CONTRIBUTIONS FROM THE UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 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 CONTRIBUTIONS FROM THE UNIVERSITY OF MICHIGAN HERBARIUM ISSN 0091-1860 Copyright 1997 University of Michigan Herbarium All rights reserved Printed in the United States of America Volume 21 Editor: Christiane Anderson For information about the availability and prices of previous volumes of the Contributions, address the Director, University of Michigan Herbarium, North University Building, Ann Arbor, Michigan 48109-1057, U.S.A. —" 20 October 1997 CONTENTS Revision of Prerandra (Malpighiaceae) CHRISTIANE ANDERSON Excentradenia, a new genus of Malpighiaceae from South America WILLIAM R. ANDERSON Notes on neotropical Malpighiaceae—VI WILLIAM R. ANDERSON The summit flora of Mt. Murud, Sarawak, Malaysia JOHN H. BEAMAN & CHRISTIANE ANDERSON Aristolochia davilae (Aristolochiaceae), a new species from Nayarit, Mexico ISMAEL CALZADA J., GABRIEL FLORES F. & OSWALDO TELLEZ V. Miscellaneous notes on Sphagnum—10 HowARD CRUM Catalog of the Acanthaceae of Belize with taxonomic and phytogeographic notes THOMAS F. DANIEL A revision and redefinition of Pseudabutilon (Malvaceae) PauL A. FRYXELL New species and new records for the Malvaceae of Baja California PAUL A. FRYXELL, JOSE Luis LEON DE LA Luz & MIGUEL DOMiNGUEz L. Revision and phylogenetic study of Anomodon and Herpetineuron (Anomodontaceae, Musci) INIGO GRANZOW-DE LA CERDA Species delimitation, and new taxa and combinations in Leucaena (Leguminosae) OLIN HUGHES Two new species of Carex (Cyperaceae) from Mexico A. A. REZNICEK & M.S. GONZALEZ-ELIZONDO Carex molestiformis (Cyperaceae), a new species of section Ovales from the Ozark Mountain region A. A. REZNICEK & PAUL E. ROTHROCK Nomenclatural changes in Mexican Dioscoreaceae and Leguminosae OswaLpo TELLEZ V. A new species of ingens (Orchidaceae) from Mt. Murud, Sarawak, Borne JEFFREY J. Woop Taxonomic and nomenclatural notes on the Delesseriaceae MICHAEL J. WYNNE ZiT 20 299 OS Contr. Univ. Michigan Herb. 21: 1-27. 1997. 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 Prerandra 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 Brazilian 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 1 2 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 40° Pterandra © andersonii @ arborea A colombiana {1 * egleri oO ° ® pyroidea ¥ sericea * ultramontana FIG. 1. Distribution of Pterandra. 1932. The many collecting efforts in Central and South America launched since World War II have greatly increased our knowledge of that flora and revealed many novelties. Since 1953, eight more species of Prerandra were recognized, and another four are added here (P. andersonii, P. colombiana, P. hirsuta, and P. mephersonil). INFRAGENERIC RELATIONSHIPS Although 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 little 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 Prerandra. 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: 0 = large shrubs to trees; | = small shrubs. Future collections may allow finer distinctions in habit. Prerandra andersonii is known from only two collections, both described as a low shrub, and thus clusters with P. hatschbachu 1997 C. ANDERSON: PTERANDRA 3 TABLE 1. Data matrix for fourteen species of Pferandra and the outgroup Acmanthera. Acmanthera 0&1 0 0&1 0 0 | 0 0 0 0 0 0 P. andersoni | 0 0 1 1 0 0 1 0 1 0 0 P. arborea 0 0 0 1 0 0 1 ! | 0 0 P. colombiana 0 0 | 1 0 0 1 1 0 1 0 0 P. egleri 0 0 0 l 0 0 0 | 0 l 0 0 P. evansii 0 | 0 l 0 0 0 1 0 1 ) 0 P. flavescens 0 0 0 1 0 0 0 l 0 0 0 0 P. guianensis 0 0 0 | 0 0 ! l l 0 0 0 P. hatschbachii I 0 | 1 1 0 0 l 0 l ! ! P. hirsuta 0 1 0 1 0 0 0 1 0 1 0 0 P. isthmica 0 0 2 1 0 0 0 1 l 1 0 1 P. mcphersonii 0 0 l 1 0 0 ] | I | 0 0 P. pyroidea I 0 O&1 1 1 0 0 1 0 l 1 1 P. sericea 0 0 2 1 0 0 0 ! 1 0 0 0 P. ultramontana 0 0 0 ] 0 0 0 l 0 1 0 0 and P. pyroidea, but it is possible that this species, like some others, varies from shrub to small tree. 2. Attachment of hairs of the abaxial laminar vesture: 0 = medifixed; 1 = basifixed. The common condition in Pterandra and Acmanthera 1s 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. u/tramon- 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: 0 = 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: 0 = absent; 1 = present. All species of Prerandra 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: 0 = 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: 0 = 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 Prerandra 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. . Abaxial stipular surface: 0 = 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: 0 = pseudoraceme; | = 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: 0 = 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. mephersonii, 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: 0 = 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: 0 = 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. 1997 C. ANDERSON: PTERANDRA 3) 12. Size of limb of petal: 0 = 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. tsthmica. 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”].— Lecrotype, 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 elliptical 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 6 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Acmanthera Acmanthera andersonii 3(1) = hatschbachii : — -L_ hirsuta ultramontana flavescens i flavescens Acmanthera Acmanthera andersonii hatschbachii 7 2(1) evansii [ hirsuta ultramontana a — evansii “aa L_ hirsuta 4 flavescens ultramontana Tree 3 Consensus tree FIG. 2. Three most parsimonious cladograms and the consensus tree for 14 species of Prerandra and the outgroup Acmanthera. 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 i 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 elliptical 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 consistent with numbers reported for the Byrsonimoideae, which are n = 6, 12, or ca. 24 (see W. R. Anderson, 1993). KEY TO THE SPECIES OF PTERANDRA 1. Petals pink; petioles of even the largest leaves only 0.1-0.5 cm long; small shrubs to 2 m. 2. Laminas abaxially tomentose to sparsely so (rarely glabrate); stipules 3.5-8 mm long, 4-7 mm wide; petals sparsely to densely sericeous on claw and center of limb; Brazil (Distrito Federal, Goids, and Minas Gerais; Mato Grosso?). (ie ee ea 2. Laminas abaxially glabrous or with some scattered hairs, especially on the major veins stipules 2-2.5 mm long and wide; petals abaxially glabrous or very sparsely sericeous claw and center of limb; Brazil (Mato Grosso). 2oPan baa 1. Petals white or cream or greenish yellow, often becoming yellow in age; petioles up to 3.5 cm long, well over 0.5 cm long in larger leaves (except to 1 cm long in P. andersonii); shrubs or trees to 30 m 3. Abaxial laminar pubescence composed mostly of erect basifixed hairs and a few scattered medifixed hairs; Brazil (Amazonas, Rond6onia) and Bolivia (Beni). Ll. P. hirsuta. 3. Abaxial laminar pubescence composed entirely of medifixed hairs (in P. evansii 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), or the eee abaxially glabrate to glabrous 4. Stipules abaxially glabrous even in bud (but often with a sericeous patch at point of attachment to petiole in P. guianensis, with a sericeous patch at the apex in P. colombiana). 5. Bracts and bracteoles EEE aE densely pubescent; laminas abaxially sparsely seri- ceous, the hairs evenly distributed 6. Petals white; pedicels 0.4-0.6 mm in diameter, the pubescence white or silvery: leaf apex obtuse or acute or acuminate with a tip to | cm long; pellucid cells on abaxial surface of lamina abundant to sometimes absent; Brazil (Amapa, Amazo- nas, Para). 4. P. arborea. Petals greenish yellow; pedicels 0.8-1 mm in diameter, the pubescence bite yellow; leaf apex acuminate to caudate, the tip up 1.5 cm long; pellucid cells in abaxial surface of lamina very few to absent; Guyana. 5. P. guianensis. a 8 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 5. Bracts and bracteoles abaxially glabrous or with some hairs on the costa and/or a tuft at the ale laminas abaxially glabrate or very sparsely sericeous and the hairs widely scatte 7. Fascicles ower sessile, the pedicels 1.4-2 cm long; petioles up to 3.5 cm long: olombia i Stdum 6. P. colombiana. 7. ses les of flowers borne on short projections (up to 3 mm long), the pecine els ae petioles up to 1.3. cm long; Panama (Col6én 7. P. mephersonii. 4. Stipules pears densely sericeous, sometimes sparsely so in age (especially in P. ultramontana). aminas 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. egleri. 8. pee abaxially glabrate or sparsely to densely sericeous and the pubescence appressed (in P. evansii 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), the costa and the secondary veins or only the costa prominent abaxially, the tertiary veins somewhat raised or n aminas abaxially very sparsely sericeous aie a ey widely scattered, ferrugine- ous, appressed hairs (the hairs sometimes faded to white or yellowish in age), only the costa prominent abaxially, the secondary veins at most very slightly raised; Venezuela (Amazonas). 12. P. flavescens. . Laminas abaxially aie sericeous, the hairs white to yellowish (letrugineous in P. chile the costa and the secondary veins prominent abaxia 10.Laminas abaxially cies sericeous with Hae ie hairs (in ae — 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) \O 13. P. ultramontana. 10. Laminas abaxially sparsely to isis sericeous with white to yellowish hairs, the hairs terete; stipules 2. m long, 2.5-3.8 mm wide. 11. Laminas abaxially ne sericeous, the hairs closely spaced and overlap- ping, each hair not readily discernible; pedicels 0.6—1 mm in diameter, the pubescence golden-yellow 12. Filaments adaxially hirsute; limb of lateral petals 3.5-4.3 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 e ere discernible, not at all to slightly touching and aera ae 0.3-0.4 (-0.5) mm in diameter, the pubescence white to 13, ca lanceolate to narrowly elliptical, the nee 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 acute to acuminate with a tip to 1.8 cm long; Brazil (Amazonas, Ron- donia). 10. P. evansii. eee 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. a ioe) 1. Pterandra pyroidea Adr. Jussieu in Saint-Hilaire, Fl. bras. merid. 3: 74. 1833 “1832”].—Type: Brazit. Minas Gerais: “in campis,” Saint- ae B(1) 1376 (holotype: P!; isotype: P!). Pterandra psidiifolia Adr. Jussieu in Saint-Hilaire, Fl. bras. merid. 3: 73. 1833 [“1832”]. Prerandra pyroidea f. psidiifolia (Adr. Jussieu) Niedenzu, Arbei- ten Bot. Inst. Kénigl. Lyceums Hosianum Braunsberg 5: 33. 1914.—Typr: BraziL. Minas Gerais: “in monte Serra da Canastra,” Saint-Hilaire C(/) 395 (holotype: P!; isotype: P!). 1997 C. ANDERSON: 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 slightly 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, limb 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, Goids, and Minas Gerais; one collection purportedly from Mato Grosso); cerrado, campo sujo, and campo; 650- Om. REPRESENTATIVE SPECIMENS. Brazil. DistRITO FEDERAL: Brasilia, bacia do Rio Sao Bartolomeu Rebrotamento, 14 May 1980, Heringer et al. 4771 (IBGE, MICH); Chapada da Contagem, ca. 15 km E of Brasilia, 700-1000 m, 18 Aug 1964, /rwin & 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); Cérrego Jeriva,, ca. 10 km E of Brasilia, 975 m, 15 Sep 1965, /rwin et al. 8342 (NY, UB); near Cérrego Taquari, E of Lagéa Paranod, 975 m, 28 Apr 1966, Irwin et al. 15404 (NY); 30 km N of Brasilia, Cérrego Sobradinho, Farm No. 19, 4 Oct 1963, Maguire et al. 57015 (MICH, MO, NY).— Golds: Serra Geral do Parana, 3 km by rd § of Sao Joao da Alianga, 1040 m, 23 Mar 1973, Anderson 7750 (NY, UB); ca. 15 km (straight line) N of Corumbé, 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 Caiap6, ca. 35 km S of Caiap6nia on rd to Jataf, 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 hare 15 Jul 1969, Saddi RFAI2314 (MICH).—Mimnas GERAIS: 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, oo 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, Bales etal. 24857 (F, MBM, MICH, RB, UB); Araxd, Rod. BR-262, 12 a 1982, Hatschbach 45623 (MBM, ICH); 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). Pterandra pyroidea is a common plant of the Brazilian Planalto, where it forms little-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 RFA12314) 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 unlikely 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. 993.—TyPe: Brazi_. 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 mm long, pellucid cells along margin of abaxial surface or absent, costa and secondary veins prominent abaxially, tertiary veins usually slightly raised as well: petioles |- 2.5 mm long, densely sericeous; each pair of stipules entirely connate or near ly 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 I1—1.8 mm long, 0 644 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—) 55.5 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 1997 C. ANDERSON: PTERANDRA 1] FIG. 3. Pterandra hatschbachii. a. Habit, x0.5. b. Portion of stem with leaf bases and les x5. c. Base of flower cluster with bracts and bracteoles, the entire cluster subtended by a ay bract (composed of a cee leaf and stipules), x5. d. Flower, posterior petal at upper left, x2.5. e. Posterior petal, abaxial view rae sparse pubescence on costa, x3.5. f. Distal portions ve stamens, adaxial (left) and abaxial (right) views, x7.5. g. Distal portion of stamen, lateral view, x7.5. (Based on: .d-g, Hatschbach 35085; c, cee 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. i Brazil (Matro Grosso); forming dense carpets in grassy campos; ca. 800 m. ADDITIONAL SPECIMENS EXAMINED. Brazil. Mato Grosso: Mpio. Alto Araguaia, vicinity of Ribe- rao Claro, NW of Alto Araguaia, ca. 800 m, 15 Feb 1975, Anderson 11400 (MBM, MICH): Mpio. Alto Araguaia, Alto Araguaia, 15 Nov 1973, Hatschbach 33276 (MBM, NY) IZ CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Pterandra hatschbachii is known only from the vicinity of the type locality in Mato Grosso, close to the border with Goids. It shares many characters with P. pyroidea, acommon species of the cerrado and campo of Goids 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- bachit 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.—T ype: BRaAziL. Maranhao: Grajau, 140 m, 3 Aug 1909, Lisbéa 2507 (holotype: 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 ellipticae, 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 subsessilibus constans, floribus cujusque fasciculi 4-6, sine foliis 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. Styli 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 FIG. 4. Pterandra andersonii, P. colombiana, P. hirsuta, and P. mcephersonii. a-e, P. andersonii. a. Terminal portion of shoot with four leaves, x0.5. b. Terminal portion of shoot with two inflores- cences, x0.5. c. Petal, abaxial view, x5. d. Dist tal portions of stamens, adaxial (left) and lateral (right) views, 7.5. e. Coccus, lateral view, x5. f-i, P. colombiana. f. Terminal portion of shoot with infructes- cence, x0.5. g. Leaf, x0.5. h. Base of leaf with cia oa (basally connate but free in the distal 1/2), adaxial view, x1.5. i. Petal, abaxial view, x5. , P. hirsuta. }. Leaf, x0.5. k. Detail showing abaxial laminar ean composed of barined es x10. 1. Base of leaf with connate stipules, adaxial etal, abaxial view, x5. n—p, P. oe n. Terminal aries of shoot with one leaf and old Aa (only one fascicle remaining), x0 . Fascicle of flowers, borne on a short a a x2.5. p. Petal, pal view, x5. (Based on: a— os heban 200756: Poker 12; f-1, Callejas et . 4656; j-l, Moraes M. & Takana G. 1857; m, Anderson 12298; n—p, McPherson 11745; drawn by Karin Douthit, ) mn 1997 C. ANDERSON: PTERANDRA 13 14 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 with the apex acute, abaxially 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 ligulate), 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, abaxially 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, limb 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. | mm high; mature seed not seen. Chromosome number unknown. ADDITIONAL SPECIMENS EXAMINED. Brazil. MARANHAO: Imperatriz, proximo do campo de Avia- cao, 31 Jul 1976, Pinheiro 12 (MICH). 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 abaxially sericeous rather than tomentulose, and it has smaller stipules and smaller flowers. Prerandra pyroidea and the similar P. hatschbachii have pink petals. Those of P. andersonii are noted on the label as ‘“amarella” (Lisbéa 2507) and “avermelhada” (Pinheiro 12; in 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), sér. 2, 4: 736. 1932.— Type: Brazit. 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, abaxially sparsely sericeous (densely so on 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 abundant in abaxial surface to absent, costa and secondary veins prominent abaxially, tertiary veins usually slightly raised as well; petioles 0.8-2 cm long, densely sericeous; 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, abaxially glabrous but with a sericeous band at the base. Inflorescence of 4—6-flowered fascicles, sessile or borne on a projection up to 1.4 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 0.8—2.2 cm long, 0.4-0.6 mm in 1997 C. ANDERSON: PTERANDRA Abs) 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 limb 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, limb (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 Pree 70-300 m, 4 Nov 1954, Cowan 38123 (MICH, NY); Mazagao, area do Felipe VI, 26 Aug 1985, N. T. Silva 5524 (MG).—Amazonas: Reserva Florestal Ducke, Manaus-Itacoatiara, Km 26, 02°53’ S, pene 14 Aug 1996, Assungdo 365 (MICH); Dist. ee ee eae 1501 (Km 41) of the WWF, INPA MCS Project, 02°24'-02°25'S, 59°43'-59° . 50-125 m, 25 Nov 1988, Boom et al. 8649 (F, NY): Manaus, margem do igarap6 do oe ‘ al eee Coélho INPA 4009 (IAN, INPA); estrada Manaus— Caracaraf, Km 58, Reserva Biol6gica INPA-SUFRAMA, Sep 1976, Coélho & Damido 855 (INPA); Reserva Florestal Ducke, Manaus— eee — 262022538: 99°58 We 2 elu ood. ee 1468 (MICH); Dist. Agropecudrio, 90 km NNE de M , Reserva 1501 (Km 41), Projeto Dinamica Biol6gica de Fragmentos eigcectaia ‘gente S, 30°43 we W, 50-125 m, 11 Aug 1991, Oliveira et al. 144 (NY); Dist. Agropecudrio, 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 on 49 (MBM, NY); Reserva Florestal Ducke, Km 26, i Aug 1 ieee Prance et al. 2122 (F, MG, MICH, NY, R, US); estrada Manaus— Caracarai, Km 39, Reserva Experimental de Silvicultura Tropical sae 14 Sep 1977, Ramos 221 (INPA): Reserva Florestal Ducke, Manaus-—Itacoatiara, Km 26, 02°53'S, 59°58'W, 21 Jul 1994, Ribeiro 1346 (MICH), Reserva Florestal Ducke (P-69), 21 Aug 1963, Rodrig 5444 (F); Manaus, estrada Manaus-Itacoatiara, Km 160, 15 Oct 1965, Rodrigues 7244 (INPA); Manaus, estrada Manaus-Ita- coatiara, Km 125, 24 Sep 1965, Rodrigues 7577 (INPA); ee Florestal Ducke, perto da estagao metereoldégica, 9 oe 1964, Rodrigues 32 (F); Reserva cums Ducke, 21 Jul 1964, Rodrigues & Loureiro 5961 (F, US); Manaus, estrada Manaus-Itacoatiara, Km 73, 30 Aug 1965, Rodrigues & Loureiro 7058 (INPA); between Missao Salesiana and esi tet Rio Maturaca, 400-800 m, 13 Jan 1966, N. T. Silva & Brazdo 60825 (F, MG, MICH, NY).—ParA: Estacdo Ecologica do ieee Projeto Reserva Genetica, SEMA, 00°75'S, 52°30'W, 14 Oct 1987, Beck 98 (NY Pterandra arborea is a common tree in the area of Manaus and thus more frequently collected than other species, except P. pyroidea. 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, Tillett et al. 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. | 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 44.3 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.—T ype: Cotompia. Antioquia: Zara- goza, Cor to 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-1. Arbor usque ad 22 m alta. Laminae 8-19.5 cm longae, 2.7-8 cm latae, obova- tae vel ellipticae, supra glabrae, subtus sparsissime sericeae vel glabratae; petioli 0.8-3.5 cm longi, juniores dense sericei, vetustiores glabrati; stipulae 2.8-4 mm longae, 3-4 mm latae, connatae sed distaliter liberae, 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, 3.2-4.5 mm longo, 2.8-4 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, linear, 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 ligulate), 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 E] 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 Samané—-Rio Claro, puente ae 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, ee os (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 Prerandra 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 mephersonii C. Anderson, sp. nov.—Typre: PANAMA. Colon, Santa Rita ridge, SE of Colén, 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 sessilibus vel brevistipitatis constans; floribus cujusque fasciculi 4 (—6?), sine foliis juvenilibus per anthesin: pedicelli 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-) 4.24.8 mm lati, limbus petali 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 ignot. | 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 slightly 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 ligulate), apex obtuse, recurved, glands 2.2—2.3 mm long, ca. | 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. Prerandra 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"]—Tyee: 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”].—TypeE: GuyAna. Upper Mazaruni River, small island across river from Isla Casa- be, 175 m, 9 Dec 1951, Maguire 32715 (holotype: MICH!; isotypes: F! K! NY! US!). Pigs. Shrubs or trees to 15 m. Laminas 3-8.5 cm long, 1.6—4.3 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, 20 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 FIG. 5. Prerandra sericea. a. Flowering branch. b. Portion of stem with leaf bases and stipules. c. Hairs from abaxial surface of lamina. d. Fascicle of flowers in leaf axil (all but one flower removed). e. Three stamens, stamen at left and right each opposing a sepal, stamen in center opposing a petal. f. Immature fruit (one carpel aborted). (Based on: a-d, f, Maguire 32715; e, Wurdack 34405; drawn by Annette Seidenschnur Mahler.) 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, 3-4 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 ol 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). Wenezuela (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); Rio 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 (MQ); Aripao, Rio Caura, aguas abajo de] 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 Abacapa 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.—T ype: BRAziL. Amaz6- s: Rio Marmelos (headwaters), Aug 1948, Schultes & Lopez 10332 (holo- ae 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, 2.8-4 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, limb 4-5 mm long, 3-3.5 mm wide; posterior petal: claw 1.5—-1.8 mm long, limb 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, Rond6nia); in “campo natural” and forest, along rivers. ADDITIONAL SPECIMENS EXAMINED. Brazil. AMAZONAS: Novo Aripuana, BR-230, 150 km ao N de Humaita e 30 km para o § na rod. do Estanho, 16 Apr 1985, 08°20'S, 61°45'W, Cid Ferreira 5596 p.p. (INPA, MG); Novo Aripuana, pa Proj. RADAM SC-20-XD-Ponto 22, margen do Rio Preto, 30 Aug 1975, Cordeiro 704 (MG, MICH); Novo Aripuana, Igarap6 Preto, SC-20-XD-Ponto 30, 30 Jun 1975, da Silva 109 (MG).—Ronbonla: 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. hirsuta but differs most notably in the abaxial laminar pubescence. In P. hirsuta that vesture is composed of erect basi- fixed hairs mixed — a few medifixed hairs. In P. evansii the laminas are abaxially sparsely sericeous, 1.e., the hairs are appressed and medifixed; however, they usu- ally also have tufts of Gasiaxed 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.—Typr: Bo.ivia. Beni: Prov. Vaca Diez, 19.4 km al N de Guayaramerin camino a Cachuela Esperanza, 10°33'S, 65°38'W, 140 m, 18 Sep 1994, Moraes M. & Takanad 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, adaxialiter hirsutae, abaxialiter sericeae. Inflorescentia ex fasciculis sessilibus constans, floribus cu- jusque fasciculi 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, limbo 1997 C. ANDERSON: PTERANDRA 23 elliptico vel late obovato; limbi petalorum lateralium 4.2-5 mm longi, 3-4 mm lati, limbus petali postici 5—S.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, 0.50.8 (-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 (Amaz6nas, Rond6énia) and adjacent Bolivia (Beni); in forest at riverside and in “pampa-monte”; 140-220 m. ADDITIONAL SPECIMENS EXAMINED. Bolivia. BENI: Prov. Vaca Diez, Guayaramerin hacia Riberalta, 23 Sep 1993, de Michel & Beck 218] (MICH). Brazil. AMAzOnas: Novo Aripuana, BR-230, 150 km ao L de Humaita e 30 km para o S na rod. do Estanho, 08°20'S, 61°45'W, 16 Apr 1985, Cid Ferreira 5596 p.p. (MG, MICH).—Ronponia: along Rio dos Pacadés Novos, just above and below the first cachoeira, ca. 220 m, 28 Mar 1978, Anderson 12298 (INPA, MICH). Pterandra 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. Gard. 8: 128. 1953.— Type: VENEZUELA. Amazonas: Cerro Sipapo (Paraque), banks of Lower Cafio 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 slightly 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 limb; 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 Cafio on the Cerro Sipapo; in savanna and marshes along the river; 1500 m. ADDITIONAL SPECIMENS EXAMINED. Venezuela. AMAZonas: Cerro Sipapo (Pardque), Cafio Negro. 1500 m, 15 Dec 1948, Maguire & Politi 27692 (NY, US), 27692A (NY), 25 Dec 1948, 27946 (F, GH, MO, NY). 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 leaf. 13. Pterandra ultramontana Riley ex Cuatrecasas, Webbia 13: 557. 1958.—T ype: Co.omsiA. Narifio: 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, limb 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 a 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, Concesi6n 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: Braziv. Para, Alto Tapajos, Rio Cururt, 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 | 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 45.2 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. | 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 Missdo Velha, ca. 2 km N of Rio Cururt, 07°45'S, 57°20'W, ca. 200 m, 13 Feb 1974, Anderson 10895 (LAN, 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. Prerandra pyroidea, of Goids 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. Prerandra 1997 C. ANDERSON: PTERANDRA Dy hirsuta 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. ExcLUDED NAMES Pterandra sect. Acmanthera Adr. Jussieu, Ann. Sci. Nat. Bot., sér. 2, 13: 328. 1840. = Acmanthera (Adr. Jussieu) Grisebach in Martius, Fl. bras. 12(1): 29. 1858 Pterandra latifolia Adr. Jussieu in Delessert, Icon. selec. 3: 19. 1838 [*1837"]. = Acmanthera latifolia (Adr. Jussieu) Grisebach in Martius, FI. bras. 12(1): 29. 1858 Pterandra opulifolia Rusby, Descr. S. Amer. pl. 38. 1920. = Hiraea opulifolia Rusby) Niedenzu in Engler, Pflanzenreich IV. 141(1): 144. 1928. ACKNOWLEDGMENTS I thank William R. Anderson for his comments and for sharing his insights on Pterandra. Karin Douthit drew Figs. 3 and 4, and Annette Seidenschnur Mahler drew Fig. 5. Figure 5 is reprinted with permission from Memoirs of the New York Botanical Garden 32: 34, Fig. 8, copyright 1981, The New York Botanical Garden. I am grateful to the et of the following herbaria for permitting me to study their collections: A, BR, F, G, GH, GOET, HB, HUA, IAN, IBGE, INPA, K, MBM, MEDEL, MG, MICH, MO, NY, P, R, RB, UB, US, VEN, W LITERATURE CITED Anderson, W. R. 1975. The taxonomy of Acmanthera (Malpighiaceae). Contr. Univ. Michigan Herb. 11: 41-50. . 1978 [“1977”]. Byrsonimoideae, a new subfamily of the Malpighiaceae. Leandra 7: 5-18. . 1981. Malpighiaceae. In “The botany of the Guayana Highland—Part XI” by Bassett Maguire and 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. 19: 41-354. Cuatrecasas, J. 1958. Prima Flora Colombiana. 2. eee ee Webbia 13: 343-664. Grisebach, A. 1858. Malpighiaceae. In Flora brasiliensis by K. F. P. von Martius, 12(1): 1-12 Jussieu, Adr. de. 1833 - 1832”]. Malpighiaceae. In Flora ee meridionalis by A. a Saint- Hilaire, 3: 5-86, t. 161-180. Paris: A. Belin . 1838 [1837 “ ete latifolia. In [eaten selectae plantarum, ed. B. Delessert, 3: 19. Swofford, D. 1993. PAUP, phylogentic analysis using parsimony, version 3.1.1. Center for Biodiversity, Illinois er History Survey. Contr. Univ. Michigan Herb. 21: 29-36. 1997. 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 IJ 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. Excentradenia W. R. Anderson, gen. nov.—Tyee: Excentradenia adenophora (Sand- with) W. R. Anderson. Lianae lignosae. Lamina foliorum eglandulosa vel margine glandulis parvis instructa, venis tertiarus 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 fertilia. Styli 3, stigmate introrso, apice dorsaliter 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 | 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 | 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 F. 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 1s 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 circular 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: . Stipules very small, triangular, borne on petiole at very base or on adjacent stem, or absent; inflorescence an axillary raceme of 3-7 (-9) 4-flowered umbels; one of each pair of bracteoles bearing a large bulging eccentric abaxial gland toward center of umbel; pedicel circinate in bud; lateral wing of samara incised to nut at apex but not at base, producing a single basally continuous wing, or (in £. adenophora) sometimes incised to nut at both apex and base. Excentradenia. — . Stipules elongated, borne on petiole at least somewhat above base; inflorescence an axillary cyme of 3-7 4-flowered umbels or a single umbel of 5—many flowers; bracteoles eglandular; pedicel straight in bud; lateral wing of samara incised to nut at both apex and base, producing 2 completely separate wings. Hiraea. The eccentric 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. | 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 (EF. 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, £. 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. 1997 ANDERSON: EXCENTRADENIA ail 80° TAOhe 60° 50° Bi 10° 10° ae . © ve °o 0° ey Excentradenia @ adenophora @ boliviana M@ primaeva © propinqua 105 10° 80° 70° 60° 50° FIG. 1. Distribution of Excentradenia. KEY TO THE SPECIES OF EXCENTRADENIA |. Leaves abaxially persistently velutinous, the hairs erect and bifurc FE. primaeva. 1. Leaves abaxially sericeous to glabrate, the hairs sessile and tightly ee 2. Leaves abaxially densely and persistently sericeous, even in age. FE. propinqua. 2. Leaves glabrate at maturity, with the abaxial midrib and often the lateral veins + persis- tently sericeous and otherwise only sparse, scattered hairs persistent on the lamin 3. Calyx bearing 8 glands on the 4 lateral sepals; small stipules present on base of petiole: lamina of larger leaves 8-15.5 cm long, the petiole 10-20 mm long; lateral eae of samara with the sides not meeting at apex, leaving an evident gap. E. adenophora. 3. Calyx completely eglandular; stipules absent from vegetative leaves; lamina Ot larger leaves 14-19 cm long, the petiole 20-29 mm long; lateral wing of samara with a sides meeting or overlapping at apex, leaving little or no evident gap. . 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 Vig K), photos MICH, WRA negs. 81-5-17 [sheet I] & 18 [sheet II]). Pel, 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 a2 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 FIG. 2. Excentradenia primaeva and E. adenophora. a-g, E. primaeva: a) leat and old infructes- cence, x().5; b) stipules, xl; c) detail of adaxial surface of lamina, minus hairs, x0.5; d) detail of abaxial surface of lamina, minus hairs, x0.5; e) hair from abaxial surface of lamina, x0.5; f) bracts and bracteoles of umbel, with eccentric glands on 4 bracteoles, x7.5; g) samara, abaxial view, x0.5. h-n, E adenophora: h) large leaf, adaxial view, x0.5, and base of petiole with stipules, x2.5; i) flowering branch, x0.5;}) umbel of 4 circinate buds, x2.5; k) base of umbel enlarged to show eccentric glands on 4 bracteoles, x5: 1) flower from above, with posterior petal uppermost, x2.5 (in nature the stamens would be less spreading, probably nearly erect); m) anthers, adaxial view (left) and side view (right), x10; n) distal portion of style, side view, x10. Drawn by Karin Douthit, a-g from Nascimento et al. 135, h-n from Fanshawe in Forest Dept. 5604, the drawings of E. primaeva were originally published as part of Fig. 50, Mem. New York Bot. Gard. 32: 241. 1981, under the basionym, Hiraea primaeva. 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 ong, 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 EXAMINED: Guyana. Bartica—Potaro Road, 51st mile, wallaba bush on white sand, fl [without date], Dawson in ey 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 ae ‘pepe 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 & Gonzdlez 5954 (MO, VEN).—Detta Amacuro: E side of Rio Cuyubini, Cerro La Paloma, Sierra Imataca, vicinity of large granitic boulders, 100-200 m, Nov fr, Steyermark 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 fruits in November. I 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.—T ype: Bo.ivia. 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 laterales; 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. tems 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 sericeous. ADDITIONAL SPECIMEN EXAMINED: Bolivia. BENi: Cachuela Esperanza, Rio Beni, Nov 1923 fr. Meyer 378 (NY). 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 £. 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 £. propinqua but not EF. adenophora. When E. boliviana is collected with flowers, those may strengthen the morphological separation of these transamazonian sisters. Excentradenia primaeva (W. R. Anderson) W. R. Anderson, comb. nov. Hiraea primaeva W. R. Anderson, Mem. New York Bot. Gard. 32: 240. 1981.— 1997 ANDERSON: EXCENTRADENIA 35 Type: BRAZIL. Amazonas: S. Gabriel, QO. NA-19-ZC, 12 Apr 1975 fr, Nasci- mento, Pires & 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 limb 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.—Typr: 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 at 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—l 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 lon slender divisions somewhat glandular-thickened at apex. Stamens with filaments 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 fine 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. 5I (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 I thank the curators of the following herbaria for sending MICH gifts and loans that made this study possible: CAY, IAN, K, MO, NY, U, US, and VEN. Karin Douthit drew the two species shown in Fig. 2, more than 15 years apart in time but in the same beautiful style. LITERATURE CITED Anderson, W. R. 1994. ow species of Hiraea (Malpighiaceae) from the Guianas and adjacent Brazil. Brittonia 46: 126-13 Contr. Univ. 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 IJ 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” species. Bunchosia cauliflora W. R. Anderson, sp. nov.—Type: Ecuapor. Pastaza: Rio Pastaza, between Destacamento Chiriboga and Apachi Entza, 2°20-32'S, 76°55'-77°08'W, 24 Jul 1980 fr, B. Mllgaard, 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 0.51.3 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 !-1.5 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 peltate 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 Catoélica station, 0°41'S, 76°28'W, Acevedo-Rodriguez & Cedefo 7555 (US): Reserva Etnica Huaorani, S of Rio Tivacuno, 0°50'S, 76°14'W, Aulestia & Ima 1583 (MICH) & between Rio Tivacuno and Rio Yasunt, 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°31'W, Pitman & ee 369 (MICH).—Pastaza: Rio Papayacu at Rio Curaray, 1°29'S, 76°42'W, Holm-Nielsen et al. 22567 (AAU).—Sucumpios: tea ufindi (Nueva Loja), a (Puerto Francisco de Orellana)—Lago en 50 km NE of Coca, Harling & Andersson 12004 (MICH): Lago Agnio, Lugo S. 3/52 (MICH): Rio Aguarico, 5 km § of Lago nee Lugo S. 3184 (MICH); Las Sachas, ane 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. pseudonitida Cuatrec., which O97. 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 Inajé, Reserva Biol6gica de Serra Negra, ascent to the forest, 8 Mar 1995 fl, E. Menezes, E. M. Villarouco, S. S. Lira & 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 lateralia 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; styli 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 | (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), 40 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 ovoid, 3-lobed, glabrous, smooth (i.e., not granulate) with the reticulum of veins prominent in dried fruit. ADDITIONAL SPECIMENS EXAMINED. Brazil. PERNAMBUCO: Buique, estrada Buique—Catimbau, 8°37'S, 37°10'W, 790 m, shrubby vegetation on sandy soil, May fr, Laurénio et al. 40 (MICH); type locality, open area, Mar fl, Tscha et al. 12 (MICH) & inside the forest, Feb fr, Tschd et al. 15 (MICH Bunchosia pernambucana is a new, northern addition to the Bunchosia mariti- ma complex, whose other members (B. acuminata Dobson, B. macilenta Dobson, and B. maritima (Vell.) 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, unlike 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, 7. 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, liberi, 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 1997 W.R. ANDERSON: MALPIGHIACEAE 4] O B. macilenta @ B. maritima m B. pernambucana 45° 40° 35° FIG. 1. Distribution of the Bunchosia maritima complex. densely sericeous, adaxially glabrous; peduncle 1.5-2.5 mm long, thinly hispid like inflorescence axis; bracteoles apical or slightly 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 42 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 FIG. 2. Leaf 7 urs of the reeks mollis complex. a, b, B. mollis (Gentry & Berry 14926): ae) whole hae. a x x96. c, d, B. thaumatothrix (Morillo P Manara 1172): c, whole hair (note x rudiment of missing branch sae base 2; d, base of a hair enlarged to show rounded basal projections, x320. e-g, B. petraea Plowman & Gudnehes 13762): e & f, whole hairs, x72; g, base of a hair enlarged to ne rudimentary basal projections, 320. All vouchers at MICH; SEM photos by Beverly Walte 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 limb 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 significantly, 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 Rincén—Las Tunitas, SE de Maiquetia, vertiente norte de la Cordillera de La Costa, bosque trop6filo, 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 44 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., 1-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 23.5 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 light 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, loculis apice extensionibus sterilibus 0.5-1 mm longis ornatis, connectivo partem fertilem loculorum ca 0.5 mm superanti. Ovarium 2 carpellis 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 | 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 tomentose, adaxially glabrous, strongly revolute in the distal half in anthesis, accrescent to ca 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. ADDITIONAL SPECIMENS EXAMINED: Wenezuela. AMAZONAS: Selvas pluviales del Medio Rio Emoni, 1/2 dia en bongo abajo del Cafio Bocén, 2°10'N, 66°17'W, 150 m, Jan fr, Stergios et al. 9955 (MICH, 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 | mm. Byrsonima formosa W. R. Anderson, sp. nov.—Typr: 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 distali 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 | 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 |-1.2 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. ADDITIONAL SPECIMENS EXAMINED: Guyana. Gunn's, Essequibo River, 1°39'N, 58°38'W, savanna, 240-260 m, Sep fr, Jansen-Jacobs et al. 1890 (MICH). Brazil. AMAzOnaAs: Mun. Presidente Figueiredo, Rio Uatuma, canteiro de obras da Usina Hidrelétrica de Balbina, 1°30'-2°00'S, 59°30'-60°00'W, campina aberta, solo arenoso branco, Jul fl/imm fr, Cid Ferreira 7559 (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 (IAN, UB). 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: 48 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 80° 70° 60° 50° 40° FIG. 3. Distribution of selected species of Byrsonima series Platylepts. 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. L. Inflorescence axis glabrous, with flowers mostly restricted to the distal 1/2; pedicels (12-) 1 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.—T ype: VENEZUELA. Amazonas: Depar- tamento Atures; Sierra Maigualida, NW sector, 5°30'N, 65°15'W, small valley along an upper tributary of Cafio Iguana, 2000 m, 28 Feb-3 Mar 1991 imm fl, P. E. Berry, O. Huber & J. Rosales 4529 (holotype: MICH!:; isotypes: MO! MYF!). Arbor 3 m alta, ramis pertinaciter tomentosis. Lamina foliorum majorum 5S— 6.8 cm longa, 2.6—3.8 cm lata, elliptica vel paulo ovata, basi cuneata vel rotundata, 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 mm diametro. 1997, W. R. ANDERSON: MALPIGHIACEAE 49 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 | 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- lally 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. huberi will certainly correct and complement my very tentative de- scription. 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 era on) CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 suite of characteristics, one most cross the Amazon basin and go to Minas Gerais in the Planalto of Brazil. There Byrsonima macrophylla (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 | 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, S—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: Brazit. 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 explicitly 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 oy 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 bracteoles 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. . 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 bracteoles 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 B. 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. K6nigl. 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.—TypeE: BRAZIL. Pernambuco: Brejo da Madre de Deus, Bituri, roadside, 4 Feb 1995 fl, M. J. N. Rodal & M. F. Sales 459 (holotype: MICH!; isotype: NY!). Arbor 7 m alta. Lamina foliorum 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, 2 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 proximally sericeous and somewhat glabrescent, distally (among flowers and on peduncles and pedicels) persistently tomentose; bracts 4-5 mm long, I-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 biglandular, 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 limb 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- sis. Fruit unknown. 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.—T ype: BRazi_. Rond6énia (“Mato Grosso”): 66 km W of Vilhena, Brasilia-Acre Highway, forest along iga- rapé, 3 Sep 1963 fl, B. Maguire, J. Murca 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, connectivo 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 | 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, 3.5-4 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 S 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 Solim6es, Lago de Alvarades, 3°13'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 Humaité—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 Rondé6nia state line on BR364 from Vilhena to Caceres, 13°22'S, 59°56'W, Thomas et al. 4769 (MICH).—Ronponia: 2-4 km E of Mutumparana on road to Pérto 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: 54 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 1. Shrubs or small trees 1.5—5 (—7) m tall; petioles 2-5 mm long. B. melanocarpa. 1. Trees (4—) 8-25 m tall; petioles 12-25 mm long 2. Lamina mostly acuminate at apex, occasionally eee supules 1-1.5 mm long; bracts 1-1.5 mm long, triangular, often revolute; flowers 1-2 per bract; sepals revolute in anthesis, accrescent in fruit, to 2.5—3 mm long; anther (osiles not or hardly winged, the wing up to 0.1 mm wide. B. eee ii) . Lamina broadly obtuse to rounded at apex; stipules 2.3-5 mm long; bracts 0.5—0.7 mm long, broadly obtuse or rounded, erect; flowers never more than | per bract; sepals ap- pressed in anthesis, not or hardly accrescent in fruit, to 1.5 mm long; anther locules bearing a well-developed wing 0.2—0.4 mm wide. B. piresi. Dicella aciculifera W. R. Anderson, sp. nov.—Typer: 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 sericcous 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 | 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 limb 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 1997 W.R. ANDERSON: MALPIGHIACEAE 5D FIG. 4. Dicella aciculifera. a) flowering branch, x0.5, with enlargement of edge of abaxial surface of lamina, x10; b) flower bud with pedicel, bracteoles, sates and subtending bract, x2.5; c) flower from above, with ae: (flag) petal uppermost, x1. side and adaxial views of anthers opposite petals, x5; e) side and adaxial views of anthers ono sepals, x5; f) Ee x5; g) style-apex, x15; h) fruit Pi above, with anterior sepal pointing downward, x0.5. Drawn by Karin Douthit, a—-g from Angulo 716, h from Angulo 594. 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 3.44 cm long and 1.6-2 cm wide. ADDITIONAL SPECIMEN EXAMINED. Costa Rica. PUNTARENAS: Cantén de Osa, Peninsula de Osa, R. F. Golfo Dulce, Cerro de Oro, alrededor del Albergue “Unioro,” Sendero Termo, 8°39'20"N, §3°26'30" W, 270 m, ripario, Mar fl, Angulo 116 (MICH). Dicella aciculifera is assignable to sect. Macropterys Chase (Chase, 1981), and resembles, in both leaves and fruits, its geographically closest neighbor, D. judianti (J. F. Macbr.) W. R. Anderson. Dicella julianii 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 Sy species of the genus, as shown in the figures cited above, the styles are erect and parallel, not at all 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 S—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., Sér. 2, 13: 274. 1840. Banisteria cotinifolia (Adr. Juss.) C. B. Rob. ex Small, N. Amer. FI. 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., Sér. 2, 13: 274. 1840. Banisteria gayana (Adr. Juss.) C. B. Rob. ex Small, N. Amer. FI. 25: 135. 1910.—Typr. 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. FI. 25: 135. 1910.—Type: Mexico. Jalisco: Barranca near Guadalajara, E. Palmer 712 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 Culiacan, 7. 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) Standl., Contr. U.S. Natl. Herb. 23: 577. 1923.—Type: Mexico. Oaxaca: San Geronimo, Jul 1914 fl/fr, C. A. Purpus 7262 (holo- type: UC!; 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 hoffmanii W. R. Anderson, sp. nov.—Type: GuyANna. 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!). Liana ramis tomentosis demum glabrescentibus. Lamina follorum majorum 6-9 cm longa, 4.5-6 cm lata, ovata, basi cordata, margine ciliata vel glandulis sessilibus instructa, utrinque tomentosa pilis tortuosis demum glabrata vel costa pertinaciter tomentosa; petiolus 3-5.5 mm longus, basi glandulifer; stipulae non visae. Inflorescentia floribus in umbellis 4-floris sessilibus vel subsessilibus dis- positis; pedunculus nullus; 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 truncati vel brevissime apiculatl. 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-S.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 /ato 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 | 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, limb 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- aliform,” 1.e., with a flattened elliptical 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. L997 W.R. ANDERSON: MALPIGHIACEAE 59 (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. hoffmanti would distinguish it at once from its sister species, even if it were not geographically disjunct. Heteropterys molesta W. R. Anderson, sp. nov.—TyPeE: 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 Félix), 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 (-4) mm longus; bracteolae apicales, 1 bracteola cujusque paris 1 glandula abaxiali excentrica ().7—0.8 mm diametro instructa; pedicellus 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 | 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 | 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 I 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 tor 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. Herteropterys 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. | 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 Heteropterys! 1997 W. R. ANDERSON: MALPIGHIACEAE 61 Heteropterys sylvatica Adr. Juss., Ann. Sci. Nat. Bot., Sér. 2, 13: 277. 1840.— Type: Bottvia. 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.—Tyre: BrRa- 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 OrAdn, Salta. Heteropterys velutina W. R. Anderson, sp. nov.—T ype: 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 9,5-11.7 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 slightly 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 like 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: floriferous 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 limb 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 clongated pseudoracemes rather than umbels. Its closest relative is probably H. subhelicina Nied. (=H. 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. subhelicina 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- licina 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 17. velutina 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 fi/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 umbellis 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 | or 2 small glands, the lateral 4 biglandular, the glands 0.9-1.5 mm long, circular or elliptical, separated from each other, slightly 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—4.5 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 like 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).—Zutia: Distr. Miranda-Bolivar, via El Consejo, La Tabla—Quirdés—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.—T ype: 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, J. 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— 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, elliptical, 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).—SantTanper: Mpio Charala, Corrig. Virolin, “Cafiaverales,” liana in tree at edge of pasture, 1680 m, Jan fr, A/bert de Escobar et al. 3060 (MICH). Venezuela. TAcuira: Distr. Uribante, Complejo Hidroelectrico Uribante-Caparo, Sector El Cedral, inmediaciones Presa La Honda, 8°01'N, 71°37'W, low montane forest, 1100-1250 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 has 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.—T ype: VENEZUELA. Amazonas: Depto. Rio Negro, Rio Baria (=Rio Mawarinuma) just upstream from Base Camp, SW side of Cerro de la Neblina, 0°49'S0"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 foliorum 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 petioli portatae. Cyma axillaris plerumque ex 3 umbellis 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 styli postici apice dorsaliter breviunci- nati unco ca 0.3 mm longo rotundato. Samara immatura alis lateralibus semicircu- laribus, ala dorsali 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 limb 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: Ecuapor. Napo: Canton Tena, Estacién Biolégica Jatun Sacha, Rio Napo, 8 km al E de Misahualli, 1°04'S, 77°36'W, 400 m, bosque muy himedo tropical, bosque primario, colinas, 3-6 Sep 1989 fl, W. Palacios 4384 (holotype: MICH!). Liana lignosa ramis velutinis. Lamina foliorum 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 umbellis 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 slightly 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-like 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, axillary, 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 | mm wide, ovate or elliptical, abaxially sericeous, adaxially glabrous, eglandular, persistent; bracteoles like bracts but short- er and narrower, narrowly elliptical or linear; 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-S 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 swollen, 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. hypoleuca), 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 1997 W.R. ANDERSON: 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. nov.—Type: VENEZUELA. Bolivar: Reser- va Forestal “La Paragua,” margenes del Rio Asa, Jun 1970 imm fr, C. Blanco 803 (holotype: F!). 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 erose, 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 70 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 they enlarge, the hairs 0.5-1 mm long, medifixed, straight and appressed; dorsal wing 1-3 mm wide, 1.8—2 mm high, triangular to rectangular, mostly dissected into several coarse teeth, sericeous to glabrate. ADDITIONAL SPECIMEN EXAMINED. Venezuela. Botivar: Altiplanicie de Nuria; forest at base, etween Rancho Alegre and base of Altiplanicie, on trail to Quebrada Cabeza Burro, 5 km E of Las Chicharras, 47 km N of Tumeremo, 100-250 m, Feb sterile, Steyermark 89349 (NY). I name this species in honor of my friend, the late Julian 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 1s incomplete and will have to await additional collection for supplementation. Among the species of Hiraea known to occur in Venezuela, H. steyermarkii bears 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 Mig., 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 enuire or subentire. Large glands on the leaf margin are found in two species of French Guiana, /7/. 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. steyermarki. Hiraea longipes also has a long pedicel and long stipules like those of H. steyermarku. 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.—Typre: Ecuapor. Napo: Cant6n Tena, Estacion Bioldégica Jatun Sacha, 8 km al E de Misahualli, 1°04'S, 77°36'W, 400 m, bosque muy humedo tropical, bosque primario cerca al riachuelo Chinquipino, parcela permanente 03, 20 Jan 1990 fr, C. E. Cerén, C. Iguago & E. Saldumbide 8370 (holotype: MICH!), 1997 W. R. ANDERSON: MALPIGHIACEAE aa Liana lignosa ramis sericeis mox vel demum glabratis, quoque nodo cristis interpetiolaribus munito. Lamina foliorum majorum 12-22.5 cm longa, 8-15 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 | 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 alis 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 retuse 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 ys 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 mm 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 humedo tropical, primario, Nov imm fr, Gudifio 194 (MICH).—Pastaza: Pastaza, pozo petrolero Villano 2 de Arco, 1°25'S, 77°20'W, 400 m, bosque htimedo tropical, primario, Dec fl, Hurtado 2820 (MICH); entre los rios Iquino y Villano, 1°29'S, 77°27'W, 350 m, bosque htimedo tropical, primario, Aug fl buds, Tirado etal. 55 (MICH). Peru. AMAzonas: Lugar Aintami, monte, Jan fr, Kayap 152 (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, /. 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 #/. 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 //. 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 1S Mascagnia haenkeana W. R. Anderson, sp. nov.—T ype: Ecuapor. Guayas: Guaya- quil, 7. Haenke 2256 in 1790 (holotype: F!). Liana lignosa, ramis velutinis. Lamina foliorum majorum 7—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 3.5-4.5 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 74 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 flange on underside of lateral wing, extending from apex of nut to apex of lateral wing, ca 0.5 mm wide; ventral areole ovate, 1.5-2 mm long, 1-1.3 mm wide; torus ca 2 mm high, surrounded by a narrow, glabrous, 3-lobed, disciform outgrowth of the receptacle. ADDITIONAL SPECIMEN EXAMINED. Ecuador. Guayas: 14 km ESE of Montero. elev. 10 m, fence shrubbery, Dec fr, MacBryde & Simmons 951 (QCA) Mascagnia haenkeana is named in honor of Thaddacus 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. Ka) 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. Kénigl. Lyceums Hosi- anum Braunsberg 3: 14. 1908. Hiraea lilacina S. Watson, Proc. Amer. Acad. Arts 17: 333. 1882.—Typr: Mexico. Coahuila: Caracol Mountains 21 mi SE of Monclova, Aug fl, FE. 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 —Typr: Ecuapor. 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. Inflores- centia minute velutina pilis brunneis; bracteae bracteolaeque eglandulosae, per- sistentes; pedicellus 3.57 mm longus, velutinus vel tomentosus. Petala plerumque rosea, lateralia ungue 1.5—2.7 mm longo, limbo 3-4 mm longo, 3-5 mm lato, posti- cum ungue 2.5—3.5 mm longo, limbo 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 lateralis 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, promiqulous or prominent below; petiole 9-18 mm long, 1997 W.R. ANDERSON: MALPIGHIACEAE i/5) 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 420 (—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, ciliate 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 slightly 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, Krukoff 4639 (MICH). Ecuador. Napo: Rio Payamino, tributary of the Rfo Napo, vicinity of Payamino-Capihuara, Lugo 2895 (GH, MICH).—Sucumsios: near ferry crossing over Coca River, 69 km S of Lago Agrio, Gentry 12556 (GB, MICH, MO, QCA, S): Can6én de los Monos, road Coca (Puerto Francisco de Orellana)-Lago Agrio, 12 km N of Coca, binee 2947 (GB, MICH); road Lago Agrio-El Chaco, Lugo 3484 (GB, MICH); environs of Limoncocha, Madi- son et al. 5484 (MICH, OCA, 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. Hudnuco, Tingo Maria, Asplund 13004 (G).—Lorero: Alto Amazonas Prov., Washintsa and vicinity, Rio Huasaga, 3°20'S, 76°20'W, Lewis et al. 11225 (MO).—Mabnre DE Dios: Parque Nacional del Manu, Rio Manu, Cocha Cashu Station, Foster 54/1 (F, MICH); Tambopata Province, Las Piedras, Cusco Amazonico, 12°29'S, 69°03'W, Timand 2009, 2336, 2414 (all MO).—SaAn Martin: Prov. Mariscal Caceres, Distr. Tocache Nuevo, Rio Cafiuto, Schunke V. 10957 (MICH). The epithet of Mascagnia lugoi honors Hélguer 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.—T ype: FRENCH GUIANA. Saiil, La Fumée Mountain Trail, 3°37'N, 53°12'W, 17 Sep 1989 fl, S. Mori, C. Gracie & M. Rothman 20945 (holotype: MICHI; 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 laterali omnino connatae. Petala omnia abaxialiter tomentosa, margine eglandulosa, lateralia 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. Styli 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 lateralis 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, light- to dark- brown-sericeous or eventually glabrescent; stalk of the umbel 6-16 mm long, bear- ing | pair of deciduous sterile bracts 2-8 mm above the base, well below to 1997 W. R. ANDERSON: MALPIGHIACEAE TE 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- mally sericeous, nearly terete, pedaliform at apex (i.e., with a short, broad abaxial extension resembling 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. ADDITIONAL SPECIMENS EXAMINED. Brazil. AMAZONAS: Distr. Agropecudrio, Reserva 1501 (Km 41) da WWF/INPA Projeto da Dinamica Biolégica dos Fragmentos Florestais, 2°24'26"-2°25'31"S, 59°43'SO"W, Lepsch da Cunha et al. 342 (MICH); Reserva Florestal Ducke, Manaus-—Itacoatiara Km 78 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 1997 W.R. ANDERSON: MALPIGHIACEAE 79 26, 2°53'S, 59°S8'W, Vicentini 1014 (MICH).—ParA: Rio Jari, Monte Dourado, Cavalcante 3329 (MG, NY); Tucuruf, ee eas da PA-149 até o Km 50, Revilla et al. 8310 (INPA): Santarém, Km 70 da estrada do Palhao, mal do Caetetu, M. Silva 2604 (MG, MICH, NY); Rio Jari, estrada entre Planalto Ae See ‘Km 13, N. 7. Silva 2785 (LAN, UB); Rio Jari, N. T. Silva 2830 (UB). French Guiana. Saiil, La Fumée Mountain Trail, 3°37'N, 53°12'W, Mori et al. 21012 (CAY, MICH, NY). Guyana. Kamoa River, Clarence Hill, 1°31'N, 58°50'W, Jansen-Jacobs et al. 1738 (MICH, : , 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. araujoi Nied., the type of the genus, which occurs far to the south of M. angelica in Espirito 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. . Mezia angelica. a) leafy stem, with enlargement of abaxial base of lamina to show peretene ve 0.5; b) portion of inflorescence, x0.5; c) umbel of 4 flower buds, each bud enclosed by 2 bracteoles and subtended by | bract, the stalk of the umbel bearing a pair of sterile bracts, x2; d) flower bud about to open, with the 2 bracteoles being forced apart by the enlarging sepals, x2; e) pen flower with petals removed, with the 2 subtending bracteoles intact (above) and with the bracteoles half cut away (below) to show sepals, with the eglandular anterior sepal in center, x2; f) abaxial view of | lateral sepal bearing a large double gland formed from 2 nearly connate glands, x4; g) open flower, from above with posterior (flag) petal uppermost (left) and from the side with 2 petals removed and the posterior petal to the right (right), x2; h) adaxial view of complete androeci- m, laid out with the shortest stamen (fifth from right) that opposite the posterior petal, x4; i) side and adaxial views of anthers, the pair on left opposite petals, those on right opposite sepals, x8; j) gynoecium with anterior style in center, x4, and | style-apex enlarged, x8; k) samaras, abaxial view (above) and adaxial view (below), x0.5; 1) samara in cross section, showing large lateral wing, smaller dorsal wing (pointing straight up), single winglets between dorsal and lateral wings, and single winglets outside lateral wing, x0.5. Drawn by Bobbi Angell, a-j from Mori et al. 20945, k & | from Silva 2830 Reproduced with permission of Dr. Scott Mori and the = York Botanical Garden. 80 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 80° 70° 60° 50° ‘i m M. angelica @M. araujoi —,,. a M. huberi e M. mariposa v M. tomentosa FIG. 6. Distribution of selected species of Mezia. Mezia mariposa W. R. Anderson, sp. nov.—T ype: 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 | 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 alis lateralibus 22-33 mm latis, 31-52 mm altis, ala dorsali 7-12 mm lata, 18-28 mm alta, alis intermedus nullis. 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, (S—) 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 | large gland at base on each side of midrib (these glands often sunken in crypts) and a single row of impressed glands several mm L997 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 SPECIMENS EXAMINED: Brazil. AcrEe: Vizinhanga de Sena Madureira, 9°05'S, 68°40'W, Nelson et al. 523 (INPA, MICH, MG); Colocagdo Boa Unido, Sothers & Santos 8 (MICH).—AMaAzo- NAS: Track from S40 Paulo to Terra Firme, Rio Purus, opposite Béca do Acre, Prance et al. 2586 A, NY).—Para: Upper Cupari River, plateau between the Xingt and Tapajés Rivers, Krukoff 1135 (A, G, K, NY, P); Tucurutf, Lima & Silva 83 (INPA); Boa Vista on the Tapajos River, Monteiro da ar 56 (F); Tucuruf, Rio Caraipé, Revilla et al. 8272 (INPA) & PA-149, aes et al. 8425 (INPA).—Ronponia: Estrada Belmonte, Cordeiro 766 (MICH); Rio Jari, BR-29, Porto Velho, Duarte 7158 (MICH); Pérto Velho, estrada Belmonte, Mota & Coélho 100 (INPA): eastern bank of Rio Madeira at Misericérdia between Cities 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, | km arriba del pueblo de Tournavista o unos 20 km arriba de la confluencia con el Rio Ucayali, Schunke V. 2062 (F, NY, US), 2179 (F, NY, US), 2/86 (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, c.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: Ecuapor. 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 2-4 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, ca 8-10 mm latae. 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 like 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 himedo tropical, primario, May 1990 fr, Gudifio 396 (MICH, MO). 84 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 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. curranii 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 Anderson, W. R. 1981. Siete ll In The botany of the Guayana Highland—Part X1. Mem. New York Bot. Gard. 32: 21-- —__—_—. 1993. Notes on Da Malpighiaceae—IV. Contr. Univ. Michigan Herb. 19: 355-392. Chase, M. W. 1981. A revision of Dicella (Malpighiaceae). Syst. Bot. 6: 159-171. Cuatrecasas, . Prima Flora Colombiana. 2. Malpighiaceae. Webbia 13: 343-664. Niedenzu, F. 8. Malpighiaceae. In Das Pflanz -enreich, ed. A. Engler, IV. 141: 1-870. Leipzig: Wilhelm sie ae O’Donell, C. A., & A. Lourteig. 1943. Malpighiaceae argentinae. Lilloa 9: 221-316, pl. 1-18. ACKNOWLEDGMENTS always, my work is made possible by the cooperation and kind assistance of the curators of many hexbaria 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, QCA, U, UB, UC, US, and VEN. I would also like to acknowledge several iadinidivals who helped in 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, Scott Mori, Basil Stergios, Stephen Tillett, and Beverly Walters. Contr. Univ. Michigan Herb. 21: 85-141. 1997. 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 Mjéberg, 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. Mjéberg (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 85 56 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 Mj6berg 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.” Mjéberg 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, Mjéberg (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 delicate plant-like orchids of the Liparis variety up to the Spathoglottis- and Vanda-like types that grow in bushes almost up to a man’s head.” Mjéberg’s expedition to Mt. Murud resulted in extensive collections of a wide range of plants and animals, many of which were published 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 Mjéberg to Ames, but the Mjéberg 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 Mjéberg collections was annotated by Ames. BOTANICAL COLLECTORS ON MT. MURUD In addition to the Mjéberg 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 Mjéberg’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 Mjéberg 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°31'15"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°51'54"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 LOoF 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 climb 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'S0"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°29'19"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 Rafflesia 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 Mjéberg submitted to Bonaparte are now in the herbarium of the Muséum National d’Histoire Naturelle, Paris (P). Merrill (1928) stated that types of the species he described based on the Mjéberg 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 Myjéberg 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 supplied 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 duplicates 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 Mjdberg 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. murudense), or elevations were given pertaining to two different mountains in Sarawak. Labels on the relevant Mjéberg 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 Mjéberg collections we thought desirable to record. Future additions to the Murud summit flora might appropriately be limited 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 locality 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 allied 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 ABLE 1. Locations of camps used during the expedition of B. L. Burtt, A. M. Martin, and the Sarawak Forest Department (Ilias Paie) in 1967. Camp Elevation Coordinates Location I 4000 ft ? Near the Komap River, ca. 4 km W of Ba Kelalan II 3800 ft = 3°58'N, 115°36'E = On the Bor River at Long Rapata, ca. 4.2 km SW of Camp 1 Il 5600 ft 3°56'N, 115°32'E __N side of the summit ridge, ca. 4.5 km S of Camp I IV 6300 ft = 3°S56'N, 115°32'E _ S side of the summit ridge, ca. 1.7 km W of Camp III Vv 6800 ft = 3°SS'N, 115°31'E _ S side of the summit ridge, ca. 4.2 km WSW of Camp IV Of 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 Mjdberg collections in AMES, P, and UC, but at least those in P and UC have been listed by Bonaparte and Merrill. PHYTOGEOGRAPHY Mjéberg (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 Miller 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 published on the Dulit 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 salient 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 ‘ef.’, ‘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 Mjéberg 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- etics. Twenty-seven new taxa of vascular plants have been described from the Murud summit area, of which 16 are currently recognized. 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 95 FERN ALLIES 1. LYCOPODIACEAE 1.1. HUPERZIA 1.1.1. Huperzia phlegmaria (L.) Rothm. [listed by Merrill (1928) as Urostachys phlegmaria (L.) Herter]. Collections. Mt. Murup: Mjéberg 73 p.p. (UC); Mr. Murup prince, SE side: 1700- 2000 m, Beaman 11489 (K, MSC, UNIMAS). 1.1.2, Huperzia serrata (Thunb. ex Murray) Trevis. Collection. Mt. MuRUD BETWEEN Camps IT AND HI: 1500 m, Burtt & Martin B. 5220 (E). 1.1.3. Huperzia verticillata (L. f.) Trevis. [listed by Merrill (1928) as Urostachys verticillata (L. f.) Herter]. Collections. Mr. Murub: 1900 m, Mjéberg 73 p.p. (UC); Mr. Murub, PATH TO SUMMIT: 1700 m, Nooteboom & Chai 1934 (L). 1.2, LYCOPODIELLA 1.2.1. Lycopodiella cernua (L.) Pic. Serm. Collection. Mr. Murup Camp IV: 1900 m, Burtt & Martin B. 5385 (K). 2. SELAGINELLACEAE 2.1. SELAGINELLA 2.1.1. Selaginella rugulosa Ces. Collections. Mr. Murup Camp IV: 2000 m, Burtt & Martin B. 5415 (K); Mr. Murup N OF SUMMIT: 2100 m, Burtt & Martin B. 5496 (K). FERNS 3. ASPLENIACEAE 3.1. ASPLENIUM 3.1.1. Asplenium nidus L. [listed by Bonaparte (1923a)]. Collection. Mr. Murup: 1500-1800 m, Mjéberg 204 p.p. (P). 96 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 4. BLECHNACEAE 4.1. BLECHNUM 4.1.1. Blechnum sp. [listed by Bonaparte (1923a) as Blechnum capense var. pro- cerum f. integrum Bonap.]. Collection. Mr. Murup: 1500-1800 m, Mjéberg 2/4 (P, holotype of B. capense var. procerum f. integrum Bonap.). 5. CYATHEACEAE 5.1. CYATHEA 5.1.1. Cyathea capitata Copel. [listed by Bonaparte (1923a)]. Collections. Mr. Murup: 1500-1800 m, Mjéberg 95 (P); Mr. Murupb N sipe: Burtt & Martin B. 5379 (E). 5.1.2. Cyathea indet. Collection. Mr. Murup summit: 2400 m, Nooteboom & Chai 2009 (L). 6. DENNSTAEDTIACEAE 6.1. HISTIOPTERIS 6.1.1. Histiopteris incisa (Thunb.) J. Sm. [listed by Bonaparte (1923a)]. Collection. Mr. Murup: 1500-1800 m, Mjéberg 108 (P). 7. DIPTERIDACEAE 7.1. DIPTERIS 7.1.1. Dipteris conjugata Reinw. [listed by Bonaparte (1923a) as D. conjugata var. alpina Christ]. Collection. Mr. Murup summit: 2400 m, Mjoberg s.n. (P). 7.1.2. Dipteris lobbiana Moore Collection. Mr. Murup: 1500-1800 m, Mjéberg s.n. (P) [doubtfully occurring as high as indicated, but so listed by Bonaparte (1923a)]. 7.1.3. Dipteris novoguineensis Posth. Collections. Mr. Murup: 2200 m, Nooteboom & Chai 2039 (L); Mr. Murup N oF SUMMIT: Burtt & 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 a form of the polymorphic D. conjugata Reinw.]. Collection. Mr. Murup: 1500-1800 m, Mjéberg s.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. Collections. Mr. Murup: 2300 m, Nooteboom & Chai 2015 (L), 2400 m, 2036 (L); Mr. Murubp N oF summit: 2100 m, Burtt & Martin B. 5465B (K). 9.1.2. Gleichenia peltophora Copel. var. peltophora Collection. Mr. MuruD N oF summit: 2100 m, Burtt & Martin B. 5465A (K). 9,2. STICHERUS 9.2.1. Sticherus hirtus (Blume) Ching var. paleaceus (Baker) Parris [listed by Bonaparte (1923a) as Gleichenia hirta Blume]. Collection. Mr. Murup: 1500-1800 m, Mjéberg s.n. (P). 10. GRAMMITIDACEAE 10.1. CTENOPTERIS 10.1.1. Ctenopteris taxodioides (Baker) Copel. [listed by Bonaparte (1923a) and Merrill (1928) as Polypodium taxodioides Baker]. Collections. Mr. Murup: 2400 m, Mjéberg 7 (K), 70 (UC), 1500-1800 m, 256 (P). 10.2. PROSAPTIA 10.2.1. Prosaptia contigua (G. Forst.) C. Pres] [listed by Bonaparte (1923a) as Davallia contigua (G. Forst.) J. Sm.]. Collections. Mr. Murup: 1500-1800 m, Mjéberg s.n. (P); Mr. Murup summit: 2400 m, Mjoberg s.n. (P). 98 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 10.3. SCLEROGLOSSUM 10.3.1. Scleroglossum minus (Fée) C. Chr. Collection. Mr. MuRuD NEAR 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 pluma Hook. ]. Collections. Mr. Murup: above 1200 m, Mjdberg 67 (UC), 2300 m, Nooteboom & Chai 2014 (L). 11.1.2. Macroglena schlechteri (Brause) Copel. Collection. Mr. Murup: 2100 m, Burtt & Martin B. 5356 (K). 11.2. PLEUROMANES 11.2.1. Pleuromanes album (Blume) Parris Collection. Mr. Murup: 1900 m, Burtt & Martin B. 5392 (E). 12. OLEANDRACEAE 12.1. OLEANDRA 12.1.1. Oleandra neriiformis Cav. [listed by Bonaparte (1923a)]. Collection. Mr. Murup: 1500-1800 m, Mjéberg 182 (P). 13. PLAGIOGYRIACEAE 13.1. PLAGIOGYRIA 13.1.1. Plagiogyria egenolfioides (Baker) Copel. var. latipinna (Copel.) Zhang & Noot. Collection. Mr. Murub, PATH To SUMMIT: 1800 m, Nooteboom & Chai 1945 (L). 13.1.2. Plagiogyria pycnophylla (Kunze) Mett. [listed by Bonaparte (1923a)]. Collection. Mr. Murupb: 1500-1800 m, Mjéberg 22 (P, holotype of P. pycnophylla var. integra subvar. stenophylla Bonap.). 13.1.3. Plagiogyria tuberculata Copel. [listed by Bonaparte (1923a) as P. rotundi- pinnata Bonap.]. Collections. Mr. Murup: 2400 m, Mjdberg s.n. (P, holotype of P. rotundipinnata Bonap.); Mt. Murup N or summit: Burtt & Martin B. 5474 (E). 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 2g 14. POLYPODIACEAE 14.1. SELLIGUEA 14.1.1. Selliguea albidosquamata (Blume) Parris [listed by Bonaparte (1923a) and Merrill (1928) as Polypodium albidosquamatum Blume]. Collection. Mr. Murup: 1900 m, Mjéberg 77 (UC). 14.1.2. Selliguea enervis (Cav.) Ching [listed by Merrill (1928) as Polypodium triquetrum Blume]. Collection. Mt. Murupb: Mjéberg 68 (UC). 15. SCHIZAEACEAE 15.1. SCHIZAEA 15.1.1. Schizaea malaccana Baker var. malaccana Collections. Mr. MurupD N sine: 2100 m, Burtt & Martin B. 5483 (K); Mr. Murup SUMMIT: 2400 m, Nooteboom & Chai 2064 (L). 16. THELYPTERIDACEAE 16.1. CoRYPHOPTERIS 16.1.1. Coryphopteris pubirachis (Baker) Holttum var. sulawesica Holttum Collection. Mr. Murup 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. VITTARIA 17.1.1. Vittaria indet. Collection. Mr. Murup: 2200 m, Nooteboom & Chai 2028a (L). 18. WOODSIACEAE 18.1. DipLAzium 18.1.1. Diplazium porphyrorachis (Baker) Diels Collection. Mr. Murub: Sarawak Museum Native Collector 2937 (PNH) [fide M. G. Price, Gard. Bull. Singapore 36: 29 (1983)]. 100 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 GYMNOSPERMS 19. ARAUCARIACEAE 19.1. AGATHIS 19.1.1. Agathis kinabaluensis de Laub. Collections. Mr. MURUD BETWEEN IST AND 2ND SUMMITS: 2300 m, Yii S. 44467 (K); Mr. Murupb Just BELOW summit: 2400 m, Nooteboom & Chai 2013 (K, L); Mt. MurRuD SUMMIT RIDGE: 2300-2400 m, Beaman 11464 (K, MSC, UNIMAS). 20. PHYLLOCLADACEAE 20.1. PHYLLOCLADUS 20.1.1. Phyllocladus hypophyllus Hook. f. Collections. Mr. Murup Camp IV: 2000 m, Burtt & Martin B. 5437 (E); Mr. Murup SUMMIT RIDGE: 2300-2400 m, Beaman 11466 (K). 21. PODOCARPACEAE 21.1. DACRYCARPUS 21.1.1. Dacrycarpus imbricatus (Blume) de Laub. [listed by Merrill (1928) as odocarpus javanicus (Burm. f.) Merr.]. Collection. Mr. Murup: 1500 m, Mjdberg 99 (UC). 21.2. DACRYDIUM 21.2.1. Dacrydium beccarii Parl. in DC. [listed by Merrill (1928)]. Collection. Mr. Murup: 1200 m, Mjéberg 100 (UC). 21.2.2. Dacrydium gibbsiae Stapf Collections. Mr. MURUD NEAR SUMMIT: 2100 m, //ias S. 26505 (IK); Mt. MuRUD SUMMIT: 2400 m, Anderson & Ilias S. 26471 (K); Mr. MuRuD SUMMIT RIDGE: 2300-2400 m, Beaman 11440 (K, MSC, UNIMAS). 21.2.3. Dacrydium xanthandrum Pilger Collections. Mr. MURUD BETWEEN IST AND 2ND SUMMITS: 2200 m, Yii S$. 44608 (IK); Mr. MuRuD SUMMIT RIDGE: 2300-2400 m, Beaman 11487 (K, MSC, UNIMAS). 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. Mr. Murub, PATH TO SuMMIT: 1700 m, Nooteboom & Chai 1937 (B, L, US). 23. ARECACEAE 23.1. CALAMUS 23.1.1. Calamus gibbsianus Becc. Collections. Mr. Murup summit: 2400 m, Nooteboom & Chai 1998 (L); Mt. Murup SUMMIT RIDGE: 2300-2400 m, Beaman 11471 (K, UNIMAS). 23.1.2. Calamus mattanensis Becc. [listed by Merrill (1928) as C. ferrugineus Becc., a synonym fide Dransfield (1992)]. Collection. Mr. Murup: 1900 m, Mjéberg 126 (UC). 23.1.3. Calamus pilosellus Becc. [listed by Merrill (1928); not recorded by Drans- field (1992) as occurring in the Kelabit Highlands]. Collection. Mr. Murup: 1900 m, Mjéberg 128 (UC). 23.2. PINANGA 23.2.1. Pinanga capitata Becc. Collection. Mr. Murup 2ND summit: 2300 m, Yii S. 44442 (K, L). 24. CENTROLEPIDACEAE 24.1. CENTROLEPIS 24.1.1. Centrolepis philippinensis Merr. Collection. Mr. Murup N sipe: 2100 m, Burtt & Martin B. 5494 (E). 25. CYPERACEAE 25.1. CAREX 25.1.1. Carex cruciata Wahl. [listed by Merrill (1928)]. Collection. Mr. Murup: Mjdberg 113 (UC). 102 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 25.1.2. Carex filicina Nees Collection. Mr. MuRUD, RIDGE ABOVE CAmP IV: 2000 m, Burtt & Martin B. 5383 (E). 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. Mr. Murup Camp V: 2100-2300 m, Burtt & Martin B. 5475 (E); Mr. Murupb summit: 2400 m, Nooteboom & Chai 198] (L); Mr. MurubD SUMMIT RIDGE: 2300- 2400 m, Beaman 11465 (K, MSC, UNIMAS). 25.3. OREOBOLUS 25.3.1. Oreobolus kiikenthalii Steenis Collections. Mr. Murup Camp V: 2100-2300 m, Burtt & Martin B. 5482 (E, L); Mr. Murub summit: 2400 m, Nooteboom & Chai 2023 (L). 26. IRIDACEAE 26.1. PATERSONIA 26.1.1. Patersonia lowii Stapf Collections. Mr. Murup: 2200 m, Nooteboom & Chai 2025 (L); Mr. MuRUD BETWEEN Camp V AND SUMMIT: 2300 m, Burtt & Martin B. 5455 (E). 27, MELANTHIACEAE 27.1. PETROSAVIA 27.1.1. Petrosavia stellaris Becc. Collection. Mr. MuRuUD NEAR Camp IIT: 1800 m, Burtt & Martin B. 5277 (E). 28. ORCHIDACEAE 28.1. APPENDICULA 28.1.1. Appendicula bilobulata J. J. Wood Collection. Mr. Murup NEAR Camp III: 1800 m, Burtt & Martin B. 5287 (E, holotype of A. bilobulata; SAR, isotype). 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 103 28.1.2. Appendicula congesta Rid. Collection. Mr. Murup Camp III: 1700 m, Burtt & Martin B. 5233 (E, SAR). 28.1.3. Appendicula foliosa Ames & C. Schweinf. Collection. Mr. Murup: 1600 m, Yii S. 4442/7 (K). 28.1.4. Appendicula longirostrata Ames & C. Schweinf. Collection. Mr. Murup ABOVE Camp IV: 2000 m, Burtt & Martin B. 5434 (E, SAR). 28.1.5. Appendicula indet. Collections. Mr. Murup: 1600 m, Yii S. 4440] (K); Mr. Murub, PATH TO SUMMIT: 1700 m, Nooteboom & Chai 1922 (L). 28.2. BROMHEADIA 28.2.1. Bromheadia crassiflora J. J. Sm. Collections. Mt. Murupb: 2200 m, Nooteboom & Chai 2054 (L); Mr. Murup N SIDE: 2100 m, Burtt & Martin B. 5463A (SAR); Mr. MuRUD BETWEEN IST AND 2ND SUMMITS: 2300 m, Yii S. 44482 (L, SAR). 28.3. BULBOPHYLLUM 28.3.1. Bulbophyllum anguliferum Ames & C. Schweinf. Collection. Mr. Murup Camp III: 2000 m, Burtt & Martin B. 5252 (E). 28.3.2. Bulbophyllum sopoetanense Schltr. Collection. Mr. Murup NEAR Camp IV: 1900 m, Burtt & Martin B. 5404 (E, SAR). 28.3.3. Bulbophyllum aff. teres Carr Collection. Mr. Murup ABOVE Camp III: 1800 m, Burtt & Martin B. 5262 (E). 28.4. CALANTHE 28.4.1. Calanthe speciosa (Blume) Lindl. Collections. Mr. Murup: 1800 m, Nooteboom & Chai 1969 (L, SAR); Mr. Murup NEAR Camp III: 1800 m, Burtt & Martin B. 5271 (E, SAR). 28.4.2. Calanthe tenuis Ames & C. Schweinf. Collections. Mt. Murup ABOVE Camp IV: 2000 m, Burtt & Martin B. 5352 (E); Mr. MurubD BELOW Camp IV: 1800 m, Burtt & Martin B. 5368 (E, SAR). 104 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 28.4.3. Calanthe indet. Collections. Mr. Murup: 1700 m, Nooteboom & Chai 1910 (L), 1800 m, 1969 (L). 28.5. CHELONISTELE 28.5.1. Chelonistele lamellulifera Carr Collection. Mr. MuRuD NEAR Camp III: 1800 m, Burtt & Martin B. 5272 (E, SAR). 28.6. COELOGYNE 28.6.1. Coelogyne craticulaelabris Carr Collection. Mr. Murup N sipe: 2300 m, Burtt & Martin B. 5454 (E, SAR). 28.6.2. Coelogyne hirtella J. J. Sm. Collections. Mr. Murup: 2200 m, Nooteboom & Chai 2019 (SAR); Mr. Murup Camp III: 1800 m, Burtt & Martin B. 5256 (E). 28.6.3. Coelogyne kinabaluensis Ames & C. Schweinf. Collection. Mr. Murup Camp III: 1700 m, Burtt & Martin B. 5236 (EB). 28.6.4. Coelogyne moultonii J. J. Sm. Collections. Mr. Murup Camp III: 1600 m, Burtt & Martin B. 5258 (SAR); Mr. Mu- RUD NEAR Camp III: 1600 m, Burtt & Martin B. 5258 (E). 28.6.5. Coelogyne planiscapa Carr var. planiscapa Collection. Mr. Murup Camp II: 1700 m, Burtt & Martin B. 5243 (E). 28.6.6. Coelogyne radioferens Ames & C. Schweinf. Collections. Mt. MURUD NEAR Camps III AND IV: 1800-2000 m, Burtt & Martin B. 5274 (E, SAR); Mr. Murup/Dapvr River: 1900 m, Ilias S. 26519 (SAR). 28.6.7. Coelogyne tenompokensis Carr Collection. Mr. Murup Camp III: 1700 m, Burtt & Martin B. 5245 (E, SAR). 28.6.8. Coelogyne indet. Collections. Mr. Murup: 2200-2300 m, Nooteboom & Chai 2055 (L); Mt. Murup/Ba KeELALAN: 1700 m, Burtt & Martin B. 5247 (SAR). 28.7. CyMBIDIUM 28.7.1. Cymbidium elongatum J. J. Wood, Du Puy & Shim 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 105 Collections. Mr. Murupb Camp V: 2100-2300 m, Burtt & Martin B. 5460 (E, SAR); Mr. MuruD ABOVE Camp IV: 2000 m, Burtt & Martin B. 5428 (E, SAR). 28.8. DENDROBIUM 28.8.1. Dendrobium alabense J. J. Wood Collections. Mr. Murup N sipe: 2100 m, Burtt & Martin B. 5480 (E); Mr. Murup ABOVE Camp IV: 1600 m, Burtt & Martin B. 5354 (E). 28.8.2. Dendrobium cymbulipes J. J. Sm. Collection. Mr. MuruD NEAR Camp III: 1800 m, Burtt & Martin B. 5333 (E). 28.8.3. Dendrobium piranha C. L. Chan & P. Cribb Collection. Mr. Murup: 2200 m, Nooteboom & Chai 2032 (L, SAR). 28.8.4. Dendrobium sp. (sect. Rhopalanthe) Collection. Mr. MuruD sUMMIT RIDGE: 2300-2400 m, Beaman 11482 (K). 28.8.5. Dendrobium indet. Collection. Mr. Murup: 2200 m, Nooteboom & Chai 2033 (L). 28.9. DENDROCHILUM 28.9.1. Dendrochilum crassifolium Ames Collections. Mr. Murup: 2400 m, Nooteboom & Chai 1995A (L); Mr. Murup Camp III: 1700 m, Burtt & Martin B. 5241 (E, K, SAR); Mr. Murup SW oF Camp III: 1800 m, Burtt & Martin B. 5319 (E). 28.9.2. Dendrochilum dewindtianum W. W. Sm. var. dewindtianum Collection. Mt. MurRuD SUMMIT RIDGE: 2300-2400 m, Beaman 11448 (K, MSC, UNIMAS). 28.9.3. Dendrochilum galbanum J. J. Wood Collections. Mt. MuruD 2ND suMMIT: 2300 m, Yii S. 44430 (K, SAR); Mr. Murup SW oF Camp III: Burtt & Martin B. 5328 (E, K). 28.9.4. Dendrochilum gibbsiae Rolfe Collections. Mr. Murup: 1600 m, Yii S. 44402 (K); Mr. Murup BELOW Camp III: 1600 m, Burtt & Martin B. 5299 (E, SAR). 28.9.5. Dendrochilum imbricatum Ames Collection. Mr. Murup: 1900-2400 m, Mjéberg 64 (AMES). 106 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 28.9.6. Dendrochilum lancilabium Ames Collections. Mr. Murup: 2100 m, Ilias S. 26455 (E, K, L), 1900-2400 m, Mjéberg 52 (AMES); Mr. Murup SW oF Camp III: 1800 m, Burtt & Martin B. 5319 (EB). 28.9.7. Dendrochilum longipes J. J. Sm. Collections. Mr. Murup: 1900-2400 m, Mjoberg 66 (AMES); Mr. Murupb 2ND SUMMIT: 2300 m, Yii S. 44432 (K, L, SING); Mr. Murup NEAR Camp IV: 1900 m, Burtt & Martin B. 5399 (E); Mr. Murup, PATH TO suMMIT: 1800 m, Nooteboom & Chai 1954 (L). 28.9.8. Dendrochilum muluense J. J. Wood Collections. Mr. Murup Camp IIE: 1700 m, Burtt & Martin B. 5244 (E, SAR); Mr. MURUD BETWEEN IsT AND 2ND SUMMITS: 2200 m, Yii S. 44616 (K). 28.9.9. Dendrochilum murudense (J. J. Wood) J. J. Wood Collections. Mr. Murupb: 2400 m, Nooteboom & Chai 1995 (K, holotype of D. crassi- folium var. murudense J. J. Wood); Mr. Murup suMMIT RIDGE: 2300-2400 m, Beaman 11459 (K, UNIMAS). 28.9.10. Dendrochilum simplex J. J. Sm. Collection. Mr. Murup: 1900-2400 m, Mjéberg 49 (AMES). 28.10. DILOCHIA 28.10.1. Dilochia cantleyi (Hook. f.) Rid. Collections. Mr. Murup: 1700 m, Yii S. 44627 (K); Mr. Murup N SIDE NEAR Camp V: 2100 m, Burtt & Martin B. 5443 (E, SAR); Mr. Murup, Dapur RIVER HEADWATERS: 2400 m, lias S. 26539 (K). 28.10.2. Dilochia rigida (Ridl.) J. J. Wood Collections. Mr. Murupb 2ND summit: 2200 m, /lias S$. 26408 (K); Mr. MURUD BETWEEN IsT AND 2ND SUMMITS: 2300 m, Yii S. 44486 (K, SAR); Mr. MuRuD SUMMIT RIDGE: 2300-2400 m, Beaman 11455 (K); Mr. Murup, Dapur RIVER HEADWATERS: 2400 m, Ilias S. 26538 (K). 28.11. EPIGENEIUM 28.11.1. Epigeneium indet. Collection. Mr. Murub: 2200-2300 m, Nooteboom & Chai 2044 (L). 28.12. ERIA 28.12.1. Eria aff. brookesii Ridl. Collections. Mr. Murup Camp HI: 1700 m, Burtt & Martin B. 523] (E, SAR); Mr. Murub SW or Camp III: 1800 m, Burtt & Martin B. 5327 (E, SAR). 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 107 28.12.2. Eria crassipes Rid. Collections. Mr. Murup: 2400 m, Nooteboom & Chai 2065 (SAR), 2300 m, Yii S. 4443] (K). 28.12.3. Eria robusta (Blume) Lind. Collections. Mr. Murup N sipe: 2100 m, Burtt & Martin B. 5466 (E); Mr. Murub ABOVE Camp III: 1800 m, Burtt & Martin B. 5263 (E); Mr. MuruD BETWEEN IST AND 2ND summits: 2300 m, Yi S$. 44497 (K); Mr. MuRuUD SUMMIT RIDGE: 2300-2400 m, Beaman 11485 (K, MSC, UNIMAS) 28.12.4. Eria indet. Collections. Mr. Murup: 1700 m, Rena, Rantai et al. S. 60336 (K); Mr. Murub, PATH TO SUMMIT: 1700 m, Nooteboom & Chai 1927 (L). 28.13. Liparis 28.13.1. Liparis compressa (Blume) Lindl. var. compressa Collection. Mr. Murup: 1700 m, Rena, Rantai et al. S. 60335 (K). 28.13.2. Liparis pandurata Ames Collection. Mr. Murup Camp III: 1700 m, Burtt & Martin B. 5237 (E, SAR). 28.14. NABALUIA 28.14.1. Nabaluia exaltata de Vogel Collections. Mr. Murup: 2100 m, Ilias S. 26466 (K), 2400 m, Rena, Rantai et al. S. 60394 (K); Mr. Murup Camp III: 1600 m, Burtt & Martin B. 5260 (E); Mr. MuruD ABOVE Camp III: 1800 m, Burtt & Martin B. 5259 (E, SAR). 28.15. OBERONIA 28.15.1. Oberonia aff. griffithiana Lindl. Collection. Mr. MuruD NEAR CAmp IV: 1900 m, Burtt & Martin B. 5393 (E). 28.16. PHAIUS 28.16.1. Phaius aff. pauciflorus (Blume) Blume Collection. Mr. MURUD NEAR Camp III: 1800 m, Burtt & Martin B. 5329 (E, SAR). 28.17. PHOLIDOTA 28.17.1. Pholidota gibbosa (Blume) de Vriese Collections. Mt. Murup Camp III: 1800 m, Burtt & Martin B. 5269 (E); Mt. Murupb Camp IV: 1900 m, Burtt & Martin B. 5390 (SAR). 108 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 28.17.2. Pholidota mediocris de Vogel Collection. Mr. MURUD NEAR Camp III: 1800 m, Burtt & Martin B. 5338 (E). 28.18. TRICHOTOSIA 28.18.1. Trichotosia cf. annulata Blume Collection. Mr. Murub/BELABAN RIVER HEADWATERS: 1700 m, Ilias S. 26353 (K). 28.18.2. Trichotosia aff. ferox Blume Collection. Mr. MuruD N sipdeE: 2100 m, Burtt & Martin B. 5464 (E). 28.18.3. Trichotosia poculata (Ridl.) Kraenzl. Collection. Mr. Murub, PATH TO SUMMIT: 1800 m, Nooteboom & Chai 1946 (L, SAR). 28.18.4. Trichotosia sarawakensis Carr Collections. Mr. Murup Camp III: 1700 m, Burtt & Martin B. 5229 (E, SAR), 1700 m, B. 5230 (E, SAR). 29. PANDANACEAE 29.1. FREYCINETIA 29.1.1. Freycinetia kinabaluana B. C. Stone Collection. Mr. Murupb, PATH TO SUMMIT: 1700 m, Nooteboom & Chai 1939 (L). 29.1.2. Freycinetia rigidifolia Hems!l. Collection. Mr. Murup/BELABAN River: 1700 m, /lias S$. 26337 (K, L). 29.2. PANDANUS 29.2.1. Pandanus papilio B. C. Stone Collection. Mr. Murup Camp IV: 2000 m, Burtt & Martin B. 5431 (E). 30. PHORMIACEAE 30.1. DIANELLA 30.1.1. Dianella ensifolia (L.) DC. [listed by Merrill (1928)]. Collections. Mr. Murup: 1500 m, Mjéberg 123 (UC), 2200 m, Nooteboom & Chai 202041), 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 109 30.1.2. Dianella javanica (Blume) Kunth Collection. Mr. Murup summit: 2400 m, Nooteboom & Chai 1999 (L). 31. POACEAE 31.1. ISACHNE 31.1.1. Isachne kinabaluensis Merr. Collections. Mr. Murup N sipe: 2100 m, Burtt & Martin B. 5463 (E); Mr. Murup SUMMIT: 2400 m, Nooteboom & Chai 2029 (L). 31.2. RACEMOBAMBOS 31.2.1. Racemobambos glabra Holttum Collection. Mr. Murup N sipe: 2100 m, Burtt & Martin B. 5495 (E). 32. SMILACACEAE 32.1. SMILAX 32.1.1. Smilax lanceifolia Roxb. Collections. Mr. Murup: 2400 m, Nooteboom & Chai 1987 (K); Mr. MuruD N sIDE: 2300 m, Burtt & Martin B. 5453 (E); Mt. Murup/BELABAN River: 1700 m, lias S. 26332 (K). 32.1.2. Smilax sp. 1 Collection. Mr. Murub, PATH TO suMMIT: 1700 m, Nooteboom & Chai 1906 (L). 32.1.3. Smilax sp. 2 Collection. Mt. Murupb, 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. Mr. Murup Camp V: 2100-2300 m, Burtt & Martin B. 5484 (E). 34. ZINGIBERACEAE 34.1. ALPINIA 110 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 34.1.1. Alpinia glabra Rid. Collections. Mt. MuRUD BELOW Camp III: 1500-1600 m, Burtt & Martin B. 5300 (E); Mr. MurRuD BETWEEN CAmps IT AND III: 1500 m, Burtt & Martin B. 5221 (E). 34.1.2. Alpinia indet. Collection. Mr. MURUD, PATH TO SUMMIT: L800 m, Nooteboom & Chai 1959 (L). 34.2. AMOMUM 34.2.1. Amomum anomalum R. M. Sm. Collections. Mr. Murup BELOW Camp III: 1500-1600 m, Burtt & Martin B. 5303 (E); Mr. MuruD NEAR Camp III: 1800 m, Burtt & Martin B. 5337 (E, L). 34.2.2. Amomum flavoalbum R. M. Sm. Collection. Mr. ieee Camp III: 1800 m, Burtt & Martin B. 5283 (E). 34.2.3. Amomum aff. flavoalbum R. M. Sm. Collection. Mr. Murup Camp IV: 2000 m, Burtt & Martin B. 5351 (BE). 34.2.4. Amomum luteum R. M. Sm. Collection. Mr. Murup BELOW Camp IV: 1800 m, Burtt & Martin B. 5372 (E). 34.3. BURBIDGEA 34.3.1. Burbidgea nitida Hook. f. [listed by Merrill (1928)]. Collection. Mr. Murup: Mjéberg 122 (UC). 34.3.2. Burbidgea schizocheila Hackett Collection. Mr. MuruD BELOW Camp III: 1500-1600 m, Burtt & Martin B. 5314 (E). 34.4. ETLINGERA 34.4.1. Etlingera fimbriobracteata (K. Schum.) R. M. Sm. Collection. Mr. Murup Camp III: 1800 m, Burtt & Martin B. 5342 (BE). 34.4.2. Etlingera longipetiolata (B. L. Burtt & R. M. Sm.) R. M. Sm. Collection. Mr. MuruD BELOW Camp IV: 2000 m, Burtt & Martin B. 5343 (E). 34.4.3. Etlingera aff. muluensis R. M. Sm. Collection. Mr. Murup Camp III: 1800 m, Burtt & Martin B. 5341 (BE). 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 111 34.4.4, Etlingera pubescens (B. L. Burtt & R. M. Sm.) R. M. Sm. Collection. Mr. Murup BELOw Camp IV: 1800 m, Burtt & Martin B. 5366 (E). 34.4.5. Etlingera punicea (Roxb.) R. M. Sm. Collection. Mr. Murup Camp III: 1800 m, Burtt & Martin B. 5279 (E). 34.5. HEDYCHIUM 34.5.1. Hedychium cylindricum Ridl. [listed by Merrill (1928) as H. myjdbergii Collections. Mr. Murup: 1900-2400 m, Mjdberg 118 (UC, holotype of H. mjébergii Merr.), 1900-2400 m, /21 (UC); Mr. Murup Camp IV: 2000 m, Burtt & Martin B. 5382 (E). 34.5.2. Hedychium indet. Collection. Mr. Murup, PATH To SUMMIT: 1700 m, Nooteboom & Chai 1938 (L). 34.6. HORNSTEDTIA 34.6.1. Hornstedtia incana R. M. Sm. Collection. Mr. Murup Camp IV: 2000 m, Burtt & Martin B. 5402 (E). 34.7. PLAGIOSTACHYS 34.7.1. Plagiostachys bracteolata R. M. Sm. Collection. Mt. MURUD 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. Murup, PATH TO SUMMIT: 1700 m, Nooteboom & Chai 1933 (L). 36. ACTINIDIACEAE 36.1. SAURAUIA 36.1.1. Saurauia amoena Stapf Collections. Mt. Murup: 1600 m, Yi S. 44405 (K); Mr. Murup BELOw Camp III: 1600 m, Burtt & Martin B. 5294 (E); Mt. MuRuD/BELABAN RIVER HEADWATERS: 1700 m, Ilias S. 26303 (K). 112 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 37. ANACARDIACEAE 37.1. TOXICODENDRON 37.1.1. Toxicodendron borneense (Stapf) Gillis Collections. Mt. MuRUD BETWEEN lsT AND 2ND suMMITS: 2300 m, Yii S. 44488 (K, SAR), 2200 m, 446/72 (K, SAR); Mr. MurupD sumMIT RIDGE: 2300-2400 m, Beaman 11454 (K, UNIMAS). 38. APIACEAE 38.1. HYDROCOTYLE 38.1.1. Hydrocotyle javanica Thunb. Collection. Mr. Murub BELow Camp IV: 1800 m, Burtt & Martin B. 5376 (E). 39. APOCYNACEAE 39.1. ALYXIA 39.1.1. Alyxia oleifolia King & Gamble Collection. Mr. Murup: 2200 m, Nooteboom & Chai 2053 (L). 39.2. LEUCONOTIS 39.2.1. Leuconotis eugenifolia (Wall. ex G. Don) A. DC. [listed by Merrill (1928) as Leuconotis sp.]. Collection. Mr. Murup: 1900 m, Mjoberg 124 (UC). 40. AQUIFOLIACEAE 40.1. ILEx 40.1.1. Nex harmsiana Loesn. Collections. Mr. Murup: 2100 m, /lias S. 26453 (K, L), 2200 m, Nooteboom & Chai 2020 (K, L), 2200 m, 2042 (L); Mr. MuruD BETWEEN IsT AND 2ND SUMMITS: 2300 m, Yii S. 44489 (IK); Mr. MURUD SUMMIT RIDGE: 2300-2400 m, Beaman 11468 (K, UNIMAS). 40.1.2. Hex havilandii Loes. [listed by Merrill (1928) as /. confertifolia Merr.]. Collections. Mr. Murup: 2200 m, Nooteboom & Chai 2037 (L, SAR); Mr. Murup 2Nnp SUMMIT: 2300 m, Yii S. 44425 (K, L); Mr. Murup Camp V: 2100-2300 m, Burtt & Martin B. 5468 (E); Mr. MuruD BETWEEN Ist AND 2ND summits: 2200 m, Yii S. 44618 (K, L); Mr. MuruD NEAR SUMMIT: 2400 m, Mjéberg 93 (UC, holotype of /. confertifolia Merr.); Mr. MURUD SUMMIT RIDGE: 2300-2400 m, Beaman 11444 (K, MSC, UNIMAS), 2300-2400 m, //483 (K, UNIMAS). 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 113 41. ARALIACEAE 41.1. ARTHROPHYLLUM 41.1.1. Arthrophyllum collinum Philipson Collections. Mr. Murup: 2300 m, /lias S. 26486 (L), 2200 m, Nooteboom & Chai 2048 (L); Mr. Murup SUMMIT RIDGE: 2300-2400 m, Beaman 11470 (K, MSC, UNIMAS). 41.2. SCHEFFLERA 41.2.1. Schefflera mjébergii Merr. [listed by Merrill (1928)]. Collections. Mr. Murup: 1900 m, Mjéberg 117 (UC, holotype of S. mjébergii Merr.); Mr. MuruD 2ND summit: 2000 m, Ilias S. 26386 (K, SAR); Mr. Murup Camp III: 1700 m, Burtt & Martin B. 5247 (E); Mr. MuruD BETWEEN IsT AND 2ND suMMITS: 2300 m, Yii S. 44487 (K, SAR); Mr. MuruD SUMMIT RIDGE: 2300-2400 m, Beaman 11479 (K, MSC, UNI- MAS); Mr. Murup, PATH To suMMiIT: 1800 m, Nooteboom & Chai 1961 (L, SAR); Mr. Murub/Dapur RIVER HEADWATERS: 2100 m, /lias S. 26467 (SAR). 41.2.2. Schefflera aff. remotiserrata Merr. Collection. Mr. Murup: 2200 m, Nooteboom & Chai 2052 (L, SAR). 41.2.3. Schefflera sp. Collections. Mr. MURUD BETWEEN IsT AND 2ND sUMMITs: 2200 m, Yii S. 44619 (K, L, SAR); Mr. Murup, LONG Rapata/BA KELALAN: 1700 m, Ilias S. 26557 (K, L); Mr. Murup, PATH TO SUMMIT: 1800 m, Nooteboom & Chai 1958 (L, SAR). 42. ASCLEPIADACEAE 42.1. DiscHIDIA 42.1.1. Dischidia nummularia R. Br. Collection. Mr. MuruD Camp III: 1700 m, Burtt & Martin B. 5238 (E). 43. ASTERACEAE 43.1. VERNONIA 43.1.1. Vernonia phanerophlebia Merr. var. dulitensis Koster Collections. Mr. Murup: 2200 m, Nooteboom & Chai 2017 (L); Mr. Murub Camp IV: 1900 m, Burtt & Martin B. 5422 (E). 44. BALANOPHORACEAE 44.1. BALANOPHORA 114 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 44.1.1. Balanophora papuana Schltr. Collections. Mr. Murup: 2200 m, Nooteboom & Chai 2034 (L); Mt. MURUD BETWEEN Camp V AND SUMMIT: 2300 m, Burtt & Martin B. 5447 (EB). 44.1.2. Balanophora indet. Collection. Mr. Murub, 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. Mt. Murup: 1900-2400 m, Mjéberg 119 (UC, holotype of B. murudensis Merr.). 45.1.2. Begonia sp. Collection. Mr. Murub, PATH To SUMMIT: 1700 m, Nooteboom & Chai 1920 (L). 46. CAPRIFOLIACEAE 46.1. ViBURNUM 46.1.1. Viburnum hispidulum Kern Collections. Mt. MURUD BETWEEN IST AND 2ND SUMMITS: 2200 m, Yii S. 4462/7 (KK); Mr. Murupb summit: 2400 m, Nooteboom & Chai 2061 (L). 47. CELASTRACEAE 47.1. MicROTROPIS 47.1.1. Microtropis valida Rid. Collection. Mr. Murub, PATH TO SUMMIT: 1800 m, Nooteboom & Chai 1943 (L). 47.1.2. Microtropis wallichiana Wight ex Thwaites Collection. Mt. Murup N oF summit: 2300 m, Nooteboom & Chai 2043 (L). 47.2. PERROTTETIA 47.2.1. Perrottetia alpestris (Blume) Loesn. Collection. Mr. Murup: 1600 m, Yii S$. 44408 (K, L). 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 115 48. CLUSIACEAE 48.1. GARCINIA 48.1.1. Garcinia indet. Collections. Mr. Murup Camp IV: 2000 m, /lias S. 26394 (L); Mr. Murup summit: 2400 m, Nooteboom & Chai 2010 (L); Mr. Murub, PATH TO SUMMIT: 1700 m, Nooteboom & Chai 1930 (L), 1800 m, 1964 (L); Mt. MurupD/BELABAN River: 1700 m, /lias S$. 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. Mr. 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 Collections. Mr. MuRuD ABOVE Camp IV: 2000 m, Burtt & Martin B. 5423 (E); Mr. Murub/BELABAN River: 1700 m, /lias S. 26315 (K), 1800 m, S. 26365 (K). 51. DIPTEROCARPACEAE 51.1. VATICA 51.1.1. Vatica granulata Slooten subsp. sabaensis Ashton Collections. Mt. Murup: 1600 m, Yi S. 44403 (K); Mr. Murupb/BELABAN RIver: 1700 m, Ilias S. 26312 (K). 51.1.2. Vatica indet. Collection. Mr. Murup: 2400 m, Nooteboom & Chai 2069 (L). 52. ELAEOCARPACEAE 52.1. ELAEOCARPUS 52.1.1. Elaeocarpus glaberrimus R. Knuth 116 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Collections. Mr. Murup: 2100 m, Ilias S. 26452 (K), 2100 m, S. 26495 (K), 2300 m, Yii S. 44427 (KK); Mt. MuruD NEAR Camp V: 2100 m, /lias S$. 26510 (IK, L); Mt. MuRUD SUMMIT: 2400 m, Nooteboom & Chai 2003 (L); Mr. MurupD SUMMIT RIDGE: 2300-2400 m, Beaman 11484 (K, UNIMAS). 52.1.2. Elaeocarpus murudensis Merr. [listed by Merrill (1928)]. Collections. Mr. Murup: 1900-2400 m, Mjdberg 53 (BM, isotype; UC, holotype of F. murudensis Merr.), 2200 m, Nooteboom & Chai 1976 (L); Mr. MuURUD BETWEEN IST AND 2ND SUMMITS: 2300 m, Yii S. 44471 (K, L); Mr. Murup summit: 2400 m, Nooteboom & Chai 2068 (K, L); Mr. MurupD SUMMIT RIDGE: 2300-2400 m, Beaman 11449 (K, MSC, UNIMAS). 52.1.3. Elaeocarpus nanus Corner subsp. congestifolius (R. Knuth) Coode Collection. Mr. Murupb summit: 2400 m, Nooteboom & Chai 1986 (L). 52.1.4. Elaeocarpus sp. nov. aff. glaberrimus R. Knuth (fide Coode, pers. comm.) Collection. Mr. Murup summit: 2400 m, Nooteboom & Chai 1988 (L). 53. EPACRIDACEAE 53.1. STYPHELIA 53.1.1. Styphelia malayana (Jack) J. J. Sm. Collections. Mt. Murup: 2100 m, /lias S. 26516 (K, L); Mr. Murup Camp V: 2100- 2300 m, Burtt & Martin B. 5481 (E); Mr. Murup summit: 2400 m, Nooteboom & Chai 2022 (L). 54. ERICACEAE 54.1. DipLycosia 54.1.1. Diplycosia acuminata Becc. Collections. Mr. Murup Camp III: 1800 m, Burtt & Martin B. 5288 (E); Mr. Murup Camp IV: 1900 m, Burtt & Martin B. 5410 (E). 54.1.2. Diplycosia barbigera Sleumer Collection. Mr. MuruD/BELABAN River: 1900 m, /lias S$. 26373 (E, L, SAR). 54.1.3. Diplycosia fimbriata Sleumer Collections. Mr. Murup 2NpD summit: 2000 m, Ilias S. 26384 (E, K, L); Mr. Murup ABOVE Camp IV: 2000 m, Burtt & Martin B. 5421 (E); Mr. MURUD BETWEEN IsT AND 2ND suMMITs: 2300 m, Yii S. 44478 (E, K, L); Mr. Murup summit: 2400 m, Nooteboom & Chai 2002B (L, SAR), 2400 m, 2063 (L); Mr. Murup suMMIT RIDGE: 2300-2400 m, Beaman 11457 (K, MSC, UNIMAS). 54.1.4. Diplycosia cf. fimbriata Sleumer Collection. Mr. Murub ABOVE Camp III: 2000 m, Burtt & Martin B. 5266A (E). 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 117 54.1.5. Diplycosia aff. fimbriata Sleumer Collection. Mr. Murup SW oF Camp IV: 2100 m, Burtt & Martin B. 5358 (E). 54.1.6. Diplycosia microsalicifolia Argent Collections. Mr. MuruD ABOVE Camp IV: 2000 m, Burtt & Martin B. 5419 (E); Mr. MuRUD BETWEEN IST AND 2ND suMMITs: 2300 m, Yii S. 44493 (K); Mr. Murup, DapuR RIVER HEADWATERS: 2000 m, Ilias S. 26447 (E, K, isotypes; SAR, holotype of D. microsalicifolia Argent). 54.1.7. Diplycosia punctulata Stapf Collection. Mr. Murub 2ND summit: 2300 m, Yii 8. 44447 (E, K, SAR). 54.1.8. Diplycosia aff. saurauioides J. J. Sm. Collections. Mt. Murup 2NpD summit: 2000 m, /lias S. 26387 (L); Mr. MuRuD ABOVE Camp III: 2000 m, Burtt & Martin B. 5267 (E), 2000 m, 5268 (E, SAR); Mr. Murup BETWEEN IST AND 2ND SUMMITS: 2300 m, Yii S. 44490 (E, K, SAR); Mt. MurupD summit: 2400 m, Nooteboom & Chai 1992 (K, L, SAR); Mr. MurupD SUMMIT RIDGE: 2300-2400 m, Bea- man 11469 (K, MSC, UNIMAS). 54.1.9. Diplycosia cf. scabrida Becc. Collection. Mr. Murup ABOVE Camp IV: 2000 m, Burtt & Martin B. 5430 (E, SAR). 54.1.10. Diplycosia cf. urceolata Stapf Collections. Mr. Murup: 2300 m, /lias S. 26487 (E, K); Mr. Murup 2ND summit: 2000 m, /lias S. 26381 (E, K, SAR), 2200 m, S. 26416 (E, K, SAR), 2300 m, Yii S. 44433 (K); Mr. Murup N sive: 2300 m, Burtt & Martin B. 5452 (E, SAR); Mt. Murup ABove Camp III: 2000 m, Burtt & Martin B. 5266 (E, SAR). 54.1.11. Diplycosia indet. Collections. Mr. Murup: 2200-2300 m, Nooteboom & Chai 2056 (L); Mr. Murup ABOVE Camp III: 1800 m, Burtt & Martin B. 5253 (E). 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. Mr. Murup: 1900 m, Mjéberg 98 (UC); Mr. Murupb 2ND summit: 2200 m, Ilias S. 26403 (K); Mt. Murup ABovE Camp IV: 1800 m, Burtt & Martin B. 5251A (E); Mr. MURUD BETWEEN IsT AND 2ND SUMMITS: 2300 m, Yui S. 44460 (K, L). 54.2.2. Rhododendron burttii P. Woods Collection. Mr. Murup ABOVE LonG Rapata: 1700 m, Burtt & Martin B. 5549 (E, holotype of R. burttii P. Woods). 118 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 54.2.3. Rhododendron buxoides Sleumer Collections. Mr. MURUD NEAR suUMMIT: 2300 m, Nooteboom & Chai 1973 (K, isotype fragment; L, holotype of R. buxoides Sleumer); Mt. Murup summit: 2400 m, Burtt & Martin B. 5446 (E). 54.2.4. Rhododendron crassifolium Stapf [listed by Merrill (1928) as R. murudense Collections. Mr. Murup: 1900 m, Mjéberg 106 (L, fragment; UC, holotype of R. murudense Merr.); Mr. Murub SW or Camp II: 1700 m, Burtt & Martin B. 5321 (E); Mr. URUD SUMMIT RIDGE: 2300-2400 m, Beaman 11474 (K), 2300-2400 m, 7/478 (K); Mr. MUuRUuD, PATH TO SUMMIT: 1800 m, Nooteboom & Chai 1940 (L). 54.2.5. Rhododendron durionifolium Becc. [listed by Merrill (1928) as R. mjdbergii Merr. ]. Collections. Mr. Murup: 2300 m, /lias S. 26493 (K, L), 1900-2400 m, Mjdéberg 105 (L, fragment; UC, holotype of R. mjébergii Merr.); Mr. MuruD ABOVE Camp III: 1800 m, Burtt & Martin B. 5264 (E); Mr. Murup summit: 2400 m, Nooteboom & Chai 1989 (L); Mr. MURUD SUMMIT RIDGE: 2300-2400 m, Beaman 11447 (K, MSC, UNIMAS). 54.2.6. Rhododendron exuberans (Sleumer) Argent Collection. Mr. Murup Camp IV: 1800 m, Burtt & Martin B. 5351A (E). 54.2.7. Rhododendron himantodes Sleumer Collection. Mr. Murub Camp III: 1800 m, Burtt & Martin B. 5251 (E). 54.2.8. Rhododendron micromalayanum Sleumer Collections. Mr. Murup Camp IV: 1900 m, Burtt & Martin B. 5396 (E); Mr. Murup, LonG Rapata/BA KELALAN: 1700 m, Ilias S. 26555 (K); Mr. Murub, PATH TO suMMIT: 1800 m, Nooteboom & Chai 1955 (L). 54.2.9. Rhododendron moultonii Rid. Collection. Mr. Murub summit: 2400 m, Nooteboom & Chai 2006 (L). 54.2.10. Rhododendron orbiculatum Rid. Collections. Mr. MurRuD BETWEEN Ist AND 2ND sumMMITS: 2200 m, Yii S. 44602 (K, SAR); Mr. MuruD NEAR SUMMIT: 2300 m, Nooteboom & Chai 1972 (L); Mr. Murub suM- MIT RIDGE: 2300-2400 m, Beaman 11486 (K, MSC, UNIMAS): Mr. Murup, LonGc Rapata/ Ba KELALAN: 1700 m, /lias S. 26556 (K); Mt. Murub, PATH TO SUMMIT: 1900 m, Nooteboom & Chai 1967 (L). 54.2.11. Rhododendron pneumonanthum Sleumer [listed by Merrill (1928) as R. jasminiflorum Hook. ]. 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 119 Collections. Mr. Murup: 2300 m, /lias S. 26492 (K, L), 1900-2400 m, Mjéberg 103 (UC), 1900-2400 m, 704 (UC); Mr. Murup NEAR sumMIt: 2400 m, Nooteboom & Chai 1970 (L). 54.2.12. Rhododendron polyanthemum Sleumer Collection. Mr. Murup SW or Camp III: 1700 m, Burtt & Martin B. 5317 (E). 54.2.13. Rhododendron retivenium Sleumer Collection. Mr. Murup BELow Camp III: 1600 m, Burtt & Martin B. 5306 (E). 54.2.14. Rhododendron rugosum Low ex Hook. f. Collection. Mr. Murup summit: 2400 m, Burtt & Martin B. 5450 (E). §4.2.15. Rhododendron stenophyllum Hook. f. ex Stapf subsp. angustifolium (J. J. Smith) Argent, A. L. Lamb & Phillipps Collection. Mr. Murup SW or Camp III: 1700 m, Burtt & Martin B. 5322(E) 54.2.16. Rhododendron yongii Argent Collections. Mr. Murup: 2100 m, /lias S. 26462 (K); Mt. Murup SW oF Camp IV: 2100 m, Burtt & Martin B. 5365 (E). 54,.2.17. Rhododendron sp. nov. ? Collection. Mr. Murup SUMMIT RIDGE: 2300-2400 m, Beaman 11441 (IK, MSC, UNIMAS). 54.2.18. Rhododendron sp. (subsect. Pseudovireya) Collections. Mt. MuRUD SUMMIT RIDGE: 2300-2400 m, Beaman 11450 (E, K, UNI- MAS), 2300-2400 m, 1/467 (E, K, UNIMAS). 54,.2.19. Rhododendron indet. Collections. Mr. Murup: 2100 m, /lias S. 26464 (K); Mt. Murup SW oF Camp IV: 2100 m, Burtt & Martin B. 5363 (E); Mr. Murup aBove Camp III: 1800 m, Burtt & Martin B. 5250 (E), 1800 m, B. 5255 (E); Mt. Murup/BELABAN River: 1700 m, Ilias S. 26363 (K). 54.3. VACCINIUM 54.3.1. Vaccinium cf. bancanum Miq. Collections. Mt. MuRuD/BELABAN River: 1700 m, lias S. 263171 (K), 1800 m, S. 26364 (K). 54.3.2. Vaccinium bigibbum J. J. Sm. Collections. Mr. Murup 2ND summit: 2200 m, lias S. 26420 (K); Mr. MuRuD SW oF Camp III: 1700 m, Burtt & Martin B. 5326A (E); Mr. MuruD/BELABAN River: 1700 m, //ias S. 26310 (K). 120 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 54.3.3. Vaccinium cf. bigibbum J.J. Sm. Collections. Mr. Murup Camp IV: 1400 m, /ias S. 26433 (K); Mr. Murup SW or Camp III: 1700 m, Burtt & Martin B. 5323 (E). 54.3.4. Vaccinium claoxylon J. J. Sm. Collections. Mr. MurubD 2ND summit: 2300 m, Yii S. 44437 (K, L); Mr. Murup Camp IV: 2000 m, Ilias S. 26393 (K, L); Mr. Murup summit: 2400 m, Nooteboom & Chai 1990 (L), 2400 m, 2005 (L); Mr. Murub suMMIT RIDGE: 2300 m, Burtt & Martin B. 5491 (E); Mr. Murub/BELABAN River: 1700 m, /lias S. 26326 (K). 54.3.5. Vaccinium clementis Merr. Collections. Mr. Murup: 2200 m, Nooteboom & Chai 1975 (L); Mr. Murup 2np suMMIT: 2200 m, /lias S. 26410 (K, L), 2300 m, Yii 5S. 44436 (K); Mr. MuRUD SUMMIT RIDGE: 2300-2400 m, Beaman 11445 (K, MSC, UNIMAS) 54.3.6. Vaccinium moultonii Merr. Collection. Mr. Murub, PATH To suMMIT: 1800 m, Nooteboom & Chai 1944 (1), 54.3.7. Vaccinium pachydermum Stapf [listed by Merrill (1928)]. Collections. Mr. Murup: 2100 m, Ilias S. 26457 (K, L), 2300 m, S. 26482 (K, L), 1900-2400 m, Mjéberg 108 (K, UC); Mr. Murup 2Np summit: 2200 m, Ilias S. 26414 (K, L); Mt. Murup BETWEEN IsT AND 2ND SUMMITS: 2300 m, Yii S. 44476 (KK, L); Mr. Murup summit: 2400 m, Nooteboom & Chai 2002A (L); Mr. Murub/BELABAN River: 1900 m, Ilias S. 26378 (K, L). 54.3.8. Vaccinium aff. stapfianum Sleumer Collection. Mr. Murup Camp IV: 2000 m, Ilias S. 26398 (K). 54.3.9. Vaccinium tenerellum Sleumer Collections. Mr. Murup SW or Camp III: 1700 m, Burtt & Martin B. 5323A (E); Mr. MURUD ABOVE Camp IV: 2000 m, Burtt & Martin B. 5424 (E 54.3.10. Vaccinium cf. tenerellum Sleumer Collection. Mr. Murup Camp IV: 1900 m, Burtt & Martin B. 5411 (E). 54.3.11. Vaccinium sp. | [listed by Merrill (1928)]. Collection. Mr. Murup: Mjéberg 110 (UC). 54.3.12. Vaccinium sp. 2 [listed by Merrill (1928)]. Collection. Mr. Murup: Mjéberg 109 (UC). §4.3.13. Vaccinium indet. 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD IAL Collections. Mr. Murup Camp III: 1800-2000 m, Burtt & Martin B. 5240 (E); Mr. Murup Camp IV: 1800 m, Burtt & Martin B. 5349 (E); Mr. Murup SW or Camp IV: 2100 m, Burtt & Martin B. 5362 (E). 55. ESCALLONIACEAE 55.1. POLYOSMA 55.1.1. Polyosma mjébergii Merr. [listed by Merrill (1928); probably not distinct from P. bracteosa Stapf]. Collections. Mt. Murup: 2300 m, Ilias S. 26489 (K, SAR), 1900-2400 m, Mjéberg 107 (K, isotype; UC, holotype of P. mjébergii Merr.); Mt. MuRuD 2ND summit: 2300 m, Yu S. 44426 (K, L). 55.1.2. Polyosma sp. | Collections. Mr. Murup: 2200 m, Nooteboom & Chai 2046 (L); Mr. MurubD/BELABAN River: 1900 m, /lias S. 26375 (E, K, SAR). 55.1.3. Polyosma sp. 2 Collection. Mr. Murup: 2000 m, /lias S. 26388 (E, K). 55.1.4. Polyosma indet. Collection. Mr. MURUD BETWEEN IST AND 2ND suMMITs: 2200 m, Yii S. 44615 (SAR). 56. EUPHORBIACEAE 56.1. AUSTROBUXUS 56.1.1. Austrobuxus nitidus Miq. Collection. Mr. Murub/BA KELacan: 1700 m, Ilias S$. 26521 (SAR). 56.2. HOMALANTHUS 56.2.1. Homalanthus grandifolius Rid. Collection. Mr. Murup, PATH TO SUMMIT: 1700 m, Nooteboom & Chai 1912 (L). 57. FABACEAE 57.1. WHITFORDIODENDRON 57.1.1. Whitfordiodendron indet. 122 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Collection. Mr. Murupb RIDGE, SE sipe: 1700-2000 m, Beaman 1149] (K, MSC, UNI- MAS). 58. FAGACEAE 58.1. LirHOCARPUS 58.1.1. Lithocarpus luteus Soepadmo Collections. Mr. MuRuD/BELABAN River: 1700 m, /ias S. 26336 (K, SAR), 1700 m, S. 26325 (K). 58.2. QUERCUS 58.2.1. Quercus kinabaluensis Soepadmo Collections. Mr. MURUD NEAR sUMMIT: 2100 m, Ilias S. 26508 (K, SAR), 2100 m, /lias S. 26515 (K, SAR). 58.2.2. Quercus valdinervosa Soepadmo Collection. Mr. MuruD/BELABAN River: 1700 m, Ilias S. 26330 (K). 59. GENTIANACEAE 59.1. GENTIANA 59.1.1. Gentiana borneensis Hook. f. Collections. Mr. Murup: 2200 m, Nooteboom & Chai 2047 (L); Mr. Murup Camp V: 2100-2300 m, Burtt & Martin B. 5457 (E). 60. GESNERIACEAE 60.1. AESCHYNANTHUS 60.1.1. Aeschynanthus magnificus Stapf [listed by Merrill (1928) as Trichosporum mjObergti Merr.]. Collections. Mr. Murup: 1900 m, Mjéberg 87 (UC), 1900 m, 89 (BM, isotype: UC, holotype of Trichosporum mjébergii Merr.); Mt. Murup Camp III: Burtt & Martin B. 5226 (E); Mr. Murupb BELOow Camp III: 1500-1600 m, Burtt & Martin B. 5312 (Ei) 60.1.2. Aeschynanthus parvifolius R. Br. Collection. Mr. Murup Camp IH: 1800 m, Burtt & Martin B. 5227 (E). 60.1.3. Aeschynanthus speciosus Hook. Collection. Mr. Murup BELOW Camp III: 1500-1600 m, Burtt & Martin B. 5298 Cys 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 123 60.2. AGALMYLA 60.2.1. Agalmyla tuberculata Hook. f. Collections. Mr. Murup: 2400 m, Nooteboom & Chai 2007 (L, SAR); Mr. Murup N sIDE: 2100 m, Burtt & Martin B. 5459 (SAR); Mr. Murup aBovE Camp III: 1800 m, Burtt & Martin B. 5249 (SAR). 60.3. CYRTANDRA 60.3.1. Cyrtandra basiflora C. B. Clarke Collection. Mr. Murup: 1600 m, Yii S. 44414 (E). 60.3.2. Cyrtandra aff. cuprea B. L. Burtt Collection. Mr. Murup NEAR Camp III: 1800 m, Burtt & Martin B. 5270 (E, SAR). 60.3.3. Cyrtandra dolichopoda B. L. Burtt Collection. Mr. Murup BELow Camp III: 1600 m, Burtt & Martin B. 5307 (E, SAR). 60.3.4. Cyrtandra horizontalis B. L. Burtt Collection. Mr. Murup RIDGE, SE ripGe: 1700-2000 m, Beaman 11493 (K, MSC, UNIMAS). 60.3.5. Cyrtandra aff. lacerata B. L. Burtt Collection. Mt. MuRUD BELOW Camp III: 1600 m, Burtt & Martin B. 5309 (E). 60.3.6. Cyrtandra aff. pedicellata B. L. Burtt Collection. Mr. MuRuD BELOW Camp IV: 1800 m, Burtt & Martin B. 5377 (E). 60.3.7. Cyrtandra aff. trisepala C. B. Clarke Collections. Mr. Murup: 1700 m, Nooteboom & Chai 1914 (L), 1600 m, Yii S. 44416 (E, L, SAR); Mr. Murup BELow Camp IV: 1800 m, Burtt & Martin B. 5375 (E, SAR); Mr. Murup NEAR Camp II: 1800 m, Burtt & Martin B. 5280 (E). 60.3.8. Cyrtandra sp. (sect. Decurrentes) Collection. Mr. MuruD BELOW Camp III: 1500-1600 m, Burtt & Martin s.n. (E). 60.4. DIDYMOCARPUS 60.4.1. Didymocarpus simplex Kraenzl. Collections. Mr. MurupD NEAR Camp III: 1700 m, Burtt & Martin B. 5246 (E); Mr. Murupb NEAR Camps III AND IV: 1800-2000 m, Burtt & Martin B. 5275 (E, SAR); Mr. MurRubD, PATH TO SUMMIT: 1800 m, Nooteboom & Chai 1950 (L). 124 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 60.4.2. Didymocarpus indet. Collection. Mr. Murup: Mjéberg 204 p.p. [UC; specimen too imperfect to warrant identification beyond genus, fide Merrill (1928)]. 60.5. LoxocaRPus 60.5.1. Loxocarpus indet. Collections. Mr. Murup BeELow Camp III: 1500-1600 m, Burtt & Martin B. 5313 (E. SAR); Mr. Murup NEAR Camp II: 1800 m, Burtt & Martin B. 5276 (E). 61. HYDRANGEACEAE 61.1. DicHROA 61.1.1. Dichroa febrifuga Lour. Collections. Mr. Murup: 1600 m, Yii S$. 44422 (i); Mr. Murup Camp IV: 1400 m, /lias S. 26421 (K); Mt. Murub NEAR Camp III: 1800-2000 m, Burtt & Martin B. 5308 (a. 62. ILLICIACEAE 62.1. ILLictum 62.1.1. Ilicium cauliflorum Merr. [listed by Merrill (1928)]. Collections. Mr. Murup: 1900-2400 m, Mjéberg 114 (UC, holotype of 1. cauliflorum Merr.); Mr. Murup BeLow Camp [V: 1800 m, Burtt & Martin B. 5371 (E). 62.1.2. Ilicitum tenuifolium A. C. Sm. Collection. Mr. Murub, PATH To suMMiIT: 1700 m, Nooteboom & Chai 1923 Cee 63. LAURACEAE 63.1. LINDERA 63.1.1. Lindera bibracteata (Blume) Boerl. var. rufa (Stapf) Kosterm. Collections. Mt. MuRUD SUMMIT RIDGE: 2300-2400 m, Beaman 11473 (K); Mr. Murup, PATH TO SUMMIT: 1800 m, Nooteboom & Chai 1952 (L). 63.1.2. Lindera subumbelliflora (Blume) Kosterm. Collection. Mr. Murup, PATH To summit: 1900 m, Nooteboom & Chai 1963 (ig). 64. LOGANIACEAE 64.1. FAGRAEA 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 125 64.1.1. Fagraea blumei G. Don Collection. Mr. MuRUD NEAR suMMIT: 2200 m, Nooteboom & Chai 1974 (L). 64.1.2. Fagraea oreophila Wong & Sugau [listed by Merrill (1928) as F. obovata Wall. in Roxb.]. Collections. Mr. Murup: 1900 m, Mjéberg 94 (UC), Yii S. 44474 (K); Mr. Murup SUMMIT: 2400 m, Nooteboom & Chai 20/1 (L). 64.2. GENIOSTOMA 64.2.1. Geniostoma rupestre Forst. Collections. Mr. Murup: 2200 m, Nooteboom & Chai 205] (L); Mr. Murup Camp IV: 2000 m, /lias S. 26397 (L). 65. LORANTHACEAE 65.1. HELIXANTHERA 65.1.1. Helixanthera cf. cylindrica (Jack) Danser Collections. Mr. MURUD BETWEEN IsT AND 2ND suMMITs: 2200 m, Yii S. 44620 (SAR); Mr. MuRubD, PATH TO SUMMIT: 1900 m, Nooteboom & Chai 1965 (L, SAR). 65.2. MACROSOLEN 65.2.1. Macrosolen cochinchinensis (Lour.) Tieghem Collections. Mr. Murup SW oF Camp IV: 2100 m, Burtt & Martin B. 5364 (E); Mr. Murup summit: 2400 m, Nooteboom & Chai 1977 (K, L). 65.2.2. Macrosolen flammeus Danser Collection. Mr. Murup summit: 2400 m, Nooteboom & Chai 2062 (L). 66. MELASTOMATACEAE 66.1. MEDINILLA 66.1.1. Medinilla crassifolia (Reinw. ex Blume) Blume Collection. Mr. Murup: 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 homoeandrum Stapf]. Collection. Mr. Murup: 1900-2400 m, Mjéberg 96 (K, UC). 126 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 66.2. MELASTOMA 66.2.1. Melastoma pulcherrimum Koord. Collections. Mr. Murup: 2200 m, Nooteboom & Chai 2024 (L); Mr. Murup Camp IV: 2000 m, Ilias S. 26399 (K). 66.3. PHYLLAGATHIS 66.3.1. Phyllagathis brookei Nayar Collections. Mr. Murup: 1900 m, /lias S. 26518 (L); Mr. MurRuD NEAR Camp IIT: 1800 m, Burtt & Martin B. 5278 (E). 66.4. PLETHIANDRA 66.4.1. Plethiandra hookeri Stapf Collections. Mr. Murub 2NpD sumMIt: Ilias S. 26407 (K); Mr. MURUD BETWEEN IST AND 2ND suMMiITS: 2300 m, Yii S. 44483 (K); Mr. Murup SUMMIT RIDGE: 2300-2400 m, Beaman 11475 (K, MSC, UNIMAS). 66.5. SARCOPYRAMIS 66.5.1. Sarcopyramis napalensis Wall. Collections. Mr. Murup: 1600 m, Yii S. 444/77 (K, L); Mr. Murup BeLtow Camp III: 1600 m, Burtt & Martin B. 5297 (BE). 66.6. SONERILA 66.6.1. Sonerila crassiuscula Stapf Collection. Mr. Murup summit: 2400 m, Nooteboom & Chai 1982 (L); Mr. Murupb SUMMIT RIDGE: 2300-2400 m, Beaman 11460 (FLAS). 66.6.2. Sonerila macrantha Merr. [listed by Merrill (1928); probably not distinct from S. tenuifolia Blume]. Collections. Mr. Murup: 2300 m, Ilias S. 26484 (L), 1900-2400 m, Mjdberg 74 (NY, UC), 1900-2400 m, 75 (UC, holotype of S$. macrantha Merr.), 1900-2400 m, 76 (UC), 1900-2400 m, 78 (UC); Mr. Murupb 2Np summit: 2300 m, Yii S$. 44434 (SAR); Mr. Murup N sive: 2300 m, Burtt & Martin B. 5449 (E, SAR); Mr. Murup aBpove Camp IV: 2000 m, Burtt & Martin B. 5345 (E); Mr. MuruD BETWEEN LoNnG Rapata AND Camp III: 1500 m, Burtt & Martin B. 5223 (E). 66.6.3. Sonerila nervulosa Rid. Collection. Mr. MurupD NEAR Camp IIT: 1800 m, Burtt & Martin B. 5254 (E, SAR). 66.6.4. Sonerila pulchella Stapf 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 27, Collection. Mt. MURUD RIDGE, SE sipE: 1700-2000 m, Beaman 11494 (K). 66.6.5. Sonerila sp. ? Collection. Mt. Murup: Mjéberg 77 (UC; noted by Merrill (1928) to be a single imperfect specimen]. 67. MORACEAE 67.1. Ficus 67.1.1la. Ficus deltoidea Jack var. deltoidea Collections. Mr. Murup, PATH TO suMMIT: 1700 m, Nooteboom & Chai 1916 (L); Mr. Murub/PARAMUSU River: 1500 m, Yii S. 44646 (L). 67.1.1b. Ficus deltoidea Jack var. intermedia Corner Collections. Mr. Murup: 2200 m, Nooteboom & Chai 2028 (L); Mr. Murup Camp 4: 2000 m, Ilias S. 26400 (K); Mr. MurRUD BETWEEN IST AND 2ND SUMMITS: 2200 m, Yii S. 4460] (K); Mr. MuruD BETWEEN CAMP V AND summit: Burtt & Martin B. 5492 (E); Mr. Murup SUMMIT: 2400 m, Nooteboom & Chai 2066 (L); Mr. Murupb, PATH To sUMMIT: 1800 m, Nooteboom & Chai 1957 (L). 67.1.1c. Ficus deltoidea var. kinabaluensis Corner Collections. Mr. Murup: 2100 m, Ilias S. 26450 (K); Mr. Murup 2np summit: 2200 m, lias §. 26417 (K); Mr. Murup Camp IV: 2000 m, /lias S. 26392 (K); Mt. MuRUD BETWEEN Camp V AND suMMIT: Burtt & Martin B. 5489 (E); Mr. MurRuD NEAR Camp V: 2100 m, /lias S. 26514 (KK); Mr. Murup summit: 2400 m, Nooteboom & Chai 2001 (L). 67.1.2. Ficus oleaefolia King var. valida Corner Collections. Mr. Murup Camp III: 2000 m, Burtt & Martin B. 5254 (E); Mr. Murup BETWEEN Ist AND 2ND SUMMITS: 2200 m, Yii S. 44604 (K); Mt. MuRuD/BELABAN River: 1900 m, flias S. 26376 (K). 67.1.3. Ficus recurva Blume var. ribesioides (Wall. ex Miq.) King Collection. Mr. MuruD, PATH TO SUMMIT: Nooteboom & Chai 1929 (L). 68. MYRICACEAE 68.1. MyrIca 68.1.1. Myrica javanica Blume Collections. Mr. Murup 2np summit: 2000 m, /lias S. 26382 (L); Mr. Murup suMMIT: 2400 m, Anderson & Ilias S. 26477 (L), 2400 m, Nooteboom & Chai 1994 (L). 128 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 69. MYRSINACEAE 69.1. ARDISIA 69.1.1. Ardisia lepidotula Merr. Collections. Mr. Murup Camp III: 1600 m, Burtt & Martin B. 5261 (E); Mr. Murup, RIDGE SW oF Camp IV: 2100 m, Burtt & Martin B. 5353 (E). 69.1.2. Ardisia mjébergii Merr. [listed by Merrill (1928)]. Collection. Mr. Murup: 1900-2400 m, Mjdéberg 80 (UC, holotype of A. mjébergii Merr.). 69.1.3. Ardisia obovatifolia Merr. [listed by Merrill (1928)]. Collections. Mr. Murup: 1900-2400 m, Mjéberg 79 (K, isotype; UC, holotype of A. obovatifolia Merr.); Mr. MuRUD, RIDGE ABOVE CAMP IV: 1900 m, Burtt & Martin B. 5412 (E). 69.1.4. Ardisia virens Kurz Collection. Mr. MuRUD BETWEEN Camps II anp HI: 1500 m, Burtt & Martin B. 5222 (EB). 69.1.5. Ardisia indet. Collections. Mr. Murup: 2200 m, /lias S. 26496 (L); Mr. Murup summit: 2400 m, Nooteboom & Chai 1978 (L); Mr. Murup, PATH To suMMIT: 1700 m, Nooteboom & Chai POOR 1700 a, 2915 (eT 700 mi, 7918 (1) L700 mi 2919 (a 69.2. EMBELIA 69.2.1. Embelia minutifolia Stapf Collections. Mr. Murup ABove Camp IV: 2000 m, Burtt & Martin B. 5346 (E); Mr. MuRUD BETWEEN IST AND 2ND suMMITS: 2200 m, Yii S. 44609 (K, L), 2200 m, 44610 (K, L); Mr. Murup SUMMIT RIDGE: 2300-2400 m, Beaman 11456 (K, UNIMAS); Mr. Murub/BELA- BAN River: 1900 m, /lias S$. 26379 (L). 69.2.2. Embelia spiraeoides Stapf Collections. Mr. Murup, Dapur River HEADWATERS: 2400 m, /lias S. 26532 (KK, L); Mr. Murup, N sIDE NEAR Camp V: 2100 m, Burtt & Martin B. 5445 (E). 69.2.3. Embelia tortuosa Stapf Collections. Mt. Murup N sipe: 2100 m, Burtt & Martin B. 5473 (E); Mr. Murup BETWEEN IST AND 2ND SUMMITS: 2300 m, Yii S. 44487 (K); Mr. MuruD SUMMIT RIDGE: 2300- 2400 m, Beaman 11472 (K, MSC, UNIMAS). 69.2.4. Embelia indet. Collections. Mr. Murup: 2200 m, /lias S. 2650/7 (L); Mr. Murup summit: 2400 m, Nooteboom & Chai 2067 (L). 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 129 69.3. MAESA 69.3.1. Maesa indica A. DC. Collection. Mr. Murup Camp III: 1800 m, Burtt & Martin B. 5334 (E). 69.4. MyRSINE 69.4.1. Myrsine cruciata (Philipson) Pipoly Collections. Mr. MuRuUD ABOvE Camp IV: 2100 m, Burtt & Martin B. 5357 (E); Mr. MURUD BETWEEN IsT AND 2ND SUMMITS: 2300 m, Yii S. 44468 (K), 2300 m, S. 44473 (K); Mr. MURUD SUMMIT RIDGE: 2400 m, Burtt & Martin B. 5490 (E). 70. MYRTACEAE 70.1. BAECKEA 70.1.1. Baeckea taxifolia Merr. [listed by Merrill (1928)]. Collections. Mr. Murup: 2400 m, Mjdberg 84 (UC, syntype of B. taxifolia Merr.), 2400 m, /// (K, isosyntype; UC, syntype of B. taxifolia Merr.), 2200-2300 m, Nooteboom & Chai 2016 (L); Mr. Murup, Camp V To summit: Burtt & Martin B. 5493 (E). 70.2. LEPTOSPERMUM 70.2.1. Leptospermum javanicum Blume [listed by Merrill (1928)]. Collections. Mr. Murup: 1500 m, Mjéberg 92 (UC); Mr. Murup Camp IV: 1900 m, Burtt & Martin B. 5425 (E); Mr. Murup NEAR summit: 2100 m, /lias S$. 26506 (L); Mr. Murub summit: 2400 m, Anderson & Ilias S. 26480 (L), 2400 m, Nooteboom & Chai 1983 (L); Mr. Murub sSuMMIT RIDGE: 2300-2400 m, Beaman 11442 (K, MSC, UNIMAS). 70.3. SYZYGIUM 70.3.1. Syzygium aff. ampullarium (Stapf) Merr. & Perry Collections. Mr. Murup: 2100 m, /lias S. 26458 (K, L); Mt. Murup 2NpD summit: 2000 m, Hias S. 26385 (K, L), 2200 m, S. 26406 (K, L), 2300 m, Yii S. 44428 (K); Mr. MuruD ABOVE Camp IV: 2100 m, Burtt & Martin B. 5360 (E); Mr. MuRUD BETWEEN IST AND 2ND SUMMITS: 2300 m, Yi S. 44467 (K), 2300 m, S. 44469 (K); Mr. Murup summit: 2400 m, Anderson & Ilias S. 26472 (K, L), 2400 m, Nooteboom & Chai 1984 (L), 2400 m, 1985 (L), 2400 m, 2072 (L); Mr. MURUD SUMMIT RIDGE: 2300-2400 m, Beaman 11446 (K, MSC, UNIMAS). 70.3.2. Syzygium castaneum (Merr.) Merr. & Perry Collections. Mt. Murup Camp IV: 2000 m, /lias S. 26389 (K, L); Mr. MurRuD SUMMIT RIDGE: 2300-2400 m, Beaman 11462 (K, MSC, UNIMAS): Mt. Murub/BELABAN RIVER: 1900 m, Ilias S. 26380 (K, L). 130 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 70.3.3. Syzygium indet. Collections. Mt. Murub, PATH TO SUMMIT: 1700 m, Nooteboom & Chai 1931 (L), 1800 m, 1941] (L); Mr. Murup/BELABAN RIVER HEADWATERS: 1700 m, /lias S. 26308 (L). 70.4. TRISTANIOPSIS 70.4.1. Tristaniopsis elliptica (Stapf) Peter G. Wilson & J.T. Waterh. Collections. Mr. MuruD NEAR Camp V: 2100 m, /lias S. 26507 UK); Mr. Murup: 2300 m, Ilias 8. 26483 (IK); Mr. Murub, PATH TO SUMMIT: 1800 m, Nooteboom & Chai 1966 (K, L). 70.4.2. Tristaniopsis indet. Collection. Mr. Murup: 2200 m, Nooteboom & Chai 2041 (L). 70.5. XANTHOMYRTUS 70.5.1. Xanthomyrtus flavida (Stapf) Diels Collections. Mr. Murup: 2400 m, /lias S. 26534 (L); Mr. Murup summit: 2400 m, Nooteboom & Chai 1997 (L); Mr. Murub, RIDGE ABOVE Camp IV: 1900 m, Burtt & Martin iB, 3401 1) 70.5.2. Xanthomyrtus moultonii (Merr.) Merr. Collection. Mt. Murup summit: 2400 m, Nooteboom & Chai 2000 (L). 71. NEPENTHACEAE 71.1. NEPENTHES 71.1.1. Nepenthes lowii Hook. f. [listed by Merrill (1928)]. Collections. Mr. Murub: Mjéberg 1/5 (UC); Mr. Murub Came V: 2100-2300 m, Burtt & Martin B. 5477 (E); Mr. Murup NEAR Camp IIT: 1800 m, Burtt & Martin B. 5325 (E); Mr. MuruD SUMMIT RIDGE: 2300-2400 m, Beaman 11476 (K, UNIMAS); Mr. Murub, PATH TO SUMMIT: 1900 m, Nooteboom & Chai 1962 (L). 71.1.2. Nepenthes murudensis Culham ex Jebb & Cheek Collections. Mr. Murup Camp IV: 2000 m, Burtt & Martin B. 5427 (E); Mr. Murup Camp V: 2100-2300 m, Burtt & Martin B. 5467 (E), 2100-2300 m, B. 5477 (E); Mr. MurRup BETWEEN IST AND 2ND SUMMITS: 2200 m, Yii S. 44623 (SAR); Mr. MuRUD NEAR CAMP V: 2100 m, Ilias §. 26513 (SAR); Mr. Murup NEAR suMMIT: 2400 m, Nooteboom & Chai 2035 (SAR); Mr. MuRuD SUMMIT RIDGE: 2300-2400 m, Beaman 11461 (K, MSC, UNIMAS). 71.1.3. Nepenthes indet. Collection. Mr. Murup: Mjdéberg 125 [UC; noted by Merrill (1928) to be only a detached inflorescence]. 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 134 72. OLEACEAE 72.1. CHIONANTHUS 72.1.1. Chionanthus enerve (Steenis) Kiew Collections. Mr. Murup Camp V: 2100-2300 m, Burtt & Martin B. 5478 (E); Mr. MuRUD BETWEEN IST AND 2ND suMMiITS: 2200 m, Yii S. 44617 (K, L); Mr. MuruD SUMMIT RIDGE: 2300-2400 m, Beaman 114517 (K, MSC, UNIMAS); Mr. Murup, N oF summit: 2300 m, Nooteboom & Chai 2049 (L). 72.2. JASMINUM 72.2.1. Jasminum oreophilum Kiew Collections. Mr. Murup Camp III: 1700 m, Burtt & Martin B. 5234 (E); Mr. Murup/ BELABAN River: 1700 m, /lias S. 26324 (K). 73. PENTAPHRAGMATACEAE 73.1. PENTAPHRAGMA 73.1.1. Pentaphragma cf. acuminatum Airy Shaw Collections. Mr. Murup SW oF Camp III: 1700 m, Burtt & Martin B. 5316 (E); Mr. MuRUD RIDGE, SE sipe: 1700-2000 m, Beaman 11490 (K, MSC, UNIMAS). 73.1.2. Pentaphragma aurantiaca Stapf [listed by Merrill (1928) as P. obtusifolium eae Collections. Mr. Murup: 2300-2400 m, Burtt & Martin B. 5451 (E), 2400 m, Ilias S. 26537 (L), 1900 m, Mjéberg 112 (UC, holotype of P. obtusifolium Merr.); Mr. Murupb suMMIT: 2400 m, Nooteboom & Chai 2008 (L); Mt. MurupD SUMMIT RIDGE: 2300-2400 m, Beaman 11443 (K, MSC, UNIMAS). 73.1.3. Pentaphragma longisepalum Kiew Collections. Mr. Murup Camp IV: 1900 m, Burtt & Martin B. 5426 (E); Mr. Murup BETWEEN Camps IT AND II: 1500 m, Burtt & Martin B. 5225 (E). 74, PIPERACEAE 74.1. PEPEROMIA 74.1.1. Peperomia laevifolia (Blume) Mig. Collection. Mr. MURUD NEAR Camp III: 1800 m, Burtt & Martin B. 5331 (E). 74.1.2. Peperomia indet. Collection. Mr. MuruD, PATH TO SUMMIT: 1700 m, Nooteboom & Chai 1924 (L). 132 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 74.2. PIPER 74.2.1. Piper cf. caninum Blume Collections. Mr. Murup BELOW Camp III: 1600 m, Burtt & Martin B. 5310 (E); Mr. MUuRUD RIDGE, SE sipE: 1700-2000 m, Beaman 11492 (K, UNIMAS). 75. POLYGALACEAE 75.1. POLYGALA 75.1.1. Polygala oreotrephes B. L. Burtt [listed by Merrill (1928) as Polygala sp.]. Collections. Mr. Murup: 1500 m, Mjdberg 91 (UC), 2200 m, Nooteboom & Chai 1971 (L); Mr. Murup NEAR Camp ITI: 1800-2300 m, Burtt & Martin B. 5324 (E). 76. RHAMNACEAE 76.1. RHAMNUS 76.1.1. Rhamnus borneensis Steenis Collections. Mr. Murup: 2300 m, [lias S. 26485 (K); Mr. MuRUD 2ND suMMIT: 2200 m, Ilias S. 26419 (K); Mr. Murup N sipe: 2100 m, Burtt & Martin B. 546] (E); Mr. Murup SW oF Camp IV: 2100 m, Burtt & Martin B. 5361 (E); Mr. Murup summit: 2400 m, Nooteboom & Chai 1996 (K). 77. ROSACEAE 77.1. PRUNUS 77.1.1. Prunus arborea (Blume) Kalkman var. densa (King) Kalkman Collections. Mr. Murup summit: 2400 m, Nooteboom & Chai 1991 (L); Mr. Murupb SUMMIT RIDGE: 2300-2400 m, Beaman 11452 (K, MSC, UNIMAS); Mr. Murupb, PATH TO SUMMIT: L900 m, Nooteboom & Chai 1951 (L). 77.2. RUBUS 77.2.1. Rubus alpestris Blume Collection. Mr. Murup summit: 2400 m, Nooteboom & Chai 1993 (L). 78. RUBIACEAE 78.1. ACRANTHERA 78.1.1. Acranthera aff. atropella Stapf Collections. Mr. MuruD NEAR Camp III: 1800 m, Burtt & Martin B. 5336 (E); Mr. MUuRUD RIDGE, SE sipe: 1700-2000 m, Beaman 11488 (K, MSC, UNIMAS). 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 133 78.2. ARGOSTEMMA 78.2.1. Argostemma borragineum Blume ex DC. Collection. Mr. Murup: 1600 m, Yii S. 44470 (K). 78.2.2. Argostemma hameliifolium Wernham Collection. Mr. Murup: 1600 m, Yii S. 44409 (K). 78.2.3. Argostemma moultonii Ridl. [listed by Merrill (1928) as A. murudensis Merr.]. Collections. Mt. Murup: 1900-2400 m, Mjéberg 82 (K, isotype; UC, holotype of A. murudensis Merr.), 1900-2400 m, 88 (UC); Mr. Murup NEAR Camp ITT: 1800 m, Burtt & Martin B. 5326 (E); Mr. Murup, PATH TO summit: 1700 m, Nooteboom & Chai 1935 (L). 78.3. CEPHAELIS 78.3.1. Cephaelis stipulacea Blume [this species probably belongs in the genus Chassalia]. Collection. Mr. MuRuD NEAR Camp III: 1800 m, Burtt & Martin B. 5285 (E). 78.4. DiscoSsPERMUM 78.4.1. Discospermum abnorme (Korth.) S. J. Ali & Robbr. Collection. Mt. Murup BELow Camp IV: 1800 m, Burtt & Martin B. 5373 (EB). 78.5. HEDYOTIS 78.5.1. Hedyotis pulchella Stapf Collections. Mr. MuruD BELOW Camp III: 1600 m, Burtt & Martin B. 5291 (E); Mr. Murup, Dapur RIVER HEADWATERS: 1900 m, Ilias S. 26517 (K). 78.5.2. Hedyotis sp. 1 Collections. Mt. Murup: Burtt & Martin B. 5448 (E), 2100 m, Ilias S. 26465 (K, L). 78.5.3. Hedyotis indet. Collection. Mr. Murup: 2200 m, Nooteboom & Chai 2021 (L). 78.6. IxORA 78.6.1. Ixora sessililimba Merr. [listed by Merrill (1928)]. Collections. Mt. Murup: 1900-2400 m, Mjéberg 116 (UC, holotype of I. sessililimba Merr.); Mr. Murup NEAR Camp III: 1800 m, Burtt & Martin B. 5340 (BE). 134 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 78.7. LUCINAEA 78.7.1. Lucinaea pentacme Stapf Collections. Mr. Murub NEAR Camp IV: 1900 m, Burtt & Martin B. 5409 (E); Mr. Murub/BELABAN River: 1900 m, /lias S$. 26374 (K). 78.8. OPHIORRHIZA 78.8.1. Ophiorrhiza aff. fibrillosa Ridl. Collections. Mr. Murup: 1600 m, Yii S. 44412 (K); Mr. Murupb BELow Camp III: 1600 m, Burtt & Martin B. 5296 (E). 78.9, PSYCHOTRIA 78.9.1. Psychotria densifolia Stapf Collections. Mr. Murub summit: 2400 m, Nooteboom & Chai 1979 (L); Mr. Murup SUMMIT RIDGE: 2300-2400 m, Beaman 11481 (K, MSC, UNIMAS). 78.9.2. Psychotria sp. Collection. Mr. Murub, PATH TO suMMIT: 1700 m, Nooteboom & Chai 1925 emp 78.10. RUBIA 78.10.1. Rubia cordifolia L. Collection. Mr. MuRub, PATH TO SUMMIT: 1700 m, Nooteboom & Chai 1926 (L). 78.11. STREBLOSA 78.11.1. Streblosa urticina Stapf Collection. Mr. MuruD NEAR Camp ILE: 1800 m, Burtt & Martin B. 5335 (EB). 78.12. TIMONTUS 78.12.1. Timonius esherianus W. W. Sm. Collection. Mr. Murub, PATH TO suMMIT: 1700 m, Nooteboom & Chai 1928 (L). 79. RUTACEAE 79.1. EVODIA 79.1.1. Euodia indet. Collection. Mr. Murup: 2200-2300 m, Nooteboom & Chai 2050 (L). 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 135 80. SYMPLOCACEAE 80.1. SympLocos 80.1.1. Symplocos adenophylla Wall. ex G. Don var. adenophylla Collections. Mr. Murup Camp IV: 1400 m, Ilias S. 26434 (K); Mr. Murub/BELABAN RIVER HEADWATERS: 1700 m, Ilias S. 26306 (K, SAR), S. 26352 (K), 1700 m, S. 26353 (SAR). 80.1.2. Symplocos anomala Brand 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.] Collections. Mt. Murup: 1900-2400 m, Mjdberg 120 (UC, holotype of S. dolichantha Merr.); Mt. Murub, PATH TO summit: 1700 m, Nooteboom & Chai 1932 (L). 80.1.4a. Symplocos laeteviridis Stapf var. mjébergii (Merr.) Noot. [listed by Mer- rill (1928) as S. mjébergii Merr.]. Collections. Mr. Murup: 1900-2400 m, Mjdéberg 97 (K, isotype; UC, holotype of S. mjébergii Merr.); Mt. MURUD, PATH TO SUMMIT: 1500 m, Nooteboom & Chai 1909 (K, L). 80.1.4b. Symplocos laeteviridis Stapf var. pauciflora Noot. Collections. Mt. Murup: 2000 m, /lias S. 26391 (K, L), 1700-2000 m, S. 26392 (?); Mr. Murup Camp IV: 1900 m, Burtt & Martin B. 5389 (E); Mr. Murup/BELABAN River: 1700 m, Ilias S. 26334 (K, L). 80.1.5. Symplocos pendula Wight var. hirtistylis (Clarke) Noot. Collections. Mt. Murup: Nooteboom & Chai 2058 (K); Mt. Murup aBove Camp IV: 2000 m, Burtt & Martin B. 5442 (E). 80.1.6. Symplocos tricoccata Noot. Collection. Mt. MuRUD/BELABAN RIVER HEADWATERS: 1700 m, /lias S. 26305 (K, L, SAR). 81. THEACEAE 81.1. ADINANDRA 81.1.1. Adinandra clemensiae Kobuski Collections. Mt. Murup: 1600 m, Yii S. 44499 (K); Mr. MuruD BETWEEN IsT AND 2ND suMMITS: 2200 m, Yii S. 446/71 (K). 81.1.2. Adinandra cf. dumosa Jack Collection. Mr. Murup SUMMIT RIDGE: 2300-2400 m, Beaman 11477 (K, MSC, UNIMAS). 136 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 81.2. SCHIMA 81.2.1. Schima wallichii (DC.) Choisy subsp. brevifolia (Hook. f.) Bloemb. Collections. Mr. Murup: 2200 m, Nooteboom & Chai 2045 (L); Mr. Murup N sIDE: 2100 m, Burtt & Martin B, 5469 (E). 81.3. TERNSTROEMIA 81.3.1. Ternstroemia beccarii Stapf Collections. Mr. Murup: 2300 m, /lias S. 26490 (K), 2200 m, Nooteboom & Chai 2031 (L); Mr. Murub 2npd summit: 2300 m, Yii S. 44439 (K), 2300 m, S. 44448 (K); Mr. Murup ABOVE Camp IV: 2000 m, Burtt & Martin B. 5438 (E); Mr. MuruD BETWEEN IST AND 2ND SUMMITS: 2300 m, Yii S. 44463 (L); Mr. Murupb suMMIT RIDGE: 2300-2400 m, Beaman 11463 (K, MSC, UNIMAS); Mr. Murupb, PATH TO suMMiT: 1800 m, Nooteboom & Chai 1948 (L). 81.3.2. Ternstroemia denticulata (Pierre) Rid. Collections. Mr. Murup: 2200 m, Ilias S. 26402 (L), 2100 m, S. 26454 (L); Mt. Murup BETWEEN Ist AND 2ND SUMMITS: 2300 m, Yii S. 44484 (K), 2300 m, S. 44485 (K), 2200 m, S. 44605 (K); Mr. Murup summit: 2400 m, Nooteboom & Chai 1980 (L); Mt. Murub, PATH TO SUMMIT: 1800 m, Nooteboom & Chai 1956 (L). 81.3.3. Ternstroemia lowii Stapf Collections. Mr. Murup ABOvE Camp IV: 2000 m, Burtt & Martin B. 5436 (E); Mr. MURUD SUMMIT RIDGE: 2300-2400 m, Beaman 11453 (K, MSC, UNIMAS). 81.3.4. Ternstroemia indet. Collection. Mr. MurupD/BELABAN River: 1700 m, Ilias S. 26309 (L). 82. THYMELIACEAE 82.1. WIKSTROEMIA 82.1.1. Wikstroemia brachyantha Merr. Collections. Mr. Murup: 1600 m, Yii S. 44420 (K); Mr. MurupD BELOW Camp IV: 1800 m, Burtt & Martin B. 5374 (E). 82.1.2. Wikstroemia cf. indica (L.) C. A. Mey. Collections. Mr. Murup: 2100 m, Ilias S. 26459 (K, L); Mr. Murup N oF summit: 2300 m, Nooteboom & Chai 2038 (L), 2300 m, 2040 (L). 83. URTICACEAE 83.1. ELATOSTEMA 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 137 83.1.1. Elatostema acuminatum (Poir.) Brongn. Collection. Mr. Murup BELOW Camp III: 1600 m, Burtt & Martin B. 5311 (E). 83.1.2. Elatostema cf. lineare Stapf Collection. Mt. Murup, Camp III/LoneG Rapata: 1500 m, Burtt & Martin B. 5224 (E). 83.1.3. Elatostema penibukanense Gibbs Collections. Mt. Murup Camp III: 1800 m, Burtt & Martin B. 5282 (E); Mr. Murupb ABOVE Camp IV: 2000 m, Burtt & Martin B. 5347 (E). 83.1.4. Elatostema sp. | Collections. Mr. Murup Camp IV: 1900 m, Burtt & Martin B. 5386 (E); Mr. Murup ABOVE Camp III: 1800 m, Burtt & Martin B. 5257 (E). 83.1.5. Elatostema sp. 2 Collection. Mr. Murup Camp IV: 1900 m, Burtt & Martin B. 5387 (E). 83.1.6. Elatostema indet. Collection. Mr. Murup summit: 2400 m, Nooteboom & Chai 2059 (L). 83.2. PILEA 83.2.1. Pilea sp. Collection. Mr. Murup BELow Camp IV: 1800 m, Burtt & Martin B. 5367 (BE). 84. VERBENACEAE 84.1. CLERODENDRUM 84.1.1. Clerodendrum kinabaluense Stapf Collection. Mr. Murup Camp [V: 1900 m, Burtt & Martin B. 5413 (BE). 85. WINTERACEAE 85.1. Drimys 85.1.1. Drimys piperita Hook. f. [listed by Merrill (1928)]. Collections. Mr. Murup: Mjéberg 101 (UC), 102 (UC), 2200 m, Nooteboom & Chai 2018 (L); Mr. Murup Camp IV: 2000 m, Burtt & Martin B. 5348 (E); Mr. Murup summit: 2400 m, Nooteboom & Chai 2008A (L); Mt. Murup suMMIT RIDGE: 2300-2400 m, Beaman 11458 (K); Mr. Murup, PATH TO suMMIT: 1700 m, Nooteboom & Chai 1915 (L), 1800 m, 7949 (L). 138 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 INDEX TO NUMBERED COLLECTIONS CITED [Collection-number prefixes have been omitted. ] Anderson & Ilias 26471 (21.2.2); 26472 (70.3.1); 26477 (68.1.1); 26480 (70.2.1). — ae (21.2.2): 11441 (542.18), 11442 (70.2.1); 11443 (73.1.2); 11444 (40.1.2); 11445 (54.3.5): 46 (70.3.1); 11447 (54.2.5); 11448 (28.9.2a); 11449 (52.1.2): 11450 (54.2.17): 11451 an D Ps (77.1.1a); 11453 (81.3.3); pes (37.1.1); 11455 (28.10.2); 11456 (69.2.1); 11457 (54 11458 (85.1.1); 11459 (28.9.9); 11460 (66.6.1); 11461 (71.1.2); 11462 (70.3.2); 11463 a ae (19.1.1); 11465 (25.2.1); 11466 (20.1.1); 11467 (54.2.17); 11468 (40.1.1); 11469 (54.1.8); 11470 (41.1.1); 11471 (23.1.1); 11472 (69.2.3); 11473 (63.1.1a); 11474 (54.2.4); 11475 (66.4.1); 11476 (71.1.1); 11477 (81.1.2); 11478 (54.2.4); 11479 (41.2.1); 11480 (21.3.1); 11481 (78.9.1); 11482 (28.8.4); 11483 (40.1.2); 11484 (52.1.1); 11485 (28.12.3); 11486 (54.2.10); 11487 (21.2.3); 11488 (78.1.1); 11489 (1.1.1); 11490 (73.1.1); 11491 (57.1.1); 11492 (74.2.1); 11493 (60.3.4): 11494 (66.6.4) Burtt & Martin 5220 (1.1.2); 5221 (34.1.1); 5222 (69.1.4); 5223 (66.6.2); 5224 (83.1.2); 5225 (73.1.3): 5226 (60.1.1); 5227 (60.1.2); 5229 (28.18.4):; 5230 fu 18.4); 5231 (28.12.1); 5233 (28.1.2); 5234 (72.2.1); 5236 (28.6.3); 5237 (28.13.2); 5238 (42.1.1); 5240 (54.3.13); 5241 (28.9.1); 5243 (28.6.5); 5244 (28.9.8): 5245 (28.6.7); 5246 (60.4.1); 5247 a 41.2.1); 5249 (60.2.1); 5250 (54.2.19): 5251 (54.2.7); S251A (54.2.1a); 5252 (28.3.1); 5253 (54.1.11); 5254 (67.1.2); 5255 (54.2.19); 5256 (28.6.2); 5257 (83.1.4); 5258 (28.6.4); 5259 (28.14.1); 5260 (28.14.1); 5261 (69.1.1); 5262 (28.3.3): 5263 (28.12.3); 5264 (54.2.5); 5266 (54.1.10); 5266A (54.1.4); 5267 (54.1.8); 5268 (54.1.8); 5269 (28.17.1); 5270 (60.3.2); 5271 (28.4.1); 5272 (28.5.1); 5274 (28.6.6); 5275 (60.4.1); 5276 (60.5.1): 5277 (27.1.1); $278 (66.3.1); 5279 (34.4.5); 5280 (60.3.7); 5282 (83.1.3); 5283 (34.2.2); 5284 (66.6.3); 5285 (78.3.1); 5287 (28.1.1); 5288 (54.1.1); 5291 (78.5.1); 5294 (36.1.1); 5296 (78.8.1); 5297 (66.5.1); 5298 (60.1.3); 5299 (28.9.4); 5300 (34.1.1); 5303 (34.2.1); 5306 (54.2.13); 5307 (60.3.3); 5308 (61.1.1); $309 (60.3.5); 5310 (74.2.1); 5311 (83.1.1); 5312 (60.1.1); 313 (60.5.1); 5314 (34.3.2): 5316 (73.1.1); 5317 (54.2.12); 5319 (28.9.1, 28.9.6); 5321 (54.2.4); 5322 (54.2.15a): 5323 (54.3.3); 5323A (54.3.9); 5324 (75.1.1); 5325 (71.1.1); 5326 (78.2.3); 5326A (54.3.2); 5327 (28.12.1); 5328 (28.9.3); 5329 (28.16.1); 5331 (74.1.1); 5333 (28.8.2); 5334 (69.3.1); 5335 (78.1.1): 5336 (78.1.1); 5337 (34.2.1); 5338 (28.17.2); 5340 (78.6.1); 5341 (34.4.3); 5342 (34.4.1); 5343 (34.4.2); 5345 (66.6.2); 5346 (69.2.1); 5347 (83.1.3); 5348 (85.1.1); 5349 (54.3.13); 5351 (34.2.3): S351A (54.2.6); $352 (28.4.2); 5353 (69.1.1); 5354 (28.8.1); 5356 (11.1.2); 5357 (69.4.1): 5358 (54.1.5): $360 (70.3.1); 5361 (76.1.1); 5362 (54.3.13); 5363 (54.2.19); 5364 (65.2.1): 5365 or 16): 5366 (34.4.4); 5367 (83.2.1); 5368 (28.4.2); 5371 (62.1.1); 5372 (34.2.4); 5373 (78.4.1): 5 (82.1.1); $375 (60.3.7); 5376 (38.1.1); 5377 (60.3.6); 5379 (5.1.1); 5382 (34.5.1); 5383 ae 5385 (1.2.1); 5386 (83.1.4); 5387 (83.1.5); 5389 (80.1.4b); 5390 (28.17.1); 5392 (11.2.1); 5393 (28.15.1): ote (54.2.8); 5399 (28.9.7); 5401 (70.5.1); 5402 (34.6.1); 5404 (28.3.2): 5409 (78.7.1); 5410 (54.1.1); 5411 (54.3.10); 5412 (69.1.3); 5413 (84.1.1); 5415 (2.1.1); 5419 (54.1.6); 5421 (54.1.3): 5422 (43.1.1a); $423 (50.1.1a); 5424 (54.3.9): 5425 (70.2.1); 5426 (73.1.3); 5427 (71.1.2); 5428 (28.7.1); $429 (34.7.1); $430 (54.1.9); 5431 (29.2.1); $434 (28.1.4); 5436 (81.3.3); 5437 (20.1.1); 5438 (81.3.1); 5442 (80.1.5a); 5443 (28.10.1); 5444 (10.3.1); 5445 (69.2.2); 5446 (54.2.3): 5447 (44.1.1); 5448 (78.5.2); 5449 (66.6.2); 5450 (54.2.14); 5451 (73.1.2); 5452 (54.1.10); 5453 (32.1.1): 5454 (28.6.1); 5455 (26.1.1); 5457 (59.1.1); 5459 (60.2.1); 5460 (28.7.1); 5461 (76.1.1); 5462 (16.1.1a): 5463 (31.1.1); 5463A oe 1); 5464 (28.18.2); 5465A (9.1.2a): 5465B (9.1.1); 5466 (28.12.3); 5467 (71.1.2); 5468 (40.1.2); 5469 (81.2.1a); 5471 (71.1.2); 5472 (7.1.3); 5473 (69.2.3): 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 (67.1.1): ee ee 5491 (54.3.4); 5492 (67.1.1b); 5493 (70.1.1); 5494 (24.1.1); $495 (31.2.1); 5496 (2.1.1): 5 (54.2.2). Ilias 26303 (36.1.1); 26305 (80.1.6); 26306 (80.1.1a); 26308 (70.3.3); 26309 (81.3.4); 26310 (54.3.2): 54.3.1); 26312 (51.1.1a); 26315 (50.1.1a); 26317 (48.1.1); 26320 (48.1.1); 26322 (48.1.1); 26324 (72.2.1); 26325 (58.1.1); 26326 (54.3.4); 26330 (58.2.2); 26332 (32.1.1); 26333 (48.1.1); 26334 (80.1.4b); 26336 (58.1.1); 26337 (29.1.2); 26342 (48.1.1); 26352 (80.1.1a); 26353 (80.1.la, ae 1); 26357 (80.1.2); 26363 (54.2.19): 26364 (54.3.1); 26365 (50.1.1a); 26373 (54.1.2, 54.1.11); 263 (78.7.1); 26375 (55.1.2); 26376 (67.1.2a); 26378 (54.3.7); 26379 (69.2.1); 26380 (70.3.2): eat (54.1.10); 26382 (68.1.1); 26384 (54.1.3); 26385 (70.3.1); 26386 (41.2.1): 26387 (54.1.8): 26388 poo — 1997 BEAMAN & ANDERSON: SUMMIT FLORA OF MT. MURUD 139 (55.1.3); a0%88 (70.3.2); 26391 (80.1.4b); 26392 (67.1.1c, 80.1.4b); 26393 (54.3.4); 26394 (48.1.1); 26397 (64.2.1); 26398 (54.3.8); 26399 (66.2.1); 26400 (67.1.1b); 26402 (81.3.2); 26403 (54.2.1a): 26406 (70.3.1): 26407 (66.4.1): 26408 (28.10.2); 26410 (54.3.5); 26414 (54.3.7): 26416 (54.1.10): 26417 (67.1.1c); 26419 (76.1.1); 26420 (54.3.2); 26421 (61.1.1); 26433 (54.3.3); 26434 (80.1.1a); 26447 (54.1.6); 26450 (67.1.1c); 26452 (52.1.1); 26453 (40.1.1); 26454 (81.3.2); 26455 (28.9.6): 26457 (54.3.7); 26458 (70.3.1); 26459 (82.1.2); 26462 (54.2.16); 26464 (54.2.19); 26465 (78.5.2): 26466 (28.14.1); 26467 (41.2.1); 26482 (54.3.7); 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): 6 (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 one 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 (67.1.1¢); 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 es 1.5); 44402 (28.9.4); 44403 (51.1.1a); 44405 (36.1.1); 44408 (47.2.1): 44409 kes 44410 (78.2.1); 44411 (66.5.1); 44412 (78.8.1); 44414 (60.3.1); 44416 (60.3.7); 44420 (82.1. ae 44422 (61.1.1); 44425 (40.1.2); 44426 (55.1.1); 44427 (52.1.1); 44428 (70.3. i sie (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 140 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 (66.4.1); 44484 (81.3.2); 44485 (81.3.2); 44486 (28.10.2); 44487 (41.2.1); 44488 (37.1.1); 44489 (40.1.1); 44490 (54.1.8); 44491 (28.12.3); 44493 (54.1.6); 44499 (81.1.1); 44601 (67.1.1b); 44602 (54.2.10); 44604 (67.1.2a); 44605 (81.3.2); 44608 (21.2.3); 44609 (69.2.1); 44610 (69.2.1); 44611 (81.1.1); 44612 (37.1.1); 44615 (55.1.4); 44616 (28.9.8); 44617 (72.1.1); 44618 (40.1.2); 44619 (41.2.3); 44620 (65.1.1); 44621 (46.1.1); 44623 (71.1.2); 44627 (28.10.1); 44633 (66.1.1); 44646 (67.1.1a). 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 ae and a number of persons from the Herbarium ee Botanic Gardens, Kew, including S. M. Andrews, M. ; , E. J. Cowley, J. Dran sfield, . Edwards, L. L. Forman, M. J. Sands, and J. J. Wood. Others who helped with identifications en G. Argent (E), N. “Callineee (FLAS), G. Clausing (MJG), J. J. Pipoly (BRIT), R. Kiew (UPM), M. G. Price of Jackson, Michigan, and L. G. (KEP). We have also been helped by B. A. Barlow (CANB), C. C. nie (BG), S. Kofman (L). Ce. 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 — State University. We greatly appreciate the apenas resources made available to us by the Sarawak st Department Herbarium, Kuchin (SAR), the Royal Botanic Gardens, Kew (K), the Sata Leiden (L), and the Royal Botanic Garden, te (E). LITERATURE CITED Anderson, J. A. R., and P. P. K. Chai. 1982. A preliminary list of angiosperms and gymnosperms recorded within Gunung Mulu National Park. In shih Mulu National Park: A Nese el and Development Plan, by J. A. R. Anderson, A. C. Jermy, and The Earl of Cranbrook, App dix HI, 233-271. Royal Geographical Society, London Anonymous. 1952. Preface. Sarawak Papers: Scientific Rasiilis of the Oxford University Expedition to Sarawak (Borneo) in 1932, i-viil. 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 oe ] Beaman, J. H. 1997. Preliminary enumeration of the summit flora, Mount Murud, Kelabit Highlands, Garewal : A scientific ae 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. a. 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 speci- pees database for the flora of Mount Kinabalu. Taxon 38: 27-42. Bonaparte, R. B. 1923a. Récoltes du Dr. E. MjOberg, Herbier du Prince Bonaparte. Notes Ptérido- logiques 14: 477-488 .1923b. Plusieurs régions, plusieurs collecteurs, Herbier du Prince Bonaparte. Notes Ptérido- 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- 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. . C. Jermy, v-xv. Kuching, Sarawak: Sarawak Forest Departm ee P. i 1977. The Altitudinal Zonation a Forests Along ie ‘West Ridge of Gunong Mulu. Kuching, Sarawak: Sarawak Forest Department Merrill, E. D. 1928. A collection of plants from Sc aoak Sarawak Museum J. 3: 513-557 Mjéberg, 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 rae 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 ae Kinabalu. 1. Ferns and Fern Allies. Kew: Royal Botanic Gardens Price, M. G. 1983. Several unusual Malesia Diplazia. Gard. Bull. Singapore 36: 25-2 Yin Ee pene 1992. Geological Map of Sarawak, 2d ed., updated by V. Hon and ae S. K. Kuala mpur: Director-General Geological Survey of Natacoe Wood, 4 J. 1984. A preliminary annotated checklist of the orchids of Gunung Mulu National Park with a key to genera. Studies on the Flora of Gunung ae National Park, Sarawak, ed. A. C. Jermy, 1-39. Kuching, Sarawak: Sarawak Forest Departm Wood, J. J., R. S. Beaman, and J. H. Beaman. 1993. The re eee Kinabalu. 2. Orchids. Kew: Royal Botanic Gardens. ADDENDUM While this paper was in page proof we learned from J. J. Wood about two additional species of orchids collected on Mt. Murud by Burtt and Martin in 1967. These are added here, but data concern- ing number of taxa, taxon numbers, etc., have not been changed in the text ORCHIDACEAE KUHLHASSELTIA Kuhlhasseltia javanica J. J. Sm. Collection. Mr. MuruD NEAR Camp IV: 1900 m, Burtt & Martin B. 5408 (E). ROBIQUETIA Robiquetia pinosukensis J. J. Wood & A. L. Lamb Collection. Mr. Murup Camp II: 1700 m, Burtt & Martin B. 5232 (E). o aes SS -_ Se ¥ a ee — i “7 a 7 - 7 4 - 7 a a —_ oe oe ~— oe = - -_ 7 - - a - 7 - - 7 a 7 a _ 7 a > Oo BC _ . areata = Sse = = _ a 7 Emus = oe -_— 7 ne - 7 ~~ - Oo 7 7 Oo 7 an 13 So ee - re = on — ca ee 7 a 7 - a > : 7 a en : 7 : . Contr. Univ. Michigan Herb. 21: 143-146. 1997. ARISTOLOCHIA DAVILAE (ARISTOLOCHIACEAE), A NEW SPECIES FROM NAYARIT, MEXICO Ismael Calzada J. Gabriel Flores F. Oswaldo Téllez V. Herbario Nacional Departamento de Botanica Instituto de Biologia UNAM Apartado Postal 70-367 04510 México, D.F., Mexico This contribution complements previously published papers dealing with the Flora of Nayarit Project (Mendez & Téllez 1995; Téllez 1995), carried out for several years by the Instituto de Biologia UNAM. Aristolochia davilae Calzada, Flores & O. Téllez, sp. nov.—T ype: Mexico. Naya- rit: Mpio. Nayar, cafiada en el ejido Carretones de Cerrito, ca. 4 km al E de la cortina del Proyecto Hidroeléctrico Aguamilpa, 21°50'30"N, 104°45'SO" W, bosque tropical caducifolio, suelo negro, rocoso, 180 m, 20 Aug 1993, Calzada, Flores & Solis 18596 (holotype: MEXU!; isotype, MICH!). Big. 7. 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, 143 144 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 FIG. 1. Aristolochia davilae. a. Habit. b. Root. c. Section of flower showing the staminal column. d. Capsule. e. Immature seed. (Based on Calzada et al. 18596, MEXU.) 1997 CALZADA, FLORES & TELLEZ: ARISTOLOCHIA DAVILAE TABLE |. Characters of D. davilae and its relatives. A. davilae A. oaxacana A. bracteosa A. socorroensis Habit erect to climbing prostrate prostrate prostrate Lamina shape linear-lanceolate ovate to ovate, or triangu- _hastate-trilobate triangular lar to lanceolate length (2.6-) 3.7-7.7 cm 4.0 cm 2.5-10.0 cm 5.0 cm width (0.3-) 0.5-2.6 cm 2.0 cm 2.0-5.0 cm 4.0 cm base auriculate cordate deeply cordate deeply cordate indumentum glabrous to spreading- — subvelutinous to puberulent to strigulose to hispid, tuberculate hispidulous subvelutinous densely hirsutulous Petiole length 3.0-5.0 mm 0.3 mm 3.0-5.0 mm 10.0 mm Calyx length 4.0 mm 3.0-4.5 mm 4.0-6.0 mm 6.0 mm Stamens shape columnar columnar coroniform coroniform length 1.5 mm 3.5 mm 3.5 mm 2.5 mm Capsule shape subspherical to oblate subspherical to subspherical to spherical spherical spherical length 1.0-1.5 cm 1.0 cm 2.0-3.0 cm 2.75 cm diameter 1.2 cm 1.2 cm 1.0-1.5 cm 1.75 cm length 4.0 mm 3.0 mm 5.0 mm 5.0 mm width 3.0-4.0 mm 3.0 mm 4.0 mm 5.0 mm Distribution Nayarit Oaxaca Nayarit, Jalisco, Colima Revillagigedo Islands 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 Comisién 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 We thank Dr. José Luis Villasefior Rios for reviewing the seas gk and for providing the Latin diagnosis. We are grateful for the financial support provided by CONACYT (grants PCCNC- NA-031529, P218CCOC880112, P218CCOC8277 792, 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. Téllez V. 1995. Una nueva especie de Russelia a a aa del estado de Nayarit, México. Anales Inst. Biol. Univ. Nac. México, Ser. Bot. 66(2): 18 Pfeiffer, H. W. 1970. A taxonomic revision of the pentandrous species of yuan Univ. Con- necticut Pub. Ser. 134 p Téllez V., O. 1995. New species of Leguminosae from Nayarit, Mexico. Contr. Univ. Michigan Herb. 20: 231-237 Contr. Univ. Michigan Herb. 21: 147-159. 1997. MISCELLANEOUS NOTES ON SPHAGNUM—10 Howard Crum University of Michigan Herbarium North University Building Ann Arbor, Michigan 48109-1057 NEW SPECIES Sphagnum [sect. Sphagnum] amoenoides Crum, sp. nov.—T ype: BRaAzIL. Sao Paulo: erra 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!). Rigs 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 laterali 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 quing ia, 1.5- 1.8 mm longa, oblongo-lanceolata, apice cucullata; cellulae hyalinae poris et fibril- lis ut in foliis caulinis; cellulae chlorophylliferae in sectione transversali orciformes utroque latere folil 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: Braziv. 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, Schdfer-Verwimp & Verwimp 15214 (holo- type: MICH!; isotype: Schafer-Verwimp!). Fig. 2. 147 148 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 FIG. |. Sphagnum amoenoides. a. Branch leaves, x24. b. Portion of branch leaf in section, x390. c. Upper cells of branch leaf, outer surface, x390. d. Upper cells of branch leaf, inner surface, x390. e. Stem leaves, x24. f. Upper cells of stem leaf, outer surface, x390. g. Upper cells of stem leaf, inner surface, x390. 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 pendenti); 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, lingulate, 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 | or sometimes 2 large, round pores, 1997 CRUM: SPHAGNUM 149 FIG. 2. Sphagnum atroligneum. a. Branch leaves, x24. b. Portion of branch leaf in section, x390. c. Upper cells of branch leaf, outer surface, x390. d. Upper cells of branch leaf, inner surface, x390. e. Stem leaves, x24. f. Upper cells of stem leaf, outer surface, x390. g. Upper cells of stem leaf, inner surface, x390. often with a few small, ringed, elliptic pseudopores as well. Branches in fascicles of 2 or 3 (2 spreading, 1 pendent); cortex delicately 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 | 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 cylinders are interesting. 150 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 2] Sphagum [sect. mace billbuckii Crum, sp. nov.—T ype: BRAZIL. Minas Gerais: spinhago, Mpio. Santa Barbara, Parque Natural do Caraga, 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!). Piey, 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 foltis 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 hyaline 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 lIcaf 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 included. Sphagnum [sect. Cuspidata|] subditivam Crum, sp. nov.—Typer: 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) 80/1 (holotype: MICH}; isotype: CHR!). Fig. 4. Plantae plus minusve graciles, laete viridi-luteae. Epidermis caulina ex stratis 2 composita; cylindrus lignosus pallide luteus. Folia 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 folii 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 discontinuis ad commissuras 1997 CRUM: SPHAGNUM 151 FIG. 3. Sphagnum billbuckii. a. Branch leaves, x24. b. Portion of branch leaf in section, x390. c. Upper cells of branch leaf, outer surface, x390. d. Stem leaves, x24. e. Upper cells of stem leaf, outer surface, x390. 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 152 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 SONI SAE SEOUL DAKO) nea FIG. 4. Sphagnum subditivum. a. Branch leaves, x24. b. Portion of branch leaf in section, x390. c. Upper cells of branch leaf, outer surface, x390. d. Upper cells of branch leaf, inner surface, x390. e. Portion of stem in section, x390. f. Stem leaves, x24. g. Upper cells of stem leaf, outer surface, x390. h. Upper cells of stem leaf, inner surface, x390. Pe 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 Subsecunda. As commonly seen in the section Cuspi- O97: CRUM: SPHAGNUM its}s) 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.—T ype: BRaziL. Sao Paulo: Serra da Bocaina bei SAo José do Barreiro, an quelligem Hang an der Strasse bei “Shangra-la,” ca. 22°47'S, 44°38'W, 1500 m, 16 Apr 1988, Schafer-Verwimp & Verwimp 9588 (holotype: MICH!). Fig25: Plantae 7-8 cm altae, pallide virides vel aurantiacae. Epidermis caulina strato uno, sine poris; cylindrus lignosus pallide 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. Folia 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 | 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: Brazi_. 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, Schéfer-Verwimp & Verwimp 8591 (holotype: MICH!). hig 6: Plantae tenues, usque ad 10 cm altae, capitulo nullo sed apice aliquantum bulboso-geminato. Cellulae epidermidis strato uno, sine poris; cylindrus 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 154 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 a FIG. 5. Sphagnum bocainense. a. Branch leaves, x24. b. Portion of branch leaf in section, 390. c. Upper cells of branch leaf, outer surface, x390. d. Upper cells of branch leaf, inner surface, x390. e. Stem leaves, x24. commissuras dispersis, interiore poris annulatis in seriebus commissuralibus con- tinuis. Rami fere solitari 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 liberae. 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, elliptic pores in continu- ous commissural rows. Branches mostly single, laxly foliate (but occasionally 2 or rarely 3, with | short, weak pendent branch. Branch leaves I-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 1997 CRUM: SPHAGNUM 155 FIG. 6. Sphagnum laxiramosum. a. Branch leaves, x24. b. Portion of branch leaf in section, x390. c. Upper cells of branch leaf, outer surface, x390. d. Upper cells of branch leaf, inner surface, x390. e. Stem leaves, x24. f. Upper cells of stem leaf, outer surface, x390. g. Upper cells of stem leaf, inner surface, x390. 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. Suwbsecunda] rio-negrense Crum, sp. nov.—T yee: CoLomsia. Cun- dinamarca: Paramo de Palacio, caveceras Rio Negro, 3375 m, 22 May 1972, A. M. Cleef 4001 (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. Folia 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 folii parietibus incrassa- tis liberae. Plants brown or yellow-brown, tumid, ca. 6 cm high. Stems simple, very slen- der, brown; epidermal cells often porose, 1-2-layered; wood cylinder brown or 156 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 FIG 7. Sphagnum rio-negrense. a. Habit of plant, x3. b. Leaves, x24. c. Portion of leaf in section, x390. d. Upper cells of leaf, outer surface, x390. e. Upper cells of leaf, inner surface, x390. 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. ADDITIONAL SPECIMEN EXAMINED. COLOMBIA. Cundinamarca: Pdaramo de Palacio, Lagunas de Butriago, 3350 m, 27 Apr 1973, A. Cleef 9580 (NY). The stems are simple, and the plants have much the same appearance as the North American Sphagnum cyclophyllum 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. Subsecunda| subhomophyllum Crum, sp. nov.—Tyre: BraziL. io 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; cylindrus lignosus brunneus. Folia caulina 1-2 mm longa, oblongo-triangularia, concavo-acuta; cellulae hyalinae dimidio su- periore fol fibrillosae, non septatae, utroque superficie poris numerosis in seriebus commissuralibus discontinuis et superne apice etiam poris vel pseudoporis 1997 CRUM: SPHAGNUM lay) , ee FIG. 8. Sphagnum subhomophyllum. a. Branch leaves, x24. b. Portion of branch leaf in section, x390. c. Upper cells of branch leaf, outer surface, x390. d. Upper cells of branch leaf, inner surface, x390. e. Stem leaves, x24. f. Upper cells of stem leaf, outer surface, x390. g. Upper cells of stem leaf, inner surface, x390. 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. ants 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 homophyllum Crum and S. pluri- porosum Crum have median pores rather than pseudopores. In addition, $. homo- phyllum 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.—T ype: CoLosia. Meta: Péramo 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 altae, tumidae, subsimplices. Caules tenues, obscure fusci; cellulae corticis stratis duobus, saepe uniporosae; cylindrus lignosus obscure fuscus. Folia caulina 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 commissuralibus continuis dispositis; cellulae chlorophylliferae in sectione trans- versali 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 (-I5) 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. : undinamarca: Paramo de Sumapaz, Chisaca, La- guna N of Laguna Grande, 3650 m, 15 Nov 1971, A. M. Cleef & R. Jaramillo-M. 41, (MICH, NY).— Without locality: R. E. Schultes 11475 (MICH, mae 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 II or 15 per cell, but in the other collections they vary from one to seven or may even be absent. The hyaline 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. 1997 CRUM: SPHAGNUM 159 CRO AP AGU IG. 9. Sphagnum sumapazense. a. Leaves, x24. b. Portion of stem in section, x390. c. Portion of leaf in section, x390. c. Upper cells of leaf, outer surface, x390. d. Upper cells of leaf, inner surface, x390. e. Cortical cells of stem, in surface view, x390. f. Portion of stem in section, x390. ADDITIONAL NOTE Sphagnum frahmii Crum, Contr. Univ. Michigan Herb. 20:130. 1995.—T ype: 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: ALTA!). I am indebted to Marshall Crosby for pointing out that this species was not validly published 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. Buchanania) leucobryoides sect. et sp. nov. from Tasmania. J. Bryol. 6: 45-54 Yamaguchi, T., R. D. Seppelt, and Z. Iwatsuki. 1992. Buchanania (Sphagnum, Sphagnaceae) sect. nov. Hikobia 11: 139-140. Contr. Univ. Michigan Herb. 21: 161-174. 1997. 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) list 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 trme. 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 (1.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, 1s 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 (1.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%, 1.e., 13 endemics/128 species; totals based on Gibson, 1974, with cor- rections in Daniel, 1995b, and various geographic updates) and Chiapas, Mexico (11%, 1.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 and southern/moister regions that could be treated along with adjacent portions of environmentally similar surrounding regions of Mexico and Guatemala (cf. Wendt 1993: 596). Collections have been made of at least seven species of Acanthaceae (Androgra- phis gangetica, Hemigraphis alternata, Ruellia coerulea, Thunbergia alata, T. erecta, T. fragrans, T. grandiflora) that have been introduced into Belize from other parts of the world. These exotics include cultivated and/or naturalized species. Additional 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 of habitats created in southern Belize by the Maya Mountains and the presence of 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 is enriched, particularly in Corozal, by “dry forest species” or species that are more common in the northern portion of the Yucatan peninsula (e.g., Bravaisia berlandieriana, Carlowrightia myriantha, and Justicia campechiana). Spellman et al. (1975) remarked on the West Indian floristic influence in Belize. There are no Acanthaceae known from the West Indies that occur on the American continent only in Belize. Although 10 of the 40 (25%) native Acan- 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). or each species included in the following annotated catalog, the district(s) of 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 of 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 (OTT: DANIEL: ACANTHACEAE OF BELIZE 163 COROZAL 10 (3) ORANGE WALK 6 (0) p Le o>? 88° 16°30’ TOLEDO 26 (7) 20 km FIG. 1. Map of Belize showing political districts. Numbers indicate number of species of Acan- thaceae in each district (and number of species of Belizean Acanthaceae known only from that district). 164 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 country, the following species might be expected to occur in Belize: Elytraria bromoides Oerst., Justicia carthagenensis Jacq., J. colorifera V.A.W. Graham, Pseu- deranthemum alatum (Nees) Radlk., P. cuspidatum (Nees) Radlk., Ruellia inundata Kunth, R. paniculata L., Stenandrium dulce (Cav.) Nees, S. pedunculatum (Donn. Sm.) Leonard, and S. subcordatum Standl. CATALOG OF THE ACANTHACEAE OF BELIZE Aphelandra aurantiaca Lindl. (including A. repanda Nees). Cayo: Schipp 625 (F).—To.epo: Daniel & Butterwick 5890 (CAS); Gentle 4288 (MICH), 7/76 (MO, US): Schipp 1063 (F, MICH, MO, NY, UC). 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, Hort.” 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 likely that the origin of Lindley’s synonym (..e., the horticultural nomen novum) was somehow derived from Scheidweiler’s name, Lindley was clearly unaware of Scheidweiler’s publication 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. in Dwyer & Spellman, 1981). LizeE: Croat 23905 (F, NY, US); Gentle 1185 (DS, F, MICH, NY, US), /304 (F, MICH, NY): MeDaniel & Clark 14348 (F): po 23942 (RSA).—Cayo: Arrigo 5 (F); Bartlett 11884 (MICH), 11942 (MICH, US), 1/947 (MICH, US); Chanek 38 (MICH, US); Gentle 2392 (MICH, NY), 8889 (CAS, F, US); Lundell 439 (CAS, _ US): Meave 1025 (MO); Meave & Howe 1164 (MO).—CoroZAL: Crane 506 (CAS); Gentle 202 (MICH, US); Kinloch 3 (F).—ORANGE WALK: Lundell 504 (DS, F, NY, US); Karling 12 (F), 51 (F).—Stann Creek: Daniel & Butterwick 5877 (CAS); Molina R. 18 (F); Schipp 37 (F, MICH, NY, UC, US).—To.epo: Gentle 3774 (MICH, NY, US), 3775 (F, MICH, NY, US); Whitefoord 1488 (MO), 1493 (MO), 1/877 (MO).—Districr Unknown: Bartlett 11349 (MICH); Stocker 13 (F). a Asystasia gangetica (L.) Anders. STANN CREEK: Gentle 7866 (F, US). This Old World species has not been heretofore reported from Belize. It is 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); Bartlett 11478 (CAS, MICH, US). The taxonomy and distribution of this species were discussed by Daniel (1995a). Blechum pyramidatum (Lam.) Urb. (as B. brownei Juss. in Dwyer & Spellman 1981). Beuize: 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).—Corozat: 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).—Totepo: 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). BeEuizE: Gentle 1193 (A, BM, F, G, GH, K, LL, MICH, MO, NY, US); McDaniel 13080 (MO).— Corozat: 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 I (F).—ORANGE WALK: Arnason & Lambert 17152 (MO); Daniel 8263 (CAS); Winzerling ViII-14 (US).—Districr UNKNown: Campbell I (K); Stevenson s.n. (US) A monograph of this genus was provided by Daniel (1988). Bravaisia grandiflora Donn. Sm. ToLepo: Gentle 4555 (LL, MO); Lamb 58 (F); Peck 730 (GH; type of B. proxima S. F. Blake): Schipp 1259 (A, BM, F, G, GH, MICH, MO, NY); Whitefoord 3219 (BM, MEXU, MO, NO). Carlowrightia myriantha (Standl.) Stand. Corozac: Crane 516 (LL, MO): Daniel 8267 (BR, C, CAS, ENCB, F, K, MEXU, MICH, MO, NY, US), 8268 (CAS). 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 lip 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. Corozat: Ramamoorthy et al. 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 1n height is probably erroneous; trees are unknown in the genus. This is the first report of the species in Belize. Dicliptera sexangularis (L.) Juss. (as D. assurgens (L.) Juss. in Dwyer & Spellman 1981) Beiize: Daniel & Butterwick 5901 (CAS); Gentle 1119 (F, MO, NY); fs (GH, eee es — & Butterwick 5899 (CAS), 5900 (CAS); Gentle 9062 (CAS, F, MO, US), 9698 (CAS, F, MO, Y, US); Lundell 4101 (MICH, US), 4/02 (MICH, US), 4703 (MICH, US); Pa a a piney Daniel 8271 (CAS); Pelly 24 (F).—OrRANGE WALK: pele & Lambert 1727 (MO); Daniel 7040 (CAS), 8215 (BR, C, CAS, K, MEXU, MICH, MO, . US).—STANN CREEK: Gentle 7582 (CAS, F, MO, US). Dicliptera sumichrastii Lindau. Totepo: Schipp S-684 (F, MO, NY). 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 likewise 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. suwmichrastii. Elytraria imbricata (Vahl) Pers. Cayo: Bartlett 11497 (MICH). Hemigraphis alternata (Burm. f.) Anders. Be.ize: Bartlett 11355 (MICH; type of Blechum cordatum 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. S08 (F); Gentle 8627 (CAS, F, MO, US); McDaniel 12999 (MO).—ToLepo: 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).—To.tepo: Daniel & Butterwick 5889 (CAS); Schipp S-601 (F, MO, NY). Justicia bartlettii (Leonard) D. N. Gibson (including J. pilifera D. N. Gibson). Beuize: Liesner & Dwyer 1486 (MO).—Cayo: Arvigo & Shropshire 203 (MO, US); Bartlett 11477 (US; type of Beloperone bartlettii Leonard).—To.tepo: Davidse & Brant 32157 (CAS). Justicia breviflora (Nees) Rusby (including Pseuderanthemum tetrasepalum (S. F. Blake) S. F. Blake). Bewize: 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).—To epo: Cosentino 86 (F); Croat 24317 (F, NY, RSA, US); Daniel & Butterwick 5895 (CAS); Gentle 4507 (MICH), 5274 (MO, US); Holst 4025 (CAS),4245 (CAS); McDaniel 12884 (F, MO); Peck 552 (GH; type of Evran- themum tetrasepalum S. F. Blake); Peck 722 (GH; type of 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 J. breviflora and correspond to the remainder of Blake’s description. Four-parted calyces are not known in Pseuderanthemum but are often present in J. 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 VOLUME 21 Justicia campechiana Stand. Coroza: Castillo 44 (F); Pelly 3 (F). Justicia candelariae (Oerst.) Leonard. Cayo: Gentle 9011 (F, MO, US).—StTann Creek: Gentle 9316 (CAS, F, MO, US). This species was not listed for Belize by Dwyer and Spellman (1981) but was noted to occur in the country by Daniel (1995b). Justicia comata (L.) Lam. Beuize: Gentle 913 (F, NY); pe 1957 (MICH, US); Whitefoord 2354 (MO).—Cayo: Lundell 4151 (F, MICH); Contreras 7145 (F, MO, US).—Stann Creek: Dwyer et al. 504 (MO); Schipp 622 (F, MICH, MO, NY, UC).—Districr eRe Usher 7 (MICH). Justicia ensiflora (Standl.) D. N. Gibson. STANN CREEK: Gentle 2115 (DS, F, MICH, NY, US); Gentry 7931 (MO); Schipp 354 (F; type of Jacobinia ensiflora Standl.). Justicia fimbriata (Nees) V. A. W. Graham (as J. magniflora (S. F. Blake) D.N. Gibson in Dwyer & Spellman 1981). Totebo: Balick et al. 2538 (US); Cosentino 90 (F); Daniel & Butterwick 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 Standl.); Stevenson 82 (F, NY, US). Justicia pectoralis Jacq. Bexize: Gentle 1530 (MICH, US).—Stann Creek: Daniel & Butterwick 5880 (CAS); Gentle 1893 °H, NY, US).—To.epo: Gentle 4419 (MICH), 5278 (MO, US), 7591 (MO); Peck 979 (GH); Proctor 36011 (MO) Justicia spicigera Schltdl. Be.ize: Peck 430 (GH; type of Jacobinia scarlatina S. F. Blake).—Cayo: Chanek 40 (MICH, US).— ORANGE WALK: Winzerling VIII-10 (F, US).—STANN CREEK: Gentle 3305 (A, MICH, MO, NY, US); wey 523 (A, F, GH, MICH, MO, NY, UC).—To evo: Gentle 7334 (CAS, F, MO, US): Téllez et al. 98 (F, GH). 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). eee alopecuroidea (Vahl) R. Br. ex Griseb. (as Teliostachya alopecuroidea (Vahl) Nees in Dwyer & Spellman 1981). Cayo: Bartlett 11754 (MICH, NY, US), 13024 (CAS, MICH, US).—StTann Creek: Daniel & Butter- wick 5879 (CAS); Schipp S-135 (F).—To.epo: Schipp 1342 (F, MICH, MO, NY) The generic placement of this species was discussed by Daniel (1995a). 1997 DANIEL: ACANTHACEAE OF BELIZE 169 Louteridium chartaceum Leonard. BeE.ize: 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 aniel & 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 donnell-smithii S. Watson. Cayo: Dwyer & Liesner 12313 (MO, NY); Dwyer et al. 360 (MEXU, MO); Gentry 7791 (MO).— To.epo: Daniel & Butterwick 5891 (CAS): Davidse & Brant 32089 (CAS, US), 32300 (CAS); Gentle 6355 (MO, UC, US); Holst 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 96/ (F; type of M. belizensis Standl.). Mendoncia retusa Turrill. STANN CREEK: Gentle 3472 (MICH, MO, NY).—To.epo: Gentle 4218 (MICH), 63177 (CAS, F, MO, US); Schipp 1051 (F, MICH, MO, NY, UC).—Districr Unknown: Gentle 3981 (F, MICH, MO, NY). Odontonema albiflorum Leonard. 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).—To epo: Balick et al. 2547 (US); Croat 24383 (F, ie ae ee (MO); Peck 786 (GH, NY), 787 (GH, NY); Proctor 35876 (MO); Whitefoord 1615 (M Odontonema callistachyum (Schltdl. & Cham.) Kuntze. Cayo: Dwyer & 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), 3507 (A, F, MICH, NY), 8202 (F, MO, US), 9250 (CAS, F, MO, US).—To.epo: 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. Bewize: Daniel 8295 (CAS, MICH, MO, US); Liesner & Dwyer 1462 (MO); Peck 278 (GH; type of O. paniculiferum S. F. Blake)—Cayo: Bartlett 11442 (MICH, US), 12947 (MICH, US); Gentle 2336 170 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 (MICH, NY), 8835 (CAS, F, MO, US), 9004 (CAS, F, MO).—STANN CREEK: Dwyer et al. 571 (MO, US); Gentle 2136 (A, DS, F, K, LL, MICH, NY, US), 2/58 (MICH), 2742 (MICH), 2793 (MICH, NY, ae 3195 (MICH), 3496 (F, MICH, NY): Molina R. 276 (F); Schipp 222 (A, BM, F, G, GH, MICH, NY, C, US).—Totepo: Daniel & Butterwick 5892 (CAS, MICH); Davidse & Brant 31921 (CAS): Gentle 4506 oe MICH, MO), 6704 (F, MO, US): Holst 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 Bewize: Daniel 8276 (CAS, MICH); Daniel & Butterwick 5904 (CAS): Gentle 1538 (GH, LL, MICH, MO, — eb Dwyer 1466 (MO): Peck 394 (GH, NY).—Cavyo: Bartlett 11942a (MICH), 11952 (CA H, US); Chanek 47 (MICH, US); Daniel & Butterwick 5897 (CAS); nhac et al. 178 ‘Mo. aaa 2371 (F, NY), 2439 (MICH); sled 1430 (MO).—STANN CREEK: Schipp 146 (F, NY, UC, US).—To.epo: Croat 24228 (F, GH, NY): Davidse & Brant 32010 (CAS); Gentle 3079 (MICH, NY); Gentry 8/68 (MO); Kinloch 33 (F); Schipp ee (K; type of O. amicorum). The taxonomic disposition of O. amicorum was addressed by Daniel (1995c). Pseuderanthemum verapazense Donn. Sm. Totepo: Peck $30 (GH; type of Eranthemum adenocarpum S. F. Blake); Schipp S-462 (F), S-669 (F), S-671 (F, K, NY Ruellia coerulea Morong. Beiize: Daniel 8278 (CAS).—Corozar: Gentle 347 (MICH). 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 5278 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 Kunth. Cayo: Bartlett 11814 (MICH); Dwyer et al. 323 (F, MO); Lundell 6602 (MICH, US); Wiley 368 (CAS).—STANN CREEK: Schipp 761 (MICH, MO, NY, UC). 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). Beuize: Dwyer 11317 (MO).—Cavyo: Bartlett 13063 (CAS, MICH, NY, a Gentle 8598 (F, MO, US): Lundell 322 (DS, F, UC), 6578 (DS, MICH, NY, US), 6667 (MICH, NY, US); Spellman 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 Standl.).—To.epo: Gentle 7056 (LL); Peck 871 (GH; type of R. obtusata). 2 Ruellia hookeriana (Nees) Hemsl. Cayo: Casentino 33 (F). 1997 DANIEL: ACANTHACEAE OF BELIZE 171 This species was not listed for Belize by Dwyer and Spellman (1981) but was noted to occur in the country by Daniel (1995b). Ruellia matagalpae Lindau. Cayo: Balick et al. 3346 (US); Gentle 2300 (MICH, NY, US); Liesner & Dwyer 1552 (MO); Spellman 1371 (MO).—To Lebo: Davidse & Brant 32422 (CAS); Gentle 3984 (F, MICH, MO, N Ruellia nudiflora (Engelm. & Gray) Urb. Cayo: Arvigo 56 (US); Arvigo 1987-17 (F); Balick et al. 1797 (US); Daniel 8272 (CAS).—CorROZAL: Crane 89 (CAS): Croat 24586 (MO); Dwyer 14504 (MO); Gentle 178 (MICH, US).—ORANGE WALK: Arnason & Lambert 17150 (MO); Daniel 7033 (CAS); Lundell 36 (F, MICH, US), 365 (F, MICH), 634 (DS, F, MO, NY, US).—District Unknown: Lundell 4979 (MICH, MO, US). Ruellia pereducta Stand. Cayo: Bartlett 12900 (CAS, MICH, US); Chanek 39 (MICH, US); Dwyer et al. 179 (MO), 376 (MO); Gentle 2170 (F, MICH, NY); Lundell 6/28 (MICH, NY, US); Molina R. 129 (F); Whitefoord 2888 (MO).—To evo: Schipp S-645 (F, NY). Ruellia pygmaea Donn. Sm. To.Lepo: Schipp S-668 (F, NY), 1357 (F, MICH, MO, NY). Thunbergia alata Bojer ex Sims. Cayo: Chanek 200 (F, MICH). This African species is widely cultivated and naturalized in the American tropics. Thunbergia erecta (Benth.) T. Anderson. Beuize: Daniel 8277 (CAS).—Cayo: Balick 2278 (US). This African species, which was not listed by Dwyer and Spellman (1981), is cultivated in Belize and elsewhere in the New Wor Thunbergia fragrans Roxb. ToLepo: Gentle 6371 (CAS, F, MO, NY, US). This Asian species was not reported by Dwyer and Spellman (1981). It is often cultivated and has become naturalized in various parts of the neotropics. Thunbergia grandiflora Roxb. Beuize: Whitefoord 2453 (MO).—Corozat: Gentle 50 (F); Lundell 4786 (MICH, NY).—STANN CREEK: Daniel 8289 (CAS). This Asian species is widely cultivated and sometimes persists or becomes locally naturalized in tropical regions. sie 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 of A. 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. sexangularis (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 magniflora (S. F. Blake) D. N. Gibson.—Synonym of J. fimbriata (Daniel Justicia sp.—Dwyer and Spellman (1981) listed Croat 23899 as an undetermined species of Justicia. The whereabouts of this collection are not known (T. Croat, pers. comm.). Odontonema flagellum (Oerst.) Kuntze.—Synonym of O. tubaeforme (Daniel QO. 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 listed 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 above). 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. Ruellia obtusata S$. F. Blake.—Synonym of R. harveyana (Daniel 1995b). 1997 DANIEL: ACANTHACEAE OF BELIZE Ai) 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 Jacqg.”—-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.—AlIl 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). Teliostachya alopecuroidea (Vahl) Nees.—Synonym of Lepidagathis alopecuroti- dea (Daniel 1995b). 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), a 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 Odontonema (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: 111-132. 174 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 1991. A revision of Aphelandra (Acanthaceae) in Mexico. Proc. Calif. Acad. Sci. 47: 235-274. .1993. Taxonomic and geographic notes on Central American Acanthaceae. Proc. Calif. Acad. — 48: 119-130 995a. New and reconsidered Mexican Acanthaceae. VI. Chiapas. Proc. Calif. Acad. Sci. 48: 284 1995b. Acanthaceae. In Flora of Chiapas, by D. E. Breedlove, pt. 4: 1- eee Revision of Odontonema (Acanthaceae) in Mexico. Contr. a Vion Herb. 20: 147- Durkee, . = 1978. Acanthaceae. In Flora of Panama, ed. R. Woodson et al., Ann. Missouri Bot. Gard. 65: 155-284. . 1986. Acanthaceae. In ghee ion ed. W. Burger, Fieldiana, Bot. 18: 1-92. Dwyer, J. D., and D. L. Spellman. 1981. st of the Dicoivledonsae of Belize. Rhodora 83: 161-236. Famighetti, R., . 1995. The world cen a 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. “ 258: 1-77. Gibson, D. N. 1974. Acanthaceae. In Flora of Guatemala, ed. P. C. Standley et al. Fieldiana, Bot. 24 (10): 328-461. euter, W. et al. es International Code of Botanical Nomenclature. Regnum Veg. 131: 1-389. Hampshire, R. J. 1989. Belize. In Floristic sea of Tropical Countries, ed. D. G. Campbell and H.D Hammond 286- 289. Bronx: New York Botanical Garden Leonard, E. Ma . The Acanthaceae of te Yucatan Peninsula. Publ. Carnegie Inst. Wash. 461: 191-23 Lindley, J. ie. oo aurantiaca. Bot. Reg. 31: no. 12. Lunde I, C. Lh. . The vegetation and natural resources of British Honduras. In Plants and plant nce in ae America, ed. F. Verdoorn, 270-273. Scheidweiler. M. 1842. Notice sur quelques nouveaux generes et espéces de plantes. Bull. Acad. Roy. Sci. Bruxelles 9: 19-26. Spellman, D. L., J. D. Dwyer, and G. Davidse. 1975. A list of the Monocotyledoneae of Belize including a historical introduction to plant collecting in ate Rhodora 77: 105-140. Standley, P. C., et al. 1958-1977. Flora of Guatemala. Fieldiana, Bo Wasshausen, D. C. 1975. The genus Aphelandra (Acanthaceae). neice 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. —680. Williams, L. O. 1981. The useful plants of Central America. Ceiba 24: 1-381. APPENDIX OVERALL DISTRIBUTION PATTERNS OF ACANTHACEAE NATIVE TO BELIZE a. (62.5%): Aphelandra aurantiaca, A. scabra; Barleria oenotheroides, Blechum pyramida- 1; Bravaisia berlandieriana, Dicliptera inutilis, D. sexangularis, D. sumichrastii; Elytraria imbri- cata, Hygrophila costata; Justicia aurea, J. breviflora, J. candelariae, J. comata spicigera, Lepidagathis alopecuroidea; Mendoncia lindavii, M. retusa: i iit callistachyum, Oz oe Ruellia geminiflora, R. hookeriana, R. matagalpae, R. nudiflora. 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%): Justicia ensiflora, Ruellia pygmaea. Endemic (2.5%): Louteridium chartaceum. Contr. Univ. Michigan Herb. 21: 175-195. 1997. A REVISION AND REDEFINITION OF PSEUDABUTILON (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 of 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 delimited 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 1s 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 of Abutilon 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 VOLUME 21 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 Pseudabutilon with two subgenera, subg. Wissadulas- trum (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. Wissadulastrum (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.— Lecrorype, designated by Fryxell, 1988: Pseuwdabutilon scabrum (C. Presl) k..E. Pr, Abutilon sect. Anasida Griseb., Fl. Brit. W.I. 78. 1859.—Tyee: Abutilon um- bellatum (L.) Sweet. Wissadula sect. Abutilastrum Baker f., J. Bot. 31: 71. 1893.—Type: 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 linear, 1-11 mm long. Flowers solitary 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, pallid, 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. 1997 FRYXELL: PPEUDABUTILON 177 1. —_ 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 to small stellate hairs (or stellate hairs absent 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- uras. 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. thurberi. 3. Carpels 6-8; petals 7- a mm long; leaf venation palmate; stem vesture eR understory of stellate hairs; Argent pee ee na. . Upper surface of leaf pie -pubescent or with bifurcate hairs, these never more than mm long, the pubescence sometimes farinose; stems minutely stellate-pubescent (or ce spreading hairs sometimes present). 4. Calyx as pedicel) with minute glandular hairs; carpels usually 5, 5-6 mm long; seeds glabrou ah ane aE hairs sometimes present on stems; mericarps with spines ca. 1.5 mm long, the endoglossum present; Mexico, Guatemala. 13. P. orientale. 5. Long simple hairs lacking on stems; mericarps with spines 3—3.5 mm long, the ae sum absent; Bolivia, Paragua Pap ene 4. Calyx (and pedicel) usually HST -pubescent, lacking glandular hairs; carpels 5 ef ve: 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 . P. cymosum. 6. Mericarps 4-9 mm long, with spines obsolete or up to 4 mm long; endoglossum present or absent. 7. Flowers short- Paes (less than 5 mm) or subsessile; carpels 8-10. . Caly inflorescence racemiform or paniculate; endoglossum com- pletely covering ms lowermost seed; spines of mericarps obsolete; Mexico, Gua- temala, Honduras 15. P. scabrum. 8. Calyx 7-8 mm eilone. 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 eolitary: Brazil a 14. P. pintoi. 7. Flowers pare pedicellate Veen often more than 1 cm long); carpels 5—LO. 10. Stems with long spreading simple 11. Flowers and fruits in axillary Heate endoglossum absent; eee ad. 8. P. umbellatum. 11. Flowers and fruits solitary or paired in the leaf axils; endoglossum present; South Americ 12. Stipules 7 12 mm long; calyx pilose and stellate-pubescent; ee leaf surface with minute often bifurcate hairs. 4. P. cinereum. 12: ee 3-5 mm ne calyx stellate- eee upper leaf surface with nute stellate hair 2. P. callimorphum var. See 10. Stems ee long ae 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 ee ca. 1 mm long; stipules 4-6 mm long; endoglossum absent; Per 5. P. cowanii. 14. Leaves broadly ovate; calyx 5-8 mm long, half-divided or less, hirsute; rpels 5-8, with divergent apical spines 1-4 mm long; stipules 8-10 mm long; eas present or absent; Mexico, West Indies, Central and outh Americ 178 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 15. Hairs of calyx | mm long or less, twisted and whitish; carpels 5, the endoglossum absent; inflorescences of axillary racemes with subumbel- late elements forming an open terminal inflorescence; Bolivi “to. P. leucothrix. 15. Hairs of calyx 3-4 mm long, straight and brownish; carpels 5-8, the endoglossum present or absent; inflorescences axillary umbels, some- times forming dense terminal inflorescences. 16. Carpels with divergent apical spines 2-4 mm long; calyx 6-8 mm long, hirsute, the hairs 3-4 mm long; endoglossum absent; widespread. 18. P. umbellatum. 16. Carpels with apical spines 1-2.5 mm long; calyx 5-6 mm long, stellate- pubescent; endoglossum present or absent; Galapagos Archipelago. 7. P. de ie ade 13. Flowers usually solitary or in axillary fascicles; carpels 5—10, the ad spine ually 2 mm long or less; South America. i Endoglossum absent; flowers solitary or in axillary fascicles, often aggre- gated into dense inflorescences; minute hairs on upper leaf surface often bifurcate. 19. P. virgatum. 17. Endoglossum present or absent; flowers usually solitary (or paired) in e leaf axils; minute hairs on upper leaf surface usually stellate. 18. Carpels 5—6; petals 5-6 mm long; stipules 1-2 mm long; Peru. 2. P. nigropunctulatum. 18. Carpels 5-10; petals 8-12 mm long [unknown in P pintoi|; stipules 4-10 mm long. 19. Carpels 8-10. 20. Pedicels 0.3—0.4 cm long; Brazil. 14. P. pintoi. 20. Pedicels 1.5—6 cm long; Paraguay, Argentina. . P. callimorphum var. intermedium. ra oO . Carpels 5-8. 21. Carpels 7-8, with spine (or apiculum) less than | mm long: endoglossum present; Paraguay, Bolivia, Argentir 16. P. stuckerti. 21. Carpels 6-8, with spine 1-2.5 mm long; endoglossum present or absent; Galapagos Archipelago. 7. P. depauperatum. 1. Pseudabutilon 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: Bo.ivia. 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 sér 2. 7: 458. 1907, non Wissadula balansae Baker f., J. Bot. 31: 69. 1893 —Typre: PARAGUAY. San Bernardino, Hassler 390 (syntype: NY!); PARAGUAY. Sierra de Amambay, Esperanza, Hassler 10564 (syntypes: CTES! G!). Subshrubs to | 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 solitary 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. ruits 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. Loo FRYXELL: PPEUDABUTILON 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 2627 (US).—Santa Cruz: 22 km N de Gutiérrez (camino Camiri-Santa Cruz), 19°20'S, 63°30'W, Krapovickas & Schinini 3145] (CTES, MO). The Venezuela specimen cited above (Aristeguieta 2021) 1s 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, Kong]. Sven- a Vetenskapsakad. Handl. 43(4): 105. 1908. Sida callimorpha Hochr., Bull. Herb. Boissier, sér. 2. 5: 295. 1905.Wissadula callimorpha (Hochr.) Hassl., Bull. Herb. Boissier, sér. 2. 7: 455. 1907.—T ype: 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- mately 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 | 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 pubescent. Distribution. Paraguay, Bolivia, and Argentina. ey ae CIMENS 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, ae Vélaz, aie et al. 2093 (CTES, NY); Depto. 1° de Mayo, Colonia Benitez, Schulz 9043 (CTES, NY): Depto. 12 de Octubre, Estancia Valverde, : W de General Pinedo, el limite con ee del ae Schulz 15346 (CTES, NY).—Corrienres: Corrientes, ren 43692 (CTES, N 3. Pseudabutilon callimorphum var. intermedium Hassl., ie Spec. Nov. Regni Veg. 7: 76. 1909. —Type: PARAGUAY. Gran Chaco: ad ripam occidentam flum. Paraguay prope Santa Rita, latit. S 23°20! "93°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 | cm 180 CONTR. UNIVERSITY 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 | 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 ae Enciso, ruta Trans-Chaco, Schinini & Bordas 16484 (CTES, NY); Ruta Trans-Chaco, 21°30'S, 61°1S'W, Schinini & Bordas 16505 (CYTES, NY). Argentina. Formosa: Riacho Monte Lindo, cae ey etal. 2151 (CTES, NY). 4. Pseudabutilon cinereum (Griseb.) Krapov., Bol. Soc. Argent. Bot. 24: 206. 1985. Abutilon cinereum Griseb., Abh. Konigl. Ges. Wiss. Géttingen 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, palmately 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 like 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 | 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. Bolivia. SANTA Cruz: Prov. Cordillera, 7 km W de Boyuibe (20°20'S, 63°20'W), Krapovickas & peg fe 308 (CTES, F, NY, US).—Tarua: Ruta Tarija-Villa Montes, Bajada de Chimeo, Krapovickas et al. 19232 (CTES, NY); Prov. O’Connor, Puerto Margarita, pe Pilcomayo, 21°10'S, 63°50'W, Krapovickas - eee 39096 (CTES, MO). Argentina. CaTra-mMarca: Depto. Andal- gala, 3 km SE of Andalgala, Cantino 774 (CTES); Depto. Belén, 11.7 km E of Rt. 40 on road to Pozo de Piedra, Spooner & Clausen 4602 (NY).—Corpbosa: Depto. Isehilin, Dean Funes, Cuezzo 706 (MO).— seri Depto. 1° de Mayo, Colonia Benitez, Schulz 9204 (CTES, NY).—Formosa: Depto. Formosa, acho Monte Lindo, Cristobal et al. 2121 (CTES, NY).—La Riosa: Depto. Capital, near Carrizal, Peders- en Tee (CTES).—Menpboza. Depto. Godoy Cruz, Cacheuta, O’Donell 1134 (MO).—Sa.ta: Depto Metan, Rio Juramento, Ruta 34, Krapovickas et al. 18597 (CTES, NY), ainda et al. 18599 (CTES, NY); Metan, O’Donell 4628 (MO).—SantTiaco beL Estero: entre Monte Quemado y Campo Gallo, Castiglioni-Ragonese 7144 (CTES); Depto, Rio Hondo: Los Naranjitos, Legname 705 (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.—Typr: 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: PPEUDABUTILON 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 material. 6. Pseudabutilon cymosum (Triana & Planch.) Fryxell, comb. nov. Abutilon cymosum Triana & Planch., Ann. Sci. Nat. Bot. ser. 4. 17: 185. 1862.—Tyre: PANAMA. 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, pallid. 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. ADDITIONAL SPECIMENS EXAMINED. Colombia. “de Pandi a Fusagasuga,” Goudot s.n. (syntype: P as photo F-35450). 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 alliance 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. Abutilon depauperatum (Hooker f.) Andersson ex B. L. Rob., Proc. Amer. Acad. Arts 38: 173. 1902.—T ype: GALAPAGOS 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. | 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— | times length of blade), with pubescence like that of stem; stipules 7-10 mm long, 1-2 mm wide, narrowly elliptic. Flowers solitary 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 547 (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 B Howell 9352 (CAS).—CHATHAM ISLAND: Snodgrass & Heller S07 (CAS).—DUNCAN ISLAND: Snodgrass & Heller 702 (NY).—GARDNER ee Snodgrass & Heller 632 (CAS).—INDEFATIGABLE IsLAND (ISLA SANTA CRUuz): 3 mi N of Academy Bay, Bowman 55 (CAS); Academy Bay, Fosberg 44721 (MO), Schimpff 48 (MO).—JAMES TSCA AND: James Bay, Howell 9681 (CAS).—Tower IsLANnb: 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 of A. umbellatum. Bates’s view seems more reasonable and is adopted here. 8. Pseudabutilon ellipticum (Schltdl.) Fryxell, comb. nov. Abutilon ellipticum Schit- dl., Linnaea 11: 368. 1837. Sida elliptica (Schltdl.) Steud., Nomencl. Bot. ed. 2. 2: 577. 1840.—Tyerre: Mexico. Michoacan: Tlalpujahua, 1830, Keerl s.n. (holotype: HAL!; isotype: BR!). 1997 FRYXELL: PPEUDABUTILON 183 Abutilon sidoides Hemsl., Diagn. pl. nov. Mex. 2: 24. 1879, non Abutilon sidoides Dalzell & Gibson, 1861. Abutilon hemsleyanum 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. 893.—T yee: 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!). ads membranaceum Baker f. ex Rose, Contr. U.S. Natl. Herb. 3: 312. 1895.—Tyre: Mexico. Nayarit: Tepic, 1892, Palmer 1959 (holotype: BM!; isotypes: F! GH! NY! US!). Shrubs 1-2 (-3) m tall, the stems coarsely stellate-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, l-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).—Cuiapas: Mpio. de Bochil, 3 km al SO de Puerto Cate, Garcia 594 (CHAPA, NY); Pinada, Siltepec, Matuda 1905 (ARIZ, NY, TEX).—Cottima: Rancho El Jabali, 22 km NNW of Colima, 19°27'N, 103°42'W, Sanders et al. 11000 (NY, UCR). ae at FEDERAL: Cerro de Santa Catarina, cerca de Santa Catarina, Rzedowski 26052 (ENCB, LL, NY); Almella cree delegacién de Tlahuac, Ventura 3786 (ENCB, NY).—DurRanco: 2 mi W of Revoeaers Breedlove 18926 (CAS, NY); Cerro de los Remedios, Ciudad de Durango, Patoni 289 (MEXU, NY).—Guanasuato: Mpio. de Cortazar, 2 km al NW de La Gavia, Rzedowski 40946 (ENCB, IEB); Mpio. de Jerécuaro, 3 km al E de Jerécuaro, Rzedowski 41713 (JEB, NY).—Guerrero: Montes de Oca, Hinton et al. 11692 (NY); ae Chilpancingo, camino a Tejocote, 17 km al S de Chilpancingo, Koch 5 Fryxell 83266 (CHAPA, —HIpatco: vicinity of Tepeji del Rio, Fryxell & Bates 2174 (BH, NY); Cafiada del Salitre, Dae See & Cristébal 23475 (CTES, NY); near Tula, Pringle 9689 re 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 fs 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 Se Pringle 4583 (MEXU, MICH, NY).—MeExico: Temascaltepec, Ocotepec, Hin- ton el al. 2916 (NY); Cerro de Santa Cruz, Sierra de Guadalupe, Matuda et al. 29520 (MEXU, NY); o. de ee cerca de San Juan Ixhuatepec, Rzedowski 23296 (ENCB, LL, TEX); San Andrés, m al SW de San Cristébal Ecatepec, Rzedowski 27642 (ENCB, NY).—MICHOACAN: Coalcoman, Hinton 12913 (MICH, NY); Mpio. de Zinapécuaro, SE del Lago Cuitzeo, 1-2 km al E de Coro, Koch & Fryxell 77397 (CHAPA, LL, NY); Cerro Grande de Cujaruato, al SW de la Piedad, Rzedowski & McVaugh 555 (LL, MICH, NY); Mpio. de Morelia, Cafiada del Rio Grande, cerca de Cointzio Rzedowski 45377 (ENCB, IEB, MEXU, TEX).—Nayaarit: Tepic, en 1892, Palmer 1959 (BM, F, GH, 184 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 2] US); Mpio. Tepic, 4 km al SW del Cuarentenio, brecha al Cora (21°26'N, 104°56'W), Tenorio et a 16961 (MEXU, NY).—Nuevo Leon: 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).—Puesca: Cerro de la Yerba, vic. of San Luis Tultitlana apa, eae 2604 (NY); pine Caltepec, Cerro El Gavilan, al SE de een es Tenorio & Romero de Tenorio 7734 (MEXU, NY, TEX oe roa La Trinidad, opales, Arguélles 2007 (MEXU. NY): Mpio. de ee 3 km al S de San Javier, Fernan- a 3053 (ENCB, IEB, NY).—SINALOa: naa a Tamiapa, eovilhe 5840 aaa I miS of EI Palmi- to, 23°36'N, LOS°S1'W, Sanders et al. 4372 (NY, UCR).—Zac : 55 m1 N of Fresnillo, 5 mi N of Rio sea Rollins & Tryon 58264 (LL). Guatemala. pean TENANGO: near Parramos, Standley 59874 (F, NY).—QuicHE: Nebaj, Skutch 1729 (NY); El Molino, 4 km from Chichicastenango, Molina et al. anus (NY).—SaAcaTEPEQUEZ: 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. reas ae 25919 (NY). 9. Pseudabutilon glomeratum Fryxell, sp. nov.—T ype: Botivia. 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 cellulis diviso, cellula infe- rior lateraliter reticulata. Subshrubs ca. | 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. ADDITIONAL SPECIMEN EXAMINED. Bolivia. CHUouISACco: Prov. Oropeza, de Sucre pasando Yotala, entrando por la quebrada de Nujchu, Beck 888/ (US). The new species is notable for its obsolete pedicels and hence glomerulate inflorescence, and for its fimbriate endoglossum. 10. Pseudabutilon leucothrix Fryxell, sp. nov—Type: Botivia. Santa Cruz: Nuflo de Chavez Prov., Lomerio, 12 km al N de la comunidad 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: MO! USZ!). 1997 FRYXELL: PPEUDABUTILON 185 Suffrutex 1-2 m altus, caulibus et foliis sparse et minutissime puberulentibus; calycibus et pedicellis fructibusque pilis albis tortis arachnoideis; corollis albis; inflorescentiis axillaribus racemiformibus apertis; mericarptiis 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-S 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). The new species is named for the distinctive pubescence found in the inflores- cence. The open structure of the terminal inflorescence is also distinctive. 11. Pseudabutilon longepilosum R. E. Fr., Kongl. Svenska Vetenskapsakad. Hand. 43(4): 107. 1908.—T ype: ARGENTINA. Catamarca: Puntilla de Villavil, White s.n. (syntype: BM!); Prov. Rioja: Stuckert 934] (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 like that of stem; stipules filiform, 7-8 mm long, pilose. Flowers axillary, solitary (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 (slightly 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. ] 186 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Distribution. Northwestern Argentina, apparently known only from the two syntypes. 12. Pseudabutilon nigropunctulatum (Ulbr.) R. E. Fr., Kongl. Svenska Vetens- kapsakad. Hand. ser. 3, 24(2): 11. 1947. Abutilon nigropunctulatum Ulbr., Jahrb. Bot. Syst. 54 (Beibl. 117): 57. 1916.—Type: Peru. Ad viam ferream a Lima oppido ad Oroyam gerentem prope vicum San Bartolomé dictum, Weberbauer 5301 (holotype: B as photo F—9281!; isotypes: F! GH! US!). Shrubs 1.5 m tall, 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 I-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).—Lma: carr. Lima a Oroya, Km 70, Krapovickas 8260 (CTES, NY); San Bartolomé, Lopez 3759 (CTES): Matucanos, L6pez 426] (CTES).—CaJAMarca: Prov. Cajamarca, road to Chilete, 5.5 km below San Pablo, Hutchison & Wright 5086 (UC, NY); Prov. Contumasa: Cascas, Ratmondi 7573 (CTES).—La Liserrap: Proy. Trujillo: Samne, Plaza Pampa, de Cevasco s.n. (CTES). 13. Pseudabutilon orientale (Standl. & Steyerm.) Fryxell, comb. nov. Abutilon orientale Standl. & 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 | 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: PPEUDABUTILON 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. Cotima: 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).—Jatisco: 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 Estacién 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 al NO de Puerto Escondido por la carretera a Pinotepa Nacional, Koch et 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. Marfa Huatulco: carretera a Salina Cruz, Koch et al. 79538 (CAS, CHAPA, NY).— SINALOA. Mazatlan, Gonzdlez-Ortega 6785 (US). 14. Pseudabutilon pintoi Monteiro, Portugal. Acta Biol. B, 12: 142. 1973.—Type: BraziL. Bahia: Cruz das Almas, Pinto 50-35 (holotype: RBR n.v.; iso- types: CTES [fragment]! IAL 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°11'S, 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, Relig. 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. Hand. 43(4): 104. 1908—T ype: 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.—Tyre: MExIco. Jalisco: Guadalajara, La Barranca, Jones 27162 (lectotype, designated by Fryxell, 1988: POM-—192430!; isolectotypes: BM-2! NY! POM-193309! UC! US! Pseudabutilon inornatum Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 15. 1943.—T ype: 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, pallid. 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. CuiApas: Mpio. Amatenango Frontera, along Rio Cuilco between Nuevo Amatenango and Frontera Comalapa, erie e 41774 (CAS, NY); Mpio. Arriaga, at La Mina microwave station, Sete 56295 (CAS, NY); Mpio. Tuxtla Gutiérrez, on road to Chicoasen, Fryxell & Lott 3236 (M NY).—Co ta: forested hills 2— a i SE of La Manzanilla, above Bahia Tenacatita, McVaugh nie (MICH, N Y); 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 Unién por la terraceria a Coahuayutla de Guerrero, Koch & Fryxell 83119 (CHAPA, NY, TEX): eee La Union, 86.5 km al NE de oe camino a Cd. Altamirano, Martinez & Barrie 5479 (MEXU, NY).— Jatisco: 2.5 km al W de laque, Mpio. Cihuatlan, Cuevas et al. 555 (IBUG, NY); ae Tonala: Barranca de San Gaspar, ie et al. 392 ler eee ee Mexia 1239 (BM, DS, F, US); L Barranca de Guadalajara, Pringle 172] (BM, K, MICH, NY, S, TEX, US).—Nayarir: Hae San Bilas, Km 19 de la autopista Tepic—Mazatlan (21°34! N, 105°04' . “Téllez et al. 12703 (MEXU, NY); Mpio. mee Km 12 del camino de Xalisco al Malinal (21°37'N, 104°58'W), Téllez & Bojorquez 12619 (MEXU).—Oaxaca: 71.4 mi NW of Salina Cruz on Hwy 200, Fryxell & Lott 3397 (MEXU, NY): Mpio. ie rae Chimalapa: entre 12 y 15 km de la carretera “Juchitan —~Tuxtla gee pe 160-220 m, Koch & . ed 75349 (CHAPA, NY, TEX): ee eae as al S de Mazahua, 13 km al NE de Ventosa, 16°40'N °S4'W, Martinez 1260 greet NY); 5 | E de Teotitlan del Gaaiho. eae la carr. a Foautla. Rzedowski 37072 (ENCB, NY): ladera S a c. rro de Guiengola, Torres et al. 673 (MEXU, NY).— Puesia: Coxcatlan, Purpus 4186 (NY).—SINALOA: 23 km W of San Ignacio, Fryxell & bates ce (BH, NY); Imala, Gentry 4966 (ARIZ, NY); Cerro oan - of Pericos, Gentry 5726 (AR EX).— Sonora: Mawarichi Arroyo, Rio Mayo, Gentry 1317 (ARIZ, TEX); 2.7 mi W of Santa = e oe 28°23'N, 109°19'W, Goldberg 77--53 (ARIZ); Sierra ae : Cebollita, 1 mi S of Nuri on the road to Tesopaco, 28°6'N, 109°19'W, pata 3703 (NY, UCR); La Higuera, ca. 7 mi N of Alamos on the San Bernardo road, 27°05'N, 108 . Van Devender & Van Devender 90-591 va NY).—VERACRUZ: Zacuapan and eae on te Aes Maria, Purpus 2239 (GH, MO, NY, UC); Mpio. Dos Rios, Plan del Rio, Ventura 2655 (A ENCB, LL, NY, TEX); Mpio. Dos Rios: La Cumbre, eae 12032 (CHAPA, NY). a EL PARAISO: Sierra de la Villa Santa, Rio California, Molina et al. 31252 (NY). 16. zecudapualon stuckertii R. E. Fr., Kongl. Svenska Vetenskapsakad. Handl. ‘ . 1908.—SyYNTYPES: Agen TINA. Tucumdan: Vipos, Lillo 2299; La Rioja: Diana: Schmidke 13225 (CORD); Cérdoba, Stuckert 1029 (CORD, S); Cordoba, Stuckert 12983; Cordoba, Fielding s.n. (BM). 1997 FRY XELL: PPEUDABUTILON . 189 Pseudabutilon callimorphum var. friesii (Hassl.) R. E. Fr., Kongl. Svenska Vetenskapsakad. Hand. 43(4): 106. 1908. Wissadula callimorpha var. friesii Hassl., Bull. Herb. Boissier, sér 2, 7: 457. 1907. Wissadula pedunculata R. E. Fr., Arkiv Bot. 6(2): 12. t. 2. figs. 1-5. 1906.—Type: Botivia. 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. Leat 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. TaruA: Prov. Gran Chaco, 10 km S de Palmar Grande, camino Yacuiba-Villa Montes, Krapovickas & Schinini 31102 (CTES, NY). Paraguay. BOQUERON 14 km N de Filadelfia, Krapovickas & Cristébal 44251 (CTES, TEX).—Cuaco: Agua Dulce, 20°14'S, 60°7'W, Schinini & Bordas 18082 (CTES, MO).—Nueva Asuncion: Ruta Trans-Chaco, 21°26'S, 61°25'W, Schinini & Bordas 16407 (CTES, MO). Argentina. CATAMARCA: Depto. La Paz, El Bello, Brizuela 836 (CTES); Depto. Andagalé, 9 km SE of Andagala, via the road to Cuesta de la Chilca, Cantino 305 (ARIZ, GH).—Cuaco: 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 José, General Pinedo, Schulz 11288 (CTES).—Corposa: Depto. Santa Maria, Cuesta San Roque, de la Sota 1395 (MO).—CorrienTEs: Depto. Mercedes, Paso Lucero, Rio Corrientes, Krapovickas & Cristébal 14763 (CTES).—Formosa: Depto. Pirani, Palo Santo, Marufak 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 Banado de Cautana, Hunziker & Cocucci 14867 (CTES).—La Ruosa: Depto. Gob. Cordillo, Campo Experimental del INTA, Biurron 1433 (CTES); entre Patquia y Capital, Dawson & Guarrera 3197 (CTES); Depto. Rosario, Vera Pefialoza, Rio Totoral, a 5 km de Chelco, Hunziker et al. 14163 hee Se Depto. Giiemes, Juramento, Krapovickas a eat 30431 (CTES, MO) —SANTIAGO DEL Estero: Depto. Robles, Beltran, Garcia 719 (NY).—TucuMAN: entre Las Tejas y Tucuman. Castiglioni A Ragonese 672 (CTES); Depto. Trancas, Vipos, a & Crist6bal 18377 (Cres, NY). 17. Pseudabutilon thurberi (A. Gray) che comb. nov. Abutilon thurberi A. Gray, Mem. Amer. Acad. Arts n.s. 5 (Pl. Thurb.): 307. 1854.—Type: Mex- Ico. Sonora: ae Thurber 911 (holotype: GH as photo US!; iso- types: K! MO! N Pseudabutilon sonorae ee 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 190 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 stem; stipules 3-11 mm long, | mm wide, linear, prominently |-nerved, sparsely setose, more or less spreading, persistent. Flowers solitary in the leaf axils; pedicels ().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. hallf- 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. 2mm 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 900-1100 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, 11.7°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 Pseud- abutilon 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.—T ype: Jamaica. Browne s.n. (holotype: LINN-866.16!). Shrubs 1-2 m tall, 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 IsLANb: Clarence Town, Correll 48150 (FTG, LL, NY).—Granpb 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. Mare, Ekman 6669 (NY, S); Depto. de l’Artibonite, vicinity of Gros Morne, Leonard 9964 (NY). Dominican Republic. 1997 FRYXELL: PPEUDABUTILON 191 Prov. Azua: entre la entrada a Puerto Tortuguero y Hatillo, 18°25'N, 70°27'W, Garcia et al. 66 (NY, TEX); Prov. Peravia, NE of Galen, 18°19N, 70°14'W, Mejia & Zanoni 7008 (NY, TEX); Prov. Azua, 3 km W of Cruce de Ocoa, 18°21'N, 70°29'W, Thompson et al. 9397 (CM, NY). Jamaica. vicinity of Kingston, Britton & Hollick 1725 (NY); Port Royal, Proctor 31091 (LL, NY); Clarendon, | 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. Sr. pea near Charlotte Amalie, Rose 3176 (NY, US).—Sr. Croix: Are- ria's Hope, Thompson 193 (NY ).—ViRGIN Ee Valley, Fishlock 183 (NY), Fishlock 264 (NY).— TorTova: Reef Island, Britton & ae 0 (NY). Lesser Antilles. GRENADA: Grand Anse, Dec 1904, Broadway s.n. (NY); Carriacou, oe 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). Curagao. Soto, Arnoldo- aa 3645 (NY); without locality, Boldingh 4811 (NY): Banks, ee Rica & pee an Y). xico. JALisco: Sierra de Manantlan, 2-4 km E of Zapotillo, 19°38'20"N, 104° , Cochrane et a 12312 (WIS); ee near Guadalajara, Rose & Painter 7355 ae ore ny 20-40 del camino de terraceria hacia la Presa de Aguamilpa, Téllez 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 § 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, See 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, ae 14296 (A, F, NY, UC); Mpio. Puente Nacional: Pachuquilla, Ventura 9425 (ENCB, MEXU, NY).— Yucatan: Cueva de Sihunchen, 15 mi N of Muna, Butterwick 13] (LL). Nicaragua. GRANADA: 8 km N de Granada on road to Paso de Panaloya, Stevens 4623 (MO, NY).—Manacua: Km 14 carr. a ce de Jiloa, 12°10'N, 86°21'W, Moreno 3566 (MO, NY); lado O de Laguna de Jiloa, Sandino 265 er NY).—Este i: sobre el camino a Limay, Pee 86°36'W, Moreno 2014 (MO, NY); on road to Pueblo Nuevo, 13°25'N, 86°25'W, Stevens Eee (MO, NY).—MataGacpa: SW slopes of Cerro El Pilon, 12°37! N. 86°02'W, Stevens 9430 (MO, NY).— oe 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); Pefias Blancas, 1 km N of Costa Rican boundary, Atwood 1808 (MO, NY). Wenezuela. Ista Marcarita, Miller & Johnston 152 (NY).—ARAGuA: Maracay, Krapovickas 15571 (CTES , NY).— Fatcon: Coro, Wingfield 5033 (CORO, NY).—Botivar: Ciudad Bolivar, Holt & Gehringer 17 (NY).— Distriro FepeRAL: Pto. La Cruz, Hacienda Panarigua, Pittier 11652 (NY).—Corpes: Depto. El Pao, Paraima, Trujillo 5438 (MY, NY).—Lara: Depto. Palovecino Agua Viva via Terepaima, Burandt 2434 (NY); Barquisimeto, Saer 207 (NY).—MeEnripa: Distr. Sucre, Las Gonzales—San José road, ca. 20 km SW of Mérida, 8°25'N, 71°20'W, 9-10 Nov 1990, Dorr & Barnett 7653 (NY, TEX).—Miranpa: SO del Valle de Caracas, 10°30'N, 66°53'W, Ramirez 1049 (NY).—Zutta: Jardin Botanico de Maracaibo, se ce 13108 (NY). Colombia. Bocivar: vicinity of Turbaco, Killip & Smith 14213 (NY).—MAGDALE- of Bonda along road from Santa Marta to Rio Hacho, 11°15'N, 74°06'W, Kirkbride 2492 (NY); ae Marta, Smith 490 (LL, NY). Norte DE SANTANDER: between Chindcota and La Esm alda, see & Smith 20924 (NY). Ecuador. EL Oro: Santa Rosa, Asplund 15707 (S); Huaquillas. frontier of Pert, Jaramillo & lees oe (QCA).—Guayas: | km NW of Chanduy towards Atahualpa, Holm-Nielsen 2154 (AAU, F, O, S); 20 km § of Manglaralto, near Palmar, Holm-Nielsen et al. 2483 (AAU, F, GB, NY); Playas, ee 166 (F, QCA, US); La Puntilla, Salinas, Svenson 11253 (GH, NY, UC, US).—Losa: 10 km W of Vilcabamba, Harling & Andersson 21774 (QCA).—Manasi: El Recreo, Eggers 15778b (F, K); Portoviejo, Mille 1963 (F). Peru. Cajamarca: 2 km E of Chilete, ee & Keil 2325 (NY). 19. arabes virgatum (Cav.) Fryxell, comb. nov. Sida virgata Cav., Icon. 1: . 73. 1791. Abutilon virgatum (Cav.) Sweet, Hort. Brit. ed. 1. 1: 53. anne 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).] 192 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: Bo.ivia. 1847, Bridges s.n. (BM! CTES [fragment]! K!). Abutilon saltense Hassl., Repert. Nov. Sp. Regni Veg. 12: 498. 1913.—Typ ARGENTINA. Salta: in dumento Rosario de la Frontera, Lillo 3877 (LIL n.v., “et herb. Hassler” G7). 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, pallid. 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. Losa: 4-6 km N of Yangana, on road to Vilcabamba, Harling & Andersson 21650 (GB); Mollococha, ca. 10 km W of Vilcabamba, Harling & 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, I/tis et al. 619 (MO).—Huanuco: 7.8 km NE of Hudnuco at Hda. Colcuy, Bird 1092 (MO, NY). Bolivia. Cotafia am IIlimani, Nov 1911, Buchtien 3215 (MO, NY).—Cocuapampa: Panduro, near Rio Caine, Cardenas 2434 (NY, US); Cochabamba, eee 3784 (CAS, DS).—La Paz: Inquisivi, 4 km SE of Inquisivi, 16°55'S, 67°6'W, Lewis 35093 (MO sia Ruta Tarija-Villa Montes, 15 km E de Entre Rios, ae et . 19129 lane LL); Prov. Arce, 31 km § of road to Entre Rios on road to Padayca 9'S, 64°40'W, Solomon 10573 (M MO, NY). Paraguay. BoOUERON: 5 km NW de Mariscal Se aera Pa & Cristébal 44374 (CTES, TEX).—Cuaco: Chaco Paraguayo, F. Olimpo, Rojas 13617 (MO, TEX); aioe Leon, 20°26'S, 60°15'W, Schinini & Bordas 17830 (MO).—CENTRAL: Jardin Botanico y Zooldgico, dad, Asuncion, 25°20'S, 57°28'W, Pérez 97 (AS, MO); Cerro Kof-Aregua, Schinini 4019B (CTES, a Argentina. CATAMarca: Depto. La Paz, El Rio de la Dorada, Brizuela 959 (TEX): Depto. Ancasti, La Brea, Brizuela 1302 (MO); ee een 16 km SE of Andalgala on noes to Cuesta de la Chilca, Cantino 766 (ARIZ, GH).—Co Depto. Union, Bell Ville, Parque Tau, cee 584 (CTES, NY); Depto. Punilla, San en Vellofane ye (CAS).—Corrikntes: Corri , Kra kas 43693 (CTES, NY); Depto. Berén de Astrada, 46 km W de Ita Ibaté, Valencia, ee 14051 (CTES, F, NY).—Jusuy: Depto. Tumbaya, acceso a Purmamarca, Krapovickas & Crist6bal 17636 (CTES, NY).—LaA Pampa: Depto. Lihuel Calel, Sierra de Lihuel Calel, =o Wea et al. 22625 (CTES, NY).—MEnpoza: Depto. San Rafael, Serrito, Spegazzini 22556 (NY).—SALTA: Dapio, Chi- coana, Chicoana, Los Los, Krapovickas et al. 28260 (CTES, NY); Depto. tae de la Frontera, 1997 FRYXELL: PPEUDABUTILON 193 Almirante Brown, O’Donell 5392 (CAS, TEX).—TucuMAn: Depto. Leales, La Florida, Ruta Nac. 9, Krapovickas & Crist6bal 17335 (LL, NY); Depto. Trancas, Vipos, Ruta 9, Krapovickas & Cristébal 17742 (CTES, LL, MO, NY). ExcCLUDED NAMES Pseudabutilon hitchcockii Ulbr., Notizbl. Bot. Gart. Berlin 11: 522. 1932. = Abutilon dispermum (Hochr.) Fryxell. Pseudabutilon langlassei Hochr., Annuaire Conserv. Jard. Bot. Genéve 20: 120. 1917. = Allosidastrum hilarianum (C. Presl) Krapov., Fryxell & D. M. Bates. Pseudabutilon lozanii (Rose) R. E. Fr., Kongl. Svenska Vetenskapsakad. Hand. 43(4): 100. 1908. = Allowissadula lozanii (Rose) D. M. Bates. Pseudabutilon pringlei (Rose) R. E. Fr., Kongl. Svenska Vetenskapsakad. Hand. 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. Genéve 20: 119. 1917. = Allosidastrum interruptum (DC.) Krapov., Fryxell & D. M. Bates. Pseudabutilon spicatum (H. B. K.) R. E. Fr., Kongl. Svenska Vetenskapsakad. Hand. 43(4): 98. 1908. = Briquetia spicata (H. B. K.) Fryxell Pseudabutilon weberbaueri Ulbr., Jahrb. Bot. Syst. 54 (Beibl. 117): 60. 1916.— yPE: PERU. Cajamarca. Prov. Jaén: 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, comb. nov. Sida dictyocarpa Morong & Britton, Ann. New York Acad. Sci. 7: 56. 1892, nomen nudum, non Sida dictyocarpa F. Muell. ex Benth. (1862), nec Sida dictyo- carpa Griseb. ex K. Schum. (1891). Sida dictyocarpa var. cordobensis Baker f. ex Morong & Britton, Ann. N. Y. Acad. Sci. 7: 56. 1892, nomen nudum. LITERATURE CITED Bates, D. e 1966. Chromosome numbers in the Malvales. I. Gentes Herb. 10: 39-46. —. 1971. Malvaceae. In Flora of the ane Islands, ed. 1. L. Wiggins and D. M. Porter, 666- 694, Cae Stanford University Pres . 1976. Chromosome numbers in the eae III. Miscellaneous counts from the Byttneriaceae and Malvaceae. Gentes Herb. 11: 143-150. .1978a. Bastardiastrum, a segregate from Wissadula (Malvaceae). Gentes Herb. 11: 311-328. . 1978b. Allowissadula, a new genus of North American Malvaceae. Gentes Herb. 11: 329-354. 194 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Slee A. 1974. Recuentos cromosémicos en Malvaceas. Bol. Soc. Argent. Bot. 15: 403-410. Fries, R. E. 1908. Entwurf einer setae ea der Gattungen Wissadula und Pseudabutilon. Kongl. Sve ae Vetenskapsakad. Handl. 43(4): 1-114 + 10 plates Fryxell, P. A. 1976. New species and new combinations in Briquetia and Hochreutinera, 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. ears P. A., and D. M. Stelly. 1993. Additional chromosome counts in the Malvaceae. Sida 15: 639- Goi R. 1993. ein Cavanillesianum. Fontqueria 38: 1-248. ochreutiner, B. P. G. 1920. Organes carpiques nouveaux ou méconnus chez les Malvacées. Annuaire Conserv. Jard. Bot. Genéve 21: 347-387 Hutchinson, J. 1967. Malvaceae. In The genera of flowering plants by J. Hutchinson, 2: 536-567. Kearney, T 951. The American genera of Malvaceae. Amer. Midl. Natural. 46: 93-131. i. OS2: Notes on Malvaceae HI. Abutilon and Poeudobuntent in the Galapagos Islands. Madrono 11: 285-289. 1955. A tentative key to the North American species of Abutilon Miller. Leafl. W. Bot. 7: 241-254 -1958. A tentative key to the South American species of Abutilon Miller. Leafl. W. Bot. 8: Krapovickas, e% 1957. Numeros prea de Malvaceas americanas de la tribu Malveae. Rev. Agronom. pres Argent. 2: 245-2 Krapovickas, A., P. Fryxell, and D. M. ie 1988. oe un nuevo género de Malvaceae de los an Bol. Soc. Bot. México 48: 23-34. Robyns, A. 1966. Flora of Panama. Family 115. see Ann. Missouri Bot. Gard. 52: 497-578. INDEX TO SCIENTIFIC NAMES Abutilon Allowissadula anderssonianum Garcke 182 lozanii (Rose) D. M. Bates 193 attenuatum B. L. Rob. & Seaton 183 pringlei (Rose) D. M. Bates 193 balansae (Hassl.) Hass]. 178 rosei (R. E. Fr.) D. M. Bates 193 barrancae M.E. Jones 188 Briquetia benense (Britton) Baker f. 178 spicata (H. B. K.) Fryxell 193 bridgesii Baker f. 192 Pseudabutilon cinereum Griseb. 180 benense (Britton) Fryxell 178 cymosum Triana & Planch. 180 callimorphum (Hochr.) R. E. Fr. var. demissum Fryxell 186 callimorphum 179 depauperatum (Hook. f.) Andersson ex B. L. callimorphum var. friesil (Hassl.) R. E. Fr. 189 dispermum (Hochr.) Fryxell 193 callimorphum var. intermedium Hassl. 179 ellipticum Schltdl. 182 cinereum ee ) Krapov. 180 grevilleanum (Gillies) Walp. 192 cowant Fryxell 1 hemselyanum Rose | 83 cymosum Lean : Planch.) Fryxell 181 membranaceum Baker f. ex Rose 183 depauperatum (Hook. f.) Kearney 182 mendocinum Phil. 192 ellipticum (Schltdl.) Fryxell 182 nigropunctulatum Ulbr. 186 glomeratum Fryxell 184 orientale Standl. & Steyerm. 186 hitchcockti Ulbr. 193 paranthemoides ee 192 inornatum Standl. & Steyerm. 188 saltense Hassl. 19 langlasset Hochr. 193 sidotdes Dalzell i a 183 leucothrix Fryxell 184 renee Hemsl.18 longepilosum R. E. Fr. 185 thurberi A. Gray | . lozanii (Rose) R. E. Fr. 193 oe (L.) Sweet 190 nigropunctulatum (Ulbr.) R. E. Fr. 186 virgatum (Cav.) Sweet 191 orientale (Standl. & Steyerm. Fryxell 186 Allosidastrum paniculatum (Rose) R. E. Fr. 187 hilarianum (C. Presl) Krapov., Fryxell & D. pintoi Monteiro M. Bates 193 pringlei Sey = E. Fr, 193 interruptum (DC.) Krapov., Fryxell & D. M. roseiR. E ale Bates 193 scabrum es Las R. E. Fr. 187 1997; FRYXELL: PPEUDABUTILON 195 Pseudabutilon smithii Hochr. 193 sonorae Wiggins 189 Ce be (H. B. K.) - E. Fr. 193 uckertii R. E. Fr. 18 ae (A. Gray) ei 189 umbellatum (L.) Fryxell 190 virgatum (Cav.) Fryxell 191 weberbaueri Ulbr. 193 Sida benensis Britton 178 callimorpha Hochr. 179 depauperata Hook. f. 182 dictyocarpa Griseb. ex. K. Schum. 193 dictyocarpa Morong & Britton 193 var. cordobensis Baker f. ex Morong & Britton 193 Tetra a dictyocarpa F. Muell. ex Benth. 193 elliptica (Schltdl.) Steud. 182 ee ie 192 umbellata virgata ae ey sida serrulata Fryxell & Fuertes 193 weberbaueri (Ulbr.) Fryxell & Fuertes 193 Wissadula balansae Hassl. 178 alansae Baker f. 178 callimorpha (Hochr.) Hassl. 179 callimorpha var. sae Hassl. 189 paniculata Ros pedunculata R. Bs 189 scabra C. Presl 187 ne er i: pie - 7 7 7 7 7 7 o 7 a =) . oO 7 - 7 _ 7 i 7 : oe a ee _ Bier _ So 7 Sn - : : : : ee - - a — a - - a= ; an a - : _ Oo a . : a - oe or a OS _ Oo 7 ~ 7 7 _ sa -_ - ; ee _ a — 7. _ SO 7 a 7 a 7 7 7 7 : _ - —_ 7 7 oe _ Oo 7 — : - 7 7 —_ Oo OS 7 a _ — a - _ 7 7 - 7 > 7 _ _ - 7 - Oo _ a _ - a oe — -_ 7 7 7 _ oO a — a 7 _ 7 7 a > oe >. Contr. Univ. Michigan Herb. 21: 197-204. 1997. NEW SPECIES AND NEW RECORDS FOR THE MALVACEAE OF BAJA CALIFORNIA SUR Paul A. Fryxell Department of Botany University of Texas Austin, Texas 78713 José Luis Leon de la Luz Miguel Dominguez L. Centro de Investigaciones Bioldgicas 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 localities 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. (unpublished 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 107 198 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Biosphere Reserve of the Sierra de La Laguna, Baja California Sur (see Fig. 1). Above the forest, a woodland community is found, where climate and flora are well delimited 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- eus 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, Leén 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!). big 2. Frutex caulibus rubellis sparse stellato-pubescentibus; laminis foliorum ovatis 5-8 cm longis; inflorescentiis spiciformibus; calycibus 8-13 mm longis ciliatis; 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. | 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 199 RACIFIC OCEAN Cabo San Lucas Tropical Deciduous Forest ees eee em 0 5O Km. ! FIG. 1. Collection sites cited. Site 1, La Burrera Canyon, is a typical locality for tropical decidu- ous forest. Malvastrum hillii and Sidastrum burrerense were collected here, as well as Sida alamosana (S. glabra), S. acuta, and Bastardia bivalvis. Site 2, Cerro Aguas Escondidas lies between the tropica deciduous forest and the sarcocaul scrub of the Sonoran Desert, and is the locality where Sida hyalina as found. 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 200 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 cc ke —— Malvastrum hillii FIG. 2. Malvastrum hillii. Left-hand branch and left hand mericarp (Dominguez 472); right-hand branch, fruit, and right-hand mericarp (Dominguez 77). 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 SPECIMENS EXAMINED. Mexico. Baja California Sur: “Los Gallos,” Rancho La Bur- rera NE de Todos Santos, Mpio. La Paz, 23°20'N, 109°59'W, 750 m, 2 Dec 1992, M. Dominguez L. 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 allied with this species or if it is of a different affinity. Sidastrum burrerense Fryxell, Leén 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. 1257 (holotype: TEX!; isotype: HCIB!). Fig. 3. Suffrutex caulibus atque foliis sparsissime atque minutissime puberulentis gla- brescentibus; laminis foliorum 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—4 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 202 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Ra Or, DO Sear Sidastrum burrerense t ~ FIG. 3. Sidastrum burrerense. Branch and flower (Dominguez 1257); fruit and mericarp (Dominguez 125) long, glabrous; calyx 44.5 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 purplish, 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. 1997 FRYXELL, LEON DE LA LUZ & DOMINGUEZ: MALVACEAE 203 ADDITIONAL SPECIMENS EXAMINED. Mexico. Baja California Sur: “La Testera,” Canon La Burrera, NE de Todos Santos, 23°30'N, 109°59'W, 620 m, 3 Jan 1991, M. Dominguez L. 125 (HCIB, TEX). 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 §. burrerense than in S. tehuacanum), the pubescence of the staminal column (hirsute in S. burrerense, glabrous in S. tehuacanum), the size of the calyx (44.5 mm in §. 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 1. Inflorescences racemiform; leaves narrowly lanceolate-elliptical, obscurely serrate, pubescent above; stems densely stellate-pubescent; corolla yellow; mericarps 5 S. strictum (Standley) Fryxell. 1. Inflorescences paniculate, often with capillary pedicels; leaves narrowly lanceolate-elliptical to ovate, glabrous or pubescent above; stems densely to sparsely pubescent or glabrous corolla yellow, lavender, or purplish; mericarps 5-8. 2. Leaves narrowly lanceolate-elliptical, obscurely serrate, glabrous above; corolla yellow; mericarps 5. S. lodiegense (E. G. Baker) Fryxell. 2. Leaves lanceolate to ovate, serrate, pubescent or glabrous above; corolla lavender, pur- plish or yellowish; mericarps 5-8. Corolla purplish, reflexed; stems densely stellate-pubescent; leaves serrate, stellate- pubescent above. S. paniculatum (L.) Fryxell. 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. Les) os) S. burrerense Fryxell, Leon de la Luz & Dominguez. 4. Corolla yellowish (sometimes fading pale lavender), rotate; staminal column gla- brous; calyx 2-4 mm long; mericarps reticulate laterally, smooth dorsally S. tehuacanum (Brandegee) Fryxell. 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. Sida alamosana §. Watson (or S. glabra Miller?). Mpio. de La Paz, ca. Rancho La Burrera, 23°29'N, 110°01'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, Ledn 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 himedo, 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 (Figs. 2, 3). LITERATURE CITED Axelrod, D. I. 1979. Age and origin of Sonoran Desert vegetation. Occas. Papers Cal. Acad. Sci. 132: 1-74. Brandegee, T. S. 1892. Distribution of the flora of the Cape Region of Baja California. Zoe 3: 223- 2315 Fryxell, P. A. 1978. Neotropical acon: from Sida L. (Malvaceae). Brittonia 30: 447-462. 1988. Malvaceae of Mexico. Syst. Bot. Monogr. 25: 1-588. [Sidastrum on pp. 416-421] Hill, S. R 1982. A monograph ee 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) [figo Granzow-de la Cerda University of Michigan Herbarium North University Building Ann Arbor, MI 48109-1057 INTRODUCTION The genus Anomodon was segregated from Neckera Hedw. by Hooker and Taylor (1818). The two species in 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 Giimbel (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 Heterocladium 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 (Mill. 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 M6nkemeyer (1927), who recognizes also the Theliaceae (close to the Fabroniaceae), Crum and Ander- son (1981), and also Ireland (1982), although he merged the Theliaceae 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 Theliaceae 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). 205 206 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Anomodon and its closely related genera do not fit clearly 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 ts 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 paraphyllia 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 Theliaceae (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, Herpetineuron, 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 Prerogonium 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 (Thér.) 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 Prerogonium gracile, following Akiyama (1988) and Anderson et al. (1990), who include Prerogonium in Anomodontaceae. The taxa included in the analysis are all eleven species of Anomodon: |A. abbreviatus Mitt., A. attenuatus (Hedw.) Hiib., A. dentatus Gao, A. giraldii Mill. Hal., A. longifolius (Brid.) Hartm., A. minor (Hedw.) Lindb., A. rostratus (Hedw.) Schimp., A. rugelii (Mill. Hal.) Keissl., A. solovjovii Lazarenko, A. thraustus Mill. Hal., A. viticulosus (Hedw.) Hook. & Taylor] and five species of Haplohymenium, considered as part of the ingroup [H. longinerve (Broth.) Broth., H. pseudotriste (Mill. 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), Sulli- 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; Kallersj6 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. 208 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Fig. 1. Fides NM - A \ (yh mI AM Aa ne FIGS. 1-7. Schematic representation of states in some characters, as coded for the cladistic analysis. Fig. 1. Character 1, branching pattern. Fig. 2. Character 9, shape of the distal region of the lamina of branch leaves, above the shoulders. Fig. 3. Character 11, shape of the branch leaf apex. Fig. 4. Character 13, length of the costa. Fig. 5. Character 14, bifurcation of the costa near the apex. Fig. 6. Character 16, papillae of dorsal surface of the costa. Fig. 7. Character 21, position of gametangial buds relative to most recent (distal) branching points. RESULTS Three most parsimonious trees, 325 steps long, with consistency index of 0.44, were obtained when scaling was applied (analysis |). Their consensus tree is given in Fig. 8a. When equal unit weight was applied, without scaling (analysis 2), four trees were generated, their length 121 steps and consistency index 0.45. The strict 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 209 consensus tree 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 Prerogo- 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, 1.e., a monophyletic clade, disappears; Bremer 1988, 1994; Donoghue et al. 1992; Kallersj6 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. lwatsuki (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, 210 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Figure 8. . Viticulosus . rugelii . minor . abbreviatus A A A A A. solovjovii A. dentatus H H H H H A . tristis . sleboldii . pseudotristis . flagelliformis . longinervis . thraustus A. rostratus A. attenuatus eae A. giraldii is coe A. longifolius = Herp. acutifolium es Herp. toccoae P. gracile . viticulosus . rugelii . minor . abbreviatus . Solovjoviil . dentatus . tristis . sleboldii . pseudotristis . flagelliformis . longinervis . thraustus A A A A A A eee en H H H H A A . rostratus A. attenuatus ee ae A. giraldii A. longifolius —- Herp. acutifolium ees Herp. toccoae P. gracile IG. 8. Strict consensus trees: a, consensus tree for analysis | (scaling at 4) of the three most parsimonious trees (325 steps): b, consensus tree for analysis 2 (no scaling) of the four most parsimo- nious trees (121 steps). Figure 9. ANOMODON HERPETINEURON = ! 1 [ | Ne) Subgenus Anomodon Subgen. Pseudoanomodon ~~ ; ; ; sect. | ! l section Anomodon section Haplohymenium Thrausti [ps subsect. subsect.| | 1 I ] subsect. Anomodon Abbreviati Dentati = ta = rc r rm See eee i wer so os faa Sy (ep) t ' 1 ~ S Z fe) fi ia Qo Ww = 6 ¢ © iE oc ke tee a oe) > oc = <= n Qa = o no re a -E o < (G) af ale ate o Q v2) > Z 7. 1->0 S m= 16° 0->1 if ee mig: 453 is wt22, 122 7 aca 5 eae a6: 0->2 = ett: 253 > polis 41 12 0-1 @ 22: 0-71 mw ttt 051 m wow? 0-1 v0) 18> 0->1 2 Bl © on>1 we 21) 0->1 ae 4 0->1 oe Lf 1->2 (9: 0/1->1) 430: 152 Toe > 4 - Z 16 ‘e) = 1->0 © L414: 2->3 7) wid: 1->2 g 18° 0->4 = 21: 0->4 a 24. 0->1 @ ; met 1>2 m wa 27. 1->0 Ly 2 ds: 0-4 m 10: O->1 2 aes oo osipa ee 12. 1->0 13: 1->2 wt: 0-1 26: 1->0 w+19: 0->4 30: 1->2 G. 9. One of the most parsimonious trees in analysis 2, which assigned equal weights (no scaling). Only unequivocal changes for this particular tree are represented (in addition to character 9). All of them, except for characters 31 and 10 (boxed), are homoplasies. Transformations with a black square show characters with consistency index of 0.5 or higher for this tree. Proposed taxonomic arrangement is also given above the species names. — — — 212 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Figure 10. 7 A. viticulosus ; A. rugelii 4 trees A. minor length = 121 steps . pse A. flagelliformis A. thraustus A. rostratus A. longifolius | ee rp Herp. acutifolium — i eee Herp. toccoae P. gracile A. viticulosus A . minor 114 trees A. abbreviatus length = 122 steps A. solovjovii A. viticulosus c. A. rugelii A. minor A. abbreviatus 1065 trees length = 123 A. solovjovii FIG. 10. Bremer support analysis of the four most parsimonious trees resulting of analysis 2 (no scaling). Strict consensus trees become progressively more unresolved as parsimony is relaxed. a, boxed numbers denote how many steps parsimony had to be relaxed to cause that clade to collapse (become unresolved) as an indication of the decay index: clades marked 3 are the most supported (robust), and those marked | the least supported; b, tree resulting from parsimony relaxed one step; c, tree resulting from parsimony relaxed two steps. Length and number of trees are given. 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 213 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. rugelii, 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. /ongifolius, 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 (1.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, 1 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 | 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. dentatus, 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 of 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 localities 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 within 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-DE LA CERDA: 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 acutifolium 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. CONCLUSIONS All the species of Anomodon (s.1.) 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 (Miill. Hal.) Kindb., and A. tristis (Ces.) Sull. & 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. rugelit, 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, Musc. Brit. 79, pl. 3. 1818.—Type: 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. Paraphyllia absent. Stem leaves minute and scalelike, 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. /ongifolius); 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) uum in diameter. 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. tristis are extremely rare in North America (only one fertile speci- men of A. viticulosus seen, none of A. tristis). In Europe, however, A. viticulosus 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. rugelii, may grow on the same tree, each usually 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. Although a high rate of sporophyte production is observed in some species, no hybrid sporophytes have been recorded. KEY TO THE SPECIES OF ANOMODON I. Plants very slender, delicate, stems <0.5 mm thick when dry, leaves <1.2 mm long, differenti- ated basal cells few, not reaching the margins; calyptra hirsute. 2. Laminal cells with numerous short, mostly branched papillae, costa short, not extending a the proximal 1/3 of the lamina. . Leaves complanate when moist, — of leaves on non-flagelliform branches obtuse to rounded, papillae inconspicuous. 9. A. pseudotrists. . Leaves not complanate oe co apex of leaves acute to acuminate, occasionally obtuse, papillae conspicuou 4. Plants extremely fete ane slender, stems ca. 0.2 mm wide, some becoming ee liform, <0.1 mm wide at the apex; leaves squarrose, leaf apex tapering 7 a lon awn (plants rare; Manchuria). 10. A. flage liformis . Plants + delicate, stems ca. 0.4 mm wide throughout; leaves erect to slightly spread- ing, leaf apex acute to acuminate, occasionally obtuse 5. Leaves invariably broken off for more than 1/2 their length, with an obvious constriction, apex acute to acuminate, occasionally obtuse; costa ending near the leaf constriction; laminal cells with strong, high, branching papillae to mammil- lose, margin crenulate near the apex owing to protruding papillae. 7. A. tristis. ee) a 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 217 5. Leaves not broken off, with a faint constriction, apex acute but not acuminate; costa reaching well beyond the leaf constriction; laminal a with delicate, nched but not high papillae, never mammillose, margin smoo 8. A. sieboldii. Pye esha cells with 1-2 (-3) rather high mammilla-like, acute, eee: papillae, costa ending beyond the proximal 2/3 of the lamina. 11 sida 1. oe somewhat slender to robust, >0.8 w dry, leaves usually >1.4 mm long Ga A. thrat tus as small as 1.2 mm long), ee basal cells almost reaching the margins ee extending to + 1/3 of the lamina; calyptra smooth Cells of the lamina with a age long conical papilla, never branched, but sometimes with 0-3 short “spurs” at the base. 7. One single papilla on ae 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, long-acuminate, somewhat subulate, with cae ee at least at the transition to 16. A. the narrower distal portion; pseudoparaphyllia pre longifolius. 7. Papillae with 0-3 basal spurs, sharply conical, ee foward 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; pseudoparaphyllia 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. abbreviatus. 8. Leaves <2.0 mm long, the distal portion (beyond constriction) lingulate, flat and with rounded apex; seta ca. 1 mm long, oa enclosed within the oi ia leaves, which are longer than vegetative leaves. 5. A. solovjovil. 6. Cells of the lamina with multiple, often more or ne foeeeed papillae. 9. Plants cages profusely and irregularly branched, branches julaceous, aa leaves 8 mm long, appressed, the apex extended into a long awn. 3. A. rostratus. 9. Plants more or less dark green to rusty brown, with secondary branches ae 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 10. Branch leaves lanceolate, acute at apex, with unbranched ee cell walls con- sly sinuose throughout the lamina; margins revolut 15. A. giraldii. 10. en leaves lingulate (or lanceolate in A. attenuatus), ries or rounded at apex, with branched papillae that obscure the cells; cell walls straight but sometimes sinuose in the basal hyaline cells; margins plane. 11. Plants dark green to rusty-brown; leaves strongly incurved-contorted (ca. | full rn) when dry, auriculate at the base, stem and costa dark brown to reddish; eee ee ent. 2. A. rugelii. 1 ie dull-green to See leaves only slightly incurved to falcate when dry, more or less decurrent, not eee stem light brown, costa yellowish to light green; pseudoparaphyllia ab 12.Leaves narrowly lin ie i constriction, this distal portion broken off, distal portion of the costa obscured by laminal cells, never ch fid. 13.Secondary branching irregularly pinnate; leaves >2 mm long; costa vee current; leaf apex (when leaves are intact) coarsely serrate owing to prom inent whole cells. 6. A. dentatus. 13.Secondary branches few; leaves <1.7 mm long; costa reaching 2/3-3/4 of the lamina; leaf apex (when leaves are intact) entire. 12. A. thraustus. 12. Leaves constricted, but the distal portion not much narrower than the proxi- mal one, intact; costa generally not obscured by laminal cells, sometimes shortly bifid at the en 14. Plants robust, branches ca. 1.3 mm wide or more when dry; leaves flexu- ose, falcate-secund, spreading to reflexed when moist, not Se when 2mm lon 1. A. viticulosus. 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; irene present on terminal branches, beyond the most distal branching eae leaf constriction well defined, the distal pornen lingulate; secondary nches not attenuate; back of costa ee lar seriate papillae on as Si 3. A. minor. 218 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 15. Leaf apex acute or occasionally obtuse, and almost always apiculate, often dentate; secondary branching profuse, irregularly pinnate: inflorescences never present on terminal branches beyond the most distal branching points: leaf constriction poorly defined, the distal portion tapering to triangular: apex of secondary branches attenuate; back of costa not papillose. 14. A. attenuatus. I. Anomodon subgenus Anomodon. Stems and primary branches branching in a simple pattern. Leaf apex rounded to broadly obtuse (acute to acuminate in section Haplohymenium); laminal cells isodiametric, hexagonal, quadrate or round in all taxa. Ia. 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. dentatus), 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. l.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. Musc. 209. 1801. Hypnum viticulosum (Hedw.) L. ex With., Syst. Arr. Brit. Pl. 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, sér. 3, 41: 84. 1894.—T ype: VieTNAM. Tonkin, Hanoi, Mt. Chua, 12-3-1888, Bon 3789 (isotype: NY!). Anomodon subintegerrimus Broth. & Paris, Rev. Bryol. 27: 77, 1900.—Tyee: VietNam. “Tonkin, inter Thai Van et Pho Lu”, 18-3-1901, Moutier (iso- types: G! JE! NICH! NY!). Anomodon decurrens Cardot, Bull. Soc. Bot. Genéve, sér. 2, 3: 281. 1911. SYNTYPES: JAPAN. Mitoyoshi, J. Cardot 1190; Korea. Seoul, Cardot 313 (Po); Plants robust, forming thick mats; branches 6-8 cm long, occasionally more, and I-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-lingulate, 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 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE PANS) = = 0.1mm FIG. 11. Anomodon viticulosus (Hedw.) Hook. & Taylor. a, b, branch leaves; c, d, branch leaf insertions (showing decurrency); e, portion of peristome. Based on: a, c, e, G.-de la Cerda 2546, Girona, Vidra, Rio Gés, 18-1-1988 (BCB): b, d, Thomson 1112, India, Simla, Oct 1849 (NY). 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. Perichactial 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 obliquely 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) pm, in others 15-16 um, densely papillose. Chromosome number: n = 10 + x/y (in meiosis, Spain: Granzow-de la Cerda, 1988b). 220 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Distribution (Figs. 12-16). Anomedon 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, Québec, 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. | 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. ee oe NY). China. Hsien: Lichuan Xian, Lojiaba valley, Exp. Sino-Amer., 6-10-1980 (FH). Hubei: Lichuan Xian, ee det 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, in Chang Qing 2953, 4- 1964 (IFSBH). Szechuan: Chungking, Siao lung kan, Chen 4-1940 (as A. thraustus, JE). ig seen ae Menking to I-wu, a ay a et al. 33960, 24-12-1986 (NY). Yunnan: He fanaa r. Longtan, P. Redfearn, 7 es & YIG 53, 27-7-1984 (NY). Georgia. Swanetia Dadianosum, Sommier Py Taian. 29-7- nr ae cel Pradesh: re Thom- son 1112, 1116, 10/49 (as A. obtusifolius Mitt., n. eer ae c. “type” in herb. Mitten NY); Kangra, Kulu, rae 7510, 24/27-11-1933 (c. fr.. DUKE, MICH). se Bombay, enh Marchal, S-1869 (c. fr., FI). Punjab: Murree, Stewart, 9-1948 (NY). Uttar Pradesh: Tihri Garwal, Kidarkanta, ee is 6-1904 (as A. negra c. fr., NY); Musoorie, Dhanoutti, Bahadru, 10-12-1903 (as . integerrimus, NY). URAN. Mazanderan, 30 km S of Amal, P. Goldblatt ace May 1973 (FH, io Japan (as A. decurrens). Hondo: Akiyoshi, Nagato, S$. Matsumoto, Mar 1 2 (NICH). Hon- shu: Jaisaku pref., Bingo, A. Noguchi, 7-7-1934 (NICH); Okayama pref., eed ri Noguchi 3-11— 1954 (NICH): Shiga pref., Sakata-gun, Kaminyd, 7. Nakajima, 4-4-1978 (MICH). Kyushu: Fukuoka pref., Hisaodai, 7. Osada, 2-5-1955 (MICH), Kaharu Mt., A. Noguchi, 13-9-1953 (NY), Kawaradake, A. Noguchi, Sep 1953 (MICH); Kumamoto pref., Kuma-gun, Koonose, Takaki 2302, 11-8-1946 (NY), K. Mayebara, Dec. 1947 (MICH), Itsuki-mura, Sangaura, A. Noguchi & Imae, 23-4-1956 (NICH); nobori-mura, A. Noguchi, 23-11-1953 (NICH); “center of back bone, Mt. Kumimi,” Kuwahara 2180, 2 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., 7. Elias et al. B-65178, 65180, 17-8-1978 (NY). Taiwan. Tienhsiang, Tailuko rvr., Z. Iwatsuki & A. ples 1787, 27-3-1965 (FH). Thailand. Phu khieo, E of Chaiyapun, Lorsen, Nielsen & Santisuk 31305, 8- i (NY). Turkey. oe as Magka, Pontus Mts., Godfrey 728, 24-9-1952 (NY). anada. Nova Scotia: Cumberland Co., Wentworth, R. /reland 10673, 7-9-1967 (as A. minor, i Ontario: Bruce eo 2 km N of Colpoys Bay, R. Ireland 20290, 12-7-1982 (MICH), ao Bluff, R. Ireland 20396, 14-7-1982 (FH); Grey Co., Inglis Falls, H. Crum 11008, 3-8-1961 (DUK MICH), Jones Falls, R. /reland 20198, 11-7-1982 (FH); Lincoln Co., betw. Vineland and Sean HM. Crum 31, 42, 5/7-10-1964 (MICH); Ottawa, Bauer, 31-3-1902 (FH). Québec: Gatineau Co., King Mt, Gatineau Park, R. Ireland 15853, 27-4-1972 (MICH); Hull, J. Macoun, 22-4-1896, 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 Destine U.S.A. Arkansas: Newton Co., junc. Leatherhood Creek and Buffalo Rvr., P. Redfea Feb 2-2-1972 (MICH). Michigan: Alpena Co., SW of Posen, Leer Road, Diill 1152/3. 18-8- poe (M H): Delta Co., Burnt Bluff, H. Gleason 2390, 22-8- iy (MICH), W. Steere 760, Aug. 1934 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 201 (FH); Keweenaw Co., gorge of Manganese Rvr., W. Steere, 1-9-1936 (MICH); Mackinac Isl., E. & S. Harper, 18-7-1899 (MICH), /. Schnooberger 2749, 21-7-1939 he 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 & ene MICH); Clinton Co., Aus- able Chasm, L. Anderson 20608, 5-9-1968 (DUKE, MICH). Vermont: Glebe Mtn., Newf Grout, ae 1 900 (as A. viticulosus var. microphyllus, FH). Virginia: Marion, A. Vail & E. Britton, 13 5- 1892 (MICH); Giles Co., E. Byers 16, 4-7-1949 (MICH), Newport, J. G.-de la Cerda 1892, 1898, 7-10- 1892 OEE Tonnes. Anderson Co., Savage ani A, Sharp, 13-1-1957 (MICH, TENN), /. G.-de la Cerda 1906 & A. Sharp, 8-10-1984 (DUKE Austria. Salzburg, pr. Seas J ee 6-5-1868 (c. fr., FH); “Traunfall, ce Schwan- enstadt,” Matouschek (c. fr., FH, FI); Bergwalder, Klaus, Jul 1910 (‘“var. latifolius,” c. fr., NY). Tirol: Windish Matrai, Klamm, F. a 7217, Aug 1911 (MICH); Innsbruck Sillschlucht, Luisier 22-3- 1904 (c. fr. INA). Belgium. Auderg hem, Piré (c. fr., FI); betw. Rochehaut & Ucimont, /.L. De Sloover 16439, 1-7-1973 onan Louvaine, pr. Pournay, Marissal, (c. fr., FH); Orval, J. Cardot (FH). Croatia. Istra, above Ici , pr. Opatija, ee Frahm 10034, 26-3- 1975 (c. fr., MICH). Czech Republic. Bohemia: “Karlste se 244-1890 (c. fr., FH). “Méhr-Weisskirchen,” Surkow, F. Petrak, 9-4-1913 (FH). Dea ae Mariager dd, K. Holmen & E. Warncke 237 (MICH); M6en, Klinteskoven, H. Mdller, 1/3-6-1893 (c. fr., FH). Sjoeland: Farum, N¢grreskov, G. Smith, 24— 10-1967 (FH); Soré, Suserup, F. Hermann 1588S. 28-5— 1960 (MICH). Finland. Tohmajirvi, Kb: Havukkavaara, S of Vuorenpaa, R. Fagerstén & M. Haapassari 4097 dup., 2-10- 1984 (FH, MICH); Riikola, R. Fagerstén & M. Hapasaari 4120, 3-10-1984 (FH). “Karelia eee Juuka, Polvijarvi, A. Huuskonen, 19-8-1965 (FH). France. Ariége: Bagnéres de Luchon, C. Casas, 20-9-1956 (BCB). asses-Pyrenés: St. Etienne-de-Baigorry, M. Fleischer, 10-5-1908 (FED. Calvados: Pont des Vers, Husnot, (c. fr., FH). Gard: Meance Le-Clap, J. L. De Sloover 44850, 12—7-1988 aay ae, Savoie: Saléve, Archenes, R. Blanche, 15-2-1954 (c. fr., MICH). Isére: Bains-la-Romaine, J. L. Sloover 34593 10-7—-1981 (MICH). Savoie: Saint Claude, “prés de Chambéri,” pier oie: ee iG fr., FI). Ger pe Giessen, Wanschoff, (c. fr., FH); Miinchen, Dodel-Port., 20-2-1869 (c. fr., FH); “Rhinefall,” n& es 4-3-1887 (c. fee Ed): nie Ioannina: Epirus, pr. Emin Agha, K. en ee 5-1961 (MA). Thessalia: Lutropigi, Smokovo, K. Rechinger, 8-5-1961 8g (MA). Ireland. Sligo: een H. Conrad, 4-8-1935 (FH). rae (Gaaeie. as Varesse, E. Corti, Mar. 1905 (c. fr., FH). Lavione: “pr. Florentiam,” J. Arcangeli, 23-12-1877, (c. fr., FH). Lom bardia: Parco di Monz eee 19-9- 1878 (c. fr., FI); Piemonte: Campello- Monti Boner: Levier, 30- 7 1906 (FI): C mo, Torno, Lago di Como, Artaria, 2-1902 (FH). San Marino: Borgo, Ornella, Pampanini, Ree (c. fr., FI). Trento: Lago Tadel, Serra del Lago, Cortini 973, 16-7-1958 (FI). Venezia: Padova, Fiori, 234-1893 (c. fr., FI). Macedonian Republic. W of Tetoro, Popova Sapka., exp. Rheno-Trai 68—/526, 10-7-1968 (MEXU). Norway. Sédra Trondheim, Hakan, J. Hagen, Jul 1899 (MICH); Lysaker, pr. Oslo, 7. Hagen 18-10 (FH). Poland. Krosno: Kurnikoéw 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 a (Krakov): Zabiezowic, Z. Zmuda, 22-4-1911 (FH). Russia. Mos- “Wald von Kunzowo,” Heyden, 1-8-1900 (FH). Slovakia. Gau Kaschau, Snina, Vinicna hora, Baler 31-7-1925 (FH); hen ete Tscheckengrund valley, /. Gyérffy 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: eee La Cénia, barranc de la Fou, C. Sérgio et al. 4.1, 14-2-1984. Girona: Sta. Maria de Besora, Serra de Caraups, J. G.-de la Cerda 2554 18-1-1988 (BCB). Vidra, rio Ges, rec de l’Espirau, /. G.-de la Cerda 2316, 2322 & M. Sanz, 10-1-1987, I. G.-de la Cerda 2546, 18- 1—1988 (c. fr., BCB). Leén: Posada de Valdeon, Garganta del rio Cares, .G.-de la Cerda, 1-1-1990 (BCB). ee Pallars Sobira, “entre Rialp i Llesui,” C. Casas 6-10-1981 (BCB). Lugo: Lézara, 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- a (MICH). Oland: Borgholm, £. Ekstrand, 8-7-1975 (c. fr., MICH), H. Méller, 16-4-1908 (FH). Skane: Kargeréd, YImesaken, C. Stenholm, 25-5-1934 (FH). Uppsala: eee S of Fabo- darna, F. Osterlind, 3-5-1951 (FH, MICH). Switzerland. “Switzerland” ? (DUKE). Mt. Saléve, M. MacFarlane, Feb 1911 (c. fr., FH). Spliigen, po eyees Jun 1861 (c. fr., ae one Chatelaine, Boissier, Mar.1821, (c. fr., MICH). Ztirich, P. Culmann, 11-3-1880 (c. fr., FH). United Kingdom. Scotland: W. Wilson, 320 9-7-1861 (c. fr., FH a nglesey [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, FE. 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, Morl Brook, /. G.-de la Cerda 2018, Jul 1884 (DUKE). Zee CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 | O A. attenuatus | @ A. viticulosus L_ Ses FIG. 12. Distribution of Anomodon attenuatus and A. viticulosus in the New World. 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. rugelii, or A. minor. In depauperate populations of A. viticulosus, this species can be distinguished from A. rugelii by its decurrent 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. thraustus. The branches are not complanate, as in A. rugelii or A. minor. The atter 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 1997 Dim. == —S ae r { | i Sar ——— J ia eee | | y L we, ee | . _) << \ / / | / / / | = / )2Q / | \ ; : / / | / \ ) \ p> | \ / f i LM / / | | | y / i / | < 4 / \ >A / / YS [> af ‘ 3 © A. attenuatus }__/ | | ( ; Oo 1 | @ A. viticulosus| / ~~~ ) | / es), O° Se | e | FIG, 13. Distribution of Anomodon attenuatus and A. viticulosus in Europe. central Mexico). Anomodon viticulosus is sometimes mistaken for A. attenuatus, 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 of 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 taxonomic consideration. 224 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 A, = 4 a” Nc 1 l rly y o i “ te \ ‘ 4 i - i> ae Pa fv J lard ¢ ) \ ane \ ( rie ian : : \ \ Bo = Ore \ mt i ° Oo —~_—— OA longifolius f ' | LS + 2 @ A viticulosus « a ? cae vy J J @ L ee 2 Oo lo | | | | _o @ ) | " ( @ A. viticulosus id nd e f @ A. rugelii | Ne | | _ *on P roe o Oo ° ’ g if + o & on FIG. 15. Distribution of Anomodon attenuatus, A. longifolius, A. rostratus, A. rugeliit, and A. viticulosus in eastern Europe and central Asia. 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 225) FIG. 16. Distribution of Anomodon rugelii and A. viticulosus in central and eastern Asia. Anomodon viticulosus is the type species forAnomodon 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 Mill. Hal., Syn. 2: 472. 1851.—Type: U.S.A. Tennessee, Smoky Mountains, Rugel s.n. (holotype: not located). Figs 1 7a.b. Anoectangium ferrugineum Besch., Ann. Sci. Nat. Bot. 17: 329. 1893. Anomo- don ferrugineus (Besch.) Nog., J. Jap. Bot. 20: 260. 1944. Anomodon rug- elli 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. Genéve, sér. 2, 3: 281 1911. Anomodon uematsui Broth., Ofver. Finsk. Vet. Soc. Férh. 62: 33. 1919- 1920.—T ype: Japan. Hokkaido, Iwate pref., Rikuchu Nat. Park, Mt. Hay- achine, 27—7-1906, Uematsu 3390 (isotype?: NY! 226 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 0.1 mm FIG. 17. Anomodon rugelii (Mull. Hal.) Keissl. and A. attenuatus. (Hedw.) Hiib. a, b, A. rugelit. a, branch leaves; b, stylized representation of a branch leaf. c-e, A. attenuatus. c, branch leaf; d, leaf apex; e, leaf insertion. Based on: a, b, G-de la Cerda 2512 & es USA, North Carolina, Joyce Kilmer Mem. Park, 3-10-1987 (DUKE, MA); c-e, Standley, Guatemala, Alta Verapaz, Rio Carché, 12-4-1941 (NY 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 foliose, often narrow to uniseriate, golden-yellow pseudoparaphyllia, generally with abundant rhizoids. Branch leaves incurved-contorted (ca. | full turn) when dry, erect-spreading, some- what secund when moist, (1.2—) 1.4-2.3 (-2.5) mm long, broadly oblong-lingulate 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 um 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 um long, with multiple high and branched papillae; 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. Pert- 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; exostome 0.15-0.30 mm high, teeth regular, papillose 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE Day. 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 «tm 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 rugelii 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. = a eee = =e E | | yo | aN eee | A . | Zw ; e / ET ja \ ] / / | | mes Wi / | a j \ \ me / / \ ae fi +> x / | a4 ee: We | ¢ YS ) | | | / e | { L me | |\ @ | [ ) ea By i \ ( = | 4) | ( 4 @ a eee | A | aT, #Y | 8s a es = \ | ; \ w® ae | Q / ) | i] \ f | | —— Se cane, ‘ Ee# |) / 7 _ mn / TS \ o : Z aS e < \ - A we e = < @ ey i e ae ‘oe } : | ‘ _ / — Te | o | : 2| Saas y i \ | if Bs A ay oO} Fa AL m7: t ) ' / | ney e A \Y fh J = = \ w°/ ad 7 | Oren ~Oy 3 \ r— ge x ) e pe He Ue 2 / y = ~ age | / / eo — (Oo) id ae : / ac ean / / ( Were ans! < \ Die NN } / \ \ Be = } oO -- % . : ( j 2 o | fo | / sf — / \ | ~ | } [ if / I { Sal H ee 4 / saat [ ( ‘ / \ = \ an / * | : | me _ | ws =a | r ia f ? | 1 | [one _ ees 2 | =o: FIG. 18. Distribution of Anomodon longifolius, A. rostratus, and A. rugelii in Europe. 228 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 ) \ 3 ) / of ee eae iy od ES | = A. rugeli SGPT aad a, ae | ; cmt vm si py ; : 4S eae | eB a | ry a Se M@ A.thraustus | ete et | — —* a ee ~ Po yi ae A 7 eal / _ ~% FIG. 19. Distribution of Anomodon minor, A. rugelii, and A. thraustus in the New World REPRESENTATIVE SPECIMENS. Azerbaijan. Bakuriani, Y). China. Hubei: Shennongjia eee Distr., 655c, 3-9-1980 (NY). Jilin: Chang Bai 1 (NY). Liaoning: ae monte Chien,” Chen Gao 1889, 19-8— (as A. ferrugineus, asia Heilo aes distr. Hailin Dahai, paomdings shan, sd Qian 9147, 28-8-1973 (IFSBH). India. “Tonglo top 10000” 7 A, planatus, “syntype herb. Mitten NY). Uttar Pradesh: Tehri Garwal, ae ee May 1904 (NY : Gollan, A. 5-1904 (c. fr., FH, NY). “India orientalis”: Bell 367 (as A. minor, Mitten NY). Japan. Hokkaido: Hoa Mt. ee wa, N. lw , 17- ic TOUS (62 1r IN). Son Nantaizan, Bisset (as A. ramulosus, syntype, NY); A 1 pref. ee be Seal 1-9-1954 (as A. ees NICH); Eti Sekida- is (Sek) ras ee pe 1-8 —1935 (as A. uwematsul, NY); Kyo . Kita- awed. gun, Sasari-to T. Nakazima 5553, 2-5-1954 (as A. aa NICH); 7. a Hirakura forest, 7. horene: 11940. 6-5-1961 (as A. ferrugineus, NICH); Nagano ref.. Aumo-mura, Mizushimako, 28-9 4 (c. fr., NY), Kami-ina-gun, ae aimee ah 5-8- 1933 (c. fr., NY), Nakajima, 22-8-1973 Ren — ek. Yoshimo-gun, Nosegawa-mura, 7. Nakazi- ma 8708, 3-5-1958 (as A. ferrugineus, NICH), M . Odaigahara, 7. Nakazima 1907, 16-10-1948 (as A. ferrugineus, NICH); Okayama pref., Maniwa-gun, Mt. Kani-hirsen, C. /gi 58/2, 12-11-1957 (c. fr., as A. Lazarenko & Abramov, “from Quiujiajing oe . Guo Qian 22137, 24-9-19 17-9-1956 > Sino-Amer. Exp. a9 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 229 A. ferrugineus, NICH); Saitama pref., Chichibu, ee dani, 1. 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. [watsuki, 13 - 1955 (NY); Kurabe, Keyakidaira, A. Nogu- chi, 13-10-1953 (as A. ferrugineus, NICH); Komagataka, Faurie 3493, 1905 (c. fr., NY). ee Oita pref., Mts. Kuju, Kurodake, M. Miyamoto 2431, Jul 1960 (c. fr., as A. ferrugineus, ieee Nepal. E. Himalaya: Kathmandu, Mt. Phulchoki, 2. /watsuki 2470, 5 7-1972 (NY). Russia. Reg. austro- ussuriensis, “fontem flumen Ssputinka,” A. Lazarenko, Oct 1934 (NY). anada. New Brunswick: Grand Falls, H. Habeeb ie 1-11-1945 (NY), Aroostook Falls, J. Colne 2770, 17-7-1902 ee Victoria Co., Kilburn, R. Ireland 12758, 9-7-1970 (MICH). Nova Scotia: Digbi Co., Acaciaville, R. Ireland 17890, 28-7-1974 (MICH, NY). Ontario: Carleton Co., Fallowfield, #. on 290, 28-10-1964 (MICH). cages - Muskoka Distr, NW of Chaffey Twp., R. ace 22021, 23-7-1986 Cae Algonquin Park, J. Macoun, 8-6-1900 (MICH). Manintulin , Sheguindah, Howland Twp., R. freland an 30-7- ee (NY). Muskoka Distr. on rv TERE Twp. (Foots Bay), R. es 24032, 7-9-1982 (NY). Québec: Terrebonne Cox E of Nie 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-S, ae (NY). Pontiac Co., Park de la Vérendrye, Lac Serpent, R. Ireland 15969 & T. Koponen, 13-6-1972 (NY). U.S.A. Maine: Pleasant Ridge, Bean Pond Ridge, J. Coury 2455, 21-7-1901 (as A. eee MICH). Michigan: Keweenaw Co., “on ledge at Delaware,” W. Steere, Sep 1936 (as A. viticulosus, oe ae York: Cayuga Lake Basin, Dudley, 16-5-1884 i . fr., mie ); Lake Placid, Britton, 9-8-1898 (DUKE); Adirondack, Essex Co., North Elba, Hermann 14776, 10-8-1958 (DUKE); Catskills, ne Lawrence Co., Cranberry Lake, N. Slack ae 25-6-1976 (DUKE). North Carolina: Roan Mountain, Weatherby, 24-2-1894 (as A. minor, c. fr., DUKE); Graham Co., Joyce Kilmer Mem. Park, J. G.-de la Cerda & K. ee 3- 10-1987 (c. fr., as BCB); ee n Co., Chatooga river, /. G.-de la Cerda 1454, 1-7-1984 (DUKE); McDowell Co., Linville Falls, 7. 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., ee meadows, /. G.- de la Cerda 1925, 20-10-1984 (DUKE). Pennsylvania: McKean Co., Ben rook, Bur on 7- 11-1897 (FH). Vermont: Willoughby, Humphry Brook, Kennedy, 17-10-1897 a obtusifolius, FH). Germany. Bayern: eee “Sauwald ober eae Holler, 22-8-1892 (COI), Holler, 29-8- 1891 (NY); “Monte Rhén, oe 1870 (c. fr., NY), “Rhéngebirge,” G. Geheeb, 1869 (NY), Geisa Wald, G. Geheeb, 1873 i , NY); H cheleuene bei Geisa, in Vorder-Rh6n, G. Geheeb, 16- 11-1868 (c. fr., NY). Hessen: ee Vogelsberg, Rdth, 9-8-1903 (herb. Bartram FH, NY), Roth, 10-6-1885 (ex herb. Uggla NY). Poland. Silesia: Gralen bei Gdbersdorf (Kreis Waldenburg), /. & H. Milde, 8- ie (c. fr., NY). Norway. Gaset i Asker, Kiaer, 7-6-1878 (ex herb. Uggla NY). Aker- shus: Asker distr., Skau suas Fridts, 10-9-1886 (c. fr., BCB); Grossaetdal, Asker, pr. Christiania [Oslo], Kaalaas, a 5-1883 (c. fr., NY). Telemark: Tinn Distr., Rollag, Kiaer, 4-8-1890 (BCB). Slovakia. “Slovakia orientalis: montes oe pohorie,” Vana, 29-5-1973 (NY). Sweden. Vastergétland, Billingen, Mdller, 9-8-1914 (herb. Bartram FH); Mosseberg, Lindberg 18.23/6, 22-6-1880 (NY), Flink, 25-6-1880 (MA); Vamb Billingen, Hii/phers, Jun 1916 (c. fr., NY). Smaland: 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 pseudoparaphyllia, but, mostly, they are completely uniseriate, as Ireland’s (1982) later figures show. Multiseriate pseudoparaphyllia are uncommon in this species. Even though they are nomina nuda, the names Anomodon apiculatus B.S.G. and to some extent also Leskea apiculata Schimp. in Mill. Hal. have been exten- sively used in older literature 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 applied to specimens named A. uwematsui 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. rugel/ii 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. rugelit 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), [ 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. Musc. 210. 1801. Neckera minor P.-Beauv., Prodr. 78. 1805. Neckera flaccida Brid., Sp. Musc. 2: 137. 1812.—Tvype: 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- on 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: INpIA. 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: Cuina. Chekiang, Ningpo, Oldham (lectotype, here designated: NY!). Anomodon platyphyllus Kindb., Ottawa Naturalist 7: 19. 1893.—Typre: CANADA. Can. Musc. 256; Macoun Cat. Pt. VI, 171. Anomodon sinensis Mill. Hal., Nuovo Giorn. Bot. Ital., n. ser., 3: 118. 1896.— Type: Cuina. Shaanxi, “China bor. or., Si-Ku-tzui-san,” Giraldi, 1894 (iso- type?: FH!). Anomodon leptodontoides Mill. Hal., Nuovo Giorn. Bot. Ital., n. ser., 4: 275. 1897.—Type: Cuina. Prov. Shen-si sept., Schan-kio, Giraldi, Aug 1895 (FI). Anomodon minor var. inaequalifolius Bartr., Bryol. 50: 207. 1947.—Typr: 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 um), 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 um 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) um 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. SSENTATIV E SPECIMENS. Bhutan. Oongar, Griffith (as A. integerrimus, herb. Mitten NY). Bchina: Heilonejian: Manchuria, Muling, Sinzew (NY). Chihli: I Yuan K’on, Clemens 5098, 28- 41913 (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 ae integerrimus, JE). Liaoning: distr. Feng Tong Yuan Bao, me 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. Ter 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, ie 10-12-1903 (as A. ee erpuE Levier NY), “Botanical Garden,” Gollan, 7-8-1900 ie ieee C Y); “below Old Bot. Garden,” W. Gollan, 26-11-1903 (as A. bee JE). ‘ a orienta bee Te 120, 367 (ex herb. Mitten NY). ener Honshu: Aichi pref., Mt. Mika eh es Pete 3898, 11-9-1947 (NY); Bungo, “Kan a,” Iwasaki ae 10-9— Sere thraustus, Lone Higuo pref., Sibara-machi, Yatu- siro-gun, Tasivo 3340, 16-1-1940 (NY); Mie siee Mt. Fujiwara, 7. Hoguro 34, 19-10-1965 (c. fr., as A. subintegerrimus, NICH); caval Misugi, 7. Nakajima, ou 8 1963 (as oe integerrimus, eXICc. Noguchi 952, JE, MICH, NY), Isejingu, (N. “Takaki 4467, 28-3-1948 (as A. integerrimus, NICH); Mikawa, Miwamura, Chichiiwa, (N. Takaki 11333. ¥ . Horikawa, 26-3- eae aaa NICH), “South Alps,” (N. Takaki 11787, 20-7-1952 (as A. planatus, NICH); Nikko, Kamiji 85, Aug/Sep 1947 (herb. Bartram FH); Okayama pref., RashéOmo, Niimi, /gi, 15—1-1958 (as A. Pere ae: exicc. 22 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 . 20. Distribution of Anomodon attenuatus, A. flagelliformis, A. longinervis, A. minor, A. sieboldii, and A. solovjovii in central and eastern Asia Noguchi 654, NY); Osaka Pref., Takatsuki, Hara, 7. Nakajima, 29-4-1960 (as A. minor oe integer- rimus, HIRO, MICH, NY); Tok yo pref., Nishitama-gun, Nippara, R. Watanabe 13928, 8-8-1967 (as A. ramulosus, NICH). Kyushu: Fukuoka pref., Kawara- dangle usar 2771, 26-8- ies (NY); Kumamoto pref., Hitoyoshi, Mayebara, Jan 1948 (as A. ramulosus, JE, MICH); Miyazaki pref., Nishi- mera-mura, 2. /watsuki 4806, 5-2-1980 (as subsp. integerrimus, NY); Yabakei, Buzan, [wasaki 534, 29-8-1916 (NY). Shikoku: Kochi, Toja, S. Okamura, 10-12-1905 (FI, herb. Bartram F osa, sayoma, Gono 247b, Jan 1904 (FI, herb. Bartram FH). Korea. B. Sauer, 1924 (JE); Fusan: Faurie a Oct 1901 (FI). Hysyeng Yang: Faurie 80, Jun 1901 (FI); Mt. Chiak, W. Hong 5877, 5887, 16/17-7- 1961 (as A. integerrimus, NICH); Mt. Chii, W. Hong 1978, 2082, 16, 17- 8-1960 (as A. integerrimus, NICH); Mt. Sam-sung, W. Hong /508, 3-4 1960 (as A. integerrimus, NICH); Mt. Sokri, W. Hong 2816, 21-12-60 (as A. integerrimus, NICH). Russia. “Regio Iskutkensis, distr. Czeremchovo, Mts. bia er orientales,” L. Bardunov & S. Lipin, 17-7-1959 (as A. eee Rees a. Manitoba: Red River, St. Norbert, Bird 3269, 28-5-1959 (c. fr., DUKE); Parkland, N of a, C Bird 5953, 13-5-1961 tc fr.. FH, MICH); Winnipeg, SE ae Mueller- oo 9-4-1961] Lee Ontario: Edmonton, White, 10-11-1893 (as A. platyphyllum, c. fr., NY); Ottawa, J. Macoun, 3-10-1892 (as A, pe, DUKE, c. fr., FH); Thunder Bay, Sturgeon Bay, Blake Twp., C. Garton Lae 5-5-1981 (MICH); “about Falls of Niagara,” Aig (MICH). Québec: Com. Montcalm, Park du Mont Tremblant, F. Hermann 16612, 19-7-19 ae Chelsea, J. Macoun, 17—5—1907 (MICH). Mexico. Chiapas: Las Casas, A. Sharp 35/5, er reas KE, MEXU, MICH, NY). Chihuahua: Parque Nac. Cascada de Basageachi, A. Whittmore 2857 et ie 4-10-1986 (MEXU). oR: 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE pss) Guerrero: Chilpancingo, 25 km W, A. Sharp 1052, 22-10-1944 (MEXU, MICH), ca. 22 km me A. eee 1096, 27-10-1944 (MICH), Omiltemi, A. Sharp 1132 & 1133b, 27-10-1944 (MICH lala XU, NY). Jalisco: La Cumbre, 10 mi SW of Autlan, H. Crum 458a, Jun 1949, H. Crum oe Aug ie (MICH). Morelos: Cerro Tepozteco, C. Delgadillo, 28-9-1991 (MEXU). Oaxaca: San Pablo hae 70 km E of Oaxaca, A. Sharp 4609 et al. (MEXU). U.S.A. Alabama: Jackson Co., Pint Rock, 1. G.-de la Cerda 1109 & E. Lyons, 4-2-1984 (DUKE). Arkansas: Marion Co., Flippen, L. ee 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, ae f Hartman, 1879 . fr., JE); Chicago, RGll, 1822 (as A. Si sia FH). Indiana: Parke Co. key Run St. Park, W. Welch 11482, 13-11-1937. Iowa: 1 oO cent S. pean 7109, 18—6- a (c. fr., FH). Kansas: Salina, Henry (as A. "ohio FH). M ee Cove, E. Merrill 126, Oct 1898 ( (FH): Spencer’s Woods, Anthony 2413, 888 (c. DUKE): ous (oe Norway, Bacon 8942, 5-5-1929 (DUKE). Massachussetts: he wed ahaha (JE); West Townsend, Thomson, 24—9-1935 (FH). Michigan: Miner’s W. Steere 594, Jul 1933 (FH). Minnesota: Winona Bluffs, J. Holzinger, 2-12-1899 (c. fr., FI); Benton Co., Henry, 9-1885 (JE); Chippewa Co., pr. Montevideo, J. Holzinger 156-1901 (c. fr., FH). Missouri: Hannibal, Mark Twain’s Cave, Davis, 7-11-1911 (FH); Cooper Co., S of Blackwater, < Sharp, 7-10- 1938 (FH); Franklyn Co., Merramec Rvyr., 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: a 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, 20-4-1952 (DUKE); Mitchell Co., Roan Mtn., A. eeees 1984 (DUKE); Orange Co., Eno River State Park, /. G.-de la Cerda 1938, 31-11-1984, (c. fr., DUKE), 1. G.-de la Cerda 1942, 1943, 7-12-1984 (DUKE). Ohio: Columbus, Schraeder (c. fr., as a ee JE); ibid. Sullivant, 1850 (c. fr., JE). Pennsylvania: Huntingdon Co., Spruce Creek, A. & P. Felton & I. G.-de la a 1853, 30-9-1984 (DUKE); McKean Co., Burnett 2831, 7-11-1897 (c. fr., FH). South Dakota: Day. Co. Wanbay Nat. Windlife Ref., §. 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, A. Grout, 23-4-1892 (as A. obtusifolius, c. fr., DUKE Newfane, A. Grout 2416, 25-7-1902 (DUKE). Virginia: Marion, A. Vail & E. Britton, 6-6-1892 (as A. ae 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 Po tomac, sia 29-5-1938 (DUKE). Wisconsin: Richland Co., Hub City, Hermann 28308, 21-5-1978 (DUKE). Some slender specimens of A. minor with branch 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- en off, as in A. thraustus. The delicate, narrowly lingulate 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. rugelii 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. ruge/ii; 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 1n 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 slightly 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 A. 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 of A. 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 4557, 29-12-1945, type, herb. Bartram in FH!), is almost identical with Iwatsuki’s subspecies. I consider all these variations within the confines of A. minor. Anomodon ramulosus Mitt. 1s 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 “foliage on some stems Is subcompressed,” I consider it appropriate to designate the Ningpo specimen of 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. 1s not to be confused with A. obtusifolius Mitt. (a nomen nudum), which has been applied to some East Asian specimens of A. viticulosus. La.ii. Anomodon subsection Abbreviati (Z. Iwats.) Granzow, comb. nov. Anomo- don sect. Abbreviati Z. Iwats., J. Hattori Bot. Lab. 26: 51. 1963. 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 29 ial 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 Mill. Hal., Nuovo Giorn. Bot. Ital., n. ser., 3: 117. 1896.—Type: Cuina. 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 um long, 23 um 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 um, 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 um 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- ne (NY). Shaanxi: /baraki, 25-5—1900 (as A. pera ex herb. Paris FI). Japan. ee 25-5- 900 (as A. ramulosus, ex herb. Paris FI); Bungo, Mt. Tsurumi, N. /wasaki 1229, 25-9-1921 (NY); re , Mt. Tsurugi, M. Kamimura 2627, Aug (ose (NICH); Higo, Gokanoso, H. Kaneda 212, 30-7- 1935 (NICH): Kii, Odaigahara, N. Takaki 4885, 3-8-1948 (NICH); Musashi, Mt. Mitsumine, NV. Takaki 4535, 9-4-1948 (NICH). Hokkaido: Iburi, nr lake Shikotan, N. /wasaki, 18-10-1914 (NICH). Honshu: Fukushima pref., Tamura-gun, Mt. Ootakine, 7. Higuchi 1564, 13-6-1953 (NICH); Hiroshima pref., M t. Kammuri, S. Nakanishi, 8-11-1960 (JE); Hyogo pref., Mt. Sappico, N of Hmayi, 7. Kodama 5—4- 1962 (c. fr., NICH); Kinkwasan, A. Noguchi, 8-6-1929 (herb. ae FH); Kanagawa pref., Hakone Mts., eee 768-J—54, 16-5-1954 (DUKE); Kotuke (?), Mt. Okeci, A. Esumoda (?), 15-11-1912 (Herb. Bartram FH); Mie pref., lishi-gun, Misugi-mura, eee Nakajima, 4-3-1963 (c. fr., JE), Z. Iwatsuki 93, 2/5-12-1963 (NICH); Miyagi pref., isl. Kiuzakan, 7. Higuchi 1195 4-3-1953 (NICH), Oshika-gun, Mt. Kinka, 7. Higuchi 1/95, 4-5-1953 (NICH); Musasaki, Titibu-me, M. Yano 5193, 24- 5 cane eee Sasaoka NY); Nagano pref., lida, Matsu river, M. Higuchi 9-8-1980 (NICH), Kamiina- Mt. Senjogadake, R. Watanabe fe 24-7-1973 (NICH), Kisofuku-shima, pica 52003, 17- 10-1972 (DUKE), Takaki 6402, 27-6-1949 (NY), Minami Zumi-gun, Azu De R. Watanabe 11425, 13-8-1964 (NICH), Shimoina-gun, Mt. Toyokuchi, R. ne 9389, —1963 ic: Sea pref., Mt. Yahiko, Y. /kewani, Mar 1948 (JE); Saitama pref., “Japonia enn Chichibu, H. May 9-5-1890 (ex herb. Broth. FH, FI, NY), Mt. Mitsumina, N. Takaki 4535, 9-4-1948 (NY), Chichibu. gun, Icirikanon, R. Watanabe 14400, 20-8-1968, (c. fr., NICH); Shimano, “Mt. Yatsu,” £. fishiba, 26- 7-1908 (DUKE); Shizuoka pref., Honkawane-cho, Kanzoo, Z. /watsuki, 15—9-1959 (NICH); Wakaya- ma pref., Mt. Gomadan, M. Micutani, 12-9-1949 (DUKE, NY); Yamanashi pref., Ashiyasu, S. Jnoue, 20-8-1974 (JE), Mt. Kita-dake, R. Watanabe 16701, 12-8- a (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. ae 2316, 6-5-1976 (NY), Nishiusuku-gun, 236 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 \2] o N Gi ry 6S2s EAA 60° 4 8 (Yes ae REGS (2 —> => FESS OP QE™ ee se LS) © SELQBS OQpDw & oie (og g ESOSS) ODED les OO g) 100 ;1m, SSS OS Nese ec 100 tum GOGSOOAGYL VY) IG. 21. Anomodon abbreviatus Mitt. a-c, branch leaves; d, cross section of stems without ditferenciated central strand; e, cross sections of branch leaves; f, apex of a branch leaf; g, h, leaf cells; 1, leaf in cross section. Based on: a, d, Japan (loc. ignot., DUKE); b, e, f, E. lishiba, Japan, Shimano, Mt. Yatsu, 26-7-1908 (DUKE); c, i, Yano, 5193, Hondo, “prov. Musasi, Titibu-me,” 245-1920 (NY); gh, Schofield 52033, Nagano pref., Kisofuku-shima, 17-10-1972 (DUKE). Mt. Shiroiwa, N. Suzuki & M. Kakuchi 12 (c. fr., NICH). Shikoku: Ehime pref., Kamiukena-gun, Omogo-kei, Rk. 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 solovjovil could be mistaken for A. abbreviatus, but the latter is more robust, and the papillae are higher (more than the laminal 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 237 | i SEN ey hh ; Perey deen _ Cy ae j a . Ms 4 fi’ } mx y) oR A. abbreviatus A. dentatus H. acutifolium —— FIG. 22. Distribution of Anomodon abbreviatus, A. dentatus, and Herpetineuron acutifolium in central and eastern Asia. 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 A. solovjovii. 5, Anomodon solovjovii Lazarenko, Rev. Bryol. Lichénol., 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. [AI] 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 mm thick when dry, scarcely branched; secondary branches erect. Central strand not differ- entiated. Branch leaves erect when dry, 1.4-2.0 mm long, secund when moist, 238 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 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 um), 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 um 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. istribution (Fig. 20). The Russian Far East (south of the Ussuri river, region of Javarovsk), Henan province in China, and Korea. Anomodon solovjovi grows on tree trunks at medium elevations. ADDITIONAL SPECIMENS EXAMINED. Korea. Mt. Odal, W. Hong 2563, 2627, 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 um 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 um). A disjunct population from the interior of eastern China was recently described as A. solovjovu 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 variability 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. Lai. Anomodon subsection Dentati Granzow, subsect. nov.—lIyree: Anomodon dentatus C. H. Gao Folia ramealia aliquantum grandia, oblongo-lingulata, nonnihil spatulata; apex margine valde serrato, plerumque effractus; costa prope apicem evanescens, sub- ter laminae cellulis occulta. Branch leaves narrowly oblong-lingulate to spatulate, abruptly constricted near the middle into a very narrow upper portion; apex strongly and irregularly serrate, often broken off; costa fading well below the apex, obscured by laminal cells; laminal cells with multiple branching papillae. 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 239 6. Anomodon dentatus C. H. Gao, Flora Muscin. China Boreali Orientale, 380. 1977.—Type: Cuina. 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 EXAMINED. CHINA. “Kirin [Jilin]: Tchangbaishan,” G. Qian 22/08, 24—9- 1981 (as A. attenuatus, NY). 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. attenuatus) from the same mountain range as the holotype, I conclude that A. dentatus is a well-defined species. Lb. Anomodon section Haplohymenium (Dozy & Molk.) Granzow, comb. nov. Haplohymenium Dozy & Molk., Musci Frond. Ined. Archip. Indici, 127. 1946.—Typre: 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. fristis), 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. 240 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 - C YH Se PRINT Nees etey: J Teh, {7 asa SS ‘ Reena PEATE oa af] 8 Ww z ery Reseegiate Baers HG ie Pood i oo a cote, Hy > pS aT = x By lh ae \& Jy Oo ave Ko TA RSI Oxy] pavers. Roos SOAS US | a i oO an, ies Cf AX) a saishe nal (ae STN ee ca ST , 0 URIS Re ‘ ANS i i NON SA FIG. 23. Anomodon dentatus C. H. Gao. a, b, branch leaves; c, d, apex of branch leaves; e, cross section of branch; f, branch leaves in cross section; g, basal cells of branch leaf. Based on: a, c, e-g Guo Quian 22108 (NY); b, d, Chen Gao 1231] (type), China, province of Kirin, Chang Bai-shan, (IFSBH). 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 241 7. Anomodon tristis (Ces.) Sull. & Lesq., Musci Bor. Amer. 52, 1856. Leskea tristis Ces. De Not., Syll. Musc. 67. 1838. Hypnum triste (Ces.) Miill. Hal., Syn. 2: 478. 1851. Haplohymenium triste (Ces.) Kindb., Rev. Bryol. 26: 25. 1899.—TypeE: ITALY. Lago Maggiore, Monte di S. Crescenzio, Verbanum, 1837, C. Cesati (holotype: RO!). Anomodon fragilis Wilson, London J. Bot. 7: 275. 1848. Hypnum fragile (Wil- son) Mill. Hal., Syn. 2: 471. 1851.—Tyre: Cuina. Chusan (n.v.). Anomodon sinensi-tristis Mull. Hal., Nuovo Giorn. Bot. Ital., n. ser., 3: 118. 1896.—TypeE: 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: Bo.i- vIA. Santa Cruz, Cerro Ambor6, Herzog (holotype: JE!; isotypes: NY! US!). Haplohymenium scabrum Broth., Rev. Bryol. Lichénol. 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: TAtIwan. 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.—T ype: Cuina. 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 um, 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 242 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 < | Bet = PSO pan “| # —H, toccoae Fig 24. Distribution of Anomodon tristis, A. pseudotrists, and Herpetineuron toccoae. rare in Asia. Anomodon tristis grows on bark of trees in deciduous forests in rather montane regions (except in eastern North America). REPRESENTATIVE SPECIMENS. China. Hubei: Shenongjia Forest Distr. Quiujiaping to Mt. Laojun, Sino-Amer Exped. 663A (as H. formosanum, NY). Jilin: An-tu, NW Mt. oa ng bai, 7. Koponen 3719], 24-9-1981 (NY). Szechuan: Pe pei, pr. Chun king, mt. Tsin yun Shan, P. C. Chen, May 1940 (JE). Japan. “Japonia meridionalis,” Omura (Hijeu), J. Ferrié, 1S—1—-1898 (FI). Honshu: Hiroshima pref., Toj6-shi, Hiva gun, W. Schofield 54588 & T. Seki, 18-4-1973 (DUKE), Miyajima Isl., Hiroshi- ma Bay, P. & A. Redfearn 33632, \&— 985 (as H. eal DUKE); Kanagawa pref., Hakone Mtns. Nat. Park, P. Redfearn 720-J-54, 16-5-1954 (DUKE); Miyagi pref., Sendai, Y. ishiba: Nov. 1906 (JE); Osaka pref., Takasuki, she T. ah 29-4-1960 (JE, MICH); Yamanashi pref., Kofu-shi, Maizuru oe T. Osada, 6-4-1978 (MICH). Kyushu: Fukuoka pref., Mt. Hikasan, Y. Kuwa- hara 2594, 30-7-1952 (NY); Miyazaki pref., Minaminaka, Obi, S. Hattori & . Kurata, May 1947 (JE, MICH). Shikoku: Kochi pref. (prov. Tosa), 7. Makino, 1887 (JE), Mt. Honokawa, S. Okamura, 31- 12-1907 (FI). Taiwan. Taitu Hsien, Yehyu Tsun, S. Lin 169, 22-8-1975 (as A. pseudotristis, MICH) Taipei: Wulai Hsiang, betw. Fushan Tsun and Pling, $. Lin, 21/24—7—-1973 (as H. formosanum, MICH); eee Po, Son Tcheou, Courtois, Feb 1908 (JE). i. Maui: Haleakala Nat. Park, E of Kaupo Trail, below Paliku Pasture, W. Hoe 4474, 5—6- 1977 nee Costa Rica. Heredia: S slope of Volcan Barva, 1 km WNW Rio Ciruelas, 8.2 km from S. José de la Montana, M. Crosby 9894, 9923, 3-5-1975 (MO). Mexico. Jalisco: Laderas de la Ferreria, Manantlan, H. Crum 9/4A, 23-7-1949 (as A. minor, NY). Nuevo Leén: Cerro de : Silla, pr. Monter- y, L. Harvey 1022b, 23-6-1939 (MICH). Sonora: Bavispe, ears del Temblar, F. Phillips 2 - 3. 1940 (MICH). Jalapa: pr. S. Miguel el Soldado, above Jalapa, C. eae et al., 8-7-1966 (MEXU). U.S.A. Ohio: mere es (WE): Se lan Philadelphia, 7. James, 1879 (JE), sis hickon Creek, G. Kaiser, 7-12-1911 (DUKE); M an Co., Quintuple (?), D. Burnett 2956, 1898 (DUKE). South Carolina: ene Co., Reedy ee Falls, pr. Nimmons, L. Anderson ee as 8-1949 (DUKE); Oconee Co., Chatooga Rvr. E fork, L. Anderson 8372, 18-8-1949 (DUKE). Virginia: — ssh Nat. Park, Big Flat Mtn., /. ane & F. Wynne, 26-6-1944 (MICH). Aus ent Wald, pr. Goriziam, Loitlesberger & Rudez (C); Meran: Vellanerthal, tc & ee peer 4 (INA, herb. Luisier). Italy. Prov. Cuomo: Borgnana, A. Artaria, 13-11-1899 (C, JE); “Gallivago in ee mo,” ae 2-7-1868 UE): Verbanum, Valle di Bieno, 5, De Notaris a 1865 (C, JE); “Luganer See, Cuasso al Piano,” A. Artaria, 9-5-1897 (C, FH), 9-10-1898, 13-11- 1899 (C). Novara: Avola, L. ce. Aue 56 (FI). Piamonte: Lago Maggiore, pr. Mergozzo, De Notaris, Aug 1869 (C), Montorfano, De Notaris Aug 1859 (FI, RO). Switzerland. Ticino: aes B. Weber, Apr. 1883 (JE), pr. Lugano, L. Mari, ae 1889 (JE), “Selva pr. Porya.” L. Mart, Jul 883, Jul. 1889 (FI); Locarno, De Notaris 19, 1863 (R); pr. Locarno, M. Jdglli, 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. sér. 3, 2: 310. 1844. Neckera sieboldii (Dozy & Molk.) Mill. Hal., Syn. 2: 81. 1850. Haplohymenium sieboldii (Dozy & Molk.) Dozy & Molk., Musci Frond. Arch. Indici 4: 127. 1846.—T ype: JAPAN. von Siebold (holotype: L!, c. fr.). Haplohymenium brachycladum $. 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, sér. 2, 3: 281. 1911.—Tyee: Japan. Shikoku, Tosa, mont Yokogura, Okamura (PC). Haplohymenium gonoi Broth., Ofver. Finsk. Vet. Soc. Férh. 62: 33. 1919- 1920.—TypeE: JAPAN. Shikoku, Tosa, M. Gono 189 (n.v.). Haplohymenium pellucens Broth., Ann. Bryol. 1: 21. 1928 —Type: Tatwan. 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) um in diameter, densely papillose. Calyptra with thick papillae and hyaline hairs. Chromosome number unknown. 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 34] (as H. pellucens, c. fr., NY). 9. Anomodon pseudotristis (Mill. Hal.) Kindb., Enum. Bryin. Exot. 7. 1888. Hyp- num pseudotriste Mill. Hal., Bot. Zeit. 13: 786. 1855. Haplohymenium pseudotriste (Mill. Hal.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907.—TyPeE: SouTH AFrRica. 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.—T yee: 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.—SynryPeEs: 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.—T ype: AUSTRALIA. Queensland, Helidon, Dec 1888, Wild /6 (n.v.). Anomodon mithouardii Paris & Broth., Rev. Bryol. 29: 97. 1902. ae mithouardii (Paris & Broth.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907.— Type: VIETNAM. Entre Lang Son et Dong Dong, Oct. 1901, ae (ni: Anomodon submicrophyllus Cardot, Beih. Bot. Centralbl. 19(2): 128. 1905. Haplohymenium submicrophyllum (Cardot) Broth., Nat. Pflanzenfam. 1(3): 986. 1907.—Synrypes (n.v.): TAIWAN. Maruyama, Faurie 6 (14, 15); Kelung., Faurie 9S; Tamsui, Faurie 102, 107; Hokuto, Faurie 115 [all in 1903}. Anomodon stenoglossus Cardot & Thér., Bull. Acad. Int. Géogr. Bot. 18: II. 1908.—T ype: JAPAN. Liou-Kiou, Naze, 1898-99, Ferrié (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. Férh. 62: 32. 1921.—Type: Japan. Hondo, Chiba, Gono IV.25. (n.v.) Haplohymenium pellucens var. obtusifolium Broth., Ann. Bryol. 1: 21. 1928.— Type: TArwan. Prov. Taihoku, Hill Senjyo, Sasaoka (n.v.) Haplohymenium tenerrimum Broth., Rev. Bryol. Lichénol. 2: 10. 1929.—Type: JAPAN. Prov. Ise, Ujiyamada, Tutiga (Herb. Sasaoka, TNS, H?). Haplohymenium mithouardii var. viride Thér., Ann. Crypt. Exot. 5: 179. 1932.— Type: CHINA. Prov. Fukien, Yanping, Buong kang, 23-6-1925, Chung B.101 (isotypes: MICH! NY!). Haplohymenium fasciculare Nog., J. Jap. Bot. 13: 791. 1937.—Type: JAPAN. Sikoku, Mt. Isiduti, prov. Iyo, 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 hyaline 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. EPRESENTATIVE SPECIMENS. South Africa. Cape Prov.: Groenkop Frst. Resv., M. & C. Crosby 8080, 25-1-1973 (DUKE); Transkei, rd. Dwesa to Idutywa, J. van Rooy 2083 (NY). Natal: St. Lucia Estuary Forest, J. van Rooy 209 (NY). Transvaal: Marieskop Plantation Natr. Resv., M. & C. Crosby 7595, 11-1- aie hacen Japan. Honshu: Miyagi Pref., Sendai, /ishiba, 16-12-1909 (as H. triste, DUKE); Mie Pref., Nabori shi, Akame, Ww 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. se ee 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, 1-11-1983 (as A. huttonii, MICH). Matakana, J. Kirk (as A. huttonii, NY). Anomodon pseudotristis is similar to A. tristis 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. ¢ristis or A. sieboldti. 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. Haplohymenium flagelliforme Savicz-L., Bot. Mater. Inst. Sporov. Rast. Glavn. Bot. Sada RSFSR. 1: 98, 101. 1922.—Type: Russia. “Asia Orientalis: regio Austro- WES OES in valli fluminis Souczan, Lag Frolovka in flumini Pen- saou,” 3-8-1913, Boulavkina (holotype: L Haplohymenium cristatum Nog., J. Jap. Bot. ae 146. 1945.—TypeE: 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, cells 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. ADDITIONAL SPECIMENS EXAMINED. Russia. “Asia Orientalis: in regio austro-ussurensis, in p. sup. 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.- SynryPEs v.): JAPAN. Kiu-diu, Ichifusa, Jun 1900, Faurie 1194; Tsu-Shima, May 1901, Faurie 1604. Haplohymenium biforme Broth., Bull. Soc. Bot. Geneve sér. 2., 3: 281. 1911.— Type: JAPAN. Mont Koya, Nakanishiki (Herb. Brotherus). Haplohymenium piliferum Broth. & M. Yasuda, Ofver. Finsk. Vet. Soc. Férh. 62: 33. 1919-1920.—Tvpr: JAPAN. Hondo, Mt. Akagi, Tsunoda (Herb. Yasuda nr. 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 spinosum 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, 7. 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 H. 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. [wat- suki & K. Minamidani, 28-10-1976 (MICH). The leaf papillae of A. longinervis are similar to those of A. abbreviatus and A, solovjovil, although in A. longinervis papillae are more numerous (to 3-4 per cell), not as high, and often branched. I.c. Anomodon section Thrausti Granzow, sect. nov.—Tyre: Anomodon thraustus Mill. Hal. Planta tenuis; folia ramealia angusta, tenuia ac/vel fragilia; apex plerumque effractus; costa evanescens, brevis, dimidium laminae vix attingens, subter lami- nae cellulis 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: Cuina. “China interior, Shen-si sept., in monte Tuikio-san,” Sept 1896, Giraldi (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 1997; GRANZOW-DE LA CERDA: ANOMODONTACEAE 247 se. & F350 wiaere age a S. LY ae G. 25. Anomodon thraustus Mill. Hal. a, branch leaf; b, cross sections of stem with cells of the central strand differentiated: c, cross sections of branch leaves; d, basal cells of the lamina; e, end o the costa, fading at the middle of the leaf; f, cells of the base of leaf at the insertion. Based on: a, e, Exp. Sino-Amer 274 A, China, “Western Hubei Prov., Shenongija Forest District, Along Miaogou 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, 259-1981 (NY); f, Lishiba, Japan, Rikuhf, Mt. Hayoshine (?), Jul 1909 (NY). side, ending far below the apex (more than 15 um 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 um 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 A. 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. ITIONAL SPECIMENS EXAmiNeD. China. Heilongjian: “Lesser Khingan Mtns,” Shuan-zi-he Guo oi 6672, 15-7-1963 (NICH). Hubei: me eng Forest distr., sy aoe Sino-American xp. 274A, 28-9-1980 (as A. viticulosus, NY); Jizigou, ile American Exp. , 25-8-1980 (as A. minor, NY). Jilin: “Chang Bai, An-tu, Er-do Ba 1 Xi Zien,” 7. Koponen pis 25-9-1981 (as A. minor subsp. integerrimus, NY); Jian, Dayu shan, Guo Qian 7802, 8-9-1963 (as A. rugelii, IFSBH); ang Qing Xian “Wang-chiu,” Daxinancha, Nangou, Guo Qian 8575, 16-8-1973 (IFSBH). Tibet: Ya-Dong, Zang Mu 56, 29-5- 1975 (NICH). India. Pamacied Pradesh: Simla “7000,” 7. Thomson 1172, 1847 (as A. obtusifolius var., Wils., syntype for A. integerrimus, NY-herb. Mitten), Simla “7000, 8000,” 7. Thomson 1126, Oct 1849 (mixed with syntype of A. planatus, herb. Mitten, NY). Kumaon: Dahli valley, /. etna 7-8-1886 (as A. planatus, MICH). Uttar Pradesh: “Western Hymalaya. Seven Oaks estate,” Mussoorie, W. Gollman, 2-10-1900 (as A. glossophyllus, NY). “India Oriental,” Bell 187 (as A. ee NY). Japan. Honshu: Hiroshima pref., Tojo-cho, Taishaku, M. Higu- chi, 7-10-1978 (HIRO); Iwate pref., Rikuchu: Mt. Hayachine, F. eae Jul 1909 (as A. minor. NY). Korea. Keisyo nando, Mitsuyo, N. Maki 2985, 31-7-1931 (NICH); Sauer, 1924 (as A. minor, JE); Mt. Ciak, W. Hong 5874, 16/17-7-1961 (as A. oo NICH); Mt. Sokri, W. Hong 2801, 21-12-1960. Fusan, Faurte 241, 1906 (ex herb. Cardot NY). Nepal. “In Himalayae reg. temp.,” Wallich (mixed with hl for A. integerrimus, 7 ve NY). Russia. Far ee (Primorsk) : Ussurusk, rvyr. Sit , L. Bardunov, 6-9-1962 (NICH), ibid., V. Ardeieva, 3-9-1968 (NICH), Tetjukhe, L. Bardunov, ot 11-1961 (NICH), pr. Partizansk, L. sehen & Y, renardaniees a, 16-9- 1964 (NICH), Schkotovskyi, pr. Podiapolsk, L. Bardunov, 9-9-1974 (NICH); “regio Ussuriensis,” Vladivostok, P. Sinzer, 25-04-1905 (as A. minor, ex herb. Sait FI), Nikolok, P. Sinzer, 3/1905 (as A, minor, FH), Utsisheibo (Manchuria), Vladivostok, P. Sinzer, 13-03-1905 (as A. minor, c. herb. Brotherus FI); “Far East, Lake Khanka, pr. village of Lusanovo,” L. Bardunov, 23-8— ite (NICH); “Sibiria: Primorskaja, Okaenskaja,” E. Sinova, Sep 1926 rene 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 of A. thraustus, in the following foliar characters. 1)The leaves of 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. 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 249 a ——— s \ Fi ( - af ) (Nee as, See \ BR ; —~— lee bls eA \ | eh ae os = cent 1 tre pi w LEY are ( J 3 C \ / a ae ») , ¥ a_i jw— VA | i : a oe C “—s 7 ae [eae ) ¢ { 7 arg ee) 4 Levi = as \ Nf —— eT Oe ay \ \4 ) /\ AN \ (i; nora a = ae 3 \ J ran FIG. 26. Distribution of Anomodon giraldii and A. thraustus in central and eastern Asia. 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 A. minor, not of A. thraustus. Mitten’s mixed types are probably the cause of much of the confusion surrounding the identity of both taxa. II. Anomodon subgenus Pseudoanomodon Limpr., Laubm. Deutschl. 2: 774. 1895.—Tyrpe: Anomodon attenuatus (Hedw.) Hib. Stems and primary branches branching in a stairway fashion, or with second- ary branches irregularly pinnate, profusely fasciculate. Leaf apex acute to acumi- nate; laminal cells rhombic in most taxa. 250 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 13. Anomodon rostratus (Hedw.) Schimp., Syn. Musc. Eur. 448. 1860. Leskea rostrata Hedw., Sp. Musc. 226. 1801. Hypnum rostratum (Hew.) P.-Beauv., rodr. 69. 1805.—T ype: 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 um, often somewhat flexuose above, with high papillae in one line 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- limes 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-I.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 um 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- macker et al. (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. longifolius (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., “Svania Dadianosum, Hippi, inter Muri et Lentechi,” Sommier & Levier, 29-7-1890 (NY anada. British Columbia: Eagle Pass Mtns., Big Eddy of Columbia Rvr., nr. Revelstoke, 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 § Distr. SE of Parsons Pond, J. Hancock & C. rae 468, 3-7-1972 ( apne Nova Scotia: Colchester Co., Five Islands, R. Ireland 17222, 10-7-1974 H). Oniane: Grey Co., Inglis Falls, nr. Owen Sound, H. Crum 11027, 3 3-8-1961 (MICH). Ottawa 7 Macoun, 20-10-1893. Manitoulin Isl., Tamarack Point, C. Grass! 6219, 19-7-1932 (MICH) 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE Pioul G. 27. Anomodon rostratus Cerone) Schimp. a-c, branch leaves; d, apex of branch leaves; e, portion of peristome. Based on: a, d, e, F. Artaria, Italy, Como prov., Bucca del Pertuso, Blevio, 15— 7-1897 (NY): b, Hermann 26045, Mexica: Nuevo Leon, Monterrey, Sierra Madre Oriental, 22—7-1975 (NY); c, Boros, Hungary, “Comit. Veszprém, rivi Gerence, pr. Baconybél,” 3-4-1959 (NY). Québec: Armand, $ of Montreal, K. Holmen, 24-9-1959 (NY). Gaspé Sud, Percé, H. Crum & H. Williams 10882. 25-7-1960 (MICH). Guatemala. Alta Verapaz: Rio Carcha, entre Coban y S. Pedro Coban, P. Standley 90129, 26-3-1941 ae Bartram FH), 90/30 (MICH). El Quiché: A. Sharp ee 6-2-1945 (herb. Bartram FH). Mexico. Chiapas: Las Casas, A. Sharp 3521, 25—4-1945 (US,M NY); S ai San Cristdébal, R. Diill 16, 8- He 1966 (NY). Hidalgo: Chapulhuacan, A. Sharp 1541a, oo 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). Puebla: pr. Zacapaxtla, A. Sharp ogee 17-10-1945 (MICH). San Luis Potosi: “above Xilitla,” A. Sharp 5866, 5868, 21-4-1946 (M Y, US). Tamaulipas: Sierra Madre, W of Victoria, H. Crum 1831, Aug 1950 (US). Jalapa: evidence, 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). ae Sta. Cruz Co., Sta. Rita Mtns., White use Canyon, i Haring 3472 & H. Haskell, 7-5-1945 (MICH). Arkansas: Drew Co. , Wilmar, D. he: 24239, 10-4-1943 (MICH). Colorado: Chandle acne T. Brandegee, 1876 ee Mon- trose Co., W Paradox Creek, W. Weber et al. B—43564, 1-6-1973 (N ee Connecticut: Eas Allen, 7-9-1882 (MICH). aoe Jackson Co., Marianna Caverns, R. Schornherst on fe 31940 (MICH). Georgia: Stone Mtn., J. Small, Jan 1894 (MICH); Demorest, O. VanHilse, Feb 1913 (MICH). Illinois: Pope Co., Bell Smith Springs, NW of Eddyville, D. Vitt 7407, 25-5-1973 (MICH). lowa: Fayette Co., Brush ne Canyon St. Park, S. Gradstein & D. Horton 6392, 16-4-1987 ee Kansas: Douglas Co., 14 km S-SE of iegnence S. Churchill 9370, 11-4-1978 (MICH). Riley Co., km SW of ae S. Churchill 9401, 14-4-1978 (MICH). Kentucky: Trigg Co., Cadiz Mill, . Harvill 2000, 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 o., Port Austin, 1. Schnooberger 1754, 8-10-1938 (MICH). Marquette Co., Presque Isl., F. Hermann pie) CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 FIG, 28. Distribution of Anomodon rostratus and Herpetineuron acutifolium in the New World. 28757, 4-10-1978 (MICH). Missouri: ages View Glades St. Nat. Area, A. Whittemore 3408 et al. 2-3-1989 (MICH). Nebraska: Cass Co., Weeping Water, J. Sheldon B25, 29-8-1898 8 (NY). New York: Clinton Co., Valcour Isl., J. Smith 38084 & L. Lowinger, 10-6-1965 (MICH). Suffolk Co., L.I., Peconic, R. Latham 36740, 25-3-1947 (MICH). North Carolina: Macon Co., Chatooga River, /. G.-de la Cerda ee 16-6-1984 (DUKE); Orange Co., New Ho ope Creek, 1. G.-de la Cerda 1939, 3-11-1984 f Wake Co., Neuse River, Wake Forest, L. Anderson 11159, 23-11-1959 (c. fr., DUKE); Chowan Co. a nton airport, Albermale Sound, A. Weakley 3540-3549, 21-11-1983 (c. fr., DUKE). South Carolina: Berkeley Co., Cypress Gdns. E. Taylor 2211, 7-4-1960 (NY). South Dakota: Lawrence Co., Black Hills, cee Canyon, FE. Lawton 2177, 29-9-1953 (NY). Meade Co., Black Hills. Stage- rm canyon area, 2 Churchill 9216, 26-S-1977 (MICH). Tennessee: Wayne Co., Glenrock Branch, a eke 8539 & L. Jackson, 20-1-1989 (MICH), Sevier Co., Elkmont, A. Sharp 405, 252-1940 H). Texas: Dallas Co., nr. Dallas, J. Boll, 1874-80 (NY). Liberty Co., 21 mi SE of — 2 aie & R. Wilbur 668, 3-7-1959 (MICH, NY). Virginia: Smyth Co., Mount Jefferson Natl. For 1. G.-de la Cerda 1887, 6-10-1984 (c. fr., DUKE). West Vir rginia: Greenbrier Co., S of White ae Springs, L. Anderson 24752, 16-7-1986 (DUKE). Wisconsin: Richland Co., Richland Ctr., M. Nee 12537, 4-7-1974 (MICH). 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 233 Bermuda. Harrington House, S. Brown 558 (NY). Abbots cliff: &. G. Britton 1877, 13-9-1813 (US). Haiti. Mtns above Kenskoff, F. Mackaness 153, 19-1-1944 (MICH); btw. Savanne 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 ° Y). Jamaica. fee Green: W. Maxon ee Jul 1926 (NY); vicinity of Arntully, Orcutt 3185, 3-10-1927 (herb. Bartram FH); vicinity of Abbey Green, W. Maxon 10083, Jul 1926 (US). Blue Mountains G. Orcu it 2854, 5-9-1927 (US, FH). “Westhalia: (Ceddar Hurst),” C. Orcutt 3762, 1-11-1927 (US). Austria. Baden: Wien, J. Benoger (?), 1871 (NY). Salzburg, D. Sauter (NY). Tirol: Merau, Milde (NY). “Austria inferor”: Isaden (?), Juratzka, 30-6-1867 (NY). “Niederésterreich”: Baumgartner, 30-4-1891 (NY). France. Haute-Garonne: Prat de Joueou, “énvirons de Luchon,” Dismier 28/7, 14— 8-1928 (MA). Germany. Bayern: “Nagelfluh, pr. Shafflarn,” pads aor (NY), 1. Familler, Aug 1906 (MICH). Hungary. Comit Veszprém: “rivi Gerence, pr. onybél,” A. Boros, 3-4-1954 (NY), Nagy-Pénzlik cave, mt. Somhegy, A. Boros, 4-10-1937 ay a Como: Buca del Pertuso, Blevio, F. Artaria, 15-—7-1897 ‘ Y); “Nino, inter Blevio et Como,” F. Artaria, 20-9-1897 (NY). Romania. Transylvania: distr. Fagaras, M. Péterfi, 2 8-1921 (NY). Spain. Girona: vall de Vidra, Les Llances, /. G.-de la Carth 2323a & M. Sanz, 10-1-87 (c. fr., BCB, MA), Rec de l’Espirau, M. Sanz, 7-3-1986 (BCB), Vilallonga de Ter, F. Lloret 33.3.7, 13-3-1986 (BCB). Huesca: Ordesa, R. Diill, 235-1960 (BCB) (see Granzow-de la Cerda, 1988a). Switzerland. Ticino: Lugano, Gandria, 7-9-1885 (c. fr., ex herb. Boissier NY); Campione-Bissone, P. Conti, Oct. 1893 (MICH). Anomodon rostratus 1s 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. Jongifolius 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 (1.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.) Htib., Musc. Germ. 562. 1833. Leskea attenua- ta Hedw., Spec. Musc. 230. 1801. Hypnum attenuatum (Hedw.) Sm., FI. Brit. 3: 1279. 1804.—Type: GERMANY, sine loc., c. fr. (syntypes: G-herb. Hedw.-Schwaegr:!). Fig. 17c-e. Hypnum fulvum Brid., Musc. 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 Mill. Hal., Hedwigia 37: 262. 1898.—Type: Cusa. “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 hyaline 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 um long, sometimes up to 35 um long; hyaline 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 um, 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 um in diameter, densely papillose. Chromosome number: n = 10 + x/y (in meiosis, North Carolina: 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. REPRESENTATIVE SPECIMENS. India. Kashmir: Ferozenpur Nullah, er R. R. Stewart 10572a, 2-9- ae MO): a: ssia. belo Garis Teberda, Abramov, 21-8-1955 Canada. New wick: Restigouche Co., St. Jean-Baptiste-de- ee R. Ireland 14478, 14-8- 1970 ce oe St. Barbe, Doctor’s Hill, R. Tuomikoski 4618, 29-7-1948 ee cea ewe Rvr., R. Tuomikoski 5412, 15-8-1949 (MICH). Nova Scotia: Annapolis Co., shor Keyimkuyjik Lake, a Ireland 12554, 3-8-1968 (MICH); Cumberland Co., Wentworth, i Ireland eal 7-9-1967 (MICH). Ontario: Grey Co., Sydenham Rvr., Inglis Falls, H. Crum 11034, 3- 8-1961 (MICH); Lake See Thunder Bay District, W. B. Schofield 27244, 1-7-1965 (as A. minor, E), W. end of Albert Lake, C. Garton 14878, 18-4-1972 (MICH). Québec: Pontiac corner of Pare de la Vérendrye, R. Ireland 16125 & T. Koponen, 15-6-1972 (MICH). Guatemala. Alta Verapaz: Rio Carcha, pr. San Pedro Carcha, P. ee 92104, 12-4-1941 (NY, MICH). Baja Verapaz: pr. Patal, A. Sharp 2947, 3-3-1945 (MEXU). El Quiché: Neboj, A. Sharp 2429, 6-2-1945 (MEXU, Jamaica. Arntully, C. R. Orcutt 3840, 12-11-1927 (DUKE). Mexico. Chiapas: San José 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, 0 3-1979 (NY). Durango: El Salto, A. Sharp 1889, 31-12-1944 (MICH). Hidalgo: Chapulhuacan, FE. Frye 2874a (MICH). Nuevo Leon: Hacienda Vistahermosa, 35 mi S of Monterrey, L. Harvey 1052, 1055, 25-6-1939 (MICH). Oaxaca: a S. Pablo Ayutla, A. Sharp et al., 23-12-1972 (MEXU). Puebla: Rio Necaxa, pr. Huachinango, A. Sharp 3143, 27-3-1945 (MICH). San Luis Potosi: Xilitlé, A. Sharp 5836, 21-4-1946 (US), C. Sat etal., 17— 12-1976 (MEXU). Sonora: Bavispe, Cafién del Temblar, -. Phillips 596, 19-8-1940 (MICH). aes pas: pr. Rancho del Cielo, Gomee Farias, C. Delgadillo, 1-5-1967, ibid. A. Sharp, 1-8-1974 (MEX Jalapa: “19 km NW Jalapa.” W. Rauh & R. Diill 31, 15-10-1966 (JE); NW of Caotepec, pr. ae lapan, L. Gil Juarez 60, 23-5-1975 (NY); gee Orizaba- me a Eggers & Frahm 38, 23-3 1979 (NY). U.S.A. Alabama: Hale Co., Rock Hollow, Havana P.O., H. Wagner et al. 71167, 30 pe 1971 (MICH). Ravonts Apache Co., Apache paeaeies 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: ae nee Upper Glen, W. Stair & W. Allen, 17-6- 1939 (MICH). Georgia: Stephen Co., Toc Falls, R. Lampton 1804, 15-5-1965 (MICH). pear LaSalle Co., French Canyon, Starved Roc k ae ton, Apr 1970 (MICH). pee Turkey Run Hollow, D. Richards et al. 5896, 23-11- 4947 (MICH). Kansas: Doniphan Co., Lege , 5. cab 10860, 30-6-1979 (MICH). Louisiana: Natchitoches Par., Cunning- ham Brake, J. se 643, 3-3-1978 (MICH). Maryland: Prince Georges Co., Beltsville, Paint Branch, PF. peas 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 ,2 mi N of Eagle Rock, P. Redfearn 26059, 4-5-1969 (as A. viticulosus, MICH); Franklin ae votes Rvr., B. Allen 6108, 30-1-1988 (MICH). ee Goodhue Co., B. Johnson, 3-5-19 (MICH). New York: Columbia Co., Old Chatham, S. Smith 3119 & B. Rey: 15-5-1961 (M Fain North Carolina: Transylvania Co., Horsepasture River, /. G. uae 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., I. G.-de la Cerda 1236, 8-6-1984 (DUKE), Nantahela River headwaters, I. G.-de la Cerda 1260, 11-6-1984 (DUKE), Crow Creek, I. G.-de la Cerda 1399. 19-6-1984 (DUKE); Orange Co., Eno River St. Park, 1. G.-de la Cerda 1999, Mar 1985 (DUKE). Pennsylvania: Hungtin- don Co., Spruce oon A. & P. Felton & I. G.-de la Cerda 1854, 30-9-1984, eres are ae Singlebranch Creek, L. Stair, a slg 1944 (MICH). Vermont: Wallingford, G. Kennedy, 3-8-1907 (MICH). Virginia: Madison Co., Fisher’s ne and Rose Rvr., . Schnooberger 4740 & ie 15- 6-1944 (MICH). West Virginia: ae r Co., H. Burtt, Tua 1971 (MICH). Wisconsin: Willburn, J. R6Il 1635, 16-12-1888 (as A. attenuatus var. Areviolias: G). Richland Co., Richland Center, M. Nee oe 13-6-1974 (MICH). Cuba. Pinar del Rio: Bafos San Vicente, Britton 7446, Aug-Sep 1910 (as A. wrightii, NY). Dominican Republic. ae orillas del Guanajuma, San Cristébal, J. Jiménez M-7, 16—-4- 1946 (NY). La Vega: Salto de Aguas Blancas, SW of Constanza, W. R. Buck 7900, 27-4-1982 (NY). Austria. Kiistenland: Gérz, K. Loitlesberger, May 1911 (c. fr., NY). Byelorussia. L. /. Savicz (NY). Denmark. Ganloese: “near Copenhagen,” W. C. Steere & rete 21-7-1954 (c. fr., NY). France. Haute Garonne: Prat de Joueou, Dismier 286, 3-7-1928 (MA). Haute Savoie: Douvaine, “Boisé Ballaison,” E. Bonnot 714, 31-12-1962 pete Germany. Oberayer: Bayerbrunn, Lewis, 9-11- 1845 (NY). Hungary. “Praeillycum, Sopianicum: Transdanubia Mts csek, Valle Nagymélyvoel- gy,” 1. Gyérffy, 25-7-1923 (BCB). Italy. * Sie ene: (NY); ‘ ve eee Monte Baldo” (ex herb. Thiimen NY). Novocomo: Flaglearo, supra Varese, F. A. Artaria, 114-1897 (NY). Norway. Akers- hus: “Emmestad in Vestby,” P. St¢rmer, Gare (BCB). Poland. “Western Carpathians”: Cieszyn, Dziegielow, Jedrzejko & Zarnowiec, 7-10-1981 (BCB). Kraké6w-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). fencint Fuente La Reina, Herrero-Borgofién 341, 16-5-1982 (MA). Girona: vall de Vidra, 7. G.-de la Cerda 2345 & F. Lloret, 28-3-87 (BCB). Huesca: Cafién de Anisclo, Fanlo, I. 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 manda, hayedo de Tobia, M. Brugués, 28-6-1977 (BCB) (see Granzow-de la Cerda, 1988a). Sweden. Uppsala: Mosén, Oct 1866 (NY). @land: St, Dalby, No. 974 3/6-8-1969 (MICH). Seitveiana: Zug, Bamberger (NY). Genéve: “bois de Frontenoy, prés de Genéve,” 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 in A. attenuatus also lacks the 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 Mill. Hal., Nuovo Giorn. Bot. Ital., n. ser., 3: 91. 1896.— Type: CHINA. “China interior, provincia Shen-si septentr.,” 1894, Giraldi, det. C. Miiller, 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-like 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 >11um long, 17 um 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) um 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 (/ocus classicus). 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). REPRESENTATIVE 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, FE. Uematsu, May 1908 (JE): 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE Diag 0.1 mm 0.1 mm FOS0EQ @ os. Sey a? a os, 10 ym G. 29. Anomodon giraldii Miill. Hal. a—c, branch leaves; d, leaf apex; e, cross section of stem, without differenciated central strand; f, cross sections of leaves; h, group of hyaline basal cells of the leaf; i, cells of costa at mid-leaf; g, cells of leaf margin near the apex. Based on: a, /watsuki 9574, Japan, “Kyushu, 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 austro-ussuriensis, fl. Ssputinka” Oct. 1934 (FH); e, Jishiba, Japan, Sendai, 7-9-1907 (DUKE); f-h, S. Murai, Japan, “Akita pref., Mt. Taiheizan, Ninbetsu Nat. Forest,” 1-4-1968 (NY). 258 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Miyadzu, Faurie 1518, 1901 (NY); Sendai, E. lishiba, 7-9-1907 (DUKE). Hokkaido: Ishikari distr., Sapporo, Tenguyama, 7. Koponen 21488, 8-07-1971 (herb. Chamberlain FH), Kamuikotan, A. Nogu- 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, 18-5-1977 (NICH), “Yunosawa valley,” K. Kakizaki, 27-7-1976 (DUKE); Ettyu, Mt. Kurobe, H. Sasaoka, 7-6-1930 (herb. Chamberlain FH); Gifu pref., Ono-gun, Nomugi-toge pass, /7. Katsuyama 13166, 25-5-1968 (NICH); Hiroshima pref., Iwaidani valley, R Watanabe 23496, 24—-8-1976 (NICH); Inaba, Mt. Kyushu-zan, H. Ochi, 17-4-1955 (herb. Bartram FH); Iwate pref., Mt. Aoba, S. Side s -7—1960 (JE), Shinoehei-gun, Iwaizumi-cho, R. Watanabe 17035, 23-8- 1972 (NICH), pete Iwate, G. Joba 224, 29-6-1928 (NY): Kyoto Oecho, Naigu, M. Mizutani, 7-10-19 WR “m 1E MICH); Mt. Komona, K. Mureta, 9-6— 1918 (herb, Sasaoka DUKE); Mie pref., ae Mt. Fujiwara-dake, H. Katsuwyama 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., Y. [kegami Mar 1948 (JE, MICH); Saitama pile: Chichibu- ey Ashi-ga-kubo, R. Wg ek 11-10-1976 (NICH); Shimane pref., lishi-gun, Akaana Shrine, M. Higuchi 13939, 17-5-1987 (c. fr., HIRO); Tot- tori pref., Mt. Daisen, 7. Koponen 21841, 20-07- 197] eee Chamberlain FH); Yamagesh pref., Iwakuni, Shiroyama, Schofield, rane & Seki 54512, 16-4-1973; * izand, Goka,” H. Sasaoka, Jul 1921 (herb. Brotherus FH); * a, Mt. Kotor,” H. Sk on i (herb. Brotherus FH); T yu, A. Noguchi, Jun 1932 hone Bartram FH). Kyushu: Bungo, Mt. Kudy, A. Noguchi, 4— wets - (herb. Bartram FH); Fukuoka pref., Asakura-gun, Y. pes oe 23-11-1951 (NY); Kumamoto pref., Mt. Fukabayama, . Takaki 2888, 23-4-1932 (NY), Kuma, Mt. Taiheizan, K. Mayebara, 4-4— 1952 (DUKE), Kohnox, K. Maebara, 3810 20-9-1955 (NICH), Mt. Aso, H. ee 1575, 25-4-1970 (NICH); Miyazaki pre net usuki-gun, Z. [watsuki 9574, 26-5-1981 (NY); Nagasaki, Wichura 1461 face A, pei iG a pref., Mt. Kurodeke, pr. Kakushimizu, Z. /watsuki 290a, 30-10-61 (c. fr., NICH). Shikoku: ae a Kami-ukena-gun, Tengu-Kogen platteau, Z. Jwatsuki 5245 & H. Kigwohi, 23/24—8-1978; Tokushima pref., Miyoshi-gun, Mt. Tsurugi, R. Watanabe 19/34, 22-8-1973 (NICH). Korea. Kong Quen 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), Mt. Sokri, W. Hong 2795, 2817, 21-12-1960 (NICH), Isl. Dagelet, W. Hong 6121, 6123, 26/29-7-1961 (NICH). sane = sf on ” Faurie 303, 1906 (NY). Russia. Ussuriisk, “Regio austro ussurensis: flumen Ssput ex herb. Verdoorn 254, Oct 1934 (FH, JE, MICH), ibid., L. Bardunov, 9- 9- ce pn oom see 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 Miiller with the type specimen. 16. Anomodon longifolius (Brid.) Hartm., Handb. Skand. FI. ed. 3, 300. 1838. Pterigynandrum longifolium Schleich. ex Brid., Mant. Musc. 128. 1819. Leskea longifolia (Brid.) Spruce, Musc. Pyren. n. 87. 1847. Hypnum longi- folium (Brid.) Mill. Hal., Syn. 2: 474. 1851.—Type: unknown. Fig, 30. atte armatus Broth., Hedwigia, 38: 243. 1899.—Type: JAPAN. Hokkaido, 29-4-1894, Oiwake, Mivabe & 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 pseudoparaphyllia, rhizoids abundant above (especially in Japanese 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 259 G { C\ wn OOD h SE ID M7 Poe NET My gj Fe ¥/ | Vase totah nase Popes aces Wk) pisces Rah RB HLL Vt ez cc iS C63 oat SSS= FIG. 30. Anomodon longifolius (Brid.) Hartm. a, branch leaves; b, surface of a leaf; c, branch leaf apices; d, e, portions of exostome; f, branch “bud” showing pseudoparaphyllia. Based on: a-f, from herb. J. Angtrém, Sweden, 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 cells 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 um); 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 um 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 um 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, ie Bakuriani, Lazarenko & Abramov, 5-9-1956 (NY). Georgia. “Tzenis eat Svania Dadia n, Hippi, inter Muri et Lentec,” Sommier & Levier, 29-7-1890 (NY). Japan. Asorezan, uate 2095, 1902 (c. fr., NY); Borezan, raul I1- i 1902 (herb. Bartram FH). Hokkaido: Kamuikotan, A. Noguchi, Aug 195] Se kuku, Mt. Hakkoda, E. Jishiba, Jul 1907 (MICH), Mutsu, 5. tas 31-7-1913 (NICH); Ishicari, Yamabe SS Twatsuki, 7-8-1971 (exicc. Noguchi 1202, MIC Y); Sapporo, Isiskaki, N. fe 4704, 16-10-1914 (NY); Si-Kari, Kamikawa-gun, Takenauti oe 0-03-1919 (NY). Honshu: Kominato, Faurie 49, 9-12-1889 (FI); Mt. Shirouma, N. Takaki, 1-08-1949 (NY); Fukushima pref., Otakine, Oniana, 7. 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 ri oe Mt. Mitoku, H. Ochi, 12-6-1960 (NY). . Markenstein, E. Be , 2-5-1869 (MICH). Wien, E. Berrover, 16-3-1869 (NY). eae. se Hallstadt, J. Prachi e 3- 1869 (NY). semen Avene en, Th. Suse, 12-8-1910 (ex herb. Uggla NY). Belgium. Frahan, Delogne 82, se 1869 (NY); ene sti me 1874 (NY). eee de Yvoir: Bauche, De oe over 30206, 1-4-1979 (NY). Byelorussia. , L. Savicz, 23- -1923 (NY). Finland. arg Pohjanmaa, Tervora, ee , TF. Ulvinen, 16-7 or pe ‘Ostrobot- — kajonensis”: Paltamo, M. Lindvquist, 18-7-1931 (NY). Pohjois-Saavo: Kuopio, pew. AL. Huuskonen, 3—9-19661 any). ‘Savonia borealis”: Kaavi, Likosaari, A. Terésvuori, 22-6-1917 (NY). Tohmajarvi, Havukkavaara, S of Vuorenpaa, R. Fagerstén & M. ee seed sn Wea = 10-1984 (as A. viticulosus, FH). Siilinjirvi: Iso-Pettéinen, Kourulah R. Fagerstén 776, 30-7 1974 (MICH). France. Haute Savoie: Reignier, Puejet, Nov oe poe Balonsdla a-Romaine, Es Sloover 35014, 25—7-1981 (MICH, NY). Jura: Jouhe, Foulotiére, P. ee 116, 5—5— io naregel Savoie: Aux-les-Bains, J. Miller (NY). Germany. Pr. Baierbrunn, P. G. Lorentz, 1860 (c. fr. herb. Uggla NY). Allgéu: Hinterstein, cee Bach, De Sloover ae 8-9-1973 (NY). have Berchtesgaden, Th. Suse, 6-9-1895 (ex herb. Uggla NY); Nagel fluhbléchen, Bayernbrunn, Mtinch A. Holler, Oct 1861 (NY). Mont Rhén: Geheeb, 1879 (NY). Thuringia: Miis oo i Hi. Moller, re: 1859 (ex herb. Uggla NY). Wartburg, V. Riisse (NY). Hungary Ceuat Pest, Madencia, pr. Ocsa, Boros, 14-10-1928 (NY). Comit. Bereg: pr. tarpa, A. Boros, 15- ee 1961 ccs. Comit. ae Mte. 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 261 Agasvar, pr. Nagybatony, poe 6-5-1928 (NY). Poland. Montes Gorce, Kamienica, Lisowski, 18— 4-1959 (Bryot. Polon. 1114 ne Kielce: Montes Gory, Swietokrzyskie, Lisowski 64989, 1-6-1964 (NY). Krosno: “W Bieszc on mnts., Kurnik6w Beskid,” R. one 7 7-1977 (BCB, MICH). Bialys- tok: Puscza Augustowska, ee cei 30-10-1958 (NY, MICH). Schlesien [Silesia]: Breslau, J. Milde (NY). Suwalki: Smolniki, Lake Jaczno, R. Ochyra, 17-5-1979 (MICH), Lake Haneza, R. Ochyra, 18-5-1979 (MICH). Russia. Gdbersdorf, Friedland [Pravdinsk], J. Milde, 30-7-186 oN Karelia: “Karelia ladogensis, Suistamo, Saarisuonvaara, J. Hueskonen, 24-8-1935 (NY); “Karelia onegen- sis, Uksjarvi, Kossalmi,” M. J. Kotilainen, 9-6— io (NY). Moscow : Ruza distr., Ozhigovo, M. [gna- tov, 11-6-1988 (MICE), Kunzowo, “pr. Mosquam,” Meyden, 224-1899 (NY). Kursk: Karotscha Dean, Lawdanski, 19-9-1902 (NY). “Sibiria,” Jenisei, Nikulina, H. Arnell, 18-7-1876 (NICH). Spain. Bar- celona: Sta. Maria de ee C. Casas, Dec 1964 (BCB). Navarra: Sierra de Urbasa, A. Ederra, 13- 3-1982 (BCB, NAU). Sw n. Angermanland: Sabra, Grovhall, H. W. Arnell, 1-6-1870 (NY). Narke: Svenshyttan, C. ease a Fie 1874 (NY). Oland: Hogbyn, Vyksj6, Hj. Méller, 25-7-1917 (DUKE), Mariehamn, H. Moller 7-1928 (NY). Vestetad Lund, K. Holmen, 8-6-1969 (NY). Ostergétland: Marmosbruket, P. a on 6-1875 (NY). Sala: “sub Pterog” (c. fr., ex herb. Sullivant NY). Skane: Oveds su, Fruelid, G. Alberg, 15-6-1919 (ex herb. Ugela NY). Smaland: Greunabruket, Vretaholm, A. Arvén, 18-5-1914 (NY); Talberg, N. Schentz (herb. Thittmen NY). Torshunda, H. Moéller, Apr 1859 (herb. aoc NY). Uppsala: Gottsundabergen, J. Zettersstedt 95, 7-5-1878 (MICH), Uppsala, 1844 (herb. en we Vistmanland: Sala Gruva, 31-5-1879 (DUKE). Switzerland. Pr. Ziirich, P. Culmann, Wake 5 (NY). Valais: Loneche- les-Bains, H. Fossell, Sep 1887 (NY). Zug: Prope Zug, Bamberger 6522, 1855 (NY). Ukraine. pr. Kiev, HoLauee: N. Wagner, 16-3-1919 (NY). United Kingdom. G ee 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 pseudoparaphyllia (shared only with A. rugelii), 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 delicate branches, with leaves 1.2-1.6 mm long, julaceous, and more imbricate when dry. Anomodon armatus Broth., described from Japan, coincides in every feature with western material and is here included in A. longifolius. Herpetineuron (Miill. Hal.) Cardot, Beih. Bot. Centralbl. 19(2): 127. 1905. Ano- modon sect. Herpetineuron Mill. 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]. Plants rather robust, forming open to thick mats. Stems prostrate, sparsely branching; branches rather arcuate, circinate 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 elliptic 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 1-4 cells high, segments 0.2—-0.4 mm high, papillose. Calyptra smooth. Chromosome number unknown. Herpetineuron 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 trom 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 I. Costa strongly flexuose-sinuose in the distal 2/3 of the lamina; lamina smooth ai ce . H. toccoae. 1. Costa straight throughout; back of the distal 1/2 of the lamina with prorulate cells. 2. H. acutifolium. 1. 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.— Synrypes: Cuina. Hong Kong, Bowring; INp1A. 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 $. 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 um 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, 4. 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: J. C. Bowring (as Anomodon devolutus, syntype, herb. Mitten Be ee oon, Panlong wan, Y. W. Taam 2007, 9-6-1941 (DUKE); New aimoshan, Chunloong, Y. W. Taam, 1948, 16-3-1941 (c. fr., MICH). India. Kumaon, J. pie 6367, 22-8-1886 (MICH). ee cine 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. sts 3046, 20-10-1907 (DUKE). Miyazaki pref., Nao, S of Nichinan, W. se ad igi 3, 9-1- 1973 (DUKE); Oita pref., Takeda-machi, A. Noguchi, 1-12-1947 (DUKE). Japan. Goto. Fai 596 1901 (as e wichurae, NY). P Higo: Konase, Kuma-gun, K. Mayebara 96, 14-10-1928 (as H. es NY). Honshu: Mie pref., ee “fall N. Takaki 529] (as H. Ses NY). 264 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Brazil. Santa Catarina, Bom Retiro, BR 282, Schdfer-Verwimp 10483 & Verwimp, 21-12-1988 (MICH). Rio de Janeiro, Glaziou 9155, 9216, 9222 (as Anomodon brasiliensis, NY). Guatemala. Huehuetenango: W of Aguacatan, P. Standley 8/215, 27-12-1940 (MICH). Honduras. Comayagua: Rio Selguapa, 4 km SE of Siguatepeque, B. Allen 1/800, 11-5-1992, 13403, 2-3-1993 (MO). Olancho: nr. Catacamas, R. Olson 83-8, 22—9-1983 (MO). Nicaragua. Esteli: Salto de Estanzuela, 6 km S of steli, D. Stevens 11754, 1-10-1979 (MO). Mexico. Chiapas: “above Las Casas,” A. Sharp 3365, 20- - 1945 (MICH). Chihuahua: es Hf. LeSueur A4b, 20-8-1937 (MICH). Durango: El Salto, A Sharp 1895a, 31-12-1994 (MICH). CmerierG: Omiltemi, 30 km W of Chilpancingo, A. Sharp 1135b, 27-10-1944 (MICH). ss La Fer above Manatlan, H. Crum 984a, 790, Jul 1949 (MICH). México: Temascaltepec, Ixtapan, G. ee 2937a, 14-12-1932 (MICH). Morelos: ee Tepozteco, C. Delgadillo, 28—9- he (MEXU). Nuevo Leon: elena Madre Or., Monterrey, SW of El Cercado, F. Hermann 26082, (MICH). Oaxaca: Tamazulapan Mixé, A. Sharp 2990 et al., 23- 1972 (MICH). Puebla: “below Mecana? Rio Tenango, A. Sharp 3124, 26- 3-1945 (MICH); Rio Necaxa, pr. Huanchi- nango, A. Sharp 3142, 27-3-1945 (MICH). San Luis Potosi: Sapa Sharp 5911, 22-4-1946 (MICH). Sinaloa: W of Las Palmitas, D. Norris 20358 et al., 25-5-1971 (MICH). Sonora: pr. El Tigre, Rancho del Roble, 8S. White 4229, Sep 1941 (MICH). Tamaulipas: Sierra de Guatemala, S$ of Rancho del Cielo, H. Crum 1664, 1690, Aug 1950 (MICH). U.S.A. Alabama: Winston Co., Bankhead Nat. For., W. ee 21869 (MICH, NY). Arizona: Sycamore Canyon, nr. Ruby, 1. Haring 3297 & H. Skell, 11-3- : . Arkansas: Cleburne Co., nr. Greer’s Ferry ee P. Redfearn 18729 & F. Bowers, 12-3- 1966 Ae. NY). Garland Co., Hot Springs Nat. Pk., L. Anderson 11347, 5-10-1953 (DUKE, MICH). ney lee Co., J. Small, 8-8-1893 (G, DUKE, MICH), L. Anderson 22078, 28-7- 1975 (DUKE. CH). Illinois: Union Co., Panter’s Den, P. Redfearn 23407, 24-1-1968 (MICH). Louisiana: res Par., Tangipahoa Rvr., E of Bonchatonla: W. Reese & G. Landry 17011, 30- 10-1986 (as ee deal triste, MICH). West Feliciana Par., pr. St. Francisville, W. Reese 10843, 3-4-1969 (MICH). Mississippi: Oktibbeha Co., S of Starkville, $8. McDaniel 5542, 4-1-1965 (MICH, NY). North Carolina: Sak ell Co., Linville Caverns, L. Anderson 11024, 14-7-1952 (DUKE, MICH). Jackson Co., Whitewater Falls, 1. G.-de la Cerda 1297, 13-6-1984 (BCB). Tennessee: Sevier Co.., trail from Greenbrier to Brushy Mt., A. Sharp & P. Voth 349, 29-3-1936 (MICH). Dominican Republic. Aztia: Canada Miguel Martin, M. Ven & T. Zanoni 8267, 189-1980 (MO). Herpetineuron 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 Herpetineuron (“snake costa”); however, the second species lacks a sinuose costa. 2. Herpetineuron acutifolium (Mitt.) Granzow, Bryol. 92: 385. 1989. Anomodon acutifolius Mitt., J. Linn. Soc. Bot., suppl. 1: 126. 1859. Bryonorrisia acuti- folia (Mitt.) Enroth, J. Bryol. 16: 407. 1991.—T ype: INp1a. Uttar Pradesh, Kumaon 6000, Sarkito Shayuk valley, 800 (lectotype, designated by Enroth, 1991: NY-herb. Mitten!). Fi Forrstroemia secunda Dixon & Badhw., Rec. Bot. Surv. India 12: 176. 1938, Bryonorrisia secunda (Dixon & Badhw.) Stark & Buck, Brittonia 38: 52. 1986.—TyPeE: INp1A. “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, |.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 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 265 pe) sang ash S35 ORO) SSorcan ROCHE p AS x i PHOS | Wi ERIS Ni DAR AAI |) Ye) Ue )\ Wen 7 aR A 1 ] :" ZN AS Ba SSS 32 Bra 3 2QOAHANSe f°) ORO f foPyAGg by OS 0 °<8 e Boy O85 ) )d Seo Oe Oat OveAvLod Grove Mosca sO Nose wy 2 08%) 1 ny a : es : Hs FIG. 31. Herpetineuron acutifolius (Mitt.) Granzow. a, b, c, branch leaves; d, e, apex of branch leaves: f, cross section of stem with differenciated cells in the center; g, h, branch leavers in cross section; i, portion of peristome with well-developed endostome; j, spores. Based on: a, f, h, Falconer & Royle, India, NW Himalaya (NY); b, d, g, j, Koeltz 7529, India, Punjab, Kangra, Bathad, (FH); c, e, i, Strachey & 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 unknown. 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) a ee Kangra, Kulu, Bathad, W. Koeltz 7529, 24-11-1933 (FH). Uttar Pradesh: Kumaon, Naintal, A. Norkett 11851, 4-10-1966 (as Herpetineuropsis secunda, NY); Kumaon, above Jallet, 7500’ coe & Winterbottom 70/13 (syntype of H. acutifolium: NY-herb. Mitten!), Kathi 7000’, Siachens & Winterbottom ie (syntype of #7. a ad ae herb. Mitten!); Rampur-Bushahr, Dorkali, W. Koeltz 3185d, 29-11-1931 (NY); “NW Himalaya, ee & Royle (NY). Japan Honshu: Saitama pref., rats 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. /watsuki ie aren (NY, HIRO). Katmandu: Mt. Phulchoki, “summit area,” Z. [watsuki 2670, 15—7-1972 (N ex as Chihuahua: “51 mi of Creel,” La Junta to Creel, F. Bowers et al., ae a toe CWitht Af, toccoae), 4-1-1974 (MEXU); “30 min. N de San Juanito” C. Delgadillo, fe i. 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 Hf. 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. EXLUDED NAMES Anomodon californicus Lesq., Mem. California Acad. 1: 30. 1968. = Triquetrella californica (Lesq.) Grout. Anomodon brasiliense Hampe, Vidensk. Meddel. Dansk Naturhist. Foren. Kjaben- 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. Musc. 2: 292. 1878. Anomodon lagoensis Hampe, Vidensk. Meddel. Dansk Naturhist. Foren. Kjaben- havn, ser. 3, 2: 238. 1870. = Leskea lagoensis (Hampe) Gier, J. Bryol. 10: 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE 2G) 211. 1978. = Pseudoleskea subcatenulata Besch., Mém. 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. Linképing. 1896. = Leskea moseri Kindb., Ottawa Nat. 7: 19. 1893. = Leskeella nervosa (Brid.) Loeske, Moosfl. Harz. 255. 1903. Anomodon nervosus (Brid.) Hiib., Musc. Germ. 561. 1833. = Leskeella nervosa (Brid.) Loeske, Moosfl. Harz. 255. 1903. Anomodon perarmatus Williams, Bryol. 31: 112. 1928. = Philonotis scabrifolia (Hook. f. & Wilson) Braithw., Brit. Moss Fl. 2: 215. 1893 (see Granzow- de la Cerda, 1989a: 383). Anomodon scaberrimus Broth., Bih. Kongl. Svenska Vetensk.-Akad. Hand. 21, Afd. 3: 65. 1895. = Macrocoma scaberrima (Broth.) Vitt, Bryol. 82: 16. 1979. Anomodon tasmanicus Broth., Ofver. Finsk. Vet. Soc. Forh. 42: 137. 1900. = Tri- guetrella curvifolia Dixon & Sainsb., J. Bot. 71: 217. 1933. = Triquetrella tasmanica (Broth.) Granzow-de la Cerda, Bryol. 92: 383. 1989. Anomodon xanthophyllum Hampe, Bot. Zeit. 26: 816. 1868. = Stereophyllum mex- icanum Williams, Bryol. 26: 34, pl. 4. 1923 (fide Ireland, 1982). = Entodon- topsis mexicana (Williams) Buck & Ireland, Nova Hedwigia 41: 105. 1985. Haplohymenium densum Schimp., Mém. Soc. Sci. Nat. Cherbourg 16: 233. 1872. = Leskea mexicana Besch., Mém. Soc. Sci. Nat. Cherbourg 16: 233. 1872. = Lindbergia mexicana (Besch.) Cardot, Rev. Bryol. 37: 51. 1910. TAXA NOT AVAILABLE FOR STUDY Anomodon grandiretis Broth., Akad. Wiss. Wien Sitzungsber., Math.-Naturwiss. KIL, Abt. 1. 133: 578. 1924. 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). Haplohymenium magnirete Sakurai, Bot. Mag. (Tokyo) 64: 196, Ic, d. 1951. ACKNOWLEDGMENTS This research was partially supported by a postdoctoral fellowship from the Comissié Interde- partamental de Recerca i Innovaci6é Tecnoldgica (CIRIT) of the Conselleria de Ensenyament, Gen- eralitat de Catalunya. The work was mainly carried out at the Laboratori de Briologia, Facultat de Ciéncies, Universitat Autonoma de Barcelona, and also at the Department of Botany, Duke Univer- sity. | am thankful above all for the understanding and unconditional support given by Creu Casas, Montserrat Brugués, Rosa Maria Cros, and Francisco Lloret, all of the Universitat Autonoma de 268 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Barcelona (BCB), and to Lewis Anderson for his comments on the manuscript. The very valuable comments and advice of William Buck, and the help of Brent Mishler and Efrain de Luna regarding the phylogenetic analysis were of enormous help. I am greatly indebted to Molly McMullen and to the staffs at MICH and the Real Jardin Botanico of Madrid for their help with loans, and especially to Mariano Sanchez for his ie with the line drawings, Gonzalo Nieto for his comments, Daniela Gobetti and Michael Bonner for checking the Latin diagnoses, and to the curators and assistant curators of pias sk geo mB, BCR, C, COI, DURE, FH, FL-G, FH, HIRO, FP) INA, JE, L, LISU, MA, MA H, MO, NAU, NICH, NY, PO, RO, TENN, US, WELT. I thank Howard Crum ae eer ae for reviewing the manuscript, as well as William and Christiane Anderson for all their support and patience. LITERATURE CITED Akiyama, H. 1988. 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Die europaischen ee ite Arten mit besonder PB CHUCRSCUEUNE ihrer R userpic Verbreitung und ihrer Verwandtschaft. Hedwigia 76: 191-298 Sainsbury, G. O. 955. A handbook of the New Zealand mosses. Roy. Soc. New Pealnad Bull. 5: 1-409. 200) CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Sakurai, K. 1954. Muscologia japonica. Tokyo. Sanderson, M. J., and M. J. Donoghue. 1989. Patterns of variation in levels of homoplasy. Evolution 1-1795. Schimper, W. P. 1860. Synopsis muscorum europaeorum. Stuttgart. Schumacker, R., P. De Zuttere, and L. Leclerq. 1982. Anomodon rostratus (Hedw.) Schimp. (Thuidi- aceae), new for the Belgian ervatlora, in the south of the Ardenne massif (Rochehaut, prov. Luxembourg, Belgium). J. Bryology 12: 171-177. Smith, A. J. E. 1978. The Se eed Britain and Ireland. Cambridge: Cambridge University Press. Stark, L. R., and W. R. 986. Corrections to Forsstroemia including Bryonorrisia, gen. nov. | aes 38: vee 4. Sullivant, W. S. 1864. [cones Muscorum. Cambridge, Massachussets: Sever and Francis. Swotford, D. L. 1990. PAUP: Phylogenetic analysis using parsimony, version 3.0r. Computer program ap ene es the Illinois Natural History Survey, Champaign, linois. n, B. C., D. E. Bouf “he ., Liu B., Wang Z., X1 Y., Ying T., Zhang C., and Zhang X. 1996. A aids to the moss flora of Henan province, China. Acta Bot. Yunnanica 18: 67-71. Vitt, D. H., 1984. Classification of the Bryopsida. In New manual of bryology, ed. R. M. Schuster, 2: 696-759. Nichinan: Hattori Botanical Laboratory. Vitt, D. H., oe P. Lee. 1984. Anomodon minor (Musci: Leskeaceae) in North America. Bryologist 87: 338-336 ae | i A revision of the family Thuidiaceae in Japan and adjacent areas. J. Hattori Bot. Lab. 36: 171-320. Wik, R. van der, W. E. Margadant, and P. A. Florschiitz. 1959 Index Muscorum, Vol. 1. Regnum Veg. 17: 1-548. 17 ———.. 1969. Index Muscorum, Vol. 5. Regnum Veg. 65: 1-922. Yano, K. 1957. Cytological studies on Japanese mosses. Memoirs Takada Branch, Niigata University 1: 85-127. APPENDIX | CHARACTER CODING The list of characters used in the cladistic analyses is given below, with an explanation for each of the states coded. The two species of Herpetineuron and Pterigoneuron gracile were used for character polarization. Undetermined states (ae as *?”) represent one of the three following scenarios: the state is not known for a particular taxon; the character does not apply (i.e., the taxon lacks the structure which is referred oy or the taxon shows polymorphism for a given character (when not resolvable by including more than one state in the matrix, using the notation, for example, “1&2”), Whether the multiple-state ees were ordered or not is indicated for each. Character 11 was “partially ordered” using a predefined state matrix. ranching pattern (unordered; Fig. 1). 0 = Primary stem with simple branches in a loose erste arrangement. 1 = Primary stem with secondary branches bearing tertiary branches in a pseudofasciculate fashion, tuftlike; the tertiary branches are erect and arise from the secondary pine ies very close to each other, appearing whorled (e.g., A. /ongifolius and A. rostratus). 2 = As in sta . but also with the secondary branches bearing branches, usually arcuate, of several successive ne in a stairstep ae (e.g., A. attenuatus i giraldit). 2. Attenuate and/or flagelliform branches. 0 = Absent. | = Present. 3. Cross section of the stem. 0 = Central vas of conspicuously smaller cells well differentiated. Le ee stem seg 2 cross sree without a differentiated central strand of smaller cells. Ba aes ei aca = Absent. | = Present. 5. Branch leaf Ai aa dry ee 0 = Erect to julaceous. 1 = Secund. 2 = Conspic- se incurved. 6. Orientation of branch leaves when moist (ordered). 0 = Not flattened on either the ventral or the aes side of secondary branches. | = Flattened only on the dorsal surface of secondary branches, while secund and pointing to the substrate on the ventral surface. 2 = Flattened on both surfaces, especially towards the end of ea branche eaf constriction. 0 = eS constriction; leaves lanceolate to triangular-ovate (e.g., A. giraldit). 1 = eee oa present (“shoulders”), more or less abruptly dividing the leaf into a broad lanceolate proximal region and a narrow ata one. 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE Dag 8. Leaf length. 0 = Small: <0.7 mm long. | = Large: >1.4 mm lon 9. Shape of the distal region of the lamina of branch leaves, above the shoulders (Fig. 2). 0 = Tapering, i.e., with margins never parallel but progressively narrowing between shoulders and apex. | = 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. viticulosus the character presents a intone within mats. 10. Leaf margin. 0 = Plane throughout. 1 = Revolute in part. 11. Shape of branch leaf apex (partially ordered; Fig. 3). 0 = Leaves tapering into a multiseriate subulate apex, in A. longifolius. 1 = ae 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 a 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; therefore, it is not at all similar to that of A. rostratus. f margin near the apex. 0 = Entire, although sometimes slightly crenulate due to the presence of papillae. 1 = With teeth. Transformation Series (Character Tree) Step Matrix O< Sr Dc So < >4 0 I Z, 3 4 A 0 : I 2 - 2 1 | - Vv 3 2 2 1 - 1 4 3 3 2 1 - 13. Length of the costa (ordered; Fig. 4). 0 = Not reaching beyond the lower 1/3 of the leaf lamina. | = Reaching to 2/3-3/4 of the length of the leaf. 2 = Reaching ca. 9/10 of the leaf length to subpercurrent. 14. Costa bifid near the apex (Fig. 5). 0 = 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. 0 = 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 abov 16. Papillae on cells of dorsal surface of the costa (unordered; Fig. 6). 0 = Cells smooth or with very discrete vesicle-like bumps. 1 = Papillae minute, numerous and randomly arranged on each cell (e.g., A. giraldii). 2 = 4-7-seriate, tall papillae per cell, not branched. ? = Cannot be scored for Anomodon tristis because of its small size and obscured costa. apillae on basalmost margin cells, at the insertion of the leaf (ordered). 0 = Lacking papillae, or Earth 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 oem papillae, larger than those on laminal cells, usually single and sometimes branched, resembling spin 18. Shape of leaf cells, at least those toward the middle of the lamina. 0 = Elongate: ee or rectangular. 1 = Isodiametric: hexagonal, quadrate or round. 19. Leaf cell papillae (unordered). 0 = 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 ce 4= Multiple, scarcely to strongly branched papillae. 20. Position of gametangial buds relative to most recent (distal) branching points (ordered; Fig. 7). 0 = The most recent gametangial buds appear distally, relative to the last branching points (e., perichaetia or perigonia develop on the youngest branches). 1 = Only flagelliform branches are produced beyond gametangial buds (e.g., A. longifolius and ae oe ee 2 = Gamet- angial buds never found distally relative to latest branching points but only on older branches. 21. Cells of ar leaves. 0 = Smooth throughout. 1 = Papillose, at ee near the leaf apex. 22. Length of seta (ordered). 0 = >4 mm. 1 = 1.5-3 mm. 2 = <1 mm (capsule immersed). ? = a ab unknown for A. en and x eS eae ength of urn. 0 = <1.1 mm. 1 = = Sporophytes are unknown for A. dentatus and A. ea length is highly ees in w nee 7 rt) pap CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 24. Stomata. 0 = Present. | = Absent. ? = No information is available for A. dentatus, A. solovjovu, and A. foe last Aelia 25. Ann . 0 = Well developed. | = Poorly differentiated. ? = Sporophytes unknown for A. dentatus and ma Vian. 26. Exostome structure. ne = Teeth more or less reduced, irregular or incomplete, <0.20 mm long. 1 = Teeth splitting more ss symmetrically, >0.25 mm long. ? = Polymorphic in A. pseudotristis; unknown for A. dentatus ian . flagelliformis. 27. Striola Scat ae rks at base of exostome. = = Absent. | = Present. ? = No information is available for A. dentatus, A. flagelliformis, and A. giralc 28. Papillose ornamentation at the base of exostome. 0 = Absent. 1 = Present. ? = No informa- tion is available for A. dentatus, A. giraldti, and A. flagelliformis, and there is a polymorphism in A. ae art } . Ornamentation at upper part of exostome teeth (unordered). 0 = Verrucose, with thick ee unevenly arranged. | = Papillose, with papillae evenly distr ea 2 = Smooth or finely rugose. ? = Sporophytes are unknown for A. dentatus and A. flagelliformis. p a 30. Segments of endostome (ordered). 0 = Absent or almost absent (0-1 cells high). | = Present but short (3 or more cells high). 2 = Well developed, almost as high as exostome teeth. ? = Sporo- phytes are unknown for A. dentatus and A. flagelliformus. 31. Calyptra. 0 = Glabrc | = With a few long and scattered hairs. ? = Sporophytes and calyptra are cee for A. ene and A. flagelliformis. LIST OF SYNONYMS Anoectangium ferrugineum Besch., Ann. Sci. Nat., sér. 7, Bot. 17: 329. 1893. = A. rugelii. Anomodon aculeatus Broth, & Paris, Rev. Bryol. 31: 56. 1904. = A. longinervis. A. acutifolius Mitt., J. Linn. Soc. Bot., suppl. 1: 126. 1859, = H. acutifolium. A, apiculatus B.S.G., Bryol. Eur., fasc. 44-45. 1850 (nomen nudum) = A. rugelii. armatus Broth., Hedwigia, 38: 243. 1899. = A. longifolius. _ aspertfolius Mill. Hal., Nuovo Giorn. Bot. Ital., n. ser., 3: 117. 1896. = A. abbreviatus. . attenuatus var. brevifolius Ren. & Cardot, Hedwigia 32: 302. 1893 (nomen nudum) = A. attenuatus A. attenuatus. brevinervis Broth., Ofver. Finsk. Vet. Soc. Férh. 33: 107. 1891. = A. preudotristis. decurrens Cardot, Bull. Soc. Bot. Genéve, sér. 2, 3: 281. 1911. = A. viticulosus. devolutus Mitt., J. Linn. Soc. Bot., suppl. 1: 127. 1859. = H. toccoae. exilis Mitt., J. Linn. Soc. Bot. 13: 309. 1873. = A. preudotristis. ferrugineus (Besch.) Nog., J. Jap. Bot. 20: 260. 1944. = A. rugelii. filiformis Thw. & Mitt., J. Linn. Soc. Bot. 13: 308. 1873. = A. preudotristis. fragilis Wilson, London J. Bot. 7: 275. 1848. = A. tristis. . fragillimus Herzog, Beih. Bot. Certralbl. 26: 82. 1909. = A. tristis. glossophyllus Mill. Hal. in Paris, Ind. Bryol. suppl. 14. 1900 (nomen nudum) = A. minor. A. huttonii Mitt., J. Linn. Soc. Bot. 13: 309, 1873. = A. preudotristis. A. integerrimus Mitt., J. Linn. Soc. Bot., suppl. 1: 126. 1859. = A. minor. A, leptodontoides Mill. Hal., Nuovo Giorn. Bot. Ital., n. ser., 4: 275. 1897. = A. minor. A. microphyllus Broth. & Paris, Rev. Bryol. 31: 56. 1904. = A. tristis. A. minor (P.-Beauv.) Lindb., Bot. Not. 1865: 126. 1865 (nom. illeg.) = A. minor. A. minor subsp. infegerrimus (Mitt.) Z. Iwats., J. Hattori Bot. Lab. 26: 41. 1963. = A. minor. A. minor var. inaequalifolius Bartr., Bryologist 50: 207. 1947. = A. minor. A. mithouardii Paris & Broth., Rev. Bryol. 29: 97. 1902. = A. preudotristis. A. obtusifolius Bruch & Schimp., London J. Bot. 2: 668. 1843 (nom. superfl.) = A. minor. A. obtusifolius Mitt. (nomen nudum) [non A. obtusifolius Bruch & Schimp.]| = A. viticulosus. bs BeBe SP oe ee ee A. obtusifolius var. w Mitt., (nomen nudum) = A. thraustus. . planatus Mitt., J. Linn. Soc. Bot., suppl. 1: 126. 1859. = A. minor. A, platyphyllus Kindb., Ottawa Naturalist 7: 19. 1893. = A. minor. > APPENDIX 2. Data matrix used for the in Appendix 1. cladistic analysis: 19 taxa, 31 characters. (ord.) = unordered character; (p. ord.) = partially ordered, s tep matrix given Characters Taxa 1 2 3 4 De Ove k tl iy SON ae, LOSS G2 aS ala i Gre SS ee sl9s S208 22 3 and ee ae O Re Oia S20 08 BOUL ol A. viticulosus 0 0 | 0 1 0 | 1 0&1 0 3 0 2 | OP ee? 1 1 4 0 | 0 1 I 0 0 0 2 1 i) A. rugelii 0 0 1 1 22 al | 1 1 0 4 O 2 1 On eee 2 1 4 0 | 0 1 0 l 0 0 1 2 0 0 A. minor 0 0 0&1 0 Or 2 1 1 1 OPV 5a4 eNO ee? l O&1 2 1&2 1 4 0 L 0 1 i 0 0 ) 1 l 0 0 thraustus 0 0 0 0 Orert0) 1 1 1 0 4 #4O i OL 2 22 1.0) 1 4 0 I ) 1 1 0 1 0 1 1 1 0 A. dentatus 0 0 1 0 0 0 1 1 I 0 3 l 2. 10ha* 2 0 O 1 4 0 I ? z ? ll Be 2 ? 2 i ? abbreviatus 0) 0 I 0 Le 20): 1 1 1 0 3 Ope 2 FeSO 40s re ee 0 1 1 0 I 1 1 1 0 1 0 0 0) 0 0 A. solovjovil 0 0 ! 0 | 0 1 1 1 0 4 0 2 0 1 0 0 1 | 0 0 2 0 ? 0 1 0 0 1 0) 0 A. rostratus 1 0 0 0) 0 0 1 Ove0 | | 0 | O&1 1 2 Oy OE T4030 5 10a) 0 0 0 1 l 0 l 2 0 A. attenuatus a) 1 | 0 | 2 l l 0 l 3 I 2: 1 1 0 1 1 4 2 1 0 1 0 i l l 0 1 2 0 A. giraldii 2 1 | 0 LrenOns 10 1 ? | Poe alle Gua ee) 1 1 Of 302-3 2 0 0 2 0 1 1 ? 2 1 0 0 A. longifolius | 1 0 1 163 2.0) l 1 0 | On ea0s e222 520 1 Ores Os © OS 22 | OF) 220 1 1 1 0 1 ? ) 1 0 Ha. triste 0 0 | 0 Ory 0 1 OURO sO gee OCS 0s Sal se ee 0 O 1 4 0 | 1 0 1 0) 1 0 0 0 0 1 Ha. sieboldii ) l 0 0 0 1 OTP ri) tess One seen kOe eC) ea 0 O 1 4 2 1 0 0 1 0 1 ) 0 0 0 1 Ha. pseudotriste 0 | 0 Oe 2: 1 0 0 O&1 3 0 0 0 2 0 0 1 4 0 I i 0 | 0 ? 0 0 0) 0 1 Ha. flagelliforme 0 1 | ] Ore O08 at Ob, XO ORs ars? 1 ! OF? 0 0 1 5 0 | 2 ze ? ? ? Z ? ? ? 2 Ha.longinerve 0 0 I 0 0 1 1 OR OF Oss 3 Fei) 1 OS Seay eer a6) 1 l 0 L 0) 0 1 0 l 0 0 0 0 1 He. acutifolium = 1 1 0 0 One: 0%es <0. 1 ee) 8 ay 1 De Ose aOnea Ors 02e4 04720 1 OPO 1 0 0 0 l 1 2 2 0 He. toccoae 2 1 0 0 ibe <0) se 40) L [eee Oe) I 2 atm RO ee 0 ess Sr) Oe 505 enix) 1 0 0 0 0) 1 0 2 0 P. gracile 1 1 0 0 OF 0-5 0 Il tee Od pes) 1 0 ? Or 20%. 20 202 F0l" 0, 202 Fa0 l 0 0 l 1] it 1 1 0 1. branching pattern ll. leaf apex (p. ord.) 21. perichaetial leaf cells papillose 2. flagelliform branches 12. margin outline near apex 22. length of seta 3. cross section of stem 13. costa relative length (ord.) 23. length of urn 4. pseudoparaphyllia 14. costa bifid at apex 24, stomata 5. leaf arrangement when dry (ord.) 15. costa fading 25. annulus cells differentiated 6. orientation of leaves when moist (ord.) 16. papillae back of costa 26, exostome structure 7. leaf constriction 17. basalmost If margin cells (ord.) 27. base of t triolat 8. leaf length 18. shape of leaf cells 28. base of exostome papillose 9. shape of leaf distal portion 19. leaf cells papillae 29. ornamentation of upper teeth 10. margin section 20. relative position of gametangial buds 30. segments 31. calyptra L661 AVAHOVLNOGOWONY ‘VduesD WT AG-MOZNVUYD 274 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 A. plicaefolius Dixon (not validly published) = A. minor. A. ramiutosus Mitt., Trans. Linn. Soc. London, ser. 2, 3: 187. 1891. = A. minor. A. rubiginosulus Mill. Hal., Flora 73: 496. 1890 (nom. superfl.) = H. toccoae. A. rugelii var. ferrugineus (Besch.) Z. Iwats., J. Hattori Bot. Lab. 26: 51. 1963. = A. rugelil. A. sinensi-tristis Mull. Hal., Nuovo Giorn. Bot. Ital., n. ser., 3: 118. 1896. = A. tristis. A. sinensis Miill. Hal., Nuovo Giorn. Bot. Ital., n. ser., 3: 118. 1896. = A. minor. A. solovjovii var. henaensis Tan, Boufford & Ying, Acta Bot. Yunnanica 18: 69. 1996. = A. solovjovii. A. stenoglossus Cardot & Thér., Bull. Acad. Int. Géogr. 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. toccoae Sull. & Lesq., Musci Bor. Amer. 52. 1856. = H. toccoae. A. tonkinensis Besch., Bull. Soc. Bot. France, sér. 3, 41: 84. 1894. = A. viticulosus. A. uematsui Broth., Ofver. Finsk. Vet. Soc. Férh. 62: 33. 1919-1920. = A. rugelii. A. uyematsui Broth., Bull. Soc. Bot. Genéve, sér. 2, 3: 281. 1911. = A. rugelii. A. wichurae Broth., Hedwigia 38: 244. 1899. = H. toccoae. A. wrightii Mull. Hal., Hedwigia 37: 262. 1898. = A. attenuatus. Bryonorrisia 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. Genéve, sér. 2., 3: 281. 1911. = A. longinervis. Af. brachycladum §. Okamura, Bot. Mag. Tokyo 25: 31. 1911. = A. sieboldii. HA. brevinerve (Broth.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907. = A. preudotristis. H. crassum Sakurai, Bot. Mag. Tokyo 47: 337. 1933. = A. longinervis. H. cristatum Nog., J. Jap. Bot. 20: 146. 1945. = A. flagelliformis. H. fasciculare Nog., J. Jap. Bot. 13: 791. 1937. = A. preudotristis. H. filiforme (Thw. & Mitt.) Broth., Nat. Pflanzenfam. 1(3): 1236. 1909. = A. preudotristis. H. flagelliforme Savicz-L., Bot. Mater. Inst. Sporov. Rast. Glavn. Bot. Sada RSFSR. 1: 98. 101. 1922. = A. flagelliformis. H. formosanum Nog., Trans. Nat. Hist. Soc. Taiwan 26: 43. 1936. = A. tristis. H. fragiliforme Nog., J. Jap. Bot. 13: 410, f. 2. 1937. = A. tristis. H. gonoi Broth., Ofver. Finsk. Vet. Soc. Férh. 62: 33. 1919-1920. = A. sieboldii. H. huttonti (Mitt.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907. = A. preudotristis. H. longiglossum P. C. Chen, Feddes Repert. Spec. Nov. Regni Veg. 58: 31. 1955. = A. tristis. H. longinerve (Broth.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907. = A. longinervis. H. microphyllum (Broth. & Paris) Broth., Nat. Pflanzenfam. 1(3) : 986. 1907. = A. tristis. H. mithouardii (Paris & Broth.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907. = A. preudotristis. A, mithouardii var. viride Thér., Ann. Crypt. Exot. 5: 179. 1932. = A. preudotristis. H. nakajii S. Okamura, J. Coll. Sci. Imp. Univ. Tokyo 36(7): 26. 1915. = A. preudotristis. H. obsoletinerve Broth., Ofver. Finsk. Vet. Soc. Férh. 62: 32. 1921. = A. preudotristis. A. okamurae Cardot, Bull. Soc. Bot. Genéve, sér. 2, 3: 281. 1911. = A. sieboldii. A. pellucens Broth., Ann. Bryol. 1:21. 1928. = A. sieboldii. Ai. pellucens var. obtusifolium Broth., Ann. Bryol. 1: 21. 1928. = A. preudotristis. H. piliferum Broth. & M.Yasuda, Ofver. Finsk. Vet. Soc. Férh. 62: 33. 1919-1920. = A. longinervis. H. pinnatum Nog., J. Jap. Bot. 13: 410. 1937. = A. sieboldi. Hf. pseudotriste (Mull. Hal.) Broth., Nat. Pflanzenfam. 1(3): 986. 1907. = A. preudotristis. Hf. sasaokae Broth., Classif. Moss. Japan: 149. 1932 (nomen nudum) = A. preudotristis. H. scabrum Broth., Rev. Bryol. Lichénol. 2: 10. 1929. = A. tristis. H. sieboldti (Dozy & Molk.) Dozy & Molk., Musci Frond. Arch. Indici 4: 127. 1846. = A. sieboldii. H. spinosum Nog., J. Jap. Bot. 13: 789. 1937. = A. longinervis. H. submicrophyllum (Cardot) Broth., Nat. Pflanzenfam. 1(3): 986. 1907. = A. preudotristis. 1997 GRANZOW-DE LA CERDA: ANOMODONTACEAE e9h8) H. tenerrimum Broth., Rev. Bryol. Lichénol. 2: 10. 1929. = A. preudotristis. H. triste (Ces.) Kindb., Rev. Bryol. 26: 25. 1899. = A. tristis. Haptymenium viticulosum (Hedw.) Firnr., Flora 10(2), Beilage 1: 81. 1827. = A. viticulosus. Herpetineuron attenuatum 8. Okamura, J. Coll. Sci. Imp. Univ. Tokyo 38(4): 54. 1916. = H. toccoae. H. formosicum Broth., Ann. Bryol. 1: 22. 1928. = H. toccoae. H. wichurae (Broth.) Cardot, Beih. Bot. Centralbl. 19(2): 128. 1905. = H. toccoae. Aypnum attenuatum (Hedw.) Sm., Fl. Brit. 3: 1279. 1804. = A. attenuatus. H. attenuatum var. fulvum (Brid.) Steud., Nomencl. Bot. 2: 252. 1824. = A. attenuatus. H. fragile (Wilson) Mull. Hal., Syn. 2: 471. 1851. = A. tristis. H. fulvum Brid., Musc. Rec. 2(2): 188. 1801. = A. attenuatus. H. longifolium (Brid.) Mill. Hal., Syn. 2: 474. 1851. = A. longifolius. Hi. pseudotriste Mull. Hal., Bot. Zeit. 13: 786. 1855. = A. pseudotristis. A. rugelii Mill. Hal., Syn. 2: 472. 1851. A. = rugelii. H. triste (Ces.) Mull. Hal., Syn. 2: 478. 1851. = A. tristis. H. viticulosum (Hedw.) With., Syst. Arr. Brit. Pl. ed. 4, 3: 830. 1801. = A. viticulosus. H. viticulosum var. brachycarpum Mill. Hal., Syn. 2: 473. 1851. = A. minor. Leptohymenium longifolium Hampe, Syn. 1: 475. 1848 (nomen nudum) = A. longifolius. L. sieboldii Dozy & Molk., Ann. Sci. Nat. Bot. sér. 3, 2: 310. 1844. = A. sieboldii. Leskea apiculata Schimp. in Mill. Hal., Syn. 2: 473. 1851 (nomen nudum) = A. rugelii. L. attenuata Hedw., Spec. Musc. 230. 1801. = A. attenuatus. L. longifolia (Brid.) Spruce, Musc. Pyren. n. 87. 1847. = A. longifolius. L. tristis Ces. in De Not., Syll. Musc. 67. 1838. = A. tristis. Neckera flaccida Brid., Sp. Musc. 2: 137. 1812. = A. minor. N. minor P.-Beauv., Prodr. 78. 1805. = A. minor. N. sieboldii (Dozy & Molk.) Miill. Hal., Syn. 2: 81. 1850. = A. sieboldii. N. viticulosa Hedw., Spec. Musc. 209. 1801. = A. viticulosus. N. viticulosa var. minor Hedw., Spec. Musc. 210. 1801. = A. minor. Prerigynandrum longifolium Schleich. ex Brid., Mant. Musc. 128. 1819. = A. longifolius. a 7 Oe > eee rans > _ rn a i oe a es ee = * = —- -— 4 | > a a : oe Oo _ => — -_ _ : - 7 a — a | | a — : - oe oe : 7 ee _ ee 7 7 : _ a 7 _ _ : —_ — - a = 7 : | — oe a a ne es " aoa Contr. Univ. Michigan Herb. 21: 277-290. 1997. SPECIES DELIMITATION, AND NEW TAXA AND COMBINATIONS IN LEUCAENA (LEGUMINOSAE) Colin Hughes Department of Plant Sciences University of Oxford South Parks Road Oxford, OX1 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. /empirana, 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 unreliable, 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 sampling, 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 Pages 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 delimitation that relied 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 difficulties. 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 case 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 applied to delimit species should be a requirement for any taxonomic revision. My aim in delimiting species within Leuwcaena 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 ‘traditional’ 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 explicit 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.—Typre: Honpuras. Yoro: 6 km SE of E] 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 foliolis numerosioribus minoribusque, ramulis capitula ferentibus terminalibus 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 280 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 = Md Wl CLA, WRN Lie leat SSUSSSSSSESS MM xX (A SS Me KS CS MOM COG He KG S Wg (UM < Mh wS eC Me Gs SSSSS NZ woe me WS Mv SS ee ale Ml SS MS SS le < a Mel mee KRSS x a ile BESS S Mt WW que ll CK (] it Me WS le WS ate SSS Ml W ( Ke ( W — at S ri BSS ASX (i \\ ia pga RRS MT SU PTL SEU ( Ss SV iF { me 1 q dd =" Ue > x f WUE MU Mlle M : WW wil US Sc 4 Wd. Sse CMU I ( AU Ns add ea \ uv AN FIG. 1. Leucaena lempirana. Leat, flower wit pod, flowering shoot, petiolar and rachis nectaries, leaflet, h peltate bract, seed, longitudinal section of flower and anther. (Based on: Hughes /4172, FHO, isotype.) 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 mucro, 2.5—-3.5 mm long, curling 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 ciliate 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 plano-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 Jocén, lower slopes 7 a S side of the Aguan Valley, 15°22'N, 86°51'W, 16 Feb 1991, Hellin & Hughes 4 (EAP, FHO, K NY); 9 km SSE of Arenal, rd towards Jocén and Yoro, 15°22'N, 86°51'W, Feb 1989, ene Be (HEH); 15 km NW of Jocén, rd to the Agudn Valley in tributary valley of the Rio Macora, between Macora and Sabana Larga, about 8 km NW of Macora, 15°21'N, 86°59'W, 17 Feb foo Hellin & Hughes 14 (EAP, FHO, K, MEXU); rd E from Santa Rita to Yoro, 4 km E of Negrito immediately E of Valle Olomdn 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°S8'W, 17 Feb 1991, Hellin & ihe II (EAP, FHO, K, MEXU, NY); 3 km f 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 2 km W of Olanchito, 15°28'N, 86°28'W, 17 Feb 1991, Hellin & Hughes 8 (EAP, F NY); rd E from Santa Rita to Yoro, nr Cuyamapa, 15°16'N, 87°27'W, 15 Feb 1991, Helin. & aches vi (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 Gaspar 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 Leuwcaena 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 Standl. 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. Leucaena 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 locality 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. nelsoni 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, 97°10'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, pedunculis 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 slightly lustrous or with dense velutinous pubescence. Seeds 4.9-6.6 mm long, 3.5—4.6 mm wide. 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 $ 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°21'N, 98°03'W, 21 Feb 1987, Hughes 873 (FHO, K, MEXU); rd N from Juchitan to Acayucan 5 km N of Matias Romero, low windswept hills, Tehuantepec isthmus, 16°56'N, 95°01'W, 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'W 284 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 3 Mar 1982, Zarate 641 (MEXU); secre 16 km NW of Palomares, rd to Tuxtepec, 17°10'N 95°11'W, 23 Feb 1978, Sousa 9240 (MEXU, MO, UC); “EI Tigre,” 2 km N of San oe Chayuco, Jamiltepec, ip N, 97°49'W, 6 Apr aoe pune 234 (CAS, MEXU, MO); 1.2 km N of jet to Vivero Guapinol towards San Meee Chayuco, 24 km SE of Pinotepa Nacional, 16°19'N, 97°51'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, a eb 1976, Sousa 5306 (MEXU); Acatlan, | km W of Tetela, Dtto. Tuxtepec, 18°29'N, 96°27'W, 22 Dec 1978, Sousa 10306 (CAS, MEXU, MO); 1 km E of Atoyaquillo, Putla, 16°49'N, 97°47'W, 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); 11 km N of Matias Romero, 4 km S of Piedra Blanca, 16°55'N, 95°03'W, 23 Feb 1978, Sousa 9225 (MEXU, MO, UC); 6 km Ae of Pinotepa Nacional, Dtto. Jamiltepec, 16°23'N, 97°S1'W, 22 Oct 1977, Sousa 8453 (CAS, MEXU, UC); fe see 3 km NE of San Juan Guichicovi, Juchitan, 16°58'N, 95°04'W, 25 Mar 1988, ee 12018 (MEXU): 3 km N : Santa Maria Chimalapa, track to Paso veel del Rio del Corte, 16°55'N, eal W, 14 Nov 1984, Hernandez 589 (CAS, MEXU, MO); 3 km E of Ojo de Agua S of Sola de Vega, 16°24'N. 97°0S'W, 9 Dec 1978, Sousa 9986 (CAS, MEXU, MO).—Veracruz: N side of Cerro de los Metates, 25 Jun 1972, Danes 1017 (F, MEXU); Salto de Agua, 11 km S of ae Tuxtla, 18°23'N, 95°20'W, 21 Mar 1967, Sousa 3007 (MEXU); nr Plan del Rio and Emiliano Zapata, 19°20'N, 96°38'W, 16 Sep 1982, Villanueva 246 (NY); Salto de Eyipantla, 8 km from Sihuapan, nr San Andrés 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 Andrés Tuxtla in southern Veracruz. It is essentially a lowland 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, have dense 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.—Typr: 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, little-branched, less seedy variant designated the ‘Giant’ 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 ‘Giant 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. /eucocephala corresponding to the shrubby ‘Common? or ‘Hawaiian’ type and var. glabrata corresponding to the ‘Giant 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 ‘Hawatian 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 /eucocephala 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. /eucocephala, but is glabrous like subsp. glabrata. This material is here described as a third subspecies. Leucaena leucocephala subsp. ixtahuacana C. E. Hughes, subsp. nov.—T Pe: Gua- TEMALA. Huehuetenango: 1 km ENE of San Miguel, track running WSW from Ixtahuacdén 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, foliolis 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 ciliate 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. Curapas: 2 km SE of Toliman rd to Ojo de Agua and Berriozdbal, 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 Ixtahuacdn, above Rio Cuilco, 15°25'N, 91°48'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 286 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 suggest. Subspecies ixtahuacana is named with reference to its very localized dis- tribution centered on the small town of Ixtahuacdn. Leucaena magnifica (C. E. Hughes) C. E. Hughes, comb. et stat. nov. Lewcaena shannonii subsp. magnifica C. E. Hughes, Kew Bull. 46: 547. 1991.—Typr: GUATEMALA. Chiquimula: Llano Grande, 1.5 km SW of Quetzaltepeque. 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 alliance. 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 (Sessé & Mocino ex DC.) Benth. subsp. matudae S. Zarate, Anales Inst. Biol. Univ. Nac. Auton. México, Bot. 65: 134. 1994.—Type: MExIco. Guerrero: Casa Verde, nr Venta Viejo in the Canon 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 alliance, 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 unlike 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 alliance (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 L. esculenta. Leucaena confertiflora S. Zdrate, Anales Inst. Biol. Univ. Nac. Auton. México, 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, Zdrate 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 15/2), 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 $. Zarate, Anales Inst. Biol. Univ. Nac. Auton. México, 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!). Leucaena 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!). 288 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME ZI 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 8. Zarate, Anales Inst. Biol. Univ. Nac. Auton. México, 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. /anceolata 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 841, 587, 835, 536), 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 Marqués, and Santiago Lachigu- iri also has wider pods than typical for var. Janceolata, 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 his hypothesis that it is of hybrid origin between L. macrophylla (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 Gaspar Alvarado for drawing my attention to L. /empirana. I also thank Richard Palmer for providing the Latin descriptions, aeee 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 aoe Polhill for na advice, ae and encouragement, and the curators of A, BM, CAS, EAP, F, FHO, GH, HEH , MEXU, MO, NY, OXF, TEX, UC, UNAH, US, and W for Senet material ei frciliaes a cial 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 Baum, R. B. 1992. Phylogenetic ae concepts. Trends Ecol. Evol. 7: 1-2. Brewbaker, J. L. 1980. What is ‘giant’ Lewcaena? Leucaena Res. Rep. 1: 43-44. . 1987. Species in the ae es Leucaena Res. Rep. 7: 6-20. Brewbaker, J. L., and G. M. 980. Taxonomic studies of the genus Leucaena. Leucaena News. |: 41-42. Brewbaker, 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. Brewbaker, J. L., D. L. Plucknett, and V. Gonzdlez. 1972. Varietal variation and ine trials of Leucaena leucocephala (Koa haole) in Hawaii. Hawaii Agric. an es Res. Bull. 166: 1-29 Britton, N. L., and J. N. Rose. 1928. Mimosaceae. N. Amer. FI. 23: SHle Chamberlain, J. R., C. E. Hughes, and N. Galwey. 1996. ee a isozyme variation in the Leucae- na shannonii alliance (Leguminosae: Mimosoideae). Silvae Genet. 45: Cracraft, ss 1983. Species concepts and speciation analysis. Current Ornith. 1: 159- 187. ———. 1992. The species of the Birds of Paradise er era applying the phylogenetic species eae to a complex pattern of diversification. Cladistics 8: 1-43. Cronquist, A. 1988. The evolution and classification of eis plants. 2d ed. Bronx, New York: ew York Botanical Garden Davis, J. 1. 1995. Species concepts and phylogenetic analysis - introduction. Syst. Bot. 20: 555-559. vis, J. I., and K. C. Nixon. 1992. Populations, genetic variation and the delimitation of phylogenetic species. ae Biol. 41: 421-435. Donoghue, M. J. 1985. A critique of the aoe species concept and recommendations for a phylogenetic alternative. Bryologist 88: 172-1 Eldredge, N., and J. Cracraft. 1980. Phylogenetic ee and the evolutionary process. New York: Columbia ae Gonzalez, V., . Bre ee and D. E. Hamill. 1967. Leuwcaena cytogenetics in relation to the bree ee a ie mimosine lines. Crop Sci. 7: 140 Gray, S. G. 1967. Inheritance of growth habit and ME eae characters in intervarietal crosses in Laie ae Austral. J. Agric. Res. 18: 63- Harris, S. A., C. E. Hughes, R. Ingram, and R. J. pie 1994 A phylogenetic analysis of Leucaena (Leguminosae: Nimosbideae): Pl. Syst. Evol. 191: Hughes, i E. 1986. A new Leucaena from Guatemala. eee Res. Rep. 7: 110-113 ———. 1991. Two new subspecies of Leucaena (Leguminosae: Mimosoideae) from Guatemala Kew Bull. 46: 547-557. . Monograph of Leuwcaena (Leguminosae: Mimosoideae). Systematic Botany Monographs; in press. Hughes, C. E., and S. A. Harris. 1994. The characterisation and identification of a naturally occurring hybrid in Leucaena Benth. (Leguminosae: Mimosoideae). Pl. Syst. Evol. 192: 177-197. 290 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 Hutton, E. M., and S. G. Gray. 1959. Problems of adapting Leucaena glauca as a forage for the Australian ae Empire J. Exp. Agric. 27: 187-196 Luckow, M. 1995. Species concepts: assumptions, methods ana applications. Syst. Bot. 20: 589-605. aac R. 1987. Flora Novo-Galiciana: a descriptive account of the vascular plants of western co. Vol. 5. Leguminosae. An bor: University of Michigan Press. n Ar Mishler, ‘: D., and A. F. Budd. 1990. Species and evolution in clonal organisms - introduction. Syst Bot. 15: 79-85. Nelson, G., and N. I. Platnick. 1981. nae ane? and biogeography: cladistics and vicariance. New York: Columbia University Pre ~ aud 2, D. Wheeler 1990. An amplification of the phylogenetic species concept. Cladis- Panchen, A. L. 1992. Classification, evolution and the nature of biology. Cambridge: Cambridge University Press. Rieseberg, L. H., and L. Brouillet. 1994. Are many plant species paraphyletic? Taxon 43: 21-32 Rosen, D. E. 1979. Fishes from the uplands and intermontane basins of Guatemala. Revisionary studies and comparative biogeography. Bull. Amer. Mus. Nat. Hist. 162: 267-376 Wendell, K. 1976. Honduras in pictures. New York: Sterling Publishing Zarate P., S. 1984. Taxonomic revision of the genus Leucaena from Mexico. Bull. Intern. Group aes sper eice 12: 24-34. 987. nomic identity of Leuwcaena leucocephala (Lam. ) de Wit with a new combination. Poon a 304-306. 994. Revision aa género Leucaena Benth. en México. Anales Inst. Biol. Univ. Nac. Auton. eel Bot. 65: 83-162. Contr. Univ. Michigan Herb. 21: 291-297. 1997. 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-Elizondo CHDIR 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 ts related to C. flexirostris Reznicek, from Oaxaca. Carex queretarensis Reznicek & S. Gonzalez, sp. nov.—T ype: 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, 21°15'50"N, 99°10'13"W, 1590-1600 m, 14 Jul 1996, Gonzdlez & Reznicek 10254, M. Pinedo (holotype: IEB!; isotypes: ANSM! CIIDIR! ENCB! GH! MEXU! MICH! MO! QMEX! TEX! UAMIZ! US!). Fig. 1. Plantae cespitosae; culmi 30-80 cm alti; 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 29) 202 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 |.1-7 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-) Q.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 em 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, Carranza G. 3421 (CIIDIR, IEB); 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, Gonzdlez & Reznicek 10283, M. Pinedo (CAS, CHAP, CHAPA, CHDIR, ENCB, F, GENT, GH, IBUG, IEB, 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 qgueretarensis refers to the species being appar- ently endemic to the state of Querétaro. 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 1997 REZNICEK & GONZALEZ: NEW SPECIES OF CAREX 293 1. Carex queretarensis. Habit of a flowering plant in the field. Photograph taken in popula- tion represented by Gonzalez & Reznicek 10283. Plant is ca. 35 cm tall. 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 ligules. Differences among the four known species of the complex are summarized in Table 1. A revised key to this group is presented below. 294 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 TaB_Le |. Comparison of characters in the Carex mackenziana complex. Character C. mackenziana C. caeligena C. queretarensis C. pubighima Basal sheaths smooth hispidulous hispidulous smooth Ligule length 0.8-3.5 (S—) 5S.5-14 1.1-5 (-6.4) 1.9-8.6 (mm) Uppermost 3-5 separated strongly + separated separated spikes ageregated to somewhat ageregated Terminal spike, 4-17 1.8-4.8 (-7) (2-) 4-9 (-17) 4,5-13 peduncle length (mm) Pistillate scale no no yes yes pubescence Pistillate scale apex Perigynium beak apex Perigynium length (mm) Anther length (mm) hene transverse section obtuse to acuminate obscurely bidentate flattened- trigonous with concave sides acute to acuminate - awned oblique or irregularly bidentate 2.9-4.3 1.3-2.5 eae se with + flats acuminate - awned oblique to irregularly bidentate (3.6-) 4.1-5.6 2.3-3 yay pn with + flat obtuse to acute oblique, strongly flared 3.6-4.6 2.3-3.8 pee with + fla —_ d. Lower leaf sheaths hispidulous with red prickle-hairs; pistillate scales acute to acuminate- awne 2: seats ae bodies glabrous or scabrous on the midvein; ligules (5 anthe 3-2.5 mm long. 2. Sree scale bodies pubescent; ligules 1.1— ~) 5.5-14 mm long; C. caeligena. (—6.4) mm long; anthers 2.3—-3 mm long. C. queretarensis. . Lower leaf sheaths glabrous; at least some pistillate scales obtu 3. Pistillate and staminate scales pubescent, the margins area perigynium beaks se ay a hya- ine, flared, ciliate apex a ella 3. Pistillate and staminate scales glabrous or scabrous on the midvein, the margins not ciliate perigynium beaks more or less bidentulate at apex. C. mac ee er Carex pinophila Reznicek & S. Gonzalez, sp. nov.—Typre: Mexico. Querétaro: 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, 2700 m, 15 Jul 1996, Gonzalez & Reznicek 10293, M. Pinedo (holotype: IEB!; isotypes: 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 1997 REZNICEK & GONZALEZ: NEW SPECIES OF CAREX 295 FIG. 2. Carex pinophila. A. Inflorescence. B. Perigynium, side view. C. Perigynium, front view. D. Achene, front view. A, scale bar = ; B-D, scale bar = 1 mm. Drawn by Susan Reznicek from the type. 296 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 2.5-5.5 mm longae. Inflorescentiae 7-11 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. 11-15, 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; ligules 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 lon Carex pinophila is known only from the type, collected in Querétaro in deep pine duff in a moist forest of Pinus 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 20 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—) 10-41 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-S 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. flexirestris 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 Jones, S. D., and A. A. Reznicek. 1992. Carex rhynchoperigynium (Cyperaceae), a new species from Hidalgo, Mexico, and a key to Mexican and Central American Hymenochlaenae. Sida 15: 215— PDN Reznicek, A. A. 1986. The taxonomy of Carex sect. Hymenochlaenae (Cyperaceae) in Mexico and Central America. Syst. Bot. 11: 56-87. . 1990. Four new species of Carex (Cyperaceae) from Mexico, with notes on the Mexican Carex Flora. Contributions Univ. Michigan Herb. 17: 279-293. : =P Sow Te A ee at ae les ; © toes : ; - _ » oe Oo : : : : Oo a a 7 OO - 7 _ : a ra : as _ 1, " a Oo _ 7 ty - 7 > a 7 a oO : a 7 = i — 7 y : 7 > _ 7 a _ : - —_ - ; : ; : > a : 7 _ a : _ ; a _ o 7 7 _ * % 7 7 7 7 : 7 = 7 ee _ _ 7 Oo ; — a ee _ - - ; — OS _ ; Se _ : a . ; _ — -— a oe ae ome eae . a 7 a tae a - os : _ 7 - : > _ - -_ : : oe Oo - aa oe 2 o a a - 7 7 7 a _ — : a 7 7 7 ; a : > - _ 7 Oo : ; - - 7 7 : 7 - - _>- _ - 7 a 7 a — a _ Oo > *s ae _ oe —— a a es a * 7 — _ a a - 7 - a i= Oo _ _ tu en > 7 rr ee i 7 7 ? = a - Contr. Univ. Michigan Herb. 21: 299-308. 1997, 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 existence of 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. mo/lesta 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 299 300 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 clavate due to its prominent staminate base, and perigynia that are nerveless or nearly so adaxially. Because of its distinctive morphology, we describe this entity as a new species, a locally frequent regional endemic. Carex molestiformis Reznicek & P. Rothr., sp. nov—Type: 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, TI9N 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, hyalinae, 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, plicate, 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, ().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—40-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, l-nerved. Staminate scales 2.6-5 mm long, |.3-1.8 mm wide, ovate, obtuse or rarely acute, pale brown with hyaline margins, l-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, TI8N 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 11W, 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 aie Seer 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 Boe & Amason (ctb - herbarium of Charles T. Br Deas CARROLL Co.: SW 1/4 of NW 1/4 section 27, TI8W R23W, 24 May 1992, ae 4388.08 (MICH, UARK). CLEBURNE Co.: Sandiff, SW 1/4 of NE 1/4 section 25, T12N R12 1993, Hyatt 5272 & Hyatt, Beer (MICH, UARK). Futon Co.: Vidette Community, S of center ie section 2, T20N R11W, 22 May 1993, Hyatt 5489 (MICH, MO, UARK); Salem, at Civic Center Lake (=about | mi W of the U.S. Post Office), SE 1/4 section 21, T20N R8W, 12 May 1992, Hyatt 4308.25 (MICH, UARK). GarLanp 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, a, 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 S557 & ee (MICH, UARK). Izarp 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, TI7N R11W, 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 on Ark. Hwy 5, SW 1/4 section 6, TI7N R11W, 10 May 1993, Hyatt 5309 (MICH, UARK); Sylamore, Ark. Hwy 9 brides at White River, 27 May 1993, Hyatt 5550 (MICH, MO, NO); about 4.5 mi west of Ark. Hwy. 9 on Knob Creek Rd at bridge over Knob Creek, E 1/2 section 31, TI7N R9W, 4 June 1993, Hyat 5596 (MO, UAM); Moccasin a about 1 mi S of Ark. Hwy 5, NE 1/4 of NW 1/4 section 7, TL7N R11W, 7 June 1993, Hyatt 5623 (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 free and Ark. 58, section 22, 21 May 1994, Thomas 139187 aoe (ctb). ee Co.: Around pond next to Ark. Hwy 295, 1.5 mi SW of Crosses, 20 May 1989, Stewart 59-30 (UARK). ae Co.: Buffalo River State Park, Hills above Buffalo River, es Yellville, 14 May rane Demaree 64832 (MO); near the Buffalo River ca. 1.8 m ESE of the town of Rush, SW 1/4 section 11, TI7N R15W, 13 May 1977, Smith 3156 (UARK): Buffalo National River, SW 1/4 of SW 1/4 section 31, TI8N 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, TI7N 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, TI6N R22W, 27 April 1977, Smith 3142 (UARK). PoLk Co.: 50 m N on Hwy 71 from its jct. with Polk County Rd 414, N of Hatfield, 13 May 302 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 1990, Jones & Jones 4653 (MICH); 0.4 mi NE of center of Hatfield on both sides of US 71, SW 1/4 section 13, T3S R32W, 10.8 mi SW of junct. with Ark. Hwy 88 at Mena, 20 May 1993, Reznicek 9368 & Reznicek, Rothrock (ctb, MICH, UARK, VPI). PuLaski Co.: Little Rock, along E-bound off-ramp from I-440 (Exit 5: Little Rock River Port/Fourche Dam Pike exit), 17 May 1993, Rothrock 290] & Reznicek, Reznicek, Bryson, Hyatt (MICH, chromosome voucher n = 37), Hyatt 5369 & Rothrock, Reznicek, Bryson (UARK). Scorr Co.: 6.2 mi NE of Polk Co. line, 2 mi SW of Y City on N side US 71, 20 May 1993, Reznicek 9371 & Reznicek, Rothrock (MICH, UARK). SEARCY Co.: Maumee Access on §S side of Buffalo River, SW 1/4 section 12, TI6N R16W, 28 May 1995, Hyatt 6463 (MICH); reed (KNK); Leshe, 2 mi S on U.S. Hwy 65, NE 1/4 sa 2, TI3N RISW, 28 May 1995, Hyatt 6473 [CH). Sevier Co.: DeQueen, 1.9 mi N on U.S. Hwy 71, section 17, T8S R31W, 23 April 1995, ao 6306 & Hyatt (MICH). SHARP Co.: Evening a about 2-3 air mi NE, at NW 1/4 of SE 1/4 section 24, TI7N R6W at Ark. Game & Fish Commission Access to Strawberry River, 25 May 1996, Hyatt 7149 (MICH, UARK). Stone Co.: Mountain View, Sylamore Ranger Station of the Ozark National Forest, 19 May 1993, Hyatt 5454 i MICH); Ozark National Forest, Sylamore Ranger District, logging rd in valley due W of Livingston Creek/Partee Springs pull-off along Ark. Hwy 5 center of NW 1/4 of section 26, T16 N R1IW, 26 May 1993, Hyatt 5526 (MICH, UARK); Ozark National Forest, Sylamore Ranger District, along Sylamore Trail at first fields balbi campgrounds, SW 1/4 of SW 1/4 section 3, TISN R11W, 27 May 1993, Hyatt 5547 (MICH, MO). VAN Buren Co.: E side Ark. Hwy 27 just N of junct. with Co. Rd. 234 (at ee 1.5 mi S of Searcy Co. line, near center of section 10, TI2N R17W, 20 May 1993, Reznicek 9392 & Reznicek, Rothrock (MICH); Fairfield Bay, SW 1/4 section 9, TIIN R12W, 8 May 1993, Hyatt 5295 & Hyatt, Beer (UARK): 5298 (MICH, MO, VSC). WASHINGTON Co.: 2 mi NW of U. Campus, 28 May 1921, J.N.B. (UARK); Fayetteville, 23 May 1949, Moore (UARK); Farmington, 9 May 1950, French 477 (UARK): just W of the Art Dept. Annex Aas: Lot 19), U. of A. Campus in Fayetteville, 17 May 1982, Smith 3678 (UARK); Springdale, E side of northbound ramp onto I-71 off coy ae 26 May 1994, Reznicek 9793 & ae Jones, Jones (BRCH, DAO, KANU, KNK, MICH, MO, UARK, VDB). County UNKNown: NW Arkansas, June 1880, Haney (UARK); May 1883, eh (UARK). —Missourl. Howe. Co.: on the Eleven Point River, T25N R7W, section 2, on farm rd 200 bridge crossing, | ae 1990, Summers 3265 (MO): 4 mi S of Caulfield, T22N R1OW section 19, 6 June 1990, eae 3276 (M OREGON Co.: 3 mi S of Billmore, Billmore Quad., T22N R2W section 29, 13 May 1990, Summers 3191 (MO) 4 1/2 mi N of Greer, T25N R4W section 23/24, 20 May 1992, Summers 5034 (MO). PuLAsK! Co.: 3 mi S of Big Piney, T34N RIIW, section 24, 3 June 1991, Summers & Ryan 4450 (MO). REYNOLDS Co.: along County Rd 700 on E side of Dry Valley, ca. 3 rd mi NW of junction with State Hwy 21, ca. 4 air mi NW of Ellington, T30N ROLE NW 1/4 SW 1/4 SW 1/4 section 7, 16 May 1993, Yatskievych & Yatskievych 93-67 (MO). aoe Co.: On Hwy 19 at Currant River Bridge, E eos . are 1974, ned 3654 & Bevard (WARM).—OKLAHOMA. ADAIR Co.: N side Okla. Hwy 62, i NW of 1 jct., Eldon, ca. 8 mi cy of Ep a N edge section 27, TI7N R23E, May 26 Boe Re znicek O78 : Reznicek, ae Jones (MICH). DELAware Co.: US 59, 3.6 mi N of rt 20 in Jay, 13 May 1980, Huft 1109 & Goodman (OKL): sa stream access rd along N shore of Lake Eucha, 1.6 mi E of Okla. Hwy 59/10, SE 1/4 section 25, aed R23E, May 25 1994, Reznicek 9770 & Reznicek, Jones, Jones (BH, BRCH, MICH, OKL, OKLA). Mays Co.: Spavinaw State Park, along N bank of Spavi- w Creek on S outskirts of Spavinaw, SW 1/4 section 15, T22N R21E, May 25 1994, Reznicek 9766 & ey Jones, Jones (BRCH, DUR, MICH (chromosome voucher n = 37), OKL). McCurtain Co.: Niprbaitiaase Fork aan N side Okla. Hwy 4 ca. | mi SE of Smithville, section 13, T1S R25E, ca. 1 W of Watson, May 20 1993, Reznicek 9357 & Reznicek, Rothrock, Jones, Jones (BRCH, ctb, 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 re i to C. molesta. An immature collection from Arkansas: Little River Co., White Cliffs, Moore SIU? (UARK) (not mapped) may be this species, but it is too young to identify with certainty. Also, the habitat (chalk cliffs) 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. 1997 REZNICEK & ROTHROCK: CAREX MOLESTIFORMIS 303 FIG. 1. Distribution of Carex molestiformis (dots) in Arkansas, Missouri, and Oklahoma. The hatched area represents the distribution of C. molesta within the area covered by this map. Carex brevior and C. festucacea both occur essentially throughout the region shown on this map (though C. festucacea is absent from the far western edge). DISCUSSION Even before the recognition of Carex molestiformis, the immediate allies 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 Nore: Fully mature perigynia are a prerequisite for keying. Perigynium measurements are best taken from perigynia between 1/3 to 2/3 the wa e spikes. Perigynia become distinctly narrower near the top of the spikes, and the lowermost few ean 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 ea produced by the first flush of growth in pring. Many species continue to flower sporadically through the summer, but these later Sener tend to be congested, with more numerous spikes with shorter staminate bases, and w e lowermost bracts longer an leaflike. These late-season inflorescence forms are not sebincctes for in the key and descriptions, and plants collected late in the season, with all the spring inflorescences missing or shattered, are quite difficult to identify and may be impossible to key. 1. Plants colonial from short-creeping rhizomes; vegetative culms numerous and ee tai strongly tristichous with 15-35 leaves when fully developed; achenes 1.6—-2 times as long a wide (and 0.9-1.2 mm wide); larger spikes with 5—25 (—30) ee Plants definitely clumping (though rhizomes may elongate witl inconspicuous, and usually with fewer than 15 1—1.6 times as long as wide (an Sean 1. age); vegetative culms few leaves, thus not ee tristichous; achenes d 0.9-2.2 mm wide): larger spikes with ca. 15-80 perigy ynia. 2. Perigynia finely granular- papillose (30- 40x), the body reniform, 0.6-0.9 times as long as N reniformis. oadly ovate, broadly elliptic, + orbicular, or rarely slightly obovate, (0.7—) 0.9-1.6 times as a as wide (and 1.5 ) e to acuminate. oe aes perigynia 2.5-5.5 mm long, 1.5—-3.6 mm wide, with beaks 0.8—1.5 (-1.9) mm long; sit plumply planoconvex or concavo-convex, the bulge Bea by the achene Soca nt only on the dorsal face (except in C. bicknellii var. bicknellii, with papillose sheat —6.1 mm wide); lower pistillate scales oe sheaths finely eae at a ee (30-40x); perigynia membrana- ceous and transluc the dark achene + visible through the adaxial face of the perigynium; i, at i some perigynia with the wings and base of b ie + erose, scalloped, or even with an irregular tooth and not ih cp tapere a: sa distinctly 4—7-nerved over the achene axial face, os 5-) 5-5.5 mm long, wings eae strongly coppery-tinged; te pa usually reddish ies anthers (2.5—) 2.9-4.1 mm long. C. bicknellii var. bicknellii. 5. Perigynia nerveless or faintly and irregularly 1-5-nerved over the achen h .6-4.8 (-5.2) m adaxial face, 3 mm long; wings and pistillate scales lee tinged; anthers 1.3—2.6 mm lon merritt-fernaldii. Leaf sheaths smooth; perigynia leathery, opaque, with the wings and base of beak usually sae and evenly serrulate and symmetrically tapere 6. Spikes ¢ ect culms (4—) 5—7 (-10), tapered at the base: the terminal one with a conspicu minate base; inflorescences typically 2.5—4.5 m long with the lowest Coane (3—) 5-13 (-23) mm long; perigynium beds 7-) 0.9-1.2 (—1.3) times as long as wide. 7. Larger achenes 1.4-1.8 mm wide, (1.6—) 1.7-2 mm long; larger perigynia 3.2- mm oe 2.5—3.3 (-3.5) mm wide, nerveless or occasionally faintly 1—5- cened ventrally. *. brevior. 7. Larger achenes 11.35 mm wide, 1.2-1.7 mm long; larger perigynia 2.54 (—4.2) mm long, 1.5—2.4 (—2.6) mm wide, mostly 2-4 (—6)-nerved ventr C. festucacea. 6. Spikes on larger culms 2-4 (—5) (rarely more), rounded at the base, the terminal one lacking a conspicuous staminate base; inflorescences 1.2—2.7 (—3.6) cm long with the lowest internodes 1.5-6 (-9) mm long; perigynium body (0.9-) 1-1.6 times as long as wide. 1997 REZNICEK & ROTHROCK: CAREX MOLESTIFORMIS 305 8. Achenes of larger eee 0.9-1.3 mm wide, elliptic to narrowly oblong, 1.3- .6 times as long as wide, larger perigynia 1.8-2.8 (-3) mm wide, squarrose- spreading at ee (25—) 30-80 per spike; widespread. C. molesta. . Achenes of ee 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 ea 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 .1 mm wide ee aes in C. bicknellii var. bicknellii with papillose sheaths), 7 cere (1.4-) 1.6-2.5 (-3.4) mm long; perigynia thin and + waferlike, + biconvex around the achene, ie 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 oats 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 2.9-4.1 mm long; plants in small clumps (usually <10 culms) in dry to mesic habitats. C. bicknellii var. bicknellii. . Leaf sheaths smooth; perigynia 2 thickened and opaque, with greenish or brown wings, the larger (3.1-) 3.5-4.8 mm wide; pistillate scales pale yellowish brown; anthers 1.8—3.5 mm long; nee in dense, large clumps (up to 200 culms) in wetlands. C. bicknellii var. pe . Larger perigynia 4-6.1 mm wide, nerveless over achene ventrally, or nearly so; staminate scales (and sometimes the lowermost pistillate scales) with the midi excurrent as a scabrous awn 0).1—-0.9 (—2.4) mm long; larger culms with (2—) 3 es. C. tetrastachya (=C. a ee oe) — = Oo \o 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 chromosome 306 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 FIG. 2. Perigynia and achenes (above and to the right) of 1) Carex brevior (from Bryson 11416, MICH): 2) C. festucacea (from Reznicek 9778, MICH); 3) C. ae (from Reznicek 9776, MICH); and 4) C. molesta (from Wilson 4046, MICH). Scale bar = 11 count of C. molestiformis was determined as n = 37 in plants from Garland Co., Arkansas (Rothrock 2904, reported as “C. cf. brevior” by Rothrock & Reznicek, 1996b); Pulaski Co., Arkansas (Rothrock 2901); and Mays Co., Oklahoma (Reznicek 766). In contrast, the chromosome count for C. molesta is n = 34 (Wahl, 1940; Love & Léve, 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 reconize Carex molestiformis in Arkansas. He collected a large proportion of the known specimens of this plant and also commented on our manuscript. Stanley and Gretchen Jones accompanied us in the 1997 REZNICEK & ROTHROCK: CAREX MOLESTIFORMIS 307 TaBLe 1. Morphological comparison of Carex brevior, C. festucacea, C. pee and C. molesta. The smaller measurements in parentheses in this table were mostly not u the key, ich emphasizes measuring the larger inflorescences, perigynia, and achenes on a es Also, the exceptionally large measurements for perigynium length in C. molestiformis and C. molesta each came from single exceptional specimens and are not accounted for in the Character C. brevior C. festucacea C. molestiformis C. molesta Perigynium length (mm Perigynium width (mm) Perigynium body shape Perigynium body L/W Adaxial nerves on perigynia Achene width (mm) Staminate and lower pistillate scale x and color Anther length (mm) Perigynium maturity) Number of spikes per inflorescence Length of staminate ase on terminal spikes (mm) Inflorescence density and aspect owes nflorescence node length (mm Inflorescence length (cm) (2.95) 3047 (25) O53 24 CSS) mostly orbicular, rarely slightly (0.7-) 0.9-1.1 (-1.3) 0 (-5, faint) (1.2-) 1.4-1.8 acute, rust-brown tinged Ce nrs-30 appressed— ascending (252) 3-914) usually + open, sometimes arched or even nodding 3 \614 (93) (32) 25445. C65) 3505-4 43 15-36) ae inte to orbicular, rarely ay S ocaie 0.9-1.2 (-1.3) (0-) 2-4 (-6) (0.95-) 1-1.35 acute, white— hyaline 1-2.1 spreading + open, some- times et (3-) 5213 220} (i5)1215=45 (26) 4-5.4 (-6.1) (2.4-) 2.6-3.4 roadly ovate to ee elliptic, sometimes + orbicular (0.9-) 1-1.6 (3-) 4-7 1.4-1.8 obtuse, pale b (1.4-) 1.6-2.6 appressed— ascending 1.5-5 (-9.5) compact, stiffly 1.5-6 (-9) (3-) 3.3-4.8 (-5.7) iS 3) broadly ovate to broadly elliptic, sometimes + orbicular 1-1.6 (O—) 3-6 0.9-1.3 obtuse, pale brown 1.1-2.2 squarrose— spreading 3-4 (-6) 1-4.5 (6) compact, stiffly erect 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, K, and WARM, for lending us material and oe us to scour their holdings for specimens of C. molestiformis. We are grateful o Susan Reznicek for drawing Figure 2, and to Sarah Cleveland and George Yatskievych for their we with the distribution map. Sc >: LITERATURE CITED Gleason, H. A., and A. Cronquist. 1991. Manual of vascular plants i Northeastern United States and adjacent anen 2d ed. New York: New York Botanical Garde Love, A., and D. Léve, 1981. Chromosome number reports LX XIII. es 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 (Cyperaceae). Rhodora 93: 51-66. Rothrock, P. E., and A. A. Reznicek. 1996a. A new species of a section Ovales (Cyperaceae) occurring in the Ozark Mountain Region. Brittonia 48: 104-110. . 1996b. Documented chromosome numbers 1996: 1. a ee numbers in Carex section 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: 458-470. Zager, S.C. 1991. Multivariate ora of the Carex brevior group in Iowa. M.A. Thesis, University of Northern Iowa. xiii + Contr. Univ. Michigan Herb. 21: 309-313. 1997. NOMENCLATURAL CHANGES IN MEXICAN DIOSCOREACEAE AND LEGUMINOSAE Oswaldo Téllez V. Herbario Nacional Instituto de Biologia UNAM Apartado Postal 70-367 04510 México, D.F., Mexico Recent work on the Mexican and Central American species of Dioscorea (Dioscoreaceae; Téllez & 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. Téllez, nom. et stat. nov. Dioscorea spiculiflora var. chiapasana G6mez-Pompa, Ciencia (México) 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!). REPRESENTATIVE SPECIMENS. Cuiapas: Amatenango de la Frontera, between Amatenango and Frontera Comalapa, along Rio Cuilco, 4 Oct 1976, Breedlove 40671 (MEXU); Mpio. Acala, near Acala, 6 Nov 1980, Breedlove 47156, 47157 (MEXU); El Chorreadero, sobre el Km 25 de la carretera Tuxtla Gutiérrez—San Cristébal de las Casas, 26 Nov 1982, Cabrera & de Cabrera 3862 (MEXU); Rancho La Esperanza, Tuxtla Gutiérrez, Gomez P. 71 (MEXU); Puente Belisario Dominguez, Rio Grijalva, al E de Tuxtla Gutiérrez, 11 Sep 1958, Gémez 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 § 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 Tuxtla 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, Téllez 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 309 310 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, Bidtica 11(3): 187. 1986.—Type: Mexico. Veracruz: Mpio. Juchiqué de Ferrer, el Cerro de Villa Rica cerca de Mundo Nuevo, 1500 m, 19°48'N, 96°46'W, 7 May 1981, Castillo, Cortés & Becerra 1836 (holotype: XAL!). R FE SPECIMENS. PueBLa: Villa de Juarez, 19 Apr 1959, Gomez P. 168 (MEXU): 5 km al sur ae Villa ee 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. pl. 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. Martius ex Griseb., 1842.—Type: Mexico. Oaxaca: Costa del Pacifico, Yolotepéquez, 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.—Tyre: 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!). REPRESENTATIVE SPECIMENS. DisrRtro FEDERAL: Ixtapalapa, 1911, Altamirano s.n. (MEXU); | km al O de Santa Clara, | Sep 1953, Barajas M. s.n. (MEXU); Pedregal de San Angel, 18 Oct 1965, Diego 173 (MEXU); Pedregal de San Angel, 18 Oct 1965, Diego 174 (MEXU); Pedregal de San Angel, | Feb 1966, Diego 393 Bikes XU): Del. de Coyoacan, 200 m al N de la Sala Netzabnaleoyotl, 9 Aug 1986, Garcia M. 26859 (MEXU); pe ie de San Angel, Gomez P. 34 (MEXU),; Ciudad Universitaria, 27 May 1980, a Nahe M. 4389 (MEXU); Pedregal de San Angel, 13 Aug 1945, Hernandez X. 308 (MEXU); Pedregal, Aug 1924, Lyonnet 33 (MEXU); Lomas, 17 Oct 1937, Lyonnet E. 1742 (MEXU); Pedregal de San Angel, 10 Sep 1950, Matuda 19537 (MEXU); Cerro Zacayuca, 11 Sep 1980, Panti M. 332 (MEXU); Ejidos de Padierna, 30 Jul 1980, Panti M. 536 (MEXU); Pedregal, Tizapan, 19 Aug 1896, Pringle s.n. (MEXU); Pedregal de San Angel, cerca de los Reyes, 28 Sep 1952, Rzedowski 1848, 1997 TELLEZ: NOMENCLATURAL CHANGES Si 1849 (MEXU); Delegaci6n de Xochimilco, desviacién del Conejo, 8 Aug 1976, Ventura A. 1839 (MEXU); Delegacién 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, Villasenor et al. 942 (MEXU).—MeExico: Mpio. Huixquilucan, 2 km al SW de San Bartolito, 18 Sep 1982, Galvan R. 1276 (MEXU); Mpio. Texcoco, Bafios de Netzahualcoyotl (Cerro Tetzcotcingo), 7 km al E de Texcoco, 3 Aug 1976, Koch & Garcia P. 76145 (MEXU); Mpio. Amecameca, en loma de Panteo6n, 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: ae o. Erongaricuaro, 5 km al W de Erongaricuaro, 22 Aug 1993, Chdazaro 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. Pétzcuaro—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 desviaci6n a Santiago Undameo, 16 km al SW de Morelia, carr. a Tiripetio, 9 Sep 1979, Soto N. 991 (MEXU).—MokreE Los: autopista Mexico—Cuernavaca, Km 53.5, 9 Oct 1960, Espinosa J. 351 (MEXU): Mpio. Tepoztlén, 2-3 km al N de Tepoztlan, sobre la canada de la zona arqueolégica 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 ( ); Dtto. San Juan Mixtepec, Yuu Tzaan (Cafiada de Cazuelas), a 9 km al S de San Juan Mixtepec, 3 Sep 1989, Reyes J. 1937 (MEXU).—QuereEtaro: El Batan, 27 May 1982, Argiielles E. 1794 (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 Pavén). 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.—Tyre: Mexico. Guerrero: Acapulco, 1842, Hinds s.n. (holotype: K!). Tephrosia smythiae McVaugh, Flora novo-galiciana 5: 726. 1987.—Typr: MEx- co. Jalisco: Mpio. Cabo Corrientes, rocky stream valley, ca. 5 km N of El Tuito, McVaugh 25487 (holotype: MICH!; isotype: MEXU!). REPRESENTATIVE SPECIMENS. GUERRERO: Potrerillos del Rincén, 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, Téllez V. & Novelo ee (MEXU).—MicHoacan: La Mada, Feb 1899, Peat 851, 85] bis (F).—NAyYaRrIT: 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 re 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 Blas, Km 3-5 sobre el camino a Pintadeno, a sale en el Km 16 de la carr. Tepic-Miramar, 12 Sep 1985, Téllez V. 9294 (MEXU); Mpio. Tepic, 3 k S12 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 al E de Cora, terraceria al Cuarentefio, 29 Mar 1987, Téllez V. 10050 (MEXU); Mpio. Tepic, Km 20- 30 camino de terraceria a el Cuarenteho—Cora—Jalcocotan, en la serrania de San Juan, 14 Jan 1988, Téllez V. 11134 (MEXU); Mpio. Compostela, Km 60 carr. Tepic—Pto. Vallarta Jal., 15 Feb 1980, Zarate S. 469 (MEXU).—Oaxaca: Dtto. Juchitan, Mpio. Sta. Marfa Chimalapa, ca. 13 km al E de Sta. Maria, filo we lava en la vereda a Chimalapilla y Monte Rico, al NE del Rio del Corte, 27 Oct 1984, eles 529 (CHAPA, MEXU); Dtto. Pochutla, Pluma Hidalgo, San Rafael Toltepec, a 17 km al N de aan 14 Feb 1976, Sousa S. et al. S347 (MEXU); Dtto. Putla de Guerrero, 15 km al SE de Putla, 5 Aug 1976, Sousa S. et al. 5858 (MEXU); Dtto. Pochutla, Mpio. Pluma Hidalgo, Toltepec, a 2 km de la carr. Pochutla—Miahuatlan, en la carr. a Pluma Hidalgo, 23 Oct 1976, Sousa S. et al. 6470 (MEXU); Dtto. oe = Guerrero, Santa Cruz del castillo, a 13 km al S de Putla, 4 Feb 1977, Sousa S. et al. 7031 (M MO); Dtto. Pochutla, a 6 km al NE de Chacalapa, en la desv. a Finca Pilas, 7 Feb 1977, Sousa “ et ie 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 Decl1978, 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 a rather than elliptic-obovate, and acute rather than obtuse or rounded at apex.” Examination of collections of 7. crassifolia from throughout its range, Sinaloa to Chiapas, showed that these differences are included in the variation of the foliar morphology. Consequently, 7. smythiae can no longer be maintained. Tephrosia leucantha Kunth in H.B.K., Nov. gen. sp. 6[folio]: 360. 12 Jul 1824; 6[quarto]: 460. Sep 1824.—Typr: 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: Exico. 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 aaa Garcia, 11 Oct 1899, Townsend & Barber 367 (MO).—DurRAanco: 21-27 Aug 1906, E. Palmer 78 (MO).—Nayarit: Mpio. Nayar, Sierra de Las Palomas eae de Alica), 9 Oct 1963, eel Sousa 2041 (MEXU); Mpio. La Yesca, 22.3 km al W de Huajimic, brecha a Tepic, 27 O Tenorio L. & Flores F. 16746 (MEXU). cies Del Cio al cerro de la Mesa, 20 are see Altamirano F. 1564 (MEXU).—Sonora: Alamos, 3 km al S La Lobera, Ejido Zahuarivo, 25 Aug 1986, Lezama T. & Frame 11925 (MEXU). DAG ATECAS: ae zea of La Bufa, 10 Aug 1948, Dressler 113 (MO 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 7. leucantha by type of pubescence, bracts length, and the degree of adnation of the vexillar stamen. I believe only the last is note- worthy; the stamen is adnate to the staminal tube in 7. leuwcantha but not in T. feddemana. Yet, the adnation of the stamen to the staminal tube is not a constant and reliable 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 7. feddemana only 1997 TELLEZ: NOMENCLATURAL CHANGES 33 the type collection is available for study. Because in all other features the type of T. feddemana agrees with the morphological variation shown by T. leucantha throughout its range, 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- a Centre (United Kingdom), and British Airways for financial support for visits to herbaria in many and Switzerland. The National Geographic Society provided financial support for fieldwork ee 4349-89), and W. Berendsohn (B) gave assistance with ee 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 Goémez—Pompa, A. 1959. Una nueva variedad de Dioscorea spiculiflora Hemsl. Ciencia (México) 18(11-12): 242-244. Matuda, E. 1953. Nuevas Dioscoreas de México. Bol. Soc. Bot. México 15: 23-25. McVaugh, R. 1987. Leguminosae. In Flora novo-galiciana by R. McVaugh (general editor W. R. Anderson), vol. 5. Ann Arbor: The University of Michigan Press. Téllez V., O., and B. G. Schubert. 1994. Dioscoreaceae. In Flora mesoamericana, ed. G. Davidse, M. Sousa, and A. Chater, 6: 54-65. Mexico City: Universidad Nacional Aut6noma de México; St. Louis: Missouri Botanical Garden; London: The Natural History Museum. 17, ’ i 7 a 7 ; ea oa = : : oe —— a . _ —_ 7 : ee a -_ 7 Oo a 7 - OO OO ; Be a a oe - oe ee - " re oe - - a 7 7 > 7 - - : : a - 7 - a : : : : - - _ 7 7 : _ : 7 7 a ; - a 7 - > 7 7 7 - _ : : 7 * ; a : 7 : : - a : : a - 7 7 - _ 7 - So . oO : a : 7: = a — _ 7 Ss > a Soe ane > oO a OS Oo i oe 7 _ 7 a a Oo : 7 7 7 - oo 7 7 — 7 - OO - - - OS 7 - 7 oe _ : 7 - a — OS a - a oe a _ — OO : _ 7 7 a - a : + ee 7 - 7 - 7 7 7 7 oe ae _ Contr. Univ. Michigan Herb. 21: 315-317. 1997. A NEW SPECIES OF APPENDICULA (ORCHIDACEAE) 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: MALaAysIA. Sarawak: route from Ba Kelalan to Mt. Murud, near Camp III, 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 obliquely 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 S15 316 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 FIG. 1. Appendicula bilobulata. A, B. Habit. C. Leaf apex, back view. D. Leaf apex, front view. E. Flower, side view. F. Lateral sepals and column. G. Dorsal sepal and petals. H. Lip, flattened. I. Lip, natural position, side view. J. Base of lip showing callus. K. Pedicel with ovary and column, anther-cap removed, side view. L. Pedicel with ovary and column, anther-cap removed, back view. M. Anther- cap. N. Pollinarium. Scale: single bar = 1 mm; double bar = | cm. Drawn by Susanna Stuart-Smith. 1997 WOOD: APPENDICULA BILOBULATA a7. 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. Opera Botanica 89: 135-142. Wood, J. J., R. S. Beaman, and J. H. Beaman. 1993. The Plants of Mount Kinabalu 2. Orchids, p. 89, fig. 7. Be a a. ee SS oe : - ; : 7 _ _ 7 a - - : 7 -_ — a _ : : _ 7 : _ _ : 7 7 7 : : : - a _. a _ 7 7 : - a - - : . a ee = 7 Oo Be aan EO - : - SS - 7 : - a - : : 7 : 7 : 7 7 7 ae ee eS! ea | Oo er ae 7 OS — Be - SO Oo 7 7 a Oo 7 a — a > >= _ Oo ey reer = : _ - rae : a: oy ete Contr. Univ. Michigan Herb. 21: 319-334. 1997. TAXONOMIC AND NOMENCLATURAL NOTES ON THE DELESSERIACEAE (RHODOPHYTA) Michael J. Wynne Department of Biology and Herbarium University of Michigan Ann Arbor, Michigan 48109 PANTONEURA AND THE STATUS OF GIGARTINA FABRICIANA LYNGB. Gigartina fabriciana Lyngb. has had a complicated history. The species was described by Lyngbye (1819, pl. 11D) 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 (6159'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.1.: D. rostrata (type locality: Greenland), D. baerii Rupr. (type locality: a site on the coast of Russian Lapland), D. corymbosa J. Agardh (type locality: 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) listed the names Rhodomenia eae 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 enn) \ 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. eH) 320 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. alata 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 earlier 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 Jénsson (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. montagne Kjellm. [var.] rostrata (Lyngb.) osenv. 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, pl. 83) initially recognized “Delesseria angustissima Griffiths msc.” as a 1997 WYNNE: DELESSERIACEAE opal 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. alata (Huds.) J. V. Lamour. Two years later, in his discussion ac- companying D. alata, Harvey (1849, pl. 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. angustissuna. 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 Liining (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; Bgérgesen & 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 sublittoral 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-type” apex (Kylin 1924). A portion of the axis of this specimen proximal to the apex is depicted in 322 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 21 (Xt = Lt 4, KON Ae oe ¥ raed ies ATS > SERSG Sans ——