


Catalog of the Acanth 
phytogeographic r 


aceae of Belize with taxonomic and 

otes Thomas F. Daniel 1 


A revision 


and redefin 


tion of Pseudabutilon (Malvaceae) Paul A. Fryxell 1 


Newspeci 
f 


AUL A. Fry 


ecords for the Malvaceae of Baja California 

XELL, Jose Luis Leon de la Luz & Miguel Dominguez L. 1 



Colin Huc;he 
. S. Gonzalez-Elizondc 



and Leguminos 

OSWAL 


DO Tellez V. 


Mt. Murud, 

Jef 


™eW.Wooo 


riaceae 

Mich 


ahU.Wvnne 



REVISION OF PTERANDRA (MALPIGHIACEAE) 

Christiane Anderson 
University of Michigan Herbarium 

North University Building 
Ann Arbor, Michigan 48109-1057 

Pterandra comprises 14 species of trees and shrubs of Central and South 
America, which are readily recognized by the nature of the inflorescence as well 
as details of floral and vegetative morphology. Its closest relative appears to be 
Acmanthera Griseb., with which it shares winged anthers, a gynoecium of three 
essentially free carpels with subapically inserted styles, and a fruit composed of 
three subspherical dry cocci. The type species of Acmanthera was first described 
by Adrien de Jussieu (1838) in Pterandra though assigned to its own section, 
which was later elevated to generic level (Grisebach 1858). The two genera differ 
most strikingly in their inflorescences and stipules. In Pterandra the inflorescence 
consists of 2-6-flowered fascicles borne in the axils of bracts or leaves that are 
crowded at branch apices. The intra- and epipetiolar stipules are entirely to partly 
connate and less than 1 cm long. In Acmanthera the flowers are crowded on an 
elongate unbranched pseudoraceme terminating an axillary branch. The four 
stipules are fused into a deciduous sheath enclosing the stem and range from 1.5 
to 11 cm long. W. R. Anderson (1978), in his delineation of the subfamily Byrso- 
nimoideae, assigned Pterandra and Acmanthera, along with the monotypic genus 
Coleostachys Adr. Juss., to his tribe Acmanthereae. Currently, he and M. W. Chase 
are investigating the phylogeny of the Malpighiaceae by comparing evidence from 
chloroplast DNA and morphology, and preliminary results confirm this grouping, 
with Pterandra and Acmanthera as sister groups. The third genus of Malpighiaceae 
with winged anthers and a fruit composed of cocci is Lophanthera Adr. Juss. It is 
included in the Galphimieae (W. R. Anderson 1978), because its carpels are con- 
nate and have apical styles; it does not cluster with Pterandra and Acmanthera in 
the molecular analyses. For further comments about Acmanthera and Lophan- 
thera, see W. R. Anderson's accounts of these genera (1975, 1981, 1983). 

Only one species of Pterandra is well collected, P. pyroidea, a small shrub 
with clusters of showy pink flowers common throughout central Brazil; the equally 
large- and pink-flowered P. hatschbachii is known from only three collections, all 
from the type locality. The other species have inconspicuous and thus easily over- 
looked flowers and are known from one to few collections from localities only 
rarely visited or from several collections but mostly from the same area {P. arborea, 
P. ultramontana). As Fig. 1 shows, most species appear isolated from each other 
(only P. evansii and P. hirta are known to be at least partly sympatric), but future 
collections may fill the gaps. Little is known about habitats other than that most 
species were collected in forests. The exceptions are P. hatschbachii and P. pyroi- 
dea, which occur in the campos and cerrado of the Brazihan Planalto, and P. 
egleri, of campo in Para (Brazil); P. flavescens was said to be growing in savanna 
bordering a river. The habitat is unknown for P. andersonii. 

Compared to most other genera of Malpighiaceae, Pterandra is still only poorly 
known. Jussieu (1833) based the genus on Saint-Hilaire's collections of P. pyroi- 
dea from central Brazil; the second species, P. arborea, was not described until 



CONTR. UNIVERSITY OF MICHK 













^i:^_^«;^ 




-^ 


♦ arborea 


/o ^''^^""i^ 


1 




>v5 


^Yl 
> 




iA 




O guianensis 


° / 


/\ 




4!^ 


n hirsuta 

V isthmica 0° 


i'^ 


®® 


• pyroidea 


\ ' 


■ * 




• ultramontana 




X,o- 


'-■ 


^-' 'i 


.^«'^^ r 


U- 


^^,.j'i 


. \ 1 

/t" -20° 


80= 


70" 


50° 40° 



■\y collecting efforts in Central and South America launched si 
have greatly increased our knowledge of that flora and revet 
s. Since 1953, eight more species of Pterandra were recognized. 



INFRAGENERIC RELATIONSHIPS 



Akhough the genus is readily circumscribed, the species are separated by 
unique combinations of characters otherwise shared in the genus as well as by 
characters unique to particular species. Because there is so httle material avail- 
able for all species except P. pyroidea, it is very difficult to assess the degree of 
variation of a particular character within a species, and future collections may 
show that the present circumscriptions are too narrow. It is therefore with some 
hesitancy that the consensus cladogram presented here is proposed as an estima- 
tion of the relationships within Pterandra. 

The cladistic analysis was conducted using PAUP 3.1.1 (Swofford 1993). Acman- 
thera was designated as the outgroup for the 14 species of Pterandra and 12 
characters were used (Table 1). Too few mature fruits were available to include 
characters pertaining to the cocci, seeds, and embryos. A chromosome count has 
been reported only for P. egleri (W. R. Anderson 1993). 

1. Habit: = large shrubs to trees; 1 = small shrubs. Future collections may 
allow finer distinctions in habit. Pterandra andersonii is known from only two 
collections, both described as a low shrub, and thus clusters with P. hatschbachii 



C. ANDERSON: PTERANDRA 



and p. pyroidea, but il is possible that this species, Uke some others, varies from 
shrub to small tree. 

2. Attachment of hairs of the abaxial laminar vesture: = medifixed; 1 = 
basifixed. The common condition in Pterandra and Acmanthera is for these hairs 
to be medifixed. Pterandra hirsuta differs in that the abaxial vesture of the lamina 
is composed almost entirely of basifixed hairs. In P. evansii it consists of medi- 
fixed hairs except for some tufts of basifixed hairs in the axils of the secondary 
veins at the costa and scattered along the costa. Other aspects of abaxial vesture, 
except for density, are too little known to be included in the analysis. For exam- 
ple, in P. flavescens and P. ultramontana the hairs are ferrugineous; yet, the color 
fades to yellowish with age. Thus, it is possible that in other species these hairs are 
actually ferrugineous and that the whitish to yellowish color is a result of age or 
drying procedures. In P. egleri these hairs vary from golden to ferrugineous. The 
shape of the hairs also varies. In most species they are terete, but in P. ultramon- 
tana they are flat and scalelike. Some such flat hairs are intermixed with the terete 
hairs in the abaxial vesture of some leaves of P. flavescens, P. mcphersonii, and P. 
sericea. Pterandra egleri stands alone in that the vesture is tomentose and com- 
posed of wavy to crisped, subsessile to stalked hairs. 

3. Abaxial surface of laminas: = sparsely to moderately pubescent (the 
epidermis readily visible); 1 = glabrate to glabrous; 2 = densely pubescent (the 
epidermis mostly hidden). Only P. colombiana, P. mcphersonii, and P. hatsch- 
bachii are characterized by laminas nearly lacking abaxial laminar vesture; both 
Acmanthera and P. pyroidea were coded as polymorphic for this trait. Pterandra 
egleri was scored as moderately pubescent, but the tomentose vesture is sloughed 



4 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

off in age. Pterandra isthmica and P. sericea share abaxially densely sericeous 
laminas. More collections may reveal that the abaxial leaf vesture in at least some 
species is as variable as in P. pyroidea. 

4. Pellucid cells: = absent; 1 = present. All species of Pterandra have pellucid 
cells in the abaxial leaf surface, but Acmanthera lacks them. The species for which the 
most material is available, P. arborea and P. pyroidea, show that the presence of 
these pellucid cells varies greatly from abundant to a few along the margin. Thus, 
only the presence but not the disposition of these cells was scored for the analysis. 

5. Length of petiole of the larger leaves: = more than 0.5 cm long; 1 = up to 
0.5 cm long. The leaves of P. hatschbachii and P. pyroidea are noteworthy for the 
short petioles, even in the largest leaves not more than 0.5 cm long. In the two 
collections of P. andersonii the petioles of the largest leaves are also mostly very 
short though they rarely reach 1 cm. This species was scored as having short 
petioles, but additional collections may show that length of the petioles is variable. 

6. Fusion of stipules: = fused across the petiole, at a node resulting in two 
deltate to cordate structures; 1 = all four stipules fused into a sheath enclosing the 
node and part of the internode. Acmanthera is characterized by the second condi- 
tion, and Pterandra by the first. In Pterandra, the degree of fusion appears to be 
variable from entirely connate to nearly so to connate only in the basal 1/3 to 1/2; 
however, too little material is available to evaluate the constancy of degree of 
fusion within a species. All degrees of fusion occur in the two better-collected 
species, P. arborea and P. pyroidea. 

7. Abaxial stipular surface: = pubescent; 1 ^ glabrous. Only four species, P. 
arborea, P. colombiana, P. mcphersonii, and P. guianensis, have the stipules abax- 
ially glabrous even when very young. In Acmanthera and the remaining species of 
Pterandra, the stipular structures are abaxially pubescent, though the vesture is 
sometimes abraded in age. 

8. Inflorescence structure: = pseudoraceme; 1 = fascicles crowded along an 
axis. All species of Acmanthera have a pseudoraceme, a common type of inflores- 
cence in the family, but Pterandra is unique in having aggregates of fascicles in the 
axil of bracts, each composed of a pair of stipules and a rudimentary leaf, or, less 
commonly, in the axil of a young leaf. 

9. Attachment of flowers to axis: = sessile or at most subsessile; 1 = fascicles 
borne on a short projection up to 3 mm long. In Acmanthera and most species of 
Pterandra, the first condition is characteristic, but in P. arborea, P. guianensis, P. 
mcphersonii, and P. sericea at least some of the fascicles on an axis are elevated 
on a short projection. The degree of plasticity of this feature can only be assessed 
with the examination of more material. 

10. Number of flowers: = 3 or fewer per fascicle; 1 = 4-6 per fascicle. In most 
species of Pterandra the fascicles are composed of 4-6 flowers, but in P. flaves- 
cens, P. guianensis, and P. sericea of only 2-3 flowers. One fascicle of P. egleri 
appeared 3-flowered, which may indicate a broader range than 4-6 in that species. 

11. Color of petals: = white to cream; 1 = pink. Only P. hatschbachii and P. 
pyroidea have pink petals. In the other species, it is often noted on labels that the 
petals are yellow, a common condition in age. Petals are persistent even in fruit, 
and care must be taken to note the age of the flower when assessing petal color. 
On the label of one collection of P. andersonii the "flowers" are said to be pink, 
which, if true, would be another attribute shared with P. hatschbachii and P. 
pyroidea; however, that collection is in mature fruit, and thus the description of 
color was based on very old petals. 



PTERANDRA 5 

12. Size of limb of petal: = less than 6 mm in diameter; 1 = 6 mm or more in 
diameter. Pterandra hatschbachii and P. pyroidea also share large petal size in 
addition to petal color. The only other species with large petals is the otherwise 
dissimilar P. isthmica. 

The analysis yielded three most parsimonious trees of 22 steps each (CI = 
0.73; RI = 0.71; RC = 0.52). Figure 2 shows the trees and the strict consensus tree. 
As can be seen from the latter, the position of five species is particularly unstable. 
Pterandra evansii and P. hirsuta are similar in many characters, but P. egleri, P. 
flavescens, and P. ultramontana differ from these two as well as each other in a 
number of characters unique to each species. As noted above, future collection of 
most species may well result in different scoring of characters. For example, the 
position of P. andersonii would change if it were found that its habit is not re- 
stricted to "small shrub" and that large leaves vary in the length of the petiole. 
Yet, it seems likely that the two cerrado species, P. hatschbachii and P. pyroidea, 
as well as the pairs P. evansii/P. hirsuta, P. colombiana/P. mcphersonii, and P. 
isthmica/P. sericea will continue to be considered sister-species. Pterandra arborea 
and P. guianensis morphologically are more similar to each other than to other 
species; however, P. guianensis is known only from the type collection. 

Many more collections of all species from additional localities, except for P. 
pyroidea, should reveal more characters suitable for use in proposing a phylogeny. 
Unfortunately, given that most species of Pterandra are easily overlooked by 
casual collectors and are found in areas little visited, it is unlikely that such addi- 
tional material will be forthcoming in the near future. The results of the analysis 
presented here may be considered a first step in understanding the links between 
these fourteen species. 

TAXONOMY 

Pterandra Adr. Jussieu in Saint-Hilaire, Fl. bras, merid. 3: 72. 1833 ["1832"].— 
Lectotype, designated by Cuatrecasas, 1958: Pterandra pyroidea Adr. Juss. 

Shrublets, shrubs, or trees to 30 m. Stems with a pattern of a long internode 
followed by a series of very short internodes, branchlets densely sericeous when 
young, becoming glabrous. Leaves opposite; laminas elhptical or narrowly so to 
obovate or oblanceolate, apex obtuse (-mucronate) to acute (-mucronate) to acumi- 
nate to sometimes caudate, base cuneate or sometimes rounded or slightly trun- 
cate, adaxially pubescent when young but soon becoming glabrous or sometimes 
glabrate, abaxially sparsely to densely sericeous with medifixed hairs or glabrate 
to glabrous (tomentose in P. egleri, hirsute with basifixed hairs in P. hirsuta), 
eglandular but abaxially commonly sprinkled with pellucid cells or these concen- 
trated along the margin, costa and secondary veins prominent abaxially (in P. 
flavescens only the costa, in P. pyroidea and P. egleri the tertiary veins as well); 
petioles densely sericeous or sparsely so in age; stipules intra- and epipetiolar, 
each pair entirely connate or distally notched or only the proximal 1/4-3/4 con- 
nate and distally free, adaxially hirsute (becoming glabrous in P. sericea), abaxially 
glabrous or densely sericeous (sometimes sparsely so in age). Inflorescence a 
series of 2-6-flowered fascicles, these sessile or borne on very short projections on 
the new growth, usually in the axil of a deciduous bract composed of a pair of 
stipules and a rudimentary leaf in the stem regions with very short internodes (in 
P. flavescens and P. sericea usually in the axils of leaves), the fascicles thus crowded 



v'ERSITY OF MICHIGAN HERBARIl 



9(1) 


3{0&1) 


2(1) 


' L_ 



Acmanthera 

andersonii 

hatschbachii 

pyroidea 

arborea 



egleri 
evansii 
hirsuta 



I — andersonii 

^Toyijri— hatschbachii 





Consensus tree 



below the developing shoot apex or below a flush of new leaves; flowers borne on 
slender pedicels, each subtended by a pair of bracteoles and one bract, peduncles 
absent; calyx, corolla, and androecium persistent in fruit. Sepals 5, narrowly trian- 
gular but the distal 1/2-2/3 often ligulate, apex recurved to revolute during anthe- 
sis but erect in fruit, each sepal bearing two elliptical glands (glands sometimes 
irregularly present and/or irregular in size or sometimes entirely absent in P. 
pyroidea and P. sericea, each sepal sometimes bearing only one gland in P. egleri). 
Petals 5, clawed with the limb ± flat, the posterior one slighter larger than the 



1997 C. ANDERSON: PTERANDRA 7 

lateral four and with a slightly thicker claw, white or cream ("greenish yellow" in 
P. guianensis), becoming yellow in age, or pink, adaxially glabrous, abaxially sparsely 
to densely sericeous on the claw and center of limb (often glabrous in P. hatsch- 
bachii) or sometimes the abaxial pubescence nearly covering the entire limb ex- 
cept for a narrow marginal band, the limb eUiptical to broadly obovate or subor- 
bicular (elliptical to oblong in P. guianensis) and often decurrent on the claw, the 
margin erose or sometimes subentire, commonly with few to numerous pellucid 
cells abaxially. Androecium of 10 free stamens, those opposing the sepals shorter 
than those opposing the petals; filaments flat, glabrous except for a basal adaxial 
tuft (adaxially hirsute in P. sericea); anthers equal, glabrous, 4-locular, the connec- 
tive enlarged and exceeding the locules, the outer two locules each with a longitu- 
dinal wing; pollen tri- and tetracolporate. Receptacle with a ring of basifixed 
hairs, these forming tufts adhering adaxially at the base of the petals, sepals, and 
filaments. Gynoecium of 3 uniovulate, distinct carpels; styles free, inserted subapi- 
cally, subulate, glabrous or with scattered hairs adaxially in the proximal 1/5-3/4, 
the stigma apical, minute; ovary hirsute-tomentulose, the hairs mostly basifixed. 
Fruit a schizocarp of three cocci borne on a small torus; coccus subspherical, dry, 
indehiscent, hirsutulose-tomentulose (glabrescent in P. colombiana), often the 
areole with spongy tissue or only with a margin of scarious or cartilaginous tissue, 
carpophore absent; seed subspherical, outer cotyledon slightly larger than the 
inner, both folded back on themselves and the inner cotyledon nested within the 
outer cotyledon. 

The single chromosome count known for the genus, n - 12 for P. egleri, is 
with numbers reported for the Byrsonimoideae, which are n = 6, 12, or 
:W. R. Anderson, 1993). 

Key to the Species of Pterandra 



Petals pink 


; petioles of even 


the largest leav 


'es only 0.1-0..^ 


; cm 1 


long; small shrubs to : 


>m. 




5 abaxially tomen^ 


tose to sparse!; 


, so (rarely gla 


bratc 


5); stipules 3.5-8 mm 


long, . 


mm wid 


e; petals sparsely 


to densely seri 


ceous on claw 


and. 


center of limb: Brazil 


(Dist! 


Federal. 


, Goias, and Mina; 


s Gerais; Mato Grosso?). 






\.P.p 


2. Lamina: 


5 abaxially glabrous or with son 


le scattered h, 


airs. 


especially on the ma 


jor ve 


stipules 


2-2.5 mm long a: 


nd wide; petals abaxially glal 


5rous 


; or very sparsely ser 


iceous 


claw and center of limb; E 




-osso). 




2. P. hatsi 


Petals whit 


e or cream or gre. 


enish yellow, o 


ften becoming 


yelk 


,w in age; petioles up 


to 3.5 



ng, well over 0.5 cm long in larger leaves (except to 1 cm long in P. andersomi); shrub: 

Abaxial laminar pubescence composed mostly of erect basifixed hairs and a few scatte 
medifixed hairs; Brazil (Amazonas, Rondonia) and Bolivia (Beni). 1 1. f 

Abaxial laminar pubescence composed entirely of medifixed hairs (in P. evansii often ; 
with tufts of basifixed hairs in the axils formed by the costa and secondary veins, 
sometimes with basifixed hairs also scattered along the costa), or the laminas abaxi 
glabrate to glabrous. 

4. Stipules abaxially glabrous even in bud (but often with a sericeous patch at poin 
attachment to petiole in P. guianensis, with a sericeous patch at the apex in P. Colombia. 
5. Bracts and bracteoles abaxially densely pubescent; laminas abaxially sparsely s 
ceous, the hairs evenly distributed. 

6. Petals white: pedicels 0.4-0.6 mm in diameter, the pubescence white or silv 
leaf apex obtuse or acute or acuminate with a tip to 1 cm long; pellucid cell; 
abaxial surface of lamina abundant to sometimes absent; Brazil (Amapa, Am: 



and secondary 


veins 


, and sometimes with basifixed hairs al 


so scattered 


: costa), the cos 


;ta an 


d the secondary veins or only the cost; 


a prominent 


the tertiary veil 


nsson 






as abaxially ver 


yspar 


scly sericeous with a few widely scattered 


d, ferrugine- 


ppressed hairs { 


the h, 


airs sometimes faded to white or yellowish in age), 


he costa promii 


lent a 


ibaxially, the secondary veins at most ■ 


^ery slightly 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



tuft at the apex; laminas abaxially glabrate or very sparsely sericeous and 
widely scattered. 

7. Fascicles of flowers sessile, the pedicels 1.4-2 cm long; petioles up to 3.5 

Colombia (Antioquia). 6. / 

7. Fascicles of flowers borne on short projections (up to 3 mm long), the pedicels 

1-1.5 cm long; petioles up to 1.3 cm long; Panama (Colon). 7. P. mcphers 

Stipules abaxially densely sericeous, sometimes sparsely so in age (especially in P. 

8. Laminas abaxially tomentose with subsessile to stalked hairs, the pubescence not 
appressed and shed in patches in older leaves, the tertiary veins as well as the costa 
and secondary veins prominent abaxially; Brazil (Para). 14. P. e^ 

8. Laminas abaxially glabrate or sparsely to densely sericeous and the pubescence 

the costa and second 
along the costa), the 
abaxially. 



a (Amazonas). 12. P. flavescens. 

s abaxially evenly sericeous, the hairs white to yellowish (ferrugineous in 
osta and the secondary veins prominent abaxially. 
10. Laminas abaxially finely sericeous with ferrugineous hairs (in old leaves the 
hairs sometimes faded to yellow or golden), the hairs flattened and scalelike; 
stipules 4.5-6 mm long, 4-5 mm wide; Colombia (Valle, Isla Gorgona). 

10. Laminas abaxially sparsely to densely sericeous with white to yellowish hairs, 
1 1. Laminas abaxially densely sericeous, the hairs closely spaced and overlap- 
pubescence golden-yellow. 

12. Filaments adaxially hirsute; limb of lateral petals 3.5^.3 mm long, 2.8- 
3.6 mm wide; Guyana, Venezuela (Bolivar). 9. P. sericea. 

12. Filaments glabrous except for a basal tuft; limb of lateral petals 6.5-7 
mm long, 5.2-5.5 mm wide; Panama (Panama). 8. P. isthmica. 

11. Laminas abaxially sparsely sericeous, the hairs well separated and each 
readily discernible, not at all to slightly touching and overlapping; pedicels 
0.3-0.4 (-0.5) mm in diameter, the pubescence white to silvery. 

13. Laminas lanceolate to narrowly elliptical, the base acute or rounded, 
the petiole of larger leaves more than 1 cm long, abaxial vesture com- 
posed mostly of medifixed hairs but often also with tufts of basifixed 
hairs in the axils formed by the costa and secondary veins, and some- 
times with basifixed hairs also scattered along the costa), apex of lamina 

13. Laminas obovate to broadly elliptical, the base cuneate, the petiole of 
even the larger leaves only up to 1 cm long, abaxial pubescence com- 
posed only of medifixed hairs, apex of lamina rounded (-mucronate) to 
acute; Brazil (Maranhao). 3. p, andersonii. 

Pterandra pyroidea Adr. Jussieu in Saint-Hilaire, Fl. bras, merid. 3: 74. 1833 
["1832"].— Type: Brazil. Minas Gerais: "in campis," Saint-Hilaire B(l) 
1376 (holotype: P!; isotype: P!). 
Pterandra psidiifolia Adr. Jussieu in Saint-Hilaire, Fl. bras, merid. 3: 73. 1833 
["1832"]. Pterandra pyroidea f. psidiifolia (Adr. Jussieu) Niedenzu, Arbei- 
ten Bot. Inst. Konigl. Lyceums Hosianum Braunsberg 5: 33. 1914.— Type: 
Brazil. Minas Gerais: "in monte Serra da Canastra," Saint-Hilaire C(l) 
395 (holotype: P!; isotype: P!). 



PTERANDRA 9 

Shrublets of unbranched or little-branched stems to 2 m arising from a woody 
base. Laminas 6.5-16 cm long, 3.3-9 cm wide, elliptical to obovate, apex obtuse 
(-mucronate) to acute (-mucronate) or briefly acuminate, acumen up to 8 mm 
long, base rounded to truncate, adaxially tomentulose to sparsely so to glabrate to 
glabrous, abaxially tomentose to sparsely so to sometimes glabrate, the hairs 0.6- 
1.7 mm long, medifixed, terete, wavy to crisped to curled, white to yellowish, 
subsessile or with a stalk to 0.2 mm long, overlapping, pellucid cells abundant to 
absent in abaxial surface or sometimes only along the margin, costa and second- 
ary and tertiary veins prominent abaxially; petioles 2.5-5 mm long, densely seri- 
ceous; each pair of stipules entirely connate or sometimes nearly so but the apex 
notched or connate in the proximal 3/4 but the distal 1/4 free, 3.5-8 mm long, 4-7 
mm wide, adaxially hirsute, abaxially sericeous. Inflorescence of sessile to sub- 
sessile, 4-6-flowered fascicles borne below young or mature leaves or distal new 
leaves not yet developed at anthesis, each fascicle subtended by a deciduous bract 
composed of a pair of stipules and a rudimentary leaf; pedicels 0.7-2.5 cm long, 
0.5-0.7 mm in diameter, densely white-sericeous; bracts 1.8-3 (-4) mm long, 1-2.5 
mm wide, triangular, bracteoles 1.4-2.5 mm long, 0.4-1 mm wide, narrowly trian- 
gular to linear, bracts and bracteoles with the apex acute, abaxially with hairs concen- 
trated on and adjacent to the costa and apex. Sepals 2.7-4.2 mm long, 2.5-4.3 mm 
wide, triangular, apex obtuse, erect or sHghtly recurved, glands 0.8-2 mm long, 
0.5-1.2 mm wide, or glands absent. Petals pink, the limb broadly obovate or 
elliptical to orbicular, margin erose, abaxially very sparsely to densely sericeous 
on claw and center of limb or over most of limb except for a band along the 
margin ca. 0.5 mm wide or sometimes glabrate to glabrous; lateral petals: claw 
0.8-1.5 mm long, limb 6-7 (-8) mm long, 4.5-6 (-7.5) mm wide; posterior petal: 
claw 1-1.5 mm long, hmb 6.5-8.5 mm long, 5.5-7.5 mm wide. Filaments glabrous 
except for an adaxial basal tuft of hairs, those of stamens opposing sepals 1.8-2.5 
mm long, those of stamens opposing petals 2-3.2 mm long; anthers 1.8-2.3 mm 
long, wing of outer locules 0.3-0.5 mm wide. Styles 3.8-5.4 mm long, 0.3-0.4 mm 
in diameter, glabrous. Coccus 4-4.5 mm high, 4-5 mm in diameter, hirsutulose- 
tomentulose; torus up to 0.5 mm high; embryo globose, ca. 3 mm long, radicle ca. 
1.3 mm long, outer cotyledon ca. 5.7 mm long, ca. 2.5 mm long, folded at 1/2 its 
length and the distal tip folded over the inner cotyledon, inner cotyledon ca. 4.5 
mm long, ca. 2.5 mm wide, folded three times. Chromosome number unknown. 

Phenology. Collected in flower from April through December, in fruit from 
September to December. 

Distribution (Fig. 1). Brazil (Distrito Federal, Goias, and Minas Gerais; one 
collection purportedly from Mato Grosso); cerrado, campo sujo, and campo; 650- 



Representative Specimens. Brazil. Distrjto Federal: Brasilia, bacia do Rio Sao Bartolomeu 
Rebrotamento, 14 May 1980, Heringer et al. 4771 (IBGE, MICH); Chapada da Contagem, ca. 15 iim 
E of Brasilia, 700-1000 m, 18 Aug 1964, Irwin & Soderstrom 5274 (NY, UB, US); ca. 35 km SW of 
Brasilia on rd to Anapolis, 700-1000 m, 5 Sep 1964, Irwin & Soderstrom 6032 (F, MICH, MO, NY, 
UB); Cdrrego Jeriva,, ca. 10 km E of Brasilia, 975 m, 15 Sep 1965, Irwin et al 8342 (NY, UB); near 
Corrego Taquari, E of Lagoa Paranoa, 975 m, 28 Apr 1966, Irwin et al. 15404 (NY); 30 km N of 
Brasilia, Corrego Sobradinho, Farm No. 19, 4 Oct 1963, Maguire et al. 57015 (MICH, MO, NY).— 
GoiAs: Serra Geral do Parang, 3 km by rd S of Sao Joao da Alian^a, 1040 m, 23 Mar 1973, Anderson 
7750 (NY, UB); ca. 15 km (straight line) N of Corumba, 1230 m, 16 May 1973, Anderson 10421 
(MICH, NY, UB); Ipameri, Rio Corumba, 4 Oct 1976, Hatschbach 38930 (MBM, MICH); Alto 
Paraiso, 2-5 km Oeste, 1200 m, 15 Oct 1990, Hatschbach 54571 (MBM, MICH); Serra do Caiapo, ca. 
35 km S of Caiaponia on rd to Jatai, 17°12'S, 51°47'W, 800-1000 m, 19 Oct 1964, Irwin & Soderstrom 
7055 (F, MICH, NY); Chapada dos Veadeiros, 24 km NW of Veadeiros, rd to Cavalcante, 14°S, 



10 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

47°W, 1200 m, 22 Oct 1965, Irwin et al. 9511 (F, MICH, NY, UB).— Mato Grosso (perhaps an error 
for Minas Gerais, see below): without locality, 15 Jul 1969, Saddi RFA12314 (MICH).— Minas Gera[s: 
5 km by rd NE of Rio Manso and Coute de Magalhaes, 17°53'S, 44°15'W, 960-1000 m, 13 Apr 1973, 
Anderson 8762 (F, MICH, NY, W); Jaguara, 28 Oct 1965, Goodhmd 122 (F, MICH, MO, NY): 
Buenopolis, Serra do Cabral, a 6-7 km da cidade, 17°53'S, 44°15'W. 760 m. 12 Oct 1988, Harley et al. 
24857 (F, MBM, MICH, RB, UB); Araxa, Rod. BR-262, 12 Oct 1982, Halschbach 45623 (MBM, 
MICH); S'Ana do Riacho, 25 Oct 1974, Hatschbach 35325 (MBM, MICH); Belo Horizonte, Morro 
das Pedras, 1000 m, 18 Sep 1945, Williams & Assis 7574 (BR, F, GH, MO, NY, R, US). 

Ptemndra pyroidea is a common plant of the Brazilian Planalto, where it 
forms Httle-branched shrubs bearing showy clusters of large pink flowers. It is 
similar to P. hatschbachii of Mato Grosso, a weak shrublet (less than 0.5 m tall) 
also with large pink petals, which differs most strikingly in its glabrous or nearly 
glabrous herbage and flowers. It has very small stipules, only 2-2.5 mm long and 
wide; those of P. pyroidea are 3.5-8 mm long, 4-7 mm wide. The only record of P. 
pyroidea from Mato Grosso {Saddi R FA 1 23 14) is questionable. The label only 
records the collector, date, and "Mato Grosso." Perhaps the specimen was collected 
in Minas Gerais, and the abbreviation "MG" misinterpreted as "Mato Grosso." It 
seems unhkely that such a common and conspicuous plant would have been over- 
looked by all other collectors who visited Mato Grosso. 

2. Pterandra hatschbachii W. R. Anderson, Contr. Univ. Michigan Herb. 19: 388. 
1993.— Type: Brazil. Mato Grosso: Alto Araguaia, Rib. Claro, 22 Sep 
1974, Hatschbach 35085 (holotype: MBM!; isotype: MICH!). Fig. 3. 

Shrublets less than 0.5 m tall arising from woody, trailing, underground stems. 
Laminas 4.2-12.5 cm long, 1.5-4.4 cm wide, obovate or oblanceolate to elliptical, 
apex acute (-mucronate) to obtuse (-mucronate) to emarginate (-mucronate), base 
cuneate or rounded to truncate, adaxially glabrous, abaxially entirely glabrous or 
sparsely sericeous on the costa and major veins or glabrate, the hairs 0.3-1.3 mm 
long, medifixed, terete, straight or wavy, white and golden, sessile or subsessile or 
with a stalk up to 0.1 mm long, rarely also with a few basifixed hairs ca. 0.4 min 
long, pellucid cells along margin of abaxial surface or absent, costa and secondary 
veins prominent abaxially, tertiary veins usually shghtly raised as well; petioles 1- 
2.5 mm long, densely sericeous; each pair of stipules entirely connate or nearly so 
but with the apex notched, or connate in the proximal 1/3-1/2 and distally free, 2- 
2.5 mm long and wide, adaxially hirsute, abaxially sericeous. Inflorescence of 
sessile to subsessile, ca. 6-flowered fascicles below a flush of new leaves or some- 
times the distal new leaves not yet developed at anthesis, each fascicle subtended 
by a deciduous bract composed of a pair of stipules and a rudimentary leaf; 
pedicels 0.9-2.5 cm long, 0.3-0.4 mm in diameter, densely yellowish to white- 
sericeous; bracts 1-1.7 mm long, 0.5-1 mm wide, triangular, bracteoles 0.7-1.2 mm 
long, 0.2-0.5 mm wide, linear to narrowly triangular, bracts and bracteoles with 
the apex acute, abaxially with scattered hairs especially along the costa and on the 
apex. Sepals 2.3-2.5 mm long, 1.8-2.7 mm wide, triangular, apex obtuse, revolute 
or recurved, glands 1-1.8 mm long, 0.6-1 mm wide. Petals pink (fading to white in 
age), the limb orbicular or broadly obovate, margin erose, abaxially glabrous to 
sparsely sericeous on claw and center of limb; lateral petals: claw (0.8-) 1-1.3 mm 
long, limb 6-7 (-7.5) mm long, (4.5-) 5-5.5 mm wide; posterior petal: claw 1.2-1.5 (- 
1.7) mm long, limb 6.5-7.5 mm long and wide. Filaments glabrous except for an 
adaxial basal tuft of hairs, those of stamens opposing sepals 2-2.5 mm long, those 




FIG. 3. Pterand. 
c. Base of flower cluster with bracts and bracteoles, the e 
(composed of a rudimentary leaf and stipules), x5. d. Flow 
Posterior petal, abaxial view showing sparse pubescence on 
adaxial (left) and abaxial (right) views, x7.5. g. Distal portion of stamen, la 
a, b, d-g, Hatschbach 35085; c, Hatschbach 33276; drawn by Karin Douthit 



of Stamens opposing petals 2.5-3.3 mm long; anthers 1.4-2 mm long, wing of outer 
locules 0.2-0.3 mm wide. Styles 4.5-5.2 mm long, 0.3 mm in diameter, glabrous or 
with scattered hairs adaxially in the proximal 1/5. Coccus ca. 3.5 mm high and in 
diameter, hirsutulose-tomentulose; torus up to 0.5 mm high (?); mature seed not 
seen. Chromosome number unknown. 

Phenology. Collected in flower in September, November, and February, in 
young fruit in November. 

Distribution (Fig. 1). Brazil (Matro Grosso); forming dense carpets in grassy 
campos; ca. 800 m. 



AoDniONAL Specimens Examined. Brazil. Mato Gross 
rao Claro, NW of Alto Araguaia, ca. 800 m, 15 Feb 1975. 
Alto Araguaia. Alto Araguaia, 15 Nov 



to Araguaia, vicmity ot KiCie- 
11400 (MBM, MICH); Mpio. 
33276 (MBM, NY). 



12 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Pterandra hatschbachii is known only from the vicinity of the type locahty in 
Mato Grosso, close to the border with Goias. It shares many characters with P. 
pyroidea, a common species of the cerrado and campo of Goias and Minas Gerais. 
Both are low shrubs bearing showy inflorescences composed of large flowers with 
pink petals. In all other species the petals are white or cream. Pterandra hatsch- 
bachii is readily separated from P. pyroidea by its low stature, small stipules (only 
2-2.5 mm long and wide), and glabrous or at most very sparsely pubescent vegeta- 
tive parts and petals. 

3. Pterandra andersonii C. Anderson, sp. nov.— Type: Brazil. Maranhao: Grajau, 
140 m, 3 Aug 1909, Lisboa 2507 {hoXoiy^Q: NY!; isotypes: MG! RB!). 

Fig. 4a-e. 

Frutex usque ad 3 m altus. Laminae 6.2-12.5 cm longae, 3.5-7 cm latae, obo- 
vatae vel eUipticae, supra glabrae, subtus sericeae vel sparsim sericeae sed costa 
dense sericea; petioli 0.3-1 cm longi, dense sericei; stipulae (2.5-) 3-3.5 mm lon- 
gae lataeque, connatae, adaxialiter hirsutae, abaxialiter sericeae. Inflorescentia ex 
fasciculis sessilibus vel subsessihbus constans, floribus cujusque fasciculi 4-6, sine 
fohis juvenilibus per anthesin; pedicelli 1.2-1.7 cm longi, 0.4-0.5 mm diametro, 
dense albo-sericei. Petala cremea (?), limbo late elliptico vel late obovato vel 
suborbiculari; limbi petalorum lateralium 3.5-4 mm longi, ca. 3 mm lati, limbus 
petali postici ca. 4.5 mm longus, 3.5-4 mm latus. Filamenta praeter caespitem 
basalem glabra. Styh ca. 4.3 mm longi. Cocci 3-3.5 mm alti et diametro, hirsutulo- 
tomentosi. 

Shrubs to 3 m. Laminas 6.2-12.5 cm long, 3.5-7 cm wide, obovate to elliptical, 
apex obtuse (-mucronate) to acute, base cuneate, adaxially glabrous, abaxially 
sericeous or sparsely so but densely so on the costa, the hairs (0.1-) 0.2-0.7 (-0.9) 
mm long, medifixed, terete, straight, mostly white or some golden, sessile or sub- 
sessile or sometimes with a stalk up to 0.1 mm long, slightly touching or overlap- 
ping or not, pellucid cells present in abaxial surface, costa and secondary veins 
prominent abaxially, tertiary veins usually slightly raised as well; petioles 0.3-1 cm 
long, densely sericeous; each pair of stipules connate, (2.5-) 3-3.5 mm long and 
wide, adaxially hirsute, abaxially sericeous. Inflorescence of sessile or subsessile, 
4-6-flowered fascicles, each fascicle subtended by a deciduous bract composed of 
a pair of stipules and a rudimentary leaf, distal new leaves not yet developed at 
anthesis; pedicels 1.2-1.7 cm long, 0.4-0.5 mm in diameter, densely white-seri- 
ceous; bracts 1.3-2.5 mm long, 0.8-1.5 mm wide, triangular, bracteoles 1-1.5 mm 
long, 0.3-0.6 mm wide, very narrowly triangular to linear, bracts and bracteoles 

a. 'lerminal portion of shoot with four leaves. x().5. b. Torminal portion of shoot with two inflores- 
cences, x().5. c. Petal, abaxial view. x5. d. Distal portions of stamens, ada.xial (left) and lateral (right) 
views, x7.5. e. Coccus, lateral view, x5. f-i, P. cohmbiana. f. Terminal portion of shool with infructcs- 
cence, x0..5. g. Leaf. xO.5. h. Base of leaf with stipules (basally connate but free in the distal 1/2), 



C. ANDERSON: PTERANDRA 




14 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

with the apex acute, abaxialiy glabrous except for tuft at the apex and appressed 
often scattered hairs on and adjacent to the costa. Sepals ca. 2.3 mm long, ca. 1.7 
mm wide, narrowly triangular (the distal 2/3 hgulate), recurved, glands 1.6-1.8 
mm long, ca. 0.7 mm wide. Petals "yellow" (?; probably creamy white and becom- 
ing yellow or perhaps pink in age), the limb broadly elliptical or broadly obovate 
to suborbicular, margin erose, abaxialiy sparsely sericeous on claw and center of 
limb or only on limb and the claw glabrous to glabrate; lateral petals: claw 1.2-1.5 
mm long, limb 3.5-4 mm long, ca. 3 mm wide; posterior petal: claw 1.5-1.6 mm 
long, hmb ca. 4.5 mm long, 3.5-4 mm wide. Filaments glabrous except for an 
adaxial basal tuft of hairs, those of stamens opposing sepals ca. 2 mm long, those 
of stamens opposing petals ca. 3 mm long; anthers ca. 1.2 mm long, wing of outer 
locules 0.2 mm wide. Styles ca. 4.3 mm long, ca. 0.3 mm in diameter, with scat- 
tered hairs adaxially in the proximal 1/4. Coccus 3-3.5 mm high and in diameter, 
hirsutulose-tomentulose; torus ca. 1 mm high; mature seed not seen. Chromosome 
number unknown. 



?ao, 31 Jul 1976, i 

Pterandra andersonii is known from only two collections from Maranhao, 
Brazil (Fig. 1). It resembles P. pyroidea of central Brazil in that its leaves have a 
cuneate base and a very short petiole (only up to 1 cm long); however, the laminas 
of P. andersonii are abaxialiy sericeous rather than tomentulose, and it has smaller 
stipules and smaller flowers. Pterandra pyroidea and the similar P. hatschhachii 
have pink petals. Those of P. andersonii are noted on the label as "amarella" 
{Lishoa 2507) and "avermelhada" {Pinheiro 12; m fruit); however, most likely the 
petals are creamy white at the onset of anthesis and turn yellow or reddish in age, 
as in other species with white petals. The laminas of the type collection are strik- 
ingly obtuse at the apex (Fig. 4a), yet larger laminas of Pinheiro 12, from a 
vegetative branch with long internodes, are acute. 

This species is named for William R. Anderson, student of Malpighiaceae, 
who first noted its distinctness. 

4. Pterandra arborea Ducke, Bull. Mus. Hist. Nat. (Paris), ser. 2, 4: 736. 1932.— 
Type: Brazil. Amazonas: Manaus, 3 Aug 1929, Ducke s.n. [RB23649] 
(holotype: RB!; isotypes: G! K! P! US!). 



Tree to 30 m. Laminas 7.5-18.5 cm long, 3.5-8.5 


cm wide, elliptical to obo- 


vate, apex obtuse to acute but commonly acuminate. 


acumen up to 10 mm long. 


base cuneate, adaxially glabrate to glabrous but often 


with appressed hairs on the 


major veins, abaxialiy sparsely sericeous (densely so o 


n the major veins), the hairs 


0.1-0.4 mm long, medifixed, terete, straight, white to 


yellowish to golden, sessile. 


not or only slightly overlapping, pellucid cells abur 


idant in abaxial surface to 


absent, costa and secondary veins prominent abaxialiy. 


tertiary veins usually slightly 


raised as well; petioles 0.8-2 cm long, densely seric 


eous; each pair of stipules 


connate or the distal 1/3-1/2 free, 2.5-3.5 mm long 


, ca. 3 mm wide, adaxially 


hirsute, abaxialiy glabrous but with a sericeous band 2 


It the base. Inflorescence of 


4-6-flowered fascicles, sessile or borne on a projectior 


I up to 1.4 mm long below a 



flush of new leaves, each fascicle subtended by a deciduous bract composed of i 
pair of stipules and a rudimentary leaf; pedicels 0.8-2.2 cm long, 0.4-0.6 mm ir 



1997 C. ANDERSON: PTERANDRA 15 

diameter, densely white- or silvery-sericeous; bracts 1-1.6 mm long, 1.2-1.5 mm 
wide, bracteoles 1.2-1.3 mm long, 1.2-1.3 mm wide, bracts and bracteoles triangu- 
lar, the apex acute to mucronate, abaxially densely white-sericeous. Sepals 1.8-3.2 
mm long, 1.8-2.2 mm wide, narrowly triangular (the distal 2/3 ligulate), apex 
obtuse, recurved to revolute, glands 1.8-2.5 mm long, 1-1.2 mm wide. Petals white 
(turning pale yellow in age), the limb elliptical to obovate to suborbicular, margin 
erose, abaxially densely sericeous on claw and center of hmb or only along the 
central vein; lateral petals: claw 0.7-1.3 mm long, limb (2.8-) 3.5-4.2 mm long, 
(2.2-) 2.5-2.8 mm wide; posterior petal: claw (0.8-) 1-1.5 mm long, hmb (3.5-) 
4.2-4.3 mm long, (2.5-) 2.8-3 mm wide. Filaments glabrous except for an adaxial 
basal tuft of hairs or sometimes adaxially with scattered hairs in the proximal 1/2, 
those of stamens opposing sepals 1.2-2.1 mm long, those of stamens opposing 
petals 1.8-3.5 mm long; anthers 0.6-0.9 mm long, wing of outer locules 0.2 mm 
wide. Styles (2.5-) 3.5 mm long, 0.2-0.4 mm in diameter, with scattered hairs 
adaxially in the proximal 1/4-3/4. Coccus 3-3.7 mm high, 3.2-3.8 mm in diameter, 
hirsutulose-tomentulose or sparsely so; torus 1.5-2 mm high; embryo globose, ca. 
3 mm in diameter, radicle ca. 1.8 mm long, outer cotyledon ca. 5.7 mm long, ca. 
1.8 mm wide, folded at 1/2 of its length and the tip folded over the inner cotyle- 
don, inner cotyledon ca. 4 mm long, ca. 1.6 mm wide, folded at 1/2 of its length. 
Chromosome number unknown. 

Phenology. Collected in flower in March and from July through October, in 
fruit in January and from July through November. 

Distribution (Fig. 1). Brazil (Amapa, Amazonas, Para); in forest; 50-800 m. 

Additional Specimens Examined. Brazil. Amapa: Serra do Navio, Rio Amapari, 70-300 m, 4 
Nov 1954, Cowan 38123 (MICH, NY); Mazagao, area do Felipe VI, 26 Aug 1985, N. T. Silva 5524 
(MG).— Amaz6nas: Reserva Florestal Ducke, Manaus-Itacoatiara, Km 26, 02°53'S, 59°58'W, 14 Aug 
1996, Assungdo 365 (MICH); Dist. Agropecuario, Reserva 1501 (Km 41) of the WWF, INPA MCS 
Project, 02°24'-02°25'S, 59°43'-59°45'W, ca. 50-125 m, 25 Nov 1988, Boom et al. 8649 (F, NY); 
Manaus, margem do igarap6 do Bind^, 30 Jul 1956, Coelho INPA 4009 (IAN, INPA); estrada Manaus- 
Caracarai, Km 58, Reserva Biologica INPA-SUFRAMA, Sep 1976, Como & Damiao 855 (INPA); 
Reserva Florestal Ducke, Manaus-Itacoatiara, Km 26, 02°53'S, 59°58'W, 27 Jul 1994, Hopkins 1468 
(MICH); Dist. Agropecuario, 90 km NNE de Manaus, Reserva 1501 (Km 41), Projeto Dinamica 
Biologica de Fragmentos Florestais, 02°24'-25'S, 59°43'^5'W, 50-125 m, 11 Aug 1991, Oliveim et al. 
144 (NY); Dist. Agropecuario, Fazenda Dimona of the WWF, INPA MCS Project, 72 kms N of 
Manaus, 02°19'S, 60°05'W, ca. 50-125 m, 6 Nov 1988, Pacheco et al. 49 (MBM, NY); Reserva Florestal 
Ducke, Km 26, 31 Aug 1966, Prance et al. 2122 (F, MG, MICH, NY, R, US); estrada Manaus- 
Caracarai, Km 39, Reserva Experimental de Silvicultura Tropical, Ribamar, 14 Sep 1977, Ramos 221 
(INPA); Reserva Florestal Ducke, Manaus-Itacoatiara, Km 26, 02°53'S, 59°58'W, 21 Jul 1994. Riheiro 
1346 (MICH); Reserva Florestal Ducke (P-69), 21 Aug 1963, Rodrigues 5444 (F); Manaus. cslrada 
Manaus-Itacoatiara, Km 160, 15 Oct 1965, Rodrigues 7244 (INPA); Manaus, estrada Manaus-Ila- 
coatiara. Km 125, 24 Sep 1965, Rodrigues 7577 (INPA); Reserva Florestal Ducke, perto da eslacao 
metereologica, 9 Sep 1964, Rodrigues 32 (F); Reserva Florestal Ducke, 21 Jul L964, Rodrigues & 
Loureiro 5961 (F, US); Manaus, estrada Manaus-Itacoatiara, Km 73, 30 Aug 1965, Rodrigues & 
Loureiro 7058 (INPA); between Missao Salesiana and Serra Pirapucu, Rio Maturaca, 400-800 m, 13 
Jan 1966, N. T. Silva & Brazao 60825 (F, MG, MICH, NY).— Para: Estacao Ecologica do Jari, 
Projeto Reserva Genetica, SEMA, 00°75'S, 52°30'W, 14 Oct 1987, Beck 98 (NY). 

Ptemndra arborea is a common tree in the area of Manaus and thus more 
frequently collected than other species, except P. pyroldea. It had been consid- 
ered endemic to the Manaus region but in the last 30 years has also been found in 
Amapa, Para, and northwestern Amazonas. It is one of four species in which the 
stipules are glabrous abaxially even in bud. Pterandra arborea is most similar to 



16 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

P. guianensis of Guyana, which differs most strikingly in its 2-3-flowered fascicles 
and stout, golden-yellow pedicels. In P. arborea the abaxial surface of the laminas 
commonly is sprinkled with abundant pellucid cells; in P. guianensis such cells are 
few or absent. 

5. Pterandra guianensis W. R. Anderson, Mem. New York Bot. Garden 32: 35. 
1981.— Type: Guyana. Mt. Ayanganna, below 762 m, 3 Aug 1960, Tilktt 
etal 45011 (holotype: MICH!; isotypes: F! K! NY! US!). 

Trees to 20 m. Laminas 8.5-14 cm long, 3.6-6 cm wide, oblanceolate to ellipti- 
cal, apex acuminate to subcaudate, acumen up to 1.5 cm long, base cuneate, 
adaxially glabrous or sometimes with appressed hairs on the costa, abaxially sparsely 
sericeous, the hairs 0.1-0.6 (-0.7) mm long, medifixed, terete, straight, yellowish 
to golden, sessile, not or only slightly touching or overlapping, pellucid cells in 
abaxial surface very few to absent, costa and secondary veins prominent abaxially, 
tertiary veins not raised or sometimes very slightly so; petioles 0.8-1.5 cm long, 
densely sericeous; each pair of stipules entirely connate or mostly connate but 
notched at the apex, 3-3.7 mm long, 2.5-3 mm wide, adaxially hirsute, abaxially 
glabrous but often with a sericeous patch at point of attachment to petiole. Inflo- 
rescence of 2-3-flowered fascicles borne on short projections ca. 0.5-3 mm long 
below a flush of new leaves, each fascicle subtended by a deciduous bract com- 
posed of a pair of stipules and a rudimentary leaf; pedicels (0.8-) 1.3-1.7 (-2.3) cm 
long, (0.6-) 0.8-1 mm in diameter, densely golden-sericeous; bracts 1.3-1.5 mm 
long and wide, bracteoles 1.2-1.3 mm long, ca. 1 mm wide, bracts and bracteoles 
triangular, the apex acute, abaxially densely sericeous. Sepals 2.5-2.7 mm long, 2.2- 
2.5 mm wide, narrowly triangular (the distal 1/2 ligulate), apex obtuse, recurved, 
glands 1.6-1.8 mm long, 0.8-0.9 mm wide. Petals "greenish yellow," the limb 
oblong to elliptical, margin subentire, abaxially sericeous on claw and more sparsely 
so on center of limb; lateral petals: claw 0.8-1.1 mm long, limb 4-4.3 mm long, 
(2.5-) 3-3.3 mm wide; posterior petal: claw 1.3-1.4 mm long, limb 4.5-4.7 mm 
long, 3.5-4 mm wide. Filaments glabrous except for an adaxial basal tuft of hairs, 
those of stamens opposing sepals 1.7-1.8 mm long, those of stamens opposing 
petals ca. 2.1 mm long; anthers 1.1 mm long, wing of outer locules 0.2 mm wide. 
Styles ca. 2.6 mm long, ca. 0.2 mm in diameter, glabrous. Mature fruit not seen. 
Chromosome number unknown. 

Pterandra guianensis is known only from the type collected in mixed ever- 
green forest on Mt. Ayanganna in Guyana (Fig. 1). Like the somewhat similar 
Amazonian P. arborea, it has the stipules abaxially glabrous and the bracts and 
bracteoles abaxially densely pubescent. Pterandra guianensis is readily separated 
by its stout and golden-yellow pedicels and fascicles of only 2-3 flowers. 

6. Pterandra colombiana C. Anderson, sp. nov.— Type: Colombia. Antioquia: Zara- 
goza, Corregimiento Saltillo en la via Zaragoza-Segovia, 3.7-16 km S de 
Zaragoza, 07°26'N, 74°50'W, 150-200 m, 13 Jul 1987, Callejas et al. 4656 
(holotype: MICH!; isotypes: HUA! MO! NY!). Fig. 4f-i. 

Arbor usque ad 22 m alta. Laminae 8-19.5 cm longae, 2.7-8 cm latae, obova- 
tae vel elhpticae, supra glabrae, subtus sparsissime sericeae vel glabratae; petioh 
0.8-3.5 cm longi, juniores dense sericei, vetustiores glabrati; stipulae 2.8-4 mm 
longae, 3-4 mm latae, connatae sed distaliter Hberae, adaxialiter hirsutae, abaxi- 
aliter glabrae sed apice sericeae. Inflorescentia ex fasciculis sessilibus constans. 



1997 C. ANDERSON: PTERANDRA 17 

floribus cujusque fasciculi 4-6, sine foliis juvenilibus per anthesin; pedicelli 1.4-2 
cm longi, 0.5-0.6 mm diametro, dense aureo-sericei vel albo-sericei. Petala alba, 
limbo late obovato vel suborbiculari, 3.2^.5 mm longo, 2.8^ mm lato. Filamenta 
praeter caespitem basalem glabra. Styli 3.5-4.8 mm longi. Cocci ca. 3.5 mm alti et 
diametro, glabri vel glabrati sed margine areolae pubescentes. 

Tree to 22 m. Laminas 8-19.5 cm long, 2.7-8 cm wide, elliptical to obovate, 
apex acuminate to caudate, acumen up to 2 cm long, base cuneate, adaxially 
glabrous, abaxially very sparsely sericeous to glabrate, the hairs 0.1-0.9 mm long, 
medifixed, terete, straight, white to yellowish, sessile, widely spaced and rarely 
touching, pellucid cells present in abaxial surface, costa and secondary veins prom- 
inent abaxially, tertiary veins usually slightly raised as well; petioles 0.8-3.5 cm 
long, densely sericeous when young, becoming glabrate; each pair of stipules con- 
nate in the proximal 1/3-1/2 and distally free, 2.8-4 mm long, 3-4 mm wide, 
adaxially hirsute, abaxially glabrous except for a sericeous patch at the apex. 
Inflorescence of sessile, 4-6-flowered fascicles, each fascicle subtended by a decid- 
uous bract composed of a pair of stipules and a rudimentary leaf, usually the 
lowestmost cluster at a node above a pair of mature leaves, distal new leaves not 
yet developed at anthesis; pedicels 1.4-2 cm long, 0.5-0.6 mm in diameter, densely 
yellowish or white-sericeous; bracts 1.8-2.4 mm long, 1-1.5 mm wide, triangular, 
bracteoles 1.8-2 mm long, 0.4-0.6 mm wide, hnear, bracts and bracteoles with the 
apex acute, abaxially glabrous or pubescent on the costa in the distal 1/2 and/or 
with a tuft of hairs at the apex. Sepals 2.5-3 mm long, 1.8-2.3 mm wide, narrowly 
triangular (the distal 1/2 hgulate), apex obtuse, recurved, glands 1.6-2 mm long, 
0.8-1 mm wide. Petals white (becoming yellow in age), the limb broadly obovate 
to suborbicular, margin subentire to slightly erose, abaxially densely sericeous on 
claw and center of limb or over most of limb except for a marginal band ca. 0.5 
mm wide; lateral petals: claw (0.8-) 1-1.2 mm long, limb 3.2-4.5 mm long, 2.8-4 
mm wide; posterior petal: claw 1.3-1.5 mm long, limb 3.2-4.5 mm long, 2.8-4 mm 
wide. Filaments glabrous except for an adaxial basal tuft of hairs, those of stamens 
opposing sepals 1.1-1.2 (-1.5) mm long, those of stamens opposing petals 2-2.5 
mm long; anthers 0.9-1.1 mm long, wing of outer locules 0.2-0.3 mm wide. Styles 
3.5-4.8 mm long, 0.2-0.3 mm in diameter, glabrous or with scattered hairs adaxially 
in the proximal 1/5-1/4. Coccus ca. 3.5 mm high and in diameter, glabrous or 
glabrate except for a ring of hairs around the areole; torus up to 0.5 mm high; 
mature seed not seen. Chromosome number unknown. 

Phenology. Collected in flower in September, December, January, and July, 
in fruit in July. 

Distribution (Fig. 1). Colombia (Antioquia); in wet forest; 150-800 m. 

Additional Specimens Examined. Colombia. Antioquia: Segovia, Corregimiento Fraguas, via 
Zaragoza-Segovia. 25.5-40 km S de Zaragoza, 07°24'N, 74°50'W, 250 m, 13 Jul 1987, Callejas & 
Betancur 4667 (HUA, MICH, MO, NY); San Luis, Corregimiento de El Prodigio, Finca Dormene y 
Serranias, 06°06'N, 74°48'W, 350-400 m, 25 Jun 1990, Cardenas et al. 2863 (MICH); San Luis, auto- 
pista Medellin-Bogota, 2 km de Rio Claro hacia Rio Samana, 790 m, 13 Jan 1983, Cogollo & Brand 
354 (MO); San Luis, autopista Medellin-Bogota, sector Rio Samana-Rio Claro, puente sobre la 
quebrada Cristalina, 790 m, 19 Dec 1982, Cogollo & Estrada 317 (MO); Anori, Tirana Creek, at the 
confluence with Rio Anori, 5 km upriver from Providencia, 400-900 m, 6-12 Sep 1973, Soejarto 4258 
(HUA, MEDEL). 

Pterandra colombiana is distinctive for its relatively large laminas that are 
nearly glabrous or at most very sparsely sericeous below. The only other species 
with glabrate laminas is P. mcphersonii, which also has abaxially glabrous stipules. 



18 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

In P. mcphersonii the fascicles of flowers are borne on small projections, whereas 
those of P. colombiana are always sessile. Of the species of Pterandra known in 
fruit, P. colombiana is the only one in which the mature cocci are glabrous except 
for a ring of hairs at the areole; in others the cocci are hirsutulose-tomentulose. 

7. Pterandra mcphersonii C. Anderson, sp. nov.— Type: Panama. Colon, Santa 
Rita ridge, SE of Colon, about 12 rd-mi from Trans-isthmian Hwy, 09°25'N, 
79°40'W, 500 m, 17 Sep 1987, McPherson 11745 (holotype: MO!). 

Fig. 4n-p. 

Arbor usque ad 11 m alta. Laminae 7-13 cm longae, 3-6 cm latae, ellipticae, 
supra glabrae, subtus sparsissime sericeae vel glabratae; petioli 0.7-1.3 cm longi, 
juniores dense sericei, vetustiores glabrati; stipulae 3-4.5 mm longae, 3-4 mm 
latae, connatae sed distaliter liberae, adaxialiter hirsutae, abaxialiter glabrae sed 
apice margineque sericeae. Inflorescentia ex fasciculis sessihbus vel brevistipitatis 
constans; floribus cujusque fasciculi 4 (-6?), sine foHis juvenilibus per anthesin; 
pedicelh 1-1.5 cm longi, 0.6-0.7 mm diametro, dense aureo-sericei. Petala alba, 
limbo late obovato vel suborbiculari; limbi petalorum lateralium 5-5.2 mm longi, (4-) 
4.2-4.8 mm lati, limbus petah postici 5.5-6 mm longus, 4.2-4.5 mm latus. Filamen- 
ta praeter caespitem basalem glabra. Styli ca. 3.5 mm longi. Cocci maturi ignoti. 

Tree to 11 m. Laminas 7-13 cm long, 3-6 cm wide, elliptical, apex acuminate, 
acumen up to 1.5 cm long, base cuneate, adaxially glabrous, abaxially very sparsely 
sericeous to glabrate, the hairs 0.1-0.7 mm long, medifixed, terete or also some 
flattened, straight, golden, sessile, rarely touching, pellucid cells very few in abaxial 
surface, costa and secondary veins prominent abaxially, tertiary veins not raised 
or sometimes shghtly so; petioles 0.7-1.3 cm long, densely sericeous when young, 
becoming glabrate; each pair of stipules connate in the proximal 1/2, distally free, 
3-4.5 mm long, 3-4 mm wide, adaxially hirsute, abaxially glabrous except for hairs 
along the margin and apex. Inflorescence of 4 (-6?)-flowered fascicles, these sub- 
sessile or borne on short projections up to 3 mm long, each fascicle subtended by 
a deciduous bract composed of a pair of stipules and a rudimentary leaf, the 
lowestmost cluster at a node above a pair of mature leaves, distal new leaves not 
yet developed at anthesis; pedicels 1-1.5 cm long, 0.6-0.7 mm in diameter, densely 
golden-sericeous; bracts ca. 1.8 mm long, ca. 1 mm wide, triangular, bracteoles 
1.5-1.7 mm long, ca. 0.8 mm wide, narrowly triangular, bracts and bracteoles with 
the apex acute, abaxially glabrous or with hairs on the costa in the distal 1/3 and 
at the apex. Sepals ca. 2.5 mm long, ca. 2.2 mm wide, narrowly triangular (the 
distal 1/2 hgulate), apex obtuse, recurved, glands 2.2-2.3 mm long, ca. 1 mm wide. 
Petals white (becoming yellow in age), the limb broadly obovate to suborbicular, 
margin subentire, abaxially sericeous on claw and center of limb; lateral petals: 
claw 0.9-1 mm long, limb 5-5.2 mm long, (4-) 4.2-4.8 mm wide; posterior petal: 
claw 1-1.1 mm long, limb 5.5-6 mm long, 4.2-4.5 mm wide. Filaments glabrous 
except for an adaxial basal tuft of hairs, those of stamens opposing sepals 1.4-1.5 
mm long, those of stamens opposing petals 2.1-2.2 mm long; anthers 1.4-1.5 mm 
long, wing of outer locules 0.4 mm wide. Styles ca. 3.5 mm long, 0.3 mm in 
diameter, glabrous. Mature fruit not seen. Chromosome number unknown. 

Pterandra mcphersonii is known only from the type, collected in primary rain 
forest (now disturbed) in Panama (Fig. 1). It is readily separated from the other 
Panamanian species, P. isthmica, by the lack of abaxial pubescensce on the lami- 
nas and stipules. The laminas and stipules of P. isthmica are abaxially densely 
sericeous. Pterandra colombiana, the only other species with abaxially glabrous 



1997 C. ANDERSON: PTERANDRA 19 

leaves, differs in its longer petioles and pedicels, and in having the fascicles sessile 
instead of borne on a short projection. 

This species is named for Gordon D. McPherson, student of Euphorbiaceae 
and discerning collector. 

8. Pterandra isthmica Cuatrecasas & Croat, Ann. Missouri Bot. Gard. 67; 918. 

1981 ["1980"].— Type: Panama. Panama: NE of town of Cerro Azul, 20 
km by rd from Inter- American Hwy, 8 Mar 1975, Mori & Kallunki 5007 
(holotype: MO!; isotypes: MICH! US!). 

Small trees to 4 m. Laminas 4-6.1 cm long, 2-3.5 cm wide, elliptical to obo- 
vate, apex acute (-mucronate) to short-acuminate, acumen up to 0.3 cm long, base 
cuneate, adaxially glabrate to glabrous but with appressed hairs on the major 
veins, abaxially sericeous, the hairs 0.4-1.1 mm long, medifixed, terete, wavy, 
mostly white but a few golden, sessile to subsessile, overlapping, pellucid cells 
only along margin or rarely also in abaxial surface or absent, costa and secondary 
veins prominent abaxially, tertiary veins not raised or sometimes slightly so; peti- 
oles 0.4-1 cm long, densely sericeous; each pair of stipules proximally connate but the 
distal 1/2 free, 2.8-3 mm long, 2.5-2.8 mm wide, adaxially hirsute, abaxially densely 
sericeous. Inflorescence of 4-6-flowered fascicles borne on short projections 1-1.8 
mm long below a flush of new leaves, each fascicle subtended by a deciduous 
bract composed of a pair of stipules and a rudimentary leaf; pedicels 1.6-2 cm 
long, 0.8-1 mm in diameter, densely yellowish to golden-sericeous; bracts 1.6-1.8 
mm long, 0.9-1 mm wide, bracteoles 2-2.2 mm long, ca. 1 mm wide, bracts and 
bracteoles narrowly triangular, the apex acute, abaxially glabrous except for ap- 
pressed hairs on and adjacent to the costa. Sepals 3.5-3.7 mm long, 3 mm wide, 
narrowly triangular (the distal 1/2 ligulate), apex obtuse, slightly recurved, glands 
1.5-1.7 mm long, 0.6-0.7 mm wide. Petals "white" (fide Cuatrecasas and Croat), 
the limb obovate to elliptical, margin erose, abaxially densely sericeous on claw 
and center of limb; lateral petals: claw 1.5-1.8 mm long, limb 6.5-7 mm long, 5.2- 
5.5 mm wide; posterior petal: claw ca. 2 mm long, limb 7-7.5 mm long, 5.5-5.8 mm 
wide. Filaments glabrous except for an adaxial basal tuft of hairs, those of stamens 
opposing sepals ca. 2 mm long, those of stamens opposing petals ca. 3 mm long; 
anthers ca. 1.5 mm long, wing of outer locules 0.4 mm wide. Styles ca. 4 mm long, 
ca. 0.3 mm in diameter, glabrous, inserted at apex. Coccus ca. 3.7 mm high, ca. 4 
mm in diameter, hirsutulose-tomentulose; torus ca. 1.5 mm high; mature seed not 
seen. Chromosome number unknown. 

Pterandra isthmica is known only from the type, collected in premontane rain 
forest in Panama (Fig. 1). The dense abaxial pubescence of the laminas is similar 
to that found in P. sericea, but that species differs in its 3-flowered fascicles, 
smaller petals (the limb 3-4 mm wide), and hirsute filaments. 

9. Pterandra sericea W. R. Anderson, Brittonia 28: 407. 1977 ["1976"].— Type: 

Guyana. Upper Mazaruni River, small island across river from Isla Casa- 
be, 175 m, 9 Dec 1951, Magiure 32715 (holotype: MICH!; isotypes: F! K! 
NY! US!). Fig. 5. 

Shrubs or trees to 15 m. Laminas 3-8.5 cm long, 1.6^.3 cm wide, elliptical to 
obovate, apex acute (-mucronate) to sometimes briefly acuminate, acumen up to 
0.5 cm long, base cuneate, adaxially glabrate to glabrous but with appressed hairs 
on the costa, abaxially densely sericeous, the hairs 0.4-1.2 mm long, medifixed, 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 




terete or rarely some flattened, wavy, white to yellowish, sessile to subsessile, 
overlapping, costa and secondary veins prominent abaxially, tertiary veins not 
raised or sometimes slightly so; petioles 0.3-1.3 cm long, densely sericeous; each 
pair of stipules entirely connate or mostly connate but notched at the apex, 3^ mm 
long, 3-3.8 mm wide, adaxially hirsute but becoming glabrous with maturity, abaxially 
densely sericeous but becoming glabrate to glabrous with maturity, sometimes 



1997 C. ANDERSON: PTERANDRA 21 

only glabrescent in the distal 1/2. Inflorescence of 3-flowered fascicles, each fascicle 
subsessile or borne on short projections to 1.5 mm long in the axil of a new leaf, 
or sometimes each cluster subtended by a deciduous bract composed of a pair of 
stipules and a rudimentary leaf; pedicels 0.9-1.6 cm long, 0.6-0.8 mm in diameter, 
densely golden-sericeous; bracts 2-2.4 mm long, 0.7-0.8 mm wide, bracteoles 1.8- 
3 mm long, 0.4-1 mm wide, bracts and bracteoles linear, the apex acute, abaxially 
densely sericeous. Sepals 2-3 mm long, 2-2.4 mm wide, triangular (the distal 1/2 
sometimes ligulate), apex obtuse, glands 1-1.7 mm long, 0.5-1.2 mm wide, or 
glands absent. Petals white to greenish white (becoming pale yellow in age), the 
limb broadly elliptical to suborbicular, margin erose, abaxially densely sericeous 
on claw and center of limb; lateral petals: claw 1-1.4 mm long, limb 3.5-4.1 mm 
long, 3-4 mm wide; posterior petal: claw 1.4-1.7 mm long, limb 3.8—4.3 mm long, 
3-4 mm wide. Filaments with an adaxial basal tuft of hairs and also adaxially 
hirsute, those of stamens opposing sepals 1.8-2 mm long, those of stamens oppos- 
ing petals 2.4-2.7 mm long; anthers 1-1.3 mm long, wing of outer locules 0.2 mm 
wide. Styles 3-3.2 mm long, ca. 0.2 mm in diameter, glabrous or with a few scat- 
tered hairs at the base. Mature fruit not seen. Chromosome number unknown. 

Phenology. Collected in flower from February through May, and in August, 
October, and December. 

Distribution (Fig. 1). Venezuela (Bolivar) and adjacent Guyana; in forest; 
100-800 m. 

Additional Specimens Examined. Venezuela. Bolivar: Dtto. Piar, aonda de Mayupa, via a la 
Isla Orquidea, 5 May 1979, Benitez de Rojas 2573 (F); Isla de las Orquideas, Rio Carrao, cerca del 
Cerro Auyantepui, 700-800 m, 9 Oct 1984, Bono 4399 (VEN); Ri'o Caura from foot of gorge below 
Salta Para, downstream 8 km, 06°03'N, 65°04'W, 250 m, 14 Aug 1985, Horner et al. 271 (MO); 
Aripao, Rio Caura, aguas abajo del Campamento de EDELCA, 06°19'52"N, 64°31'44"W, 100 m, 20 
Aug 1995, Rosales C. et al. 1596 (MICH); Chimanta Massif, Rio Apacara over igneous rock, 1.4 mi 
downstream from mouth of Rio Abacapd to mouth of Rio Abacapa, W side of Apacara-tepui, 415 m, 
29 Mar 1953, Steyermark 74677 (F, MICH, NY); slopes above Morrison-Knudsen Camp and Pilot 
Plant, 700-750 m, 26 Feb 1953, Wurdack 34405 (F, MICH, NY). 

Pterandra sericea superficially resembles the Panamanian P. isthmica, with 
which it shares a dense abaxial laminar pubescence. It differs from that species in 
its 3-flowered fascicles and smaller petals; P. isthmica has 4-6-flowered fascicles 
and petals with the limb 6.5-7 mm long and 5.2-5.5 mm wide. Pterandra sericea is 
the only species in the genus in which the filaments are adaxially hirsute rather 
than bearing only a basal tuft of hairs. 

10. Pterandra evansii Cuatrecasas, Brittonia 11: 170. 1959.— Type: Brazil. Amazo- 
nas: Rio Marmelos (headwaters), Aug 1948, Schultes & Lopez 10332 (holo- 
type: US!; isotypes: GH! IAN!). 

Shrubs or small trees to 4 m. Laminas 5.6-15 cm long, 2-5.7 cm wide, lan- 
ceolate to narrowly elliptical, apex acute (-mucronate) to acuminate, acumen up 
to 1.8 cm long, base acute, adaxially glabrous but sometimes with appressed hairs 
on the costa, abaxially sericeous, the hairs 0.1-0.4 (-0.6) mm long, medifixed, 
terete, straight, white to yellowish, sessile, slightly or not at all touching or over- 
lapping, often also with tufts of basifixed hairs in the axils formed by the costa and 
secondary veins, and sometimes with basifixed hairs also scattered along the costa, 
pellucid cells present in abaxial surface, costa and secondary veins prominent 
abaxially, tertiary veins not raised or sometimes slightly so; petioles 0.5-1.4 cm 
long, densely sericeous; each pair of stipules entirely connate or nearly so but the 
apex notched or connate in the proximal 3/4 but the distal 1/4 free, 2.8^ mm long, 



22 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

3-3.5 mm wide, adaxially hirsute, abaxially sericeous (the pubescence abraded 
from stipules on old growth). Inflorescence of sessile, 4-5-flowered fascicles be- 
low a flush of new leaves or sometimes the distal new leaves not yet developed at 
anthesis, each fascicle subtended by a deciduous bract composed of a pair of 
stipules and a rudimentary leaf; pedicels 1.2-1.8 (-2.8) cm long, 0.3-0.4 mm in 
diameter, densely white-sericeous; bracts 2-3 mm long, 0.6-1.4 (-1.8) mm wide, 
triangular, bracteoles 1.5-2.5 mm long, 0.3-0.7 (-1.1) mm wide, linear, bracts and 
bracteoles with the apex acute, abaxially sericeous or with hairs concentrated on 
the costa. Sepals 2-2.4 mm long, 1.8-2 mm wide, narrowly triangular (the distal 1/2 
ligulate), apex obtuse, recurved, glands 1.4-1.7 mm long, 0.8-1 mm wide. Petals 
white to cream, the limb elliptical to obovate or suborbicular, margin erose, abaxi- 
ally sericeous on the claw and most of limb except for a marginal band 0.3-0.5 mm 
wide; lateral petals: claw (0.8-) 1-1.2 mm long, hmb 4-5 mm long, 3-3.5 mm wide; 
posterior petal: claw 1.5-1.8 mm long, hmb 5-5.5 mm long, 4-4.5 mm wide. Fila- 
ments glabrous except for an adaxial basal tuft of hairs, those of stamens opposing 
sepals 1.5-1.8 mm long, those of stamens opposing petals 2.7-3 mm long; anthers 
1.1-1.3 mm long, wing of outer locules 0.2 mm wide. Styles 3.2-3.5 mm long, 0.2 
mm in diameter, with a few scattered hairs adaxially in the proximal 1/5. Coccus 
2.8 mm high and in diameter, hirsutulose-tomentulose; well-developed torus not 
seen; mature seed not seen. Chromosome number unknown. 

Phenology. Collected in flower from June through August, in fruit in August. 

Distribution (Fig. 1). Brazil (Amazonas, Rondonia); in "campo natural" and 
forest, along rivers. 

Additional Specimfns Examined. Brazil. Amazonas: Novo Aripuana, BR-230, 150 km ao N de 

(INPA, MG); Novo Aripuana, sub-base Proj. RADAM SC-20-XD-Ponto 22, margen do Rio Preto, 
30 Aug 1975, Cordeiro 704 (MG, MICH); Novo Aripuana, Igarapo Preto, SC-20-XD-Ponto 30, 30 
Jun 1975, da Silva 109 (MG).— Rondonia: track between Mutumparana and Rio Madeira, 5 Jul 1968, 
Prance et al. 5554 (INPA, MG, MICH, MO, NY, VEN). 

Pterandra evansii is very similar to P. hlrsula but differs most notably in the 
abaxial laminar pubescence. In P. hirsuta that vesture is composed of erect basi- 
fixed hairs mixed with a few medifixed hairs. In P. evansii the laminas are abaxially 
sparsely sericeous, i.e., the hairs are appressed and medifixed; however, they usu- 
ally also have tufts of basifixed hairs in the axils formed by the costa and second- 
ary veins and sometimes also along the costa. Cid Ferreira 5596 from Amazonas, 
Brazil, is a mixed collection of both species. 

11. Pterandra hirsuta C. Anderson, sp. nov.— Type: Bolivia. Beni: Prov. Vaca 
Diez, 19.4 km al N de Guayaramerin camino a Cachuela Esperanza, 
10°32'S, 65°38'W, 140 m, 18 Sep 1994, Moraes M. & Takana G. 1857 
(holotype:MICH!). Fig. 4j-m. 

Arbor vel frutex usque ad 12 m altus. Laminae 5-12.5 cm longae, 2.5-6 cm 
latae, ellipticae, supra glabrae vel costa sparsissime sericeae, subtus pilos erectos 
et paucos medifixos ferentes; petioli 0.5-1.2 cm longi, dense sericei; stipulae 4-7 
mm longae, 4-5 mm latae, connatae sed distaliter liberae, adaxiahter hirsutae, 
abaxialiter sericeae. Inflorescentia ex fasciculis sessilibus constans, floribus cu- 
jusque fascicuh 4-6, sine foliis juvenilibus per anthesin; pedicelli (0.8-) 1.2-2.3 cm 
longi, 0.4-0.5 mm diametro, dense albo-sericei. Petala alba vel cremea, hmbo 



1997 C. ANDERSON: PTERANDRA 23 

elliptico vel late obovato; limbi pelalorum lateralium 4.2-5 mm longi, 3-4 mm lati, 
limbus petali postici 5-5.5 mm longus, 4-5 mm latus. Filamenta praeter caespitem 
basalem glabra. Styli 3-4.3 mm longi. Cocci maturi ignoti. 

Shrubs or trees to 12 m. Laminas 5-12.5 cm long, 2.5-6 cm wide, elliptical, 
apex acute (-mucronate) to obtuse, base rounded to cuneate, adaxially glabrous 
or with a few appressed hairs on the costa, abaxially hirsute with erect basifixed 
hairs and also some medifixed hairs, all terete, white to golden, basifixed hairs 
0.1-0.7 mm long, medifixed hairs 0.4-1.1 mm long, subsessile or with a stalk up to 
0.1 mm long, pellucid cells abundant to very few in abaxial surface or mostly along 
margin, costa and secondary veins prominent abaxially, tertiary veins usually slightly 
raised as well; petioles 0.5-1.2 cm long, densely sericeous; each pair of stipules 
entirely connate or nearly so but the apex notched or the distal 1/4 free, 4-7 mm 
long, 4-5 mm wide, adaxially hirsute, abaxially sericeous. Inflorescence of sessile, 
4-6-flowered fascicles, each fascicle subtended by a deciduous bract composed of 
a pair of stipules and a rudimentary leaf, distal new leaves not yet developed at 
anthesis; pedicels (0.8-) 1.2-2.3 cm long, 0.4-0.5 mm in diameter, densely white- 
sericeous; bracts 2-4 mm long, 1-2 mm wide, triangular, bracteoles 1.4-3 mm long, 
0.5-0.8 (-1.2) mm wide, narrowly triangular to linear, bracts and bracteoles with 
the apex acute, sericeous on the costa and adjacent region but glabrous along the 
margin. Sepals 2.3-3.3 mm long, (1.7-) 2-2.4 mm wide, triangular, apex obtuse or 
acute, recurved to revolute, glands 1.3-1.8 mm long, 0.7-1 mm wide. Petals white 
to cream, the limb elliptical to broadly obovate, margin erose, abaxially densely 
sericeous on claw and most of limb except for a marginal band ca. 0.5 mm wide; 
lateral petals: claw 1.2-1.4 mm long, limb 4.2-5 mm long, 3-4 mm wide; posterior 
petal: claw 1.2-1.5 mm long, limb 5-5.5 mm long, 4-5 mm wide. Filaments gla- 
brous except for an adaxial basal tuft of hairs, those of stamens opposing sepals 
1.8-2.6 mm long, those of stamens opposing petals 3-3.4 mm long; anthers 1-1.3 
mm long, wing of outer locules 0.2-0.3 (-0.4) mm wide. Styles 3-4.3 mm long, ca. 
0.3 mm in diameter, with scattered hairs adaxially in the proximal 2/3-3/4. Mature 
fruit not seen. Chromosome number unknown. 

Phenology. Collected in flower in September, in young fruit in March. 

Distribution (Fig. 1). Brazil (Amazonas, Rondonia) and adjacent Bolivia 
(Beni); in forest at riverside and in "pampa-monte"; 140-220 m. 



ao L de Humaita e 30 km para o S na rod. do Estanho, 08°20'S, 6r45'W. 16 Apr 1985, Cid Ferreira 
5596 p.p. (MG, MICH).— Rondonia: along Rio dos Pacaas Novos, just above and below the first 
cachoeira, ca. 220 m, 28 Mar 1978, Anderson 12298 (INPA, MICH). 

Pteranclra hirsuta differs from all other species in that the abaxial laminar 
pubescence is composed almost entirely of basifixed hairs. In other aspects, it is 
most similar to P. evansii, which has the laminas abaxially sericeous but usually 
with tufts of basifixed hairs in the axils of the secondary veins at the costa and also 
along the costa. 

12. Pterandra flavescens Maguire, Mem. New York Bot. Card. 8: 128. 1953.— 
Type: Venezuela. Amazonas: Cerro Sipapo (Paraque), banks of Lower 
Cano Negro, 1500 m, 25 Dec 1948, Maguire & Politi 28104 (holotype: 
NY!; isotypes: BM, F! RB! US! VEN). 



24 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Shrub or small tree to 10 m. Laminas 5-9.8 cm long, 2-5 cm wide, elliptical to 
slightly obovate, apex obtuse-mucronate, base cuneate, adaxially glabrate but dense- 
ly pubescent on the costa, abaxially very sparsely sericeous but densely sericeous 
on the costa, the hairs 0.1-0.7 mm long, medifixed, mostly terete but some flat- 
tened, straight, ferrugineous (or fading to yellowish in age), sessile, widely spaced 
and never touching, pellucid cells abundant to sparse in abaxial surface or some- 
times absent, costa prominent but the secondary veins only very slightly if at all 
raised; petioles 1.5-2.7 cm long, densely sericeous; each pair of stipules entirely 
connate or nearly so but the apex notched or the distal 1/3 free, 3.5-4.5 mm long 
and wide, adaxially hirsute, abaxially densely sericeous. Inflorescence of subsessile, 
3-flowered fascicles, each fascicle borne in the axil of a new leaf; pedicels 1.5-2.7 
cm long, 0.6-0.8 mm in diameter, densely ferrugineous-sericeous; bracts 1.7-2.7 
mm long, 0.7-1.3 mm wide, narrowly triangular, bracteoles 1.8-3 mm long, 0.6-0.9 
mm wide, linear, bracts and bracteoles with the apex acute, abaxially densely 
sericeous. Sepals 2-2.8 mm long, 2.3-3 mm wide, narrowly triangular or the distal 
1/2 ligulate, apex obtuse, erect or shghtly recurved, glands 1.7-2.5 mm long, 1-1.4 
mm wide. Petals cream or pale yellow (probably in age), the limb broadly obovate 
to suborbicular, margin erose, abaxially densely sericeous on claw and center of 
hmb; lateral petals: claw 1-1.2 (-1.5) mm long, limb 4.5-5 mm long, 3.7-4.5 mm 
wide; posterior petal: claw 1-1.5 mm long, limb 5-5.2 mm long, 4-5 mm wide. 
Filaments glabrous except for an adaxial basal tuft of hairs, those of stamens 
opposing sepals 2.2-2.5 mm long, those of stamens opposing petals 2.5-3 mm 
long; anthers 1.1-1.4 mm long, wing of outer locules 0.4 mm wide. Styles 2.8-3.2 
mm long, 0.3-0.4 mm in diameter, glabrous. Coccus ca. 4.5 mm high, 3.8-3.9 mm 
in diameter, hirsutulose-tomentulose; torus ca. 1.5 mm high; immature embryo 
with the inner cotyledon folded four times and enveloped by the outer cotyledon 
(folded at 1/2 its length and the distal tip folded over the tip of the inner cotyle- 
don). Chromosome number unknown. 

Phenology. Collected in flower and young fruit in December. 

Distribution (Fig. 1). Venezuela, Amazonas, along Rio Caiio on the Cerro 
Sipapo; in savanna and marshes along the river; 1500 m. 



Pterandra flavescens has very distinctive leaves. Abaxially only the costa is 
prominent, and the secondary veins are only very slightly if at all raised. In all 
other species, the costa as well as the secondary veins are prominent. The laminas 
are abaxially very sparsely flecked with ferrugineous hairs (fading in age). The 
flowers are grouped into 3-flowered clusters, each borne in the axil of a new leaf. 
In most other species the flowers number 4-6 per cluster and are usually borne in 
the axil of a deciduous bract composed of a pair of stipules and a rudimentary 



13. Pterandra ultramonfana Riley ex Cuatrecasas, Webbia 13: 557. 1958.— Type: 
Colombia. Narino: Gorgona Island, 20 Nov 1924, Collenette 707 (holo- 
type: K!; isotypes: F! NY! US!). 



1997 C. ANDERSON: PTERANDRA 25 

Tree to 25 m. Laminas 6.5-19 cm long, 3-6.7 cm wide, elliptical or narrowly so 
to obovate, apex acute-mucronate to acuminate, acumen up to 1.5 cm long or 
sometimes obtuse-mucronate, base cuneate, adaxially glabrous or glabrate, abaxi- 
ally sericeous, the hairs 0.1-0.2 (-0.4) mm long, medifixed, flattened and scalelike, 
straight, ferrugineous (sometimes faded to yellowish or white), sessile, rarely touch- 
ing or overlapping, costa and secondary veins prominent abaxially, tertiary veins 
slightly raised; petioles 1-2 cm long, sericeous; each pair of stipules entirely con- 
nate or the apex notched or connate in the proximal 1/3-1/2 and distally free, 4.5- 
6 mm long, 4-5 mm wide, adaxially hirsute, abaxially sericeous but sparsely so in 
age. Inflorescence of sessile 4-6-flowered fascicles, each fascicle subtended by a 
deciduous bract composed of a pair of stipules and a rudimentary leaf, the lowest- 
most cluster at a node above 1-2 pairs of mature leaves (or these already lost), 
distal new leaves not yet developed at anthesis; pedicels 1.2-2.8 cm long, 0.4-0.6 
mm in diameter, densely ferrugineous-sericeous; bracts 1.8-2.7 mm long, 1-1.8 
mm wide, triangular, bracteoles 1.1-2.2 mm long, 0.2-0.8 mm wide, narrowly tri- 
angular to linear, bracts and bracteoles with the apex obtuse to acute, abaxially 
glabrous or pubescent on the costa. Sepals 2-2.5 mm long and wide, narrowly 
triangular (the distal 1/2-3/4 ligulate), apex obtuse, recurved to revolute, glands 
1.4-2.3 mm long, 1-1.2 mm wide. Petals white to cream, the limb broadly elliptical 
or oblong to suborbicular, margin subentire to slightly erose, abaxially sericeous 
or sparsely so on claw and center of limb; lateral petals: claw 0.8-1.2 mm long, 
limb (2.5-) 3-3.5 mm long, 2.5-3 mm wide; posterior petal: claw 1-1.2 mm long, 
hmb 3.2-4 mm long, 2.8-3 mm wide. Filaments glabrous except for a sparse adaxial 
basal tuft of hairs, those of stamens opposing sepals 1.1-1.6 mm long, those of 
stamens opposing petals 2-2.3 mm long; anthers 1-1.2 mm long, wing of outer 
locules 0.3 mm wide. Styles 3.2-3.5 mm long, 0.3 mm in diameter, glabrous. Coc- 
cus ca. 2.8 mm high and in diameter, hirsutulose-tomentulose; torus up to 1 mm 
high (?); mature seed not seen. Chromosome number unknown. 

Phenology. Collected in flower February through April and in September; 
date of immature fruiting collection unknown. 

Distribution (Fig. 1). Colombia (Valle and Gorgona Island, Narifio); transi- 
tion between tropical wet forest and pluvial forest; 50-100 m. 

Additional Specimens Examined. Colombia. Narino: Gorgona Island, 15 Oct 1924, Collenette 
589 {F, NY).— Valle: Bajo Calima, 10-20 m, 9 Apr 1961, Cabrera R. 554 (F, MICH); Bajo Calima 
Concession, ca. 1.5 km from end of Gasolina Rd, in Juanchaco area, BV-82, ca 16 km NW of 
Buenaventura, 03°50'N, 77°10'W, 50 m, 9 Jun 1987, Faber-Langendoen et al 818 (MO); Bajo Calima, 
concession ca. 20 km N of Buenaventura, 03°40'N, 77°00'W, 50 m, 4 Jul 1987, Faber-Langendoen 
1197 (MICH); Bajo Colima, ca. 1.5 km NW of Buenaventura, 03°59'N, 77°05'W, 50 m, 12 Apr 1987, 
Gentry et al. 56807 (MO); Bajo Calima, Lijal-Gasolina rd bifurcation, 03°58'N, 77°W, 50 m, 10 Jun 
1988, Gentry 62804 (MICH); Bajo Calima, ca. 15 km N of Buenaventura, 03°56'N, 77°08'W, 18 Feb 
1983, Gentry & Juncosa 40472 (MICH, MO); Bajo Calima, ca. 10 km due N of Buenaventura, 
03°56'N, 77°08'W, ca. 50 m, 1982, Mazuera 34 (MO); Bajo Calima, Concesion Pulpapel, Buenaventura, 
03°55'N, 77°W, 100 m, 27 Mar 1985, Monsalve B. 825 (MICH, MO). 

Pterandra ultramontana is distinguished by the unique abaxial pubescence of 
the laminas, composed of ferrugineous, flattened and scalelike hairs 0.1-0.2 (-0.4) 
mm long. In older laminas the color sometimes fades to yellow, but this may also 
be a result of drying procedures. Pterandra ultramontana has been confused with 
P. colombiana, but in that species the abaxial surface of the laminas and stipules is 
glabrous. 



26 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

14. Pterandra egleri W. R. Anderson, Contr. Univ. Michigan Herb. 19: 386. 1993.— 
Type: Brazil. Para, Alto Tapajos, Rio Cururu, Erereri, 25 Jul 1959, Egler 
1033 (holotype: MG!; isotypes: HB! IAN! MICH! NY! R!). 

Shrubs or small trees to 5 m. Laminas 4.5-9.4 cm long, 2-4.4 cm wide, ellipti- 
cal or narrowly so, apex acute (-mucronate), base rounded, adaxially glabrous, 
abaxially tomentose but becoming glabrate to glabrous in age, the vesture shed in 
patches, the hairs 0.5-1.4 mm long, medifixed, terete, wavy and crisped, golden to 
ferrugineous, subsessile or with a stalk up to 0.2 mm long, pellucid cells only along 
margin or rarely also in abaxial surface or absent, costa and secondary and tertiary 
veins prominent abaxially; petioles 0.5-1.1 cm long, densely sericeous; each pair 
of stipules entirely connate or sometimes notched at the apex, (2.5-) 3-4.5 mm 
long, 3-4 mm wide, adaxially hirsute, abaxially densely sericeous (the pubescence 
abraded from stipules on old growth). Inflorescence of sessile, (3-) 4-6-flowered 
fascicles below a flush of new leaves or sometimes the distal new leaves not yet 
developed at anthesis, each cluster subtended by a deciduous bract composed of a 
pair of stipules and a rudimentary leaf, sometimes the lowestmost cluster at a 
node above a pair of mature leaves; pedicels 1.1-1.7 cm long, 0.5-0.6 mm in 
diameter, densely yellowish to white-sericeous; bracts 1.5-1.8 mm long, 0.7-1 mm 
wide, narrowly triangular, bracteoles 1.2-2 mm long, 0.3-0.5 mm wide, linear, 
bracts and bracteoles with the apex acute, abaxially sericeous. Sepals 1.7-3.2 mm 
long, 1.2-2.2 mm wide, triangular or narrowly so (the distal 1/2 sometimes ligu- 
late), apex obtuse, slightly recurved to revolute, glands 0.8-1.3 mm long, 0.5-0.8 
mm wide; sometimes with only 1 gland per sepal. Petals white (becoming yellow 
in age), the limb orbicular or suborbicular, margin erose, abaxially densely seri- 
ceous on claw and on limb in the proximal 2/3 except for marginal band ca. 0.5 mm 
wide, the distal 1/3 glabrous; lateral petals: claw 0.7-1 mm long, limb 3.3-5 mm 
long, 3.5-4.5 mm wide; posterior petal: claw 0.8-1.2 mm long, limb 4-5.2 mm long, 
3.5-5 mm wide. Filaments glabrous except for an adaxial basal tuft of hairs, those 
of stamens opposing sepals 1.7-2 mm long, those of stamens opposing petals 2.7-3 
mm long; anthers 0.3 mm long, wing of outer locules 0.3 mm wide. Styles 3.5-4.5 
(-4.7) mm long, 0.2-0.3 mm in diameter, with a few scattered hairs adaxially in the 
proximal 1/4-1/3. Coccus 2.6-3 mm high and in diameter, hirsutulose-tomentu- 
lose; torus ca. 1 mm high; embryo globose, ca. 2.3 mm in diameter, radicle 1.2 mm 
long, outer cotyledon ca. 3.7 mm long, ca. 1.5 mm long, folded at 1/2 of its length 
and the tip folded over the inner cotyledon, inner cotyledon ca. 3 mm long, ca. 1.5 
mm wide, folded twice. Chromosome number: n = 12 (based on Anderson 10895- 
W.R.Anderson 1993). 

Phenology. Collected in flower and fruit in February, July, and October. 

Distribution (Fig. 1). Brazil (Para); in campo; ca. 200 m. 

Additional Specimens Examined. Brazil. Para. Region of Missao Velha, ca. 2 km N of Rio 
Cururu, 07°45'S, 57°20'W, ca. 200 m, 13 Feb 1974, Anderson 10895 (IAN, MICH. NY); Bara de S. 
Manoel, Rio Tapajos, 12 Oct 1973, Ribeiro 348 (MICH). 

Pterandra egleri is readily recognized by the tomentose abaxial laminar vesture, 
which is shed patchily in age. In all other species, except P. hirsuta and P. pyroi- 
dea, the abaxial pubescence is appressed. Pterandra pyroidea, of Goias and Minas 
Gerais, is also similar in that the tertiary veins of the lamina are prominent abaxi- 
ally; it is most readily separated by its larger flowers with pink petals. Pterandra 



nn. Sci. Nat. Bot., 


ser. 2, 13: 328. 1840. 


Dbach in Martius 


, Fl. bras. 12(1): 29. 



: PTERANDRA 27 

hirsiita differs from all other species in that the abaxial laminar pubescence con- 
sists of erect basifixed hairs mixed with some medifixed hairs. In P. egleri all hairs 
composing the leaf pubescence are medifixed. 



Pterandra latifolia Adr. Jussieu in Delessen 
Acmanthera latifolia (Adr. Jussieu) 
29. 1858. 

Pterandra opulifolia Rusby, Dcscr 
(Rusby) Niedenzu in Engle: 



ACKNOWLEDGMENTS 

I thank William R. Anderson for his comments and for sharing his insights on Pterandra. Kc 
Douthit drew Figs. 3 and 4, and Annette Seidenschnur Mahler drew Fig. 5. Figure 5 is reprinted v 
permission from Memoirs of the New York Botanical Garden 32: 34, Fig. 8, copyright 1981, The > 
York Botanical Garden. I am grateful to the curators of the following herbaria for permitting m< 



LITERATURE CITED 

;on. W. R. 1975. The taxonomy oi Acmanthera (Malpighiaceac). Contr. Univ. Michigan Hei 

. 1978 ["1977"]. Byrsonimoideae, a new subfamily of the Malpighiaceac. Leandra 7: 5-18. 

. 1981. Malpighiaceae. In "The botany of the Guayana Highland— Part XI" by Bassett Magu: 

id collaborators. Mem. New York Bot. Gard. 32: 21-305. 

. 1983. Lophanthera, a genus of Malpighiaceae new to Central America. Brittonia 35: 37-41. 

. 1993. Chromosome numbers of neotropical Malpighiaceae. Contr. Univ. Michigan Herb. ; 

11-354. 

casas, J. 1958. Prima Flora Colombiana. 2. Malpighiaceae. Webbia 13: 343-664. 

ach, A. 1858. Malpighiaceae. In Flora brasiliensis by K. F. P. von Martius, 12(1): 1-123. 

I, Adr. de. 1833 ["1832"]. Malpighiaceae. In Flora brasiliensis meridionalis by A. de Sail 



EXCENTRADENIA, A NEW GENUS OF MALPIGHIACEAE 
FROM SOUTH AMERICA 

William R. Anderson 
University of Michigan Herbarium 

North University Building 
Ann Arbor, Michigan 48109-1057 

When I described Hiraea propinqua in 1994 I discussed the problems created 
in Hiraea by the inclusion of H. adenophora Sandwith and its relatives, but hesi- 
tated to segregate that complex as a distinct genus (Anderson 1994, pp. 132-133). 
Now I have come to the conclusion that segregation is inevitable and desirable, so 
that Hiraea can return to the homogeneity that distinguished it before these spe- 
cies were described. 

-Type: Excentradenia adenophora (Sand- 



Lianae lignosae. Lamina foliorum eglandulosa vel margine glandulis parvis 
instructa, venis tertiariis scalariformibus; petiolus biglandulosus; stipulae parvae, 
triangulares, basi petioli vel caule juxta petiolum portatae, vel nullae. Inflorescen- 
tia axillaris racemus 3-7 (-9) umbellarum 4-florarum; bracteae floriferae eglandu- 
losae; pedunculus brevis vel nullus; 1 bracteola cujusque paris eglandulosa, altera 
uniglandulifera glandulo versus centrum umbellae excentrico; pedicellus in ala- 
bastro circinatus. Sepala triangularia vel ovata, apice acuta revolutaque. Petala 
lutea, glabra. Stamina 10, omnia fertiha. Styh 3, stigmate introrso, apice dorsahter 
truncati, apiculati, vel breviuncinati. Samarae ala lateralis membranacea, plerum- 
que subcircularis, apice usque ad nucem incisa, basi continua vel interdum usque 
ad nucem incisa, nuce subglobosa. 

Woody vines. Leaves opposite, subopposite, or alternate; petiole biglandular 
at or above middle, the lamina eglandular or bearing small glands on margin, the 
tertiary veins strongly parallel (scalariform); stipules small, triangular, borne on 
very base of petiole or on adjacent stem, or absent. Inflorescence a single axillary 
raceme of 3-7 (-9) 4-flowered umbels, with 1 umbel terminal and the other 1-4 
pairs axillary to bracts bearing stipules and often petiole glands; floriferous bracts 
small, persistent, eglandular; floriferous peduncle short or absent; bracteoles small, 
persistent, one of each pair bearing 1 bulging eccentric abaxial gland toward 
center of umbel, the gland 0.7-1.1 mm long, circular or elliptical in outline; pedicel 
well developed, circinate in bud as far as known. Sepals 5, triangular or ovate, 
acute at apex and revolute in anthesis, all eglandular or the lateral 4 abaxially 
biglandular, the glands broadly elliptical to orbicular. Corolla bilaterally symmet- 
rical, the posterior petal strongly differentiated from the lateral 4; petals yellow, 
glabrous. Receptacle glabrous. Androecium bilaterally symmetrical; stamens 10, 
all fertile, glabrous; filaments briefly connate at base; anthers ± alike. Gynoecium 
bilaterally symmetrical, the anterior style shorter than the posterior styles; ovary 
with the 3 carpels nearly distinct, all fertile; styles inserted low on ventral face of 
carpels, the apex with a large internal stigma and dorsally truncate, apiculate, or 

29 



30 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

bearing a hook up to 0.3 mm long. Fruit dry, breaking apart into 3 samaras 
separating from a short pyramidal torus; samara with a large, membranous, usually 
subcircular lateral wing borne on upper edge of nut, continuous at base and incised to 
nut at apex, or (in E. adenophora) sometimes incised to nut at both apex and base; 
dorsal wing small; intermediate winglets absent; nut subglobose; ventral areole circu- 
lar, 1.5-2 mm in diameter, partially surrounded (on the sides but not around the base) 
by an irregular callose thickening 0.5-1 mm thick; carpophore absent. 

Excentradenia is probably a close sister to the genus Hiraea Jacquin. They are 
linked at least by scalariform tertiary veins in the leaf, lamina glands borne only on 
the margin, flowers borne ultimately in umbels, nearly distinct carpels, a subglobose 
nut in the samara, and a callose thickening along the sides of the samara's circulai^ 
ventral areole; having a glandular-fimbriate posterior petal may eventually prove to 
be another feature of their common ancestor. Until the phylogeny of the genera with 
mascagnioid samaras can be resolved, it is impossible to say which of these char- 
acter-states are synapomorphies and which are symplesiomorphies. The differences 
between Hiraea and Excentradenia are summarized in the following couplet: 

1. Stipules very small, triangular, borne on petiole at very base or on adjacent stem, or absent; 
illary raceme of 3-7 (-9) 4-flowered umbels; one of each pair of bracteoles 
large bulging eccentric abaxial gland toward center of umbel; pedicel circinate in 
al wing of samara incised to nut at apex but not at base, producing a single basally 
s wing, or (in E. adenophora) sometimes incised to nut at both apex and base. 

longated, borne on petiole at least somewhat above base; inflorescence an axillary 
-7 4-flowered umbels or a single umbel of 5-many flowers; bracteoles eglandular; 
-aight in bud; lateral wing of samara incised to nut at both apex and base, producing 



; bracteole glands found in all species of Excentradenia, to which 
the name refers, are most likely a synapomorphy of those species, as are the 
elongated epipetiolar stipules of Hiraea, so it seems probable that both genera are 
monophyletic in the strictest sense. The difference between their inflorescence 
branching deserves further comment. In those specimens of Excentradenia in which 
there are more than three umbels, they are arranged as two to four successive 
pairs originating from the same axis, followed by a terminating umbel. In Hiraea 
this never happens. If the inflorescence is branched, it is strictly cymose — it termi- 
nates in an umbel, and the two lateral branches are axillary to a single pair of 
bracts on the stalk below. Those branches terminate in an umbel, and may be 
subtended by another pair of lateral branches. Thus, as far as I can tell from the 
imperfect material now available, there seems to be a fundamental difference. I 
do not know which branching pattern is ancestral in the clade containing both 
Excentradenia and Hiraea. 

Excentradenia is a genus of northern South America (Fig. 1). Three of the 
species (E. adenophora, E. boliviana, and E. propinqua) form a tight complex, 
and the fact that one of them is disjunct across the Amazon from the others is 
intriguing. The fourth species, E. primaeva, is both morphologically and geo- 
graphically quite isolated. 

Excentradenia is a nice example of how poorly lianas are known. For the four 
species I am recognizing here, we have a total of only 12 collections, an average of 
three per species. The only conclusion I can draw from such a number is that an 
incredible wealth of wonderful plants is still out there in tropical America waiting 
to be discovered — if those forests are spared for another generation or two. 



ANDERSON: EXCENTRADENIA 






Key to the S 


PECIES OF EXCENTRADENIA 








_.eaves abaxially pers 


sternly velutinous 


the hairs erect and bifurcate. 




E. prin 


aeva. 


.eaves abaxially seric 


sous to glabrate, t 


le hairs sessile and tightly appressed. 








. Leaves abaxially densely and persiste 


ntly sericeous, even in age. 




E.prop 


nqua. 


. Leaves glabrate at 












d otherwise only s 


Darse, scattered hairs persistent on the 


lam 


na. 




3. Calyx bearing 8 


glands on the 4 la 


teral sepals; small stipules present on 


base 


of petiole 




lamina of large 


leaves 8-15.5 cm 


long, the petiole 10-20 mm long; la 


eral 


wing(s) o 












E. adenophora. 


3. Calyx complete 


y eglandular; stip 


ules absent from vegetative leaves; 1 


min 


a of large 






long, the petiole 20-29 mm long; lateral wmg of samar 


w, 


h the side 





meeting or overlapping at apex, leaving little or no evident gap. E. boliviana 

Excentradenia adenophora (Sandwith) W. R. Anderson, comb. nov. Hiraea adeno- 
phora Sandwith, Kew Bull. 1951: 33. 1951.— Type: Guyana. Dukalikuru Creek, 
Berbice River, 31 Mar 1938 fl, Fanshawe in Forest Dept. 2682 (holotype: K! 
photos MICH, WRA negs. 81-5-17 [sheet I] & 18 [sheet II]). Fig. 2 



Stems densely and persistently sericeous. Leaves opposite, subopposite, or 
alternate; lamina of larger leaves 8-15.5 cm long, 4.5-10.3 cm wide, ovate or 
elliptical to orbicular, rounded or truncate at base, eglandular or bearing a few 
tiny buttonlike glands along margin, obtuse or rounded and abruptly short-acumi- 
nate at apex, adaxially glabrate at maturity with the midrib usually persistently 
sericeous, abaxially persistently sericeous on midrib and major lateral veins and 
otherwise nearly to quite glabrate, the lateral veins 7-10 pairs and connected by 
parallel cross-veins 3-6 mm apart, the veins and reticulum prominent below and 



CONTR. UNI VERS n 




l.nl o[ cidaxidl suifcice ol Idiniiid. minus hairs xO 5 d) detail of abaxial 

liic glands on 4 bracteolcs, x7 5: g) samara abaxial view xO S h-n, £. 
\ial view xO 'i and base of petiole with stipules x2 5 i) flowering 
latc buds x2 S k) base of umbel enlarged to show eccentiic glands on 









1997 ANDERSON: EXCENTRADENIA 33 

visible above in dried leaves; petiole 10-20 mm long, 1.2-2 mm in diameter, densely 
and persistently sericeous, bearing between middle and apex a pair of glands 1.4-2 
mm long; stipules 0.4-0.8 mm long, triangular, borne on petiole at very base. 
Inflorescence densely and persistently sericeous, 2-5 cm long, comprising (3-) 5-7 
(-9) umbels, each borne on a stalk 5-10 mm long; floriferous bracts 2-2.5 mm 
long, triangular or ovate, abaxially sericeous, adaxially glabrous; floriferous pe- 
duncle 1-2.5 mm long; bracteoles 1-1.5 mm long, ovate or triangular, abaxially 
sericeous, adaxially glabrous, borne at apex of peduncle; pedicel 7-10 mm long, 
0.5-1 mm in diameter, up to 2.5 mm in diameter at apex, sericeous. Sepals 3.5-4 
mm long, 2-2.5 mm wide, abaxially sericeous, adaxially glabrous, the anterior 
eglandular, the lateral 4 biglandular with the glands 1.5-2.2 mm long. Lateral 4 
petals with the claw 2.5-3 mm long, the limb 4-5 mm long and wide, dentate to 
laciniate, eglandular; posterior petal with the claw 4.5 mm long, the limb 4.5 mm 
long, 5 mm wide, fimbriate with the long slender divisions somewhat glandular- 
thickened at apex. Stamens with filaments opposite sepals 3-3.5 mm long, erect 
and straight or curved distally toward posterior petal, filaments opposite petals 2- 
2.5 mm long, erect, straight; anthers 0.7-0.9 mm long, the connective yellow. 
Ovary sericeous; styles bowed outward (i.e., curved outward from base and then 
back toward center of flower), sparsely sericeous in proximal half, dorsally apicu- 
late at apex with the projection 0.1-0.2 mm long, the anterior style 3.4-3.7 mm 
long, the 2 posterior styles 4.5 mm long. Samara depressed-circular with the nut 
positioned below the center, 57-66 mm wide, ca 50 mm high, the lateral wing 
continuous at base and incised to nut at apex or butterfly-shaped with the lateral 
wings separate (incised to nut at both apex and base), each lateral wing 25 mm 
wide and 30 mm high; sides of lateral wing(s) not meeting at apex, leaving an 
evident gap, sinuous at margin, thinly sericeous with very fine white appressed 
hairs; dorsal wing 2-3 mm wide, 5-6 mm high, coarsely dentate; nut ca 4 mm in 
diameter, finely sericeous. 

Additional Specimens ExAMrwEo: Guyana. Bartica-Potaro Road, 51st mile, wallaba bush on 
white sand, fl [without date], Dawson in Forest Dept. 2010 (K); Groete Creek, Essequibo River, 
mixed "ropy" forest on lateritic soil, Mar fr, Fanshawe in Forest Dept. 4485 (K); Barabara Creek, 
Mazaruni River, Mar fl, Fanshawe in Forest Dept. 5604 (K, NY, US). Venezuela. Bolivar: 20-35 
km SW of El Manteco, road to San Pedro de las Dos Bocas, 7°10'N, 62°55'W, 200 m, disturbed 
primary forest, Aug fl buds, Liesner & Gonzalez 5954 (MO, VEN).— Delta Amacuro: E side of Rio 
Cuyubini, Cerro La Paloma, Sierra Imataca, vicinity of large granitic boulders, 100-200 m, Nov fr, 
Steyerrnaric 87644 (MICH, NY, VEN). 

This was the first species of the genus to be described, and it is the best 
known, with a total of six collections from northern Guyana and adjacent Venezu- 
ela (Fig. 1). It is known from forests at low elevations (to 200 m), and has been 
collected with flowers and fruits in March, with flower buds in August, and with 

1 have seen only two collections with fruits. One of them, Steyermark 87644, 
has subcircular samaras with the lateral wing continuous at the base, like the 
other species of Excentradenia. In the other, Fanshawe in Forest Dept. 4485, the 
samaras, although not well preserved, seem to show clearly that there are two 
separate lateral wings, as in Hiraea. Only the accumulation of additional collec- 
tions of E. adenophora in fruit will permit assessment of the variation in this 
important character in this species. 



34 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Excentradenia boliviana W. R. Anderson, sp. nov.— Type: Bolivia. Beni: Cachue- 
la Esperanza, Rio Beni, Yuta road, Jan 1924 fr, Meyer 381 (holotype: 
MICH!;isotype:NY!). 

Lamina foliorum majorum 14-19 cm longa, 7-10.4 cm lata, ovata ellipticave, 
margine glandulis parvis instructa, utrinque glabrata praeter costam et nervos 
lateraies; petiolus 20-29 mm longus; stipulae non visae in foliis vegetativis. Sepala 
omnia eglandulosa. Samara 40-60 mm lata, 35-50 mm alta; ala lateralis basi con- 
tinua, lobis supra plus minusve superpositus. 

Stems densely and persistently sericeous. Leaves opposite or subopposite; 
lamina of larger leaves 14-19 cm long, 7-10.4 cm wide, ovate or elliptical, cuneate 
to rounded at base, bearing a series of tiny impressed buttonlike glands along 
margin, abruptly short-acuminate at apex, adaxially glabrate at maturity with the 
midrib sometimes persistently sericeous, abaxially persistently sericeous on mid- 
rib and sometimes on major lateral veins and otherwise nearly to quite glabrate, 
the lateral veins 9-11 pairs and connected by parallel cross-veins 3-7 mm apart, 
the veins prominent below and impressed above in dried leaves, the reticulum 
visible on both sides; petiole 20-29 mm long, 1.5-2.5 mm in diameter, densely and 
persistently sericeous, bearing between middle and apex a pair of glands 1.7-2.8 
(-4) mm long; stipules not found on vegetative leaves, but present as tiny triangles 
on bracts of inflorescence axis. Inflorescence densely and persistently sericeous, 
1.5-2.5 cm long, comprising 3-7 umbels, each borne on a stalk 1-7 mm long; 
floriferous bracts 1.5-2 mm long, triangular or ovate, abaxially sericeous, adaxially 
glabrous; floriferous peduncle 1-1.5 mm long; bracteoles 1-1.5 mm long, ovate or 
triangular, abaxially sericeous, adaxially glabrous, borne at apex of peduncle; 
pedicel 14-15 mm long in fruit, 0.5-0.8 mm in diameter, up to 2 mm in diameter at 
apex, sericeous. Flowers not seen. Sepals 4.5 mm long and 3 mm wide in fruit, 
abaxially sericeous, adaxially glabrous, all eglandular. Samara depressed-circular 
with the nut positioned below the center, 40-60 mm wide, 35-50 mm high; lateral 
wing continuous at base, incised to nut at apex with the two sides meeting or 
overlapping, leaving little or no evident gap, sinuous at margin, thinly sericeous 
with very fine white appressed hairs; dorsal wing 5-8 mm wide, 5-12 mm high, 
coarsely dentate; nut 5 mm in diameter, finely s 



This species is known only from the fruiting specimens cited, which were 
collected by G. Meyer two months apart at the same location in Bolivia (Fig. 1); 
the species is to be expected in adjacent Brazil. The epithet refers, of course, to 
the country of origin, which is of interest because this is the only species of 
Excentradenia known from southern Amazonia. Excentradenia boliviana is very 
similar to E. adenophora in the characters for which I have information now, but 
it has estipulate leaves with longer petioles and longer laminas, its sepals are all 
eglandular, and the lobes of the lateral wing of the samara meet and overlap 
above the nut, as in E. propinqua but not E. adenophora. When E. boliviana is 
collected with flowers, those may strengthen the morphological separation of these 



ANDERSON: EXCENTRADENIA 



Amazonas: S. Gabriel, Q. NA-19-ZC, 12 Apr 1975 fr, Nasci- 
c Coradin 135 (holotype: IAN!; isotype: MICH!). Fig. 2. 



Stems velutinous with the hairs up to 0.6 mm long, erect, fusiform, generally 
short-bifurcate. Leaves opposite; lamina (except apex) 23-32 cm long, 16-25 cm 
wide, broadly elliptical, rounded or subcordate at base, revolute and bearing many 
small glands at margin, rounded and very abruptly caudate at apex with the tip 8 
mm long, persistently velutinous on both sides with the hairs erect, stalked, and 
bifurcate, the lateral veins 9-11 pairs, the parallel tertiary veins mostly 5-7 mm 
apart, the veins impressed above and prominent below; petiole 21-30 mm long, 4- 

5 mm in diameter, velutinous, bearing near middle a pair of glands 2-3.5 mm in 
diameter; stipules 1-1.5 mm long, triangular, borne on petiole at very base (?) or 
interpetiolar. Inflorescence velutinous, up to 8 cm long, comprising 3-7 umbels, 
each borne on a stalk 9-13 mm long; floriferous bracts 1.5-2 mm long, triangular, 
abaxially subvelutinous, adaxially glabrous; peduncle absent; bracteoles 1-1.5 mm 
long, ovate, abaxially subvelutinous, adaxially glabrous; pedicel 11-15 mm long in 
fruit, 0.8-1.5 mm in diameter, up to 2.5 mm in diameter at apex, subvelutinous. 
Sepals 2.5-3 mm long beyond glands, 2.5-3 mm wide, up to 3.5 x 4 mm in fruit, 
abaxially velutinous, adaxially glabrous, the anterior eglandular, the lateral 4 biglan- 
dular with the glands 2-2.5 mm in diameter. Lateral petals with the limb denticu- 
late and eglandular, the posterior with the hmb long-glandular-fimbriate. Stamens 
with filaments 3-5 mm long, straight; anthers 1-1.5 mm long. Ovary velutinous; 
styles (in fruit) 4.5-5.5 mm long, straight, sericeous at base, with a dorsal hook ca 
0.3 mm long at apex. Samara subcircular or transversely broadly elliptical with the 
nut positioned near center, 50-65 mm wide, 40-50 mm high; lateral wing continu- 
ous at base, incised to nut at apex with the sides not overlapping and leaving an 
evident gap, entire and sinuate at margin, puberulous; dorsal wing 5-6 mm wide, 
5-8 mm high, dentate; nut 5 mm in diameter, shortly velutinous. 

The above description is translated and modified from the original. In the 
protologue I described the inflorescence as a cyme, but the type shows it to have 
the kind of raceme of umbels described above in the discussion of the genus. The 
species is still known only from the type collection, so my information about its 
flowers is very incomplete. With its huge velutinous leaves, Excentradenia primaeva 
is quite unlike the other three species of the genus, and it is far-disjunct from 
them, too (Fig. 1). 

Excentradenia propinqua (W. R. Anderson) W. R. Anderson, comb. nov. Hiraea 
propinqua W. R. Anderson, Brittonia 46: 131. 1994.— Type: French Gui- 
ana. Riviere Tampoc ["Tampok"], dans une ile, 6 Apr 1977 fr, Moretti 
696 (holotype: MICH!; isotype: CAY, not seen). 

Stems densely and persistently sericeous. Leaves opposite or subopposite; 
lamina of larger leaves 10-14.3 cm long, 6-9.7 cm wide, ovate, rounded or trun- 
cate at base, usually bearing a series of tiny buttonlike glands on distal half of 
margin and occasionally 1-2 somewhat larger glands on margin near base, obtuse 
or abruptly short-acuminate at apex, adaxially sparsely sericeous to glabrate at 
maturity with the midrib usually persistently sericeous, abaxially densely and per- 
sistently sericeous, the lateral veins 7-9 pairs and connected by parallel cross- veins 3- 

6 mm apart, the veins and reticulum prominent below and visible above in dried 
leaves; petiole 12-20 mm long, 1.5-2 mm in diameter, densely and persistently 



36 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

sericeous, bearing al or slightly below apex a pair of glands 1.1-1.8 mm long; 
stipules 0.4-0.6 mm long, triangular, borne on petiole at very base or on stem 
beside petiole. Inflorescence densely and persistently sericeous, 2-5 cm long, usu- 
ally comprising 7 umbels, each borne on a stalk 5-10 mm long; floriferous bracts 
2-2.5 mm long, triangular or ovate, abaxially sericeous, adaxially glabrous; florif- 
erous peduncle 0.5-2.5 mm long; bracteoles 1-1.5 mm long, ovate, abaxially seri- 
ceous, adaxially glabrous, borne at apex of peduncle; pedicel 7-10 mm long in 
flower, 12-14 mm long in fruit, 0.5-1 mm in diameter, up to 2.5 mm in diameter at 
apex, sericeous. Sepals 3.5-4.5 mm long, 2.5-3 mm wide, abaxially sericeous, adax- 
ially glabrous, all eglandular or the anterior eglandular and the lateral 4 biglandu- 
lar with the glands 1.5-2.3 mm long. Lateral 4 petals with the claw 3-3.5 mm long, 
the limb 4.5-5 mm long and wide, laciniate, eglandular; posterior petal with the 
claw 5 mm long, the limb ca 6 mm long, 7 mm wide, fimbriate with the long 
slender divisions somewhat glandular-thickened at apex. Stamens with fllaments 
opposite sepals ca 4 mm long, erect but curved distally toward posterior petal, 
filaments opposite petals 2.5-3 mm long, erect, straight; anthers 1-1.3 mm long, 
the connective yellow. Ovary sericeous; styles bowed outward (i.e., curved out- 
ward from base and then back toward center of flower), sparsely sericeous in 
proximal half, dorsally acute or apiculate at apex with the projection up to 0.1 mm 
long, the anterior style 4 mm long, the 2 posterior styles 4.7 mm long. Samara 
depressed-circular with the nut positioned below the center, 48-56 mm wide, 42- 
50 mm high; lateral wing continuous at base, incised to nut at apex with the two 
sides overlapping, leaving little or no evident gap, sinuous at margin, thinly seri- 
ceous with very flne white appressed hairs; dorsal wing 3-5 mm wide, 5-7 mm 
high, entire or coarsely dentate; nut ca 3 mm in diameter, finely sericeous. 

Additional Specimens Examined: Guyana. Rupununi Distr., Kumukowau River, Camp 3, 2°56'N. 
59°02'W, 160 m, forest along river, Feb fl, Jansen-Jacobs et al. 3774 (MICH). Suriname. Forest near 
Brownsweg, Distr. Brokopondo. Feb fl, Lindeman, Mennega, et al. 51 (MICH, NY). 

As I noted in the protologue, this species is very similar to E. adenophora, but 
its densely and persistently sericeous leaves distinguish it immediately. The two 
species are also geographically disjunct (Fig. 1). Excentradenia propinqua has 
been collected in flower in February, and in fruit in April. 

ACKNOWLEDGMENTS 

study possible: CAY, IAN, K, MO, NY, U, US, and VEN. Karin Douthit drew the two species shown 



LITERATURE CITED 



'. Michigan Herb. 21: 37-84. 1997. 



NOTES ON NEOTROPICAL MALPIGHIACEAE— VI 

William R. Anderson 
University of Michigan Herbarium 

North University Building 
Ann Arbor, Michigan 48109-1057 

Like the preceding installments in this series, this is a miscellany. Many of the 
new species described here are needed for forthcoming floristic treatments. Others 
are included now, rather than later, just because they took my fancy as being 
somehow more interesting than the usual undescribed species, of which there are 
many more in the Malpighiaceae than I can treat in the limited time available to 
me. I have long assumed that, as my collaborators and I described novelties in the 
family, we would find fewer and fewer until they became unusual. I suppose that 
time must come eventually, but I now realize that it will not arrive any time soon. 
Collectors continue to find obviously new species, but the more important source 
is careful study of the specimens we already have, which never fails to reveal 
subtle new species lurking unrecognized in the piles of widespread, "well-known" 

Bunchosia cauliflora W. R. Anderson, sp. nov.— Type: Ecuador. Pastaza: Rio 
Pastaza, between Destacamento Chiriboga and Apachi Entza, 2°20-32'S, 
76°55'-77°08'W, 24 Jul 1980 fr, B. 0llgaard, E. Asanza C, J. Brandbyge, 
S. Roth & C. Sperling 35203 (holotype: AAU!; isotype: MICH!). 

Frutex (1-) 2-3 m longus vel arbor 3-10 m alta, ramis glabris. Lamina foli- 
orum majorum 7-12 (-13) cm longa, (2.5-) 3-5 cm lata, elliptica vel ovata, apice 
acuminata acumine 8-20 mm longo, glabra, abaxialiter utrinque costae (2-) 3-5 (- 
7) glandulis seriatis munita; petiolus 4-7 (-9) mm longus; stipulae 0.5-1.3 mm 
longae. Inflorescentiae caulinae, 1-3 (-4) cm longae, ex 6-30 floribus decussatis 
constantes. Petala omnia limbo toto circuito glanduloso-dentato vel petalum exti- 
mum distaliter eglandulosum. Filamenta sepalis opposita 2-3 mm longa, petalis op- 
posita 1.5-2.5 mm longa; antherae 0.7-1.3 mm longae. Gynoecium 3-carpellatum, 
glabrum; stylus ut videtur 1, ex 3 connatis constans. Fructus siccus 8-10 mm lon- 
gus et diametro, glaber, granulatus. 

Shrub (1-) 2-3 m tall or tree 3-10 m tall; stems mostly quite glabrous, very 
rarely sparsely sericeous at first, soon glabrate. Lamina of larger leaves 7-12 (-13) 
cm long, (2.5-) 3-5 cm wide, elliptical or somewhat ovate, cuneate to almost 
truncate at base, acuminate at apex with the acumen 8-20 mm long, glabrous, 
bearing abaxial glands in a row of (2-) 3-5 (-7) on each side of midrib, proximally 
near midrib but distally often diverging toward margin, the principal lateral veins 
few (4-7 pairs) and arching to fuse well within margin; petiole 4-7 (-9) mm long, 
glabrous or bearing a few appressed hairs and soon glabrate, narrowly winged for 
much of its length; stipules 0.5-1.3 mm long. Inflorescences borne on old stems after 
leaves have fallen, never axillary to current leaves, simple or occasionally basally 
ternate, without vegetative leaves, single at nodes or clustered with up to 6 or more in 
a cluster, slender and fragile in flower, becoming stouter in fruit; pseudoraceme 



38 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

1-3 (-4) cm long, containing 6-30 crowded decussate flowers, ± persistently 
sericeous on axis, peduncles, and pedicels; bracts mostly 0.7-1 mm long, triangu- 
lar, up to 2.5 mm long and linear at lowest nodes; peduncle 1-2 (-3.5) mm long; 
bracteoles 0.5-1 mm long, triangular, one of each pair bearing 1 flat basal abaxial 
gland 0.5-0.8 mm in diameter, this half or more below free part of bracteole on 
peduncle; pedicel 5-8 mm long in flower, up to 12 mm long in fruit. Sepals ca 1- 
1.2 mm long beyond glands and 1-1.5 mm wide, broadly obtuse or rounded, 
abaxially thinly sericeous, ciliate on margin, adaxially glabrous, pressed against 
filaments in anthesis; glands 8 (actually 10, apparently 8 through connation of 4 in 
pairs), 1.7-4 mm long, the anterior 2 shortest and the posterior 2 longest, obovate, 
compressed, glabrous, decurrent onto the pedicel. Petals yellow, glabrous, all 5 
with the limb glandular-dentate or glandular-fimbriate all around the margin or 
the outermost with the distal divisions not or hardly glandular; lateral petals with 
claw 1-1.7 mm long and limb 3.5-6 mm long, 3-7 mm wide, the outermost limb 
largest and cupshaped, the others nearly flat; posterior petal with the thick claw 
1.5-3 mm long, the limb 3.5-4.5 mm long and wide. Stamens glabrous; filaments 
2-3 mm long opposite sepals, 1.5-2.5 mm long opposite petals, ca 1/2 connate; 
anthers 0.7-1.3 mm long, pressed against styles, the connectives glandular-swol- 
len. Gynoecium 3-carpellate, glabrous; ovary ca 1 mm high, cylindrical, 3-locular; 
style apparently 1, actually 3 completely connate, 2-3 mm long, reaching higher 
than the anthers; 3 stigmas distinct but held together in a tight triangle, each 
pehate with a ventral indentation and a moderate dorsal extension. Fruit yellow, 
8-10 mm long and in diameter (dried), ovoid or globose or depressed-globose, 3- 
lobed, glabrous, granulate. 



Additional Specimens Examined. Ecuador. Napo: Yasuni Forest Reserve, 10 km E of Pontificia 
Universidad Catolica station, 0°41'S, 76°28'W, Acevedo-Rodriguez & Cedeno 7555 (US); Reserva 
Etnica Huaorani, S of Rio Tivacuno, 0°50'S, 76°14'W, Aulestia & Ima 1583 (MICH) & between Rio 
Tivacuno and Rio Yasuni. 0°50'S, 76°18'W. Aulestia & Gonti 1735 (MICH); Parque Nacional Yasuni, 
S of Rio Tiputini, 0°37'S, 76°29'W, Aulestia 1689 (MICH), 0°36'S, 76°29'W, Pitman & Aulestia 206 
(MICH) & 0°34'S, 76°3rW, Pitman & Dik 369 (MICH).— Pastaza: Rio Papayacu at Rio Curaray, 
]°29'S, 76°42'W, Holm-Nielsen et al. 22567 (AAU).— Sucumbios: Shushufindi (Nueva Loja), Coca 
(Puerto Francisco dc Orellana)-Lago Agrio, 50 km NE of Coca, Hading & Andersson 12004 (MICH); 
Lago Agrio, Lugo S. 3152 (MICH); Rio Aguarico, 5 km S of Lago Agrio, Lugo S. 3184 (MICH); Las 
Sachas, Coca-Lago Agrio, 30-40 km NE of Coca, Lugo S. 3390 (MICH); Guamanyacu, Coca-Lago 
Agrio, 40 km NE of Coca, Lugo S. 3423 (MICH). 



Collected in moist tropical forests at elevations of 235-400 m, with flowers 
from November to March and with fruits in June and July. 

Most species of Bunchosia bear their inflorescences axillary to leaves of the 
current year's growth, but in this one they appear only on leafless stems at least 
one year old; the epithet cauliflora refers to that peculiarity. In some specimens 
several pseudoracemes are clustered at swollen nodes on stems that must be 
several years old, which I take to mean that the stems produce these cauline 
inflorescences not just once at a given node, but several times, perhaps for many 
years. Bunchosia cauliflora is also notable for the glabrous stems, the small, gla- 
brous, long-acuminate leaves with a row of 3-5 abaxial glands on each side of the 
midrib, the short slender pseudoracemes of crowded decussate flowers, the glan- 
dular-dentate petals, and the glabrous tricarpellate gynoecium with the styles com- 
pletely connate. The species most like this one is B. pseiidonitida Cuatrec. which 



1997 W. R. ANDERSON: MALPIGHIACEAE 39 

is known from western Colombia and western Ecuador (Los Rios and Guayas). In 
B. pseudonitida the leaf glands are fewer (usually 1 or 2 per side) and the inflores- 
cence is borne axillary to current leaves. 

Bunchosia pernambucana W. R. Anderson, sp. nov. —Type: Brazil. Pernambuco: 
Floresta Inaja, Reserva Biologica de Serra Negra, ascent to the forest, 8 
Mar 1995 fl, E. Menezes, E. M. VHlaroiico, S. S. Eira & E. Rodrigues 33 
(holotype: MICH!). 

Frutex 0.8-1 m longus, ramis sericeis mox vel demum glabratis. Lamina foli- 
orum majorum 6.5-9 cm longa, 2.3-3.6 cm lata, elliptica, mox glabrata, abaxialiter 
eglandulosa vel 2 glandulis basalibus munita; petiolus 4-6 mm longus; stipulae 1- 
1.8 mm longae. Inflorescentia 2.5-5 cm longa, ex 6-10 floribus decussatis con- 
stans. Petala lateraha limbo eroso eglanduloso, petalum posticum limbo proxi- 
maliter glanduloso-eroso distaliter eglanduloso. Filamenta sepalis opposita 3 mm 
longa, petalis opposita 2.3-2.5 mm longa; antherae 1-1.3 mm longae. Gynoecium 
3-carpellatum, glabrum; styh 3, usque ad medium connati vel cohaerentes, dis- 
taliter liberi. Fructus siccus 8-10 mm longus, 7-8 mm diametro, glaber, laevis (i.e., 
non granulatus) reticulo prominenti. 

Shrub 0.8-1 m tall; stems initially thinly to moderately densely sericeous, soon 
or eventually glabrescent to quite glabrate. Lamina of larger leaves 6.5-9 cm long, 
2.3-3.6 cm wide, elliptical, cuneate and often somewhat decurrent at base, obtuse 
or abruptly short-acuminate at apex, initially thinly sericeous but nearly or quite 
glabrate at maturity, eglandular or bearing 2 abaxial glands, 1 on each side near 
base (up to 7 mm above base) and beside midrib or between midrib and margin, 
the very fine reticulum and 7-10 pairs of lateral veins usually prominent on both 
sides in dried leaves but more strongly so above than below; petiole 4-6 mm long, 
soon glabrate; stipules 1-1.8 mm long. Inflorescence axillary, simple, without veg- 
etative leaves, loosely sericeous to glabrate, 2.5-5 cm long, containing 6-10 decus- 
sate flowers; bracts 1-2.5 mm long, ovate, often acuminate; peduncle 0.5-2 mm 
long; bracteoles 1-1.5 mm long, ovate, one of the pair (sometimes both) bearing 1 
(sometimes 2) raised eccentric basal-abaxial glands 0.5-1 mm in diameter; pedicel 
2.5-5 mm long, sericeous to glabrate. Sepals 1-1.8 mm long beyond glands, 1.5-1.8 
mm wide, broadly obtuse or rounded, glabrous except for the ciliate margin; 
glands 8 (actually 10, apparently 8 through connation of 4 in pairs), 1.5-3 mm 
long, the anterior 2 shortest and the posterior 2 longest, elliptical or obovate, 
compressed, glabrous, the posterior 2 decurrent further on pedicel than the oth- 
ers. Petals yellow, glabrous; lateral petals reflexed, the posterior erect; outermost 
(anterior-lateral) petal with claw 3 mm long and limb cupshaped, 5 mm long, 6 
mm wide, erose, eglandular; other 3 lateral petals with claw 2.5-3 mm long, limb 
4-4.5 mm long and wide, ± flat, erose and eglandular; posterior petal with the 
thick claw 3.5 mm long, the limb 4.5 mm long and wide, ± flat, erose with the 
proximal divisions glandular. Stamens glabrous; filaments 3 mm long opposite 
sepals, 2.3-2.5 mm long opposite petals, ca 1/3-1/2 connate; anthers 1-1.3 mm 
long, pressed against styles, the connectives glandular-swollen. Gynoecium 3-car- 
pellate, glabrous; ovary 2 mm high, ovoid, 3-locular; styles 3, 1/2 connate or coher- 
ent, distally distinct, 2 mm long, held erect and together, reaching to or just above 
tops of anthers; stigmas capitate with a ventral indentation and a dorsal extension. 
Fruit orange-red at maturity, 8-10 mm long and 7-8 mm in diameter (dried), 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



Additional Specimens Examined. Brazil. Pernambuco: Buique, estrada Buique-Calimbau, 8°37'S, 
37°10'W, 790 m, shrubby vegetation on sandy soil, May fr, Laurenio et al. 40 (MICH); type locality, 
open area. Mar fl, Tschd et al. 12 (MICH) & inside the forest, Feb fr, Tschd et al. 15 (MICH). 

Bunchosia pemambucana is a new, northern addition to the Bunchosia mariti- 
ma complex, whose other members (B. acuminata Dobson, B. macilenta Dobson, 
and B. maritima (Veil.) J. F. Macbr.) occur well south of Pernambuco (see Fig. 1). 
The new species has the small leaves, short inflorescences, glabrous ovary, and 
small fruit of B. macilenta, but the paired leaf glands, alternating long and short 
filaments, short styles, and smooth fruit of B. acuminata. Its petals are nearly 
eglandular, unhke those of both B. acuminata and B. macilenta. 

Bunchosia petraea W. R. Anderson, sp. nov.— Type: Venezuela. Amazonas: Dept. 
Atures, 56 km NE of Puerto Ayacucho on road to El Burro, extensive 
granitic outcrop ("laja") west of road, 28 Apr 1984 fl, T. Plowman & F. 
Gudnchez 13762 (holotype: MICH!; isotypes: F! VEN!). Fig. 2e-g. 

Frutex fragilis 1.5 m longus, ramis primo hispidis demum glabratis, pilis plerum- 
que V-formibus sessilibus vel subsessilibus basi interdum calcaratis. Lamina foli- 
orum majorum 3-4 cm longa, 1.6-2 cm lata, margine costaque hispida; petiolus 2- 
4 mm longus, hispido-tomentosus; stipulae 0.4-0.7 mm longae. Inflorescentia 2-3 
cm longa, ramum bifoliatum terminans, ex 8-12 floribus decussatis constans. Peta- 
lum posticum limbo toto circuitu glanduloso-dentato. Antherae connectivis tumi- 
dis. Ovarium 2- vel 3-carpellatum, glabrum; styli 2 vel 3, hberi, 3 mm longi. 

"Open fragile shrub" 1.5 m tall; vegetative internodes initially densely hispid 
but many hairs deciduous as maturation occurs, the stems moderately to sparsely 
hispid by the end of the first year, quite glabrate in later years; hairs colorless, 
mostly V-shaped with ± straight ascending arms 0.5-1 mm long, sessile or sub- 
sessile, the very short stalk apparently smooth in some hairs, bearing 1-several 
sharp basal spurs in others. Leaves deciduous promptly at end of each growing 
season, the new leaves emerging as flowers develop. Lamina of larger leaves (not 
fully expanded?) 3-4 cm long, 1.6-2 cm wide, elliptical, cuneate at base, obtuse or 
acute or slightly acuminate and usually bearing a convex gland at apex, bearing 2 
large, raised, button-like glands abaxially near base beside midrib and often a 
distal row of smaller glands on each side between margin and midrib but closer to 
margin, with the 6-9 pairs of lateral veins unusually prominent below, densely 
hispid on the margin and moderately hispid on both sides of midrib, with only 
scattered hairs elsewhere on lamina, the hairs like stem hairs but with arms more 
widely diverging, to horizontal; petiole 2-4 mm long, eglandular, hispid-tomen- 
tose with a mixture of hairs like stem hairs and short, fat, ± appressed. wormlike 
hairs, the latter occurring also on node between petioles and on abaxial midrib; 
stipules 0.4-0.7 mm long, narrowly triangular, perhaps distally fleshy or glandular, 
borne on inner face of petiole slightly above base. Inflorescence a pseudoraceme 
terminating a lateral shoot 1.5-3 cm long and bearing a single pair of full-sized 
vegetative leaves, or rarely terminating a principal shoot with several nodes; inflo- 
rescence axis 2-3 cm long, hispid like stems but with hairs shorter and less dense, the 
8-12 flowers decussate from base to apex; bracts 1.5-2 mm long, ovate, abaxially 



. R. ANDERSON: MALPIGHIACEAE 



D B. acuminata 
O B. macilenta 
• B. maritima 
■ B. pernambucana 



densely sericeous, adaxially glabrous; peduncle 1.5-2.5 mm long, thinly hispid like 
inflorescence axis; bracteoles apical or sHghtly subapical, 1-1.2 mm long, triangu- 
lar, abaxially sericeous, adaxially glabrous, one of each pair bearing an eccentric 
abaxial gland 1.2-1.5 mm in diameter, the gland actually developing mostly below 
bracteole on peduncle; pedicel 4.5-5.5 mm long, thinly hispid especially on adaxial 
side. Sepals 1.7-2 mm long beyond glands, 1.7-2 mm wide, rounded, glabrous or 
bearing a few hairs abaxially or on margin, appressed in anthesis, the glands 10 
(but appearing fewer due to fusion of adjacent glands on anterior sepals), 2.2-3.5 
mm long, elliptical or obovate, the posterior glands longer and more decurrent 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



■■0 



1997 W. R. ANDERSON: MALPIGHIACEAE 43 

than anterior ones, some detached at apex, glabrous. Petals yellow, glabrous; 
lateral petals reflexed, with the claw 3.5 mm long, the hmb 6.5-8.5 mm long and 
wide, concave (especially the outermost), nearly orbicular, denticulate or bearing 
a few small glands near base; posterior petal with the claw erect, 5 mm long, very 
thick, constricted at apex, the limb erect or distally reflexed, 7 mm long and wide, 
flat, suborbicular, glandular-dentate all around the margin with the glands much 
larger proximally than distally. Stamens glabrous; filaments 2.5-4 mm long, longer 
opposite sepals than opposite petals, connate in the proximal 0.5-1 mm; anthers 
(1.3-) 1.6-1.8 mm long, pressed against styles in anthesis, the connective yellow or 
light brown, swollen and extended at right angles to anther to produce an abaxial 
gland 0.5-1 mm long, this much larger on anthers opposite sepals than opposite 
petals. Gynoecium glabrous; ovary 2 mm high, ovoid, 2- or 3-carpellate; styles as 
many as carpels, distinct, 3 mm long, reaching about as high as tops of anthers or 
slightly higher, very slender, straight and erect, the stigmas small, capitate with a 
very short abaxial extension. Fruit unknown. 

This species is known only from the type, which was collected on top of a 
large boulder in the shade of trees. Its name means "rock-dweller" and refers to 
the fact that this seems to be another of the many species endemic to the granitic 
outcrops along the Rio Orinoco that are known locally as "lajas." 

Bunchosia petraea is closely related to B. mollis Benth., which is widespread 
in Venezuela. Bunchosia mollis is a variable species, and for a long time I tried to 
avoid the problem presented by Plowman & Gudnchez 13762 by treating it as an 
aberrant representative of that species, but I have reluctantly concluded that it 
probably represents a species that merits recognition. In B. mollis the larger leaves 
are usually 6-18 cm long and 3-12 cm wide. Perhaps more signiflcantly, the hairs 
on those leaves are abundant and evenly distributed over the whole abaxial sur- 
face, and the individual hair usually has a well-developed stalk longer than the 
stalk (if any) in a hair of B. petraea (Fig. 2). The inflorescence in B. mollis is also 
somewhat to much larger, 4-12 cm long and containing 10-30 flowers. 

Bunchosia petraea bears a superficial resemblance to B. postuma Nied., a species 
of dry habitats on the north coast of Venezuela, but that has straight, sessile, 
appressed hairs. The spurs at the base of the hairs in B. mollis, B. petraea, and B. 
thaumatothrix constitute a synapomorphy that marks those species as a clade. 

Bunchosia thaumatothrix W. R. Anderson, sp. nov. — Type: Venezuela. Distrito 
Federal: Camino El Rincon-Las Tunitas, SE de Maiquetia, vertiente norte 
de la Cordillera de La Costa, bosque tropofilo, 200-600 m, 4 Jun 1971 fl, 
G. Morillo & B. Manara 1172 (holotype: MICH!). Fig. 2c-d. 

Frutex 2-3 m longus, ramis primo hispidis demum glabratis, pilis ut videtur 
simplicibus basi tuberculis rotundatis instructis. Lamina foliorum majorum 5-6.3 
cm longa, 2.2-2.6 cm lata, supra hispida demum glabrescens, subtus pertinaciter 
hispida; petiolus 3-4 mm longus, hispidus; stipulae non visae. Inflorescentia 2-3.5 
cm longa, ramum bifoliatum terminans, ex 6-10 floribus decussatis constans. Peta- 
la limbo toto circuitu glanduloso-dentato. Antherae connectivis non tumidis. Ova- 
rium 3-carpellatum, glabrum; styli 3, liberi, 4-4.5 mm longi. 

Shrub 2-3 m tall; stems initially densely hispid, the stiff spreading hairs golden 
fading to whitish, persistent during first year, deciduous in later years. Lamina of 
larger leaves 5-6.3 cm long, 2.2-2.6 cm wide, elliptical, cuneate at base, acute to 
obtuse and eglandular at apex, bearing an abaxial row on each side of 2-4 glands 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 



between midrib and margin, with the 7-9 pairs of lateral veins prominent below 
but not above, initially hispid on both sides with stiff erect hairs ca 1 mm long, 
those of the abaxial surface whitish, very dense, persistent, those of the adaxial 
surface golden, less dense, eventually deciduous except from midrib and margins, 
the hairs mostly "basifixed," i.e., l-branched, occasionally with a second very 
short branch from the base, and subtended at very base by several (ca 4-5) short 
bulbous tubercles; petiole 3-4 mm long, eglandular, densely and persistently gold- 
en-hispid; stipules not found. Inflorescence a pseudoraceme terminating a lateral 
shoot 0.6-1 cm long and bearing a single pair of full-sized vegetative leaves; inflo- 
rescence axis 2-3.5 cm long, hispid like stems but with hairs often somewhat 
shorter and less dense, the 6-10 flowers decussate from base to apex; bracts 1.5- 
2.2 mm long, ovate, abaxially densely sericeous, adaxially glabrous; peduncle 2-4 
mm long, thinly hispid, the hairs much less dense than on axis; bracteoles apical or 
slightly subapical, 0.8-1 mm long, ovate, abaxially sericeous, adaxially glabrous, 
one or both bearing an eccentric abaxial gland 1-1.5 mm in diameter, the gland 
actually developing mostly below bracteole on peduncle; pedicel 2.5-4 mm long, 
thinly hispid with the hairs mostly on the adaxial side. Sepals 1-2.2 mm long 
beyond glands, 2-2.5 mm wide, rounded, minutely denticulate and sometimes 
bearing tiny hairs on margin, otherwise glabrous or bearing a few scattered abaxial 
hairs, appressed in anthesis, the glands 10, 3-4 mm long, obovate and somewhat 
decurrent onto pedicel, sometimes detached at apex, mostly connate in pairs, 
glabrous. Petals yellow, glabrous; lateral petals reflexed, with the claw 2.5-3 mm 
long, the limb 5.5-7 mm long, 4-7 mm wide, concave, obovate, glandular-dentate 
or glandular-fimbriate all around the margin but with the glands larger on posteri- 
or-lateral petals than on anterior-lateral petals; posterior petal erect, the claw 3.5- 
4 mm long, very thick, constricted at apex, the limb 5.5 mm long, 4 mm wide, flat, 
elliptical, glandular-dentate all around the margin with the glands much larger 
proximally than distally. Stamens glabrous; filaments 3-4 mm long, longer oppo- 
site sepals than opposite petals, 1/2-2/3 connate; anthers 0.9-1.2 mm long, pressed 
against styles in anthesis or somewhat spreading, the connective hght brown, not 
or only slightly swollen at apex. Gynoecium glabrous; ovary 2-2.5 mm high, ovoid, 
3-carpellate; styles 3, distinct, 4-4.5 mm long, exceeding the anthers by ca 1.5 mm, 
very slender, straight and erect or slightly spreading, the stigmas capitate with a 
short abaxial extension. Fruit unknown. 

This species is known only from the holotype; attempts to locate additional 
sheets of that number at VEN have failed. The epithet means "wonderful hair" 
and refers to the very peculiar hairs that distinguish the species from its closest 
relative, Bunchosia mollis Benth. In B. mollis the leaf hairs have both branches 
well developed and borne horizontally or at a wide angle from a vertical stalk, 
which may be rather short; the result is a hair that is T-shaped or widely V- or Y- 
shaped (Fig. 2a-b). In B. thaumatothrix there is no stalk at all and one of the two 
branches has been lost completely or reduced to a very short branch from the 
base, while the main branch is nearly or quite erect, giving the impression of a 
simple basifixed hair. Moreover, in B. mollis each hair has a basal ring of several 
pointed outgrowths; very short basal outgrowths are present on the hairs of B. 
thaumatothrix too, but they are rounded, not pointed (Fig. 2c-d). Bunchosia mol- 
lis also differs from B. thaumatothrix in the following ways: the leaves are mostly 
larger, the stipules are well developed, the inflorescence is usually longer and 
contains more flowers, the lateral petals are eglandular or bear only a few small 
glands proximally on the posterior-lateral pair, the anther connectives are notably 



1997 W. R. ANDERSON: MALPIGHIACEAE 45 

swollen at the apex, and the styles are shorter, up to 3 mm long, barely exceeding 
the anthers. Bunchosia mollis is not known from the Distrito Federal, but it has 
been collected in the adjacent state of Miranda. 

Byrsonima basiliana W. R. Anderson, sp. nov.— Type: Venezuela. Amazonas: 
Selvas pluviales cerca y en las orillas del Rio Casiquiare, entre Curimacare 
y Buena Vista, 2°0'N, 66°30'W, 150 m, 20-30 Oct 1986 fl, B. Stergios, H. 
Martinez & O. Martinez 9654 (holotype: MICH!; isotypes: MO! PORT! 
VEN!). 

"Suffrutex" vel "arbor parva," ramis vegetativis primo tomentosis vel laxe 
sericeis demum glabratis. Lamina foliorum majorum 9.5-13.5 cm longa, 3.5-5.7 
cm lata, abaxialiter pertinaciter tomentosa vel laxe subsericea vel demum glabre- 
scens; petiolus 8-12 mm longus, primo tomentosus vel subsericeus; stipulae 1.5- 
2.5 mm longae, omnino connatae. Inflorescentia floribus 1 in axilla cujusque brac- 
teae portatis; bracteae 1.2-1.5 (-2) mm longae, in fructu persistentes vel deciduae; 
pedunculus (0-) 0.5-2 (-3) mm longus; pedicellus 6.5-9 mm longus, circinatus in 
alabastro, decurvatus vel tortus in fructu. Petala rosea et alba. Antherae sericeae, 
locuhs apice extensionibus sterilibus 0.5-1 mm longis ornatis, connective partem 
fertilem loculorum ca 0.5 mm superanti. Ovarium 2 carpelhs fertilibus. Fructus 
rubescens, ovoideus apice rostratus, 8-9.5 mm longus, 6-7 mm diametro (siccus). 

"Subshrub" or "small tree"; vegetative internodes loosely sericeous or appressed- 
tomentose, glabrescent during the first year with the simultaneous peeling and 
eventually complete loss of the cuticle. Lamina of larger leaves 9.5-13.5 cm long, 
3.5-5.7 cm wide, elliptical, cuneate and sometimes somewhat decurrent at base, 
flat at margin, mostly rounded-acute or narrowly obtuse at apex, appressed-to- 
mentose to glabrate above with some hairs persistent on the midrib, persistently 
appressed-tomentose or loosely subsericeous below or eventually patchily gla- 
brescent, the hairs sinuous to twisted, appressed to erect, the principal lateral 
veins 8-10 pairs, the lateral veins and reticulum prominulous above, prominent 
below; petiole 8-12 mm long, appressed-tomentose or subsericeous to eventually 
glabrescent; stipules 1.5-2.5 mm long, completely and smoothly connate, the pair 
rounded at apex, abaxially sericeous, adaxially glabrous. Inflorescence 10-15 cm 
long, densely tomentose or subsericeous to incompletely glabrescent in age, the 
flowers borne 1 per bract; bracts 1.2-1.5 (-2) mm long, ovate and rounded at 
apex, abaxially appressed-tomentose, adaxially glabrous, appressed or spreading 
but not revolute, persistent or deciduous in fruit; peduncle (0-) 0.5-2 (-3) mm 
long, variable in the same inflorescence; bracteoles like bracts but smaller, 0.7-1 
mm long, persistent or deciduous in fruit; pedicel 6.5-9 mm long, densely tomen- 
tose to belatedly glabrescent, circinate in bud, decurved to twisted in fruit. Flowers ca 
16 mm in diameter. Sepals all biglandular, ca 2-2.3 mm long beyond glands, ca 1.5 
mm wide, triangular and obtuse or rounded at apex, abaxially densely 1 
adaxially glabrous, strongly revolute in the distal half in anthesis, 
3.5 mm long and wide in fruit; glands 1.6-2.2 mm long. Petals pink and white, 
glabrous; lateral petals strongly recurved in anthesis, the claw ca 3 mm long, the 
limb 4.5-5 mm long and wide; posterior petal with the thick claw erect, 3.5 mm 
long, the limb ca 4 mm long, 5 mm wide. Filaments ca 2-2.5 mm long, straight, 
distinct, abaxially glabrous, adaxially tomentose at base; anthers 2.5-3 mm long, 
caducous; locules linear, densely sericeous, with the fertile part 1.7-2 mm long, 
drawn out at apex into slender, sterile extensions 0.5-1 mm long; connective 



46 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

extended beyond fertile part of locules ca 0.5 mm, tapering distally, straight or 
recurved. Ovary ca 1.2 mm high, conical, glabrous or tomentose, only 2 of the 
locules fertile; styles 3.5-4 mm long, bent at the apex in bud, partially straighten- 
ing in anthesis. Fruit reddish, ovoid with a notable apical beak, 8-9.5 mm long and 
6-7 mm in diameter (dried), glabrous or tomentose to glabrate with the hairs 
persisting longest on the apical beak. 



MO, PORT, VEN). 

This species is named in honor of Dr. Basil G. Stergios (b. 1940), who, with 
the help of his collaborators from the "Universidad Nacional Experimental de los 
Llanos Ezequiel Zamora" in Portuguesa, has made a very significant contribution 
to our knowledge of the flora of Amazonian Venezuela, especially in the lowlands 
where life is often uncomfortable and collecting plants is far from easy. There can 
be little doubt that Byrsonima basiliana is most closely related to B. japurensis 
Adr. Juss., whose extensive range includes the area in Amazonian Venezuela 
where B. basiliana has been collected. The two species are linked by the connate 
stipules, the pink and white petals, the sericeous anthers whose locules are drawn 
out into long slender apical extensions and whose connective exceeds the fertile 
part of the locules, and the reddish fruit with an apical beak and only two fertile 
locules. Byrsonima japurensis has short, straight, strongly appressed, soon decidu- 
ous hairs on its leaves and stems; its pedicel is sessile or raised on a peduncle that 
is at most 0.5 mm long; and its bracts are very short, rarely exceeding 1 mm. 

Byrsonima formosa W. R. Anderson, sp. nov. — Type: Guyana. Kamoa River, 
1°32'N, 58°50'W, swamp between camp and Toucan Mountain, 260 m, 19 
Sep 1989 fl, M. J. Jansen-Jacobs 1701 (holotype: MICH!). 

Arbor 5-15 m alta, ramis vegetativis glabris. Lamina foliorum majorum 4.5- 
10 cm longa, 2.5-5.6 cm lata, elliptica, basi cuneata, apice acuta vel breviter acumi- 
nata, glabra; petiolus 7-10 mm longus, glaber; stipulae 1-2 (-2.5) mm longae, 
liberae. Inflorescentia 4.2-8 cm longa, glabra, floribus in dimidio distah congestis; 
bracteae bracteolaeque 0.5-1.2 mm longae lataeque, persistentes; pedicellus (12-) 
15-17 mm longus, circinatus in alabastro, rectus et rubescens in fructu. Sepala in 
fructu accrescentia usque ad 5.5 mm longa, rubescentia. Petala alba vel limbo albo 
et ungue roseo. Antherae 1.6-2.2 mm longae, glabrae, loculis 0.9-1.3 mm longis, 
dorsiventaliter complanatis, non vel vix alatis, connectivo globoso, loculos 0.7-1 
mm superanti. Ovarium glabrum, loculis omnibus fertilibus. Fructus 4 mm longus, 
4.5-5 mm diametro, nuce laevi debilique. 

Tree 5-15 m tall; vegetative internodes glabrous except hispid in axils of 
stipules. Lamina of larger leaves 4.5-10 cm long, 2.5-5.6 cm wide, elliptical, cu- 
neate and sometimes somewhat decurrent at base, nearly or quite flat at margin, 
acute or abruptly short-acuminate at apex, glabrous, the principal lateral veins 8-10 
pairs, prominulous on both sides, poorly differentiated from parallel lesser veins 
and reticulum; petiole 7-10 mm long, glabrous; stipules 1-2 (-2.5) mm long, free, 
ovate, broadly obtuse or rounded, abaxially glabrous, adaxially densely appressed- 
hispid. Inflorescence 4.2-8 cm long, all or most of the flowers crowded in the 
distal 2-5 cm and the proximal 2-4 cm consisting of a naked stalk, the axis gla- 
brous (except for hairs associated with bracts and bracteoles), the flowers borne 



1997 W. R. ANDERSON: MALPIGHIACEAE 47 

1 per bract; bracts 0.5-1.2 mm long and wide, broadly ovate to rotund, abaxially 
glabrous or bearing a few hairs, ciliate on the margin, adaxially glabrous except 
for a ring of hairs at base and between bract or bracteole and pedicel, appressed 
or spreading but not reflexed or revolute, persistent; peduncle none; bracteoles 
like bracts or smaller, especially narrower; pedicel (12-) 15-17 mm long, bearing a 
few scattered hairs to glabrate, pink in flower and turning red in fruit, circinate in 
bud, straight in fruit. Flowers ca 10-12 mm in diameter. Sepals all biglandular, 1-2 
mm long beyond glands, 1.2-2 mm wide, broadly rounded, glabrous except for the 
ciliate margin, usually revolute in anthesis, accrescent to 3.5-5.5 mm long and 3- 
4.5 mm wide in fruit and turning red; glands 1.5-2.5 mm long, pink, sometimes 
slightly decurrent. Petals white or white with pink claws, glabrous; lateral petals 
strongly recurved in anthesis, the claw 2.5-3 mm long, the limb oblate, 3-3.5 mm 
long, 4-5.3 mm wide; posterior petal with the thick claw erect, 3-4 mm long, the 
limb reflexed, 3-4 mm long, ca 5 mm wide, crumpled. Filaments 1.7-2.5 mm long 
opposite sepals, 2.2-3 mm long opposite petals, straight, basally connate with 
some usually connate for ca I mm, abaxially glabrous, adaxially tomentose in 
proximal half; anthers 1.6-2.2 mm long, glabrous; locules 0.9-1.3 mm long, rounded at 
apex, the outer locules flattened but unwinged or bearing a barely perceptible 
wing less than 0.1 mm wide; connective globose, exceeding locules by 0.7-1 mm. 
Ovary 1-1.2 mm high, glabrous, all 3 locules fertile; styles 3-3.6 mm long, bent at 
the apex in bud, straightening or not in anthesis. Fruit depressed-globose, 4 mm 
long, 4.5-5 mm in diameter when dried (said by collector to be 7 mm in diameter), 
green (submature?), glabrous, the stone nearly smooth and (at least in submature 
fruit) relatively thin-walled and easily broken open. 

mined: Guyana. Gunn's, Essequibo River, 1°39'N, SS^SS'W, savanna, 
et al. 1890 (MICH). Brazil. Amaz6nas: Mun. Presidente Figueiredo, 
Rio Uatuma, canteiro de obras da Usina Hidreletrica de Balbina, 1°30'-2°00'S, 59°30'-60°00'W, 
campina aberta, solo arenoso branco, Jul fl/imm fr, Cid Ferreira7559 (MICH) & Sep fl/imm fr, Cid 
Ferreira et al. 8189 (MICH): Rio Negro, Preto, Matupiry, t.f. high land, caatinga, Piassabal, Nov fl, 
Froes 22804 (\A}<,VB). 

The distribution of this species is shown in Fig. 3, except for the collection by 
Froes, for which I have been unable to discover the exact provenance on the Rio 
Negro. The epithet formosa [handsome, beautiful] refers to the display of white 
petals contrasting with reddish sepals and pedicels; this must be a very attractive 
tree. Byrsonima formosa is assignable to Niedenzu's series Platylepis. Within that 
series Byrsonima formosa can be compared to B. umbellata Adr. Juss. and B. densa 
(Poir.) DC. In most characters of its inflorescence and flowers it resembles B. 
umbellata, a species with a much more southern range (see Fig. 3); even the 
glabrous inflorescence axis, while atypical for B. umbellata, is found in some 
populations of that species. However, B. umbellata has the leaves very broadly 
obtuse or rounded at the apex, with the petiole only 2-3.5 mm long and the 
stipules 2-4 mm long; the lamina is occasionally broadly cuneate at the base, but 
usually it is rounded to cordate. Byrsonima umbellata is generally a smaller plant 
than B. formosa, a shrub or small tree seldom exceeding 5 m. In its habit, leaves, 
and stipules B. formosa resembles B. densa, a species of northeastern Amazonia 
(see Fig. 3), except that the vegetative parts are quite glabrous from the beginning 
in B. formosa, while in B. densa they usually bear some hairs when first formed, 
which are soon lost. The following couplet summarizes the most consistent differ- 
ences between the two species in their reproductive structures: 



VERSITY OF MICHIGAN HERBARIUM 




A B. densa 

▲ B. formosa 

V B. garcibarrigae 

O B. melanocarpa 



A^^ 



FIG. 3. Dislributi 



1. Inflorescence axis hairy, bearing flowers for about 3/4 of its length, to within 1-2 cm of the 
base; pedicels up to 11 mm long; petals usually reported as pink; filaments nearly or quite 
distinct; stone of fruit bearing prominent ribs, extremely hard. B. densa. 

1. Inflorescence axis glabrous, with flowers mostly restricted to the distal 1/2; pedicels (12-) 15- 
17 mm long; petals white or white with pink claws; some or all filaments usually connate in 
the basal 0.5-1 mm; stone of fruit smooth, easily broken open. B. formosa. 

It also seems likely that the sepals are much more accrescent and redder in 
fruit in B. formosa than in B. densa, but better field notes are needed to enable 
one to assess the consistency of that difference. 

Byrsonima huberi W. R. Anderson, sp. nov. — ^Type: Venezuela. Amazonas: Depar- 
tamento Atures; Sierra Maigualida, NW sector, 5°30'N, 65°15'W, small 
valley along an upper tributary of Cano Iguana, 2000 m, 28 Feb-3 Mar 
1991 imm fl, P. E. Berry, O. Hiiber & J. Resales 4829 (holotype: MICH!; 
isotypes: MO! MYF!). 



Arbor 3 m alta, r< 
cm longa, 2.6-3.8 c 






folio 






n lata, elliptica vel paulo ovata, basi cune; 
apice late obtusa vel rotundata, coriacea, abaxialiter dense et pertinaciter tomen- 
tosa; petiolus 8-12 mm longus, pertinaciter tomentosus; stipulae 3-4 mm longae, 
connatae, abaxialiter tomentosae, adaxialiter glabrae. Bracteae bracteolaeque 
persistentes; pedicellus 8 mm longus in fructu. Sepalorum glandulae roseae. Antherae 
glabrae, connectivo loculos non superanti. Ovarium glabrum. Fructus siccus 10 



1997 W.R.ANDERSON: 

Tree 3 m tall; stems densely and persistently tomentose, the hairs turning 
from ferrugineous or reddish brown to dark brown and eventually fading to gray 
or white. Lamina of larger leaves 5-6.8 cm long, 2.6-3.8 cm wide, elliptical or 
slightly ovate, cuneate to rounded at base, somewhat revolute in age, broadly 
obtuse to rounded at apex, coriaceous, adaxially initially ferrugineous-tomentose 
but the hairs fading and gradually deciduous in age, with some gray hairs usually 
persistent at least on midrib, abaxially densely and persistently tomentose with 
the hairs so dense as to completely hide the epidermis, the hairs turning from 
reddish brown to dark brown to gray, most hairs strongly twisted and tightly held 
but the midrib bearing also an admixture of straight spreading hairs 0.8-1.4 mm 
long; lateral veins 7-9 pairs, the lateral veins and reticulum sunken adaxially to 
produce a rugose effect, prominent abaxially; petiole 8-12 mm long, densely and 
persistently tomentose with the same mixture of hair types as the abaxial midrib, 
fading from brown to gray; stipules 3-4 mm long, completely and smoothly con- 
nate with the pair rounded or slightly emarginate at apex, abaxially densely and 
persistently appressed-tomentose, adaxially glabrous. Inflorescence up to 8 cm 
long, densely and persistently brown- to gray-tomentose with a mixture of twisted 
appressed hairs and straight spreading hairs, the flowers restricted to the distal 1/2-2/3 
of the axis and borne 1 per bract; bracts 2.5-3 (-3.5) mm long, 1.5-2 mm wide, 
triangular or ovate, abaxially densely tomentose, adaxially glabrous, spreading to 
revolute, persistent; peduncle none; bracteoles like the bracts but smaller; pedicel 
8 mm long (1 seen, on a detached fruit), densely and persistently tomentose, 
straight or slightly nodding (?) in fruit. Sepals all biglandular, 3.5-5 mm long and 
3-4 mm wide in fruit, abaxially densely and persistently tomentose, adaxially 
glabrous, the glands pink, 1.5-2 mm long in fruit. Petals glabrous. Filaments abax- 
ially glabrous, adaxially hirsute at base; anthers glabrous, the locules cylindrical, 
the connective not or hardly exceeding the locules. Ovary glabrous. Fruit "dull 
orange," 10 mm in diameter (dried), globose or slightly ovoid, glabrous. 

This most interesting species is named in honor of my friend Dr. Otto Huber 
(b. 1944), in recognition of his contributions to the botany of the Venezuelan 
Guayana. The type is the only known collection, and it is grossly inadequate, as it 
bears only extremely immature flower buds (so young that the pedicel is not at all 
elongated and the petals are not yet visible), one old inflorescence axis, and one 
detached fruit. I am therefore uncertain about many of the characters necessary 
for the placement of the species in the genus Byrsonima. The buds are so young 
that it is impossible to say whether or not the pedicel becomes circinate as it 
elongates. The pink calyx glands, noted by the collectors, tell us that the petals are 
almost certainly white or pink, most likely white turning pink in age. The anthers 
are so immature that it is possible, but not very probable, that the connective 
enlarges and exceeds the locules at maturity. The pedicel of the single detached 
fruit is straight, not curved or twisted, but the fruit itself seems to face somewhat 
downward, so I suspect this will prove to be one of those species in which the 
pedicels are not strongly decurved but the fruits are not held erect. Future collec- 
tions of B. hiiberi will certainly correct and complement my very tentative de- 
No species known to me from northern South America could be mistaken for 
Byrsonima huberi, which is distinguished by the following combination of charac- 
ters: Lamina densely and persistently tomentose below, stipules connate, white/ 
pink petals, glabrous anthers with cylindrical locules and the connective not or 
hardly exceeding the locules, and a glabrous ovary. To find other species with that 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

ne most cross the Amazon basin and go to Minas Gerais 
in the Planalto of Brazil. There Byrsonima macwphylla (Pers.) W. R. Anderson 
[= B. nervosa DC] and B. variabilis Adr. Juss. share all of the above features with 
B. huberi, except that in the same flower of B. macrophylla there is a series of 
anthers from some with the connective equalling the locules to others with the 
connective distinctly exceeding the locules. Both B. macrophylla and B. variabilis 
differ from B. huberi in having the bracts and bracteoles mostly deciduous during 
anthesis or at least before maturation of the fruit, and in both species the pedicel 
is strongly decurved in fruit, becoming twisted in B. variabilis. Byrsonima macro- 
phylla strongly resembles B. huberi in its rugose coriaceous leaves, whose tomen- 
tum undergoes the same aging from dark brown to gray. However, the leaves of 
B. macrophylla are much larger (laminas 10-23 x 4.5-12 cm), the inflorescence is 
longer (10-27 cm), the flowers are often borne 2 (-3) per bract, and the fruit is 
much larger, 15-20 mm in diameter when dried, up to 25 mm in diameter when 
fresh. In B. variabilis the lamina is comparable in size to that of B. huberi, the 
inflorescence is about the same length, and the flowers seem to be borne 1 per 
bract. However, B. variabilis has a very short petiole, usually 1-4 mm long, short- 
er than the stipules to slightly longer, never twice as long. The bracts are long (3-5 
mm) and narrow, and the fruit is smaller, 5-6 mm in diameter when dried. 

Given their deciduous bracts and bracteoles and decurved pedicels, I hesitate 
to claim too much for B. macrophylla and B. variabilis as putatively closest rela- 
tives of B. huberi. However, the three species do share a number of significant 
character-states, and all three grow in rather similar habitats, open vegetation on 
nutrient-poor soils among rock outcrops at elevations of 1000-2000 m, so they 
may really be sister species. If so, that raises the intriguing question of how their 
disjunct distribution came about. 

Byrsonima melanocarpa Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 180. 1922.— 
Type: Brazil. Para: Campinas do Achipica, Baixo [Rio] Trombetas, 20 
Sep 1910 fl/fr, A. Ducke s.n. [Museu Goeldi 10948] (lectotype, here desig- 
nated: MG! F neg. 45549). 

When Ducke described this species he correctly placed it in Niedenzu's series 
Platylepis but did not exphcitly compare it to any of the other species in the series. 
He simply stated that one could recognize the species by the fact that the petals 
are white throughout anthesis, the inflorescence turns red in fruit, and the very 
small fruit is black at maturity. Unfortunately, all of those character-states are 
found also in Byrsonima umbellata Adr. Juss., 1840, and for some years I have 
been unsure that B. melanocarpa would survive careful comparison with B. um- 
bellata. Both species are shrubs or small trees, usually growing in periodically 
inundated sandy campos or near water, especially at the margin of woods or 
gallery forests. In both the glabrous lamina is very broadly obtuse or rounded at 
the apex, and the lamina is often glaucous abaxially when young. The lamina in B. 
melanocarpa is cuneate to rounded or rarely subcordate at the base, while that of 
B. umbellata is typically subcordate or cordate at the base, but in some popula- 
tions it is merely truncate or even cuneate. Similarly, the lamina is mostly narrower in 
B. melanocarpa than in B. umbellata, but the two overlap considerably in this 
character. One easy course would be to sink B. melanocarpa into synonymy under 
B. umbellata, but study of the collections at MICH suggests that there do exist two 
similar but separable taxa, and that they are geographically disjunct. Therefore, I 



1997 W. R. ANDERSON: MALPIGHIACEAE 51 

shall continue to recognize B. melanocarpa, at least for now. The couplet below 
summarizes the differences that most reliably distinguish these two species, and 
their distributions are shown in Fig. 3. 

1. Stipules 2-4 mm long, the petiole 0.5-1.2 (-1.4) times as long as stipules; most inflorescences 
with all or most flowers borne well above middle of inflorescence axis, occasionally with an 
isolated pair of flowers near or below the midpoint; bracts and bracteoies mostly appressed 
or spreading, occasionally revolute at the apex; claw of lateral petals 3.3-4 mm long; claw of 
posterior (flag) petal 3-4.5 mm long; styles 3.3-4.2 mm long. B. umbellata. 

1. Stipules 1-2 (-2.4) mm long, the petiole (1.5-) 2-3 times as long as stipules; flowers usually 
borne down to or below the midpoint of the inflorescence axis; bracts and bracteoies mostly 
strongly revolute; claw of lateral petals 1.7-3 mm long; claw of posterior (flag) petal 1.7-2.5 
mm long; styles 2.2-3.2 mm long. B. melanocarpa. 

For those using Niedenzu's 1928 treatment of Byrsonima, a note of clarifica- 
tion is in order. He listed B. melanocarpa as a synonym of B. coniophylla Adr. 
Juss., which he included in his series Platylepis. He did this because the two 
specimens to which he assigned the name B. coniophylla represented B. melano- 
carpa (indeed, one was a syntype), and he never saw the type of 5. coniophylla. In 
fact B. coniophylla is superficially similar to B. melanocarpa but has very different 
anthers and leaf venation, and while I am not certain where it belongs in Byrsoni- 
ma, I can say that it is not a good candidate for inclusion in series Platylepis 
because its stipules are often partly to fully connate; see my treatment for the 
Guayana Highland (1981). 

Niedenzu's name Byrsonima series Platylepis (Arbeiten Bot. Inst. Konigl. 
Lyceums Hosianum Braunsberg 1: 33. 1901) was described with several syntypes, 
not all of which I would include in the group. I here select Byrsonima densa 
(Poir.) DC. as the lectotype of that name. 

Byrsonima pedunculata W. R. Anderson, sp. nov.— Type: Brazil. Pernambuco: 
Brejo da Madre de Deus, Bituri, roadside, 4 Feb 1995 fl, M. /. N. Rodal & 
M. F. Sales 459 (holotype: MICH!; isotype: NY!). 

Arbor 7 m aha. Lamina fohorum majorum 9.5-12.5 cm longa, 3.5-5.4 cm lata, 
costa tomentosa; petiolus 9-11 mm longus; stipulae 3-4 mm longae, omnino con- 
natae vel distaliter liberae. Inflorescentiae bracteae 4-5 mm longae, 1-1.5 mm 
latae, caducae; pedunculus (2-) 3-7 mm longus, 1-2-florus; bracteolae 1.5-2.5 mm 
longae lataeque, caducae; pedicellus 9-10 mm longus. Calycis glandulae 2.6-3.1 
mm longae. Petala alba. Filamenta 2-2.2 mm longa; antherae 2.5-3.1 mm longae, 
sericeae inter loculos, connectivo loculos aequanti. Ovarium conicum, sulcatum, 
sparsim sericeum, 3 loculis omnibus fertilibus. 

Tree 7 m tall; stems persistently sericeous in the first year and glabrate in 
subsequent years, the hairs brown, straight and appressed or somewhat spreading. 
Lamina of larger leaves 9.5-12.5 cm long, 3.5-5.4 cm wide, elliptical or somewhat 
ovate, cuneate at base, bluntly acute or slightly acuminate at apex, nearly glabrate 
on both sides at maturity except for the ± persistently tomentose midrib, the 
principal lateral veins 8-9 pairs, prominent below; petiole 9-11 mm long, persis- 
tently tomentose or eventually glabrescent; stipules 3-4 mm long, completely con- 
nate or distinct just at apex, the pair smooth or sometimes somewhat sulcate 
abaxially, abaxially densely and persistently sericeous, adaxially glabrous. Inflo- 
rescence up to 14 cm long with flowers absent from the proximal 3 cm of the axis. 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 7 



proximally sericeous and somewhat glabrescent, distally (among flowers and on 
peduncles and pedicels) persistently tomentose; bracts 4-5 mm long, 1-1.5 mm 
wide, linear or very narrowly triangular, somewhat to strongly revolute or re- 
flexed, abaxially loosely sericeous, adaxially glabrous or bearing a few hairs, cadu- 
cous; peduncle (2-) 3-7 mm long, bearing 1-2 flowers; bracteoles 1.5-2.5 mm long 
and wide, triangular, mostly revolute at apex, abaxially loosely sericeous to to- 
mentose, adaxially nearly glabrous, caducous; pedicel 9-10 mm long, straight in 
bud (or slightly circinate?). Flowers 15 mm in diameter. Sepals all bigiandular, 2- 
2.2 mm long beyond glands, 1.7-2.5 mm wide, triangular and rounded at apex, 
appressed in anthesis but revolute at apex, abaxially sericeous, adaxially glabrous; 
glands 2.6-3.1 mm long, obovate. Petals white, glabrous; lateral petals reflexed, 
with the claw 2.5-3 mm long, the limb 6-7 mm long, 6-8 mm wide, cupshaped, 
entire or erose; posterior petal erect, the claw 3.2 mm long, stout, eglandular, the 
hmb 3.5 mm long, 4.5 mm wide, corrugated, erose. Filaments 2-2.2 mm long, 
nearly distinct, abaxially glabrous or sparsely hirsute at base, adaxially hirsute on 
proximal half with the hairs reaching to base of anther; anthers 2.5-3.1 mm long, the 
locules linear, unwinged, seemingly glabrous or bearing a few straight appressed 
hairs but densely sericeous between locules, the connective equalling the locules. 
Ovary 1.5 mm high, conical, sulcate, sparsely sericeous, all 3 locules fertile; styles 
3.2-3.6 mm long, bent over at apex in bud and retaining some curvature in anthe- 

This species is named for its well-developed peduncles, which are rare in 
Byrsonima. It is probably most closely related to B. cacaophila W. R. Anderson, 
which resembles it not only in having long peduncles often bearing more than one 
flower, but also in its similarly shaped bracts and bracteoles, its white petals, and 
the sericeous anthers with connectives that do not exceed the locules. Byrsonima 
cacaophila has substantially larger leaves than B. pedunculata (13-24 x 5.5-10 cm) 
with straight, more or less appressed hairs, and its petiole is longer (12-32 mm). 
Its bracts and bracteoles persist past maturity of the fruit, its pedicels are only 5-7 mm 
long, and its calyx glands are longer (3-4 mm), as are its filaments (2.5-3 mm). 

Byrsonima piresii W. R. Anderson, sp. nov.— Type: Brazil. Rondonia ("Mato 
Grosso"): 66 km W of Vilhena, Brasilia-Acre Highway, forest along iga- 
rape, 3 Sep 1963 fl, B. Maguire, J. Murqa Pires, C. K. Maguire & N. T. 
Silva 56531 (holotype: MICH!; isotype: NY!). 

Arbor 8-25 m alta, ramis vegetativis glabris. Lamina foliorum majorum 6- 
12.3 cm longa, 3-5.7 cm lata, elliptica vel obovata, apice late obtusa vel rotundata, 
glabra; petiolus 13-24 mm longus, glaber; stipulae 2.3-5 mm longae, liberae. Inflo- 
rescentia 8-16 cm longa, sericea vel subvelutina; bracteae bracteolaeque minimae, 
0.5-0.7 mm longae, rotundatae, erectae, persistentes; flores singuli in axilla cujus- 
que bracteae; pedicellus 5-7 mm longus, circinatus in alabastro, rectus in fructu. 
Sepala per anthesin appressa, in fructu vix accrescentia. Petala alba. Filamenta 
liberae; antherae 1.3-1.6 mm longae, glabrae, loculis 0.7-1 mm longis, dorsiven- 
traliter complanatis alatis ala 0.2-0.4 mm lata, connective globoso, loculos 0.4-0.6 
mm superanti. Ovarium glabrum, 3 loculis omnibus fertilibus. Fructus usque ad 6 
mm longus et diametro, nuce tuberculata. 

Tree 8-25 m tall; vegetative internodes glabrous except hispid in axils of 
stipules. Lamina of larger leaves 6-12.3 cm long, 3-5.7 cm wide, elliptical or 
somewhat obovate, cuneate and often somewhat decurrent at base, flat at margin. 



1997 W. R. ANDERSON: MALPIGHIACEAE 53 

broadly obtuse to rounded and sometimes slightly apiculate at apex, glabrous, the 
principal lateral veins 9-12 pairs, prominulous above, prominent below, well dif- 
ferentiated from lesser veins and reticulum; petiole 13-24 mm long, glabrous; 
stipules 2.3-5 mm long, free, ovate, broadly obtuse, abaxially glabrous, adaxially 
appressed-hispid. Inflorescence 8-16 cm long with the proximal 2-3 (-4) cm of the 
stalk lacking flowers, the axis persistently sericeous or subvelutinous and turning 
red in fruit, the flowers borne 1 per bract; bracts 0.5-0.7 mm long, 0.7-1 mm wide, 
broadly ovate to rounded, glabrous abaxially and on margin or bearing a few 
hairs, adaxially glabrous except for a ring of hairs at base and between bract and 
pedicel, erect, persistent; peduncle none; bracteoles like bracts or smaller, espe- 
cially narrower; pedicel 5-7 mm long in flower, up to 9 mm in fruit, persistently 
tomentose, circinate in bud, straight in fruit. Flowers ca 9-10 mm in diameter. 
Sepals all biglandular, 0.7-1.5 mm long beyond glands, 1.3-1.5 mm wide, broadly 
rounded, sparsely sericeous abaxially just above glands, short-ciliate on margin, 
otherwise glabrous, not or only slightly accrescent in fruit and probably turning 
red; glands white turning red, 1.2-1.9 mm long, the posterior pair longer than 
others and often decurrent. Petals white turning pink or red in age, glabrous; 
lateral petals spreading to strongly recurved in anthesis, the claw 1.5-2 mm long, 
the limb concave-oblate, 2.5-3 mm long, 4-4.5 mm wide; posterior petal with the 
thick claw erect, 2-2.5 mm long, the limb reflexed, 2.5-3.5 mm long, 3.5^ mm 
wide, crumpled. Filaments 1.3-1.5 mm long opposite sepals, 1.7-2 mm long oppo- 
site petals, straight, distinct, abaxially glabrous, adaxially sericeous proximally; 
anthers 1.3-1.6 mm long, glabrous; locules 0.7-1 mm long, rounded or short- 
apiculate at apex, the outer locules flattened and bearing a dark, well-developed 
wing 0.2-0.4 mm wide; connective globose, exceeding locules by 0.4-0.6 mm. Ova- 
ry 0.8-1.2 mm high, glabrous, all 3 locules fertile; styles 2-2.7 mm long, bent at the 
apex in bud, straightening or not in anthesis. Immature fruit ovoid and notably 
beaked, ca 5 mm long and 3.5 mm in diameter (dried), becoming 5-6 mm long and 
in diameter and nearly globose at maturity, glabrous, the stone bearing prominent 
horizontally oriented protuberances, very hard. 

Additional Specimens Examined. Brazil. Amazonas: Mpio Alvaraes, Rio Solimoes, Lago de 
Alvaraes, 3n3'S, 64°50'W, Amaral et al. 668 (MICH); Rio Negro, Santa Izabel, Black 48-2849 (IAN); 
Mpio Humaita, near Livramento on Rio Livramento, Krukoff 6934 (G, MICH, NY); Mpio Humaita, 
estrada Humaita-Porto Velho km 38, 8°S, 63°W, Teixeira et al. 277 (MICH). — Mato Grosso: Mpio 
Vila Bela da Santissima Trindade, 58 km S of Rondonia state line on BR364 from Vilhena to 
Caceres, 13°22'S, 59°56'W, Thomas et al. 4769 (MICH).— Rondonia: 2-4 km E of Mutumparana on 
road to Porto Velho, Prance et al. 8799 (MICH, NY); 4 km de Vilhena, 12°45'S, 60°10'W, Vieira et al. 
810 (INPA, MICH). 

Forests on terra firme, collected with flowers from September to November, 
and with fruits in October, November, and May. 

I am happy to name this beautiful plant in honor of Joao Murga Pires (1917- 
1994), a fine botanist who was very kind to me when I started working in Brazil 
some 25 years ago. Byrsonima piresii is a species of Niedenzu's series Platylepis. 
Among the other species of that difficult group, it is notable for its relatively large 
stipules, obtuse to rounded leaf tips, long and very dense inflorescence with a 
hairy axis, extremely short bracts and bracteoles, and anther locules bearing wide 
longitudinal wings. As Fig. 3 shows, Byrsonima piresii is a species of western 
Amazonia, where it is to be distinguished from B. garcibarrigae Cuatrec. and B. 
melanocarpa Ducke. The following key should facilitate their separation: 





CONTR. UNIVERSITY OF 


MICHIG. 


\N HERBARIUM 




.rubs 


or small trees 1 


.5-5 (-7) m tall; pe 


t,oles2-5mmlo 


ng. 


D. 


•ees (. 


4-) 8-25 m tall; 


petioles 12-25 mn- 


, long. 








Lam 


lina mostly acuminate at apex, occ 


:asion£ 


illy acute; : 


stipules 1-1.5 mm Ion 


ig; bra( 




long, triangular, often revolute; 




rs 1-2 per 


bract; sepals revolu 




accr 




to 2.5-3 mm long; 


anthei 


- locules n( 


3t or hardly winged. 


the wi 


0.1 r 


nm wide. 










B.i 


Lam 


lina broadly ob 


>tuse to rounded a 


It ape> 


c; stipules 


2.3-5 mm long; brae 


;ts 0.5- 


long 














sed in anthesis 


;, not or hardly a 


cc"resc( 


2nt in frui 


it, to 1.5 mm long; 


anther 


bear 


■ing a well-deve 


loped wing 0.2-0.4 


mmw 


'ide. 







Dicella aciculifera W. R. Anderson, sp. nov.— Type: Costa Rica. Puntarenas: 
Canton de Osa, Peninsula de Osa, Rio Termo 2 km arriba de la Estacion 
Cerro de Oro, 8°32'50"N, 83°30'45"W, 220 m, primary forest, 14 Apr 
1996 fr, L. Angulo 594 (holotype: MICH!; isotype: INB!). Fig. 4. 

Dicellae julianii similis, sed petalis lateralibus limbo 9-13 mm longo et 7.5-10 
mm lato, petalo postico limbo 7.5-8 mm longo et 6-7 mm lato, stylis divergentibus 
teretibus dorsaliter apiculatis, et nuce pilis aciculiformibus dense vestita differt. 

Woody vine; stems tightly sericeous to glabrate. Lamina of larger leaves 11- 
15 cm long, 4.5-8 cm wide, ovate or elliptical, broadly cuneate or rounded at base, 
abruptly acuminate at apex, bearing several small impressed glands on margin, 
sericeous to glabrate at maturity with the midrib and sometimes the lateral veins 
± persistently sericeous and sometimes with scattered hairs persistent elsewhere 
on the abaxial surface, the principal lateral veins 4-6 pairs, prominent below but 
not above, the intricate reticulum visible and ± prominent on both sides in dried 
leaves; petiole 11-13 mm long, persistently sericeous, eglandular; stipules inter- 
petiolar, triangular, minute (0.2-0.3 mm long), difficult to find and perhaps not 
always present. Inflorescence terminal and axillary, densely and persistently seri- 
ceous, a compound panicle composed of strictly decussate pseudoracemes, each 
pseudoraceme 0.5-1.5 cm long and containing 2-8 flowers; bracts 3-3.5 mm long, 
2.4-2.8 mm wide, boatshaped, abaxially sericeous, adaxially glabrous except to- 
mentose near apex, deciduous before or during anthesis; peduncle 5-6 mm long, 
persistently sericeous; bracteoles apical, 2.5-3 mm long, 1.5-2 mm wide, obovate 
and rounded at apex, concave, abaxially sericeous, adaxially glabrous, eglandular, 
persistent during anthesis, mostly deciduous in fruit; pedicel 9-10 mm long in 
flower, 16-18 mm long in fruit, persistently sericeous. Sepals 2-2.2 mm long be- 
yond glands, 2-2.5 mm wide, broadly rounded at apex, abaxially densely and 
evenly golden-sericeous, adaxially glabrous, pressed against stamens in anthesis, 
the anterior sepal eglandular or rarely bearing 1 small gland, the lateral 4 biglan- 
dular with the glands 1.8-2.5 mm long, elliptical or obovate, detached at apex. 
Petals yellow, flat or nearly so, obovate or orbicular, cuneate or truncate at base, 
broadly rounded at apex, erose or denticulate at margin with the proximal teeth 
sometimes glandular especially in posterior petal; 4 lateral petals probably spread- 
ing to reflexed, abaxially densely golden- or silvery-sericeous with the hairs com- 
pletely covering claw and limb to margin or nearly to margin in posterior-lateral 
pair, the claw 2.5 mm long, the limb 9-13 mm long, 7.5-10 mm wide, larger in 
anterior pair than in posterior pair; posterior petal probably erect, abaxially densely 
appressed-tomentose on claw and in center of limb and glabrous toward margin, 
the claw 3 mm long, the hmb 7.5-8 mm long, 6-7 mm wide. Stamens with the 
filaments 2.5-2.7 mm long, connate for ca 2 mm, abaxially sericeous, adaxially 



MALPIGHIACE\E 



«■ 



V 








\i 



=4: 



FIG 4 Dice! la 



56 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

glabrous; anthers strongly reflexed in anthesis, all fertile; locules 0.8-1.3 mm long, 
pilose on anthers opposite sepals, glabrous or bearing a few hairs on anthers 
opposite petals; connective red, ventrally sulcate from locules to apex, exceeding 
locules by 0.9-1.1 mm in anthers opposite sepals, 0.1-0.6 mm in anthers opposite 
petals. Ovary ca 1 mm high, very densely hispid, only the 2 posterior locules 
fertile; 2 posterior styles well developed, 2.3 mm long, strongly divergent from 
base, proximally sericeous, distally glabrous, nearly straight, and nearly terete 
with an internal stigma and extended at the apex into an acute dorsal projection 
ca 0.1-0.2 mm long; slender rudimentary style 0.7-1 mm long present on sterile 
anterior carpel, hidden among hairs. Fruit with the nut spherical, 11-14 mm in 
diameter, completely enclosed in a dense layer of sub-basifixed, stiff, somewhat 
spreading, needle-like hairs 1.5-2.2 mm long, and beneath the needle-like hairs 
densely sericeous with soft, appressed, ± medifixed hairs only 0.5-0.7 mm long; 
wings formed by enlargement of the sepals elliptical or obovate, abaxially thinly 
sericeous to glabrate, with the veins and reticulum prominent on both sides, the 
anterior smallest (2.4-2.5 cm long, 1.3-1.6 cm wide), the anterior-lateral pair 3-3.2 cm 
long and 1.3-1.6 cm wide, the posterior-lateral pair 3.4^ cm long and 1.6-2 cm wide. 

Additional Specimen Examined. Costa Rica. Puntarenas: Canton de Osa, Peninsula de Osa, R. 
F. Golfo Dulce, Cerro de Oro, alrededor del Albergue "Unioro," Sendero Termo, 8°39'20"N, 
83°26'30" W, 270 m, ripario, Mar fl, Anguto 116 (MICH). 

Dicella aciculifera is assignable to sect. Macropterys Chase (Chase, 1981), and 
resembles, in both leaves and fruits, its geographically closest neighbor, D. jidianii 
(J. F. Macbr.) W. R. Anderson. Dicella juliami differs from D. aciculifera in sever- 
al details, such as its longer pseudoracemes and its larger bracts that persist during 
anthesis, but the two differ most dramatically in their fruit hairs, styles, and petals. 
Dicella aciculifera is immediately distinguished by the dense covering of stiff needle- 
like hairs that completely conceal the nut. Nothing like them is found in any other 
species of Dicella; indeed, the underlayer of soft appressed hairs found on D. 
aciculifera fruits resembles the vesture found on the ovary and young fruits of its 
congeners, so it seems likely that the ancestor of D. aciculifera started with a 
sericeous fruit and added the dense coat of formidable needles. The epithet acicu- 
lifera (bearing little needles) refers to those hairs. The collector of the type noted 
them, describing them in Spanish as "urticantes." Many descriptions of Malpighi- 
aceae accuse members of the family of having "stinging" or "urticating" hairs, but 
they do not. The ability of those hairs to irritate is entirely mechanical — they 
inject no chemical substance like the truly stinging hairs of some Urticaceae and 
Euphorbiaceae. Of course, when the hairs are pointed on both ends, as in Dicella 
aciculifera, they have a doubled potential for inflicting pain— they get you coming 
and going, so to speak. 

Dicella aciculifera is also remarkable for its styles. In other species of the 
genus the style is laterally flattened and longitudinally rolled, such that the stigma 
faces toward the posterior petal and the dorsal sterile portion looks like a round- 
ed or angled shoulder or rostrum; see Chase, 1981, Fig. 2, and Anderson, 1981, 
Fig. 58. Nothing of the sort occurs in D. aciculifera. The style is terete or only very 
slightly flattened. The stigma is internal, and there is a short, acute, apical-dorsal 
extension; the style looks just like that found in many species of wing-fruited 
Malpighiaceae, in genera such as Heteropterys and Hiraea. Moreover, in D. acicu- 
lifera the two styles are strongly divergent from the base, then ascending; in other 



1997 W. R. ANDERSON: MALPIGHIACEAE 57 

Species of the genus, as shown in the figures cited above, the styles are erect and 
parallel, not at ail divergent. The new species has unusually large petals for a 
Dicella; in D. julianii, the limb of the lateral petals is only 7-9 x 5-6.5 mm, and 
that of the posterior petal only 4-6 x 3-5 mm. 

This species is known only from the type and paratype, which are the first 
collections of Dicella from Central America; the six species treated by Chase 
(1981) are all South American. 

Heteropterys cotinifolia Adr. Juss., Ann. Sci. Nat. Bot., Ser. 2, 13: 274. 1840. 
Banisteria cotinifolia (Adr. Juss.) C. B. Rob. ex Small, N. Amer. Fl. 25: 

134. 1910.— Type: Mexico. Oaxaca: Totolapa, Aug 1834 fl, G. Andrieux 
492 (holotype: P!; isotypes: G! K! P! P-JU!). 

Heteropterys gayana Adr. Juss., Ann. Sci. Nat. Bot., Ser. 2, 13: 274. 1840. 

Banisteria gayana (Adr. Juss.) C. B. Rob. ex Small, N. Amer. Fl. 25: 135. 

1910.— Type. Mexico: Oaxaca: Guichilona inter Tehuantepec et Guazacual- 

cos, Sep 1834 fr, G. Andrieux 494 (holotype: P!; isotypes: F! G! K! P! 

US!). 
Heteropterys portillana S. Watson, Proc. Amer. Acad. Arts 22: 402. 1887. 

Banisteria portillana (S. Watson) C. B. Rob. ex Small, N. Amer. Fl. 25: 

135. 1910. — Type: Mexico. Jahsco: Barranca near Guadalajara, E. Palmer 
112 in Jun 1886 fl (holotype: GH!; isotypes: NY! US!). 

Heteropterys arborescens Brandegee, Zoe 5: 203. 1905. Banisteria arborescens 
(Brandegee) Small, N. Amer. Fl. 25: 135. 1910.— Type: Mexico. Sinaloa: 
Cerro Colorado, vicinity of CuUacan, T. S. Brandegee s.n., 28 Oct 1904 fr 
(holotype: UC!; isotypes: GH! NY! US!). 

Heteropterys pallida Brandegee, Univ. Calif. Publ. Bot. 6: 182. 1915. Banisteria 
pallida (Brandegee) Standi., Contr. U.S. Natl. Herb. 23: 577. 1923.— Type: 
Mexico. Oaxaca: San Geronimo, Jul 1914 fl/fr, C. A. Purpus 7262 (holo- 
type: UCI; isotypes: A! F! GH! NY! MO! US!). 

Banisteria nemorum Brandegee, Univ. Calif. Publ. Bot. 10: 410. 1924.— Type: 
Mexico. Chiapas: Jalisco, Aug 1923 fr, C. A. Purpus 9185 (holotype: UC!; 
isotypes: A! F! GH! MEXU! MO! NY! UC! US!). 

This species is widespread in Mexico, but seems not to occur outside that 
country's borders. Within Mexico it is variable, which probably explains why it 
has been described so many times. My reason for giving its synonymy here is to 
establish that, when they are treated as the same species, Jussieu's name Hetero- 
pterys gayana is to be considered a synonym for his H. cotinifolia. 

Heteropterys hofiPmanii W. R. Anderson, sp. nov. — Type: Guyana. Upper Takutu- 
Upper Essequibo Region: southern Pakaraima Mountains, 5 km E of 
Tipuru village, Ureisha Mt. summit, 4°11'N, 59°31'W, 994 m, low forest, 4 
Mar 1992 fl, B. Hoffman 1194 (holotype: BRG!; isotypes: MICH! US!). 

nentosis demum glabrescentibus. Lamina foliorum majorum 
6 cm lata, ovata, basi cordata, margine ciliata vel glandulis 
utrinque tomentosa pilis tortuosis demum glabrata vel costa 
osa; petiolus 3-5.5 mm longus, basi glandulifer; stipulae non 
I floribus in umbelhs 4-floris sessilibus vel subsessilibus dis- 
tiullus; bracteae bracteolaeque 0.5-1 mm longae, eglandulosae 




58 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

vel bracteae (1-) 2 glandulis abaxialibus instructae; pedicellus 6-8 mm longus. 
Sepala per anthesin appressa. Petala lutea, glabra, in alabastro exposita. Stamina 
glabra, petalis posticolateralibus opposita maxima. Styli apice pedaliformes dor- 
saliter trmicati vel brevissime apiculati. 

Woody vine in upper canopy; stems initially densely tomentose with strongly 
twisted, dark brown hairs, eventually glabrescent. Lamina of larger leaves 6-9 cm 
long, 4.5-6 cm wide, ovate, cordate at base, mostly obtuse or rounded and some- 
times apiculate at apex, bearing all around the margin many sessile glands or, 
especially on smaller leaves in inflorescence, vascularized cilia up to 2.5 mm long, 
initially densely brown-tomentose on both sides but glabrate in age or persistently 
tomentose on midrib, especially below, the hairs with a short to long stalk and a 
long, twisted crosspiece, the 8-10 pairs of lateral veins sunken above and promi- 
nent below and connected by prominent + scalariform tertiary veins; petiole 3-5.5 
mm long, densely tomentose to eventually glabrescent, bearing 2 swollen glands 
1.3-1.8 mm in diameter at very base; stipules not found. Inflorescence densely 
brown-tomentose, paniculate sensu lato with the ultimate units 4-flowered umbels, 
each umbel sessile or subsessile in the axil of a much-reduced bract bearing 2 
large abaxial glands; peduncle absent, each pedicel subtended by a cluster of 1 
bract and 2 bracteoles, these appressed around base of pedicel, 0.5-1 mm long, 
broadly ovate, abaxially tomentose, adaxially glabrous, eglandular or the bracts (es- 
pecially those subtending the lower pair of flowers in each umbel) bearing (1-) 2 
abaxial glands, persistent; pedicel 6-8 mm long, tomentose, somewhat thicker 
distally. Sepals 2.5 mm long, 1.2-1.5 mm long beyond glands, 1.7-2 mm wide, 
ovate and rounded at apex, pressed in against filaments in anthesis, abaxially 
densely and uniformly appressed-tomentose, adaxially glabrous, the anterior eg- 
landular, the lateral 4 biglandular, the glands 1.6-1.8 mm long, elliptical or slightly 
obovate, separated on sepal, attached or slightly detached at apex, glabrous. Pet- 
als exposed in the enlarging bud, yellow, glabrous, abaxially keeled but not winged, 
spreading to reflexed; lateral petals with claw 1-1.5 mm long, Hmb 3-3.5 mm long 
and wide, orbicular or obovate, sometimes slightly lobed at base, decurrent onto 
claw, entire or minutely denticulate or erose, eglandular or with a small glandular 
area on margin at base; posterior petal similar to lateral 4 but with the claw longer 
(ca 2 mm) and the limb distinctly lobed at base with the lobes glandular at tip. 
Stamens glabrous; filaments connate for ca 0.5 mm, 1.8-2.5 mm long, longest 
opposite sepals and posterior-lateral petals, very stout opposite posterior-lateral 
petals, otherwise slender, especially opposite lateral sepals; anthers 1.3-1.7 mm 
long, largest opposite posterior-lateral petals, the connective dark red, equalling 
but not exceeding the locules. Ovary ca 1.5 mm high, densely brown-sericeous; 
styles 2.1-2.5 mm long, the anterior shorter, slenderer, and straight or slightly 
reflexed, the posterior 2 longer, stouter, somewhat sigmoid and distally reflexed, 
all 3 laterally flattened distally, with a distinctly internal stigma and apically "ped- 
ahform," i.e., with a flattened eUiptical top, this dorsally truncate or slightly apicu- 
late. Fruit unknown. 

The epithet of this species honors Bruce Hoffman (b. 1962), collector (with H. 
Jacobs and C. Capellaro) of the type and only known collection. 

Heteropterys hoffmanii is assignable to series Holopetalon (Griseb.) Nied., 
even when that group is defined narrowly as was done by Niedenzu in Das Pflan- 
zenreich (1928). This is the first collection I have seen of the series from the 
Guianas, nor has any previous author reported such a distribution. The species 
most like H. hoffmanii are H. ciliata Nied. and H. thyrsoidea (Griseb.) Adr. Juss. 



1997 

(including H. cordifolia Moric. ex Adr. Juss.). That complex, whose taxonomy is 
not well resolved, is known from Sao Paulo, Rio de Janeiro, Bahia, and near 
Manaus. Those plants always have the hairs on the abaxial side of the lamina 
stiffly erect, Y- or T-shaped with quite straight stalks and arms, and the hairs are 
persistent. The twisted, deciduous leaf hairs of H. hoffmanii would distinguish it 
at once from its sister species, even if it were not geographically disjunct. 

Heteropterys molesta W. R. Anderson, sp. nov.— Type: Venezuela. Bolivar: Cam- 
pamento "La Yagua" aproximadamente a 24 km NE del caserio Los 
Rosos, este ultimo a 17 km de Upata (sobre la carretera nueva Upata- 
San Felix), 16-25 Jun 1965 fl, C Blanco 134 (holotype: VEN!; isotypes: 
MO! US!). 

Liana ramis sericeis demum glabrescentibus. Lamina foliorum majorum 7.8- 
11.3 cm longa, 4-5.6 cm lata, elliptica ovatave, basi obtusa rotundatave, apice 
obtusa mucronulataque, margine glandulis parvis instructa, supra sericea demum 
glabrescens, subtus pertinaciter sericea; petiolus 7-9 mm longus, basi biglanduli- 
fer; stipulae minutae, interpetiolares. Inflorescentia pseudoracemus axillaris seri- 
ceus 8-17 cm longus, 20-60-florus; bracteae eglandulosae; pedunculus (0.5-) 1-3 
{-A) mm longus; bracteolae apicales, 1 bracteola cujusque paris 1 glandula abaxiali 
excentrica 0.7-0.8 mm diametro instructa; pediceilus 4-7 mm longus. Sepala per 
anthesin apice revoluta. Petala lutea, glabra, in alabastro exposita; petalum posti- 
cum limbo toto circuitu glanduloso-dentato. Stamina petalis posticolateralibus op- 
posita maxima. Styli apice dorsaliter truncati vel brevissime apiculati. 

Vine, the stems initially sericeous with fine, straight, white and golden hairs, 
eventually glabrescent. Lamina of larger leaves 7.8-11.3 cm long, 4-5.6 cm wide, 
elliptical or ovate, obtuse or rounded at base, mostly obtuse and mucronulate at 
apex, bearing many small bordered glands on very margin or on adaxial side of 
the eventually slightly revolute margin, adaxially loosely sericeous to eventually 
glabrescent with the hairs stramineous, 1-1.5 mm long, sessile or short-stalked, 
abaxially tightly and persistently sericeous but never so densely that the epidermis 
is completely concealed, the hairs on midrib and principal veins silvery, the hairs 
on lamina between veins golden, strongly parallel to each other, 0.2-1.2 mm long; 
lateral veins 6-8 pairs; petiole 7-9 mm long, persistently golden-sericeous, bearing 
2 glands ca 1 mm in diameter at very base; stipules ca 0.3 mm long, triangular, 
borne on stem beside petiole, often deciduous. Inflorescence an unbranched axil- 
lary pseudoraceme 8-17 cm long, sericeous with the hairs straight or sinuous, 
proximally silvery, distally golden, the flowers 20-60; bracts 0.9-1.5 mm long, 
narrowly triangular, abaxially sericeous, adaxially glabrous, eglandular, persistent; 
peduncle (0.5-) 1-3 (-4) mm long, golden-sericeous to glabrescent; bracteoles like 
bracts but ovate and 0.5-0.7 mm long, borne at apex of peduncle, one of each pair 
bearing 1 large eccentric abaxial gland 0.7-0.8 mm in diameter; pedicel 4-7 mm 
long, golden-sericeous. Sepals 1.5-2 mm long, ca 1 mm long beyond glands, 1-1.5 
mm wide, triangular, revolute at apex and sides in anthesis, abaxially densely and 
uniformly golden-sericeous, adaxially glabrous, the anterior eglandular, the lateral 
4 biglandular, the glands 1.5-2 mm long, narrowly elliptical, slightly detached but 
not reflexed at apex, glabrous. Petals exposed in the enlarging bud, yellow, gla- 
brous, abaxially smooth; lateral petals spreading to reflexed, with claw 1-1.5 mm 
long and limb 3-3.5 mm long, 3-3.2 mm wide, subrotund, eglandular and entire or 
slightly erose or denticulate at the margin; posterior petal erect or spreading, with 



60 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

claw 1.5-1.8 mm long and limb 3-3.2 mm long, 2.5-2.8 mm wide, elliptical, glandular- 
dentate all around the margin. Stamens glabrous; filaments connate at base, 1.5- 
2.3 mm long, longest opposite anterior sepal, shortest opposite posterior petal, very 
stout opposite posterior-lateral petals, very slender opposite lateral sepals; an- 
thers 0.8-1.2 mm long, largest opposite lateral petals, smallest opposite lateral 
sepals. Ovary ca 1 mm high, densely sericeous with white or white and brown 
hairs; styles 1.5-1.8 mm long, subequal, glabrous, erect, the anterior straight, the 
posterior 2 somewhat bowed, all 3 laterally flattened especially distally, dorsally trun- 
cate or more commonly very briefly apiculate at apex. Fruit unknown; enlarging 
carpels of oldest flowers apparently with a dorsal wing and without lateral wings. 

This species is known only from the type. Similar plants have been collected 
in Mato Grosso, Brazil, but they probably deserve recognition as a distinct spe- 
cies; I shall defer describing that in the hope of seeing fruits. 

The epithet of this peculiar species refers to the vexation it has caused me for 
the last 20 years. I have never doubted that it represents an undescribed species, 
but 1 have been unable to decide in which genus it should be described. The two 
obvious possibilities are Mascagnia and Heteropterys. Several characters argue for 
Mascagnia: glands at the base of the petiole, marginal lamina glands, interpetiolar 
stipules, long unbranched axillary pseudoracemes, one large gland on one of the 
bracteoles, and revolute sepals. All of these character-states occur in various spe- 
cies of Mascagnia, but never all together. For example, the sepals are only revo- 
lute in species in which the petals are completely concealed in bud, not exposed as 
they are here. The species whose lamina glands are most like those in Blanco 134, 
M. sinemariensis (Aubl.) Griseb., has petiole glands absent or borne well above 
the base, short crowded compound inflorescences, eglandular bracteoles, appressed 
sepals, and densely hairy petals. Search as I might, I cannot find a Mascagnia 
(even construing that polymorphic "genus" broadly) that I can advance as a close 
relative for Blanco 134. 

Heteropterys presents similar difficulties. Revolute sepals are always present 
in the subgenus in which the petals are concealed in bud, but very rare in the rest 
of the genus. Bracteoles with glands are rare in Heteropterys, and nearly unknown 
in species with marginal lamina glands. Heteropterys leschenaultiana Adr. Juss., of 
southeastern Brazil and adjacent Argentina, has glands at the base of the petiole, 
lamina glands that are marginal or submarginal, petals exposed in bud, and revo- 
lute sepals, but its stipules are epipetiolar, the inflorescence is compound with the 
flowers mostly in few-flowered umbels or corymbs, and the bracteoles are eglan- 
dular. Heteropterys sylvatica Adr. Juss., a species of Mato Grosso do Sul, Brazil, 
Santa Cruz, Bolivia, and Salta, Argentina, is the other most convincing candidate 
for a close relative to Blanco 134. In that the lamina glands, if they are present at 
all, are marginal, the inflorescence is a long simple axillary pseudoraceme, and the 
bracteoles bear glands (indeed, both are biglandular). However, the petiole is 
biglandular above the middle, the stipules are epipetiolar, and the sepals are not 
revolute. 

Fruits would decide the matter, but fruits are not available. The oldest flowers 
on Blanco 134 show enlarging carpels with the beginning of a dorsal wing that 
seems to be thicker on the abaxial edge, and no sign of lateral wings; that is clear 
support for Heteropterys. Therefore, I have decided to describe the plant in Het- 
eropterys. I suppose its relationships lie with H. leschenaultiana and its relatives or 
with H. sylvatica, but as I have tried to show above, H. molesta is rather isolated 
in Heteropterys — assuming it really does belong in Heteropterysl 



1997 W. R. ANDERSON: MALPIGHIACEAE 61 

Heteropterys sylvatica Adr. Juss., Ann. Sci. Nat. Bot., Ser. 2, 13: 277. 1840.— 
Type: Bolivia. Santa Cruz: "In sylvis interioribus juxta Santa-Crux de la 
Sierra," Mar fr, A. d'Orbigny (holotype: P! F neg. 35593, WRA negs. 81- 
27-4 & 5). 
Heteropterys tenuifolia (Nied.) Nied. in Engler, Pflanzenr. IV. 141: 333. 1928. 
Clonodia tenuifolia Nied., Meded. Rijks-Herb. 19: 74. 1913.— Type: Bra- 
zil. Mato Grosso do Sul: Corumba, 18 Dec 1902 fl/fr, A. Robert 736 
(lectotype, here designated: BM!; isolectotype: K!) 

In his 1928 treatment in Das Pflanzenreich, Niedenzu recognized his own 
species Heteropterys tenuifolia while placing Jussieu's name H. sylvatica among 
the species of whose identity he was uncertain, because he never had the opportu- 
nity to see Jussieu's type. Comparison of Niedenzu's syntypes to my photographs 
of and notes on Jussieu's type shows that the two names represent the same very 
distinctive species, of which I have now seen specimens from Santa Cruz, Bolivia, 
adjacent Mato Grosso do Sul, Brazil, and Salta, Argentina. The following diag- 
nostic characters make H. sylvatica easy to recognize: Ovate, glabrescent lamina 
raised on a well-developed petiole with two knobby glands near the apex; lamina 
eglandular or with small glands on the margin, not on the surface of the lamina; 
inflorescence a simple or ternate pseudoraceme in the axil of current leaves, the 
long narrow bracts deciduous in fruit; peduncle ca 5-6 mm long, bearing both 
bracteoles at its very apex; both bracteoles bearing 1 or 2 abaxial glands; pedicel 
only 1-2 mm long, much shorter than the peduncle; sepals erect; petals yellow; 
samara 3.5-4 cm long, the nut spherical and smooth-sided, the wing straight and 
distally flared to 1.5 cm wide. 

Niedenzu described a sister species, Heteropterys mollis (Nied.) Nied., also 
from lowland Bolivia. That seems to differ from H. sylvatica only in having the 
lamina abaxially very densely and persistently tomentose. It remains to be seen 
whether those two species will stand as distinct as more collections of the two 
accumulate. The one collection I have seen from Argentina {Abbiatti & Claps 205, 
from Vespucio, Oran, Salta [MO]) is somewhat intermediate in its vesture, but for 
now I am considering that to represent H. sylvatica. The name Heteropterys syl- 
vatica was misapplied by O'Donell & Lourteig (1943) to a very different species 
that is common in Argentina, Heteropterys dumetorum (Griseb.) Nied., of which 
the type came from Oran, Salta. 

Heteropterys velutina W. R. Anderson, sp. nov.— Type: Guyana. Siparuni-Potaro 
Region; Iwokrama Rainforest Reserve; Burro Burro River, between Sand- 
stone and confluence with Sipariparu River, 4°23'N, 58°55'W, riparian 
vegetation, 65 m, 31 Mar 1996 fr, D. Clarke 1541 (holotype: MICH!). 

Arbor 6 m alta ramis dense et pertinaciter velutinis. Lamina foliorum ma- 
jorum 9.5-1 1.7 cm longa, 2.5-3.3 cm lata, anguste elliptica, basi cuneata vel subro- 
tundata, apice acuminata, supra glabrata, subtus + pertinaciter tomentosa pilis T- 
formibus; petiolus 4-5 mm longus, velutinus. Inflorescentia panicula ex pseudora- 
cemis 6-14-floris constans, velutina; bracteae deciduae; pedunculus 0.5-2 mm lon- 
gus, velutinus; bracteolae 2.5-3.5 mm longae, 1.8-2.2 mm latae, eglandulosae, 
reticulo adaxialiter visibili; pedicellus 5-7 mm longus (in fructu), velutinus vel 
subtomentosus. Sepala omnia eglandulosa, apice revoluta. Samara 15-23 mm longa, 
8-10 mm lata, saepe reflexa, ala dorsali proximaliter ultra nucem producta. 



62 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Tree 6 m tall; stems densely and persistently velutinous, the hairs brown, erect 
and straight, 0.4-1.2 mm long, many basifixed but some Y-shaped or bearing a 
rudimentary second branch near the middle. Lamina of larger leaves 9.5-11.7 cm 
long, 2.5-3.3 cm wide, narrowly elliptical or widest sUghtly above or below the 
middle, cuneate to nearly rounded at base, mostly long-acuminate at apex, bear- 
ing in the abaxial surface, set in 1-3 mm from both margins, a row of 3-7 small 
impressed glands, adaxially quite glabrate at maturity or persistently tomentose 
on margin or base of midrib, abaxially densely and persistently tomentose or 
eventually patchily glabrescent with the hairs T-shaped with the stalk 0.1-0.15 mm 
long and the sinuous crosspiece 0.6-1 mm long, the 6-9 pairs of lateral veins 
somewhat sunken above between raised sections of lamina and prominent below, 
the reticulum prominent on both sides; petiole 4-5 mm long, persistently veluti- 
nous hke stem, eglandular; stipules not found, to be sought among hairs on inter- 
petiolar ridge. Inflorescence a terminal panicle, with the flowers borne in decus- 
sate pseudoracemes 2-5 cm long and containing 6-14 flowers, all the axes densely 
and persistently velutinous like stem; florifcrous bracts deciduous in fruit and 
mostly not seen, apparently 3.5-4 mm long, 3-3.5 mm wide, broadly ovate, con- 
cave, eglandular, appressed-tomentose on both sides but more densely so on abaxial 
surface, the reticulum visible on adaxial side; peduncle 0.5-2 mm long, velutinous; 
bracteoles apical, resembling bracts but smaller (2.5-3.5 x 1.8-2.2 mm), more 
narrowed at base and thus elliptical, nearly flat, deciduous in fruit or irregularly 
persistent; pedicel 5-7 mm long in fruit, velutinous or subtomentose. Flowers not 
seen; sepals all eglandular, ca 3 mm long and 2 mm wide, strongly revolute distally. 
abaxially appressed-tomentose, adaxially glabrous; old petals glabrous, the claw 
ca 3 mm long, the hmb 4.2-5 mm long, 3-4 mm wide, broadly elliptical to rotund, 
erose; filaments glabrous, partially connate; ovary densely velutinous; styles 3.5-4 
mm long in fruit, with an internal stigma and a dorsal hook ca 0.5 mm long at 
apex, the hook pedaliform (i.e., flattened and elliptical when viewed from above). 
Samara immature but probably full-sized, 15-23 mm long, 8-10 mm wide, ellipti- 
cal or somewhat wider distally than proximally, borne horizontally or reflexed; 
dorsal wing extended proximally over apex of nut; nut 6-9 mm long, 4-5 mm high, 
mostly bent downward, without lateral wings or crests. 

This distinctive species is named for its velvety stem and inflorescence. It is 
known only from the type. Heteropterys velutina belongs in subgenus Parabanisteria 
(C. V. Morton) C. V. Morton, and there it is to be compared to other species that 
bear elongated pseudoracemes rather than umbels. Its closest relative is probably 
H. subheUcina Nied. (=//. catoptera W. R. Anderson), a shrub or woody vine of 
savannahs and riverine forests in southern Guyana and adjacent Roraima, Brazil. 
They are especially similar in the samara, which in both species has the nut bent 
downward, producing an often reflexed orientation of the whole samara, and the 
dorsal wing in both extends over the apex of the nut as a rounded flange. They 
also have similar large bracteoles with the reticulum visible adaxially. However, in 
H. subheUcina the vegetative axes are sparsely sericeous to glabrate and the inflo- 
rescence, peduncles, and pedicels are persistently sericeous— this dramatic differ- 
ence in vesture gives the two species very different aspects. The leaf in H. subhe- 
Ucina is sparsely sericeous to quite glabrate, the base of the lamina is more rounded, 
and the apex is obtuse, acute, or only slightly acuminate. When H. velutina is 
found with flowers, additional differences may be evident. 

The label with the type says that the plant was a tree 6 m tall. Habit is 
variable in other species in this subgenus, so it will not surprise me if H. veUitina is 
also found growing as a shrub and woody vine. 



1997 W. R. ANDERSON: MALPIGHIACEAE 63 

In the holotype, one side of the stem is persistently velutinous, while the 
other side has lost most of the hairs. I have never seen such one-sided glabres- 
cence in a malpighiaceous stem before, and suspect that in this case the side with 
few hairs may have had the vesture scraped off during collection. If H. velutina 
really loses its stem hairs in that pattern, that will be yet another way in which it 
differs from its congeners. 

Hiraea buntingii W. R. Anderson, sp. nov. — Type: Venezuela. Trujillo: Distr. 
Betijoque, carretera Agua Viva-Carora, entre Agua Viva y Valerita, en 
bosque seco premontano, 300 m, 1 Nov 1977 fl/fr, G. S. Bunting 5845 
(holotype: MICH!; isotype: VEN!). 

Frutex vel liana ramis sericeis. Lamina foliorum majorum 6.5-16.5 cm longa, 
3-7.3 cm lata, abaxialiter sericea, margine aliquot glandulis parvis instructa; petio- 
lus 5-11 mm longus, sericeus, distaliter biglandulosus; stipulae (2-) 3-4 mm lon- 
gae. Cyma axillaris ex (1-) 3-7 umbelhs 4-floris constans; bracteae 2.5-5 mm 
longae, 1.3-2 mm latae; bracteolae 1.5-2.5 mm longae, 1-1.5 mm latae; pedicellus 
9-23 mm longus, gracilis. Glandulae calycis saepe brevistipitatae. Petala eglandu- 
losa. Antherae 0.5-0.9 mm longae. Samara alis lateralibus 12-15 mm latis, 16-21 
mm altis, ala dorsali 1.5-3 mm lata, 3.5-5.5 mm alta, in dentibus acutis dissecta. 

Shrub 1.6 m tall or woody vine in treetops; stems densely sericeous with a 
persistent mixture of silvery and golden or brownish hairs, eventually glabrescent 
as stem enlarges through secondary growth. Lamina of larger leaves 6.5-16.5 cm 
long, 3-7.3 cm wide, elliptical or widest slightly above middle, broadly cuneate, 
rounded, or slightly cordate at base, obtuse or rounded and usually short-apicu- 
late at apex, bearing several small button-like glands evenly distributed on distal 
2/3 of margin, initially sericeous above but soon glabrescent and eventually quite 
glabrate or with some hairs persistent proximally on midrib, moderately to dense- 
ly and persistently sericeous below or eventually glabrescent, the principal lateral 
veins 8-10 pairs, prominent below but not above, interconnected by scalariform 
tertiary veins; petiole 5-11 mm long, sericeous like stem, biglandular between 
middle and apex; stipules (2-) 3-4 mm long, subulate, sericeous, borne well above 
base of petiole but mostly below middle, oriented at right angles to petiole and 
parallel to each other, straight or decurved. Inflorescence a sericeous axillary 
cyme of (1-) 3-7 4-flowered umbels, each cyme usually solitary in its axil, occa- 
sionally subtended by a second smaller one; umbel without a gland in center, each 
borne on a stalk 4-15 (-22) mm long; bracts 2.5-5 mm long, 1.3-2 mm wide, ovate 
or lanceolate and somewhat boatshaped, abaxially sericeous, adaxially glabrous or 
very sparsely sericeous, eglandular, persistent; bracteoles similar to subtending 
bract but only about half as long, 1.5-2.5 mm long, 1-1.5 mm wide, elliptical; 
pedicel 9-19 mm long (-23 mm in fruit), very slender, thinly sericeous to glabrate. 
Sepals 2-2.5 mm long, 1.5-2 mm wide, ovate or elliptical, rounded at apex, ap- 
pressed in anthesis, abaxially sericeous, adaxially glabrous, the anterior eglandu- 
lar or bearing 1 or 2 small glands, the lateral 4 biglandular, the glands 0.9-1.5 mm 
long, circular or elliptical, separated from each other, shghtly detached all around 
or (especially in age) short-stalked. Petals yellow, glabrous, the limb + orbicular, 
erose or short-laciniate, eglandular; lateral 4 petals spreading to reflexed, with the 
claw 2-2.5 mm long and the limb 5-6 mm long and wide; posterior petal similar to 
the lateral 4 but more nearly erect, with a slightly longer claw (2.5-3 mm long) 
and a smaller limb (4-5 mm long and wide), the margin more deeply dissected. 
Stamens glabrous; filaments connate for 1-1.5 mm, the anterior 3 longest (3.5^.5 



64 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

mm long), the posterior 7 2.5-3.5 mm long, shortest opposite the posterior petal; 
anthers 0.5-0.9 mm long, similar to each other, the locules separated on a massive 
glandular connective. Ovary ca 1.5 mm high, densely hispid; styles with a few hairs 
proximally or glabrous, short-apiculate dorsally at apex; anterior style 2.5-3 mm 
long, erect and straight or slightly bowed toward posterior petal; posterior styles 
ca 3 mm long, strongly bowed outward and then back toward center of flower. 
Samara butterfly-shaped; lateral wings membranous, 12-15 mm wide, 16-21 mm 
high, roughly tetrahedral to semicircular, subentire or sinuous to rounded-erose, 
thinly but persistently sericeous on both sides with fine, sessile, appressed hairs; 
dorsal wing 1.5-3 mm wide, 3.5-5.5 mm high, widest near style, irregularly but 
deeply dissected into sharp teeth, hispid hke nut; nut globose, 2.5-3 mm in diame- 
ter, hispid with long spreading sub-basifixed hairs or belatedly glabrescent. 

As often happens in this genus, the filaments are not straight or parallel, but 
bent and even twisted in a complex but bilaterally symmetrical pattern. Because 
all flowers now available are rather far along in anthesis, I shall not try to describe 
the bending of the filaments here, but defer that until younger material is available. 

Additional Specimens Examined: Venezuela. Trujillo: Agua Viva, carretera de Maracaibo, dry 
steep slopes, Aug fl, Tamayo 1767 (US, VEN).— Zulia: Distr. Miranda-Bolivar, via El Consejo, La 
Tabla-Quiros-El Pensado, en Corral de Nava (2 km al NE de La Tabla), in gallery forest and 
adjacent chaparral near stream, 140 m, Feb fl. Bunting 8633 (MICH). 

The epithet of this interesting species honors George S. Bunting (b. 1927), 
who collected the type and one of the paratypes during the course of many years 
of botanizing in northern Venezuela. 

This species belongs in subg. Hiraea (see discussion below under H. hypoleuca). 
The most obvious feature distinguishing Hiraea buntingii is the unusually long 
bracts and bracteoles, larger than any others I have seen in the genus. In addition, 
the species differs from others in the following combination of characteristics: 
lamina abaxially sericeous, with a rounded or slightly cordate base and a row of 
marginal glands; axillary cyme relatively open (for Hiraea) with the umbels raised 
on well-developed stalks; pedicel long and slender; calyx glands detached to short- 
stalked; posterior petal eglandular; anthers small; dorsal wing of samara dissected 
into sharp teeth. Hiraea buntingii bears a resemblance to H. sanctae-marthae C. V. 
Morton, from nearby Colombia, because that too has the lamina abaxially seri- 
ceous, but H. sanctae-marthae has the lamina shape and veins of H. reclinata Jacq. 
and the compact cyme and short bracts and bracteoles typical of the genus. 

Bunting 8633 was a vine, whereas the other two collections were shrubs, and 
its leaves are substantially larger than theirs. However, the three agree well in 
characters of the inflorescence, flowers, and fruits, so I think they must all repre- 
sent the same species. 

Hiraea hypoleuca W. R. Anderson, sp. nov.— Type: Venezuela. Tachira: region 
of Cerro Las Minas, S of main road from Santa Ana, 17 km SE of Santa 
Ana, pastured ridge with bordering forest, 1150-1250 m, 11 Nov 1979 fr, 
/. A. Steyermark, R. Liesner & A. Gonzalez 119901 (holotype: MICH!; 
isotype: VEN!). 

Liana ramis sericeis. Lamina foliorum majorum 8.5-13.5 cm longa, 5-9 cm 
lata, elliptica vel late elliptica vel paulo obovata, basi rotundata vel subcordata, 
apice rotundata vel latissime obtusa, adaxialiter glabrata, abaxialiter appresso- 



1997 W. R. ANDERSON: MALPIGHIACEAE 65 

tomentosa glaucaque; petiolus 10-25 mm longus sericeus; stipulae 1-1.5 mm lon- 
gae. Umbella axillaris 10-30-flora; bracteae bracteolaeque ca 1 mm longae, rotun- 
datae; pedicellus 18-25 mm longus. Petalum posticum distaliter glanduloso-fim- 
briatum. Styli postici in fructu arcuati. Samara alis lateralibus 13-18 mm latis, 20- 
30 mm altis, ala dorsali 1-3 mm lata, 3-6 mm alta. 

Woody vine; stems densely and persistently sericeous. Lamina of larger leaves 
8.5-13.5 cm long, 5-9 cm wide, elliptical or broadly elliptical or somewhat obo- 
vate, rounded or subcordate at base, rounded or very broadly obtuse and some- 
times apiculate at apex, eglandular or bearing several tiny button-like glands just 
within margin on abaxial side, initially appressed-tomentose above but glabrate at 
maturity or with some hairs persistent proximally on midrib, persistently appressed- 
tomentose below or belatedly glabrescent except for the ± persistently sericeous 
midrib and lateral veins, thinly to densely glaucous below, the principal lateral 
veins 7-9 pairs, prominent below but not above, interconnected by scalariform 
tertiary veins; petiole 10-25 mm long, sericeous like stem, bearing 2 (-4) small (ca 
0.5 mm diam.) glands between middle and apex, mostly at apex or just above apex 
against midrib of lamina; stipules 1-1.5 mm long, subulate, sericeous, borne above 
base of petiole but below middle, mostly curved away from stem, parallel or bent 
toward each other. Inflorescence axillary, an umbel of 10-30 flowers borne on a 
sericeous stalk 11-33 mm long, 1.5-2 mm in diameter, jointed 1-12 mm below 
umbel and bearing a pair of deciduous or persistent bracts or much-reduced leaves 
at the joint; floriferous bracts and bracteoles ca 1 mm long, wider than long, 
broadly rounded, abaxially sericeous, adaxially glabrous, eglandular, persistent; 
pedicel 18-25 mm long, 1-1.5 mm in diameter (-2 mm at apex), persistently seri- 
ceous. Good flowering material not seen. Sepals 2-2.5 mm long and 1.5-2 mm 
wide in flower, up to 3.5 x 2.5 mm in fruit, ovate or triangular, obtuse to rounded 
at apex, appressed in anthesis, abaxially sericeous, adaxially glabrous, the anterior 
eglandular, the lateral 4 biglandular, the glands 1.3-2.5 mm long, elhptical, sepa- 
rated from each other or compressed. Petals yellow, glabrous, the lateral 4 erose 
or dentate, the posterior glandular-fimbriate in the distal 2/3 of the limb. Stamens 
glabrous; filaments connate at base, ca 2.7-3.2 mm long. Ovary densely hispid; 
posterior 2 styles strongly bowed (at least eventually), laterally slightly flattened, 
dorsally truncate or apiculate at the apex. Samara butterfly-shaped; lateral wings 
membranous, 13-18 mm wide, 20-30 mm high, trapezoidal to flabellate, subentire 
or sinuous to rounded-erose, persistently thinly sericeous or appressed-tomentose 
on both sides; dorsal wing 1-3 mm wide, 3-6 mm high, entire or irregularly dis- 
sected, usually not extending to base of nut, tomentose; nut globose, 4-5 mm in 
diameter, tomentose or hispid with straight spreading hairs. 

Additional Specimens Examined: Colombia. Antioquia: Mpio Anori, Vereda El Carmen, 18- 
20.5 km NW de Anori en la via a "Dos Bocas," 7°15'N, 75°12'W, 820-910 m, Nov fr, Callejas et al. 
8802 (MICH). — Santander: Mpio Charala, Corrig. Virolin, "Canaverales," liana in tree at edge of 
pasture, 1680 m, Jan fr, Albert de Escobar et al. 3060 (MICH). Venezuela. Tachira: Distr. Uribante, 
Complejo Hidroelectrico Uribante-Caparo, Sector El Cedral, inmediaciones Presa La Honda, S^Ol'N, 
7r37'W, low montane forest, 1100-12.50 m, Nov fl, Rivero et al. 1920 (MICH, PORT). 

Hiraea hypoleuca is named for the glaucescence on the abaxial surface of its 
lamina, which is much thicker in the Venezuelan specimens than in the Colombi- 
an specimens cited. I know of no other species of Hiraea with many-flowered 
umbels and glaucous leaves. It is also notable for the relatively small leaf that is 
rounded at the apex, short stipules, very short rounded bracts and bracteoles, and 
glandular-fimbriate posterior petal. 



66 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

I divide Hiraea into two subgenera. Subgenus Hiraea lias the flowers borne in 
umbels of four, with the umbels often borne in a compound, cymose, axillary 
inflorescence, or several such inflorescences in the same axil. The other subgenus 
is Archihiraea Nied., to which I would assign all species in which the axillary 
inflorescence is unbranched and bears a single umbel of 5-60 flowers (but see the 
discussion below under H. perplexa). Cuatrecasas (1958) distributed the species of 
subg. Archihiraea between sect. Archihiraea (Nied.) Cuatrec. and sect. Polyactinia 
Nied., on the basis of the shape of the mature styles. The lack of open flowers 
makes it difficult for me to assign //. hypoleuca to one or the other with confi- 
dence, but the strong curvature of the posterior styles in fruit suggests that it 
probably belongs in sect. Polyactinia. 

Hiraea neblinensis W. R. Anderson, sp. nov.— Typk: Venezuela. Amazonas: Depto. 
Rio Negro, Rio Baria (=Rio Mawarinuma) just upstream from Base Camp, 
SW side of Cerro de la Neblina, (r49'50"N, 66°09'40"W, tropical lowland 
evergreen forest along gravelly and silty banks of river, 140 m, 17 Feb 
1985 fl, M. Nee 30920 (holotype: MICH!; isotypes: NY! VEN!). 

Liana lignosa ramis sericeis mox glabratis. Lamina foHorum majorum 13.5-31 
cm longa, 8.5-17 cm lata, late elliptica vel obovata, basi paulo cordata, apice 
rotundata vel late obtusa vel abrupte breviacuta, margine eglandulosa, utrinque 
mox plus minusve glabrata; petiolus 9-19 mm longus; stipulae 2-3 mm longae, 
super medium petioh portatae. Cyma axillaris plerumque ex 3 umbelhs 4-floris 
constans; umbella sine glandula centrali; bracteae bracteolaeque 1-1.2 mm lon- 
gae, 0.8-1.2 mm latae, eglandulosae; pedicellus 11-20 mm longus. Sepala adaxi- 
aliter sparsim sericea. Petala omnia limbo transverse late elliptico, fimbriato fim- 
briis 0.5-1 mm longis eglandulosis. Filamenta sepalis opposita 3.1-3.6 mm longa, 
petalis opposita 2.3-3 mm longa; antherae 0.9-1.3 mm longae. Stylus anticus apice 
dorsaliter obtusus ca 0.2 mm productus, 2 styh postici apice dorsahter breviunci- 
nati unco ca 0.3 mm longo rotundato. Samara immatura ahs lateralibus semicircu- 
laribus, ala dorsaU bene evoluta. 

Woody vine in trees; stems initially sericeous, soon glabrate. Lamina of larger 
leaves 13.5-31 cm long, 8.5-17 cm wide, broadly elliptical or somewhat obovate, 
shallowly cordate at base, rounded or broadly obtuse at apex or abruptly nar- 
rowed to a very short acute tip, eglandular on margin, initially sericeous but soon 
nearly glabrate, often with some hairs persisting on abaxial midrib and sometimes 
with sparse short appressed hairs scattered on abaxial surface, the principal lateral 
veins 8-10 pairs, prominent below but not above, interconnected by many closely 
spaced scalariform tertiary veins; petiole 9-19 mm long, abaxially sericeous, adax- 
ially velutinous, glabrescent in age, bearing 2 large glands at apex or just above apex 
against midrib of lamina; stipules 2-3 mm long, subulate, sericeous, borne slightly 
above middle of petiole, oriented at right angles to petiole and parallel to each 
other, distally ascending. Inflorescence a sericeous axillary cyme of (1-) 3 4-flow- 
ered umbels, the cymes solitary in the axil or 2-3 in a vertical array; umbel with- 
out a gland in the center, each borne on a stalk 2-5 mm long; bracts 1-1.2 mm 
long and wide, triangular or ovate, abaxially sericeous, adaxially glabrous, eglan- 
dular, persistent; bracteoles similar to subtending bract but smaller, ca 1 mm long 
and 0.8 mm wide; pedicel 11-20 mm long, loosely sericeous. Sepals 2-2.5 mm 
long, 1-1.5 mm beyond glands, 1-1.5 mm wide, ovate, obtuse or acute at apex, 
appressed in anthesis, abaxially densely sericeous, adaxially sparsely sericeous, the 



1997 W. R. ANDERSON: MALPIGHIACEAE 67 

anterior eglandular, the lateral 4 biglandular, the glands 1.3-2 mm long, elliptical, 
distally detached. Petals yellow, glabrous, the limb transversely broadly elliptical 
and long-fimbriate with the divisions 0.5-1 mm long and eglandular; lateral 4 
petals with the claw 2.5-3 mm long, the Hmb 4.5-6 mm long, 5.5-7.5 mm wide; 
posterior petal with the claw 3.5 mm long, the limb 4.5-5 mm long, 5-5.5 mm 
wide. Stamens glabrous; filaments connate for 0.7-1 mm at base, mostly bowed, 
3.1-3.6 mm long opposite sepals with the longest opposite the anterior sepal, 2.3- 
3 mm long opposite petals with the shortest opposite the posterior petal; anthers 
0.9-1.3 mm long, slightly longer opposite petals than opposite sepals, with the 
connective proximally red and distally yellow in dried material. Ovary ca 1.2 mm 
high, densely hispid; styles tomentose on proximal half; anterior style ca 3.5 mm 
long, moderately bowed, with an obtuse dorsal projection at apex ca 0.2 mm long; 
posterior 2 styles longer (ca 4 mm) but more strongly bowed, bearing a rounded 
dorsal hook at apex ca 0.3 mm long. Immature samara with the lateral wings 18 
mm wide, 30 mm high, semicircular to almost reniform with the rounded edges 
meeting or overlapping at base and apex, entire or slightly sinuous, appressed- 
tomentose with most hairs medifixed; dorsal wing 5 mm wide, 7 mm high, rectan- 
gular, entire or subentire, appressed-tomentose; nut slightly depressed-globose 
(?), ca 5 mm in diameter, tomentose with many hairs medifixed. 

Additional Specimens Examined: Venezuela. Amazonas: Depto. Rio Negro, at west or lower 
end of Rio Mawarinuma, 4 km by air west of Cerro de la Neblina Base Camp, 0°50'N, 66°12'W, 140 
m, Mar imm fr, Liesner & Funk 16380 (MICH, MO, VEN). 

Hiraea neblinensis is named for the Cerro de la Neblina, at the base of which 
were found the only known collections. It is assignable to subgenus Hiraea; see 
the discussion above under H. hypoleuca. It is especially notable for its large 
leaves and petals with a long-fimbriate limb that is wider than long. It may be 
compared to two other species of southern Venezuela in which the lamina is large 
and has an eglandular margin, H. celiana W. R. Anderson and H. affinis Miq. 
Hiraea celiana is known from only three collections from Cerro Cuao-Sipapo. Its 
lamina is cuneate at the base, its sepals are adaxially glabrous, and its styles are 
dorsally rounded or truncate at the apex. Its fruit is not known. Hiraea affinis is 
an extremely variable species that is fairly well collected in the Guianas but has 
been found only twice in Venezuela, in Delta Amacuro. Its petiole is shorter, up 
to 8 mm long, its sepals are adaxially glabrous, and its petals are dentate to short- 
fimbriate, with the divisions up to 0.2 mm long. The samara of H. affinis lacks a 
dorsal wing but is similar in outline to that of H. neblinensis, with rounded lateral 
wings that meet or overlap at the base and apex, which leads me to think that they 
may be closely related. 

Hiraea perplexa W. R. Anderson, sp. nov.— Type: Ecuador. Napo: Canton Tena, 
Estacion Biologica Jatun Sacha, Rio Napo, 8 km al E de Misahualli, 
1°04'S, 77°36'W, 400 m, bosque muy hiimedo tropical, bosque primario, 
colinas, 3-6 Sep 1989 fl, W. Palacios 4384 (holotype: MICH!). 

Liana lignosa ramis velutinis. Lamina fohorum majorum 10.5-13.5 cm longa, 
5-7.2 cm lata, abaxialiter pertinaciter velutina; petiolus 9-12 mm longus, veluti- 
nus, prope apicem biglandulosus; stipulae 3-5 mm longae. Cyma axillaris basi 
ternata, ex 3 umbelhs 4-8-floris constans, saepe 2 floribus proximalibus additis; 



68 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

pedicellus 15-22 mm longus, gracilis. Sepala eglandulosa, abaxialiter hispida. Pet- 
alum posticum margine distali glandulis parvis globosis instructum. Ovarium den- 
sissime hispidum pilis brunneis, 1.5-3 mm longis, rectis, ut videtur basifixis sed re 
vero plerumque calcari basali instructis. 

Woody liana; stems densely and persistently velutinous with the hairs erect, 
1-1.5 mm long, mostly bifurcate but some simple. Lamina of larger leaves 10.5- 
13.5 cm long, 5-7.2 cm wide, elliptical or widest shghtly above middle, broadly 
cuneate or rounded at base, obtuse or rounded at apex with the midrib sometimes 
extended slightly beyond lamina, bearing several small button-hke glands on mar- 
gin or just within margin on abaxial side, initially velutinous above with erect 
bifurcate hairs but glabrate at maturity except for midrib, margin, and scattered 
hairs elsewhere, densely and persistently velutinous below with most hairs erect 
and bifurcate, the hairs even denser on midrib and on the 8-10 pairs of major 
lateral veins than elsewhere, the veins, including the scalariform tertiary veins, 
sunken above and prominent below; petiole 9-12 mm long, velutinous like stem, 
biglandular near apex; stipules 3-5 mm long, subulate, velutinous to loosely seri- 
ceous, borne between base and middle of petiole, curved back toward stem. Inflo- 
rescence a velutinous, axihary, basally ternate cyme of 3 umbels, the lateral um- 
bels occasionally not developing; umbel without a gland in center, comprising 4-8 
flowers plus often an additional pair on the stalk well below the rest, the stalk 2- 
10 mm long; bracts 1.5-2.5 mm long, ca 1 mm wide, ovate or elliptical, abaxially 
sericeous, adaxially glabrous, eglandular, persistent; bracteoles like bracts but short- 
er and narrower, narrowly elliptical or hnear; pedicel 15-22 mm long, very slen- 
der, subvelutinous or appressed-tomentose. Sepals ca 2 mm long, 1.5 mm wide, 
ovate, obtuse or acute at apex, appressed in anthesis, abaxially densely appressed- 
hispid, adaxially glabrous, all eglandular. Petals yellow, glabrous; lateral 4 petals 
with the claw 2-3 mm long, the limb 4.5-5 mm long and 4-4.5 mm wide, ovate or 
obovate, erose or denticulate, eglandular; posterior petal with the claw 3-3.5 mm 
long and the limb 4-5 mm long, 3.5-4 mm wide, elliptical or obovate, dentate or 
short-laciniate with the divisions on the distal 1/2-2/3 each terminating in a tiny 
globose gland. Stamens glabrous; filaments connate at very base, longer opposite 
sepals (3.5-4.5 mm) than petals (2.7-3.5 mm), mostly bowed, especially opposite 
the posterior-lateral sepals; anterior 7 anthers 0.9-1.1 mm long with the glandular 
connective swohen, posterior 3 anthers 0.7-0.9 mm long with the connective glan- 
dular but not much enlarged. Ovary almost 1 mm high but completely hidden by 
the very dense covering of straight brown hairs 1.5-3 mm long, these apparently 
basifixed but most with a tiny basal spur representing the missing branch; styles 
bearing scattered long hairs on proximal half, the apex with a short acute dorsal 
projection ca 0.2 mm long; anterior style 3.5-4 mm long, moderately bowed; pos- 
terior 2 styles longer (4.5-5.5 mm) but more strongly bowed so that all 3 stigmas 
are at same level. Fruit not seen. 

Hiraea perplexa is known only from the type. Its epithet refers to the puzzling 
architecture of the inflorescence. Its basally ternate structure suggests that the 
species belongs in subgenus Hiraea (see discussion above under H. hypoleiica), 
but most species of that subgenus are very consistent in that each umbel contains 
exactly four flowers, whereas they usually contain more in this species, up to 
eight. Moreover, in many species of the subgenus the two bracts near the middle 
of the stalk of each umbel subtend an additional pair of umbels; in H. perplexa 
they generally subtend a pair of flowers, each with two bracteoles just like the 
other flowers in the umbel. It seems likely that the inflorescence in subg. Hiraea is 



MALPIGHIACEAE 69 

ancestral in the genus, because the sister genus Excentradenia (see separate article 
in this volume) also bears four-flowered umbels. Perhaps H. perplexa indicates 
how that cyme of four-flowered umbels gave rise to the unbranched multiflow- 
ered umbel of subg. Archihiraea — through increase in the number of flowers in 
each umbel, followed by suppression of the lateral umbels. Hiraea perplexa is also 
notable for the velutinous stems and leaves, the eglandular hispid sepals, the small 
petals, the small globose glands around the margin of the posterior petal, and the 
very densely hispid ovary. 

Hiraea steyermarkii W. R. Anderson, sp. no 
va Forestal "La Paragua," margem 
Blanco 803 {\io\oiypQ:¥\). 

Liana lignosa ramis sericeis vel velutinis demum glabratis. Lamina foliorum 
majorum 14-23.5 cm longa, 7-13.7 cm lata, obovata, basi cuneata vel obtusa, apice 
abrupte acuminata, margine glandulis prominentibus munita; petiolus 14-20 mm 
longus; stipulae 4.5-6 mm longae, plerumque inter basim et medium petioli porta- 
tae. Cyma axillaris ex 3-7 umbellis 4-floris constans, laxe sericea vel subtomentosa 
pilis cinereis vel stramineis; umbella sine glandula centrali; pedicellus 14-28 mm 
longus. Petala ut videtur lutea; petala lateralia limbo integro vel eroso eglandu- 
loso; petalum posticum limbo fimbriato fimbriis ± glandulosis. Styli glabri. Samara 
immatura ala dorsali bene evoluta, grosse dentata. 

Woody vine; stems sericeous or velutinous to glabrate. Lamina of larger leaves 
14-23.5 cm long, 7-13.7 cm wide, obovate, cuneate to obtuse at base, abruptly 
rounded and acuminate at apex to a short tip 5-10 mm long, bearing a series of 
large prominent button-like glands on distal 1/2-2/3 of margin, initially sericeous 
but soon glabrate except for ± persistently sericeous abaxial midrib, or with some 
hairs persistent on abaxial surface as well, the principal lateral veins 8-12 pairs, 
prominent below but not above, interconnected by scalariform tertiary veins; peti- 
ole 14-20 mm long, sericeous or velutinous, bearing 2 large bulging glands at or 
just below apex; stipules 4.5-6 mm long, subulate, borne between base and middle 
of petiole or slightly higher, generally reflexed. Inflorescence a cyme of 3-7 4- 
flowered umbels, loosely sericeous or subtomentose with gray or gray and straw- 
colored hairs, the cymes solitary in the axil or 2 in a vertical array, each cyme 
borne on a stalk 3-7 mm long; umbel without a gland in the center, each borne on 
a stalk 7-22 mm long; bracts 1-1.5 mm long and wide, triangular or ovate, abaxial- 
ly sericeous, adaxially glabrous, eglandular, persistent; bracteoles similar to sub- 
tending bract but usually smaller, 1-1.2 mm long, ca 0.8 mm wide; pedicel 14-22 
mm long in flower, up to 28 mm long in immature fruit, loosely sericeous. Flowers 
known only in age, with enlarging fruits. Sepals ca 2 mm long and 2-2.5 mm wide, 
broadly triangular or ovate, obtuse at apex, appressed in anthesis, abaxially densely 
sericeous except glabrous near margin, adaxially glabrous, all eglandular. Petals 
apparently yellow, glabrous; lateral 4 petals with the claw 2.5 mm long, the limb 
4.5-5 mm long, 5.5-6 mm wide, concave, entire or arose, eglandular; posterior 
petal with the claw 3-3.5 mm long, the limb 4.5-5 mm long and wide, flat or 
crumpled, fimbriate all around the margin with the divisions ca 0.5 mm long and 
glandular-thickened, especially proximally. Stamens glabrous; filaments bowed; 
anthers 1-1.4 mm long. Styles strongly bowed, bearing a rounded dorsal projec- 
tion at apex only 0.1 mm long. Immature samara with the lateral wings elliptical 
or semicircular, entire or slightly sinuous, initially sericeous but glabrescent as 



. UNIVERSITY OF MICHIGAN HERBARIUM 



they enlarge, the hairs 0.5-1 mm long, medifixed, straight and appressed; do 
wing 1-3 mm wide, 1.8-2 mm high, triangular to rectangular, mostly dissected i 
several coarse teeth, sericeous to glabrate. 



Ad 


iDiTFONAL Specimen Examined. Venezuela. Bolivar: Altiplanicie de 


between 




Chichar 


ras, 47 km N of Tumeremo, 100-250 m, Feb sterile, Steyermark 89349 ( 



I name this species in honor of my friend, the late Juhan A. Steyermark 
(1909-1988), whose accomplishments in Venezuelan botany need no description 
by me. The holotype and paratype have leaves that are similar in shape and 
glands, but those of Steyermark 89349 are substantially larger than those of Blanco 
803 (22-23.5 x 12.7-13.7 cm in the Steyermark collection vs. up to 14.5 x 7.4 cm in 
the Blanco collection). Moreover, the stems of the Steyermark collection are 
velutinous, while those of the type are sericeous. The two collections may repre- 
sent different species, but it is not rare for lianas to have smaller leaves in the 
inflorescence than lower on the stems, so the Steyermark collection's large leaves 
may simply reflect the fact that they were collected from a sterile plant; I cannot 
assess the importance of the difference in vesture with so little material. Although 
the type is fertile, it is in an intermediate stage, with the remnants of old flowers and 
very immature fruits, such that our knowledge of characters of both flowers and fruits 
is incomplete and will have to await additional collection for supple: 

Among the species of Hiraea known to occur in Venezuela, H. j 
a strong resemblance to H. celiana W. R. Anderson, because their laminas are similar 
in shape, cuneate at the base, not at all rounded or cordate, and they both have 
unusually long pedicels. However, the two are easy to distinguish, because H. celiana 
has the leaf margin quite eglandular, its stipules are only 1-1.5 mm long, its inflores- 
cence is rufotomentose, its petals are all fimbriate and eglandular, and its styles are 
hairy. In spite of the leaf shape, I think those two species are not especially closely 
related. Hiraea steyermarkii is also easily distinguished from H. affinis Miq., in which 
the lamina margin is eglandular, the petioles, stipules, and pedicels are shorter, all the 
petals are eglandular, and the samara lacks a dorsal wing. Hiraea neblinensis W. R. 
Anderson has shallowly cordate leaves with an eglandular margin, shorter stipules, 
petals that are all long-fimbriate and eglandular, and the dorsal wing of the samara 
entire or subentire. Large glands on the leaf margin are found in two species of 
French Guiana, H longipes W. R. Anderson and H. morii W. R. Anderson; both 
have the lateral petals eglandular and the posterior petal glandular-dentate, as in H. 
steyermarkii. Hiraea longipes also has a long pedicel and long stipules like those of H. 
steyermarkii. However, its lamina is rounded to cordate at the base, producing a shape 
like that of H. reclinata Jacq., and it has shorter petioles and longer bracts and 
bracteoles. In H. morii the lamina is similar in shape to that of H. steyermarkii, the 
marginal leaf glands are even more prominent, and the stipules are almost as long, 
but the inflorescence is much less open, with the cyme subsessile and the umbels 
raised on stalks only 4-10 mm long. Its pedicels are short, up to 17 mm long, and its 
petals are red in anthesis. Young flowers and mature fruits of H. steyermarkii should 
help to sharpen these distinctions. 

Hiraea valida W. R. Anderson, sp. nov.— Type: Ecuador. Napo: Canton Tena, 
Estacion Biologica Jatun Sacha, 8 km al E de Misahualli, 1°04'S, 77°36'W, 
400 m, bosque muy hiamedo tropical, bosque primario cerca al riachuelo 
Chinquipino, parcela permanente 03, 20 Jan 1990 fr, C E. Ceron, C. 
Iguago & E. Saldumbide 8370 (holotype: MICH!). 



1997 W. R. ANDERSON: MALPIGHIACEAE 71 

Liana lignosa ramis sericeis mox vel demum glabratis, quoque nodo cristis 
interpetiolaribus munito. Lamina foliorum majorum 12-22.5 cm longa, 8-L5 cm 
lata, obovata, basi obtusa, rotundata, vel subcordata, apice late rotundata vel 
saepius retusa vel subcordata, margine glandulosa, supra mox glabrata, subtus 
pertinaciter sparsim vel densius sericea; petiolus 15-23 mm longus, apice biglan- 
dulosus glandulis 2-3 mm longis; stipulae (2.5-) 3-5 mm longae, 0.9-1.5 mm latae, 
complanatae, in quarta distali petioli portatae. Inflorescentia ex 4-10 cymis in 
serie verticali axillari constans, quaque cyma plerumque ex 1 umbella 4-flora 
constanti; pedicellus 10-16 mm longus, usque ad 24 mm in fructu. Petala lateralia 
eglandulosa vel dentibus distalibus paulo glandulosis; petalum posticum limbo 
glanduloso-fimbriato. Styli apice dorsaliter acuti. Samara alls lateralibus 25-40 
mm latis, 40-60 mm altis, ala dorsali 4-10 mm lata, 10-14 mm alta. 

Woody vine; stems originally sericeous with very short, brown, strongly ap- 
pressed hairs, soon or eventually glabrate, the nodes marked by a prominent 
corky ridge connecting the bases of opposite petioles. Lamina of larger leaves 12- 
22.5 cm long, 8-15 cm wide, obovate, obtuse, rounded, or subcordate at base, 
broadly rounded or more often refuse to obcordate at apex and often apiculate, 
bearing several non-prominent, often recessed glands on distal 1/2 or more of 
margin, initially sericeous on both sides, soon glabrate above or with some hairs 
persistent on midrib, sparsely to fairly densely sericeous below at maturity with 
the hairs 0.1-0.3 mm long, very straight and tightly appressed, the principal lateral 
veins 8-11 pairs, very prominent below but not above, interconnected by scalari- 
form tertiary veins; petiole 15-23 mm long, densely sericeous to glabrate, bearing 
2 glands 2-3 mm long at or just above apex, pressed against petiole and/or abaxial 
midrib; stipules (2.5-) 3-5 mm long, 0.9-1.5 mm wide, flattened, sericeous to 
glabrate, arising from upper edge of petiole and pressed together at base, borne 
on distal 1/4 of petiole. Flowers borne above current leaves and, often, on old 
leafless stems. Inflorescence a dense vertical array of 4-10 cymes in an elongated 
zone on the stem above the axil, each cyme usually with only the central branch 
developing and flanked on each side by dormant buds (but the lateral branches 
occasionally developing), bearing a single 4-flowered umbel; umbel without a 
gland in the center, borne on a sericeous stalk 2-11 mm long; bracts 0.7-1 mm 
long and wide, triangular or ovate, abaxially sericeous to glabrate, adaxially gla- 
brous, eglandular, persistent; bracteoles similar to subtending bract or smaller, 
especially narrower; pedicel 10-16 mm long in flower, up to 24 mm long in fruit, 
sericeous. Sepals 1-1.7 mm long, hardly exceeding glands when they are present 
(ca 0.5 mm), 1.5-2.5 mm wide, broadly ovate, obtuse to rounded at apex, ap- 
pressed in anthesis, abaxially densely sericeous, adaxially glabrous, the anterior 
eglandular, the lateral 4 all eglandular or all biglandular, the glands 1.4-2 mm 
long, broadly elliptical, borne partly below sepals on receptacle. Petals "orange," 
glabrous, thick-textured; lateral 4 petals with the claw 1.5-2 mm long, the limb 
4.5-6.5 mm long and wide, the anterior pair ± concave and undulate or erose, 
eglandular, the posterior pair ± flat and subrectangular, irregularly dentate or 
short-laciniate, eglandular or with the distal teeth slightly glandular; posterior 
petal with the claw ca 3.5 mm long and held erect, the limb 3.5-4 mm long, 3.5-4.5 
mm wide, bent forward and usually conduplicate, glandular-fimbriate all around 
margin or at least on the proximal 2/3. Stamens glabrous; filaments nearly free to 
connate up to 0.7 mm at base, straight or somewhat bowed, 2.2-2.5 mm long 
opposite sepals, ca 1.5 mm long opposite petals; anthers 0.7-1.1 mm long, mostly 
somewhat longer opposite sepals than opposite petals, with the connective proxi- 
mally dark red and distally yellow in dried material. Ovary 1.5 mm high, densely 



72 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

sericeous; styles glabrous, with an acute dorsal projection at apex 0.1-0.2 mm 
long; anterior style 2-2.5 mm long, moderately bowed; posterior 2 styles 2.5-3 ram 
long, strongly bowed. Samara butterfly-shaped; lateral wings membranous, 25-40 
mm wide, 40-60 mm high, broadly flabellate to nearly semicircular but with an 
evident gap between their edges, at least above, sinuous or erose, persistently 
thinly sericeous or appressed-tomentose on both sides; dorsal wing 4-10 mm wide, 
10-14 mm high, rectangular or irregularly dissected, erose or coarsely toothed, 
extending forward at apex between lateral wings, appressed-tomentose; nut glo- 
bose, 5-7 mm in diameter, tomentose with short medifixed hairs, the ventral 
areole circular, 2.5-3 mm in diameter, partially surrounded (on the sides but not 
around the base) by an irregular callose thickening 0.5-1 mm thick. 

Additional Specimens Examined. Ecuador. Napo: confluence of Quiwado and Tiwaeno Rivers, 
primary forest, Apr fl, Davis & Yost 1038 (ECON, F, MICH, QCA); Canton Orellana, Sector Huash- 
ito, 20 km al N de Coca, propiedad de Palmoriente, 0°20'S, 77°05'W, 250 m, bosque huinedo tropical, 
primario, Nov imm fr, Gudiho 194 (MICH).— Pastaza: Pastaza, pozo petrolero Villano 2 de Arco, 
1°25'S, n°20'W, 400 m, bosque humedo tropical, primario, Dec fl, Hiirtado 2820 (MICH): entre los 
rios Iquino y Villano, 1°29'S, 77°27'W, 350 m, bosque humedo tropical, primario, Aug fl buds. Tirado 
et al. 55 (MICH). Peru. Amazonas: Lugar Amtami, monte, Jan fr, Kayap 182 (MO). 

The epithet of this species, which means strong, refers to its stout woody 
stems and large leaves, and its habit of growing in the canopy of primary forests. 
Hiraea valida is distinguished by the ridge across its nodes, the large leaf with a 
retuse to subcordate apex and marginal glands, the flattened stipules borne on the 
distal fourth of the petiole, the elongated vertical array of umbels above each axil, 
the glandular-fimbriate posterior petal, the dorsally acute style apex, and the 
large samara with a well-developed dorsal wing. Perhaps it is best compared to H. 
affinis Miq., because that also has a vertical array of umbels, the lamina often 
retains some short hairs on the abaxial surface, and it usually has a large membra- 
nous samara. However, H. affinis has no interpetiolar ridge, its stipules are much 
shorter, not especially flattened, and borne lower on the petiole, its lamina is 
more pointed and lacks marginal glands, its petals are all eglandular, the dorsal 
projection at the style apex is rounded, and the samara lacks a dorsal wing. A 
closer sister species is probably H. morii W. R. Anderson, of French Guiana, 
which differs from H. valida in lacking prominent interpetiolar ridges and in 
having the leaf apex short-acuminate to rounded, the leaf glands prominent, the 
petiole shorter, the stipules nearly terete and borne lower on the petiole, the 
inflorescence a single cyme of three umbels in each axil, and longer filaments and 
styles. The petals in H. morii were described by the collector as red in anthesis, 
those of H. valida as "orange." 

There are two sterile specimens at MICH that probably represent this species. 
They are Gentry et al. 27924 & 42992, both collected in Maynas, Loreto, Peru, in 
the Yanamono Explorama Tourist Camp near the confluence of the Rio Napo 
and Rio Amazonas, 3°28'S, 72°50'W. The lamina is enormous, up to 44 cm long 
and 22 cm wide, and more gradually tapered proximally, and the stipules are very 
long, up to 10 mm. Nevertheless, the specimens have a ridge across the node 
between the petiole bases as in H. valida, and they also resemble the latter in the 
shape and position of the stipules, in the very large petiole glands, in the thinly 
sericeous abaxial surface of the lamina, and in the marginal glands of the lamina. 
Moreover, there is an elongated zone above the axil, suggesting the area that 
produces the inflorescences in H. valida. This is most likely a case of sterile stems 
having leaves that are much larger than those found on fertile branches. 



1997 W, R. ANDERSON: MALPIGHIACEAE 73 

Mascagnia haenkeana W. R. Anderson, sp. nov.— Type: Ecuador. Guayas: Guaya- 
quil, T. Haenke2256 in 1790 (holotype: F!). 

Liana lignosa, ramis velutinis. Lamina foliorum majorum 1-11.5 cm longa, 4- 
6.5 cm lata, ovata ellipticave, subtus pertinaciter velutina; petiolus 11-18 mm 
longus, pertinaciter velutinus, adaxialiter aliquot glandulis parvis instructus. Inflo- 
rescentia paniculata, velutina pilis albis V-formibus; bracteae eglandulosae, per- 
sistentes; pedunculus 0.8-2 (-2.5) mm longus; bracteolae eglandulosae, persis- 
tentes, in medio pedunculi portatae; pedicellus 7-9 mm longus, velutinus vel to- 
mentosus. Antherae 0.8-1 mm longae, tomentosae. Styli 1.8-2.1 mm longi, apice 
dorsaliter vix apiculati. Samara 13-18 mm alta, 14-19 mm lata; ala lateralis basi 
apiceque continua, membranacea; ala dorsalis 3.5^.5 mm alta, 1.5-3 mm lata; nux 
super medium samarae posita. 

Woody vine; stems persistently velutinous with the hairs very short, V-shaped, 
mixed white and brown, eventually deciduous. Lamina of larger leaves 7-11.5 cm 
long, 4-6.5 cm wide, ovate or elliptical, broadly cuneate to rounded at base, acute 
(?) or short-acuminate at apex, initially densely tomentose or velutinous above 
but soon or eventually glabrate, densely and persistently velutinous below with V- 
shaped hairs, bearing 1-several impressed glands abaxially near base on each side 
of midrib and several more distally between midrib and margin, the principal 
lateral veins few (ca 5 pairs), visible but not raised above, prominent below; 
petiole 11-18 mm long, densely and persistently velutinous, bearing several small 
glands distributed along the adaxial edges; stipules 0.5-0.8 mm long, triangular, 
sericeous, borne on horizontal ridges on stem beside base of petiole. Inflorescence 
a large open panicle, the axes and peduncles densely and persistently velutinous 
like stem but the hairs all white, the ultimate pseudoracemes 5-40 mm long, 
bearing 4-30 flowers ± evenly distributed except often absent from the proximal 
3-8 mm, proximally decussate but borne in no regular order distally; bracts 0.8- 
1.3 mm long, narrowly triangular, abaxially subvelutinous to glabrescent, adaxially 
glabrous, eglandular, persistent; peduncle 0.8-2 (-2.5) mm long; bracteoles like 
bracts but smaller, 0.5-0.7 mm long, borne at or below middle of peduncle; pedicel 
7-9 mm long in flower, up to 11 mm long in fruit, densely and persistently veluti- 
nous or tomentose. Lateral 4 sepals bearing 8 glands 1.1-2 mm long, the sepals 1- 
1.5 mm long beyond glands, ca 1.2 mm wide, ovate and rounded at apex, abaxially 
white-sericeous, ciliate on margin, adaxially glabrous, appressed in anthesis. Pet- 
als glabrous, exposed in enlarging bud, abaxially very narrowly keeled on claw; 
lateral petals with the claw ca 1.5 mm long, the limb ca 3 mm long, 2 mm wide, 
subentire or denticulate; posterior petal with the claw ca 2 mm long, the limb 
somewhat larger than in lateral petals, erose. Filaments ca 2-2.5 mm long, gla- 
brous; anthers 0.8-1 mm long, densely tomentose. Ovary ca 0.8 mm high, densely 
white-hirsute; styles glabrous, stigmatic on inner angle of apex and dorsally mini- 
mally apiculate with the extension up to 0.1 mm long, the distance from face of 
stigma to dorsal tip 0.3-0.4 mm; anterior style ca 1.8 mm long, erect and straight; 
posterior styles 2.1 mm long, divergent at base, then erect and straight. Samara 
13-18 mm high, 14-19 mm wide, broadly ovate, sparsely sericeous to nearly gla- 
brate at maturity, the lateral wing membranous, continuous at base and apex, 
entire or very shallowly emarginate at apex, entire or sinuous and often denticu- 
late around margin; nut positioned above middle of samara; dorsal wing 3.5-4.5 
mm high, 1.5-3 mm wide, rounded or quadrangular, entire or slightly erose, aris- 
ing near base of nut and extending about halfway from apex of nut to apex of 
lateral wing, connate with lateral wing beyond nut; ventral winglet present as a 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

nge on underside of lateral wing, extending from apex of nut to apex of lateral 
ig, ca 0.5 mm wide; ventral areole ovate, 1.5-2 mm long, 1-1.3 mm wide; torus 
2 mm high, surrounded by a narrow, glabrous, 3-lobed, disciform outgrowth of 
; receptacle. 



shrubbery, Dec fr, MacBryde & Simmons 951 (QCA). 

Mascagnia haenkeana is named in honor of Thaddaeus Haenke (1761-1817), 
the collector of the holotype. The only flowers available, those on the holotype, 
are badly damaged by insects, so the estimates of sizes given above are rough 
approximations. The petals will surely prove to be pink or violet. In most charac- 
ters M. haenkeana is very similar to the widespread species Mascagnia divaricata 
(H. B. K.) Nied., which has long been misidentified as M. ovatifolia (H. B. K.) 
Griseb. (see discussion in Anderson, 1993, pp. 380-381). However, in M. divaricata 
the lamina is thinly sericeous to glabrate, the hairs when present nearly straight 
and appressed. I have never seen specimens of it with the leaves densely and 
persistently velutinous as they are in M. haenkeana. 

Mascagnia lilacina (S. Watson) Nied., Arbeiten Bot. Inst. Konigl. Lyceums Hosi- 
anum Braunsberg 3: 14. 1908. Hiraea lilacina S. Watson, Proc. Amer. 
Acad. Arts 17: 333. 1882.— Type: Mexico. Coahuila: Caracol Mountains 
21 mi SE of Monclova, Aug fl, E. Palmer 124 in 1880 (lectotype, here 
designated: GH!; isolectotypes: G! NY! US!). 

This species is mentioned here for the sole purpose of designating a lectotype. 

Mascagnia lugoi W. R. Anderson, sp. nov.— Type: Ecuador. Napo: Tierra Colo- 
rada, ca 3 km N of Coca (Puerto Francisco de Orellana), 22 Jan 1973 fl, 
H. Lugo S. 2920 (holotype: GB!; isotype: MICH!). 

Liana lignosa, ramis viridibus sericeis mox glabratis. Lamina foliorum majo- 
rum 7-12 cm longa, 2.5-6.5 cm lata, ovata ellipticave, sparsim sericea mox glabrata; 
petiolus 9-18 mm longus, sparsim sericeus vel glabratus, eglandulosus. Inflorcs- 
centia minute velutina pilis brunneis; bracteae bracteolaeque eglandulosae, per- 
sistentes; pcdicellus 3.5-7 mm longus, velutinus vel tomentosus. Pctala plcrumque 
rosea, lateralia ungue 1.5-2.7 mm longo, hmbo 3-4 mm longo, 3-5 mm lato, posti- 
cum ungue 2.5-3.5 mm longo, hmbo 4-4.5 mm longo, 5-6 mm lato. Antherae 0.9- 
1.4 mm longae, sericeae vel tomentosae. Stylus anticus 2-3 mm longus, fere rec- 
tus, styli postici 2.5-4.5 mm longi valde arcuati, omnes 3 styli apice dorsaliter 
apiculati apiculo 0.2-0.3 mm longo. Samara 19-30 (-38) mm alta lataque; ala 
laterahs basi apiceque continua, membranacea; ala dorsalis (3.5-) 5-9 mm alta, 2- 
4 mm lata; nux super medium samarae posita. 

Woody vine; stems green the first year, sericeous to glabrate, the hairs when 
present brown, straight, appressed. Lamina of larger leaves 7-12 cm long, 2.5-6.5 
cm wide, ovate or elliptical, cuneate or rounded at base, usually slightly revolute 
at margin, acuminate at apex, sparsely sericeous to glabrate on both sides, bearing 
4-9 impressed glands abaxially in a row on each side of midrib, from base to apex 
between midrib and margin, the 5-7 pairs of principal lateral veins visible but 
hardly raised above, promi^iulous or prominent below; petiole 9-18 mm long. 



1997 W. R. ANDERSON: MALPIGHIACEAE 75 

sparsely sericeous to glabrate, eglandular; stipules 0.3-0.8 mm long, triangular, 
sericeous to glabrate, borne on stem beside base of petiole. Inflorescence panicu- 
late with the branches subtended by successively more reduced vegetative leaves, 
the axes and peduncles minutely velutinous or eventually glabrescent with the 
hairs mostly brown, the ultimate pseudoracemes 10-50 mm long, bearing 4-20 (-28) 
flowers ± evenly distributed above the proximal 5-15 mm or denser toward apex, 
often decussate proximally but borne in no regular order distally; bracts 0.5-1.2 
mm long, narrowly triangular, abaxially sericeous to glabrescent, adaxially gla- 
brous, eglandular, persistent; peduncle 0.5-3.5 mm long; bracteoles like bracts but 
smaller and more elliptical, 0.4-0.7 mm long, borne well below apex of peduncle; 
pedicel 3.5-7 mm long in flower, up to 10 mm long in fruit, velutinous or tomen- 
tose. Lateral 4 sepals bearing 8 glands, the sepals 0.8-1.5 mm long beyond glands, 
1-1.3 mm wide, obtuse or rounded, abaxially sericeous with the hairs white or 
golden, ciUate on margin, adaxially glabrous, appressed in anthesis; glands 1.1-2 
mm long, detached at apex. Petals "pink" or "reddish," rarely "lilac," glabrous, 
exposed in enlarging bud, usually abaxially narrowly keeled on claw and base of 
limb, especially the lateral petals; lateral petals with the claw spreading and the 
limb often reflexed, the claw 1.5-2.7 mm long, the limb 3-4 mm long, 3-5 mm 
wide, nearly rotund to oblate or roughly triangular with rounded lobes at base, 
flat or slightly concave, proximally erose, distally entire; posterior petal with the 
claw nearly erect and the limb strongly reflexed, the claw 2.5-3.5 mm long, stout, 
the limb 4-4.5 mm long, 5-6 mm wide, oblate, somewhat crumpled, erose. Fila- 
ments 1.5-3.5 mm long, longest opposite anterior sepal and posterior-lateral pet- 
als with the latter stouter, shortest opposite posterior petal, connate at base for up 
to 0.7 mm, glabrous; anthers 0.9-1.4 mm long, longest opposite posterior-lateral 
petals, sericeous or tomentose. Ovary ca 1 mm high, densely hirsute; styles with a 
few hairs at base, distally glabrous; anterior style 2-3 mm long, erect and straight 
or curved shghtly toward posterior petal, posterior styles 2.5-4.5 mm long, very 
strongly bowed outward from base and then back toward center of flower, elon- 
gating and twisting outward in age; all 3 styles with a sharp well-developed apical- 
dorsal extension 0.2-0.3 mm long, the distance from internal stigma to tip of 
dorsal extension 0.5-0.8 mm. Samara 19-30 (-38) mm high and wide, broadly 
ovate to rotund, persistently sericeous on nut, otherwise sparsely sericeous to 
glabrate at maturity, the lateral wing membranous, continuous at base and apex, 
broadly notched ca 2-3 mm deep at apex, sinuous around margin; nut positioned 
above middle of samara; dorsal wing (3.5-) 5-9 mm high, 2-4 mm wide, semicircu- 
lar or quadrangular or irregular in shape, entire or sinuous, arising at or below 
base of nut and extending almost halfway from apex of nut to apex of lateral 
wing, connate with lateral wing beyond nut; ventral winglet absent or present only 
as a rudimentary flange ca 0.4 mm wide extending from apex of nut about halfway 
to apex of lateral wing; ventral areole ovate, 2.5-3.3 mm long, 1-1.3 mm wide; 
torus 1.7-2.5 mm high, surrounded by a wide, reddish, glabrous, 3-lobed, disci- 
form outgrowth of the receptacle. 

Additional Specimens Examined. Brazil. Amazonas: near mouth of Rio Embira, tributary of 
Rio Tarahuaca, 7°30'S, 70°15'W. Krukoff4639 (MICH). Ecuador. Napo: Rio Payamino, tributary of 
the Rio Napo, vicinity of Payamino-Capihuara, Lugo 2895 (GH, MICH).— Sucumbios: near ferry 
crossing over Coca River, 69 km S of Lago Agrio, Gentry 12556 (GB, MICH, MO, QCA, S); Canon 
de los Monos, road Coca (Puerto Francisco de Orellana)-Lago Agrio, 12 km N of Coca, Lugo 2947 
(GB, MICH); road Lago Agrio~El Chaco, Lugo 3484 (GB, MICH); environs of Limoncocha, Madi- 
son et al. 5484 (MICH, QCA, SEL). Peru. Amazonas: Rio Cenepa, vicinity of Huampami, 5 km E 



76 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

of Chavez Valdivia, 4°30'S, 78°30'W, Ancuash 1247 (MICH, MO).-~Huanuco: Prov. Huanuco, Tingo 
Maria, Asplund 13004 (G).— Loreto: Alto Amazonas Prov., Washintsa and vicinity, Rio Huasaga, 
3°20'S, 76°20'W, Lewis et al 11225 (MO).— Madre de Dios: Parque Nacional del Manu, Rio Manu, 
Cocha Cashu Station, Foster 5411 (F, MICH); Tambopata Province, Las Piedras, Cusco Amazonico, 
12°29'S, 69°03'W, Timand 2009, 2336, 2414 (all MO).-San Martin: Prov, Mariscal Caceres, Distr. 
Tocache Nuevo, Rfo Canute, Schunke V. 10957 (MICH). 

The epithet of Mascagnia lugoi honors Holguer Lugo Sanchez (b. 1941), the 
collector of the type and many other excellent specimens from Amazonian Ecua- 
dor. This species inhabits Amazonian tropical wet forests at elevations of 185-500 
m. It has been collected with flowers in January, February, May, June, August, 
and September, and with fruits in June, September, October, and November. 
Mascagnia lugoi is closely related to M. divaricata (H. B. K.) Nied., but that 
widespread species has petioles bearing several small glands on the adaxial edges, 
densely white-velutinous inflorescences, smaller petals that are usually described 
as violet, and shorter styles that bear only a very short dorsal apiculum at the 
apex, ca 0.1 mm long. 

Mezia angelica W. R. Anderson, sp. nov.— Type: French Guiana. Saul, La Fumee 
Mountain Trail, S^^S^N, 53°12'W, 17 Sep 1989 fl, S. Mori, C. Grade & M. 
Rothman 20945 (holotype: MICH!; isotypes: CAY! K! NY! U! US!). Fig. 5. 

Liana lignosa. Lamina foliorum majorum 12-22 (-24) cm longa, 4.5-8 (-9.8) 
cm lata, primo sericea mox glabrescens vel glabrata. Bracteolae 5-7 (-8) mm 
longae, eglandulosae. Glandulae 2 in quoque sepalo lateraH omnino connatae. 
Petala omnia abaxialiter tomentosa, margine eglandulosa, lateraha citrina, posti- 
cum distaliter citrinum proximaliter rubrum. Filamenta tomentosa, 2-4.2 mm lon- 
ga, tantum basaliter connata, valde heteromorpha; antherae basi tomentosae, prae- 
cipue illae sepalis oppositae. Styh apice pedaliformes, anticus 3-3.5 mm longus ± 
rectus, postici 3.5-4 mm longi lyrati vel sigmoidei. Samara oblata, 55-80 mm lata, 
45-60 mm alta; ala laterahs basi continua; 3 alae dorsales planae, parallelae; nux 
utrinque inter areolam ventralem et alam lateralem 1 crista vel alula 1-9 (-15) 
mm lata et 11-23 mm alta instructa. 

Woody vine, the stems initially brown-sericeous but mostly soon glabrescent, 
initially flattened or quadrangular but soon or eventually becoming terete. Lami- 
na of larger leaves 12-22 (-24) cm long, 4.5-8 (-9.8) cm wide, elliptical or some- 
what ovate or obovate, truncate or cuneate at base, flat and not or only slightly 
thickened at margin, abruptly short-acuminate at apex with the acumen 5-10 mm 
long, initially sericeous adaxially but very soon glabrate or thinly sericeous proxi- 
mally on and near midrib, thinly sericeous abaxially to glabrescent with the hairs 
dark brown fading to white, sessile, straight, tightly appressed, and hardly notice- 
able without a lens, abaxially eglandular or bearing 1 large gland at base on each 
side of midrib (these glands often sunken in crypts) and occasionally several small 
impressed glands distally in a single row several mm inside margin, the reticulum 
and 5-8 pairs of lateral veins visible but hardly raised adaxially, prominent abaxi- 
ally; petiole 15-30 mm long, sericeous to glabrate, eglandular; stipules none or 
minute triangular rudiments up to 0.3 mm high, borne on an interpetiolar ridge. 
Inflorescence open with spreading flattened branches 2-13 cm long, hght- to dark- 
brown-sericeous or eventually glabrescent; stalk of the umbel 6-16 mm long, bear- 
ing 1 pair of deciduous sterile bracts 2-8 mm above the base, well below to 



1997 W. R. ANDERSON: MALPIGHIACEAE 77 

slightly above the middle; floriferous bracts 3.5-6.5 mm long, obovate and broadly 
rounded at apex, concave, eglandular, abaxially densely sericeous, adaxially gla- 
brous or proximally sericeous, deciduous before or during anthesis; peduncle 7-14 
mm long in flower, thickened and often somewhat elongated in fruit, sericeous or 
subvelutinous; bracteoles 5-8 mm long, eglandular, abaxially densely sericeous 
with the hairs light to fairly dark brown, adaxially glabrous, emarginate or bifid at 
apex, with the midrib usually raised abaxially, mostly persistent past maturity of 
the fruit; pedicel 0-0.8 mm long in flower, up to 2 mm long in fruit, hirsute with 
the hairs spreading or appressed. Sepals 4-7 mm long beyond glands, 2-3 mm 
wide, spatulate, often strongly revolute along sides, abaxially densely brown-to- 
mentose, adaxially glabrous, the anterior eglandular, the lateral 4 biglandular with 
the glands 2-3 mm long, nearly or completely connate, the pair 1.8-3 mm wide, 
obovate. Petals abaxially loosely white-tomentose in center; lateral petals pure 
lemon-yellow, the claw 3-4 mm long, the limb 6-11 mm long and wide, signifi- 
cantly larger in anterior pair than in posterior pair, orbicular or suborbicular or 
obovate, slightly crumpled, erose, eglandular; posterior petal distally lemon-yel- 
low, proximally red in the center, the claw 3.5 mm long, very thick, constricted at 
apex, the limb 6-7 mm long and wide, orbicular, often crumpled and folded back- 
ward, distally erose, proximally dentate or short-fimbriate but eglandular. Fila- 
ments connate only in the basal 0.5-1 mm, tomentose especially distally, erect and 
straight or (especially opposite lateral sepals) bent toward center of flower, strongly 
heteromorphic, 2-4.2 mm long, shortest opposite posterior petal, then progres- 
sively longer opposite anterior-lateral petals, lateral sepals, and posterior-lateral 
petals (these 2 much stouter than all others), longest opposite anterior sepal; 
anthers 1-1.8 mm long, tomentose at base, with the connective adaxially dark red 
its whole length and abaxially dark red proximally and yellow distally, heteromor- 
phic, those opposite sepals more abundantly tomentose than those opposite petals 
and having the connective widened so as to displace the locules laterally and 
elongated so as to exceed the locules. Ovary 1.5 mm high, sericeous; styles proxi- 
, nearly terete, pedaliform at apex (i.e., with a short, broad abaxial 
nbling from above the sole of a shoe); anterior style 3-3.5 mm 
long, nearly straight and erect or inclined slightly toward posterior petal; posterior 
styles 3.5-4 mm long, lyrate or sigmoid, bent outward in proximal third, then erect 
or bent inward, and distally often bent outward, with the stigmas twisted toward 
posterior petal. Samara oblate, 55-80 mm wide, 45-60 mm high, sericeous on the 
nut, thinly sericeous on the wings; lateral wing 25-34 mm wide, continuous at 
base, incised to nut at apex, membranous, apparently nearly flat or somewhat 
corrugated, entire or repand at margin; central dorsal wing 7-14 mm wide, 18-33 
mm high, semicircular or irregularly repand, flat; 1 flat winglet 3-7 mm wide and 
10-18 mm high present on each side of and parallel to central dorsal wing, entire 
or irregularly repand; 1 flat crest or winglet 1-9 (-15) mm wide and 11-23 mm 
high present on each side of nut, outside of and parallel to lateral wing, irregularly 
repand or coarsely dentate; nut with the ventral areole 9-13 mm high, 2-4 mm 
wide, narrowly ovate or elliptical, bordered by 2 ribs that remain on samara. 
Pyramidal receptacle 1-2 mm high. Embryo circular in cross section and elliptical in 
longitudinal section, the cotyledons equal, straight, semicircular in cross section. 



; Reserva Florestal Ducke, Manai 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 7 




1997 W. R. ANDERSON: MALPIGHIACEAE 79 

26, 2°53'S, 59°58'W, Vicentini 1014 (MICH).— Para: Rio Jari, Monte Dourado, Cavalcante 3329 
(MG, NY); Tucurui, margens da PA-149 ate o Km 50, Revilla et al. 8310 (INPA); Santarem, Km 70 da 
estrada do Palhao, Ramal do Caetetu, M. Silva 2604 (MG, MICH, NY); Rio Jari, estrada entre 
Planalto A e Tinguelin, Km 13, N. T. Silva 2785 (IAN, UB); Rio Jari, N. T. Silva 2830 (UB). French 
Guiana. Saul, La Fumee Mountain Trail, 3°37'N, 53°12'W, Mori et al. 21012 (CAY, MICH, 
NY). Guyana. Kamoa River, Clarence Hill, r31'N, 58°50'W, Jansen-Jacobs et al. 1738 (MICH, 
MO, NY, US); Rupununi Distr., between Kuyuwini Landing and Kassikaityu River, 2°00'N, 59°15'W, 
Jansen-Jacobs et al. 3019 (MICH). 

Moist non-flooded forests at elevations of 50-360 m, in eastern Amazonia 
from eastern Amazonas and northern Para, Brazil, north to central French Gui- 
ana and southern Guyana (Fig. 6); to be expected in Suriname. Collected with 
flowers and fruits from August to October. The epithet of this species honors 
Bobbi Angell (b. 1955), the talented artist who has contributed so much to plant 
taxonomy in recent years through her graceful drawings. 

Mezia angelica is probably most closely related to M. araiijoi Nied., the type 
of the genus, which occurs far to the south of M. angelica in Espfrito Santo, Rio de 
Janeiro, and eastern Minas Gerais (see Fig. 6). Both species have all five petals 
abaxially loosely white-tomentose, tomentose filaments that are connate only at 
the base, tomentose anthers, and slender styles of which the posterior two are 
lyrate. They differ most dramatically in their fruits. In M. araujoi the samara bears 
a complex of many winglets with diverse orientations on each side of the central 
dorsal wing, some of them at right angles to the lateral and dorsal wings. In M. 
angelica there is only one flat intermediate crest or winglet on each side of the 
dorsal wing, parallel to it; there is also a flat winglet on each side of the nut 
outside the large lateral wing. A similar external winglet occurs sometimes in M. 
araujoi, associated with ribs oriented at right angles to the lateral wing. Mezia 
huberi, a species of southern Venezuela (Fig. 6), has a smaller but somewhat 
similar samara, with only two flat intermediate winglets (but without additional 
winglets outside the lateral wing); it differs from M. angelica in many characters, 
including its persistently sericeous leaves, glabrous posterior petal, glabrous fila- 
ments 1/3-2/3 connate, glabrous anthers, and straight, erect, stout styles. Mezia 
angelica is further distinguished from both M. araujoi and M. huberi in having the 
two glands on each sepal nearly or completely connate. 



FIG. 5. Mezia angelica, a) leafy stem, 
istent hairs, xO.5; b) portion of inflorescence, xO.5; c) umbel of 4 flower buds, each bud enclosed 

bracteoles and subtended by 1 bract, the stalk of the umbel bearing a pair of sterile bracts, x2; d) 
'er bud about to open, with the 2 bracteoles being forced apart by the enlarging sepals, x2; e) 
a flower with petals removed, with the 2 subtending bracteoles intact (above) and with the 
:teoles half cut away (below) to show sepals, with the eglandular anterior sepal in center, x2; f) 

(left) and from the side with 2 
e posterior petal to the right (right), x2; h) adaxial view of complete androeci- 
shortest stamen (fifth from right) that opposite the posterior petal, x4; i) side 
mthers, the pair on left opposite petals, those on right opposite sepals, x8; j) 
;tyle-apex enlarged, x8; k) samaras, abaxial view 
ew (below), xO.5; 1) samara in cross section, showing large lateral wing, smaller 
traight up), single winglets between dorsal and lateral wings, and single winglets 
0.5. Drawn by Bobbi Angell, a-j from Mori et al. 20945, k & 1 from Silva 2830. 
Dr. Scott Mori and the New York Botanical Garden. 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 




mariposa W. R. Anderson, sp. nov. — Type: Brazil. Acre: Basin of Rio 
Purus, near mouth of Rio Macauhan (tributary of Rio Yaco), 9°20'S, 
69°W, 11 Aug 1933 fl B. A. Krukoff 5452 (holotype: MICH!; isotypes: A! 
F! G! K! MO! NY! U! US!). 



Liana lignosa. Lamina foliorum majorum 13-22 cm longa, (5-) 6-10.5 (-12) 
cm lata, primo sericea vel subsericea permox glabrata. Bracteae floriferae 1.5-3.5 
mm longae; bracteolae 5-8 mm longae, exterior 1 glandula abaxiali excentrica 
instructa. Petala glabra, lateralia citrina eglandulosaque, posticum distaliter citri- 
num proximaliter rubrum margine eglandulosum vel proximater glanduloso-den- 
tatum. Antherae sepalis oppositae tomentosae, petalis oppositae glabrae vel paene 
glabrae. Styli complanati, apice dorsaliter breviuncinati. Samara papilionacea alls 
lateralibus 22-33 mm latis, 31-52 mm altis, ala dorsali 7-12 mm lata, 18-28 mm 

Woody liana climbing to tops of tall trees, the stems initially sericeous with 
brown and cinereous hairs but very soon glabrate, initially flattened or quadran- 
gular but soon or eventually becoming terete. Lamina of larger leaves 13-22 cm 
long, (5-) 6-10.5 (-12) cm wide, narrowly to broadly elliptical or somewhat ovate 
or obovate, rounded, truncate, or cuneate at base, flat and not thickened at mar- 
gin, abruptly short-acuminate at apex with the acumen 9-17 mm long, initially 
sericeous or subsericeous but very soon glabrate (usually before lamina is fully 
expanded), abaxially bearing 1 large gland at base on each side of midrib (these 
glands often sunken in crypts) and a single row of impressed glands several mm 



1997 W. R. ANDERSON: MALPIGHIACEAE 81 

inside margin on distal 1/2-3/4, the reticulum and 5-8 pairs of lateral veins visible 
but only moderately raised adaxially, prominent abaxially; petiole 8-14 (-17) mm 
long, initially sericeous but very soon glabrate, eglandular; stipules none or minute, 
caducous, triangular rudiments ca 0.2 mm high, borne on an interpetiolar ridge. 
Inflorescence open, often branched, up to 50 cm long, the major axes indetermi- 
nate, the short lateral branches cymes of 1-3 (-7) 4-flowered umbels, all axes 
dark-brown-sericeous or appressed-tomentose, the major axes often glabrescent, 
the ultimate axes and peduncles with the vesture persistent or patchily deciduous 
in fruit; stalk of the umbel or cyme 5-13 mm long, bearing 1 pair of persistent or 
deciduous sterile bracts above middle to (usually) near the apex, 0-2 (-4) mm 
below umbel; floriferous bracts 1.5-3.5 mm long, ovate, rounded at apex, often 
concave or conduplicate, eglandular, abaxially loosely sericeous, adaxially gla- 
brous, deciduous during anthesis; peduncle (9-) 11-18 mm long in flower, thick- 
ened and elongated in fruit; bracteoles 5-8 mm long, the outer bearing 1 circular 
or elliptical gland 0.5-1.1 mm long sunken in abaxial tissue at base between mid- 
dle and margin, the inner eglandular, both abaxially densely sericeous or ap- 
pressed-tomentose with the hairs reddish brown to dark brown, adaxially glabrous 
or sparsely sericeous, broadly rounded or truncate but not emarginate at apex, the 
midrib not raised abaxially, persistent past maturity of fruit; pedicel 0-1 mm long 
in flower, up to 3 mm long in fruit, hirsute with the hairs spreading or appressed 
to glabrate. Sepals (4-) 5-6.5 mm long beyond glands, 2-3.4 mm wide, spatulate 
or narrowly oblong, reflexed in anthesis, strongly revolute along sides, abaxially 
densely appressed-tomentose or subsericeous with the hairs reddish to fairly dark 
brown, adaxially glabrous, the anterior usually eglandular (rarely biglandular), 
the lateral 4 biglandular, the glands 2-3.3 mm long, distinct or partially to almost 
completely connate, the pair 1.5-2.7 mm wide, quadrate or obovate. Petals gla- 
brous; lateral petals yellow, the claw 2.5-3.5 mm long, the limb 9-16 mm long, 9- 
15 mm wide, significantly larger in anterior pair than in posterior pair, orbicular 
or obovate, flat or slightly crumpled, coarsely erose, eglandular; posterior petal 
distally yellow, proximally red in the center, the claw 2.7-3.5 mm long, ca 1 mm 
wide, constricted at apex, the limb 7-9 mm long, 5-9 mm wide, orbicular or 
elliptical, often crumpled and folded backward, proximally dentate with the teeth 
glandular or eglandular, distally erose. Filaments connate in the proximal 0.5-1.2 
mm, those opposite sepals glabrous or sparsely pilose, those opposite petals gla- 
brous, erect and straight or (especially opposite lateral sepals) bent toward center 
of flower, strongly heteromorphic, 1.8-4 mm long, shortest opposite posterior 
petal, then progressively longer opposite anterior-lateral petals, lateral sepals, and 
posterior-lateral petals (these 2 much stouter than all others), longest opposite 
anterior sepal; anthers (1-) 1.3-2.4 mm long, shortest opposite the posterior petal, 
with the connective adaxially reddish and abaxially dark red proximally and yel- 
low distally, heteromorphic, those opposite sepals abundantly tomentose the whole 
length of their locules and having the connective widened so as to displace the 
locule laterally and elongated so as to exceed the locules, those opposite petals 
nearly or quite glabrous, the locules extending to apex of connective. Ovary 1.5- 
1.8 mm high, loosely sericeous; styles loosely sericeous on proximal half, flattened 
laterally in distal half, not or only very narrowly pedaliform at apex, dorsally 
short-hooked at apex with the projection 0.1-0.3 mm long; anterior style 2.5-3.2 
mm long, straight and erect or inclined slightly toward posterior petal; posterior 
styles 3-4 mm long, lyrate, bent outward in proximal third, then erect or bent 
inward, and distally often bent outward, with the stigmas twisted toward posterior 



82 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

petal. Samara butterfly-shaped, the whole oblate, 56-70 mm wide, 31-52 mm high, 
densely short-tomentose or subsericeous on the nut, the vesture thinner on wings, 
especially distally, or patchily deciduous; lateral wings 22-33 mm wide, dissected 
to nut at base and apex, roughly trapezoidal but rounded at base and obtuse at 
apex, relatively firmly membranous or subcoriaceous (compared to the very frag- 
ile samaras of other species of the genus), nearly flat, entire or repand at margin; 
dorsal wing 7-12 mm wide, 18-28 mm high, crescent-shaped, triangular, or rectan- 
gular, usually widest near apex, entire or coarsely erose, flat; nut devoid of winglets 
or other outgrowths between dorsal and lateral wings, and lacking crests or winglets 
outside lateral wings; nut inflated, broadly cylindrical, 10-13 mm across, 14-17 
mm high, with the ventral areole 10-13 mm high, 4-6 mm wide, elliptical or obovate, 
bordered by 2 ribs that remain on samara. Pyramidal receptacle 2-3 mm high. 

Additional Spectmens Examined: Brazil. Acrh: Vizinhanga de Sena Madureira, 9°05'S, 68°40'W, 
Nelson et al. 523 (INPA, MICH, MG); Coloca^ao Boa Uniao, Sothers & Santos <S (MICH).— Amazo- 
NAs: Track from Sao Paulo to Terra Firme, Rio Purus, opposite Boca do Acre, France et al. 2586 
(INPA, NY).— Para: Upper Cupari River, plateau between the Xingu and Tapajos Rivers, Krukoff 
1135 (A, G, K, NY, P); Tucurui, Lima & Silva 83 (INPA); Boa Vista on the Tapajos River, Monteiro 
da Costa 56 (F); Tucurui, Rio Caraipe, Revilla et al. 8272 (INPA) & PA-149, Revilla et al. 8425 
(INPA).— Rondonia: Estrada Belmonte, Cordeiw 766 (MICH); Rio Jarii, BR-29, Porto Velho, Duarte 
7158 (MICH); Porto Velho, estrada Belmonte, Mota & Coelho 100 (INPA); eastern bank of Rio 
Madeira at Misericordia between Cachoeiras Madeira and Misericordia, Prance et al. 6633 (INPA, 
MG, MICH, NY). Peru. Huanuco: Prov. Pachitea, Distr. Honoria, Bosque Nacional de Iparia, Rio 
Pachitea cerca del campamento Miel de Abeja, 1 km arriba del pueblo de Tournavista o unos 20 km 
arriba de la confluencia con el Rio Ucayali, Schunke V. 2062 (F, NY, US), 2779 (F, NY, US), 2186 (F). 

Collected with flowers from June to October and with fruits in September 
and October, in high terra firme forests at elevations up to 400 m, in Amazonian 
Brazil and Peru (Fig. 6). The epithet mariposa is the Portuguese and Spanish 
word for butterfly. It refers to the shape of the samara of Mezia mariposa, which 
is unique in the genus in having two lateral wings completely distinct at the base 
as well as at the apex; all other species have a single lateral wing continuous at the 
base. Another unique characteristic is the single gland present on one side of the 
outer bracteole; other species of Mezia have both bracteoles eglandular. In many 
other characters M. mariposa resembles M. angelica, e.g., the relative lengths of 
the filaments and shapes of the anthers, but M. angelica has hairy petals, all the 
anthers are tomentose, and the samara bears additional winglets or crests between 
the dorsal and lateral wings and outside the lateral wings. See the description and 
discussion of M. angelica above. 

Mezia tomentosa W. R. Anderson, sp. nov.— Type: Ecuador. Pastaza: Pastaza 
Canton; Pozo petrolero "Masaramu" de UNOCAL, 40 km al NE de Mon- 
talvo, 76°52'W, 00°44'S, 390 m, bosque humedo tropical, primario, 1-17 
May 1990 fr, S. Espinoza 244 (holotype: MICH!; isotype: MO!). 

Liana lignosa. Lamina foliorum majorum 11.5-17 cm longa, 5-9.4 cm lata, 
abaxialiter dense et pertinaciter velutina vel tomentosa, eglandulosa; petiolus 15- 
23 mm longus, eglandulosus. Umbellae pedunculus 5-11 mm longus; bracteae 
floriferae 2-A mm longae; pedunculus florifer in fructu 10-14 mm longus; bracteolae 
6-8 mm longae, eglandulosae. Glandulae 2 in quoque sepalo laterali distinctae. 
Filamenta ca 1/2-connata, glabra. Styli apice dorsaliter rotundati vel obtusi. Samara 



1997 W. R. ANDERSON: MALPIGHIACEAE 83 

oblata vel subcircularis, 70-90 mm lata, 60-75 mm alta; ala lateralis basi continua; 
alae dorsales et intermediae numerosae, dissectae, interconnexae, non parallelae, 
ca8-10mmlatae. 

Woody liana, the stems persistently subsericeous or appressed-tomentose with 
dark reddish-brown hairs to eventually glabrate, initially quadrangular but soon 
or eventually becoming terete. Lamina of larger leaves 11.5-17 cm long, 5-9.4 cm 
wide, elliptical or slightly ovate or obovate, cuneate at base, somewhat revolute at 
margin, abruptly acuminate at apex with the acumen 7-15 mm long, probably 
initially hairy adaxially but at maturity glabrate or tomentose proximally on mid- 
rib and often shiny as if lacquered, abaxially densely and persistently appressed- 
tomentose on midrib and elsewhere velutinous or tomentose with the hairs dark 
brown or reddish brown, often fading in age and sometimes unevenly abraded 
from oldest leaves, a mixture of sessile V-shaped hairs and Y- or T-shaped hairs 
with a short but definite stalk and the branches straight to sinuous, the lamina 
apparently eglandular, the reticulum and 5-8 pairs of lateral veins sunken or at 
most prominulous adaxially, prominent adaxially; petiole 15-23 mm long, persis- 
tently subsericeous hke stem, eglandular; stipules not found, but perhaps initially 
present on stem beside petiole where apparent scars ca 0.5 mm wide may some- 
times be found. Inflorescence known only in late fruit, such that only a very 
imperfect idea of its branching and structures is possible, but apparently a lateral 
panicle with vesture like that of stems on decussate branches and presumably 
bearing bracts or small leaves at the nodes, subtending ultimately single umbels of 
4 flowers; stalk of the umbel 5-11 mm long, bearing 1 pair of deciduous sterile 
bracts 1-3 mm above base, well below the middle; floriferous bracts 2-4 mm long, 
elliptical or obovate and rounded at apex, concave, eglandular, abaxially densely 
sericeous, adaxially glabrous, deciduous before maturation of fruit or persistent; 
peduncle 10-14 mm long in fruit, subsericeous or subvelutinous; bracteoles 6-8 
mm long, eglandular, abaxially densely sericeous or subsericeous with the hairs 
light to dark brown, adaxially glabrous, truncate or somewhat emarginate at apex, 
with the midrib sometimes raised abaxially, mostly persistent past maturity of 
fruit; pedicel 1-2.5 mm long in fruit, hirsute with the hairs spreading. Flowers not 
seen. Sepals 6.5-7.5 mm long beyond glands in fruit, 2.5-3 mm wide, spatulate, 
slightly revolute all around margin, abaxially densely and loosely subsericeous 
with the hairs more or less spreading, adaxially glabrous, the anterior eglandular, 
the lateral 4 biglandular with the glands 1.5-2 mm long, 1-1.3 mm wide, obovate, 
distinct. Filaments ca 1/2 connate, glabrous. Styles 2.4-3.2 mm long in fruit, straight, 
terete or laterally flattened distally, densely sericeous on proximal 1/2-2/3, dorsal- 
ly rounded or obtuse at apex. Samara oblate to nearly circular, 70-90 mm wide, 
60-75 mm high, tomentose on nut, tomentose to subsericeous on wings; lateral 
wing 30-43 mm wide, continuous at base, incised to nut at apex, membranous, 
distally flat but strongly corrugated near nut, repand at margin; dorsal wing and 
several intermediate wings indistinguishable, highly dissected and interconnected 
by transverse winglets, all ca 8-10 mm wide, together forming a ruffled complex 
of winglets of diverse orientations; nut with the ventral areole 7-10 mm high, 2-3 
mm wide, narrowly ovate, bordered by 2 ribs that remain on samara. Pyramidal 
receptacle 2-3.5 mm high. 

Additional Specimens Examined. Ecuador. Pastaza: Pastaza Canton; Pozo petrolero "Masara- 
mu" de UNOCAL, 40 km al NNE de Montalvo, 76°52'W, 00°44'S, 400 m, bosque hiimedo tropical, 
primario, May 1990 fr, Gudino 396 (MICH, MO). 



84 CONTR. UNIVERSITY OF MICHIGAN 

This species is named for its most distinctive feature, the persistently veluti- 
nous or tomentose leaves. Those stalked hairs are unique in the genus and give 
Mezia tomentosa a very different look from its congeners, whose hairs are sessile, 
straight, and tightly appressed. The large, membranous samara with a ruffled 
complex of dorsal winglets suggests that its closest relatives are M. includens 
(Benth.) Cuatrec. and M. curmnli W. R. Anderson. When it is collected with 
flowers, those should supply additional evidence to use in assessing its relation- 
ships. Mezia tomentosa is known only from the two collections cited above, which 
were made at the same time in the same place (Fig. 6). 



LITERATURE CITED 

Lnderson, W. R. 1981. Malpighiaceae. In The bukiny of the Guayana Highland— Part XL Mem. New 

York Bot. Card. 32: 21-305. 

. 1993. Notes on neotropical Malpighiaceae— IV. Contr. Univ. Michigan Herb. 19: 355-392. 

'base, M. W. 1981. A revision of Dicella (Malpighiaceae). Syst. Bot. 6: 159-171. 

'uatrecasas, J. 1958. Prima Flora Colombiana. 2. Malpighiaceae. Webbia 13: 343-664. 

fiedenzu, F. 1928. Malpighiaceae. In Das Pflanzenreich, ed. A. Engler, IV. 141: 1-870. Leipzig: 

Wilhelm Engelmann. 
)'Donell, C. A., & A. Lourteig. 1943. Malpighiaceae argentinae. Lilloa 9: 221-316, pi. 1-18. 



ACKNOWLEDGMENTS 

As always, my work is made possible by the cooperation and kind assistance of the curators of 
many herbaria. I send my thanks to those in charge of all the following collections: A, AAU, BM, 
BRG, CAY, F, G, GB, GH, IAN, INB, INPA, K, MEXU, MG, MO, MYF, NY, P, PEUFR, PORT, 
OCA, U, UB, UC, US, and VEN. I would also like to acknowledge several individuals who helped m 
different ways, all much appreciated. They are Bobbi Angell, David Bay, Paul Berry, Karin Douthit, 
Carol Gracie, Michael Grayum, Barry Hammell, Otto Huber, Fernand Jacquemoud, ScoU Mori, 
Basil Stergios, Stephen Tillett, and Beverly Walters. 



THE SUMMIT FLORA OF MT. MURUD, 
SARAWAK, MALAYSIA 

John H. Beaman 

Department of Botany and Plant Pathology 

Michigan State University 

East Lansing, Michigan 48824-1312 

Christiane Anderson 
University of Michigan Herbarium 

North University Building 
Ann Arbor, Michigan 48109-1057 



INTRODUCTION 

Mt. Murud, elev. 2,424 m, is the highest mountain in Sarawak. It is located on 
the boundary of the Limbang and Miri Divisions, in the Kelabit Highlands of 
northeastern Sarawak. The geology of Mt. Murud (as well as most of the Kelabit 
Highlands) is a white or yellowish sandstone of the Miocene Meligan Formation 
(Yin Ee Heng 1992). The long, narrow ridge constituting Mt. Murud proper, 
above 2,000 m, is about 4 km long, running in an ENE-WSW direction. The two 
highest points are fairly close together and nearly equal in elevation, the higher 
exceeding the lower by only 15 m. A panoramic view of the ridge is provided by 
Mjoberg (1925, Fig. 10), and a color photograph of Mt. Murud as seen from Batu 
Lawi is included in Plate 25 of Briggs (1988). 

Since 1982 or earlier the Murud area has been proposed as a national park, 
but political considerations have not yet allowed that to happen. Until now the 
Kelabit Highlands have been one of the last remaining accessible but culturally 
and environmentally unspoiled areas in Sarawak; however, changes seem immi- 
nent. From the summit of Mt. Murud the sounds of a logging operation to the 
north are now within earshot. Bario, the largest settlement in the Kelabit High- 
lands, is getting a new airstrip, and a golf course and other tourist attractions have 
been proposed for the area. Bario featured prominently in Tom Harrisson's (1959) 
book. World Within: A Borneo Story, which chronicles the Australian penetration 
behind Japanese lines during World War II and gives a dramatic and intimate 
view of the Kelabit Highlands before they were affected by Western culture. 

The first successful expedition to Mt. Murud was undertaken by the Swedish 
zoologist Eric Mjoberg, at that time Curator of the Sarawak Museum, who ap- 
proached the mountain in October, 1922, and spent six days on its summit in early 
November. He made a large museum collection of animals in diverse groups and 
obtained a considerable number of herbarium specimens as well. Subsequently 
eight other botanical collectors or collecting teams have been to the summit. 

Mjoberg (1925) published a detailed account of his 1922 expedition and a 
summary of previous exploration in the Kelabit Highlands. In discussing earlier 
expeditions he noted that the first attempt to explore Mt. Murud was initiated in 
1914 by J. C. Moulton, then Curator of the Sarawak Museum. Moulton left on an 
expedition in October but returned on November 19; he noted that "failure was 



86 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

due in the first place to the lack of food in the district, which prevented us 
obtaining sufficient natives for transport, and secondly to the alarming reports of 
a Dayak invasion which soon after our start robbed us of the few natives we had 
managed to get together" (Moulton 1915). Moulton made a second attempt to 
reach the mountain in 1920, but again failed. 



THE MJOBERG EXPEDITION 

Mjoberg submitted his plan of exploration to Rajah Charles Vynar Brooke on 
July 13, 1922, and received immediate approval. He spent five weeks traveling up 
the Baram River and into the Kelabit Highlands, at some points with more than 
70 porters and guides (his description of the personnel situation is somewhat 
ambiguous). He left for Baram Station (Marudi) on September 6, and, after vari- 
ous difficulties, arrived in the Kelabit region on October 5. He first saw Mt. 
Murud on October 10, about a two-hour hike before his party arrived at Pa Trap. 

He persuaded seven local Kelabits to show him the route to the summit of 
Mt. Murud. As he tells it, they hacked a path for him climbing steadily upwards, 
often walking on undergrowth only hanging together by roots. On October 19, he 
reached the foot of Mt. Murud, and on November 2, "perhaps my most strenuous 
day during the whole journey," he climbed to the top ridge. His characterization 
of the summit follows: "We found ourselves in a strange landscape where low 
bushes with thick leathery leaves constituted the predominating vegetation. Here 
and there smaller trees were seen, among them a conifer with trunk and larger 
branches practically covered with the yellow blossoms of a small, richly flowering, 
epiphytic orchid. Bright scarlet or snow-white flowers of rhododendron and simi- 
lar plants were met with everywhere; and most noticeable were the enormous and 
characteristically shaped pitchers of Nepenthes lowii, hitherto recorded only from 
Kinabalu and Batu Lawi." Mjoberg spent six days on the summit, until November 
13. It should be noted that the labels of his botanical collections from the summit 
area bear the erroneous date of "October 1922." By November 27 he had re- 
turned to Pa Trap and departed the Kelabit Highlands on December 4; he arrived 
back at Baram Station on December 16 or 17. 

In his book Forest Life and Adventures in the Malay Archipelago, Mjoberg 
(1930) gave an even more picturesque account of the summit than that quoted 
above, as follows. "I shall not soon forget the sight that met my eye when I 
succeeded, after tremendous exertions, in reaching the dim cloud-wreathed heights 
of the unexplored Mt. Murud. The whole of the vast forest was, as it were, illumi- 
nated by the newly opened orchids, which produced such a glory of colour that we 
could not turn our eyes away from all their beauty. Some of the trees were 
actually turned into pale yellow or mauve under their covering veil of orchids. 
From the boughs there hung in literal garlands the yard-long clusters of the flow- 
ering Coelogyne, and even the ground, which at other times is usually dull and 
bare, was now gay with the dehcate plant-like orchids of the Liparis variety up to 
the Spatho glottis- and Vanda-WkQ types that grow in bushes almost up to a man's 

Mjoberg's expedition to Mt. Murud resulted in extensive collections of a wide 
range of plants and animals, many of which were pubUshed on by various authors 
in volume 3 of the Sarawak Museum Journal (1928). Most of his fern collections were 
reported on by Bonaparte (1923a). A few pteridophytes and the gymnosperms 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 87 

and angiosperms were listed by Merrill (1928), who described 19 new species from 
the Murud collection (eight of which are no longer recognized). Merrill noted that he 
sent a first lot of orchids to Oakes Ames, and a second collection was sent directly 
by Mjoberg to Ames, but the Mjoberg orchids apparently were not reported 
upon. Jeffrey Wood of Kew has recently had on loan the Sarawak specimens of 
Dendrochilum from AMES, and none of four Mjoberg collections was annotated 
by Ames. 

BOTANICAL COLLECTORS ON MT. MURUD 

In addition to the Mjoberg expedition and our own field work, other botani- 
cal collectors who have visited the summit of Mt. Murud are as follows: B. L. 
Burtt of the Royal Botanic Garden, Edinburgh, with A. M. Martin, a plantsman 
from the Garden, collected there from September 26 to October 13, 1967. Their 
expedition was part of a joint activity with the Sarawak Forest Department, for 
which Ilias Paie was the principal collector. His collections from the summit area 
are dated from September 27 to October 13, 1967. Around October 10 he was 
joined briefly by J. A. R. Anderson, the Sarawak Forest Botanist, who had just 
come from Mt. Kinabalu, and independently left, exiting to Bario. Although the 
Burtt-Ilias expedition used the same camps and probably covered much of the 
same area, they worked independently, with Burtt and Martin concentrating on 
the shrub and herb flora and Ilias, with a team of climbers, concentrating on the 
trees. H. P. Nooteboom of the Rijksherbarium, Leiden, with Paul Chai of the 
Sarawak Forest Department, collected in the summit area from April 4-9, 1970; 
Yii Puan Ching of the Sarawak Forest Department collected on the summit ridge 
from September 10-13, 1982. Rena George and Rantai Jawa of the Sarawak 
Forest Department were on Mt. Murud between August 7 and 16, 1991, but we 
have seen only a few orchids collected by them; other collections they made 
probably are not yet processed. Jumaat Adam of Universiti Kebangsaan Malaysia 
was on Mt. Murud in 1994, but we have seen none of his collections. One Murud 
record of a collection by a Sarawak Museum Native Collector (Diplazium por- 
phyrorachis, based on a report by Price, 1983) is recorded in the enumeration. We 
do not know when this collection was made. 

Bonaparte (1923b) cited specimens of several species of ferns collected by J. 
C. Moulton from the summit and foot of Mt. Murud in December, 1914. Because 
Moulton (1915) indicated that his expedition did not reach Mt. Murud, these 
records appear to be in error. Furthermore, the numbers attributed by Bonaparte 
to Moulton do not agree with the numbering system otherwise used by Moulton. 
These specimens have not been included in the enumeration. 

OUR TRIP TO MT. MURUD 

Between April 10 and 23, 1995, a large component of the Universiti Malaysia 
Sarawak (UNIMAS) staff as well as scientists from other universities traveled to 
Bario, the largest settlement in the Kelabit Highlands, for the first major Bornean 
expedition sponsored by UNIMAS. The principal purpose of the expedition was 
to gather baseline information on various environmental aspects, including cultural, 
anthropological, and socio-economic values of the local communities, water quality, 
climate, geology, flora and fauna, and the river systems in the area. 



88 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Our trip to Mt. Murud was considerably more modest than Mjoberg's, start- 
ing with four botanists (one of whom left the group after the second day because 
of the rigors of the trail) and four porter-guides. We flew into Bario on a Malaysia 
Airlines Twin Otter rather than spending a month getting up the Baram River. 
We would agree with Mjoberg that attaining the summit of Mt. Murud requires 
^'tremendous exertions," but along the summit ridge the path cleared by his guides 
over 70 years ago may have helped us. In any case, during the Confrontation 
there was a Gurkha post on the summit, with the remains of a hut still in evidence 
in 1967 (Burtt, pers. comm.). The helicopter fuel and other supplies they left 
behind were equitably shared by the local population of Kelabits and neighboring 
Indonesians. This travel undoubtedly helped impress a track into the summit 
ridge, and further clearing was done in 1967 by the team of B. L. Burtt and Ilias 
Paie. In more recent years the ridge-crest trail has become reasonably well worn 
as something of a "tourist" route. 

We attended the Expedition Opening Ceremony in Bario on the morning of 
April 10, and, after feasting on a cow sacrificed for the occasion, found the local 
representative from Pa Lungan (3°48'32"N, 115°3ri5"E), the kampung (village) 
furthest from Bario toward Mt. Murud, who took us through a soaking rain on a 
four-hour hike to his house for the evening. The next morning he arranged for our 
porter-guides and by 9 a.m. we were off, seeing no further signs of current civiliza- 
tion until we returned to Pa Lungan six days later. What a privilege it was, one we 
had never before experienced, to walk through unbroken primary forest (or in 
some cases old secondary forest) for six straight days, notwithstanding that we 
were rained on much of the time and constantly under attack by voracious leeches. 

The first day's journey out of Pa Lungan took us over a fairly high ridge 
(Sekelun Hill, elev. ca. 1350 m, 3°50'42"N, 115°33'07"E) to Long Rapung (elev. 
ca. 1140 m, 3°5r54"N, 115°33'58"E), now a grove of giant bamboo [Dendrocala- 
mus asper (Schult. f.) Backer ex K. Heyne, called buluh betung in Kelabit and 
Malay] on the Dapur River floodplain, but formerly a kampung that was evacuated 
during the Confrontation. Our night in the open shelter, constructed entirely of 
bamboo with rattan lashings, was comfortable except that we had to position 
ourselves to avoid the drips that continued with the all-night rain. By 9 o'clock the 
next morning we were underway again and were able to cross the Dapur River on 
a huge tree trunk that had been undercut by the river and fallen across it. We 
were hardly 15 minutes out of camp before one of the first exciting discoveries of 
the expedition was made, a large vine of a vanilla orchid {Vanilla kinabaluensis 
Carr), the first time it had been recorded in Sarawak. We marked the spot and left 
the plant to be collected on the return. In spite of the great size of the plant, we 
could find only one beautiful large yellow flower. 

A slightly ridiculous nightly routine of the trip was the attempt to dry one's 
shoes and socks over the campfire. This was a particularly futile effort on the first 
day out of Long Rapung, because the Dapur River had to be crossed three more 
times (and the near-equal-sized Belaban River once), these crossings without the 
aid of a fallen tree, and it was not only shoes that were wet, but trousers up to the 
crotch or higher. These rivers were so swift and swollen by the recent heavy rains 
that our guides had to help us across. 

The next day's journey brought us to the Belaban Camp (elev. ca. 1160 m), at 
the confluence of the Dapur and Belaban Rivers. Here there was no bamboo hut, 
but we were lucky to have a tent that developed only minor leaks during another 
night of constant rain. Our porter-guides made do (and stayed about as dry) with 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 89 

a lean-to made of a large piece of nylon fabric suspended between two poles, and 
a ground-covering of bark pulled off the common tristania (Tristaniopsis) trees in 
strips 8 m long. Surely this campsite must be one of the most beautiful anyone has 
ever enjoyed, and, along with the soothing river sounds, came complete with a 
bath site on the mossy rocks of the Belaban. 

The following day's hike was probably the easiest of the entire trip, requiring 
only three to four hours to the final base camp (elev. ca. 1775 m, known by the 
local people as Punang Pa Bermusuh) before the ascent of Mt. Murud. This day 
was less strenuous in part because there were no large rivers to ford, the cHmb 
was relatively gentle, going up a total of about 500 m, and there were not so many 
deep, slushy root-enclosed puddles as we had to slog through along the Dapur 
River floodplain on the previous days. Fortunately we arrived at this base camp 
early enough to get the tent and lean-to set up before the rain resumed (and 
continued all night). Tristania bark again provided the floor mat for the porter- 
guides. This camp served us two nights, the one before the final ascent and upon 
descent. We continued to marvel at how excellent a fire our porter-guides could 
build and maintain in spite of the firewood's being sopping wet. They achieved 
wonderful fires with the bamboo at Long Rapung, and with fresh green tristania 
wood at the other camps. 

The next morning we left the base camp early, with the objective of getting to 
the summit as quickly as possible, which turned out to be nearly five hours later. 
The trail ascended steeply, first up through an oak-laurel lower montane forest, 
which then turned into a lower stature mossy forest just below the crest of the 
summit ridge at about 2,000 m (3°54'18"N, 115°30'50"E). We made it to this point 
in less than an hour, and initially thought that once on the ridge, the going would 
be easy. The actual situation was just the opposite. Along the ridge, the trail was 
deeply worn into tree roots and branches, and went up and down interminably 
with climbs and drops of 3-6 m, so that in 2.5 hours we had gained hardly more 
than 300 m in elevation through this spectacular dwarfed mossy elfin forest with 
the trees sometimes only 3-4 m high and rarely attaining more than 8 m. At about 
11 a.m. we thought we were approaching the highest point along the ridge, but 
this was not actually reached until more than an hour later. 

The final ascent to Mt. Murud's highest point (3°54'17"N, 115°29T9"E) was 
achieved without the soaking to which we had become accustomed, but we had 
been there less than half an hour when the rain started again. On reaching the 
summit we immediately began the collecting activities. After about an hour we 
started our way slowly back toward base camp (along that terrain one can only go 
slowly, but also to collect as much as possible along the summit ridge). The rain 
was incessant and hampered collecting, and the clouds obscured many of the 
beautiful views we should have been able to enjoy. 

We continued collecting all the way back to the base camp, where we arrived 
just as it was getting dark. Had we not reached it before dark we would have had 
to spend the night in the open in the rain, because the trail was too difficult and 
dangerous to negotiate in darkness. Again it rained all night, but our tent provided 
some escape from the water. 

One of the nicest discoveries of the journey was the vanilla plant noted above, 
but we were also able to document with a specimen the occurrence of Raffiesia 
pricei Meijer in Sarawak. This species was previously collected only from the Mt. 
Kinabalu area in Sabah (and more recently in Brunei). The plant was growing as a 
parasite on a huge Tetrastigma vine near the small stream Pa Parabao, about 45 



90 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

minutes walking distance from Long Rapung toward Sekelun Hill. Another worth- 
while discovery was the slipper orchid, Paphiopedilum javanicum var. virens (Rchb. 
f.) Stein, near the top of Sekelun Hill. In total our collections amounted to a little 
over 70 numbers. We arrived back in Bario just after midday on April 17, almost 
exactly one week after setting out on the trip. 

METHODS 

The enumeration provided below is incomplete but represents all collections 
reported by Bonaparte (1923a), Merrill (1928), our own collections, those encoun- 
tered in the Sarawak Forest Department Herbarium (SAR), the Edinburgh Bo- 
tanic Garden (E), the Royal Botanic Gardens, Kew (K), and the Rijksherbarium, 
Leiden (L). The nomenclature and taxonomic concepts applied by Bonaparte and 
Merrill have been updated to the extent possible without a detailed study of all 
the relevant material. The fern specimens Mjoberg submitted to Bonaparte are 
now in the herbarium of the Museum National d'Histoire Naturelle, Paris (P). 
Merrill (1928) stated that types of the species he described based on the Mjoberg 
collections are in the Herbarium of the University of California (Berkeley) (UC). 
Presumably the first set of non-types is also deposited there, but he noted addi- 
tionally that a set was deposited in the Natural History Museum, London (BM). 
The latter specimens have not been examined in the present study, but some 
Mjoberg collections, including isotypes, have been found in K. 

The principal set of the Burtt and Martin collections is in E, with some dupli- 
cates in SAR. Burtt supphed to us the data for many of his collections and provid- 
ed a loan of others that have been studied at K. The first set of Nooteboom and 
Chai collections is in L, and some duphcates have been found in K and SAR (a 
few specimens have been located in K that could not be found in L). The first set 
of Sarawak Forest Department collections is in SAR, with many duplicates in K 
and/or L. The specimens collected by Beaman and Anderson are deposited at 
UNIMAS, K, and MSC, with additional duplicates still to be distributed. 

Only collections from above 1,500 m are included in the enumeration (there- 
fore, the collections of Vanilla, Rafflesia, and Paphiopedilum noted above are not 
listed). For some specimens recorded from below that elevation it cannot be 
ascertained if the materials actually came from Mt. Murud or from nearby locali- 
ties. The most interesting physiognomic and biogeographic aspects of the Murud 
flora are encountered above 1,500 m (mostly above 2,000 m), so this seems an 
appropriate lower limit. The elevation data provided by Bonaparte (1923a) and 
Merrill (1928) for Mjoberg collections are not in all cases readily convertible into 
a computer database, particularly when such statements as "at high altitudes" 
(e.g., Carex cruciata) and "from above an altitude of 1900 m" (e.g., Rhododen- 
dron crassifolium, reported as R. munidense), or elevations were given pertaining 
to two different mountains in Sarawak. Labels on the relevant Mjoberg specimens 
we have seen give the elevation as 1900 to 2400 m. 

It might have been preferable to use 1800 m or 6000 ft as the lower boundary 
for collections to be included in this study, because this elevation more strictly 
defines the Mt. Murud summit ridge, whereas the 1500 m or 5000 ft contour 
extends rather widely from it. To choose the higher limit, however, would have 
resulted in excluding a number of Mjoberg collections we thought desirable to 
record. Future additions to the Murud summit flora might appropriately be hmited 



1997 BEAMAN& ANDERSON: SUMMIT FLORA OF MT.MURUD 91 

to the 1800 m contour and above. If one attempted to document the Murud flora 
by including species below 1500-1800 m, the mountain and its flora would become 
ill-defined. 

Locality data provided by the different collectors have been variously ex- 
pressed, making it necessary to standardize them. The latitude-longitude coordi- 
nates we recorded were obtained with a Magellan Trailblazer™ GPS unit, but this 
instrument could not always be used because of obstructing vegetation cover. 
Localities have been stated as precisely and briefly as possible, based on informa- 
tion provided by the labels. Sometimes this is no more precise than just "Mt. 
Murud," but if accompanied by an elevation figure, the locahty becomes some- 
what more precise. During the 1967 expedition of B. L. Burtt, A. M. Martin, and 
Ilias Paie five camps were used, of which Camps III, IV, and V were above 1500 
m (5000 ft) and were along (not on the crest of) the summit ridge. Because many 
of their collections are located relative to these camps, the following information 
supplied by Burtt or interpreted from a map he provided is indicated in Table 1. 

Achieving a complete set of determinations in a flora as poorly known as that 
of Mt. Murud and the rest of Borneo is a daunting task. For this project, as for 
many floristic efforts, the specimens are scattered in different herbaria and have 
had to be examined at different times, and have not all been seen. The enumera- 
tion therefore includes a rather high percentage of incompletely determined speci- 
mens and taxa named with varying levels of uncertainty. When the expression 'cf.' 
separates a generic name and specific epithet, this means that we think the speci- 
mens so identified might be a particular species but are uncertain of the identifica- 
tion; this is equivalent to putting a query (?) after the name. When the expression 
'aff.' separates a generic name and specific epithet, it means that we are fairly sure 
that the taxon so identified is not the one named, but is alHed to it. We have used 
the expression 'sp.' (or sp. 1, sp. 2) when we are unable to identify a specimen(s) 
but believe it (them) to be different from other taxa in the list. The expression 
indet.' has been used when material is undetermined or insufficiently studied. 

The overall concept of a floristic enumeration of the type provided in this 
paper was outlined by Beaman and Regalado (1989) for the flora of Mt. Kinabalu 
in Sabah, Malaysia. An integrated system of computer programs used for data 
editing and printing enumerations (e.g., Parris et al. 1992; Wood et al. 1993; Bea- 
man & Beaman 1993) was written in the dBASE IV programming language by 
Reed Beaman. These programs allow access to any aspect of the database through 



Sarawak Forest Depart 



Near the Komap River, ca. 4 km W of Ba Kelalan 
5°36'E On the Bor River at Long Rapata, ca. 4.2 km SW of Camp I 
5°32'E N side of the summit ridge, ca. 4.5 km S of Camp II 
5°32'E S side of the summit ridge, ca. 1 .7 km W of Camp III 
5°31 'E S side of the summit ridge, ca. 4.2 km WSW of Camp IV 



92 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

a menu system. Six principal relational data files were employed. Two of these 
contain data on specimens, including types. Taxonomic, nomenclatural, and bib- 
liographic information is linked from other files. Menus facilitate entering and 
editing specimen and taxon data, globally replacing various expressions, such as 
changing an author's name or abbreviation, indexing and querying the database, 
computing a summary of elevation ranges for taxa, numbering taxa, making an 
index to numbered collections, and printing enumerations of all taxa in the data- 
base or of selected families or genera. 

An earlier paper on the Murud summit flora by Beaman (1997) has been 
prepared, but that account was written before there was opportunity to record 
any of the specimens in E, L, and most of those that we have now been able to 
examine in K. The present enumeration is considerably more extensive than the 
previous account, but many species still may have been missed. It is also unfortu- 
nate that we have not been able to examine the Mjoberg collections in AMES, P, 
and UC, but at least those in P and UC have been listed by Bonaparte and Merrill. 



PHYTOGEOGRAPHY 

Mjoberg (1925) noted a strong biogeographic relationship between Mt. Mu- 
rud and Mt. Kinabalu. He indicated that he "had not been working many days in 
the new field of exploration before I realized the great affinity between the fa- 
mous fauna of Kinabalu and that of Mt. Murud situated more than a hundred 
miles farther south. Among the more conspicuous forms of life, the birds afforded 
clear evidence. Altogether I found nine birds more than one hundred miles far- 
ther south of their previously known home. To this we may add quite a number 
previously recorded only from Kinabalu and Mt. Dulit, and I feel confident that 
many more of what hitherto were considered exclusively Kinabalu birds would 
have been found not only on Mt. Murud but also in the intervening regions 
between Kinabalu and Mt. Murud and Mt. Dulit and even much farther south, 
proving that we have a continuous avifauna right through the central mountains 
of Borneo from Kinabalu in the north to Mt. Poi in the west and possibly with 
many forms south to the Miiller and Schwaner Mountains." 

Mt. Mulu (2377 m), 65 km WSW of Mt. Murud, is a close geographical neigh- 
bor and is only about 50 m lower. Considering their proximity and the fact that 
both mountains have a sandstone geology, one might expect a strong phytogeo- 
graphic relationship, which our present data do not support. B. L. Burtt noted 
some years ago (unpubl. report) that Mt. Mulu is a different type of mountain, 
rising rapidly from 60 m to the summit. There is no area of surrounding highlands 
as at Mt. Murud, which would affect climatic conditions. 

Gunung Mulu National Park was the subject of intensive botanical explora- 
tion in 1977-78 by the Royal Geographic Society and the Sarawak Government. 
During that time 115 scientists and assistants spent over 10,000 man-days in the 
Park (Jermy 1984). Various accounts based on this research have been published. 
A complete inventory of the Mulu flora was contemplated but has not appeared, 
although a preliminary list of angiosperms and gymnosperms was compiled by 
Anderson and Chai (1982). A comparison of the summit flora of Mt. Murud with 
the entire Mulu Park flora, an area of 52,864 ha embracing a great diversity of 
habitats from lowland peat-swamp forest, limestone outcrops, and kerangas vege- 
tation, would not be meaningful. 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 93 

Martin (1977) provided two tables listing common shrubs (Table 21) and 
miscellaneous collections (Table 22) from the summit of Mt. Mulu. These lists 
include just 33 taxa, of which 20 (61%) are common to Mt. Murud. We have 
examined the Mulu species lists for pteridophytes (Parris et al. 1984) and orchids 
(Wood 1984) to see how many species in these more detailed lists are common to 
the two mountains. In the case of fully determined pteridophytes, 19 taxa out of 
31 total (61%) recorded from Mt. Murud are also recorded on Mt. Mulu. Among 
the orchids only 8 fully determined taxa out of 42 total (19%) from Mt. Murud 
are also recorded for Mt. Mulu. 

Another mountain system about 150 km SW of Mt. Murud that could be 
considered for floristic comparison is Mt. Dulit in central Sarawak. The Dulit 
range, composed of Miocene limestone rising to an elevation of ca. 1460 m, was 
the subject of what must have been one of the most successful university student 
expeditions ever undertaken (Anonymous 1952). Notwithstanding that some 2,500 
plants were collected (Harrisson 1933) and many papers subsequently pubhshed 
on the Duht vegetation and flora, no list of species has been produced. 

While identifying the plant collections from Mt. Murud and searching the 
herbaria for specimens from there, we have come to believe that each mountain 
in northern and central Sarawak has its own unique constellation of species. Apart 
from the studies of Mt. Mulu and Mt. Dulit, most other neighboring mountains 
have not been the subject of published reports, although many specimens have 
been accumulated that would permit the writing of florulas or development of 
computer databases. 

One of the more intensively collected adjacent locations is Batu Lawi, twin 
columnar sandstone peaks only 12 km WSW of Mt. Murud. The summit of the 
higher peak apparently is still botanically unexplored. Some species are common 
to Mt. Murud and Batu Lawi, but frequently one notes that a species has been 
found on one mountain and not the other. Additionally, even with species com- 
mon to the two areas, minor differences in the specimens are sometimes apparent. 
The first expedition to Batu Lawi was made by J. C. Moulton in 1911. His report 
(Moulton 1912) includes appendices by H. N. Ridley on general plant collections, 
by J. J. Smith on orchids, and by E. B. Copeland on ferns. Although of historical 
interest, especially because of the new species described, these accounts do not 
provide a satisfactory basis for a phytogeographic comparison of Batu Lawi with 
Mt. Murud. The Moulton expedition required about 2.5 months from Kuching 
and back, but only two days were spent around the base of Batu Lawi. 

Sarawak Forest Department collectors have visited the Kelabit Highlands 
many times and assembled extensive collections from relatively high elevations in 
the Tama Abu Range, Apo Duat on the Indonesian border, Apad Keruma, Batu 
Buli, Batu Lawi, and other areas. The species that have been collected sometimes 
are represented in the Murud summit flora, but often they have not been found 
on Mt. Murud. 

The most sahent phytogeographic relationship of Mt. Murud appears to be 
with Mt. Kinabalu in northern Sabah. A major motivation for our trip to Mt. 
Murud was to obtain a personal view of similarities in the floras of these two 
mountains. The database we are preparing for a full enumeration of the flora of 
Mt. Kinabalu provides a basis for comparison, although, as with Mt. Mulu, com- 
paring a summit flora with a far greater elevational range and edaphic diversity 
than that of Mt. Murud complicates the consideration. The Kinabalu database we 
have assembled includes species from elevations as low as about 300 m to the 



94 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

summit at 4101 m. Likewise, Mt. Kinabalu has various geological substrates not 
found on Mt. Murud. Parris (1997) has recently reported on the phytogeography 
of Mt. Kinabalu pteridophytes, but she does not consider Mt. Murud per se, and 
the phytogeographical elements she recognizes, e.g., Borneo endemics and Sabah 
endemics, are too inclusive for use in the present analysis. 

In the enumeration below, 260 fully determined taxa (i.e., those not listed as 
'cf.', 'aff.' or 'sp.') can be used in a comparison with the Kinabalu flora. Of this 
number 184 taxa (71%) are common to Mt. Kinabalu and Mt. Murud. Many of 
the species are found in other localities as well, so it cannot be said that the 
Kinabalu-Murud relationship is unique. Nevertheless, the similarity of the flora of 
Mt. Murud to that of Mt. Kinabalu appears stronger than to the floras of Mt. 
Murud's geographically closer neighbors. As indicated above, only 61% of the 
Murud pteridophytes and 19% of the Murud orchids are also common to Mt. 
Mulu. In contrast, 24 pteridophyte taxa (77%) and 31 orchid taxa (74%) are 
common to Mt. Murud and Mt. Kinabalu (Kinabalu pteridophyte data from Par- 
ris et al., 1992; Kinabalu orchid data from Wood et al., 1993). The phytogeograph- 
ic data thus support the observation of Mjoberg that there are strong similarities 
in the species composition of the biota of Mt. Murud and Mt. Kinabalu. 

Among particularly interesting disjuncts between Mt. Kinabalu and Mt. Murud 
are Centrolepis philippinensis Merr., Patersonia lowii Stapf, and Gentiana bor- 
neensis Hook, f., taxa that occur at very high elevations on Mt. Kinabalu. Some of 
the more unusual discoveries in the Murud flora, including these three species, 
come from a basin on the north side a little below the summit in an area of 
extensive exposure of nearly bare sandstone. Noteworthy Murud disjunctions also 
occur in other directions, as for example Oreobolus kiikenthalii Steenis, otherwise 
known only in northern Sumatra and the Malay Peninsula, and Polygala oreotrephes 
B. L. Burtt in the Malay Peninsula. 

ENUMERATION 

The enumeration includes 35 pteridophytes (5 fern allies and 30 ferns), 7 gymno- 
sperms, 96 monocotyledons, and 207 dicotyledons. The total number of taxa is 
345, representing 85 families and 173 genera, based on 945 specimen records from 
ca. 700 collections; 85 taxa (25%) are incompletely determined or determined 
with uncertainty. The Orchidaceae are the largest family, with 49 species and 
varieties, and the largest genus is Rhododendron, with about 18 species and vari- 
eties. Twenty-seven new taxa of vascular plants have been described from the 
Murud summit area, of which 16 are currently recognized. 



BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 

FERN ALLIES 
L LYCOPODIACEAE 



L.Ll. Huperzia phlegmaria (L.) Rothm. [listed by Merrill (1928) as Urostachys 
phlegmaria (L.) Herter]. 

Collections. Mt. Murud: Mjoberg 73 p.p. (UC); Mt. Murud r[dgh, SE side: 1700- 
!000 m, Beaman 11489 (K, MSC, UNIMAS). 

L.1.2. Huperzia serrata (Thunb. ex Murray) Trevis. 

Collection. Mt. Murud between Camps J I and III: 1500 m, Burtt & Martin B. 5220 (E). 

.1.3. Huperzia verticiilata (L. f.) Trevis. [listed by Merrill (1928) as Urostachys 
verticillata (L. f.) Herter]. 



1.2. Lycopodiella 

ua (L.) Pic. Serm. 
Collection. Mt. Murud Camp IV: 1900 m, Burtt & Martin B. 5385 ( 

2. SELAGINELLACEAE 
2.1. Selaginella 
2.1.1. Selaginella rugulosa Ces. 



FERNS 

3. ASPLENIACEAE 
3.1. Asplenium 
.1. Asplenium nidus L. [listed by Bonaparte (1923a)]. 
Collection. Mt. Murud: 1500-1800 m, Mjoberg 204 p.p. (P). 



CONTR. UNIVERSITY OF MICHIGAN HE 

4. BLECHNACEAE 

4.1. Blechnum 

Blechnum sp. [listed by Bonaparte (1923a) as Blec, 
cerum f. integrum Bonap.]. 



itype of B. capense 



5. CYATHEACEAE 
5.1. Cyathea 
5.1.1. Cyathea capitata Copel. [listed by Bonaparte (1923a)]. 



5.1.2. Cyathea indet. 

Collection. Mi. Murdd summit: 2400 m, Nooteboom & Chai 2009 (L). 

6. DENNSTAEDTIACEAE 
6.1. HisnoPTERis 
6.1.1. Histiopteris incisa (Thunb.) J. Sm. [listed by Bonaparte (1923a)]. 
Collection. Mr. Murud: 1500-1800 m, Mjoberg 108 (P). 

7. DIPTERIDACEAE 

7.1, DiPTERlS 

7.1.1. Dipteris conjugata Reinw. [listed by Bonaparte (1923a) as D. conjugata \ 

alpina Christ]. 

Collection. Mr. Murud summit: 2400 m, Mjoberg s.n. (P). 

7.1.2. Dipteris lobbiana Moore 

Collection. Mr. Murud: 1500-1800 m, Mjoberg s.n. (P) [doubtfully occurring as 1 
as indicated, but so listed by Bonaparte (1923a)]. 

7.1.3. Dipteris novoguineensis Posth. 

Collections. Mt. Murud: 2200 m, Nooteboom & Chai 2039 (L); Mt. Murud N 
summit: Burn & Martin B. 5472 (E). 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 97 

7.1.4. Dipteris quinquefurcata Christ [listed by Bonaparte (1923a), but probably £ 
form of the polymorphic D. conjugata Reinw.]. 

Collection. Mr. Murud: 1500-1800 m, Mjdbergs.n. (P). 

8. DRYOPTERIDACEAE 
8.1. Dryopsis 
8.1.1. Dryopsis indet. 

Collection. Mr. Murud, path to summit: 1800 m, Nooteboom & Chai 1960 (L). 

9. GLEICHENIACEAE 
9.1. Gleichenia 
9.1.1. Gleichenia dicarpa R. Br. 



9.1.2. Gleiclienia peitophora Copel. var. peltophora 

Collection. Mt. Murud N of summit: 2100 m, Burn & Martin B. 5465 A (K). 



9.2.1. Sticherus hirtus (Blume) Ching var. paleaceus (Baker) Parris [listed by 
Bonaparte (1923a) as Gleichenia hirta Blume]. 

Collection. Mt. Murud: 1500-1800 m, Mjdbergs.n. (P). 

10. GRAMMITIDACEAE 
10.1. Ctenopteris 

10.1.1. Ctenopteris taxodioides (Baker) Copel. [listed by Bonaparte (1923a) and 
Merrill (1928) as Poly podium taxodioides Baker]. 

Collections. Mt. Murud: 2400 m, Mjoberg 7 (K), 70 (UC), 1500-1800 m, 256 (P). 



10.2. Prosaptia 

10.2.1. Prosaptia contigua (G. Forst.) C. Presl [listed by Bonaparte (1923a) . 
Davallia contigua (G. Forst.) J. Sm.]. 

Collections. Mt. Murud: 1500-1800 m, Mjdbergs.n. (P); Mt. Murud summit: 2400 i 



98 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

10.3. SCLEROGLOSSUM 

10.3.1. Scleroglossum minus (Fee) C. Chr. 

Collection. Mr. Murud ni:ar Camp V: 2100 m, Burtt & Martin B. 5444 (E). 

11. HYMENOPHYLLACEAE 

11.1. Macroglena 

11.1.1. Macroglena meifolia (Bory ex Willd.) Copel. [listed by Merrill (1928) as 

Trichomanes pliima Hook.]. 

Collections. Mt. Murud: above 1200 m, Mjoherg 67 (UC), 2300 m, Nooteboorn & 
Chai2014(L). 

11.1.2. Macroglena schlechteri (Brause) Copel. 
Collection. Mr. Murud: 2100 m. Burn c& Martin B. 5356 (K). 

11.2. Pleuromanes 
11.2.1. Pleuromanes album (Blume) Parris 

Collection. Mt. Murud: 1900 m, Burn & Martin B. 5392 (E). 

12. OLEANDRACEAE 
12.1. Oleandra 
12.1.1. Oleandra neriiformis Cav. [listed by Bonaparte (1923a)]. 
Collection. Mt. U\\\<v\r. 1500-1800 m, Mjoherg 182 (P). 

13. PLAGIOGYRIACEAE 
13.1. Plagiogyria 

13.1.1. Plagiogyria egenolfioides (Baker) Copel. var. latipinna (Copel.) Zhang & Noot. 
Collection. Mr. Murud, path to summit: 1800 m, Nooteboorn & Chai 1945 (L). 

13.1.2. Plagiogyria pycnophylla (Kunze) Mett. [listed by Bonaparte (1923a)]. 



13.1.3. Plagiogyria tuberculata Copel. [listed by Bonaparte (1923a) ; 
pinnata Bonap.]. 



1997 BEAMAN& ANDERSON: SUMMIT FLORA OF MT.MURUD 99 

14. POLYPODIACEAE 

14.1. Selliguea 

14.1.1. Selliguea albidosquamata (Blume) Parris [listed by Bonaparte (1923a) and 

Merrill (1928) as Polypodium alhidosquamatum Blume]. 

Collection. Mi. Murud: 1900 m, Mjoherg 71 (UC). 

14.1.2. Selliguea enervis (Cav.) Ching [listed by Merrill (1928) as Polypodium 

triquetrum Blume]. 

Collection. Mt. Murud: Mjoherg 68 (UC). 

15. SCHIZAEACEAE 

15.1. SCHIZAEA 

15.1.1. Schizaea malaccana Baker var. malaccana 

Collections. Mr. Muri 
summit: 2400 m, Nooteboofr. 

16. THELYPTERIDACEAE 

16.1. CORYPHOPTERIS 

16.1.1. Coryphopteris pubirachis (Baker) Holttum var. sulawesica Holttu 

Collection. Mt. Murud N of summit: Burtt & Martin B. 5462 (E). 

16.1.2. Coryphopteris viscosa (Baker) Holttum 
Collection. Mt. Murud summit ridge: Burtt & Martin B. 5488 (E). 

17. VITTARIACEAE 
17.1. VlTTARIA 

17.1.1. Yittaria indet. 

Collection. Mt. Murud: 2200 m, Nooteboom & Chai 2028a (L). 

18. WOODSIACEAE 

18.1. DiPLAZIUM 

18.1.1. Diplazium porphyrorachis (Baker) Diels 



100 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

GYMNOSPERMS 
19. ARAUCARIACEAE 
19.1. Agathis 
19.1.1. Agathis kinabaluensis de Laub. 

Collections. Mt. Murud between 1st and 2nd summits: 2300 m, Yii S. 44461 (K); Mt. 
MuRUD JUST BELOW SUMMIT: 2400 m, Nooteboom & Chai 2013 (K, L); Mt. Murud summit 
ridge: 2300-2400 m, Beaman 11464 (K, MSC, UNIMAS). 

20. PHYLLOCLADACEAE 
20.1. Phyllocladus 
20.1.1. Phyllocladus hypophyllus Hook. f. 



21. PODOCARPACEAE 
21.1. Dacrycarpus 

21.1.1. Dacrycarpus imbricatus (Blume) de Laub. [listed by Merrill (1928) ; 
Podocarpiis javaniciis (Burm. f.) Merr.]. 

Collection. Mt. Murud: 1500 m, Mjoberg 99 (UC). 

21.2. Dacrydium 

21.2.1. Dacrydium beccarii Pari, in DC. [listed by Merrill (1928)]. 
Collection. Mt. Murud: 1200 m, Mjoberg 100 (UC). 

21.2.2. Dacrydium gibbsiae Stapf 

Collections. Mt. Murud near summit: 2100 m, lUas S. 26505 (K); Mt. Murud summi 
2400 m, Anderson & Ilias S. 26471 (K); Mt. Murud summit ridge: 2300-2400 m, Beamc 
//^4fO(K, MSC, UNIMAS). 

21.2.3. Dacrydium xanthandrum Pilger 



21.3. Podocarpus 
21.3,1. Podocarpus neriifolius D. Don 

Collection. Mt. Murud summit ridge: 2300-2400 m, Beaman 11480 (K, UNIMAS). 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 101 

ANGIOSPERMS: MONOCOTYLEDONS 
22. ARACEAE 
22.1. SCINDAPSUS 

22.1.1. Scindapsus borneensis Engl. 

Collection. Mt. Murud, path to summit: 1700 m. Nooteboom & Chai 1937 (B, L, US). 

23. ARECACEAE 
23.1. Calamus 
23.1.1. Calamus gibbsianus Becc. 



23.1.2. Calamus mattanensis Becc. [listed by Merrill (1928) as C. fermglneus Becc, 
. fide Dransfield (1992)]. 



Collection. Mt. Murud: 1900 m, Mjoberg 126 (UC). 

23.1.3. Calamus pilosellus Becc. [listed by Merrill (1928); not recorded by Dr< 
field (1992) as occurring in the Kelabit Highlands]. 

Collection. Mt. Murud: 1900 m, Mjoberg 128 (UC). 

23.2. PiNANGA 
23.2.1. Pinanga capitata Becc. 

Collection. Mt. Murud 2nd summit: 2300 m, Yii S. 44442 (K, L). 

24. CENTROLEPIDACEAE 
24.1. Centrolepis 
24.1.1. Centrolepis philippinensis Merr. 

Collection. Mt. Murud N side: 2100 m, Bunt & Martin B. 5494 (E). 

25. CYPERACEAE 
25.1. Carex 
25.1.1. Carex cruciata Wahl. [listed by Merrill (1928)]. 
Collection. Mt. Murud: Mjoberg 113 (UC). 



1 02 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 

25.1.2. Carex filicina Nees 

Collection. Mr. Mdrud, ridge above Camp IV: 2000 m. Bum & Martin 

25.1.3. Carex sp. (sect. Mitratae) 

Collection. Mr. Murud summit: 2400 m, Nooteboom & Chai 2030 (L). 

25.2. Gahnia 

25.2.1. Gahnia javanica Zoll. & Moritzi ex Moritzi 

Collections. Mt. Murud Camp V: 2100-2300 m, Burtt & Martin B. 
Murud summit: 2400 m, Nooteboom & Chai 1981 (L); Mr. Murud su.mmi 
2400 m, Seaman 11465 (K, MSC, UNIMAS). 

25.3. Oreobolus 
25.3.1. Oreobolus kiikenthalii Steenis 



26. IRIDACEAE 
26.1. Patersonia 



26.1.1. Patersonia lowii Stapf 



27, MELANTHIACEAE 
27.1. Petrosavia 
27.1.1. Petrosavia stellaris Becc. 

Collection. Mt. Murud near Camp III: 1800 m, Bunt & Martin 1 

28. ORCHIDACEAE 
28.1. Appendicula 
28.1.1. Appendicula bilobuiata J. J. Wood 

of A. bilobuiata: SAR, isotype). 



1997 BEAMAN& ANDERSON: SUMMIT FLORA OF MT.MURUD 1 

28.1.2. Appendicula congesta Ridl. 

Collection. Mt. Murud Camp III: 1700 m, Bum & Martin B. 5233 (E, SAR). 

28.1.3. Appendicula foliosa Ames & C. Schweinf. 
Collection. Mt. Murud: 1600 m, Yii S. 44421 (K). 

28.1.4. Appendicula longirostrata Ames & C. Schweinf. 

Collection. Mt. Murud above Camp IV: 2000 m. Burn & Martin B. 5434 (E, SAR). 

28.1.5. Appendicula indet. 



28.2. Bromheadia 

28.2.1. Bromheadia crassiflora J. J. Sm. 

Collections. Mt. Murud: 2200 m, Nooteboom & Chai 2054 (L); Mt. Murud N side: 
2100 m, Burtt & Martin B. 5463 A (SAR); Mt. Murud between 1st and 2nd summits: 2300 
m, Yii S. 44482 (L, SAR). 

28.3. BULBOPHYLLUM 

28.3.1. Bulbophyllum anguliferuni Ames & C. Schweinf. 
Collection. Mt. Murud Camp III: 2000 m, Burtt & Martin B. 5252 (E). 

28.3.2. Bulbophyllum sopoetanense Schhr. 

Collection. Mt. Muirud near Camp IV: 1900 m, Burtt & Martin B. 5404 (E, SAR). 

28.3.3. Bulbophyllum aff. teres Carr 

Collection. Mt. Murud above Camp III: 1800 m, Burtt & Martin B. 5262 (E). 



28.4. Calanthe 
28.4.1. Calanthe speciosa (Blume) Lindl. 



28.4.2. Calanthe tenuis Ames & C. Schweinf. 

Collections. Mt. Murud above Camp IV: 2000 m, Burtt & Man 



104 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUM: 

28.4.3. Calanthe indel. 

Collections. Mr, Murdd: 1700 m. Nooteboom & Chai 1910 (L), 1800 m, 1969 (L). 

28.5. Chelonistele 
28.5.1. Chelonistele lamellulifera Carr 

Collection. Mt. Murud near Camp III: 1800 m. Burn & Martin B. 5272 (E, SAR). 

28.6. COELO(iYNE 

28.6.1. Coelogyne craticulaelabris Carr 

Collection. Mr. Murud N side: 2300 m, Bunt & Martin B. 5454 (E, SAR). 

28.6.2. Coelogyne hirtella J. J. Sm. 

Collections. Mr. Murud: 2200 m, 
III: 1800 m, Burtt & Martin B. 5256 (E; 

28.6.3. Coelogyne kinabaluensis Ames & C. Schweinf. 
Collection. Mt. Murud Camp III: 1700 m, Burtt & Martin B. 5236 (E). 

28.6.4. Coelogyne moultonii J. J. Sm. 

Collections. Mt. Murud Camp III: 1 
RUD NEAR Camp III: 1600 m, Burtt & Mart 

28.6.5. Coelogyne planiscapa Carr var. planiscapa 
Collection. Mt. Murud Camp III: 1700 m, Burtt & Martin B. 5243 (E). 

28.6.6. Coelogyne radioferens Ames & C. Schweinf. 

Collections. Mt. Murud near 
(E, SAR); Mt. Murud/Dapur Riv 

28.6.7. Coelogyne tenompokensis Carr 
Collection. Mt. Murud Camp III: 1700 m, Burtt & Martin B. 5245 (E, SAR). 

28.6.8. Coelogyne indet. 



28.7. Cymbidium 
28.7.1. Cymbidium elongatum J. J. Wood, Du Puy & Shii 



EAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 



28.8. Dendrobium 

28.8.1. Dendrobium alabense J. J. Wood 

Collections. Mt. Murud N side: 2100 m, Burtt & Martin B. 5480 (E); Mt. Murud 
ABOVE Camp IV: 1600 m, Burtt & Martin B. 5354 (E). 

28.8.2. Dendrobium cymbulipes J J. Sm. 

Collection. Mt. Murud near Camp III: 1800 m, Burtt & Martin B. 5333 (E). 

28.8.3. Dendrobium piranha C. L. Chan & P. Cribb 

Collection. Mt. Murud: 2200 m, Nooteboom & Chai 2032 (L, SAR). 

28.8.4. Dendrobium sp. (sect. Rhopalanthe) 

Collection. Mt. Murud summit ridge: 2300-2400 m, Beaman 11482 (K). 

28.8.5. Dendrobium indet. 

Collection. Mt. Murud: 2200 m, Nooteboom & Chai 2033 (L). 

28.9. Dendrochilum 

28.9.1. Dendrochilum crassifolium Ames 

Collections. Mt. Murud: 2400 m, Nooteboom & Chai 1995A (L); Mt. Murud Camp 
III: 1700 m, Burtt & Martin B. 5241 (E, K, SAR); Mt. Murud SW of Camp III: 1800 m, 
Burtt & Martin B. 5319 (E). 

28.9.2. Dendrochilum dewindtianum W. W. Sm. var. dewindtianum 

Collection. Mr. Murud summit ridge: 2300-2400 m, Beaman 11448 (K, MSC, UNIMAS). 

28.9.3. Dendrochilum galbanum J. J. Wood 
4430 (K, SAR); Mt. Murud SW 



: Rolfe 



. Mt. Murud: 1600 m, Yii S. 44402 (K); 
in B. 5299 (E, SAR). 



^.9.5. Dendrochilum imbricatum Ames 
Collection. Mt. Murud: 1900-2400 m, Mjobei 



106 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 

28.9.6. Dendrochilum lancilabium Ames 



28.9.7. Dendrochilum longipes J. J. Sm. 

Collections. Mt. Murud: 1900-2400 m, Mjoberg 66 (AMES); Mr. Murud 2nd summit: 
2300 m, Yii S. 44432 (K, L, SING); Mt. Murud near Camp IV: 1900 m, Burtt & Martin B. 
5399 (E); Mt. Murud, path to summit: 1800 m, Nootehoom & Chai 1954 (L). 

28.9.8. Dendrochilum muluense J. J. Wood 

t & Martin B. 5244 (E, SAR); Mr. 
}6]6 (K). 

28.9.9. Dendrochilum murudense (J.J. Wood) J. J. Wood 

Collections. Mt. Murud: 2400 m, 
folium var. murudense J. J. Wood); 
11459 (K, UNIMAS). 

28.9.10. Dendrochilum simplex J. J. Sm. 

Collection. Mt. Murud: 1900-2400 m, Mjoberg 49 (AMES). 

28.10. DlLOCHlA 

28.10.1. Dilochia cantleyi (Hook, f.) Ridl. 

Collections. Mt. Murud: 1700 m, Yii S. 44627 (K); Mt. Murud N 5 
2100 m, Burtt & Martin B. 5443 (E, SAR); Mt. Murud, Dapur River 
m, Ilias S. 26539 (K). 

28.10.2. Dilochia rigida (Ridl.) J. J. Wood 

Collections. Mt. Murud 2nd summit: 2200 m. Bias S. 26408 (K); Mi 
1st and 2nd summits: 2300 m, Yii S. 44486 (K, SAR); Mt. Murud summit 
m, Beaman 11455 (K); Mt. Murud, Dapur River headwaters: 2400 m, . 

28.11. Epigeneium 
28.11.1. 

Collection. Mr. Murud: 2200-2300 m, Noc 



28.12. Eria 
28.12.1. Eria aff. brookesii Ridl. 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 107 

28.12.2. Eria crassipes Ridl. 

Collections. Mr. Murud: 2400 m, Nooteboom & Chai 2065 (SAR), 2300 m. Yil S. 
44431 (K). 

28.12.3. Eria robusta (Blume) Lindl. 

Collections. Mr. Murud N side: 2100 m, Bum & Martin B. 5466 (E); Mt. Murud 
ABOVE Camp III: 1800 m, Burtt & Martin B. 5263 (E); Mt. Murud between 1st and 2nd 
summits: 2300 m, Yii S. 44491 (K); Mt. Murud summit ridge: 2300-2400 m, Beaman 11485 
(K, MSC, UNIMAS). 

28.12.4. Eria indet. 



28.13. LiPARis 

28.13.1. Liparis compressa (Blume) Lindl. var. compressa 

Collection. Mt. Murud: 1700 m, Rena, Rantai et al. S. 60335 (K). 

28.13.2. Liparis pandurata Ames 

Collection. Mt. Murud Camp III: 1700 m, Burtt & Martin B. 5237 (E, SAR). 

28.14. Nabaluia 

28.14.1. Nabaluia exaltata de Vogel 

Collections. Mt. Murud: 2100 m, llias S. 26466 (K), 2400 m, Rena, Rantai et al. S. 
60394 (K); Mt. Murud Camp III: 1600 m, Burtt & Martin B. 5260 (E); Mt. Murud above 
Camp III: 1800 m, Burtt & Martin B. 5259 (E, SAR). 

28.15. Oberonia 
28.15.1. Oberonia aff. griffithiana Lindl. 

Collection. Mt. Murud ni;ar Camp IV: 1900 m, Burtt & Martin B. 5393 (E). 

28.16. Phaius 
28.16.1. Phaius aff. pauciflorus (Blume) Blume 

Collection. Mt. Murud near Camp III: 1800 m, Burtt & Martin B. 5329 (E, SAR). 

28.17. Pholidota 
28.17.1. Pholidota gibbosa (Blume) de Vriese 



108 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

28.17.2. Pholidota mediocris de Vogel 

Collection. Mt. Murud near Camp III: 1800 m. Burn & Martin B. 5338 (E). 

28.18. Trichotosia 

28.18.1. Trichotosia cf. annulate Blume 

Collection. Mt. Murud/Belaban River headwaters: 1700 m. Ilias S. 26353 (K). 

28.18.2. Trichotosia aff. ferox Blume 

Collection. Mt. Murud N side: 2100 m. Burn & Martin B. 5464 (E). 

28.18.3. Trichotosia poculata (Ridl.) Kraenzl. 

Collection. Mt. Murud, path to summit: 1800 m, Nooteboom & Chai 1946 (L, SAR). 

28.18.4. Trichotosia sarawakensis Carr 



29. PANDANACEAE 
29.1. Freyciinetia 

29.1.1. Freycinetia kinabaiuana B. C. Stone 

Collection. Mt. Murud, path to sl>mmh: 1700 m, Nooteboom & Chai 1939 (L). 

29.1.2. Freycinetia rigidifoiia Hemsl. 

Collection. Mt. Murud/Belaban River: 1700 m, Ilias S. 26337 (K, L). 

29,2. Pandanus 
29.2.1. Pandanus papilio B. C. Stone 

Collection. Mt. Murud Camp IV: 2000 m. Bum & Martin B. 5431 (E). 

30. PHORMIACEAE 

30.1. DiANELLA 

30.1.1. Dianella ensifolia (L.) DC. [listed by Merrill (1928)]. 

Murud: 1500 m, Mjbberg 123 (UC), 2200 m. Nooteboom & Chai 



2026 (L). 



: ANDERSON: SUMMIT FLORA OF MT. MURUD 
30.1.2. Dianella javanica (Blume) Kunth 

Collection. Mt. Murud summit: 2400 m, Nooteboom & Chai 1999 (L). 

31. POACEAE 

31.1. ISACHNE 

31.1.1. Isachne kinabaluensis Merr. 



31.2. Racemobambos 
31.2.1. Racemobambos glabra Holttum 

Collection. Mt. Murud N side: 2100 m, Burtt & Martin B. 5495 (E). 

32. SMILACACEAE 
32.1. Smilax 



32.1.1. Smilax lanceifolia Roxb. 



32.1.2. Smilax sp. 1 

Collection. Mt. Murud, path to summit: 1700 m, Nooteboom & Chai 1906 (L). 

32.1.3. Smilax sp. 2 

Collection. Mt. Murud, path to summit: 1700 m, Nooteboom & Chai 1936 (L). 

33. XYRIDACEAE 
33.1. Xyris 
33.1.1. Xyris capensis Thunb. var. schoenoides (Mart.) Nilss. 

Collection. Mt. Murud Camp V: 2100-2300 m, Burtt & Martin B. 5484 (E). 



34. ZINGIBERACEAE 
34.1. Alpinia 



. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 



34.1.1. Alpinia glabra Ridl. 



Collections. Mt. Murud belo 


:w Camp III: 1500-1600 m, 


Burtt & Martin B. 53 


[t. Murud between Camps II anc 


ail: 1500 m, Biirtt & Martir 


I B. 5221 (E). 


4.1.2. Alpinia indet. 






Collection. Mt. Murud, pai h 


m SUMMIT. IHOO m, Nootebc 
34.2. Amomum 


wm & Chai 1959 [L). 




M. Sm. 





34.2.2. Amomum flavoalbum R. M. Sm. 

Collection. Mt. Murud Camp III: 1800 m. Burtt & Martin B. 5283 (E). 

34.2.3. Amomum aft. flavoalbum R. M. Sm. 

Collection. Mt. Murud Camp IV: 2000 m, Burtt & Martm B. 5351 (E). 

34.2.4. Amomum luteum R. M. Sm. 

Collection. Mr. Murud hi:i.ow Camp IV: ISOO m, Burtt & Martin B. 537 

34.3. BURBIDGEA 

34.3.1. Burbidgea nitida Hook. f. [listed by Merrill (1928)]. 
Collection. Mr. Murud: Mjoberg 122 (UC). 

34.3.2. Burbidgea schizocheila Hackett 

34.4. Etlingera 

34.4.1. Etlingera fimbriobracteata (K. Schum.) R. M. Sm. 
Collection. Mt. Murud Camp III: 1800 m, Burtt & Martin B. 5342 (E). 

34.4.2. Etlingera longipetiolata (B. L. Burtt & R. M. Sm.) R. M. Sm 
Collection. Mr. Murud hflow Camp IV: 2000 m, Burtt & Martin B. 534. 

34.4.3. Etlingera aff. muluensis R. M Sm. 

Collection. Mt. Murud Camp III: 1800 m. Burtt & Martin B. 5341 (E). 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT.MURUD 111 

34.4.4. Etiingera pubescens (B. L. Burtt & R. M. Sm.) R. M. Sm. 
Collection. Mt. Murud below Camp IV: 1800 m, Burtt & Martin B. 5366 (E). 

34.4.5. Etiingera punicea (Roxb.) R. M. Sm. 

Collection. Mt. Murud Camp III: 1800 m, Btirtt & Martin B. 5279 (E). 

34.5. Hedychium 
34.5.1. Hedychium cylindricum Ridl. [listed by Merrill (1928) as H. mjobergii 



34.5.2. Hedychium indet. 

Collection. Mt. Murud, path to summit: 1700 m, Nooteboom & Chai 1938 (L). 

34.6. HORNSTEDTIA 
34.6.1. Hornstedtia incana R. M. Sm. 

Collection. Mt. Murud Camp IV: 2000 m, Burtt & Martin B. 5402 (E). 

34.7. Plagiostachys 
34.7.1. Plagiostachys bracteolata R. M. Sm. 

Collection. Mt. Mlirud near Camp IV: 2000 m, Burtt & Martin B. 5429 (E, K). 

ANGIOSPERMS: DICOTYLEDONS 

35. ACANTHACEAE 
35.1. Strobilanthes 

35.1.1. Strobilanthes indet 

Collection. Mt. Murud, path to summit: 1700 m, Nooteboom & Chai 1933 (L). 

36. ACTINIDIACEAE 
36.1. Saurauia 

36.1.1. Saurauia amoena Stapf 



112 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

37. ANACARDIACEAE 

37.1. Toxicodendron 

37.1.1. Toxicodendron borneense (Stapf) Gillis 

Collections. Mr. Murud between 1st and z 
SAR), 2200 m, 44612 (K, SAR); Mr. Murud sum 
{K,UNIMAS). 

38. APIACEAE 
38.1. Hydrocotyle 
38.1.1. Hydrocotyle javanica Thunb. 

39. APOCYNACEAE 
39.1. Alyxia 
39.1.1. Alyxia oleifolia King & Gamble 

Collection. Mr. Murud: 2200 m, Nootehoom & Chai 2053 

39.2. Leuconotis 



Collection. Mr. 



40. AQUIFOLIACEAE 
40.1. Ilex 

40.1.1. Ilex harmsiana Loesn. 

Collections. Mt. Murud: 2100 m, Ilias S. 26453 (K, L), 2200 m, Nooteboom & Chai 
2020 (K, L), 2200 m, 2042 (L); Mt. Murud between 1st and 2nd summits: 2300 m, Yii S. 
44489 (K); Mt. Murud summit ridge: 2300-2400 m, Beaman 11468 (K, UNIMAS). 

40.1.2. Ilex havilandii Loes. [listed by Merrill (1928) as /. confertifolia Merr.]. 

Collections. Mt. Murud: 2200 m, Nooteboom & Chai 2037 (L, SAR); Mt. Murud 2nd 
summit: 2300 m, Yii S. 44425 (K, L); Mt. Murud Camp V: 2100-2300 m, Burn & Martin B. 5468 
(E); Mt. Murud between 1st and 2nd summits: 2200 m, Yii S. 44618 (K, L); Mt. Murud near 
summit: 2400 m, Mjoberg 93 (UC, holotype of /. confertifolia Merr.); Mt. Murud summit ridge: 
2300-2400 m, Beaman 11444 (K, MSC, UNIMAS), 2300-2400 m, 11483 (K, UNIMAS). 



BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 
41. ARALIACEAE 
41.1. Arthrophyllum 
. Arthrophyllum collinum Philipson 



41.2. SCHEFFLERA 

41.2.1. Schefflera mjobergii Merr. [listed by Merrill (1928)]. 

Collections. Mt. Murud: 1900 m, Mjoberg 117 (UC, holotype of S. mjobergii Merr.); 
Mt. Murud 2nd summit: 2000 m, Ilias S. 26386 (K, SAR); Mt. Murud Camp III: 1700 m, 
Burtt & Martin B. 5247 (E); Mt. Murud between 1st and 2nd summits: 2300 m, Yii S. 
44487 (K, SAR); Mt. Murud summit ridge: 2300-2400 m, Beaman 11479 (K, MSC, UNI- 
MAS); Mt. Murud, path to summit: 1800 m, Nooteboom & Chai 1961 (L, SAR); Mt. 
Murud/Dapur River headwaters: 2100 m, Ilias S. 26467 (SAR). 

41.2.2. Schefflera aff. remotiserrata Merr. 

Collection. Mt. Murud: 2200 m, Nooteboom & Chai 2052 (L, SAR). 

41.2.3. Schefflera sp. 

Collections. Mt. Murud between 1st and 2nd summits: 2200 m, Yii S. 44619 (K, L, 
SAR); Mt. Murud, Long Rapata/Ba Kelalan: 1700 m. Was S. 26557 (K, L); Mt. Murud, 
path to summit: 1800 m, Nooteboom & Chai 1958 (L, SAR). 



42. ASCLEPIADACEAE 

42.1. DiSCHlDIA 

ria R. Br. 

:amp III: 1700 m, Burtt & Martir, 



43. ASTERACEAE 
43.1. Vernonia 
, Vernonia phanerophlebia Merr. var. dulitensis Koster 

. Murud Camp IV: 



44. BALANOPHORACEAE 
44.1. Balanophora 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



44.1.1. Balanophora papuana Schllr. 



44.1.2, Balanophora indet. 

Collection. Mi. Muruo, path to summit: 1800 m, Nooteboom & Chai 1968 (L). 

45. BEGONIACEAE 
45.1. Begonia 

45.1.1. Begonia murudensis Merr. [listed by Merrill (1928)]. 

Collection. Mr. Murud: 1900-2400 m, Mjoberg 119 (UC, holotypc of B. muntdnms Merr.). 

45.1.2. Begonia sp. 

Collection. Mt. Murud, path to summit: 1700 m, Nooteboom & Chai 1920 (L). 

46. CAPRIFOLIACEAE 

46.1. Viburnum 

46.1.1. Viburnum hispidulum Kern 

Collections. Mt. Murud between 1st and 2nd summits: 2200 m, 7/7 S. 44621 (K); Mt. 
Murud summit: 2400 m, Nooteboom & Chai 2061 (L). 

47. CELASTRACEAE 

47.1. MiCROTROPIS 

47.1.1. IVIicrotropis valida Ridl. 

Collection. Mt. Murud, path to summit: 1800 m, Nooteboom & Chai 1943 (L). 

47.1.2. Microtropis wallichiana Wight ex Thwaites 

Collection. Mt. Murud N of summit: 2300 m, Nooteboom & Chai 2043 (L). 

47.2. Perrottetia 
47.2.1. Perrottetia alpestris (Blume) Loesn. 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 115 

48. CLUSIACEAE 

48.1. Garcinia 

48.1.1. Garcinia indet. 

Collections. Mr. Murud Camp IV: 2000 m, Ilias S. 26394 (L); Mt. Murud summit: 
2400 m, Nooteboom & Chai 2010 (L); Mt. Murud, path to summit: 1700 m, Nooteboom & 
Chai 1930 (L), 1800 m, 1964 (L); Mt. Murud/Belaban River: 1700 m, Ilias S. 26317 (L), 
1700 m, S. 26320 (L), 1700 m, S. 26322 (L), 1700 m, S. 26333 (L), 1700 m, S. 26342 (L). 

49. CUNONIACEAE 
49.1. Weinmannia 
49.1.1. Weinmannia aphanoneura Airy Shaw 

Collection. Mt. Murud summit ridge: 2400 m, Burtt & Martin B. 5487 (E, SAR). 

50. DAPHNIPHYLLACEAE 
50.1. Daphniphyllum 
50.1.1. Daphniphyllum glaucescens Blume subsp. borneense (Stapf) Huang 



51. DIPTEROCARPACEAE 
51.1. Vatica 

51.1.1. Vatica granulata Slooten subsp. sabaensis Ashton 

Collections. Mt. Murud: 1600 m, Yii S. 44403 (K); Mt. Murui 
m, Ilias S. 26312 (K). 

51.1.2. Vatica indet. 

Collection. Mt. Murud: 2400 m, Nooteboom & Chai 2069 (L). 

52. ELAEOCARPACEAE 
52.1. Elaeocarpus 
52.1.1. Elaeocarpus glaberrimus R. Knuth 



116 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 

Collections. Mt. Murud: 2100 m, Ilias S. 26452 (K), 2100 m, S. 26495 (K), 2300 m, Yii 
S. 44427 (K); Mt. Murud nhar Camp V: 2100 m, Ilias S. 26510 (K, L); Mt. Murud summit: 
2400 m, Nooteboom & Chai 2003 (L); Mt. Murud summit ridge: 2300-2400 ni, Beaman 
11484 (K, UNIMAS). 

52.1.2. Elaeocarpus murudensis Merr. [listed by Merrill (1928)]. 

Collections. Mt. Murud: 1900-2400 m, Mjoberg 83 (BM, isotype; UC, holotype of E. 
murudensis Merr.), 2200 m, Nooteboom & Chai 1976 (L); Mt. Murud between 1st and 
2nd summits: 2300 m, Yii S. 44471 (K, L); Mt. Murud summit: 2400 m, Nooteboom & Chai 
2068 (K, L); Mt. Murud summit ridge: 2300-2400 m, Beaman 11449 (K, MSC, UNIMAS). 

52.1.3. Elaeocarpus nanus Corner subsp. congestifolius (R. Knuth) Coode 
Collection. Mt. Murud summit: 2400 m, Nooteboom & Chai 1986 (L). 

52.1.4. Elaeocarpus sp. nov. aff. glaberrimus R. Knuth (fide Coode, pers. comm.) 
Collection. Mt. Murud summit: 2400 m, Nooteboom & Chai 1988 (L). 



53. EPACRIDACEAE 
53.1. Styphelia 
. Styphelia malayana (Jack) J. J. Sm. 



54. ERICACEAE 

54.1. DiPLYCOSIA 

54.1.1. Diplycosia acuminata Becc. 

Collections. Mt. Murud Camp III: 1800 m, Burtt & Martin B. 5288 (E); Mt. Murud 
Camp IV: 1900 m, Burtt & Martin B. 5410 (E). 

54.1.2. Diplycosia barbigera Sleumcr 

Collection. Mt. Murud/Belaban River: 1900 m, lUas S. 26373 (E, L, SAR). 

54.1.3. Diplycosia Ombriata Sleumer 

Collections. Mt. Murud 2nd summit: 2000 m, lUas S. 26384 (E, K, L); Mt. Murud 
ABOVE Camp IV: 2000 m, Burtt & Martin B. 5421 (E); Mt. Murud between 1st and 2nd 
summits: 2300 m, Yii S. 44478 (E, K, L); Mt. Murud summit: 2400 m, Nooteboom & Chai 
2002B (L, SAR), 2400 m, 2063 (L); Mt. Murud summit ridge: 2300-2400 m, Beaman 
11457 (K, MSC, UNIMAS). 



54.1.4. Diplycosia cf. fimbriata Sleumer 
Collection. Mt. Murud above Camp III: 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 117 

54.1.5. Diplycosia aff. fimbriata Sleumer 

Collection. Mt. Murud SW of Camp IV: 2100 m. Bum & Martin B. 5358 (E). 

54.1.6. Diplycosia microsalicifolia Argent 

Collections. Mt. Murud above Camp IV: 2000 m, Burtt & Martin B. 5419 (E); Mt. 
Murud between 1st and 2nd summits: 2300 m, Yii S. 44493 (K); Mt. Murud, Dapur River 
headwaters: 2000 m, Ilias S. 26447 (E, K, isotypes; SAR, holotype of D. microsalicifolia 
Argent). 

54.1.7. Diplycosia punctulata Stapf 

Collection. Mt. Murud 2nd summit: 2300 m, Yii S. 44447 (E, K, SAR). 

54.1.8. Diplycosia aff. saurauioides J. J. Sm. 

Collections. Mt. Murud 2nd summit: 2000 m, Ilias S. 26387 (L); Mt. Murud above 
Camp III: 2000 m, Burtt & Martin B. 5267 (E), 2000 m, 5268 (E, SAR); Mt. Murud 
between 1st and 2nd summits: 2300 m, Yii S. 44490 (E, K, SAR); Mt. Murud summit: 2400 
m, Nooteboom & Chai 1992 (K, L, SAR); Mt. Murud summit ridge: 2300-2400 m, Bea- 
man 11469 (K, MSC, UNIMAS). 

54.1.9. Diplycosia cf. scabrida Becc. 

Collection. Mt. Murud above Camp IV: 2000 m, Burtt & Martin B. 5430 (E, SAR). 

54.1.10. Diplycosia cf. urceolata Stapf 

Collections. Mt. Murud: 2300 m, Ilias S. 26487 (E, K); Mt. Murud 2nd summit: 2000 
m, Ilias S. 26381 (E, K, SAR), 2200 m, S. 26416 (E, K, SAR), 2300 m, Yii S. 44433 (K); Mt. 
Murud N side: 2300 m, Burtt & Martin B. 5452 (E, SAR); Mt. Murud above Camp III: 
2000 m, Burtt cfe Martin B. 5266 (E, SAR). 

54.1.11. Diplycosia indet. 



54.2. Rhododendron 

54.2.1. Rhododendron borneense (J. J. Sm.) Argent, A. L. Lamb & Phillipps 

subsp. villosum (J. J. Sm.) Argent, A. L. Lamb & Phillipps [listed by 
Merrill (1928) as R. cuneifolium Stapf var. subspathulatum Ridl.]. 

Collections. Mt. Murud: 1900 m, Mjoberg 98 (UC); Mt. Murud 2nd summit: 2200 m, 
Ilias S. 26403 (K); Mt. Murud above Camp IV: 1800 m, Burtt & Martin B. 5251 A (E); Mt. 
Murud between 1st and 2nd summits: 2300 m, Yii S. 44460 (K, L). 

54.2.2. Rhododendron burttii P. Woods 

'e Long Rapata: 1700 m, Burtt & Martin B. 5549 (E, 



118 CONTR. UNIVERSITY OF MICHIGAN HERBARIlJf 

54.2.3. Rhododendron buxoides Sleumer 



54.2.4. Rhododendron crassifoHum Slapf [listed by Merrill (1928) as R. miirudense 

Mcrr.]. 

Collections. Mi. Murud: 1900 m, Mjoberg 106 (L, fragment; UC, holotype of R. 
mumdense Merr.); Mt. Murud SW of Camp III: 1700 m, Burtt & Martin B. 5321 (E); Mt. 
Murud summit ridge: 2300-2400 m, Beaman 11474 (K), 2300-2400 m, 11478 (K); Mt. 
Murud, path to summit: 1800 m, Nooteboom & Chai 1940 (L). 

54.2.5. Rhododendron durionifoUum Becc. [listed by Merrill (1928) as R. mjohergii 

Mcrr.]. 

Collections. Mt. Murud: 2300 m, llias S. 26493 (K, L), 1900-2400 m, Mjoberg 105 (L, 
fragment; UC, holotype of R. mjobergii Merr.); Mt. Murud above Camp III: 1800 m, Burtt 
& Martin B. 5264 (E); Mt. Murud summit: 2400 m, Nooteboom & Chai 1989 (L); Mt. 
Murud summit ridge: 2300-2400 m, Beaman 11447 (K, MSC, UNIMAS). 

54.2.6. Rhododendron exuberans (Sleumer) Argent 

Collection. Mt. Murud Camp IV: 1800 m, Burtt & Martin B. 5351 A (E). 

54.2.7. Rhododendron himantodes Sleumer 

Collection. Mt. Murud Camp III: 1800 m, Burtt & Martin B. 5251 (E). 

54.2.8. Rhododendron micromalayanum Sleumer 

Collections. Mr. Murud Camp IV: 1900 m, Burtt & Martin B. 5396 (E); Mt. Murud, 
Long Rapata/Ba Kelalan: 1700 m, Bias S. 26555 (K): Mt. Mi;rud, path to summit: 1800 
I & Chai 1955 (L). 



54.2.10. Rhododendron orbiculatuni Ridl. 

Collections. Mt. Murud between 1st and 2nd summits: 2200 m, Yii S. 44602 (K, 
SAR); Mt. Murud near summit: 2300 m, Nooteboom & Chai 1972 (L); Mt. Murud sum- 
mit ridge: 2300-2400 m, Beaman 11486 (K, MSC, UNIMAS); Mt. Murud, Long Rapata/ 
Ba Kelalan: 1700 m, Uias S. 26556 (K); Mt. Murud, path to summit: 1900 m, Nooteboom 
& Chai 1967 (L). 

54.2.11. Rhododendron pneumonanthum Sleumer [listed by Merrill (1928) as R. 
jasminiflorum Hook.]. 



MAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 



Collections. Mt. Murud: 2300 r 
1900-2400 m, 104 (UC); Mt. Murud 

54.2.12. Rhododendron polyanthemum Sleumer 

Collection. Mr. Murud SW of Camp III: 1700 m, Bunt & Martin B. 5317 (E). 

54.2.13. Rhododendron retivenium Sleumer 

Collection. Mr. Murud below Camp III: 1600 m, Burtt & Martin B. 5306 (E). 

54.2.14. Rhododendron rugosum Low ex Hook. f. 

Collection. Mt. Murud summit: 2400 m, Burtt & Martin B. 5450 (E). 

54.2.15. Rhododendron stenophyllum Hook. f. ex Stapf subsp. angustifolium (J. J. 

Smith) Argent, A. L. Lamb & Phillipps 

Collection. Mt. Murud SW of Camp III: 1700 m, Burtt & Martin B. 5322{E) 

54.2.16. Rhododendron yongii Argent 
ud SW OF Camp IV: 2100 

54.2.17. Rhododendron sp. nov. ? 

Collection. Mt. Murud summit ridge: 2300-2400 m, Beaman 11441 (K, MSC, UNIMAS). 

54.2.18. Rhododendron sp. (subsect. Pseudovireya) 



Collections. Mt. Murud summi 
MAS), 2300-2400 m, 11467 (E, K, U: 


T ridge: 2300-2400 m, Beaman 11450 (E, K, UNI- 
NIMAS). 


54.2.19. Rhododendron indet. 




Collections. Mt. Murud: 2100 n^ 
m, Burtt & Martin B. 5363 (E); Mt. 
5250 (E), 1800 m, B. 5255 (E); Mt. V 


1, Ilias S. 26464 (K); Mt. Murud SW of Camp IV: 2100 
Murud above Camp III: 1800 m, Burtt & Martin B. 
Iurud/Belaban River: 1700 m, Ilias S. 26363 (K). 



54.3. Vaccinium 

54.3.1. Vaccinium of. bancanum Miq. 

Collections. Mt. Murud/Belaban River: 1700 m, Ilias S. 26311 (K), 1800 m, S. 26364 (K). 

54.3.2. Vaccinium bigibbum J. J. Sm. 



Collections. Mt. Murud 2^ 


TO summit: 2200 m, ; 


Hias S. 26420 (K); Mt. Murud SW of 


Camp III: 1700 m, Burtt & Marh 


in B. 5326A (E); Mt. 


Murud/Belaban River: 1700 m, Ilias 


S. 26310 (K). 







1 20 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 2 1 

54.3.3. Vaccinium cf. bigibbum J. J. Sm. 

Collections. Mr. Murud Camp IV: 1400 m, Ilias S. 26433 (K); Mr. Murud SW of 
Camp III: 1700 m, Burtt & Martin B. 5323 (E). 

54.3.4. Vaccinium claoxylon J. J. Sm. 

Collections. Mr. Murud 2nd summit: 2300 m, Yii S. 44437 (K, L); Mt. Murud Camp 
IV: 2000 m, Ilias S. 26393 (K, L); Mr. Murud summit: 2400 m, Nooteboom & Chai 1990 
(L), 2400 m, 2005 (L); Mt. Murud summit ridge: 2300 m, Burtt & Martin B. 5491 (E); Mt. 
Murud/Belaban River: 1700 m, Ilias S. 26326 (K). 

54.3.5. Vaccinium dementis Merr. 

Collections. Mt. Murud: 2200 m, Nooteboom & Chai 1975 (L); Mt. Murud 2nd 
summit: 2200 m, Ilias S. 26410 (K, L), 2300 m, Yii S. 44436 (K); Mt. Murud summit ridge: 
2300-2400 m, Beaman 11445 (K, MSC, UNIMAS). 

54.3.6. Vaccinium moultonii Merr. 

Collection. Mt. Murud, path to summit: 1800 m, Nooteboom & Chai 1944 (L). 

54.3.7. Vaccinium pachydermum Stapf [listed by Merrill (1928)]. 

Collections. Mt. Murud: 2100 m, Ilias S. 26457 (K, L), 2300 m, S. 26482 (K, L), 1900-2400 
m, Mjoberg 108 (K, UC); Mt. Murud 2nd summit: 2200 m, Ilias S. 26414 (K, L); Mt. Murud 
between 1st and 2nd summits: 2300 m, Yii S. 44476 (K, L); Mt. Murud summit: 2400 m, 
Nooteboom & Chai 2002A (L); Mt. Murud/Belaban River: 1900 m, Ilias S. 26378 (K, L). 

54.3.8. Vaccinium aff. stapfianum Sleumer 

Collection. Mr. Murud Camp IV: 2000 m, Ilias S. 26398 (K). 

54.3.9. Vaccinium tenerellum Sleumer 



54.3.10. Vaccinium cf. tenerellum Sleumer 
Collection. Mt. Murud Camp IV: 1900 m, Burtt & 1 

54.3.11. Vaccinium sp. 1 [listed by Merrill (1928)]. 
Collection. Mt. Murud: Mjoberg 110 (UC). 

54.3.12. Vaccinium sp. 2 [listed by Merrill (1928)]. 
Collection. Mt. Murud: Mjoberg 109 (UC). 

54.3.13. Vaccinium indet. 



997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 121 

Collections. Mt. Murud Camp III: 1800-2000 m, Burtt & Martin B. 5240 (E); Mr. 
/[URUD Camp IV: 1800 m, Burtt & Martin B. 5349 (E); Mt. Murud SW of Camp IV: 2100 
Q, Burtt & Martin B. 5362 (E). 



55. ESCALLONIACEAE 

55.1. POLYOSMA 

55.1.1. Polyosma mjobergii Merr. [listed by Merrill (1928); probably not distinct 

from P. bracteosa Stapf]. 

Collections. Mt. Murud: 2300 m, Bias S. 26489 (K, SAR), 1900-2400 m, Mjoberg 107 
(K, isotype; UC, holotype of P. mjobergii Merr.); Mt. Murud 2nd summit: 2300 m, Yii S. 
44426 (K, L). 

55.1.2. Polyosma sp. 1 



55.1.3. Polyosma sp. 2 

Collection. Mt. Murud: 2000 m, llias S. 26388 (E, K). 

55.1.4. Polyosma indet. 

Collection. Mr. Murud between 1st and 2nd summffs: 2200 m, Yii S. 44615 (SAR). 

56. EUPHORBIACEAE 

56.1. AUSTROBUXUS 
56.1.1. Austrobuxus nitidus Miq. 

Collection. Mr. Murud/Ba Kelalan: 1700 m, llias S. 26521 (SAR). 

56.2. HOMALANTHUS 
56.2.1. Homalanthus grandifolius Ridl. 

Collection. Mt. Murud, path to summit: 1700 m, Nooteboom & Chai 1912 (L). 

57. FABACEAE 
57.1. Whitfordiodendron 
57.1.1. Whitfordiodendron indet. 



. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

D RIDGE, SE side: 1700-2000 m, Beaman 11491 (K, MSC, UNI- 



MAS). 



58. FAGACEAE 

58.1. LiTHOCARPUS 

58.1.1. Litliocarpus liiteus Soepadmo 

Collections. Mr. Murud/Bfi aban River: 1700 m. Urns S. 26336 (K, SAR), 1700 m, S. 
26325 (K). 

58.2. QuERcus 

58.2.1. Quercus kinabaluensis Soepadmo 

5. 265/5 (K, SAR). 

58.2.2. Quercus valdinervosa Soepadmo 

Collection. Mt. Murhd/Bi:eaban River: 1700 ra, llias S. 26330 (K). 

59. GENTIANACEAE 
59.1. Gentiana 

59.1.1. Gentiana borneensis Hook. f. 

Collections. Mt. Murud: 2200 m, Nooteboom & Chai 2047 (L); Mr. Murud Camp V: 
2100-2300 m, Bum & Martin B. 5457 (E). 

60. GESNERIACEAE 
60.1. Aeschynanthus 

60.1.1. Aeschynanthus magnificus Stapf [listed by Merrill (1928) as Trichospornm 

mjobergii Merr.]. 

Collections. Mt. Murud: 1900 m, Mjoberg 81 (UC), 1900 m, 89 (BM, isotype; UC, 
holotype of Trichospornm mjobergii Merr.); Mt. Murud Camp III: Burtt & Martin B. 5226 
(E); Mt. Murud below Camp III: 1500-1600 m, Burtt & Martin B. 5312 (E). 

60.1.2. Aeschynanthus parvifoiius R. Br. 

Collection. Mt. Murud Camp III: 1800 m, Burtt & Martin B. 5227 (E). 

60.1.3. Aeschynanthus speciosus Hook. 

Collection. Mt. Murud below Camp III: 1500-1600 m, Burtt & Martin B. 5298 (E). 



1997 BEAMAN& ANDERSON: SUMMIT FLORA OF MT. MURUD 123 

60.2. ACALMYLA 

60.2.1. Agalmyla tuberculata Hook. f. 

Collections. Mt. Murud: 2400 m, Nootehoom & Chai 2007 (L, SAR); Mt. Murud N 
side: 2100 m, Bum & Martin B. 5459 (SAR); Mt. Murud above Camp III: 1800 m, Burtt & 
Martin B. 5249 (SAR). 

60.3. Cyrtandra 

60.3.1. Cyrtandra basiflora C. B. Clarke 

Collection. Mt. Murud: 1600 m, Yii S. 44414 (E). 

60.3.2. Cyrtandra aff. cuprea B. L. Burtt 

Collection. Mt. Murud near Camp IJI: 1800 m, Burtt & Martin B. 5270 (E, SAR). 

60.3.3. Cyrtandra dolichopoda B. L. Burtt 

Collection. Mt. Murud bi:low Camp III: 1600 m, Burtt & Martin B. 5307 (E, SAR). 

60.3.4. Cyrtandra horizontalis B. L. Burtt 

Collection. Mt. Murud ridge, SE ridge: 17(){)-2()()() m, Bcaman 11493 (K, MSC, 
UNIMAS). 

60.3.5. Cyrtandra aff. lacerata B. L. Burtt 

Collection. Mt. Murud below Camp III: 1600 ni. Burn & Mariin B. 5309 (E). 

60.3.6. Cyrtandra aff. pedicellata B. L. Burtt 

Collection. Mt. Murud below Camp IV: 1800 m, Burtt & Martin B. 5377 (E). 

60.3.7. Cyrtandra aff. trisepala C. B. Clarke 

Collections. Mt. Murud: 1700 m, Nooteboom & Chai 1914 (L), 1600 m, Yii S. 44416 
(E, L, SAR); Mt. Murud below Camp IV: 1800 m, Burtt & Martin B. 5375 (E, SAR); Mt. 
Murud near Camp III: 1800 m, Burtt & Martin B. 5280 (E). 

60.3.8. Cyrtandra sp. (sect. Decurrentes) 

Collection. Mt. Murud below Camp III: 1500-1600 m, Burtt & Martin s.n. (E). 

60.4. DlDYMOCARPUS 

60.4.1. Didymocarpus simplex Kraenzl. 

Collections. Mt. Murud near Camp III: 1700 m, Burtt & Martin B. 5246 (E); Mt. 
Murud near Camps III and IV: 1800-2000 m, Burtt & Martin B. 5275 (E, SAR); Mt. 
Murud, path to summit: 1800 m, Nooteboom & Chai 1950 (L). 



124 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 

60.4.2. Didymocarpus indet. 



60.5. Lo: 

60.5.1. Loxocarpus indet. 

Collections. Mr. Murud below Camp III: 1500-1600 m, 
SAR); Mr. Murud near Camp III: 1800 m, Bunt & Martin B. 

61. HYDRANGEACEAE 

61.1. DiCHROA 
61.1.1. Dichroa febrifuga Lour. 



62. ILLICIACEAE 
62.1. Illicium 

62.1.1. Illidum cauliflomm Merr. [listed by Merrill (1928)]. 



62.1.2. Illicium tenuifolium A. C. Sm. 

Collection. Mr. Murud, path to summit: 1700 m, Nooteboom & Chai 19. 

63. LAURACEAE 

63.1. LiNDEKA 

63.1.1. Lindera bibracteata (Blume) Boerl. var. rufa (Stapf) Kosterm. 



1.2. Lindera subumbelliflora (Blume) Kosterm. 

Collection. Mt. Murud, path to summit: 1900 m, Noot 

64. LOGANIACEAE 
64.1. Fagraea 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 

64.1.1. Fagraea blumei G. Don 

Collection. Mt. Murud near summit: 2200 m, Nooteboom & Chai 1974 (L). 



64.2. Geniostoma 
64.2.1. Geniostoma rupestre Forst. 

Collections. Mt. Murud: 2200 m, Nooteboom & Chai 2051 (L); Mt. Murud Camp IV: 
2000 m, Ilias S. 26397 (L). 

65. LORANTHACEAE 

65.1. Helixanthera 

65.1.1. Helixanthera cf. cylindrica (Jack) Danser 

Collections. Mt. Murud between 1st and 2nd summits: 2200 m, Yii S. 44620 (SAR); 
Mt. Murud, path to summit: 1900 m, Nooteboom & Chai 1965 (L. SAR). 

65.2. Macrosolen 
65.2.1. Macrosolen cochinchinensis (Lour.) Tieghem 

' B. 5364 (E); Mt. 



65.2.2. Macrosolen flammeus Danser 

Collection. Mt. Murud summit: 2400 m, Nooteboom & Chai 2062 (L). 

66. MELASTOMATACEAE 
66.1. Medinilla 

66.1.1. Medinilla crassifolia (Reinw. ex Blume) Blume 

Collection. Mt. Murud: 1500 m, Yii S. 44633 (AAU, fide G. Clausing, pers. comm.). 

66.1.2. Medinilla homoeandra (Stapf) Nayar [listed by Merrill (1928) as Anplec- 

trum homoeandnim Stapf]. 

Collection. Mt. Murud: 1900-2400 m, Mjoberg 96 (K, UC). 



126 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

66.2. Melastoma 

66.2.1. Melastoma pulcherrimum Koord. 

Collections. Mt. Murud: 2200 m, Nooteboom & Chai 2024 (L); Mt. Murud Camp IV: 
2000 m, llias S. 26399 (K). 

66.3. Phyllagathis 
66.3.1. Phyllagathis brookei Nayar 



66.4. Plethiandra 

66.4.1. Plethiandra hookeri Stapt 

Collections. Mt. Murud 2nd summit: llias S. 26407 (K); Mt. Murud between 1st and 
2nd summits: 2300 m, Yii S. 44483 (K); Mt. Murud summit ridge: 2300-2400 m, Beaman 
11475 (K, MSC, UNIMAS). 

66.5. Sarcopyramis 
66.5.1. Sarcopyramis napalensis Wall. 



66.6. So 
66.6.1. Sonerila crassiuscula Stapf 



Chai 1982 (L); Mt. Mu 



66.6.2. Sonerila macrantha Merr. [listed by Merrill (1928); probably not distinct 

from S. tenuifolia Blume]. 

Collections. Mt. Murud: 2300 m, llias S. 26484 (L), 1900-2400 m, Mjoherg 74 (NY, 
UC), 1900-2400 m, 75 (UC, holotype of S. macrantha Merr.), 1900-2400 m, 76 (UC), 
1900-2400 m, 78 (UC); Mt. Murud 2nd summit: 2300 m, Yii S. 44434 (SAR); Mt. Murud 
N side: 2300 m, Burtt & Martin B. 5449 (E, SAR); Mt. Murud above Camp IV: 2000 m, 
Burn & Martin B. 5345 (E); Mt. Murud between Long Rapata and Camp III: 1500 m, 
Burtt & Martin B. 5223 (E). 

66.6.3. Sonerila nervulosa Ridl. 

Collection. Mt. Murud near Camp 111: 1800 m, Burtt & Mariin B. 5284 (E, SAR). 

66.6.4. Sonerila pulchella Stapf 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 127 

Collection. Mt. Murud ridge, SE side: 1700-2000 m, Beaman 11494 (K). 

66.6.5. Sonerila sp. ? 

Collection. Mt. Murud: Mjdberg 77 [UC; noted by Merrill (1928) to be a single 
imperfect specimen]. 

67. MORACEAE 

67.1. Ficus 
67.1.1a. Ficus deltoidea Jack var. deltoidea 

ATH TO summit: 1700 m, Nooteboom <& Chai 1916 (L); Mt. 



67.1.1b. Ficus deltoidea Jack var. intermedia Corner 

Collections. Mt. Murud: 2200 m, Nooteboom & Chai 2028 (L); Mt. Murud Camp 4: 
2000 m, Ilias S. 26400 (K); Mt. Murud between 1st and 2nd summits: 2200 m, Yii S. 44601 
(K); Mt. Murud between Camp V and summit: Burtt & Martin B. 5492 (E); Mt. Murud 
summit: 2400 m, Nooteboom & Chai 2066 (L); Mt. Murud, path to summit: 1800 m, 
Nooteboom & Chai 1957 (L). 

67.1.1c. Ficus deltoidea var. kinabaluensis Corner 

Collections. Mt. Murud: 2100 m, Bias S. 26450 (K); Mt. Murud 2nd summit: 2200 m, 
Bias S. 26417 (K); Mt. Murud Camp IV: 2000 m, Ilias S. 26392 (K); Mt. Murud between 
Camp V and summit: Burtt & Martin B. 5489 (E); Mt. Murud near Camp V: 2100 m, Ilias 
S. 26514 (K); Mt. Murud summit: 2400 m, Nooteboom & Chai 2001 (L). 

67.1.2. Ficus oleaefolia King var. valida Corner 

Collections. Mt. Murud Camp III: 2000 m, Burtt & Martin B. 5254 (E); Mt. Murud 
between 1st and 2nd summits: 2200 m, Yii S. 44604 (K); Mt. Murud/Belaban River: 1900 
m, Ilias S. 26376 (K). 

67.1.3. Ficus recurva Blume var. ribesioides (Wall, ex Miq.) King 
Collection. Mt. Murud, path to summit: Nooteboom & Chai 1929 (L). 



68. MYRICACEAE 
68.1. Myrica 



128 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 

69. MYRSINACEAE 
69.1. Ardisia 

69.1.1. Ardisia lepidotula Merr. 

Collections. Mt. Murud Camp III: 1600 m, Burn & Martin B. 5261 (E); 
RIDGE SW OF Camp IV: 2100 m, Burtt & Martin B. 5353 (E). 

69.1.2. Ardisia injobergii Merr. [listed by Merrill (1928)]. 

Collection. Mt. Murud: 1900-2400 m, Mjoberg 80 (UC, holotype of A. mjobe 

69.1.3. Ardisia obovatifolia Merr. [listed by Merrill (1928)]. 

Collections. Mt. Muri 
obovatifolia Merr.); Mt. Mui 

69.1.4. Ardisia virens Ku 



69.1.5. Ardisia indet. 

Collections. Mt. Murud: 2200 m, Ilias S. 26496 (L); Mt. Murud summit: 2400 m, 
Nooteboom & Chai 1978 (L); Mt. Murud, path to summit: 1700 m, Nooteboom & Chai 
1908 (L), 1700 m, 1913 (L), 1700 m, 1918 (L), 1700 m, 1919 (L). 

69.2. Embelia 

69.2.1. Embelia minutifolia Stapf 

Collections. Mt. Murud above Camp IV: 2000 m, Burtt & Martin B. 5346 (E); Mt. 
Murud between 1st and 2nd summits: 2200 m, Yii S. 44609 (K. L), 2200 m, 44610 (K, L); 
Mt. Murud summit ridge: 2300-2400 m, Beaman 11456 (K, UNIMAS); Mt. Murud/Bela- 
ban River: 1900 m, Ilias S. 26379 (L). 

69.2.2. Embelia spiraeoides Stapf 

Collections. Mt. Murud, Dapuf 
Mt. Murud, N side near Camp 

69.2.3. Embelia tortuosa Stapf 

Collections. Mt. Murud N side: 2100 m, Burtt & Martin B. 5473 (E); Mt. Murud 
BETWEEN 1st AND 2nd SUMMITS: 2300 m. Yii S. 44481 (K); Mt. Murud summit ridge: 2300- 
2400 m, Beaman 11472 (K, MSC, UNIMAS). 

69.2.4. Embelia indet. 



BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 

69.3. Maesa 



69.3.1. Maesa indica A. DC. 

Collection. Mr. Murud Camp III: 1800 n 



69.4. Myrsine 
69.4.1. Myrsine cruciata (Philipson) Pipoly 

Collections. Mt. Murud above Camp IV: 2100 m. Burn & Martin B. 5357 (E); Mt. 
Murud between 1st and 2nd summits: 2300 m, Yii S. 44468 (K), 2300 m, S. 44473 (K); Mt. 
Murud summit ridge: 2400 m, Burtt & Martin B. 5490 (E). 

70. MYRTACEAE 

70.1. Baeckea 

70.1.1. Baeckea taxifolia Merr. [listed by Merrill (1928)]. 

Collections. Mt. Murud: 2400 m, Mjoberg 84 (UC, syntype of B. taxifolia Merr.), 2400 
m. 111 (K, isosyntype; UC, syntype of B. taxifolia Merr.), 2200-2300 m, Nooteboom & 
Chai 2016 (L); Mt. Murud, Camp V to summit: Burtt & Martin B. 5493 (E). 

70.2. Leptospermuivi 

70.2.1. Leptospermum javanicum Blume [listed by Merrill (1928)]. 

Collections. Mt. Murud: 1500 m, Mjoberg 92 (UC); Mt. Murud Camp IV: 1900 m, 
Burtt & Martin B. 5425 (E); Mt. Murud near summit: 2100 m, Bias S. 26506 (L); Mt. 
Murud summit: 2400 m, Anderson & llias S. 26480 (L), 2400 m, Nooteboom & Chai 1983 
(L); Mt. Murud summit ridge: 2300-2400 m, Beaman 11442 (K, MSC, UNIMAS). 

70.3. Syzygium 

70.3.1. Syzygium aff. ampullarium (Stapf) Merr. & Perry 

Collections. Mr. Murud: 2100 m, llias S. 26458 (K, L); Mt. Murud 2nd summit: 2000 m, 
llias S. 26385 (K, L), 2200 m, S. 26406 (K, L), 2300 m, Yii S. 44428 (K); Mt. Murud above 
Camp IV: 2100 m, Burtt & Martin B. 5360 (E); Mt. Murud between 1st and 2nd summits: 2300 
m, Yii S. 44467 (K), 2300 m, S. 44469 (K); Mt. Murud summit: 2400 m, Anderson & llias S. 
26472 (K, L), 2400 m, Nooteboom & Chai 1984 (L), 2400 m, 1985 (L), 2400 m, 2012 (L); Mt. 
Murud summit ridge: 2300-2400 m, Beaman 11446 (K, MSC, UNIMAS). 

70.3.2. Syzygium castaneum (Merr.) Merr. & Perry 

Collections. Mt. Murud Camp IV: 2000 m, llias S. 26389 (K, L); Mt. Murud summit 
ridge: 2300-2400 m, Beaman 11462 (K, MSC, UNIMAS); Mt. Muruf^/Belaban River: 
1900 m, Was S. 26380 (K, L). 



130 CONTR. UNIVERSITY OF MICHIGAN HERBARIl 

70.3.3. Syzygium indet. 



70.4. Tristaniopsis 
70.4.1. Tristaniopsis elliptica (Stapf) Peter G. Wilson & J. T. Waterh. 



70.4.2. Tristaniopsis indet. 

Collection. Mr. Murdd: 2200 m, Nootehoom & Chai 2041 (L). 

70.5. Xanthomyrtus 

70.5.1. Xanthomyrtus flavida (Stapf) Diels 

Collections. Mr. Murud: 2400 m, Ilias S. 26534 (L); Mt. Murud summit: 2400 i 
Nootehoom & Chai 1997 (L); Mr. Murud, ridge aikwe Camp IV: 1900 m, Burtt A Mart 
B. 5401 (E). 



70.5.2. Xanthomyrtus moultonii (Merr. 

Collection. Mr. Muirud summit: 2400 n 



71. NEPENTHACEAE 
71.1. Nepenthes 

71.1.1. Nepenthes lowii Hook. f. [listed by Merrill (1928)]. 

Collections. Mx. Murud: Mjoberg 115 (UC); Mt. Murud Camp V: 2100-2300 m, B 
& Martin B. 5477 (E); Mt. Murud near Camp III: 1800 m, Burtt & Martin B. 5325 
Mt. Murud summit ridge: 2300-2400 m, Beaman 11476 (K, UNIMAS); Mt. Murud, p 
TO summit: 1900 m, Nootehoom & Chai 1962 (L). 

71.1.2. Nepenthes murudensis Culham ex Jebb & Cheek 

Collections. Mt. Murud Camp IV: 2000 m, Burtt & Martin B. 5427 (E); Mt. Mu 
Camp V: 2100-2300 m, Burtt & Martin B. 5467 (E), 2100-2300 m, B. 5471 (E); Mt. Mu 
BETWEEN 1st AND 2nd SUMMITS: 2200 m, Yii S. 44623 (SAR); Mt. Murud near Cami 
2100 m, Ihas S. 26513 (SAR); Mt. Murud near summit: 2400 m, Nootehoom & Chai 2 
(SAR); Mt. Murud summit ridge: 2300-2400 m, Beaman 11461 (K, MSC. UNIMAS). 

71.1.3. Nepenthes indet. 

Collection. Mt. Murud: Mjoberg 125 [UC; noted hy Merrill (1928) to be on 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT.MURUD 131 

72. OLEACEAE 

72.1. Chionanthus 

72.1.1. Chionanthus enerve (Steenis) Kiew 

Collections. Mt. Murud Camp V: 2100-2300 m, Burtt & Martin B. 5478 (E); Mt. 
MuRUD BETWEEN IsT AND 2nd SUMMITS! 2200 m, Yii S. 44617 (K, L); Mt. Murud summit 
ridge: 2300-2400 m, Beaman 11451 (K, MSC, UNIMAS); Mt. Murud, N of summit: 2300 
m, Nooteboom & Chai 2049 (L). 

72.2. Jasminum 
•philum Kiew 



73. PENTAPHRAGMATACEAE 
73,1. Pentaphragma 

73.1.1. Pentaphragma cf. acuminatum Airy Shaw 

Collections. M 
Murud ridge, SE s 

73.1.2. Pentaphragma aurantiaca Stapf [listed by Merrill (1928) as P. obtusifoliu 

Merr.]. 

Collections. Mt. Murud: 2300-2400 m, Burtt & Martin B. 5451 (E), 2400 m, Ilias 
26537 (L), 1900 m, Mjoberg 112 (UC, holotype of P. ohtusifolium Merr.); Mt. Muri 
summit: 2400 m, Nooteboom & Chai 2008 (L); Mt. Murud summit ridge: 2300-2400 
Beaman 11443 (K, MSC, UNIMAS). 

73.1.3. Pentaphragma longisepalum Kiew 

CoUectio 

between Cam 



74. PIPERACEAE 
74,1. Peperomia 

74.1.1. Peperomia laevifolia (Blume) Miq. 

Collection. Mt. Murud near Camp III: 1800 m, Burtt & Martin B. 5331 (E). 

74.1.2. Peperomia indet. 

Collection. Mt. Murud, path to summit: 1700 m, Nooteboom & Chai 1924 (L). 



132 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME! 

74.2. Piper 

74.2.1. Piper cf. caninum Blume 

Collections. Mt. Murud below Camp III: 1600 m, Burn & Martin B. 5310 (E); M 
MuRUD RIDGE, SE SIDE: 1700-2000 m, Beaman 11492 (K, UNIMAS). 

75. POLYGALACEAE 

75.1. POLYCiALA 

75.1.1. Polygala oreotrephes B. L. Burtt [listed by Merrill (1928) as Polygala sp.] 
Collections. 



3. RHAMNACEAE 
76.1. Rhamnus 



1.1. Rhamnus borneensis Steenis 

Collections. Mt. Murud: 2300 m, Ilias S. 26485 (K); Mt. Murud 2nd summit: 2200 r 
s S. 26419 (K); Mt. Murud N side: 2100 m, Burtt & Martin B. 5461 (E); Mt. Muri. 
' OF Camp IV: 2100 m, Burtt & Martin B. 5361 (E): Mt. Murud summit: 2400 r 
i&Chail996{Y.). 



11. ROSACEAE 

77.1. Prunus 

1.1. Prunus arborea (Blume) Kalkman var. densa (King) Kalkman 

Collections. Mt. Murud summit: 2400 m, Nooteboom & Chai 1991 (L); Mt. Mu 
IMIT ridge: 2300-2400 m, Beaman 11452 (K, MSC, UNIMAS); Mt. Murud, patf 
imit: 1900 m, Nooteboom & Chai 1951 (L). 



77.2.1. Rubus alpestris Blume 

Collection. Mt. Murud summit: 2400 m. Nooteboom 

78. RUBIACEAE 

78.1. ACRANTHERA 

78.1.1. Acranthera aff. atropeila Stapf 



,SEs 



1997 beaman & anderson: summit flora of mt. murud 

78.2. Argostemma 

78.2.1. Argostemma borragineum Blume ex DC. 

Collection. Mt. Murud: 1600 m, Yii S. 44410 (K). 

78.2.2. Argostemma hameliifolium Wernham 

Collection. Mt. Murud: 1600 m, Yii S. 44409 (K). 



Collections. Mt. Murud: 1900-2400 m, Mjoberg 82 (K, isotype; UC, holotype of A. 
murudensis Merr.), 1900-2400 m, 88 (UC); Mt. Murud near Camp III: 1800 m, Burtt & 
Martin B. 5326 (E); Mt. Murud, path to summit: 1700 m, Nooteboom & Chai 1935 (L). 



78.3. Cephaelis 

78.3.1. Cephaelis stipulacea Blume [this species probably belongs in the gen 
Chassalia]. 

Collection. Mt. Murud near Camp III: 1800 m, Burtt & Martin B. 5285 (E). 



78.4. DiSCOSPERMUM 

78.4.1. Discospermum abnorme (Korth.) S. J. Ali & Robbr. 

Collection. Mt. Murud below Camp IV: 1800 m, Burtt & Martin B. 5373 (E). 

78.5. Hedyotis 

78.5.1. Hedyotis pulchella Stapf 

Collections. Mt. Murud below Camp III: 1600 m, Burtt & Martin B. 5291 (E); ^ 
Murud, Dapur River headwaters: 1900 m, Ilias S. 26517 (K). 

78.5.2. Hedyotis sp 1 

Collections. Mt. Murud: Burtt & Martin B. 5448 (E), 2100 m, Hias S. 26465 (K, L). 

78.5.3. Hedyotis indet. 

Collection. Mt. Murud: 2200 m, Nooteboom & Chai 2021 (L). 



78.6. IxoRA 
78.6.1. Ixora sessililimba Merr. [listed by Merrill (1928)]. 



VERSITY OF MICHIGAN HERBARIUM 
78.7. LUCINAEA 



78.7.1. Lucinaea pentacme Stapf 



78.8. Opiiiorrhiza 
78.8.1. Ophiorrhiza aft. fibrillosa Ridl 



78.9. Psvchotria 
78.9.1. Psychotria densifolia Stapt 



Collection. Mt. Murud, path to summit: 1700 m, Nootchoom & Chai 1925 (L). 

78.10. RuBiA 
78.10.1. RubiacordifoliaL 

Collection. Mr. Mi,ri;d, path to summit: 1700 m, Nootehoom & Chai 1926 (L). 

78.11. Streblosa 
78.11.1. Streblosa urticina Stapf 

Collection. Mt. Murud near Camp III: 1«()0 m, Burtt & Martin B. 5335 (E). 

78.12. TiMONius 
78.12.1. Timonius esherianus W. W Sm. 



79. RUTACEAE 
79.1. EuoDiA 



79.1.1. Eiiodia indet. 

Collection. Mt. Muirud: 2200-23i 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 

80. SYMPLOCACEAE 
80.1. Symplocos 
80.1.1. Symplocos adenophylla Wall, ex G. Don var. adenophylla 



80.1.2. Symplocos 

Collection. Mr. Murud/Belaban River headwaters: 1700 m, Ilias S. 26357 (L, SAR). 

80.1.3. Symplocos henschelii (Moritzi) Benth. ex C. B. Clarke var. henschelii [listed 

by Merrill (1928) as S. dolichantha Merr.] 



80.1.4a. Symplocos laeteviridis Stapf var. mjobergii (Merr.) Noot. [listed by Mer- 
rill (1928) as S. mjobergii Merr.]. 

Collections. Mt. Murud: 1900-2400 m, Mjoberg 97 (K, isotype; UC, holotype of S. 
mjobergii Merr.); Mt. Murud, path to summit: 1500 m, Nooteboom & Chai 1909 (K, L). 

80.1.4b. Symplocos laeteviridis Stapf var. pauciflora Noot. 

Collections. Mt. Murud: 2000 m, Ilias S. 26391 (K, L), 1700-2000 m, S. 26392 (?); Mt. 
Murud Camp IV: 1900 m, Burn & Martin B. 5389 (E); Mt. Murud/Belaban River: 1700 
m, Ilias S. 26334 (K, L). . 

80.1.5. Symplocos pendula Wight var. hirtistylis (Clarke) Noot. 



80.1.6. Symplocos tricoccata Noot. 

Collection. Mt. Murud/Belaban Rivhr headwaters: 1700 m, Ilws S. 26305 (K, L, SAR). 



81. THEACEAE 
81.1. Adinandra 

81.1.1. Adinandra clemensiae Kobuski 

Collections. Mt. Murud: 1600 
summits: 2200 m, Yii S. 44611 (K). 

81.1.2. Adinandra cf. dumosa Jack 

Collection. Mt. Murud summit ridge: 2300-2400 m, Beaman 11477 (K, MSC, UNIMAS). 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

81.2. SCHIMA 

/allichii (DC.) Choisy subsp. brevifolia (Hook, f.) Bloemb. 

• (L); Mt. Murud N side: 

81.3. Ternstkoemia 

81.3.1. Ternstroemia beccarii Slapf 

Collections. Mt. Murud: 2300 m, Ilias S. 26490 (K), 2200 m, Nooteboom & Chai 2031 
(L); Mt. Murud 2nd summit: 2300 m, Yii S. 44439 (K), 2300 m, S. 44448 (K); Mt. Murud 
ABOVE Camp IV: 2000 m, Bunt & Martin B. 5438 (E); Mt. Murud between 1st and 2nd 
summits: 2300 m, Yii S. 44463 (L); Mt. Murud summit ridge: 2300-2400 m, Seaman 11463 
(K, MSC, UNIMAS); Mt. Murud, path to summit: 1800 m, Nooteboom & Chai 1948 (L). 

81.3.2. Ternstroemia denticulata (Pierre) Ridl. 

Collections. Mt. Murud: 2200 m, Ilias S. 26402 (L), 2100 m, S. 26454 (L); Mt. Murud 
between 1st and 2nd summits: 2300 m, Yii S. 44484 (K), 2300 m, S. 44485 (K), 2200 m, S. 
44605 (K); Mt. Murud summit: 2400 m, Nooteboom & Chai 1980 (L); Mt. Mi;rud. path to 
summit: 1800 m, Nooteboom & Chai 1956 (L). 

81.3.3. Ternstroemia lowii Stapf 



81.3.4. Ternstroemia indet. 

82. THYMELIACEAE 

82.1. WlKSTKOEMIA 

82.1.1. Wikstroemia brachyantha Merr. 



82.1.2. Wikstroemia cf. indica (L.) C. A. Mey. 

Collections. Mt. Murud: 2100 m, Was S. 26459 (K, L 
m, Nooteboom & Chai 2038 (L). 2300 m, 2040 (L). 

83. URTICACEAE 
83.1. Elatostema 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 

83.1.1. Elatostema acuminatum (Poir.) Brongn. 

Collection. Mt. Murud below Camp III: 1600 m, Burft & Martin B. 531 

83.1.2. Elatostema cf. lineare Stapf 
Collection. Mt. Murud, Camp II I/Long Rapai 

83.1.3. Elatostema penibukanense Gibbs 



Bum & Martin B. 5224 (E). 



Colle. 



5282 (E); 



■IV:: 



83.1.4. Elatostema sp. 



83.1.5. Elatostema sp. 2 



' IV: 1900 m, Burtt & Martin B. 5387 (E). 



83.1.6. Elatostema indet. 
Collection. Mt. Murud 



83.2.1. Pilea sp. 

Collection. M 



. Mur 



/ Camp IV: 1800 m, Burtt & Martin B. 5367 (E). 



84. VERBENACEAE 
84.1. Clerodendrum 
1.1. Clerodendrum kinabaluense Stapf 
Collection. Mt. Murud Camp IV: 1900 m, Burtt & Martin B. 5413 (E). 



85. WINTERACEAE 
85.1. Drimys 



j.1.1. Drimys piperita Hook. f. [listed by Merrill (1928)]. 



Collections. Mt. Murud: Mjoberg 101 (UC), 102 (UC), 2200 m, Nooteboom & Chai2018 
(L); Mt. Murud Camp IV: 2000 m, Burtt & Martin B. 5348 (E); Mr. Murud summit: 2400 m, 
Nooteboom & Chai 2008A (L); Mt. Murud summit ridge: 2300-2400 m, Beaman 11458 (K); 
Mt. Murud, path to summit: 1700 m, Nooteboom & Chai 1915 (L), 1800 m, 1949 (L). 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



INDEX TO NUMBERED COLLECTIONS CITED 

[Collection-number prefixes have been omitted.] 



lerson & Ilias 26471 (21.2.2); 26472 (70.3.1 


.); 26477 i 


man 11440 (21.2.2); 11441 (54.2.18); 11442 


: (70.2.1); 


11446 (70..1. 1); 11447 (54.2.5); 11448 (28,' 


9.2a): 114 


11452 (77. 1.1 a); 11453 (81. 3 J); 11454 (37 


M.l);114 






(19.1.1); 11465 (25.2.1): 11466 (20.1.1): 1 


1467 (.54.: 


(41.1.1); 11471 (23.1.1); 11472 (69.2.3): i: 




(71.1.1); 11477 (81.1.2); 11478 (,S4.2.4): 11 


1479(41.: 


(28.8.4); 11483 (40.1.2); 11484 (52.1 1). 1] 


1485(28.1 



(66.6.3); 5285 (78.3.1); 5287 (28.1.1); 5288 (54.1.1 
5297 (66,5.1); 5298 (60.1.3); 5299 (28.9.4); .5300 (; 
(60.3.3); 5.3(J8 (61.1.1); 5309 (60.3,5); 5310 (74,2.1 
5314 (34.3,2); 5316 (73,1,1); 5317 (54.2.12): 5319 



4 (83, 


1.2);, 


5225 (73.1.3); 


4U98 


133 C 


^8,1,2); 5234 


60.2,1; 


); 5250 (54,2.19); 5: 








l'8)-^5 


,1,1) 


54,1,8); 5269 
5276(60.5.1); 


iSl. 


(34,2 


,2); 5284 



'(66.6,2); 54.50 (54.2.1^ 


0:5451(7.3.1,2); .54.52 (54,1,10); 5 


5457 (59,1.1); .5459 (6 


0,2, 1);.S460 (28,7.1); .5461 (76, 1,1 


3A (28,2,1); 5464 (28, 


18,2);5465A (9.1.2a): 5465B (9,1, 


8 (40.1.2); 5469 (81.2,1 


.a); .5471 (71.1,2); 5472 (7.1,3); 54 



5474 (13.1,3); 5475 (25.2,1); 5477 (71.1.1); 5478 (72,1,1); 5480 (28.8.1); 5481 (53.1,1); 5482 
(25.3.1); 5483 (15.1.1a); 5484 (33.1.1a); 5487 (49.1.1); 5488 (16.1.2); 5489 (67,1.1c); 5490 (69,4 
5491 (54.3.4); 5492 (67.1.1b); 5493 (70,1.1); 5494 (24.1,1); 5495 (31.2,1); 5496 (2.1.1); 5549 



80,1,2); 26363 (54.2,19); 26364 (54,3,1); 26365 (50.1.1a); 26373 (54.1 
; 26375 (55.1.2); 26376 (67.1.2a); 26378 (54.3.7); 26379 (69.2.1); 263 
I); 26382 (68.1.1); 26384 (54.1.3); 26385 (70.3,1); 26386 (41.2.1); 263 



.3); 26389 


(70.3 


.2);2639 


eaoil)! 


2640- 


7 (66.4.1) 


7 (67.1.1c; 


); 264: 


19 (76.1.1 


7 (54.1.6); 




3 (67.1.1c 


6(28:M^r 




B (70.3.1) 



BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 139 

2.1); 26400 (67.1.1b); 26402 (81.3.2); 26403 (54.2.1a); ' 

10.2); 26410 (54.3.5); 26414 (54.3.7); 26416 (54.1.10); 

t.3.2); 26421 (61.1.1); 26433 (54.3.3); 26434 (80.1.1a); 

l.l.l)- 26453 (40.1.1); 26454 (81.3.2); 26455 (28.9.6); 

1.2); 26462 (54.2.16); 26464 (54.2.19); 26465 (78.5.2); 

1.3.7); 26483 (70.4.1); 26484 (66.6.2); 26485 (76.1.1); 
26486 (41.1.1); 26487 (54.1.10); 26489 (55.1.1); 26490 (81.3.1); 26492 (54.2.11); 26493 (54.2.5); 
26495 (52.1.1); 26496 (69.1.5); 26501 (69.2.4); 26505 (21.2.2); 26506 (70.2.1); 26507 (70.4.1); 26508 
(58.2.1); 26510 (52.1.1); 26513 (71.1.2); 26514 (67.1.1c); 26515 (58.2.1); 26516 (53.1.1); 26517 
(78.5.1); 26518 (66.3.1); 26519 (28.6.6); 26521 (56.1.1); 26532 (69.2.2); 26534 (70.5.1); 26537 
(73.1.2); 26538 (28.10.2); 26539 (28.10.1); 26555 (54.2.8); 26556 (54.2.10); 26557 (41.2.3). 

Mjoberg 7 (10.1.1); 22 (13.1.2); 49 (28.9.10); 52 (28.9.6); 64 (28.9.5); 66 (28.9.7); 67 (11.1.1); 68 

(14.1.2); 70 (10.1.1); 71 (14.1.1); 73 (1.1.1, 1.1.3); 74 (66.6.2); 75 (66.6.2); 76 (66.6.2); 77 (66.6.5); 
78 (66.6.2); 79 (69.1.3); 80 (69.1.2); 81 (60.1.1); 82 (78.2.3); 83 (52.1.2); 84 (70.1.1); 88 (78.2.3); 89 
(60.1.1); 91 (75.1.1); 92 (70.2.1); 93 (40.1.2); 94 (64.1.2); 95 (5.1.1); 96 (66.1.2); 97 (80.1.4a); 98 
(54.2.1a); 99 (21.1.1); 100 (21.2.1); 101 (85.1.1); 102 (85.1.1); 103 (54.2.11); 104 (54.2.11); 105 
(54.2.5); 106 (54.2.4); 107 (55.1.1); 108 (6.1.1, 54.3.7); 109 (54.3.12); 110 (54.3.11); 111 (70.1.1); 
112 (73.1.2); 113 (25.1.1); 114 (62.1.1); 115 (71.1.1); 116 (78.6.1); 117 (41.2.1); 118 (34.5.1); 119 
(45.1.1); 120 (80.1.3a); 121 (34.5.1); 122 (34.3.1); 123 (30.1.1); 124 (39.2.1); 125 (71.1.3); 126 
(23.1.2); 128 (23.1.3); 182 (12.1.1); 204 (3.1.1, 60.4.2); 214 (4.1.1); 256 (10.1.1). 

Nooteboom & Chai 1906 (32.1.2); 1908 (69.1.5); 1909 (80.1.4a); 1910 (28.4.3); 1912 (56.2.1); 1913 
(69.1.5); 1914 (60.3.7); 1915 (85.1.1); 1916 (67.1.1a); 1918 (69.1.5); 1919 (69.1.5); 1920 (45.1.2); 
1922 (28.1.5); 1923 (62.1.2); 1924 (74.1.2); 1925 (78.9.2); 1926 (78.10.1); 1927 (28.12.4); 1928 
(78.12.1); 1929 (67.1.3a); 1930 (48.1.1); 1931 (70.3.3); 1932 (80.1.3a); 1933 (35.1.1); 1934 (1.1.3); 
1935 (78.2.3); 1936 (32.1.3); 1937 (22.1.1); 1938 (34.5.2); 1939 (29.1.1); 1940 (54.2.4); 1941 
(70.3.3); 1943 (47.1.1); 1944 (54.3.6); 1945 (13.1.1a); 1946 (28.18.3); 1948 (81.3.1); 1949 (85.1.1); 
1950 (60.4.1); 1951 (77.1.1a); 1952 (63.1.1a); 1954 (28.9.7); 1955 (54.2.8); 1956 (81.3.2); 1957 
(67.1.1b); 1958 (41.2.3); 1959 (34.1.2); 1960 (8.1.1); 1961 (41.2.1); 1962 (71.1.1); 1963 (63.1.2); 
1964 (48.1.1); 1965 (65.1.1); 1966 (70.4.1); 1967 (54.2.10); 1968 (44.1.2); 1969 (28.4.3, 28.4.1); 1970 
(54.2.11); 1971 (75.1.1); 1972 (54.2.10); 1973 (54.2.3); 1974 (64.1.1); 1975 (54.3.5); 1976 (52.1.2); 
1977 (65.2.1); 1978 (69.1.5); 1979 (78.9.1); 1980 (81.3.2); 1981 (25.2.1); 1982 (66.6.1); 1983 
(70.2.1); 1984 (70.3.1); 1985 (70.3.1); 1986 (52.1.3a); 1987 (32.1.1); 1988 (52.1.4); 1989 (54.2.5); 
1990 (54.3.4); 1991 (77.1.1a); 1992 (54.1.8); 1993 (77.2.1); 1994 (68.1.1); 1995 (28.9.9); 1995A 
(28.9.1); 1996 (76.1.1); 1997 (70.5.1); 1998 (23.1.1); 1999 (30.1.2); 2000 (70.5.2); 2001 (67.1.1c); 
2002A (54.3.7); 2002B (54.1.3); 2003 (52.1.1); 2005 (54.3.4); 2006 (54.2.9); 2007 (60.2.1); 2008 
(73.1.2); 2008A (85.1.1); 2009 (5.1.2); 2010 (48.1.1); 2011 (64.1.2); 2012 (70.3.1); 2013 (19.1.1); 
2014 (11.1.1); 2015 (9.1.1); 2016 (70.1.1); 2017 (43.1.1a); 2018 (85.1.1); 2019 (28.6.2); 2020 
(40.1.1); 2021 (78.5.3); 2022 (53.1.1); 2023 (25.3.1); 2024 (66.2.1); 2025 (26.1.1); 2026 (30.1.1); 
2028 (67.1.1b); 2028a (17.1.1); 2029 (31.1.1); 2030 (25.1.3); 2031 (81.3.1); 2032 (28.8.3); 2033 
(28.8.5); 2034 (44.1.1); 2035 (71.1.2); 2036 (9.1.1); 2037 (40.1.2); 2038 (82.1.2); 2039 (7.1.3); 2040 
(82.1.2); 2041 (70.4.2); 2042 (40.1.1); 2043 (47.1.2); 2044 (28.11.1); 2045 (81.2.1a); 2046 (55.1.2); 
2047 (59.1.1); 2048 (41.1.1); 2049 (72.1.1); 2050 (79.1.1); 2051 (64.2.1); 2052 (41.2.2); 2053 
(39.1.1); 2054 (28.2.1); 2055 (28.6.8); 2056 (54.1.11); 2058 (80.1.5a); 2059 (83.1.6); 2061 (46.1.1); 
2062 (65.2.2); 2063 (54.1.3); 2064 (15.1.1a); 2065 (28.12.2); 2066 (67.1.1b); 2067 (69.2.4); 2068 
(52.1.2); 2069 (51.1.2). 

Rena, Rantai et al. 60335 (28.13.1a); 60336 (28.12.4); 60394 (28.14.1). 

Sarawak Museum Native Col 2937 (18.1.1). 

Yii 44401 (28.1.5); 44402 (28.9.4); 44403 (51.1.1a); 44405 (36.1.1); 44408 (47.2.1); 44409 (78.2.2); 44410 
(78.2.1); 4441 1 (66.5.1); 44412 (78.8.1); 44414 (60.3.1); 44416 (60.3.7); 44420 (82.1.1); 44421 
(28.1.3); 44422 (61.1.1); 44425 (40.1.2); 44426 (55.1.1); 44427 (52.1.1); 44428 (70.3.1); 44430 
(28.9.3); 44431 (28.12.2); 44432 (28.9.7); 44433 (54.1.10); 44434 (66.6.2); 44436 (54.3.5); 44437 
(54.3.4); 44439 (81.3.1); 44442 (23.2.1); 44447 (54.1.7); 44448 (81.3.1); 44460 (54.2.1a); 44461 
(19.1.1); 44463 (81.3.1); 44467 (70.3.1); 44468 (69.4.1); 44469 (70.3.1); 44471 (52.1.2); 44473 
(69.4.1); 44474 (64.1.2); 44476 (54.3.7); 44478 (54.1.3); 44481 (69.2.3); 44482 (28.2.1); 44483 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 



(40.1.1); 44490 (54.1.8 


); 44491 (28.12 


(54.2.10); 44604 (67.1 


2a); 44605 (81 


(81.1.1); 44612 (37.1.1 


); 44615 (55.1 


(41.2.3); 44620 (65.1.1 


); 44621 (46.1 



ACKNOWLEDGMENTS 

When one considers that Mt. Murud is the highest mountain in Sarawak, it is remarkable that 
we have been able to discuss personally its flora with most of the major botanical collectors who have 
been to its summit. This account benefits enormously from the specimen data, information, and 
advice provided by B. L. Burtt, H. P. Nooteboom, and Yii Puan Ching. The other botanist on our trip 
was Harry Lohok, who is in charge of the Poring Orchid Garden of Sabah Parks. Our head guide was 
Walter Parran, and the other porter-guides were Marran Pu'un, Shimpson Kores, and Edwin Radu, 
all from Pa Lungan. They got us to Mt. Murud and back without mishap. Robert Yong of Kuching 
contributed substantially in the organization and equipping of our trip. The organizers of the Bario 
Expedition, Prof. Dr. Laily bin Din, Prof. Dr. Ghazally Ismail, and Dr. Petrus Bulan (all of UNI- 
MAS) provided advice and support essential for the trip. The friendly people of Bario, especially the 
Penghulu (head man of the Bario area), and the Ketua Kampung and Assistant Ketua Kampung 
(head and assistant head) of Pa Lungan, were extremely hospitable and helpful. The involvement of 
these persons, together with the many other UNIMAS participants in the Bario Expedition, contributed 
to the success of our trip. Help in identification of the collections was received from staff of the 
Herbarium of the Sarawak Forest Department, Hj. Othman Ismawi and Yii Puan Ching, and a 
number of persons from the Herbarium, Royal Botanic Gardens, Kew, including S. M. Andrews, M. 
J. E. Coode, E. J. Cowley, J. Dransfield, P. J. Edwards, L. L. Forman, M. J. Sands, and J. J. Wood. 
Others who helped with identifications include G. Argent (E), N. Cellinese (FLAS), G. Clausing 
(MJG), J. J. Pipoly (BRIT), R. Kiew (UPM), M. G. Price of Jackson, Michigan, and L. G. Saw 
(KEP). We have also been helped by B. A. Barlow (CANB), C. C. Berg (BG), S. Kofman (L), C. 
Pendry (E), and J. L. Strother (UC). The computer programming was done by R. S. Beaman (FLAS), 
and data for the orchids in K and SAR were recorded by T. E. Beaman (MSC). Research support for 
this project was provided by a short-term research grant from Universiti Malaysia Sarawak and 
National Science Foundation Grant DEB-9400888 to Michigan State University. We greatly appreciate 
the herbarium resources made available to us by the Sarawak Forest Department Herbarium, Kuching 
(SAR), the Royal Botanic Gardens, Kew (K), the Rijksherbarium, Leiden (L), and the Royal Botanic 
Garden, Edinburgh (E). 



LITERATURE CITED 

Anderson, J. A. R., and P. P. K. Chai. 1982. A preUminary list of angiosperms and gymnosperms 
recorded within Gunung Mulu National Park. In Gunung Mulu National Park: A Management 
and Development Plan, by J. A. R. Anderson, A. C. Jermy, and The Earl of Cranbrook, Appen- 
dix III, 233-271. Royal Geographical Society, London. 

Anonymous. 1952. Preface. Sarawak Papers: Scientific Results of the Oxford University Expedition to 
Sarawak (Borneo) in 1932, i-viii. Published for the Oxford University Exploration Club by the 
Oxford University Press, London. [A compiled reprint of 48 papers reporting on the results of 
this expedition.] 

Beaman, J. H. 1997. Preliminary enumeration of the summit flora. Mount Murud, Kelabit Highlai 
Sarawak. In A scientific journey through Borneo: Bario-The Kelabit Highlands of Sarawak, ed. 
Ghazally Ismail and Laily Din, 51-81. Kuala Lumpur: Pelanduk Publications. 

Beaman, J. H., and R. S. Beaman. 1993. The gymnosperms of Mount Kinabalu. Contr. Univ. Michi- 
gan Herb. 19: 307-340. 

Beaman, J. H., and J. C. Regalado. 1989. Development and management of a microcomputer spi 
men-oriented database for the flora of Mount Kinabalu. Taxon 38: 27-42. 

Bonaparte, R. B. 1923a. Recoltes du Dr. E. Mjoberg, Herbier du Prince Bonaparte. Notes Pteri 
logiques 14: 477-488. 

.1923b. Plusieurs regions, plusieurs collecteurs, Herbier du Prince Bonaparte. Notes Pteri 

logiques 14: 447-475. 



1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 141 

Briggs, J. 1988. Mountains of Malaysia — a Practical Guide and Manual. Longman Malaysia Sdn. Bhd. 
Dransfield, J. 1992. The Rattans of Sarawak. Kew: Royal Botanic Gardens; Kuching, Sarawak: Sara- 
wak Forest Department. 
Harrisson, T. 1933. The Oxford University Expedition to Sarawak, 1932. Geographical J. 82: 385-410. 

. 1959. World Within: A Borneo Story. London: Cresset Press. 

Jermy, A. C. 1984. Introduction. In Studies on the Flora of Gunung Mulu National Park, Sarawak, ed. 

A. C. Jermy, v-xv. Kuching, Sarawak: Sarawak Forest Department. 
Martin, P. J. 1977. The Altitudinal Zonation of Forests Along the West Ridge of Gunong Mulu. 

Kuching, Sarawak: Sarawak Forest Department. 
Merrill, E. D. 1928. A collection of plants from Sarawak. Sarawak Museum J. 3: 513-557. 
Mjoberg, E. 1925. An expedition to the Kalabit country and Mt. Murud, Sarawak. Geographical Rev. 

15: 411-427. 
. 1930. Forest Life and Adventures in the Malay Archipelago, translated from the Swedish by 

A. Barwell. London: George Allen & Unwin Ltd. 
Moulton, J. C. 1912. An expedition to Mount Batu Lawi. J. Straits Branch Roy. Asiat. Soc. 63: 1-104. 

. 1915. Thirteenth Report on the Sarawak Museum, 1914. 

Parris, B. S. 1997. The ecology and phytogeography of Mount Kinabalu pteridophytes. Sandakania 9: 

89-102. 
Parris, B. S., A. C. Jermy, J. M. Camus, and A. M. Paul. 1984. The pteridophyta of Gunung Mulu 

National Park. In Studies on the Flora of Gunung Mulu National Park, Sarawak, ed. A. C. 

Jermy, 145-233. Kuching, Sarawak: Sarawak Forest Department. 
Parris, B. S., R. S. Beaman, and J. H. Beaman. 1992. The Plants of Mount Kinabalu. 1. Ferns and Fern 

Allies. Kew: Royal Botanic Gardens. 
Price, M. G. 1983. Several unusual Malesian Diplazia. Gard. Bull. Singapore 36: 25-29. 
Yin Ee Heng. 1992. Geological Map of Sarawak, 2d ed., updated by V. Hon and Lam S. K. Kuala 

Lumpur: Director-General Geological Survey of Malaysia. 
Wood, J. J. 1984. A preliminary annotated checkHst of the orchids of Gunung Mulu National Park 

with a key to genera. Studies on the Flora of Gunung Mulu National Park, Sarawak, ed. A. C. 

Jermy, 1-39. Kuching, Sarawak: Sarawak Forest Department. 
Wood, J. J., R. S. Beaman, and J. H. Beaman. 1993. The Plants of Mount Kinabalu. 2. Orchids. Kew: 

Royal Botanic Gardens. 



ADDENDUM 

)er was in page proof we learned from J. J. Wood a 
1 Mt. Murud by Burtt and Martin in 1967. These are 



ORCHIDACEAE 

KUHLHASSELTIA 



ARISTOLOCHIA DAVILAE (ARISTOLOCHIACEAE), 
A NEW SPECIES FROM NAYARIT, MEXICO 

Ismael Calzada J. 
Gabriel Flores F. 
Oswaldo Tellez V. 
Herbario Nacional 
Departamento de Botanica 
Institute de Biologia UNAM 

Apartado Postal 70-367 
04510 Mexico, D.F., Mexico 

This contribution complements previously published papers dealing with the 
Flora of Nayarit Project (Mendez & Tellez 1995; Tellez 1995), carried out for 
several years by the Instituto de Biologia UNAM. 

Aristolochia davilae Calzada, Flores & O. Tellez, sp. nov. — Type: Mexico. Naya- 
rit: Mpio. Nayar, cafiada en el ejido Carretones de Cerrito, ca. 4 km al E 
de la cortina del Proyecto Hidroelectrico Aguamilpa, 21°50'30"N, 
104°45'50"W, bosque tropical caducifoHo, suelo negro, rocoso, 180 m, 20 
Aug 1993, Calzada, Flores & Solis 18596 (holotype: MEXU!; isotype, 
MICH!). Fig. 1. 

Herbae perennes 40.0-60.0 cm altae, erectae vel scandentes. Laminae (2.6-) 
3.7-7.7 cm longa, (0.3-) 0.5-2.6 cm lata, linearia vel lineari-lanceolata vel hastata, 
basi auriculata. Flores ca. 5.0 cm longi, flavovirides, solitarii, axillares; stamina 5, 
columna ca. 1.5 mm longa. Capsulae ca. 1.5 cm longae, ca. 1.2 cm latae, sphaeri- 
cae, laevigatae, cinereae; semina ca. 4.0 mm longa, ca. 3.0-4.0 mm lata, triangularia. 

Perennial herbs 40.0-60.0 cm tall, erect to scandent; stems puberulent. Main 
root 20.0-25.0 cm long, the bark divided in small quadrangular plates, brownish. 
Laminas (2.6-) 3.7-7.7 cm long, (0.3-) 0.5-2.6 cm wide, linear-lanceolate to hastate 
but the base auriculate, linear in the distal parts of branches, apex acuminate to 
acute, adaxially densely tuberculate, mainly on the nerves, abaxially glabrous to 
spreading-hispid and tuberculate, margin tuberculate but appearing ciliate; primary 
nerves 3, secondary venation reticulate, inconspicuous; petiole 0.3-0.5 cm long, 
puberulent. Flowers solitary, axillary, yellowish green, the throat red, puberulent; 
calyx ca. 4.0 mm long, straight, densely puberulent; corolla ca. 5.0 cm long, the 
limb ca. 1.4 cm wide; stamens 5, joined in a column ca. 1.5 mm long, 5-lobed at the 
apex. Capsules 1.0-1.5 cm long, ca. 1.2 cm in diameter, spherical to subspherical, 
smooth, puberulent, greenish, with three dehiscence lines; immature seeds ca. 4.0 
mm long, 3.0-4.0 mm wide, triangular, the encircling wing not well developed. 

Aristolochia davilae superficially resembles, especially in leaf shape, several 
species in subsect. Pentandrae (e.g., A. acanthophylla, A. palmeri, A. porphyro- 
phylla), but it appears most closely related to small group of endemic species from 
Western Mexico, particularly to A. bracteosa, A. oaxacana, and A. socorroensis 
(table 1). Pfeiffer (1970) in his key divides subsect. Pentandrae into two groups, 
but without formally recognizing them. One group is characterized by straight, 



. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 




CALZADA, FLORES & TELLEZ: ARISTOLOCHIA DAVILAE 



erect to subarcuate flowers, with an angle of less than 90°, and includes the three 
species considered closest to A. davilae. It is further divided into two parts; A. 
davilae is allied with the species that have a wide, 6-nerved floral limb with a 
rounded to acuminate apex. Species listed in the other part have a long-acuminate 
floral limb. Pfeiffer's second group comprises species that have geniculate flowers, 
with an angle of at least of 90° and the limb reflexed to the utricle. 

Aristolochia davilae inhabits tropical deciduous forests at ca. 180 m, associated 
with Acacia tenuifolia, Karwinskia latifolia, Diphysa suberosa, Chamaesyce umbel- 
lata, Hilaria ciliata, Jatropha cordata, etc. Flowering and fruiting occur during the 
rainy season in August. 

This species was collected on the islands created by the construction of the 
dam in the Aguamilpa Hydroelectric Project, as part of a flora and fauna rescue 
program carried out by the Instituto de Biologia UNAM and the Comision Federal 
de Electricidad (CFE) during 1993. We consider this species a restricted endemic 
to this region of Nayarit, and at the same time as an endangered species, because 
some of these islands have already disappeared below water level. 

The name for this new species honors Dr. Patricia Davila Aranda, researcher 
associated with the National Herbarium (MEXU), in recognition of her leader- 
ship in promoting botanical research in Mexico. 



146 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

ACKNOWLEDGMENTS 

Latin diagnosis. We are grateful for the financial support provided by CONACyT (grants PCCNC- 
NA-()31529, P218CCOC880112, P218CCOC827792, D112-903588, 0455-N9108; 1988-1993), and to 
the National Geographic Society (grant 4349-89) to the Flora de Nayarit Project. We also thank Biol. 
Eduardo Avalos Guzman, head of the Ecology and Environment Group of CFE in the City of Tepic, 
Nayarit, for logistic support during the rescue program. Elvia Esparza drew for the excellent illustration. 

LITERATURE CITED 



Mendez L., I., and O. Tellez V. 1 


995. Una nu 


eva especie de R 


usselia (Scrophula 


iriaceae) del estado de 


Nayarit, Mexico. Anales Ins 


;t. Biol. Uni' 


J. Nac. Mexico, S 


er. Bot. 66(2): 183 


-188. 


Pfeiffer, H. W. 1970. A taxonor 


nic revision 


of the pentandn 


ous species of Ar 


istolochia. Umv. Con- 


necticut Pub. Ser. 134 pp. 










Tellez v., O. 1995. New species . 


of Leguminc 


3sae from Nayari 


t. Mexico. Contr. 


Univ. Michigan Herb. 


20: 231-237. 











MISCELLANEOUS NOTES ON SPHAGNUM— 10 

Howard Crum 
University of Michigan Herbarium 

North University Building 
Ann Arbor, Michigan 48109-1057 

NEW SPECIES 

[sect. Sphagnum] amoenoides Crum, sp. nov. — Type: Brazil. Sao Paulo: 
Serra da Mantiqueira, Pindamonhangaba, Pico do Itapeva, ca. 6 km SE of 
Campos do Jordao, 22°46'S, 45°35'W, ca. 2000 m, 18 Oct 1994, Buck 
26422 (holotype: MICH!; isotype: NY!). Fig. 1. 

Plantae usque ad 4 cm altae. Cellulae epidermidis caulinae sine fibrillis et 
poris. Folia caulina 1.8 mm longa, anguste oblongo-ovata, apice cucullata, mar- 
gine lateral! sulco resorpto; cellulae hyalinae usque ad basim fibrillosae, dorso 
poris parvis annulatis ad angulos instructae, superficie interiore poris nullis. Fasci- 
culi ramorum ramis tribus (1 pendenti). Folia ramulina madida quinquefaria, 1.5- 
1.8 mm longa, oblongo-lanceolata, apice cucullata; cellulae hyalinae poris et fibril- 
lis ut in foliis cauhnis; cellulae chlorophylliferae in sectione transversali orciformes 
utroque latere folii parietibus incrassatis brunneis liberae. 

Plants small and slender (up to 4 cm high), pale green tinged with brown. 
Stem cortex efibrillose, without pores; wood cylinder brown. Stem leaves 1.8 mm 
long, narrowly oblong-ovate, cucullate-concave, bordered by a resorption furrow; 
hyaline cells fibrillose nearly to the base, on the outer surface with small, strongly 
ringed pores at angles (some of them in 3's), on the inner surface without pores. 
Branches in fascicles of 3 (1 ± pendent); cortex efibrillose. Branch leaves 5-ranked 
when moist, 1.5-1.8 mm long, oblong-lanceolate; hyaline cells as in stem leaves; 
green cells barrel-shaped, exposed on both surfaces with a thickened brown wall, 
the hyaline cells convex on both surfaces. 

The plants are small, with short branches and spreading leaves. They show 
some similarity to S. amoenum Warnst. but have stems with a brown wood cylin- 
der and larger stem and branch leaves of different porosity. Both stem and branch 
leaves have small, ringed pores on the outer surface but none on the inner, and 
the hyaline cells of stem leaves are fibrillose nearly throughout. Sphagnum amoe- 
num has more differentiated leaves, those of the stems efibrillose, with pores 
none or few on the outer surface and small to large membrane gaps on the inner, 
and the branch leaves have numerous pseudopores on the outer surfaces and both 
pseudopores and membrane pleats on the inner. 

Sphagnum [sect. Sphagnum] atroligneum Crum, sp. nov. — Type: Brazil. Parana: 
Bergland bei Lapa, 70 km SW von Curitiba, sickerfeuchte Sandsteinfelsen 
und Quellmulden bei der Gruta do Monge, ca. 25°47'S, 49°42'W, 590- 
1000 mm NN, 17-18 Dec 1991, Schafer-Verwimp & Verwimp 15214 (holo- 
type: MICH!; isotype: Schafer- Verwimp!). Fig. 2. 




FIG. 1. Sphagnum am 
c. Upper cells of branch le 
Stem leaves, x24. f. Uppe: 
surface, x390. 



b. Portion c 

r surface, x390. d. Upper cells of branch leaf, inner surface, x390. < 
if stem leaf, outer surface, x390. g. Upper cells of stem leaf, inne 



Plantae pusillae, fuscae. Caules plus minusve subnigri; cellulae epidermidis 
delicate fibrillosae, uniporosae; cylindrus lignosus brunneo-niger. Folia caulina 
1.8-1.9 mm longa, concava, lingulata, apice fimbriatula, inferne sulco resorpto 
marginata; cellulae hyalinae omnino fibrillosae, dorso 2-5 poris rotundo-ellipticis, 
superficie interiore poris 1-2 magnis, rotundis, saepe etiam margine pseudoporis 
commissuralis. Fasciculi ramorum ramis 2-3 (1 pendemi); cellulae epidermidis 
fibrillosae, uniporosae. Folia ramulina 1.8-2 mm longa, oblongo-ovata, cucullata, 
dorso poris magnis ad angulos cellularum, interiore 0-3 poris parvis ad angulos et 
commissuras dispositis; cellulae chlorophylliferae in sectione transversali orciformes, 
utroque latere foliorum liberae, superficie exteriore plus minusve latius crassiusculae. 

Plants small, brown. Stems very dark, even black; cortex (2- ) 3 (-4)-layered, 
delicately fibrillose, uniporose; wood cylinder blackish brown. Stem leaves 1.8-1.9 
mm long, Ungulate, bordered by a resorption furrow below, somewhat fringed at 
the broad apex, concave, fibrillose nearly to the base, on the outer surface with 2- 
5 moderately large, ringed, elliptic or rounded-elliptic pores, in 3's at adjacent 
angles, on the inner surface many cells with 1 or sometimes 2 large, round pores. 



CRUM: SPHAGNUM 






Je J^O~Y g MniOJ 



m 



FIG. 2. Sphagnum atroligneum. a. Branch leaves, x24. b. Portion of branch leaf in sectior 
.Upper cells of branch leaf, outer surface, x390. d. Upper cells of branch leaf, inner surface, > 
, x24. f. Upper cells of stem leaf, outer surface, x390. g. Upper cells of stem leal 




, x390. 



often with a few small, ringed, elliptic pseudopores as well. Branches in fascicles 
of 2 or 3 (2 spreading, 1 pendent); cortex dehcately fibrillose, uniporose. Branch 
leaves 1.8-2 mm long, oblong-ovate, cucullate-concave, on the outer surface with 
large, ringed pseudopores in 3's at adjacent corners, on the inner surface with 0-3 
small, ringed pseudopores at commissures and side angles and sometimes 1 or few 
round pores at angles; green cells barrel-shaped, narrowly exposed on both sur- 
faces because of thickened cell ends (but rather more broadly so on the outer 
surface). 

The stem and branch leaves are quite similar, except that the stem leaves 
have a fair number of pores (2-5) at side angles and commissures in addition to 
those in 3's at adjacent corners on the outer surface and 1-2 large round pores on 
the inner, while the branch leaves have large, ringed pseudopores in 3's at adja- 
cent corners on the outer surface and fewer smaller, ringed pseudopores on the 
inner. The dark stems and their blackish wood cyhnders are interesting. 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

[sect. Sphagnum] billbuckii Crum, sp. nov. — Type: Brazil. Minas Gerais; 
Serra do Espinha^o, Mpio. Santa Barbara, Parque Natural do Cara^a, 
along trail from Santuario do Carapuga to Pico da Carapuga, 20°02'S, 
43°30'W, alternately moist and dry sandstone, 1300-1460 m, 22 Oct 1994, 
Wm. R. Buck 26642 (holotype: MICH!; isotype: NY!). Fig. 3. 

Plantae minutae, 1-2 cm altae, pallide virides. Cellulae epidermidis caulinae 
efibrillosae, porosae; cylindrus lignosus pallide fulvus. Folia caulina 1.5-1.8 mm 
longa, ovata, cucullata, margine sulco resorpto; cellulae hyalinae omnino fibrillo- 
sae, superficie exteriore poris plus minusve parvis ad angulos, interiore poris mag- 
nis ad margines foliorum dispositis. Fasciculi ramorum ramis 2-3 (1-2 pendenti- 
bus), cellulis epidermidis ramorum patentium efibrillosis, pendentium distincte 
fibrillosis. Folia ramulina 2 mm longa, ovato-acuminata, apice patentia; cellulae 
hyalinae ut in foliis caulinis; cellulae chlorophylliferae in sectione transversali 
anguste triangulares, latere foliorum interiore liberae. 

Plants very small and slender, 1-2 cm high, pale green. Cortical cells of stems 
efibrillose at the surface, porose; wood cylinder pale brown. Stem leaves 1.5-1.8 
mm long, ovate, cucullate-concave, bordered by a resorption furrow; hyaline cells 
fibrillose throughout, on the outer surface with rather small, elliptic pores at 
angles (in 3's at adjacent corners), on the inner surface with large, round pores 
toward the leaf margins, otherwise very few or none. Branches in fascicles of 2-3 
(1-2 very slender and pendent), the spreading branches tapered; cortical cells of 
spreading branches efibrillose at the surface, those of the pendent branches dis- 
tinctly fibrillose, porose. Branch leaves 2 mm long, ovate-acuminate, spreading at 
the tips; hyaline cells as in stem leaves; green cells very narrowly triangular, some- 
what exposed on the inner surface, the hyahne cells bulging on both surfaces, with 
side walls in well-formed leaves very minutely and faintly papillose. 

The plants are small and have tapered branches, spreading leaf tips, isophyl- 
lous stem and branch leaves, sides of hyaline cells very minutely papillose (at least 
in well-formed leaves), and green cells narrowly triangular with exposure on the 
inner surface. The species is somewhat like S. brevirameum Hampe, except for 
smaller size, leaves larger and isophyllous (with resorption furrows in both stem 
and branch leaves), fewer branches per fascicle, and green cells not central and 



Sphagnum [sect. Cuspidata] subditivum Crum, sp. nov.— Type: New Zealand. 
Canterbury: emergent at edge of tarn in hollow among Sphagnum aus- 
trale, moist, subalpine Notofagus solandri forest, 900 m, S bank of upper 
reaches of Bealey River near Margaret's Tarn (Bealey Glacier Track), 
Arthur's Pass National Park, 2 Dec 1986, A. Fife (with C. D. Meurk) 8011 
(holotype: MICH!; isotype: CHR!). Fig. 4. 

Plantae plus minusve graciles, laete viridi-luteae. Epidermis caulina ex stratis 
2 composita; cylindrus lignosus pallide luteus. Foha caulina 1.4-1.7 mm longa, 
oblongo-ovata vel elliptica, superne concava, acuta, anguste limbata, limbo deor- 
sum non dilatato; cellulae hyalinae fibrillosae ad fohi basin, dorso apice pseudoporis 
annulatis et poris veris non annulatis in seriebus commissuralibus brevibus dis- 
positis, superficie interiore poris magnis rotundis non annulatis pluribus. Folia 
ramulina ca. 1.7 mm longa, ovato-lanceolata; cellulae hyalinae dorso pseudoporis 
annulatis et poris veris non annulatis in seriebus dis 



^UM: SPHAGNUM 




dispositis, superficie interiore poris magnis rotundis pluribus in seriebus duobus; 
cellulae chlorophylliferae in sectione transversali triangulatae, utroque latere folii 
liberae, cum pariete longiore superficie exteriore sitae. 

Relatively slender plants in loose, bright, yellow-green mats. Stem cortex well 
differentiated in 2 layers, without pores; wood cylinder light yellow. Stem leaves 
1.4-1.7 mm long, oblong-elliptic to oblong-ovate, gradually narrowed to a slightly 
concave, acute apex, narrowly bordered; hyaline cells fibrillose throughout, on the 
outer surface near the apex undivided, with numerous ringed, elliptic pseudopores 
in discontinuous commissural rows and a scattering of small, round, thin-mar- 
gined pores, also at commissures, toward the leaf middle occasionally 2-divided, 
without pores or pseudopores or occasionally with a small, ringed pseudopore at 
cell angles, on the inner surface with numerous well-spaced, large, round, thin- 
margined pores in 2 rows. Branches in crowded fascicles of 4 (3 stouter and ± 
deflexed, 1 slender and pendent); retort cells with inconspicuous necks. Branch 
leaves about as long as stem leaves, flattened out and slightly undulate-margined 
when dry, distinctly concave when moist, ovate-lanceolate, narrowly bordered, 
often somewhat serrulate toward the apex; hyaline cells on the outer surface with 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 




I-I(. 4 SphnKintin Mihditu uni. a. Branch leaves, x24. b. Portion of branch 1. 
Uppci cells of blanch leal, outei surface, x390. d. Upper cells of branch 
'ortion or stem in section. x3yO 1 Stem leaves, x24. g. Upper cells of stei 
Upper cells ol stem leaf mnei surface, x390. 



many ringed, elliptic pseudopores and a scattered few round, thin-margined pores 
in rather short commissural rows, often between well-spaced fibrils, also with a 
very few tiny, round, thin-margined pores in a more median position, on the inner 
surface with many well-spaced, large, round, thin-margined pores near the com- 
missures; green cells in section triangular, the lumen submedian in position, broadly 
exposed on the outer surface and variously exposed by wall thickening on the 
inner, the hyaline cells bulging on the inner surface, plane on the outer. 

The plants are curiously bright and greenish yellow, with branches crowded 
and deflexed. The specific epithet, signifying false, was chosen because of a super- 
ficial resemblance to section Siibsecunda. As commonly seen in the section Cuspi- 



1997 CRUM: SPHAGNUM 153 

data, the leaves are flattened out and somewhat wavy-margined when dry. The 
well-marked cortex of the stem suggests that the plants grew in moist hollows but 
not normally submerged. The arrangement of ringed pseudopores and non-ringed 
pores in discontinuous commissural rows on the outer surface of the hyaline cells 
of branch leaves (and the apical cells of stem leaves) are unusual in the Cuspidata, 
but the large, round, thin-margined pores on the inner surface and triangular 
green cells with broader exposure on the outer give evidence of a relationship 
there. 

Sphagnum [sect. Subsecunda] bocainense Crum, sp. nov.— Type: Brazil. Sao Paulo: 
Serra da Bocaina bei Sao Jose do Barreiro, an quelligem Hang an der 
Strasse bei "Shangra-la," ca. 22°47'S, 44°38'W, 1500 m, 16 Apr 1988, 
Schdfer-Verwimp & Verwimp 9588 (holotype: MICH!). Fig. 5. 

Plantae 7-8 cm altae, pallide virides vel aurantiacae. Epidermis caulina strato 
uno, sine poris; cylindrus lignosus palHde fulvus. Folia caulina 1.1 mm longa, 
oblongo-ovata, concavo-obtusa; cellulae hyalinae fere usque ad basim fibrillosae, 
interdum 1-septatae, superfice exteriore poris valde annulatis, permultis in serie- 
bus ad commissuras dispositis, interiore poris multis vel margine paucis. Fasciculi 
ramorum ramis duobus, brevibus. FoHa ramulina 1 mm longa, ovato-lanceolata; 
cellulae hyalinae superficie exteriore poris valde annulatis seriebus commissurali- 
bus continuis, interiore poris vel pseudoporis paucis vel nullis; cellulae chloro- 
phylliferae in sectione transverali orciformes vel triangulo-trapezoideae, pariete 
exteriore longiore. 

Plants 7-8 cm high, light green or tinged with orange brown. Epidermis of the 
stem in 1 layer, without pores; wood cylinder pale yellowish brown. Stem leaves 
oblong-ovate, concave-obtuse, 1.1 mm long; hyaline cells fibrillose nearly to the 
base, not or sometimes 1-divided, on the outer surface with strongly ringed, ellip- 
tic pores in continuous commissural rows, on the inner surface with pores none or 
occasional, more numerous toward the margins. Branches short, in fascicles of 2 
(both spreading, 1 slightly smaller and somewhat decurved). Branch leaves 1 mm 
long, ovate-lanceolate; hyaline cells fibrillose, on the outer surface with heavily 
ringed, elliptic pores in continuous commissural rows, on the inner surface with 
elliptic pores or pseudopores few or none; green cells in section barrel-shaped to 
triangular-trapezoidal, more broadly exposed on the inner surface, the hyaline 
cells convex on both surfaces but more strongly so on the outer. 

The plants are tinged with a soft shade of orange brown. The branches are in 
fascicles of two, both spreading; the stem and branch leaves are similar in struc- 
ture, having an abundance of strongly ringed pores on the outer surface but few 
or none on the inner, and green cells are more broadly exposed on the inner face. 

Sphagnum [sect. Subsecunda] laxiramosum Crum, sp. nov. — Type: Brazil. Mato 
Grosso: Chapada dos Guimaraes NE Cuiaba, Schluchtwald beim Cachoe- 
rinha, an sickerfeuchter Felswand, 600 m NN, 15°28'S, 55°49'W, 4 Jul 
1987, Schdfer-Verwimp & Verwimp 8591 (holotype: MICH!). Fig. 6. 

Plantae tenues, usque ad 10 cm altae, capitulo nullo sed apice aliquantum 
bulboso-geminato. Cellulae epidermidis strato uno, sine poris; cyhndrus lignosus 
luteo-aurantiacus. Folia caulina 1 mm longa, ovata, cucullata; cellulae hyalinae ad 
basim fibrillosae, non septatae, superficie exteriore poris 7-8, non annulatis, ad 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 




commissuras dispersis, interiore poris annulatis in seriebus commissuralibus con- 
tinuis. Rami fere solitarii sed interdum 2 vel 3 (1 pendens). Folia ramulina 1-1.1 
mm longa, ovato-lanceolata, acuminata; cellulae hyalinae superficie exteriore poris 
multis parvis annulatis in seriebus commissuralibus discontinuis, superficie interi- 
ore poris parvis, annulatis, in seriebus commissuralibus discontinuis; cellulae chlo- 
rophylliferae in sectione transversali lenticulares, utroque latere foliorum anguste 

Slender plants in loose tufts as much as 10 cm high, without a capitulum but 
ending in a prominent terminal bud. Epidermal cells of stems in 1 layer, without 
pores; wood cylinder orange-yellow. Stem leaves 1 mm long, ovate, concave (simi- 
lar to branch leaves), rounded-cucullate at the apex; hyaline cells fibrillose through- 
out, not divided, on the outer surface with 7-8 thin-margined pores scattered 
along the commissures, on the inner surface with ringed, elhptic pores in continu- 
ous commissural rows. Branches mostly single, laxly foliate (but occasionally 2 or 
rarely 3, with 1 short, weak pendent branch. Branch leaves 1-1.1 mm long, ovate- 
lanceolate, acuminate; hyaline cells on the outer surface with many small, ringed 
or thin-margined, elliptic pores in discontinuous commissural rows, on the inner 
surface with pores small, ringed, and rounded-elliptic in nearly continuous com- 
missural rows; green cells in section lenticular, very narrowly and equally exposed 
on both surfaces, the hyaline cells somewhat convex on both surfaces. 

The plants have well-spaced, laxly foliate branches that are mostly single, very 
similar stem and branch leaves with ringed commissural rows on the inner surface 
but rather few and often unringed on the outer, and green cells lenticular in 
section. Sphagnum delamboyense, which was collected at the same locality has 
stems with a dark red-brown wood cylinder, stem leaves triangular with fewer 



CRUM: SPHAGNUM 




FIG. 6. Sphagnum laxiramosum. a. Branch leaves, x24. b. Portion of branch leaf in section, x39( 
:. Upper cells of branch leaf, outer surface, x390. d. Upper cells of branch leaf, inner surface, x390. i 
;tem leaves, x24. f. Upper cells of stem leaf, outer surface, x390. g. Upper cells of stem leaf, inne 



pores on both surfaces, those on the inner surface more or less irregularly arranged, 
branches in threes, branch leaves larger and ovate with pores in nearly continuous 
commissural rows on both surfaces, and green cells barrel-shaped. 

Sphagnum [sect. Suhsecunda] rio-negrense Crum, sp. nov. — Type: Colombia. Cun- 
dinamarca: Paramo de Palacio, caveceras Rio Negro, 3375 m, 22 May 
1972, A. M. Cleef4001 (holotype: NY!). Fig. 7. 

Plantae tumidae, ca 6 cm altae, brunneae vel fulvae. Caules simplices, valde 
tenues; cellulae epidermidis 1-2-stratosae, saepe porosae; cylindrus lignosus fus- 
cus vel rufo-brunneus. FoUa 2 mm longa, ovato-elliptica, profunde concava; cellu- 
lae hyalinae non septatae, superficie exteriore fibrillis vestigialibus, poris parvis, 
annulatis, in seriebus commissuralibus continuis, interiore pseudoporis fibrillosis 
annulatis paucis vel numerosis, in seriebus commissuralibus discontinuis; cellulae 
chlorophylliferae lenticulares, inclusae vel utroque latere fohi parietibus incrassa- 



Plants brown or yellow-brown, tumid, ca. 6 cm high. Stems simple, very si 
der, brown; epidermal cells often porose, 1-2-layered; wood cylinder brown 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME / 




red-brown. Stem leaves 2 mm long, ovate-elliptic, deeply concave; hyaline cells 
not divided, fibrillose on the inner surface but with fibril stumps on the outer, on 
the outer surface with small, ringed, rounded-elliptic pores crowded in commis- 
sural rows, on the inner surface with few scattered to rather numerous ringed 
pseudopores in discontinuous commissural rows; green cells lenticular, included 
or narrowly exposed on both surfaces by thickened end walls or more broadly 
exposed on the outer surface, the hyaline cells plane. 



SPECi 



Butriago, 3350 rr 



JED. Colon 
' Apr 1973, A. Cleef9580 (NY), 






: Palacio, Lagunas de 



The stems are simple, and the plants have much the same appearance as the 
North American Sphagnum cyclophylliim Sull., which has much larger leaves with 
minute, rounded, well-spaced pores on the outer surface and only a few corner 
pores, or none at all, on the inner surface, and trapezoidal green cells that in 
section show a broader exposure on the inner surface. 

Sphagnum [sect. Siibsecimda] subhomophyllum Crum, sp. nov.— Type: Brazil. 
Rio de Janeiro: Serra de Itatiaia bei Visconde de Maua, Triefstelle an der 
Strasse nach Maromba, 1400 m, 17 Apr 1987, Schdfer-Verwimp & Ver- 
wimp 8323 (holotype: MICH!; isotype: Schafer-Verwimp!). Fig. 8. 

Plantae ca 7 cm altae, pallido-brunneae, superne aeneae. Caules brunnei; 
cellulae corticis unistratosae, sine poris; cyhndrus lignosus brunneus. Folia caulina 
1-2 mm longa, oblongo-triangularia, concavo-acuta; cellulae hyalinae dimidio su- 
periore folii fibrillosae, non septatae, utroque superficie poris numerosis in 
seriebus commissuralibus discontinuis et superne apice etiam poris vel pseudoporis 



CRUM: SPHAGNUM 




:G. 8. Sphagnum subhomophyllum. a. Branch leaves, x24. b. Portion of bran 
. Upper cells of branch leaf, outer surface, x390. d. Upper cells of branch k 
. Stem leaves, x24. f. Upper cells of stem leaf, outer surface, x390. g. Upper 



in medio cellulae medio instructis. Fasciculi ramorum ramis tribus (1 pendenti). 
Folia ramulina 1.8-2 mm longa, oblongo-lanceolata; cellulae hyalinae dorso poris 
ellipticis in seriebus commissuralibus continuis, apice folii in medio cellularum 
poris 2-6 (-12), rotundis, superficie interiore poris in seriebus commissuralibus 
fere continuis; cellulae chlorophylliferae in sectione transversali ellipticae vel len- 
ticulares, utroque latere peranguste liberae, praesertim superficie exteriore. 

Plants up to 7 cm tall, pale brown, bronze above. Stems brown; cortex 1- 
layered, without pores; wood cylinder brown. Stem leaves 1-2 mm long, oblong- 
triangular, concave-acute; hyaline cells fibrillose in the upper half or two-thirds, 
undivided, on both surfaces with numerous rounded-elliptic pores in discontinuous 
commissural rows and near the apex 2-3 or more round pores or pseudopores in a 
median row. Branches in fascicles of 3 (2 spreading, 1 weak and pendent). Branch 
leaves 1.8-2 mm long, oblong-lanceolate; hyaline cells on the outer surface with 



158 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

elliptic pores in continuous commissural rows and toward the apex 2-6, rarely 12 
round, median pseudopores, on the inner surface with pores in nearly continuous 
commissural rows; green cells elliptic to lenticular, very narrowly exposed on one 
or both surfaces, especially the outer. 

The species is interesting because of median pseudopores on the outer surfac- 
es toward the apex of both stem and branch leaves and also on the inner surface 
quite near the apex of stem leaves. Sphagnum homophyUum Crum and S. pluri- 
porosum Crum have median pores rather than pseudopores. In addition, S. homo- 
phyUum has shorter branch leaves with the commissural pores on the outer sur- 
face indistinctly partitioned off and green cells more exposed on both surfaces. 
Sphagnum pluriporosum also has shorter branch leaves and green cells central 
and included. 

Sphagnum [Sect. Subsecunda] sumapazense Crum, sp. nov.— Type: Colombia. Meta: 
Paramo de Sumapaz, Hoya de la Quebrada Sitiales, 2.5 km SW Laguna 
La Primavera, 3500 m, 26 Jan 1972, A. M. Cleef 1052 (holotype: MICH!; 
isotype: NY!). Fig. 9. 

Plantae ca 10 cm akae, tumidae, subsimplices. Caules tenues, obscure fusci; 
cellulae corticis stratis duobus, saepe uniporosae; cyUndrus lignosus obscure fuscus. 
Folia cauhna 2 mm longa, profunde concava, late ovata; cellulae hyalinae parce 1- 
septatae, fibrillis nullis vel tantum praesentibus, dorso poris annulatis in seriebus 
commissuralibus continuis, etiam 0-7 (-15) poris minutis, non annulatis in medio 
cellulae, superficie interiore poris et pseudoporis rotundis, annulatis, in seriebus 
commissurahbus continuis dispositis; cellulae chlorophylliferae in sectione trans- 
versah anguste orciformes, utroque superficie liberae propter crassitunicas. 

Plants brown, tumid, nearly simple (the branches scarcely emergent beyond 
stem leaves), up to 10 cm high. Stems slender, dark brown; cortex in 2 layers, the 
cells often 1-porose; wood cylinder dark brown. Stem leaves 2 mm long, deeply 
concave, broadly ovate; hyaline cells not or rarely 1-divided, with fibrils reduced 
or lacking, on the outer surface with rounded-elliptic, ringed pores in continuous 
commissural rows and also with 0-7 (-15) minute, round, unringed pores in a 
median position, on the inner surface with ringed, rounded-elliptic pores and 
pseudopores in continuous commissural rows; green cells narrowly barrel-shaped, 
with central lumina, exposed on both surfaces by thickened cell ends, the hyaline 
cells plane on both surfaces. 

Additional Specimens Examined. Colombia. Cundinamarca: Paramo de Sumapaz, Chisaca, La- 
guna N of Laguna Grande, 3650 m, 15 Nov 197L A. M. Cleef & R. Jaramillo-M. 41, (MICH, NY).— 
Without locality: R. E. Schultes 11475 (MICH, NY, TENN). 

The plants bear a considerable resemblance to S. cyclophyllum Sull. but have 
much smaller leaves with pores on the outer surface crowded in commissural rows 
and also well-spaced, small median pores of a variable number. In the type collec- 
tion the median pores are rather consistently numerous, often as many as 11 or 15 
per cell, but in the other collections they vary from one to seven or may even be 
absent. The hyahne cells of the stem leaves are virtually lacking in fibrils. Sphag- 
num rio-negrense Crum, also from Colombia, is rather similar, but its leaves have 
no median pores at all and fibrils on the inner surface. 



CRUM: SPHAGNUM 



\ \ 











FIG. 9. Sphagnum sumapazense 
leaf in section, x390. c. Upper cells c 
x390. e. Cortical cells of stem, in surf: 



■s, x24. b. Portion of stem in section, x390 c Portion c 
Iter surface, x390. d. Upper cells of leaf, inner surface 
x390. f. Portion of stem in section, x390. 



ADDITIONAL NOTE 

mii Crum, Contr. Univ. Michigan Herb. 20:130. 1995.— Type: Brazil. 
Sao Paulo: an der Kiistenstrasse SP55 zwischen Peruibe und Itahaem, feuchter 
Restingawald und anschliessende offene Sandflachen, ca. 5 m.s.m., 17 Jul 
1977, J.-P. Frahm 1866, 1869, 1874 (holotype: MICH!; isotype: ALTAI). 



I am indebted lo Marshall Crosby for pointing out that this species was not 
validly pubhshed because the type was cited from both MICH and ALTA. The 
specimen in the Herbarium of the University of Michigan is hereby designated as 
the holotype. 



LITERATURE CITED 



Crum, H. 1984. Sphagnopsida, Sphagnaceae. North Amer. Fl., ser. 2, pt. 11: 1-180. 
Yamaguchi, T., R. D. Seppelt, Z. Iwatsuki, and A. M. Buchanan. 1990. Sphagnum (sect. L 

leucobryoides sect, et sp. nov. from Tasmania. J. Bryol. 6: 45-54. 
Yamaguchi, T., R. D. Seppelt, and Z. Iwatsuki. 1992. Buchanania {Sphagnum, Sphagn 



CATALOG OF THE ACANTHACEAE OF BELIZE WITH 
TAXONOMIC AND PHYTOGEOGRAPHIC NOTES 

Thomas F. Daniel 

Department of Botany 

California Academy of Sciences 

San Francisco, California 94118 



INTRODUCTION 

Belize is the second smallest country in Central America, with an area of 
about 23,000 square kilometers. It has the lowest population density in Central 
America with about 9 people per square kilometer (Famighetti 1995). The land- 
scape comprises a low, often swampy Caribbean coastal plain, numerous cays, and 
a relatively low mountain range in the southern portion of the country. Approxi- 
mately 3,200 species of flowering plants occur there (Spellman et al. 1975; Dwyer 
& Spellman 1981). The comprehensive Flora of Guatemala (Standley et al. 1958- 
1977) included Belize within its geographic scope, thus making it the best source 
of information for plants in this small nation. 

Dwyer and Spellman's (1981) hst of dicotyledonous plants in Belize provided 
the most recent account of Acanthaceae for the country. They listed 50 species of 
the family and cited collections for some of them. Daniel (1993) documented the 
occurrence in Belize of two species {Carlowrightia myriantha and Justicia albo- 
bractea) previously excluded or not known from the country. Daniel (1995b) sub- 
sequently noted the presence in Belize of several additional members of Acan- 
thaceae (e.g., Justicia candelariae and Ruellia hookeriana), but did not document 
their occurrence by citing collections. Given the recent renewed interest in the 
botanical resources of Belize (e.g., the Flora Mesoamericana project; the New 
York Botanical Garden's project to produce a checklist of the flora of Belize), a 
verified account of the Acanthaceae in Belize is warranted. 

In the following catalog, 47 species in 18 genera are documented from Belize, 
some for the first time. These comprise 40 native species and seven introduced 
species. Native Belizean Acanthaceae (Appendix) can be classified according to 
their overall distributions as widespread (i.e., occurring over a broader area than 
defined below), regional (i.e., restricted to the region from the Isthmus of Tehuan- 
tepec in southern Mexico eastward through Guatemala, Belize, and the Yucatan 
Peninsula to the lowlands of northwestern Honduras), local (i.e., restricted to 
Belize and adjacent regions of Guatemala, Honduras, and/or the Yucatan Penin- 
sula of Mexico), and endemic (i.e., restricted to Belize). The appendix shows that 
the greatest proportion of native species are widespread. Indeed, for the Belizean 
flora as a whole, Lundell (1945) had noted that the majority of the species there 
are widely distributed ones of the West Indies and the Caribbean slope of Mexico 
and Central America. A sizable percentage of Belizean Acanthaceae (30% of the 
native species) is regional in distribution, that is, restricted to the northern Meso- 
american region. If this region were extended slightly to include some of the moist 
to wet forests encountered further to the north in Veracruz, at least one additional 
species here treated as widespread (i.e., Dicliptera sumichrastii) would be added 

161 



162 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

to this category. A single species of Acanthaceae, Louteridium chartaceum, is 
endemic to Belize. This species was discussed by Daniel (1993) who, at that time, 
noted three other endemic Acanthaceae in Belize. Each of the other three was 
subsequently (Daniel 1995b) treated as also occurring outside of the country. The 
percentage of endemic Acanthaceae with respect to all Belizean members of the 
family (i.e., native and introduced) is 2%. This percentage is considerably lower 
than the estimate of 4.6-6% endemism provided by Hampshire (1989) for the 
Belizean flora as a whole. The percentage of endemism among Belizean Acan- 
thaceae is also much lower than that found among Acanthaceae in nearby larger 
regions with considerably greater diversities of climate and habitat, e.g., Guate 
mala (10%, i.e., 13 endemics/128 species; totals based on Gibson, 1974, with cor 
rections in Daniel, 1995b, and various geographic updates) and Chiapas, Mexicc 
(11%, i.e., 15 endemics/131 species; based on Daniel, 1995b). For more meaning 
ful phytogeographic data, particularly regarding endemism, Belize should be re 
garded as part of a greater Yucatan region or divided into northern/drier anc 
southern/moister regions that could be treated along with adjacent portions o 
environmentally similar surrounding regions of Mexico and Guatemala (cf. Wend 
1993: 596). 

Collections have been made of at least seven species of Acanthaceae {Androgra 
phis gangetica, Hemigmphis alternata, Ruellia coerulea, Thunbergia data, T. erecta, J 
fragrans, T. grandiflora) that have been introduced into Belize from other parts o: 
the world. These exotics include cultivated and/or naturalized species. Additiona 
non-native Acanthaceae almost certainly are cultivated in the country. 

Within the country, the greatest concentrations of species of Acanthaceae are 
found in the southern three districts (Fig 1). This is likely due to the diversity oi 
habitats created in southern Belize by the Maya Mountains and the presence 
tropical rain forest there. In spite of the greater diversity of species in the south, 
particularly species associated with moist to wet forests (e.g., Aphelandra auranti 
aca, Bravaisia grandiflora, Justicia albobractea, J. aurea, J. fimbriata, Louteridium 
donnell-smithii, and Mendoncia spp.), the acanthaceous flora of Belize 
particularly in Corozal, by "dry forest species" or species that are mc 
in the northern portion of the Yucatan peninsula (e.g., Bravaisia berlandieria. 
Carlowrightia myriantha, and Justicia cainpechiana). 

Speliman et al. (1975) remarked on the West Indian floristic influence 
Belize. There are no Acanthaceae known from the West Indies that occur on i 
American continent only in Belize. Although 10 of the 40 (25%) native Ac; 
thaceae of Belize also occur in the West Indies, all ten are widely distributed 
species, and none could be considered as primarily West Indian in distribution. 
Interestingly, no Acanthaceae have been reported from any of the numerous 
small islands (cays) off Belize (Fosberg et al. 1982). 

For each species included in the following annotated catalog, the district(s) o 
occurrence and one or more collections that I have examined and identified are 
cited in order to voucher the occurrences and to provide a general indication o: 
distribution within the country. Identification keys to most of these species can be 
found in the floristic accounts of Gibson (1974) for Guatemala and Daniel (1995b 
for Chiapas, Mexico. Distinguishing features of species not treated in those 
accounts are provided herein. Because many Acanthaceae thrive in disturbed 
habitats and because portions of Belize remain inadequately collected, it seems 
likely that additional members of the family will be found in the country. Based 
on their overall distributions and the likelihood of appropriate habitats in the 



lEL: ACANTHACEAE OF BELIZE 




164 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

country, the following species might be expected to occur in Belize: Elytraria 
bromoldes Oerst., Justlcia carthagenensis Jacq., /. colorifera V.A.W. Graham, Pseu- 
deranthemum datum (Nees) Radlk., P. cuspldatum (Nees) Radlk., Ruellia inundata 
Kunth, R. paniculata L., Stenandrium duke (Cav.) Nees, S. pedunculatum (Donn. 
Sm.) Leonard, and S. subcordatum Standi. 



CATALOG OF THE ACANTHACEAE OF BELIZE 
I Lindl. (including A. repanda Nees). 



The author of A. aurantiaca is usually cited as "(Scheidw.) Lindl." (e.g., Gib- 
son 1974; Wasshausen 1975; Durkee 1978, 1986; Dwyer & Spellman 1981; Daniel 
1991, 1995b). As pointed out to me by Mike Grayum in a letter, Hemisandra 
aurantiaca Scheidw. is a heterotypic synonym of, rather than the basionym of, A. 
aurantiaca Lindl. for the following reason. In 1842, Scheidweiler published Hemis- 
andra aurantiaca. In Lindley's publication of A. aurantiaca in 1845, he cited the 
synonym "Hesemasandra aurantiaca, //orr." He further noted (Lindley 1845) that 
plants with this name had been exhibited at a horticultural meeting, but that the 
name was "not to be found in any Botanical books in our possession." Although it 
is very hkely that the origin of Lindley's synonym (i.e., the horticultural nomen 
novum) was somehow derived from Scheidweiler's name, Lindley was clearly 
unaware of Scheidweiler's pubHcation and, indeed, utilized a different generic 
spelling for the name he cited. Thus, Scheidweiler and Lindley described the same 
species, utilizing the same epithet, in different genera based on different speci- 
mens. Scheidweiler's name is older, but, if transferred to Aphelandra, becomes a 
later homonym of A. aurantiaca Lindl. 

The name A. repanda applies to a sporadic form of A. aurantiaca with narrow, 
sinuately margined leaves (see Daniel, 1991: 251). 

Aphelandra scabra (Vahl) Sm. (as A. deppeana Schltdl. & Cham, m Dwyer & 
Spellman, 1981). 

Belize: Croat 23905 (F, NY, US); 
McDaniel & Clark 14348 (F); Worth. 
11942 (MICH, US), 11947 (MICH, US); Chanek 38 (MICH, US); Gentle 2392 (MICH, NY), 888 
(CAS, F, US); Lundell 439 (CAS, F, US); Meave 1025 (MO); Meave & Howe 1164 (MO).— Corozai 
Crane 506 (CAS); Gentle 202 (MICH, US); Kinloch 3 (F).— Orange Walk: Limclell 504 (DS, F, N\ 
US); Karling 12 (F), 51 (F).— Stann Creek: Daniel & Butterwick 5877 (CAS); Molina R. 18 (F); Schipp 3 
(F, MICH, NY, UC, US).— Toledo: Gentle 3774 (MICH, NY, US), 3775 (F, MICH, NY, US); Whitefoor, 
1488 (MO), 1493 (MO), 1811 (MO).— District Unknown: Bartlett 11349 (MICH); Stocker 13 (F). 

Asystasia gangetica (L.) Anders. 

Stann Creek: Gentle 7866 (F, US). 

This Old World species has not been heretofore reported from Belize. It i 
cultivated and has become naturalized in various parts of tropical America (e.g. 
Costa Rica and Panama). The genus is not included in the treatments of Acan 



1997 DANIEL: ACANTHACEAE OF BELIZE 165 

thaceae for Guatemala (Gibson 1974) or Chiapas (Daniel 1995b). It may be dis- 
tinguished from all other Acanthaceae occurring in Belize by the combination of 
its herbaceous habit, infundibular corolla, four didynamous stamens with 2-the- 
cous anthers, and four or fewer seeds lacking hygroscopic trichomes. 

Barleria oenotheroides Dum. Cours. (as B. micans Nees in Dwyer & Spellman 1981). 

Cayo: Balick et al. 1990 (US); Bartiett 11478 (CAS, MICH, US). 

The taxonomy and distribution of this species were discussed by Daniel (1995a). 
Urb. (as B. brownei Juss. in Dwyer & Spellman 



Belize: Dieckman 187 (MO); Liesner & Dwyer 1441 (MO); Lundell 4212 (MICH).— Cayo: Croat 
23701 (MO); Daniel & Butterwick 5876 (CAS); Dwyer et al. 65 (MO); Lundell 4154 (F, MICH); Utley 
803 (F, MO, NY).— Corozal: Daniel 8270 (CAS).— Orange Walk: Daniel 7015 (CAS), 8256 (CAS); 
Egler 42-95 (F); Lundell 24 (F, US).— Stann Creek: Daniel & Butterwick 5881 (CAS); Schipp S-134 
(F), 875 (F, MICH, MO, NY, UC).— Toledo: Croat 24117 {F, MO); Whitefoord 1549 (MO). 

The correct name for this species was discussed by Daniel (1995a). 

Bravaisia berlandieriana (Nees) T. F. Daniel (as B. tubiflora Hemsl. in Dwyer & 
Spellman 1981). 

Belize: Gentle 1193 (A, BM, F, G, GH, K, LL, MICH, MO, NY, US); McDaniel 13080 (MO).— 
Corozal: Balick et al. 3306 (US); Crane 47 (LL), 321 (LL); Daniel 8265 (BR, C, CAS, ENCB, K, 
MEXU, MICH, MO, NY, US); Davidse & Brant 32596 (CAS); Gentle 380 (CAS, F, MICH, US); 
Stevenson 1 (F).— Orange Walk: Arnason & Lambert 17152 (MO); Daniel 8263 (CAS); Winzerling 
VlII-14 (US).— District Unknown: Campbell 1 (K); Stevenson s.n. (US). 

A monograph of this genus was provided by Daniel (1988). 

Bravaisia grandiflora Donn. Sm. 



Carlowrightia myriantha (Standi.) Standi. 



This species, reported from Belize by Daniel (1993), was not treated by Gib- 
son (1974) or Daniel (1995b). It differs from other Belizean Acanthaceae by the 
combination of its stenotribal flowers; white to pink corollas 9-12 mm long with 
the upper hp lacking a rugula and having maroon lines within; two fertile stamens 
with dithecous anthers; parallel and more or less evenly inserted thecae lacking 
basal appendages; lack of staminodes; and glabrous, lenticular seeds 2-3 mm long 
that have entire margins. 

Recent field observations reveal that this rarely collected species is common 
in the seasonal evergreen forests of Corozal, where it flowers during the dry 
season (e.g., March-April). 



166 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Dicliptera inutilis Leonard. 

Corozal: Ramamoorthy et ai 2735 (MEXU). 

Among taxa in Mexico and Central America, this collection most closely 
resembles D. inutilis. It has relatively long (4 cm), apparently pinkish corollas and 
large (15-20 x 11-18 mm), broadly ovate to broadly trullate, and abaxially gla- 
brous outer cymule bracteoles. The notation on the specimen label that the plant 
was a tree four to five meters in height is probably erroneous; trees are unknown 
in the genus. This is the first report of the species in Belize. 

. (as D. assurgens (L.) Juss. m Dwyer & Spellman 



Belize: Daniel & Butterwick 5901 (CAS); Gentle 1119 (F, MO, NY); Peck 351 (GH, NY).-Cayo; 
Daniel & Butterwick 5899 (CAS). 5900 (CAS); Gentle 9062 (CAS, F, MO, US), 9698 (CAS, F. MO, 
NY, US); Lundell4Wl (MICH, US), 4102 (MICH, US), 4103 (MICH, US); McDaniel 13070 (MO).— 
Corozal; Daniel 8271 (CAS); Felly 24 (F).-Oranc;e Walk: Arnason & Lambert 7727 (MO); Daniel 
7040 (CAS), 8215 (BR, C, CAS, K, MEXU, MICH, MO, NY, US).-Stann Creek: Gentle 7882 
(CAS, F, MO, US). 



The specimen at F was annotated by Leonard as D. acuminata (Ruiz & Pav.) 
Juss. and was so treated by him (Leonard 1936). Subsequently, Gibson annotated 
the same specimen as "D. sumichrasti ?" and hkewise treated it (Gibson 1974). 
Various species and complexes of species of North American Dicliptera remain to 
be adequately circumscribed. The collection cited above appears to represent a 
taxon closely related to, if not conspecific with, D. sumichrastii. 

Elytraria imbricata (Vahl) Pers. 

Cayo; Bartlett 11497 (MICH). 

Hemigraphis alternata (Burm. f.) Anders. 

Belize; Bartlett 11355 (MICH; type of Bleehum cordatnm Leonard). 

This native of tropical Asia is naturalized in parts of tropical America. 
Hygrophila costata Nees (as H. guianensis Nees in Dwyer & Spellman 1981 ). 

Cayo: Croat 24852 (MO); Dwyer & Dieckman 13012 (CAS); Dwyer & Liesner 12059 (MO).— Stann 
Creek: Daniel & Butterwick 5884 (CAS); Dwyer et al. 508 (F); Gentle 8627 (CAS, F, MO, US); 
McDaniel 12999 (MO).— Toledo: Peck 759 (GH); Schipp 1109 (F, MICH, MO, NY, UC). 

Justicia albobractea Leonard. 

Toledo: Proctor 36627 (F); Schipp 1277 (F; type). 



1997 DANIEL: ACANTHACEAE OF BELIZE 167 

This species was not listed by Dwyer and Spellman (1981), nor was it treated 
by Gibson (1974) for lack of flowering material. The species was fully described 
by Daniel (1993). It differs from other species of Justicia in Belize by the combi- 
nation of its inflorescence of axillary pedunculate spikes to 65 mm long, ovate to 
elliptic bracts 8.5-16 mm long and 4.5-9.5 mm wide, 5-lobed calyces, corollas 29- 
31 mm long, superposed and parallel thecae 1.6-2 mm long (the lower with a 
basal appendage 0.5 mm long), and 3-aperturate pollen. 

Justicia aurea Schltdl. 

Cayo: Bartlett 12937 (CAS, MICH. US).— Toledo: Daniel & Butterwick 5889 (CAS); Schipp S-601 
(F, MO,NY). 

Justicia bartlettii (Leonard) D. N. Gibson (including/ pilifera D. N. Gibson). 

Belize: Liesner & Dwyer 1486 (MO).— Cayo: Arvigo & Shropshire 203 (MO, US); Bartlett 11477 
(US; type of Beloperone bartlettii Leonard).— Toledo: Davidse & Brant 32157 (CAS). 

Justicia breviflora (Nees) Rusby (including Pseuderanthemum tetrasepalum (S. F. 
Blake) S. F. Blake). 

Belize: Daniel 8296 (CAS).— Cayo: Arvigo et al. 101 (US); Bartlett 11948 (TEX, US), 12017a (MICH), 
13096 (MICH); Cowan et al. 5149 (CAS), 5169 (CAS); Daniel 8273 (CAS, K, MICH); Gentle 2207 
(MICH, NY), 2209 (F, MO, MICH, NY, RSA), 9077 (CAS, F, MO, US), 9645 (F, MO), 9687 (F, MO, 
US); Lundell 6124 (MICH, NY, RSA, US), 6209 (MICH, US), 6290 (F, MICH, NY, US).— Stann 
Creek: Long 3290 (CAS, MO); Molina R. 331 (F); Schipp 542 (F, MICH, MO, NY, UC).— Toledo: 
Cosentino 86 (F); Croat 24317 (F, NY, RSA, US); Daniel & Butterwick 5895 (CAS); Gentle 4507 (MICH), 
5274 (MO, US); Hoist 4025 (CAS),4245 (CAS); McDaniel 12884 (F, MO); Peck 552 (GH; type of Eran- 
themum tetrasepalum S. F. Blake); Peck 722 (GH; type oi Dianthera peckii S. F. Blake); Schipp S-473 (F). 

Some of the morphological variation exhibited by this species in southern 
Mexico and northern Central America was discussed by Daniel (1995b). Pseuder- 
anthemum tetrasepalum is herewith included in the synonymy of this species for 
the first time. In the protologue of P. tetrasepalum, Blake (1917) described the 
corollas as about 2.4 cm long and noted the presence of staminodes. Both of these 
features are more suggestive of Pseuderanthemum than Justicia. In a packet on 
the type, there are flowers of two taxa. Two of these flowers match Blake's descrip- 
tion in length of the corolla and presence of staminodes. These flowers indeed 
belong to a species of Pseuderanthemum. The plant mounted on the sheet and the 
other, smaller flowers in the packet (with anthers pubescent and with thecae 
superposed) pertain to /. breviflora and correspond to the remainder of Blake's 
description. Four-parted calyces are not known in Pseuderanthemum but are often 
present in /. breviflora (Daniel 1995b). Pollen from the mounted specimen and from 
one of the small flowers in the packet on the type of P. tetrasepalum (i.e., 2-porate 
with a row of insulae on either side of each aperture) also resembles that of other 
specimens referred to J. breviflora. The description in Blake's protologue was thus 
derived from species of two genera. In accordance with Article 9.9 of the Internation- 
al Code of Botanical Nomenclature (Greuter et al. 1994), a lectotype for J. breviflora 
is herewith designated as that portion of Peck 552 mounted on the sheet and the 
flowers in the packet with the anthers pubescent and the thecae superposed only, 
thereby excluding the two flowers of Pseuderanthemum also in the packet. 



168 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 

Justicia campechiana Standi 
Corozal: Castillo 44 (F); Pelly 3(¥). 

Justicia candelariae (Oerst.) Leonard. 

Cayo: Gentle 9011 (F, MO, US).— Stann Creek: Gentle 9316 (CAS, F, MO, US). 

This species was not listed for Belize by Dwyer and Spellman 
noted to occur in the country by Daniel (1995b). 

Justicia coniata (L.) Lam. 

Belize: Gentle 913 (F, NY); Liindell 1957 (MICH, US); Whitefoord 2354 (MO).-' 
(F, MICH); Contreras 7145 (F, MO, US).-Stann Creek: Dwyer et al. 504 (M 
MICH, MO, NY, UC).-DiSTRiCT Unknown: Usher 7 (MICH). 

Justicia ensiflora (Standi.) D. N. Gibson. 

Gentry 7931 (MO): Schi 



Justicia fimbriata (Nees) V. A. W. Graham (as J. magniflora (S. F. Blake) D. N. 
Gibson in Dwyer & Spellman 1981). 

Toledo: Balick et al. 2538 (US); Cosentino 90 (F); Daniel & Biitterwick 5893 (CAS, MICH); Davidse 
& Brant 32153 (CAS); Gentle 5060 (MO, US); Peck 622a (GH; type of Dicliptera magniflora S. F. 
Blake); Schipp S-694 (F; type of Beloperone crenata Standi.); Stevenson 82 (F, NY, US). 

Justicia pectoralis Jacq. 

Belize: Gentle 1530 (MICH, US).— Stann Creek: Daniel & Biitterwick 5880 (CAS); Gentle 1893 
(MICH, NY, US).— Toledo: Gentle 4419 (MICH), 5278 (MO, US), 7591 (MO); Peck 979 (GH); 
Proctor 36011 (MO). 



> (GH; type oiJacobinia scarlatina S. F. Blake).— Cayo: Chanek 40 (MICH. US).- 
mnzerling VllI-10 (F, US).— Stann Creek: Gentle 3305 (A, MICH, MO, NY, US 
, GH, MICH, MO, NY, UC).— Toledo: Gentle 7334 (CAS, F, MO, US); Tellez et a 



Daniel (1995b) discussed the distinctions between this species and the similar 
J. colorifera, both of which are cultivated for use as a bluing agent in laundering 
fabric in Central America (Williams 1981). 

Lepidagathis alopecuroidea (Vahl) R. Br. ex Griseb. (as Teliostachya alopecuroidea 
(Vahl) Nees in Dwyer & Spellman 1981). 



1997 DANIEL: ACANTHACEAE OF BELIZE 169 

Louteridium chartaceum Leonard. 

Belize: Daniel 8294 (BR, CAS, MEXU, MICH, MO, US); Daniel & Butterwick 5905 (C, CAS, K, 
MICH, MO. NY); Dwyer 10959 (LL, MO); Gentle 1526 (US; type); Liesner & Dwyer 1485 (BM, 
DUKE, MO, NY, TEX). 

Daniel (1993) noted that this endemic species was known from a single locality 
that was threatened with destruction. Indeed, quarrying activities have since 
destroyed the population at the site (at or near the type locality) from which 
Daniel & Butterwick 5905 was collected. However, other populations have recently 
been located on several of the isolated limestone hills in the southern portion of 
Belize District (e.g., Daniel 8294). These hills persist as islands of relatively undis- 
turbed vegetation among settlements and cultivated lands. 

Louteridium donneil-smithii S. Watson. 

Cayo: Dwyer & Liesner 12313 (MO, NY); Dwyer et al. 360 (MEXU, MO); Gentry 7791 (MO).— 
Toledo: Daniel & Butterwick 5891 (CAS); Davidse & Brant 32089 (CAS, US), 32300 (CAS); Gentle 
6355 (MO, UC, US); Hoist 4045 (CAS); Peck 780 (GH, NY); Proctor 36148 (MO); Schipp 1110 
(MICH, MO, NY, UC). 

Mendoncia lindavii Rusby. 

Stann Creek: Gentle 3344 (F, MICH, MO, NY), 3523 (MICH, MO. NY, US); Schipp 961 (F; type of 
M. belizensis Standi.). 

Mendoncia retusa Turrill. 



Cayo: Bartlett 13065 (CAS, MICH, US); Gentle 2440 (A, F, K, LL, MICH, NY).— Stann Creek: 
Schipp S-278 (A, F, G, GH. K, MICH, MO, NY, UC).— Toledo: Balick et al. 2547 (US); Croat 24383 
(F, MO); Gentle 6567 (MO); Peck 786 (GH, NY), 787 (GH, NY); Proctor 35876 (MO); Whitefoord 
1615 (MO). 

Odontonema callistachyum (Schltdl. & Cham.) Kuntze. 

Cayo: Dwyer c& Dieckman 13014 (CAS); Gentle 2441 (MICH), 8724 (CAS, F, MO, US).— Stann 
Creek: Croat 24523 (F, MO); Daniel & Butterwick 5878 (CAS), 5882 (CAS), 5887 (CAS); Gentle 
2142 (MICH), 3501 (A, F, MICH, NY), 8202 (F, MO, US), 9250 (CAS, F, MO, US).— Toledo: Croat 
24312 (MO); Daniel & Butterwick 5894 (CAS); Davidse & Brant 32067 (CAS). 

This species was treated as a heterogeneous complex by Gibson (1974). Dis- 
cussion of this and the other species of Odontonema noted here can be found in 
Daniel (1995c). 

Odontonema hondurense (Lindau) D. N. Gibson. 



170 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

(MICH, NY), 8835 (CAS, F, MO, US), 9004 (CAS, F, MO).— Stann Creek: Dwyer el al. 571 (MO. 
US); Gentle 2136 (A, DS, F, K, LL, MICH, NY, US), 2158 (MICH), 2742 (MICH), 2793 (MICH, NY, 
US), 3195 (MICH), 3496 (F, MICH, NY); Molina R. 276 (F); Schipp 222 (A, BM, F, G, GH, MICH, NY, 
UC, US).— Toledo: Daniel & Butferwick 5892 (CAS, MICH); Davidse & Brant 31921 (CAS); Gentle 4506 
(LL, MICH, MO), 6704 (F, MO, US); Hoist 3873 (CAS); Proctor 35928 (MO); Whitefoord 1565 (CAS). 

Odontonema tubaeforme (Bertol.) Kuntze (as O. flagellum (Oerst.) Kuntze in 
Dwyer & Spellman, 1981, and including O. amicorum V. M. Baum). 

BiM/i nan,il82-'fU{ \S MICH) Dunul A. BultLr^MLk -^904 {(. h^) (nntk 1518 (C,U LL MICH 



The Uixonomic disposition ol () anmoiuw was additsscd b\ DaniJ (IWx) 
Pseuderanthemuiii >erapa/ciise Donn Sni 

S 671 (h K NY) 

Ruellia coerulea Morong. 

Belize: Daniel 8278 (CAS).— Corozal: Gentle 347 {M\CU). 

This species was not included by Dwyer and Spellman (1981). It is often 
cultivated under the name R. brittoniana Leonard, and it may sometimes become 
naturalized. Daniel 8278 was cultivated in a garden. It is unclear from the label 
data on Gentle 347 whether the collection was cultivated or not. It is not likely 
that R. coerulea is indigenous in Belize. Gentle's collection was identified by 
Leonard in 1936 as R. malacosperma Greenm., a species described from Mexico 
that may be conspecific with R. coerulea. The distribution, taxonomy, and nomen- 
clature of this species were discussed by Daniel (1995b). 



Ruellia geminiflora Kun 



Schipp's collection differs from more typical representatives of the species by 
the inconspicuous punctate glands on the calyx and by the lack of (or very incon- 
spicuous) punctate glands on the corolla. 

Ruellia harveyana Stapf (including R. obtusata S. F. Blake). 

Belize: Dwyer 11317 (MO).— Cayo: Bartlett 13063 (CAS, MICH, NY, US); Gentle 8598 (F, MO, US); 
Limdell 322 (DS, F, UC), 6518 (DS, MICH, NY, US), 6667 (MICH, NY, US); Spelhnan 1577 (MO).— 
Stann Creek: Dwyer et al. 569 (MO); Gentle 7957 (F, MO); Schipp 976 (F, MICH, MO, NY, UC); Stocker 
20 (F; type of R. longipila Standi.).— Toledo: Gentle 7056 (LL); Peck 871 {CA\: type of R. obtusata). 

Ruellia hookeriana (Nees) Hemsl. 



DANIEL: ACANTHACEAE OF BELIZE 



Ruellia matagalpae Linda 



(o: Balick et al. 3346 (US); Gentle 2300 (MICH, N^ 
1 (MO).— Toledo: Davidse & Brant 32422 (CAS); ^ 



Ruellia nudiflora (Engelm. & Gray) Urb. 



): Arvigo 56 (US); Arvigo 1987-17 (F); Balick et al. 1797 (US); Daniel 8272 (CAS).— Corozal: 
? 89 (CAS); Croat 24586 (MO); Dwyer 14504 (MO); Gentle 178 (MICH, US).— Orange Walk: 
son & Lambert 17150 (MO); Daniel 7033 (CAS); Lundell 36 (F, MICH, US), 365 (F, MICH), 
DS. F, MO. NY. US).— District Unknown: Liaulell 4979 (MICH, MO, US). 



Cayo: Bartlett 12900 (CAS, MICH, US); Chanek 39 (MICH 
Gentle 2170 (F, MICH, NY); Lundell 6128 (MICH, NY, L 
(MO).— Toledo: Schipp S-645 (F, NY). 

Ruellia pygmaea Donn. Sm. 

Toledo: Schipp S-668 (F, NY), 135! (F. MICH. MO. NY). 

Thunbergia alata Bojer ex Sims. 

Cayo: C/ume/t 200 (F.MICH). 

This African species is widely cultivated and naturalized in the American tropics. 



Thunbergia erecta (Benth.) T. Ander 
Belize: Daniel 8277 (CAS).— Cayo: Balick 22 



Thunbergia fragrans Roxb. 

Toledo: Gentle 6371 (CAS. F, MO, NY. US). 

This Asian species was not reported by Dwyer and Spellman (1981). It i: 
often cultivated and has become naturalized in various parts of the neotropics. 

Thunbergia grandiflora Roxb. 

Belize: Whitefoord 2453 (MO).— Corcjzal: Gentle 50 (F); Lundell 4786 (MICH, NY).— Stann Creek 



This Asian species is widely cultivated and : 
locally naturalized in tropical regions. 



172 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

EXCLUDED TAXA AND NAMES 

The following names were listed as species occurring in Belize by Dwyer and 
Spellman (1981) but are excluded from the list above. Their respective disposi- 
tions are noted below. For those that are now treated as synonyms of other names 
a reference in which the synonymy was accepted is provided. 

Aphelandra repanda Nees. — Synonym of A. aurantiaca (Daniel 1991). 

A. deppeana Schltdl. & Cham.— Synonym oiA. scabra (Daniel 1991). 

Barleria micans Nees.— Synonym of B. oenotheroides (Daniel 1995a). 

Blechum brownei Juss.— Synonym of B. pyramidatum (Daniel 1995a). 

Bravaisia tubiflora Hemsl. — Synonym of B. berlandieriana (Daniel 1988). 

Dicliptera assurgens (L.) Juss.— Synonym of D. sexangidaris (Daniel 1995b). 

Hygrophila guianensis Nees.— Synonym of H. costata (Daniel 1995b). 

"Justicia assurgens (L.) Juss."— This "name" represents an error in Dwyer and 
Spellman (1981), apparently based on mislabeled specimens at MO, for 
Dicliptera assurgens \=D. sexangularis). 



Justicia sp.—DwyQT and Spellman (1981) listed Croat 23899 i 
species of Justicia. The whereabouts of this collect: 
(T. Croat, pers. comm.). 



O. glabrum Brandegee.— This species was listed by Dwyer and Spellman (1981) 
but was not noted as occurring in Belize by either Baum (1982) or Daniel 
(1995c). Odontonema glabrum was hsted by Leonard (1936) for Belize 
based on Bartlett 12947, which here is assigned to O. hondurense. 

Pseuderanthemum tetrasepalum S. F. Blake.— Synonym of Justicia breviflora (see 

Ruellia biolleyi Lindau.— Dwyer and Spellman (1981) included this species, known 
from lowland rain forests of Costa Rica and Panama, on the basis of 
Dwyer 12061. No specimens of this collection have been located, and it is 
unlikely that the species occurs in Belize. 

RuelHa ohtusata S. F. Blake.— Synonym of R. harveyana (Daniel 1995b). 



1997 DANIEL: ACANTHACEAE OF BELIZE 173 

Ruellia paniculata L.— Dwyer and Spellman (1981) included this species and cited 
Kellman 511. Although the species might be expected to occur in Belize, I 
have not been able to locate this collection. 

"Ruellia pilosa."— Dwyer and Spellman (1981) were presumably referring to R. 
pilosa (Nees) Pav. ex Hemsl. (a later homonym of R. pilosa L.f.), among 
the various taxa given this name. This species was described from Peruvi- 
an and Mexican collections; its circumscription is not well understood. 
Dwyer and Spellman (1981) included this species on the basis of Dwyer 
10155. I have not been able to locate specimens of this collection. 

"Ruellia sarmentosa Jacq."— Dwyer and Spellman's (1981) reference to this "name" 
is unclear. I find no record of such a name used by Jacquin. There is an 
Asian species that was described as Ruellia sarmentosa Nees and is, accord- 
ing to Index Kewensis, Hemigraphis hirta T. Anders. Dwyer and Spellman 
(1981) cited Kellman 583 as representing this taxon; no specimens of this 
collection have been located. 

Ruellia stemonacanthoides (Oerst.) Hemsl. — All specimens from Belize identified 
with or cited under this name are referable to R. pereducta; however, I 
have not located Proctor 30096. which was cited by Dwyer and Spellman 
(1981). It is unlikely that this species of the Pacific escarpment occurs in 
the Caribbean lowlands. 

Stenandrium pedunculatum (Donn. Sm.) Leonard.— This species was included by 
Dwyer and Spellman (1981) without citation of a voucher or source. It 
was not noted as occurring in Belize by Gibson (1974) or Daniel (1985, 
1995b). 

Tehc 



ACKNOWLEDGMENTS 

I am grateful for the assistance of M. Butterwick, D. Wasshausen, M. Grayum, B. Winning, M. 
Balick, the Belize Ministry of Tourism and the Environment (Department of Archaeology), and the 
Belize Ministry of Natural Resources (Department of Forestry). Partial financial support for field 
studies was provided by Oceanic Society Expeditions and the California Academy of Sciences In- 
house Research Fund. I thank curators of the following herbaria for making specimens available for 
study: A, BM, CAS, DS, DUKE, ENCB, F, G, GH, K, LL, MEXU, MICH, MO, NO, NY, RSA, 
TEX, UC, and US. 



LITERATURE CITED 

Baum, V. M. 1982. A revision of the genus Odonlonema (Acanthaceae). Unpublished M.Sc. Thesis, 

University of Maryland, College Park. 
Blake, S. F. 1917. Descriptions of new spermatophytes, chiefly from the collection of Prof. M. E. Peck 

in British Honduras. Contr. Gray Herb. 52: 59-106. 
Daniel, T. F. 1985. A revision of Stenandrium (Acanthaceae) in Mexico and adjacent regions. Ann. 

Missouri Bot. Gard. 71: 1028-1043. 
. 1988. A systematic study of Bravaisia DC. (Acanthaceae). Proc. Calif. Acad. Sci. 45: 1 1 1-132. 



OF MICHIGAN HERBARIUM VOLUME 21 

laceae) in Mexico. Proc. Calif. Acad. Sci. 47: 235-274. 
on Central American Acanthaceae. Proc. Calif. Acad. 

Mexican Acanthaceae. VI. Chiapas. Proc. Calif. Acad. Sci. 48: 

.1995b. Acanthaceae. In Flora of Chiapas, by D. E. Breedlove, pt. 4: 1-158. 

. 1995c. Revision of Odontonema (Acanthaceae) in Mexico. Contr. Univ. Michigan Herb. 20: 

147-171. 
Durkee, L. H. 1978. Acanthaceae. In Flora of Panama, ed. R. Woodson et al., Ann. Missouri Hot. 

Card. 65: 155-284. 

. 1986. Acanthaceae. In Flora Costaricensis, ed. W. Burger, Fieldiana, Bot. 18: 1-92. 

Dwyer, J. D., and D. L. Spellman. 1981. A list of the Dicotyledoneae of Belize. Rhodora 83: 161-236. 

Famighetti, R., ed. 1995. The world almanac and book of facts 1996. 

Fosberg, F. R., D. R. Stoddart, M.-H. Sachet, and D. L. Spellman. 1982. Plants of the Belize Cays. 

Atoll Res. Bull. 258: 1-77. 
Gibson, D. N. 1974. Acanthaceae. In Flora of Guatemala, ed. P. C. Standley et al. Fieldiana, Bot. 24 

(10): 328-461. 
Greuter, W. et al. 1994. International Code of Botanical Nomenclature. Regnum Veg. 131: 1-389. 
Hampshire, R. J. 1989. Belize. In Floristic Inventory of Tropical Countries, ed. D. G. Campbell and 

H. D. Hammond, 286-289. Bronx: New York Botanical Garden. 
Leonard, E. C. 1936. The Acanthaceae of the Yucatan Peninsula. Publ. Carnegie Inst. Wash. 461: 

Lindley, J. 1845. Aphelandra aurantiaca. Bot. Reg. 31: no. 12. 

Lundell, C. L. 1945. The vegetation and natural resources of British Honduras. In Plants and plant 

science in Latin America, ed. F. Verdoorn, 270-273. 
Scheidweiler, M. 1842. Notice sur quelques nouveaux generes et especes de plantes. Bull. Acad. Roy. 

Sci. Bruxelles 9: 19-26. 
Spellman, D. L., J. D. Dwyer, and G. Davidse. 1975. A hst of the Monocotyledoneae of Belize 

including a historical introduction to plant collecting in Belize. Rhodora 77: 105-140. 
Standley, P. C, et al. 1958-1977. Flora of Guatemala. Fieldiana, Bot. 24. 

Wasshausen, D. C. 1975. The genus Aphelandra (Acanthaceae). Smithsonian Contr. Bot. 18: 1-157. 
Wendt, T. 1993. Composition, floristic affinities, and origins of the canopy tree flora of the Mexican 

Atlantic slope rain forests. In Biological diversity of Mexico: origins and distribution, ed. T. P. 

Ramamoorthy et al., 595-680. 
Williams, L. O. 1981. The useful plants of Central America. Ceiba 24: 1-381. 



APPENDIX 

Overall Distribution Patterns of Acanthaceae Native i. 

Widespread (62.5%): Aphelandra aurantiaca, A. scabra; Barleria oenotheroides; 1 
turn; Bravaisia berlandieriana; Dicliptera inutilis, D. sexangularis, D. sumichra 
cata; Hygrophila costata; Justicia aurea, J. breviflora, J. candelariae, J. comt 
spicigera; Lepidagathis alopecuroidea; Mendoncia lindavii, M. retusa; Odontor 



Regional (30%): Bravaisia grandiflora; Carlowrightia myriantha; Justicia albobractea, J. bartlettii, J. 
campechiana, J. fimbriata; Louteridium donnell-smithii; Odontonema albiflorum, O. hondurense; 
Pseuderanthemum verapazense; Ruellia harveyana, R. pereducta. 



Local(5%):/im; 
Endemic (2.5%): 



A REVISION AND REDEFINITION OF PSEUD ABUTILON 
(MALVACEAE) 

Paul A. Fryxell 
Department of Botany 
University of Texas 
Austin, Texas 78713 



INTRODUCTION 

Fries (1908) defined Pseudabutilon R. E. Fr. on the basis of the presence of an 
endoglossum, an internal partition that divides the carpel into two chambers, one 
above the other. This structure, however, is now known to occur in many relatively 
unrelated malvaceous genera, including some or all species oi Allosidastrum (Hochr.) 
Krapov., Fryxell & D. M. Bates, Allowissadula D. M. Bates, Anisodontea C. Presl, 
Batesimalva Fryxell, Dirhamphis Krapov., and Gaya H. B. K., among others. The 
endoglossum of these various genera exhibits a variety of structures that are evi- 
dently not homologous, although a careful comparative anatomical study has not 
yet been undertaken, other than the discussion presented by Hochreutiner (1920) 
wherein the term "endoglossum" was proposed. Furthermore, several genera (e.g., 
Malvastrum A. Gray, Allosidastrum, Fuertesimalva Fryxell, etc.) have what is best 
regarded as a vestigial endoglossum in at least some species. Moreover, Pseud- 
abutilon as here dehmited has the endoglossum present or absent depending upon 
the species. This polymorphism, too, is known in other genera. The lack of impor- 
tance of the endoglossum as a generically defining character in Pseudabutilon is 
emphasized by its variable expression in P. depauperatum (present or absent; cf. 
Kearney, 1952; Bates, 1971, p. 370) and by its partial development in P. glomera- 
tum, where it is a reduced, fimbriate structure that is easily overlooked. These are 
simply intermediate expressions of the range from a well-developed endoglossum 
(e.g., P. callimorphum, P. orientale) to no endoglossum (e.g., P. benense, P. virga- 
tum) among species that are evidently congeneric. In addition to the distinctive 
chromosome number (x - 8, differing from most species oiAbutilon or of Wissad- 
ula, which have x = 7), Pseudabutilon as circumscribed here is characterized by 
distinctive and relatively uniform fruit morphology (sometimes including the pres- 
ence of an endoglossum); similar pubescence patterns, usually including small 
stellate hairs and sometimes long simple hairs, rarely glandular hairs; small rounded 
and ecostate calyx; and other characters including a distinctive chromosome num- 
ber. Fryxell and Stelly (1993) reviewed the cytotaxonomical problems and noted 
the need for realignment of generic boundaries within this group of species. Broader 
questions of generic limits are discussed by Fryxell (1997). 

Kearney (1951, 1955, 1958) and Hutchinson (1967, who more or less followed 
Kearney, 1951) accepted Fries's definition of the genus. They included within 
Pseudabutilon those species with an endoglossum and excluded those species that 
lack an endoglossum, leaving the latter in Abutilon. Most recent floristic works 
have conformed to this view, except Fryxell (1988), who reduced Pseudabutilon sensu 
Fries to synonymy with Abutilon. With respect to the large genus Abutilon, Kearney 
(1955) noted: "The genus is a very difficult one and much further investigation 

175 



176 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



will be required to solve all of the problems." Some of these problems involve 
also the genus Wissadula and related genera, and have been addressed by Fryxell 
(1976), Bates (1978a, 1978b), and Krapovickas et al. (1988). The present contribu- 
tion is intended as a continuation of these studies. The recognition of Pseud- 
abutilon, with the revised circumscription that follows, is intended to solve another of 
the problems alluded to by Kearney. 

Fries (1908) constituted Pseudabiitilon with two subgenera, subg. Wissadulas- 
tnim (K. Schum.) R. E. Fr. and subg. Abutilastrum (Baker f.) R. E. Fr., both 
originally treated as sections of Wissadula, but neither alluded to by Fries as 
"typical" of his new genus. The typification of these two subgenera was discussed 
by Fryxell (1988). Here Pseudabutilon will be redefined by exclusion of subgen. 
Wissadulastnim (following Bates, 1978b) and by enlarging the number of species 
included in subgen. Abutilastrum, the additional species drawn from Abutilon, 
and including the description of three new species. The species transferred from 
Abutilon are those that lack an endoglossum (Fries's defining character) but which 
are clearly congeneric with those that possess an endoglossum. The circumscrip- 
tion of the enlarged genus will thus need an amended description. Those species 
of this group that are known cytologically all have chromosome numbers of 2n = 
16, including Abutilon anderssonianum Garcke, A. ellipticum Schltdl., A. thurberi A. 
Gray, A. umbellatum (L.) Sweet, A. virgatum (Cav.) Sweet, and Pseudabutilon calli- 
morphum (Hochr.) R. E. Fr. (Bates 1966, 1976; Fernandez 1974; Krapovickas 1957). 

TAXONOMY 

Pseudabutilon R. E. Fr., Kongl. Svenska Vetenskapsakad. Handl. 43(4): 96. 1908.— 
Lectotype, designated by Fryxell, 1988: Pseudabutilon seabrum (C. Presl) 
R. E. Fr. 

Abutilon sect. Anasida Griseb., Fl. Brit. W.I. 78. 18.59.— Typi-,: Abutilon um- 
bellatum (L.) Sweet. 

Wissadula sect. Abutilastrum Baker f., J. Bot. 31:71. 1893.— Tyi'i.: Wissadula 
scabra C. Presl. 

Shrubs or subshrubs, the stems usually densely stellate-pubescent (rarely gla- 
brescent), sometimes also with long simple hairs. Leaf blades ovate to elliptic, 
basally cordate to truncate, crenate-serrate (rarely subentire), acute or acuminate, 
minutely pubescent (rarely glabrescent), sometimes with long appressed simple 
hairs, at least on upper surface; petioles much shorter than the blades to subequal 
to the blades; stipules subulate or hnear, 1-11 mm long. Flowers sohtary or glom- 
erate in the axils, or borne in small axillary umbels or racemes, sometimes aggre- 
gated into terminal inflorescences, the pedicels short or long and slender; calyx 
basally rounded, small, ecostate, about half-divided or less (rarely more deeply 
divided), stellate-pubescent and sometimes also with long simple hairs, sometimes 
with glandular hairs or with twisted subarachnoid hairs; petals yellow (sometimes 
white), 4-15 mm long; staminal column shorter than the corolla, palUd, usually 
more or less stellate-pubescent, sometimes glabrous, the filaments short, anthers 
yellow; styles 5-10, with capitellate stigmas. Fruits 5-10 mm in diameter, stellate- 
pubescent or hirsute, sometimes with twisted subarachnoid hairs; mericarps 5-10, 
apically acute to spinescent, with or without an endoglossum (of dorsal origin), 
the lateral walls sometimes striate-reticulate basally, especially in species with a 
well-developed endoglossum, 3-seeded. Base chromosome number: x = 8. 



Key to the Species of Pseudabutilon 

Upper surface of leaf with appressed simple hairs 1-2 mm long in addition to small stellate 
hairs (or stellate hairs absent); stems with long (2-4 mm) simple spreading hairs in addition 

2. Mericarps 13-14 mm long, with divergent spines 1-4 mm long; petals 8-15 mm long; 
inflorescence a terminal raceme or panicle; endoglossum absent; Mexico, Guatemala, Hon- 
duras. 8. P. ellipticum 

2. Mericarps 5-6 mm long, with spines 2-3 mm long; petals 4-9 mm long; flowers solitary in 
the leaf axils; endoglossum present. 
3. Carpels 5; petals 4-6 mm long; leaf venation more or less pedate; stem vesture lacking 

understory of stellate hairs; Mexico. 17. P. thurben 

3. Carpels 6-8; petals 7-9 mm long; leaf venation palmate; stem vesture with understory 
of stellate hairs; Argentina. U.P. longepilosum 

Upper surface of leaf stellate-pubescent or with bifurcate hairs, these never more than 0.5 

mm long, the pubescence sometimes farinose; stems minutely stellate-pubescent (or long 

spreading hairs sometimes present). 

4. Calyx (and pedicel) with minute glandular hairs; carpels usually 5, 5-6 mm long; seeds 
glabrous. 
5. Long simple hairs sometimes present on stems; mericarps with spines ca. 1.5 mm long, 

the endoglossum present; Mexico, Guatemala. 13. P. orientak 

5. Long simple hairs lacking on stems; mericarps with spines 3-3.5 mm long, the endoglos- 
sum absent; Bolivia, Paraguay. 1. P. benenst 

4. Calyx (and pedicel) usually stellate-pubescent, lacking glandular hairs; carpels 5-11, 4-11 
mm long; seeds glabrous to minutely pubescent or verruculate. 

6. Mericarps 10-11 mm long, with spines up to 2 mm long; endoglossum absent; Panama 
(Colombia?). 6. P. cymosum 

6. Mericarps 4-9 mm long, with spines obsolete or up to 4 mm long; endoglossum present 

7. Flowers short-pedicellate (less than 5 mm) or subsessile; carpels 8-10. 

8. Calyx 3-6 mm long; inflorescence racemiform or paniculate; endoglossum com- 
pletely covering the lowermost seed; spines of mericarps obsolete; Mexico, Gua- 
temala, Honduras. 15. P. scabrum 
8. Calyx 7-8 mm long; flowers solitary or inflorescence glomerulate or spiciform; 
endoglossum completely or partially covering the lowermost seed; spines of the 
mericarps 1-2 mm long; South America. 

9. Endoglossum fimbriate, partially covering lowermost seed; inflorescence glom- 
erulate or spiciform; Bolivia. 9. P. glomeratum 
9. Endoglossum completely covering lowermost seed; flowers solitary; Brazil 
(Bahia). 14. P. pintOi 
7. Flowers manifestly pedicellate (pedicels often more than 1 cm long); carpels 5-10. 
10. Stems with long spreading simple hairs. 

11. Flowers and fruits in axillary umbels; endoglossum absent; widespread. 

18. P umbellatum 
11. Flowers and fruits solitary or paired in the leaf axils; endoglossum present; 
South America. 
12. Stipules 7-12 mm long; calyx pilose and stellate-pubescent; upper leaf 

surface with minute often bifurcate hairs. 4. P. cinereum 

12. Stipules 3-5 mm long; calyx stellate-pubescent; upper leaf surface with 

10. Stems lacking long spreading hairs, except sometimes present in the inflorescence. 
13. Flowers in few-flowered axillary inflorescences, these sometimes aggregated 

into a terminal compound inflorescence; carpels 5-8. 

14. Leaves lanceolate-ovate, twice as long as wide; calyx 5-6 mm long, deep- 
ly divided, stellate-pubescent; carpels 8, with apiculum ca. 1 mm long; 
stipules 4-6 mm long; endoglossum absent; Peru. 5. P. cowani, 

14. Leaves broadly ovate; calyx 5^8 mm long, half-divided or less, hirsute; 
carpels 5-8, with divergent apical spines 1-4 mm long; stipules 8-10 mm 
long; endoglossum present or absent; Mexico, West Indies, Central and 
South America. 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



ninal inflorescence; Bolivi 



t or absent; inflorescences 
terminal inflorescences, 
ergent apical spines 2-4 n 
hairs 3-4 mm long; endoglof 



, Flowers usually solitary or in axill 

usually 2 mm long or less; South / 

17. Endoglossum absent; flowers 

gated into dense inflorescence 



Carpels 8-10. 
20. Pedicels 0.3- 


-0.4 cm long; Brazil. 


20. Pedicels 1.5- 

Carpels 5-8. 

21. Carpels 7-8 
endoglossum p 


-6 cm long; Paraguay, Argentina. 
3. P. callimorphu 

, with spine (or apiculum) less th 
resent; Paraguay, Bolivia, Argenti; 


21. Carpels 6-8, 
or absent; Gala 


, with spine 1-2.5 mm long; endog 
pagos Archipelago. 



benense (Britton) Fryxell, comb. nov. Sida benensis Britton, 
Bull. Torrey Bot. Club 16: 153. 1889. Abutilon benense (Britton) Baker f., 
J. Bot. 31: 338. 1893.— Type: Bolivia. Junction of Rivers Beni and Madre 
de Dios, Rusby 1455 (holotype: NY!; isotypes: CTES! GH! K! MICH! 
NY-2! P as photo! CTES! PH! US!). 
Abutilon balansae (Hassl.) Hassl., Repert. Spec. Nov. Regni Veg. 7: 376. 1909, 
Wissadula balansae Hassl., Bull. Herb. Boissier ser 2. 7: 458. 1907, non 
Wissadula balansae Baker f., J. Bot. 31; 69. 1893.— Type: Paraguay. San 
Bernardino, Hassler 390 (syntype: NY!); Paraguay. Sierra de Amambay, 
Esperanza, Hassler 10564 (syntypes: CTES! G!). 

Subshrubs to 1 m tall, the stems minutely stellate-pubescent. Leaf blades up 
to 9 cm long, 6.5 cm wide, ovate, basally cordate, crenate, acute or (usually) 
acuminate, minutely and obscurely stellate-pubescent above and beneath; petioles 
0.5-0.75 times as long as blade; stipules 1-1.5 (-5) mm long, subulate. Flowers 
sohtary in the axils or aggregated apically into a raceme or panicle; pedicels 1-2 
cm long, slender, viscid; calyx 5 mm long, shallowly lobed, densely viscid; petals 
6-8 mm long, yellow, pubescent on margins of claw; staminal column 2-3 mm 
long, pallid, pubescent, the filaments 1 mm long; styles 5, surpassing the androecium. 
Fruits 5-9 mm in diameter, hispid-tomentulose; mericarps 5, 3-seeded, with apical 
spines 3-3.5 mm long; endoglossum absent; seeds 2 mm long, glabrous. 

Distribution. Paraguay and Bolivia, possibly also in Venezuela. 



1997 FRYXELL: PSEUDABUTILON 179 

Additional Specimens Examined. Venezuela. Yaracuy; al pie de la Sierra de Aroa, San Felipe, 
Aristeguieta 2021 (NY). Paraguay. GuairA: Cerro Nelville, 5 km E de Mbocayaty, 25°42'S, 56°25'W, 
Schinini et al. 27898 (TEX). Bolivia. Without locality. Bang 2621 (US).— Santa Cruz: 22 km N de 
Gutierrez (camino Camiri-Santa Cruz), 19°20'S, 63°30'W, Krapovickas & Schinini 31451 (CTES, MO). 

The Venezuela specimen cited above {Aristeguieta 2021) is geographically 
disjunct from the remainder of the species and differs in having larger fruits, but is 
tentatively included here. 

2. Pseudabutilon callimorphum (Hochr.) R. E. Fr. var. callimorphum, Kongl. Sven- 

ska Vetenskapsakad. Handl. 43(4): 105. 1908. Sida callimorpha Hochr., 
Bull. Herb. Boissier, ser. 2. 5: 295. 1905. Wissadula callimorpha (Hochr.) 
Hassl., Bull. Herb. Boissier, ser. 2. 7: 455. 1907.— Type: Paraguay. Prope 
Concepcion, Hassler 7558 (lectotype, here designated: G!; isotypes: BM! 
P as photo F-35532! UC as photo! CTES!). 

Subshrubs 0.8-1.2 m tall, the stems stellate-tomentulose and with scattered 
long (2 mm) simple hairs intermingled. Leaf blades to 4.5 cm long, to 3.5 cm wide, 
ovate to broadly ovate (rarely subtrilobed), basally cordate, crenate, acute, pal- 
mateiy 5-7-nerved, with appressed stellate hairs above and beneath; petioles 1-3 
cm long, with pubescence like that of stem; stipules 3-5 mm long, narrowly linear, 
minutely stellate-pubescent. Flowers axillary, solitary; pedicels 3-5.5 cm long, slen- 
der, with pubescence like that of stem; calyx 7-8 mm long, the lobes 4 mm long, 3 
mm wide, tomentulose; petals 9-11 mm long, yellow, pubescent on margins of 
claw; staminal column ca. 2.5 mm long, the filaments 1.5-2 mm long; styles 7. 
Fruits 6-8 mm in diameter, stellate-pubescent, the mericarps 7, 7-8 mm long 
(including 1 mm apiculum), 3 mm wide, 3-seeded, divided into two cells by an 
endoglossum (one seed in lower cell, two in upper); seeds ca. 3 mm long, minutely 

Distribution. Paraguay, Bolivia, and Argentina. 

Additional Specimens Examined. Bolivia. Santa Cruz: Prov. Caballero, 5.7 km al SE de San 
Isidro, 18°4'S, 64°24'W, Solomon & Nee 17977 (MO, NY). Argentina. Chaco: Depto. Rio Bermejo, 
8 km S de Puerto Velaz, Cristobal et al. 2093 (CTES, NY); Depto. 1° de Mayo, Colonia Benitez, 
Schulz 9043 (CTES, NY); Depto. 12 de Octubre, Estancia Valverde, al W de General Pinedo, sobre 
el h'mite con Santiago del Estero, Schulz 15346 (CTES, NY).— Corrientes: Corrientes, Krapovickas 
43692 (CTES, NY). 

3. Pseudabutilon callimorphum var. intermedium Hassl, Repert. Spec. Nov. Regni 

Veg. 7: 76. 1909. —Type: Paraguay. Gran Chaco: ad ripam occidentam 
flum. Paraguay prope Santa Rita, latit. S 23°20'-23°30', Rojas [for Has- 
sler] 2362 (isotypes: G as photo F-23752! MO! NY-2! P as photo CTES!). 

Subshrubs, the stems minutely stellate-pubescent, the hairs yellowish or fer- 
rugineous, 0.1-0.3 mm long. Leaf blades 3-7.5 cm long, ca. as wide as long, broad- 
ly ovate, basally cordate, coarsely crenate-serrate, acute or acuminate, palmately 
(or sometimes pedately) 5-7-nerved, minutely pubescent above, the hairs often 
bifurcate, antrorsely oriented, ca. 0.3 mm long, minutely stellate-pubescent be- 
neath, the hairs ca. 0.3 mm long; petioles 0.5-1 times the length of the blades, with 
pubescence like that of the stem; stipules 3-5 mm long, subulate. Flowers solitary 
in the leaf axils; pedicels slender, 1.5-6 cm long, with pubescence like that of the 
stem, articulated 2-3 mm below the flower; calyx 7-8 mm long, densely and even- 
ly stellate-pubescent, the hairs coarser toward the base, ferrugineous; petals 1 cm 



OF MICHIGAN HERBARIUM VOLUME 21 

long, yellow, minutely pubescent on margins of claw, otherwise glabrous; staminal 
column 3 mm long, sparsely stellate-pubescent, the filaments 1-2 mm long; styles 
8-10, exceeding the stamens. Fruits 8-9 mm in diameter, densely and evenly 
stellate-pubescent; mericarps 8-10, ca. 7 mm tall, the apiculum 1 mm long or less, 
3-seeded, laterally smooth, each with an endoglossum; seeds 2 mm long, minutely 
and sparsely pubescent. 

Distribution. Paraguay and northern Argentina at low elevation. 

Additional Specimens Examined. Paraguay. Fortin Tte. Enciso, ruta Trans-Chaco, Schinini & 
Bordas 16484 (CTES, NY); Ruta Trans-Chaco, 21°30'S, 61°15'W, Schinini & Bordas 16505 (CTES, 
NY). Argentina. Formosa: Riacho Monte Undo, Cristobal et al. 2151 (CTES, NY). 

4. Pseudabutilon cinereum (Griseb.) Krapov., Bol. Soc. Argent. Bot. 24: 206. 

1985. Abutilon cinereum Griseb., Abh. Konigl. Ges. Wiss. Gottingen 24: 
45. 1879.— Type: Argentina. Salta: Pasage del fl. Juramento, Lorentz & 
Hieronymus 291 (holotype: GOET!; isotype: UC as photo! CTES!). 

Shrubs 0.5-2 m tall, the stems minutely stellate-puberulent and with spread- 
ing simple hairs 1-3 mm long. Leaf blades 3-8 (-13) cm long, 2.5-6 (-10) cm wide, 
ovate, basally cordate, prominently serrate, acuminate, palmateiy 5-7-nerved, con- 
colorous, densely and softly pubescent, the hairs frequently bifurcate above, stel- 
late beneath; petioles 1-3 cm long, with pubescence like that of stem; stipules 7- 
12 mm long, linear, ciliate. Flowers solitary in the leaf axils, sometimes geminate, 
often aggregated in a terminal inflorescence more or less above the leaves; pedicels 
1.5-3 cm long [or flowers and fruits subsessile in Krapovickas et al. 19142, 19281], 
with pubescence hke that of stem; calyx 7-8 mm long, half-divided, stellate-pubes- 
cent and pilose; petals 8-11 mm long, yellowish to whitish, pubescent on margins 
of claw; staminal column 2-3 mm long, the filaments 1.5 mm long; styles 7-9, 
exserted from staminal column. Fruits stellate-pubescent (and with glandular 
hairs?), 8-9 mm in diameter, exceeding the calyx; mericarps 7-9, apically apicu- 
late (apiculum 1 mm long), 6-7 mm long, 3-seeded, with an endoglossum covering 
the lowermost seed; seeds 1.5-2 mm long, minutely pubescent. 

Distribution. Bolivia and northwestern Argentina. 

Additional Specimens Examined. BoUvia. Santa Cruz: Prov. Cordillera, 7 km W de Boyuibe (20°20'S, 
63°20"W), Krapovickas & Schinini 31308 (CTES, F, NY, US).— Tarija: Ruta Tarija-Villa Montes, Bajada 
de Chimeo, Krapovickas et al. 19232 (CTES, NY); Prov. O'Connor, Puerto Margarita, Rio Pilcomayo, 
21°10'S, 63°50'W, Krapovickas & Schinini 39096 (CTES, MO). Argentina. Cata-marca: Depto. Andal- 
gala, 3 km SE of Andalgala, Cantino 774 (CTES); Depto. Helen, 11.7 km E of Rt. 40 on road to Pozo de 
Piedra, Spooner & Clausen 4602 (NY).— C6rdoba: Depto. Isehilin, Dean Funes, Cuezzo 706 (MO).— 
Chaco: Depto. 1° de Mayo, Colonia Benitez, Schidz 9204 (CTES, NY).— Formosa: Depto. Formosa, 
Riacho Monte Lindo, Cristobal et al. 2121 (CTES, NY).— La Rioja: Depto. Capital, near Carrizal, Peders- 
en 11812 (CTES).^Mendoza. Depto. Godoy Cruz, Cacheuta, O'Donell 1134 (MO).— Salta: Depto. 
Metan, Rio Juramento, Ruta 34, Krapovickas et al. 18597 (CTES, NY), Krapovickas et al. 18599 (CTES, 
NY); Metan, O'Donell 4628 (MO).— Santiago del Estero: entre Monte Quemado y Campo Gallo, 
Castiglioni-Ragonese 7144 (CTES); Depto. Rio Hondo: Los Naranjitos, Legname 105 (NY).— San Juan: 
Depto. Zonda, Sierra Alta de Zonda, Km 29 del camino a Calingaste, Nicora et al. 8449 (CTES). 

5. Pseudabutilon cowanii Fryxell, sp. nov. — Type: Peru. Amazonas: Chachapoyas, 

Nogal Cucho, camino de Balsas hacia Chachapoyas, 1830 m, 6 Nov 1984, 
Cowan, Canne & Torrel 4290 (holotype: TEX!; isotypes: CPUN! NY! 
TEX! USM!). 



1997 FRYXELL: PSEUDABUTILON 181 

Frutices dense stellato-pubescentes; laminis foliorum anguste lanceolati-ova- 
tis (duplo longioribus quam latioribus); floribus atque fructibus in umbellis axil- 
laribus paucifloribus longi-pedunculatis; calycibus profunde divisis; fructibus grosse 
hirsutis, mericarpiis 8 unumquidque apiculo 1 mm longo sine endoglosso. 

Shrubs 1 m tall, the stems densely and minutely stellate-pubescent. Leaf blades 
5-7 cm long, 2.5-3.5 cm wide, narrowly lanceolate-ovate (about twice as long as 
wide), basally cordate, crenate-dentate, acute, very finely pubescent above (the 
hairs stellate and bifurcate), densely and coarsely pubescent beneath (the hairs 
stellate and up to 0.5 mm long); petioles 2-4.5 cm long, with pubescence like that 
of stem; stipules 4-6 mm long, subulate, minutely pubescent, persistent. Flowers 
and fruits borne on axillary few-flowered umbels, the umbels subtended by an 
obsolete leaf and stipules; peduncles up to 7 cm long, the pedicels 1-1.5 cm long; 
calyx [in fruit] 5-6 mm long, deeply divided, the lobes 2-3 mm wide (or narrower); 
flowers otherwise unknown. Fruits 6-7 mm long, 7-9 mm in diameter, densely and 
coarsely hirsute (the hairs mostly 0.5 mm long); mericarps 8, apiculate, the apicu- 
lum ca. 1 mm long, 3-seeded, lacking an endoglossum; seeds 2 mm long, sparsely 
and minutely pubescent. 

Distribution. Known only from the type collection at 1800 m elevation. 

The new species is distinctive for its long-pedunculate inflorescences, and for 
its relatively deeply divided calyx with narrow lobes. Its shows a resemblance to 
P. umbellatum in its umbellate inflorescence, to P. cymosum and P. virgatum in its 
narrowly lance-ovate leaves, but differs from each of these as indicated in the key. 
The specific epithet is chosen to honor Clark P. Cowan, collector of the type 



Veraguas, Seemann 1628 (lectotype, designated by Robyns, 1966: K!). 

Shrubs 1-2 m tall, the stems stellate-pubescent. Leaf blades up to 9 cm long, 
ovate (ca. twice as long as wide), basally cordate, serrate, acuminate, somewhat 
discolorous, stellate-pubescent; petioles up to 3 cm long, stellate-pubescent; stipules 
subulate, ca. 3 mm long. Flowers and fruits in terminal racemes or panicle; pedicels 
5-12 mm long; calyx ca. 7 mm long, stellate-pubescent; petals ca. 8 mm long, 
yellow; staminal column 4-5 mm long, stellate-pubescent, the filaments 3-4 mm 
long; styles 7-8, slender, palhd. Fruits 10-11 mm long, coarsely stellate-pubescent; 
mericarps 7-8, apically spinescent (the spine up to 2 mm), 3-seeded; seeds ca. 1.8 
mm long, glabrous or papillose. 

Distribution. Known only from the two collections cited, presumably from 
intermediate elevations. 

. Colombia, "de Pandi a Fusagasuga," Goudot s.n. (syntype: P 

In the original description, Triana and Planchon noted a similarity to Abutilon 
umbellatum, here treated as Pseudabutilon umbellatum. This view was reiterated 
by Kearney (1958). However, the relatively large fruits and narrow leaves (broad- 
er in the Goudot specimen) indicate a closer alHance to P. ellipticum, in my 
opinion. Robyns (1966) recognized the species as distinct, but cited no specimens 
other than the type. No modern collections are known. 



182 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

7. Pseudabutilon depauperatum (Hooker f.) Kearney, Madrono 11: 287. 1952. 

Sida depauperata Hooker f., Trans. Linn. Soc. 20: 232. 1847. Abiitilon 
depauperatum (Hooker f.) Andersson ex B. L. Rob., Proc. Amer. Acad. 
Arts 38: 173. 1902.— Type: Gai,apagos Archipelago. Charles Island, Sep 
1835, Darwin s.n. (holotype: CGE as photo ex CAS!). 
Abutilon anderssonianum Garcke in Andersson, Kongl. Svenska Vetenskapsakad. 
Handl. 1853: 230. 1855.— Type: Galapagos Archipelago. Andersson 178 
(holotype: S!; isotypes: GOET! K!). 

Shrubs or subshrubs to ca. 1 m tall, the stems minutely stellate-pubescent. 
Leaf blades 3-11 cm long, 2-8 cm wide (often smaller in stressed plants), ovate, 
basally cordate, crenate-serrate to subentire, acute to acuminate, palmately 5-7- 
nerved, minutely stellate-pubescent above and beneath; petioles 1-9 cm long (0.5- 
1 times length of blade), with pubescence like that of stem; stipules 7-10 mm long, 
1-2 mm wide, narrowly elliptic. Flowers sohtary in the leaf axils, sometimes in 
axillary umbels, sometimes these in terminal compound inflorescences with reduced 
leaves; pedicels 1-2 cm long, slender; calyx densely stellate-pubescent, 5-6 mm 
long; petals 6-9 mm long, yellow, ciliate on margins of claw; staminal column ca. 3 
mm long, stellate-pubescent, the filaments borne apically, 1-3 mm long; styles 6- 

8. Fruits 6-7 mm in diameter, densely stellate-pubescent; carpels 6-8, the carpel 
body 6-7 mm long with spine 1-2.5 mm long, the lateral wall narrowly rectangular 
(6 X 2.5 mm), basally reticulate or smooth, 3-seeded, the endoglossum sometimes 
present or more commonly absent; seeds 2 mm long, obscurely and minutely 
pubescent. 

Distribution. Confined to the Galapagos Archipelago, occurring generally at 
low elevation. 

Additional Specimens Examined. Galapagos Archipelago. Abingdon Island: Snodgrass & Heller 
847 (CAS).— Albemarle Island: Iguana Cove, Snodgrass & Heller 76 (CAS).— Barrington Island: 
Snodgrass & Heller 479 (CAS).— Charles Island: Post Office Bay, Howell 8823 (CAS); near Black 
Beach, Howell 9352 (CAS).— Chatham Island: Snodgrass & Heller 507 (CAS).— Duncan Island: 
Snodgrass & Heller 702 (NY),— Gardner Island: Snodgrass & Heller 632 (CAS).— Indefatigable 
Island (Isla Santa Cruz): 3 mi N of Academy Bay, Bowman 55 (CAS); Academy Bay, Fosherg 
44721 (MO), Schimpff 48 (MO).— James Island: James Bay, Howell 9681 (CAS).— Tower Island: 
Snodgrass & Heller 794 (CAS). 

Kearney (1952) and Bates (1971) note that this species is polymorphic and 
"closely related to if not conspecific with... Abutilon umbellatum (L.) Sweet." Bates 
finds polymorphism in fruit morphology, in that some plants have mericarps with 
a small endoglossum and mericarp lateral walls that are minutely striate basally; 
other plants have neither an endoglossum nor striate lateral walls. He concludes 
that these differences are not sufficient for the recognition of two distinct taxa. 
Kearney (1952), on the other hand, restricted P. depauperatum to those species 
possessing an endoglossum and basal reticulations, and relegated specimens with- 
out an endoglossum to Abutilon anderssonianum, which he regarded as a syn- 
onym oi A. umbellatum. Bates's view seems more reasonable and is adopted here. 

8. Pseudabutilon ellipticum (Schltdl.) Fryxell, comb. nov. Abutilon ellipticum Schlt- 
dl., Linnaea 11: 368. 1837. Sida elliptica (Schltdl.) Steud., Nomencl. Bot. 
ed. 2. 2: 577. 1840.— Type: Mexico. Michoacan: Tlalpujahua, 1830, Keerl 
s.n. (holotype: HAL!; isotype: BR!). 



1997 FRYXELL: PSEUDABUTILON 183 

Abutilon sidoides HemsL, Diagn. pi. nov. Mex. 2: 24. 1879, non Abutilon 

sidoides Dalzell & Gibson, 1861. Abutilon hemsleyamim Rose, Contr. U.S. 

Natl. Herb. 10: 123. 1906.— Type: Mexico. San Luis Potosi: Parry & Palmer 

80 (holotype: K!; isotypes: NY! US!). 
Abutilon attenuatum B. L. Rob. & Seaton, Proc. Amer. Acad. Arts 28: 104. 

1893.— Type: Mexico. Jalisco: near Lake Chapala, Pringle 4354 (holotype: 

GH!; isotypes: BM! BR! F as photo F-56142! GOET! K! MASS! MEXU! 

MIN! MO! MSC! MU! NY-2! P as photo, CTES! PH! TEX! UC! US! VT!). 
Abutilon membranaceum Baker f. ex Rose, Contr. U.S. Natl. Herb. 3: 312. 

1895.— Type: Mexico. Nayarit: Tepic, 1892, Palmer 1959 (holotype: BM!; 

isotypes: F!GH! NY! US!). 

Shrubs 1-2 (-3) m tall, the stems coarsely stehate-pubescent, the hairs 0.5 mm 
long or less. Leaf blades 5-11 cm long, about half as wide, ovate to elliptic, basally 
truncate (rarely subcordate), serrate, acute or acuminate, palmately 5-7-nerved, 
with simple appressed hairs above, stellate hairs beneath; petioles 1/4-1/2 the 
length of the blades, with pubescence like that of stem; stipules 5-9 mm long, 
subulate or narrowly lanceolate, 1 -nerved, often more or less spreading. Flowers 
in terminal racemes or panicles, these dense or open, more or less above the 
leaves; pedicels 2-15 mm long; calyx 6-8 mm long, stellate-pubescent, basally 
rounded, more than half-divided; petals 8-15 mm long, yellow or yellow-orange; 
staminal column 4-6 mm long, pallid, stellate-pubescent at the base, the filaments 
ca. 2 mm long; styles 5-7, slender, pallid. Fruits 13-14 mm long, 8-10 mm in 
diameter, coarsely stellate-pubescent, closely invested by and exceeding the calyx; 
mericarps 5-7, apically spinose, the spines divergent, 1-4 mm long, 3-seeded, 
lacking an endoglossum; seeds ca. 2 mm long, minutely squamose. 

Distribution. Generally found between 1200-2600 m elevation in Mexico, Gua- 
temala, and Honduras. 



Representative Specimens Examined. Mexico. Aguascalientes: near Aguascalientes, Rose & 
Painter 7783 (US).— Chiapas: Mpio. de Bochil, 3 km al SO de Puerto Gate, Garcia 594 (CHAPA, 
NY); Pinada, Siltepec, Matuda 1905 (ARIZ, NY, TEX).— Colima: Rancho El Jabali, 22 km NNW of 
Colima, 19°27'N, 103°42'W, Sanders et al. 11000 (NY, UCR).— Distrito Federal: Cerro de Santa 
Catarina, cerca de Santa Catarina, Rzedowski 26052 (ENCB, LL, NY); Almella, Tlaltengo, delegacion 
de Tlahuac, Ventura 3786 (ENCB, NY).— Durango: 2 mi W of Revolcaderos, Breedlove 18926 (CAS, 
NY); Cerro de los Remedios, Ciudad de Durango, Patoni 289 (MEXU, NY).— Guanajuato: Mpio. 
de Cortazar, 2 km al NW de La Gavia, Rzedowski 40946 (ENCB, lEB); Mpio. de Jerecuaro, 3 km al 
E de Jerecuaro, Rzedowski 41713 (lEB, NY).— Guerrero: Montes de Oca, Hinton et al. 11692 (NY); 
Mpio. Chilpancingo, camino a Tejocote, 17 km al S de Chilpancingo, Koch & Fryxell 83266 (CHAPA, 
NY).— Hidalgo: vicinity of Tepeji del Rio, Fryxell & Bates 2174 (BH, NY); Canada del Salitre, 
Krapovickas & Cristobal 23475 (CTES, NY); near Tula, Pringle 9689 (MEXU, NY).— Jalisco: 15-30 
km N of Mascota on road to San Sebastian, Anderson & Anderson 5957 (MICH); Sierra de Manantlan, 
en el Rancho Lamial, Calzada & Nieves 9468 (NY, XAL); E of Tizapan El Alto, ca 4 km W of 
Michoacan state line, on S shore of Lake Chapala, Fryxell & Bates 2153 (BH, CTES, ENCB, NY); 
Guadalajara, Jones 27156 (DS, POM, UC); 12-25 mi SE of Autlan, McVaugh & Koelz 904 (MICH, 
NY); near Magdalena, Pringle 4583 (MEXU, MICH, NY).— Mexico: Temascaltepec, Ocotepec, Hin- 
ton et al. 2916 (NY); Cerro de Santa Cruz, Sierra de Guadalupe, Matuda et al. 29520 (MEXU, NY); 
Mpio. de Texcoco, cerca de San Juan Ixhuatepec, Rzedowski 23296 (ENCB. LL, TEX); San Andres, 

5 km al SW de San Cristobal Ecatepec, Rzedowski 27642 (ENCB, NY).— Michoacan: Coalcoman, 
Hinton 12913 (MICH, NY); Mpio. de Zinapecuaro, SE del Lago Cuitzeo, 1-2 km al E de Coro, Koch 

6 Fryxell 77397 (CHAPA, LL, NY); Cerro Grande de Cujaruato, al SW de la Piedad, Rzedowski & 
McVaugh 555 (LL, MICH, NY); Mpio. de Morelia, Caiiada del Rfo Grande, cerca de Cointzio, 
Rzedowski 45377 (ENCB, lEB, MEXU, TEX).— Nayarit: Tepic, en 1892, Palmer 1959 (BM, F, GH, 



184 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

NY, US); Mpio. Tepic, 4 km al SW del Cuarenteno, brecha al Cora (21°26'N, 104°56'W), Tenorio et 
al. 16961 (MEXU, NY).— Nuevo Le6n: Zaragoza, Hinton et al. 23481 (TEX).— Oaxaca: Mpio. 
Tlacolula, 5 km N of Diaz Ordaz, on road to Cuajiimoloyas, Hill 1852 (NY); Mpio. San Juan Mixte- 
pec. Cerro Metate a 17 km NO de San Juan Mixtepec, Reyes 1376 (MEXU, NY).— Puebla: Cerro de 
la Yerba, vie. of San Luis Tultitlanapa, Purpus 2604 (NY); Mpio. Caltepec, Cerro El Gavilan, al SE 
de Caltepec, Tenorio & Romero de Tenorio 7734 (MEXU, NY, TEX).— Queretaro: La Trinidad, 
mina de opales, Arguelles 2007 (MEXU, NY); Mpio. de Cadereyta, 3 km al S de San Javier, Fernan- 
dez 3053 (ENCB, lEB, NY).— Sinaloa: Puerto a Tamiapa, Gentry 5840 (ARIZ); 1 mi S of El Palmi- 
to, 23°36'N, 105°5rW, Sanders et al. 4372 (NY, UCR).— Zacatecas: 55 mi N of FresniUo, 5 mi N of 
Rio Grande, Rollins & Tryon 58264 (LL). Guatemala. Chimaltenango: near Parramos, Standley 
59874 (F, NY).— Quiche: Nebaj, Skutch 1729 (NY); El Molino, 4 km from Chichicastenango, Molina 
et al. 30299 (NY).— Sacatep^quez: Cerro de la Cruz, above Antigua, Standley 63314 (F, NY).— 
SololA: 3-5 km W of Panajachel, Williams et al. 25299 (F, NY). Honduras. Morazan: Mt. Uyuca, 
Mo/ma 25979 (NY). 

9. Pseudabutilon glomeratum Fryxell, sp. nov. — Type: Bolivia. Chuquisaca: Luis 
Calvo Prov., 80 km al E de Boyuibe, 20°30'S, 62°10'W, Killeen, Vargas, & 
Mostacedo 4198 (holotype: TEX!; isotypes: LPB! MO!). 

Suffrutices dense stellato-pubescentes; laminis foliorum ovatis; floribus igno- 
tis; fructibus in fasciculis densis axillaribus quasi sessilibus; calycibus ad dimidium 
divisis; fructibus dense pubescentibus; mericarpiis 8 unumquidque apiculo 1-2 
mm longo, per endoglossum partialem fimbriatum in 2 celiulis diviso, cellula infe- 
rior lateraliter reticulata. 

Subshrubs ca. 1 m tall, the stems densely stellate-pubescent, the hairs ca. 0.5 
mm long. Leaf blades ovate, basally truncate to cordate, crenate, palmately 7-9- 
nerved, densely stellate-pubescent beneath, the hairs 0.5-1 mm long, more sparsely 
pubescent above, the hairs often bi- and trifurcate and antrorsely oriented, ca. 0.5 
mm long; petioles 1-2.5 cm long, densely pubescent like stems; stipules subulate, 
2-5 mm long, pubescent. Flowers unknown. Fruits in dense axillary clusters, some- 
times forming a terminal spike-like inflorescence; pedicels obsolete, the fruits 
essentially sessile; calyx 7-8 mm long, densely stellate-pubescent, ca. half-divided, 
the lobes triangular or somewhat acuminate. Fruits 7-8 mm in diameter, densely 
pubescent; mericarps ca. 8, ca. 6 mm long (excluding spine), divided by a constric- 
tion and also by a partial, fimbriate endoglossum into a lower and an upper cell, 
the lower cell 1 -seeded and laterally reticulate (somewhat similar to mericarps of 
Sphaeralcea), the upper cell smooth and 2-seeded, with an apical spine 1-2 mm 
long; seeds 2 mm long, sparsely verruculose. 

Distribution. Known only from two collections from Bolivia. 



ntrando por la quebrada de Nujchu, Beck 8881 (US). 

; notable for its obsolete pedicels and hence glomerulate 
s fimbriate endoglossum. 

10. Pseudabutilon ieucothrix Fryxell, sp. nov.— Type: Bolivia. Santa Cruz: Nuflo 
de Chavez Prov., Lomerio, 12 km al N de la coniunidad Las Trancas Area 
de estudio del proyecto "BOLFOR," Las Trancas-95, 16°31'13"S, 
61°50'47"W, 450 m, 13 Jun 1995, Mamani 778 (holotype: TEX!; isotypes: 



1997 FRYX 

Suffrutex 1-2 m altus, caulibus et foliis sparse et minutissime puberulentibus; 
calycibus et pedicellis fmctibusque pilis albis tortis arachnoideis; coroUis albis; 
inflorescentiis axillaribus racemiformibus apertis; mericarpiis 5, sine endoglosso, 
spinis apicalis 2 mm longis. 

Subshrubs 1-2 m tall, the stems very minutely and sparsely puberulent except 
in the inflorescence, where in addition there are scattered straight setae 2-5 mm 
long and dense curly subarachnoid whitish hairs ca. 0.5 mm long. Leaf blades up 
to 13 cm long, 11 cm wide, but gradually reduced upwards and very small (to 
bractlike) in the inflorescence, broadly ovate, basally cordate, crenate or serrate, 
short-acuminate, palmately 5-7-nerved, somewhat discolorous, with extremely 
minute stellate hairs (less than 0.1 mm diameter), those of the upper surface 
sparser than those of the lower surface; petioles 1-6 cm long, with pubescence like 
that of the stems; stipules 8-9 mm long, 1 mm wide, liguliform, very minutely 
stellate-pubescent. Flowers borne in inflorescences in the axils of the upper leaves, 
these becoming aggregated into a compound terminal inflorescence, which is made 
up of axillary racemes with subumbellate parts; pedicels 3-15 mm long, densely 
covered with fragile subarachnoid whitish hairs and with scattered straight setae 
2-5 mm long; calyx rounded at base, ca. 5 mm long, covered with subarachnoid 
hairs, less than half-divided, the lobes apiculate; corolla white, flowers otherwise 
unknown. Fruits 6-7 mm in diameter, yellowish, densely covered with whitish 
subarachnoid hairs; mericarps 5, the mericarp body 4 mm long, each with two 
spines 2 mm long, 3-seeded, lacking an endoglossum; seeds 2 mm long, obscurely 
pubescent. 

Distribution. Bolivia at elevations of 250-750 m in semideciduous forest. 

Additional Specimens Examined. Bolivia. Santa Cruz: Velasco, Parque Nacional Noel Kempff, 
1 km al S del Rio Itenez, 13°37'0"S, 60°54'0"W, 250-750 m, Toledo 28 (MO, TEX). 



11. Pseudabutilon longepilosum R. E. Fr,, Kongl. Svenska Vetenskapsakad. Handl. 
43(4): 107. 1908.— Type: Argentina. Catamarca: Puntilla de Villavil, White 
s.n. (syntype: BM!); Prov. Rioja: Stuckert 9341 (syntype: S!). 

Subshrubs, the stems minutely tomentose and with long simple spreading hairs. 
Leaf blades to 5 cm long, 3.5 cm wide, ovate or rotund-ovate, basally cordate, 
serrate, acute, palmately 7-nerved, stellate-pubescent above and beneath and with 
long appressed simple hairs on upper surface; petioles ca. 2 cm long (half length 
of blades), with pubescence hke that of stem; stipules fihform, 7-8 mm long, 
pilose. Flowers axillary, sohtary (or paired); pedicels 1 cm long (in flower) to 1.5- 
4 cm long (in fruit), with pubescence like that of stem; calyx 6-8 mm long (shghtly 
accrescent in fruit), tomentulose and pilose, the lobes 4-5 mm long, 2.5 mm wide; 
petals 7-9 mm long, whitish; staminal column 1.5 mm long, sparsely pilose, the 
filaments 1-1.5 mm long. Fruits 7 mm in diameter, whitish stellate-pubescent; 
mericarps 6-8, ca. 6 mm long, with apiculum 2 mm long, 3-seeded, divided into 
two cells by an endoglossum; seeds 1-1.5 mm long, subreniform. [Adapted from 
original description.] 



^ MICHIGAN HERBARIUM 



nigropunctulatum (Ulbr.) R. E. Fr., Kongl. Svenska Vetens- 
kapsakad. Handl. ser. 3, 24(2): 11. 1947. Abutllon nigropunctulatum Ulbr., 
Jahrb. Bot. Syst. 54 (Beibl. 117): 57. 1916.— Type: Peru. Ad viam ferream 
a Lima oppido ad Oroyam gerentem prope vicum San Bartolome dictum, 
Weberbauer 5301 (holotype: B as photo F-9281!; isotypes: F! GH! US!). 

Shrubs 1.5 m tail, the stems very minutely and sparsely farinose-pubescent 
(hairs 0.1 mm in diameter or less). Leaf blades 5-8 cm long, 2.5-4.5 cm wide, 
ovate, basally truncate to cordate, serrate, acuminate, palmately 5-7-nerved, fari- 
nose-pubescent above and beneath; petioles 1-2.5 cm long; stipules subulate, 1-2 
mm long. Flowers axillary and aggregated into terminal panicles; pedicels 1-3 cm 
long, slender, spreading; calyx 5 mm long, ca. half-divided or less, densely brown- 
ish puberulent; petals white, 5-6 mm long, glabrous except pubescent on claws; 
staminal column 4 5 mm long, the filaments short; styles 5-6. Fruits 6-7 mm in 
diameter, with pubescence like calyx; mericarps 5-6, apically apiculate (the spines 
1-1.5 mm long), 6 mm tall, 3-seeded, with prominent endoglossum separating 
lowermost seed from upper two seeds; seeds 2 mm long, obscurely pubescent. 

Distribution. Peru at elevations of 1300-2300 m. 

Additional Specimens Examined. Peru. Apurimac: Prov. Abacay: Rio Pachachaca, 20 km N of 
Abancay, Stork et al. 10524 (MO).— Lima: carr. Lima a Oroya, Km 70, Krapovickas 8260 (CTES, NY); 
San Bartolome, Lopez 3759 (CTES); Matucanos, Lopez 4261 (CTES).— Cajamarca: Prov. Cajamarca, 
road to Chilete, 5.5 km below San Pablo, Hutchison & Wright 5086 (UC, NY); Prov. Contumasa: Cascas, 
Raimondi 7573 (CTES).— La Libertad: Proy. Trujillo: Samne, Plaza Pampa, de Cevasco s.n. (CTES). 



orientale (Standi. & Steyerm.) Fryxell, comb. nov. Abutilon 
orientale Standi. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 
173. 1944. — Type: Guatemala. Zacapa: between Agua Blanca and Cum- 
bre de Chiquimula, Standley 74420 (holotype: F!). 
Abutilon demissum Fryxell, Brittonia 32: 263. 1980.— Type: Mexico. Oaxaca; 
Mpio. de Pochutla: 1-2 km al oeste de Puerto Angel, Koch & Fryxell 
78402 (holotype: CHAPA!; isotypes: BH! BM! CAS! CTES! ENCB! F! 
K! MEXU! MO! NY! TEX! WIS!). 

Arching or sprawling subshrubs to 1 m tall, sometimes scandent, the branch tips 
drooping, the stems minutely stellate-pubescent, sometimes also with long spread- 
ing hairs. Leaf blades 6-8 cm long (occasionally longer, progressively reduced 
upward), 4-7 cm wide (occasionally wider), ovate, basally cordate, crenate or 
serrulate, acuminate, with minute stellate and bifurcate hairs above, with slightly 
larger and denser hairs beneath, palmately 5-nerved; petioles ca. half the length of 
the blade on lower leaves, progressively shorter upward, with pubescence similar 
to that of stem; stipules 4-8 mm long, subulate, pubescent. Flowers usually soli- 
tary in the leaf axils; pedicels 4-15 mm long, articulated ca. 2 mm below the 
flower, with both stellate and some glandular hairs; calyx 4 mm long, basally 
rounded, shallowly 5-lobed, with both stellate and glandular hairs; petals 5-7 mm 
long, yellow; staminal column 2-3 mm long, pallid, stellate-pubescent, the fila- 
ments 1.5 mm long; styles 5 (-6), slender, glabrous. Fruits 5-6 mm long, about as 



1997 FRYXELL: PSEUDABUTILON 187 

wide, minutely stellate- and glandular-pubescent; mericarps 5 (-6), apically spines- 
cent, the spine ca. 1.5 mm long, 3-seeded, somewhat constricted basally, completely 
divided internally by a broad double endoglossum, the lower cell 1-seeded, the 
upper cell 2-seeded; seeds ca. 1.6 mm long, glabrous. 

Distribution. Western coastal Mexico and Guatemala at elevations below 200 m, 
in seasonally dry deciduous forest. 

Additional Specimens Examined. Mexico. Colima: Mpio. Manzanillo: 1.5 km al NE de Las 
Juntas de Abajo, brecha a Las Juntas de Arriba, Santana & Cervantes 1168 (IBUG).— Guerrero: 
Mpio. Petatlan: 28 km al SO de Zihuatanejo, Koch & Fryxell 82150 (CHAPA, F, NY); Mpio. Acapul- 
co: terraceria a Ocotillo, 2 km al O del Km 26, carr. Acapulco-Chilpancingo, Koch & Fryxell 82232 
(CHAPA, NY).— Jalisco: Mpio. La Huerta: Rancho Cuixmala, 19°27'N, 104°59'W, Lott et al. 4058 
(NY, UCR), Lott et al. 4146 (CAS, NY, TEX, UCR); near Estacion de Biologia Chamela, Magallanes 
3917 (MEXU, NY).— Oaxaca: Mpio. Chahuites: 2.5 mi W of Chiapas state line (W of Arriaga), 
Fryxell & Lott 3376 (CAS, MEXU, NY); Mpio. Juquila: 34 km a! NO de Puerto Escondido por la 
carretera a Pinotepa Nacional, Koch el al. 79454 (CHAPA, NY); Mpio. San Pedro Mixtepec, terrace- 
ria a San Gabriel Mixtepec, 6.5 km al N de Puerto Escondido, Koch et al. 79460 (CHAPA, NY); 
Mpio. Sta. Maria Huatulco: carretera a Salina Cruz, Koch et al. 79538 (CAS, CHAPA, NY).- 
Sinalga. Mazatlan, Gonzalez-Ortega 6785 (US). 

14. Pseudabutilon pintoi Monteiro, Portugal. Acta Biol. B, 12: 142. 1973.— Type: 

Brazil. Bahia: Cruz das Almas, Pinto 50-35 (holotypc: RBR n.v.; iso- 
types: CTES [fragment]! lAL n.v.). 

Subshrubs 0.5-1 m tall, the stems stellate-tomentulose. Leaf blades 3-5 cm 
long, 2.5-4 cm wide, ovate to broadly ovate, basally cordate, irregularly crenate, 
acute or acuminate, stellate-pubescent above and beneath, the hairs of the upper 
surface smaller and sparser; petioles 1-1.8 cm long (1/3 length of blade), densely 
stellate-tomentose; stipules narrowly linear, 5-7 mm long. Flowers and fruits axil- 
lary, solitary; pedicels (in fruit) 3-4 mm long, stellate-pubescent; calyx (in fruit) 8 
mm long, the lobes 5 mm long, 4 mm wide, yellowish tomentose; corolla, staminal 
column, and styles unknown. Fruits globose, yellowish tomentose; mericarps 10, 8 
mm long (including 1 mm spine), divided into two cells by an endoglossum (1 seed 
in lower cell, 2 in upper cell); seeds reniform, sparsely and minutely pubescent. 
[Adapted from original description.] 

Distribution. With certainty known only from the type collection, from the 
state of Bahia, Brazil. 

In describing this species, Monteiro relates it to P. callimorphum, but distin- 
guishes it in having very short pedicels and 10-parted fruits. It is also distinguished 
by its geographical isolation from other species of Pseudabutilon. On geographical 
grounds, another collection from Bahia (Rio Sao Francisco frente a Ibotirama, 
12°irS, 43°13'W, Krapovickas et al. 37884, CTES, NY) may belong here, although it 
was originally placed in P. virgatum. 

15. Pseudabutilon scabrum (C. Presl) R. E. Fr., Kongl. Svenska Vetenskapsakad. 

Handl. 43(4): 103. 1908. Wissadula scabra C. Presl, Reliq. Haenk. 2: 117. t. 
69. 1835.— Type: Mexico, sine loc, Haenke s.n. (holotype: PR!). 
Wissadula paniculata Rose, Contr. U.S. Natl. Herb. 5: 178. 1899. Pseudabutilon 
paniculatum (Rose) R. E. Fr., Kongl. Svenska Vetenskapsakad. Handl. 
43(4): 104. 1908.— Type: Mexico. Sinaloa: Ymala, 1891, Palmer 1743 (holo- 
type: US!; isotypes: BM! NY! US!). 



188 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Abutilon barrancae M. E. Jones, Contr. W. Bot. 18: 57. 1933.— Type: Mexico. 
Jalisco: Guadalajara, La Barranca, Jones 27162 (lectotype, designated by 
Fryxell, 1988: POM-192430!; isolectotypes: BM-2! NY! POM-1 93309! UC! 
US!). 

Pseudabutilon inomatum Standi. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. 
Ser. 23: 15. 1943.— Type: Guatemala. Guatemala: near Amatitlan, Stand- 
ley 61425 (holotype: F as photos F-56242! F-56243!; isotype: US!). 

Shrubs 1-2 (-3) m tall, minutely stellate-pubescent, the older branches be- 
coming glabrate. Leaf blades 8-12 cm long, 2.5-10 cm wide, ovate, basally trun- 
cate to somewhat cordate, obscurely serrate, acute to acuminate, minutely pubes- 
cent, the hairs stellate and bifurcate; petioles 1/3-1/2 as long as the blades; stipules 
ca. 3 mm long, subulate. Inflorescence racemiform or paniculate, simple or more 
often branched; flowers short-pedicellate or subsessile; calyx 3-6 mm long, ca. 
half-divided, basally rounded and enclosing the base of the fruit, the lobes trian- 
gular; petals 5-10 mm long, yellow; staminal column 3-5 mm long, sparsely pubes- 
cent to glabrous, the filaments almost as long as the column; styles 8-10, slender, 
paUid. Fruits 4-6 mm long, about as wide, stellate-pubescent; mericarps 8-10, 
usually 3-seeded, apically rounded to acute, basally constricted, sometimes later- 
ally reticulate at base, with double internal partition (endoglossum); seeds 2 mm 
long, papillate. 

Distribution. Mexico to Honduras at elevations of 200-1200 m, in seasonally 
dry deciduous forests or in more open habitats. 

Representative Specimens. Mexico. Chiapas: Mpio. Amatenango Fronlera, along Rio Cuilco 
between Nuevo Amatenango and Frontera Comalapa. Breedlove 41774 (CAS, NY); Mpio. Arriaga, at La 
Mina microwave station, Breedlove 56295 (CAS. NY); Mpio. Tuxtla Gutierrez, on road to Chicoasen, 
Fryxell & Lott 3236 (MEXU, NY).— Colima; forested hills 2-6 km SE of La Manzanilla, above Bahia 
Tenacatita, McVaugh 25042 (MICH, NY); Mpio. Manzanillo, 1.5 km al NE de Las Juntas de Abajo, 
Santana & Cervantes 1174 (IBUG, NY, TEX).— Guerrero: Mpio. La Union, 8 km al N de La Union por 
la terraceria a Coahuayutla de Guerrero, Koch & Fryxell 83119 (CHAPA, NY, TEX); Mpio. La Union, 
86.5 km al NE de Zihuatanejo camino a Cd. Altamirano, Martinez & Barrie 5479 (MEXU, NY).— 
Jalisco: 2.5 km al W de Melaque, Mpio. Cihuatlan, Cuevas et al. 555 (IBUG, NY); Mpio. Tonala: 
Barranca de San Caspar, Guerrero et al 392 (WIS); vicinity Quimixto, Mexia 1239 (BM, DS, F, US); La 
Barranca de Guadalajara, Pringle 1721 (BM, K, MICH, NY, S, TEX, US).— Nayarit: Mpio. San Bias, 
Km 19 de la autopista Tepic-Mazatlan (2r34'N, 1()5°()4'W), TcUez et al. 12703 (MEXU, NY); Mpio. 
Xalisco, Km 12 del camino de Xalisco a) Malinal (2I°37'N, H)4°58'W), Tellez & Bojorquez 12619 
(MEXU).— Oaxaca: 71.4 mi NW of Salina Cruz on Hwy 200, Fryxell & Lott 3397 (MEXU, NY); Mpio. 
San Miguel Chimalapa: entre 12 y 15 km de la carretera Juchitan-Tuxtla Gutierrez, 160-220 m, Koch & 
Fryxell 78349 (CHAPA, NY, TEX); Mpio. Ixtaltepec, al S de Mazahua, 13 km al NE de Ventosa, 16°40'N, 
94°54'W, Martinez 1260 (MEXU, NY); 5 km al E de Teotitlan del Camino, sobre la carr. a Huautla, 
Rzedowski 37072 (ENCB, NY); ladera S del Cerro de Guiengola, Torres et al. 673 (MEXU, NY).— 
Puebla: Coxcatlan, Purpus 4186 (NY).— Sinai .oa: 23 km W of San Ignacio, Fryxell & Bates 2117 (BH, 
NY); Imala, Gentry 4966 (ARIZ, NY); Cerro Tecomate, W of Pericos, Gentry 5726 (ARIZ, TEX).— 
Sonora: Mawarichi Arroyo, Rfo Mayo, Gentry 1317 (ARIZ, TEX); 2.7 mi W of Santa Fe de Yecora, 
28°23'N, 109°19'W, Goldberg 77-53 (ARIZ); Sierra de la Cebollita, 1 mi S of Nuri on the road to 
Tesopaco, 28°6'N, 109°19'W, Sanders 3703 (NY, UCR); La Higuera, ca. 7 mi N of Alamos on the San 
Bernardo road, 27°05'N, 108°57'W, Van Devender & Van Devender 90-591 (ARIZ, NY).— Verac ri:z: 
Zacuapan and vicinity, Rfo de Santa Maria, Purpus 2239 (GH, MO, NY, UC); Mpio. Dos Ri'os, Plan del 
Rio, Ventura 2655 (ARIZ, ENCB, LL, NY, TEX); Mpio. Dos Ri'os: La Cumbre, Ventura 120.12 (CHAPA, 
MEXU, NY). Honduras. El Paraiso: Sierra de la Villa Santa. Rio California, Molina el al. 31252 (NY). 

16. Pseudabutilon stuckertii R. E. Fr., Kongl. Svenska Vetenskapsakad. Handl. 
43(4): 107. 1908.— Syntypes: Argentina. Tucuman: Vipos, Lillo 2299; La 
Rioja: Diana, Schmadke 13225 (CORD); Cordoba, Stuckert 1029 (CORD, 
S); Cordoba, Stuckert 12983; Cordoba, Fielding s.n. (BM). 



PSEUDABUTILON 189 

Pseudabutilon callimorphum var. friesli (Hassl.) R. E. Fr., Kongl. Svenska 
Vetenskapsakad. Handl. 43(4): 106. 1908. Wissadula callimorpha yar.friesii 
Hassl., Bull. Herb. Boissier, ser 2, 7: 457. 1907. Wissadula pedunculata R. 
E. Fr., Arkiv Bot. 6(2): 12. t. 2. figs. 1-5. 1906.— Type: Bolivia. Gran 
Chaco, Fortin Crevaux ad Rio Pilcomayo, 16 Apr 1902, Fries 1572 (holo- 
type:S!). 

Subshrubs 0.5-1.5 m tall, the stems minutely and densely stellate-pubescent. 
Leaf blades 3-7 cm long, 2-5 cm wide, ovate, basally cordate, prominently serrate, 
acute, palmately 5-7-nerved, softly stellate-pubescent above and beneath; petioles 
1-2.5 cm long, with pubescence like that of the stem; stipules narrowly lanceolate, 
4-10 mm long, pubescent, persistent. Flowers solitary or paired in the leaf axils, 
sometimes forming a terminal inflorescence; pedicels 2-5 cm long, slender, stel- 
late-pubescent; calyx 5-7 mm long, about half-divided, densely stellate-pubescent 
and with a few pilose hairs 3 mm long; petals 8-10 mm long, yellow; staminal 
column 2-3 mm long, minutely pilose, the filaments ca. 1.5 mm; styles 6-7. Fruits 
7-9 mm in diameter, densely stellate-pubescent; mericarps 7-8, each 6 mm long, 
apically apiculate (apiculum 0.5-1 mm long or shorter), 3-seeded, with endoglos- 
sum above the lowermost seed; seeds ca. 2 mm long, minutely pubescent. 

Distribution. Paraguay, Bolivia, and northern Argentina. 

Additional Specimens Examined. Bolivia. Tarija: Prov. Gran Chaco, 10 km S de Palmar Grande, 
camino Yacuiba-Villa Monies, Krapovickas & Schinini 31102 (CTES, NY). Paraguay. Boquer6n: 
14 km N de Filadelfia, Krapovickas & Cristobal 44251 (CTES, TEX).— Chaco: Agua Duke, 20°14'S, 
60°7W, Schinini & Bordas 18082 (CTES, MO).— Nueva Asuncion: Ruta Trans-Chaco, 2r26'S, 
61°25'W, Schinini & Bordas 16407 (CTES, MO). Argentina. Catamarca: Depto. La Paz, El Belle, 
Brizuela 836 (CTES); Depto. Andagala, 9 km SE of Andagala, via the road to Cuesta de la Chilca, 
Cantino 305 (ARIZ, GH).— Chaco: Depto. Tapenaga Enrique Urien, Campo Bonazzola, Rodrigo 
2676 (NY); Depto. 1° de Mayo, Colonia Benitez, Schulz 11121 (CTES, NY); Depto. 12 de Octubre, 
Estancia San Jose, General Pinedo, Schulz 11288 (CTES).— Cordoba: Depto. Santa Maria, Cuesta 
San Roque, de la Sola 1395 (MO).— Corrientes: Depto. Mercedes, Paso Lucero, Rio Corrientes, 
Krapovickas & Cristobal 14763 (CTES).— Formosa: Depto. Pirani, Palo Santo, Maruhak et al. 423 
(CTES, NY); Depto. Pilcomayo, Laguna Vera, Morel 5007 (CAS); Depto. Patifio, Las Lomitas, 
Schinini & Pire 24214 (CTES, NY).-Junin: San Luis, Quebrada del Tigre, entre Santa Rose y 
Bariado de Cautana, Hunziker & Cocucci 14867 (CTES).— La Rioja: Depto. Gob. Cordillo, Campo 
Experimental del INTA, Biurron 1433 (CTES); entre Patquia y Capital, Dawson & Guarrera 3197 
(CTES); Depto. Rosario, Vera Penaloza, Rio Totoral, a 5 km de Chelco, Hunziker et al. 14163 
(CTES).— Salta: Depto. Guemes, Juramento, Krapovickas & Schinini 30431 (CTES, MO).— Santiago 
DEL Estero: Depto. Robles, Beltran, Garcia 719 (NY).— Tucuman: entre Las Tejas y Tucuman, 
Castiglioni & Ragonese 672 (CTES); Depto. Trancas, Vipos, Krapovickas & Cristobal 18377 (CTES, NY). 

17. Pseudabutilon thurberi (A. Gray) Fryxell, comb. nov. Abutilon thurberi A. 
Gray, Mem. Amer. Acad. Arts n.s. 5 (PI. Thurb.): 307. 1854.— Type: Mex- 
ico. Sonora: Magdalena, Thiirber 911 (holotype: GH as photo US!; iso- 
types: K! MO! NY!). 
Pseudabutilon sonorae Wiggins, Contr. Dudley Herb. 3: 70. 1940.— Type: Mex- 
ico. Sonora: along Rio de los Alisos, 31 mi S of Nogales, Wiggins 7034 
(holotype: DS!). 

Erect subshrubs (sometimes decumbent), the stems sparsely setose (the hairs 
2-3 mm long), otherwise glabrescent. Leaf blades 3-10 cm long, 3-6 cm wide, 
ovate, coarsely crenate-serrate, basally shallowly cordate, acuminate, pedately 7- 
nerved, above with widely scattered appressed simple hairs (1-2 mm) and beneath 
with even fewer such hairs (primarily on the veins), occasionally with stellate 
hairs; petioles 1/2-1 times the length of the blade, with pubescence like that of 



1 90 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Stem; stipules 3-11 mm long, 1 mm wide, linear, prominently 1-nerved, sparsely 
setose, more or less spreading, persistent. Flowers solitary in the leaf axils; pedicels 
0.5-2.5 cm long, slender, pilose, articulated ca. 2 mm below the flower; calyx 4-5 
mm long in flower, to 7 mm long in fruit, basally rounded, with minute glandular 
and a few stellate hairs and prominently setose (hairs 1-3 mm long), ca. half- 
divided, the lobes ecostate, ovate-acuminate; petals 4-6 mm long, yellowish or 
yellow-orange; staminal column 2-3 mm long, glabrous, the filaments ca. 2 mm 
long; styles 5. Fruits subequal to calyx, 5-6 mm long and wide, hispidulous, straw- 
colored at maturity; mericarps 5, usually 3-seeded, with an endoglossum, with 
scattered stellate pubescence, apically spinescent, the spine 2-3 mm long; seeds 2- 
3 mm long, sparsely stellate-puberulent. 

Distribution. Southern Arizona and northern and central Sonora at elevations 
of 900-1 100 m. 

Additional Specimens Examined. U.S.A. Arizona: Pima County, Baboquivari Canyon, 3500 ft, 
Gilman 35 (ARIZ), Kearney & Peebles 14970 (ARIZ, CAS, NY); Fresnal, Peebles 8987 (ARIZ), 
Peebles 9060 (ARIZ). Mexico. Sonora: 10 mi S of Mazatan, between Colorado and Mazatan, Wig- 
gins & Rollins 369 (ARIZ, LL, MO, NY, US); 15.3 mi by road S of Sasabe, 31.3°N, n.7°W, 900 m, 
Hastings & Turner 72-23 (ARIZ). 

In describing Abutilon thurberi, Gray did not note the presence of an endo- 
glossum, but this structure was described and illustrated by Wiggins for Pseiid- 
abutllon sonorae. Presumably, Gray merely overlooked a feature not then known 
to be significant. 

18. Pseudabutilon umbellatum (L.) Fryxell, comb. nov. Sida umbellata L., Syst. 
Nat. ed. 10. 1145. 1759. Abutilon umbellatum (L.) Sweet, Hort. Brit. ed. 1, 
1: 53. 1826.— Type: Jamaica. Browne s.n. (holotype: LINN-866.16!). 

Shrubs 1-2 m tah, the stems densely stellate-pubescent, sometimes also with 
long simple hairs. Leaf blades 6-12 cm long, broadly ovate to weakly 3-lobulate, 
basally cordate, serrate, acute or acuminate, minutely pubescent with stellate hairs 
beneath and frequently bifurcate hairs above, more densely pubescent beneath, 
palmately 7-9-nerved; petioles 1/2-1 times the length of the blade; stipules 8-10 
mm long, lanceolate. Flowers in few-flowered axillary umbels, these often aggre- 
gated into a terminal compound inflorescence; peduncles 1-5 cm long, the pedicels 
usually 1-2 cm long; calyx 6-8 mm long, basally rounded, half-divided, the lobes 
triangular, stellate-pubescent and sometimes prominently hirsute, the hairs 3-4 
mm long; petals 6-8 mm long, yellowish; staminal column 3-4 mm long, the fila- 
ments 1-2 mm long; styles 6-8. Fruits 5-9 mm long and wide, hirsute, with diver- 
gent apical spines 2-4 mm long; mericarps 6-8, 3-seeded, lacking an endoglossum; 
seeds 2 mm long, papillate. 

Distribution. Widely distributed from Mexico and the West Indies to Central 
America and northern South America, generally at relatively low elevation. 

Representative Specimens. Bahamas. Long Island: Clarence Town, Correll 48150 (FTG, LL. 
NY).— Grand Turk: Cockburn Town, Correll 46476 (FTG, NY).— Great Inagua: about Matthew 
Town, Correll 47439 (FTG, NY). Cuba. Guantanamo Bay, Oriente, Britton 2169 (NY); Loma del 
Gato, Cobre Range of Sierra Maestra, Bro. Leon et al. 9807 (NY); Santa Clara, Maria Antonia at Rio 
Banao, Ekman 16223 (NY, S). Haiti. Massif des Matheux, St. Marc, Cap St. Marc, Ekn 
(NY, S); Depto. de I'Artibonite, vicinity of Gros Morne, Leonard 9964 (NY). " • • ' 



lyy/ FRYXELL: PSEUD ABUTILON 191 

Prov. Azua: entre la entrada a Puerto Tortuguero y Hatillo, 18°25'N, 1Q°2TV^, Garcia et al. 66 (NY, 
TEX); Prov. Peravia, NE of Galeon, 18°19N, 70°14'W, Mejia & Zanoni 7008 (NY, TEX); Prov. Azua' 
3 km W of Cruce de Ocoa, 18°2rN, 70°29'W, Thompson et al. 9397 (CM, NY). Jamaica, vicinity of 
Kingston, Britton & Hollick 1725 (NY); Port Royal, Proctor 31091 (LL, NY); Clarendon, 1 mi SE of 
Portland Cottage, Proctor 31131 (LL); Lititz, 17°55'N, 77°36'W, Aug 1847, Wolle s.n. (CM, 
NY). Puerto Rico. Island of Culebra, Britton & Wheeler 136 (NY); Mona Island, Britton et al. 1818 
(NY); Coamo Springs, Britton & Britton 9478 (NY); Maruca, Guanica, Liogier et al. 33666 
(NY). Virgin Islands. St. Thomas: near Charlotte Amalie, Rose 3176 (NY, US).— St. Croix: Are- 
ria's Hope, Thompson 193 (NY).-~Virgin Gorda: Valley, Fishlock 183 (NY), Fishlock 264 (NY).— 
Tortola: Reef Island, Britton & Shafer 870 (NY). Lesser Antilles. Grenada: Grand Anse Dec 
1904, Broadway s.n. (NY); Carriacou, Howard 10827 (NY).-St. Barthelemy: Camaruche, Questel 
731 (NY). Trinidad. W end of Monos Island, Britton et al. 2722 (NY); Chachacare, 25 Jan 1921, 
Broadway s.n. (NY). Curasao. Soto, Arnoldo-Broeders 3645 (NY); without locality, Boldingh 4811 
(NY); Banks, Patrick, Britton & Shafer 3068 (NY). Mexico. Jalisco: Sierra de Manantlan, 2-4 km E 
of Zapotillo, 19°38'20"N, 104°24-25'W, Cochrane et al. 12312 (WIS); barranca near Guadalajara, 
Rose & Painter 7355 (US).— Nayarit: Km 2(M0 del camino de terraceria hacia la Presa de Aguamilpa, 
Tellez 11193 (MEXU, NY).— Oaxaca: 74 mi SE of Oaxaca, Fryxell 1148 (CAS, CTES, NY, TEX); 
Mpio. Ixtaltepec, 25 km al N de Juchitan, Koch et al. 78289 (CHAPA, NY).— San Luis Potosi: Mpio.' 
de Rioverde, S of San Diego near Presa San Diego, Fryxell 3789 (NY).— Tamaulipas: Mpio. San 
Fernando: 5-10 km S of Carboneras on road to Punta Piedras, Fryxell 3664 (CHAPA, ENCB, K, 
MEXU, MICH, MO, NY, TEX, US); Mpio. Soto La Marina, 27 km E of Soto La Marina on road to 
La Pesca, Fryxell 3686 (CHAPA, CTES, ENCB, MEXU, MICH, MO, NY, TEX, US); Mpio. Aldama: 
ca. 40 km NE of Aldama near Barra de Tordo, Fryxell 3695 (CHAPA, ENCB, GB, MEXU, MICH, 
MO. MSC. NY, TEX, US).— Veracruz: 9 km de Tampico hacia Cd. Valles, Chiang 269 (MEXU); 
near Puente Nacional Fryxell & Bates 858 (BH, CTES, NA, NY); Rancho Remudadero, Piirpus 
14296 (A, F, NY, UC); Mpio. Puente Nacional: Pachuquilla, Ventura 9425 (ENCB, MEXU, NY).— 
Yucatan: Cueva de Sihunchen, 15 mi N of Muna, Butterwick 131 (LL). Nicaragua. Granada: 8 km 
N de Granada on road to Paso de Panaloya, Stevens 4623 (MO, NY).— Managua: Km 14 carr. a 
Laguna de Jiloa, 12°10'N, 86°21'W, Moreno 3566 (MO, NY); lado O de Laguna de Jiloa, Sandino 265 
(MO, NY).— EsTELi: sobre el camino a Limay, 13°12'N, 86°36'W, Moreno 2014 (MO, NY); on road to 
Pueblo Nuevo, 13°25'N, 86°25'W, Stevens 5747 (MO, NY).— Matagalpa: SW slopes of Cerro El 
Pilon, 12°37'N, 86°02'W, Stevens 9430 (MO, NY).— Leon: Lago de Managua, Isla Momotombito, 
Araquistain & Moreno 1057 (MO, NY); El Transito, Estero El Guineo y alrededores, 12°03'N, 86°42'W, 
Moreno 4797 (MO, NY).— Rivas: 18 km al S de San Juan del Sur, Araquistain & Moreno 1264 (MO, 
NY); Peiias Blancas, 1 km N of Costa Rican boundary, Atwood 1808 (MO, NY). Venezuela. Isla 
Margarita, Miller & Johnston 152 (NY).— Aragua: Maracay, Krapovickas 15571 (CTES, NY).— 
Falc6n: Coro, Wingfield 5033 (CORO, NY).— Bolivar: Ciudad Bolivar, Hoh & Gehringer 17 (NY).— 
DiSTRiTO Federal: Pto. La Cruz, Hacienda Panarigua, Pittier 11652 (NY).— Cojedes: Depto. El Pao, 
Paraima, Trujillo 5438 (MY, NY).— Lara: Depto. Palovecino Agua Viva via Terepaima, Burandt 
2434 (NY); Barquisimeto, Saer 207 (NY).— Mi^rida: Distr. Sucre, Las Gonzales-San Jose road, ca. 20 
km SW of Merida, 8°25'N, 7P20'W, 9-10 Nov 1990, Dorr & Barnett 7653 (NY, TEX).— Miranda: SO 
del Valle de Caracas, 10°30'N, 66°53'W, Ramirez 1049 (NY).— Zulia: Jardin Botanico de Maracaibo, 
Bunting 13108 (NY). Colombia. Bolivar: vicinity of Turbaco, Killip & Smith 14213 (NY).— Magdale- 
na: 1 km E of Bonda along road from Santa Marta to Rio Hacho, in5'N, 74°06'W, Kirkbride 2492 
(NY); Santa Marta, Smith 490 (LL, NY). Norte de Santander: between Chinacota and La Esmer- 
alda, Killip & Smith 20924 (NY). Ecuador. El Oro: Santa Rosa, Asplund 15707 (S); Huaquillas- 
frontier of Peru, Jaramillo & Narvdez 784 (QCA).— Guayas: 1 km NW of Chanduy towards Atahualpa, 
Holm-Nielsen 2154 (AAU, F, GB, MO, S); 20 km S of Manglaralto, near Palmar, Holm-Nielsen et al. 
2483 (AAU, F, GB, NY); Playas, Mille 166 (F, OCA, US); La Puntilla, Salinas, Svenson 11253 (GH, 
NY, UC, US).— Loja: 10 km W of Vilcabamba, //ar/mg & Andersson 21774 (QCA).— Manabi: El 
Recrco, Eggers 15778b (F, K); Portoviejo, Mille 1963 (F). Peru. Cajamarca: 2 km E of Chilete, 



19. Pseudabutilon virgatum (Cav.) Fryxell, comb. nov. Sida virgata Cav., Icon. 1: 
53. t. 73. 1791. Abutilon virgatum (Cav.) Sweet, Hort. Brit. ed. 1. 1: 53. 
1826.— Type: in Hort. Bot. Madrid ex Peru (holotype: MA-476302!; iso- 
types: CTES [fragment]! K! P as photo F-35549!). [The type is noted by 
Garilleti (1993: 141).] 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 



Sida grevilleana Gillies ex Hook, f., Bot. Misc. 3: 154. 1832. Abutilon grevil- 

leanum (Gillies ex Hook, f.) Walp., Repert. Bot. Syst. 1: 324. 1842.— Type: 

Argentina. Mendoza: Gillies 139 (holotype: K as photo CTES!; isotypes: 

G as photo F-8000! GH! OXF!). 
Abutilon mendocinum Phil., Anal. Univ. Chile 36: 164. 1870.— Type: Argentina. 

Mendoza: Philippi 1874 (isotypes: GOET!, W as photo F-32633!). 
Abutilon paranthemoides Griseb., Goett. Abh. 24: 46. 1879.— Type: Argentina. 

Catamarca: Quebrada de Choya, Schickendantz 208b (holotype: GOET!). 
Abutilon bridgesii Baker f., J. Bot. 31: 338. 1893.— Type: Bolivia. 1847, Bridges 

s.n. (BM! CTES [fragment]! K!). 
Abutilon saltense Hassl., Repert. Nov. Sp. Regni Veg. 12: 498. 1913.— Type: 

Argentina. Salta: in dumento Rosario de la Frontera, Lillo 3877 (LIL 

n.v., "et herb. Hassler" G?). 

Shrubs 0.5-2 m tall, the stems moderately to densely stellate-pubescent. Leaf 
blades 3-8 cm long, 2-5 cm wide, ovate or weakly 3-lobed, basally truncate to 
cordate, coarsely crenate-serrate, acute, palmately 7-nerved, softly and minutely 
stellate-pubescent, more densely so beneath, with often bifurcate hairs above; 
petioles 1/3-1 times as long as the blade, with pubescence like that of the stem; 
stipules 4-10 mm long, linear, pubescent. Flowers solitary or fasciculate in the leaf 
axils, sometimes in terminal racemes, with the leaves much reduced; pedicels 1-35 
mm long, pubescent; calyx 5-8 mm long, ca. half-divided, densely stellate-pubes- 
cent, sometimes with long (1-2 mm) simple hairs in addition, especially at base; 
petals 4-8 mm long, yellow, pubescent on margins of claw, otherwise glabrous; 
staminal column 2 mm long, pallid, distally pubescent, the filaments 1.5 mm long; 
styles 7-9, paUid. Fruits 6-9 mm long (exceeding the calyx), 5-9 mm in diameter, 
stellate-pubescent; mericarps 6-10, with apical spines 1-2 mm long, 3-seeded, with- 
out an endoglossum; seeds 2 mm long, sparsely and minutely pubescent. 

Distribution. Ecuador, Peru, Bolivia, Paraguay, and Argentina, at elevations 
up to 2000 m. 



Additional Specimens Examined. Ecuador. Loja: 4-6 km N of Yangana, on ro 
Marling & Andersson 21650 (GB); Mollococha, ca. 10 km W of Vilcabamba, Marling & Andersson 
21774 (GB, NY). Peru. Apurimac: 15 km NW of Chalhuanca. below village of Pakayca, Gentry et 
al. 23322 (MO); SW of Abancay on road to Pachachaca, litis el al. 619 (MO).— Huanuco: 7.8 km NE 
of Huanuco at Hda. Colcuy, Bird 1092 (MO, NY). Bolivia. Cotana am Illimani, Nov 1911, Buchtien 
3215 (MO, NY).— Cochabamba: Panduro, near Rio Came, Cardenas 2434 (NY, US); Cochabamba, 
Cardenas 3784 (CAS, DS).— La Paz; Inquisivi, 4 km SE of Inquisivi, 16°55'S. 67%'W, Lewis 35093 
(MO, NY).— Tarua: Ruta Tarija-Villa Montes, 15 km E de Entre Ri'os, Krapovickas et al. 19129 
(CTES, LL); Prov. Arce. 31 km S of road to Entre Rios on road to Padayca, 21°49'S. 64°40'W, 
Solomon 10573 (MO. NY). Paraguay. Booueron: 5 km NW de Mariscal Estigarribia. Krapovickas 
& Cristobal 44374 (CTES, TEX).— Chaco: Chaco Paraguayo, F. Olimpo, Rojas 13617 (MO. TEX); 
Cerro Leon, 20°26'S, 60°15'W, Schinini & Bordas 17830 (MO).— Central: Jardin Botanico y Zoologico, 
Trinidad, Asuncion, 25°2(yS, 57°28'W, Perez 97 (AS, MO); Cerro Koi-Aregua. Schinini 4019B (CTES, 
NY). Argentina. Catamarca: Depto. La Paz, El Rio de la Dorada. Brizuela 959 (TEX): Depto. 
Ancasti, La Brea, Brizuela 1302 (MO); Depto. Andalgala, 16 km SE of Andalgala on road to Cuesta de la 
Chilca, Cantino 766 (ARIZ, GH).— C6rdoba: Depto. Union, Bell Ville, Parque Tau, Ferriicci 584 
(CTES, NY); Depto. PuniUa, San Esteban, Vellofane 443 (CAS).— Corrientes: Corrientes, Krapovic- 
kas 43693 (CTES, NY); Depto. Beron de Astrada, 46 km W de Ita Ibate, Valencia, Schinini 14051 
(CTES, F, NY).— Jujuy: Depto. Tumbaya, acceso a Purmamarca, Krapovickas & Cristobal 17636 
(CTES, NY).— La Pampa: Depto. Lihuel Calel, Sierra de Lihuel Calel, Krapovickas et al. 22625 
(CTES, NY).— Mendoza: Depto. San Rafael, Serrito, Spegazzini 22556 (NY).— Salta: Depto. Chi- 
coana, Chicoana, Los Los, Krapovickas et al 28260 (CTES, NY); Depto. Rosario de la Frontera, 



FRYXELL: PSEUD ABUTILON 

irante Brown, O'Donell 5392 (CAS, TEX).— Tucuman: Depto. Leales, 
ovickas & Cristobal 17335 (LL, NY); Depto. Trancas, Vipos, Ruta 9, i 
2 (CTES, LL, MO, NY). 



Pseudabutilon hitchcockli Ulbr., Notizbl. Bot. Gart. Berlin 1 1: 522. 1932. = Abutilon 
dispermum (Hochr.) Fryxell. 

Pseudabutilon langlassei Hochr., Annuaire Conserv. Jard. Bot. Geneve 20: 120. 
1917. - Allosidastrum hilarianum (C. Presl) Krapov., Fryxell & D. M. 
Bates. 

Pseudabutilon lozanii (Rose) R. E. Fr., Kongl. Svenska Vetenskapsakad. Handl. 
43(4): 100. 1908. = Allowissadula lozanii (Rose) D. M. Bates. 

Pseudabutilon pringlei (Rose) R. E. Fr., Kongl. Svenska Vetenskapsakad. Handl. 
43(4): 99. 1908. = Allowissadula pringlei (Rose) D. M. Bates. 

Pseudabutilon rosei R. E. Fr., Kongl. Svenska Vetenskapskad. Handl. 43(4): 102. 
1908. = Allowissadula rosei (R. E. Fr.) D. M. Bates. 

Pseudabutilon smithii Hochr., Annuaire Conserv. Jard. Bot. Geneve 20: 119. 1917. 
= Allosidastrum interruptum (DC.) Krapov., Fryxell & D. M. Bates. 

Pseudabutilon spicatum (H. B. K.) R. E. Fr., Kongl. Svenska Vetenskapsakad. 
Handl. 43(4): 98. 1908. = Briquetia spicata (H. B. K.) Fryxell. 

Pseudabutilon weberbaueri Ulbr., Jahrb. Bot. Syst. 54 (Beibl. 117): 60. 1916.— 
Type: Peru. Cajamarca. Prov. Jaen: Tal des Flusses Shumba, eines rechten 
Nebenflusses des Chinchipe, 5°30'S, Weberbauer 6169 (holotype: B?; iso- 
types: CTES fragment! GH!). = Tetrasida serrulata Fryxell & Fuertes, nom. 
superfl.; correct name: Tetrasida weberbaueri (Ulbr.) Fryxell & Fuertes, 



Sida dictyocarpa Morong & Britton, Ann. Nt 
nudum, non Sida dictyocarpa F. Mu 
carpa Griseb. ex K. Schum. (1891). 



LITERATURE CITED 

es, D. M. 1966. Chromosome numbers in the Malvales. I. Gentes Herb. 10: 39-46. 

. 1971. Malvaceae. In Flora of the Galapagos Islands, ed. I. L. Wiggins and D. M. Porter, . 

694. Stanford: Stanford University Press. 
. 1976. Chromosome numbers in the Malvales. HI. Miscellaneous counts from the Byttneria 

— . 1978a. Bastardiastrum, a segregate from Wissadiila (Malvaceae). Gentes Herb. 11: 311-32 
— . 1978b. Allowissadula, a new genus of North American Malvaceae. Gentes Herb. 11: 329-; 



194 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Fernandez, A. 1974. Recuentos cromosomicos en Malvaceas. Bol. Soc. Argent. Bot. 15: 403-410. 

Fries, R. E. 1908. Entwurf einer Monographie der Gattungen Wissadida und Pseudahutilon. Kongl. 
Svenska Vetenskapsakad. Hand). 43(4): 1-114 + 10 plates. 

Fryxell, P. A. 1976. New species and new combinations in BriqueUa and Hochreutiiwra. and a discus- 
sion of the Briquetia generic alliance (Malvaceae). Brittonia 28: 318-325. 

. 1988. Malvaceae of Mexico. Syst. Bot. Monogr. 25: 1-522. 

. 1997. The American genera of Malvaceae— II. Brittonia 49: 204-269. 

Fryxell, P. A., and D. M. Stelly. 1993. Additional chromosome counts in the Malvaceae. Sida 15: 639- 
647. 

Garilleti, R. 1993. Herbarium Cavanillesianum. Fontqueria 38: 1-248. 

Hochreutiner, B. P. G. 1920. Organes carpiques nouveaux ou meconnus chez les Malvacees. 
Annuaire Conserv. Jard. Bot. Geneve 21: 347-387. 

Hutchinson, 1. 1967. Malvaceae. In The genera of flowering plants by J. Hutchinson, 2: 536-567. 

Kearney, T. H. 1951. The American genera of Malvaceae. Amer. Midi. Natural. 46: 93-131. 

. 1952. Notes on Malvaceae III. Abutilon and Pseudahutilon in the Galapagos Islands. Madrono 

. 1955. A tentative key to the North American species of Abutilon Miller. Leafl. W. Bot. 7: 

. 1958. A tentative key to the South American species of Abutilon Miller. Leafl. W. Bot. 8: 

Krapovickas, A. 1957. Niimeros cromosomicos de Malvaceas americanas de la tribu Malveae. Rev. 

Agronom. Noroeste Argent. 2: 245-260. 
Krapovickas, A., P. A. Fryxell, and D. M. Bates. 1988. Allosidastrum, un nuevo genero de Malvaceae 

de los neotropicos. Bol. Soc. Bot. Mexico 48: 23-34. 
Robvns. A. 1966. Flora of Panama. Family 115. Malvaceae. Ann. Missouri Bot. Card. 52: 497-578. 



INDEX TO SCIENTIFIC NAMES 



iitton) Baku t 17S spicata (H. B. K.) Fryxell 193 

ikei 1 192 Pseudabutilon 

,nsch ISO benense (Britton) Fryxell 178 

1 1 ina cV Planch 1 <S0 callimorphum (Hochr.) R. E. Fr. var. 

rwcll 186 callimorphum 179 

urn (Hook t ) Andersson e\ B L. calHmorphum var. friesii (Hassl.) R. E. I 



1 Hassl. ; 



nistlMuuim RosL 1 Si 




cymosum (Triana & Planch.) Fryxell 181 


uubnuHuulm\U\.^^^i .x Rose IS^^ 




depauperatum (Hook, f.) Kearney 182 
ellipticum (Schltdl.) Fryxell 182 


'^lopmnlulanim I'lbi 18(i 




glomeratum Fryxell 184 


icniak Slandl c^Skxum 1S6 






uuntlumouh'.^ms^b 192 




inornatum Slandl. & Steyerm. 188 


'luns, Hassl 192 




lar^glassei Hochv.m^ 








douhs Hunsl IS^i 




longepilosum R. E. Fr. 185 








idHlUimn, (1 ) Sweet 190 




nigropunctulatum (Ulbr.) R. E. Fr. 186 


I'^auun (( a\ ) Sweet 191 




orientale (Standi. & Steyerm.) Fryxell 186 


isidaslium 




paniculatum (Rose) R. E. Fr. 187 


lananum(r Picsl) Kiapov Fiyxell 


&D 


pintoi Monteiro 187 






pringlei (Rose) R. E. Fr. 193 


teiruptum(DC ) K.apox FiyxcllA. 


D M. 


rosei R.E.FrA93 


Bates 193 




scabrum (C. Presl) R. E. Fr. 187 



(H. B. K.) R. E. Fr. 
R. E. Fr. 188 

;A. Gray) Fryxell 1^ 

jm (L.) Fryxell 190 

(Cav.) Fryxell 191 

Ulbr. 193 



semilata Fryxell & Fuertes 193 
weberbaueri (Ulbr.) Fryxell & Fuerl 
Wissadula 

balansae Hassl. 178 
balansae Baker f. 178 
callimorpha (Hochr.) Hassl. 179 
callimorpha var. friesii Hassl. 189 
paniculata Rose 187 
peduncidata R. E. Fr. 189 



NEW SPECIES AND NEW RECORDS FOR THE 
MALVACEAE OF BAJA CALIFORNIA SUR 

Paul A. Fryxell 
Department of Botany 
University of Texas 
Austin, Texas 78713 

Jose Luis Leon de la Luz 

Miguel Dominguez L. 

Centro de Investigaciones Biologicas de Baja California Sur 

Apartado Postal 128 

La Paz, Baja California Sur, Mexico 



INTRODUCTION 

The tropical dry forest of Baja California Sur is in one of the driest areas in 
which this subtropical type of vegetation is found (Fig. 1). It is a disjunct plant 
community, most similar to that found in the foothills and canyons of the Sierra 
Madre Occidental of southern Sonora (Shreve 1934, 1937), where it extends about 
4° of latitude farther north. Both locahties meet the Sonoran Desert at their 
northern boundaries, but the peninsular one has been developed in isolation at 
least since the last expansion of a xerothermic period, some thousands of years 
ago (Axelrod 1979). Both the Gulf of California and the desert seem to have 
played an effective role as geographic barriers since that time. This dry-season 
deciduous forest covers an approximate surface area of 1750 km^ and occupies 
foothills, hills, and middle mountain slopes in the range of 400-1000 m. The best 
expression of the community is located in the canyons of the Sierra de La Laguna; 
outside it, the composition and structure of the forest gradually degrades. This 
community has been floristically documented by several authors in the last centu- 
ry. Leon de la Luz et al. (unpubhshed data), based on their own collections, have 
been updating early information from Brandegee (1892) and Wiggins (1980); at 
present 575 species of vascular plants have been reported for this community, of 
which 42 (7%) can be considered as endemics. According to the present flora, the 
ratio of families/genera is 1/3.5, that of genera/species is 1/1.8, values that resem- 
ble ratios found in island floras. Recent collections in La Burrera Canyon, in the 
vicinity of Todos Santos, B. C. S., have brought to light two new species of Mal- 
vaceae (cf. Wiggins 1980) that are very distinct, and which are described and 
illustrated below. Following that, some range extensions and noteworthy recent 
collections of Malvaceae from the region described are presented. 

LA BURRERA CANYON 

All of the localities included in this report (except one, the report for Sida 
hyalina) come from La Burrera Canyon (23°29'N, 110°00'W), which is within the 
only tropical dry forest in all the Baja California peninsula. It is located within the 



198 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Biosphere Reserve of the Sierra de La Laguna, Baja CaUfornia Sur (see Fig. 1). 
Above the forest, a woodland community is found, where cHmate and flora are 
well dehmited by a sharp boundary in the mountain gradient. In the alluvial plains 
of the lowlands, below 400 m, the dry forest mingles with the xerophytic vegetation of 
the Sonoran Desert in a transition of gradual change over tens of kilometers. 

Families with the greatest richness prior to this report are Leguminosae, Com- 
positae, Polypodiaceae, Convolvulaceae, and Malvaceae, the last with 8 genera 
and 23 species and infraspecific taxa (no endemics recognized yet). Physiognomi- 
cally dominant species in La Burrera Canyon are Lysiloma divaricata, Pachycer- 
eiis pecten-aboriginum, Tecoma stans, Adelia virgata, Karwinskia humboldtiana, 
Jatropha vernicosa, and Albizzia occidentalis. 

According to the official Mexican climatic chart (SPP-INEGI, 1981), La Burrera 
Canyon and most of the tropical dry forest of the Sierra de La Laguna are located 
in the Koeppen BS climatic types (subtypes dry and semi-hot to semi-dry and 
semi-hot, winter precipitation less than 10% of the annual total). It is possible to 
consider a single rainy season, which goes from July to January, but two periods 
are generally recognized: September and January. Some climatic stations report 
up to 600 mm of annual precipitation, but temperature is probably the most 
important limiting factor, since in the summer soil-surface temperatures can reach 
60°C and air temperatures 45°C. Soils are derived from the decomposition of 
granitic rocks. All of them are immature without differentiated layers; the rocky 
character is variable. Landforms are basically arroyos and grassy slopes. The high- 
est diversity of plant life-forms and density of individuals is found alongside the 
arroyos. Nevertheless, in a study of endemism, Leon de la Luz et al. (1995) found 
that 75% of the endemics grow on the slopes of the range, where water and shade 
are less available than in the arroyos. 

NEW SPECIES 

Malvastrum hillii Fryxell, Leon de la Luz & Dominguez, sp. nov. — Type: Mexico. 
Baja California Sur: El Chilicote, 2 km S de Corral Gde. La Burrera, 
Todos Santos, 23°30'N, 110°02'W, 970 m, 18 Feb 1990, Dominguez L. 77 
(holotype: TEX!; isotype: HCIB!). Fig. 2. 

Frutex caulibus rubellis sparse stellato-pubescentibus; laminis foliorum ovatis 
5-8 cm longis; inflorescentiis spiciformibus; calycibus 8-13 mm longis cihatis; pet- 
alis luteis 8-10 mm longis; columna staminalis glabris, 3-4 mm longis; mericarpiis 
ca. 13 in apiculo hirsutis, cuspide apicalis brevis (0.5 mm) atque cuspidibus dorsa- 
lis duabus (1 mm), parietibus lateralis fere laevibus. 

Perennial shrubs ca. 1 m tall, the stems reddish, sparsely stellate-pubescent 
(becoming glabrescent), the hairs 3-5-armed, shortly stipitate, without preferen- 
tial orientation. Leaf blades ovate, basally truncate, coarsely crenate-serrate, acute, 
palmately 5-nerved, 5-8 cm long, 3-5.5 cm wide, the upper surface with sparse 
hairs 0.5-1 mm long that are stellate, bifurcate, or simple, the lower surface with 
only stellate hairs; petioles 1-3 cm long, with pubescence denser than that of the 
stem; stipules broadly linear, 7-8 mm long (but very early deciduous, leaving only 
scars). Inflorescence usually a terminal spiciform raceme, but the earliest flowers 
solitary in the leaf axils; pedicels solitary, 0.5-1.5 cm long, with pubescence similar 
to that of the stem, reduced upward and subsessile at the summit; bracts of the 



1997 FRYXELL. LEON DE LA LUZ & DOMINGUEZ: MALVACEAE 




PACIFIC OCEAN 



WM Tropical Decidu 
Forest 



Cobo San Lucas 



FIG. I. Collection s 



ted. Site 1, La Burrera Canyon, is a typical loc£ 
id Sidastrum burrerense were collected here, as 
irdia bivalvis. Site 2, Cerro Aguas Escondidas li 
caul scrub of the Sonoran Desert, and is the loc 



involucel 3, liguliform, 7-12 mm long, 1 mm wide; calyx 8-13 mm long, stellate- 
pubescent and ciliate on the margin; petals yellow, 8-10 mm long, glabrous; staminal 
column 3-4 mm long, pallid, glabrous; filaments apical, 1-2 mm long, the anthers 
numerous; styles ca. 13, exceeding the androecium, the stigmas capitellate. Fruits 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 




^£^— - 



1997 FRYXELL. LEON DE LA LUZ & DOMINGUEZ: MALVACEAE 201 

oblate or disciform, 7 mm in diameter, reddish brown; mericarps ca. 13, horse- 
shoe-shaped with a prominent ventral notch, 2.5 x 4 mm, with a small apical cusp 
(0.5 mm) and two dorsal cusps (1 mm), apically hirsute (hairs to 0.5 mm long), 
glabrous basally and laterally, the lateral walls nearly smooth. 

Additional Specime 
rera NE de Todos Santo: 
472 (HCIB, TEX). 

Malvastrum hillii is found in low deciduous forest at elevations of 750 to 970 
m, where it is fairly abundant. It flowers in December; fruiting follows in February. 

The specific epithet honors S. R. Hill, whose noteworthy monograph of the 
genus Malvastrum (Hill 1982) is a model of a good monograph and the principal 
source of our knowledge of the genus. 

Wiggins (1980) recorded only Malvastrum coromandelianum (L.) Garcke for 
Baja California, but Hill (1982) and Fryxell (1988) also reported M. bicuspidatum 
(S. Watson) Rose subsp. bicuspidatum for the peninsula, but farther to the north 
than M. hillii. The relationship of the new species is problematical. Geographically 
one might compare it to M. bicuspidatum (with various subspecies), but it does 
not appear to be of this affinity. In general aspect, especially in the large flowers, 
there are similarities to the Bolivian species, M. grandiflorum Krapovickas (see 
Hill 1982: 256, fig. 56). There is support for this affinity in the presence of an 
apical cusp on the mericarp, in addition to the two dorsal cusps, and in the size of 
the leaves, which are relatively large. The flowers and the fruits are also relatively 
large, but not as large in M. hillii as they are in M. grandiflorum. The mericarps of 
M. hillii do not have the prominent ribs on the lateral walls that are characteristic 
of M. grandiflorum (see Hill 1982: 254, fig. 55). Clearly, M. hillii is distinct from 
M. grandiflorum, but it is not certain if it is aUied with this species or if it is of a 
different affinity. 

Sidastrum burrerense Fryxell, Leon de la Luz & Dominguez, sp. nov.— Type: 
Mexico. Baja California Sur: "Los Gueribos," Canon La Burrera, 25 km al 
NE de Todos Santos, 23°29'N, 109°57'W, 2 Nov 1994, 550 m, R. Dominguez 
C. 7257 (holotype: TEX!; isotype: HCIB!). Fig. 3. 

Suffrutex cauhbus atque foliis sparsissime atque minutissime puberulentis gla- 
brescentibus; laminis fohorum ovato-lanceolatis, valde serratis; inflorescentiis pan- 
iculatis terminalibus pedicellis gracilis; calycibus ecostatis; petalis purpurascenti- 
bus, reflexis; columna staminalis hirsutis; mericarpiis 6-7, valde atque subtiliter 
reticulatis. 

Perennial subshrub, the stems terete, sometimes reddish, very sparsely and 
minutely puberulent becoming glabrescent. Leaf blades narrowly ovate to lan- 
ceolate, basally truncate or subcordate, with a weak tendency to be hastately 
lobed, palmately 7-nerved, coarsely and irregularly crenate-serrate, gradually acumi- 
nate, 4-9 cm long, 1.5-4.5 cm wide, sparsely and inconspicuously pubescent, the 
upper surface with simple hairs 0.3 mm long and a few stellate hairs, the lower 
surface with stellate hairs 0.2 mm in diameter dispersed uniformly; petioles 0.5^ 
cm long with pubescence similar to that of the young stems and sometimes with a 
few simple hairs to 1 mm long; stipules filiform, 7-8 mm long. Flowers solitary in the 
leaf axils or on lateral branches, forming a terminal paniculiform inflorescence, 
more or less exceeding the leaves; pedicels slender (almost capillary), 2-3 cm 



CONTR. UNIVERSITY OF f 



HERBARIUM VOLUME 21 




Sidastrum burrerense 



long, glabrous; calyx 4-4.5 mm long, minutely stellate-pubescent, ecostate, divided 
more or less to the middle; petals 5-6 mm long, reflexed, lavender or purplish; 
staminal column 4 mm long, sometimes purphsh, with prominent white hairs 0.3 
mm long; filaments apical, few (5-10), 1.5 mm long, the anthers whitish; styles 6- 
7, exceeding the androecium, the stigmas capitellate. Fruits oblate, 4.5-5 mm in 
diameter; mericarps 6-7, ca. 3 mm long, finely reticulate in the basal part, with a 
small apical rostrum, smooth; seeds solitary, ca. 2 mm long. 



FRYXELL, LEON DE LA LUZ & DOMINGUEZ: 



The type locality is described as a wet arroyo with low deciduous forest and 
riparian vegetation. Sidastrum burrerense is found between 550 to 620 m eleva- 
tion, where it is rare to abundant. It flowers in November and fruits in January. 

The specific epithet notes that this species is characteristic of the La Burrera 
Canyon. Sidastrum burrerense is the fifth species of the genus known from Mexico 
(Fryxell 1978, 1988) and is isolated from the others geographically (see the maps 
in Fryxell 1988: 411, 420). It has a very similar aspect to that of S. tehuacanum but 
has reflexed petals similar to those of S. paniculatum. The color of the corollas is 
intermediate between these species. Similarities between S. burrerense and S. 
tehuacanum include the form and dentition of the leaves, the type of pubescence, 
and the number of the mericarps. Differences include the size of the leaves (larger 
in S. burrerense than in S. tehuacanum), the pubescence of the staminal column 
(hirsute in S. burrerense, glabrous in S. tehuacanum), the size of the calyx (4^.5 
mm in S. burrerense, 2-4 mm in S. tehuacanum), the reticulation of the mericarp 
(dorsally reticulate in S. burrerense, dorsally smooth in S. tehuacanum), and char- 
acteristics of the corolla (purplish and reflexed in S. burrerense, pale rose and 
rotate in S. tehuacanum). 

Key to the Mexican Species of Sidastrum 



S. strictum (Standley) Fryxell. 
iflorescences paniculate, often with capillary pedicels; leaves narrowly lanceolate-elliptical 
ovate, glabrous or pubescent above; stems densely to sparsely pubescent or glabrous; 
irolla yellow, lavender, or purplish; mericarps 5-8. 
Leaves narrowly lanceolate-elliptical, obscurely serrate, glabrous above; corolla yellow; 
mericarps 5. S. lodiegense (E. G. Baker) Fryxell. 

Leaves lanceolate to ovate, serrate, pubescent or glabrous above; corolla lavender, pur- 
plish or yellowish; mericarps 5-8. 

3. Corolla purplish, reflexed; stems densely stellate-pubescent; leaves serrate, stellate- 
pubescent above. S. paniculatum (L.) Fryxell. 
3. Corolla lavender or yellowish, reflexed or rotate; stems sparsely pubescent to glabrate; 
leaves notably serrate, subglabrous above. 

4. Corolla lavender, reflexed; staminal column hirsute; calyx 4-4.5 mm long; mericarps 
finely reticulate laterally and dorsally. 

5". burrerense Fryxell, Leon de la Luz & Dominguez. 



1 dorsally. 



RANGE EXTENSIONS 

Recent collections in Baja California Sur have brought to light the following 
range extensions to the peninsula of Baja California of Malvaceae previously 
known only from mainland Mexico. 

1. Slda alamosana S. Watson (or S. glabra Miller?). Mpio. de La Paz, ca. Rancho 
La Burrera, 23°29'N, llO^Ol'N, 480 m, herbacea anual, margenes del arroyo, selva 
baja caducifolia, 3 Dec 1990, M. Dominguez 29 (HCIB, TEX). 



204 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

2. Sida hyalina Fryxell. Mpio. de La Paz, Cerro Aguas Escondidas, E de La 
Paz, 24°08'N, 110°08'W, 100 m, herbacea perenne, ladera rocosa, matorral sarco- 
caule, 29 Oct 1990, Leon de la Luz 4722 (HCIB, TEX). 

3. Bastardia bivalvis (Cav.) H. B. K. Mpio. de La Paz, Rancho La Burrera, 
"El Garbancillo," 23°29'N, 110°01'W, 600 m, herbacea perenne, en ladera, selva 
baja caducifolia, 2 Dec 1992, M. Dominguez 467 (HCIB, TEX). 

4. Sida acuta Burm. f. Mpio. de La Paz, Sierra de la Laguna, "El Palmillar," 
23°28'N, 110°00'W, 900 m, arroyo humedo, selva baja caducifolia, 6 Nov 1990, 
Leon de la Luz 4821 (HCIB, TEX). 

ACKNOWLEDGMENTS 

The authors are grateful to Benjamin P. Shaw for his skill in illustrating the two new species 



LITERATURE CITED 

Axelrod, D. I. 1979. Age and origin of Sonoran Desert vegetation. Occas. Papers Cal. Acad. Sci. 132: 

Brandegee, T. S. 1892. Distribution of the flora of the Cape Region of Baja California. Zoe 3: 223- 

Fryxell, P. A. 1978. Neotropical segregates from Sida L. {Malvaceae). Brittonia 30: 447^62. 

. 1988. Malvaceae of Mexico. Syst. Bot. Monogr. 25: 1-588. [Sidastrum on pp. 416-421] 

Hill, S. R. 1982. A monograph of the genus Malvastrum A. Gray (Malvaceae: Malveae). Rhodora 84: 

1-83, 159-264, 317-409. 
Leon de la Luz, J. L., A. Breceda, and R. Coria. 1995. A floristic criterion for conservation purposes. 

Poster Session of the 46th AIBS Annual Meeting, BSA Symposium, 6-10 August 1995, San 

Diego, California. 
Shreve, F. 1934. Vegetation of the northwestern coast of Mexico. Bull. Torrey Bot. Club 61: 373-380. 

. 1937. Lowland vegetation of Sinaloa. Bull. Torrey Bot. Club 64: 605-613. 

SPP-INEGI. 1981. Carta de Climas 1:1,000,000. Hoja La Paz. 

Wiggins, I. L. 1980. Flora of Baja California. Stanford: Stanford University Press. 



Contr. Univ. Michigan Herb. 21: 205-275. 1997. 



REVISION AND PHYLOGENY OF ANOMODON AND 
HERPETINEURON (ANOMODONTACEAE, MUSCI) 



inigo Granzow-de la Cerda 

University of Michigan Herbarium 

North University Building 

Ann Arbor, MI 48109-1057 



INTRODUCTION 



egregated from Neckera Hedw. by Hooker and 
n Hooker and Taylor's new genus, formerly in- 
cluded in Neckera, are A. viticulosus (Hedw.) Hook. & Taylor and A. curtipendu- 
lus (Hedw.) Hook. & Taylor, the latter transferred a year later to Antitrichia by 
Bridel (1819: 136). 

Bruch, Schimper, and Gtimbel (1851-55) assigned Anomodon, together with 
Leskea, to the family Leskeaceae, as a segregate family from the Hypnaceae, 
which still included Pseudoleskea BSG, Thuidium BSG, and Heterodadium BSG. 
Brotherus (1909) recognized five subfamilies within the Leskeaceae: Thelieae, 
Heterocladieae, Leskeae, Thuidieae, and Anomodontae. Schimper (1860) had pre- 
viously recognized the Thuidiaceae at the category of family (as "Thuidiae"), 
although he included Anomodon in the Leskeaceae ("Leskeae"), along with Myrinia 
Schimp., Myurella BSG, and Leskea Hedw. In the second edition of Pflanzenfami- 
lien, Brotherus (1925) recognized the Thuidieae, Leskeae, and Thelieae, all with 
family rank and, based on gametophytic characters, placed Anomodon in the 
Thuidiaceae. Many modern workers (Gangulee 1978; Iwatsuki 1963; Noguchi 1972, 
1974; Nyholm 1960; Smith 1978; Watanabe 1972, among others) have followed 
Brotherus's treatment by recognizing the three families, sometimes together with 
the Fabroniaceae, and also acknowledging the order Thuidiales. These authors 
include Anomodon and closely related genera [Haplohymenium Dozy & Molk., 
Herpetineuron (Mull. Hal.) Cardot, and sometimes others] within the Thuidiaceae, 
often in the subfamily Anomodontoideae (or the tribe Anomodontae). In their 
list of European mosses, Corley et al. (1981), for example, segregate Thuidiaceae 
and Leskeaceae, and include Anomodon and Haplohymenium in the former. 

Among the authors who have included Anomodon and Haplohymenium in 
the Leskeaceae (although the Thuidiaceae are segregated) are Monkemeyer (1927), 
who recognizes also the Theliaceae (close to the Fabroniaceae), Crum and Ander- 
son (1981), and also Ireland (1982), although he merged the Thehaceae with the 
Leskeaceae. The decision to recognize Leskeaceae is most likely based on the 
poorly branched leskeaceous mosses, devoid of paraphyllia, with erect and sym- 
metric capsules and an imperfect peristome, whereas the Thuidiaceae have much- 
branched stems covered with abundant paraphyllia and pluripapillose, obscure 
leaf cells. Neither family (three, if the Thehaceae are considered) seem to be very 
well defined, but the Thuidiaceae have been segregated from the Leskeaceae on 
the basis of their inclined, slightly asymmetric capsule with a perfect, hypnaceous 
peristome (Crum 1976). 



206 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Anomodon and its closely related genera do not fit dearly into either family. The 
gametophytic characters lead to the placement of these genera among the Thuidi- 
aceae, whereas the sporophytic ones suggest a closer relationship to the Leskeaceae. 
The capsules are erect and the peristome is not perfect, although in some species the 
exostome is horizontally striate at the base. The segments, on the other hand, are not 
keeled, the cilia are rudimentary or absent, and the basal membrane is no more than 3 
cells tall or, in most species, absent. These sporophytic characters and the lack of 
paraphylHa suggested to Crum and Anderson (1981) and Ireland (1982) a relationship 
with the Leskeaceae sensu stricto. According to Brotherus (1925), the profusely 
branched stems, the pluripapillose and obscure leaf cells, and the presence of horizon- 
tal striae on the exostome indicate a closer relationship with the Thuidiaceae. This 
opinion has been followed by Crosby and Magill (1978, 1981), Smith (1978), Corley et 
al. (1981), Dull (1985), and many others. Other authors, like Limpricht (1895), 
Dixon (1924), Grout (1903, 1932-1934), Bartram (1947, 1949), and Crum (1976), 
have adopted a more conservative classification, considering the Leskeaceae sensu 
lato, with the Thuidiaceae subordinated within. 

Crosby (1980), Vitt (1984), Buck and Vitt (1986) and, Buck and Crum (1990) 
propose different arrangements for the genera within the Thuidiaceae and the 
related families Leskeaceae and Thehaceae (including Anomodon and closely 
related genera). Watanabe (1972) recognized the subfamily Anomodontoideae, in 
which he included Miyabea Broth., Haplohymenium, Anomodon, and Herpetineu- 
ron. Buck and Vitt (1986) excluded the Anomodontaceae (comprising Anomo- 
don, Herpetineiiron, Thelia SulL, Myurella BSG, and Lindbergia Kindb.) from 
either family and placed them closer to the Leptodontaceae and Cryphaeaceae 
(superfamily Cryphaeacanae). The three families share the character ("homopla- 
sic") of having a costa, which distinguishes them from the Leucodontaceae and 
Hedwigiaceae. However, the straightforward rearrangement of the Thuidiaceae 
and Leskeaceae by Buck and Crum (1990) definitively removes the Anomodonta- 
ceae from the Leskeaceae alliance, and places it close to the Cryphaeaceae, in the 
Leucodontales, based on peristome morphology, presence of a creeping primary 
stem, and leaf areolation. The most representative characters of the Anomodon- 
taceae are absence of paraphyllia and presence of papillae in the lamina cells 
(probably secondarily lost in Herpetineuron and other taxa), as well as epiphytic 
and epilithic habitats. In addition, the peristome of the Leskeaceae, although 
somewhat reduced, is essentially hypnaceous and differs from that of the Anomo- 
dontaceae. Akiyama (1988), and more recently Anderson et al. (1990), present in 
their North American list a rather similar position for the family Anomodontaceae. 
They include it in the order Leucodontales, but they place Theliaceae, Leskeaceae, 
and Thuidiaceae quite close to each other within the Hypnales. The novelty within 
Anomodontaceae is the addition of Pterogonium as a fourth genus, with Haplohy- 
menium, Anomodon and Herpetineuron. Enroth (1992) included two other mono- 
typic genera in the Anomodontaceae, transferred from the Neckeraceae: Chileo- 
bryon callicostelloides (Ther.) Enroth and Bissetia lingulata (Mitt.) Broth., the 
latter species lacking papillae and with a very short costa. 

The most complete taxonomic work for the genus Anomodon is by Iwatsuki 
(1963), for the East Asiatic taxa. Iwatsuki reduced the 14 taxa then recognized for 
the region (Sakurai 1954) to 9 species and 2 subspecies. He included Anomodon 
in the family Thuidiaceae. 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 207 

The purpose of this paper is to provide a comprehensive taxonomic treatment 
for all the species in Anomodon and a phylogenetic analysis for the genus as it is 
known today, but including also the other taxa in the family Anomodontaceae, at 
least those represented by a sufficient number of collections. The revision of the 
genus Haplohymenium by Noguchi (1957) has been the starting point for the 
analysis regarding the species in this group. 

PHYLOGENETIC ANALYSIS 

Materials and Methods 

In preliminary analyses in which Haplohymenium and Herpetineuron were 
used as outgroups, Haplohymenium consistently turned out to nest within the 
ingroup. Consequently, three outgroup taxa were used: Herpetineuron toccoae 
and H. acutifolius, in addition to Pterogonium gracile, following Akiyama (1988) 
and Anderson et al. (1990), who include Pterogonium in Anomodontaceae. 

The taxa included in the analysis are all eleven species of Anomodon: [A. 
abbreviatus Mitt., A. attenuatus (Hedw.) Htib., A. dentatus Gao, A. giraldii Mull. 
Hal., A. longifolius (Brid.) Hartm., A. minor (Hedw.) Lindb., A. rostratus (Hedw.) 
Schimp., A. rugelii (Miill. Hal.) Keissl., A. solovjovii Lazarenko, A. thraustus Miill. 
Hal., A. viticulosus (Hedw.) Hook. & Taylor] and five species of Haplohymenium, 
considered as part of the ingroup [H. longinerve (Broth.) Broth., H. pseudotriste 
(Miill. Hal.) Broth., H. sieboldii (Dozy & Molk.) Dozy & Molk., H. triste (Ces.) 
Kindb., H. flagelliforme Savicz-L.]. The outgroup comprises Herpetineuron toc- 
coae (Sull.) Cardot, H. acutifolium (Mitt.) Granzow-de la Cerda, Pterogonium 
gracile (Hedw.) Sw. 

Characters were coded based on an examination of herbarium specimens; 
additional information was obtained from descriptions of Brotherus (1925), Crum 
and Anderson (1981), Grout (1932-1934), Iwatsuki (1963), Noguchi (1957), SuUi- 
vant (1864), and Watanabe (1972). Some characters were considered unordered, 
and character 11 (shape of leaf apex) was assigned a stepmatrix (see Appendix 1). 
Schematic drawings for some characters are provided in Figs. 1-7. Trials with 
different character weighting criteria were run: (1) a character scaling was applied 
in order to assign the same weight to all characters, regardless of number of states 
(base weight = 4, as characters present had 2, 3, or 5 states); (2) equal weight (= 1; 
weight becomes dependent on the number of character states). The 31 characters 
used are discussed below (Appendix 1); the data matrix is shown in Appendix 2. 
Autapomorphic characters were not included in the analyses. 

The cladistic analyses were conducted using PAUP (Version 3.0r, for the 
Macintosh computer, Swofford, 1990) on the 19 taxa and 31 characters. Heuristic 
searches were carried out using random stepwise addition option and the TBR 
(tree bisection reconnection), branch swapping option, keeping all most parsimo- 
nious (shortest) trees found. Strict consensus trees were computed for all trees 
obtained at the end of each analysis. As a measure of support to evaluate the 
relative robustness of the clades, "Bremer support" (Bremer 1988, 1994; Kallersjo 
et al. 1992) or the "decay index" (Mishler et al. 1991; Donoghue et al. 1992) was 
determined. Heuristic searches retaining trees progressively longer by one step 
were run, until their strict consensus rendered a tree that was totally unresolved. 





fi\ [^fl\ 




Results 

Three most parsimonious trees, 325 steps long, with consistency index of 0.44, 
were obtained when scahng was apphed (analysis 1). Their consensus tree is given 
in Fig. 8a. When equal unit weight was appHed, without scaling (analysis 2), four 
trees were generated, their length 121 steps and consistency index 0.45. The strict 



for analysis 2 is given in Fig. 8b, and Fig. 9 shows one of its four 
most parsimonious trees. Within the ingroup, two major clades can be distin- 
guished in these trees. One consists of the taxa A. attenuatus, A. giraldii, and A. 
longifolius, and in half of the trees A. rostratus is also included (these species 
constitute the subgenus Pseudoanomodon Limpr.) The other clade is made up of 
the remaining species of subgenus Anomodon: A. viticulosus, A. rugelii and A. 
minor (subsection Anomodon), in addition to A. abbreviatus and A. solovjovii 
(subsection Abbreviati Iwats.), A. dentatus, the species of the genus Haplohymenium 
{-A. section Haplohymenium), and A. thraustus. Anomodon rostratus appears as 
a sister species in the other half of the trees. This particular species presents an 
ambiguous position: in two of the trees is part of the basal clade (with A. attenua- 
tus, A. giraldii, and A. longifolius) and in the other two as the sister taxa for the 
Anomodon s. str. clade. The Haplohymenium clade always appears as a mono- 
phyletic clade nested within Anomodon s. str., as in all analyses performed previ- 
ously (Granzow-de la Cerda 1988b, 1989a). Whether character scaling (analysis 1) 
or equal weighting of all characters was used (analysis 2), the topologies, as well 
as consistency indices, remained rather similar. The outgroup {Herpetineuron and 
Pterogonium) always remain external. 

The values for the Bremer support, or "decay index", for each monophyletic 
group are given in the strict consensus tree of Fig. 10a. This value indicates how 
many parsimony steps must be loosened before that branch disappears (becomes 
unresolved). Figs. 10b and 10c. show trees for which parsimony has been relaxed one 
and two steps, respectively. At a length of 124 steps (parsimony relaxed by 3 steps) all 
branches collapsed, therefore no single clade had a support greater than three. 

Discussion 

Haplohymenium appeared in all cladograms as part of the ingroup, always 
between two groups of Anomodon species. The differences in the cladograms 
obtained here, as compared to the ones from a previous analyses (in which two 
species of Thelia, instead of Pterogonium gracile, were used along with Herpetineuron 
as outgroups; Granzow-de la Cerda 1988b, 1990), are the result of using Pterogo- 
nium as an outgroup, which seems a more likely close relative to the Anomodon- 
taceae (Akiyama 1988). According to these analyses, Anomodon is monophyletic 
when Haplohymenium is included within it. 

Although the consistency indices, 0. 44, and 0.45, for analyses 1 and 2 respec- 
tively, may be seen as rather poor, they are considerably larger than would be 
expected given random data for the same number of taxa and characters (Sander- 
son & Donoghue 1989; Klassen et al. 1991). Moreover, the most parsimonious topol- 
ogies are quite robust, as they basically remain unmodified under different charac- 
ter weighting procedures; the two main clades are well supported as measured by 
their decay index or Bremer support (number of steps that parsimony must be 
relaxed before a group, i.e., a monophyletic clade, disappears; Bremer 1988, 1994; 
Donoghue et al. 1992; Kallersjo et al. 1992), as shown in Fig. 10. The clades 
corresponding to the whole genus Anomodon, and to the subgenus Anomodon 
are the most robust, or best supported, as parsimony needs to be relaxed three 
steps in order to make those branches collapse). 

Taxonomic implications. Iwatsuki (1963), in his monograph of Eastern Asian 
species of Anomodon, places A. viticulosus, A. rugelii, A. minor, A. thraustus, A. 
solovjovii, and A. abbreviatus within subgenus Anomodon. This group of species, 



. UNIVERSITY OF MICHIGAN HERBARIUM 




Subgen.Pseudoanomodon 




■ 


rE 


»- 






+« ,... ■ 








+ 11 3->2 ■ 


,19 4->l 












■ 


.3.0.H1 . 


i'.'t, 




— 




;s in analysis 2, wh 
.em, except for chai 
. Proposed taxonom 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME . 

Figure 10. 



rQ 




4P^-g:: 




FIG. 10. Bremer support analysis of the four most parsimoni 
scaling). Strict consensus trees become progressively more unre 
boxed numbers denote how many steps parsimony had to be reh 
(become unresolved) as an indication of the decay index: clades 
(robust), and those marked 1 the least supported; b, tree resultinj 



s trees resulting of analysis 2 (no 
Ived as parsimony is relaxed, a, 
d to cause that clade to collapse 
arked 3 are the most supported 
-om parsimony relaxed one step; 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 



if A. dentatus (described 14 years after Iwatsuki's treatment) and Haplohymenium 
are included, coincides exactly with what I have come to call the subgenus Anom- 
odon clade. According to the analysis 2 (Figs. 8b, 9), subsection Abbreviati Iwats., 
within subgenus Anomodon, is composed of those species with a solitary high 
papilla on each leaf cell, A. abbreviatus and A. solovjovii. These two taxa appear 
in the cladograms as the sister group of the terminal clade {A. viticulosus, A. 
nigelii, and A. minor). Anomodon thraustus, now in its own section, consistently 
appears at the base of the subgenus Anomodon clade, which indicates a closer 
relationship to Haplohymenium rather than to A. minor, a taxon with which it is 
often confused. The monotypic section Auriculati Iwats. (with A. rugelii) is super- 
fluous, given that in all analyses (including preliminary ones) A. rugelii appears as 
a sister species of A. minor in a rather robust clade. Subgenus Pseudoanomodon 
Limpr. consists of A. attenuatus, type species for the subgenus, A. longifolius, and 
A. rostratus (Podpera 1954) in addition to, according to Iwatsuki (1963), A. giral- 
dii and A. acutifolius {=Herpetineuron acutifolium; see Granzow-de la Cerda, 
1989a). This subgenus coincides with the other monophyletic clade, when A. ros- 
tratus is included. Such dual positioning of A. rostratus at the base of either clade, 
however, introduces some ambiguity regarding its relationship with one or the 
other group. 

It can be inferred that according to the traditional treatment of Anomodon 
(i.e., excluding Haplohymenium), the genus is paraphyletic. Consequently, only a 
group of species of Anomodon (i.e., subsections Anomodon, Abbreviati, and Den- 
tati. Fig. 9) is monophyletic, with Haplohymenium and A. thraustus as part of a 
sister group. Therefore, in agreement with the cladistic orthodoxy, the taxonomic 
alternative available for the Anomodontaceae is to include the species of Haplo- 
hymenium within Anomodon; the thus enlarged Anomodon does form a mono- 
phyletic clade. 

Herpetineuron acutifolium (Granzow-de la Cerda 1989a) was recently pro- 
posed by Enroth (1991) as belonging to Bryonorrisia Stark & Buck. The resulting 
phylogeny, nonetheless, ratifies the close relationship between Bryonorrisia and 
Herpetineuron; H. acutifolium consistently appeared in all analyses paired together 
with H. toccoae. When Stark and Buck (1986) described Bryonorrisia {B. secunda 
Stark & Buck), they placed it in the Anomodontaceae and acknowledged a very 
close relationship to Herpetineuron. In spite of Enroth's emphasis on "the yellow- 
ish colour of the exostomes" of Bryonorrisia, I do not see relevant differences 
between these two genera and therefore regard both as one genus. The lack of 
leaf cell papillae in Herpetineuron, as well as in Pterogonium, is not sufficient for 
not considering these genera to be the closest sister group(s) of Anomodon, or 
part of the Anomodontaceae. 

Considerations about the characters. There are no real synapomorphies for the 
Herpetineuron clade, and the lack of leaf papillae is a plesiomorphy. Within the 
ingroup, the hairiness of the calyptra (character 31) is the only true synapomor- 
phic character shared by all species in the Haplohymenium clade in all analyses. 
No other character is consistently synapomorphic, although character 10 (margins 
flat or revolute) appears as a synapomorphy only in some trees. Unambiguous 
characters for this particular tree, with their corresponding transformation, are 
indicated in Fig. 9; those with a consistency index of 0.5 or over are marked with a 
black square. 



214 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Other shared characters for the Haplohymenium clade, besides the hirsute 
calyptra (character 31, Appendix 1 and Fig. 9), are the more reduced habit of the 
gametophyte (this character not included in these analyses) and the smaller leaves, 
shorter than 0.7 mm (character 8, although not an actual synapomorphy because 
it also appears in A. rostratus). These characters could explain the predisposition 
to segregate these taxa into a different genus, Haplohymenium, on the basis of 
what seems a purely phenetic criterion. The state of having the end of the costa 
obscured by laminal cells for a good part of its upper portion (character 15, state 
2), is shared by A. thraustus, A. dentatiis, and, although not so strikingly, by the 
Haplohymenium clade. However, the condition of having a delicate leaf with the 
apex broken off (excluded from the analyses represented in Figs. 8-10; its inclu- 
sion has no effect on the number of trees, their topologies, or consistency) seems 
to have appeared independently in A. thraustus, H. triste, and A. dentatus. 

Geographic distribution. The Asian Far East (Manchuria, Korea, Japan) and 
the Himalayas are the regions where Anomodon is best represented (nine spe- 
cies); only five species occur in Europe and six in the New World. 

Subgenus Pseudoanomodon clade (Fig. 9). Anomodon attenuatus and A. rostra- 
tus are not found in East Asia but occur in Europe, east to the Caucasus {A. attenua- 
tus present also in Kashmir), and in North and Central America. Both are the only 
species in the genus in the Caribbean islands and Bermuda. The ranges oi A. giraldii 
and A. attenuatus, on the other hand, apparently do not overlap, although in all 
topologies generated they appear as sister species. Anomodon longifolius has a wide 
distribution throughout Eurasia, yet it is absent from the American continent. 

Subgenus Anomodon clade (Fig. 9). Little correlation between phylogenetic 
position and distribution can be seen in this clade. Two of the terminal species, A. 
viticulosus and A. rugelii, are the most cosmopolitan; they both occur in Eurasia 
as well as in North America, but are more northern than any of the extra-Asiatic 
species. Anomodon viticulosus has been reported from Algeria and the Canary 
Islands (Dixon 1924), but these reports are most likely erroneous (for the SW 
Mediterranean distribution, see Granzow-de la Cerda, 1988a). In America, A. 
rugelii does not reach farther south than Georgia and Arkansas. I have seen 
specimens of A. viticulosus from southern Mexico; Gier (1980) also indicates its 
presence in Mesoamerica but gives no further information. Those locahties are 
probably the southernmost limit of its range. 

The other species in the section Anomodon are restricted to the Asian Far 
East: A. solovjovii from Manchuria, Henan province in eastern China, and Korea, 
and A. abbreviatus restricted also to Manchuria and Korea, as well as to Japan. 
Anomodon dentatus is endemic to Manchuria. Very different distribution patterns 
are seen in the taxa from the Haplohymenium clade. Anomodon sieboldii and A. 
longinervis are known from Eastern Asia, and A. pseudotristis occurs in South 
Africa and Madagascar, East and Southeast Asia, Australia, New Zealand. Anomo- 
don tristis is a cosmopolitan species, present in Asia, South Pacific, Europe (although 
rare), and North and Central America (U.S.A., Mexico, Costa Rica). It has been 
recorded from the Bolivian Andes under the synonym Anomodon fragillimus Herz. 
(!; Granzow-de la Cerda 1988b, 1989a; ). The Haplohymenium clade is, therefore, the 
only one mihin Anomodon with representatives in the Southern Hemisphere (includ- 
ing southern Africa, South America, Australia, and New Zealand). 

The basalmost species in the subgenus Anomodon clade, A. thraustus, shows 
a disjunct Asian-Mesoamerican distribution, quite similar to that of A. minor. 
Anomodon thraustus, however, has a much more restricted range; it is found, in 



1997 GRANZOW-DELACERDA: ANOMODONTACEAE 215 

the New World, only in Mexico, whereas A. minor is common and occurs from 
Canada to Guatemala. 

Outgroups. Within the Herpetineuron clade, H. toccoae is quite widely distrib- 
uted: South and East Asia, from the Himalayas to New Caledonia and the Philip- 
pines, south-east Africa, and throughout the American continent, but is absent in 
Europe (see Kis, 1985, and Norris & Sharp, 1961). Herpetineuron acudfolium is 
more restricted, occurring only in the Himalayas and East Asia (Iwatsuki 1963; 
Gangulee 1973; Granzow-de la Cerda 1988b, 1989a), but is also present in northern 
Mexico. It shares with A. thraustus a similar disjunct distribution pattern. 

All the species of Anomodon (s.l.) are grouped in two main clades (Fig. 9) that 
correspond to the traditional grouping into two subgenera, Anomodon and Pseudo- 
anomodon Limpr. Anomodon rostratus occupies a somewhat ambiguous position, 
because in some topologies it appears as a sister species for subgenus Anomodon. 

The genus Anomodon constitutes a monophyletic group only when species of 
Haplohymenium are included. Haplohymenium Dozy & Molk. (Musci frond, ined. 
Archip. Indici 4: 127. 1846) therefore needs to be subordinated to Anomodon, as 
supported by the cladistic analysis. All five species of Haplohymenium have been 
studied, and they form a monophyletic clade with well-defined synapomorphies. 
A change in the taxonomic category to treat these species as a section is appropri- 
ate and Anomodon section Haplohymenium (Dozy & Molk.) Granzow is here 
proposed (including A. sieboldii, the type species for Haplohymenium). Three of 
the species had previously been placed into Anomodon: A. longinervis Broth., A. 
pseudotristis (Mull. Hal.) Kindb., and A. tristis (Ces.) SuU. & Lesq. (as recognized 
by Noguchi,1957, in his revision of Haplohymenium). Haplohymenium formo- 
sanum is considered here a synonym for Anomodon tristis Ces. Two new combi- 
nations are proposed for the remaining taxa: Anomodon sieboldii (Dozy & Molk.) 
Granzow and Anomodon flagelliformis (Savicz-L.) Granzow. 

The remaining sections within subgenus Anomodon, besides Haplohymenium, 
are the monotypic section Thrausti and section Anomodon. The monotypic sec- 
tion Auriculati proposed by Iwatsuki (1963), which includes only A. rugelii, is 
superfluous (Fig. 9). On the other hand, if A. abbreviatus and A. solovjovii (which 
have been shown consistently to be monophyletic in most analyses) are to be 
grouped, the appropriate ranking, rather than at the sectional level (sect. Abbre- 
viati Iwats.), should be as subsection Abbreviati, a sister group to the subsection 
Anomodon clade, also very consistent in all analyses. Accordingly, A. dentatus, 
the basal taxon within section Anomodon (a sister species to subsections Abbre- 
viati and Anomodon, Fig. 9), must be assigned to its own subsection Dentati, in 
compliance with taxonomic orthodoxy. 

TAXONOMY 

Anomodon Hook. & Taylor, Muse. Brit. 79, pi. 3. 1818.— Typr: Anomodon 
viticulosus (Hedw.) Hook. & Taylor. 

Delicate to robust pleurocarpous mosses, forming more or less glaucous-green, 
yellowish, or brownish dense or loose mats, freely branched and irregularly pin- 
nate from a creeping stem. ParaphyUia absent. Stem leaves minute and scalehke, 
costa thick, usually pellucid, ending below the apex; secondary stem and branch 



216 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

leaves differing in size and shape. Secondary stem and branch leaves similar, 
ovate to lanceolate at base, more or less abruptly narrowed toward the middle 
(above or below), distal portion above the shoulders with parallel margins to 
tapering, sometimes narrowly so (in A. longifolius); leaf cells with one to several, 
mostly well-developed papillae on both surfaces, small, hexagonal, thin-walled 
and obscure to irregular, somewhat incrassate and pellucid (in A. giraldii), at the 
base oblong, incrassate, smooth, and pellucid. Dioicous. Perichaetial leaves differ- 
entiated, narrow and longer than vegetative leaves. Seta long, to 20 mm, flexuous 
(except for A. abbreviatus and A. solovjovii, ca. 1-2 mm), reddish. Capsule sym- 
metric, ovoid, cylindrical to oblong-cylindrical, sometimes with stomata; opercu- 
lum conic to obliquely rostrate; peristome incomplete, exostome white to pale 
brown, narrowly lanceolate, densely papillose, occasionally cross-striolate and some- 
times slightly trabeculate; endostome sometimes very reduced or absent (sect. 
Haplohymenium), usually not keeled, pale with low basal membrane, delicately 
papillose, with linear segments more or less reduced, cilia absent or nearly so. 
Calyptra cucullate, smooth or hirsute (sect. Haplohymenium). Spores 9-20 (-23) 

All species of Anomodon are dioicous. In North America, at least two, A. 
rostratus and A. attenuatus, fruit profusely. On the other hand, A. rugelii and A. 
minor fruit less abundantly and perhaps less frequently; sporophytes of A. viticu- 
losus and A. trisds are extremely rare in North America (only one fertile speci- 
men of ^. vUiciilosus seen, none oi A. trlstis). In Europe, however, A. vdiculosus 
is the most common species and often produces sporophytes. 

The species of Anomodon grow on tree trunks, including the base, and on 
logs, soil, or rock (mostly calcareous). In North America, they are an important 
element of the eastern deciduous forest. It is remarkable how often several of the 
species grow together yet remain taxonomically distinct. Three of the North Amer- 
ican species, A. attenuatus, A. rostratus, and A. minor, and sometimes a fourth, A. 
rugeUi, may grow on the same tree, each usuahy forming its own mat, but overlap- 
ping at the margins of mats. The gametophytes of several species may be inter- 
twined in mixed mats. The different species may be found growing intermixed 
only occasionally. Ahhough a high rate of sporophyte production is observed in 
some species, no hybrid sporophytes have been recorded. 

Key to the Species of Anomodon 



Plants very slender, delic; 


ale, stems <0.5 mm thici 


t when dry, le; 


aves <1.2 mn 


1 long, differenti- 


ated basal cells few, not r« 


caching the margins; calyptra hirsute. 






2. Laminal cells with nui 




anched papill; 


le, costa sho 


rt, not extending 


beyond the proximal 1 


/3 of the lamina. 








3. Leaves complanate 


when moist, apex of le^ 


wes on non-fl; 


agelliform branches obtuse to 


rounded, papillae ii 








9. A. pseudotn 




mate when dry, apex of leaves acut 






obtuse, papillae cor 


ispicuous. 








4. Plants extremely delicate and slender, st. 


sms ca. 0.2 mn 


1 wide, some 


becoming flagel- 


liform, <0.1 mm 


wide at the apex; leave 


;s squarrose, 1 


eaf apex tap. 


sring into a long 


awn (plants rare 


; Manchuria). 






10. A. flafrellifor. 


4. Plants ± delicate 


:, stems ca. 0.4 mm wide 


throughout; 1. 


eaves erect t( 


3 slightly spread- 


ing, leaf apex aci 


jte to acuminate, occasi( 


Dnally obtuse. 






5. Leaves invari 


ably broken off for more than 1/2 t 


heir length, 


with an obvious 


constriction, ; 


apex acute to acuminate 


, occasionally 




.endmg near the 


leaf constrict! 


on; laminal cells with stt 


■ong, high, branching papill 




lose, margin c 


;renulate near the apex c 


5wing to prolr 


uding papillae. 7. A. tri 



397 GRANZOW-DE LA CERDA: ANOMODONTACEAE 21 

costa reaching well beyond the leaf constriction; lamina! cells with delicate, 
branched but not high papillae, never mammillose, margin smooth. 8. A. siebold 
2. Laminal cells with 1-2 (-3) rather high mammilla-like, acute, unbranched papillae, costa 

extending beyond the proximal 2/3 of the lamina. l\. A. longinervi 

Plants somewhat slender to robust, >0.8 when dry, leaves usually >1.4 mm long (in A. thraus- 
tus as small as 1.2 mm long), differentiated basal cells almost reaching the margins and 
extending to ± 1/3 of the lamina; calyptra smooth. 

6. Cells of the lamina with a single, long conical papilla, never branched, but sometimes with 
0-3 short "spurs" at the base. 

7. One single papilla on each cell, low and rounded, without basal "spurs"; the height of 
the papilla not more than the laminar cell's diameter in cross section; leaves slender, 

the narrower distal portion; pseudoparaphyllia present. 16. A. longifolii, 

7. Papillae with 0-3 basal spurs, sharply conical, pointing toward the leaf apex; the height 
of the papillae greater or equal to the diameter of the cell in cross section; leaves 
obtuse or acute but not acuminate; pseudoparaphylHa absent. 
8. Leaves >2.8 mm long, the distal portion (beyond the constriction) tapering, apex 

acute or obtuse; seta >2.5 mm, capsule exserted. 4. A. abbreviati 

8. Leaves <2.0 mm long, the distal portion (beyond constriction) Ungulate, flat and 

with rounded apex; seta ca. 1 mm long, capsule enclosed within the perichaetial 

6. Cells of the lamina with multiple, often more or less branched papillae. 

9. Plants glaucous, profusely and irregularly branched, branches julaceous, erect; leaves 
<0.8 mm long, appressed, the apex extended into a long awn. 13. A. rostrati 

9. Plants more or less dark green to rusty brown, with secondary branches loosely spread- 
ing to erect, sometimes irregularly pinnate, branches never julaceous; leaves >(1.2-) 
1.5 mm long, appressed or not, the apex rounded, obtuse, acute, or apiculate, without 

; cell walls con- 



ceolate acute 


at apex, with unbranched papilla 


. throughout tl 


le lamina; margins revolute. 


;ulate (or lane 


eolate in A. attenuatus), obtuse c 



more or less decurrent, not auriculate, stem light brown, costa yellowish to light 

green; pseudoparaphyllia absent. 

12. Leaves narrowly Ungulate above constriction, this distal portion broken off; 

distal portion of the costa obscured by laminal cells, never bifid. 

13. Secondary branching irregularly pinnate; leaves >2 mm long; costa subper- 



dry, >2 mm long. i- ^- vnitmu. 

14. Plants of various sizes, but branches never more than 1 mm wide when 

dry; leaves complanate when moist, appressed when dry, <2 mm long. 

15. Leaf apex rounded, margin entire; secondary branches poorly devel- 
oped; inflorescences present on terminal branches, beyond the most 
distal branching points; leaf constriction well defined, the distal portion 
Ungulate; secondary branches not attenuate; back of costa with large 
seriate papillae on each cell. 3. A. mir 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



ate; secondary branching profuse, irregularly pinns 

r present on terminal branches beyond the most distal branching points; 

constriction poorly defined, the distal portion tapering to triangular; 



I. Anomodon subgenus Anomodon. 

Stems and primary branches branching in a simple pattern. Leaf apex rounded to 
broadly obtuse (acute to acuminate in section Haplohymenimn): laminal cells 
isodiametric, hexagonal, quadrate or round in all taxa. 

I.a. Anomodon section Anomodon. 

Plants rather robust to somewhat slender. Secondary branches terete, never 
attenuate or flagelliform. Branch leaves spreading to complanate when moist, 
(1.2-) 1.6-4.0 (-4.8) mm long, apex not broken off (except in A. dentatiis), obtuse 
to rounded; costa subpercurrent, or extending for ca. 90% the length of the leaf, 
not obscured by laminal cells. Sporophytes variable in size, urn >1.1 mm long (ca. 
0.8 mm in A. solovjovii). Calyptra glabrous. 

I.a.i. Anomodon subsection Anomodon. 

Branch leaves broadly oblong-lingulate, constricted near the middle into a 
wide upper portion; apex entire, apiculate or slightly denticulate in some species; 
costa often shortly bifid, not obscured in its distal portion by laminal cells; laminal 
cells with multiple branching papillae. Seta >4 mm long. 

1. Anomodon viticulosus (Hedw.) Hook. & Taylor, Muscol. Brit. 79. 1818. Neck- 
era viticulosa Hedw., Spec. Muse. 209. 1801. Hypnum viticulosum (Hedw.) 
L. ex With., Syst. Arr. Brit. PI. ed. 4, 3: 830. 1801. Haptymenium viticulo- 
sum (Hedw.) Fiirnr., Flora 10 (2, Beil. 1): 81. 1827.— Type: Germany. 
Giessen, Dillenius (as Neckera viticulosa and Hypnum viticulosum: G- 
Hed.-Schwaegr. !, sine loc, c. fr., fide Grout 1932-34). Fig. 11. 

Anomodon tonkinensis Besch., Bull. Soc. Bot. France, ser. 3, 41: 84. 1894.— Type: 
Vietnam. Tonkin, Hanoi, Mt. Chua, 12-3-1888, Bon 3789 (isotype: NY!). 

Anomodon subintegerrimus Broth. & Paris, Rev. Bryol. 27: 77, 1900.— Type: 
Vietnam. "Tonkin, inter Thai Van et Pho Lu", 18-3-1901, Moutier (iso- 
types: G! JE! NIGH! NY!). 

Anomodon decurrens Cardot, Bull. Soc. Bot. Geneve, ser. 2, 3: 281. 1911.- 
Syntypes: Japan. Mitoyoshi, ./. Cardot 1190\ Korea. Seoul, Cardot 313 
(PC). 

Plants robust, forming thick mats; branches 6-8 cm long, occasionally more, 
and 1-1.8 mm thick when dry, erect-ascending to arcuate, stem without a central 
strand of differentiated cells, pseudoparaphyllia absent. Branch leaves erect when 
dry, secund to falcate-secund and somewhat flexuose, spreading to reflexed when 
moist, oblong-Hngulate, abruptly narrowed below the middle from an ovate-lan- 
ceolate base, 2.2-4.0 mm long, moderately spinulose at the insertion, margins 
plane, tapering into an obtuse to rounded, sometimes acute apex; costa strong, 
pellucid, ending sharply (not obscured by laminal cells) near the apex (0.05-0.15 



GRANZOW-DE LA CERDA: ANOMODONTACEAE 




nomodon viticulosus (Hedw. 
\ (showing decurrency); e, portion 
'idra, Rio Ges, 18-1-1988 (BCB); b. 



mm), sometimes flexuose, rarely bifid; cells at the back of the costa long, each 
with several thick seriate papillae; dorsal laminal cells occasionally covering the 
upper end of the costa, obscuring it with short cells; basal hyaline cells of the 
lamina barely extending beyond 1/4 the length of the leaf base, each cell bearing a 
single papilla, the basalmost with walls sinuose, often perforate; cells of the median 
lamina obscure, hexagonal, with multiple branched papillae. Inflorescences appearing 
distally from the most recent branching points. Perichaetial leaves, similar in shape 
and size to vegetative leaves, with costa ending closer to the apex, cells papillose. 
Seta 10-20 mm. Capsule long-elliptic, elongate, urn (1.5-) 1.8-2.8 (-3.1) mm long, 
stomata absent, annulus compound; operculum obhquely short-rostrate, 0.7-1 mm 
long; exostome 0.3-0.5 mm high, nearly smooth, with the base yellow, faintly 
striolate, teeth irregular, not trabeculate, with inconspicuous papillae toward the 
apex; endostome with basal membrane 2-4 cells (0.07-0.09 mm) high, almost 
smooth, segments moderately developed, 0.07-0.15 (-0.2) mm, several cells high, 
smooth or variably papillose. Calyptra smooth. Spores very variable in diameter 
between capsules, in some (19-) 20.5-23 (-25) |im, in others 15-16 |im, densely 
papillose. Chromosome number: n = 10 + x/y (in meiosis, Spain: Granzow-de la 
Cerda, 1988b). 



220 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 2 1 

Distribution (Figs. 12-16). Anomodon viticulosus is the most cosmopolitan spe- 
cies in the genus. Circumboreally, it occurs in North America, Asia, and Europe 
(where it is the most common). It has been reported from the Canary Islands by 
Dixon (1908) and Madeira by Luisier (1920-1922), although I have not been able 
to confirm these records (Granzow-de la Cerda 1988a) or others from Algeria 
(Diill 1985). Anomodon viticulosus occurs in practically all of Europe, from Spain 
(including the Baleric Islands) and Great Britain to Turkey, east to the Caucasus, 
and north to Scandinavia; in the western Himalayas from Pakistan to southern 
Siberia, China, Korea, Japan, Burma, and Vietnam; in eastern North America in 
Canada (Ontario, Quebec, New Brunswick, and Nova Scotia), the northeastern 
United States to the southern Appalachians of Virginia and Tennessee, the Ozarks, 
north to Iowa; also in Mexico (Guerrero and Oaxaca). It is almost restricted to 
rather mesic montane calcareous environments of deciduous forests, on calcare- 
ous rocks or vertical walls, although sometimes it also occurs as an epiphyte. In 
Europe, A. viticulosus fruits frequently, but almost never in the New World or 
Asia, most likely because of the lack of male gametophytes in these regions. 1 
have seen just one fruiting specimen from North America, and only one sporo- 
phyte has been reported from eastern Asia, from Japan, according to Iwatsuki 
(1963). Anomodon viticulosus is not common but locally abundant. 

Representative Specimens. Burma. Pr. Now Kangi Village, Jawahir-Singh, 2-1-1904 (as A. 
integerrimus, NY). China. Hsien: Lichuan Xian, Lojiaba valley, Exp. Sino-Amer., 6-10-1980 (FH). 
Hubei: Lichuan Xian, Lojiaba, Sino-Amer. Exp. 1991, 6-10-1980 (MICH, NY); Shenongjia Forest 
Distr., pr. Raojia, S of Shi rvr., Exp. Sino-Amer. 1666, 21-9-1980 (FH, MICH). Jilin: Chang Bai shan, 
Lin Chang Qing 2953, 4-1964 (IFSBH). Szechuan: Chungking, Siao lung kan, Chen 4-1940 (as A. 
thraustus, JE). Xishuangbanna: Mengla, Menking to I-wu, P. Redfearn el ai 33960, 24-12-1986 (NY). 
Yunnan: Songming, pr. Longtan, P. Redfearn, S. He c& Yi G. Su 853, 27-7-1984 (NY). Georgia. 
Swanetia Dadianosum, Sommier & Levier, 29-7-1890 (FI). India. Himachal Pradesh: Simla, Thom- 
son 1112, 1116, 10/49 (as A. ohtu.sifolius Mitt., n. nud., c. fr., "type" in herb. Mitten NY); Kangra, 
Kulu, Koeltz 7510, 24/27-11-1933 (c. fr.. DUKE, MICH). Maharashtra: Bombay, Rachira, Marchal, 
5-1869 (c. fr., FI). Punjab: Murree. Stewart, 9-1948 (NY). Uttar Pradesh: Tihri Garwal, Kidarkanta, 
Bahadru, 12-6-1904 (as A. integerrimiLs. c. fr., NY); Musoorie, Dhanoutti, Bahadru, 10-12-1903 (as 
A. integerrimus. NY). IRAN. Mazanderan, 30 km S of Amal, P. Goldblatt 1221a, May 1973 (FH, 
MO). Japan (as A. deciirrens). Hondo: Akiyoshi, Nagato, S. Matsiimoto, Mar 1932 (NICH). Hon- 
shu: Jaisaku pref.. Bingo, A. Nogiichi, 7-7-1934 (NICH); Okayama pref., Niimi-shi, A. Noguchi 3-11- 
1954 (NICH); Shiga pref., Sakata-gun, Kaminyu, T. Nakajima, 4-4-1978 (MICH). Kyushu: Fukuoka 
pref., Hisaodai, T. Osada, 2-5-1955 (MICH), Kaharu Mt., A. Noguchi, 13-9-1953 (NY), Kawaradake, 
A. Noguchi, Sep 1953 (MICH); Kumamoto pref., Kuma-gun, Koonose, Takaki2302, 11-8-1946 (NY), 
K. Mayebara, Dec. 1947 (MICH), Itsuki-mura, Sangaura, A. Noguchi & Imae, 23-4-1956 (NICH); 
Oita pref., Kuwanobori-mura, A. Noguchi, 23-11-1953 (NICH); "center of back bone, Mt. Kumimi,'" 
Kuwahara 2180, 12-8-1951 (NY). Nepal. Longtang Himal: Dunche to Kyangjing, Langtang Khola, 
W. Weber B-99386, 21-10-1991 (NY). Russia. Gorno-Altaisk: SE of Ust-Syana, Katun rvr., T. Elias 
et al. B-65178, 65180, 17-8-1978 (NY). Taiwan. Tienhsiang, Tailuko rvr., Z. Iwatsuki & A. Sharp 
1787, 27-3-1965 (FH). Thailand. Phu khieo, E of Chaiyapun. Lorsen, Nielsen & Santisuk 31305, 3- 
8-1972 (NY). Turkey. Trabzon: Ma^ka. Pontus Mts., Godfrey 728, 24-9-1952 (NY). 

Canada. Nova Scotia: Cumberland Co.. Wentworth, R. Ireland 10673, 7-9-1967 (as A. minor, 
DUKE). Ontario: Bruce Co., 2 km N of Colpoys Bay, R. Ireland 20290, 12-7-1982 (MICH), White 
Bluff, R. Ireland 20396, 14-7-1982 (FH); Grey Co., Inglis Falls, H. Crum 11008, 3-8-1961 (DUKE, 
MICH), Jones Falls, R. Ireland 20198, 11-7-1982 (FH); Lincoln Co., betw. Vineland and Beamsville, 
H. Crum 31, 42, 5/7-10-1964 (MICH); Ottawa, Bauer, 31-3-1902 (FH). Quebec: Gatineau Co., King 
Mt, Gatineau Park, R. Ireland 15853, 27-4-1972 (MICH); Hull, /. Macoun. 22^-1896, 18-5-1901 
(MICH). Mexico. Guerrero: Omiltemi, 30 km W of Chilpancingo, A. Sharp 1118c, 1122, 27-10-1944 (as 
A. minor, MEXU, MICH). Oaxaca: S. Pablo Ayutla, 70 km E of Oaxaca, A. Sharp et al., 23-12-1972 
(MEXU). U.S.A. Arkansas: Newton Co., junc. Leatherhood Creek and Buffalo Rvr., P. Redfearn 
29261, 2-2-1972 (MICH). Michigan: Alpena Co., SW of Posen, Leer Road. Dull 1152/3, 18-8-1981 
(MICH); Delta Co., Burnt Bluff, H. Gleason 2390, 22-8-1939 (MICH), W. Steere 760, Aug. 1934 



lyy / GRANZOW-DE LA CERDA: ANOMODONTACEAE 221 

(FH); Keweenaw Co., gorge of Manganese Rvr., W. Steere, 1-9-1936 (MICH); Mackinac Is!., E. <Sc S. 
Harper, 18-7-1899 (MICH), /. Schnooberger 2749, 21-7-1939 (MICH); Presque Isle Co., nr. Long 
Rapids, R. Fidler 89, 3-8-1974 (MICH). Minnesota: Winona, Holzinger, 1890 (FH). New York: 
Buffalo, G. Clinton (MICH), Niagara Falls (ex Sullivant & Lesqureaux, MICH); Clinton Co., Aus- 
able Chasm, L. Anderson 20608, 5-9-1968 (DUKE, MICH). Vermont: Glebe Mtn., Newfane, A. 
Grout, Aug. 1900 (as A. viticulosus var. microphyllus, FH). Virginia: Marion, A. Vail & E. Britton, 23-5- 
1892 (MICH); Giles Co., E. Byers 16, 4-7-1949 (MICH), Newport, /. G.-de la Cerda 1892, 1898, 7-10- 
1892 (DUKE). Tennessee: Anderson Co., Savage Gardens, A. Sharp, 13-1-1957 (MICH, TENN), /. 
G.-de la Cerda 1906 & A. Sharp, 8-10-1984 (DUKE). 

Austria. Salzburg, pr. Furntenstein, /. Zimmerman, 6-5-1868 (c. fr., FH); "Traunfall, pr. Schwan- 
enstadt," Matouschek (c. fr., FH, FI); Bergwalder, Klaus, Jul 1910 ("var. latifolius,'' c. fr., NY). Tirol: 
Windish Matrai, Klamm, E. Bauer 1217, Aug 1911 (MICH); Innsbruck Sillschlucht, Luisier 22-3- 
1904 (c. fr., INA). Belgium. Auderg hem, Pire (c. fr., FI); betw. Rochehaut & Ucimont, J.L. De 
Sloover 16439, 1-7-1973 (MICH); Louvaine, pr. Tournay, Marissal, (c. fr., FH); Orval, / Cardot 
(FH). Croatia. Istra, above Icici, pr. Opatija, J.P. Frahm 10034, 26-3-1975 (c. fr., MICH). Czech 
Republic. Bohemia: "Karlstein," Bauer, 24-4-1890 (c. fr., FH). "Mahr-Weisskirchen," Surkow, F. 
Petrak, 9-4-1913 (FH). Denmark. Manager kridtbrud, K. Holmen & E. Warncke 237 (MICH); 
Moen, Klinteskoven, H. Moller, 1/3-6-1893 (c. fr., FH). Sjoeland: Farum, N0rreskov, G. Smith, 24- 
10-1967 (FH); Soro, Suserup, E Hermann 15885, 28-5-1960 (MICH). Finland. Tohmajarvi,' Kb: 
Havukkavaara, S of Vuorenpaa, R. Fagersten & M. Haapassari 4097 dup., 2-10-1984 (FH, MICH); 
Rukola, R. Fagersten & M. Hapasaari 4120, 3-10-1984 (FH). "Karelia borealis," Juuka, Polvijarvi, A. 
Huuskonen, 19-8-1965 (FH). France. Ariege: Bagneres de Luchon, C. Casas, 20-9-1956 (BCB). 
Basses-Pyrenes: St. Etienne-de-Baigorry, M. Fleischer, 10-5-1908 (FH). Calvados: Pont des Vers, 
Husnot, (c. fr., FH). Gard: Mejanes-Le-Clap, /. L. De Sloover 44850, 12-7-1988 (MICH). Haute- 
Savoie: Saleve, Archenes, R. Blanche, 15-2-1954 (c. fr., MICH). Isere: Bains-la-Romaine, J. L. De 
Sloover 34593 10-7-1981 (MICH). Savoie: Saint Claude, "pres de Chamberi," Songeon, 20-10-1866 
(c. fr., FI). Germany. Giessen, Wanschoff, (c. fr., FH); Munchen, Dodel-Port., 20-2-1869 (c. fr., 
FH); "Rhinefall," Mann & Cummings, 4-3-1887 (c. fr., FH). Greece. loannina: Epirus, pr. Emin 
Agha, K. Rechinger, 12-5-1961 (MA). Thessalia: Lutropigi, Smokovo, K. Rechinger, 8-5-1961 
(MA). Ireland. Sligo: Knocknarea, H. Conrad, 4-8-1935 (FH). Italy. Caiagilo, pr. Varesse, E. 
Corti, Mar. 1905 (c. fr., FH). Lavione: "pr. Florentiam," /. Arcangeli, 23-12-1877, (c. fr., FH). Lom- 
bardia: Parco di Monza, Mazzucchelli, 19-9-1878 (c. fr., FI); Piemonte: Campello-Monti, Roncaccio, 
Levier, 30-7-1906 (FI); Como, Torno, Lago di Como, Artaria, 2-1902 (FH). San Marino: Borgo, 
Ornella, Pampanini, 8-1-1916 (c. fr., FI). Trento: Lago Tadel, Serra del Lago, Cortini 973, 16-7-1958 
(FI). Venezia: Padova, Fiori, 23-4-1893 (c. fr., FI). Macedonian Republic. W of Tetoro, Popova 
Sapka., exp. Rheno-Trai 68-1526, 10-7-1968 (MEXU). Norway. Sodra Trondheim, HSkan, /. Hagen, 
Jul 1899 (MICH); Lysaker, pr. Oslo, /. Hagen 18-10 (FH). Poland. Krosno: Kurnikow Beskid, R. 
Ochyra, 7-7-1977 (BCB, MICH); Nowy Sacz, Spiskie Klippen (W. Carpathians); Zielone Skaly, 
Dunajec rvr., R. Ochyra 657, 1-6-1979 (BCB, MICH); Kramnika Klippe, Rvr. Bialka, R. Ochyra 564, 
19-6-1979 (MICH). Skata Knity (Krakov): Zabiezowic, Z. Zmuda, 22-4-1911 (FH). Russia. Mos- 
cow, "Wald von Kunzowo," Heyden, 1-8-1900 (FH). Slovakia. Gau Kaschau, Snina, Vinicna hora, 
Bauer, 31-7-1925 (FH); Tatra Magna, Tscheckengrund valley, /. Gyorffy 42, 6-8-1927 (MICH). Spain. 
Barcelona: Montserrat, Sant Geroni, C. Casas, 5-10-1947 (BCB). Cantabria: Puente Viesgo, A. Casares 
Aug. 1911 (MA). Castello: Mangraner, La Cenia, barranc de la Fou, C. Sergio et al. 4.1, 14-2-1984. 
Girona: Sta. Maria de Besora, Serra de Caraups, /. G.-de la Cerda 2554 18-1-1988 (BCB). Vidra, rio 
Ges, rec de I'Espirau, /. G.-de la Cerda 2316, 2322 & M. Sanz, 10-1-1987, /. G.-de la Cerda 2546, 18- 
1-1988 (c. fr., BCB). Leon: Posada de Valdeon, Garganta del ri'o Cares, l.G.-de la Cerda, 1-1-1990 
(BCB). Lleida: Pallars Sobira, "entre Rialp i Llesui," C. Casas 6-10-1981 (BCB). Lugo: Lozara, F 
Merino (c. fr., INA, herb. Luisier). Navarra: Alto de Lizarrusti, A. de Miguel & A. Ederra, 27-11-1983 
(c. fr., NAU). (see Granzow-de la Cerda 1988a). Sweden. Jamtland: Brunflo, Backen, C. Stenholm, 
31-7-1934 (MICH). Oland: Borgholm, E. Ekslrand, 8-7-1975 (c. fr., MICH), H. Moller, 16-4-1908 
(FH). Skane: Kirgerod, YlmesSken, C. Stenholm, 25-5-1934 (FH). Uppsala: Gottsunda,'s of Fabo- 
darna, F Osterlind, 3-5-1951 (FH, MICH). Switzerland. "Switzerland" ? (DUKE). Mt. Saleve, M. 
MacFarlane, Feb 1911 (c. fr., FH). Splugen, Areschoug, Jun 1861 (c. fr., FH). Geneve: Chatelaine, 
Boissier, Mar.1821, (c. fr., MICH). Zurich, P. Culmann, 11-3-1880 (c. fr., FH). United Kingdom. 
Scotland: W. Wilson, 320 9-7-1861 (c. fr., FH). Anglesey [Gwynedd]: Tyfry, W. Wilson, Mar. 1830 (c. 
fr., MICH). Devon: Honiton, H. Shacklette 2266 25-1-1943 (MICH), Plymouth, Holmes, 1867 (c. fr., 
FI). Gloucester: Lead, Morgan's wood, E. Elliot, 18-1-1886 (c. fr., FH). Somerset: Weston-Super- 
Mare, 2km NNW Chedar, Jury et al. J1787, 8-1-1983 (BCB). Wiltshire: Edington, pr. Bridgenorth, 
Mori Brook, /. G.-de la Cerda 2018, Jul 1884 (DUKE). 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 









V o . 
o 




O A.attenua.us 




FIG. 12. Distribution Of yl/i 


jmoclon 


atwnuatus ..nd A 






^-3V 



Anomodon viticulosus is the most robust species of the genus. It forms thick 
mats on rocks and sometimes also on tree trunks. The robust habit and the thick- 
ness of the mats make it clearly distinguishable from any other species for which it 
could be mistaken, such as A. attenuatus, A. rugelil, or A. minor. In depauperate 
populations of A. viticulosus, this species can be distinguished from A. rugelii by its 
decurrcnt instead of auriculate leaves. Also, A. rugelii has incurved leaves when dry, 
whereas those of A. viticulosus are, at most, falcate-secund, tapering toward the api- 
ces from the shoulders. The apex is never rounded, as in A. rugelii, A. minor, and A. 
(hraustus. The branches are not complanate, as in A. rugelii or A. minor. The 
latter species forms much looser mats than A. viticulosus. The endostomes of A. 
minor and A. viticulosus differ considerably; they are rudimentary in A. minor and 
better developed in A. viticulosus. Some specimens, however, have a very rounded 
leaf apex, therefore resembling A. minor (most often seen in specimens from 



GRANZOW-DE LA CERDA: ANOMODONTACEAE 







central Mexico). Anomodon viticulosus is sometimes mistaken for 
although this species has more prostrate stems and its branching pattern is differ- 
ent, mostly more profuse. Branching in A. attenuatus occurs in several successive 
orders; the terminal branches are attenuate, complanate, crowded and somewhat 
fasciculate. In A. attenuatus the back of the costa is smooth, and perichaetia do 
not develop on the last year's branches (but only in older portions of branches). 

Most specimens oi A. viticulosus from Japan have been called A. decurrens. It 
is true that some specimens from these islands may have slightly more slender 
leaves. Also, some individuals - within the same specimen along with typical 
forms of A. viticulosus - may resemble either A. attenuatus or A. minor in habit. 
Nonetheless, there are no differences to account for placement of such plants in 
any taxon other than A. viticulosus. Anomodon dentatus, a rare endemic from 
northeastern China, is a more delicate plant, although similar in habit to A. viticu- 
losus; also, the portion of the leaves above the shoulder in A. dentatus is lingulate 
to slightly spatulate, with a serrate apex, often broken off. Anomodon tonkinensis 
and A. subintegerrimus are examples of depauperate forms of A. viticulosus, with less 
crowded leaves (perhaps somewhat reminding of A. minor), which do not deserve 
: consideration. 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 




1 ' ' 


js 


-^ °B*' 1 


- 





1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 225 




Anomodon viticulosus is the type species ioxAnomodon Hook. & Taylor (1818; 
basionym: Neckera viticulosa Hedw.) The original material on which Hedwig based 
his description was Linnaeus's "'Hypnum viticulosum,'" collected by Dillenius near 
Giessen in Germany (Grout 1934). 

2. Anomodon rugelii (Miill. Hal.) Keissl., Ann. Naturh. Hofmus. Wien 15: 214. 1900. 
Hypnum rugelii Miill. Hal., Syn. 2: 472. 1851.— Type: U.S.A. Tennessee, 
Smoky Mountains, Rugel s.n. (holotype: not located). Fig. 17a, b. 

Anoectangium fermgineum Besch., Ann. Sci. Nat. Bot. 17: 329. 1893. Anomo- 
don ferrugineus (Besch.) Nog., J. Jap. Bot. 20: 260. 1944. Anomodon rug- 
elii var. ferrugineus (Besch.) Z. Iwats., J. Hattori Bot. Lab. 26: 51. 1963.— 
Type: Japan. Yezo, montagnes d'Yesashi, 6-6-1889, Faurie 3543 (FI?). 

Anomodon uyematsui Broth., Bull. Soc. Bot. Geneve, ser. 2, 3: 281 1911. 
Anomodon uematsui Broth., Ofver. Finsk. Vet. Soc. Forh. 62: 33. 1919- 
1920.— Type: Japan. Hokkaido, Iwate pref., Rikuchu Nat. Park, Mt. Hay- 
achine, 27-7-1906, Uematsu 3390 (isotype?: NY!). 



(^ONTR. UNIVERSITY OF MICHIGAN 




FIG. 17. 



nodon rugelil (Mull. Hal.) Keissl. and A. attem 
a, branch leaves; b, stylized representation of a branch leaf, c-e, . 
apex; e, leaf insertion. Based on: a, b, G-de la Cerda 2512 & Ren: 
Kilmer Mem. Park, 3^10-1987 (DUKE, MA); c-e, Standley. Gua 
12-4-1941 (NY). 



, Alta Verapaz, Rio Carche, 



Plants rather slender, forming thick, dense mats, dark green to rusty-brown. 
Stems and main branches to 3.5 (-5) cm long, 0.8-1.5 mm thick when dry, creep- 
ing, with few primary branches, erect-ascending, somewhat arcuate, poorly 
branched, dark reddish brown, without a central strand, with fohose, often narrow 
to uniseriate, golden-yellow pseudoparaphyllia, generally with abundant rhizoids. 
Branch leaves incurved-contorted (ca. 1 full turn) when dry, erect-spreading, some- 
what secund when moist, (1.2-) 1.4-2.3 (-2.5) mm long, broadly oblong-hngulate 
from an ovate, auriculate base; margins of auricles strongly spinulose, with high 
and branched papillae; apex broadly obtuse to rounded, often apiculate; costa 
strong, pellucid, golden-yellow to rusty-brown, ending sharply near the apex, gen- 
erally bifid, sometimes slightly flexuose at the end; cells of the costa elongate, 
smooth on the back, up to 70 (im long; basal cells hyaline and well differentiated, 
occupying more than half the basal portion of the lamina, their walls not papil- 
lose, somewhat irregular, often sinuose; medial laminal cells hexagonal, 7-12 )im 
long, with multiple high and branched papihae; marginal cells of the auricles with 
1-2 strong branched spinelike papillae, occasionally minutely denticulate at the 
apex. Perigonia and perichaetia at the end of terminal secondary branches. Peri- 
gonial leaves with few or no papillae, upper margins crenulate. Perichaetial leaves 
abruptly narrowed toward the apex, becoming almost subulate, cells papillose. 
Seta length variable, (5-) 9-22 mm. Urn (1.2-) 1.8-2.3 (-2.5) mm long, elongate, 
with stomata at the base; annulus absent, operculum conic to obliquely short- 
rostrate, 0.5-0.8 mm long; exoslome 0.15-0.30 mm high, teeth regular, papillose 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 227 

throughout, often inconspicuously horizontally striolate at the base, trabeculate 
above; endostome rudimentary, 0.042-0.057 mm high, with a basal membrane 2-4 
(-6) cells high, segments very reduced (1 cell high) or absent. Calyptra smooth. 
Spores 9.5-14 |im in diameter, slightly papillose. Chromosome number: n = 10 + x/y 
(in meiosis, North Carolina: Granzow-de la Cerda, 1988b). 

Distribution (Figs. 15, 16, 18, 19). Present in Asia, eastern North America and 
Central Europe; circumboreal montane, growing in deciduous forests. Anomodon 
nigelii forms extensive, dense and thick mats on tree trunks, generally, although 
not always, 1-2 m above the base. Some specimens have been collected from both 
basic and acidic rocks. Sporophytes are found in fair abundance. 




CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 






fit. 1 i^ 

T-Si5ft;.VX 




R 


LPRI:ShNTATIVl: SPLCIMl 




Azerbaijan. Baku 


iani, A. 


L«2i 


renk 


} & 




mov, 17 


-9-1956 


(NY). 


China. Hubci: Shcnnc 


ngj 


1 Forest Distr., -f 


om Qui 


ujiajing 10 


Lao 


un," 


.9//ir>/lm 


.. £.-/;. 


655r. 


-9-1980 (NY). .Iilin: Cha.iL 


Bai Shan. Gno Qi 










Y). I 






monlc 




y. 




nuginei 


V, IFS 


BH) 


He 


long. 


an: distr 


Hailin, 


Dahai 
plcmcl 


Laotudin^., ^^-^'^ 


rh" 


Miucn NY). Ullar 


(IFSBl 
PiadL'sl 


) In 

Teh. 


Ga 


To 


Kid 


.p 1 ()()()() 


cdZinu 




K)4 (NY). Colhin. 23-3- 


9(M 


(c. fr.. FII. NY). •• 


idia uri 


nlalis 


■.Be 


/.?6 


7 (as 


\. minor. 


jx lierb. 


Mit'tci 


NY). Japan. Hokkait 


o: S 


.pporo. Ml. Moivv. 


. 'V /uv 


■aki. 1 


7-11 


-191 


3 (e. 


r.. NY). 


ionslui: 


[l^A 


'lZui:'nL':^\niy.\li 


:;:; 


Sckidn-t(v-c (.Sek,) 


/'^/uT 


ri pre 


m. 


n-^ 


''\r. 


'{^4"! 


tuasin, 


NY): 


Kvc,u> prel., Kila-Kiivsa 


k'-L 


uii. Sasan-loge. r. 


Sakn:ii 


m 5.\ 


3. 2 


.5-1 


J54 ( 


s A ten 


igineus, 


Nicn 


:Mieprd-.. lliiakui-afc 
















Nagano 






A^^ 




N'Y), K; 




-till 11 


Ak 


'i&'»^ 


ira. 7Vi/c« 


/t,, 5-8- 


1933"( 


c. rr.. NY), Nakajnna. 2 


-8 


97.^ (NY); Nara p 


cf.. Yos 


illU)-g 


Lin, Nose 


gawa 


mura, T. 


\cik(izi- 


ma 87 


)8, 3-5-1958 (as A. fern 


-in, 


(/v, NlC'in. Mt.Oc 


ugahara 


7. A' 


kazi 


na 


w;7. 


6-10-194 


8 (as /I 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 229 

A. fernigineus, NICH); Saitama pref., Chichibu, Higashi-dani, /. Nagano 546, 4-11-1951 (as A. uye- 
matsui, NICH); Tottori pref., Yazu-gun, Mt. Nagi-sen, H. Ochi 1582, 26-8-1951 (as A. ferrugineus, 
NICH); Toyama pref., Mt. Tateyama, Z. Iwatsuki, 13-8-1955 (NY); Kurabe, Keyakidaira, A. Nogu- 
chi, 13-10-1953 (as A. ferrugineus, NICH); Komagataka, Faurie 3493, 1905 (c. fr., NY). Kyushu: Oita 
pref., Mts. Kuju, Kurodake, M. Miyamoto 2431, Jul 1960 (c. fr., as A. ferrugineus, NICH). Nepal. E. 
Himalaya: Kathmandu, Mt. Phulchoki, Z. Iwatsuki 2470, 15-7-1972 (NY). Russia. Reg. austro- 
ussuriensis, "fontem flumen Ssputinka," A. Lazarenko, Oct 1934 (NY). 

Canada. New Brunswick: Grand Falls, H. Habeeb 86, 1-11-1945 (NY), Aroostook Falls, /. 
Collins 2770, 17-7-1902 (MICH). Victoria Co., Kilburn, R. Ireland 12758, 9-7-1970 (MICH). Nova 
Scotia: Digbi Co., AcaciaviUe, R. Ireland 17890, 28-7-1974 (MICH, NY). Ontario: Carleton Co., 
Fallowfield, H. Crum 290, 28-10-1964 (MICH). HuntsviUe Co., Muskoka Distr, NW of Chaffey Twp., 
R. Ireland 22021, 23-7-1986 (MICH). Algonquin Park, J. Macoun, 8-6-1900 (MICH). Manintulin 
Distr., Sheguindah, Howland Twp., R. Ireland 22372, 30-7-1987 (NY). Muskoka Distr., Moon rvr., 
Medora Twp. (Foots Bay), R. Ireland 24032, 7-9-1982 (NY). Quebec: Terrebonne Co., E of Mt. 
Tremblant Lodge, H. Crum 9885, 23-8-1958 (MICH); Gattienau, nr. Kingsmere and Old Chelsea, H. 
Crum 2468, 1954 (NY); St. Hyacinthe, F. & M. Anselme A-5, 3-5-1948 (NY). Pontiac Co., Park de la 
Verendrye, Lac Serpent, R. Ireland 15969 & T. Koponen, 13-6-1972 (NY). U.S.A. Maine: Pleasant 
Ridge, Bean Pond Ridge, J. Collins 2455, 21-7-1901 (as A. viticulosus, MICH). Michigan: Keweenaw 
Co., "on ledge at Delaware," W. Sleere, Sep 1936 (as A. viticulosus, FH). New York: Cayuga Lake 
Basin, Dudley, 16-5-1884 (c. fr., DUKE); Lake Placid, Britton, 9-8-1898 (DUKE); Adirondack, 
Essex Co., North Elba, Hermann 14776, 10-8-1958 (DUKE); Catskills, St. Lawrence Co., Cranberry 
Lake, A^. Slack 2031, 25-6-1976 (DUKE). North Carolina: Roan Mountain, Weatherby, 24-2-1894 (as 
A. minor, c. fr., DUKE); Graham Co., Joyce Kilmer Mem. Park, /. G.-de la Cerda & K. Renzaglia, 3- 
10-1987 (c. fr., DUKE, BCB); Macon Co., Chatooga river, /. G.-de la Cerda 1454, 1-7-1984 (DUKE); 
McDowell Co., Linville Falls, /. G.-de la Cerda 1931, 21-10-1984 (DUKE); Watauga Co., Sim's 
Creek, Blowing Rock, L. Anderson, 27-9-1984 (c. fr., DUKE); Yancey Co., Crabtree meadows, /. G.- 
de la Cerda 1925, 20-10-1984 (DUKE). Pennsylvania: McKean Co., Bennet Brook, Burnett 2831, 7- 
11-1897 (FH). Vermont: Willoughby, Humphry Brook, Kennedy, 27-10-1897 (as A. obtusifolius, FH). 

Germany. Bayern: Allgau, "Sauwald ober Hinterstein," Holler, 22-8-1892 (COI), Holler, 29-8- 
1891 (NY); "Monte Rhon," G. Geheeb, 1870 (c. fr., NY), "Rhongebirge," G. Geheeb, 1869 (NY), 
Geisa Wald, G. Geheeb, 1873 (c. fr., NY); Hiibelkuppe bei Geisa, in Vorder-Rhon, G. Geheeb, 16- 
11-1868 (c. fr., NY). Hessen: Laubach, Vogelsberg, Roth, 9-8-1903 (herb. Bartram FH, NY), Roth, 
10-6-1885 (ex herb. Uggla NY). Poland. Silesia: Gralen bei Gobersdorf (Kreis Waldenburg), / & 
H. Milde, 8-1869 (c. fr., NY). Norway. Gaset i Asker, Kiaer, 7-6-1878 (ex herb. Uggla NY). Aker- 
shus: Asker distr., SkaugumsSsen, Fridts, 10-9-1886 (c. fr., BCB); Grossaetdal, Asker, pr. Christiania 
[Oslo], Kaalaas, 29-5-1883 (c. fr., NY). Telemark: Tinn Distr., Rollag, Kiaer, 4-8-1890 (BCB). Slovakia. 
"Slovakia orientalis: monies Cerhovske pohorie," Vdna, 29-5-1973 (NY). Sweden. Vastergotland, 
Billingen, Moller, 9-8-1914 (herb. Bartram FH); Mosseberg, Lindberg 18.23/6, 22-6-1880 (NY), Flink, 
25-6-1880 (MA); V^mb Billingen, Hulphers, Jun 1916 (c. fr., NY). SmSiand: Ljungarum, Rosenlund, 
Arnell, 9-1891 (NY). Switzerland. "Mtns. of Rhone valley," G. Geheeb (c. fr., NY). 

Anomodon rugelii is distinctive in the rusty brownish color of the plant, the 
feltlike primary branches (due to the abundance of rhizoids), the more slender 
branches with strongly incurved leaves when dry, the redness of the costa, and, 
above all, the conspicuous auricles at the leaf base. Apiculate leaves are not a 
reliable character, as there are many plants whose leaves are obtuse to rounded 
and lack an apiculum. According to Ireland (1971), A. rugelii has foliose 
pseudoparaphyiha, but, mostly, they are completely uniseriate, as Ireland's (1982) 
later figures show. Multiseriate pseudoparaphyiha are uncommon in this species. 

Even though they are nomina nuda, the names Anomodon apicidatus B.S.G. 
and to some extent also Leskea apiculata Schimp. in Miill. Hal. have been exten- 
sively used in older hterature for A. rugelii. The apiculate leaf apex is a distinc- 
tive, if inconsistent, feature, and one can understand why so many authors have 
been drawn into using these names. 

Some authors, including Iwatsuki (1963), recognize the variety ferrugineus 
(Besch.) Z. Iwats. (also apphed to specimens named A. uematsui Broth.; A. fer- 
rugineus (Besch.) Noguchi), on the basis of a more reduced habit, more delicate 



230 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

leaves, smaller auricles, and the lamina with an even more pronounced rusty tint 
(which accounts for the epithet). Leaves in these specimens are less incurved than 
in the type. In my opinion, this variability does not deserve formal taxonomic 
recognition. Leaves in many specimens of A. rugelii are not apiculate but round- 
ed. Specimens from NE China (province of Jilin) and Japan show very reduced 
and inconspicuous leaf auricles; auricles may even be absent in some of the leaves 
on either side of the leaf insertion. Some leaves are even found to be decurrent, 
and occasional large round papillae are found on the back of the costa; in A. 
rugelii the costa is usually smooth abaxially. Nonetheless, these specimens do 
have the highly raised papillose-mammillose cells on the basal leaf margin and 
also the auricles, the most characteristic feature of A. rugelii (although sometimes 
dramatically reduced). This material could very well belong to what Iwatsuki consid- 
ers subsp. ferrugineus {=A. uematsui), although this taxon is supposed to be an 
endemic of Japan. 

One of the syntypes of A. ramulosus Mitt. (Japan, Nantaizan, collected by 
Bisset) is A. rugelii. The description is rather vague; because the most distinctive 
features of A. rugelii are not mentioned (rusty color, incurved, apiculate leaves 
with auricles), I am inclined to think that Mitten considered the specimen to 
belong to A minor. 

3. Anomodon minor (Hedw.) Fiirnr., Flora 12 (Erg. 2): 49. 1829. Neckera viticu- 
losa var. minor Hedw., Spec. Muse. 210. 1801. Neckera minor P.-Beauv., 
Prodr. 78. 1805. Neckera flaccida Brid., Sp. Muse. 2: 137. 1812.— Type: 
U.S.A. Pennsylvania, Lancaster, Muhlenberg (as Neckera viticulosa var. 
minor, G-Hedw.-Schwaegr.!, c. fr.). 

Hypnum viticulosum var. brachycarpum Miill. Hal., Syn. 2: 473. 1851.— Syn- 
types: U.S.A. Drummond, Musci Amer. 163; Sullivant, Musci Allegen. 74. 

Anomodon integerrimus Mitt., J. Linn. Soc. Bot., suppl. 1: 126. 1859. Anomo- 
don minor subsp. integerrimus (Mitt.) Z. Iwats., J. Hattori Bot. Lab. 26: 
41. 1963.— Type: Nepal. Wallich (lectotype, here designated: NY-herb. 
Mitten!). 

Anomodon planatus Mitt., J. Linn. Soc. Bot., suppl. 1: 126. 1859.— Type: India. 
Himachal Pradesh, Simla, "7000, 8000," Thomson 1126, Oct 1849 (lecto- 
type, here designated: NY-herb. Mitten !). 

Anomodon ramulosus Mitt., Trans. Linn. Soc. London, ser. 2, 3: 187. 1891.— 
Type: China. Chekiang, Ningpo, Oldham (lectotype, here designated: NY!). 

Anomodon platyphyllus Kindb., Ottawa Naturalist 7: 19. 1893.— Type: Canada. 
Can. Muse. 256; Macoun Cat. Pt. VI, 171. 

Anomodon sinensis Mull. Hal., Nuovo Giorn. Bot. Ital., n. ser., 3: 118. 1896.— 
Type: China. Shaanxi, "China bor. or., Si-Ku-tzui-san," Giraldi, 1894 (iso- 
type?: FH!). 

Anomodon leptodontoides Mull. Hal., Nuovo Giorn. Bot. Ital., n. ser., 4: 275. 
1897.— Type: China. Prov. Shen-si sept., Schan-kio, Giraldi, Aug 1895 
(FI). 

Anomodon minor n^lx. inaequalifolius Bartr., Bryol. 50: 207. 1947.— Type: Gua- 
temala. Huehuetenango, Rio Ocheval, pr. S. Miguel Acatlan, 29-12-1945, 
Sharp 4857 (isotypes: FH-Bartram! MICH!). 

Plants of median size, forming loose mats, dull green when dry, bright green 
when moist. Main stems creeping, branches to 2.5 cm long, exceptionally to 6 cm, 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 231 

(0.6-) 1.0-1.7 mm thick when dry (leaves included), erect-ascending to arcuate, 
julaceous when dry, primary branches not or seldom branched; central strand of 
stems poorly or not differentiated, pseudoparaphyllia absent. Branch leaves broadly 
decurrent, imbricate, sometimes slightly crisped when dry, erect to spreading, 
complanate, not secund when moist, broadly oblong-lingulate, (1.0-) 1.5-2.3 (-3.0) 
mm long, abruptly narrowed from an ovate base with undulate margins, some- 
times dentate to spinulose at the insertion because of very high papillae and 
sometimes infolded into a rounded apex, generally widened; costa moderately 
robust and prominent on the back, pellucid, ending at (0.04-) 0.08-0.15 (-0.20) 
mm from the apex, often bifid, not obscured by laminal cells near the apex; cells 
on the back of the costa elongate (40-68 |im), with seriate globulose papillae; 
basal hyaline cells extending just 1/3 of the basal leaf portion or less, mostly 
papillose; medial laminal cells hexagonal, 7-12 ^im wide, densely papillose, with 
high and branched papillae. Inflorescences occurring on terminal branches, be- 
yond the distalmost branching points; perichaetial leaves with apical cells papil- 
lose. Seta 4-16 mm. Capsule ovoid; urn 1.5-2.2 mm long, without stomata, annu- 
lus well developed; operculum obliquely short-rostrate, 0.35-0.6 mm; exostome 
reduced, irregular, up to 0.34 mm high, teeth papillose above, not striolate below, 
often trabeculate; endostome very reduced, consisting of a papillose basal mem- 
brane, no more than 2 cells high (ca. 0.035 mm) and segments ca. 0.045 mm high, 
irregular and fragmentary. Calyptra smooth. Spores (11-) 13-18 (-21) |im in dia- 
meter, almost smooth to faintly papillose. Chromosome number: n = 11 (in meio- 
sis. North Carolina: Granzow-de la Cerda, 1988b). 

Distribution (Figs. 19, 20). South and East Asia, from far eastern Russia, 
Manchuria, Korea, and Japan to Burma and the Himalayas; all of eastern North 
America to New Mexico, highlands of Mexico and Guatemala; absent in Europe. 
Anomodon minor occurs in deciduous forests, where it grows on calcareous rocks 
as well as on tree bark and forms loose, thin, somewhat discontinuous mats. In 
North America it requires the most mesic habitats and fruits infrequently. 

Representative Specimens. Bhutan. Oongar, Griffith (as A. integerrimus, herb. Mitlen 
NY). China. Heilongjian: Manchuria, Muling, Sinzew (NY). Chihh: I Yuan K'on, Clemens 5098, 28- 
4-1913 (NY). Hubei: Shennonjia Forest Distr., Jizigon, Sino-Amer. Exp. 882, 25-8-1980 {as A. thraustus, 
NY). Jiangsu: Nanjing, Oixia Mt., Li Zihua 59, 24-6-1963 (NY, MICH), "Kiang Su," S. Henry, 4-11- 
1907 (NY). Jilin: Chang Bai-shan, Guo Qian 22235, 23-9-1981 (as A. integerrimus, c. fr., NY); distr. 
Linjian, Hualazi shan, Guo Qian 7764, 8-9-1963 (as A. planatus, IFSBH); An tu, Erdo Bai xi zien, 
Koponen 36559, 20-9-1981 & 37252, 25-9-1981 (as subsp. integerrimus, JE). Liaoning: distr. Feng 
Cheng, Tong Yuan Bao, Guo Qian 6016 & Nan Man Ling, 6-7-1961 (IFSBH). Xishuangbanna: 
Mengla Bot. Trop. Garden, P. Redfearn et al 33987, 25-12-1986 (as A. viticulosus, NY). Yunnan: Pe yen 
tsin, Sommer 1921 (as A. thraustus, JE); Kweitschou, "ad viam Tschenning-Huang-tsanba," Handel- 
Mazzetti 2037 (10.424), 23-6-1917 (JE). India. Nubra, Thomson, 27-3-1848 (NY). Himachal Pradesh: 
Simla, Thomson 1105, Oct 1849 (NY). Kumaon: Kapkate, Walker 459, Sep 1899 (as A. planatus, c. fr., 
herb. Bartram FH, MICH). Madras: Palni hills, Kodaikanal area, P. Foureau 1268 (as A. plicaefolius, 
NY). Mussoorie: NW Himalaya, Dhanoulti, Bahadru, 10-12-1903 (as A. integerrimus, c. fr., Brioth. 
Levier NY); "Botanical Garden," Gollan, 7-8-1900 (as A. glossophyllus, c. fr., NY); "below Old Bot. 
Garden," W. Gollan, 26-11-1903 (as A. planatus JE). "India orientalis": Bell 119, 120, 367 (ex herb. 
Mitten NY). Japan. Honshu: Aichi pref., Mt. Mikawa-Hongusan, Takaki 3898, 11-9-1947 (NY); 
Bungo, "Kannawa," Iwasaki 1740, 10-9-1916 (as A. thraustus, NY); Higuo pref., Subara-machi, Yatu- 
siro-gun, Tasiro 3340, 16-1-1940 (NY); Mie pref., Mt. Fujiwara, T. Hoguro 34, 19-10-1965 (c. fr., as 
A. subintegerrimus, NICH); Kawakani, Misugi, T. Nakajima, 22-8-1963 (as subsp. integerrimus, exicc. 
Noguchi 952, JE, MICH, NY), Isejingu, {N. Takaki 4467, 28-3-1948 (as A. integerrimus, NICH); 
Mikawa, Miwamura, Chichiiwa, (M Takaki 11333, Y. Horikawa, 26-3-1950 (as A. integerrimus, NICH), 
"South Alps," {N. Takaki 11787, 20-7-1952 (as A. planatus, NICH); Nikko, Kamiji 85, Aug/Sep 1947 
(herb. Bartram FH); Okayama pref., Rashomo, Niimi, Igi, 15-1-1958 (as A. integerrimus, exicc. 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



^^S^'^&'-' 






"A 



feLtt 



Noguchi 654, NY); Os 


ikaPrcr.Takatsu 


i. Hara. T. Naka 


m«, 29-4-1960 (as /I. /M//;r;; subs 


, ,„[ei>er- 


rimus. 


HIRO, MICH 


NY): Tokyo prcf. 


Nishitama-gun, 


Nippara. R. WnUmnhr LW8. H-h 


-1967 (as 


A. ran 


ulosus, NICH 


. Kyushu: Fukuo 


a p,cl.. Kawaia 




Kuman 


oto pref.. Hitc 


voshi, Mavehnrn. . 


an 1948 (as y1 ru 


//(f/as».s. JE. MICH); Mi\a/aki p 


ef., Nishi- 






kl 4806. 5-2-1980 


(as subsp. huegcr 


rimus. NY): Yabakei. Bu/an. /m 


nak, 5.U. 


29-8-1 




Oku: Kochi. Toja 


S. Okamura. 1 


)-12 1905 (Fl. he.b. Bartram 1 




Tosavo 




, Jan 1904 (Filler 


b. Barlram FH). 


Korea.R.SVmcv. 1924(JL):lus 


ni: Faur.c 


72. Ocl 


1901 (Fl). Hv 


ycng Yang: Fcu.ric 














8. 2082. 16,17-8-1960 (as A mi 




NICH) 


Ml. Sam-sun^, W. Hong 1508. 


^_4_ 1 g60 (as A. 




W. Hong 


2816. 2 


-12-60 (as A 


miegcrrmms. NIC 


H). Russia. 'R 


^eio Iskutkensis distr C/eremcl 


ovo Mis 


Sajanc 


SOS orientals. 


L. Barditnov & S 


Lipin. 17-7-195 


(as/l./;/rt/)rtfiM,NICH). 




c 


nada. Manitob 


V. Rod River. Sl > 


orbert. Bird .?26^ 


, 28-5-1959 (c. Ir., DUKF); Park 


and. N ol 


Glcnbc 
9-4-19 


ro, C. Bird 595 


, l.V.5-1961 (c. fr.. 


FH. MICH): Wii 
iw. 10 11-1893 


iipeg, SE of Hadashville, Miwlk-r 
as A. pkKvphvllum. c. fr., NY); 


;;;:;:;;;7 


Mdcou 


,3-10-1892 (; 


s A. obliisifolius. L 


UKE. c. fr.. FH 


Thunder Bav. Sturgeon Bay. Bl 


Ike Tup., 






1981 (MICH); -a 


out Falls of Niagara." Drummond (MICH). Que 




Moiitci 


Im. Park du IV 


ont Tremblanl. F 




19-7-1961 (MICH); Chelsea. ./ 


.Vlaamn. 


17-5-1 


07 (MICH). Mexico, (^lianas: L 




^.?5/.5. 26^-1 945 (DUKE. M EX 


I.MICH. 


NY). Chihuahua: Parquc Nac. Cascada de Basageachi, A. 




MEXU). 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 233 

Guerrero: Chilpancingo, 25 km W, A. Sharp 1052. 22-10-1944 (MEXU, MICH), ca. 22 km W, A. 
Sharp 1096, 27-10-1944 (MICH), Omiltemi, A. Sharp 1132 & 1133b, 27-10-1944 (MICH, & FH, 
MEXU, NY). Jalisco: La Cumbre, 10 mi SW of Autian, H. Crum 458a, Jun 1949, H. Crum 514a, Aug 
1949 (MICH). Morelos: Cerro Tepozteco, C. Delgadillo, 28-9-1991 (MEXU). Oaxaca: San Pablo 
Ayutla, 70 km E of Oaxaca, A. Sharp 4609 et al. (MEXU). U.S.A. Alabama: Jackson Co., Pint 
Rock, /. G.-de la Cerda 1109 & E. Lyons, 4-2-1984 (DUKE). Arkansas: Marion Co., Flippen, L. 
Anderson 12317, 12-6-1953 (DUKE); Stone Co., Ozark Mts., White River, L. Anderson 11596, 18-5- 
1953 (DUKE), Blanchard Springs, D. Demaree 28855, 30-4-1950 (DUKE). Illinois: Laramie River (c. 
fr., as A. obtusifolius, JE); Athens, Hall & Hartman, 1879 (c. fr., JE); Chicago, Roll, 1822 (as A. 
obtusifolius, FH). Indiana: Parke Co., Turkey Run St. Park, W. Welch 11482, 13-11-1937. Iowa: 
Pottawatamie Co., W of Crescent S. Churchill 7109, 18-6-1976 (c. fr., FH). Kansas: Salina, Henry (as 
A. obtusifolius, FH). Maine: Pea Cove, E. Merrill 126, Oct 1898 (FH); Spencer's Woods, Anthony 
2413, 16-4-1888 (c. fr., DUKE); Oxford Co., Norway, Bacon 8942, 5-5-1929 (DUKE). Massachussetts: 
Amesbury, Huntingdon (JE); West Townsend, Thomson, 24-9-1935 (FH). Michigan: Miner's Castle, 
W. Steere 594, Jul 1933 (FH). Minnesota: Winona Bluffs, /. Holzinger, 2-12-1899 (c. fr., FT); Benton 
Co., Henry, 9-1885 (JE); Chippewa Co., pr. Montevideo, /. Holzinger 15-^1901 (c. fr., FH). Missouri: 
Hannibal, Mark Twain's Cave, Davis, 7-11-1911 (FH); Cooper Co., S of Blackwater, A. Sharp, 7-10- 
1938 (FH); Franklyn Co., Merramec Rvr., B. Allen 6109, 30-1-1988 (MICH). New York: McLean 
Bog, Cornell Univ., Kauffman, 25-10-1902 (as A. obtusifolius, c. fr., FH). New Jersey: Closter, Austin 
(FH). North Carolina: Alleghany Co., Roaring Gap, L. Anderson 1273, 18-8-1933 (DUKE); Grand- 
father Mtn., Small & Heller 20, 25-7-1891 (c. fr., DUKE); Jones Co., Coastal Plain, Trent river, 
Trenton, L. Anderson 10819, 20-^1952 (DUKE); Mitchell Co., Roan Mtn., A. Weakley, 1984 (DUKE); 
Orange Co., Eno River State Park, /. G.-de la Cerda 1938, 31-11-1984, (c. fr., DUKE), /. G.-de la 
Cerda 1942, 1943, 7-12-1984 (DUKE). Ohio: Columbus, Schraeder (c. fr., as A. obtusifolius, JE); ibid. 
Sullivant, 1850 (c. fr., JE). Pennsylvania: Huntingdon Co., Spruce Creek, A. & P. Felton & I. G.-de la 
Cerda 1853, 30-9-1984 (DUKE); McKean Co., Burnett 2831, 7-11-1897 (c. fr., FH). South Dakota: 
Day. Co. Wanbay Nat. Windhfe Ref., S. Churchill 7826, 24-6-1976 (FH); Robert Co., Harford Beach 
St. Park, S. Churchill 7807, 23-6-76 (FH). Tennessee: Knox Co., Island Home, A. Sharp 3532, 3-3- 
1935 (FH). Texas: Culberson Co., Guadalupe Mtns., McKittrick Canyn, Moore & Steyermark, 2-7- 
1931 (FH). Vermont: Manchester, Lye Brook, ^. Grout, 23^-1892 (asyl. obtusifolius, c. fr., DUKE); 
Newfane, A. Grout 2416, 25-7-1902 (DUKE). Virginia: Marion, A. Vail & E. Britton, 6-6-1892 (as A. 
viticulosus, MICH); Giles Co., Mountan Lake, Blomquist 3261 , 3-8-1934 (FH); Smyth Co., Whiteface 
Mountain, M. Bucher & A. Weakley, Oct 1984 (DUKE). West Virginia: Pendleton Co., South Branch 
of Potomac, Boardman. 29-5-1938 (DUKE). Wisconsin: Richland Co., Hub City, Hermann 28308, 
21-5-1978 (DUKE). 

Some slender specimens of A. minor with brancti leaves that are not complan- 
ate when moist and a costa that becomes obscured by dorsal laminal cells near its 
end (the latter is the case of Hedwig's type material from Pennsylvania) resemble 
those of A. thraustus. In A. minor the upper part of the costa remains pellucid for 
most of its length and often becomes bifid before ending, whereas in A. thraustus 
the costa is obscured by dorsal laminal cells for the upper 1/3 of the leaf or more. 
Occasionally, in such specimens the upper portion of the leaves may also be bro- 
ken off, as in A. thraustus. The delicate, narrowly Ungulate upper leaf portion of A. 
thraustus is still distinct, whereas that of A. minor is wider and proportionally 
shorter. Larger specimens of A. minor with non-complanate leaves can be told 
apart from A. viticulosus by their more slender habit and smaller leaves, imbricate 
when dry and not secund when moist. Anomodon viticulosus also has more con- 
gested, secund branch leaves. 

Certain specimens, mostly from East Asia, seem intermediate between A. 
minor and A. rugelii. Often specimens of A. minor have strongly spinulose basal 
marginal cells, like those of the auricles of A. rugelii. The lack of auricles and the 
thick papillae on the back of the costa remain good characters for separating both 
species, although it is also true that the presence of obvious papillae on the back 
of the costa in A. minor is not absolutely consistent. Characters that traditionally 



234 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

have been taken as of great taxonomic value for both A. minor and A. nigelii may 
be found mixed, and therefore such value is questionable. Apparently, the only 
characters that remain congruent with each other are the auricles (sometimes 
extremely reduced) and foliose pseudoparaphyllia of A. rugelii; both structures 
absent in all specimens of A. minor. Therefore, these two characters always keep 
both species distinct, in spite of the presence of superficially intermediate forms 
that seem to indicate a blending of these two taxa. 

Iwatsuki (1963) recognized A. minor subsp. integerrimus (Mitt.) Z. Iwats., as 
the taxon occurring in Asia and restricted subsp. minor to eastern North America 
and Central America. According to Iwatsuki, subsp. integerrimus differs from the 
subsp. minor in having very slightly larger spores, some irregularity in the pinnate 
branching pattern of primary branches, as well as asymmetric branch leaves caused 
by a narrowing of the lamina toward the leaf base, therefore becoming shghtly 
falcate-secund. In practice, however, the distinctions barely stand in this rather 
variable species. I have chosen to recognize only a single taxon, A. minor. 

The name A. planatus Mitt, has been used for some smaller forms of ^. minor 
with leaves up to 1.2 mm long. Of all segregates of A. minor this is probably the 
most clear-cut and its recognition understandable. The name has been used, how- 
ever, mostly for specimens of A. minor collected in the southern Himalayas, for 
no apparent morphological reason. Iwatsuki (1963) included A. planatus Mitt, in 
his A. minor subsp. integerrimus. However, mixed in Mitten's type material (NY!) 
are specimens that correspond to A. thraustus (for lectotypification oiA. integerri- 
mus and A. planatus, see the discussion for A. thraustus below). Another taxon, A. 
minor var. inaequalifolius Bartr. (Bartram 1947, 1949), from Guatemala (Huehue- 
tenango: Rio Ocheval, near S. Miguel Acatlan, A. Sharp 4857, 29-12-1945, type, 
herb. Bartram in FH!), is almost identical with Iwatsuki's subspecies. 1 consider 
all these variations within the confines of A. minor. Anomodon ranmlosus Mitt, is 
a robust form that somewhat resembles A. rugelii in habit because of its narrower 
and longer leaves (leaves are slightly incurved, or "curly"). Only the syntype from 
China, Ningpo, collected by Olden is A. minor (one of the others is actually A. 
rugelii). Because the only discriminating character provided in the protologue is 
"fohage on some stems is subcompressed," I consider it appropriate to designate 
the Ningpo specimen oi A. minor (Oldham's collection, NY!) as the lectotype for 
A. ramulosus. 

The name Anomodon obtusifolius Bruch & Schimp., has been extensively 
used until the turn of this century, mostly for North American specimens of A. 
minor; however, because Bruch and Schimper (1848) based it on Hedwig's Neck- 
era viticulosa var. minor, which had already been elevated to specific status 19 
years earlier, it is a superfluous name. Anomodon obtusifolius Bruch & Schimp. is 
not to be confused with A. obtusifolius Mitt, (a nomen nudum), which has been 
applied to some East Asian specimens of ^4. viticulosus. 



Branch leaves broadly oblong-lingulate, constricted near the middle into a 
wide upper portion, apex entire; costa not obscured in its distal portion by laminal 
cells; laminal cells with high mammillae, acute, spiny, generally single, or at least 
with one large central mammilla, with 1-3 smaller, unbranched papillae at its 
base. Capsules immersed or exserted on a seta 2-3 mm long. 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 235 

4. Anomodon abbreviatus Mitt., Trans. Linn. Soc. Bot., ser. 2, 3: 187. 1891. — Type: 

Japan. Joshim, Bisset, Apr. 1886 (holotype: NY!). Fig. 21. 

Anomodon asperifolius Mull. Hal., Nuovo Giorn. Bot. Ital., n. ser., 3: 117. 

1896.— Type: China. Shaanxi, "Shen-si septentr., reg. montanae," Giraldi, 

1894 (isotypes: Fl-herb. Biondi! G!). 

Plants robust, dark green, branching pattern simple, consisting of a creeping 
stem with primary branches 4-5 (-8) cm long, occasionally secondarily branched. 
Central strand of the stem not differentiated. Branch leaves erect-spreading, secund 
when moist, 2.8-3.5 (-4.8) mm long, abruptly narrowed at the middle; leaf base 
oval-lanceolate, upper 1/2 to 2/3 tapering to an acute, occasionally obtuse apex; 
costa strong and prominent at the back, with elongate cells, ca. 30 \xm long, 23 jim 
or less in the distal portion, flexuose toward the apex, ending a few cells below, 
neither bifid nor obscured by laminal cells at the end. Lower cells hyaline, well 
differentiated, extending 1/4 up the basal portion of lamina; laminal cells 13-18 
jim, incrassate and with round lumen in cross section, with a single, unbranched, 
tall, sharply conic central papilla on either surface, somewhat arcuate and point- 
ing toward the leaf apex, as high or higher than the cell's lumen in cross section, 
with 2-3 short, sharply conic, basal papillae. Perichaetial buds on terminal branch- 
es; perichaetial leaves, somewhat longer, more lingulate, and with more elongate 
cells than the branch leaves, with papillose cells. Seta (1-) 2-3 mm; urn (1.1-) 1.3- 
1.8 mm long, ovoid to ovoid-elliptic, annulus present, stomata absent; exostome 
teeth ca. 0.27 mm long, sometimes trabeculate, verrucose above, smooth at the 
base or very slightly striolate; endostome absent or reduced to a few thin seg- 
ments without a basal membrane. Spores 14-24 |im in diameter. Chromosome 
number unknown. 

Distribution (Fig. 22). Japan (in the four larger islands), central and north- 
eastern China, and Korea. 

Representative Specimens. China. Liaoning: Kuandian, Bai shi laji Mtns, W. Buck 23778, 21-8- 
1993 (NY). Shaanxi: Ibaraki, 25-5-1900 (as A. ramulosus, ex herb. Paris FI). Japan. Ibaraki, 25-5- 
1900 (as A. ramulosus, ex herb. Paris FI); Bungo, Mt. Tsurumi, N. Iwasaki 1229, 25-9-1921 (NY); 
Awa, Mt. Tsurugi, M. Kamimura 2621, Aug 1936 (NICH); Higo, Gokanoso, //. Kaneda 212, 30-7- 
1935 (NICH); Kii, Odaigahara, N. Takaki 4885, 3-8-1948 (NICH); Musashi, Mt. Mitsumine, A^. Takaki 
4535, 9^-1948 (NICH). Hokkaido: Iburi, nr lake Shikotan, N. Iwasaki, 18-10-1914 (NICH). Honshu: 
Fukushima pref., Tamura-gun, Mt. Ootakine, T. Higuchi 1564, 13-6-1953 (NICH); Hiroshima pref., 
Mt. Kammuri, S. Nakanishi, 8-11-1960 (JE); Hyogo pref., Mt. Sappico, N of Hmaji, T. Kodama 5^^- 
1962 (c. fr., NICH); Kinkwasan, A. Noguchi, 8-6-1929 (herb. Bartram FH); Kanagawa pref., Hakone 
Mts., Redfearn 768-J-54, 16-5-1954 (DUKE); Kotuke (?), Mt. Okeci, A. Esumoda (?), 15-11-1912 
(Herb. Bartram FH); Mie pref., lishi-gun, Misugi-mura, Kawakami, Nakajima, 4-3-1963 (c. fr., JE), 
Z. Iwatsuki 93, 2/5-12-1963 (NICH); Miyagi pref., isl. Kiuzakan, T. Higuchi 1195 4-3-1953 (NICH), 
Oshika-gun, Mt. Kinka, T. Higuchi 1195, 4-5-1953 (NICH); Musasaki, Titibu-me, M. Yano 5193, 24- 
5-1920 (Herb. Sasaoka NY); Nagano pref., lida, Matsu river, M. Higuchi 9-8-1980 (NICH), Kamiina- 
gun, Mt. Senjogadake, R. Watanabe 18808, 24-7-1973 (NICH), Kisofuku-shima, Schofield 52003, 17- 
10-1972 (DUKE), Takaki 6402, 27-6-1949 (NY), Minami Zumi-gun, Azumi-mura, R. Watanabe 11425, 
13-8-1964 (NICH), Shimoina-gun, Mt. Toyokuchi, R. Watanabe 9389, 9-8-1963 (NICH); Niigata 
pref., Mt. Yahiko, Y. Ikewani, Mar 1948 (JE); Saitama pref., "Japonia centralis," Chichibu, H. Mayr, 
9-5-1890 (ex herb. Broth. FH, FI, NY), Mt. Mitsumina, A^. Takaki 4535, 9-4-1948 (NY), Chichibu- 
gun, Icirikanon, R. Watanabe 14400, 20-8-1968, (c. fr., NICH); Shimano, "Mt. Yatsu," E. lishiba, 26- 
7-1908 (DUKE); Shizuoka pref., Honkawane-cho, Kanzoo, Z Iwatsuki, 15-9-1959 (NICH); Wakaya- 
ma pref., Mt. Gomadan, M. Micutani, 12-9-1949 (DUKE, NY); Yamanashi pref., Ashiyasu, S. Inoue, 
20-8-1974 (JE), Mt. Kita-dake, R. Watanabe 16701, 12-8-1972 (NICH). Kyushu: Kumamoto pref., 
Mt. Kurobaru, K. Mayebara 131, May 1984 (JE, MICH, NY); Gokagoe (Mumimi), Y. Kuwahara 
2163, 13-8-1951 (NY); Miyazaki pref., Koyu-gun, Z. Iwatsuki 2316, 6-5-1976 (NY), Nishiusuku-gun, 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 




FIG. 21. Anomodon abbreviatus Mitt, a-c, branch leaves; d, cross section of stems without 

af in cross section. Based on: a, d, Japan (loc. ignot., DUKE); b, e, f, E. lishiba, Japan, Shimano, 
Yatsu, 26-7-1908 (DUKE); c, i, Yano, 5193, Hondo, "prov. Musasi, Titibu-me," 24-5-1920 (NY); 
, Schofield 52033, Nagano pref., Kisofuku-shima, 17-10-1972 (DUKE). 



Mt. Shiroiwa, N. Suzuki & M. Kakuchi 12 (c. fr., NICH). Shikoicu: Ehime pref., Kamiukena-gun, 
Omogo-kei, R. Watanabe 20205, 25-8-1973 (NICH); Tosa, Mt. Yokogura, M. Kamimura 5100, 24-7- 
1937 (NICH). Korea. Mt. Chii, 1250 m, W. Hong 1225, 16/23-8-1959 (NICH). 

The habit and leaf outline of A. abbreviatus resemble that of A. viticulosus, 
but the characteristic large sharp central papilla on leaf cells of A. abbreviatus 
prevents confusion. Anomodon solovjovii could be mistaken for A. abbreviatus, 
but the latter is more robust, and the papillae are higher (more than the laminal 



GRANZOW-DE LA CERDA: ANOMODONTACEAE 





r— 


^Ir< 


— _r 


/; 


■3^ 




" 






'^ 


^-$ 




.^^CV^r-^ 


^ ▼ H fcuWolium 


„ 


^r^ 








^P 




'4 
















^ 


b\ 


u 


C- 


















'-t 


^■^ 


E;] 


\^J 


D:^^ 


^ 


h 














"^JAx 


- 












^' 









cell's diameter in cross section). Setae are short in A. abbreviatus, but the capsules 
are exserted, whereas in A. solovjovii they are immersed, partly owing to the long 
perichaetial leaves of ^. solovjovii. 

5. Anomodon solovjovii Lazarenko, Rev. Bryol. LichenoL, 5: 45. 1933. — Syntypes 
(n.v.): Russia. "Siidussurisches Gebiet, Murawjow-Amurkij's Halbinsel, 
Umgegenden der Stadt Wladiwostok": "Eisenbahnstation Sedanska," Solov- 
jov, 28-8-1930, "Eisenbahnstation Okeanskaja," Solovjov, 19-8-1930; 
Schamara-Bucht, Schkotowo- Bezirk, Maj-che rivers, Solovjov, 26-8-1930; 
"Im Walde an dem Korjawaja-Fluss," Solovjov, 7-9-1930. [All collected 
on stems of Acer mono and Carpinus cordatus.] 
Anomodon solovjovii var. henaensis Tan, Boufford & Ying, Acta Bot. Yun- 
nanica 18: 69, 1996.— Type: China. Henan Prov., Neixian Xian: N of 
Xiaguan, Boufford et al. 26458-B, 4-6-1994 (holotype: FH!). 



Plants moderately robust; primary branches 3-6 mm long, 1.0-1.2 
when dry, scarcely branched; secondary branches erect. Central strand r 
entiated. Branch leaves erect when dry, 1.4-2.0 mm long, secund wh 



thick 



238 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 2 1 

crowded and spreading at branch apices, broadly oblong-lingulate, abruptly nar- 
rowed at the middle from a broadly ovate base into an obtuse, more often rounded 
apex; hyaline cells at the base reaching less than 1/4 the height of the portion of 
the lamina below the shoulders. Costa strong, ending near the apex (less than 200 
|im), upper portion becoming diffuse, obscured by short laminal cells on the dor- 
sal surface. Laminal cells with a single papilla on both surfaces; papillae arcuate, 
pointing toward the apex, their height less than the cell lumen in cross section, 
never branched but sometimes with 2-3 smaller papillae at the base. Inflorescences 
on terminal branches. Perichaetial leaves much longer than branch leaves, also 
more lingulate and the cells more elongate, with papillose cells. Seta usually less 
than 1 mm long. Capsules immersed, obovate, urn ca. 0.8 mm long, annulus well 
differentiated; peristome teeth papillose above, almost smooth below; segments 
of inner peristome rudimentary on very low basal membrane, finely papillose; 
spores 16-30 |im in diameter, minutely papillose; calyptra cucullate with globulose 
papillae in the upper half. [Sporophytes not seen; description taken from the proto- 
logue (Lazarenko 1933) and from Iwatsuki (1963)]. Chromosome number unknown. 

Distribution (Fig. 20). The Russian Far East (south of the Ussuri river, region 
of Javarovsk), Henan province in China, and Korea. Anomodon solovjovi grows 



Additional Specimens Examined. Korea. Mt. Odai, W. Hong 2563, 262L 24 Oct. 1960 (only 
male plants, NICH). Russia. "Asia Borealis Orientale: reg. austro ussuriensis, fl. Ssputinka," A. 
Lazarenko, Oct 1934 {ex Verdoorn-255, JE, MICH, NY). 

The outline and size of leaves are like those of A. minor, although the two 
species cannot be mistaken, given the tall, arcuate central papilla of leaf cells 
found in A. solovjovil. The areolation and papillae morphologies of A. solovjovii 
closely resemble those of A. abbreviatus. Anomodon solovjovii differs from this 
species by its smaller leaves, smaller laminal cells (14.3 \xm at the most), and by 
having the end of the costa covered with dorsal lamina cells. Thus, the costa 
becomes papillose on the back; it does not appear as clearly defined as in A. 
abbreviatus and ends farther from the apex (more than 200 |im). 

A disjunct population from the interior of eastern China was recently described 
as A. solovjovii var. henanensis (Tan et al. 1996). The variety, which was distin- 
guished by the smaller size of the single papilla of the leaf cells, seems to be well 
within the variabihty of an otherwise poorly collected species, and in my opinion 
lacks sufficient merit to be segregated at the varietal level. 

The species is named after the assistant at the Forestry Department at a 
Russian Far East University, K. P. Solovjov, whom made the first collections. 



Folia rameaha ahquantum grandia, oblongo-lingulata, nonnihil spatulata; apex 
rgine valde serrato, plerumque effractus; costa prope apicem evanescens, sub- 
laminae cellulis occulta. 

Branch leaves narrowly oblong-lingulate to spatulate, abruptly constricted 
ir the middle into a very narrow upper portion; apex strongly and irregularly 
rate, often broken off; costa fading well below the apex, obscured by laminal 
Is; laminal cells with multiple branching papillae. 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 239 

6. Anoniodon dentatus C. H. Gao, Flora Muscin. China Boreali Orientale, 380. 
1977.— Type: China. Kirin, Tchangbaishan, "in truncis arborum silvati- 
carum/' Gao 1231, 28-6-1958 (holotype: IFP!). Fig. 23. 

Plants rather robust; branches pinnate, erect to arcuate, ascending. Leaves 
julaceous and somewhat flexuose when dry, secund or widely spreading to reflexed 
when moist, 2.2-4.0 mm long, narrowly oblong-lingulate, somewhat spatulate, 
abruptly constricted near the middle, base ovate, apex strongly and irregularly 
serrate, bistratose, often broken; margins flat. Costa reaching just short of the 
apex, obscured by laminal cells just above the constriction, becoming hardly visi- 
ble toward the apex. Basal hyaline cells well differentiated, smooth, elongate, with 
more or less sinuose walls, forming an extensive group that occupies half the leaf 
base, gradually becoming shorter and rhomboidal and merging with the upper 
cells; median laminal cells pluripapillose, the papillae weakly branched. Pericha- 
etia present on the most terminal branches, perichaetial leaves with cells papil- 
lose. Sporophytes unknown. Chromosome number unknown. 

Distribution (Fig. 22). Endemic to the northeastern region of the China, known 
only from the provinces of Kirin (or Jilin) and Liaoning, in the mountain range of 
Tchangbaishan, somewhat close to the border with North Korea. According to 
the label of the type collection, this species grows on tree trunks. 

Additional Specimen Examini-d. CHINA. "Kirin [Jilin]: Tchangbaishan," G. Qian 22108, 24-9- 



Anomodon dentatus is a distinctive species, which clearly differs from any 
other species of Anomodon. Although more delicate, it resembles A. abbreviatus 
and A. viticulosus in its leaf shape (lamina abruptly constricted, with a wide, ovoid 
basal portion and a long, narrow upper part). Its leaves, reflexed when dry, and 
the loss of the leaf apices remind of A. thraustus, although it is unlikely to be 
mistaken for any of these species or any other in the genus. The most characteris- 
tic features of A. dentatus are the pinnate branching pattern (more similar to 
species of section Haplohymenium) and shape of the leaves: linear above the 
constriction, wider and strongly serrate at the apex. 

Anomodon dentatus was known only from the holotype (!) and the paratype 
(not seen; from province of Liaoning: "Chean-Schan, ad arborum radices," 1958). 
Upon examination of the type as well as another specimen (originally misidenti- 
fied as A. attemiatus) from the same mountain range as the holotype, I conclude 
that A. dentatus is a well-defined species. 



ion Haplohymenium (Dozy & Molk.) Granzow, comb. nov. 
Haplohymenium Dozy & Molk., Musci Frond. Ined. Archip. Indici, 127. 
1946. — Type: Anomodon sieboldii (Dozy & Molk.) Granzow. 

Plants well-differentiated by the slender, very delicate habit. Secondary branches 
often flagelliform. Branch leaves spreading when moist, <1.2 mm long, apex not 
broken off (except in A. tristis), obtuse to acuminate; costa ending well below the 
apex, 35-70% the length of the leaf, in some species obscured above the shoulders 
by laminal cells. Sporophytes small, urn < 1.0 mm long. Calyptra hirsute. 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME : 




1231 (type), China, province of Kirin, Chang Bai 



GRANZOW-DE LA CERDA: ANOMODONTACEAE 241 

stis (Ces.) Sull. & Lesq., Musci Bor. Amer. 52, 1856. Leskea tristis 
Ces. De Not., Syll. Muse. 67. 1838. Hypnum triste (Ces.) Miill. Hal., Syn. 
2: 478. 1851. Haplohymenium triste (Ces.) Kindb., Rev. Bryol. 26: 25. 
1899. — Type: Italy. Lago Maggiore, Monte di S. Crescenzio, Verbanum, 
1837, C. Cesad (holotype: RO!). 

Anomodon fragilis Wilson, London J. Bot. 7: 275. 1848. Hypnum fragile (Wil- 
son) Miill. Hal., Syn. 2: 471. 1851.— Type: China. Chusan (n.v.). 

Anomodon sinensi-tristis Mull. Hal, Nuovo Giorn. Bot. Ital., n. ser., 3: 118. 
1896.— Type: unknown. 

Anomodon microphyllus Broth. & Paris, Rev. Bryol. 31: 56. 1904. Haplohy- 
menium microphyllum (Broth. & Paris) Broth., Nat. Pflanzenfam. 1(3): 
986. 1907.— Syntypes: Japan. Tsuruga, 14 Aug 1902, Dampax & F. Gaultier 
(n.v.); Tsu-Shima, May 1901, Faurie 1662 (n.v.). 

Anomodon fragillimus Herzog, Beih. Bot. Certralbl., 26: 82. 1909.— Type: Boli- 
via. Santa Cruz, Cerro Amboro, Herzog (holotype: JE!; isotypes: NY! 
US!). 

Haplohymenium scabrum Broth., Rev. Bryol. Lichenol. 2: 10. 1929. — Type: 
Russia. Siberia, prov. Primorskaja, Nikoljsk-Ussurijsk, pr. fl. Panicheza, 
Ssemin (n.v.). 

Haplohymenium formosanum Nog., Trans. Nat. Hist. Soc. Taiwan 26: 43. 
1936.— Type: Taiwan. Tainan, Mt. Kodama, Aug 1932, Noguchi 5867 (n.v.). 

Haplohymenium fragiliforme Nog., J. Jap. Bot. 13: 410, f. 2. 1937.— Type: 
Japan. Sikoku, Mt. Yokogura, prov. Tosa, Aug. 1934, Kamimura 648 (holo- 
type: HIRO) 

Haplohymenium longiglossum P. C. Chen, Feddes Repert. Spec. Nov. Regni 
Veg. 58: 31. 1955.— Type: China. Prov. Szetschwan, Omei schan, Da-ping, 
20-8-1842, Chen 5353 (n.v.). 

Plants very delicate and slender, filiform to wiry in appearence, forming lax 
mats, dull dark brownish green. Branches 0.5-1.5 cm long, 0.3-0.5 mm thick when 
dry; branching pattern simple, main stem creeping, sparsely branched into pros- 
trate primary branches, which rarely branch themselves; central strand not differ- 
entiated. Leaves julaceous when dry, spreading, not complanate when moist, very 
delicate, 0.5-0.9 mm long, with apex (more often ca. 1/3 of the leaf) broken off, 
quite polymorphic among individual plants, lamina narrowing near the middle 
and tapering to an acute to acuminate or obtuse apex; margin flat, not papillose at 
the insertion, crenulate toward the apex because of very prominent papillae; costa 
short and delicate, pellucid below, ending below the middle of the leaf, obscured 
by laminal cells; basal laminal cells elongate, median ones round, ca. 4 [im, with 
multiple prominent papillae, thin-walled, cells of margins becoming mammillose 
toward the apex. Perichaetia rare, on terminal branches, perichaetial leaves oblong, 
acuminate. [Sporophytes not seen, but very similar to those of A. sieboldii, according 
to Noguchi (1957)]. Chromosome number: n - 11 (n = 9+H+h, in mitosis, Japan: 
Inoue, 1965). 

Distribution (Fig. 24). Anomodon tristis is quite rare in Europe, where it is 
restricted to the eastern Alps, but rather common in eastern North America, west 
to the Ozarks and eastern Texas, with disjunct localities in eastern Arizona. It is 
present in Mexico, and reaches the highlands of South America (Bolivia, as Ano- 
modon fragillimus). It also occurs in eastern Asia (eastern China, Japan, Taiwan) 
and Hawaii. No sporophytes are reported from Europe or the New World and are 



. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 




rare in Asia. Anomodon tristis grows on bark of trees in deciduous forests in 
rather montane regions (except in eastern North America). 

Rfpresi-ntaiivi- Si'hciMi-NS. China. Hubei: Shcnongjia Forest Dislr. Ouiujiaping to Mt. Laojun, 
Sino-Amer Exped. 663A (as //. formosanum, NY). Jilin: An-tu, NW Ml. Chang bai, T. Koponcn 



2-1907 (Fl). Taiwan.- 

aipci: Wulai Hsiang. he 

sang )\), Son '["clicou, ( 

Hawaii. Mam: llalc 



umosamwuU\<:\- 



Costa Rica. Hcicciu. ,S sK 


K> of Volc.in Barva. 1 km WNW Rio Cirue 


la Montana. M. Crosin <m4 


W2.?. 3-.V1975 (MO). Mexico. .lalisLO- 


Manantlan. H. Criun 9I4A. 23 


7 1949 (as A. minor. NY). Nuc\o Leon: ( e 


rcy, L. Harvev 1022b. 23-6- ly. 


9 (MICH). Sonora: Bavispe. Canon del le 


8-1940 (MICH). Jalapa: pr. S 


. Miguel el Soldado. ab.wc .lalapa. ( /) 


(MFXU). U.S.A. Ohm: .S////a 


iiu. 1850 (Jh). Pennsylvania: Philadelphia, 


hickon Creek, C. Kaiser. 7-12 


-1911 (DUKE): McKean Co., Ouintuple ('. 


1898 (DUKE). South Carolina 


Pickens Co.. Reedv Cove Falls, pr. Nimmo 


8-1949 (DUKE); Oconee Co.. 


lialooua Km. E loik. /.. Amk-rson Hm. 18- 



Austria. Panowilzer Wald. pr. (Joii/.iam, Luillcshcr^cr ct- Rude: (C): Meran: Vellanerthal. i 
Nicholson, Aug 1904 (INA. herb. Luisicr). Italy. Prov. Cuomo: Borgnana. A. Ariaria. 13-11 - 
JE); "Gallivago in val Giacomo,- PJeiJJer, 2-7-1868 (JE): Verbanum, Valle di Bieno. De Note, 



Nt)vara: Avola, /.. Calvesi. Aug 19.56 (FI). Piamonte: 

(C). Monlorfano. De Notaris Aug 18.59 (FI. RO). 

(JE), pr. Lugano. L. Mari. 3 10 1889 (JE), "Sclva pr. Porya." L. Mari, Jul 8^ 

Notaris 19. 1863 (R); pr. Locarno, M. Jaglli. Apr 1934 (DUKE, JE, MICH). 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 243 

8. Anomodon sieboldii (Dozy & Molk.) Granzow, comb. nov. Leptohymenium 

sieboldii Dozy & Molk., Ann. Sci. Nat. Bot. ser. 3, 2: 310. 1844. Neckera 
sieboldii (Dozy & Molk.) Mull. Hal., Syn. 2: 81. 1850. Haplohymenium 
sieboldii (Dozy & Molk.) Dozy & Molk., Musci Frond. Arch. Indici 4: 
127. 1846.— Type: Japan, von Siebold (holotype: L!, c. fr.). 

Haplohymenium brachycladum S. Okamura, Bot. Mag. Tokyo 25: 31. 1911.— 
Type: Japan. Hondo, Prov. Ecchu, Tsukiokashin-mura in Kaminiikawa- 
gun, Sasaoka, 9-3-1910 (NICH?). 

Haplohymenium okamurae Cardot, Bull. Soc. Bot. Geneve, ser. 2, 3: 281. 
1911.— Type: Japan. Shikoku, Tosa, mont Yokogura, Okamura (PC). 

Haplohymenium gonoi Broth., Ofver. Finsk. Vet. Soc. Forh. 62: 33. 1919- 
1920.— Type: Japan. Shikoku, Tosa, M. Gono 189 (n.v.). 

Haplohymenium pellucens Broth., Ann. Bryol. 1: 21. 1928.— Type: Taiwan. 
Prov. Taihoku, Toihoku, Sasaoka (n.v.). 

Haplohymenium pinnatum Nog., J. Jap. Bot. 13: 410. 1937. — Type: unknown. 

Plants delicate and slender, forming lax to rather thick mats, dull brownish 
green. Branches 0.5-1.5 cm long, 0.3-0.5 mm thick when dry; branching pattern 
simple, main stem creeping, sparsely branched into prostrate primary branches, 
which irregularly branch themselves; central strand not differentiated. Leaves jula- 
ceous when dry, spreading when moist, delicate, ovate-lanceolate with a very faint 
constriction above the middle of the leaf, acute; laminal cells inconspicuously 
pluripapillose; papillae very short, barely branched. Seta 3.5-5 mm. Capsule erect, 
symmetrical, oblong-ovate; urn 0.7-1.0 (-1.1) mm long, stomata absent; annulus 
well differentiated, operculum obliquely long-rostrate; exostome ca. 0.26 mm high, 
smooth below, teeth verrucose; endostome consisting of a basal membrane 3-4 
cells high, segments absent. Spores (15-) 17-21 (-23) jim in diameter, densely 
papillose. Calyptra with thick papillae and hyaline hairs. Chromosome number 

Distribution (Fig. 20). Restricted to Eastern Asia: Japan, Korea, and Taiwan. 

Representative Specimens. Japan. Hondo: Prov. Ettyu, Kaminiikawa-gun, H. Sasaoka, 30-3- 
1916 (as H. brachycladum, NY). Kyushu: Prov Higo, Aida-m., Kuma-gun. K. Mayebara 159, 25-11- 
1928 (NY); Kagoshima pref., Yakushima Is., Suzukawa, W. Schofield 53404, 15-6-1973 (DUKE); 
Miyazaki pref., Minaminaka obi, S. Hattori, Oct 1946 (MICH); Nagasaki Wichura 1472 (c. fr., 
NY). Korea. Cheju, Faurie 331, 1906 (NY). Taiwan, "prov. Sintiku: Kyuko," Y. Simada 341 (as H. 
pellucens, c. fr., NY). 

9. Anomodon pseudotristis (Mull. Hal.) Kindb., Enum. Bryin. Exot. 7. 1888. Hyp- 

num pseudotriste Miill. Hal., Bot. Zeit. 13: 786. 1855. Haplohymenium 
pseudotriste (MiiU. Hal.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907.— Type: 
South AFRica. Cape of Good Hope, Ecklon (n.v.). 

Anomodon filiformis Thw. & Mitt., J. Linn. Soc. Bot. 13: 308. 1873. Haplohy- 
menium filiforme (Thw. & Mitt.) Broth., Nat. Pflanzenfam. 1(3): 1236. 
1909.— Type: Sri Lanka. "Ceylon: Kandy," Thwaites c256 (holotype: NY!). 

Anomodon huttonii Mitt., J. Linn. Soc. Bot. 13: 309. 1873. Haplohymenium 
huttonii (Mitt.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907.— Syntypes: New 
Zealand. North Island, Great Barrier Isl., Hutton & Kirk 66 & 144 (MICH! 
NY!). 

Anomodon exilis Mitt., J. Linn. Soc. Bot. 13: 309. 1873.— Type: South Africa. 
Natal, Attercliffe, 1860, Sanderson (isotypes: NY-herb. Mitten!). 



244 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Anomodon brevinervis Broth., Ofver. Finsk. Vet. Soc. Forh. 33: 107. 1891. 

Haplohymenium brevinerve (Broth.) Broth., Nat. Pflanzenfam. 1(3): 986. 

1907.— Type: Australia. Queensland, Helidon, Dec 1888, Wild 16 (n.v.). 
Anomodon mithouardii Paris & Broth., Rev. Bryol. 29: 97. 1902. Hapiohymenuuu 

mithouardii (Paris & Broth.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907.— 

Type: Vietnam. Entre Lang Son et Dong Dong, Oct. 1901, Mithoiiard (n.v.). 
Anomodon submicrophyllus Cardot, Beih. Bot. Centralbl. 19(2): 128. 1905. 

Haplohymenium submicrophyllum (Cardot) Broth., Nat. Pflanzenfam. 1(3): 

986. 1907.— Syntypes (n.v.): Taiwan. Maruyama, Faurie 6 (14, 15); Kelung. 

Faurie 98; Tamsui, Faurie 102, 107; Hokuto, Faurie 115 [all in 1903]. 
Anomodon stenoglossus Cardot & Ther., Bull. Acad. Int. Geogr. Bot. 18: II. 

1908.— Type: Japan. Liou-Kiou, Naze, 1898-99, Ferrie (n.v.). 
Haplohymenium nakajii S. Okamura, J. Coll. Sci. Imp. Univ. Tokyo 36(7): 26. 

1915.— Type: Japan. Hondo, Prov. Hoki, Mt. Awashima, 31-10-1913, Na- 

kaji (n.v.). 
Haplohymenium obsoletinerve Broth., Ofver. Finsk. Vet. Soc. Forh. 62: 32. 

1921.— Type: Japan. Hondo, Chiba, Gono IV.25. (n.v.) 
Haplohymenium pellucens var. obtusifolium Broth., Ann. Bryol. 1: 21. 1928.— 

Type: Taiwan. Prov. Taihoku, Hill Senjyo, Sasaoka (n.v.) 
Haplohymenium tenerrimum Broth., Rev. Bryol. Lichenol. 2: 10. 1929.— Type: 

Japan. Prov. Ise, Ujiyamada, Tutiga (Herb. Sasaoka, TNS, H?). 
Haplohymenium mithouardii var. viride Ther., Ann. Crypt. Exot. 5: 179. 1932.— 

Type: China. Prov. Fukien, Yanping, Buong kang, 23-6-1925, Chung B.lOl 

(isotypes: MICH! NY!). 
Haplohymenium fasciculare Nog., J. Jap. Bot. 13: 791. 1937.— Type: Japan. 

Sikoku, Mt. Isiduti, prov. lyo, Aug 1935, Noguchi 1169 (holotype: HIRO). 

Plants dark green to brownish, slender, forming lax, pendulous mats. Branches 
0.5-1.5 cm long, 0.3-0.5 mm thick when dry, scarcely branching; central strand not 
differentiated. Leaves somewhat complanate when moist, delicate, ca. 0.5 mm 
long, with apex intact, obtuse to rounded; margin entire, not crenulate; costa 
length variable, from 1/3 to 2/3 of the lamina, covered by laminal cells when 
reaching beyond the middle; lamina cells pluripapillose, papillae short and incon- 
spicuous, rarely or faintly branching. Seta 2-3 mm long. Capsule erect, symmetri- 
cal, oblong-ovate; urn 0.7-1.0 mm long, stomata absent; annulus differentiated; 
exostome smooth below, teeth verrucose. Calyptra with papillae and hyahne hairs. 
Chromosome number unknown. 

Distribution (Fig. 24). Anomodon pseudotristis is rather widely distributed in 
the regions around the Indian Ocean and the SW Pacific; it is one of the few 
species that reaches to the Southern Hemisphere: South Africa, New Zealand, 
probably in Mauritius, SE Asia, including Thailand and Sri Lanka, in addition to 
eastern Asia (E China, Korea, Japan); also in Hawaii. 

Representativh Specimens. South Africa. Cape Prov.: Groenkop Frst. Resv.. M. & C. Crosby 
8080, 25-1-1973 (DUKE); Transkei, rd. Dwcsa to Idutywa, J. van Rooy 2083 (NY). Natal: St. Lucia 
Estuary Forest, J. van Rooy 209 (NY). Transvaal: Marieskop Plantation Natr. Resv., M. & C. Croshv 
7595, 11-1-1973 (DUKE). 

Japan. Honshu: Miyagi PreL, Sendai, lishiha, 16-12-1909 (as H. triste, DUKE); Mie Pref., Nabori 
shi, Akame, W. Schofield 52214, 22-10-1972 (DUKE), Shrine naigu, W. Schofield 52082, 21-10-1972 
(DUKE). Prov. Ettyu, Toyama, H. Sasaoka 4060, 6-7-1928 (as H. sasaokae, DUKE, MICH, NY). 
Kyushu: Fukuoka Pref., Nogata, Y. Kuwahara 2647, (as H. sieboldii, NY); prov. Hyuga, Mt. Aoiga- 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 245 

dake, H. Sasaoka 2379, 26-10-1925 (as H. submicmphyllum, NY). Korea. Cheju, Faurie 246 (ex 
herb. Cardot, as H. submicrophyllum, NY). Thailand. Udawn, Phu Luanh, A. Touw 10379, 7-1- 
1966 (MICH). 

Hawaii. Hawaii: Kohala Mtns., Waipio Valley, Hiilawe Str., W. Hoe 2399, 11-6-1972 (as H. 
triste, MICH). Oahu: N. Waianae Mtns, Waialua Distr., Pahole Gulch SW of Peacock Flat, W. Hoe 
2923, 23-7-1973 (as H. triste, MICH). 

New Zealand. North Island: N Aukland distr., S of Wangaparaoa Pen., Okura Bush, J. Beever 
21-3, 1-1 1-1983 (as A. huttonii, MICH). Matakana, /. Kirk (as A. huttonii, NY). 

Anomodon pseudotristis is similar to A. trisds but differs in having the leaves 
of mature branches mostly with a non-fragile, obtuse to rounded apex. Also, the 
leaf cell papillae are not as prominent as in A. trisds or A. sieboldii. The names A. 
mithouardii and A. exilis have been applied to specimens that seem depauperate 
or etiolated. 



10. Anomodon flagelliformis (Savicz-L.) Granzow, comb. nov. Haplohy 

flagelliforme Savicz-L., Bot. Mater. Inst. Sporov. Rast. Glavn. Bot. Sada 
RSFSR. 1: 98, 101. 1922.— Type: Russia. "Asia Orientalis: regio Austro- 
Ussurensis, in valh fluminis Souczan, pagum Frolovka in flumini Pen- 
saou," 3-8-1913, Boulavkina (holotype: LE). 
Haplohymenium cristatum Nog., J. Jap. Bot. 20: 146. 1945.— Type: unknown. 

Plants dark green, very delicate, pendulous; main stem with regular branches, 
as well as numerous flagelliform ones more than 30 mm long, very slender (less 
than 0.1 mm in diameter), often branching, bearing much reduced leaves. Stems 
without a differentiated central strand; pseudoparaphyllia present, foliose but nar- 
row to uniseriate. Leaves oblong-lanceolate, tapering, only slightly constricted 
above the middle; apex acute to acuminate; costa 0.11-0.19 mm long, extending 
into the proximal ca. 2/3 of the lamina, cehs smooth, covered by lamina cells 
beyond 1/3-1/2 its length, cell walls thick; margin crenulate, dentate near the apex 
owing to protruding marginal cells near the apex, smooth at base; laminal cells 
quadrate to rounded, 6.5-8 |um in diameter, with 4-5 very short to hardly percepti- 
ble, non-branching papillae. Perichaetia abundant, present on terminal branches, per- 
ichaetial leaves longer than branch leaves but similar to them in areolation and 
papillae. Sporophytes not seen. Chromosome number unknown. 

Distribution (Fig. 20). Restricted to the Russian Far East and Manchuria. 



fluminis Majche," Sep 1933, A. Lazarenko (exicc. Verdoorn, 2 nr. 70, DUKE, MICH, NY). 

11, Anomodon longinervis Broth., Hedwigia 38: 243. 1899. Haplohymenium longi- 

nerve (Broth.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907.— Type: Japan. 

Shikoku, Tosa, Makino (H-Br). 
Anomodon aculeatus Broth. & Paris, Rev. Bryol. 31: 56. 1904.- Syntypes 

(n.v.): Japan. Kiu-diu, Ichifusa, Jun 1900, Faurie 1194; Tsu-Shima, May 

1901, Faurie 1604. 
Haplohymenium biforme Broth., Bull. Soc. Bot. Geneve ser. 2., 3: 281. 19n.— 

Type: Japan. Mont Koya, Nakanishiki (Herb. Brotherus). 
Haplohymenium piliferum Broth. & M. Yasuda, Ofver. Finsk. Vet. Soc. Forh. 62: 

33. 1919-1920.— Type: Japan. Hondo, Mt. Akagi, Tsunoda (Herb. Yasuda 

m. 34). 



246 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Haplohymenium crassum Sakurai, Bot. Mag. Tokyo 47: 337. 1933.— Type: 

Japan. Yakushima, Sakurai 2840 (holotype: TNS). 
Haplohymenium splnosum Nog., J. Jap. Bot. 13: 789. 1937.— Type: Japan. 

Kyusyu, Ins. Yakusima, prov. Osumi, Jul 1933, Hasimoto 8640 (holotype: 

HIRO). 

Plants dark green to rusty reddish. Leaves lanceolate, apex acute to obtuse; 
margin flat, crenulate due to the high papillae; costa extending to near the apex, 
strong; laminal cells round, bulging, with 1-3 (-4) acute, spinelike high papillae. 
Perichaetia and perigonia borne at the end of secondary branches. Sporophyte 
delicate; seta ca. 25 mm long; urn globose, 0.8-0.9 mm long, stomata not seen; 
exostome yellowish, 0.2 mm high, endostome absent. Calyptra with hyaline hairs. 
Chromosome number unknown. 

Distribution (Fig. 20). Known only from Japan. 

Representative Specimens. Japan. Kanto, E. lishiba 3045, 8-8-1911 (DUKE). Honshu: Izushi- 
chito Islands, Mt. Hachijo, T. Osada, 30-12-1971, Nakamura, 3-3-1979 (c. fr. as H. pseudotriste, 
MICH); Gifu pref., Kamo gun, Kashao dani, M. Mitzutani 16177, 29-10-1992 (as //. flagelliforme, 
NY); Nagano pref., Kisafukushima, W. Schofield 51943, 17-10-1972 (DUKE). Kyushu: Kagoshima 
pref., Yakushima Is., Onakaido area, W. Schofield 53526, 17-6-1973 (DUKE), Korishima, Chuami 
noike lake, W. Schofield 52917 & Iwatsuki, 20-12-1972 (DUKE); Miyazaki pref., Mt. Oyaji' Z. Iwat- 
siiki & K. Minamidani, 28-10-1976 (MICH). 

The leaf papillae of A. longlnervls are similar to those of A. abbrevlatus and 
A. solovjovif although in A. longlnervls papillae are more numerous (to 3-4 per 
cell), not as high, and often branched. 



Planta tenuis; folia ramealia angusta, tenuia ac/vel fragilia; apex plerumque 
effractus; costa evanescens, brevis, dimidium laminae vix attingens, subter lami- 
nae celluhs occulta. 

Plants slender. Secondary branches terete. Branch leaves spreading when moist, 
delicate, 1.2-1.8 (-2.1) mm long, apex rounded, usually broken off; costa ending 
well below the apex, extending for 60% the length of the leaf, fading far below the 
apex or just above the shoulders obscured by laminal cells on its dorsal side. Urn 
1.3-1.8 mm long. Calyptra smooth. 

12. Anomodon thraustus Miill. Hal., Nuovo Giorn. Bot. Ital., n. ser., 5: 207. 1898.— 
Type: China. "China interior, Shen-si sept., in monte Tuikio-san,"' Sept 
1896, Glraldl (holotype: FI!). Fig. 25. 

Plants rather slender, in loose mats, dark when dry, bright green when moist; 
stems creeping, primary branches erect to ascending, julaceous when dry. some- 
what densely branched and sometimes irregularly pinnate, with secondary branches 
to 20 mm long. Central strand of the stem differentiated. Branch leaves slightly 
crisped when dry, secund, reflexed when moist, rarely complanate, 1.2-1.8 (-2.1) 
mm long, delicate, abruptly narrowing at the middle from an ovate base to an 
(often broken-off) obtuse to rounded apex; margin flat, not papillose-crenulate at 
the insertion. Costa obscured above the shoulders by laminal cells on its dorsal 




Hal. a, branch leaf; 
central strand differentiated; c, cross sections of branch leaves; d, basal cell; 
the costa, fading at the middle of the leaf; f, cells of the base of leaf at the 
Exp. Sino-Amer 274 A, China, "Western Hubei Prov., Shenc 
Canyon," 28-8-1980 (NY); b, c, Exp. Sino-Amer 93 A, id., "Jizigou Canyon," 25-8-1980 (NY); d 
Koponen 37301, China, prov. Jilin, Chang Bai, An-tu, Er-do Bai Xi Zien, 25-9-1981 (NY); f, lishiba, 
Japan, Rikuhii, Mt. Hayoshine (?), Jul 1909 (NY). 

side, ending far below the apex (more than 15 (im below), often just above the 
shoulders, occasionally bifid, with in-line large papillae on the dorsal side, where 
exposed larger than those of the laminal cells; basal pellucid cells extending up to 
half the basal portion of the lamina, or above; laminal cells with multiple high and 
branched papillae. Inflorescences on terminal branches, beyond the distalmost 
branching points. Seta 5-13 mm. Urn 1.3-1.8 mm long, stomata absent, annulus 
differentiated; operculum short-rostrate, ca. 0.7 mm long. Exostome teeth well 
developed, ca. 0.3 mm high, papillose to the base; endostome papillose, basal 



248 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

membrane very low (2-3 cells, or ca. 0.045 mm high), with very reduced segments 
(not more than 0.035 mm long). Spores green, 16-19.5 ^m in diameter, finely 
papillose. Calyptra glabrous. Chromosome number unknown. 

Distribution (Figs. 19, 26). Previously, A. thraustus has been considered re- 
stricted to Japan and eastern China. Gier (1980) cited a specimen from Mexico, 
and I have also found herbarium material of v4. thraustus from that country (named 
as A. minor). Its distribution area, therefore, becomes dramatically enlarged. Such 
disjunction agrees with that of A. minor, although in the New World A. thraustus 
occurs more to the south than does A. minor. 

Additional Specimens Examined. China. Heilongjian: ^Lesser Khingan Mtns," Shuan-zi-he, 
Guo Qian 6672, 15-7-1963 (NICH). Hubei: Shennongjian Forest distr., Miaogou, Sino-American 
Exp. 274A, 28-9-1980 (as A. viticulosus, NY); Jizigou, Si,w-American Exp. 93A, 25-8-1980 (as A. 
minor, NY). Jilin: "Chang Bai, An-tu, Er-do Bai Xi Zien," T. Koponen 37301, 25-9-1981 (as A. 
minor subsp. integerrimus, NY); Jian, Dayu shan. Guo Qian 7802, 8-9-1963 (as A. rugelii, IFSBH); 
Wang Qing Xian "Wang-chiu," Daxinancha, Nangou, Guo Qian 8575, 16-8-1973 (IFSBH). Tibet: 
Ya-Dong, Zang Mu 56, 29-5-1975 (NICH). India. Himachai Pradesh: Simla "7000," T. Thomson 
1172, 1847 (as A. obULsifolius var., Wils., syntype for A. integerrimus, NY-herb. Mitten), Simla "7000, 
8000," T. Thomson 1126, Oct 1849 (mixed with syntype of A. phmatus, herb. Mitten, NY). Kumaon: 
Dahli valley, /. Duthie 6359, 7-8-1886 (as A. plcmaUis, MICH). Uttar Pradesh: "We.stern Hymalaya^ 
Seven Oaics estate," Mussoorie, W. Gollman, 2-10-1900 (as A. glossophyUus, NY). "India Oriental," 
Bell 187 (as A. integerrimus, NY). Japan. Honshu: Hiroshima pref.. Tojo-cho. Taishaku. M. fJigu- 
chi, 7-10-1978 (HIRO); Iwate pref., Rikuchu: Mt. Hayachine, E. lishiba, Jul 1909 (as A. minor. 
NY). Korea. Keisyo nando, Mitsuyo, /V. Maki 2985, 31-7-1931 (NICH); Sauer. 1924 (as A. minor. 
JE); Mt. Ciak, W. Hong 5874, 16/17-7-1961 (as A. integerrimus, NICH);'Mt. Sokri, W. Hong 2801 
21-12-1960. Fusan, Faurie 241, 1906 (ex herb. Cardot NY). Nepal. "In Himalayae reg. temp.." 



Tetjukhe, L. Bardunov, 29-1 1-1961 (NICH), pr. Partizansk, L. Bardunov & V. Tchardantzeva 16-9- 
1964 (NICH), Schkotovskyi, pr. Podiapolsk, L. Bardunov, 9-9-1974 (NICH); "regio Ussuriensis," 

(as A. minor, FH), Utsisheibo (Manchuria), Vladivostok, P. Sinzer, 13-03-1905 (as A. minor, c. fr., ex 
herb. Brotherus FI); "Far East, Lake Khanka, pr. village of Lusanovo," L. Bardunov. 23-8-1962 
(NICH); "Sibiria: Primorskaja, Okaenskaja," E. Sinova, Sep 1926 (as A. minor, MICH). 

Mexico. Hidalgo: Jacobo, V. Chase 7405, 11-7-1939 (Bartram herb. FH). Jalisco: Slopes of La 
Ferreria, above Manantlan, H. Crum 914, 926a, 942, Jul 1949 (as A. minor, MICH). San Luis Potosi; 
45 km E Rio Verde, betw. Ciudad Valles and S. Luis Potosi, C. Delgadillo 4-12-1978 (as A minor 
MEXU). 

Anomodon thraustus has often been reduced to synonymy within A. minor; 
however, Sakurai (1954), Iwatsuki (1963), Iwatsuki and Noguchi (1973), Noguchi 
(1974) and other Japanese authors recognize it as a species. Unquestionably, A. 
thraustus is closely related to A. minor, but they differ, in addition to the more 
delicate habit oiA. thraustus, in the following foliar characters. l)The leaves oi A. 
thraustus are almost invariably not complanate when moist but clearly reflexed, 
spreading in all directions. 2)The costa is short and entirely obscured by laminal 
cells beyond the shoulders, ca. 1/3 from the apex. 3)The leaf apices are often 
broken off, as in A. tristis or A. dentatus. 4)The cells are papillose-crenulate at the 
insertion, almost spinulose, in A. minor but entire in A. thraustus. Also, the cen- 
tral strand of the stem is differentiated in A. thraustus, but not in A. minor. The 
exostome is made of well-developed and regular teeth, and the endostome is 
papillose with segments ca. 3 cells high in A. thraustus, whereas in A. minor teeth 
of the exostome are irregular, and the endostome is smooth and practically with- 
out segments. 




Further confusion resulted from Mitten's having specimens of A. thraustus 
mixed with his type material for A. integerrimus and A. planatus (lectotype for A. 
integerrimus: Nepal, Wallich, Mitten Herbarium, NY; lectotype for A. planatus: 
Simla, 1126, 10/49, "A. obtusifolius small barren state W." Mitten Herbarium, NY, 
here designated). His protologues for these two taxa do not describe the morphol- 
ogy of A. thraustus, nor do the sketches of leaves drawn on the herbarium packets 
containing the syntypes of A. integerrimus and A. planatus depict the leaves of A. 
thraustus but rather those of A. minor. Anomodon integerrimus and A. planatus 
are synonyms of ^. minor, not oi A. thraustus. Mitten's mixed types are probably 
the cause of much of the confusion surrounding the identity of both taxa. 



II. An 



I subgenus Pseudoanomodon Limpr., Laubm. Deutschl. 2: 774. 
-Type: Anomodon attenuatus (Hedw.) Hiib. 



Stems and primary branches branching in a stairway fashion 
ary branches irregularly pinnate, profusely fasciculate. Leaf ape?< 
nate; laminal cells rhombic in most taxa. 



250 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

13. Anomodon rostratus (Hedw.) Schimp., Syn. Muse. Eur. 448. 1860. Leskea 
rostrata Hedw., Sp. Muse. 226. 1801. Hypnum rostratum (Hew.) P.-Beauv., 
Prodr. 69. 1805.— Type: U.S.A. "circa Lancaster, Pensylvaniae," Muhlen- 
berg (holotype: G -herb. Hedw. Schwaegr.!). Fig. 27. 

Plants forming thick dense mats, glaucous, light to dull yellowish green. Stems 
prostrate, profusely branching, with primary branches 0.5-2 cm long, ca. 1 .0 mm 
thick when dry, clustered, with a fasciculate, pseudo-verticillate branching pat- 
tern, secondary branches erect, stem with a differentiated central strand of smaller 
cells, pseudoparaphyllia absent. Branch leaves julaceous, imbricate when dry, erect 
when moist; leaves ovate-lanceolate, acuminate, slightly constricted in the distal 1/3, 
up to 1.2 mm long, margins revolute, ending in a hyaline hair-point several cells 
long (to 0.2 mm or more) and one cell thick; costa pellucid, ending well below the 
apex, ca. 10 |im, often somewhat flexuose above, with high papillae in one hne at 
the back, sometimes slightly branched, almost scabrid at the base, the group of 
basal cells small, very few, with sinuose walls; medial laminal cells rhomboidal, 
scarcely isodiametric at the basal 1/2-2/3 of the lamina, multipapillose, with papil- 
lae slightly branched. Perichaetial buds beyond the last branching nodes; pericha- 
etial leaves elongate, long-acuminate, smooth. Seta dark red, 7-13 mm long, some- 
times shorter. Capsule short, elliptic; urn 1-1.5 mm long, ovoid to oval-cylindrical, 
with stomata at the base, annulus of 2 rows of cells; operculum obliquely rostrate, 
0.8-1.1 mm long; peristome well developed, exostome 0.2-0.3 mm high, teeth 
trabeculate, with a yellowish base, striolate to the middle or more, apex conspicu- 
ously papillose; endostome well developed, with tall basal membrane, 7 cells high 
or more, as high as the exostome, segments keeled, long and slender, 0.9-1.1 mm 
high, papillose; cilia absent. Spores 11.5-15 |im in diameter, densely papillose. 
Chromosome number: n = 10 + x/y (in meiosis, North Carolina: Granzow-de la 
Cerda 1988b). Additional illustrations in Crum and Anderson (1981) and Schu- 
mackeretal. (1982). 

Distribution (Figs. 15, 18, 28). Circumboreal montane, from central Europe 
south to the Pyrenees and Italian Alps, east to the Caucasus; widely distributed 
throughout eastern North America, from Newfoundland to Mexico and Guatemala, 
also in Bermuda, Hispaniola, and Jamaica. Iwatsuki (1963) excluded this species 
from Japan and, in general, from eastern Asia, given that most records are misi- 
dentifications of A. longifollus (Yano 1957) or Claopodium pellucinerve (Rei- 
mers 1936). Anomodon rostratus usually grows in deciduous forests, sometimes in 
rather open spaces, as well as secondary forests, where it forms dense and thick 
mats, often very extensively, on vertical rocks, calcareous in Europe (Schumacker 
et al. 1982) but also acidic in North America. In the eastern United States, it is 
almost always found at the base of deciduous trees or on their bark, on soil, and, 
less often, on fallen logs. 

Representative Specimens. Georgia. Tzenis Tzchali rvr.. "Svaina Dadianosum. Hippi. intLT Muri 
et Lentechi," Sommier & Levier, 29-7-1890 (NY). 

Canada. British Columbia: Eagle Pass Mtns., Big Eddy of Columbia Rvr., nr. Revelsloke, W. 
Taylor 5310, 11-8-1921 (MICH). New Brunswick: Albert Co.. Fundy Nat. Park, Dickson Falls R 
Ireland 10744, 2-7-1967 (MICH). Newfounland: Notre Dame Bay, New World Isl.. Cobbs Arm G 
Brassard 10506 et al., 19-10-1975 (NY). St. Barbe S Distr. SE of Parsons Pond, J. Hancock & C 
Butler 468, 3-7-1972 (MICH). Nova Scotia: Colchester Co., Five Islands, R. Ireland 17222, 10-7-1974 
(MICH). Ontario: Grey Co., Inglis Falls, nr. Owen Sound, H. Crum 11027, 3-8-1961 (MICH). Ottawa 
Co., / Macoun, 20-10-1893. Manitoulin Isl., Tamarack Point, C. Grassl 6219, 19-7-1932 (MICH). 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 




FIG. 27. Anomodon rostratus (Hedw.) Schimp. ; 
portion of peristome. Based on: a, d, e, F. Artaria, Italy, Como prov., Bucca del Pertuso, Blc 
7-1897 (NY); b, Hermann 26045, Mexico, Nuevo Leon, Monterrey, Sierra Madre Oriental, 22- 
(NY); c, Boros, Hungary, "Comit. Veszprem, rivi Gerence, pr. Baconybel," 3-4-1959 (NY). 



Queb 



Williams 10882. 25- 
Coban, P. Standley 
6-2-1945 (herb. Be 
NY); S of San Crisi 



r S. Pedro 
; 90729, 26-3-1941 (herb. Bartram FH), 90130 (MICH). El Quiche: A. Sharp 2434, 
artram FH). Mexico. Chiapas: Las Casas, A. Sharp 3521, 25^-1945 (US, MICH, 
tobal, R. Dull 16, 8-10-1966 (NY). Hidalgo: Chapulhuacan, A. Sharp 1541a, 1542a, 
1580, 7/8-12-1944 (DUKE, MICH, NY, US); Jacala, A. Sharp, 17-12-1962 (US). Nuevo Leon: Sierra 
Madre Oriental, Monterrey, Chipinque, F. Hermann 26043, 2-7-1975 (NY), 26037b, 27-1-1975 
(MICH); Sierra Madre, Monterrey, C Pringle 10460, 29-5-1908 (JE, US). Puebia: pr. Zacapaxtla, A. 
Sharp 4185, 17-10-1945 (MICH). San Luis Potosi: "above Xilitla," A. Sharp 5866, 5868, 21-4-1946 
(MICH, NY, US). Tamaulipas: Sierra Madre, W of Victoria, H. Crum 1831, Aug 1950 (US). Jalapa: 
Acutlzingo, pr. El Puerto, A. Sharp 629, 670, 6/13-9-1944 (MICH). U.S.A. Alabama: Madison Co., 
Monte Santo Co., T. Bryson 898, 3-1-1976 (MICH). Arizona: Sta. Cruz Co., Sta. Rita Mtns., White 
House Canyon. /. Haring 3472 & H. Haskell, 7-5-1945 (MICH). Arkansas: Drew Co., Wilmar, D. 
Demaree 24239, 10-4-1943 (MICH). Colorado: Chandler Canyon, T. Brandegee, 1876 (NY). Mon- 
trose Co., W Paradox Creek, W. Weber et al. B-43564, 1-6-1973 (NY). Connecticut: East Haven, J. 
Allen, 7-9-1882 (MICH). Florida: Jackson Co., Marianna Caverns, R. Schornherst 1508, 9-3-1940 
(MICH). Georgia: Stone Mtn.,J. Small, Jan 1894 (MICH); Demorest, O. VanHilse, Feb 1913 (MICH). 
Illinois: Pope Co., Bell Smith Springs, NW of EddyviUe, D. Vitt 7407, 25-5-1973 (MICH). Iowa: 
Fayette Co., Brush Creek Canyon St. Park, S. Gradstein & D. Horton 6392, 16-4-1987 (MICH). 
Kansas: Douglas Co., 14 km S-SE of Lawrence, S. Churchill 9370, 11-4-1978 (MICH). Riley Co., 14 
km SW of Manhattan, 5. Churchill 9401, 14-4-1978 (MICH). Kentucky: Trigg Co., Cadiz Mill, A. 
Harvill2000, 10-8-1946 (MICH). Maine: Somerset Co., Moxie Falls, B. Allen 9327, 11-7-1990 (MICH). 
Massachusetts: Weston, J. Lowe, 24-12-1903 (DUKE); Worcester Co., Bolton, H. Greenwood 155, 3- 
6-1933 (DUKE). Michigan: Cheboygan Co., Mill Creek, N. Shiskoff 10-33, 11-7-1980 (MICH). Huron 
Co., Port Austin, /. Schnooberger 1754, 8-10-1938 (MICH). Marquette Co., Presque Isl., F Hermann 



ONTR. UNIVERSITY ( 



D 









-^ > i5^ 







^ 



i .'i -2^'^- 



barn canvon area. S. Churciull 9216, 26-5-1977 (MICH). Tcincssc. Wa\n 


( o., (iienroc 


< B.anc 




\luu-p 405. : 




(MICH). Texas: Dallas Co.. nr. Dallas. ./. Boll. 1874-80 (NY) I >hcrtv Co ^ 




el-nul ( 


Webster & R. Wilbur 6(iH, 3-7-1959 (MICH. NY). Virginia: Sinvlh Co.. Moui 


t Jefferson Na 


1. 1-ores 


/. G.-de In Ccrdu IHH7. 6-10-1984 (c. Ir.. DUKE). West Virginia: Greenbrier 


Co.. SofWhil 


- Sulphi 


Springs, /.. Anderson 24752, 16-7-1986 (DUKE). Wisconsin: Richland Co. 






12537, 4-7-1974 (MICH). 







1 997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 253 

Bermuda. Harrington House, S. Brown 558 (NY). Abbots cliff: E. G. Britton 1877, 13-9-1813 
(US). Haiti. Mtns above Kenskoff, F. Mackaness 153, 19-1-1944 (MICH); btw. Savanna Zombi and 
Satnon, F. Mackaness 70, 18-3-1944 (MICH). Dept. de Sud'est: Massif de la Salle, W. R. Buck 9347, 
21-11-1982 (NY). Jamaica. Abbey Green: W. Maxon 10061, Jul 1926 (NY); vicinity of ArntuUy, 
Orcutt 3185, 3-10-1927 (herb. Bartram FH); vicinity of Abbey Green, W. Maxon 10083, Jul 1926 
(US). Blue Mountains: G. Orcutt 2854, 5-9-1927 (US, FH). "Westhalia: (Ceddar Hurst)," C. Orcutt 
3762, 1-11-1927 (US). 

Austria. Baden: Wien, 7. Benoger (?), 1871 (NY). Salzburg, D. Sauter (NY). Tirol: Merau, Milde 
(NY). "Austria inferor": Isaden (?), Jiiratzka, 30-6-1867 (NY). "Niederosterreich": Baumgartner, 
30-4-1891 (NY). France. Haute-Garonne: Prat de Joueou, "environs de Luchon," Dismier 281, 14- 
8-1928 (MA). Germany. Bayern: "Nagelfluh, pr. Shafflarn," Molendo, 1860 (NY), /. Familler, Aug 
1906 (MICH). Hungary. Comit Veszprem: "rivi Gerence, pr. Bakonybel," A. Boros, 3-4-1954 (NY), 
Nagy-Penzlik cave, mt. Somhegy, A. Boros, 4-10-1937 (MICH). Italy. Como: Buca del Pertuso, 
Blevio, F. Anuria, 15-7-1897 (NY); "Nino, inter Blevio et Como," F. Artaria, 20-9-1897 
(NY). Romania. Transylvania: distr. Fagaras, M. Peterfi, 10-8-1921 (NY). Spain. Girona: vail de 
Vidra, Les Llances, /. G.-de la Cerda 2323a & M. Sam, 10-1-87 (c. fr., BCB, MA), Rec de I'Espirau, 
M. Sanz, 7-3-1986 (BCB), Vilallonga de Ter, F. Lloret 33.3.7, 13-3-1986 (BCB). Huesca: Ordesa, R. 
Dull, 23-5-1960 (BCB) (see Granzow-de la Cerda, 1988a). Switzerland. Ticino: Lugano, Gandria, 
17-9-1885 (c. fr., ex herb. Boissier NY); Campione-Bissone, P. Conti, Oct. 1893 (MICH). 

Anomodon rostratus is well distinguished from all others in the genus by having 
the shortest branch leaves, ending in a hair point. The leaf areolation is also more 
lax; the basal portion of rhomboidal cells extends beyond half the leaf length. The 
species somewhat resembles A. longifolius in having long, acuminate leaves with 
revolute margins. Nonetheless, differences between both species are more than 
obvious: A. rostratus has leaf cells with multiple papillae, a uniseriate acumen 
throughout most of its length (or biseriate at the base) with cells completely 
distinct from those of the lamina (i.e., hyaline and much longer than wide). Also, 
A. longifolius, along with A. rugelii, are the only species that have pseudoparaphyllia. 

Anomodon rostratus fruits profusely in North America, but mostly remains 
sterile in Europe, although a specimen bearing two sporophytes was found in a 
population from NE Spain (Granzow-de la Cerda 1988a). Based on observations 
by Schumacker et al. (1982), it can be inferred that these are the only sporophytes 
ever reported from Europe. According to those authors, the absence of sporo- 
phytes is due to the absence of male gametophytes in Europe. My observations 
revealed that in North America male gametophytes are extremely scarce, even in 
populations with abundant sporophytes (Granzow-de la Cerda 1988b, 1989b). 

Iwatsuki stated that A. rostratus var. japonicus Sak. belongs to Trachypus 
humilis (Iwatsuki 1963: 58). 

14. Anomodon attenuatus (Hedw.) Hiib., Muse. Germ. 562. 1833. Leskea attenua- 

ta Hedw., Spec. Muse. 230. 1801. Hypnum attenuatum (Hedw.) Sm., Fl. 

Brit. 3: 1279. 1804.— Type: Germany, sine loc, c. fr. (syntypes: G-herb. 

Hedw.-Schwaegr.!). Fig. 17c-e. 

Hypnum fulvum Brid., Muse. Rec. 2(2): 188. 1801. Hypnum attenuatum var. 

fulvum (Brid.) Steud., Nomencl. Bot. 2: 252. 1824.— Type: Herb. Ph. Com- 

merson (not located). 
Anomodon wrightii Miili. Hal., Hedwigia 37: 262. 1898.— Type: Cuba. "Rocks in 

a ravine," Wright (isotypes: in Cuban Mosses exicc. nr. 69, MICH! NY!). 

Plants forming dense mats, dull yellowish green. Primary branches 1.5-3 cm 
long, 0.8-1 mm thick when dry, prostrate to arcuate, irregularly pinnate, secund 
when dry, complanate when wet, mostly attenuate, branching pattern often con- 
sisting of several orders of successive branching, in a stepwise fashion; without a 



254 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

differentiated central strand, pseudoparaphyllia absent. Branch leaves slightly nar- 
rowed at the middle, with the portion above the constriction mostly lanceolate, 
acute, sometimes obtuse or slightly apiculate, 1.2-2 (-2.2) mm long; margins flat, 
entire from the base up (although cells moderately mammillose at the insertion); 
apex sometimes denticulate, with cells at the tip, along the margins, generally 
hyahne and somewhat larger and less papillose than the laminal ones; costa pellucid, 
abruptly ending a few cells below the apex, sometimes fading in youngest leaves; 
cells on back of costa usually less than 23 |Lim long, sometimes up to 35 |Lim long; 
hyahne basal cells poorly differentiated, sometimes chlorophyllose, but extending 
more than half the length of leaf base; cell walls somewhat sinuose, porous near 
the costa; upper laminal cells quadrate, 6-8 |im, pluripapillose, papillae branched. 
Perichaetial buds never present beyond the last branching points; perichaetial 
leaves abruptly narrowed, costa reaching well into the distal half of lamina, end- 
ing near the apex in the innermost leaves. Seta 10-15 mm. Urn 1.6-2.8 mm long, 
with stomata at the base; annulus not differentiated; operculum obliquely short- 
rostrate, ca. 1.3 mm long; exostome striolate below, teeth irregular, 0.35 mm high, 
yellow, becoming white with age, trabeculate and more or less finely papillose 
toward the apex; endostome also papillose with high basal membrane (3-4 cells 
high), segments keeled and well developed, to 0.03 mm long, cilia absent. Spores 
10-13 |im in diameter, densely papillose. Chromosome number: n = 10 + x/y (in 
meiosis, North Carohna: Granzow-de la Cerda, 1988b). 

Distribution (Figs. 12, 13, 15, 20). Present throughout Europe (except in Ire- 
land, Portugal, and the Mediterranean islands), Turkey, and from the Caucasus to 
Kashmir; very abundant in eastern and midwestern North America, from New- 
foundland to Manitoba, south to Florida, Mexico, Guatemala, and Belize; Cuba, 
Jamaica, and Hispaniola. Anomodon attenuatus grows on bark, mostly at the base 
of trees, and also on soil and rocks. Populations in North America produce sporo- 
phytes abundantly. 

Representat[ve Specimens. India. Kashmir: Ferozenpur Nullah, Gulmarg, R. R. Stewart 10572a, 
2-9-1929 (NY). Russia. Krasnodorskyi: Teberda, Abramov, 21-8-1955 (NY). 

Canada. New Brunswick: Restigouche Co., St. Jean-Baptiste-de-Resligouche, R. Ireland 14478, 
14-8-1970 (MICH). Newfoundland: St. Barbe, Doctor's Hill, R. Tuomikoski 4618, 29-7-1948 (MICH); 
Humber, Benoit's Rvr., R. Tuomikoski 5412, 15-8-1949 (MICH). Nova Scotia: Annapolis Co., E 
shore of Kejimkujik Lake, R. Ireland 12554, 3-8-1968 (MICH); Cumberland Co., Wentworth, R. 
Ireland 10681, 7-9-1967 (MICH). Ontario: Grey Co., Sydenham Rvr., Inglis Falls. H. Crum 11034. 3- 
8-1961 (MICH); Lake Superior, Thunder Bay District, W. B. Sdiofiekl 27244, 1-7-1965 (as A. minor, 
DUKE), W. end of Albert Lake. C. Carton 14878. 18^-1972 (MICH). Quebec: Pontiac Co., SE 
corner of Pare de la Verendrye, R. Ireland 16125 & T. Koponen, 15-6-1972 (MICH). Guatemala. 
Alta Verapaz: Rio Carcha, pr. San Pedro Carcha, P. Standley 92104. 12^-1941 (NY, MICH). Baja 
Verapaz: pr. Patal, A. Sharp 2947, 3-3-1945 (MEXU). EI Quiche: Neboj, A. Sharp 2429. 6-2-1945 
(MEXU, US). Jamaica. Arntully, C. R. Orcutt 3840, 12-11-1927 (DUKE). Mexico. Chiapas: San 
Jose del Lago, A. Sharp 3572, 3-5-1945 (US); "Las Casas," A. Sharp 3516, 26-4-1945 (MEXU, NY); 
"Strasse Ocosingo," San Cristobal, Eggers & Frahm 29, 10-3-1979 (NY). Durango: El Salto, A. Sharp 
1889, 31-12-1944 (MICH). Hidalgo: Chapulhuacan, E. Frye 2874a (MICH). Nuevo Leon: Hacienda 
Vistahermosa, 35 mi S of Monterrey, L. Harvey 1052, 1055, 25-6-1939 (MICH). Oaxaca: pr. S. Pablo 
Ayutla, A. Sharp et ai, 23-12-1972 (MEXU). Puebla: Rio Necaxa, pr. Huachinango, A. Sharp 3143, 
27-3-1945 (MICH). San Luis Potosi: Xilitia, A. Sharp 5836, 21-4-1946 (US), C. Delgadillo et al, 17- 
12-1976 (MEXU). Sonora: Bavispe, Caiion del Temblar, E. Phillips 596, 19-8-1940 (MICH). Tamauli- 
pas: pr. Rancho del Cielo, Gomez Farias, C. Delgadillo, 1-5-1967, ibid. A. Sharp, 1-8-1974 (MEXU). 
Jalapa: "19 km NW Jalapa," W. Rauh & R. Diill 31, 15-10-1966 (JE); NW of Caotepec, pr. Conso- 
lapan, L. GilJudrez 60, 23-5-1975 (NY); "Strasse Orizaba-Tehuacan," Eggers & Frahm 38, 23-.3- 
1979 (NY). U.S.A. Alabama: Hale Co., Rock Hollow, Havana P.O., H. Wagner et al. 71167, 30-12- 
1971 (MICH). Arizona: Apache Co., Apache Reservation, W. Phillips, 2-9-1949 (DUKE, MICH). 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 255 

Arkansas: Newton Co., Little Buffalo Rvr., L. Anderson 11969, 29-5-1953 (DUKE, MICH). Colo- 
rado: Boulder Co., Boulder Canyon, W. Weber, 30-3-1963 (MA). Connecticut: Valley Forge, Upper 
Glen, W. Stair & W. Allen, 17-6-1939 (MICH). Georgia: Stephen Co., Toccoa Falls, R. Lampton 
1804, 15-5-1965 (MICH). Illinois: LaSalle Co., French Canyon, Starved Rock, W. Hinton, Apr 1970 
(MICH). Indiana: Turkey Run Hollow, D. Richards et al. 5896, 23-11-1947 (MICH). Kansas: Doniphan 
Co., Missouri Rvr., S. Churchill 10860, 30-6-1979 (MICH). Louisiana: Natchitoches Par., Cunning- 
ham Brake, J. Bruza 643, 3-3-1978 (MICH). Maryland: Prince Georges Co., Beltsville, Paint Branch, 
F. Hermann 16414, 27-11-1960 (MICH). Massachussets: Amherst (as A. viticulosus, FH). Michigan: 
Mackinac Co., Caffey Corner, H. Crum, 9-8-1968 (MICH); Washtenaw Co., Cascade Glen, G. Web- 
ster 577, 2-5-1959 (MICH). Mississippi: Scott Co., K. Rogers 9357, Jan 1974 (MICH). Missouri: Barry 
Co., 2 mi N of Eagle Rock, P. Redfearn 26059, 4-5-1969 (as A. viticulosus, MICH); Franklin Co., 
Meramec Rvr., B. Allen 6108, 30-1-1988 (MICH). Minnesota: Goodhue Co., B. Johnson, 3-5-1953 
(MICH). New York: Columbia Co., Old Chatham, S. Smith 3119 & B. Reilly, 15-5-1961 (MICH). 
North Carolina: Transylvania Co., Horsepasture River, /. G.-de la Cerda 1938, 31-11-1984, 1286, 13- 
6-1984 (DUKE); Graham Co., Joyce Kilmer Mem. Park, /. G.-de la Cerda & K. Renzaglia, 3-10-1987 
(c. fr., DUKE); Macon Co., /. G.-de la Cerda 1236, 8-6-1984 (DUKE), Nantahela River headwaters, 
/. G.-de la Cerda 1260, 11-6-1984 (DUKE), Crow Creek, /. G.-de la Cerda 1399. 19-6-1984 (DUKE); 
Orange Co., Eno River St. Park, /. G.-de la Cerda 1999, Mar 1985 (DUKE). Pennsylvania: Hungtin- 
don Co., Spruce Creek, A. & P. Felton & 1. G.-de la Cerda 1854, 30-9-1984, (DUKE); CUnton Co., 
Singlebranch Creek, L. Stair, 11-7-1944 (MICH). Vermont: Wallingford, G. Kennedy, 3-8-1907 
(MICH). Virginia: Madison Co., Fisher's Gap and Rose Rvr., /. Schnooberger 4740 & F. Wynne, 15- 
6-1944 (MICH). West Virginia: Greenbrier Co., H. Burtt, Jun 1971 (MICH). Wisconsin: Willburn, /. 
Roll 1635, 16-12-1888 (as A. attenuatus var. brevifolius, G). Richland Co., Richland Center, M. Nee 
12059, 13-6-1974 (MICH). 

Cuba. Pinar del Rio: Bafios San Vicente, Britton 7446, Aug-Sep 1910 (as A. wrightii, 
NY). Dominican Republic. Santiago: orillas del Guanajuma, San Cristobal, /. Jimenez M-7, 16^- 
1946 (NY). La Vega: Salto de Aguas Blancas, SW of Constanza, W. R. Buck 7900, 27-4-1982 (NY). 

Austria. Kustenland: Gorz, K. Loitlesberger, May 1911 (c. fr., NY). Byelorussia. L. /. Savicz 
(NY). Denmark. Ganloese: "near Copenhagen," W. C. Steere & Holmen, 21-7-1954 (c. fr., NY). France. 
Haute Garonne: Prat de Joueou, Dismier 286, 3-7-1928 (MA). Haute Savoie: Douvaine, "Boise de 
Ballaison," E. Bonnot 714, 31-12-1962 (BCB). Germany. Oberbayern: Bayerbrunn, Lewis, 9-11- 
1845 (NY). Hungary. "Praeillycum, Sopianicum: Transdanubia Mts., Mecsek, Valle Nagymelyvoel- 
gy," /. Gyorffy, 25-7-1923 (BCB). Italy. "Silva Montello" (NY); "Venetia, Monte Baldo" (ex herb. 
Thumen NY). Novocomo: Flaglearo, supra Varese, F. A. Artaria, 11-4-1897 (NY). Norway. Akers- 
hus: "Emmestad in Vestby," P. St0rmer, 10-4-1950 (BCB). Poland. "Western Carpathians": Cieszyn, 
Dziegielow, /e^rz<?)A:o & Zarnowiec, 7-10-1981 (BCB). Krakow-Czestochowa Upland, "Sokole Gory," 
15 km SE of Czestochowa, H. Bednarek -Ochyra & R. Ochyra, 21-6-1986 (MICH). Russia. Karelia: 
inter Petrosavodsu et Kosalma, H. Baukoh, 20-7-1973 (NY). Spain. Barcelona: Santa Fe de Mont- 
seny, Casas, 26-3-1969 (BCB). Castellon: Fuente La Reina, Herrero-Borgohon 341, 16-5-1982 (MA). 
Girona: vail de Vidra, /. G.-de la Cerda 2345 & F. Lloret, 28-3-87 (BCB). Huesca: Caiion de Anisclo, 
Fanlo, /. G.-de la Cerda 2216 et al., 15-8-1986 (MA). Lleida: val d'Aran, Portillo de Bossost, C. 
Casas, 3-6-1977 (BCB). Navarra: Selva de Irati, A. Ederra, 25-3-1982 (NAU). Rioja: Sierra de la 
Demanda, hayedo de Tobia, M. Brugues, 28-6-1977 (BCB) (see Granzow-de la Cerda, 
1988a). Sweden. Uppsala: Mosen, Oct 1866 (NY). 01and: St, Dalby, No. 974 3/6-8-1969 
(MICH). Switzerland. Zug, Bamberger (NY). Geneve: "bois de Frontenoy, pres de Geneve," 15-1- 
1853 (NY). Ukraine. "Ditio Chmelnitski, Kamenets-Podalsk," M. Kotov, 23-7-1948 (NY). "Ditio 
Kioviensis, pr. Bila Tzerkva," D. Zerov, 24-5-1926 (NY). 

Anomodon attenuatus is a polymorphic species. Plants from the Pyrenees, for 
instance, are very reduced in size. The distinctive attenuate branches, for which 
the species is named, are sometimes arcuate and seldom ascending; however, 
sometimes in some specimens the branch apices become slightly capitate and 
incurved. Robust plants of A. attenuatus might be mistaken for A. viticulosus, 
because both have acute leaf apices. They can be told apart by their branch 
morphology: ascending and robust in A. viticulosus, and prostrate and complanate 
in A. attenuatus. Also, the leaf shape is different in both species: lanceolate and 
without constriction in A. attenuatus, but lingulate and abruptly narrowed from a 
broadly ovate base in A. viticulosus. The costa ir 



256 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

thick, seriate papillae on the dorsal surface. A consistent character in fertile plants 
of A. attenuatus is the absence of gametangia beyond the most recent branching 
points (i.e., they are never formed on the youngest branches). 

The name A. wrightii has been used for specimens from the Caribbean islands 
(the type specimen collected from Cuba), but they agree with A. attenuatus. 

15. Anomodon giraldii Mtill. Hal., Nuovo Giorn. Bot. Ital., n. ser., 3: 91. 1896.— 
Type: China. "China interior, provincia Shen-si septentr.," 1894, Giraldi. 
det. C. Muller, nr. 1032 (holotype: FI!). Fig. 29. 

Plants moderately robust, forming loose mats, yellowish green to brown; plants 
irregularly pinnate; central strand not differentiated; primary branches to 6 cm 
long, 0.9-1.3 mm thick when dry, profusely branched; secondary branches attenu- 
ate, inserted close to each other on primary branches in a whorl-hke fashion, or 
successively in a stairlike fashion; branches arcuate, recurved, and often thickened 
at the end; flagelliform branches with reduced leaves present. Branch leaves erect 
to erect-spreading when moist, 1.5-2.5 mm long, 1.0-1.4 mm wide, strongly decur- 
rent, secund, somewhat falcate near the apex and concave to keeled because of a 
prominent costa, ovate-lanceolate, acute, widest at 1/3 from the base, margins 
almost straight from that point to the apex, without constriction, usually revolute, 
plicate in the basal half. Costa strong, prominent on the abaxial side, pellucid, 
flexuose in the distal half, percurrent or ending very near the apex (<0.1 mm), 
thickened at the end; cells of the costa rather short (almost as short as laminal 
cells, rarely >llp,m long, 17 |xm closer to the base), on the dorsal surface smooth 
or, mostly toward the apex, with numerous, randomly arranged minute papillae. 
Laminal cells pellucid, not isodiametric but irregularly rectangular, basal cells not 
papillose, walls strongly sinuose, often porous, gradually merging with the upper 
cells; upper cells with somewhat sinuose walls, papillae multiple, simple and short 
on both surfaces. Perichaetia few, with abundant paraphyses, absent beyond the 
most recent branching points. Perichaetial leaves long and narrow, smooth, costa 
absent or very reduced. Seta 7-20 mm. Capsules cylindrical, erect or slightly curved; 
urn 2-3 mm, with some stomata, annulus absent; operculum rostrate, ca. 1/3-2/3 
the length of the urn; exostome ca. 0.3 mm high, smooth at the base, teeth finely 
papillose above; endostome consisting of a basal membrane several cells high, 0.1 
mm, segments rudimentary. Spores (12-) 16-23 (-25) pm in diameter, smooth to 
very finely papillose. Chromosome number unknown. 

Distribution (Fig. 26). Japan (the four large islands), China (Hunan, western 
Hubei, Chensi, Manchuria, Yellow Sea coast), Korea (Cheju Island included), Far 
Southeast Russia; restricted to the Yellow Sea and Sea of Japan region as well as 
to the Chensi Mountains {locus dasslcus). According to Iwatsuki (1963), A. giraldii 
usually grows on rock substrates, acidic and basic, including stone walls, also on 
tree trunks and rotten logs. This species can be dominant in epiphytic communi- 
ties in montane deciduous forests, where it forms thick and extensive mats. Sporo- 
phytes have seldom been collected (Iwatsuki recorded only six fertile collections). 



2 Specimens. China. Hubei: "Shennongjia, Forest District," Sino-Amer. Exp. 665, 
3-9-1980 (NY, FH, HIRO, MICH). Hunan: Hsikwangschan mines, pr. Hsinhwa, Handel-Mazzetti, 
May 1918 (NY, FH, JE). Jiangxi: Mt. Lu-Shan, C K. Tung, Jun 1955 (JE). Liaoning: distr. Benxi, 
Yangchi, Guo Qian 8435 & Zhang Guang Chu, 6-7-1963 (IFSBH); distr. Feng Cheng, Feng Huang 
Shan, Guo Qian 6889, 29-5-1963. Japan. Etehu, Ohsho-mura, H. Sasaoka, Apr 1917 (JE), Ohyama, 
Kaminii kawa, Sasaoka (c. fr., herb. Chamberlain FH); Mt. Bandai, E. Uemalsu, May 1908 (JE); 



GRANZOW-DE LA CERDA: ANOMODONTACEAE 




i Mull. Hal. a-( 

ives; h, group of hyaline basal cells of the 

;ar the apex. Based on: a, Iwatsuki 9574, 

[, Miyazaki-ken, Higashi-usuki-gun" 26-5-1981 (NY); b, i, H. Sasaoka, Japan, Echizau, 

Goka, Jul 1921 (FH); c, d, Lazarenko, Russia, "Asia Bor. Orien. in regio ; 

a" Oct. 1934 (FH); e, lishiba, Japan, Sendai, 7-9-1907 (DUKE); f-h, S. Murai, Japan, ' 
, Mt. Taiheizan, Ninbetsu Nat. Forest," 1-4-1968 (NY). 



25« CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Miyadzu, Faurie 1518. 1901 (NY); Sendai, E. lishiba, 7-9-1907 (DUKE). Hokkaido: Ishikari distr., 
Sapporo, Tenguyama, T. Koponen 21488, 8-07-1971 (herb. Chamberlain FH), Kamuikotan, A. Nogii- 
chi, 21-8-1951 (JE, MICH, NY). Honshu : Akita pref.. Akita, Mt. Taiheizan, S. Murai, 1-4-1968 
(NY); Aomori pref., Shirikari, K. Kikizaki 5, 18-.'i-1977 (NICH). "Yunosawa valley," K. Kakizaki, 
27-7-1976 (DUKE); Ettyu, Mt. Kurobe. H. Sasaaka, 7-6-1930 (herb. Chamberlain FH); Gifu pref., 
Ono-gun, Nomugi-toge pass, //. Katsuyama 13166, 25-5-1968 (NICH); Hiroshima pref., Iwaidani 
valley, R. Watanabe 23496, 24-8-1976 (NICH); Inaba, Ml. Kyushu-zan, H. Ochi, 17-4-1955 (herb. 
Bartram FH); Iwate pref., Mt. Aoba, S. Nakamishi, 21-7-1960 (JE), Shinoehei-gun, Iwaizumi-cho, R. 
Watanabe 17035, 23-8-1972 (NICH), Rikutuyu, Mt. Iwate, G. Joba 224. 29-6-1928 (NY); Kyoto 
pref., Oecho, Naigu, M. Mizutani, 7-10-1967 (NY, c. fr., JE, MICH); Mt. Komona. K. Miirela, 9-6- 
1918 (herb. Sasaoka DUKE); Mie pref., Inabe-gun, Mt. Fujiwara-dakc, //. Kalsiiyama 4593, 20-4- 
1969 (NICH); Miyagi pref., Isl. Kinkazan, M. Mizutani, 6-9-1959 (NICH); Nagano pref.. N. Takaki, 
27-6-1949 (NY), Kisofukushima., Schofield 51961, 7-10-1972 (DUKE); Niigata pref., V. Ikegami 
Mar 1948 (JE, MICH); Saitama pref., Chichibu-gun, Ashi-ga-kubo, R. Watanabe 23576, 11-10-1976 
(NICH); Shimane pref., lishi-gun, Akaana Shrine, M. Higuchi 13939, 17-5-1987 (c. Jr.. HIRO); Tot- 
tori pref., Mt. Daisen, T. Koponen 21841, 20-07-1971 (herb. Chamberlain FH); Yamagushi pref., 
Iwakuni, Shiroyama, Schofield, Ando & Seki 54512, 16-4-1973; "Echizand, Goka," H. Sasaoka, Jul 
1921 (herb. Brotherus FH); "Hida, Mt. Kotori," H. Sasaoka, Aug 1918 (herb. Brotherus FH); Taisy- 
ayu, A. Noguchi, Jun 1932 (herb. Bartram FH). Kyushu: Bungo, Mt. Kudy, A. Noguchi, 4-11-1934 
(herb. Bartram FH); Fukuoka pref., Asakura-gun, Y. Kitwahara 2361, 23-11-1951 (NY); Kumamoto 

1952 (DUKE). Kohnox, K. Maebara, 3810 20-9-1955 (NICH), Mt. Aso, H. Shojima 1575, 25-4-1970 
(NICH); Miyazaki pref., Higashi-usuki-gun, Z. Iwatsuki 9574, 26-5-1981 (NY); Nagasaki, Wichura 
1461 (as A. acutifoliiis, NY); Oita pref., Mt. Kurodeke, pr. Kakushimizu, Z. Iwatsuki 290a, 30-10-61 
(c. fr., NICH). Shikoku: Ehmie pref., Kami-ukena-gun, Tengu-Kogen platteau, Z. Iwalsuki 5245 & H. 
Kiguchi, 23/24-8-1978; Tokushima pref., Miyoshi-gun, Mt. Tsurugi, R. Watanabe 19134, 22-8-1973 
(NICH). Korea. Kong Ouen to, Faurie 4052, Jul 1907 (FI), Mt. Chiak, W. Hong 5875 5876, 5879, 
5878 (c. fr.), 16/17-7-1961 (NICH), Mt. Haura, W. Hong 2246, 7-8-1960 (NICH), Mt. Kwan-ak, W. 
Hong 1518, 27-3-60 (NICH), Mt. Odai, W. Hong 2567, 2570, 2608, 24-10-1960 (NICH), Ml. Sokri, 
W. Hong 2795, 2817, 21-12-1960 (NICH), Isl. Dagelet, W. Hong 6121, 6123, 26/29-7-1961 (NICH). 
Cheju: "He Guelpaert," Faurie 303, 1906 (NY). Russia. Ussuriisk, "Regio austro ussurensis: flumen 
Ssputinka," A. Lazarenko ex herb. Verdoorn 254, Oct 1934 (FH, JE, MICH), ibid., L. Bardunov, 9- 
9-1962 (NICH); distr. Partizanskyi, pr. Lozovyi, L. Bardunov & V. Tcherdantzeva, 13-9-1974 (NICH). 

Flagelliform branches are common in A. giraldii. Some branches, however, 
bear reduced leaves or are leafless at the middle, while the leaves near the apex 
are of a regular size. Anomodon giraldii resembles A. attenuatus in its branching 
pattern and attenuate branch apices, but it is readily distinguishable from any 
other species in the genus by its ovate-lanceolate leaves without constriction, its 
lax areolation of rectangular leaf cells with sinuose walls, especially near the leaf 
base, and the multiple, unbranched, rather small papillae. Anomodon giraldii and 
A. attenuatus are, moreover, allopatric in distribution. 

This species is named after Father Giuseppe Giraldi, an active collector in 
China and Japan, who provided Carl Muller with the type specimen. 

16. Anomodon longifolius (Brid.) Hartm., Handb. Skand. Fl. ed. 3, 300. 1838. 
Pterigynandrum longifolium Schleich. ex Brid., Mant. Muse. 128. 1819. 
Leskea longifolia (Brid.) Spruce, Muse. Pyren. n. 87. 1847. Hypnum longi- 
folium (Brid.) Mull. Hal., Syn. 2: 474. 1851.— Type: unknown. Fig. 30. 
Anomodon armatus Broth., Hedwigia, 38: 243. 1899.— Type: Japan. Hokkaido, 
29-4-1894, Oiwake, Miyabe & Tokubuchi 392 (syntype: FH!). 

Plants somewhat slender, growing in rather thick mats, yellowish green, with 
branches 20-30 (-60) mm long, 0.5-1 mm thick when dry. Stems with filamentous 
and foliose pseudoparaphylha, rhizoids abundant above (especially in Japanese 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 259 




weden, Sala, "sub. Pterog." (NY). 



material); branches profuse, fasciculate or irregularly pinnate; secondary branches 
usually attenuate, often flagelliform, apices frequently club-shaped and/or recurved, 
central strand of stem well differentiated. Branch leaves 2.1 mm long (to 2.8 mm 
or more), 0.3-0.5 (-0.6) mm wide at the base, quite densely arranged, secund, 
clearly falcate, almost recurved when dry and somewhat plicate at the base, in 



260 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

robust forms; in smaller, more delicate plants the leaves 1.2-1.6 mm, julaceous 
and more imbricate when dry; branch leaves decurrent, narrowing more or less 
abruptly below the middle (0.2-0.35 mm at the widest above the shoulders) and 
tapering into a subulate apex, often 2-3 cells wide at the end; margins revolute 
near the shoulder, entire to serrulate and often with shorter and wider cehs near 
the apex; costa yellowish, somewhat decurrent, protruding and making the leaf 
carinate, often scabrous on the back in the upper half, percurrent, with dorsal 
cells noticeably shorter and incrassate, as long as those of the lamina (7-11 |am); 
basal cells near the costa generally not hyaline, smooth, longer than those of the 
rest of the lamina, the basal group extending less than 1/4 the length of the leaf 
base; medial laminal cells ca. 15 |Lim long, with a single, central, rather high papilla on 
both surfaces, which never exceeds the cell's diameter. The youngest perichaetia 
and perigonia on the most distal portions of the branches usually with minute or 
flagelliform branches extending beyond the inflorescences; perigonial leaves 
smooth, areolation lax; cells of perichaetial leaves isodiametric and smooth near 
the apex. Seta yellow, 4-8 mm; urn 1.2-1.7 (-2.0) mm, elongate, cylindrical, exothe- 
cial cells hexagonal-isodiametric, stomata absent; annulus absent; operculum conic 
to short-rostrate, ca. 0.5 mm, somewhat asymmetric, obtuse; exostome irregular, 
0.25-0.30 mm, striolate at the base; endostome rudimentary (0.08-0.13 mm), gen- 
erally reduced to a short pellucid basal membrane, 1-2 cells high, sometimes 
bearing a few scattered very slender segments of the same height. Spores 12-17 
|Lim in diameter. Chromosome number unknown. 

Distribution (Figs. 14, 15, 18). Throughout most of Europe, except in Ireland, 
Portugal, and the Mediterranean islands, to the Caucasus; also in Japan (Hok- 
kaido and Honshu) and Russia's Far East; absent from the New World. Anomo- 
don longifolius grows generally on rocks but sometimes on tree bark. 

Representative Specimens. Azerbaijan. Bakuriani, Lazarenko & Ahramov, 5-9-1956 (NY). Georgia. 
"Tzenis Tzchali, Svania Dadianosum, Hippi, inter Muri et Lentec," Sommier & Levier, 29-7-1890 
(NY). Japan. Asorezan, Faurie 2095, 1902 (c. fr., NY); Borezan, Faurie, 1 1-10-1902 (herb. Bartram 
FH). Hokkaido: Kamuikotan, A. Nogiichi, Aug 1951 (MICH); Rikuku, Mt. Hakkoda, F. lishiha, Jul 

1907 (MICH), Mutsu, S. Okamura, 31-7-1913 (NICH); Ishicari, Yamabe, Z. Jwaisuki, 7-8-1971 
(exicc. Noguchi 1202, MICH, NY); Sapporo, Isiskaki, N. Iwasaki 4704, 16-10-1914 (NY); Si-Kan, 
Kamikawa-gun, TakenaiKi 1432, 10-03-1919 (NY). Honshu: Kominato, Faurie 49, 9-12-1889 (FI); 
Mt. Shirouma, N. Takaki, 1-08-1949 (NY); Fukushima pref., Otakine, Oniana, T. Higuchi, 3-10-1958 
(exicc. Noguchi, NY); Nagano pref., Shimotakai-gun, Sakai-mura, S. Okamura, 28-10-1913 (NICH), 
S. Uto (?), 23-10-1913 (FI), Kijimadaira, K. Saito, 3-8-1972 (MICH); Shimano, Mt. Tagakushi, 1-8- 

1908 (FH); ToUori pref., Mt. Mitoku, H. Ochi, 12-6-1960 (NY). 

Austria. Markenstein, E. Berroyer, 2-5-1869 (MICH). Wien, E. Berroyer, 16-3-1869 (NY). 
Salzburg: bei Hallstadt, /. Fraehlich, 16-3-1869 (NY). Steyermark: Altenessen, Fh. Suse, 12-8-1910 
(ex herb. Uggla NY). Belgium. Frahan, Delogne 82, Mar 1869 (NY); Namur, Feyr, May 1874 (NY). 
Comunne de Yvoir: Bauche, De Sloover 30206, 1-4-1979 (NY). Byelorussia. Minsk, L. Savicz, 23- 
5-1923 (NY). Finland. Pohjois-Pohjanmaa, Tervora. Peu, F. Ulvinen, 16-7-1977 (NY). "Ostrobot- 
nia kajonensis": Paltamo, M. Findvquisl, 18-7-1931 (NY). Pohjois-Saavo: Kuopio, SiJkalahti, A.J. 
Huuskonen, 3-9-19661 (NY). "Savonia borealis": Kaavi, Likosaari, A. Ferasvuori, 22-6-1917 (NY). 
Tohmajarvi, Havukkavaara, S of Vuorenpaa, R. Fagcrsten & M. Haapa.saari 4095 (MICH), 4097. 2- 
10-1984 (as A. viticulosus, FH). Siilinjarvi: Iso-Pettainen. Kourulahti Bay, R. Fagersten 776. 30-7- 
1974 (MICH). France. Haute Savoie: Reignier, Puejet, Nov (NY). Isere: Baigns-la-Romaine, De 
Sloover 35014, 25-7-1981 (MICH, NY). Jura: Jouhe, Foulotiere, P. Cuynet 114-116, 5-5-1960 (BCB). 
Savoie: Aux-les-Bains, /. Milller (NY). Germany. Pr. Baierbrunn, P. G. Lorentz, 1860 (c. fr., ex 
herb. Uggla NY). Allgau: Hinterstein, Bargundele Bach, De Sloover 16494, 8-9-1973 (NY). Bayern: 
Berchtesgaden, Fh. Suse, 6-9-1895 (ex herb. Uggla NY); Nagelfluhblochen, Bayernbrunn, Munchen, 
A. Holler, Oct 1861 (NY). Mont Rhon: Geheeb, 1879 (NY). Thuringia: Mustberg, H. Moller, Apr 
1859 (ex herb. Uggla NY). Wartburg, V. Russe (NY). Hungary. Comit. Pest, Madencia, pr. Ocsa, 
Boros, 14-10-1928 (NY). Comit. Bereg: pr. tarpa, A. Boros, 15-3-1961 (BCB). Comit. Heves: Mte. 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 261 

Agasvar, pr. Nagybatony, Boros, 6-5-1928 (NY). Poland. Monies Gorce, Kamienica, Lisowski, 18- 
4-1959 (Bryot. Polon. 1114 NY). Kielce: Monies Gory, Swielokrzyskie, Lisowski 64989, 1-6-1964 
(NY). Krosno: "W Bieszczady mnls., Kurnikow Beskid," R. Ochyra, 7-7-1977 (BCB, MICH). Bialys- 
tok: Puscza Augustowska, Lisowsky 68685, 30-10-1958 (NY, MICH). Schlesien [Silesia]: Breslau, J. 
MiUle (NY). Suwalki: Smolniki, Lake laczno, R. Ochyra, 17-5-1979 (MICH), Lake Haneza, R. Ochyra, 
18-5-1979 (MICH). Russia. Gobersdorf, Friedland [Pravdinsk], /. Milde, 30-7-1868 (NY). Karelia: 
"Karelia ladogensis, Suislamo, Saarisuonvaara," A. J. Hueskonen, 24-8-1935 (NY); "Karelia onegen- 
sis, Uksjarvi, Kossalmi," M. J. Kotilainen, 9-6-1942 (NY). Moscow : Ruza distr., Ozhigovo, M. Igna- 
[ov, 11-6-1988 (MICH), Kunzowo, "pr. Mosquam," Meyden, 22-4-1899 (NY). Kursk: Karolscha Dean, 
Lawdanski, 19-9-1902 (NY). "Sibiria," Jenisei, Nikulina, H. Arnell, 18-7-1876 (NICH). Spain. Bar- 
celona: Sla. Maria de Besora, C. Casas, Dec 1964 (BCB). Navarra: Sierra de Urbasa, A. Ederra, 13- 
3-1982 (BCB, NAU). Sweden. Angermanland: Sabra, Grovhall, H. W. Arnell, 1-6-1870 (NY). Narke: 
Svenshyttan, C Hartman, 31-10-1874 (NY). Oland: Hogbyn, Vyksjo, Hj. Moller, 25-7-1917 (DUKE), 
Mariehamn, H. Moller, 26-7-1928 (NY). Veslelad Lund, K. Holmen, 8-6-1969 (NY). Oslergolland: 
Marmosbruket, P. Olsson, 23-6-1875 (NY). Sala: "sub Pterog" (c. fr., ex herb. Sullivanl NY). Skane: 
Oveds su, Fruelid, G. Alberg, 15-6-1919 (ex herb. Uggla NY). Smaland: Greunabruket, Vretaholm, 
A. Arven, 18-5-1914 (NY); Talberg, N. Schentz (herb. Thumen NY). Torshunda, H. Moller, Apr 1859 
(herb. Uggla NY). Uppsala: Gollsundabergen, /. Zettersstedl 95, 7-5-1878 (MICH), Uppsala, 1844 
(herb. Thumen NY). Vaslmanland: Saia Gruva, 31-5-1879 (DUKE). Switzerland. Pr. Zurich, P. 
Culmann, Apr 1885 (NY). Valais: Loneche-les-Bains, H. Fossell, Sep 1887 (NY). Zug: Prope Zug, 
Bamberger 6522, 1855 (NY). Ukraine, pr. Kiev, Holossijevo, N. Wagner, 16-3-1919 (NY). United 
Kingdom. Gloucestershire, Forest of Dean, H. N. Dixon, 20-7-1903 (NY). Wales: Somerset, C H. 
Bisnstead, 4-12-1886 (NY). 

Anomodon longifolius can hardly be mistaken for any other species in the 
genus. Distinctive characters are the presence of pseudoparaphylha (shared only 
with A. nigelii), the incrassate, isodiametric, irregular or shortly rectangular cells 
on the back of the costa, and the irregularly striolate exostome (in some speci- 
mens alternating with finely punctate areas, sometimes smooth above, sometimes 
divided into two filaments, which are generally verruculose-papillose). Specimens 
of Leskeella nervosa (Brid.) Loeske are rather often found misidentified as A. 
longifolius, probably due to their remarkably similar appearance. The leaf cells of 
Leskeella, however, lack papillae, its costa does not extend toward the apex as far 
as in A. longifolius, and its marginal cells are shorter and more elliptical; the 
leaves of L. nervosa are more crowded, julaceous (not secund), and more acumi- 
nate than subulate. 

In some specimens of A. longifolius, branches of two kinds may appear: robust 
branches with leaves to 2.1 (-2.8) mm long, crowded, secund, clearly falcate, 
almost recurved when dry and somewhat plicate at the base, as well as more 
dehcate branches, with leaves 1.2-1.6 mm long, julaceous, and more imbricate 

Anomodon armatus Broth., described from Japan, coincides in every feature 
with western material and is here included in A. longifolius. 



rpetineuron (Mull. Hal.) Cardot, Beih. Bot. Centralbl. 19(2): 127. 1905. Ano- 
modon sect. Herpetineuron Mull. Hal, Flora 73: 496. 1890.— Type: Her- 
petineuron toccoae (Sull. & Lesq.) Cardot. 
Bryonorrisia Stark & Buck, Brittonia 38: 52. 1986.— Type: Bryonorrisia secunda 
(Dixon & Badhw.) Stark & Buck [^Herpetineuron acutifolius (Mitt.) 
Granzow]. 



rather robust, forming open t 


o thick mats 


. Stems prostrate, sparsely 


branches rather arcuate, circin; 


ate or, often. 


attenuate toward the apex. 



262 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

with a well-differentiated central strand of smaller cells, without paraphyllia or 
pseudoparaphyllia. Branch leaves incurved, secund when dry, erect, not complan- 
ate when moist, ovate-lanceolate, tapering from the base, without an obvious 
constriction; margins flat, serrate in the distal 1/3-1/4; apex sharply acuminate; 
costa single, strong, pellucid, smooth on the back, not obscured by laminal cells, 
ending sharply at or near the apex, not bifid at the tip; laminal cells practically not 
differentiated throughout the lamina, rectangular to rhombic, slightly longer than 
wide, smooth, sometimes prorulate, walls somewhat thickened. Seta 8-16 mm. 
Capsule erect, cylindrical to elhptic or almost symmetrical, urn 1.5-2.8 mm, annu- 
lus well differentiated; peristome relatively well developed, exostome striolate at 
the base, endostome papillose with a basal membrane 1^ cells high, segments 
0.2-0.4 mm high, papillose. Calyptra smooth. Chromosome number unknown. 

Herpetineiiron is undoubtedly the genus most closely related to Anomodon, 
at the core of the Anomodontaceae; they both lack paraphyllia and have a rela- 
tively undeveloped peristome. The endostome in the two species of Herpetineu- 
ron here recognized is, nonetheless, somewhat more developed than those of 
most species of Anomodon. The lack of leaf cell papillae in Herpetineuron must be 
regarded as a derived trait. Stark and Buck (1986) created the genus Bryonorrisia 
when they removed B. secunda from Forrstroemia and assigned it to the Anomo- 
dontaceae, and stated that it strongly resembles Herpetineuron. Enroth (1991) 
also acknowledged the close relationship of Herpetineuron and Bryonorrisia, but 
argued that both genera belong in the Leskeaceae. I do not see a sufficient rea- 
son, based either on gametophytic or sporophytic morphology, to segregate Bry- 
onorrisia from Herpetineuron. Whether the Anomodontaceae are closest to the 
Leskeaceae or to the Thuidiaceae, or should be subordinated to either of them, 
has been an ongoing and unresolved argument (see Introduction). I feel more at 
ease with the placement suggested by Anderson et al. (1990), or at least with that 
of Buck and Crum (1990), in which the family Anomodontaceae, Herpetineuron 
included, is considered to be close to the Leucodontaceae. 

The generic name refers to the snakelike costa, characteristic of H. toccoae. 
the type species. 



Key to the Species of Herpetineuron 



, Herpetineuron toccoae (Sull. & Lesq.) Cardot, Beih. Bot. Centralbl. 19(2): 128. 

1905. Anomodon toccoae Sull. & Lesq., Musci Bor. Amer. 52. 1856.— 

Type: U.S.A. Georgia: Toccoa Falls, C. L. Lesquereux, Musci Bor. Amer. 

exsicc. 1, 240 (isotypes: G! MICH! NY!). 
Anomodon devolutus Mitt., J. Linn. Soc. Bot., suppl. 1: 127. 1859.- Syntypes: 

China. Hong Kong, Bowring; India. Mehra, "11.000 ft.," Thomson, 23-7- 

1848; Simla, "7000 ft.," Thomson 10/49 (syntypes, NY-herb. Mitten!). 
Anomodon wichurae Broth., Hedwigia 39: 244. 1899. Herpetineuron wichurae 

(Both.) Cardot, Beih. Bot. Centralbl. 19(2): 128. 1905.— Type: Japan. Kiush- 

iu, Nagasaki, Wichura 1461a (isotype: NY). 
Herpetineuron formosicum Broth., Ann. Bryol. 1: 22. 1928.— Type: Taiwan. 

Prov. Taityn, Hori, Suzuki (n.v.). 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 263 

Herpetineuron attenuatum S. Okamura, J. Coll. Sci. Imp. Univ. Tokyo 38(4): 54. 
1916. — Type: Japan. Hondo, Prov. Sagomi, Kamakura, 1-3-1914, Hisauchi 
(n.v.). 

Plants rather robust, in open, dull, dark greenish to yellowish brown mats. 
Stems prostrate, sparsely branching; branches (1-) 2-4 cm long, (0.2-) 0.9-1.3 mm 
thick when dry, rather arcuate, terete when dry, branching pattern often consist- 
ing of several orders of successive branching in a stepwise fashion, circinate or, 
often, attenuate toward the apex, with a well-differentiated central strand of smaller 
cells, without paraphyllia or pseudoparaphyllia. Branch leaves incurved, secund, 
involute in a tubular fashion when dry, erect, not complanate when moist, ovate- 
lanceolate, tapering from the base, without an obvious constriction, 1.8-2.8 mm 
long, 0.5-1.2 mm wide; margins flat, irregularly and strongly serrate in the distal 1/3- 
1/4, serrations consisting of 1 to several cells; apex sharply acuminate; costa single, 
strong, pellucid, almost always sinuose in the distal 1/3, smooth on the back, not 
obscured by laminal cells, ending sharply at or near the apex, not bifid at the tip; 
laminal cells ± homogeneous throughout the lamina, rectangular to rhombic, slightly 
longer than wide, smooth, walls somewhat thickened. Perichaetia found on youngest 
branches; perichaetial leaves 1.5-2.2 mm, flexuose-subulate, often somewhat cris- 
pate, laminal cells smooth throughout. Seta 14-16 mm, light reddish brown. Cap- 
sule erect, almost symmetrical, yellow to reddish brown; urn (1.7-) 2.0-2.8 mm, 
stomata few, cells at mouth and annulus well differentiated; operculum conic; 
peristome fairly well developed, exostome ca. 0.5 mm high, papillose and faintly 
striolate at the base and above, endostome papillose with a basal membrane to 3-4 
cells high (0.05-0.15 mm), segments 0.2-0.3 mm high, papillose throughout. Spores 
16-22 |Lim in diameter, densely papillose. Chromosome number: n = 11 (n = 9+H+h, 
in mitosis, Japan: Inoue 1965). 

Distribution (Fig. 24). Herpetineuron toccoae is widely distributed in temper- 
ate as well as tropical regions. In the New World H. toccoae is present and some- 
what common in the mountains of North and South Carolina, Georgia, and Ten- 
nessee, in the Ozarkian Highlands, in scattered localities in southern Illinois, western 
Florida, Louisiana, eastern Texas, Mexico, Guatemala, Honduras, Nicaragua, the 
Dominican Republic, and SE Brazil. In the Old World it occurs in the Himalayas 
and southern India, Sri Lanka, Manchuria and southeastern China, Korea, Japan, 
Vietnam, the Philippines, Celebes, Sumatra, Java, and New Caledonia (Norris & 
Sharp 1961). According to Brotherus (1925), H. toccoae is present in South Africa 
and West Africa, and Kis (1985) cites five references from Tanzania. The species 
is absent in Europe and Central Asia. Herpetineuron toccoae grows on trees, 
sometimes at their base, or occasionally on rocks. It forms rather thick mats in 
temperate, deciduous, mostly montane forests. It never fruits in North America. 

Representative Specimens. China. Fukien: Buong Kang, Yenping, H. Chung, 14-6-1925 (DUKE); 
Kushan, pr. Foochow, H. Chung B-240a, 17-8-1925 (c. fr., DUKE MICH). Kwangtung: Jen hwa distr., 
Man Chi Shan, Tsang W.T. 26178, Apr 1936 (c. fr., MICH). Hong Kong: / C Bowring (as Anomodon 
devohitus, syntype, herb. Mitten NY); Kowloon, Panlong wan, Y. W. Taam 2007, 9-6-1941 (DUKE); New 
Territory, Taimoshan, Chunloong, Y. W. Taam, 1948, 16-3-1941 (c. fr., MICH). India. Kumaon, /. 
Duthie 6367, 22-8-1886 (MICH). Madras distr: Pulney Hills, Kodaikanal (coll. ignot. Nr. 88), Jan 1957 
(DUKE). NW Himalayas: Mt. Nag Tiba, Rhada Sal, 7-12-1900 (as A. devolutus, DUKE). Japan. Sen- 
dai, E. lishiba 3046, 20-10-1907 (DUKE). Miyazaki pref., Nao, S of Nichinan, W. Schofleld 53094-3, 9-1- 
1973 (DUKE); Oita pref., Takeda-machi, A. Noguchi, 1-12-1947 (DUKE). Japan. Goto. Faurie 1596, 
1901 (as H. wichurae, NY). Prov. Higo: Konase, Kuma-gun, K. Mayebara 96, 14-10-1928 (as H. wichurae, 
NY). Honshu: Mie pref., Yoro fall, N. Takaki 5291 (as H. wichurae, NY). 





Brazil S 


mt I C atar 


ina Bom Retire BR 282 ScfiaUr Vein imp 1048 


3 & Vcn^imp 21 


-12-1988 


(MK H) Rio 


cIl JmciK 


) Old u 


m f^/5:) g2/6 9222 (as Aiwmodon bras 


lunsn NY) Guatemala 


Hu 


ichucttnaii 


^o Vv ol A 


i,iiacala 


n /^ Sto)J/(v Sy2/^ 27-12-1940 (MICH 


1 Honduras Com IV loui 


Ri( 


) ScUucipci 


"4 km SE 


1 S.^Liat 


sptqUL B Allen 1/ WO 11-^-1992 n40^ 


2-1 1991 (MO) 


01 incho 


ni 




s R Olson 




. dc Lst in/ucl 1 6 












79 (MO) Mexico C hup IS ibo%c I is 


C IS IS A Sluup 




4 


194^ (MICH) Chihua 


hui Ch 


upiL // UShcu> A4h 2()-S-l9^7 (MK 


H) Duiinoo LI 


Silto^/^ 


Sfu 




M 12 1994 


(MICI 




pincinyo 4 Sliai 


/) lll^h 


27 


10-L944 ( 


MICH) JaJ 


isco La 


htrreria ibovt Manall in H Cimn 9S 




(MICH) 


Mc 


xico I emascaltcpcc 


Ixlapaii 


(, Hinion 2917a 14-12-19"i2 (MK H) [ 


Vloiclos (cno k 


po/tcco 












i.LV SW otLKu 


ILldO f 


Ik 


tnuiNH 26( 


S2 (MK H 


) Oixi 


ei lama/uhpin Mixc A Slunp 2990 c 


, al 21-12-19-2 


(MICH) 


Pucbh hLlo 


W N^CIM 


Rio U 


am.o A Sluup 1124 26-->-194S (MIC H) 


Rio Net 1x1 p. 1 


Hu HKhl 


s'n 


X,V;;, 


'r//p!lnm 


is /)^'n 


fMlCII) SmluisPolosi \ilul 1 ^ Shmj 


noil pi hi li>iL 


(MICH) 


del 


KoblL S 


V\////( 4^^ 


y Sep 1 


941 (Midi) Tmuulipas Sitii 1 dt Cm 


nun il 1 S oi R . 


ncho 6l\ 


Ck 


lo H Cm 


m /664 /6' 


H) \lil 


19M)(MICII) USA. Alabama Winstoi 


tCo Bankheadf 


s'at Foi 


V^ 


Biiik2lS6 


y(Mic n ? 


^Y) Ai 


i/on 1 Svcamorc Can>on n, Rubv / Ha, 


mg 1297 & H Ski 


// 11-3- 


19 


h(NV) \ 


ikansis CI* 


buineC 


(, n. CnccrsFcirvReserv P Redfean 


1 J8729 i F Bowl 


n 12-1- 


\9( 


)6 (MIC H 


NY) i,u 


land C( 


. Hot Spiinus Nit Pk / Anderson 1 


1147 ^-10-19^3 


(DUKE 



iW2 4-1-196^ tMK H 
19S2(DIjKL MICH) 



Herpedneuron toccoae is distinguished by the sinuose costa near the leaf apex. 
It is unmistakable in its habit when dry: a large, usually epiphytic moss forming 
thick mats, with the leaves falcate -secund and strongly incurved. The distinctive 
costa inspired Cardot to name the genus Herpedneuron ("snake costa"); however, 
the second species lacks a sinuose costa. 

2. Herpetineuron acutifolium (Mitt.) Granzow, Bryol. 92: 385. 1989. Anomodon 
acudfoUus Mitt., J. Linn. Soc. Bot., suppl. 1: 126. 1859. Bryonorrisia acud- 
foUa (Mitt.) Enroth, J. Bryol. 16: 407. 1991.— Type: India. Uttar Pradesh, 
Kumaon 6000, Sarkito Shayuk valley, 800 (lectotype, designated by Enroth, 
1991: NY-herb. Mitten!). Fig. 31. 

Forrstroemia secunda Dixon & Badhw., Rec. Bot. Surv. India 12: 176. 1938. 
Bryonorrisia secunda (Dixon & Badhw.) Stark & Buck, Brittonia 38: 52. 
1986.— Type: India. "Dalhousie," Badhwar 339 (holotype: BM). 

Plants moderately robust, branches fasciculate or irregularly pinnate, arcuate, 
often attenuate and flagelliform, 0.5-0.7 mm in diameter when dry. Stem and 
primary branch leaves secund and wider, those of secondary branch leaves nar- 
rower and julaceous when dry, triangular-lanceolate, 1.3-1.8 mm long, sometimes 
somewhat secund and asymmetric, without mid-leaf constriction, decurrent, always 
acute, margin flat and entire but becoming serrulate just at the apex; costa decur- 
rent, weak, straight, ending sharply 0.2-0.35 mm, or sometimes more, from the 
leaf apex, cells of costa elongate, longer than laminal cells; lamina cells shortly 
rectangular or rhomboidal, heterogeneous in size and orientation, smooth but 
some prorulate on the abaxial surface. Male gametangial buds borne proximally 



GRANZOW-DE LA CERDA: ANOMODONTACEAE 




acutifolius (Mitt.) Granzow. a, b, c, 
jm with differenciated cells in the center; g, h, bran, 
ortion of peristome with well-developed endostome; j, spores. Based c 
dia, NW Himalaya (NY); b, d, g, j, Koeltz 7529, India, Punjab. Kangra. Bathad, (FH); 
fe Winterbottom 70/13, India, Kumaon, above Jallet, (NY). 



266 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

from the last branching points, but female gametangial buds borne proximally as 
well as distally. Perichaetial leaves acute with the apex strongly serrate. Seta 8-9 
mm. Capsules cylindrical or elliptic; urn 1.5-2.2 mm; exostome well developed, 
horizontally striolate at the base, teeth trabeculate, endostome well developed, 
basal membrane 1-3 cells high, segments long, filiform. Chromosome number 

Distribution (Figs. 22, 28). Unlike H. toccoae, H. acutifolium has a rather 
narrow, although disjunct distribution. It occurs in the Himalayas, southern China, 
Taiwan, and Japan, and is also known from two localities in the state of Chihua- 
hua, Mexico. 

Representative Specimens. China- Yunnan: Diancang Mt. Range, P. Redfearn &Y.G . Su 1520, 
18/20-7-1984. India. Himachal Pradesh: Simla, Bell 113 (NY), "7-8000 ft," Thomson, Oct 1849 
(NY) Punjab: Kangra, Kulu, Bathad, W. Koeltz 7529, 24-11-1933 (FH). Uttar Pradesh: Kumaon. 

7500', Strachey & Winterbottom 70/13 (syntype of H. acutifolium: NY-herb. Mitten!). Kathi 7000'. 
Strachey & Winterbottom 70/43 (syntype of H. acutifolium: NY-herb. Mitlcn!): Rampur-Bushahr. 
Dorkali, W. Koeltz 3185d, 29-11-1931 (NY); "N-W Himalaya," Falconer c^ Ro\le (NY). Japan. 
Honshu: Saitama pref., Chichibu, Mt. Akazawa-dake, Nagano 5021, 9-9-1953, Nagano 10991, 28-9- 
1957, Nagano 9816, 7-8-1957 (NICH). Nepal. East Nepal: "between Dor and Chauke," Z. Iwatsuki 
421, 431, 7-6-1972 (NICH); "between Topke Gola and Shewaden," Z. Iwatsuki 1899, 28-6-1972 (NY, 
HIRO). Katmandu: Mt. Phulchoki, "summit area," Z. Iwatsuki 2670, 15-7-1972 (NICH). 

Mexico. Chihuahua: "51 mi of Creel," La Junta to Creel, F. Bowers et at., 31-12-73 (with H. 
toccoae), 4-1-1974 (MEXU); "30 min. N de San Juanito" C. Delgadillo, 4-1-1974 (MEXU). 

Herpetineuron acutifolium reminds of A. attenuatus and A. girandii in its habit, 
because they too have attenuate and often flagelliform branches. However, H. 
acutifolium differs entirely in its leaf and cell morphologies from any species of 
Anomodon, including the absence of a leaf constriction and the smooth leaf cells. 
The habit and branching pattern, as well as the triangular-lanceolate leaf outline 
and serrulate apex, are very similar to that of H. toccoae. The costa, unlike that of 
H. toccoae, is not sinuose but slender and decurrent at the base, but the areolation 
is almost identical in both species. Cells are rhomboidal, sometimes with scattered 
prominent cells on the laminal surface. The peristome and especially the endo- 
stome are rather well developed in both species, more than in any species of 
Anomodon; the segments are narrow but strong, as long as the exostome teeth, 
perforate, and rising from a very low basal membrane (Granzow-de la Cerda 
1989). Enroth (1991) transferred H. acutifolium to Bryonorrisia, based on its more 
developed peristome. 

ExLiJDED Nambs 

Anomodon califomicus Lesq., Mem. California Acad. 1: 30. 1968. = Triquetrella 
californica (Lesq.) Grout. 

Anomodon brasiliense Hampe, Vidensk. Meddel. Dansk Naturhist. Foren. Kj0ben- 
havn, ser. 3, 4: 56. 1872. = Dimerodontium brasiliense (Hampe) Broth., 
Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26, Afd. 3: 50. 1900. ^ Myrinia 
brasiliense (Hampe) Schimp., Gen. Sp. Muse. 2: 292. 1878. 

Anomodon lagoensis Hampe, Vidensk. Meddel. Dansk Naturhist. Foren. Kj0ben- 
havn, ser. 3, 2: 238. 1870. = Leskea lagoensis (Hampe) Gier, J. Bryol. 10: 



.A CERDA: ANOMODONTACEAE 267 

211. 1978. = Pseudoleskea subcatenulata Besch., Mem. Soc. Sci. Nat. Cher- 
bourg 16: 234. 1872. = Rauia subcatenulata (Besch.) Broth., Nat. Pflanzenfam. 
1(3): 1005. 1907. = Rauiella subcatenulata (Besch.) Wijk. & Marg., Taxon 
11: 222. 1962. = Rauiella lagoensis (Hampe) Buck, Bryol. 94: 82. 1991. 

Anomodon moseri (Kindb.) Kindb., Spec. Eur. & N. Amer. Bryineae, pt. 1: Pleu- 
roc. Linkoping. 1896. = Leskea moseri Kindb., Ottawa Nat. 7: 19. 1893. = 
Leskeella nervosa (Brid.) Loeske, Moosfl. Harz. 255. 1903. 



Anomodon perarmatus WilUams, Bryol. 31: 112. 1928. = Philonotis scabrifolia 
(Hook. f. & Wilson) Braithw., Brit. Moss Fl. 2: 215. 1893 (see Granzow- 
de la Cerda, 1989a: 383). 



Anomodon tasmanicus Broth., Ofver. Finsk. Vet. Soc. Forh. 42: 137. 1900. = Tri- 
quetrella curvifolia Dixon & Sainsb., J. Bot. 71: 217. 1933. = Triquetrella 
I (Broth.) Granzow-de la Cerda, Bryol. 92: 383. 1989. 



Anomodon xanthophyllum Hampe, Bot. Zeit. 26: 816. 1868. = Stereophyllum mex- 
icanum Williams, Bryol. 26: 34, pi. 4. 1923 (fide Ireland, 1982). - Entodon- 
topsis mexicana (Williams) Buck & Ireland, Nova Hedwigia 41: 105. 1985. 



Haplohymenium densum Schimp., Mem. Soc. Sci. Nat. Cherbourg 16: 233. 1872. 
Leskea mexicana Besch., Mem. Soc. Sci. Nat. Cherbourg 16: 233. 1872. 
I (Besch.) Cardot, Rev. Bryol. 37: 51. 1910. 



ungsber., Math.-Naturwiss. 



Haplohymenium tenuissimum (Besch.) Broth. {Pseudoleskea tenuissima Besch.). 
According to the description and distribution, this name is most likely a 
synonym of Haplohymenium pseudotriste (see also Noguchi, 1957). 

nagnirete Sakurai, Bot. Mag. (Tokyo) 64: 196, Ic, d. 1951. 



ACKNOWLEDGMENTS 

This research was partially supported by a postdoctoral fellowship from the Comissio Interde- 
partamental de Recerca i Innovacio Tecnologica (CIRIT) of the Conselleria de Ensenyament, Gen- 
eralitat de Catalunya. The work was mainly carried out at the Laboratori de Briologia, Facultat de 
Ciencies, Universitat Autonoma de Barcelona, and also at the Department of Botany, Duke Univer- 
sity. I am thankful above all for the understanding and unconditional support given by Creu Casas, 
Montserrat Brugues, Rosa Maria Cros, and Francisco Lloret, all of the Univei 



268 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLU^ 

Barcelona (BCB), and to Lewis Anderson for his comments on the manuscript. The very va: 
comments and advice of William Buck, and the help of Brent Mishler and Efrain de Luna reg; 
the phylogenetic analysis were of ent)rmous help. I am greatly indebted to Molly McMullen i 
the staffs at MICH and the Real Jardin Botanico of Madrid for their help with loans, and espt 
to Mariano Sanchez for his help with the line drawings, Gonzalo Nieto for his comments. D 
Gobetti and Michael Bt)nner for checking the Latin diagnoses, and to the curators and ass 
curators of bryophyte collections in B. BCB. C, COI, DUKE, FH, FI, G, H. HIRO. IFF, INA, . 
LISU, MA, MAC, MEXU, MICH, MO, NAU, NICH, NY, PO, RO, TENN, US, WELT. I 



LITERATURE CITED 

i) and related genera in East i 



.vacid sequence data in angiosperm phylogenetic reconstruclic 

ul (iasslflcalion. Portland: Timber Press. 

!• ioonitarum ad norman Hedwigii. SuppI 4. Gotha: C. G. Ettinger 



tsrucn. P., W. f. bchimpcr. and 1. (rinnhel. IS.^l IS.^.i. nrvoloiiiu eitropaea sen genera muscomm 

Buck, W. R., and H. Criim. 1990. .\\\ evaluation of lanulial limits among the genera traditionally 
aligned with the Thuidiaceae and Leskeaceae. Contr. Univ. Vlichi^an Herb. 17: ,"^5-69. 

mosses. Taxon 35: 21-60. 
Corley, M. F. V., A. C. Crundwell, R. Dull, M. O. Hill, and A. J. E. Smith. 1981. Mosses of Europe 
and the Azores; an annotated list of species, with synonyms from the recent literature. J. Bryology 
1 1 : 609-689. 

R. J. Taylor and A. E. Leviton, 115-129. San Francisco: Pacific Divission A.A.A.S., California 

Academy of Science. 
Crosby, M. R., and R. E. Magill. 1978. A dictionary of mosses, 2d printing. St. Louis: Missouri 

Botancial Garden. 

. 1981. A dictionary of mosses, 3d printing. St. Louis: Missouri Botancial Garden. 

Crum, H. A. 1976. Mosses of the Great Lakes forest. Ann Arbor: University of Michigan Press. 
Crum, H. A., and L. E. Anderson. 1981. Mosses of eastern North America. Vol. 2. New York: 



21.3-220. 
Donoghue, M. J., R. G. Olmstead, J. F. Smith, and D. Palmer. 1992. Phylogenetic 
Dipsacales based on rbcL sequences. Annals Missouri Bot. Gard. 79: 333-345. 



GRANZOW-DE LA CERDA: ANOMODONTACEAE 

an Mosses (Bryophytii 

Enroth, J. 1991. A commentary on the moss genus Bryonorrisia Stark & Buck (L( 

acutifolia (Mitt.) Enroth comb. nov. J. Bryology 16: 407-412. 
. 1992. Notes on the Neckeraceae (Musci) 11-12. The taxonomic position 

coslelloides and Bissetia lingulata, with the d 

Gangulee, H. C, 1978. Mosses from Eastern India a 
Gier, L. J. 1980. A preliminary study of the Thu 

Granzow-de la Cerda, I. 1988a. Distribucion de las especies del genero Anomodon Hook. & Tayl. 

(Musci) en la Region Occidental. Fontqueria 16: 9-26. 
. 1988b. Estudios taxonomicos y biosistematicos en el genero Anomodon Hook, and Tayl. 

(Musci). Doctoral Thesis. Universitat Autonoma de Barcelona, Bellaterra (Barcelona). 
. 1989a. Notes on the taxonomic position of five species of Anomodon Hook, and Tayl., some 

with erroneous identity, and two new combinations. Bryologist 92: 381-386. 
. 1989b. Flujo gametico en un musgo pleurocarpico, Anomodon viticulosus (Hedw.) Hook, and 

Tayl., mediante un experimento de trasplantes. Botanica Complutensis 15: 91-100. 
. 1990. Aplicacion del metodo cladistico a un caso real: analisis filogenetico de Anomodon 

Hook. & Tayl. (Musci). Anales del Jardin Botanico 47: 305-325. 
Grout, A. J. 1903. Mosses with hand-lens and microscope. New York: published by the author. 
. 1932-1934. Moss flora of North America north of Mexico. Vol. 3, parts 3-4. Newfane, Ver- 
mont: published by the author.. 
Hooker, W. J., and T. Taylor. 1818. Muscologia Britannica. London: R. & A. Taylor. 
Inoue, S. 1965. Karyological studies in mosses, IV. Similarity of karyotypes in Thuidiaceae. Bot. Mag. 

Tokyo 78: 496-502. 
Ireland, R. R. 1971. Moss pseudoparaphyllia. Bryologist 74: 312-330. 
. 1982. Moss flora of the Maritime Provinces. National Museum of Natural Sciences, Publica- 

Iwatsuki, Z. 1963. A revision of the East Asiatic species of the genus Anomodon. J. Hattori Bot. Lab. 

Iwatsuki, Z., and A. Noguchi. 1973. Index muscorum japonicarum. J. Hattori Bot. Lab. 37: 299-418. 
Kiillcrsio, M., J. S. Farris, A. G. Kluge, and C. Bult. 1992. Skewness and permutation. Cladistics 8: 

275-287. 
Kis, G. 1985. Mosses of south-east tropical Africa. An annotated list with distributional data. Vacratot: 

Institute of Ecology, and Botany of the Hungarian Academy of Sciences. 
Klassen, G. J., R. D. Mooi, and A. Locke. 1991. Consistency indices and random data. Syst. Zool. 40: 

Lazarenko, A. S. 1933. Eine neue Anomodon- Art aus dem Ussuri Gebiet. Revue Bryol. Lichenol 5: 

44-45. 
Limpricht. K. G. 1895. Die Laubmoose Deutschlands Osterreichs und der Schweiz. II. In Kryptoga- 

men-Flora von Deutschland, Osterreich und der Schweiz, ed. L. Rabenhorst, 2d ed. Leipzig: 

Eduard Kummer. 
Mishler, B. D., M. J. Donoghue. and V. A. Albert. 1991. The decay index as a measure of relative 

robustness within a cladogram. (abstract). Hennig X, Toronto. 
Monkemeyer, W. 1927. Die Laubmoose Europas IV. Erganzensband. In Kryptogamen-Flora von 

Deutschland, Osterreich und der Schweiz, ed. L. Rabenhorst. Leipzig: Akademische Verlagsge- 

sellschaft. 
Noguchi, A. 1957. A revision of the genus Haplohymenium Doz. & Molk. (Musci). Kumamoto J. Sci., 

ser. B. sect. 2, 3(1): 20-35. 

. 1972. Musci japonici. IX. The Leskeaceae. J. Hattori Bot. Lab. 36: 449-529. 

. 1974. A checklist of the mosses of Japan. Kumamoto. 

Norris, D. H., and A. J. Sharp. 1961. The known distribution of Herpetineuron toccoae (Sull. & Lesq.) 

Card. J. Hattori Bot. Lab. 24: 110-114. 
Nyholm, E. 1960. Moss flora of Fennoscandia. II. Musci, Fasc. 4. Lund: C. W. K. Gleerup. 
Podpera, J. 1954. Conspectus Muscorum Europaeorum. Prague: Ceskoslovenske Akad. Ved. 
Reimers, H. 1936. Die europaischen Haplocladium-Arten mit besonder Beriicksichtigung ihrer 

aussereuropaischen Verbreitung und ihrer Verwandtschaft. Hedwigia 76: 191-298. 
Sainsbury, G. O. K. 1955. A handbook of the New Zealand mosses. Roy. Soc. New Zealand Bull. 5: 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



vels of homoplasy. Evi 



llivant, VV. S. lSf)4. hom's .Vliisrontni. ("ambridge. Massachussets: Sever and Francis. 

ol'ford, D. L. 1490. PAUI^ Fhylogcncilc analysis using parsimony, version 3. Or. Computer program 

distributed by tlie Illinois Natural History Survey. Champaign. Illinois, 
n, B. C, D. E. BoulTord, Ciheng H., Liu B.,'watig Z.. Xi Y., Ying T.. Zhang C, and Zhang X. 1996. 

A contribution to the moss flora of Hcnan province. China. Acta Bot. Yunnanica 18: 67-71. 
[t, D. H.. 19S4. Classification of the Bryopsida. In !\'ew manual of bryology, ed. R. M. Schuster, 2: 

tt, D. H.. and P. Lee. 1984. Anomodon minor (Musci: Leskeaceae) in North America. Bryologist 

87: 338-339. 
itanabe, R. 1972. A revision of the family Thuidiaceae in Japan and adjacent areas. J. Hattori Bot. 

Lab. 36: 171-320. 
jk, R. van der, W. E. Margadant, and P. A. Florschiitz. 1959 Index Muscorum, Vol. 1. Regnum 

Veg. 17: 1-548. 

. 1969. Index Muscorum, Vol. 5. Regnum Veg. 65: 1-922. 

no, K. 1957. Cytological studies on Japanese mosses. Memoirs Takada Branch, Niigata University 



APPENDIX 1 

HARACTER CODING 



. Branch leaf arrangement when dry (ordered). - Erect to julaceous. 1 = Secund. 2 = Conspic- 

. Orientation of branch leaves when moist (ordered). = Not flattened on either the ventral or 
rsal side of secondary branches. I - Flattened only on the dorsal surface of secondary branches, 
secund and pointing lo the substrate on the ventral surface. 2 = Flattened on both surfaces, 
ally towards the end of secondary branches. 

. Leaf constriction. = No distinguishable constriction; leaves lanceolate to triangular-ovate 
4. giraldii). 1 = Constriction present ("shoulders"), more or less abruptly dividing the leaf into a 
lanceolate proximal region and a narrow distal one. 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 271 

8. Leaf length. = Small: <0.7 mm long. 1 = Large: >1.4 mm long. 

9. Shape of the distal region of the lamina of branch leaves, above the shoulders (Fig. 2). = 
Tapering, i.e., with margins never parallel but progressively narrowing between shoulders and apex. 1 
= Portions of the lamina above the shoulders as narrow as the apex (or even narrower, as in A. 
minor), i.e., with margins parallel at least in some portions. ? = For species without a distinct shoul- 
der. Note: in A. viticidosus the character presents a polymorphism within mats. 

10. Leaf margin. = Plane throughout. 1 = Revolute in part. 

11. Shape of branch leaf apex (partially ordered; Fig. 3). - Leaves tapering into a multiseriate 
subulate apex, in A. longifolius. 1 = Leaves abruptly ending in a sharp and long uniseriate acumen {A. 
rostratus). 2 - Acute {A. giraldii). 3 = Acute to obtuse, in the same plant. 4 = Rounded to obtuse, but 
never acute. Note: The leaf apex is often short-apiculate in A. rugelii, although the outline of those 
leaves is essentially obtuse or rounded. Such an apiculum is not at all uniseriate and not even acute; 

; slightly crenulate due to the 
Transformation Series (Character Tree) Step Matrix 



. Length of the costa (ordered; 1 

. 9/10 of the leaf length to 

14. Costa bifid near the apex (Fig. 5). = Never bifid. 1 = Costa becomes bifid just at the apex 
(although one of the branches often much shorter), at least in some leaves on a mat. ? = Character 
polymorphic among stems within mats {A. attenuatus and P. gracile). 

15. End of the costa covered by laminal cells. = Well defined throughout, ending sharply, not 
obscured by laminal cells. 1 = Costa not ending sharply, somewhat fading as laminal cells cover it, but 
only at the end. 2 = Upper part of the costa totally obscured by overlaying laminal cells, starting as 
far down as the level of the leaf constriction, or just above it. 

16. Papillae on cells of dorsal surface of the costa (unordered; Fig. 6). = Cells smooth or with 
very discrete vesicle-like bumps. 1 = Papillae minute, numerous and randomly arranged on each cell 

Anomodon tristis because of its small size and obscured costa. 

17. Papillae on basalmost margin cells, at the insertion of the leaf (ordered). = Lacking 
papillae, or with papillae numerous, but always similar to other laminal cells. 1 = Intermediate: one 
or two somewhat mamillose cells may appear at the margin. 2 = Cells with very prominent papillae, 
larger than those on laminal cells, usually single and sometimes branched, resembling spines. 

18. Shape of leaf cells, at least those toward the middle of the lamina. = Elongate: rhombic or 
rectangular. 1 = Isodiametric: hexagonal, quadrate or round. 

19. Leaf cell papillae (unordered). = Absent. 1 = One central large papilla, pointed, arcuate, 
(sometimes with 1-3 small branches at the base), entirely covering cell lumen. 2 = One central short 
papilla, rounded, entirely covering cell lumen. 3 = Multiple, unbranched, very short papillae. 4 = 
Multiple, scarcely to strongly branched papillae. 

20. Position of gametangial buds relative to most recent (distal) branching points (ordered; Fig. 
7). = The most recent gametangial buds appear distally, relative to the last branching points (i.e., 
perichaetia or perigonia develop on the youngest branches). 1 = Only flagelliform branches are 
produced beyond gametangial buds (e.g., A. longifolius and Herpeuneuron acutifolium). 2 = Gamet- 
angial buds never found distally relative to latest branching points but only on stem or older branches. 

21 . Cells of perichaetial leaves. = Smooth throughout. 1 = Papillose, at least near the leaf apex. 

22. Length of seta (ordered). = >4 mm. 1 = 1.5-3 mm. 2 = <1 mm (capsule immersed). ? = 
Sporophytes unknown for A. dentatus and A. flagelliformis. 

23. Length of urn. = <1.1 mm. 1 - >1.3 mm. ? = Sporophytes are unknown for A. dentatus and 
A. flagelliformis; length is highly variable in A. giraldii. 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



27. Stnokiic DinanuiUanoii .ii base ui cxostome. U = Absent. 1 = Present. ? = No information is 

lion is ii\ailahlc lor A ilcntiiiiis. A i^uul/lii. and A. flagclliformis, and there is a polymorphism in A. 
longifoliiLs. 

2S). Ornamentation at upper part of exostome teeth (unordered). = Verrucose, with thick 
bumps unevenly arranged. 1 = Papillose, with papillae evenly distributed. 2 - Smooth or finely 
rugose. ? = Sporophytes arc imknown for A. dentutus and A. fUigelliforinis. 

but short (3 or more cells hieh) 2 - Well developed, almost as high as exostome teeth. ? = Sporo- 

31. Calyptra l) (llahnMis I Wuli a lew long and scattered hairs. ? = Sporophytes and 



IS r OF SYNONYMS 



i Cardot, Hedwigia 32: 302. 1893 (nomen nudum) 
Vet. Soc. Forh. 33: 107. 1891 - A preudotristis. 



illimus Herzog, Beih. Bot. Ccrtralbl. 26: 82. 1909. = A. 
sophylhis Miill. Hal. in Paris. Ind. Bryol. suppl. 14. 190 
onii Mitt., J. Linn. Soc. Bol. 13: 309. 1873. = A. preudo' 
iernmiis Mitt., J. Linn. Soc. Bot., suppl. 1: 126. 1859. = 
■jdontoidcs Mull. Hal.. Nuovo Giorn. Hot. Ital.. n. ser., • 
^■ophylhis Broth. & Paris. Rc\. Br\ol. 31: .^6. 1904. - A. 
w/-(P.-Beauv.) Lindb.. Bot. Not. 18(,S: 126. 186.S (nom. 
(;/- subsp. inicgcrriimt.s (Mill.) Z. Iwals.. J. 1 lallori Bot. 



Taxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 

A. minor O&l 2 1 1 1 4 2 1 O&l 2 1&2 1 4 1 1 1 1 

A.thraustus 1110 4 10 2 2 14 10 110 10 1 

Ha.triste 0010 01000200020014 11010 1 00 






274 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 

A. pllcaefolius Dixon (not validly published) = A. minor. 

A. ramulosus Mitt., Trans. Linn. Soc. London, ser. 2, 3: 187. 1891. = A. minor. 

A. rubiginosuhis Mtill. Hal., Flora 73: 496. 1890 (nom. superfl.) = H. toccoae. 

A. mgelii vin.fernigineiis (Besch.) Z. Iwats., J. Hattori Bot. Lab. 26: 51. 1963. = A. rugelii. 

A. sinensi-tristis Mull. Hal.. Nuovo Giorn. Bot. Ital., n. ser., 3: 118. 1896. = A. tristis. 

A. sinensis Mull. Hal., Nuovo Giorn. Bot. Ital., n. sen, 3: 118. 1896. = A. minor. 

A. solovjovii var. henaensis Tan, Boufford &Ying, Acta Bot. Yunnanica 18: 69. 1996. = A. solovjov 

A. stenoglussits Cardot & Ther., Bull. Acad. Int. Geogr. Bot. 18: II. 1908. = A. preudotristis. 

A. subintegerrimus Broth. & Paris, Rev. Bryol. 27: 77. 1900. = A. viticulosus. 

A. submicrophyllus Cardot, Beih. Bot. Centralbl. 19(2): 128. 1905. = A. preudotristis. 

A. tonkinensis Besch., Bull. Soc. Bot. France, ser. 3, 41: 84. 1894. = A. viticulosus. 
A. uematsui Broth., Ofver. Finsk. Vet. Soc. Forh. 62: 33. 1919-1920. = A. rugelii. 
A. iiyematsui Broth., Bull. Soc. Bot. Geneve, ser. 2, 3: 281. 1911. = A. rugehi. 
A. wichume Broth., Hedwigia 38: 244. 1899. = H. toccoae. 

A. wrighiii Mull. Hal., Hedwigia 37: 262. 1898. = A. attenuatus. 
Bryonorhsia acutifolia (Mitt.) Enroth, J. Bryol. 16: 407. 1991. = H. acutifolium. 

B. secunda (Dixon & Badhw.) Stark & Buck, Brittonia 38: 52. 1986. = H. acutifolium. 
Forrstroemia secunda Dixon & Badhw., Rec. Bot. Surv. India 12: 176. 1938. = H. acutifolium. 
Haplohymenium biforme Broth., Bull. Soc. Bot. Geneve, ser. 2., 3: 281. 1911. = A. longinervis. 
H. brachydadum S. Okamura, Bot. Mag. Tokyo 25: 31. 1911. = A. sieboldii. 

brevinerve (Broth.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907. = A. preudotristis. 

crassum Sakurai, Bot. Mag. Tokyo 47: 337. 1933. = A. longinervis. 

cristatum Nog., J. Jap. Bot. 20: 146. 1945. = A. flagelliformis. 

fascicidare Nog., J. Jap. Bot. 13: 791. 1937. = A. preudotristis. 

filiforme (Thw. & Mitt.) Broth., Nat. Pflanzenfam. 1(3): 1236. 1909. = A. preudotristis. 

flagelliforme Savicz-L., Bot. Mater. Inst. Sporov. Rast. Glavn. Bot. Sada RSFSR. 1: 98. 101. 19 

= A. flagelliformis. 
formosaniim Nog., Trans. Nat. Hist. Soc. Taiwan 26: 43. 1936. = A. tristis. 

gonoi Broth., Ofver. Finsk. Vet. Soc. Forh. 62: 33. 1919-1920. = A. sieboldii. 
huttonii (Mitt.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907. = A. preudotristis. 
hmgiglossum P. C. Chen, Fcddes Repert. Spec. Nov. Regni Veg. 58: 31. 1955. = A. tristis. 
hngmcrve (Broth.) Broth., Nat, Pflanzenfam. 1(3): 986. 1907. = A. longinervis. 

mithouardii (Pans & Broth.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907, = A. preudotristis. 
mithouardii var. viride Ther., Ann. Crypt. Exot. 5: 179. 1932. = A. preudotristis. 
nakajii S. Okamura, J. Coll. Sci. Imp. Univ. Tokyo 36(7): 26. 1915. = A. preudotristis. 
obsoletinerve Broth., Ofver. Finsk. Vet. Soc. Forh. 62: 32. 1921. = A. preudotristis. 
okamurae Cardot. Bull. Soc. Bot. Geneve, ser. 2, 3: 281. 1911. = A. sieboldu. 

pellucens var. obtusifolium Broth., Ann. Bryol. 1: 21. 1928. = A. preudotristis. 

pUifemni Broth. & M.Yasuda, Ofver. Finsk. Vet. Soc. Forh. 62: 33. 1919-1920. = A. longinervis. 

pinnatuni Nog., J. Jap. Bot. 13: 410. 1937. = A. sieboldii. 

pseudotriste (Miill. Hal.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907. = A. preudotristis. 

sasaokae Broth., Classif. Moss. Japan: 149. 1932 (nomen nudum) = A. preudotristis. 

scabrum Broth., Rev. Bryol. Lichenol. 2: 10. 1929. = A. tristis. 

sieboldii (Dozy & Molk.) Dozy & Molk., Musci Frond. Arch. Indici 4: 127. 1846. = A. sieboldii. 

spinosum Nog., J. Jap. Bot. 13: 789. 1937. = A. longinervis. 

submicrophyllum (Cardot) Broth., Nat. Pflanzenfam. 1(3): 986. 1907. = A. preudotristis. 



1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 

H. tenenimum Broth., Rev. Bryol. Lichenol. 2: 10. 1929. = A. preudotristis. 

H. triste (Ces.) Kindb., Rev. Bryol. 26: 25. 1899. = A. tristis. 

Haptymenium viticulosum (Hedw.) Furnr., Flora 10(2), Beilage 1: 81. 1827. = A. viticulosus 

Herpetmeuron attenuatum S. Okamura, J. Coll. Sci. Imp. Univ. Tokyo 38(4) 54 1916 = H. i 

^slcum Broth., Ann. Bryol. 1: 22. 1928. = H. toccoae. 
H. wichurae (Broth.) Cardot, Beih. Bot. Centralbl. 19(2): 128. 1905. = H. toccoae 
Hypniim attenuatum (Hedw.) Sm., Fl. Brit. 3: 1279. 1804. = A. attenuatus 
//. flr/e«tiafwrtivar./i</vwm(Brid.)Steud.,Nomencl. Bot. 2 252 1824 =A attenuatus 
H. fragile (Wilson) MUll. Hal, Syn. 2: 471. 1851. = A. tristis 
H. fulvum Brid., Muse. Rec. 2(2): 188. 1801. = A. attenuatus 
H. longifolium (Brid.) Mull. Hal., Syn. 2: 474. 1851. = A. longitohus 
H. pseudotriste Mull. Hal., Bot. Zeit. 13: 786. 1855. = A. pseudotiistis 
H. rugelii Mull. Hal., Syn. 2: 472. 1851. A. = rugelii. 
H. triste (Ces.) Mlill. Hal., Syn. 2: 478. 1851. = A. tristis. 

H. viticulosum (Hedw.) With., Syst. Arr. Brit. PI. ed. 4, 3: 830. 1801. = A. viticulosus. 
H. viticulosum var. brachycarpum Mull. Hal., Syn. 2: 473. 1851. = A. minor. 

lymenium longifolium Hampe, Syn. 1: 475. 1848 (nomen nudum) = A. longifolius. 

oldii Dozy & Molk., Ann. Sci. Nat. Bot. ser. 3, 2: 310. 1844. = A. sieboldii. 

f apiculata Schimp. in Mull. Hal., Syn. 2: 473. 1851 (nomen nudum) = A. rugelii. 

•^uata Hedw., Spec. Muse. 230. 1801. - A. attenuatus. 

is Ces. in De Not., Syll. Muse. 67. 1838. = A. tristis. 

■aflaccida Brid., Sp. Muse. 2: 137. 1812. - A. minor. 

or P.-Beauv., Prodr. 78. 1805. = A. minor. 

oldii (Dozy & Molk.) Miill. Hal., Syn. 2: 81. 1850. = A. sieboldii. 

■ulosa Hedw., Spec. Muse. 209. 1801. = A. viticulosus. 

■ulosa var. minor Hedw., Spec. Muse. 210. 1801. = A. minor. 

'nandrum longifolium Schleich. ex Brid., Mant. Muse. 128. 1819. = A. longifolius. 



SPECIES DELIMITATION, AND NEW TAXA AND 
COMBINATIONS IN LEUCAENA (LEGUMINOSAE) 

Colin Hughes 

Department of Plant Sciences 

University of Oxford 

South Parks Road 

Oxford, OXl 3RB, England 



INTRODUCTION 

Different approaches to species delimitation have resulted in widely differing 
numbers of species recognized within Leucaena Benth. by different authors. This 
has been the main cause of taxonomic confusion within the genus. In this paper, 
species delimitation in Leucaena is discussed, one new species and two new sub- 
species are described, and four new combinations proposed. These are published 
here in advance of a forthcoming monograph (Hughes, in press) in order to expe- 
dite use of these new names in a series of imminent publications and databases 
concerned with economic use of the genetic resources of Leucaena, including the 
Oxford Leucaena seed database (Hughes, unpubl.), a World Germplasm Cata- 
logue (Bray et al, unpubl.) and a Genetic Resources Handbook (Hughes, unpubl). 
Full descriptions, specimen citation lists, drawings, and distribution maps are pre- 
sented in the monograph, except for L. lempirana, which is fully described and 
illustrated here. 



SPECIES DELIMITATION 

Although the variation in numbers of species recognized and in the impor- 
tance given to infraspecific ranks by different authors is partly attributable to the 
history of collection and species discovery, it is also a function of differing views 
about what constitutes good character evidence for species or infraspecific taxa. 
Britton and Rose (1928) recognized 39 species of Leucaena and based delimita- 
tions on characters that are now viewed as unrehable, either because they present 
continuous patterns of variation across species (e.g., leaf, leaflet, and pod dimen- 
sions) and are therefore not amenable to anything but arbitrary division, or be- 
cause they vary within populations that are otherwise constant (e.g., leaf and pod 
pubescence). In the absence of rangewide samphng, they failed to detect the 
continuities and population variation that are now obvious and the result was a 
proliferation of supposed new species. Brewbaker and colleagues reduced the 
number of species initially to ten (Brewbaker et al. 1972; Brewbaker & Ito 1980) 
with gradual re-acceptance of additional species to 16 (Brewbaker 1987; Brew- 
baker & Sorensson 1994). The criteria they used to delimit species were never 
explicitly stated, and no formal taxonomic account was produced. It is clear, how- 
ever, that Brewbaker maintained a skeptical view of the distinction of any species 
until he had collected material of it himself and observed its progeny in cultiva- 
tion in the Waimanalo arboretum in Hawaii. Additional species were acknowl- 
edged only with some reluctance during the 20 years after his initial sweeping 



277 



278 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

reduction to ten. Zarate (1984, 1994), in his revision of the Mexican species, 
described two new species, four new subspecies, and proposed five new combina- 
tions, four of them based on species recognized by Britton and Rose. In so doing 
he adopted a third approach to species dehmitation that rehed on very extensive 
use of subspecies. Zarate (1994) justified this on the frequent occurrence of inter- 
specific hybridization in Leucaena, and the unusual "abundance of incipient allo- 
patric speciation" (Zarate 1994: 88), which he attributed to the complex biogeo- 
graphical history of the region. Subspecies were viewed as a solution to these 
perceived difficukies. Zarate also apparently saw subspecies as a mechanism to 
indicate relationships reflected in his belief that "a classification exclusively of 
distinct species is of no benefit either to the interested scientist, or to communica- 
tion of this knowledge to the user community" (Zarate 1994: 88, translated the 
from Spanish). Zarate further mentioned ease of identification (to binomial) with 
certainty as more than compensating for the inconvenience caused by a system 
replete with trinomials. 

The last 15 years has seen a dramatic renewal of interest in the nature of 
species, re-examination of traditional species concepts, development of new ones, 
and new methods to discover species, driven by the rise and adoption of cladistic 
methods and molecular evidence in systematics (reviewed by, e.g., Mishler & 
Budd 1990; Baum 1992; Panchen 1992; Rieseberg & Brouillet 1994; Davis 1995; 
Luckow 1995). Given this renewed debate, Luckow (1995) suggested that explicit 
justification for the species concept adopted and the criteria apphed to delimit 
species should be a requirement for any taxonomic revision. My aim in delimiting 
species within Leucaena has been to name, as species, all the diagnosable entities 
based on available character evidence. Such an objective must emphasize distinc- 
tions rather than similarities, as pursued by Zarate (1994). To do this I have used 
the explicit pattern based species concept of Rosen (1979), Eldredge and Cracraft 
(1980), Nelson and Platnick (1981), and Cracraft (1983), recently re-defined by 
Nixon and Wheeler (1990: 218) as "the smallest aggregation of populations (sexual) 
or lineages (asexual) diagnosable by a unique combination of character states in 
comparable individuals," and now widely known as the phylogenetic species con- 
cept. This concept uses populations as the units of analysis and looks for aggrega- 
tions of populations that possess constant and unique character states or unique 
combinations of character states. A logical consequence of the phylogenetic spe- 
cies concept is that species are delimited by the same criteria used to define 
characters and character states in primary homology assessment for cladistic anal- 
ysis. The advantage of the phylogenetic species concept over other morphological 
concepts such as the intuitive 'taxonomic' (species as the smallest groups that are 
consistently and persistently distinct as distinguished by ordinary means, Cron- 
quist 1988), phenetic, or 'traditionar concepts, is that it provides explicit justifica- 
tion for decisions taken. One obvious limitation is that characters that do not 
show discrete states when the total variation within a genus is viewed simulta- 
neously may be amenable to division when only two, or a small subset of species, 
are considered. 

Definitions of 29 morphological characters and their character states which 
partition variation within Leucaena are presented in Hughes (in press). These 
character states provide the basis for delimitation of the 22 species recognized in 
the forthcoming monograph of Leucaena (Hughes, in press). This result means 
that many of the species recognized by Britton and Rose are treated as conspecific. In 
contrast, some of the subspecies recognized by Zarate (1994), and one described 



1997 HUGHES: LEUCAENA 279 

by Hughes (1991), qualify as distinct species. While most of these are attributable 
to older names provided by Britton and Rose, two require new combinations 
proposed here. Although there is indeed evidence for interspecific hybridization within 
Leucaena (e.g., Hughes & Harris, 1994), infraspecific taxa provide no solution to 
that problem. Zarate's second contention that subspecies are an effective way to 
indicate groups of closely related taxa, is again little substitute for an informed 
analysis of species relationships and presentation of an exphcit hypothesis of rela- 
tionships in the form of a branching diagram (Hughes, in press). Zarate's justifica- 
tion, based as it was on process assumptions and scenarios, is a retreat from the 
morphological evidence. Furthermore, experience in other economically impor- 
tant forestry genera (e.g., Pinus L.) indicates a persistent reluctance to use trino- 
mials in practice. 

Application of the phylogenetic species concept has raised the question of the 
nature and reality, if any, of infraspecific taxa. Some have argued that the phylo- 
genetic species concept denies the existence of infraspecific taxa (Rosen 1979; 
Donoghue 1985; Cracraft 1992). After all, if species are the smallest aggregations 
of populations diagnosable by a unique combination of character states, what can 
be the nature of infraspecific taxa? Again, this depends on the criteria used to 
define characters. Use of strict criteria that demand discrete, fixed differences 
among populations leaves a large body of evidence, largely from quantitative 
characters that show continuous variation, but are not amenable to objective 
partitioning based on absolute gaps. Use of this evidence, albeit using arbitrary 
divisions, provides a viable extension of the phylogenetic species concept to in- 
fraspecific level (Nixon & Wheeler 1990; Luckow 1995). I have used variation in a 
number of quantitative leaf and pod traits that were rejected as characters, be- 
cause they vary continuously across species and show overlapping variation when 
viewed across the genus as a whole, to recognize six infraspecific taxa (Hughes, in 
press). Subspecies are used for entities which are distinguished by several quanti- 
tative traits and which are clearly correlated with geography. Varieties are used 
for entities which differ in several quantitative traits but which are not correlated 
with geography or for which the geographic limits of the variants are poorly 
known. On this basis two new subspecies are described here and two subspecies 
originally described by Zarate (1994) are demoted to varietal rank. 



NEW TAXA AND COMBINATIONS 

Leucaena lempirana C. E. Hughes, sp. nov. — Type: Honduras. Yoro: 6 km SE of 
El Negrito, on side rd to Nueva Esperanza, nr Rio Cuyamapa, on edge of 
Oloman Valley, 15°17'N, 87°40'W, 25 Feb 1991, Hughes 1412 (holotype: 
EAP!; isotype: FHO! additional isotypes to be distributed to MEXU, K, 
NY). Fig. 1. 

Species nova, L. salvadorensi proxima, sed foliis pinnis numerosioribus, pinnis 
folioHs numerosioribus minoribusque, ramuhs capitula ferentibus terminahbus erec- 
tis, determinatis, aphyllis, in apicem vegetativum abortivum desinentibus differt. 

Small slender tree 4-15 (-20) m tall, 10-40 cm bole diameter, typically with 
upright branching and a rounded crown above a short clear bole to 4 m. Bark 
light grey-brown with powdery orange-brown lenticels and shallow rusty orange- 
brown vertical fissures, inner bark green. Leafy shoots terete, pale orange-brown 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME / 




1997 HUGHES: LEUCAENA 281 

striped pale grey-brown covered in fine whitish velutinous pubescence. Stipules 
subulate with small asymmetric wings at base, hairy, 3.5-3.8 mm long, persistent. 
Leaves 19-24 (-26) cm long, 11-14 (-19) cm wide, petioles (including pulvinus) 
25-35 mm long, one or occasionally two pale yellow to orange-yellow, sessile, 
elliptic dome-shaped or truncate-conic nectaries, 3 x 1.5 mm, at distal end on 
adaxial side of petiole; rachis 11-15 cm long with two narrow adaxial ridges and a 
small elliptic conic nectary, 1 x 0.7 mm, at distal end, extending beyond the termi- 
nal pair of pinnae as a short pointed hairy macro, 2.5-3.5 mm long, curHng when 
dry; pinnae 14-19 pairs; pinnular rachis 7-10 cm long, densely covered in short 
white hairs, with a minute round nectary at distal end; leaflets (27-) 30-36 (-40) 
pairs per pinna, nearly sessile, 5-6 mm long, 1.6-2 mm wide, asymmetric, oblong, 
obtuse to rounded apically, rounded to truncate basally, densely white-pubescent, 
midrib strongly asymmetric, a few secondary veins visible. Capitula 16-18 mm in 
diameter at anthesis, each with 100-130 pale cream-white flowers, in fascicles of 
3-5 at nodes or in leaf axils on long erect terminal anauxotelic shoots on which 
leaf development is suppressed, the capitula exserted beyond the periphery of the 
tree crown; peduncles 12-25 mm long, angled, pale orange-brown with velutinous 
white pubescence and an involucre of bracts. Flowers subtended by peltate bracts, 
2.2-2.6 mm long, 0.6-0.9 mm in diameter, densely pubescent; calyx 2.4-2.7 mm 
long, glabrous, pale cream, the lobe tips tinged green; petals free, 3.5-4.5 mm 
long, glabrous, occasionally sparsely cihate on lobe tips, pale green; filaments 10- 
12 mm long, white; anthers with a tuft of terminal hairs, apiculum absent, cream- 
white; ovary 2.3-3.4 mm long, sessile, pale brown, densely covered in short white 
hairs, style 10-11 mm long, white, with a terminal tubular stigma. Pods (10-) 12- 
20 (-25) cm long, (18-) 20-26 (-32) mm wide, 1-2 (-3) per capitulum, pendulous 
on sturdy 8-11 mm long stipes, oblong to linear-oblong, constricted where seed 
abortion has occurred, apex mucronate with a beak 5-15 mm long, base cuneate, 
narrowly piano-compressed, 14-20 seeded, valves membranous, mid-orange-brown, 
pubescence variable from nearly glabrous to dense and velutinous, venation retic- 
ulate, most pronounced at margins, the margins slightly thickened, dehiscent along 
both sutures. Seeds 6.6-8.8 mm long, 3.8-5.2 mm wide, compressed, flattened, 
elliptic to ovate, deep chestnut-brown, glossy, aligned transversely in pods, pleu- 
rogram visible, deep U-shaped, symmetrical. 

Additional Specimens Examined. Honduras. Yoro: W end of the lower Aguan Valley, rd W 
from Olanchito towards Sabana Larga, 1 km S of San Lorenzo, 15°25'N, 86°57'W, 25 Mar 1991, 
Hughes 1447 (EAP, FHO, K, MEXU, NY); 3 km WSW of Arenal, rd to Jocon, lower slopes of hills S 
side of the Aguan Valley, 15°22'N, 86°51'W, 16 Feb 1991, Hellin & Hughes 4 (EAP, FHO, K, MEXU, 
NY); 9 km SSE of Arenal, rd towards Jocon and Yoro, 15°22'N, 86°5rW, Feb 1989, Alvarado G/2/89 
(HEH); 15 km NW of Jocon, rd to the Aguan Valley in tributary valley of the Rio Macora, between 
Macora and Sabana Larga, about 8 km NW of Macora, 15°2rN, 86°59'W, 17 Feb 1991, Hellin & 
Hughes 14 (EAP, FHO, K, MEXU); rd E from Santa Rita to Yoro, 4 km E of Negrito immediately E 
of Valle Oloman above the Rio Cuyamapa, 15°18'N, 87°40'W, 25 Feb 1991, Hughes 1411 (EAP, 
FHO, K, MEXU, NY); rd E from Santa Rita to Yoro, 3 km SE of El Negrito or 2 km NW of the Rio 
Cuyamapa in the Valle Oloman, 15°18'N, 87°41'W, 25 Feb 1991, Hughes 1414 (EAP, FHO, K, 
MEXU, NY); 0.5 km SW of San Lorenzo, N side of the Aguan Valley, 47 km W of Olanchito, 
15°25'N, 86°58'W, 17 Feb 1991, Hellin & Hughes 11 (EAP, FHO, K, MEXU, NY); 3 km S of 
Zapamatepe, 8 km W of Arenal, S side of the Aguan Valley, 15°22'N, 86°54'W, 16 Feb 1991, Hellin & 
Hughes 7 (EAP, FHO, K, MEXU, NY); N side of the Aguan Valley between Medina and Coyoles, rd 
12 km W of Olanchito, 15°28'N, 86°28'W, 17 Feb 1991, Hellin & Hughes 8 (EAP, FHO, K, MEXU, 
NY); rd E from Santa Rita to Yoro, nr Cuyamapa, 15°16'N, 87°27'W, 15 Feb 1991, Hellin & Hughes 1 
(EAP, FHO, K, MEXU, NY); Santa Rita, 5 km ESE of El Negrito, side rd off the main El Negrito to 
Yoro rd to Nueva Esperanza, banks of the Rio Cuyamapa, 15°17'N, 87°40'W, 4 Aug 1991, Hughes 



282 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

1479 (EAP, FHO, K, MEXU, NY); rd E from Santa Rita to Yoro, 4 km E of El Negrito, nr the 
turnoff over the Rio Cuyamapa towards Morazan, 15°17'N, 87°39'W, 7 Mar 1992, Hughes 1712 (E, 
FHO, K, MEXU, NY). 

The first herbarium specimen of this new species {Alvarado G/2/89) was col- 
lected in 1990 by Caspar Alvarado and Jon Hellin, foresters from the National 
Forestry School (Escuela Nacional de Ciencias Forestales), Siguatepeque, Hon- 
duras, and I thank them for drawing my attention to it. Leucaena lempirana is the 
only species of Leucaena endemic to Honduras and is named here in honor of the 
Indian chief Lempira, a Cacique from the Celaque region of Honduras, who was 
killed while attending a peace conference, the victim of a breach of truce imposed 
by the Spaniards (Wendell 1976) and after whom the Honduran currency is named. 
Leucaena lempirana is restricted to two areas, the Aguan Valley system and the 
valleys of Oloman and Cataguana, both in the Department of Yoro in northern 
Honduras. 

Leucaena lempirana belongs within the L. shannonii J. D. Smith alliance with 
L. shannonii, L. salvadorensis Standi, ex Britton & Rose, and L. magnifica (C. E. 
Hughes) C. E. Hughes but shows closest affinities to L. salvadorensis. It is distin- 
guished from L. salvadorensis by its long terminal anauxotelic flowering shoots on 
which the capitula and pods are borne on the periphery of the tree crown. These 
flowering shoots are strongly reminiscent of those of L. shannonii. It is distin- 
guished from both L. salvadorensis and L. shannonii by its smaller and more 
numerous leaflets and pairs of pinnae per leaf. The pods are also similar to those 
of L. shannonii and are variably glabrous or pubescent as for that species. Leucaena 
lempirana, L. salvadorensis, and L. shannonii all occur in Honduras but occupy 
distinct and virtually allopatric distributions in different, isolated valley systems at 
low to middle elevations. 



macrophylla Benth. Bot. Voy. Sulphur. 90. 1844.— Type: Mexico. Gue- 
rrero: Acapulco, 1841, Hinds s.n. (holotype: K!). 

Britton and Rose (1928) distinguished L. nelsonii Britton & Rose from L. 
macrophylla solely by its copiously pubescent leaflets and had not seen fruiting 
material at the time of their original description. Zarate (1994) placed L. nelsonii 
as a subspecies of L. macrophylla based on quantitative leaf and pod differences 
and leaflet pubescence, and assigned material from SE coastal Guerrero (type 
locality of L. nelsonii), coastal Oaxaca, and Veracruz to that subspecies. The type 
specimen of L. nelsonii does indeed have copious pilose or villous pubescence on 
both leaflet surfaces, a feature also noted by McVaugh (1987) to be present on 
trees from throughout the Balsas region from eastern Jalisco to Guerrero. As 
noted by McVaugh (1987), even on typical representatives of L. macrophylla 
from the northern part of its range, leaflets are rarely completely glabrous, but 
often have stiff white hairs on the pulvinules and along the primary and secondary 
veins. Rangewide survey of leaflet pubescence reveals that while material from 
the Balsas region is generally more pubescent, it is variable, and pubescent indi- 
viduals also occur elsewhere (e.g., coastal Michoacan). At the type locality of L. 
nelsonii leaflet pubescence also varies, as shown by some very sparsely pubescent 
individuals (e.g., Hughes 644). Material from coastal Oaxaca, assigned to subsp. 
nelsonii by Zarate (1994), is generally glabrous. Given this pattern of variation in 
leaflet pubescence and the close similarity of the material from coastal Guerrero 



1997 HUGHES: LEUCAENA 283 

to typical examples of L. macrophylla, there appears to be no basis for maintain- 
ing L. nelsonii as a separate species or subspecies within L. macrophylla. The type 
specimen of L. macrophylla was collected from 'Acapulco'; given that there is no 
record of typical material of L. macrophylla from the immediate vicinity of Aca- 
pulco, it is probable that the type was collected either along the coast (near the 
type locahty of L. nelsonii) or inland close to the route to Mexico City, further 
encouraging treatment of these two species as conspecific. Similar variation in 
leaflet pubescence has been noted within the other large-leaflet species L. lan- 
ceolata S. Watson and L. trichodes (Jacq.) Benth. 

Material of L. macrophylla from coastal Oaxaca and Veracruz, placed in 
subsp. nelsonii by Zarate (1994), does however differ from typical representatives 
of L. macrophylla in leaf and pod dimensions, habit, and chloroplast DNA restric- 
tion site variation, and merits recognition as a distinct subspecies. Because the 
type of L. nelsonii belongs with L. macrophylla and not with the distinct Oaxacan 
material, there seems no alternative but to add a new subspecies, here named 
istmensis, to account for this variation, despite the additional confusion that this 
may cause. Subspecies istmensis differs from subsp. macrophylla in its smaller 
leaflets and pods, shorter peduncles, and in its cpDNA, by 16 autapomorphic 
fragment changes, six of which are unique (Harris et al. 1994). 

Leucaena macrophylla Benth. subsp. istmensis C. E. Hughes, subsp. nov. — Type: 
Mexico. Oaxaca: 40 km W of Puerto Escondido, coast rd to Pinotepa 
Nacional, nr San Isidro Llano Grande, 2 km inland from Pacific coast, 
15°58'N, 97nO'W, 26 Mar 1989, Hughes 1338 (holotype: FHO!; isotypes 
distributed to AAU, K, MEXU, NY). 

A L. macrophylla Benth. subsp. macrophylla pinnis cujusque folii paucioribus, 
foliolis angustioribus, peduncuhs brevioribus, leguminibus angustioribus differt. 

Small, generally single-stemmed tree 3-10 (-15) m tall, bole 10-15 (-40) cm in 
diameter. Leaves: petioles (including pulvinus) 11-22 mm long, rachis (7-) 8.2- 
10.9 cm long; pinnae (2-) 3 pairs; leaflets (15-) 23-56 (-64) mm long, (6-) 17-24 
(-28) mm wide, 3 or 4 (-6) pairs per pinna. Capitula 7-9 (-10) mm in diameter at 
anthesis; peduncles (3-) 4-10 mm long. Flowers subtended by small peltate bracts, 
1.7-2.1 mm long; calyx 1.9-2.3 mm long; petals (2.5-) 2.6-3 mm long; filaments 
(3.7-) 4-5 (-5.2) mm long; style (3-) 5.5-6 mm long. Pods (9-) 12-14 (-15.3) cm 
long, (9-) 14-18 (-23) mm wide, 2-4 (-8) per capitulum, glabrous and sHghtly 
lustrous or with dense velutinous pubescence. Seeds 4.9-6.6 mm long, 3.5-4.6 mm 

Representative Specimens. Mexico. Oaxaca: 25 km S of Sola de Vega, rd to Puerto Escondido, 
Sierra Madre del Sur, 16°02'N, 97°09'W, 12 Nov 1983, Hughes 386 (FHO, K, MEXU); 2 km E of San 
Isidro Llano Grande, coast rd between Pinotepa Nacional and Puerto Escondido, 15°59'N, 97°16'W, 

9 Mar 1985, Hughes 580 (FHO, K, MEXU); 10 km N of Candelaria, 40 km N of Pochutla, rd to 
Oaxaca, 15°51'N, 96°28'W, 11 Feb 1987, Hughes 846 (FHO, K, MEXU); immediately S of El Arenal, 

10 km E of Playa Coyula, nr the Pacific coast, 15°44'N, 96°01'W, 14 Feb 1987, Hughes 854 (FHO, K, 
MEXU); 6 km W of Pinotepa Nacional, rd to Acapulco, 16°2rN, 98°03'W, 21 Feb 1987, Hughes 873 
(FHO, K, MEXU); rd N from Juchitan to Acayucan 5 km N of Mafias Romero, low windswept hills, 
Tehuantepec isthmus, 16°56'N, 95°0rW, 12 Mar 1989, Hughes 1304 (CR, FHO, K, MEXU, NY); 2 
km inland from Pacific coast, 10 km E of Coyula, 20 km E of Puerto Angel, dry valley between 
Coyula and Bahia Santa Cruz, 15°47'N, 96°07'W, 27 Mar 1989, Hughes 1340 (BR, FHO, K, MEXU, 
NY); 8 km SE of Cacahuatepec, 3 km NW of Ixcapa, 16°35'N, 98°10'W, 17 Apr 1976, Sousa 5507 
(MEXU); 14 km SE of Pinotepa Nacional, 4 km NW of Nuaxpaltepec, Jamiltepec, 16°21'N, 97°56'VV', 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

1982, Zdrale 641 (MEXU); Revolucion, 16 km NW of Palomares, rd to Tuxtepec, 17°10'N, 
W, 23 Feb 1978, Sousa 9240 (MEXU, MO, UC); "El Tigre," 2 km N of San Agustin Chayuco, 
epec, 16°25'N, 97°49'W, 6 Apr 1982, Tenorio 234 (CAS, MEXU, MO); 1.2 km N of jcl to 
Vivero Guapinol towards San Agustfn Chayuco, 24 km SE of Pinolepa Nacional, 16°19'N, 97°5r'W, 
22 Oct 1982, Torres 1637 (CAS, MEXU, MO); Rancheria La Esmeralda, 6 km S of San Gabriel 
Mixtepec, 16°04'N, 97°04'W, 12 Feb 1976, Sousa 5306 (MEXU); Acatlan, 1 km W of Tetela, Dtto. 
Tuxtepec, I8°29'N, 96°27'W, 22 Dec 1978, Sousa 10306 (CAS, MEXU, MO); 1 km E of Atoyaquillo, 
Putla, 16°49'N, 9TA1"^, 22 Dec 1987, Solano 70 (MEXU): 5 km W of Jamiltepec, 16°18'N, 97°52'W, 
7 Dec 1978, Sousa 9928 (CAS, MEXU, MO); 1! km N of Mati'as Romero, 4 km S of Piedra Blanca, 
5°55'N, 95°03'W, 23 Feb 1978, Sousa 9225 (MEXU. MO. UC): 6 km NW of Pinotepa Nacional, 
itto. Jamiltepec, 16°23'N, 97°51'W, 22 Oct 1977, Sousa 8453 (CAS, MEXU, UC); Rio Pacine, 3 km 
fE of San Juan Guichicovi, Juchitan, 16^58'N, 95°04'W, 25 Mar 1988, Torres 12018 (MEXU); 3 km N 
f Santa Maria Chimalapa, track to Paso Mactaspac del Rio del Corle, 16°55'N, 94°41'W, 14 Nov 
584, Hernandez 589 (CAS, MEXU, MO); 3 km E of Ojo de Agua S of Sola de Vega, 16°24'N. 
97°05'W, 9 Dec 1978, Sousa 9986 (CAS, MEXU, MO).— Veracruz: N side of Cerro de los Metates. 
i Jun 1972, Doranles 1011 (F, MEXU); Salto de Agua, 11 km S of Santiago Tuxtla, 18°23'N. 
i°20'W, 21 Mar 1967, Sousa 3007 (MEXU); nr Plan del Rio and Emiliano Zapata, 19'20'N, 96°38'W. 
) Sep 1982, Villanueva 246 (NY); Salto de Eyipantla, 8 km from Sihuapan, nr San Andres Tuxtla. 
18°24'N, 95°12'W, 26 Jan 1978, Calzada 4245 (F). 

The name istmensis refers to the occurrence of this subspecies from the Pacific 
foothills of coastal Oaxaca across the Isthmus (istmo) of Tehuantepec to a re- 
stricted area around San Andres Tuxtla in southern Veracruz. It is essentially a 
A/land subspecies of the dry deciduous tropical forest. Pods of subsp. istmensis 
are usually glabrous and slightly lustrous, but pods of trees from one population 
(Hughes 854, 855, 1340) from coastal Oaxaca, 20 km east of Pochutla near Coyula. 
iense velutinous pubescence. 



Leucaena leucocephala (Lam.) de Wit, Taxon. 10: 53. 1961. Mimosa leucocephala 
Lam., Encycl. 1: 12. 1783. Acacia leucocephala (Lam.) Link, Enum. hort. 
berol. 2: 444. 1822. — Type: specimen in Lamarck herbarium labeled "Mi- 
mosa latisiliqua," "Mimosa leucocephala" (holotype: P-LA, microfiche K!). 

Variation within L. leucocephala was first noted by agronomists who were 
evaluating the performance of different accessions for fodder production in field 
trials (e.g., Hutton & Gray 1959; Brewbaker et al. 1972). Two main variants, based 
primarily on habit, degree of branching, and vigor were recognized: a shrubby, low 
growing, highly branched, seedy, and often weedy, variant designated the "Com- 
mon" or 'Hawaiian' type; and an erect, arborescent, httle-branched, less seedy 
variant designated the 'Gianf or 'Salvador' type (Hutton & Gray 1959; Gray 
1967; Brewbaker et al. 1972; Brewbaker 1980; Brewbaker, 1987). 

Although it was realized from the start that the 'Gianf or 'Salvador' type 
corresponded to the type material of L. glabrata Rose {Palmer 386 from the 
vicinity of Acapulco, Guerrero, Mexico) (e.g., Gonzalez et al. 1967), Brewbaker 
(1980) initially treated L. glabrata as a synonym of L. leucocephala and only later 
alluded to the distinction of two formal varieties, var. leucocephala corresponding 
to the shrubby 'Common' or 'Hawaiian' type and var. glabrata corresponding to 
the 'Gianf or 'Salvador' type (Brewbaker 1987). The formal recognition of two 
infraspecific taxa within L. leucocephala was discussed by Zarate (1987), who 
raised their rank to subspecies and published the combination: L. leucocephala 
(Lam.) de Wit subsp. glabrata (Rose) S. Zarate. The two subspecies recognized by 
Zarate (1987) correspond directly to the agronomic 'types,' viz: subsp. leucocephala = 
'Common' or 'Hawaiian' type; subsp. glabrata = 'Giant' or 'Salvador' type. 



1997 HUGHES: LEUCAENA 285 

Zarate (1987) listed the characters, in addition to habit, degree of branching, 
and vigor, that distinguish subspecies leucocephala and glabrata. These included 
leaf size, leaflet size, pod and seed size, habit, and, most notably, vestiture of the 
shoot, leaf rachis, leaflet, and pod. Subspecies glabrata has larger leaves, leaflets, 
and pods, and is almost entirely glabrous, compared to the smaller leaves, leaflets, 
pods, and densely puberulent, canescent shoot, leaf rachis, and pods of subsp. 
leucocephala. 

During recent exploration by Hughes and collaborators in northern Guate- 
mala, an additional variant, which differs from both subspecies leucocephala and 
glabrata, was encountered in a localized area around the town of Ixtahuacan in 
the highlands of Huehuetenango. This variant has the small leaves, leaflets, and 
pods of subsp. leucocephala, but is glabrous like subsp. glabrata. This material is 
here described as a third subspecies. 

Leucaena leucocephala subsp. ixtahuacana C. E. Hughes, subsp. nov.— Type: Gua- 
temala. Huehuetenango: 1 km ENE of San Miguel, track running WSW 
from Ixtahuacan into the valley of the Rio Cuilco, 15°23'N, 91°50'W, 
Hughes et al. 1689 (holotype: FHO!; isotypes: distributed to: E, K, MEXU, 
MO, NY). 

A L. leucocephala subsp. leucocephala ramulis, rachibus, foUolis fructibusque 
glabris differt. 

Small tree, (3-) 5-7 m tall, bole 5-20 cm in diameter, arborescent with a short 
clear bole to 2 m high and a rounded compact crown. Leafy shoots glabrous. 
Leaves (10-) 13-16 cm long, 8-11 cm wide, petioles 13-23 mm long, glabrous, 
petiolar nectary 2-2.4 x 1.2-1.5 mm; rachis (5-) 7-12 long, glabrous, mucro 1.9-3.6 
mm long, very sparsely puberulent; pinnae (4-) 6-8 pairs; pinnular rachis 5-8 cm 
long, glabrous; leaflets 16-20 pairs per pinna, 9-11 mm long, 2-2.6 mm wide, 
glabrous or very sparsely ciUate along leaflet margins near base. Capitula 12-17 
mm in diameter at anthesis each with 110-120 flowers, in fascicles of 2-6; pedun- 
cles 15-19 mm long, glabrous. Flowers subtended by peltate bracts, 2.3-2.4 mm 
long; calyx 2.8-3.8 mm long; petals free at base, partially united above, 5-5.2 mm 
long; filaments 6.2-8.8 mm long; ovary 2-2.3 mm long; style 7.8-8.8 mm long. 
Pods (9-) 10-13 cm long, (13-) 15-17 mm wide, (3-) 6-12 per capitulum, linear- 
oblong, acute apically, 8-14-seeded, valves mid-reddish brown, glabrous and slightly 
lustrous. Seeds 8-9.6 mm long, 4.7-6.3 mm wide. 

Additional Specimens Examined . Mexico. Chiapas: 2 km SE of Toliman rd to Ojo de Agua and 
Berriozabal, nr Motozintla de Mendoza, 15°19'N, 92°19'W, 10 Mar 1981, Sousa 11827 (CAS, K, 
MEXU). Guatemala. Huehuetenango: Huehuetenango, 10 Nov 1934, Skutch 1631 (A); rd from 
Huehuetenango NW to La Mesilla, 10 km WNW of Colotenango, valley of Rio Selegua, 15°27'N, 
91°46'W, 30 Mar 1988, Hughes 1132 (FHO, K, MEXU); rd W from Colotenango to Cuilco, 5 km W of 
Ixtahuacan, above Rio Cuilco, 15<'25'N, 9r48'W, 9 Apr 1991, Hughes 1469 (EAP, FHO, K, MEXU, 
NY); 1 km ENE of village of San Miguel, track running WSW from Ixtahuacan, into valley of Rio 
Cuilco, 15°23'N, 91°50'W, 28 Feb 1992, Hughes 1689 (E, FHO, K, MEXU, MO, NY). 

The characteristics and distribution of this subspecies remain poorly known. 
Further field exploration will be needed to assess whether this variant is as local- 
ized—in a small area of northern Guatemala and the immediate border zone in 
Mexico around Motozintla, largely in the valleys of the Rio Cuilco and Rio Sele- 
gua, in the Department of Huehuetenango and in Chiapas — as current collections 



CONTR. UNP 


^ERSITY OF MICHIGAN HERBARIUM 


VOLUME 21 


Subspecies ixtahuaca 


ma is named with reference to its ver 


■y localized dis- 


n centered on the sma 


11 town of Ixtahuacan. 





Leucaena magnifica (C. E. Hughes) C. E. Hughes, comb, et stat. nov. Leucaena 
shannonii subsp. magnifica C. E. Hughes, Kew Bull. 46: 547. 1991. — Type: 
Guatemala. Chiquimula: Llano Grande, 1.5 km SW of Quetzahepeque. 
rd towards Esquipulas, 14°37'N, 89°27'W, 26 Feb 1988, Hughes 1093 (holo- 
type: FHO!; isotypes: K! MEXU!). 

Leucaena magnifica was first discovered in 1984 (Hughes 1986) and originally 
described as a subspecies of L. shannonii (Hughes 1991). It is here recognized as a 
species distinct from L. shannonii based on its consistently once-branched, as 
opposed to unbranched, flowering shoots. Taking this character alone L. magnifica 
is clearly diagnosable by its unique combination of character states and thus com- 
prises a "phylogenetic species." Evidence from cpDNA and isozymes also sup- 
ports recognition of L. magnifica as a species distinct from L. shannonii. Harris et 
al. (1994), in their cpDNA analysis, showed that L. magnifica was the sister spe- 
cies to L. shannonii, a relationship supported in 85% of bootstrap replicates, and 
that they were separated by four autapomorphic fragment changes, two of which 
were unique. Chamberlain et al. (1996) analyzed isozyme variation across the L. 
shannonii alhance. Using the population aggregation analysis method of Davis 
and Nixon (1992) to delimit phylogenetic species, they showed that L. magnifica 
was separated from L. shannonii by the fixed occurrence of allele A of the Per-1 
isozyme system. In addition, L. magnifica and L. shannonii differ in a number of 
quantitative characters (Hughes 1991): L. magnifica has larger leaves with more 
pairs of pinnae, more pairs of leaflets, larger leaflets, larger flowers, more flowers 
per capitulum, larger pods and seeds, pods with dense velutinous pubescence, that are 
often deep maroon when unripe, and more pods per capitulum than L. shannonii. 

Leucaena matudae (S. Zarate) C. E. Hughes, comb, et stat. nov. Leucaena escu- 
lenta (Sesse & Mociiio ex DC.) Benth. subsp. matudae S. Zarate, Anales 
Inst. Biol. Univ. Nac. Auton. Mexico, Bot. 65: 134. 1994.— Type: Mexico. 
Guerrero: Casa Verde, nr Venta Viejo in the Caiion de Zopilote, "guaje 
chismoso," 17°50'N, 99°34'W, 12 Dec 1978, Halbinger 288 (holotype: 
MEXU!, isotypes: ENCB! MO! NY!). 

Leucaena matudae was originally described as a subspecies of L. esculenta 
(Zarate 1994), but is here raised to species rank in recognition of its clear mor- 
phological and molecular distinction from L. esculenta. Although cladistic analy- 
ses of morphology (Hughes, in press) and cpDNA (Harris et al. 1994) support the 
placement of L. matudae in the L. esculenta alhance, L. matudae is distinguished 
by a number of discrete character states. Most notably, the petiolar nectary of L. 
matudae is stipitate, erect, cylindrical, and quite unhke the large sessile, elongate, 
concave, crateriform nectary of L. esculenta. In addition, L. matudae may be 
distinguished by its terete, as opposed to the angular, ridged shoots of L. esculen- 
ta, fewer pinnae pairs and leaflets per pinna, fewer flowers per capitulum, smaller, 
weakly constricted pods which are partitioned between the seeds, and oblique 
alignment of seeds in the pods. Although L. matudae shares the same bark type as 
the remaining species in the L. esculenta alhance (thick corky bark with a single 



1997 HUGHES: LEUCAENA 287 

periderm, a pale metallic grey surface, and deep blood-red inner bark), the bark 
surface pattern in L. matudae is quite distinct and unique within the genus in 
having a scalloped surface resulting from shedding of small circular plates. Zarate 
(1994) mentioned use of the bark for medicinal purposes and attributed the un- 
usual surface pattern to local harvesting of bark for medicinal use, but given that 
there is no evidence of harvesting in most areas, that the patterns are extremely 
regular, and that they are found on inaccessible branches as well as boles, it seems 
clear that this is the natural state. Finally, Harris et al. (1994) emphasized the 
clear distinction in cpDNA between L. matudae (treated by them as L. esculenta 
subsp. matudae) and L. esculenta. Recognition of L. matudae as a separate species 
is thus amply justified by a suite of discrete character states that separate it from 

Leucaena confertiflora S. Zarate, Anales Inst. Biol. Univ. Nac. Auton. Mexico, 
Bot. 65: 148. 1994. — Type: Mexico. Oaxaca: Matatlan, Cerro Nueve Pun- 
tas nr shortwave relay mast, 5 km S of Matatlan, 16°49'N, 96°21'W, 2050 
m, 22 Jan 1980, Zarate with Reid 428 (holotype: MEXU!). 

The material of L. confertiflora from San Pedro Chapulco, Puebla, Mexico, 
and the surrounding area near the village of Azumbilla and the northern fringes 
of the Sierra Zongolica was observed by Zarate (1994) to differ in the shape of 
the extrafloral petiolar and leaf rachis nectaries from material from the remainder 
of the distribution further south in Puebla and Oaxaca. This difference was used 
by Zarate (1994) to divide L. confertiflora into two subspecies: subsp. confertiflora 
with sessile, discoid or shallow crateriform, concave nectaries on the petiole and 
rachis, and subsp. adenotheloidea with stipitate, erect peg-shaped, cylindrical nec- 
taries, usually occurring at the base of each pair of pinnae. Zarate (1994) dis- 
cussed the lack of other distinguishing features between these two subspecies. 
While the nectary shape is clearly distinct at the extremes, and fixed in most 
areas, variation within populations and even within individuals has been observed. 
Firstly, material from the mountains north-west of Huajuapan de Leon, around 
Santa Catarina Zapoquila and Membrillos (Hughes 1812), has both gland types 
within populations indicating the presence of both infraspecific taxa in this area. 
Secondly, in the village of Santa Catalina Oxolotepec, in the Sierra Zongolica, 
both nectary types and intermediate forms are found within the cultivated mate- 
rial there and even on single individuals (Hughes 1616, 1731, 1799). Inconstancy 
of the nectary shape in some areas and the lack of other distinguishing features 
justify treatment at varietal rather than subspecific rank necessitating the new 
combination: 

Leucaena confertiflora S. Zarate var. adenotheloidea (S. Zarate) C. E. Hughes, stat. 
nov. Leucaena confertiflora subsp. adenotheloidea S. Zarate, Anales Inst. 
Biol. Univ. Nac. Auton. Mexico, Bot. 65: 149. 1994.— Type: Mexico. Puebla: 
San Pedro Chapulco, 13 km NE of Tehuacan on rd towards Orizaba, 18°36'N, 
97°25'W, 2100 m, 7 Dec 1981, Zarate with Reid 610 (holotype: MEXU!). 

lanceolata S. Watson, Proc. Am. Acad. Arts 21: 427. 1886.— Type: Mexico. 
Chihuahua: Batopilas, Hacienda San Miguel, SW Chihuahua, 27°53'N, 
108°26'W, Sep 1885, Palmer 6 (holotype: NY!; isotypes: UC! US! GH!). 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



Leucaena lanceolata is an extremely variable species across its extensive and 
largely continuous distribution along the Pacific coast of Mexico from Sonora to 
Chiapas, with outlying occurrences in Baja California and Veracruz. Zarate (1994) 
described the new subspecies sousae to account for the variation encountered towards 
the southern limits in Michoacan and Oaxaca. A detailed study of morphological 
variation within L. lanceolata was undertaken to investigate the basis for this 
subdivision. This study shows that there are no clear discontinuities in quantita- 
tive leaf or pod traits across the range of L. lanceolata that might be used to 
divide the species unambiguously. Nevertheless, there is some evidence to sup- 
port the recognition of subsp. sousae. Variation in leaflet and pod traits shows 
some correlation with geography and with leaflet and pod vestiture, and two 
broad groups can be distinguished as recognized by Zarate (1994). Moreover, 
Harris et al. (1994) mentioned the existence of two cpDNA plastome types within 
L. lanceolata across the seven accessions included in that study. Further analysis 
shows that cpDNA variation within L. lanceolata is not perfectly correlated with 
the morphological variation, although the two accessions of subsp. sousae did 
group together, lending some support for recognition of that taxon. Pending more 
detailed analysis and given the lack of clear discontinuities, subsp. sousae is here 
demoted to varietal rank: 

Leucaena lanceolata var. sousae (S. Zarate) C. E. Hughes, stat. nov. Leucaena 
lanceolata subsp. sousae S. Zarate, Anales Inst. Biol. Univ. Nac. Auton. 
Mexico, Bot. 65: 117. 1994.— Type: Mexico. Oaxaca: 17 km WNW of 
Puerto Escondido, Dtto. Juquila, 15°57'N, 97°13'W, 21 Oct 1976, Sousa 
6390 (holotype: MEXU!, isotype UC!). 
Leucaena rekoi Britton &. Rose, Fl. N. America. 23: 122. 1928.— Type: Mexi- 
co. Oaxaca: nr Pochutla, close to the Pacific coast, 15°44'N, 96°28'W, 28 
Sep 1917, Reko 3632 (lectotype, here designated, flowering shoot and 
leaves only: US!). 

Discontinuities in the morphological variation within L. lanceolata are blurred 
by three exceptions to the general trends in leaflet and pod size and vestiture 
identified as the basis for recognition of var. sousae. Firstly, pods which are gener- 
ally pubescent in var. lanceolata and glabrous in var. sousae, are exceptional in 
several areas. In Baja California, pods are glabrous, whereas quantitative traits 
indicate that this material belongs with var. lanceolata (see Hughes 1544). In the 
transition area between var. sousae and var. lanceolata in a well-defined zone 
around Bahia de Santa Cruz in south-central Oaxaca (see Hughes 84L 587, 835, 
836), pods are pubescent, while the large leaflets and pods indicate that this mate- 
rial belongs with var. sousae. At inland, slightly higher-elevation, drier sites in 
southern Oaxaca (see Hughes 556, 1345, 1724) pods are glabrous, with an unusual 
glossy or 'basted' surface. Secondly, this same material from inland Oaxaca from 
the Municipios of San Bartolo Yautepec, Jalapa de Marques, and Santiago Lachigu- 
irf also has wider pods than typical for var. lanceolata, and pods which are slightly 
falcate. Finally, in the coastal zone 20-40 km west of Playa Azul in Michoacan, it 
is difficult to separate the two varieties, which appear to grow together there. 

These difficulties were recognized by Zarate (1994) in his discussion of var. 
sousae (as subsp. sousae) when he stated that although readily distinguished at the 
type locality, in other areas it may be more difficult to define. Zarate (1994) 



1997 HUGHES: LEUCAENA 289 

attributed this to iiis hypothesis that it is of hybrid origin between L. macwphylla 
(L. macrophylla subsp. nelsonii sensu Zarate) and L. lanceolata var. lanceolata, 
although he did not provide conclusive evidence to confirm this hypothesis. 

ACKNOWLEDGMENTS 

I particularly thank Jon Hellin and Caspar Alvarado for drawing my attention to L. lempirana. I 
also thank Richard Palmer for providing the Latin descriptions, Rosemary Wise for the botanical 
drawing of L. lempirana, Sergio Zarate for sharing his many insights and a copy of his revision of the 
Mexican species in advance of publication, David Mabberley and Roger Polhill for their advice, 
support, and encouragement, and the curators of A, BM, CAS, EAP, F, FHO, GH, HEH, K, LAGU, 
M, MEXU, MO, NY, OXF, TEX, UC, UNAH, US. and W for providing material and/or facilities. 
Financial support was provided by the Forestry Research Programme of the Overseas Development 
Administration of the UK government (Projects R.4524 and R.6296). 



LITERATURE CITED 

m, R. B. 1992. Phylogenetic species concepts. Trends Ecol. Evol. 7: 1-2. 

wbaker, J. L. 1980. What is 'giant' Leucaenal Leucaena Res. Rep. 1: 43-44. 

— . 1987. Species in the genus Leucaena. Leucaena Res. Rep. 7: 6-20. 

wbaker, J. L., and G. M. Ito. 1980. Taxonomic studies of the genus Leucaena. Leucaena News. 1: 

41-42. 
wbaker, J. L., and C. T. Sorensson. 1994. Domestication of lesser-known species of Leucaena. In 
Tropical trees: the potential for domestication and rebuilding forest resources, ed. R. R. B. Leakey 
and A. C. Newton, 195-204. London: HMSO. 
wbaker, J. L., D. L. Plucknett, and V. Gonzalez. 1972. Varietal variation and yield trials of 
Leucaena leucocephala (Koa haole) in Hawaii. Hawaii Agric. Exp. Stn. Res. Bull. 166: 1-29. 
ton, N. L., and J. N. Rose. 1928. Mimosaceae. N. Amer. Fl. 23: 121-131. 

imberlain, J. R., C. E. Hughes, and N. Galwey. 1996. Patterns of isozyme variation in the Leucae- 
na shannonii alliance (Leguminosae: Mimosoideae). Silvae Genet. 45: 1-7. 
Cracraft, J. 1983. Species concepts and speciation analysis. Current Ornith. 1: 159-187. 

. 1992. The species of the Birds of Paradise (Paraisaedieae): applying the phylogenetic species 

concept to a complex pattern of diversification. Cladistics 8: 1-43. 
Cronquist, A. 1988. The evolution and classification of flowering plants. 2d ed. Bronx, New York: 

New York Botanical Garden. 
Davis, J. I. 1995. Species concepts and phylogenetic analysis - introduction. Syst. Bot. 20: 555-559. 
Davis, J. I., and K. C. Nixon. 1992. Populations, genetic variation and the dehmitation of phylogenetic 

Donoghue, M. J. 1985. A critique of the biological species concept and recommendations for a 

phylogenetic alternative. Bryologist 88: 172-181. 
Eldredge, N., and J. Cracraft. 1980. Phylogenetic patterns and the evolutionary process. New York: 

Columbia University Press. 
Gonzalez, V., J. L. Brewbaker, and D. E. Hamill. 1967. Leucaena cytogenetics in relation to the 

breeding of low mimosine lines. Crop Sci. 7: 140-143. 
Gray, S. G. 1967. Inheritance of growth habit and quantitative characters in intervarietal crosses in 

Leucaena leucocephala. Austral. J. Agric. Res. 18: 63-70. 
Harris, S. A., C. E. Hughes, R. Ingram, and R. J. Abbott. 1994. A phylogenetic analysis of Leucaena 

(Leguminosae: Mimosoideae). PI. Syst. Evol. 191: 1-26. 
Hughes, C. E. 1986. A new Leucaena from Guatemala. Leucaena Res. Rep. 7: 110-113. 
. 1991. Two new subspecies of Leucaena (Leguminosae: Mimosoideae) from Guatemala. Kew 

Bull. 46: 547-557. 
. Monograph of Leucaena (Leguminosae: Mimosoideae). Systematic Botany Monographs; in 

Hughes, C. E., and S. A. Harris. 1994. The characterisation and identification of a naturally occurring 



UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 



ton, E. M., and S. G. Gray. 1959. Problems of adapting Leucaena ghuica as a forage for the 
Australian tropics. Empire J. Exp. Agric. 27: 187-196. 
kow, M. 1995. Species concepts: assumptions, methods and applications. Syst. Bot. 20: 589-605. 
^augh, R. 1987. Flora Novo-Galiciana: a descriptive account of the vascular plants of western 

hler, B. D., and A. F. Budd. 1990. Species and evolution in clonal organisms - introduction. Syst. 

Bot. 15: 79-85. 
son, G., and N. I. Platnick. 1981. Systematics and biogeography: cladistics and vicariance. New 

York: Columbia University Press, 
an, K. C, and Q. D. Wheeler. 1990. An amplification of the phyiogenetic species concept. Cladis- 



Panchen, A, 


. L. 1992. Classificat 


ion, evoh 


ition and 


the nature 


of biology. 


. Cambridge 


: Cambridge 




.ity Press. 














Rieseberg,L 


.. H., and L. BrouiUet 


. 1994. Are many ph 


ant species 


paraphyleti. 


::? Taxon 43: 


21-32. 


Rosen, D. E. 1979. Fishes from 


the uplar 


ids and in 


itermontan 




Guatemala. 






and comparative biogeography 


. Bull. Am 






• 267-376. 




Wendell, K. 


1916. Honduras in pictures, ne- 


w York: Sterling Pub 


iishing. 






Zarate P., S 


. 1984. Taxonomic r. 


svision ol 


" the genu 


s Leucaem 




:ico. Bull. Ir 


itern. Group 


Study N 


4imosoideae 12: 24-3. 














. 1987 


. Taxonomic identity 


of Leuca, 


ena leucoc 


ephala (La 


m. ) de Wit 


with a new i 


combination. 


Phytologia 63: 304-306. 














. 1994 


. Revision del generc 


> Leuccwn 


a Benth. e 


n Mexico. 


Anales Inst 


. B.ol. Univ. 


Nac. Auton. 


Mexico 


, Bot. 65: 83-162. 















TWO NEW SPECIES OF CAREX (CYPERACEAE) 
FROM MEXICO 

A. A. Reznicek 
University of Michigan Herbarium 

North University Building 
Ann Arbor, Michigan 48109-1057 

M. S. Gonzalez-Ehzondo 

CllDIR y COFAA-IPN 

Apdo. Postal 738 

Durango, Dgo., 34000, Mexico 

Recent field work has disclosed two additional species in Carex section Hy- 
menochlaenae sensu lato, one of the largest sections in the Mexican Carex flora, 
comprising 18 species including the two described below. One of these is a mem- 
ber of the complex centered on C. mackenziana Weath., which is distinguished 
from other members of the section in Mexico by pubescent perigynia. This group 
is discussed in some detail by Reznicek (1990). Three species were known in the 
complex, all local endemics, and a fourth is added below. The other new species is 
related to C. flexirostris Reznicek, from Oaxaca. 

Carex queretarensis Reznicek & S. Gonzalez, sp. nov. — Type: Mexico. Queretaro: 
Mpio. Landa de Matamoros, ca. 7.5 (air) km SW of El Lobo (Km 226) on 
road (Hwy 120) to Jalpan, between El Lobo and Landa, 2ri5'50"N, 
99°10'13"W, 1590-1600 m, 14 Jul 1996, Gonzalez & Reznicek 10284, M. 
Pinedo (holotype: lEB!; isotypes: ANSM! CIIDIR! ENCB! GH! MEXU! 
MICH! MO! QMEX! TEX! UAMIZ! US!). Fig. 1. 

Plantae cespitosae; culmi 30-80 cm ahi; vaginae basales purpurascentes, sca- 
brae. Folia 5-9, plerumque basalia; laminae 6-45 cm longae, 2.5-6.2 mm latae; 
vaginae 1.5-12 cm longae, hispidulae; ligulae 1.1-5 (-6.4) mm longae. Inflorescen- 
tiae 9-41 cm longae; spicae (3-) 4-7; spica terminalis plus minusve erecta; spicae 
laterales androgynae interdum pistillatae, pendulae; bracteae infimae laminis 2.3- 
24 cm longis, 0.8-3.6 mm latis et vaginis 1.1-7 cm longis. Squamae pistillatae 
staminataeque pubescentes. Perigynia (3.6-) 4.1-5.6 mm longa, 1.4-1.6 mm lata, 
trigona, viridia, pubescentia, ascendentia, in rostrum contracta; rostra 0.3-0.6 mm 
longa. Achenium 2.4-2.8 mm longum, 1.2-1.4 mm latum. Styli marcescentes; stig- 
mata 3. Antherae 3, 2.3-3 mm longae. 

Plants loosely cespitose, with short, thick rhizomes; roots brown to dark brown, 
not densely felted with root hairs; fertile culms 30-80 cm tall, trigonous, smooth, 
with scabrous, purple, bladeless sheaths with stramineous veins. Leaves 5-9, mostly 
basal; blades 6-45 cm long, 2.5-6.2 mm wide, plicate, more or less hispidulous on 
both surfaces just above ligule, otherwise glabrous, the margins antrorsely scabrous, 
the widest leaves 4.2-6.2 mm wide; leaf sheaths 1.5-12 cm long, more or less 
tightly enveloping culms, hispidulous with red prickle-hairs, especially on lower 
sheaths, strongly purple-tinged near base; inner band of sheaths finely hispidulous 



292 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

or glabrous, stramineous to purple-mottled, the apex concave; ligules 1.1-5 (-6.4) 
mm long, obtuse to acute, the free portion entire, erose, or slightly ciliate. Vegeta- 
tive shoots ca. 15-40 cm tall; leaves 5-9, similar to those of fertile culms; pseudoc- 
ulms 2-10 cm tall. Inflorescences 9-41 cm long, with the upper 2-4 spikes some- 
what overlapping and the lowest 2 spikes 2-30 cm distant; spikes single at nodes, 
with all but the upper pendulous on filiform, flexuous, smooth to slightly sca- 
brous-angled peduncles; lowermost spikes with peduncles 2.5-17.5 cm long, the 
uppermost lateral spikes with peduncles 0.4-1.7 cm long; lowermost bracts with 
blades 2.3-24 cm long and 0.8-3.6 mm wide and sheaths 1.1-7 cm long, the upper- 
most bracts much reduced. Spikes (3-) 4-7, the terminal staminate proximally and 
distally, and pistillate in the middle, occasionally staminate, the lateral androgy- 
nous or the lower sometimes pistillate. Terminal spikes 1.4-5.2 cm long, 1.5-2.9 
mm wide, ca. 20-40-flowered if staminate, otherwise the staminate portion (0-) 
0.5-1.5 cm long and 3-8-flowered proximally, and 0.4-1.8 cm long and 15-35- 
flowered distally; pistillate middle portion 0.3-2.5 cm long, 5.5-7 mm wide, 1-40- 
flowered; peduncles (2-) 4-9 (-17) mm long. Lateral spikes 1-5.3 cm long; stami- 
nate portion (0-) 0.2-1.5 cm long, (0-) 3-25-flowered; pistillate portion 1-4.4 cm 
long, 8-35-flowered. Pistillate scales 2.8-5.2 mm long, 1.5-2.1 mm wide, ovate to 
oblong, acute to acuminate-awned with scabrous awn up to 1.8 mm long, finely 
and sparsely pubescent distally, as well as scabrous on the midvein, green with 
broad, hyaline, whitish to pale brown margins, 3-veined. Staminate scales 3.9-5.8 
mm long, 1.4-1.9 mm wide, ovate to narrowly ovate, obtuse to acuminate, occa- 
sionally with a scabrous awn up to 0.4 mm long, glabrous or finely scabrous, 
especially on and near the midvein, green with broad, hyaline, whitish to pale 
brown margins, 3-veined. Perigynia (3.6-) 4.1-5.6 mm long, 1.4-1.6 mm wide, 
ascending, trigonous with more or less flat, elliptic to obovate sides, green, pubes- 
cent over most of the body, cuneately tapered to a short-stipitate base, ca. 15-19- 
veined with 3-5 veins prominent and the rest faint, tapered into a beak; beaks 0.3- 
0.6 mm long, green, hispidulous, the apex oblique or irregularly bidentate, hya- 
line. Achenes 2.4-2.8 mm long, 1.2-1.4 mm wide, trigonous with more or less flat, 
obovate sides, tightly enveloped by the perigynium, pale brown, short-stipitate. 
Styles withering; stigmas 3. Anthers 3, 2.3-3 mm long. 

Additional Specimens Examined. Mexico. Queretaro: Mpio. Landa de Matamoros, 1.5 km al 
W del Parador de Santa Martha, 1700 m, 20 Aug 1991, Cananza G. 3421 (CIIDIR, lEB); Mpio. 
Landa de Matamoros, ca. 6 km SW of El Lobo on road (Hwy 120) to Jalpan, 21°16'35"N, 99°09'21"W, 
1750-1760 m, 14 Jul 1996, Gonzalez & Reznicek 10283, M. Pineda (CAS, CHAP, CHAPA. CIIDIR, 
ENCB, F, GENT, GH, IBUG, lEB, K, MEXU, MICH, MO, NY, TEX, US, WIS). 

Carex queretarensis is locally frequent in pine-oak and oak-pine-juniper forest 
in the Municipio of Landa de Matamoros at ca. 1590-1760 m. Plants collected July 
14 ranged from flowering to mature fruit, so fruiting presumably occurs from mid- 
July through August. The epithet queretarensis refers to the species being appar- 
ently endemic to the state of Queretaro. 

All species of this complex resemble each other rather closely, but examina- 
tion shows consistent, correlated differences in a number of features (Table 1). 
The variation appears to be neither clinal nor simply quantitative. All the taxa 
appear to be geographically isolated, local, endemic species. 

Carex queretarensis is most similar to C. pubigluma Reznicek, sharing with 
that species the pubescent pistillate and staminate scales and long anthers. It 



REZNICEK & GONZALEZ: NEW S 




1 represented by Gonzalez & Rezi 



differs from C pubigluma, however, in having hispidulous lower sheaths and 
more pointed pistillate scales; in these two characters it resembles C. caeligena 
Reznicek, which has smaller perigynia, shorter anthers, more strongly aggregated 
upper spikes, and longer hgules. Differences among the four known species of the 
complex are summarized in Table 1. A revised key to this group is presented 
below. 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



length (mm) 
Pistillate scale 



Lower leaf sheaths hispidulous with red prickle-hairs; pistillate scales aculc u 




m.uc- 


2. Pistillate scale bodies glabrous or scabrous on the midvein; ligules (5-) 5.5- 


14 mm 


long; 


anthers 1.3-2.5 mm long. 




. cm'Uf^c. 


2. Pistillate scale bodies pubescent; ligules 1.1-5 (-6.4) mm long; anthers 2.3 3 n 


im lo.u 




Lower leaf sheaths glabrous; at least some pistillate scales obtuse. 


' 




3. Pistillate and staminate scales pubescent, the margins ciliate; perigynium beak: 


> with ; 


1 hva- 


line, flared, ciliate apex. 


(^■1 


•mhi^huu 








perigynium beaks more or less bidentulate at apex. 


C. ma 


ckcn:i(ih 



Carex pinophila Reznicek & S. Gonzalez, sp. nov.— Type: Mixico. Qucrelaro: 
Mpio. Landa de Matamoros, ca. 2 km SW of Pinal de Amoles on road 
(Hwy 120) to San Juan del Rio, 21°07'47"N, 99°38'02"W. 27(H) m. 15 Jul 
1996, Gonzalez & Reznicek 10293, M. Pineda (holotype: lliB!: isolvpes: 
CIIDIR! MEXU! MICH!). Fig. 2. 

Plantae cespitosae; culmi 70-90 cm alti; vaginae basales rubescentes, glabrae. 
Folia ca. 11-15 plerumque basalia; laminae (6-) 15-60 cm longae, (1.3-) 2.1-3.6 
mm latae; vaginae 3.2-10 cm longae, stramineae, rubrotinctae, glabrae; ligulae 



REZNICEK & GONZALEZ: NEW SPECIES OF CAREX 




296 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

2.5-5.5 mm longae. Inflorescentiae 7-1 1 cm longae; spicae 3-5; spica terminalis 
plus minusve erecta; spicae laterales androgynae, infimae pendulae; bractae infi- 
mae laminis 3.1-6 cm longis, 0.7-1.3 mm latis et vaginis 1.2-1.6 cm longis. Perigy- 
nia 2.3-3 mm longa, 0.9-1.2 mm lata, ascendentia, trigona, viridia, purpureomacu- 
lata, glabra, in rostrum contracta; rostra 0.4-0.7 mm longa. Achenium ca. 1.5-1.7 
mm longum, 0.8-1.1 mm latum. Styli marcescentes; stigmata 3. Antherae 3, ca. 
1.4-1.8 mm longae. 

Plants cespitose in small clumps or solitary, with short, knotty rhizomes; roots 
brown, not densely felted with root hairs; fertile culms ca. 70-90 cm tall, trigo- 
nous, smooth, with glabrous, reddish, bladeless sheaths with stramineous veins. 
Leaves ca. 1 1-15, mostly basal; blades (6-) 15-60 cm long, (1.3-) 2.1-3.6 mm wide, 
flat to plicate, glabrous, the margins and midrib smooth or finely antrorsely sca- 
brous distally, the widest leaves ca. 3.2-3.6 mm wide; leaf sheaths 3.2-10 cm long, 
tightly enveloping the culms, glabrous, red-tinged; inner band of sheaths hispidu- 
lous near apex, stramineous and faintly red-striped, faintly veined, the apex con- 
cave, thickened and finely (and sometimes inconspicuously) ciliate; hgules ca. 2.5- 
5.5 mm long, acute, usually reduced on upper leaves, the free portion thickened 
and scabrous, reddish to stramineous. Vegetative shoots ca. 40-60 cm tall; leaves 
5-7, similar to those of fertile culms; pseudoculms ca. 6-10 cm tall. Inflorescences 
ca. 7-11 cm long, with the upper spikes strongly overlapping and the lowest 2 
spikes 2.5-4.7 cm distant; spikes single at nodes, the lower pendulous, the upper 
ascending to erect, on filiform, flexuous, smooth or sparsely scabrous peduncles; 
lowermost spikes with peduncles (1.3-) 1.9-2.6 cm long, the uppermost lateral 
spikes with peduncles ca. 0.7 cm long; lowermost bracts with blades 3.1-6 cm long 
and 0.7-1.3 mm wide, dorsally hispidulous along the veins, the sheaths 1.2-1.6 cm 
long, the uppermost bracts much reduced. Spikes 3-5, the terminal staminate 
proximally and distally, and pistillate in the middle, the lateral androgynous. Ter- 
minal spikes 3.2-4 cm long, staminate portion 9.2-13 mm long, 1.2-2.3 mm wide, 
and 10-14-flowered proximally, ca. 13-18 mm long and 55-60-flowered distally; 
pistillate middle portion ca. 6.5-10.5 mm long, 3.6-3.8 mm wide, 13-18-flowered; 
peduncles ca. (5.6-) 8.8-13.2 mm long. Lateral spikes (2.4-) 3.3-4.7 cm long; 
staminate portion 6.7-16 mm long, ca. 16-45-flowered; pistillate portion (10.5-) 
19.5-30 mm long, 3.3-4.4 mm wide, ca. (8-) 30-40-flowered. Pistillate scales 2.6- 
3.3 mm long, 1.3-1.8 mm wide, ovate, obtuse to acute, glabrous, reddish purple 
with narrow hyaline margins, faintly 5-7-veined. Staminate scales 2.9-4 mm long, 
1.1-1.8 mm wide, narrowly elliptic to obovate, obtuse to acuminate, glabrous, 
reddish purple with narrow hyaline margins, 1-3 (-9)-veined. Perigynia 2.3-3 mm 
long, 0.9-1.2 mm wide, ascending, more or less flattened-trigonous with more or 
less flat, obovate sides, green with reddish purple mottling especially distally (some- 
times uniformly purple distally), scabrous on the main veins just below the beak, 
more or less cuneately tapered to the base, ca. 11-16-veined with 2 veins promi- 
nent and the rest fainter, contracted into a beak; beaks 0.4-0.7 mm long, green 
with dense reddish purple mottling, scabrous on the main veins, the apex oblique, 
erose. Achenes ca. 1.5-1.7 mm long, 0.8-1.1 mm wide, trigonous with more or less 
flat, obovate sides, tightly enveloped by the perigynium, brown, tapered to the 
base. Styles withering; stigmas 3. Anthers 3, ca. 1.4-1.8 mm long. 

Carex pinophila is known only from the type, collected in Queretaro in deep 
pine duff in a moist forest of Finus patula Schltdl. & Cham, on a steep slope at 
2700 m. Plants were rare and localized at the site, and few individuals were fruit- 
ing, perhaps due to the heavily shaded habitat. On July 15, plants were somewhat 



1997 REZNICEK & GONZALEZ: NEW SPECIES OF CAREX 297 

overmature, so fruiting probably occurs through June and into early July. The 
epithet pinophila refers to the habitat of the type. 

This apparently very rare species is related to Carex perlonga Fernald, C. flexiros- 
tris Reznicek, and C chiapensis F. J. Herm. These four Mexican species of section 
Hymenochlaenae are relatively narrow-leaved plants of moist forests, with red 
bases, terminal spikes staminate proximally and distally and pistillate in the mid- 
dle (rarely staminate), glabrous perigynia contracted into a distinct beak (except 
in C. chiapensis), and mostly awnless pistillate and uniformly awnless staminate 
scales. From all these species, C pinophila can be separated easily by its smooth 
lower sheaths, and short perigynia 2.3-3 mm long. Carex chiapensis, C. flexirostis, 
and C perlonga have the lower sheaths sparsely to densely hispidulous with red 
prickle-hairs (except sometimes in C. chiapensis with perigynia 3.9-6 mm long). 
All three species also tend to have larger perigynia 2.7-6 mm long, and have 
longer inflorescences (7.5-) 10^1 cm long, and longer lower inflorescence bracts 
(3.6-) 6-32 cm long with sheaths (1-) 1.6-5.9 cm long. Carex pinophila has shorter 
inflorescences 7-11 cm long with short, small bracts with blades 3.1-6 cm long and 
with sheaths only 1.2-1.6 cm long. Carex chiapensis and C. perlonga have broad 
lower inflorescence bracts 1.7-5 mm wide, but C pinophila has narrow inflores- 
cence bracts 0.7-1.3 mm wide. In the last feature C. pinophila most closely resem- 
bles C. flexirostris, which has the lower bracts with blades only 1-1.9 cm wide. If a 
small individual of C flexirostris with smooth sheaths were to occur (or if the 
sometimes sparse prickle hairs were overlooked), C pinophila could be distin- 
guished by its thickened apex to the inner band of the sheath, perigynia not at all 
outcurved, and shorter achenes 1.5-1.7 mm long. Carex flexirostris has thin, frag- 
ile apices to the inner band, at least slightly outcurved perigynia, and achenes 1.8- 
2.1 mm long. 

Because of its smooth sheaths, Carex pinophila would key to C. pertenuis L. H. 
Bailey rather than C. flexirostris in the keys in Reznicek (1986) and Jones and 
Reznicek (1992). These two species can be readily distinguished: C pertenuis has 
± elongate rhizomes, all the spikes androgynous, awned pistillate and staminate 
scales, and perigynia gradually tapered to the apex, whereas C. pinophila has 
short, knotty rhizomes, the terminal spike staminate distally and proximally and 
pistillate in the middle, awnless pistillate and staminate scales, and perigynia con- 
tracted into a distinct beak. 



LITERATURE CITED 



:nicek, A. A. 1986. The taxonomy of Carex sect. Hymenochlaenae (Cyperac 

Central America. Syst. Bot. 11: 56-87. 
— . 1990. Four new species of Carex (Cyperaceae) from Mexico, with not 

Carex Flora. Contributions Univ. Michigan Herb. 17: 279-293. 



CAREX MOLESTIFORMIS (CYPERACEAE), 

A NEW SPECIES OF SECTION OVALES 
FROM THE OZARK MOUNTAIN REGION 

A. A. Reznicek 
University of Michigan Herbarium 

North University Building 
Ann Arbor, Michigan 48109-1057 

Paul E. Rothrock 

Randall Environmental Center 

Taylor University 

500 W. Reade Avenue 

Upland, Indiana 46989-1001 



INTRODUCTION 

Recent field research in Arkansas and Oklahoma disclosed the ( 
an entity in Carex section Ovales that seemed similar to both C brevior (Dewey ) 
Mack, and C. molesta Mack, ex Bright, but which differed consistently in several 
characters. Closer examination of herbarium material and more detailed field 
work revealed that this plant is a regional endemic within the C. brevior group 
restricted to the Ozark mountain region, in western Arkansas, southern Missouri, 
and eastern Oklahoma. 

Carex section Ovales, a well-defined, species-rich group of about 85 species 
most diverse in North America, is characterized by gynaecandrous spikes, wing- 
margined perigynia, distigmatic flowers, and a more or less cespitose habit with 
true vegetative culms present. The C. brevior group is one of the most difficult 
and poorly understood groups of species within eastern North American members 
of section Ovales. Diagnostic features of the C. brevior group [a major portion of 
Mackenzie's (1931) "subsection" Festucaceae] include medium to large, broadly 
winged perigynia (mostly ca. 2-6 mm wide and 3.5-8 mm long) with broadly 
elliptic, broadly ovate, orbicular, or even reniform bodies, and leaf sheaths which 
usually have a conspicuous hyaline zone on the inner band. The C. brevior group 
presently includes nine entities in eastern North America, C bicknellii Britton 
var. bicknellii, C. bicknellii var. opaca F. H. Herm., C. brevior, C. festucacea Willd., 
C hyalina Boott, C. merritt-fernaldii Mack., C molesta, C. reniformis (L. H. Bailey) 
Small, and C. tetrastachya Scheele (=C. brittoniana L. H. Bailey). Several other 
species occurring west of the Rocky Mountains and in Mexico may also belong to 
this group (Mackenzie 1931). 

Though close to C. brevior and C. molesta, this Ozark Mountain region entity 
clearly possesses a unique combination of morphological features. The inflores- 
cences bear a close resemblance to those of C. molesta, compact with few spikes 
that lack conspicuous staminate bases, but it differs from C molesta in having 
larger perigynia that are more prominently nerved and with larger achenes. The 
large achenes are similar to those of C brevior, but C brevior has more elongate 
inflorescences with more spikes, at least the terminal spike normally conspicuously 



299 



. UNIVERSITY OF MICHIGAN HERBARIUM 



nearly so adaxially. Because of its distir 

as a new species, a locally frequent regional endemic. 

Carex molestiformis Reznicek & P. Rothr., sp. nov.— Tvpt-:: U.S.A. Oklahoma. 
Adair Co.: S bank of Illinois River, just E of bridge for Okla. Hwy 59, ca. 
1.5 mi N of Watts, NE 1/4 section 18, T19N R26E, 26 May 1994, A. A. 
Reznicek 9777 & S A. Reznicek, S. D. Jones, G. D. Jones (holotype; MICH!; 
isotypes: BRCH! BRIT/SMU! CM! ctb (Herb. Charles T. Bryson)! F! 
FTG! GENT! GH! K! KANU! KNK! MIN! MO! NLU! NY! OKL! PH! 
TENN! TEX! UARK! US! VDB! WIS!). 

Plantae cespitosae; culmi fertiles 25-120 cm alti; vaginae basales brunneae. 
Culmi vegetativi erecti, annul. Folia 3-5; laminae 2.5-32 cm longae, 1.7-3.8 mm 
latae; vaginae ca. 1-10 cm longae, ventraliter membranaceae, pallide brunneae vel 
albidae, hyahnae, laeves. Inflorescentiae 1.2-2.7 (-3.4) cm longae, erectae, con- 
gestae; spicae 2-4 (-5), gynaecandrae, ovoideae vel globosae, 7-12 mm longae, 5- 
10 mm latae, basi staminata 1-5 (-9.5) mm longa. Squamae pistillatae pallide 
brunneae, obtusae vel acutae. Perigynia 4-5.4 (-6.1) mm longa, (2.4-) 2.6-3.4 mm 
lata, adpresso-ascendentia, corpibus late ovatis, late ellipticis, vel rotundis, in ros- 
trum serrulatum 1.1-1.9 mm longum contracta. Achenium 1.6-2 mm longum, 1.4- 
1.8 mm latum, biconvexum. Stigmata 2. Antherae 3, (1.4-) 1.6-2.6 mm longae. 

Cespitose in small clumps from thick, woody, very short-creeping rhizomes; 
fertile culms 25-120 cm tall, stiffly erect, trigonous, smooth except for finely- 
scabrous angles just below inflorescence; bladeless basal sheaths medium to dark 
brown, disintegrating into short, dark brown fibers. Leaves 3-5, on lower 1/5-2/5 
of the culm; blades 2.5-32 cm long, 1.7-3.8 mm wide, pHcate, glabrous or papillose 
adaxially, the margins and midrib antrorsely scabrous distally; leaf sheaths ca. 1- 
10 cm long, tightly enveloping culms, smooth, yellow-green, the intervenal areas ± 
white with scattered septa; the inner band of sheaths glabrous, its apex concave or 
truncate and reaching from the base of the blade to 1.5 mm above, light brown to 
whitish, sometimes brown-tinged at the summit, hyaline; ligules 1-2.8 mm long, 
rounded, the free portion entire and up to 0.5 mm long. Vegetative culms differ- 
ent from fertile, fully developed only after perigynia are largely shed, annual, 5- 
40 cm tall with ca. 9-14 leaves clustered near the apex of the culm. Inflorescences 
1.2-2.7 (-3.4) cm long, erect or occasionally angled above the lowest spike, the 
spikes overlapping or congested, the lowest spikes 1.5-6 (-9) mm apart, spikes 
single at nodes, sessile, lowermost bracts scalelike or setaceous, 0.4-1 .5 cm long, 
inconspicuous, sheathless, upper bracts much reduced; spikes 2-4 (-5), gynaecan- 
drous, globose to ovoid with rounded or tapered bases, 7-12 mm long, pistillate 
portion 6-10 mm long, 5-10 mm wide, (10-) 15^0-flowered, staminate portion 1- 
5 (-9.5) mm long, 1.2-2.5 mm wide, ca. 6-14-flowered. Pistillate scales usually 
reaching the base or sometimes the middle of the beak, 2.9-4 mm long, 1.1-1.6 
mm wide, lance-ovate, concave, obtuse or the upper acute, pale brown with nar- 
row yellow-green or pale brown center and hyaline margins, 1-nerved. Staminate 
scales 2.6-5 mm long, 1.3-1.8 mm wide, ovate, obtuse or rarely acute, pale brown 
with hyaline margins, 1-nerved. Perigynia 4-5.4 (-6.1) mm long, (2.4-) 2.6-3.4 mm 
wide, (1.3-) 1.5-1.9 times as long as wide, appressed-ascending, planoconvex with 
broadly ovate, broadly elliptic, or orbicular bodies, 2.7-3.5 mm long, (0.9-) 1-1.6 
times as long as wide and 1.8-2.7 times as long as beak, widest 1.2-2 mm above 



1997 REZNICEK & ROTHROCK: CAREX MOLESTIFORMIS 301 

base, broadly thin-winged, wing 0.3-0.8 mm wide, finely serrulate-margined except 
near base, gradually and symmetrically contracted into beak, deep green to brown- 
ish (overmature), with paler margins, smooth, leathery, sessile, sharply (3-) 4-7- 
nerved adaxially over achene, 6-9-nerved abaxially over achene and 1-2-nerved 
in the winged margin; beaks 1.1-1.9 mm long, strongly flattened and serrulate- 
margined to apex, the apex bidentate with scabrous-margined teeth 0.1-0.5 mm 
long, distance from summit of achene to tip of beak 1.7-2.6 (-2.9) mm long. 
Achenes 1.6-2 mm long, 1.4-1.8 mm wide, 1-1.3 times as long as wide, biconvex, 
broadly oval to orbicular, pale brown to brown, short-stipitate at base, apiculum 
0.1-0.45 mm long; style straight; stigmas 2. Anthers 3, (1.4-) 1.6-2.6 mm long. 
Chromosome number: n = 37. 

Additional Specimens Examined. U.S.A. Arkansas. Baxter Co.: from White River on Push 
Mountain Road, about 1 mi S, then left on second gravel road at this point for 2.3 mi to Rough 
Hollow Creek, section 25, T18N R13W, 14 May 1990, Hyatt 2887.03 (MICH, UARK); from Mtn. 
Home P.O. 4 mi W on U.S. Hwy. 412 to Baxter Co. Rd. 6, then 1 mi N to 100 m past Kirby-Tucker 
Cemetery, 10 May 1993, Hyatt 5310 (MO); Mountain Home, about 7 mi N on Ark. Hwy 201, Pigeon 
Creek near Lake Norfolk, SW 1/4 of SW 1/4 section 9, T20N R13W, 15 May 1993, Hyatt 5342 (GA, 
MICH); 5345 (MICH, VSC); Gamaliel Post Office, 3.3 mi E on Bx. Co. Rd 46, then 0.3 mi E on Bx. 
Co. Rd. 43 to creek, SW 1/4 of SW 1/4 section 20, T21N R IIW, 22 May 1993, Hyatt 5485 (MICH, 
MO); Norfolk, confluence of White and Norfolk Rivers then 100 m downstream, section 20, T18N 
R12W, 7 June 1993 Hyatt 5624 (MICH, UARK); Henderson Use Area on Lake Norfolk in cove 
behind causeway upstream from boat dock area, NE 1/4 of SW 1/4 section 27, T20N R12W, 24 June 
1993, Hyatt 5708 (MICH). Boone Co.: Harrison, Harrison High School Property, Baker Prairie, 3 
May 1991, Hyatt 3848.05 (MICH, UARK); right-of-way of US 62 about 2.7 mi W of the Marion 
County line and just E of Harmon, section 17, 20 May 1994, Thomas 135746 & Amason (ctb - 
herbarium of Charles T. Bryson). Carroll Co.: SW 1/4 of NW 1/4 section 27, T18W R23W, 24 May 

1992, Hyatt 4388.08 (MICH, UARK). Cleburne Co.: Sandiff, SW 1/4 of NE 1/4 section 25, T12N 
R12W, 8 May 1993, Hyatt 5272 & Hyatt, Beer (MICH, UARK). Fulton Co.: Vidette Community, S 
of center of section 2, T20N RllW, 22 May 1993, Hyatt 5489 (MICH, MO, UARK); Salem, at Civic 
Center Lake (=about 1 mi W of the U.S. Post Office), SE 1/4 section 21, T20N R8W, 12 May 1992, 
Hyatt 4308.25 (MICH, UARK). Garland Co.: In triangle of junct. Ark. Hwy 171 & 290 and large 
stream, NE corner sect 6, T4S R18W, just E of Lake Hamilton, 17 May 1993, Reznicek 9296 & Reznicek, 
Rothrock, Bryson, Hyatt (MICH, UARK); Rothrock 2904 & Reznicek, Reznicek (MICH, chromosome 
voucher n = 37). Independence Co.: Oil Trough, Ark. Hwy 932 at White River, 29 May 1993, Hyatt 
5557 & Bishop (MICH, UARK). Izard Co.: 1.1 mi E of Moccasin Creek (and Baxter Co. line) on 
Ark. Hwy 5, 29 May 1992, Hyatt 4606.33 (MICH, UARK); along gravel rd and along Moccasin 
Creek, 0.8 mi SE of Baxter Co. line and Ark. Hwy 5, NW 1/4 section 7, T17N RllW, 2 June 1992, 
Hyatt 4612.33 (MICH, UARK); Calico Rock, at White River boat access, 3 May 1992, Hyatt 4833.33 
(MICH, UARK); Calico Rock, from railroad 6.3 mi N 
10 May 1993, Hyatt 5309 (MICH, UARK); Sylamorc 

1993, Hyatt 5550 (MICH, MO, NO); about 4.5 mi wes 
over Knob Creek, E 1/2 section 31, T17N R9W, 4 J 
Creek, about 1 mi S of Ark. Hwy 5, NE 1/4 of NW 1/4 
(MICH. NA); Cedar glade beside Ark. 9, about 11.9 mi N of White River and Sylamore SW of 
Melbourne, section 16, 20 May 1994, Thomas 139152 & Amason (ctb); along dirt rd from Lovelady's 
Boat Dock and White River up wooded valley N of Guion and Ark. 58, section 22, 21 May 1994, 
Thomas 139187 & Amason (ctb). Madison Co.: Around pond next to Ark. Hwy 295, 1.5 mi SW of 
Crosses, 20 May 1989, Stewart 89-30 (UARK). Marion Co.: Buffalo River State Park, Hills above 
Buffalo River, P.O. YellviUe, 14 May 1972, Demaree 64832 (MO); near the Buffalo River ca. 1.8 mi 
ESE of the town of Rush, SW 1/4 section 11, T17N R15W, 13 May 1977, Smith 3156 (UARK); 
Buffalo National River, SW 1/4 of SW 1/4 section 31, T18N R13W, 20 May 1993, Hyatt 5459 (MICH, 
UARK); Buffalo National River, Rush, Buffalo River banks between Rush Creek and Clabber 
Creek, SW 1/4 section 11, T17N R15W, 6 June 1993, Hyatt 5602 (MICH). Montgomery Co.: 0.3 mi W 
of Caddo Gap on Ark. Hwy 8, NE 1/4 of NE 1/4 section 13, T4S R25W, 22 April 1995, Hyatt 6291 & 
Hyatt (MICH). Newton Co.: Lost Valley, park yard area and Clark Creek, 8 June 1974, Thompson 
309 (NLU); middle area of Sneed Creek, NE 1/4 section 8, T16N R22W, 27 April 1977, Smith 3142 
(UARK). Polk Co.: 50 m N on Hwy 71 from its jet. with Polk County Rd 414, N of Hatfield, 13 May 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



lones 4653 


(MICH): 




R32W. 10 


.S mi SW 


of j una. with 


''!^Tt-,n\c 


xMICII. 
Rock Ri' 


UARK,VPI) 

ver Port/Fourc 
(MICH, chro 



: Springs pull-oil' : 
Hyaii 5526 (MICI 

;^7(MICH.M0). ' 



(BRCH, DAO, ] 



i Bridge. 
; Okla. II 

i R23r:. .N 



Reznicck. Jones. .Iu>ir^ fliRCII. HI JR. MICH (chromosome vouchern = 37). OKL), M.Ct k i xix Co.: 
Mountain Fork Campground, N side Okla. Hu-y 4 ca. I mi SE of Smilliville. section 13, TIS R2.^E. ca. 
4.6 mi W of Watson. May 20 1993. Reznkck 9357 & Rcznicck, Rothrock, Jones, Jones (BRCH, clb. 
DUR. KNK. MICH. MO. OKL. TEX, TRTE). 

The range of Carex molestiformis (Fig. 1) is confined to the Ozark Mountains 
of northern Arkansas and adjacent Oklahoma and Missouri, and the Ouachita 
Mountains in west-central Arkansas and southeastern Oklahoma. Flowering oc- 
curs from late March through April, and fruits mature from mid-May to mid-June 
depending on latitude. The epithet "molestiformis" alludes to the resemblance of 
this species to C. molesta. 

An immature collection from Arkansas: Little River Co., White Cliffs, Moore 
510127 (UARK) (not mapped) may be this species, but it is too young to identify 
with certainty. Also, the habitat (chalk chffs) and the fact that this site is on the 
coastal plain both would be unusual for C. molestiformis. More material is needed 
from this site. 



REZNICEK & ROTHROCK: CAREX MOLESTIFORMIS 




DISCUSSION 



Even before the recognition of Carex molestiformis, the immediate alUes of C. 
brevior proved particularly troublesome to systematists. Divergent taxonomic treat- 
ments abound. Mackenzie (1931) and many subsequent authors have recognized 
C. brevior and its close allies C. festucacea, C. molesta, and C. merritt-fernaldii as 
distinct species. Gleason and Cronquist (1991), however, submerged C. molesta 
and C. merritt-fernaldii into a broad concept of C. brevior, while, somewhat incon- 
sistently, recognizing C. festucacea. However, studies by Zager (1991) as well as 
our own field work, morphological studies (Rothrock 1991), examination of large 
numbers of herbarium specimens, and chromosome studies (Rothrock & Reznicek 
1996b), support Mackenzie's (1931) recognition of C brevior, C. festucacea, C. 
merritt-fernaldii, and C. molesta as distinct species. In fact, though often misidenti- 
fied, C. brevior, C. festucacea, and C. molesta are widespread and common species 
over much of eastern North America, differing consistently in morphology and 
ecology throughout their ranges without apparent intergradation. Many of the 
problems botanists have in recognizing the species within this group result from 
vague and generalized descriptions, which lack clear contrasts between species, and 
from keys that strongly emphasized single features, such as perigynium nervation, 



304 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

that are not always reliable. The following key, based on a broader range of 
features, separates all eastern North American members of the C. brevior group 
and outlines the salient diagnostic features of C. molestiformis. 

Key to the Carex brevior Group in Eastern North America 

Note: Fully mature perigynia are a prerequisite for keying. Perigynium measurements are best 
taken from perigynia between 1/3 to 2/3 the way up the spikes. Perigynia become distinctly narrower 
near the top of the spikes, and the lowermost few perigynia in the spikes are often shorter and wider 
than usual and sometimes somewhat deformed. Before choosing which lead in the key to follow, 
several perigynia should be sampled to assess variation in nervation, shape, and size. Dimensions of 
the larger perigynia and achenes should be used in keying. Inflorescence characters are best taken 
from inflorescences produced by the first flush of growth in the spring. Many species continue to 
flower sporadically through the summer, but these later inflorescences tend to be congested, with 
more numerous spikes with shorter staminate bases, and with the lowermost bracts longer and more 
leaflike. These late-season inflorescence forms are not accounted for in the key and descriptions, and 
plants collected late in the season, with all the spring i 
difficult to identify and may be impossible to key. 



riaius cuiuiuai irum snori-creepmg rnizomes; vegetative culms numerous and conspicuous, 
strongly tristichous with i.S-35 leaves when fully developed; achenes 1.6-2 times as long as 
wide (and 0.9-1 .2 mm wide); larger spikes with 5-25 (-30) perigynia. C. hyalina. 

Plants definitely clumping (though rhizomes may elongate with age); vegetative culms few, 
inconspicuous, and usually with fewer than 15 leaves, thus not strikingly tristichous; achenes 
1-1.6 times as long as wide (and 0.9-2.2 mm wide); larger spikes with ca. 15-80 perigynia. 
2. Perigynia finely granular-papillose (30-40x), the body reniform, 0.6-0.9 times as long as 

wide (and 3.2-4.9 mm wide); lower pistillate scales obtuse-rounded. C reniforniis. 

2. Perigynia smooth, the body broadly ovate, broadly elliptic, + orbicular, or rarely slightly 

obovate, (0.7-) 0.9-1.6 times as long as wide (and 1.5-6.1 mm wide); lower pistillate scales 

3. Larger perigynia 2.5-5.5 mm long, 1.5-3.6 mm wide, with beaks 0.8-1.5 (-1.9) mm long; 
perigynia plumply planoconvex or concavo-convex, the bulge formed by the achene 



4. Leaf sheaths finely papillose at high magnification (30-40x); perigynia membrana- 
ceous and translucent, the dark achene ± visible through the adaxial face of the 
perigynium; usually at least some perigynia with the wings and base of beak ± erose, 
scalloped, or even with an irregular tooth and not symmetrically tapered. 
5. Perigynia distinctly 4-7-nerved over the achene on the adaxial face, (4.5-) 5-5.5 
mm long, wings usually strongly coppery-tinged; pistillate scales usually reddish 
brown; anthers (2.5-) 2.9-4.1 mm long. C. bicknellii var. bicknelUi. 

5. Perigynia nerveless or faintly and irregularly 1-5-nerved over the achene on the 
adaxial face, 3.6-4,8 (-5.2) mm long; wings and pistillate scales yellowish tinged; 
anthers 1.3-2.6 mm long. C. merritt-fernaldii. 

4. Leaf sheaths smooth; perigynia leathery, opaque, with the wings and base of beak 
usually finely and evenly serrulate and symmetrically tapered. 

6. Spikes on larger culms (4-) 5-7 (-10), tapered at the base, the terminal one with 
a conspicuous staminate base; inflorescences typically 2.5-4.5 (-6.5) cm long with 
the lowest inlernodes (3-) 5-13 (-23) mm long; perigynium body (0.7-) 0.9-1.2 
(-1.3) limes as long as wide. 

7. Larger achenes 1.4-1.8 mm wide, (1.6-) 1.7-2 mm long; larger perigynia 3.2- 
4.7 ( 5) mm long. 2..5-3.3 (-3.5) mm wide, nerveless or occasionally faintly 1-5- 
iKTxcclvcntially. C brevior. 



Spikes on larger culms 2-4 (-5) (rarely more), rounded at the base, the te 
one lacking a conspicuous staminate base; inflorescences 1.2-2.7 (-3.6) ci 
with the lowest internodes 1.5-6 (-9) mm long; perigynium body (0.9- 



1 997 REZNICEK & ROTHROCK: CAREX MOLESTIFORMIS 305 

8. Achenes of larger perigynia 0.9-1.3 mm wide, elliptic to narrowly oblong, 1.3- 
1.6 times as long as wide, larger perigynia 1.8-2.8 (-3) mm wide, squarrose- 
spreading at maturity, (25-) 30-80 per spike; widespread. C. molesta. 

8. Achenes of larger perigynia 1.4-1.8 mm wide, broadly oval to ± orbicular, 1- 
1.3 times as long as wide, larger perigynia 2.6-3.4 mm wide, appressed 
ascending at maturity, (10-)15-40 per spike; Ozark Mountain region. 

C. molestiformis. 
3. Larger perigynia 5.5-8 (-8.7) mm long, (3.1-) 3.5-6.1 mm wide (except rarely in C. 
bicknellii var. bicknellii with papillose sheaths), with beaks (1.4-) 1.6-2.5 (-3.4) mm 
long; perigynia thin and ± waferlike, ± biconvex around the achene, the bulge often 
prominent on both faces of the perigynia. 

9. Larger perigynia (2.8-) 3.2-4.8 mm wide, (2-) 4-7-nerved over achene ventrally 

(occasionally nerveless); staminate and pistillate scales obtuse to long-acuminate, 

but the midrib not excurrent as a scabrous awn; larger culms with (3-) 5-7 (-9) spikes. 

10. Leaf sheaths finely papillose, at least near the apex; perigynia thin, translucent, 

usually with coppery-tinged wings, the larger (2.8-) 3.2-4.3 mm wide; pistillate 

scales usually reddish brown; anthers 2.9^.1 mm long; plants in small clumps 

(usually <10 culms) in dry to mesic habitats. C. bicknellii var. bicknellii. 

10. Leaf sheaths smooth; perigynia ± thickened and opaque, with greenish or pale 

brown wings, the larger (3.1-) 3.5-4.8 mm wide; pistillate scales pale yellowish 

brown; anthers 1.8-3.5 mm long; plants in dense, large clumps (up to 200 culms) 

9. Larger perigynia 4-6.1 mm wide, nerveless over achene ventrally, or nearly so; 
staminate scales (and sometimes the lowermost pistillate scales) with the midrib 
excurrent as a scabrous awn 0.1-0.9 (-2.4) mm long; larger culms with (2-) 3^ (-5) 

Collections of Carex molestiformis were rare in herbaria before our recogni- 
tion of it as a distinct entity, although the earliest specimen we have seen was 
collected in 1880. Specimens in herbaria were usually misidentified either as C. 
bicknellii because of their large perigynia with prominent adaxial nerves or as C. 
molesta because of their few-spiked, congested inflorescences. Carex bicknellii, 
however, is a quite different plant; much larger, with longer, wider, and thinner 
perigynia and more spikes per inflorescence. The real affinities of C. molestiformis 
are with C. brevior and its close allies C festucacea and C. molesta. A morphologi- 
cal comparison of these three species with C. molestiformis is presented in Table 1. 
Carex molestiformis is easily distinguished from C festucacea by its larger perigy- 
nia and achenes, and by its short, congested, few-spiked inflorescences. Carex 
molestiformis shares with C. molesta perigynia that are usually distinctly longer 
than wide, several-nerved over the achene adaxially, and few-spiked, short, con- 
gested inflorescences, with the spikes lacking a conspicuous staminate base (though 
the staminate base in C. molestiformis is often longer than that of C. molesta). 
Carex molestiformis is similar to C. brevior in its large and especially wide achenes, 
large anthers, and relatively few appressed-ascending perigynia per spike; however, 
C molestiformis is in no way intermediate between C brevior and C molesta. 
Rather, it shares some features found in C. brevior and some found in C. molesta 
but also has unique characters. Among its close relatives, it has the largest perigy- 
nia but also, on average, the fewest spikes. The adaxial nervation of C. molestifor- 
mis is more prominent than in any of its close relatives, and the perigynia tend to 
be a deeper green color at full maturity. Robust culms of C. molestiformis also 
tend to be quite stout (above the uppermost leaf sheaths) though rather soft; 
occasionally as much as 3 mm wide when flattened by pressing, a feature not seen 
in any other species in the C brevior group. The perigynium and achene features 
noted for the four species treated in Table 1 are illustrated in Fig. 2. 

Additional strong evidence for the distinctness of Carex molestiformis from both 
C. brevior and C. molesta comes from chromosome numbers. The chn 



^ MICHIGAN HERBARIUM 










FIG. 2. Perigynia and achenes (a 


bove and to the right) ■ 


MICH); 2) C fesmcacea (from Rczn 


icck 9778. MICH); 3) 


MICH): and 4) C molesta (from Wils< 


.« 4^W6. MICH). Scale ^ 



count of C. molestiformis was determined as n = 37 in plants from Garland Co., 
Arkansas {Rothrock 2904, reported as "C. cf. breviof by Rothrock & Reznicek, 
1996b); Pulaski Co., Arkansas (Rothrock 2901); and Mays Co., Oklahoma (Reznicek 
9766). In contrast, the chromosome count for C. molesta is n = 34 (Wahl, 1940; 
Love & Love, 1981; unpublished data) or n = 35 (unpublished data). Carex brevior 
also has fewer chromosomes than C. molestiformis, with n = 34 (Love & Love 
1981) or lower (unpublished data). 

Carex molestiformis thrives in a wide variety of open to somewhat shaded, 
mostly ruderal, habitats with a wide range of associates, including ditches, dryish 
roadsides, river and stream banks, moist meadows, and open forests. Soils ranged 
from clays to loamy silts and sometimes coarser, sandy and gravelly soils along 
streams. Typically, colonies of C. molestiformis rarely occur far from streams, and 
the most undisturbed habitats in which the species occurs are openings in forested 
bottoms of rivers and streams with alluvial soils. This may have been the original 
natural habitat of the species, but it has certainly found disturbed sites suitable for 
colonization. In these sites, the species can occur with C. festucacea, C. brevior. 
and, in northern Arkansas, occasionally C. molesta. No intermediates were dis- 
covered at any sites where two or more species occurred. Although C. molestifor- 
mis can co-occur with any of the other three species to which it is most similar, all 
do have certain distinctive ecological preferences. Carex molesta is also commonly 
a ruderal like C molestiformis, and also often occurs near or along streams and in 
open woodland. However, it often prefers heavy, clayey soils and is definitely more of 
a calciphile. Carex brevior tends to occur in drier sites, often on sandier soils, and 
rarely in even lightly shaded settings. Carex festucacea tends to occur in at least 
moist, and sometimes quite wet soils, but also rarely in deeply shaded conditions. 

Like Carex ozarkana, another recently described species of section Ovales 
from the Ozark Mountain region (Rothrock & Reznicek 1996a), C. molestiformis 
was probably much rarer in the past, before large scale human alteration of the 
landscape generated extensive ruderal habitats. 

ACKNOWLEDGMENTS 

We are very grateful to Philip Hyatt, who accompanied us in the field, and quickly learned to 
our manuscript. Stanley and Gretchen Jones accompanied us in the 



REZNICEK & ROTHROCK: CAREX MOLESTIFORMIS 

Morphological comparison of Carex brevior, C. festucacea, C. mo, 
nts in parentheses in this table were mostly no 
the larger inflorescences, perigynia, and achenes on 
irements for perigynium length in C. molestiformis ai 
specimens and are not accounted for in the key. 



usually + open, 
or even nodding 



308 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

field during two springs and helped considerably in finding stations. We thank the curators of a 
number of herbaria, including ctb, MICH, MO, NLU, OKL, UAM, UARK, and WARM, for lending 
us material and allowing us to scour their holdings for specimens of C molestiformis. We are grateful 
to Susan Reznicek for drawing Figure 2, and to Sarah Cleveland and George Yatskievych for their 
help with the distribution map. 



LITERATURE CITED 

Gleason, H. A., and A. Cronquist. 1991. Manual of vascular plants of Northeastern United States 

adjacent Canada. 2d ed. New York: New York Botanical Garden. 
Love, A., and D. Love, 1981. Chromosome number reports LXXIII. Taxon 30: 829-861. 
Mackenzie, K. K. 1931. Cyperaceae - Cariceae. N. Amer. Flora 18(1-3): 1-168. 
Rothrock, P. E. 1991. The identity of Carex albolutescens, C. festucacea, and C. longii (Cyperaci 

Rhodora 93: 51-66. 
Rothrock, P. E., and A. A. Reznicek. 1996a. A new species of Carex section Ovales (Cyperac 

occurring in the Ozark Mountain Region. Brittonia 48: 104-110. 
. 1996b. Documented chromosome numbers 1996: 1. Chromosome numbers in Carex se( 

Ovales (Cyperaceae) from eastern North America. Sida 17: 251-258. 
Wahl, H. A. 1940. Chromosome numbers and meiosis in the genus Carex. Amer. J. Bot. 27: 458-4 
Zager, S. C. 1991. Multivariate analysis of the Carex brevior group in Iowa. M.A. Thesis, Unive 



Contr. Univ. Michigan Hej 



NOMENCLATURAL CHANGES IN MEXICAN 
DIOSCOREACEAE AND LEGUMINOSAE 

Oswaldo Tellez V. 

Herbario Nacional 

Instituto de Biologia UNAM 

Apartado Postal 70-367 
04510 Mexico, D.F., Mexico 

Recent work on the Mexican and Central American species of Dioscorea 
(Dioscoreaceae; Tellez & Schubert 1994) and Tephrosia (Leguminosae) revealed 
the necessity for some nomenclatural changes. These are presented here, along 
with discussions and citations of selected specimens. 

DIOSCOREACEAE 

Dioscorea gomez-pompae O. Tellez, nom. et stat. nov. Dioscorea spiculiflora var. 
chiapasana Gomez-Pompa, Ciencia (Mexico) 18(11-12): 242. 1959.— Type: 
Mexico. Chiapas: [Jardin Botanico de Chiapas] material procedente de la 
bajada de Suchiapa, selva baja decidua, 11 Nov 1953, Miranda 7914 (holo- 
type: MEXU!; isotype: MEXU!). 



Acala, 6 Nov 1980, Breedlove 47156, 47157 (MEXU); El Chorreadero, sobre el Km 25 de la carretera 
Tuxtia Gutierrez-San Cristobal de las Casas, 26 Nov 1982, Cabrera & de Cabrera 3862 (MEXU); 
Rancho La Esperanza, Tuxtia Gutierrez, Gomez P. 71 (MEXU); Puente Belisario Dominguez, Rio 
Grijalva, al E de Tuxtia Gutierrez, 11 Sep 1958, Gomez P. 137 (MEXU); bajada Suchiapa, 11 Nov 
1953, Lopez E. 7915 (MEXU); Nuevo Orizaba a 28 km al W del vertice del Rio Chixoy, camino a 
Chajul, 8 Nov 1985, Martinez 15259 (MEXU); Mpio. Ocozocuautla, El Yeso, a 15 km al S de Cintala- 
pa, 11 Dec 1987, Martinez & Reyes G. 22000 (MEXU); Escuintla, 5 Feb 1936, Matuda 875 (MEXU); 
Escuintla, 11 Dec 1937, Matuda 2137 (MEXU); Cacaluta, Escuintla, 12 Oct 1947, Matuda 17061 
(MEXU); Mpio. Tonala, Sta. Rosa, 17-18 Dec 1947, Matuda 17360 (MEXU); Villa Flores, Rancho 
Margarita, 2 Jul 1950, Miranda 6425 (MEXU); Barranca Pishtinbek (N Tuxtia G.), 3 Dec 1950, Miranda 
6729 (MEXU); Mpio. Huixtla, Huixtla, 3 Sep 1952, Miranda 7634 (MEXU); Cerro Vernal, 20 km al E 
de Tonala, 20 Nov 1984, Tellez et al. 8142 (MEXU). 

This taxon was described by Gomez-Pompa (1974) as D. spiculiflora var. 
chiapasana in sect. Apodostemon Uline of subg. Dioscorea. It is characterized by its 
sinistrorsely twining stems and its leaves. The leaf blades are suborbicular to 
orbicular and basally auriculate (occasionally the lobes overlapping) with the sinus 
campanulate in outline; the petioles are basally winged. It occurs in the coastal 
regions of Chiapas and the Pacific slope of Guatemala. Dioscorea spiculiflora 
Hemsl. ["var. spiculiflora"] differs in its dextrorsely twining stems, triangular leaf 
blades basally auriculate (the lobes never overlapping) with the sinus subquadrate 
to subrectangular in outline, and unwinged petioles. Its range extends, mainly 
along the Atlantic slope, from San Luis Potosi, Mexico, to Costa Rica. 

The differences in morphology and distribution warrant the recognition of 
"var. chiapasana"" at the species level; however, the name D. chiapasensis is already 



310 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

occupied (Matuda 1953). The new name Dioscorea gomez-pompae is here pro- 
posed and honors Arturo Gomez-Pompa, an expert in the study of the Mexican 
Dioscoreaceae. 

Dioscorea fasciculocongesta (Sosa & B. G. Schubert) O. Tellez, stat. nov. Dioscorea 
spiculiflora var. fasciculocongesta Sosa & B. G. Schubert, Biotica 11(3): 
187. 1986.— Type: Mexico. Veracruz: Mpio. Juchique de Ferrer, el Cerro 
de Villa Rica cerca de Mundo Nuevo, 1500 m, 19°48'N, 96°46'W, 7 May 
1981, Castillo, Cortes & Becerra 1836 (holotype: XAL!). 

Representative Specimens. Puebla: Villa de Juarez, 19 Apr 1959, Gomez P. 168 (MEXU); 5 km 
al sur de Villa Juarez, 17 Apr 1958, Gomez P. 85 (MEXU), Tenorio 8770 (MEXU).— Veracruz: 
Ventura 1135, 3503, 12668 (ENCB), Sosa et al. 163 (XAL). 

This taxon was described as a variety of D. spiculiflora, but is there misplaced 
and here elevated to the level of species. It is more closely allied with D. mexicana 
Scheidw. with which it shares ovate, chartaceous to membranous leaves with con- 
spicuous and prominent midveins, oblong capsules, and pollen of similar size. 
Dioscorea fasciculocongesta has hypogeous and smooth rhizomes; the staminate 
inflorescences bear the flowers arranged in congested fascicles that are conspicu- 
ously separated on the axis and have ovate, long-acuminate bracts. It is endemic 
to a montane forest located along the border of Puebla and Veracruz. In contrast, 
the rhizomes of D. mexicana are epigeous and have the surface broken into flat 
prismatic plates, resembling the shell of some turtles. The staminate inflorescences 
have the flowers scattered along the rachis and have short, inconspicuous bracts. 
It grows mainly in tropical forests from southern Veracruz and Oaxaca, Mexico, 
to Panama. 

Dioscorea galeottiana Kunth, Enum. pi. 5: 409. 1850. Dioscorea convolvulacea 
var. galeottiana (Kunth) Uline, Bot. Jahrb. Syst. 22: 427. 1897. Dioscorea 
grandiflora M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 9(2): 
392. 1842, non Dioscorea grandiflora C. Martins ex Griseb., 1842.— Type: 
Mexico. Oaxaca: Costa del Pacifico, Yolotepequez, 7000 ft, Sep-Nov 1840, 
Galeotti 5457 (holotype: BR, photo: MEXU!). 

Dioscorea lobata Uline, Bot. Jahrb. Syst. 22: 427. 1897— Type: Mexico. [Dis- 
trito Federal:] in vallibus prope urbem, Schmitz s.n. (holotype: B!). 

Dioscorea lobata var. lasiophylla Uline ex Knuth, Notizbl. Bot. Gart. Berlin- 
Dahlem 7: 194. 1917.— Type: Mexico. Pavon s.n. (holotype: B!). 

Dioscorea convolvulacea var. viridis Uline, Bot. Jahrb. Syst. 22: 427. 1896. — 
Type: Mexico. Oaxaca: Sierra de San Felipe, 2500 m, Pringle 4890 (holo- 
type: B!). 



Pedrcgal de San Angol. 10 Sep 195{). Maiiuh, I95.U (MEXU); (Y-rro Zacayuca. 1 1 Sep 1980. 
332 (MEXU); Ejidos de Padicrna. .-^^O Jul 1980, Pami M. 536 (MEXU); Pedrcgal. Tizapan 
1896, Pringle s.n. (MEXU); Pedregal de San Angel, cerca de los Reyes. 28 Sep 1952, Rzedow 



1997 TELLEZ: NOMENCLATURAL CHANGES 311 

1849 (MEXU); Delegacion de Xochimiico, desviacion del Conejo, 8 Aug 1976, Ventura A. 1839 
(MEXU); Delegacion de Milpa Alta, San Pedro Atoxpan, 4 Sep 1977, Ventura A. 3027 (MEXU); 
Pedregal de San Angel, Ciudad Universitaria, D.F., frente a la Facultad de Ciencias, U.N.A.M., 7 
Aug 1986, Villasehor et al. 942 (MEXU).— Mexico: Mpio. Huixquilucan, 2 km al SW de San Bartolito, 
18 Sep 1982, Galvdn R. 1276 (MEXU); Mpio. Texcoco, Bancs de Netzahualcoyotl (Cerro Tetzcotcingo), 
7 km al E de Texcoco, 3 Aug 1976, Koch & Garcia P. 76145 (MEXU); Mpio. Amecameca, en loma de 
Panteon, Amecameca, 20 Sep 1950, Matuda 19322 (MEXU); Cerro del Tigre, al NW de Atizapan, 4 
Aug 1974, Rzedowski 31993 (MEXU); Mpio. Huehuetoca, vertiente W del Cerro Sincoque, 3 Aug 
1976, Rzedowski 34347 (MEXU); lomas de Atizapan y Chiluca, 7 Sep 1980, Ventura A. 3666 (MEXU).— 
Guerrero: Mpio. Chilpancingo, 3 km al SW de Omiltemi, 4 Dec 1966, Rzedowski 23599 (MEXU).— 
Michoacan: Mpio. Erongaricuaro, 5 km al W de Erongaricuaro, 22 Aug 1993, Chdzaro B. et al. 7201 
(MICH, MEXU); Mpio. Erongaricuaro, 0.5 km al W de la hacienda de Charahuen, 27 Oct 1986, 
Espinosa G. 2314 (MEXU); Mpio. Patzcuaro, 2.5 km al SE de Ajuno, carr. Patzcuaro-Uruapan, 3 
Nov 1986, Espinosa G. 2405 (MEXU); 4 km al S de Indaparapeo, sobre el camino a Las Peras, 27 Sep 
1989, Rzedowski 49046 (MEXU); en la desviacion a Santiago Undameo, 16 km al SW de Morelia, 
carr. a Tiripetio, 9 Sep 1979, Soto N. 991 (MEXU).— Morelos: autopista Mexico-Cuernavaca, Km 
53.5, 9 Oct 1960, Espinosa J. 351 (MEXU); Mpio. Tepoztlan, 2-3 km al N de Tepoztlan, sobre la 
caiiada de la zona arqueologica del Tepozteco, 5 Oct 1986, Flores F. & Cabrera 149 (MEXU); Mpio. 
Tepoztlan, Tepozteco, 22 Sep 1938, Lyonnet 2555 (MEXU); Mpio. Cuernavaca, Salto de San Anto- 
nio, 5 Aug 1951, Matuda 21629 (MEXU); Mpio. Tepoztlan, Tepoztlan, 5 Sep 1953, Matuda 30143 
(MEXU); Mpio. Tepoztlan, Tepoztlan, 5 Oct 1953, Matuda 30151 (MEXU).— Oaxaca: Dtto. San 
Juan Mixtepec, Independencia, a 8 km al E de San Juan Mixtepec, 17 Nov 1988, Reyes J. 1329 
(MEXU); Dtto. San Juan Mixtepec, Yuu Tzaan (Caiiada de Cazuelas), a 9 km al S de San Juan 
Mixtepec, 3 Sep 1989, Reyes J. 1931 (MEXU).— Queretaro: El Batan, 27 May 1982, Argiielles E. 
7794 (MEXU). 

Review of the types of Dioscorea lobata and D. lobata var. lasiophylla, and 
comparison with other collections showed that the two names are synonyms of the 
older name D. galeottiana. The type material for both is fragmentary. The type of 
D. lobata consists of a leaf and two staminate inflorescences collected by Schmitz 
near Mexico City, and that of D. lobata var. lasiophylla of only one leaf from an 
unknown locality (the collection attributed to Pavon). In both the leaf is lobate 
and densely pubescent, and the flowers of D. lobata have a purple corolla and 
three stamens. These characters are shared with D. galeottiana, the only species of 
Dioscorea occurring in the Valley of Mexico. 



LEGUMINOSAE 

Tephrosia crassifolia Benth., Bot. Voy. Sulph. 80. 1844. — Type: Mexico. Guerrero: 
Acapulco, 1842, Hinds s.n. (holotype: K!). 
Tephrosia smythiae McVaugh, Flora novo-galiciana 5: 726. 1987.— Type: Mex- 
ico. Jahsco: Mpio. Cabo Corrientes, rocky stream valley, ca. 5 km N of El 
Tuito, McVaugh 25487 (holotype: MICH!; isotype: MEXU!). 

Representative Specimens. Guerrero: PotreriUos del Rincon, camino San Luis Acatlan-Ilia- 
tenco, 5 Feb 1986, Aguilar J. 1414 (MEXU).— Jalisco: Mpio. Cabo Corrientes, 6 km al O de El 
Tuito, camino a Chacala, cerca de Los Guacimas, 5 Feb 1979, Solis M. et al. 1526, 1532 (MEXU); 
Mpio. Tuito, 5 km al N de El Tuito, carr. Barra de Navidad-Puerto Vallarta, 6 Feb 1991, Tellez V. & 
Novelo 12923 (MEXU).— Michoacan: La Mada, Feb 1899, Langlasse 851, 851 bis (F).— Nayarit: 
Mpio. Compostela, along hwy 200 between Tepic and Puerto Vallarta, 33 mi S of Tepic, 4 mi N of Las 
Varas, 9 Jan 1979, Croat 45369 (MEXU); Mpio. Acaponeta, Tiger Mine, 1 Mar 1927, Jones & Jones 
23022 (MEXU); Mpio. Compostela, cerca de Las Varas, Km 60 de la carr., 27 Apr 1974, Vargas & 
Ochoa A. & 1335 (MEXU); Mpio. Compostela, Km 60 de la carr. Compostela-Las Varas, 22 Jan 
1994, Calzada J. et al. 19122, 19125 (MEXU); Mpio. San Bias, Km 3-5 sobre el camino a Pintadeno, que 
sale en el Km 16 de la carr. Tepic-Miramar, 12 Sep 1985, Tellez V. 9294 (MEXU); Mpio. Tepic, 3 km 



312 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

al E de Cora, terraceria al Cuarenteno, 29 Mar 1987, Tellez V. 10050 (MEXU); Mpio. Tepic, Km 20- 
30 camino de terraceria a el Cuarenteno-Cora-Jalcocotan, en la serrania de San Juan, 14 Jan 1988, 
Tellez V. II 134 (MEXU); Mpio. Compostela, Km 60 carr. Tepic-Pto. Vallarta Jal., 15 Feb 1980, 
Zdrate S. 469 (MEXU).— Oaxaca: Dtto. Juchitan, Mpio. Sta. Maria Chimalapa, ca. 13 km al E de 
Sta. Maria, file de lava en la vereda a Chimalapilla y Monte Rico, al NE del Rio del Corte, 27 Oct 
1984, Hernandez G. 529 (CHAPA, MEXU); Dtto. Pochutla, Pluma Hidalgo, San Rafael Toltepec, a 
17 km al N de Pochutla, 14 Feb 1976, Soiisa S. el al. 5347 (MEXU); Dtto. Putla de Guerrero, 15 km al 
SE de Putla, 19 Aug 1976, Sousa S. et al. 5858 (MEXU); Dtto. Pochutla, Mpio. Pluma Hidalgo, 
Toltepec, a 2 km de la carr. Pochulla-Miahuatlan, en la carr. a Pluma Hidalgo, 23 Oct 1976, Sousa S. 
et al. 6470 (MEXU); Dtto. Putla de Guerrero, Santa Cruz del castillo, a 13 km al S de Putla, 4 Feb 
1977, Sousa S. el al. 7031 (MEXU, MO); Dtto. Pochutla, a 6 km al NE de Chacalapa, en la desv. a 
Finca Pilas, 7 Feb 1977, Sousa S. et al. 7121 (MEXU, MO); Dtto. Putla de Guerrero, a 14 km al SE de 
Putla de Guerrero, 28 Jun 1977, Sousa S. et al. 7683 (MEXU); Dtto. Putla de Guerrero, Santa Cruz 
del Castillo, a 13 km al S de Putla de Guerrero, 6 Decl978, Sousa S. & Zdrate 9897 (MEXU, MO). 

When McVaugh (1987) described Tephrosia smythiae, he stated that this spe- 
cies is "very like Tephrosia crassifolia in most details, but the leaflets tending to 
be ovate rather than elliptic-obovate, and acute rather than obtuse or rounded at 
apex." Examination of collections of T. crassifolia from throughout its range, 
Sinaloa to Chiapas, showed that these differences are included in the variation of 
the foliar morphology. Consequently, T. smythiae can no longer be maintained. 

Tephrosia leucantha Kunth in H.B.K., Nov. gen. sp. 6[foHo]: 360. 12 Jul 1824; 
6[quarto]: 460. Sep 1824.— Type: Mexico. Guanajuato: cerca de Guana- 
juato, ca. 2000 m, Sep 1803, Humboldt & Bonpland s.n. (holotype: P, 
microfiche IDC 800-12: MEXU!). 
Tephrosia feddemana McVaugh, Flora novo-galiciana 15: 712. 1987.— Type: 
Mexico. Michoacan: Cerro Santa Maria, 8-10 km SW of Jiquilpan, and 
ca. 5 km NE of Quitupan, Jal, shrubby vegetation in oak forest near 
summits, ca. 2000 m, 8-9 Aug 1959, Feddema 227 (holotype: MICH!; 
isotype: MEXU!). 

Representative Specimens. Chihuahua: Sierra Madre Mts, Guayanopa Canyon, 24 Sep 1903, 
Jones & Jones s.n. (MO); near Cusihuiriachic, 28 Sep 1888, Pringle 2006 (MO); Sierra Madre near 
Colonia Garcia, 11 Oct 1899, Townsend & Barber 367 (MO).— Durango: 21-27 Aug 1906, E. Palmer 
478 (MO).— Nayarit: Mpio. Nayar, Sierra de Las Palomas (Sierra de Alica), 9 Oct 1963, Schubert & 
Sousa 2041 (MEXU); Mpio. La Yesca, 22.3 km al W de Huajimic, brecha a Tepic, 27 Oct 1989, 
Tenorio L. & Flares F. 16746 (MEXU).— Querftaro: Del Ciervo al cerro de la Mesa, 20 Aug 1905, 
Altamirano F 1564 (MEXU).— Sonora: Alamos, 3 km al S de La Lobera, Ejido Zahuarivo, 25 Aug 
1986, Lezama T. & Frame 11925 (MEXU).-Zacatecas: south slope of La Bufa, 10 Aug 1948, 
Dressier 1 13 {UO). 

When McVaugh (1987) decribed Tephrosia feddemana he did not discuss how 
it differs from other species; however, in his key to Leguminosae in the same 
work, he differentiates it from T. leucantha by type of pubescence, bracts length, 
and the degree of adnation of the vexillar stamen. I beheve only the last is note- 
worthy; the stamen is adnate to the staminal tube in T. leucantha but not in T. 
feddemana. Yet, the adnation of the stamen to the staminal tube is not a constant 
and rehable character. I have observed many times that during the development 
of the fertilized gynoecium the vexillar stamen tends to separate from the stami- 
nal tube; thus, the degree of adnation seen may depend on the stage of gynoecial 
development. I believe this phenomenon deserves further investigation before its 
taxonomic significance can be evaluated. Unfortunately, for T. feddemana only 



TELLEZ: NOMENCLATURAL CHANGES 313 

available for study. Because in all other features the type of 
; with the morphological variation shown by T. leucantha 
it is here included in T. leucantha. 



ACKNOWLEDGMENTS 

I thank Patricia Davila Aranda for reviewing the manuscript and translating it into English, and 
for her useful comments. I am grateful to Harriet Gillett (WCMC), the World Conservation Monitor- 
ing Centre (United Kingdom), and British Airways for financial support for visits to herbaria in 
Germany and Switzerland. The National Geographic Society provided financial support for fieldwork 
(grant 4349-89), and W. Berendsohn (B) gave assistance with logistics. I thank the curators of the 
following herbaria for permitting me to study their collections: B, BM, ENCB, F, GH, K. MICH, 
MO, NY, US, and Z. 



LITERATURE CITED 

eva variedad de Dioscorea spiculi flora Hemsl. Ciencia (Me) 

as de Mexico. Bol. Soc. Bot. Mexico 15: 23-25. 
In Flora novo-galiciana by R. McVaugh (general editor W 
Anderson), vol. 5. Ann Arbor: The University of Michigan Press. 
Tellez v., O., and B. G. Schubert. 1994. Dioscoreaceae. In Flora mesoamericana, ed. G. Davidse 
Sousa, and A. Chater, 6: 54-65. Mexico City: Universidad Nacional Autonoma de Mexico 
Louis: Missouri Botanical Garden; London: The Natural History Museum. 



A NEW SPECIES OF APPENDICULA (ORCHID ACEAE) 
FROM MT. MURUD, SARAWAK, BORNEO 

Jeffrey J. Wood 

Royal Botanic Gardens 

Kew, Richmond, Surrey TW9 3AE, England 

The genus Appendicula, although unlikely to be familiar to most growers, is 
sometimes encountered in a few specialist collections and in botanical gardens. 
About sixty species are distributed from tropical Asia to the Pacific islands, with 
the majority occurring in Indonesia, the Philippines, and New Guinea. Most inhabit 
lowland forest or mid-elevation forest in the mountains, particularly favoring shady 
areas with high humidity. The leaves are arranged in two rows along the entire 
length of the stem. These are often twisted at the base, so that the blade is lying in 
one plane and at right angles to the low intensity light source often available. The 
flowers are very small and usually of a whitish or greenish hue, although often 
flushed with pink or purple. In one Bornean species, A. torta Blume, these are 
subtended by colorful overlapping lilac-pink floral bracts. The lip always has a 
variably shaped, often appendage-like callus situated on the upper surface just 
above the base. Appendicula is distinguished from the closely related Podochilus 
by having six instead of four pollinia. 

Appendicula bilobulata, described below, is known only from the type material, 
which was collected in 1967 by Bill Burtt, of the Royal Botanic Garden, Edin- 
burgh, on Mt. Murud (2,438 m), the highest mountain in the Malaysian state of 
Sarawak. The general appearance is that of a Podochilus, which I thought it to be 
on first sight. Examination of the flowers, however, showed it to be a hitherto 
undescribed species of Appendicula related to A. fractiflexa J. J. Wood, also recently 
described from Borneo, and A. undulata Blume, which occurs in Peninsular Malaysia, 
Sumatra, Java, and Borneo. Appendicula bilobulata is distinguished from both by 
the distinctive bilobed lip. 

Appendicula bilobulata J. J. Wood, sp. nov.— Type: Malaysia. Sarawak: route 
from Ba Kelalan to Mt. Murud, near Camp 111, ca. 1740 m, 28 Sep 1967, 
Burtt & Martin B. 5287 (holotype: E! herbarium material only; isotypes: 
K! spirit material only, SAR! herbarium material only). Fig. 1. 

Appendiculae fractiflexae J. J. Wood atque A. undulatae Blume affinis, sed 
epichilo labelli bilobulato, hypochilo appendiculo laevi upsiloide basali instructo 
distinguitur. 

Trailing, mat-forming epiphyte. Roots filiform, wiry, elongate, simple, very 
minutely papillose, or hirsute, produced at intervals along stem. Stem 12-25 cm 
long, very slender, branching distally, internodes 2-3 mm long, enclosed in persis- 
tent leaf sheaths. Leaves 0.7-1 x 0.2-0.3 cm, narrowly oblong-elliptic, minutely 
obhquely retuse, mucronate, thin-textured, sheaths 2-3 mm long. Inflorescences 
lateral and/or terminal, one flower open successively; peduncle ca. 5 mm long, 
enclosed by 2 or 3 tubular, acute to acuminate, non-floriferous bracts; rachis 0.5- 
ca. 1.2 cm long, fractiflex; floral bracts 2.5-4 mm long, lanceolate, narrowly acumi- 
nate. Flowers white, tip of labellum pale purple. Pedicel with ovary 4.8-5 mm 

315 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 




1997 WOOD: APPENDICULA BILOBULATA 317 

long, narrowly clavate, gently curving. Sepals 3-nerved. Dorsal sepal 3 x 1.6-1.7 
mm, ovate, concave, acuminate, cuspidate. Lateral sepals 6 x 1.8-2 mm, obliquely 
oblong, acuminate, cuspidate. Mentum 5 mm long, oblong, obtuse. Petals 2.4-2.5 
X 0.9 mm, narrowly oblong, subacute, 1-nerved. Labellum hypochile: 5.8-6 mm 
long, tubelike, with erect sides, margins fleshy and sulcate, especially distally; 
provided with a smooth, fleshy, U-shaped basal appendage; labellum epichile 3 
mm long, 4.5-5 mm wide across lobules, flabellate, bilobulate, lobules each ca. 3 x 
2.5 mm, broadly oblong, rounded, margin minutely irregular. Column 0.5-0.6 mm 
long; foot 5 x ca. 0.7-0.8 mm; wings 0.4-0.5 mm long, oblong, obtuse, fleshy; apex 
truncate; rostellum acute, toothlike; anther-cap 0.8-0.9 x 0.9-1 mm, ovate, cucul- 
late, acute, smooth; pollinia 6, obliquely clavate. 



LITERATURE CITED 

Seidenfaden, G. 1986. Orchid Genera in Thailand XIII. Thirty-three epidendroid Genera. Operj 

Botanica 89: 135-142. 
Wood, J. J., R. S. Beaman, and J. H. Beaman. 1993. The Plants of Mount Kinabalu 2. Orchids, p. 89 



. Michigan Herb 



TAXONOMIC AND NOMENCLATURAL NOTES ON THE 
DELESSERIACEAE (RHODOPHYTA) 



Michael J. Wynne 
mi of Biology and Herbari 
University of Michigan 
I Arbor, Michigan 48109 



Gigartina fabriciana Lyngb. has had a complicated history. The species was 
described by Lyngbye (1819, pi. IID) on the basis of a collection from Greenland 
by the Reverend [Bishop] O. Fabricius. Ruprecht (1851) indicated that the collec- 
tion was made in the period 1768-1773 from the vicinity of the colony at Frederiks- 
haab (6r59'N, 49°42'W). Although Lyngbye's descriptions and illustrations of the 
algae were meticulously executed and the work was generally well received, C. 
Agardh of Lund, the major Scandinavian authority on algae of that time, was 
critical of Lyngbye's work, which resulted in some friction between Danish and 
Swedish phycologists (Hansen 1985). Agardh (1822, 1824) treated Gigartina fabri- 
ciana at the varietal level within his Sphaerococcus cristatus. Subsequently, Agardh's 
son, J. Agardh (1841), initially transferred G. fabriciana to Rhodymenia [as Rhodo- 
menia fabriciana] and later (1847) transferred it to Euthora [E. fabriciana], where 
it was considered a close relative of E. cristata (C. Agardh) J. Agardh. Kiitzing 
(1849) transferred Lyngbye's species to Callophyllis with a query.^ Later, J. Agardh 
(1852), after having examined a fragment of the type, realized that G. fabriciana 
did not belong in any of the genera to which he and his father had assigned it. He 
stated (p. 698) that G. fabriciana was without any doubt a member of the "Deles- 
seriae," compared it to Delesseria rostrata (Lyngb.) J. Agardh, and referred to the 
costal cells as "elongate." 

Over the years workers have struggled with the question of how to differenti- 
ate the following four species which had been placed in the genus Delesseria s.L: 
D. rostrata (type locality: Greenland), D. baerii Rupr. (type locality: a site on the 
coast of Russian Lapland), D. corymbosa J. Agardh (type locahty: Greenland), 
and D. angustissima (Turner) A. Griffiths ex Harv. (type locality: west coast of 
Scotland). These four species, along with D. juergensii (J. Agardh) J. Agardh, had 
previously been clustered in the subgenus Cryptoneura of Delesseria by J. Agardh 
(1876). In his treatment of the algae of the Arctic Sea, Kjellman (1883) recognized 
D. rostrata to include Gigartina fabriciana. He also recognized D. baerii, D. corym- 
bosa, D. angustissima, D. alata (Huds.) J. V. Lamour., D. montagnei Kjellm. (a nomen 

'Hooper and South (1974) hsted the names Rhodomenia fabriciana J. Agardh (1841) and Euthora 
fabriciana (J. Agardh) J. Agardh (1847) in the taxonomic synonymy of Callophyllis cristata. DeToni 
(1897, p. 335), while placing these same J. Agardh names in synonymy with Euthora fruticulosa 
(Rupr.) J. Agardh, qualified this assignment with the phrase "excl. syn. Lyngb.," but elsewhere (p. 
229) he stated that Gigartina fabriciana Lyngb. "est Delesseria rostrata." However, on the basis of 
ICBN Art. 10. 1 (Greuter et al. 1994), these various names based on Gigartina fabriciana Lyngb. must be 
treated as its homotypic synonyms. 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 



novum), and D. sinuosa (Gooden. & Woodw.) J. V. Lamour. The last three taxa 
can be dismissed from further discussion here, because D. data is now recognized 
as Membranoptera alata (Huds.) Stackh., D. montagnei was a superfluous and 
thus an illegitimate substitute name for D. denticulata (Mont.) Mont, [now known 
as Membranoptera spinulosa (Rupr.) Kuntze] (see Silva et al. 1996), and D. sinu- 
osa is now known as Phycodrys rubens (L.) Batters (see below). In regard to the 
first four species Kjellman (1883) admitted that they were so closely similar to one 
another that many phycologists were inclined to abandon J. Agardh's (1852) rec- 
ognition of them as separate species. Yet, Kjellman concluded that they strongly 
differ from one another and that they arose "probably not long ago, from one or 
two fundamental types" (1883, p. 133). 

Similarly, Farlow (1886) struggled with the question of taxonomic limits of 
species assigned to subgenus Cryptoneura of Delesseria. He had earlier (Farlow 
1879) reported Delesseria rostrata from Cumberland Sound in Arctic America. 
On the basis of a very large collection of 150 specimens from Ungava Bay in the 
Canadian Arctic, Farlow (1886) stated that "In general the specimens could be 
arranged in two sets [which he attributed to D. baerii and D. corymbosa], although 
transitional forms were unfortunately frequent, and I am almost forced to believe 
that they are all forms of a single species, but what to call that species it is not 
easy to decide." He admitted that the material he had earher identified as D. 
rostrata had been sent to J. Agardh, who identified it as D. baerii. He went on to 
say that he was "inclined to regard all the specimens from Ungava Bay as forms of 
a single species, because there are several specimens which connect the two sets 
already described." He concluded by asking whether D. baerii and D. corymbosa 
were really distinct. 

Rosenvinge (1893, p. 806) next addressed this same taxonomic problem of the 
relationships among these same species. He recognized Delesseria corymbosa as a 
variety within D. baerii. This view was later followed by Jonsson (1901, 1904), 
whereas Lund (1959) and Caram and Jonsson (1972) regarded D. corymbosa as 
conspecific with P. baerii. At the same time Rosenvinge (1893, p. 804) treated 
Gigartina fabriciana as a synonym of his D. montagnei Kjellm. [var.] rostrata (Lyngb.) 
Rosenv. As for G. fabriciana, the type locality of Gigartina purpurascens [var.] 
rostrata Lyngb. is also Greenland. These workers ignored that the name with 
priority at the specific rank is G. fabriciana Lyngb. (1819). 

Kylin (1924) then transferred these various species that had been assigned to 
Delesseria to the genus Pantoneura. Pantoneura had been established by Kylin (in 
Kylin & Skottsberg 1919) as a monotypic genus of Delesseriaceae, based on P. 
plocamioides Kylin from South Georgia. Pantoneura differs from its closest rela- 
tive, Membranoptera, by its terete to compressed thallus in which the marginal 
parts are polystromatic rather than monostromatic. Kylin also effected the trans- 
fer of P. juergensii (J. Agardh) Kylin, a species described from the Kamchatka 
Peninsula, Russia, and easily separated from the other North American taxa by its 
much broader axes and robust size. Kylin (1924) acknowledged that three of the 
species that he transferred to Pantoneura (namely, P. rostrata, P. baerii, and P. 
corymbosa) are closely related to one another, and he remarked that it was not 
unlikely that they are only different forms of one species. 

The next question concerns the status of Pantoneura angustissima (Turner) 
Kylin. Turner (1811) was the first to recognize this entity as a variety within his 
Fucus alatus. Citing Turner's Fucus alatus var. angustissimus as a synonym, Harvey 
(1847, pi. 83) initially recognized ''Delesseria angustissima Griffiths msc." as a 



1997 WYNNE: DELESSERIACEAE 321 

distinct species. Harvey also cited his own Gelidium (?) rostratum A. Griffiths ex 
Harvey (1841) in taxonomic synonymy. Harvey stated that he was persuaded by 
the opinions of Mr. Brodie and Mrs. Griffiths of the distinctiveness of D. angustis- 
sima from D. data (Huds.) J. V. Lamour. Two years later, in his discussion ac- 
companying D. alata, Harvey (1849, pi. 247) indicated that he was no longer 
disposed to accept the specific rank of D. angustissima. Because the character of 
the membrane was not "absolute," Harvey regarded D. angustissima to be "an 
extreme variety" of D. alata. Some workers (e.g., Debray 1899; Picquernard 1912) 
followed Harvey in treating D. angustissima as a form of D. alata. Others, however, 
have recognized D. angustissima (or as Hypoglossum angustissimum or Pantoneura 
angustissima) as a distinct species; these workers include Kiitzing (1866), Batters 
(1890), Newton (1931), Parke and Dixon (1976), Rueness (1977), Lee (1980), 
South and Tittley (1986), and Hiscock (1986) in the eastern North Atlantic. More 
recently, Maggs and Hommersand (1993, p. 199) stated that "The seaweed recorded 
for the British Isles as Pantoneura angustissima (Griffiths ex Harvey) Kylin (1924, 
p. 18) is a species of Membranoptera. It appears to be a growth form of M. alata 
but further studies, preferably of freshly collected material, are required." Earlier, 
Kiitzing (1866, 1869) recognized Hypoglossum angustissimum from the Atlantic 
Ocean and D. baerii from the Okhotsk Sea. Kjellman (1877) reported D. baerii 
from the Murmansk Sea on the west coast of Novaja Zemlya. Sinova (1929a) 
recognized both D. baerii and D. angustissima from Novaja Zemlya between the 
Barents and the Kara seas of northern Russia, and she (1929b) also recorded both 
species from the White Sea. More recently, Vinogradova (1995) included the 
record of P. baerii from Spitsbergen. 

On the coast of eastern North America Taylor (1962) and Lamb and Zimmer- 
mann (1964) recognized the presence of both Pantoneura baerii and P. angustissima. 
Wilce (1959) reported only P. baerii from Quebec, Canada, whereas South (1984) 
referred to only P. baerii in his checklist of marine algae of eastern Canada. Although 
Ltining (1990) characterized P. baerii as probably restricted to the Arctic region, 
other workers have regarded this species to be generally widespread in Cold Boreal 
(sub-Arctic) seas (Gobi 1878; Rosenvinge 1898, 1924, 1933; B0rgesen & Jonsson 1905; 
Sinova 1954; Zinova 1954; Pedersen 1976). Actual reports of P. baerii from Arctic 
regions are rare (Lund 1933). In their flora of the Canadian Maritimes, Bird and 
McLachlan (1992) noted that specimens of Membranoptera alata in the Gulf of St. 
Lawrence become filiform and outwardly resemble P. baerii, which they said was a 
rarer species and lacked the marginal wings present in M. alata. 

Perestenko (1994) reported Pantoneura baerii from the Bering and Okhotsk 
Seas and the Sea of Japan. She (Perestenko 1996) later reported this species to be 
a common component of the subhttoral vegetation from the Okhotsk Sea and the 
Shantar Islands of the northwestern North Pacific Ocean. Saunders's report (1901, 
as Delesseria baerii) of the presence of P. baerii from Sitka, Alaska, on the basis 
of "two small sterile plants," has been questioned by Wynne (1970). 

Lamb and Zimmermann (1964) distinguished Pantoneura baerii from P. angustis- 
sima by its relatively large axial midrib-cells in the upper branches, which are 
apparent in surface view. In contrast, cells in the upper branches of P. angustissima 
are uniform in surface view, small, rounded, and more or less isodiametric. A slide 
with fragments of the type of Gigartina fabriciana was received on loan from 
Copenhagen, and Fig. 1 is a camera-lucida depiction of an apical region. The 
organization is clearly delesseriacean with a ""Membranoptera-iype"" apex (Kylin 
1924). A portion of the axis of this specimen proximal to the apex is depicted in 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 




f Gigartina fabrick 

1 to apex. Figs. 2, 4. Pantoneura angustis 

, MICH). Fig. 2. Apical region. Fig. 4. Porti 



Fig. 3. The cortication of the blade demonstrates that this material belongs to 
Pantoneura, and the presence of elongate cells along the midline suggests P. baerii 
rather than P. angustissima (sensu Lamb & Zimmermann, 1964). 

Among the many collections of material from the Okhotsk and the White 
Seas and identified as Pantoneura baerii were reproductive specimens, including 
male (US 43210), tetrasporic (US 14768), and cystocarpic (MICH, leg. E. Sinova 
1932). Spermatangial sori are restricted to final-order branches (Fig. 5), which are 
broader than vegetative blades. A sterile midline separates the pair of spermatan- 
gial sori. Tetrasporangia are not in obvious sori but are scattered among the 
cortical cells (Fig. 6), and fertile axes tend to be enlarged relative to vegetative 
blades. Cystocarps are immersed in fertile axes (Fig. 7), which results in some 
blade distortion. The portion of the axis distal to the cystocarp is narrowly attenu- 
ated, giving a beaked, or rostrate, appearance. This appearance conforms closely 
to Lyngbye's (1819) figure, plate 12B, in which he depicted cystocarpic material of 
his Gigartina purpurascens [y] rostrata. This taxon was the basis of Gelidium Iros- 
tratum Harvey (1841) as well as of Rhodomenia rostrata J. Agardh (1841) and 
Delesseria rostrata J. Agardh (1852). Harvey's (1853, p. 96) translation of J. Agardh's 
description of D. rostrata, in which the conceptacles [i.e., cystocarps] are described 



^YNNE: DELESSERIACEAE 




FIGS. 5-7. 
(US 43210). Fig. 
Sinova in 1932). 



as "immersed in the terminal segments, thrice as thick as the excurrent beak-Uke 
point of the ramuius and shorter than it" conforms both to Lyngbye's depiction of G. 
purpurascens var. rostrata and also to the cystocarpic material identified by E. Sinova 
as P. baerii (Fig. 7). 



. UNIVERSITY OF MICHIGAN HERBARIUM 



/' \^. 



<J 



( ockii Scdlteicd airangcment of cyslocarpic proliteralions over hldde (US 27342) 

Fragments of material labeled Fucus alatus [y] angustissimus Turner, collected 
by Brodie from northern Scotland, are in MICH. These fragments are derived 
from Turner's type material and were sent by the Royal Botanic Gardens, Kew, 
to W. R. Taylor some years ago. The axes are somewhat compressed and bear 
cystocarps. An examination of an apex shows that the onset of cortication com- 
mences close to the apex (Fig. 2), and more proximally the axis is totally covered 
by small, isodiametric cortical cells obscuring any elongate cells of the midline 
(Fig. 4). This observation is in agreement with that made by Lamb and Zimmer- 
mann (1964) in their characterization of Pantoneura angustissima. Cystocarps are 
produced on stipitate final-order branches, which are flattened distally (Fig. 8) 
and are axillary in position. Cystocarpic bladelets of the Brodie collection match 
those (Figs. 9, 10) of a specimen from Aberdeen [Scotland] (US 34894), which 
was distributed in Hohenack. Meeralgen No. 599 (as ''Hypoglossum angustissi- 
mum"'). The structure and position of these cystocarpic proliferations are in full 
agreement with the description by Harvey (1847, pi. LXXXIII), who stated that 
the "tubercles [were] mostly immersed in small accessory ramuli, springing from 
the axils of the upper branches." Likewise, Johnstone and Croall (1859) showed 
the cystocarps in P. angustissima to be produced in small axillary leaflike expan- 
sions. Despite the specific epithet, the axes of P. angustissima are always stated to 
be broader than those of P. baerii (Ruprecht 1851; Lamb & Zimmermann 1964). 
Basal portions of axes of specimens of P. angustissima from Scotland and England 
are about 1 mm wide, whereas axes of specimens of P. baerii from the White and 
Okhotsk Seas are usually in the range of 220 to 340 jim wide. 



1997 WYNNE: DELESSERIACEAE 325 



The conclusions of the present observations are that Gigartina fabriciana, 
based on an examination of the type, belongs to the genus Pantoneura; that this 
name (G. fabriciana Lyngb., 1819) has priority; and that P. rostrata, P. baerii, and 
P, corymbosa are indistinguishable from P. fabriciana and thus are regarded as its 
taxonomic synonyms. On the other hand, P. angustissima is regarded as a distinct 
species in agreement with Lamb and Zimmermann (1964). The following suite of 
differences are proposed to delineate this pair of species: 



ively broad, robust, 346-600 |am br 



Pantoneura fabriciana (Lyngb.) M. J. Wynne comb. nov. Gigartina fabriciana 
Lyngb., Tent, hydrophytol. dan. 48, pi. IID. 1819. Sphaerococcus cristatus 
C. Agardh [var.] fabricianus (Lyngb.) C. Agardh, Species algarum 301. 
1822. Sphaerococcus fabriciana (Lyngb.) K. Sprengel, Syst. veg. 329. 1827. 
Rhodomenia fabriciana (Lyngb.) J. Agardh, Linnaea: 15: 15. 1841. Euthora 
fabriciana (Lyngb.) J. Agardh, Kongl. Vetensk. Acad. Handl. 5: 12. 1847. 
Callophyllis {!) fabriciana (Lyngb.) Kiitz., Spec. alg. 747. 1849. 

Gigartina purpurascens [var.] rostrata Lyngb., Tent, hydrophytol. dan. 46, pi. 
12B. 1819. Rhodomenia rostrata (Lyngb.) J. Agardh, Linnaea: 15: 14. 1841. 
Gelidium ? rostratum (Lyngb.) A. Griffiths ex Harv., Man. Brit. alg. 82. 
1841. Delesseria rostrata (Lyngb.) J. Agardh, Spec. gen. ord. alg. 2: 685. 
1852. Pteridium rostratum (Lyngb.) J. Agardh, Spec. gen. ord. alg. 3: 481 
1876. Membranoptera rostrata (Lyngb.) Kuntze, Rev. gen. pi. 2: 904. 1891. 
Delesseria montagnei var. rostrata (Lyngb.) Rosenv., Meddel. Gr0nland 3: 
804. 1893. Hydrolapatha rostrata (Lyngb.) Kuntze, Rev. gen. pi. 3: 410. 
1898. Pantoneura rostrata (Lyngb.) Kylin, Lunds Univ. Arsskr., N. F., 
Avd. 2, 20(6): 17. 1924. 

Delesseria baerii Rupr., Tange Ochotsk Meer. 1(2): 239. 1851. Membranoptera 
baerii (Rupr.) Kuntze, Rev. gen. pi. 2: 904. 1891. Hydrolapatha baerii 
(Rupr.) Kuntze, Rev. gen. pi. 3: 410. 1898. Pteridium baerii (Rupr.) J. 
Agardh, 1898, p. 224. Pantoneura baerii (Rupr.) Kylin, Lunds Univ. Arsskr., 
N.F., Avd. 2, 20(6): 18. 1924. 

Delesseria corymbosa J. Agardh, Spec. gen. ord. alg. 2: 482. 1852. Membran- 
optera corymbosa (J. Agardh) Kuntze, Rev. gen. pi. 2: 904. 1891. Deles- 
seria baerii Ruprecht var. corymbosa (J. Agardh) Rosenv., Meddel. Gr0n- 
land 3: 806. 1893. Hydrolapatha corymbosa (J. Agardh) Kuntze, Rev. gen. 
pi. 3: 410. 1898. Pteridium corymbosum (J. Agardh) J. Agardh, Spec. gen. 
ord. alg. 3: 225. 1898. Pantoneura corymbosa (J. Agardh) Kylin, Lunds 
Univ. Arsskr., N. P., Avd. 2, 20(6): 18. 1924. 

Specimens Examined: Russia. Sea of Okhotsk, Siberia: /. Zaks, 10. ix. 1927, epiphytic on Laminaria 
stipe (US 014769); Ul'bansky Bay, 11 km north of Zarzhetsky Cape: V. Averintsev, 22. viii. 1978 
(MICH); Cape Ugalvnoy: E. Blinova, 28. viii. 1965, male (US 43210); Bay of Ayarna: collector 
unknown, 24. vi. 1903 (US 014768); mouth of Aldoma River: /. G. Wosnessensky, 1845, epiphytic on 



326 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

Odonthalia (US 014770); Gulf of Ajan: /. G. Wosnessensky, 1845, epiphytic on Phycodrys rubens (US 
014771). White Sea, Lemny Head: E. Sinova 1930 (US 014764); E. Sinova. 8-10. viii. 1928 (US 
014763); Kandalaksha, near Gulf of Maloye: E. Sinova & A. Zinova 1932 (MICH, US014765); mouth 
of River Ponoi: G. Tzinzerling, 6. ix. 1927 (US 014766). Novaya Zemlya, Matotschkin: R R. KjeUmau. 
7-13. vii. 1875 (MICH); Russian Harbor: A. Gemir, 5. x. 1936, epiphytic on Ptilota pectinata (US 014767). 
Barents Sea, south shore of Kolguyev Island: F. Riiprechl. 1841, epiphytic on Coccotylus (nmcatns 
(US 014772); Cape Kanin Noss: F. Riiprechl, 1841 (US 014773). Canada. Labrador, Ungava Bay: L. 
M. Turner, 1884 (MICH. US 078075). Quebec. Paspebiac, Co. Bonaventure: Akpatok-Quesnel = A. 
Cardinal 2270), 8. vi. 19h5 (MICH). DEVON, N. W. T. Eastern Bay, near base camp, ca. 13 miles SW 
of Cape Si-»arbo: R. Wilce. 27. vi. 1963 (MICH). Newfoundland, Otter Rub Point, Fortune Bay: R. Hooper 
& M. Rolwri^e, b. xi. 1971, Algae Terrae Novae No. 88 (MICH, US 085053); Burnt Point, Bonne Bay: R. 
Hooper & M. Roherge, 30. x. 1971, epiphytic on Phyllophora (MICH). U.S.A. Massachusetts, Essex 
County. Halibut Point, Cape Ann: /. M. Lamb A-385 19. xi. 1961, epiphytic on Chondrus crispus (MICH); 
Pauley Cove, Gloucester: R. Wetherbee & J. Sears (Wynne 3335), 25. vii. 1971 (MICH). 

Pantoneura angustissima (Turner) Kylin, Lunds Univ. Arsskr., N. F., Avd. 2, 
20(6): 18. 1924. Fucus alatus [var.] angustissimus Turner, Fuci, 60, pi. 160, 
figs, k, 1. 1811. Delesseria alata [var.] angustissima (Turner) C. Agardh, 
Spec. alg. 1(2): 179. 1822. Delesseria angustissima (Turner) A. Griffiths ex 
Harv., Phycol. Brit. pi. 83. 1847. Hypoglossum angustissimum (Turner) 
Ktitz., Spec. alg. 877. 1849. Pteridium angustissimum (Turner) J. Agardh, 
Spec. gen. ord. alg. 3: 482. 1876. Membranoptera angustissima (Turner) 
Kuntze, Rev. gen. pi. 2: 904. 1891. Hydrolapatha angustissima (Turner) 
Kuntze, Rev. gen. pi. 3: 410. 1898. 
Hypoglossum carpophyllum Kiitz., Tab. phycol. 16: 7. pi. 17. 1866 [fide Bat- 
ters (1890)]. 

Specimens Examined: Scotland. North of Scotland: leg. Brodie (fragment of type from KEW, now in 
MICH); Aberdeen: Hohenack. Meeralgen No. 599 (US 34894); Scapa Flow, Orkneys: 28. ix. 1940 (US 
161326). England. Scarborough: without date (US 34895). Filey Brigg: vii. 1880 (MICH). Canada. 
Baffin Island, Frobisher Bay, shore of Koojesse Inlet (63" 45' N, 68" 34' W): /, Calder & H. Senn. 10-20. 
vii. 1948 (MICH). 



THE STATUS OF THE NAME NEUROGLOSSUM LIGULATUM 

The genus Neuroglossum was established by Kiitzing (1843) based on N. bin- 
derianum Kiitz. (type locality: Cape of Good Hope, South Africa). Two additional 
species have been assigned to the genus. Kylin (1924) recognized N. ligulatum 
(Reinsch) Skottsb. in Kylin and Skottsberg (1919), which was originally described 
from South Georgia (Reinsch 1888, as Delesseria ligulata) but has also been re- 
ported from the South Sandwich Islands (Wynne 1982). The third species, N. 
multilobum, was described by Baardseth (1941) from Tristan da Cunha, Nightin- 
gale, Stoltenhoff, and Inaccessible islands in the South Atlantic. 

The basionym of Neuroglossum ligulatum, Delesseria ligulata Reinsch (1888), 
is an illegitimate name, because it is a later homonym for D. ligulata Gray (1821) 
[^^Hypoglossum hypoglossoides (Stackh.) Collins & Herv.]. According to Art. 58. 3 
(Greuter et al. 1994), N. ligulatum Skottsberg is to be treated as a new name; that 
name, however, is pre-dated by the taxonomic synonym Choreocolax delesseriae 
Reinsch (1890). Reinsch (1890) had misinterpreted the early stages in the produc- 
tion of laterals as a parasite, which he described as C. delesseriae on the host D. 
ligulata. Observations by Kylin and Skottsberg (1919) and Wynne (1982) demon- 
strated that these outgrowths are not an independent (parasitic) alga but are 
merely part of the alleged host. Thus, C. delesseriae can be applied to this species, 
and the following transfer is made. 



1997 WYNNE: DELESSERIACEAE 327 

Neuroglossum delesseriae (Reinsch) M. J. Wynne, comb. nov. Choreocolax deles- 
seriae Reinsch, Zur Meeresalgenflora von Siid-Georgien 2: 398, pi. 15, 
figs. 4-6. 1890. Delesseria ligulata Reinsch, Ber. Deutsch. Bot. Ges. 6: 148. 
1888, non Delesseria ligulata Gray, 1821. Neuroglossum ligulatum Skottsb. 
in Kylin & Skottsberg, Kenntn. Meeresalg. 4(15): 37. 1919. 
Nitophyllum manginii Gain, Bull. Mus. Hist. Nat. [Paris] 17: 483. 1911. Myrio- 
gramme manginii (Gain) Skottsb., Kongl. Svenska Vetenskapsakad. Handl. 
63(8): 548, fig. 13. 1923. 

Neuroglossum Kutz. (Kutzing 1843), along with Platydinia J. Agardh (J. Agardh 
1898), produces polystromatic blades, which in cross section show a central layer 
of large medullary cells surrounded by cells that gradually decrease in size toward 
the surfaces of the blade (Kutzing 1843, pi. 65, fig. 6; Skottsberg 1923). This 
anatomy is different from that of Myriogramme, in which the organization shows 
medullary cells to be essentially of uniform size (Ricker 1987; Wynne 1996). Neu- 
roglossum delesseriae has been shown to have the blade anatomy appropriate 
for the genus (Reinsch 1890, pi. 7, figs. 7, 8, as Delesseria ligulata- Skottsberg 1923, 
fig. 19c, as N. ligulatum; Wynne 1982, figs. 30, 31, as N. ligulatum). 

Myriogramme manginii (Gain) Skottsb. is another foliose species of Deles- 
seriaceae that has been reported to occur in Antarctic waters (Gain 1911; Skotts- 
berg 1953; Papenfuss 1964; Moe & DeLaca 1976). This species has been depicted 
(Gain 1912) as having an axis with basal branching and bearing a large number of 
simple rounded blades. The basal stipe is transformed into a conspicuous midrib 
that gradually disappears in the more distal portions of the blades. Cross sections 
of the stipe and the blade (Gain 1912, figs. 62-64) show an anatomy conforming 
to Neuroglossum rather than to Myriogramme. When Skottsberg (1953) trans- 
ferred the species to Myriogramme, he admitted that the generic position was 
doubtful in that the cells of the tri-stratose blades were not of uniform size, thus 
not conforming to classic Myriogramme. The many similarities between M. manginii 
and N. delesserieae in general branching pattern, which is quite variable in both 
taxa, in internal organization, and in reproduction are offered as evidence to 
regard M. manginii as a later taxonomic synonym of N. delesserieae. 

The occurrence of Neuroglossum binderianum from Cabo Frio, Brazil, reported 
by Yoneshigue (1985) and Yoneshigue-Valentin and Valentin (1992), represents a 
significant range extension in its distribution. Although Yoneshigue (1985) stated 
that the Brazihan material corresponded well with Wagner's (1954) description of 
this South African-based species, an examination of the anatomy (Yoneshigue 
1985, fig. 366) shows medullary cells of uniform size throughout, which casts 
doubt on the correct identification of the Brazilian material. 



Delesseria hancockii was described by Taylor (1945) from the Galapagos Is- 
lands, where it was dredged off Academy Bay, at Isla Santa Cruz. Taylor regarded 
it as rare and based his description only on tetrasporangiate material. It was 
depicted as a large and handsome plant, with a height reaching 85 cm. Its taxo- 
nomic status has remained unchallenged (Silva 1966), and it continues to be re- 
garded as endemic to the Galapagos. 

Abundant collections of Delesseria hancockii made in June, 1977, by F. Anger- 
meyer and M. M. Littler from Isla Isabella are now deposited in the US. These 



328 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 

collections include both cystocarpic (Fig. 12) and tetrasporangiate (Fig. 13) speci- 
mens. The tetrasporangiate specimens are in full agreement with Taylor's original 
account. It is the discovery of cystocarpic specimens that permits the recognition 
that this species has its correct taxonomic assignment to Pseudolaingia rather than 
to Delesseria. Pseudolaingia was established by Levring (1944) on the basis of 
Delesseria larsenii Skottsb. (in Kyhn and Skottsberg 1919) and remains a mono- 
typic genus. Pseudolaingia larsenii (Skottsb.) Levring has a distribution including 
South Georgia (the type locality), the Kerguelen Islands (Levring 1944), Tierra 
del Fuego of southern Chile (Levring 1960), and southern Argentina (Mendoza 
1973). Male, female, and tetrasporangiate plants were described by Mendoza (1973), 
and Wynne (1989) proposed criteria to distinguish Ps. larsenii from superficially 
similar taxa [including Delesseria lancifolia J. Agardh, Laingia hookeri (Lyall) 
Kylin, Pseudonitophylla (Mendoza 1975), and Odontolaingia (Mendoza 1976)]. 

Delesseria and Pseudolaingia are closely related genera, both having a similar 
apical organization and monostromatic blades in regions between midribs and 
lateral nerves. These two genera are primarily separated as follows: Delesseria 
bladelets bearing procarps/cystocarps arise only from the midrib or lateral nerves 
of the parent blades (Mendoza 1974; Wynne 1982), whereas in Pseudolaingia the 
special proliferations bearing the procarps (and resultant cystocarps) are located 
randomly over the entire blade surface (Figs. 11, 12) and have no relationship 
with the midrib or lateral nerves of the parent blade (Mendoza 1973; Wynne 
1989). Additionally, the tetrasporangial sori are produced over the surface of 
ordinary blades in Pseudolaingia, but tetrasporangia are produced in special pro- 
liferations in Delesseria (Mendoza 1974; Wynne 1989). The random arrangement 
of the cystocarpic proliferations evident in the female specimens of D. hancockii 
and the pattern of tetrasporangial production both demonstrate that this species 
should be assigned to Pseudolaingia rather than Delesseria. Another generic dis- 
tinction offered by Wynne (1989) is that the vegetative branching in Delesseria, 
where it occurs, tends to be from the midrib, but the vegetative branching in 
Pseudolaingia is from the blade margins. Thus, the following transfer is effected: 

Pseudolaingia hancockii (W. R. Taylor) M. J. Wynne, comb. nov. Delesseria han- 
cockii W. R. Taylor, Allan Hancock Pacific Exped. 12: 278, pi. 90. 1945. 

On the basis of the collections of both species available at my disposal, 
Pseudolaingia hancockii can be distinguished from the congeneric Ps. larsenii by 
the following key: 

1. Thallus prominently of secondary blades, the primary and secondary blades often becoming 

each 3-7 cm wide; midrib of primary axes conspicuous but narrow, 2-3 mm wide. 

Ps. hancockii. 
I. Thallus prominently of a primary blade, this primary blade 
blades relatively broad, 4-28 cm, but often deeply lobed c 
blade well developed, 3-5 mm wide. 

Specimens ExAMiNEt 
meyer & M. Littler, 29. 
027338, 027339, 027340) 




8cm 



330 CONTR. UNIVERSITY OF MICHIGAN 

PHYCODRYS RUBENS (L.) BATTERS VERSUS 
PHYCODRYS CRENATA (S. G. GMEL.) P. C. SILVA 

Silva et al. (1996) presented arguments to dismiss the name Phycodrys rubens 
(L.) Batters, which had become generally accepted to be the correct name of the 
type of the genus Phycodrys (Dixon 1964; Maggs & Hommersand 1993). The 
taxonomically equivalent name Phycodrys crenata (S. G. Gmelin) P. C. Silva (in 
Silva et al. 1996) was proposed as the name to replace P. rubens. The crux of the 
argument offered by Silva et al. (1996) to disallow P. rubens was that Batters 
(1902: 76) listed "F. rubens Hudson" (1762: 475) as basionym rather than F. rubens 
L. (Linnaeus 1753: 1162). Hudson (1762) credited Linnaeus (1753) as author of 
the name Fucus rubens. Yet Silva et al. (1996: 906) treated "F. rubens Hudson" as a 
circumscription rather than as a name. The original account of Fucus rubens 
(Linnaeus 1753) is the following: "23. Fucus caule tereti ramoso, frondibus oblongis 
undulatis sinuatis. Roy. lugdb. 514. Alga minor suave rubens varie divisa. Mart, 
cent. 32. t. 32. Habitat in Oceane." 

As Dixon (1964) has pointed out, this first reference is taken directly from 
van Royen (1740). According to Batters (1902: 65) the evidence strongly suggests 
that Linnaeus gave this name {F. rubens) to van Royen's species of Fucus with the 
polynomial "without ever having seen a specimen of the plant." Linnaeus then, 
unfortunately, later placed in his herbarium material of two totally different spe- 
cies (now assignable to the genera Phyllophora and Palmaria) that had nothing to 
do with the van Royen alga cited. This action caused the name Fucus rubens to be 
misapplied to Phyllophora. Batters (1902) recognized that the Linnaean name 
applied to the Delesseriacean alga Phycodrys rather than to Phyllophora. The 
imphcation is that Batters (1902) did not cite Linnaeus as author of the name F. 
rubens, because he was unable to locate authentic material of van Royen's alga. 
Dixon (1964) succeeded in tracking down the specimen in question and confirmed 
that it was the delesseriacean alga that Batters had thought it likely to be, without 
having seen the actual specimen. Dixon credited Batters as having made the 
combination Phycodrys rubens and cited Linnaeus rather than Hudson as being 
the author of the basionym, i.e., Phycodrys rubens (L.) Batters. According to 
ICBN Art. 33. 3, incorrect forms of author citation do not invalidate pubhcation 
of new combinations, and Example 6 can be cited as a similar instance allowing 
for such an adjustment. This interpretation of the Code is clearly a more reason- 
able course to follow than the one presented in Silva et al. (1996). 

THE AUTHORSHIP OF THE TYPE SPECIES OF PLATYCLINIA 

When J. Agardh (1898) estabhshed the genus Platyclinia, one of the three 
species he assigned to it was P. stipitata J. Agardh. Wynne (1983) presented argu- 
ments to treat P. stipitata as the lectotype of the genus. J. Agardh cited, with a 
query, Nitophyllum stipitatum Harv. in Hooker (1860), described from Tasmania, 
as the apparent basionym. This name, however, is ihegitimate because it is a later 
homonym of N. stipitatum Suhr (1841) from Tangier. DeToni (1900) indicated 
that Suhr's N. stipitatum is perhaps identical to Stenogramme sp. Stenogramme inter- 
rupta (C. Agardh) Mont. (Montagne 1846) has a worldwide distribution, including 
several records from Morocco and elsewhere in the Mediterranean (Bornet, 1892; 
Gayral 1958; Schotter 1968; Dixon & Irvine 1977). Nonetheless, in respect to 



1997 WYNNE: DELESSERIACEAE 33 

ICBN Art. 58. 3 (Greuter et al. 1994), the authorship for P. stipitata should be . 
Agardh (1898) rather than (Harvey) J. Agardh. 



ACKNOWLEDGMENTS 

teful to Ruth Nielsen of the Botanical Museum and Library, University of Copenl 
in locating Lyngbye's material of Gigartina fabriciana, and to Barrett Brooks 
Institution for his handling a loan of specimens from US. I thank Howard A. Cr 
y of Michigan for translating a portion of Ruprecht's publication for me and Davi 
; the photographs of the artwork. Craig W. Schneider and Robert T. Wilce 



LITERATURE CITED 

Agardh, C. A. 1822-1823. Species algarum.. .Vol 1(2), pp. [1-6] + 169-398 (1822), 399-531 (1823). 

. 1824. Systema algarum. xxxviii + 312 pp. Lund. 

Agardh, J. G. 1841. In historiam algarum symbolae. Linnaea: 15: 1-50; 443-457. 

. 1847. Nya alger frSn Mexico. Kongl. Vetensk. Acad. Handl. 5: 46-49. 

. 1852. Species genera et ordines algarum... Vol. 2(2). pp. 337-720. Lund. 

. 1876. Species genera et ordines algarum.. . Vol. 3(1 ). vii + 724 pp. Lund. 

. 1898. Species genera et ordines algarum... Vol. 3(3). [vi] + 239 pp. Lund. 

Baardseth, E. 1941. The Marine Algae of Tristan da Cunha. Results of the Norwegian Scientific 

Expedition to Tristan da Cunha 1937-1938. No. 9. 173 pp. Oslo. 
Batters, E. A. L. 1890. A list of the marine algae of Berwick-on-Tweed. Hist. Berwickshire Natural- 
ists' Club 12: 221-392, pis. VII-XI. [Reprinted: 1-172 pp.] 

. 1902. A catalogue of the British marine algae. J. Bot. 40 (Suppl.). 107 pp. 

Bird, C. J., and J. L. McLachlan. 1992. Seaweed Flora of the Maritimes 1. Rhodophyta - the Red Algae. 

V + 177 pp. Biopress Ltd., Bristol, England. 
B0rgesen, F., and H. Jonsson. 1905. The distribution of the marine algae of the Arctic Sea and of the 

northernmost part of the Atlantic. Botany of the Faeroes, vol. 3, Appendix, pp. I-XXXVIIL 

Copenhagen. 
Bornet, E. 1892. Les algues de P.-K.-A. Schousboe recoltees au Maroc et dans le Mediterranee de 

1815 a 1829. Mem. Soc. Sci. Nat. Cherbourg 28: 165-376, 3 pis. 
Caram, B., and S. Jonsson. 1972. Nouvel inventaire des algues marines de I'lslande. Acta Bot. Islandi- 

ca 1: 5-31. 
Debray, F. 1899. Florule des algues marines du Nord de la France. Bull. Sci. France Belgique 32 (ser. 

DeToni, G. B. 1897. Florideae, Sectio I. In Sylloge algarum. Vol. IV, xx + Ixi, 1-386 + 387-388 

[Index]. Padua. 

-. 1900. Florideae, Sectio II. In Sylloge algarum. Vol. IV, 3^1-11'^ + 115-ll(y [Index]. Padua. 

Dixon, P. S. 1964. Taxonomic and nomenclatural notes on the Florideae, IV. Botaniska Not. 117: 56-78. 
Dixon, P. S., and L. M. Irvine. 1977. Seaweeds of the British Isles. Volume 1, Rhodophyta. Part 1, 

Introduction, Nemaliales, Gigartinales. xi + 252 pp. British Museum (Natural History), London. 
Farlow, W. G. 1879. A list of algae collected at points in Cumberland Sound during the autumn of 1877. In 

Contributions to the Natural History of Arctic America, made in Connection with the Howgate Polar 

Expedition, 1877-78 (L. Kumlien), p. 169. Government Printing Office, Washington D.C. 
. 1886. Notes on Arctic algae; based principally on collections made at Ungava Bay by Mr. L. 

M. Turner. Proc. Amer. Acad. Arts & Sci. 21: 469-477. 
Gain, L. 1911. Sur trois especes nouvelles d'algues marines provenant de la region antarctique sud- 

americaine. Bull. Mus. Hist. Nat. [Paris] 17: 482-484. 
. 1912. La flore algologique des reegions antarctiques et subantartiques. In Deuxieme Expedition 

Antarctique Franqaise (1908-1910) (J. Charcot, ed.), 218 pp., 5 pis. Science naturelles, Paris. 
Gayral, P. 1958. Algues de la Cote Atlantique Marocaine. La Nature au Maroc. Public par la Societe 

des sciences naturelles et physiqes du Maroc. Rabat. 
Gobi, C. 1878. Die Algenflora des Weissen Meeres und der demselben zunachstliegenden Theile des 

nordlichen Eismeeres. Mem. Acad. Imp. Sci. Saint Petersbourg, ser. 7, 26(1). 92 pp. 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUN 



Joy, London. 
Greuter, W. et al. (eds.) 1994. Internatior 

by the Fifteenth International Bota 

num Veg. 131: 1-389. 
Hansen, J. B. 1985. An outline of phycology in Denmark. Published in conjunction with the Second 

International Phycological Congress, 1985. Danish Nat. Sci. Res. Council. 24 pp. 
Harvey, W. H. 1841. A Manual of the British Algae... Ivii + 229 pp. John Van Voorst, London. 

. 1847. Phycologia Britannica. Pis. 73-144. London. 

. 1849. Phycologia Britannica. Pis. 217-252. London. 

. 1853. Nereis Boreali- Americana... Part II. Rhodospermeae. Smithsonian Contr. Knowl. 5(5). 

258 pp., pis. XIII-XXXVl. 
. 1860. Algae. In The Botany of the Antarctic Voyage of H.M. Discovery ships Erebus and 

Terror... III. Flora Tasmanicae. Part II. Monocotyledons and Acotyledons. (J. D. Hooker, ed.) 

Pp. 282-343, pis. 185-196. London. 
Hiscock, S. 1986. A field key to the British red seaweeds. AIDGAP, Field Studies Council, Occas. 

Publ. 13. 101 pp. ["Field Studies," Nettlecombe Court, WiUiton, Taunton, Somerset, U. K., TA4 

4HT] 
Hooper, R., and G. R. South. 1974. A taxonomic appraisal of Callophyllis and Euthora (Rhodophyta). 

Br. Phycol. J. 9: 423-428. 
Hudson, W. 1762. Flora anglica... viii + 506 pp. London. 

Johnstone, W. G., and A. Croall. 1859. The Nature-Printed British Sea-Weeds... Vol. I. Rhodosper- 
meae. Bradbury and Evans, London. 
Jonsson, H. 1901. The marine algae of Iceland. I. Rhodophyceae. Bot. Tidsskr. 24: 127-155. 

. 1904. The marine algae of East Greenland. Meddel. Gr0nland 30: 1-73. 

Kjellman, F. R. 1877. Ober die Algenvegetation des Murmanischen Meeres an der Westkuste von Nowaja 

Semlja und Wajgatsch. Nova Acta Regiae Soc. Sc. Upsal., ser. 3, vol. extra ord. (12): 1-86, 1 pi. 

general characters and the development of the flora. Kongl. Svenska Vetenskapsakad. Handl. 

20(5). 350 + 1 pp. 
Klitzing, F. T. 1843. Phycologia generalis... 458 pp., 80 pis. Leipzig. 

. 1849. Species Algarum. vi + 922 pp. Brockhaus, Lipsiac [Leipzig]. 

. 1866. Tabulae Phycologicae. Vol. 16. 35 pp., 100 pis. Nordhauscn. 

. 1869. Tabulae Phycologicae. Vol. 19. 36 pp., 100 pis. Nordhausen. 

Kyhn, H. 1924. Studien uber die Delesseriaceen. Lunds Univ. Arsskr., N. F., Avd. 2, 20(6). Ill pp. 
KyHn, H., and C. Skottsberg. 1919. Zur Kenntnis der Subantarktischen und Antarktichen Meeresal- 

gen. 11. Rhodophyceen. Wiss. Ergeb. Schwed. Sudpolar-Exped. 1910-1903. Vol. 4(15). 88 pp., 1 

Lamb, I. M., and M. H. Zimmermann. 1' 

chusetts. II. The occurring of the j 

Rhodora 66: 234-254. 
Lee. R. K. S. 1980. A catalogue of the marine algae of the Canadian Arctic. Natl. Mus. Canada Publ. 

Bot. 9. 82 pp. 
Levring, T. 1944. Meeresalgen von den Crozet-lnseln und Kerguelen. Arkiv Bot. 31(8). 31 pp. 
. 1960. Contributions to the marine algal flora of Chile. Lunds Univ. Arsskr., N.F. Avd. 2, 

56(10). 83 + [2] pp. 
Linnaeus, C. 1753. Species plantarum... Vol. 2. Pp. 561-1200 + [31]. Holmiae [Stockholm]. 
Liining, K. 1990. Seaweeds, their Environment, Biogeography, and Ecophysiology. [English language 

edition edited by C. Yarish and H. Kirkman.] xiii + 527 pp. John Wiley & Sons, Inc., New York. 
Lund, S. 1933. The marine algae. The Godthaab Expedition 1928. Meddel. Gr0nland 82(4); 1-18 . 

. 1959. The marine algae of East Greenland. I. Taxonomical part. Meddel. Gr0nland 156(1); 1-247. 

Lyngbye, H. C. 1819. Tentamen Hydrophytologiae Danicae. xxxii + 248 pp., 70 pis. Hafniae [Copenhagen]. 
Maggs, C. A. and M. H. Hommersand. 1993. Seaweeds of the British Isles. Volume 1, Rhodophyta 

Part 3A Ceramiales. xv + 444 pp. Nat. Hist. Mus., London. 
Mendoza, M. L. 1973. The masculine sexual plants of Pseudolaingia larsenii (Skottsb.) Levring (Rhodo- 
phyta) and its occurrence in Tierra del Fuego and Isla de los Estados. Physis (Buenos Aires), 

. 1974. The masculine sexual plants of Delesseria fuegiensis Skottsberg and the occurrence of 



3 Estados. Physis (Bu- 



1997 WYNNE: DELESSERIACEAE 333 

. 1975. Pseudonitophylla, a new genus of the family Delesseriaceae (Rhodophyta). Physis 

(Buenos Aires), Secc. A, 34: 59-65. 
. 1976. Odontolaingia, a new genus of the family Delesseriaceae (Rhodophyta). Bol. Soc. 

Argent. Bot. 17: 190-198. 
Moe, R. L., and T. E. DeLaca. 1976. Occurrence of macroscopic algae along the Antarctic Peninsula. 

Antarc. J. U.S. 11:20-24. 
Montagne, [J. F.]C. 1846. Note sur le genre Stenogramma Harv., de la famille des Floridees. Rev. Bot. 

Bull. Mens. 1: 481-483. 
Newton, L. 1931. A Handbook of the British Seaweeds, xiii + 478 pp. Brit. Mus., London. 
Papenfuss, G. F. 1964. Catalogue and bibliography of Antarctic and Sub-Antarctic benthic marine 

algae. Antarc. Res. Ser. 1: 1-76. 
Parke, M., and P. S. Dixon. 1976. Check-list of British marine algae - third revision. J. Mar. Biol. 

Assoc. U. K. 56: 527-594. 
Pedersen, P. M. 1976. Marine, benthic algae from southernmost Greenland. Meddel. Gr0nland 199(3): 

1-80. 
Perestenko, L. P. 1994. Red Algae of the Far-Eastern Seas of Russia. Komarov Botanical Institute, 

Russian Academy of Sciences, St. Petersburg. 331 pp. [In Russian with English abstract.] 
. 1996. Sublittoral phytocoenoses of the south-western coasts of the Okhotsk Sea and the 

Shantar Islands. Bot. Zurn. SSSR 81(7): 41-55. 
Picquernard, C.-A. 1912. Etudes sur les collections botaniques des freres Crouan. IV. Florideae. 

Trav. Sci. Lab. Zool. Physiol. Marit. Concarneau 4(4): 1-104. 
Reinsch, P. F. 1888. Species et genera nova algarum ex insula Georgia Australi. Ber. Deutsch. Bot. 

Ges. 6: 144-156. 
. 1890. Zur Meeresalgenflora von Sud-Georgien. In International Polarforschung, 1882-1883. 

Die dentschen Expeditionen und ihre Ergebnisse. Vol. 2. Beschreibende Naturwissenschaften... 

(G. Neumayer. ed.) pp. 336-449, pis. 1-19. Berlin. 
Ricker, R. W. 1987. Taxonomy and Biogeography of Macquarie Island Seaweeds, vi + [2] + 344 pp. 

Brit. Mus., London. 
Rosenvinge, L. K. 1893. Gr0nlands Havalger. Meddel. Gr0nland 3: 763-981, 2 pis. 

. 1898. Deuxieme memoire sur les algues marines du Groenland. Meddel. Gr0nland 20: 1-128, 1 pi. 

. 1924. Marine algae. In "A botanical trip to Jan Mayen" by J. Gandrup. Dansk Bot. Arkiv 

4(5): 21-23. 
. 1933. Marine algae from Kangerdlugssuak. The Scoresby Sound Committee's 2nd East Green- 
land Expedition in 1932 to King Christian IX's Land. Meddel. Gr0nland 104(8): 14 pp. 
Royen, A. van. 1740. Florae leydensis prodromus... [LXXII] + 538 + [30] pp. Lugduni batavorum 

[Leiden]. 
Rueness, J. 1977, Norsk Algflora. 266 pp. Universitetsforlaget, Oslo. Bergen, Troms0. 
Ruprecht, F. J. 1851. Tange des Ochotskischen Meeres. In Sibirische Reise by A. Th. von Middendor- 

ff, 1(2): [191]-435, 10 pis. St. Petersburg. 
Saunders, De A. 1901. Papers from the Harriman Alaska Expedition. XXV. The algae of the expedi- 
tion. Proc. Wash. Acad. Sci. 3: 391^86, 19 pis. 
Schotter, G. 1968. Recherches sur les Phyllophoracees. Bull. Inst. Oceanogr. Monaco 67(1383): 99 pp. 
Silva, P. C. 1966. Status of our knowledge of the Galapagos benthic marine algal flora prior to the 

Galapagos International Scientific project. In The Galapagos, Proceedings of the Symposia of 

the Galapagos International Scientific Project (R. I. Bowman, ed.). Pp. 149-156. University of 

California Press, Berkeley. 
Silva, P. C, P. W. Basson, and R. L. Moe. 1996. Catalogue of the benthic marine algae of the Indian 

Ocean. Univ. Calif. Publ. Bot. 79. xiv + 1259 pp. 
Sinova, E. S. 1929a. Algues de la Novaja Zemlja. Institut Hydrologique, Explorations des mers 

d'U.R.S.S. Fasc. 10, pp. 41-128. Leningrad. [In Russian, with French resume.] 
. 1929b. Les algues de la mer Blanche. Travaux de la Societe des Naturalistes de Leningrad 

59(3): 3-40. [In Russian with French resume.] 
. 1954. Vodorosli okhotskogo moria. [Seaweeds of the Okhotsk Sea]. Trudy Botan. Komarov 

Inst., Akad. Nauk S.S.S.R., ser. 2, 9: 259-310. [In Russian.] 
Skottsberg, C. 1923. Botanische Ergebnisse der Schwedischen Expedition nach Patagonien und dem 

Feuerland 1907-1909. IX. Marine algae 2. Rhodophyceae. Kongl. Svenska Vetenskapsakad. 

Handl. 63(8): 1-70. 
. 1953. On two collections of Antarctic marine algae. Arkiv Bot., ser. 2, 2: 531-566, 1 pi. 



CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 



Suhr, J. N. 1S41. Beitrage zur Algenkunde. Verhandl. Kaiserl. Leopold.-Carol. Akad. Naturf. 18, 

Taylor, W. R. 1945. Pacific marine algae of the Allan Hancock Expeditions to the Galapagos Islands. 

Allan Hancock Pacific Exped. 12: iv + 528 pp. 
. 1962. Marine Algae of the Northeastern Coast of North America. Revised Edition, second 

printing with corections. viii + [1] + 509 pp. University of Michigan Press, Ann Arbor. 
Turner, D. 1811. /-((cv... Vol. 3, pp. 1-148, pis. 135-196. John and Arthur Arch, London. 
Vinogradova, K. L. 1995. The checklist of the marine algae from Spitsbergen. Bot. Zurn. SSSR 80: 

50-61. 
Wagner, F. S. 1954. Contributions to the morphology of the Delesseriaceae. Univ. Calif. Publ. Bot. 

27: 279-346. 
Wilce, R. T. 1959. The marine algae of the Labrador Peninsula and northwest Newfoundland (ecology 

and distribution). Natl. Mus. Canada Bull. 158. iv + 103 pp. 
Wynne, M. J. 1970. Marine algae of Amchitka Island (Aleutian Islands). I. Delesseriaceae. Syesis 3: 

. 1982. Observations on four species of Delesseriaceae (Rhodophyta) from the South Sand- 
wich Islands, the Antarctic. Contr. Univ. Michigan Herb. 15: 325-337. 

. 1983. The current status of genera in the Delesseriaceae (Rhodophyta). Bot. Mar. 26: 437^ 

450. 

. 1989. Observations on Pseudolaingia larsenii (Skottsb.) Levr. (Delesseriaceae, Rhodophyta). 

Jap. J. Phycol. 37: 39-45. 



Akad. Nauk SSSR. Moscow