CONTRIBUTIONS FROM THE UNIVERSITY OF MICHIGAN HERBARIUM Volume 15 A Monograph of the Genus Peixotoa (Malpighiaceae) CHRISTIANE ANDERSON Notes on Neotropical Malpighiaceae—I WILLIAM R. ANDERSON On Meiothecium (Sematophyllaceae) WILLIAM R. BUCK The North American Distribution of the Genus Dryopteris THOMAS M. CARLSON & W. H. WAGNER, JR. The Itinerary of Mary Strong Clemens in Queensland, Australia A. M. CARTER The Genus Mirandea (Acanthaceae) THOMAS F. DANIEL Further Notes on Steccherinum crassiusculum (Hydnaceae) KENNETH A. HARRISON & CLARK OVREBO The New Synantherology vs. Eupatorium in Nueva Galicia ROGERS McVAUGH Compositarum Mexicanarum Pugillo Supplementum ROGERS McVAUGH The Ferns of Steere and Harrington MICHAEL G. PRICE Linum McVaughu, a New Species from Mexico C. M. ROGERS Comments on the Mielichhoferioideae of Central America with the Description of a New Species of Synthetodontium JONATHAN SHAW & HOWARD CRUM Pohlia Hedw. (Musci) in North and Central America and the West Indies JONATHAN SHAW Selected species of Leucocoprinus from the Southeastern United States HELEN V. SMITH & NANCY S. WEBER A New Species of Rhynchospora (Cyperaceae) from Venezuela WM. WAYT THOMAS -Botrychium rugulosum (Ophioglossaceae), a Newly Recognized Species of Evergreen Grapefern in the Great Lakes Area of North America W. H. WAGNER, JR. & F. S. WAGNER Observations on Four Species of Delesseriaceae (Rhodophyta) from the South Sandwich Islands, the Antarctic MICHAEL J. WYNNE University of Michigan Herbarium Ann Arbor, Michigan 1982 CONTRIBUTIONS FROM THE UNIVERSITY OF MICHIGAN HERBARIUM Volume 15 A Monograph of the Genus Petxotoa OT DIS Mace) CHRISTIANE ANDERSON l Notes on Neotropical Malpighiaceae—I WILLIAM R. ANDERSON — 93 On Meiothecium (Sematophyllaceae) WILLIAM R. BUCK _ 137 The North American Distribution of the Genus Dryopteris HOMAS M. CARLSON & W. H. WAGNER, JR. 141 The Itinerary of Mary Strong Clemens in Queensland, Australia A. M. CARTER — 163 The Genus Mirandea (Acanthaceae) THOMAS F. DANIEL 171 Further Notes on Steccherinum crassiusculum (Hydnaceae) KENNETH A. HARRISON & CLARK OVREBO ~ 177 The New Synantherology vs. Eupatorium in Nueva Galicia ROGERS McCVAUGH _ 181 Compositarum Mexicanarum Pugillo Supplementum ROGERS MCVAUGH _ 191 The Ferns of Steere and Harrington MICHAEL G. PRICE — 197 Linum McVaughii, a New Species from Mexico C. M. ROGERS — 205 Comments on the Mielichhoferioideae of Central America with the Description of a New Species of Synthetodontium ONATHAN SHAW & HOWARD CRUM_— 209 Pohlia Hedw. (Musci) in North and Central America and the West Indies ONATHAN SHAW) 219 Selected species of Leucocoprinus from the Southeastern United States HELEN V. SMITH & NANCY S. WEBER 297 A New Species of Rhynchospora (Cyperaceae) from Venezuela WM. WAYT THOMAS © 311 Botrychium rugulosum (Ophioglossaceae), a Newly Recognized Species of Evergreen Grapefern in the Great Lakes Area of North America W. H. WAGNER, JR. & F.S. WAGNER 315 Observations on Four Species of Delesseriaceae (Rhodophyta) from the South Sandwich Islands, the Antarctic MICHAEL J. WYNNE = 325 University of Michigan Herbarium Ann Arbor, Michigan 1982 CONTRIBUTIONS FROM THE UNIVESITY OF MICHIGAN HERBARIUM Volume 15 Editor: Anton A. Reznicek Editorial Committee: William R. Anderson, Howard Crum, Robert L. Shaffer Part of the cost of publication of this volume was provided by funds from NSF grant DEB 79-04946 to Christiane Anderson, DEB 81-03522 to William R. Anderson and DEB 80-04178 to Rogers McVaugh. Previous numbers of the Contributions are listed at the end of this issue. For information address the Director, University of Michigan Herbarium, North University Building, Ann Arbor, Michigan 48109, U.S.A. For issues previous to Volume 9, see inside back cover of Volume 14. December 13, 1982 Contr. Univ. Mich. Herb. 15:1-92. 1982. A MONOGRAPH OF THE GENUS PEIXOTOA (MALPIGHIACEAE) Christiane Anderson Herbarium The University of Michigan Ann Arbor, Michigan 48109 INTRODUCTION Peixotoa is a genus of 28 species of yellow-flowered, wing-fruited vines and small shrubs whose center of diversity is the Planalto of Brazil. It is a very homogeneous assemblage that is characterized by large, cordate stipules, com- pound inflorescences composed of 4-flowered umbels, and a uniseriate androe- cium of alternating stamens and staminodes. Within the family this is a relatively recently evolved group in which the species have not greatly diverged. Attempts at studying Peixotoa were limited in the past by a paucity of good material and complicated by the occurrence of apomixis, which has not been suspected until now. [The most recent monograph (Niedenzu, 1928) contained little information beyond that presented in Jussieu’s study of 1843 and an unsatisfactory key. It is hoped that this treatment will provide a beginning toward a better understand- ing of this difficult genus. TAXONOMIC HISTORY St. Hilaire’s collections from central Brazil included three species of Malpighiaceae which Adrien de Jussieu grouped into a new genus, Peixotoa, in his treatment of that family for St. Hilaire’s Flora Brasiliae Meridionalis in 1832. The name commemorates Domingos Ribeiro dos Guimaraens Peixoto, a physi- cian who had published “Dissertation sur les médicamens brésiliens” in Paris in 1830. Grisebach published three more species in 1839 based on Sellow collec- tions. In 1840 Jussieu, who by then had seen collections of Claussen, Gaudi- chaud, Martius, and Salzmann, added seven more names when he published his “Malpighiacearum synopsis,” which preceded the comprehensive monograph of the family of 1843. Turczaninow recognized a Blanchet collection as another new species in 1858. Kuntze described P. cordobensis from Argentina in 1898. However, his type is now included in Cordobia argentea (Griseb.) Niedenzu. Few collections and almost none from additional geographical areas had been made by the time Niedenzu published his monograph of Pezxotoa in 1912. The limited material available to him was often scanty and in poor condition. In his treat- ment of the genus for Das Pflanzenreich (1928) he recognized 11 species and one variety in addition to numerous forms. With the exception of a new name pub- lished by Ferreira in 1969, little work has been done since Niedenzu. However, extensive field work within the last 30 years in the Planalto and adjacent regions has yielded a large number of collections of Petxotoa, including an astonishing number of novelties. In this treatment I recognize twenty-eight species, of which eighteen are new. RELATIONSHIPS WITHIN THE FAMILY Niedenzu (1928) divided the Malpighiaceae into two subfamilies, the Planitorae, characterized by a flat torus and wingless fruits, and the Pyramidotorae, characterized by a pyramidal torus and winged fruits. Morton (1968) substituted names based on included genera, since Niedenzu’s names were not formed in accordance with the Code of Nomenclature. The Pyramidotorae thus became the Gaudichaudioideae. Niedenzu divided his Pyramidotorae into two tribes, the Hiraeeae, for genera in which the lateral wings are larger than the dorsal wing, and the Banisterieae in which the dorsal wing exceeds the lateral two. He placed Peixotoa into the latter next to Banisteriopsis (his “Banisteria”). Since Niedenzu based his scheme strictly on the torus and wing characters, it contains a number of unnatural assemblages. W. R. Anderson’s current investigation of the infra- familial structure of the Malpighiaceae will lead to changes in the present group- ings (see Anderson, 1978). However, no matter how the family may eventually be subdivided, Peixotoa will be retained next to Banistertopsis. This large and diverse genus is the most likely ancestor for the homogeneous, advanced Perxotoa. With the exception of the “stipules,” 1.e. the two structures resulting from the fusion of the interpetiolar stipules, and the presence of the characteristic staminodes, all generic characters of Peixotoa occur in Banisteriopsis as circum- scribed by Gates in her recent monograph (1982). While no single species shares all these characters, the following are found in the subgenus Hemiramma (sensu Gates): a vining habit; leaf glands borne on the lamina or petiole; 4-flowered umbels arranged in compound dichasia; sessile pedicels; calyx glands borne on the free part of the sepal and attached by their entire length; a corolla of yellow, glabrous petals with the posterior petal borne erect; glabrous anthers with the glandular connectives composed of large, inflated polygonal cells; a samara with a pair of lateral wings parallel to the areole; straight styles with capitate alae In some species of Banisteriopsis the glandular connective becomes greatly e larged while the size of the pollen sacs decreases, but completely sterile stami- nodes are not found. Yet, the pattern of reduction in the androecium is the same as in Peixotoa in that the change takes place in the stamens opposite the sepals. The composition of the androecium, 1.e. fertile stamens opposite the petals alternating with sterile staminodes opposite the sepals, is constant throughout Peixotoa and was probably already established in the ances monotypic genera, Cordobia, Mionandra, and ie from the southern edge of the family’s range might be considered closely related to Peixotoa. In these three taxa the “stipules” a consist of the interpetiolar supules which have fused into two cordate or triangular structures. In Cordobia and Gallardoa these may be bifid or entire with an acute or notched apex. Those of Mionandra are always bifid. In all three genera the inflorescence is reduced to a single flower with yellow, glabrous petals. Each androecium is similar to that of Peixotoa in that the stamens opposite the sepals are reduced to sterile structures. In Cordobia only tiny filaments are retained, while in Gallardoa and Mionandra the filaments, which are half as long or less than the stamen filaments, bear small apical glands. These glands are fusiform in Gallardoa and rhomboid in Mionandra. Vhe pollen of Cordobia is like that of other Banisterioid genera, while that of Mionandra and Gallardoa is highly atypical (S. R. Lowrie, pers. comm.). One reason for grouping Cordobia with Peixotoa is its Banisterioid samara with its large dorsal wing and the usually present pair of lateral winglets. In the samara of Gallardoa the dorsal wing is reduced to a crest while the lateral wings are greatly enlar ged. In Mionandra the change from a wind-dispersed samara to a detritus- fruit is reflected by the narrow, highly irregular crests that cover the mericarps. The vesture in all three genera is sericeous; the ‘T-shaped hairs typical of Peixotoa are lacking. The erect, capitate styles of Mionandra are similar to those of some — —_ 3 species of Peixotoa. Vhose of Cordobia and Gallardoa are somewhat curved and bear most unusual, internal stigmas. In Cordobia the stigma is borne laterally along the broad edge of the apically flattened style. In Gallardoa the stigma 1s associated with an appendage, similar to the condition in Stigmaphyllon. Though Cordobia, Gallardoa, and Mionandra have several characters in com- mon with Perxotoa, I do not believe that these similarities form evidence that any or all of the three genera should be included in Peixotoa or are derived from it. The atypical inflorescences and the peculiar style of Cordobia and Gallardoa point toward a different origin. It seems more probable that they also arose from ancestors in Banisteriopsis. ANATOMY AND MORPHOLOGY Hair Types. The two basic hair types found in Peixotoa are Y-shaped and T-shaped. Pubescence composed of Y-shaped hairs is termed velutinous (Fig. la) and may occur on any part of the plant other than on the lower surface of the stem leaves. In the ovary and on the samara the stalks of these hairs are very short or the hairs may be subsessile. The T-shaped hairs have an unusually long stalk, which in most species is 0.2—0.5 mm long. In P. gardneri and P. hirta the stalks are 0.5—1 mm or more long. While standard in Peixotoa these hairs are only infrequently encountered in the rest of the family. When the stalk is sinuous or twisted and the trabecula is wavy or curled the resulting vesture is called tomen- tose (Fig. Ib). If the hairs are very densely intertwined and so numerous that the epidermis is obscured, the term woolly is used. However, no term exists that describes vesture composed of T-shaped hairs in which the trabecula and stalk are straight (Fig. Ic). Little purpose is served by coining yet another term. For the sake of clarity, surfaces covered with such hairs are here described as “pubescent with T-shaped hairs.” P. adenopoda and P. sericea are the only species in the genus whose stem leaves have the true sericeous vesture that is a common condition in the rest of the family. These hairs consist of a long, straight, stiff, medifixed trabecula which is sessile or subsessile (Fig. Id). In Peixotoa such hairs are also found on stipules and less frequently on inflorescence leaves. Habit. While a small number of species are true vines with stems up to 8 m long, the majority are small shrubs ca I—2.5 m tall. Yet, many of these shrubs form ascending or scandent branches that twine at the tips or may even switch to a vining habit if a suitable support is available. This flexibility suggests that the Banisteriopsis ancestor was probably a vine. P. andersonii is unusual in that it forms procumbent stems up to 4 m long that send up erect flowering axes. The stems of both vines and shrubs are terete and generally velutinous or sometimes tomentulose or both when young. The pubescence is generally abraded from the older parts. In P. glabra, P. hatschbachii, and P. psilophylla even the relatively young vegetative branches that bear inflorescences are glabrous at the time of flowering. Stipules. The strikingly large stipules are a generic character in Peixotoa. Their presence or the characteristic stipule-scar permits even sterile material to be readily assigned to this genus. Each “stipule” is actually the product of fusion of the two opposing interpetiolar stipules. The two large, cordate structures that result from this fusion are decussate to the pair of leaves at that node. The stipules act as valvate bud scales. They may be glabrous or variously pubescent but are always finely and densely tomentulose along the margins. These mar- ginal hairs interlock and act as a kind of seal when the stipules enclose the young shoot. The sides may be flat or recurved. Each stipule is commonly entire with an acute or, especially in the larger ones, notched apex or it may be bifid. While the stipules are necessarily concave when they act as bud scales, they usually become flat as the shoot matures. Yet in some species, especially in the inflorescence, the ] > Types and eae ams of ane ence Structure and Flower (see section Anatomy and Morphology) Hairs representative of these vesture types: a) velutinous, b) tomentose, c) “pubescent with ‘T-shaped hie d) sericeous. aoc structure: e) hypothetical transition eee pedunculate to sessile flower; f) typical umbel subtended by a pair of leaves and a pair of large ules; g) “subsidiary axis’ Miseated between the secondary axis and the leaf; h-k) transition from re ieee (1- umbellate) terminal node to 12-flowered (3-umbellate) terminal node: h) all umbels 4-flowered, each umbel subtended by a pair of leaves and a pair of stipules, i-]) suppression of secondary axes cel ea saved a loss « of leaves and stipules, terminal node 12-flowered, lateral nodes with a sessile umbel xil, k) variation of condition in J: ae umbels borne on secondary peduncles but not es i leaves and stipules. Floral Diagrat ~ Jo concave shape is retained. The stipules are usually largest on the stem and become decreasingly smaller toward the distal portion of the inflorescence. In a few cases the inflorescence stipules equal those of the stem or may exceed them. They are deciduous in most species but are persistent in some. In P. cordistipula they are retained in the inflorescence even after the leaves and pedicels are shed. In very large inflorescences the stipules of the most distal nodes may be lacerated or split into narrow strips of tissue, or sometimes the stipules are fused. In these cases it is common for the inflorescence leaves to be suppressed entirely. Leaves. Peixotoa, like most Malpighiaceae, has opposite, decussate, simple, entire leaves. The petioles are terete and usually short, often less than | cm long. In some species, especially in vines, the petioles may be longer but never more than 3.5 cm long. The laminas are most commonly elliptical or ovate, but may also be obovate, rhombic, suborbicular, lanceolate or oblanceolate. While the apex 1s apiculate or sometimes acuminate, the tip is often broken off in the larger leaves, which then appear to be rounded or obtuse. The base is usually cordate or sometimes subtruncate. In P. catarinensis the larger leaves are cordate but the sinuses may be truncate. In the stem leaves of P. sericea the bases are cuneate. The margins are at least slightly recurved. Thick, almost coriaceous laminas are rugose. The veins are prominent below in all species. The vesture is variable. The laminas are sparsely to densely velutinous or glabrous above. Exceptions are P. cipoana, in which the laminas are sometimes tomentulose above, and P. hirta, in which they may bear T-shaped hairs and/or be velutinous above. Below, the laminas vary from glabrous to sparsely to densely pubescent with T- shaped hairs to sparsely to densely tomentose to woolly. P. adenopoda and P. sericea are the only species in which the stem leaves are sparsely sericeous above and more densely so below. The sericeous condition is common in the family but excep- tional in Peixotoa. Asa rule the inflorescence leaves become gradually smaller toward the distal nodes, but sometimes the change is abrupt. The most proximal are usually similar to the most distal stem leaves. They usually change in shape toward the more distal nodes from lanceolate or triangular to linear and subulate. The presence or absence of a petiole in the more distal leaves is a good character for most species. The leaf glands are always borne on the lower surface. Typically, each lamina bears two discoid or slightly oblong, prominent glands at the base, one on each side of the costa. Sometimes the glands are borne halfway on the petiole or rarely entirely on the petiole or rarely on the laminar tissue but very near the base and the costa. In the more distal inflorescence leaves the glands are usually disproportionately large and commonly cause the smallest leaves to become almost conduplicate. One to four additional smaller glands may be present. Most commonly they occur as a second pair adjacent to the larger glands. These additional glands are found especially in the inflorescence leaves. At times they become variously fused, especially so in the inflorescence leaves, where, in the most extreme cases, a band of glandular tissue extends across the base of the lamina. In P. leptoclada and P. paludosa the more distal inflorescence leaves may have a glandular tip. In P. adenopoda, P. catarinensis, P. paludosa, and P. spinensis the leaf glands are borne on the proximal ¥ of the surface of the lamina, usually approximately equidistant from the base and costa. In P. sericea the glands are borne above the base and at or very close to the costa but at different levels, rather than equidistant, from the base. In P. jussiewana the glands may be at the base or on the laminar surface. The larger leaves of P. leptoclada may have one pair at the base and a second pair on the lamina, or on the same plant some of the lar gest leaves may bear a pair of glands on the lamina only and lack the basal pair. In the more distal inflorescence leaves the glands are always basal regard- less of the position of the glands on the stem leaves. Though prominent, the 6 glands of most species are sessile. [In the stem leaves of P. adenopoda and P. hatschbachu they are stalked. ‘This stalk is ca 0.8-1.2 mm long. Glands of the more distal inflorescence leaves of these species are sessile. In some species one or both glands are sometimes lacking. Inflorescence. The basic unit of the inflorescence is a 4-flowered umbel. This is so thoroughly established in the genus that it was probably already present in the Banisteriopsis ancestor. Each umbel is an extremely reduced 4-flowered raceme [the pseudoraceme of Cuatrecasas (1958) and Anderson (1981)] in which the axis is foreshortened and in which the primary peduncle is lost (Fig. le). Uhis legacy is reflected in the maturation pattern of the four flowers: two are always at least slightly older than the other two. T ypically each umbel ts sessile or subsessile at the node and subtended by a pair of leaves and by the pair of stipules which functioned as bud scales (Fig. If). The sessile pedicel of each flower is subtended by a much reduced, non-vascularized bract and a pair of equally reduced bracteoles. These scale-like structures vary from triangular to oblong to semicircular to subrectangular and are glabrous or variously pubes- cent. ‘They are persistent though in some species occasionally absent. “The pedicel is extremely variable in size in all species. It may increase up to 4 of its length in fruit and always becomes a little stouter. [tis always pubescent even in species which are otherwise glabrous. The umbels are arranged into compound dichasia and thyrses, which are terminal and/or axillary. Solitary umbels are common only in P. axillaris and P. hispidula. In large inflorescences, especially in vines, additional axes are some- times produced between the inflorescence branch and the leaf (Fig. 1g). Such subsidiary axes are always much smaller than the axes with which they are associated, In P. adenopoda, P. catarinensis, P. parviflora, P. tomentosa and a few indi- viduals of P. reticulata, the secondary inflorescence axes with their stipules and leaves are suppressed at the more distal nodes. As a result of this change the umbel borne on such a secondary axis now becomes sessile in the axil at the branch point. At the terminal node the central umbel is flanked by two sessile umbels, which results in a three-umbellate Seen (Fig. [h-j). Phe terminal nodes of these species are thus always 12-flowered. Commonly, when such a reduction of the inflorescence branches has ae place, the umbel instead of being sessile is supported by a secondary peduncle (Fig. 1k). This secondary peduncle differs from a typical inflorescence axis in that it lacks the stipules and leaves. Its length 1s variable within an individual inflorescence. Sometimes the bracts occur down on the secondary peduncle. Peixotoa octoflora is unique in the genus in that each umbel is composed of four pairs of flowers. In each pair one pedicel is sessile while the second pedicel is borne on a short (1-2.5 mm long) primary peduncle (Fig. 21b). ‘The umbels are grouped into a compound inflorescence typical for Peixotoa. A primary peduncle is otherwise only very rarely encountered in some individuals of P. reticulata. Flower. Vhe terminology used here to describe the various floral parts fol- lows that of Niedenzu (1928), Cuatrecasas (1958), and Anderson (1981), and is illustrated in Fig. I. The flowers of Pe:xotoa are bilaterally symmetrical with the odd, fifth petal in the posterior position. The remaining petals are grouped into an anterior-lateral pair and a posterior-lateral pair. These petals are reflexed between the sepals, while the posterior petal is erect. ‘The eglandular sepal ts designated the anterior sepal, while the other four, like the lateral petals, are grouped into an anterior-lateral pair and a posterior-lateral pair. ‘The plane of symmetry passes through the posterior petal and the anterior sepal. The posi- tion of the stamens and staminodes is indicated by the perianth part they oppose. ‘Thus, the stamen opposite the posterior petal is called the posterior stamen. The i 3-carpellate ovary is placed so that the plane of symmetry passes through the anterior carpel and between the two posterior carpels. The designation of the style follows that of its carpel, i.e. the anterior style belongs to the anterior carpel. The posterior carpels and styles are alike. No distinction is made between them. Within the genus the size and to a lesser extent the shape of the floral parts vary, but the arrangement is constant and permits visits by oil-collecting bees in the manner described by Vogel (1974) and Anderson (1979). In brief, the bee lands on the center of the flower, orients itself by the posterior petal, the “flag,” whose claw it grasps with its mandibles, and with its front and middle legs reaches between the lateral petals to scrape oil from the calyx glands. Some bees also collect pollen. The oil and pollen are transferred to the hairy hind legs and carried to the nest to serve as food for the larvae. During the bee’s visit pollen is rubbed on the underside of the bee and presumably transferred to the stigmas of the next flower visited. The claw of the flag petal of a visited flower commonly shows two lateral indentations which are caused by the pinching grasp of the bee’s mandibles. Sepals. The five valvate sepals are ovate or broadly oblong with an acute apex which becomes recurved or revolute when the flower opens. They are glabrous or variously pubescent adaxially and densely velutinous abaxially. All are densely tomentulose along the margins. These hairs interlock when the sepals are closed in bud. All except the anterior sepal bear a pair of oblong glands on the proximal 1/3-1/2 of the free part of the sepal. These glands are green in nature and turn dark brown upon drying. The anterior sepal is usually a little narrower than the lateral four. Petals. The five petals are yellow, glabrous, and clawed. The four lateral petals are of equal shape and usually of equal size, though the anterior-lateral two are sometimes a little larger than the posterior-lateral two. The flat limb varies from elliptical to obovate to orbicular. P. andersonii is unique in that the limbs of its lateral petals are oblanceolate. The margin is usually fimbriate, sometimes denticulate, or sometimes both. The fimbriae and teeth commonly bear tiny glands. The posterior petal, the “flag,” differs markedly from the other four. The claw is much longer than and about twice as thick as that of the lateral petals and is continuous with the limb. The posterior limb is also flat, but usually notably smaller than and often of a different shape than the lateral limbs. The margin is either entirely capitate-glandular or only so in the proximal 1/3—4/5 with the distal part fimbriate- or denticulate-glandular. ndroecium. The uniseriate androecium is composed of five fertile stamens, opposite the petals, alternating with five sterile staminodes, opposite the sepals. The filaments are connate at the base. The four lateral stamen fila- ments are usually equally long, but sometimes the anterior-lateral two exceed the posterior-lateral two a little. They are straight or, if very long, curved in the distal 4 toward the ovary. The posterior stamen filament is usually much shorter than the lateral four and is commonly arced toward the posterior petal. The anthers are basifixed and glabrous. They are all of equal size and have longi- tudinal, introrse pollen sacs. A prominent glandular connective projects ca 0.3— 0.5 mm above the locules; it is lacking only in P. anadenanthera. The pollen is apparently released at or just prior to the opening of the flower. In apomictic plants the pollen sacs usually remain closed. Stuart R. Lowrie, who is completing a comprehensive study of pollen in the Malpighiaceae, has kindly provided me with his findings in Peixotoa. The pollen grains of Peixotoa are of the cuboidal type that Lowrie believes is unique to the family and typical of most of the genera in Niedenzu’s tribe Banisteriae. In Peixotoa each grain has 12 rugae and 6 pores (Fig. 2). The pores are not placed at the corners of the cube but slightly removed from them. Lowrie determined that they are located at the points of coincidence of an octahedron inscribed in the IG. 2. Pollen of Peixotoa. Left, P. goiana, X 1300 (Irwin et al. 17562), right, P. parviflora, x 1300 Te 9225). SEM photographs by S. R. Lowrie. cube. Though there is considerable variation in the sculpturing of the pollen grains among the species of other genera, the pollen of the various species ¢ Peixotoa is relatively uniform. ‘Vhe sizes given in the species descriptions, based on measurements of pollen grains placed in water, range from 42 to 74 ym. The slender staminode filaments are about half as thick as the stamen fila- ments and taper toward the apex. Each filament bears a basifixed, obovoid or sometimes spherical gland at the apex. The adaxial face of the gland has a groove in the proximal 1/3-3/4 (-7/8). The apex and abaxial surface are either smooth or the apex has a shallow to deep indentation which is decurrent on the abaxial face. If the indentation is very deep, the glands appear cordate. In P. irwind the apex is smooth, but the abaxial face has a shallow and broad depres- sion. Individual glands bearing one or even two small locules occur in P. magnift- ca, but are rare. Such locules are 0.7—1 mm long, contain largely aborted pollen, and remain closed. Vhe anterior staminode is usually the largest but may equal the anterior-lateral two. The posterior-lateral two are usually the smallest. While the anterior and anterior-lateral filaments are erect, the posterior-lateral two are commonly inflexed between the posterior styles. Gynoecium. The 3- -carpellate, 3-loculate ovary is densely velutinous. Each locule is fertile and contains one anatropous, pendent ov ule. The styles are free to the base and persistent in fruit. Phe anterior style is longer or shorter than or equal to the posterior two but always a little stouter. It may be erect but more commonly is arced or inclined toward the posterior petal, rarely toward the anterior sepal. The posterior styles are usually divergent and variously inclined toward the posterior-lateral petals. Dna. parviflora they are curved toward the anterior-lateral petals. All styles are glabrous or may be very sparsely velutinous, especially near the base. ‘The stigmas are terminal and in nearly all species capitate. ‘The exceptions are P. jussieuana and P. magnifica, in which the anterior stigma is oblong and borne on the abaxial surface at the apex of a long, strongly curved style. Fruits. Vhe fruits of Peixotoa ave schizocarps in which each mericarp devel- ops into a velutinous samara. The samaras are grouped around a pyramidal torus and at matur ity are suspended by a carpophore. Each samara bears a large dorsal wing which ts thickened along the curved, sigmoid, or arced, entire upper (adaxial) margin. The lower margin may be straight, sigmoid or semicircular 9 and, though at times entire, is commonly somewhat erose. The dorsal Wing Is usually broadest at the middle or beyond. Each nut also bears two lateral wing- lets, which are much smaller than the dorsal wing. These are usually rectangular, and entire, erose or coarsely dentate. In some species the lateral wings are semicircular and recurved and give the samara an inflated appearance. In P. hatschbachii the lateral wings are absent or reduced to a narrow strip of tissue less than 1.5 mm wide or to a crest. The dorsal and lateral wings are confluent at the base of the nut and sometimes extend into a tooth. The ovoid nut has a triangu- lar or cordate, concave areole. At maturity the seed fills the locule. The embryo has obspatulate, unequal cotyledons. The outer cotyledon is always larger and folded over the inner one. The inner cotyledon may also be folded or may be straight. Sometimes the apices are twisted. CHROMOSOME NUMBERS Counts of n=10 have been obtained for P. glabra (Anderson 11549) (W. R. Anderson, pers. comm.) and P. hispidula (Ormond et al., 1981). Since n= 10 is the base number for Banisteriopsis (Gates, 1982), the most probable ancestor of Peixotoa, it 18 most likely that this is also the base number for Peixotoa. Counts have also been made of pollen mother cells of the apomictic P. reticulata (Anderson 11790). Here 2n=30, but the separation at anaphase I/telophase I is highly irregular. Figures of 17 and 13, 16 and 14 as well as of 15 and 15 pairs occur (W. R. Anderson, pers. comin.). The pollen grains of the voucher, as in the great majority of the specimens of P. reticulata, are largely aborted. P. reticulata may be a triploid that resulted from a cross between a diploid and a tetraploid. N=15 is otherwise unknown in the Malpighiaceae. DISTRIBUTION Peixotoa is composed largely of small shrubs from the cerrados of the Brazil- ian Planalto and adjacent areas. Vines are commonly found also in woodlands, gallery forests and at forest margins. Three species from Bahia, P. adenopoda, P. paludosa, and P. sericea, and also P. Jussieuana are reported from caatingas. catarinensis and P. hispidula are coastal species that grow in the coastal forests, restingas, on dunes, and on beaches. Like many other Planalto groups, Peixotoa is characterized by a high degree of endemism. Twenty of the twenty-eight species are found in Minas Gerais and Goias, of which only two are common to both states. Of the twelve occurring in Minas Gerais, eight are known only from that state. Of these, seven have been reported only from the Serra do Espinhago. Five of the eight species occurring in Goias are endemics. Four species are known only from Bahia. Only one species, a coastal endemic, is found in Santa Catarina. Only five species have relatively wide ranges. P. hispidula, the other coastal species, ranges from Bahia to Rio de Janeiro. It has also been collected twice from Sao Paulo. P. parviflora occurs in central and southern Minas Gerais, Rio de Janeiro, S40 Paulo and Parana. P. magnifica is a vine of central and southern Goids, central Mato Grosso and eastern Bolivia. The range of P. jussteuana, a close relative, extends from Ceara to Piaur and Maranhao. It has been collected once in Parana (an atypical plant). The most widespread species is the apomictic P. reticulata. It is common from southern Minas Gerais, southern Goias and southern Mato Grosso to Sao Paulo, Parana, eastern Paraguay, and eastern Bolivia. APOMIXIS Apomixis has not been previously reported in Peixotoa. The findings noted here are only a preliminary report. In this treatment “suspected apomicts” will 10 simply be called “apomicts” for the sake of brevity but also with the understand- ing that only a thorough study to be undertaken in the future will reveal the mechanisms involved. Apomixis apparently occurs in five species. In P. leptoclada the apomicts are more robust than the sexual individuals but otherwise resemble them closely. There is no such gross morphologica al difference between the sexual specimens and the one known apomictic collection (Roth 1587) of P. paludosa. P. axillaris and P. octoflora, while clearly distinct taxa, are known only from apomictic material. Since both are from little-collected areas, sexual populations may well be dis- covered during future field work. The commonest Peixotoa is P. retic ulata, which doubtlessly owes its wide distribution to apom In the flowers of apomicts the anthers never open and contain largely aborted pollen grains that are 88-99% non-staining in cotton blue in lacto- phenol. ‘The few grains that do stain are often large and are perhaps unreduced pollen mother cells. Exceptions have been noted only in some individuals of P. reticulata. In Krapovickas & Cristobal 34372 almost 100% of the pollen, contained in closed anthers, does stain; yet, the grains are abnormal in that they are small and very thick-walled. In a few specimens some flowers do have open anthers which often contain a lower percentage of non-staining grains than do closed anthers. For example, in Fiebrig 4590 pollen from the open anthers is 45% non-staining, while pollen from closed anthers is 94% non-staining. However, in Hatschbach 21770 the pollen from closed as well as from the few open anthers is only ca 3% non-staining. Only two collections are known in which all flowers have all anthers open. In Mexia 5798, the staining of the pollen is highly variable. The percentage of grains that do not stain ranges from 45-89% per anther. In the second collection, Occhiont 5775, nearly all the grains do stain. Perhaps this 1s a rare sexual individual. Examination of pollen mother cells of P. reticulata (Anderson 11590) revealed that while 2n=30, the separation at anaphase I/ telophase I is highly irregular with 17 and 13, 16 and 14, or 15; ind 15 pairs segregating to the opposite poles (W. R. Anderson, pers. comm.). Such irregular meiosis would account for the high percentage of aborted pollen. Yet, if occa- sionally the separation is regular, some functional pollen could be produced. Since the anthers do open sometimes in some individuals, it may be possible for an apomict to function as a male parent. Of course, it is also possible that an apomuct may function as a female parent. In areas where the range of P. reticulata overlaps with that of sexual species, some hybridization apparently does take place. Anderson 8564 from the Gouveia- Diamantina area in Minas Gerais has the glabrous foliage and the indented staminode glands of P. glabra but otherwise is like specimens of P. reticulata in that area. A number of collections from southern Goias and the Distrito Federal are intermediate in various flower characters between P. reticulata and P. goiana. Hatschbach 2465 from ‘Verenos, Mato Grosso, has fruits similar to those of P. magnifica in that the dorsal and the lateral wings are relatively large and extend into a long (ca 0.5 cm) tooth below the nut. Other collections from southern Mato Grosso and also from Paraguay have some attributes of P. cordistipula, A species of central Mato Grosso and eastern Bolivia. The readily recognizable regional types have perhaps originated as sterile hybrids that were apomictically propagated. (he apomicts are morphologically highly variable, especially in the shape and size of the flower. On the other hand, the differences in curvature and size, especially in length, of the stamens, staminodes and the styles are minimal. In sexual species these differences are generally constant. In apomictic individuals, especially of P. reticulata, it is not unusual to encounter flowers in which the stamens, staminodes and styles are subequal and erect. Teratologies, such as flowers with extra floral parts, have been noted in apomicts but are exceedingly 1] rare in sexual species. All of these morphological changes in the apomictic plants may reflect the relative unimportance of presenting the flower to the pollinator to assure seed set. The fruit set in apomictic plants tends to be unusually high, often 100% or nearly so. Apparently maturation of the fruit begins soon after the flowers open. In /rwin 17830 it seems to begin even earlier. Here the petals do not expand but are lifted like a cap and then shed by the developing fruit. Study of the seeds shows that polyembryony is sometimes involved. Often the seed contains both a normal embryo and an additional, smaller one. This extra embryo is commonly misshapen or may be an unformed mass of tissue. TAXONOMY Peixotoa Adr. Jussieu in St. Hilaire, Fl. Bras. Mer. 3: 60. 1832 [1833]. Vines or shrubs or subshrubs with erect, ascending, scandent, pendent, or procumbent branches. Vegetative branches glabrous or velutinous or tomentose When young, the pubescence commonly abraded from older parts. Stipules large, interpetiolarly connate, the pair cordate or triangular, entire with the apex acute or notched or bifid, variously pubescent or glabrous adaxially and abaxi- ally, persistent or deciduous, margins tomentulose, flat or recurved; inflores- cence stipules often concave, enclosing immature umbels. Leaves Opposite and decussate, petiolate, the lamina simple, often rugose, ovate, elliptical, obovate, rhombic, suborbicular, lanceolate, or oblanceolate, the apex apiculate or some- times acuminate, the base cordate or subtruncate, velutinous or glabrous above (sericeous in P. adenopoda and P. sericea, sometimes tomentulose in P. cipoana), pubescent with T-shaped hairs, tomentose, woolly or glabrous below (sericeous in P. adenopoda and P. sericea), the veins prominent below, margins entire, re- curved, with a pair of glands at the base at the costa or borne in the proximal 4% of the lamina away from the base and costa, sometimes with | to 4 smaller, additional glands, these sometimes variously fused, or the glands absent; glands discoid or slightly oblong, prominent, sessile (stalked in P. adenopoda and P. hatschbachi). Inflorescence leaves smaller than the cauline, petiolate or sessile, lanceolate, oblanceolate, triangular, linear, or subulate. Inflorescence terminal and/or axillary, of 4-flowered umbels (8-flowered in P. octoflora), borne in com- pound dichasia or thyrses with axes to the 6th order (or the umbel solitary in P. axillaris and P. hispidula), with 1 sessile umbel per node, or the distal lateral branches suppressed and then the terminal nodes with 3 umbels and the sub- terminal nodes with | or 2 umbels per leaf axil, these umbels sessile or borne on secondary peduncles, primary peduncles absent (present in P. octoflora). Bracts and bracteoles present and persistent, or absent, triangular, oblong, semicircu- lar, or subrectangular, variously pubescent or glabrous. Pedicels pubescent, always stouter and commonly longer in fruit. Sepals 5, valvate, ovate or broadly oblong, apex acute, recurved or revolute, variously pubescent or glabrous adaxially, densely velutinous abaxially, margins tomentulose, the anterior sepal eglandular, the 4 lateral sepals each with a pair of oblong glands entirely borne on the free part of the sepal in the proximal 4 or 4. Petals yellow, glabrous, clawed, the limb orbicular, obovate, or elliptical (oblanceolate in P. andersoniz), the 4 lateral petals equal or subequal, the margins fimbriate or denticulate or both, glandular or eglandular, the posterior petal with a thicker, longer claw and a usually smaller limb than the lateral petals, its margin capitate-glandular or only so in the proximal part, the distal section fimbriate- or denticulate- glandular. Androecium uniseriate, the filaments connate at the base; fertile stamens 5, opposite the petals, the 4 lateral filaments equal or subequal, erect, the posterior filament usually shorter, arced or inclined toward the posterior 12 petal or erect, anthers basifixed, glabrous, with a prominent glandular con- nective (absent in P. anadenanthera), pollen sacs longitudinal, introrse, pollen cuboidal, with 12 rugae and 6 pores. Staminodes 5, sterile, opposite the sepals, the filaments ca % as wide as the stamen filaments, tapered, the anterior and anterior-lateral filaments erect, the posterior-lateral filaments usually inflexed between the posterior styles, bearing apical glands (modified ster ile connectives), the glands basifixed, obovoid or spherical, glabrous, with a groove in the proximal 1/3—3/4 (-7/8) in the adaxial face, the apex smooth or with a shallow or deep indentation decurrent on the abaxial face (in P. rwini the apex smooth, the abaxial face with a broad, shallow depression), the anterior and anterior-lateral staminodes usuallly larger than the posterior-lateral 2. Ovary densely velutinous, three-carpellate, three-loculate, with one anatropous, pendent ovule. Styles 3, free to the base, glabrous or sparsely velutinous, especially near the base, the anterior style arced or inclined toward the posterior petal or erect, longer or shorter than or equal to the posterior two, the posterior styles usually divergent, curved or inclined toward the posterior-lateral petals (toward the anterior-lateral petals in P. parviflora), stigmas terminal, capitate (in P. magnifica and P. jussieuana the anterior stigma oblong and borne at the apex but laterally on the abaxial surface). Torus pyramidal. Fruit schizocarpic, each mericarp a samara, veluti- nous, suspended by a carpophore, with a large dorsal wing and 2 much smaller lateral winglets (these much reduced or absent in P. hatschbachi), the 3 wings sometimes forming a basal tooth, areole cordate or triangular, nut ovoid, the mature seed filling the locule, the embryo with unequal, obspatulate cotyledons, the large outer cotyledon folded over the smaller, folded or straight, inner cotyledon. Chromosome number n= 10 Lectotype. Peixotoa glabra Adr. Jussieu. NOTES ABOUT THE KEY TO THE SPECIES Peixotoa is a homogeneous genus whose species are often separated on the basis of microscopic characters. To use this key it is essential to have an under- standing of the structure of the flower and of the inflorescence. The reader who is not familiar with the genus is urged to read the section Anatomy and Morphol- ogy and to consult Fig. 1. he terminology used follows that of Niedenzu (1928), Cuatrecasas (1958), and Anderson (1981). It is necessary to revive dried flowers to ascertain the shapes and sizes of the petals, stamens, staminodes, and styles. For this purpose Pohl’s solution (Pohl, 1965) was found to be a superior wetting agent. Measurements for floral parts, except for the sepals, are taken from revived material. The measurements for the wings of the samara are indicated as follows. In the dorsal wing the distance from the attachment at the nut to the farthest margin Is the length”; the maximum measurement at right angles to this length is the “width.” For each lateral winglet, the maximum measurement from the attachment at the nut to the farthest margin is the “width”; the maxi- mum measurement at right angles to this width is the “height.” Since a large number of species are endemics, geographical ranges are listed in the key. The discussion following each species description cites the most distinguishing char- acters which help to separate that species from morphologically similar species and from those within its range. The reader may also find it useful to refer to the illustrations which are provided for each species. reticulata 1s an apparently apomictic and polyphyletic assemblage. ‘The anthers of all or of nearly all flowers remain closed. ‘Vhis condition is used here to key P. reticulata. Of all the collections examined only two have all anthers open. [This condition has been omitted from the key. Since P. reticulata is highly variable and apparently hybridizes with some sexual species, structuring the key 13 to include all the possible meee variation for this exceedingly rare condition would make the key extremely complicated and difficult to use. Therefore, the reader is cautioned that if his specimen, bearing open anthers, does not key and is from the range of P. reticulata, he may have another such rare individual. In such a case, it is suggested that for correct determination the description and illustrations of P. reticulata and of the sexual species of the same range be compared. KEY TO THE SPECIES OF PEIXOTOA 1. Lower surface of the stem leaves eaten or very sparsely pubescent with the few hairs re- stricted to the major veins and margin 2. Anthers cee (Minas Gerais) 25. P. reticulata. 2. Anthers oper 3. Glands of ae stem leaves stalked. (Goias) 13. P. hatschbachii. 3. Glands of the stem leaves sessile or absent. Inflorescence commonly of 1 or 3 or 5 umbels, occasionally more, 3° axes absent; all inflorescence leaves petiolate; limb of the latera petals 13-20 mm long; anterior and anterior-lateral staminode glands wider than long. (Coastal oe from Bahia to Sao Paulo) 15. P. hispidula. 4. Inflorescence of more than 5 umbels, 3° axes usually present; the more dict al inflorescence leaves sessile or subsessile; limb of the lateral petals 7-14 (-16) mm long; anterior and anterior-lateral staminode glands longer than wide or as long as Wi 5. as of the posterior petal (4-) 5.2-7.5mm long, (3.2-) 3.84.5 mm ues oblong, elliptical, or obovate; limbs of the lateral petals with a fimbriate largin. 6. Petioles glabrous; limbs of the lateral petals orbicular; staminode glands of equal size; anterior style shorter than the posterior two. (Minas ne 11. P. glabra. 6. Petioles golden ee: limbs of the lateral petals obovate; anterior anc anterior-lateral staminode glands larger than the ea -lateral two; anterior style equal to ie posterior two. (Minas Gerais) 16. P. irwinii. 5. Limb of the posterior petal 7.5-9 mm long, 6.8-10 mm wide, orbicular or obovate; limbs of the lateral petals with a fimbriate and/or denticulate margin. 7. Pedicels and inflorescence leaves densely white velutinous; limb of the posterior petal 8.5-9 mm long, 8.5-10 mm wide, obovate: limbs of the lateral petals 11-14.5 mm wide, the margin denticulate; filaments of the ae ior-lateral staminodes 3.4-3.8 mm long, the glands 1.2-1.3 mm Ae eae mm wide; anterior style inclined toward the posterior cee Se Grosso) P. poilophyila de Pedicels and inflorescence leaves rea Sean A cuemes white vel 1tI- nous; limb of the posterior petal 7.5-8.3 m ie ng, mm_ wide, orbicular; fra of the lateral petals Q- 12 mm w the racer fimbriate and/or danneuiaee filaments of the posterior ce staminodes 2.8— an mm long, the glands 1-1.1mm_ long, 0.9-l] mm_ wide; anterio erect. (Goias 2neP: poi 1. Lower pees Bi ie stem leaves pubescent, hairs not restricted to the major veins and ee eanieel inflorescence nodes bearing more than 4 flower 9. Glands of the stem leaves borne on the surface of the lamina away from the base and costa, or absen 10. Glands of the stem leaves stalked, 0.8-1.5 mm in diameter; anterior and anterior-lateral staminode glands with an indentation across the apex an de- current on the aes face. (B aa Be seats 10. Glands of the stem leaves sessile, 0.2-0.7 mm in diameter; anterior and : terior-lateral cee glands smooth. (Santa Catarina He Ps Sete Glands of the stem leaves borne at the base of the svat at the costa or halfway on the petiole, or rarely on the petiole just below the lar Il. Each eet umbel 8-flowered, consisting of i ae of flowers, each pair with one pedicel sessile and one pedicel borne on a short primary peduncle 1—-2.5 mm long). (Mato Grosso) 21. P. octoflora. 11. Each individual umbel 4-flowered, all pedicels sessile (rarely 1 pedicel in an umbel with a primary peduncle in P. reticulata). —_ ad 12. Anthers of all or of almost all flowers closed. (Minas Gerais, Sao Paulo, Parana, southern Goias, southern and central Mato Grosso, eastern Para guay, eastern Bolivia) pasa ek ear Anthers of all eek 13. L ob of the ee pet “ eases: petioles of the stem leaves 8— 13 mm long; anterior and anterior-lateral staminode glands wa an indentation across the mane and decurrent on the abaxial fac (Minas Gerai ; onde Limb of the lateral De fee ete broadly sea Wa or broadly 0) vate; petioles of the stem leav —2(0) (-32) mm long; anterior ae anterior-lateral stamin ee glands smooth. 14. Limb of the lateral petals (4.3—-) 5.2-9.5 mm long, (4—) 5-8.5 mm wide; leaves loosely pubescent with T-shaped hairs or loosely to- mentose below; lateral umbels always borne on secondary pedun- cles; all inflorescence leaves petiolate; posterior styles curved to- ward the anterior-lateral petals; pubescence of the se Aa _ NO —_ os a — axes commonly white (sometimes golden), sparse to moderate (rarely dense). (Minas Gerais, Rio de Janeiro, Sao ele. Age 23. P. parviflora. ie Limb of the lateral petals 9.5-12.3 mm long, 10-11 mm wide; woolly, tomentose, or sometimes pubescent with ‘T-shape hairs and tomentose below; lateral umbels sessile or borne on sec- ondary pec noes all or only the distal eumemen leaves sessile or subsessile, or sometimes all petiolate; posterior styles inclined toward the ater ior-lateral petals; pubescence of aie inflorescence axes gO olden, very dense and obscuring the epidermis. (Minas xeTals 28. P. tomentosa. 8. Verminal inflorescence nodes bearing 4 flowers. 1h; Inflor rescence borne ona short axillary Posi ey than | cm long), of | or ne ° axes less than 5 mm long or absent. (Goia 4. P. axillaris. Inflor. rescence terminal or if axillary borne on a shoot more than | cm long, of at least 3 umbels, usually many more (except commonly of | or 3 umbels in P Se een a axes more than 5 mm long. nthers of all or of almost all flowers closed. Leal glands borne on the surface of the lamina away from the base and the costa, or one or both nae absent; limb of the posterior petal 5 a 7mm ng. (Minas Gerais, Bahia oe ee ee Leaf glands borne at the base of the lamina at the costa or one ont petiole (sometimes in very large leaves e P. leptoclada ae a second pair of ands, or rarely only with one pair of glands, borne on the surface of the en away from the base and a peas of the ee ior petal 7.5— 12 mm long. 18. peeuial peduncles pega inflorescence sae persistent or some- —_ Or ~I ° mes deciduous at the most proximal nodes, large, even the most stl 7-17 mm One: ‘limb of the posterior ‘petal with a one andular margin. (Minas Gerias, Goids Pauper 18. routs peduncles nA to 6 mm long or absent; inflorescence stip- rior petal with the margin capitate-glandular in the proximal 1/2-4/5, fimbriate-glandular in the distal 1/5—1/2, or sometimes entirely capi- tate-glandular. (Minas Gerais, Sao Paulo, Parana, southern Goias, southern and central Mato Grosso, eastern Paraguay, eastern Bolivia) 25. P. reticulata. 16. Anthers of all flowers open. 9. Anthers lacking a glandular connective. (Goias) 2. P. anadenanthera. 19. Anthers bearing a glandular connective. 90. All or at least the most distal inflorescence leaves sessile. 2]. Laminas sericeous below, the hairs sessile or subsessile; base of the lamina cuneate. (Bahia) 26. P. sericea. 21. Laminas pubesc ent ele T-shaped hairs or tomentose or woolly below, the hairs stalked; base of the lamina cordate or sub- trun Zo. pear and anterior-lateral staminode glands with an in- entation across the apex and decurrent on the abaxial face 23. Stem leaves sparsely to densely pubescent with T-shaped hairs below; limb of the posterior peta (-9) mm wide, orbicular or rarely broadly obovate; aiike of the lateral petals with a fimbriate or res Sa ad terior-lateral staminode glands 1-1.1 long, 0.9- 1 mm wide. (Goias) 12. P. gowana. 23. Stem leaves tomentose or woolly below; limb of the pos- terior petal 4—6.2 mm wide, elliptical or narrowly obo- vate; limbs of the lateral petals with a fimbriate ne eae ee lateral staminode glands 1.2-1.4 mm_ long, 1.1-1.3 mm wide 24. Anterior and anterior-lateral staminode glands re- —1.6 mm long, 1.3-1.5 mm wide; stem leaves velutinous or tomentulose above, woolly be- yw; anterior style shorter than the posterior ne (Minas Gerais) 8. P. cipoana. 24. Anterior and = anterior-lateral staminode glands straight, 1.2—-1.4 mm long, 1.1-1.3 mm wide; stem an velutinous above, tomentose saa? oe t equally long. (Minas Gerais) . P. barnebyr. 22 Anterior 2 anterior- cl staminode glands sm 20 wide, obovate; limbs of the ea petals ea ge mm long, 8-8.5mm _ wide; anterior and nterior- lateral staminode glands equal but larger on the posterior-lateral two; pedicels never arranged around a central glandular mass. (Minas Se Rok urea 26. Limb of the posterior petal ca 8.5 mm long, ca 9 wide, orbicular; limbs of the lateral petals TL 11.5 mm long, ca 12 mm wide; all staminode glands equally long, ave posterior-lateral two narrower than the anterior an anterior-lateral three; pedicels sometimes arranged around a central glandular mass. (Goidas) 10. P. gardneri. 25. Hairs of the stem and inflorescence axes with stalks less than 0.5 mm long. 27. Stem and inflorescence stipules 16-23 mm long, persistent; limbs of the lateral petals 16—16.6 mm long, 15-16 mm wide; posterior styles 5.4—5.5 mm long. (Bahia) a P. megatantha. 27. Stem stipules 4-13 (-19) mm long, deciduous or sometimes persistent, Aig os ‘ipules usually — maller, the most distal ca 5—6 mm long or less , de- cluows aoe of the eee petals 9 atts 2 St 5) mm long, (-15) mm _ wide; posterior styles 3.3 4.7 mm ane 28. Limb of the posterior petal 7.5-10 mm long, 6.8-9 mm wide, orbicular or sometimes broad- ly obovate, ca 2/3-3/4 as wide as the limbs of the lateral petals. (Goias) 12. P. gowana. Limb of the posterior petal 6-7 mm long, 4— 5mm wide, elliptical or renee) obovate, 1/3- 1/2 as wide as the limbs of the lateral petals. 29. Limbs of the lateral petals elliptic al « or obo- ho ge 1 shorter than the posterior two. preys eae no 5. 9. Limbs of the lateral petals orbicular, < margins fimbriate with the fimbriae up to 0.8 (-1) mm long; styles about nae long. (Minas Gerais se ee re 20. All inflorescence leaves peuolate 30. Anterior and anterior-latet sal staminode glands with an indenta- uion across the apex and ee on the abaxial face. 3 orescence commonly of | or 3 or 5 umbels, occasionally more, 3° axes absent; anterior and anterior-lateral staminode glands ae: than long; limbs of the lateral 13- 20 mm long. (Coastal regions from Bahia to Sao Paulo) 15. P. hispidula. 31. Inflorescence of more than 5 umbels, 3° axes usually present; anterior and anterior-lateral staminode glands longer than wide or as long as wide; limbs of the lateral petals 9.5-I14 mm ay Glands of the stem leaves borne on the surface of the lamina away from the base and costa. (Minas pe >. spinensts, 32. Glands of the stem leaves borne at the base of 2 I mina at the costa. 33. Petioles of the stem leaves 8-13 mm long; limbs of the lateral petals oblanceolate. (Minas Ger: ee ee "Pp. andersonit. 33. Petioles of the stem leaves 2-5 mm long; limbs of the lateral petals orbicular. (Minas Gerais) 6. P. barnebyi. 30. Anterior and anterior- — see glands smooth. Anterior style 5—9.2 mm long, tally strongly arced toward the posterior petal, aa wr ae an oblong stigma at the apex but laterally on the abaxial surface (or rarely with the apex un- cinate and bearing a capitate stigma 35. Limbs of the lateral petals | 5-17, 8 mm long ine wide, orbicular; styles 7.3-9 mm long; stem stipules 3—) 12— 23 mm long. (Goias, Mato Grosso) ie eo ee 35. Limbs of the lateral petals 10.5-12 mm long, ca 9- 10 mm wide, obovate; styles 5-6.5 mm long; stem st ip- ules ca 6-8 mm long. (Ceara, Piaur, Maranhao, Parans 17 > Jussieuana. 34. Anterior style 2.6-5.5 mm long, arced or inclined tow ed the posterior petal (or rarely toward the anterior sepal) or erect, bearing a capitate stigma, apex never uncinate. 36. Inflorescence stipules large, even the most distal 7— 17 mm long, persistent or sometimes dec a at the most proximal nodes. 37. Pedicels commonly arr anged around a central gland- ular mass up to 2.5mm in diameter; limb of the posterior petal with the margin capitate- ee in the proximal 1/2-2/3, fimbriate-glandu distal 1/3—1/2; anterior style longer than an poste- rior two; posterior-lateral staminode pee gro than the anterior and anterior-lateral three. (Mat Grosso, eastern Bolivia) 9. P. pres 37. Pedicels never arranged around a central glandular mass; limb of the posterior petal with the margin o 1e posterior two; staminode glands all equally long but the posterior-lateral two narrower than the anterior and anterior-lateral three. (Goias, Minas es ) >. leptoclada. 36. See Aaa stipules small, the most distal ee mm ae 1g r less, usually deciduous but often present at nodes panes recently opened flowers. 38. Glands of the stem leaves borne on the surface of the amina away from the base and costa, or sometimes on some leaves one or both glands Bel yout erais, Bahia >. paludosa. 38. Glands of the stem leaves borne at the ee e. ie lamina at the costa or halfway on the petiole. 39. Limb of the posterior petal 7.5-10 mm long, 6.8-9 mm wide, orbicular or sometimes broad- ly obovate, ca 2/3-3/4 as wide as the limbs of the lateral petals; anter ior style shorter than the 12. a i =~ 2 = co S a ow A o =~ fan = po P. gotana. 39. Limb of the ee petal oa mm ae 4— 5 mm wide, elliptical, ca 1/3-1/2 as wide as the limbs of the lateral petals; styles about pen) long. (Minas Gerais) . P. barnebyi. 1. Peixotoa adenopoda C. Anderson, sp. nov. Fig. 128 Liana. Stipulae vegetativae 7-16 mm longae, acutae vel bifidae, plerumque persistentes. Petioli 4-8 mm longi. Laminae 4.8-9.5 cm longae, 1.3—4.8 cm latae, ellipticae vel parum obovatae vel lanceolatae, rugosae, apice apiculatae vel acuminatae, supra sparsim aureo-velutinae vel sparsim aureo-sericeae, subtus sparsim et laxe aureo-sericeae, ferentes 2 glandulas pedicellatas 1.7-10 mm supra basim 1.2-6 mm a costa. Folia inflorescentiae petiolo brevi, distaliora linearia vel triangularia, apice acuminata vel caudata. Inflorescentia terminalis vel axillaris, umbellis singularibus in nodis subterminalibus, umbellis 3 in nodis terminalibus, pedunculis secundaris 0.5—3 mm Honels Bracteae bracteolaeque praesentes. Pedicelli 11-20 mm longi. Sepala 4-5 mm longa. Limbus petalorum lateralium 10-10.5 mm _ longus, orbicularis, margine glanduloso-fimbriato. Limbus petali posticl ca 6 mm longus, ellipticus vel anguste obovatus, margine glandulis capitatis ornato. Glandulae 3 staminodiorum anticorum apice pro- funde indentatae, gee ae in pagina abaxiali decurrenti. Styli subaequales vel anticus brevior, 2.8-3.6 mm longi, anticus parum arcuatus versus petalum posticum, postici ee inclinati versus petala postico-lateralia; stigmata cd uae Samara ign ota e. Vegetative Reaie hes purplish-brown, velutinous, pubescence abraded from aller parts. Stem stipules 7-16 mm long, 7.6—17 mm wide, cordate, entire with the apex acute, or bifid, glabrous or finely, sometimes sparsely, velutinous or tomentulose adaxially, appressed sericeous abaxially, usually persistent; in- florescence stipules gradually decreasing in size toward the distal nodes, the smallest not less than 5 mm long, 3 mm wide, commonly persistent. Stem leaves with the petiole 4-8 mm long, densely and loosely golden velutinous, laminas 4.8-9.5 cm long, 1.3-4.8 cm wide, elliptical or slightly obovate to lanceolate, apex apiculate or in the narrower acuminate, base cordate or almost truncate, rugose, sparsely golden velutinous-sericeous above, sparsely golden sericeous below, a pair of stalked glands borne on the surface of the lamina, 1.7—10 mm above the base, 1.2-6 mm from the costa, each gland 0.8-1.5 mm in diameter, stalk 0.8-— 1 mm high. Inflorescence leaves with the petiole ca 1 mm or more long, laminas abruptly smaller than the cauline, the smallest 1.3 mm long, 0.7 mm wide, the most proximal elliptical or linear-lanceolate, the more distal narrowly triangular or linear, apex acuminate or caudate, sparsely sericeous or glabrous above, sparsely sericeous below, a pair of glands at or near the base at or near the costa, each gland 0.5-1 mm in diameter, the glands stalked in the most proximal, sessile in the more distal. Inflorescence terminal or axillary, the axes loosely sericeous or velutinous, internodes of the primary axis 0.6—3.5 cm long, 2° axes 1-6 cm long, 3° axes 6.5-12 mm long, subsidiary axes absent, at the more distal nodes the 2° and 3° axes usually suppressed, with | or 2 umbels sessile or borne on secondary peduncles in the leaf axil, always with 3 umbels at the terminal nodes, or occasionally with only | umbel at the terminal node Ol a oe axis, each umbel 4-flowered, primary peduncles absent, secondary peduncles up to 3mm long or absent. Bracts 0.5-1 mm long, 0.2—-0.6 mm wide, triangular, labrous or very sparsely velutinous on both surfaces and on the margins; bracteoles 0.1—0.3 mm long and wide, oblong, triangular or semicircular, 18 glabrous, sometimes the margins velutinous. Pedicels 11-20 mm long, veluti- nous. Sepals 4-5 mm long, 1.3—2 mm wide, adaxially very sparsely velutinous or tomentulose, more densely so near the apex, glands 1.2-1.5 mm long. Claw of the lateral petals ca 2 mm long, limb 10-10.5 mm long and wide, orbicular, base attenuate, fimbriae up to 0.3 mm long, glandular, claw of the posterior petal 4—4.3 mm long, limb ca 6 mm long, ca 4 mm wide, elliptical or narrowly obovate, fimbriae up to 0.4 mm long, capitate-glandular. Filaments of the lateral stamens 3.4—3.6 mm long, posterior filament 2.5-2.6 mm long, arced toward the poste- rior petal, anthers |.1—1.2 mm long, glandular connectives ca 0.5 mm long. Pollen grains ca 42 um. Filaments of the anterior staminode 4—4.2 mm long, anterior-lateral filaments 3.3-3.7 mm long, posterior-lateral filaments 2.6— 2.8 mm long, inflexed between the posterior styles, anterior and anterior-lateral glands ca 1.2 mm long and wide, posterior-lateral glands ca 1.1 mm long, 0.9— I mm wide, anterior and anterior-lateral glands with a deep indentation across the apex and decurrent on the abaxial face, posterior-lateral glands smooth. Styles 2.8-3.6 mm long, anterior styles ca 0.4 mm wide, sometimes shorter but always a little stouter than the posterior two, slightly curved toward the posterior petal, posterior styles ca 0.3 mm wide, strongly divergent, inclined toward the posterior-lateral petals, all styles velutinous in the proximal 1/3 of their lengths, stigmas 0.4—0.5 mm in diameter, capitate. Samara not seen. Type. Jesus 392. Brazil. Bahia: Maracas, saida de Itirugu, 20 May 1969 (holotype CEPEC, isotype MICH). Paratype. Pinheiro 1870. Brazil. Bahia: Rod. Itirugu-Maracas, 8 July 1972 (CEPEC, MICH). Peixotoa adenopoda is known only from the type and the paratype (Fig. 3). It is one of four species in which the terminal inflorescence nodes bear three umbels and thus are 12-flowered. It is readily distinguished from the other three by its stalked leaf glands which are borne on the surface of the lamina away from the base and either away from the costa or adjacent to it. The leaf glands of P. catarinensis, P. parviflora, and P. tomentosa are sessile. None of these species ( \ / ( / Ly / i ( S ‘ } ( ee | i J \ ; f \ J Vie | ? bs ‘as ed / s \ ) \ \ } aN 2 M i a \ 7 \ Fs i eal es eae a” } ff \ rele - (° WW j a % | | vs \ ol ee ye ey FEED fom ¥ a ab, . fF f Peixotoa 1 @ adenopoda \ _ # anadenanthera »-~ M@ andersonii pee © axillaris a Se oe ea | FIG. 3. Distribution of P. adenopoda, P. anadenanthera, P. andersonit, and P. axillaris. 19 occurs in Bahia. The only other species with stalked but basal leaf glands is the glabrous P. hatschbachi of south-central Goias. P. adenopoda is also unusual in that its leaves are sericeous. P. sericea, also from Bahia, is the only other species in the genus with sericeous leaves. It has sessile leaf glands, and its terminal inflores- cence nodes bear a single umbel, 1.e., are 4-flowered. 2. Peixotoa anadenanthera C. Anderson, sp. nov. Hig lS. Frutex pauciramosus usque ad 1.5 m. Caules dense aureo-velutini. Stipulae vegetativae 7.3-7.5 mm longae, anguste triangulares, deciduae. Petioli 2.5- 6 mm longi. Laminae 7-11 cm longae, 3.2-5.3 cm latae, anguste ellipticae vel lineari-lanceolatae, supra aureo-velutinae, subtus aureo-tomentosae, basi costae biglandulosae vel glandulis apice petioli. Folia inflorescentiae petiolo brevi vel sessilia, linearia vel subulata, distaliora decurrentia, petiolis basi biglandulosis. Inflorenscentia axillaris, umbellis singularibus. Bracteae bracteolaeque praesentes. Pedicelli 11-26 mm longi. Sepala ca 5.2mm_ longa. Limbus petalorum lateralium 9.5-ca 10 mm longus, late obovatus, margine glanduloso- fimbriato. Limbus petali postici 6.2-6.5 mm longus, obovatus, margine 4 proximalibus glandulis capitatis ornato, “4 distali fimbriato vel denticulato, glanduloso vel eglanduloso. Antherae sine connectivo glanduloso. Stylus anticus 3.3-3.5 mm longus, erectus vel parum inclinatus versus petalum posticum, styli postic1 3.6-4 mm longi, parum divergentes, parum inclinati versus petala postico-lateralia; stigmata capitata. Samara ignota. Little-branched shrub to 1.5m. Vegetative branches densely golden velutinous, pubescence abraded from older parts. Stem stipules 7.3-7.5 mm long, 4-6.5 mm wide, narrowly triangular, apex acute, tomentulose to glabrate adaxially, densely tomentulose-velutinous abaxially, deciduous; inflorescence stipules like the cauline, sometimes smaller, the smallest not less than 3.6 mm long, 3.5 mm wide, persistent (7). Stem leaves with the petiole 2.5-6 mm long, densely velutinous, laminas 7-11 cm long, 3.2-5.3 cm wide, narrowly elliptical or linear-lanceolate, apex apiculate, base truncate, rugose, golden velutinous above, golden tomentose below, a pair of sessile glands hidden in the dense pubescence halfway on the petiole or on the petiole up to 1 mm below the lamina, each gland 0.6—1.2 mm in diameter, or sometimes the glands absent. Inflorescence leaves with a petiole 3.5 mm or less long, or subsessile or sessile, laminas abruptly smaller than the cauline, the smallest not less than 2.5 mm long, 0.1 mm wide, linear to subulate, apex acute or acuminate, the petiolate with the base decur- rent, tomentulose and pubescent with T-shaped hairs above, densely tomentu- lose-velutinous below, a pair of glands at the base of the petiole in petiolate leaves, at the base of the lamina at the costa in sessile leaves, each gland 0.4— 0.7 mm in diameter. Inflorescence terminal or axillary, internodes of the pri- mary axis 0.9-5.8 cm long, mature secondary axes not seen, tomentulose- velutinous, subsidiary axes absent; umbels 4-flowered, one umbel per node, primary and secondary peduncles absent. Bracts 0.8—1.1 mm long, ca 0.3 mm wide, narrowly triangular, glabrous; bracteoles ca 0.3 mm long, ca 0.2 mm wide, triangular, glabrous. pedicels 11-26 mm long, densely tomentulose-velutinous. Sepals ca 5.2 mm long, ca 2.3 mm wide, adaxially the proximal 43 glabrous, the distal %4 tomentulose, glands 1.8—2.2 mm long, 0.8—1.1 mm wide. Claw of the lateral petals 2-2.5 mm long, limb 9.5-10 mm long and wide, broadly obovate, base briefly attenuate, margin irregularly fimbriate- ay Breed ce up to 0.5 mm long, glandular. Claw of the posterior petal 3.5 m long, limb 6.2-6.5 mm long, 5.2-5.5 mm wide, obovate, fimbriae up to ae mm long, those of the proximal 4 capitate-glandular, the distal 4 fimbriate-denticulate, glandu- lar or eglandular. Filaments of the anterior-lateral and the posterior stamens rs 3.2-3.5 mm long, posterior filament slightly curved toward the posterior petal, 20 posterior-lateral filaments 3.6-3.7 mm long, anthers 1.5—-1.7 mm long, glandular connectives absent. Pollen grains ca 50 um. Filaments of the anterior and the posterior-lateral staminodes 3.1-3.3 mm long, posterior-lateral filaments in- flexed between the posterior styles, anterior-lateral filaments 3.5-3.6 mm long, anterior gland ca 1.4mm long and wide, anterior-lateral glands 1.2—-1.3 mn long, ca 1.2 mm wide, posterior-lateral glands I-1.1 mm long, ca 1 mm wide, all glands with the apex and abaxial face smooth. Anterior style 3.3-3.5 mm long, ca 0.4 mm wide, erect or slightly curved toward the posterior petal, sometimes with a few scattered hairs in the proximal 4 of its length, posterior styles 3.6—4 mm long, ca 0.3 mm wide, somewhat divergent, slightly inclined toward the poste- rior-lateral petals, glabrous, stigmas 0.4—0.5 mm in diameter, capitate. Samara not seen. Type. Hatschbach 36904. Brazil. — GO-12, 30 km S$ of Cavalcante, 26 May 1975 (holotype MBM, isotype MICH). Peixotoa anadenanthera is known only from the type (Fig. 3). It is unique in that its anthers lack the glandular connectives. It also differs from most other species in that the stem stipules are narrow, and in that the leaves tend to be narrowly elliptical. The internodes of the younger branches and branchlets are densely velutinous. The hairs are similar to those of P. hirta and P. gardneri in that the stalk is usually more than 0.5 mm long. 3. Peixotoa andersonii C. Anderson, sp. nov. Fig. 14. Caules procumbentes usque ad 4 m. Stipulae vegetativae 7-11 mm longae, triangulares, acutae vel bifidae, persistentes. Petioli 8-13 mm longi. Laminae 9.7-15.3 cm longae, 3.3-7.9 cm latae, anguste ellipticae, supra aureo-velutinae, subtus aureo-tomentosae, basi juxta costam biglandulosae. Folia inflorescentiae petiolo brevi vel sessilia, lineari-lanceolata vel anguste tr langularia vel subulata. Inflorescentia terminalis, umbellis singularibus, interdum ramis secundariis nodorum distalissimorum multum redactis. Bracteae bracteolaeque praesentes. Pedicelli 16-23 mm longi. Sepala (3.8-)4-5.5 mm longa. Limbus_ petalorum lateralium 12.5-14 mm longus, oblanceolatus, decurrens, margine glanduloso- fimbriato. Limbus petali postici ca 6 mm longus, subrectangularis, margine 34 proximalibus glandulis capitatis ornato, % distali glanduloso-fimbriato. Glandu- lae 3 staminodiorum anticorum apice profunde indentatae, indentatione in pagina abaxiali decurrenti. Stylus anticus 4.1-4.3 mm longus, arcuatus versus petalum posticum, styli postici 4.4—4.6 mm longi, parum divergentes, parum inclinati versus petala postico-lateralia; sugmata capitata. Samara ignota. Stems procumbent, up to 4m. Vegetative branches golden velutinous, pubescence abraded from older parts. Stem stipules 7-l11mm_ long, 7- 10.6 mm wide, triangular, entire with the apex acute, or bifid, finely tomentu- lose adaxially, tomentulose-sericeous abaxially, persistent; inflorescence stipules gradually decre easing in size toward the distal nodes, the smallest not less than 3.2mm long and wide, the most distal sometimes split into two narrow lobes, persistent. Stem leaves with the petiole 8-13 mm long, densely golden veluti- nous, laminas 6.7-15.3 cm long, 3.3-7.9 cm wide, narrowly elliptical, apex apiculate, base usually truncate, rarely cordate, golden velutinous above, golden tomentose below, a pair of sessile glands at the base at the costa, each gland (1-) 9-2.5 mm in diameter. Inflorescence leaves with the petiole | mm or more long or the most distal sessile, laminas abruptly smaller than the cauline, the smallest not less than 1.8 mm long, 0.6 mm wide, linear-lanceolate or narrowly Bee ola or subulate, apex acuminate, sparsely tomentulose above, densely elutinous-tomentulose below, a pair of glands at the base at the costa, these sometimes fused, each gland 0.7—1.2 mm in diameter, sometimes with a second, smaller pair (ca 0.4 mm in diameter) just below the larger pair; rarely the most 21 distal leaves reduced to sessile, linear or subulate, eglandular bracts, or rarely the most distal nodes bearing split stipules, the leaves absent. Inflorescence terminal or axillary, the axes densely golden velutinous, internodes of the primary axis 2.8-6.5 cm long, 2° axes 2.1-6.1 cm long, 3° axes 0.5-1.1 cm long, subsidiary axes absent, sometimes the most distal axes much reduced; umbels 4- flowered, one umbel per node, primary and secondary peduncles absent. Bracts 0.5- 0.8 mm long, 0.2-0.3 mm wide, narrowly triangular, glabrous; bracteoles 0. 1- 0.2 mm long and wide, linear, glabrous. Pedicels 16-23 mm long, densely golden velutinous. Sepals (3.8—) 4—-5.5 mm long, 2—2.3 mm wide, adaxially the proximal ’% glabrous, the distal % tomentulose, glands 1.4—1.9 mm long, 0.8—1.1 mm wide. Claw of the lateral petals 1.6—2 mm long, limb 12.5-14 mm long, 9-11 mm wide, oblanceolate, base decurrent, fimbriae up to 0.5 mm long, glandular. Claw of the posterior petal 3.7-4 mm long, limb ca 6 mm long, 44.6 mm wide, subrectangular, fimbriae up to 0.4 mm long, those of the proximal % capitate- glandular, the distal 3 fimbriate-glandular. Filaments of the lateral stamens 3.6-4 mm long, posterior filament ca 3mm long, strongly arced toward the posterior petal, anthers 1.2—-1.3 mm long, glandular connectives 0.6—0.7 mm long. Pollen grains ca 50 ym. Filament of the anterior and anterior-lateral staminodes 3.6—3.8 mm long, the anterior filament often a little longer than the anterior-lateral two, posterior-lateral filaments 3.2-3.3 mm long, inflexed be- tween the posterior styles, anterior and anterior-lateral glands 1.4—1.5 mm long, 1.3—1.4 mm wide, posterior-lateral glands 1.1—1.2 mm long, ca 1.1 mm wide, anterior and anterior-lateral glands with a deep indentation across the apex and decurrent on the abaxial face, posterior-lateral glands smooth. Anterior style 4.1-4.3 mm long, 0.3-0.4 mm wide, arced toward the posterior petal, posterior styles 4.44.6 mm long, 0.3—0.4 mm wide, slightly divergent, slightly curved toward the posterior-lateral petals, all styles with scattered hairs in the proximal ¥s of their lengths, sugmas 0.4—0.5 mm in diameter, capitate. Samara not seen. Type. Anderson 11591. Brazil. Minas Gerais: Mun. Diamantina, steep-sided valley of Biribiri, 1100 m, 25 Feb 1975 (holotype MICH, isotype NY). Peixotoa rae is known only from the type (Fig. 3). It forms procumbent stems up to 4 m long which bear erect inflorescences, an unusual habit in this genus. It is unique in that the limbs of the lateral petals are oblanceolate and decurrent on the claw. The anterior and anterior-lateral staminode glands have a deep indentation across the apex and decurrent on the abaxial face. This species is also unusual in that the stem leaves have a petiole at least 8 mm long. Vegetatively it resembles P. parviflora and the less densely pubescent forms of P. tomentosa. These two species have smooth staminode glands and lateral petals whose limbs are usually orbicular or sometimes elliptical or broadly ovate. ‘They are also distinguished by their inflorescences, in which the terminal node bears three umbels and is thus 12-flowered. In P. andersonii the terminal nodes bear one umbel, t.e., are 4-flowered. However, sometimes at some terminal nodes the 2° axes are greatly reduced, the stipules are split into several, often lacerate, strips, and the inflorescence leaves are absent. With such an arrangement the inflorescence branches resemble those of P. parviflora and P. tomentosa. This species is named for my husband, William R. Anderson, an expert student of the Malpighiaceae, to whom I am much indebted. 4. Peixotoa axillaris C. Anderson, sp. nov. Fig. 15 Frutex usque ad | m. Stipulae vegetativae 10.7—11.5 mm longae, triangu- lari-cordatae, acutae, deciduae, Petioli 4-8 mm longi. Laminae 7.1—13.4 cm longae, 5.5—9 cm latae, ellipticae vel ovatae vel suborbiculatae, supra velutinae, subtus tomentosae, basi juxta costam biglandulosae. Folia inflorescentiae petiolo brevi vel sessilia, elliptica vel linearia vel subulata. Inflorescentia ex 1-3 umbellis rp ramo axillari brevi (2-9 mm longo) constans, umbellis singularibus. Bracteae bracteolaeque praesentes. Pedicelli 10-13 mm longi. Sepala 4.2-4.7 mm longa. Limbus petalorum lateralium = 12.5-13 mm longus, late obovatus, margine glanduloso-fimbriato. Limbus petali postici ca 10 mm longus, obovatus, margine ¥3 proximali glandulis capitatis ornato, % distalibus glanduloso-fimbriato. Fila- menta staminum subaequalia. Styli subaequales, 4.4-4.6 mm longi, anticus erectus, postici divergentes, parum inclinati versus petala postico-lateralia; stigmata capitata. Ala dorsalis samarae 2.7-2.9 cm longa, 1.9-2 cm lata, alae laterales 4.6—5 mm latae, 10-11 mm altae. Shrub ca Im tall. Vegetative branches velutinous-tomentulose. Stem stipules 10.7-11.5 mm long, ca 11 mm wide, triangular-cordate, apex acute, adaxially glabrous or the proximal % finely tomentulose, the distal % glabrous, abaxially tomentulose-velutinous, deciduous; inflorescence stipules like the cauline but smaller, the smallest not less than 4.2 mm long, 5 mm wide, cordate, concave, deciduous. Stem leaves with the petiole 4-8 mm long, densely golden velutinous, laminas 7.1—13.4 cm long, 5.5-9 cm wide, elliptical or ovate or sub- orbicular, apex apiculate, base cordate, velutinous above, tomentose below, a pair of sessile glands at the base at the costa, each gland |.1—1.6 mm in diameter. Inflorescence leaves with the petiole up to 1.5 mm long or sessile, laminas abruptly smaller than the cauline, the smallest not less than 3.8 mm long, 0.8 mm wide, elliptical or linear or subulate, apex acuminate, glabrous above, densely velutinous-tomentulose below, a pair of glands at the base at the costa, each gland 0.6—-0.8 mm in diameter. Inflorescence borne on a short (2-9 mm long) axillary shoot, the terminal umbel sessile or borne on a primary axis, I-6.5 mm long, solitary or with two lateral sessile umbels, or with a pair of 2° axes 3-3.5 mm long, each bearing one umbel, subsidiary axes absent; umbels 4-flowered, primary and secondary peduncles absent. Bracts 0.6—0.7 mm long, ca 0.7 mm wide, triangular or semicircular, glabrous, sometimes with scattered hairs abaxially and/or on the margins; bracteoles 0.2—0.6 mm long, 0.2-0.3 mm wide, triangular or narrowly so, glabrous, often with scattered hairs on the margins. Pedicels 10-13 mm long, velutinous, up to % again as long in fruit. Sepals 4.2-4.7 mm long, 2.5-2.6 mm wide, adaxially the proximal 4 glabrous, the distal 3% tomentulose, glands 1.5-1.7 mm long, 0.8—0.9 mm wide. Claw of the lateral petals 1.7—2 mm long, limb ca 12.5-13 mm long, ca 11.5-12 mm wide, broadly obovate, base attenuate, fimbriae up to 0.6 mm long, glandular. Claw of the posterior petal ca 5 mm long, limb ca 10 mm long, ca 9 mm wide, obovate, fimbriae up to 0.6 mm long, those of the proximal 3 capitate-glandular, the distal % fimbriate-glandular. Stamen filaments 4.2-4.5 mm long, posterior fila- ment usually a little shorter than the lateral four, very slightly curved toward the posterior petal, anthers 1.5—1.8 mm long, glandular connectives 0.8—1 mm long. Pollen grains 57-66 ym. Filament of the anterior staminode ca 5.2 mm long, anterior-lateral and posterior-lateral filaments 4.8-4.9 mm long, the posterior- lateral two inflexed between the posterior styles, all glands 1.41.5 mm long, anterior and anterior-lateral glands 1.2-1.3 mm wide, posterior-lateral glands ca | mm wide, all glands with the apex and abaxial face smooth. tyles 4.44.6 mm long, 0.3-0.4 mm wide, anterior style usually a little longer and stouter than the posterior two, erect, posterior styles divergent, slightly inclined toward the posterior-lateral petals, all styles glabrous, anterior stigma ca 0.6 mm in di- ameter, posterior stigmas ca 0.5 mm in diameter, all stigmas capitate. Carpo- phore up to 3.5 mm long. Torus up to 5.5 mm high. Samara with the dorsal wing 2.7-2.9 cm long, 1.9-2 cm wide, upper margin gently sigmoid, lower margin arced, erose, lateral wings 4.6-5 mm wide, 10-11 mm high, rectangular, areole 5-6 mm long, 6-6.6 mm wide, nut 7-7.5 mm long, 4—5.4 mm in diameter, seed ca 8.5 mm long, outer cotyledon ca 11 mm long, ca 3.2 mm wide, folded at ca 3 of its length, inner cotyledon ca 5.6 mm long, ca 2.3 mm wide, straight. 23 Type. Irwin & Soderstrom 7443. Brazil. Goias: Serra de Caiapo, 17°12'S, 51°47'W, ca 60 km S of Caiaponia on road to Jatart, 800-1000 m, 27 Oct 1964 (holotype UB, isotypes MICH, NY Peixotoa axillaris is known only from the type (Fig. 3). It is readily distin- guished by its reduced inflorescence, borne on a short axillary shoot, which is composed of one or three umbels. ‘The terminal umbel may be sessile or borne on a tiny primary axis. The lateral umbels are sessile. The type is apparently an apomict. The closed anthers contain pollen that is mostly aborted and 88-91% non-staining in cotton blue in lactophenol. The coastal P. hispidula is the only other species whose inflorescences may consist of only one or three umbels, but these are never sessile. It is commonly glabrous or only very sparsely pubescent. The limbs of its lateral petals are 13-20 mm long, while the limb of its posterior petal is only 6.5-8.2 mm long. The staminode glands are strongly dimorphic. The anterior and anterior-lateral three have an indentation across the apex and decurrent on the abaxial face. P. axillaris is pubescent in all its vegetative parts. The limb of the posterior petal (ca 10 mm long) is only a little smaller than the limbs of the lateral petals (12.5- 13 mm long). The staminode glands are subequal and smooth. 5. Peixotoa bahiana C. Anderson, sp. nov. Fig. 14. Frutex usque ad 2.5 m. Stipulae vegetativae 7.5-13 mm longae, acutae vel bifidae, deciduae. Petioli 2.5-4 mm longi. Laminae 4.5-6.5 cm longae, 2.8- 3.8 cm latae, anguste ellipticae vel anguste ovatae, rugosae, supra velutinae, subtus in venis tomentosae, basi juxta costam biglandulosae. Folia inflorescentiae sessilia, subulata. Inflorescentia terminalis, umbellis singularibus. Bracteae bracteolaeque praesentes. Pedicelli (13-)18-23 mm longi. Sepala 4.5—5.8 mm longa. Limbus petalorum lateralium 11-12.3 mm longus, obovatus vel ellipticus, denticulatus, glandulosus vel eglandulosus. Limbus petali postici 6.2-7 mm longus, ellipticus vel parum obovatus, margine glandulis capitatis ornato. Stylus anticus 3.4—3.6 mm longus, erectus, styli postici 4.2-4.3 mm longi, erect, parum divergentes; stigmata capitata. Samara ignota. Shrub up to 2.5 m. Vegetative branches finely tomentulose-velutinous. Stem stipules 7.5-13 mm long, ca 9-13 mm wide, cordate, entire with the apex acute, or bifid, finely and sparsely velutinous or velutinous-tomentulose adaxially, densely velutinous-tomentulose abaxially, deciduous; inflorescence stipules like the cauline, usually a little smaller, deciduous. Stem leaves with the petiole 2.5— 4 mm long, densely velutinous, laminas 4.5-6.5 cm long, 2.8-3.8 cm wide, nar- rowly elliptical or narrowly ovate, apex apiculate, base cordate, rugose, veluti- nous above, tomentose especially on the veins below, a pair of sessile glands at the base at the costa, each gland 0.8—1.6 mm in diameter. Inflorescence leaves of the first and sometimes second node of the primary axis petiolate, the others sessile, laminas abruptly smaller than the cauline, the smallest not less than 4.6 mm long, 0.8 mm wide, the most proximal linear-lanceolate, the more distal subulate, apex acuminate, glabrous above, velutinous-tomentulose below, a pair of glands at the base at the costa, each gland 0.7—-1.5 mm in diameter, sometimes with a second, smaller pair (0.3—-0.6 mm in diameter) above or below the larger pair. Inflorescence terminal, the axes velutinous or velutinous-tomentulose, internodes of the primary axis 1.2-6.5 cm long, 2° axes 2.9-4.5 cm long, 3° axes 2.2-5.8 cm long, subsidiary axes absent; umbels 4-flowered, one umbel per node, primary and secondary peduncles absent. Bracts 1—-1.2 mm long, 0.7- 1.1 mm wide, oblong, with a few scattered hairs on both surfaces and along the margin; bracteoles ca 0.2 mm long, 0.1-0.2 mm wide, oblong or triangular, glabrous. Pedicels (13-) 18-23 mm long, velutinous. Sepals 4.5-5.8 mm long, 24 9.5-2.7 mm wide, adaxially glabrous or tomentulose only in the distal 1/5, glands 1.8—2.2 mm long, 0.9-1.2 mm wide. Claw of the lateral petals 2-2.3 mm long, limb 11-12.3 mm long, 8-I1.1 mm wide, obovate or elliptical, base briefly attenuate or truncate, denticulate, the teeth up to 0.3 mm long, glandular o1 eglandular. Claw of the posterior petal 4—4.3 mm long, limb 6.2-7 mm long, 4.1-4.2 mm wide, elliptical or slightly obovate, fimbriae up to 0.4 mm long, capitate-glandular. Filaments of the lateral stamens 4—4.2 mm long, posterior filament 3.2-3.6 mm long, arced toward the posterior petal, anthers 1.2—1.5 mm long, glandular connectives 0.7—1 mm long. Pollen grains 42-50 um. Filaments of the anterior and anterior-lateral staminodes 4—4.2 mm long, posterior fila- ments 3.3-3.6 mm long, slightly curved toward the posterior styles, anterior and anterior-lateral glands |.1-1.2 mm long and wide, posterior-lateral glands 0.8— | mm long, 0.7-0.8 mm wide, all glands with the apex and abaxial face smooth. Anterior style 3.4—3.6 mm long, ca 0.4 mm wide, erect or very slightly curved toward the posterior petal, posterior styles 4.2-4.3 mm long, ca 0.3 mm wide slightly divergent, erect, all styles with some scattered hairs in the proximal 4 (rarely 3) of their lengths, stigmas 0.4—0.5 mm in diameter, capitate. Samara not cr seen. Type. Irwin et al. 14889. Brazil. Bahia: Drainage of the Rio Corrente, ca 5 km S of Rio Roda Velha, ca 150 km SW of Barreiras, 900 m, 15 Apr 1966 Caeuiee UB, isotypes MICH, NY). iratype. Davidse et al. 12208. Brazil. Bahia: 39 km NE of the Goias-Bahia oe along Highway BR-020, 930 m, 8 April 1976 (MO). ‘his species is known only from the type and paratype (Fig. 4). In P. bahiana the narrowly elliptical leaves are tomentose below, the inflorescence leaves are _ Peixotoa | & bahiana - @ barnebyi ~~ @ catarinensis ‘\ @ cipoana . Distribution of P. bahiana, P. barnebyi, P. catarinensis, and P. cipoana. FIG. 4 25 sessile, and the stipules are deciduous. ‘The elliptical or slightly oblong limb of the posterior petal is relatively small. The margins of the limbs of the lateral petals are denticulate. P. bahiana i is readily distinguished from the other three relatively small-flowered species that are known from Bahia. P. adenopoda and P. sericea are both reported from the more eastern part of the state. P. adenopoda is recognized by the stalked leaf glands that are borne on the surface of the blade away from the costa and base, and P. sericea by the sericeous vesture of its leaves. In P. bahiana the leaf glands are sessile and borne at the base at the costa. P. paludosa also bears the leaf glands on the surface of the lamina rather than at the base, and has petiolate inflorescence leaves. The inflorescence supules are often persistent at the more distal nodes. 6. Peixotoa barnebyi C. Anderson, sp. nov. Fig. 16. Caules usque ad 1.51 1. Stipulae vegetativae 6.5—12 mm longae, acutae vel plerumque bifidae, plerumque persistentes. Petioli 2-5 mm longi. Laminae 5.3- 10.7 cm longae, 2.5-7.5 cm latae, anguste vel late ellipticae, ovatae vel sub- orbiculares, supra velutinae, subtus dense tomentosae, basi juxta costam bi- glandulosae. Folia inflorescentiae petiolo brevi, distaliora subsessilia, lanceolata vel linearia. Inflorescentia terminalis, umbellis singularibus. Bracteae bracteo- laeque praesentes vel interdum absentes. Pedicelli 12-25 mm longi. Sepala —)4.5—5.2 mm longa. Limbus petalorum lateralium 11—13.5 mm longus, orbicularis, margine fimbriato, glanduloso vel eglanduloso. Limbus petali postici 6-7 mm_ longus, ellipticus, margine glandulis capitatis ornato. Glandulae staminodiorum apice indentatae, indentatione in pagina abaxiali decurrenti. Styli subaequales, 3.8—4.1 mm longi, anticus posticis semper brevior, erectus vel parum inclinatus versus sepalum anticum, postici divergentes, arcuati versus petala postico-lateralia; stigmata capitata. Samaram maturam non vidi. rub to 1.5m. Vegetative branches purplish-brown, tomentulose. Stem stipules 6.5-12 mm long, 9-12 mm wide, cordate, commonly bifid, sometimes entire with the apex acute, rarely split almost to the base, sparsely tomentulose or glabrate adaxially, tomentose- velutinous abaxially, usually persistent; inflores- cence stipules gradually decreasing in size toward the distal nodes, the smallest not less than 3.5 mm long, 3.2 mm wide, glabrous or sparsely velutinous adaxi- ally, tomentulose or tomentulose-velutinous abaxially, deciduous. Stem leaves with the petiole 2-5 mm long, densely velutinous-tomentose, laminas 5.3— 10.7 cm long, 2.5-7.5 cm wide, narrowly to broadly elliptical or ovate or sub- orbicular, apex apiculate, base cordate, velutinous above, densely tomentose below, a pair of sessile glands at the tae at the costa or halfway on the petiole, each gland |-1.9 mm in diameter. Inflorescence leaves with a short petiole at the more proximal nodes, the most distal subsessile or sessile, laminas abr uptly smaller than the cauline or gradually decreasing in size toward the distal nodes, the smallest not less than 1.2 mm long, 0.8 mm wide, lanceolate or linear, veluti- nous above, densely velutinous-tomentulose below, a pair of glands halfway on the petiole, each gland (0.4—) 0.7—1.8 mm in diameter. pea es terminal, the axes tomentulose, internodes of the primary axis 2.7—10.8 mm long, 2° axes 3.4—9.6 cm long, 3° axes (0.5-) 24.8 cm long, ae 4° axes not seen, subsidiary axes absent; umbels 4-flowered, one umbel an node, puneny peduncles absent, second: ary peduncles rare, up to 1.5 mm long. Bracts 0.7— 1.3mm long, 0.3-0.5 mm wide, triangular, glabrous; bracteoles O.1—0.4 mm long and wide, broadly triangular, glabrous; sometimes bracts and bracteoles absent. Pedicels 12-25 mm long, tomentose. Sepals (4-) 4.5-5.2 mm_ long, 2-3.1 mm wide, adaxially the proximal 4—'% glabrous, the distal 4-34 tomentu- lose, glands 1.6-2.1 mm long, 0.8-1.3 mm wide. Claw of the lateral petals 2- 2.5 mm long, limb 11-13.5 mm long, 11-13.8 mm wide, orbicular, base attenu- 26 ate, fimbriae up to 0.8 (-1) mm long, glandular or eglandular. Claw of the posterior petal 4.5-5 mm long, limb 6-7 mm long, 4-4.5 mm wide, elliptical, fimbriae up to 0.5 mm long, capitate- glandular. Filaments of the lateral stamens 4—4.5 mm long, poster ior filament 3.3-3.5 mm long, arced toward the posterior petal, anthers 1.3-1.5 mm long, glandular connectives 0.8—0.9 mm long. Pollen grains ca 50 yum. Filaments of the anterior stamiode 3.9-4.7 mm long, always exceeding the anterior-lateral two, anterior-lateral filaments 3.7-4.3 mm long, always exceeding the posterior-lateral two, posterior-lateral filaments 3.5—4 mm long, inflexed between the posterior styles, glands 1.2-1.4 mm long, 1.1-1.3 mm wide, the posterior two often a little narrower than the anterior three, all glands with an indentation across the apex and decurrent on the abaxial face, the indentation sometimes very shallow in the posterior-lateral two. Styles 3.8— 4.1 mm long, anterior style ca 0.3 mm wide, always a little stouter than the posterior two, erect or slightly inclined toward the anterior sepal, posterior styles ca 0.2 mm wide, divergent, arced toward the posterior-lateral petals, all styles glabrous, stigmas ca 0.4 mm in diameter, capitate. Mature samara not seen. Type. Anderson 8623. Brazil. Minas Gerais: Serra do Espinhago, 30 km by road SW of Gouveia, at KM 60 on road to Curvelo, 1150 m, 11 Apr 1973 (holotype UB, isotypes MICH, Paratype. Anderson 11537. Brazil. Minas Gerais: Mun. Gouveia, ascent of Serra do Espinhago, on BR-259, 860-1000 m, 23 Feb 1975 (MBM CH Peixotoa barnebyi is known only from the type and paratype (Fig. 4 . [tis one of three species, all from the Serra do Espinhagco, in which all five staminode glands have an indentation across the apex and decurrent on the abaxial face. In the posterior-lateral glands the indentation may be quite shallow and difficult to detect. P. spinensis differs in that the leaf glands are borne on the surface of the blade away from the base and costa. In P. cipoana the unusually large staminode glands are recurved. In both species the posterior styles exceed the anterior one. In P. barnebyi the leaf glands are borne at the base at the costa, the staminode glands are erect, and the styles are subequal. Mature fruits have not been seen. However, the immature samaras of Anderson 8623 are similar to those of P. parviflora and P. tomentosa. This species is named for Rupert C. Barneby, a distinguished authority on many groups of plants. 7. Peixotoa catarinensis C. Anderson, sp. nov. Fig. Frutex volubilis. Stipulae vegetativae et inflorescentiae 4.5-ca 9 mm longae, deciduae. Petioli 4.5-15 mm longi. Laminae 3-10.2 cm longae, 2.2-7.6 cm latae, late vel anguste ellipticae interdum ovatae vel obovatae interdum lanceolatae, basi cordatae interdum sinu truncato, supra velutinae, subtus sericeae vel tomen- tosae, biglandulosae |.2—9.8 mm supra basim 2.3-6.5 mm a costa. Folia inflores- centiae peuolo brevi. Inflorescentia terminalis vel axillaris, umbellis singularibus in nodis subterminalibus, umbellis 3 in nodis terminalibus, pedunculis secun- darus ca I-13 mm longis. Bracteae bracteolaeque praesentes. Pedicelli 18— 25 mm longi. Sepala 4.2—5 mm longa. Limbus petalorum lateralium 11-12 mm longus, orbicularis, margine glanduloso-fimbriato. Limbus petali postici 8.5— 9 mm longus, orbicularis vel late obovatus, margine % proximalibus glandulis capitatis ornato, 3 distali glanduloso-fimbriato. Filamentum staminis postici erectum vel parum inclinatum versus petalum posticum. Glandulae stamino- diorum subaequales. Stylus anticus 3.2—3.8 mm longus, arcuatus versus petalum posticum, styli posticit 3.5-4.2 mm longi, semper anticum excedentes, diver- gentes, inclinatl versus ae postico-lateralia; stigmata capitata, Ala dorsalis samarae 2.3-2.7 cm longa, 1.1-1.2 cm lata, alae laterales 3.2-4.5 mm latae, 9- 10 mm altae. 27 Twining shrub. Vegetative branches purplish-brown, tomentulose and pubescent with T-shaped hairs or velutinous, pubescence abraded from older parts. Stem stipules 4.5-ca 9mm long and wide, cordate, apex acute, finely, usually sparsely, velutinous-sericeous or glabrate adaxially, sericeous abaxially, deciduous; inflorescence stipules like the cauline, usually more densely pubes- cent, persistent or eventually deciduous. Stem leaves with the petiole 4.5-15 mm long, loosely but densely velutinous, laminas 3-10.2 cm long, 2.2-7.6 cm wide, narrowly to broadly elliptical, sometimes ovate or obovate, apex apiculate, base cordate, often the sinuses truncate, velutinous above, pubescent with T-shaped hairs or loosely tomentose below, a pair of sessile glands borne on the surface of the lamina 1.2-9.8 mm above the base, 2.3-6.5 mm from the costa, each gland Q.2—0.7 mm in diameter, sometimes one or both glands absent. Inflorescence leaves with the petiole at least ca 1.5 mm long, laminas abruptly smaller than the cauline or gradually decreasing in size toward the distal nodes, the smallest not less than 3.7 mm long, 1.1 mm wide, elliptical or lanceolate, velutinous above, sericeous or sericeous-tomentulose below, in the proximal leaves the pair of glands borne on the surface of the lamina, in the more distal at the base at the costa, each gland 0.2-0.7 mm in diameter. Inflorescence terminal or axillary, the axes velutinous or tomentulose-velutinous, internodes of the primary axis |—4 (-6.7) cm long, 2° axes 1.7-7.3 cm long, subsidiary axes absent, at the more distal nodes the 2° axes suppressed, with | or 2 umbels borne on secondary peduncles in the leaf axil, always with 3 umbels at the terminal nodes, each umbel 4- flowered, primary peduncles absent, secondary peduncles up to 13 mm long. Bracts 0.4-1.9mm long, 0.4-0.7 mm wide, narrowly to broadly triangular, glabrous, sometimes velutinous on the margins; bracteoles 0.3-0.7 mm long, 0.2-0.3 mm wide, oblong or triangular, glabrous, sometimes velutinous on the margins. Pedicels 18-25 mm long, velutinous or tomentulose-velutinous, equally long in fruit. Sepals 4.2-5 mm long, 1.8-2.5 mm wide, adaxially velutinous or tomentulose or velutinous-tomentulose, glands 1.6-2.5 mm long, 0.8—1 mm wide. Claw of the lateral petals 2-2.5 mm long, limb 11-12 mm long, 11- 11.5 mm wide, orbicular, base attenuate, fimbriae up to 0.3 mm long, glandular. Claw of the posterior petal 44.5 mm long, limb 8.5-9 mm long and wide, orbicular or broadly obovate, fimbriae up to 0.3 (—0.4) mm long, those of the proximal % capitate-glandular, the distal “4 fimbriate-glandular. Filaments of e lateral stamens 3.8—4.4 mm long, posterior filament 3.2-3.7 mm long, erect or very slightly curved toward the posterior petal, anthers I-1.2 mm long, glandular connective 0.6-0.7 mm long. Pollen grains 50-57 wm. Filament of the anterior staminode 3.7-3.9 mm long, anterior-lateral filaments 3.4-3.6 mm long, posterior-lateral filaments 3-3.2 mm long, inflexed between the posterior styles, all glands I-1.1 mm long, ca 0.9 mm wide, sometimes the anterior gland a little larger than the lateral four, all glands with the apex and abaxial face smooth. Anterior style 3.2-3.8 mm long, ca 0.3 mm wide, always shorter and stouter than the posterior two, arced toward the posterior petal, posterior styles 3.5-4.2 mm long, ca 0.2 mm wide, divergent, inclined toward the posterior- lateral petals, all styles glabrous or with a few scattered hairs in the proximal “5 of their lengths, stigmas 0.4—0.5 mm in diameter, capitate. Carpophore up to ca 0.7 mm long. Torus up to ca 3.5 mm high. Samara with the dorsal wing 2.3- 2.7.cm long, 1.1-1.2 cm wide, upper margin sigmoid, lower margin arced, straight or sigmoid, erose, lateral wings 3.2-4.5 mm wide, 9-10 mm high, rec- tangular, areole ca 3.5 mm long, ca 3 mm wide, nut ca 6 mm long, ca 3.6 mm in diameter, mature seeds not seen. Type. Smith & Reitz 12276. Brazil. Santa Catarina: Ilha de Santa Catarina, Mun. Florianopolis, Canavieiras, 1-5 m, 28 Mar 1957 (holotype US, isotypes K, NY, P, R, RB). Distribution. (Fig. 4). Known only from the beaches, dunes and adjacent areas along the coast of Santa Catarina and its islands. 28 BRAZIL. Santa Catarina: Barclay 60 (MO, NY); Reserva Florestal dos ne Duarte 3339 Falcao (RB); Itha de Santa Catarina, Morro das Pedras, 5m, Klein et al. 5834 (P); Ilha de Santa Catarina, Pantano do Sul, 2 m, Klein & Bresolin 5988 (P); estrada a Itajar, Lutz s.n.(R icant 3); Morro da Santa La guna, Moreira & Moreira 495 (US); Raben 443 (BR); Praia Braba, Itajar, | m, Reitz 3240 (US); Ilha de Santa Catarina, Campeche, Reitz 5083 (US); Piloes, Palhoga, Reitz & Klein 2752 (P); Rio dos Pinheiros, Brago do Norte, 100 m, Reitz & Klein 6779 (P); Mun. Itajar, Praia Braba, ca 26°58'S, 48°36'W, 3m, Smith & Reitz 6085 (R, RB); Ule 711 (NY, fragment) — Peixotoa fcmlntien is one of ae fae in which the terminal inflorescence nodes bear three umbels, Le., are 12-flowered. The inflorescences are small compared to ee of other nee and ar have more than two pairs of 2° axes. 3° axes have not been seen. P. cafarinensis has unusually small leaf glands (Q0.2-0.7 mm in diameter), which are borne on the surface of the lamina away from the base and costa; sometimes they are absent. In some of the larger leaves the cordate base has truncate sinuses. P. catarinensis is the southernmost species in the genus and ts known only from Santa Catarina. It is readily differentiated from the other three species with a similar but usually larger inflorescence. P. parviflora, which ranges from Minas Gerais to Parana, and P. tomentosa, of Minas Gerais, have larger, basal leaf glands. In P. adenopoda of Bahia the leaf glands are borne on the lamina but are stalked and 0.8-1.5 mm in diameter. The range of the only other beach species, P. hispidula, extends from Bahia to Sao Paulo. It resembles P. catarinensis in its small inflorescences which bear petiolate leaves, but differs by its larger flowers and basal leaf glands. Collected in flower throughout the year; only one fruiting collection (March). 8. Peixotoa cipoana C. Anderson, sp. nov. Fig, 18. Frutex ascendens. Stipulae vegetativae 7-8.5 mm_ longae, persistentes. Petioli ca 1-5 mm longi. Laminae 3-9.5 cm longae, 2.7-6.7 cm latae, ovatae vel ellipticae interdum suborbiculares, rugosae, supra tomentulosae vel velutinae, subtus lanatae, basi juxta costam biglandulosae. Folia inflorescentiae subsessilia vel sessilia, distaliora linearia vel subulata. Inflorescentia bo aa umbellis singularibus. Bracteae bracteolaeque praesentes. Pedicelli 10-20 mm_ longi. Sepala 3.8-6 mm longa. Limbus petalorum lateralium aaa mm longus, orbicularis, margine glanduloso-fimbriato. Limbus petali postici 6-8 mm longus, ellipticus vel anguste obovatus, margine glandulis capitatis ornato. Glandula staminodit antici apice profunde indentata, indentatione in pagina abaxiali decurrenti; glandulae 4 staminodiorum lateralium apice non profunde inden- tatae. Stylus anticus 3.3-3.7 mm longus, erectus vel parum arcuatus versus petalum posticum, styli postict 3.4—4 mm longi, divergentes, parum inclinati versus petala postico-lateralia, semper anticum superantes; stigmata Capitata. Samara ignota. Scandent shrub. Vegetative branches velutinous-tomentulose. Stem stipules 7-8.5 mm long, (7.2—) 8-12 mm wide, broadly cordate, apex acute, glabrous or sometimes very sparsely velutinous or tomentulose adaxially, tomentulose- velutinous or woolly abaxially, persistent; inflorescence stipules gradually de- creasing in size toward the distal nodes, the smallest not less than 4.1 mm long, 4mm wide, concave, more densely pubescent than the cauline, persistent or sometimes deciduous. Stem leaves with the petioles 1-5 mm long, densely tomentose-velutinous, laminas 3-9.5 cm long, 2.7-6.7 cm wide, ovate or ellipti- cal, sometimes suborbicular, apex apiculate, base cordate, rugose, tomentulose or velutinous above, woolly below, a pair of sessile glands at the base at the costa, each gland 0.9-2.3 mm in diameter, sometimes with a second, smaller pair (0.6— 1.2 mm in diameter) above, below or adjacent to the larger pair. Inflorescence leaves subsessile to sessile, laminas abruptly smaller than the cauline, the smallest 29 not less than 1.4 mm long, 0.4 mm wide, shape of the more proximal like the cauline, the more distal linear to subulate, apex acuminate, tomentose- velutinous or sparsely so above, woolly below, a pair of glands at the base at the costa, each gland Q.6—2 mm in diameter, sometimes with a second, smaller pair (0.2— 0. J mm in diameter) adjacent to the larger pair, one or both pairs some- times fused, or sometimes a large gland on the costa with two large or small glands adjacent to it, or these three sometimes fused. Inflorescence terminal or axillary, the axes tomentulose-velutinous, Bes of the primary axis 1.7- 8.2 cm long, 2° axes 1.5-13.6 cm long, 3° s 1.2-2 cm long, subsidiary axes absent; umbels 4-flowered, one umbel es tone primary and secondary peduncles absent. Bracts 0.8-2.2 mm long, 0.2-0.7 mm wide, narrowly tri- angular, glabrous adaxially, tomentulose-velutinous abaxially; bracteoles 0. 1— 0.4 mm long, 0.2-0.3 mm wide, linear, oblong, semicircular, or triangular, both surfaces glabrous or with a few scattered hairs, or sometimes tomentulose- velutinous abaxially and/or on the margins. Pedicels 10-20 mm long, tomentose. Sepals 3.8-6 mm long, |.4-2.7 mm wide, adaxially the proximal %—% glabrous, the distal 4—’% tomentulose, glands (1.2-) 1.4—2.2 mm long, 0.7-1.1 mm wide. Claw of the lateral petals 2.3-3 mm long, limb 8.8-10.8 mm long, 8.5—11 mm wide, orbicular, base briefly attenuate, fimbriae up to 0.4 mm mee) glandular. Claw of the posterior petal ca 3.7-4 mm long, limb 6-8 mm long, 4.8-6.2 mm wide, elliptical or narrowly obovate, fimbriae up to 0.2 (-0.4) mm long, capitate- glandular. Filaments of the lateral stamens 4—-4.3 mm long, posterior filament 3-3.5 mm long, slightly curved toward the posterior petal, anthers 1.2—1.4 mm long, glandular connectives 0.8—1 mm long. Pollen grains 50-57 ym. Filament of the anterior staminode 4.14.5 mm long, anterior-lateral filaments 3.5-3.8 mm long, posterior-lateral filaments 3-3.5 mm long, slightly inflexed between the posterior styles, anterior and anterior-lateral glands 1.4-1.6 mm long, 1.3- 1.5mm wide, curved posterior-lateral glands 1.2—-1.4 mm long, 1.1-1.3 mm wide, slightly curved, anterior gland with a deep indentation across the apex and decurrent on the abaxial face, the anterior-lateral and posterior-lateral glands with a shallower indentation. Anterior style 3.3-3.7 mm long, 0.3-0.4 mm wide, shorter than the posterior two, erect or very slightly inclined toward the poste- rior petal, posterior styles 3.44 mm long, divergent, slightly curved toward the posterior-lateral petals, all styles glabrous or more commonly with scattered hairs in the proximal 4 of their lengths, stigmas 0.3-0.5 mm in diameter, capitate. Samara not seen. Type. Occhiont 5009. Brazil. Minas Gerais: Serra do Cipo, estrada da Conceigao, 26 Jun 1973 (holotype RFA, isotype MICH). Distribution. (Fig. 4). Known only from the Serra do Cipo of Minas Gerais. ZIL. Minas Gerais: Serra do Cipo, estrada da Représa, Occhioni 5054 (MICH, RFA); Serra do Cips. Tere 62, 63 (R), Damazio 2054 (RB), Glassman & Gomes, Jr. 8064 (SP), Vidal s.n. (R 108249). Peixotoa cipoana is unique in the genus in that its large staminode glands are recurved. All five glands have an indentation across the apex and decurrent on the abaxial face. In the posterior-lateral two the indentation is sometimes shal- low. P. cipoana 1s also the only species in which the upper surface of the lamina of the stem leaves is sometimes tomentulose. The vesture is always velutinous in all the pubescent species except for P. adenopoda and P. sericea, in which it is sericeous. P. cipoana is readily separated from the other two species in the Serra do Espinhago in which the leaves are densely tomentose or woolly below. P. tomentosa is recognized by its inflorescence in which the terminal nodes bear three umbels and thus are 12-flowered. Also, its staminode glands are straight and smooth. P. spinensis differs, in addition to its smaller, straight staminode glands, in its leaf glands which are borne on the surface of the lamina away from the margin and costa. The leaf glands of P. cipoana are basal. 30 Collected in flower in April, June, and July; two fruiting collections from April and July. 9. Peixotoa cordistipula Adr. Jussieu, Arch. Mus. Hist. Nat. Paris 3: 431. 1843. Fig. 19. Scandent shrub or subshrub or vine up to 1.5 m. Vegetative branches velutinous or velutinous-tomentose, pubescence sometimes abraded from older parts. Stem stipules 14-30 mm long, 10-20 mm wide, triangular or cordate- triangular, apex acute, notched or sometimes acuminate, velutinous adaxially, densely velutinous-tomentulose abaxially, deciduous; inflorescence stipules like the cauline or sometimes the most distal a little smaller, the smallest not less than 9mm long, 7.5 mm wide, persistent. Stem leaves with the petiole 4.5-21 mm long, densely velutinous-tomentose, lamina 7—-20.3 cm long, 3.5-14.5 cm wide, ovate or elliptical or sometimes orbicular, apex apiculate, base cordate or slightly so, velutinous above, tomentose below, usually densely so, with a pair of sessile glands at the base at the costa or sometimes halfway on the petiole, rarely up to 0.5 mm above the base at the costa, each gland 1I-3.3 mm in diameter. In- florescence leaves with the petiole (0.6—) | mm or more long, laminas abruptly smaller than the cauline, the smallest not less than (0.4—-) 1.2 mm long, (0.2-) 1 mm wide, shape of the more proximal like the cauline, the more distal ovate or narrowly elliptical or lanceolate or linear, apex acuminate, sparsely velutinous or rarely glabrous above, densely velutinous below, with a pair of glands halfway on the petiole, each gland (0.4—) 0.7-2.1 mm in diameter, sometime with a second pair of smaller glands (0.4—1.4 mm in diameter) above or adjacent to the pair of larger glands. Inflorescence terminal or axillary, the axes densely velutinous, internodes of the primary axis 0.8-10 cm long, 2° axes 2-13 cm long, 3° axes 1—4.6 cm long, 4° axes 1.7-3.2 cm long, 5° axes 0.9-1.6 cm long, subsidiary axes absent; umbels 4-flowered, one umbel per node, primary peduncles absent, secondary peduncles rarely present, up to | mm long; pedicels commonly ar- ranged around a large glandular mass up to 2.5 mm in diameter, often sparsely velutinous, or the glandular tissue absent. Bracts (0.2—) 0.4-2.3 mm long, 0.2- 0.5 mm wide, triangular or linear, glabrous or tomentulose abaxially or on the margins; bracteoles 0.1-0.4 mm long, 0.1-0.4 mm wide, triangular, linear or subulate, glabrous. Pedicels 11-32 mm long, densely velutinous, usually a little longer in fruit. Sepals 5—7.1 mm long, 2-4 mm wide, adaxially velutinous or sometimes the proximal 43-4 glabrous, the distal 4—*4 pubescent with T-shaped hairs, glands 1.5-2.5 mm long, 0.9-1.1 mm wide. Claw of the lateral petals 2.2— 3 mm long, limb 13-16 mm long, 11-15 mm wide, orbicular, base briefly attenu- ate or almost truncate, fimbriae up to 0.5 (-0.7) mm long, glandular. Claw of the posterior petal 3.8-4.5 mm long, limb 9.5-11.5 mm long, 8-11.5 mm wide, obovate, base briefly attenuate, fimbriae up to 0.6 mm long, those of the proxi- mal 12—%4 capitate-glandular, the distal 4—“% fimbriate-glandular. Lateral stamen filaments 4—5.2 mm long, posterior filament 3.5-4.5 mm long, usually shorter than the lateral filaments, very slightly inclined toward the posterior petal, anthers 1.4—-1.6 mm long, glandular connectives 0.8-1 mm long. Pollen grains 50-57 wm. Anterior staminode filament 4.5—5.3 mm long, exceeding the ante- rior-lateral two, anterior-lateral filaments 4.2—4.8 mm long, usually exceeding the posterior-lateral two, posverior-lateral filaments 3.7—4 (—4.5) mm_ long, shghtly em between the aaa styles, anterior and anterior-lateral glands (1.3-) 1.5-1.9 mm long, (1.2-) 1.5-1.8 mm wide, posterior-lateral glands 1.1—-1.5 mm ioe 1.2-1.3 mm wide, all eee with the apex and abaxial face smooth. Anterior style (4.4—) 5-5.3 mm long, 0.3-0.4 mm wide, always at least a little longer than the posterior two, slightly arced toward the posterior petal, posterior styles 4.2-5 mm long, 0.3-0.4 mm wide, divergent, slightly arced 31 toward the posterior-lateral petals, all styles with scattered hairs in the proximal 73—Y2 of their lengths, stigmas 0.4-0.5 mm in diameter, capitate. Carpophore up to 4 mm long. Torus up to 7 mm high. Samara with the dorsal wing 2.3-3.4 cm long, 1.6-1.9 cm wide, upper margin arced, lower margin slightly arced or sigmoid, slightly erose, lateral wings 4.5-5.5 mm wide, 9-10 mm high, rectangu- lar, areole 3.5—5 mm long, 4-7 mm wide, nut ca 5.8-ca 6.5 mm long, ca 3- 5.5 mm in diameter, mature seeds not seen. : Type. Martius 656. Brazil. Mato Grosso: “In morro do Ernesto prope Cujaba [Cuiaba] .. .” (holotype P!, isotypes BM! BR! F! G! GH! K! LE! M! MO! NY! S! W)). Distribution. (Fig. 5). Central Mato Grosso and eastern Bolivia; cerrado, dry woodlands, and at forest margins. BOLIVIA. Chiquitos, D’Orbigny 729 (P). BRAZIL. Mato Grosso: Mun. Rondonopolis, Serra da Petroliva, Hatschbach 34713 (MBM, MICH); Caceres, Giran, Hoehne 158 (R); Houllet cat. no. 1051 (P); Serra Azul, ca 85 km S of Xavantina, 550 m, /rwin et al. 17176 (MICH, NY, UB); Villa Maria, Kuntze s.n. (NY); inter Cuyaba [Cuiaba] et Diamantino, Lindman A3533 (A, F, G, S); Cuyaba [Cuiaba], Malme 1610, 1919 (S); Santa Ana da Chapada, Malme s.n.(S); ca 11 km W of KM 90, Xavantina-Aragarg¢as Road, Vale dos Sonhos, 700 m, Philcox & Freeman 4696 (K, NY, UB); vicinity of Buri, Chapada dos Guimaraes, 720 m, Prance et al. 19239 (MICH); Serra da Chapada et Cuyaba [Cuiaba], Riedel 1013 , S); Santa Ana da Chapada, Robert 347b (K); between Sao Luiz Caceres and Mato Grosso, Sandeman 2134 (K, NY); Rio Juruena, Aripuana, Silva 3193 (MO, NY); Cuyaba [Cutaba], da Silva Manso (Lhotzky 99) (G, W, syntype of P. macrophylla); Morro do Ernesto, Cuyaba [Cuiaba], da Silva Manso (Lhotzky 117) (BR); Santa Ana da Chapada, Sladen 347 (BM); H. Smith s.n. (R 19721); Veloso 1235, 1311, (RB). _ — The dense, white pubescence of the vegetative parts of P. cordistipula gives the plants a silvery-grey appearance. The stem stipules are 14-30 mm long and 10-20 mm wide. Those of the inflorescence are also large, at least 9 mm long and 7.5 mm wide, and persist even after the inflorescence leaves and pedicels are shed. In umbels of P. cordistipula the pedicels are often arranged around a central, glandular mass, up to 2.5 mm in diameter. The only other species with : 4 A \ a A hee a 4 A a: f Peixotoa 4 cordistipula @ gardneri | @ glabra @ goiana N r jena on FIG. 5. Distribution of P. cordistipula, P. gardnert, P. glabra, and P. gotana. o2 such glandular tissue is P. gardneri, of northwestern Goias, which has smaller, deciduous stipules. The western part of the range of P. magnifica overlaps with the range of P. cordistipula. Both species have petiolate inflorescence leaves, and large flowers and fruits. pies are readily separated by their styles. Uhose of P. magnifica are 7-9.2 mm long. The anterior one is strongly arced toward the posterior petal and bears an oblong stigma at the apex but laterally on the abaxial surface. ‘Vhe styles of P. cordistipula are all less than 5.5 mm long and all stigmas are capitate. Both species produce large inflorescences, but in P. mag- nifica the flowers all open about the same time, while in P. cordistipula only a few are open at any one time Collections of P. reticulata with large stipules and/or from the western part of Brazil, from Bolivia, and from Paraguay have often been assigned to P. cordistipula. All the ones seen are apomicts with closed anthers that contain mostly aborted pollen. These plants have smaller and deciduous stipules and shorter staminode filaments than P. cordistapula. They lack the central, glandular mass in the umbel. Collected in flower from April through August, in fruit in June and July. 10. Peixotoa gardneri C. Anderson, sp. nov. Pie. Teg, Frutex (7). Rami juniores dense aureo-velutini, vetustiores glabri; pili pes ca 0.5-1 mm longus. Stipulae vegetativae 7.5-10 mm longae, acutae vel bifidae, deciduae vel persistentes. Petioli 3-7 mm longi. Laminae 7.8-11.5 cm longae, 3.7-7 cm latae, ovatae vel anguste ellipticae vel lanceolatae, supra aureo- velutinae, subtus aureo-velutinae necnon ferentes pilos T-formes in venis, basi juxta costam biglandulosae vel interdum glandulis in lamina prope basim. In- florescentia terminalis, ramis dense aureo-velutinis, umbellis singularibus. Bracteae bracteolaeque praesentes. Pedicelli 16-24 mm longi. Sepala 5-7 mm longa. Limbus petalorum lateralium 11-11.5 mm longus, orbicularis, margine glanduloso-fimbriato. Limbus petali postici ca 8.5 mm_ longus, orbicularis, margine % proximalibus glandulis capitatis ornato 3 distali glanduloso- fimbriato vel glanduloso-denticulato. Stylus anticus 3.2-3.7 mm longus, posticis semper brevior, erectus, styli postici 3.5-4.2 mm longi, inaequales, erecti vel parum divergentes et inclinati versus petala postico-lateralia; stigmata capitata. Samara ignota. Shrub? Vegetative branches densely golden velutinous, stalks of the hairs usually 0.5-l1 mm or more long, pubes scence abraded from older parts. Stem stipules 7.5-10 mm long, 6.7-7.5 mm wide, cordate, entire with the apex notched, or bifid, glabrous adaxially, densely golden velutinous abaxially, deciduous or persistent; inflorescence stipules gradually decreasing in. size toward the distal nodes, the smallest not less than 5.5 mm long, 5.2 mm wide, apex of the more proximal notched, of the more distal acute, adaxially finely tomentulose, densely so in the proximal %—', sometimes the distal 24—%4 glabrous or glabrate, abaxially saa golden tomentose or woolly, deciduous. Stem leaves with the petiole 3-7 mm long, densely golden velutinous, laminas 7.8-11.5 cm long, 3.7—7 cm nae ovate or narrowly so or narrowly elliptical or lanceolate, apex apiculate, base cordate, golden velutinous above, golden velut- nous and pubescent with T-shaped hairs on the veins below, a pair of sessile glands at the base at the costa or sometimes on the surface of the lamina up to 2mm above the base, up to 0.5 mm from the costa. Inflorescence leaves of the proximal nodes petiolate, the more distal sessile, laminas abruptly smaller than the cauline, the smallest not less than 3.2 mm long, 0.6 mm wide, shape of the more proximal lanceolate or linear-lanceolate, of the more distal linear or sub- ulate, apex acuminate, tomentose or sparsely tomentulose or glabrous above, densely golden velutinous-tomentose below, a pair of glands at the base of the 33 smaller glands (0.4-0.5 mm in diameter) above and/or adjacent to it. In- florescence terminal, the axes golden velutinous or velutinous-tomentulose, internodes of the primary axis (0.7-) 2-9 cm long, 2° axes 0.2-8.7 cm long, 3° axes ().6—-1.5 cm long, subsidiary axes absent; umbels 4-flowered, one umbel per node, primary and secondary peduncles absent; the pedicels sometimes ar- ranged around a glandular mass up to 0.8 mm in diameter, glabrous or sparsely to densely velutinous. Bracts 0.5-1.5 mm long, 0.3-0.7 mm wide, oblong, tri- angular or semicircular, glabrous adaxially, glabrous or velutinous or tomentose abaxially and/or on the margins; bracteoles 0.2—0.4 mm long, 0.2-0.3 mm wide, triangular, glabrous adaxially, glabrous or velutinous or tomentose abaxially and/or on the margins. Pedicels 16-24 mm long, velutinous or velutinous- tomentose, pubescence often dense. Sepals 5-7 mm long, 2.6-3.4 mm wide, adaxially densely tomentulose or glabrous in the proximal ¥%, the distal % densely tomentulose, glands 1.5-1.8 mm long, I-1.3 mm wide. Claw of the lateral petals 2.2-2.5 mm long, limb 11-11.5 mm long, ca 12 mm wide, orbicu- lar, base briefly attenuate, fimbriae up to 0.5 mm long, glandular. Claw of the posterior petal 4.2-5 mm long, limb ca 8.5 mm long, ca 9 mm wide, orbicular, fimbriate or denticulate, the fimbriae or teeth up to 0.3 mm long, those of the proximal % capitate-glandular, those of the distal 4% denticulate- or fimbriate- glandular. Filaments of the lateral stamens 4.2—4.3 mm long, posterior filament ca 3.6 mm long, erect or very slightly inclined toward the posterior petal, anthers 1.5-1.4mm_ long, glandular connectives 0.7-0.8 mm _ long. Pollen grains 50-57 wm. Filaments of the anterior and anterior-lateral staminodes 3.6—-3.8 mm long, posterior-lateral filaments ca 2.5 mm long, curved toward the posterior styles but not inflexed, all glands 1.6-1.7 mm long, anterior gland ca 1.5 mm wide, anterior-lateral and posterior-lateral glands [-1.1 mm wide, all glands with the apex and abaxial face smooth. Anterior style 3.2-3.7 mm long, ca 0.4 mm wide, shorter than the posterior two, erect, posterior styles 3.5-4.2 mm long, unequal, erect or very slightly divergent and inclined toward the posterior- lateral petals, all styles glabrous, stigmas 0.4—0.5 mm wide, capitate. Samara not seen. Type. Gardner 3069 p.p. Brazil. Goias: Serra da Mangabeira [Chapada das Mangabeiras], Sep 1839 (holotype K, isotype K). eixotoa gardneri is known only from the type (Fig. 5). It resembles P. anadenanthera and especially P. hirta in that the hairs of the younger branches ave very long stalks which are at last 0.5 mm long, but commonly are | mm or more. P. anadenanthera is easily recognized, since it is the only species in which the anthers lack the glandular connectives. P. hirta differs from P. gardneri in its smaller flowers. The limb of the posterior petal is obovate. In P. gardneri the limbs of all petals are orbicular. The pedicels of P. gardneri are sometimes ar- ranged around a central, glandular mass which is often hidden in the dense pubescence. P. cordistipula of Mato Grosso is the only other species in which such glandular tissue is found in the umbel. The staminode glands of P. gardneri are unusual in that the adaxial groove extends nearly to the apex. However, this has been observed only in the two individuals of the type collection and may not be typical of the species. Extremely long adaxial grooves occur occasionally also in individuals of other species. spe costa, each gland 0.7-2 mm in diameter, sometimes each gland with | or 2 cles. Gardner 3069 is a mixed gathering. The two specimens deposited at K are P. gardneri. Vhe specimen at BM is P. goiana. This species is named for the intrepid George Gardner, one of the impor- tant early collectors of the Brazilian flora. He ventured into many remote areas, such as the Chapada das Mangabeiras where P. gardnert was collected, which have not been visited by botanists since his time. 34 11. Peixotoa soi Adr. Jussieu in St. Hilaire, Fl. Bras. Mer. 3: 60. a 172. 1832 [1833]. Fig. 17. Erect or procumbent, often virgate shrub up to 2 m. Vegetative branches shiny, purplish-brown, very sparsely velutinous when young, soon glabrous. Stem stipules 8-20 mm long, 7-10 mm wide, cordate or narrowly triangular, bifid or sometimes entire and the apex acute, glabrous adaxially, glabrous or sometimes sparsely velutinous abaxially, persistent; inflorescence stipules gradu- ally decreasing in size toward the distal nodes, the smallest not less than 5 mm long and wide, apex of the most distal acute or notched, commonly per- sistent. Stem leaves with the petiole 1.5-8.5 mm long, glabrous, lamina 5— 12.2 cm long, 2.6—7.5 cm wide, elliptical, narrowly ovate or lanceolate, apex apiculate or acuminate, base cordate, glabrous above and below or rarely with a few scattered hairs along the margins, costa and major veins, a pair of sessile glands at the base at the costa or sometimes halfway on the petiole or rarely on the surface of the lamina up to 1.5 mm above the base, up to | mm from the costa, each gland 0.8—2.4 mm in diameter, sometimes one or both glands absent. Inflorescence leaves often petiolate at the most proximal nodes, subsessile or sessile at the most distal nodes, laminas gradually decreasing in size toward the distal nodes, the smallest not less than 1.7 mm long, 0.3 mm wide, lanceolate or triangular or linear or subulate, acuminate or rarely caudate, glabrous above and below or sometimes very sparsely velutinous, a pair of glands at the base, each gland 0.5-1.5 mm in diameter, these sometimes fused, or with 3-4 separate or fused ae or with a second pair of smaller glands (0.5-0.7 mm in diameter) borne above the pair of larger glands. Inflorescence terminal, the axes glabrous or with a few scattered hairs or sometimes sparsely velutinous, internodes of the primary axis 2.7—10.7 cm long, 2° axes 1.7-9 cm long, 3° axes 0.8-5.9 cm long, 4° axes |.6-2.6 cm long, 5° axes 0.7-0.9 cm long, subsidiary axes absent; umbels 4-flowered, one umbel per node, primary and secondary peduncles absent. Bracts 0.7—1.6 mm long, 0.4—-0.9 mm wide, narrowly triangular, apex obtuse, glabrous or with scattered hairs distally and/or on the margins; bracteoles 0.1— 0.3 (-0.6) mm long, 0.2—-0.4 mm wide, triangular, apex obtuse, glabrous. Pedicels 11-25 mm long, velutinous, sometimes a little longer in fruit. Sepals (3—) 3.8- 6 mm long, (1.3-) 2-2.7 mm wide, adaxially the proximal 4 glabrous, the distal +s tomentulose, glands 1.1-1.9 (-2.7) mm long, 0.8-1.9 mm wide. Claw of the lateral petals (1—) 1.3-3.3 mm long, limb (7.5-) 9.3-14 mm long, (7.4—) 8-14 mm wide, orbicular, base truncate or attenuate, fimbriae up to 0.5 mm long, glandu- lar or eglandular. Claw of the posterior petal (2.5-) 3.3-4.6 mm long, limb (4—) 5,6-7.5 mm long, (3.2-) 3.8-4.5 mm wide, elliptical or oblong or slightly obovate, base truncate, all fimbriae up to 0.5 mm long and capitate-glandular, or those of the distal 4 up to 0.7 mm long and fimbriate-glandular. Lateral stamen fila- ments (2.8-) 3-5 mm long, posterior filament 2.5-4 mm long, usually shorter than the lateral four, slightly arced toward the posterior petal, anthers 1-1.4 (1.6) mm long, glandular connectives 0.5-0.9mm long. Pollen grains 50- 54 ym. Anterior and anterior-lateral staminode filaments (3-) 3.5—4.7 mm long, posterior-lateral filaments 3-3.9 mm long, usually shorter than the anterior three, shghtly inflexed between the posterior styles, glands 1.1-1.5 mm long, I-1.5 mm wide, the anterior gland often a little larger than the other four, the anterior and anterior-lateral glands with a deep indentation across the apex and decurrent on the abaxial face, posterior-lateral glands smooth. Anterior style 3—4 mm long, 0.2-0.3 mm wide, gently arced toward the posterior petal, poste- rior styles 3.4—4.5 mm long, 0.2-0.3 mm wide, divergent, especially the distal arced toward the posterior-lateral petals, all styles glabrous or sometimes with scattered hairs in the proximal 3 of their lengths, stigmas 0.4-0.5 mm in di- ameter, capitate. Carpophore up to 3.7 mm long. Torus up to 4.5 mm high. 35 Samara with the dorsal wing 1.6—2.7 cm long, 0.9-1.6 cm wide, upper margin gently sigmoid, lower eee straight, sometimes Sues erose, lateral wings 1.5-4 mm wide, 7-12 m igh, areole 3.5-5.5 mm long, 2.7-5.6 mm wide, nut 5.7-6.5 mm high, 4.2-6 mm in diameter, seed 5.7-6.1 mm ne outer cotyledon 7.6-7.9 mm long, 2.6-3 mm wide, folded at ca % of its length, inner cotyledon 4.8-5.1 mm long, 1.9-2.2 mm wide, folded at ca 4— of its length Type. St. Hilaire s.n. Brazil. Minas Gerais: “in pascuis juxta Tijuco [Diamantina]” (holotype P!, isotype P!) Distribution. (Fig. 5). nena only from the central part of the Serra do Espinhago of Minas Gerais, especially from the environs of Diamantina; cerrado. . Minas Gerais: ca 18 km SW of Diamantina on road to Curvelo, 1400 m, Anderson SI1IT (MIC a NY); Mun. Gouveia, 35 km SW of es on BR-259, 1250 m, Anderson 11549 (C, MICH); ca 10 km SW of Diamantina, een et al. 35189 (MO, NY); Diamantina, Brade 13722, 13775, oe 13781 (RB); Serra dos Cristais, Diamantina, Glaziou 18937 (C, ; Mun. Sérro, rod. MG-2, entre Serro e Datas, Hatschbach 28921 (NY, P); ca 20 km S of Sie Joao ida Chapada, 1200 m, hee et al. 28371 (MIC ae MO, ae );ca lO kmS bf Sho Tooaae hapada, 1200 m, Jrwin et al. 28412 (MICH, MO, NY); Serra da Lapa, Langsdorff s.n. (LE); Serra do Sao Gongalo prope Diamantina, do Linea (Herb. ode 12816) (RB); just W ' Sérro, on road from Conceicao to Diamantina, le 49144 (MICH, NY); between Diamantina and Gouveia, Maguire 49164 (NY); a Frio, ad Tejuco [Diamantina], Martius as M); Ouro Preto, Villa Rica [Ouro Preto], ee s.n. (two collections) (M); Diamantina, crest of Serra do Rio Grande, 1300 m, Mexia 5804 (A, BM, G, GH, K, MICH, MO, NY,R, 5S, US); a, Pereira 1535 (RB); Riedel XII p.p. (LE, S); Serra ie Lapa, Riedel 1156 (LE, S); ad Riacho das Varas prope Diamantina, Schwacke 8370 (RB); Tejuco [Diamantina], Vauthier 459 (G, GH, P); Morro dos Cruzeiros, Diamantina, ca 1300 m, Vidal s.n. (R 72121). = Peixotoa glabra is the only glabrous species in Minas Gerais. The very sparsely pubescent P. trwinii and some glabrous forms of P. reticulata occur in the same area, but are readily distinguished by the staminode glands. Those of P. glabra are of equal size. The anterior and anterior-lateral three have an indentation across the apex and decurrent on the abaxial face. The glands of P. irwinii are smaller, unequal, and have a smooth apex. The abaxial face of the anterior and anterior-lateral glands have a shallow, broad depression. Also, the petioles of P. irwinw are golden pubescent. Those of P. glabra are glabrous. In P. reticulata the staminode glands are smooth and usually unequal. Its flowers are usually larger than those of P. glabra. Anderson 8564 1s perhaps a hybrid between P. glabra and P. reticulata. The specimens look like P. reticulata, but the foliage and stems are glabrous and the staminode glands are indented. The flowers, as in nearly all collections of P. reticulata, have closed anthers containing aborted pollen. Peixotoa glabra has sometimes been confused with the usually glabrous or glabrate P. hispidula, which grows along the coast from Bahia to Sao Paulo. It differs in its usually large flowers born in a single umbel or a small inflorescence. (he staminode glands are strongly dimorphic. The inflorescence leaves are all petiolate. Those of P. glabra are sessile at the more distal nodes. Though P. glabra is apparently another narrow endemic, it is well repre- sented in herbaria. Diamantina was an important destination for early collectors and still is much visited today. The locality of two Martius collections in the Munich herbarium is given as Ouro Preto, another frequently collected area. While it is tempting to ascribe this to a mistake in labeling, it must be remem- bered that the flora of Minas Gerais is still only poorly known. Collected in flower from December through August, in fruit from March through August. 12. Peixotoa goiana C. Anderson, sp. nov. Bigs. 2520); Suffrutex, caules ascendentes usque ad 2 m. Stipulae vegetativae 4-19 mm longae, acutae vel bifidae, plerumque persistentes. Petioli 1—7(—9) mm longi. 36 Laminae (4.3-)5-17 cm longae, 2.3-10.6 cm latae, lanceolatae vel ellipticae vel ovatae vel suborbiculares, supra velutinae raro glabrae, subtus ferentes pilos ‘T-formes vel non nisi in venis vel raro glabrae, basi juxta costam biglandulosae vel interdum glandulis in lamina prope basim. Folia inflorescentiae petiolo brevi vel sessilia, in nodis distalibus linearia vel subulata. Inflorescentia terminalis, umbellis singularibus. Bracteae bracteolaeque plerumque praesentes interdum absentes. Pedicelli 10-34 mm longi. Sepala 4-6.5 mm longa. Limbus petalorum lateralium 9.6-12(-15) mm_ longus, orbicularis vel late ellipticus, margine glanduloso-denticulato vel glanduloso-fimbriato interdum eglanduloso. Limbus petali postici 7.5-8.3(-10) mm longus, orbicularis vel interdum late obovatus, margine glandulis capitatis ornato vel margine % proximalibus glandulis capitals ornato 1% distali glanduloso-fimbriato vel glanduloso-dentculato. Glandulae 3 staminodiorum anticorum plerumque apice profunde indentatae, indentatione in pagina abaxiali decurrenti, interdum laeves. Stylus anticus (2.6-)3.1-4.3(-4.6) mm longus, plerumque erectus raro parum inclinatus versus petalum posticum, styli postici 3.3-4.7 mm longi, divergentes, plerumque saltem parum inclinatt. versus petala postico-lateralia, semper anticum superantes; stigmata capitata. Ala dorsalis samarae |.6-2.7 cm longa, |.1-1.9 cm lata, alae laterales 0.6—2.5 mm latae, 5.5-9.8 mm altae, erosae interdum dissectae. Subshrub with one or few ascending stems to 2m. Vegetative branches densely, usually golden, velutinous or tomentulose- velutinous. rarely glabrous; the pubescence often abraded from older parts. Stem stipules 4-19 mm long, 2.3-16.5 mm wide, cordate or triangular-cordate, sometimes narrowly so, entire with the apex acute or notched, or bifid, finely velutinous or sparsely so or glabrous adaxially, densely velutinous or densely or sparsely tomentose or tomentose-velutinous abaxially, usually persistent; inflorescence stipules of the proximal nodes equal or larger than the cauline, the distal gradually decreasing in size toward the distal nodes, the smallest not less than 5.8 mm long, 5.3 mm wide, cordate, often concave, entire with the apex acute, the most proximal sometimes with the apex notched, or bifid, velutinous or sparsely so adaxially, velutinous-tomentulose abaxially, usually deciduous. Stem leaves with the petiole 1-7 (-9) mm long, densely, usually golden velutinous or rarely glabrous, laminas (4.3—) 5-17 cm long, 2.3-10.6 cm wide, lanceolate or elliptical or narrowly so or ovate or suborbicular, apex apiculate, base cordate, rugose, velutinous or rarely glabrous above, sparsely to densely pubescent with T-shaped hairs or rarely tomentose or rarely the hairs restricted to the veins or rarely glabrous below, hairs usually golden above and below, a pair of sessile glands halfway on the petiole or sometimes at the base at the costa, sometimes borne on the sur face of the lamina |-6 mm above the base, up to 3.8 mm from the costa, rarely at the margin 0.7—1 mm from the costa, each gland 0.5-1.8 mm in diameter, some- times with a second, smaller pair (0.4—0.5 mm in diameter) below or adjacent to ee larger pair. Inflorescence leaves of the proximal nodes with the petiole up to 5 (-6) mm long, the more distal sessile or very rarely with a petiole up to 1.4 mm (one, laminas abr uptly smaller than the cauline or gradually decreasing toward the distal nodes, the smallest not less than 2 mm long, 0.2 mm wide, the more proximal elliptical or narrowly so, the more distal linear, or subulate, apex acuminate, vesture of the more proximal like the cauline, the more distal glabrous above, densely velutinous below, all with a pair of glands at the base at the costa, the glands rarely fused, each gland (0.2-) 0.5-1.8 mm in diameter, sometimes with a second, smaller pair (0.3-1.2 mm in diameter) above or adjacent to the larger pair, or rarely one large gland borne on the costa with two arge glands on either side. Inflorescence terminal, the axes velutinous or tomentulose, usually densely, rarely sparsely so, internodes of the primary axes 3.2-18.4 cm long, 9° axes 1.3—16.5 cm long, 2° subsidiary axes (0.2—) 11:5=-3.5 em long, 3° axes 0.6-7.2 cm long, 4° axes I-3.8 cm long, 5° axes ().9-1.8 cm long; 37 umbels 4-flowered, one umbel per node, primary and secondary peduncles absent. Bracts 0.3-1.3 mm long, 0.3-1.2 mm wide, triangular or semicircular, glabrous, sometimes velutinous or sparsely so abaxially and/or on the margins; bracteoles 0.1-0.5 mm long, 0.2-0.4 mm wide, triangular, glabrous, rarely velutinous abaxially and/or on the margins; bracts and bracteoles sometimes absent. Pedicels 10-34 mm long, (usually golden) densely velutinous or tomen- tose, becoming up to ¥y again as long in fruit. Sepals 4-6.5 mm long, 2.2-3.3 mm wide, adaxially the proximal 4—% velutinous or sparsely so, the distal %4—¥% tomentulose, sometimes entirely sparsely velutinous, glands 1.6-2.6 mm long 0.6-1.2 mm wide. Claw of the lateral petals 2.2-3.2 mm long, limb 9.6-12 15) mm long, 9-12 (—15) mm wide, orbicular or rarely broadly elliptical, base briefly attenuate, denticulate or fimbriate or both, the teeth and fimbriae up to 0.5 (-0.8) mm long, glandular or eglandular. Claw of the posterior petal 4—5 mm long, limb (-10) mm long, 6.8-8 (-9) mm wide, orbicular or rarely broadly obovate, denticulate or fimbriate, teeth and fimbriae up to 0.6 mm long those of the proximal -*4 capitate-g elandular, the distal “’—'’3 denticulate- or fimbriate-glandular, or sometimes all capitate-g] stamens (3.3-) 3. andular. Filaments of the lateral —4.8 mm long, posterior filament 2.6-4.3 mm long, always shorter than the lateral four, erect or very slightly curved toward the posterior petal, anthers 1.5-1.7 mm long, glandular connectives 0.7-0.8 mm long. Pollen grains ca 57 wm. Filament of the anterior staminode 3.54.4 mm long, usually exceeding the anterior-lateral two, anterior-lateral filaments 3.2-3.6 mm long posterior-lateral filaments 2.8-3.5 mm long, always shorter than the anterior lateral two, inflexed between the posterior styles, anterior gland 1.2-1.5 mm long, 1.2-1.3 mm wide, anterior-lateral glands ca 1.2 mm long, 1.1-1.2 mm wide, posterior-lateral glands 1—1.1 mm long, 0.9-1 mm wide, anterior and anterior-lateral glands with a deep indentation across the apex and decurrent on the abaxial face or sometimes smooth, posterior-lateral glands smooth. Anterior style (2.6—-) 3.1-4.3 (-4.6) mm long, 0.3-0.4 mm wide, erect or rarely slightly inclined toward the posterior petal, posterior styles 3.3-4.7 mm long, ca 0.3 mm wide, always longer than the anterior style, divergent, at least slightly inclined toward the posterior-lateral petals, all styles glabrous or with scattered hairs in the pi oximal 4 of their lengths, stigmas 0.3-0.5 mm wide, capitate Carpophore up to 3.6mm long. Torus up to 6.2 mm high. Samara with the dorsal wing 1.6—2.7 cm long, |.1—1.9 cm wide, upper margin sigmoid or arced, lower margin sigmoid, arced or semicircular, slightly erose, lateral wings 0.6-2.5 mm wide, 5.5-9.8 mm high, rec tangulz ir, erose or irregularly dentate or dissected, areole 4.5-6.1 mm long, 3.5-5.5 mm wide, nut 5.5-7 mm long, 3 —5) mm in diameter, seed 6.9-8.2 mm long, outer cotyledon 7 .8-10.8 mm long, 1.6-3.3 mm wide, folded at ca *%% of its length, inner cotyledon 5.1—7.2 mm long, 1.5-—2 mm wide, folded at ca 2/3-6/7 of its length or straight. Type. Irwin et al. 12855. Brazil. Goias: Chapada dos Veadeiros, 14°S, 47°W, ca 15 km W of Veadeiros [Alto Paraiso], 1000 m, 14 Feb 1966 (holotype UB, isotypes MICH, NY). Distribution. (Fig. 5). Central Goias, especially in the Serra dos Veadeiros, Serra dos Cristais, Serra dos Pirineus, and Serra Dourada, and the Distrito Federal; cerrado and campo. BRAZIL. Goias: Chapada dos Veadeiros, ca 20 km N of Alto Pararso, ca eee m, Anderson 6201 (MICH, NY); Serra dos Cristais, 9km S of Cristalina on road to Catalao, 1170 m, Anderson 8090 (NY); Sane Dourada, ca 15 km (str. line) S of Goias Velho, 1000 m, Ander 10024 (MICH, NY); ae dos Pirineus, ca 15 km (str. line) N of Coes de Goias, 1300 m, Anderson 10282 (MICH, Y); Chapada dos Veadeiros, 14°S, 47°30'W, 2.5 km E of Alto Pararso ee & (MICH); Chapada ae Veadeiros, Ferreira 6 (UB); Serra dos pene: Ferreira 8 (UB); Serra da eae [Chapada das Seanad Gardner 3069 p.p. (BM); C isa a dos Veadaeiros, Pouso Alto, 1800 m, Ferreira 10 ; Serra Dourada, Ferreira 32 (UB); 70 km de Braslandia, Vale S. Geronimo, Ferreira 150, ce Chapada dos Veadeiros, mun. nite Paraiso, 16 km by road N of Alto Paraiso, 1600 m 152 38 14°S, 47°W,Gates & Estabrook 103 (MICH); Serra dos Pirineus, Pirenopolis, Giulietti @ Lima 725 (UB); Chapada dos Veadeiros, Pouso Alto, 1800 m, Graziela [Barroso] 609 (UB); Chapada dos Veadeiros, 65 km N of Brasilia, Harley et al. 11432 (K, NY); Serra do Rio Preto, 16°S, 47°W, ca 5 km E of Goias boundary on stags to Guarapua, M. G., 1000 m, /rwin el 10291 (MICH); Serra dos Pirineus, 16°S, 49°W, ca 14 km S of Corumba, 975 m, Irwin et al. 10773 (NY); Serra Dourada, 20 km SE of Goias Walco: 300 m, Ba et al, 11705 (MICH, NY); C hapa dos Veadeiros, 14°S, 47°W, ca 15 km W of Veadeiros [Alto Paraiso], /rwin et al. 12351 (MICH, NY); Serra dos Cristais, 17°, 48°W, ca 10 km S of Cristalina, 1200 m, Irwin et al. 13733 (MICH, NY); ca 87 km SE of Aragargas, 700 m, Irwin et al. 17562 (MICH); Serra dos Pirineus, 15-20 km N of Corumba de Goias on road to Niquelandia, ca 1150 m, /rwin et al. 18616 (MICH, NY), loa (NY); Chapada dos Veadeiros, ca 10 km W of Alto Paraiso, 1000 m, Irwin et al. 24985 (NY); Chapada dos Veadeiros, ca 19 km N of Alto Paraiso, ca 1250 m, Irwin et al. 32799 (MICH, NY); Serra dos Pirineus, ca 18-20 km E of Pirenopolis, 1000 iy er one etal. 34061 (MICH, NY), 34096 (NY); 14 km S of Niquelandia, ca 750 m, Irwin et al. 34743, 4 (NY); Niquelandia, Magds 3671 (RB); Chapada dos Veadeiros, 2 km from Veadeiros [Alto ee Prance & Silva 58200 (MICH, NY); Serra Dourada, Rizzo 3997, 4128, 4442, 4494, 4597, 4638 (RB); mun. Alto Paraiso, Cinturao Verde, Parque Nacional do Tocantins, Rosa 67 (UB). Distrito Federal: Brasilia, Andrade 318 & Emmerich 310 (R), Ferreira 2a, 46, 47, 118 (UB), Gomes 1056 ae 8394 (SP, UB), /rwin & Soderstrom 6083a (NY), Ha 407 (GH), 476 (GH, UB), Occhioni 23 (MICH), Pares et al. 9O86 (UB, US), Sucre 274 (NY, RB, UB); Braslandia, Ferreira 4 (UB); age) of Seer da Contagem, ca 10 km E of Brasilia, 1000 m, iran et al. 8221 (MICH); ca 12 km V ’ Taguatinga on road to Braslandia, 1250 m, Irwin et al. 10683 (N ae Braslandia, Macédo 23 a Chapada da Seek along highway DF-5 about 36 km W of intersection with BR-020, 1280 m, Plowman 9943 (MICH); Fazenda Agua Limpa (Univ. of Brasilia field station), near Vargem Bonita, Ratter et al. 2806, oe 3129 (MICH); Salinas, Weddell 2193 (P). Peixotoa goiana is distinguished by its flowers and inflorescence leaves. All the petals have an orbicular (rarely broadly obovate or broadly elliptical) limb. The margins of the lateral limbs are commonly denticulate or sometimes fimbriate. The anterior style is usually erect or sometimes slightly inclined toward the posterior petal. The anterior and anterior-lateral staminode glands of plants from the Distrito Federal, the Serra dos Pirineus and the Serra dos Cristais have an indentation across the apex and decurent on the abaxial face. Those of the Serra Dourada are smooth. In specimens from the Chapada dos Veadeiros the glands are either indented, often shallowly so, or smooth. ‘The inflorescence leaves of the more distal nodes are linear to subulate and sessile; only [rwin 17562 has inflorescence leaves with tiny petioles. This collection is from the extreme western part of the range at the Mato Grosso border. The abundance of hairs on the vegetative parts is highly variable. Most plants are moderately to densely pubescent, but in some the hairs are very sparsely distributed and on the leaves restricted to the major veins. Glabrous forms are rare (Anderson 8090, Irwin 10773, Macedo 3671), Vhe stem and in- florescence stipules are usually 4-12 mm long. However, in a few collections their length ranges from 9-19 mi The samaras have ebinely 6 narrow lateral wings which are 0.6—2.5 mm wide. In most specimens the wings are at least | mm wide. Only the probably closely related P. hatschbachii has more reduced wings. In that species each lateral wing is a narrow strip less than | mm wide in the proximal ' of the nut or a crest, or the lateral wings are absent. In P. govana the wings extend along the entire length of the nut. In the southern part of its range P. govana overlaps with the apomictic P. reticulata. Some plants that are intermediate, especially in floral characters, occur, but all are apparently apomictic. Collected in flower and fruit throughout the year. 13. Peixotoa hatschbachii C. Anderson, sp. noy. Fig. 15: Frutex erectus usque ad 1.2m. Caules glabri. Stipulae vegetativae 17— 19.5 mm _ longae, plerumque bifidae, deciduae vel persistentes. Petioli 6.5— 10 mm longi. Laminae 6.2—9.3 cm longae, 4.4-7.5 cm latae, ovatae, rugosae, glabrae, basi juxta costam 2— vel 4—glandulosae, glandulis pedicellis. Folia in- florescentiae petiolo brevi vel sessilia, distaliora linearia vel subulata, subtus dense velutina, basi juxta costam 2-6-glandulosa. Inflorescentia terminalis (vel axillaris’), umbellis singularibus. Bracteae bracteolaeque praesentes. Pedicelli 18.3-23 mm longi. Sepala 4.5-5.2 mm longa. Limbus petalorum lateralium 9.8— 10 mm longus, late obovatus vel suborbicularis, margine glanduloso-fimbriato. Limbus petali postici 3.5-3.8 mm longus, obovatus vel late obovatus, margine proxmali glandulis capitatis ornato, % distali glanduloso-fimbriato. Styli sub- aequales, 3.6—-4 mm longi, anticus parum inclinatus versus petalum posticum, postici divergentes, parum inclinati versus petala postico-lateralia; stigmata capitata. Ala dorsalis samarae 1.7-1.9 cm longa, |.3-1.4 cm lata, alae laterales absentes vel multo redactae, minus quam 1.5mm latae, 5.5-7.2 mm altae, extensae e medio nucis ad basim. Erect shrub to 1.2 m. Vegetative branches shiny, purple, glabrous. Stem stipules 17-19.5 mm long, 18-19 mm wide, cordate, bifid, glabrous, persistent or deciduous; inflorescence stipules gradually decreasing in size toward the distal nodes, the smallest not less than 8.3 mm long, 10.5 mm wide, entire, apex acute, tomentulose adaxially, velutinous abaxially, deciduous. Stem leaves with the petiole 6.5-10 mm long, glabrous, laminas 6.2—9.3 cm long, 4.4-7.5 cm wide, ovate, apex apiculate, base cordate, rugose, glabrous above and below, a pair of stalked glands at or just above the base at the costa, each gland 1-1.2 mm in diameter, stalk 0.8—1.2 mm long, commonly each gland with an adjacent smaller gland, 0.8—1.1 mm in diameter, stalk 0.4—1 mm long. The more proximal in- florescence leaves with a petiole up to 5 mm long, the more distal commonly sessile, laminas abruptly smaller than the cauline, the smallest not less than 5.5 mm long, 0.6 mm wide, shape of the more proximal like the cauline, the more distal linear to subulate, apex acuminate, sparsely velutinous or glabrous above, densely velutinous below, glands sessile, sometimes with a pair of glands at the base at the costa, usually with multiple glands: one pair but each large gland with one or two smaller glands adjacent and/or below it, some or all of these sometimes fused, or sometimes with one large gland on the costa and two smaller glands adjacent on each side, or sometimes with two large glands at each side of the costa, some or all of these fused, and with one or two smaller glands below them, each large gland I-1.5 mm in diameter, each small gland 0.5-1 mm in diameter. Inflorescence terminal (or axillary ?), the axes sparsely velutinous to glabrate, internodes of the primary axis 10-13 cm long, 2° axes 5.8-19 cm long, 2° subsidiary axes up to 2.7 cm long, 3° axes 3.8-7.1 cm long, 3° subsidiary axes .2-1.5 cm long, 4° axes 1.8-3.1 cm long, umbels 4-flowered, one umbel per node, primary and secondary peduncles absent. Bracts 1.8—4 mm long, 0.6— 9mm wide, narrowly triangular, glabrous adaxially, sparsely velutinous abaxi- ally; bracteoles 0.2—0.3 mm long, 0.3-0.5 mm wide, semicircular, glabrous. Pedicels 18.3—23 mm long, velutinous, slightly longer in fruit. Sepals 4.5-5.2 mm long, 2.1-2.6 mm wide, adaxially the proximal %—%4 glabrous, the distal 4-1 tomentulose, glands 1.7—2 mm long, 0.6—1 mm wide. Claw of the lateral petals 2-2.2 mm long, limb 9.8-10 mm long, 9-10 mm wide, broadly obovate to sub- orbicular, base attenuate, fimbriae up to 0.6 mm long, glandular. Claw of the posterior petal 3.5-3.8 mm long, limb 5.5-6.5 mm long, 5—5.8 mm wide, obovate or broadly so, fimbriae up to 0.4 mm long, those of the proximal % capitate- glandular, the distal 4 fimbriate-glandular. Lateral stamen filaments 3.6—4 mm long, posterior filament (3—) 3.5-3.7 mm long, arced toward the posterior petal, anthers 1.5—1.8 mm long, glandular connectives 0.7—0.9 mm long. Pollen grains 50-57 um. Anterior staminode filament 3.8—-4 mm long, anterior-lateral fila- ments 3.6-3.7 mm long, posterior-lateral filaments 3.2-3.4 mm long, inflexed between the posterior styles, anterior and anterior-lateral glands 1.3-1.4 mm long, I.1-1.2 mm wide, posterior-lateral glands ca 1.1 mm long, ca 0.8 mm wide, — 40 all glands with the apex and abaxial face smooth. Styles 3.64 mm long, equal or the anterior style a little longer or shorter than the posterior two, anterior style ca 0.4 mm wide, slightly inclined toward the posterior petal, posterior styles ca 0.3 mm wide, divergent, slightly inclined toward the posterior- -lateral petals, all with some scattered hairs in the proximal 4 of their lengths, sugmas 0.5—0.6 mm in diameter, capitate. Car se up to 3 mm long. Torus up to ca 5 mm high. Samara with the dorsal wing 1.7-1.9 cm long, 1.3-1.4 cm wide, upper margin gently arced, lower margin slightly erose or sometimes entire, lateral wings ab- sent or reduced to a crest or to a narrow, erose strip less than 1.5 mm wide, 5-7.2 mm high, usually extending from the middle to the base of the nut, Sole 5-5.3 mm long, ca 5 mm wide, nut 6-7 mm long, 4.7—5 mm in diameter, seed ca 6.8 mm long, ca 3 mm wide, outer cotyledon ca 9.7 mm long, ca 2.8 mm wide, folded at 4 of its length, inner cotyledon ca 5.2 mm long, ca 1.6 mm wide, aeot ohe 4/5 of its length. _ Hatschbach 36675. Brazil. Goias: BR-135, 10 km E of Anapolis, 22 May eee (holotype MBM, isotypes C, MICH). Peixotoa hatschbachii is known only from the type (Fig. 6). It is distinguished by its stalked leaf glands and by its glabrous vegetative branches and stem leaves. The samaras are unique in that the lateral wings are absent, or reduced to a crest or a narrow strip of tissue, less than 1.5 mm wide, which usually extends from the middle to the base of the nut. The stipules are unusually large. Vhose of the stem are 17-19.5 mm long and wide. The smallest inflorescence stipule seen ts 8.3 mm long an 10.5 mm wide. P. hatschbachii occurs within the range of P. goiana, to which it may be closely related. Plants of P. goiana are only rarely glabrous. ‘These glabrous forms as well as the glabrous or very sparsely pubes- cent P. psilophylla from east-central Mato Grosso differ from P. hatschbachi by their sessile leaf glands, smaller stipules, and larger posterior petals. This species is named in honor of Gert Hatschbach, an astute student as well as a prolific and discerning collector of the flora of central and southern Brazil. Peixotoa / ' & hatschbachii @ hirta @ hispidula N © irwinii . \ eee ‘ i : mete ie =o i % \ \ a Baa a : ( Q : A Nock OT, N ~e \ { 2 me ‘ { one { \ ¥ an \ ; ic : od / j i FIG. 6. Distribution of P. hatschbachi, P. hirta, P. huspidula, and P. irwinu. 4] 14. Peixotoa hirta Adr. Jussieu, Ann. Sci. Nat. Sér. II, Bot. 3: 279. 1840. Fig. 20. Erect, subscandent shrub, 1-2 m. Vegetative branches densely but loosely golden velutinous, stalks of the hairs usually 0.5—]1 mm or more long, pubes- cence abraded from older parts. Stem stipules 6-9 mm long, 7-9 mm wide, cordate, entire with the apex acute or notched, or bifid, glabrous or sparsely tomentulose adaxially, velutinous and pubescent a, ‘T-shaped hairs abaxially, deciduous; inflorescence stipules like the cauline or sometimes a little larger, glabrous adaxially, tomentose abaxially, deciduous. Stem leaves sessile or with the petiole up to 6.5 mm long, densely golden velutinous, laminas 7.3-9 cm long, +.7-8.2 cm wide, elliptical or ovate or suborbicular, rugose, apex apiculate, base cordate, loosely golden velutinous and pubescent with golden T- -shaped hairs above, densely pubescent with golden T-shaped hairs below, a pair of sessile glands at the base at the costa, each ag 0.6-1.2 mm in diameter. Inflorescence leaves with the petiole up to 3.2mm long, the more distal sessile, laminas abruptly smaller than the cauline, the smallest not less than 3 mm long, 0.6 mm wide, narrowly lanceolate to subulate, apex acuminate, densely golden veluti- nous or tomentulose above, very densely pubescent with golden T “shaped hairs below, a pair of glands at the base at the costa, each gland 0.6-2.5 mm in di- ameter. Inflorescence terminal, the axes densely but loosely golden velutinous, internodes of the primary axis 4.5—-10.7 cm long, 2° axes ().7-4.7 cm long, sub- sidiary axes absent; umbels 4-flowered, one umbel per node, primary and secondary peduncles absent. Bracts 0.9-1.2 mm long, 0.6—0.8 mm wide, tri- angular, glabrous; bracteoles ca 0.2 mm long and wide, oblong or triangular, glabrous. Pedicels 11-15 mm long, densely golden tomentose. Sepals 4.4— 5.8 mm long, 2.1-3 mm wide, adaxially glabrous, glands 1.5-2 mm long, 0.6— 1.1 mm wide: Claw of the lateral petals 999.4 mm long, limb 9-9.5 mm long, ca 8—8.5 mm wide, orbicular, base truncate, fimbriae up to 0.5 mm long, fimbriate- glandular, sometimes a few fimbriae of the anterior-lateral petals eglandular. Claw of the posterior petal 2.5-2.6 mm long, limb ca 6.5 mm long, ca 5 mm wide, obovate, fimbriae up to 0.6 mm long, capitate- Seaumre Lateral stamen filaments 3.3-3.4 mm long, posterior filament ca 3.2 mm long, erect or very slightly curved toward the posterior petal, anthers 1.7-2 mm long, glandular connectives |—1.2 mm long. Pollen grains 42 wm. Filaments of the anterior and anterior-lateral staminodes ca 3.4 mm long, posterior-lateral filaments ca 3 mm long, inflexed between the posterior styles, anterior and anterior-lateral glands 1.3-1.4 mm long, 1.2—1.3 mm wide, posterior-lateral glands ca 1.1 mm long, ca 0.8 mm wide, all glands with the apex and abaxial face smooth. Anterior style ca 3.2 mm long, ca 0.4 mm wide, slightly arced toward the posterior petal, posterior styles ca 2.8 mm long, ca 0.3 mm wide, divergent, arced toward the posterior- veel petals, stigmas 0.5—0.6 mm wide, capitate. Samara not seen Type. Martius s.n. Brazil. Minas Gerais: “Inter fruticeta ad pr aedium ¥, oe [Yha] Bes Minas Geraea in Goyasanae confinibus. . .” (holotype M! fra ment P-JUSS This oe is known only from the type, which was collected in north- western Minas Gerais near the Goias border (Fig. 6). Plants of P. hirta are densely golden pubescent. The hairs of the stem and vegetative branches and of the upper surface of the larger leaves have unusually long stalks, at least 0.5 mm long and commonly | mm or more long. Such hairs are also found in P. gardneri of the southern part of the Chapada das Mangabeiras of northeastern Goias. That species differs from P. Airta in its larger flowers in which the petal limbs are orbicular. Its staminode glands are all equally long. In P. hirta the limb of the posterior petal is obovate. The anterior and anterior-lateral staminode glands are longer than the posterior-lateral two. P. anadenanthera of central Goias also — 42 has long-stalked hairs, but is easily recognized by its anthers, which lack the glandular connective 15. Peixotoa hispidula Adr. Jussieu, Ann. Sci. Nat. Ser. II, Bot. 3: res ee we P. oe ead . subtomentosa Niedenzu, Verz. Vorles. Akad. ee -S 13: 1. 1912. Type. Gaudichaud 964 (lectotype G!, isotype P!). ae rene f 2. subglabrata Niedenzu, Verz. Vorles. Akad. Braunsberg W-S I9I2-1915 "44, 1912;. Type. tae 1169 (lectotype G!, isotypes BM! BR! F! GH! K! LE! M! MO! NY! P! S! W)). P. hispidula t. 3. pn Niedenzu, on Vorles. Akad. Braunsberg W-S 1912-1913: 41. 1912. Type. Salzmann 93 (holotype B, destroyed; isotype G!). Erect, scandent or prostrate shrub or subshrub or vine. Vegetative branches loosely velutinous or sparsely so or glabrous when young, older parts glabrous or glabrate. Stem stipules 3.4—13 mm long, 3-11 mm wide, cordate or sometimes triangular, entire with apex acute or notched, or rarely bifid, finely velutinous or tomentulose or glabrous adaxially, sericeous, velutinous or tomentulose, or velutinous-tomentulose abaxially or only on the margins, or glabrous abaxially, usually deciduous, sometimes persistent; inflorescence stipules like the cauline but usually more densely pubescent, eventually deciduous. Stem leaves with the petiole 2-16 mm long, loosely, sparsely or densely, velutinous, sometimes glabrous, lamina 2.3—12 cm long, 1.5-7.3 cm wide, elliptical or ovate, sometimes lanceolate, oblanceolate or orbicular, apex apiculate, base cordate or slightly so, rarely almost truncate, glabrous or glabrate or sparsely sericeous or velutinous or sparsely so above, glabrous or glabrate or loosely or sparsely pubescent with ‘T-shaped hairs or loosely or sparsely tomentose below, a pair of sessile glands at the base at the costa or halfway on the petiole or on the petiole up to 0.5 mm below the lamina or on the surface of the lamina up to 1.5 mm above the base, up to 0.5 (-2.3) mm from the costa, each gland 0.3-1.6 mm in diameter. In- florescence leaves with the petiole at least 2 mm long, laminas abruptly smaller than the cauline, the smallest not less than 3.9 mm long, 1.2 mm wide, elliptical or lanceolate or linear-lanceolate or linear, apex acuminate to caudate, finely velutinous or sparsely sericeous to glabrate above, sericeous or sparsely so below, a pair of glands at the base at the costa or halfway on the petiole or sometimes on the petiole just below the lamina, each gland (0.2—-) 0.4—1.3 (-1.8) mm in di- ameter, rarely with a second, smaller pair (ca 0.4 mm in diameter) just below the larger pair. Inflorescence terminal or axillary, the axes usually sparsely velu- tinous, sometimes sparsely tomentulose, internodes of the primary axis 0.5— 5.3.cm long, 2° axes 0.9-4 cm long, 3° axes 1.3-3.2 cm long, commonly the inflorescence with one terminal umbel only or with 2° axes only, subsidiary axes absent; umbels 4-flowered, one umbel per node, primary peduncles absent, secondary peduncles rare, up to 0.5 mm long. Bracts 0.4-1.5 mm long, 0.2— 0.5 mm wide, triangular or oblong or linear, glabrous, or sometimes velutinous or tomentose abaxially and/or on the margins; bracteoles 0.1-0.3 mm long, 0. 1- 0.2 mm wide, triangular or semicircular, glabrous, or bracteoles absent. Pedicels 17-39 mm long, finely or sparsely velutinous, usually a little longer in fruit. Sepals 4.2-6 mm long, 1.2—3 mm wide, adaxially glabrous and the extreme apex tomentulose or the proximal 1|/2—4/5 glabrous, the distal 1/2—1/5 tomentulose, lands 1.5-2.5 mm long, 0.6-1.2 mm wide. Claw of the lateral petals 2-3 mm long, limb 13-16 (—20) mm long, 11.5-15 (-18) mm wide, orbicular or broadly elliptical, base attenuate, fimbriae up to 0.5 mm long, glandular. Claw of the posterior petal 4.7—6.6 mm long, limb 6.5-8.2 mm long, 5-7 mm wide, elliptical, sometimes orbicular, fimbriae up to 0.6 mm long, capitate-glandular. Lateral stamen filaments 3.6—5 mm long, posterior filament 2.6-3.3 mm long, usually at 43 rectangular, erose or irregularly dentate, areole 3.5—5.2 mm lon , 3.2-4.2 mm 3.8 i length. Type. Martius 1169. Brazil. Rio de Janeiro: Between Cabo Frio and Espirito Santo [fide Niedenzu, 1912]. 1841 (Lectotype P!, isotypes BM! BR! F! G! GH! K! LE! M! MO! NY! S! W)). Distribution. (Fig. 6). Along the coast from Bahia to Rio de Janeiro, two collections from Sao Paulo; restingas, dunes, beaches. RAZIL. Bahia: Mun. Marau, 4 km S of Marau, Almeida 136 (CEPEC); Marau, Belém 3456 (CEPEC, MICH, NY); Salvador, Lagoa de Abaeté, Belém & Mendes 290 (IAN, NY, UB, US); Blanchet 1908 (G), 116 (W); Itapaba, at Lagoa do Abaeté, E of Salvador, 10-20 m, Davis 61059 (E); Porto Seguro, Duarte 5966 (MICH); 5 km SE of Marau at junction with new road N to Ponta do Muta, 14°08’S, 39°00’W, 0-50 m, Harley et al. 185 10 (MICH); Ilhéus, Riedel 617 (A, LE, S); Salzmann 93 (G); Wawra &: Maly 142 (W). Espirito Santo: Mun. Serra, entre Nova Almeida e Santa Cruz, Araujo 311 & Peixoto 181 (RB); Guarapari, Duarte 3655, 4173 (RB), Occhioni 7226 (MICH); inter Cabo Frio et Espirito Santo, Wied-Neuwied s.n. (BR). Rio de Janeiro: Rio de Janeiro, Gaudichaud 963 (F, P, P-JUSS, RB, syntype of P. hispidula), 964 (G, P), Lund 477c (C, GOET); restinga de Itapu, Araujo 690 & Peixoto 492 (MICH, RB); restinga de Piratininga, Brade 12014 (R); Sarangiros dans les bois des Priner, Glaziou 11819 (P); Lagoa de Piratininga, Kuhlmann s.n. (RB 26272): Santa Rosa, Netto 88 (R); restinga de Aramama, Occhioni s.n. (RB 26440); restinga da Marambaia, Occhioni 3554, 4313 (MICH); restinga de Itaipu, Schwacke 7130 (NY, R); entre Macaé e Rio das Ostras, BR-6, Km 161, Trinta 966 & Fromm 2042, Trinta 966-A & Fromm 2042-A (R); Cabo Frio, Alston-Lutz 56, 57 (R), Fromm et al. 1334 (R), Lindeman 6327 (MICH), Markgraf 3047 & Brade (RB), Mello Filho 1129 (R), Moreira & Angosto s.n. (R 72184), Netto et al. s.n. (R 19558), Restinga [-324 (GH, R, US), I-765 (US), /-1232 (US), [-1233 (NY), Riedel S Luschnath s.n. (LE), Segadas & Vianna 4210 (P), Smith 6570 (R, US), Sucre 3853, 4921 (MICH), Ule 4706 (R), Vidal s.n. (R 39291); Recreio dos Bandeirantes, Krapovickas et al. 23197 (F, G, MICH, MO, P), Lutz 653 (R), Markgraf 3788 & Brade (RB), Mello Filho 830, 980, 98 1 (R), Pereira 10525 (M), Rente 2, 238 (R), Rodrigues 92 (R), Santos 5086, 5087, 5088 (R), Smith 6367 (NY, R,S, US); Restinga de Jacarepagua, Arawjo 119 (MICH), 124 (RB), 140 (MICH), Atala 80, 90 (R), Berg 243 (U), Brade 11003, 11396 (R), Cuatrecasas SF Duarte 26636 p.p. (US), Maas & Carauta 3129 (MICH), Montalvo 17A (RB), Pereira 3569 (RB), Vianna 135 (NY), 537 (MICH); Mun. Maricd, 2km from “Amaral Peixoto,” Anderson 11650, 11653 (MICH), Lagoa de Marica, Occhioni 6389 (MICH), Vidal V-100 (R); Praia Grande, Taipu, Glaziou 1328 (BR, C, P), 3891 (C, F, K, P, R), 17815 (K, LE, US), Riedel & Luschnath 1304 (LE, S, US); Restinga da Barra da Tijuca, Anderson 11196 (MICH), Calvano 11 (R), Flaster 35 (R), Kuhlmann 6221 (RB), Machado 0H), Occhioni 650 (MICH), 1153 (RB), 3675 (MICH), Pereira 55 (R), 536 (MICH), 576 (RB), Pimentel 14 & Barbosa 15 (RB), Schott 3677 (W), Trinta 542 & Fromm 1618 (R, M), Trinta et al. 1386 (R), Vianna 15 (R). Sao Paulo: Sao Paulo, Frazdo s.n. (RB 11700); Mogu-guacu [Moji-Guacu], da Silva Manso s.n. (Lhotzky 257) BR). Co — za es = -~ — on ar = BS ~ is) 8. = S De be nr rn = ‘?) Aap, = — on a) hs ®& ~ iss} oa Q _ — Zz — — C Aas mn 44 One of the few well-collected species, P. hispidula is characterized by its large-flowered but small inflorescences. These are commonly composed of one terminal umbel and one or two pairs of secondary branches, each of which bears one terminal umbel only, or of a solitary umbel. 3° axes are rare. The flowers are among the largest in the genus. The limb of the lateral petals varies from ca 13 to 20 mm in diameter. The limb of the posterior petal is about half as long and wide as the limbs of the lateral petals. The staminode glands are strongly di- morphic. The anterior and anterior-lateral glands are wider than long, a condi- tion unique in the genus, and have an indentation across the apex and decurrent on the abaxial face. The posterior glands are smaller, as long as wide, and smooth. The inflorescence leaves are relatively large, mostly lanceolate or nar- rowly elliptical, and with a petiole commonly 4—6 mm or more long. Peixotoa hispidula is highly variable in habit and density of pubescence. In open areas it forms shrubs or subshrubs with erect, ascending or procumbent branches. Like many other species, it turns into a vine or climber if suitable support from adjacent shrubs and small trees is available. While the vegetative parts are usually glabrous or at most very sparsely pubescent, in some individuals the branches may be velutinous and the leaves velutinous above and sparsely tomentose below. This variation occurs throughout the range and does not merit infraspecific recognition. e pubescent P. axillaris, of south-central Goias, is the only other species with an inflorescence of one or three umbels. [t differs in its smaller flowers whose staminode glands are subequal and smooth. P. glabra, from central Minas Gerais, is sometimes confused with P. hispidula, but is easily distinguished. It has larger inflorescences, whose distal leaves are sessile or subsessile, and subequal staminode glands. Collected in flower and fruit throughout the year. 16. Peixotoa irwinii C. Anderson, sp. nov. Fig... Suffrutex; caules procumbentes vel ascendentes usque ad 2m. Stipulae vegetativae 7-8 mm longae, acutae vel bifidae, deciduae. Petioli 4.5—5.5 mm longi. Laminae 6.3—9.5 cm longae, 3.7—5.6 cm latae, ellipticae vel anguste ovatae, rugosae, supra sparsim aureo-velutinae, subtus in venis ferentes pilos ‘T-formes, basi juxta costam biglandulosae. Folia inflorescentiae petiolo brevi vel sessilia, linearia vel subulata. Inflorescentia terminalis, umbellis singularibus. Bracteae bracteolaeque praesentes. Pedicelli 17-26 mm longi. Sepala 3.8-4 mm longa. Limbus petalorum lateralium 8.2—9 mm longus, obovatus, margine glanduloso- fimbriato. Limbus petali postici 5.2—5.8 mm longus, obovatus, margine glandulis capitatis ornato. Glandulae 3 staminodiorum anticorum late sed non profunde indentatae in pagina abaxiali. Styli subaequales, 4.1-4.2 mm longi, anticus arcuatus versus petalum posticum, postici divergentes, parum inclinati versus petala postico-lateralia; stigmata capitata. Samara ignota. rostrate or ascending subshrub to ca 2m. Vegetative branches coarsely golden velutinous, older parts glabrous. Stem stipules 7-8 mm long, 8-9 mm wide, cordate, entire with the apex acute or notched, or bifid, finely tomentulose adaxially or pubescent only in the distal 4%, the proximal glabrous, coarsely golden velutinous-sericeous abaxially, deciduous; inflorescence stipules gradu- ally decreasing in size toward the distal nodes, the smallest not less than 3.6 mm long, ca 2.5 mm wide, the most distal sometimes split into two narrow, acute lobes, persistent. Stem leaves with the petiole 4.5—5.5 mm long, coarsely golden velutinous, laminas 6.3-9.5 cm long, 3.7-5.6 cm wide, elliptical or narrowly ovate, apex apiculate, base cordate, rugose, very sparsely golden velutinous 45 above, very sparsely pubescent with golden ‘T-shaped hairs on the major veins only below, a pair of sessile glands at the base at the costa, each gland I-1.5 mm in diameter. Inflorescence leaves of the proximal nodes petiolate, the more distal subsessile or sessile, laminas abruptly smaller than the cauline, the smallest not less than 3.2 mm long, 0.4 mm wide, linear or subulate, apex acuminate, finely velutinous above, sericeous or sericeous-velutinous below, a pair of glands at the base at the costa, each gland I-1.7 mm in diameter, sometimes the most distal leaves reduced to eglandular, subulate bracts. Inflorescence terminal, the axes coarsely golden velutinous, internodes of the primary axis 34.5 cm long, 2° axes 0.9-4.8 cm long, 3° axes 1.3-2 cm long, subsidiary axes absent; umbels 4- flowered, one umbel per node, primary and secondary peduncles absent. Bracts 0.5-1 mm long, 0.3-0.4 mm wide, linear or narrowly triangular, glabrous or sometimes sparsely velutinous on the margins; bracteoles 0.1—0.3 mm long and wide, triangular or oblong, glabrous. Pedicels 17-26 mm long, densely golden velutinous. Sepals 3.8-4 mm long, 2.2-2.6 mm wide, adaxially the proximal glabrous, the distal 24 tomentulose, glands 1.4-1.6 mm long, 0.9-1 mm wide. Claw of the lateral petals 1.8-2.3 mm long, limb 8.2-9 mm long, 7-8.7 mm wide, obovate, base briefly attenuate, fimbriae up to 0.3 mm long, glandular. Claw of the posterior petal 44.2 mm long, limb 5.2-5.8 mm long, 4.3-4.5 mm wide, obovate, fimbriae up to 0.3 mm long, capitate-glandular. Filaments of the lateral stamens 3.5-4 mm long, posterior filament 3.4-3.5 mm long, very slightly curved toward the posterior petal, anthers I-1.3 mm long, glandular connec- tives 0.6-0.7 mm long. Pollen grains 47-50 ym. Filament of the anterior stami- node 3.7-4 mm long, arced toward the posterior petal, anterior-lateral filaments 3.4-3.8 mm long, posterior-lateral filaments 3—3.2 mm long, curved toward the posterior styles, anterior gland 1.1-1.2mm long and wide, anterior-lateral glands I-I.1 mm long and wide, posterior-lateral glands (.9-1 mm long and wide, all glands with the apex smooth, anterior and anterior-lateral glands with a broad and shallow indentation on the abaxial face, posterior-lateral glands smooth. Styles 4.1-4.2 mm long, anterior style ca 0.4 mm wide, arced toward the posterior petal, posterior styles ca 0.3 mm wide, very slightly divergent, slightly inclined toward the posterior-lateral petals, stigmas ca 0.5 mm in diameter, capitate. Samara not seen. Type. Irwin et al. 282366. Brazil. Minas Gerais: Serra do Fspinhago, ca 3 km N of Sao Joao de Chapada, 1200 m, 24 Mar 1970 (holotype UB, isotype NY) Peixotoa irwinii is known only from the type (Fig. 6). It is one of the few sparsely pubescent species in the genus, and is known only from the Serra do Espinhaco of Minas Gerais. P. glabra and sparsely pubescent forms of P. reticulata also occur in the Serra do Espinhago, but are readily distinguished from P irwinti, The anterior and anterior-lateral staminode glands of P. irwinii are unique in that, though the apex of each gland is smooth, the abaxial face has a shallow depression. Its petioles are golden velutinous, the limbs of the lateral petals are obovate, and the styles are subequal. In P. glabra the anterior and anterior-lateral staminode glands have an indentation across the apex and de- current on the abaxial surface, the petioles are glabrous, the limbs of the lateral petals are orbicular, and the styles are unequal. P. reticulata has smooth stami- node glands and larger petals. The limbs of its lateral petals are ca 12- 15 mm long and wide. The limb of its posterior petal is ca 8-12 mm long and wide. The limbs of the lateral petals of P. irwinii are 8.2-9 mm long and 7- 8.7 mm wide. The limb of the posterior petal is 5.2-5.8 mm long, 4.3-4.5 mm wide. his species is named for Howard S. Irwin in recognition of his extensive collecting efforts in the Planalto of Brazil. 46 17. Peixotoa jussieuana Adr. Jussicu, Ann. Sci. Nat. Ser. II, Bot. 13: 279. 1840. P. jussieuana var. B velutina 2 ana Verz. Vorles. Akad. Braunsberg W-S 1912-1913: 36. 1912. Type. Gardner 2067 (lectotype G!, isotypes F! GH! K! P! W)). Ascending shrub or vine to ca 0.8 m. Vegetative branches loosely or densely velutinous, pubescence abraded from older parts. Stem stipules ca 6-8 mm long, ca 6-7 mm wide, cordate-triangular, apex acute, glabrous adaxially, loosely sericeous-tomentulose abaxially, deciduous; inflorescence stipules 6-—ca 9 mm long, 5.3-ca 9mm _ wide, cordate or cordate-triangular, finely or sparsely velutinous adaxially, sericeous-tomentulose abaxially, deciduous. Stem leaves with the petiole 95-9 mm long, loosely or densely velutinous or velutinous- tomentose, laminas 6-13 cm long, 3-7.5 cm wide, ovate or lanceolate, often the distal 43 attenuated toward the apiculate apex, base cordate or nearly truncate, sparsely or densely velutinous above, loosely, sparsely or densely pubescent with ‘T-shaped hairs, rarely tomentose, below, a pair of sessile glands at the base at the costa or halfway on the petiole or on the surface of the blade up to 4 mm above the base, up to 5 mm from the costa, each gland 0.5-1.5 mm in diameter, some- times one or both glands absent. Inflorescence leaves with the petiole 0.5 mm or more long, laminas abruptly smaller than the cauline, the smallest not less than 2.7 mm long, | mm wide, shape of the more proximal like the cauline, the more distal linear-lanceolate or linear, apex acuminate, velutinous or sparsely so or sparsely tomentulose or the most distal sometimes glabrous. adaxially, densely pubescent with T-shaped hairs abaxially, a pair of glands at the base at the costa or halfway on the petiole, each gland 0.3-1.5 mm in diameter. Inflorescence terminal or axillary, the axes velutinous or tomentulose- lag internodes of the primary axis 0.4—6.5 cm long, 2° axes 0.5-9 cm long, 3° axes 0.4-2.7 long, 4° axes 0.4—-1.6 cm long, aan axes absent; umbels Nee one umbel per node, iM aaa! and secondary peduncles absent. Bracts 0.2-0.7 (-3) mm ous 0.2-0.5 (-1.2) mm wide, triangular or semicircular, glabrous; bracteoles 0. 1—0.3 an 5) mm long, 0.1—0.2 mm wide, triangular, glabrous: some- times bracts and bracteoles absent. Pedicels 9-18 mm long, velutinous or veluti- nous-tomentulose, sometimes a little longer in fruit. Sepals 4.8—6 mm long, 2.5— 3 mm wide, adaxtally the proximal 4—'4 glabrous, the distal %4—'2 velutinous, glands 1.5-2.5 mm long, 0.7-1.3 mm wide. Claw of the lateral petals 2- 2.3 mm long, limb 10.5-12 mm long, ca 9-10 mm wide, obovate, base attenuate, fimbriae up to 0.9 mm long, glandular. Claw of the posterior petal 4.2-4.5 mm long, limb 7.5-8 mm long, ca 6 mm wide, obovate or elliptical, fimbriae up to 0.5 mm long, those of the proximal 74 capitate- glandular, the distal / fimbriate- glandular. Lateral stamen filaments 5.5-7 mm long, the distal 1/5-1/7 of the anterior-lateral two curved toward the posterior petal, posterior filament 4.4— 5 mm long, strongly arced toward the posterior petal, anthers 1.2—1.4 mm long, glandular connectives 0.8—l1 mm long. Pollen grains ca 50 wm. Anterior stami- node filament 5.5-6.2 mm long, always exceeding the anterior-lateral two, ante- rior-lateral filaments 4.5-5.8 mm long, always exceeding the posterior-lateral two, posterior-lateral filaments 4—5.3 mm long, inflexed between the posterior styles, glands I-1.4 mm long, 0.9-1.2 mm wide, the anterior gland sometimes a little larger than the other four, the posterior-lateral glands sometimes a little narrower than the anterior three, all with the apex and abaxial face smooth. Styles (5—) 5.2-6.5 mm long, 0.3-0.4 mm wide, often the anterior a little longer or sometimes a little shorter but always a little stouter than the posterior two, anterior style strongly arced toward the posterior petal, the stigma oblong, ca 0.4 mm long, borne laterally at the apex on the abaxial surface or very rarely capitate and the style sharply hooked at the apex, posterior styles divergent, the 47 distal 1/5—1/6 curved toward the posterior-lateral petals, all styles with scattered hairs in the proximal 4—' of their lengths, stigmas capitate, ca 0.4 mm in di- ameter. Carpophore up to 2.3 mm long. ‘Torus up to ca 4 mm high. Samara with the dorsal wing ca 2 cm long, ca 1.1 cm wide, upper margin arced, lower margin straight in the proximal 4, the distal 2% curved, erose, lateral wings 7-9 mm wide, 9-11 mm high, rectangular, extending up to ca 7 mm below the nut, erose, areole 3.5-4 mm long, 3.5-3.6 mm wide, nut ca 5mm long, ca 4mm in di- ameter, mature seed not seen. ype. Martius s.n. Brazil. Maranhao: “In sylvis sepibusque ad fluv. Itapicuru prope Caxias...” June [1819] (lectotype M! fragment P-JUSS!, isotype M!). Distribution. (Fig. 7). Ceara, Piaur, and Maranhao, one collection from Parana; caatingas, woodlands, and at forest margins. AZIL. Ceara: Allemao & Cysneiros 214 (R); Serra da [biapaba, Mello s.n. (RB 67422). oa Maranhao: Mun. Loreto, “Ilha de Balsas” region, ca 45 km S of Loreto, 7°2 45°0'W, 250-300 m, Eiten & Eiten 4835 (SP, US); Grajau, ui 2506 (MG). Parana: Mun. Parana, estrada Paranat- Panelas, Hatschbach 1413 (MICH, RB). t : Mun. Picos, ca 30 km E of Picos, 7°7'S, 41°9'W, Eiten & Eiten 4917 (US); between Boa Esperanga ae Santa Ana das Mercés, Gardner 2067 (BM, F, G, GH, K, >, W); Lagoa Comprida, oes 2498 (BM, K): Oeiras, Jobert 1045, 1053 (P, R); Chapada de Bom Jesus, Liitzelburg 275 (M), 1721 (RB); Sao Joao do Piautr, ara 1599 (RB); Oeiras, Martius s.n. (M, P-JUSS, syntype of P. ee Serra Branca, Ule 7 In P. jussieuana the anterior style is strongly arced toward the posterior petal and bears an oblong stigma at the apex but laterally on the abaxial surface. ‘The only other species with such an anterior stigma is P. magnifica from central and southern Goias, Mato Grosso and eastern Bolivia. It is probably a close relative. It differs in that its stipules are larger (12-23 mm long), and in that the limbs of its lateral petals are orbicular. ‘he samaras of P. magnifica have semicircular lateral wings 10.5-20 mm high. In P. jusseeuana the stem stipules are ca 6-8 mm long and the lateral petal limbs are obovate. The lateral wings of the samara are rectangular and 8-11 mm high. The leaves of P. jusstewana are at times at- tenuated in the distal one-third. A surprising disjunct is Hatschbach 1413 trom eastern Parana. This collec- tion 1s also atypical in that the tip of the anterior style is uncinate and bears a capitate stigma. Some intermediates between this peculiar style and the usual condition have been noted in some of the flowers. It is unlikely a the speci- mens labeled Glaziou 9680 were actually collected in Rio de Jane This spec ies was named by Martius in honor of Adrien de see, who published it. Collected in flower from January through August, in fruit in January and in May through July. 18. Peixotoa leptoclada Adr. Jussieu, Ann. Sci. Nat. Ser. I, Bot. 13: 279. 1840. Fig. 16. P. jussieuana var. a glabrescens Niedenzu, Verz. Vorles. Akad. Braunsberg W-S 1912-1913: 36. 1912. Type. Mendonga 976 (lectotype, NY!) Vine or scandent shrub. Vegetative branches tomentulose-velutinous, the older parts glabrous. Stem. stipules 10.5-20 mm long, 11-18 mm _ wide, tri- angular or broadly cordate, entire with the apex acute or notched, or bifid, finely velutinous or sericeous-velutinous or glabrate adaxially, tomentulose or seri- ceous-velutinous abaxially, persistent or deciduous; inflorescence stipules gradu- ally decreasing in size toward the distal nodes, the smallest not less than 7 mm long, 7.7 mm wide, or sometime equal to or larger than the cauline, entire with the apex acute or notched, or bifid, more densely pubescent than the cauline, persistent. Stem leaves with the petiole 5-20 mm long, densely golden 48 velutinous, laminas 6—17.5 cm long, 3-11 cm wide, elliptical or lanceolate or, especially the larger, ovate, apex apiculate, base cordate or almost truncate, golden velutinous above, loosely tomentose or pubescent with T-shaped hairs below, a pair of sessile glands at the base at the costa or on the surface of the lamina up to 4 mm above the base, up to 9mm from the costa, or sometimes halfway on the petiole, often with a second pair borne on the surface of the lamina, 0.6-17 mm above the base, up to 15 mm from the costa, each gland (0.6-) 0.9-1.8 mm in diameter. Inflorescence leaves with the petiole 0.6 mm or more long, very rarely the most distal sessile, laminas abruptly smaller than the cauline, the smallest not less than 1.3 mm long, 0.4 mm wide, lanceolate or oblanceolate or linear or subulate, apex acuminate and sometimes with a tiny gland, velutinous above, the most distal glabrous above, densely sericeous or velutinous below, a pair of glands at the base at the costa or halfway on the etiole, sometimes the glands obliquely stalked, each gland ca 0.4—1.2 mm in diameter, the larger laminas often with a second, smaller pair of glands (0.3— 0.5 mm in diameter) borne on the surface of the lamina or adjacent to or just above or just below the larger pair; sometimes the most distal leaves reduced to sessile, subulate, eglandular bracts, 2.7-3.5 mm long, 0.6-0.8 mm wide. In- florescence terminal or axillary, the axes velutinous, internodes of the primary axis 1-9.3 cm long, 2° axes 0.8-16.7 cm long, 2° subsidiary axes 2~2.3 cm long, 3° axes ().8—3.3 cm long, 4° axes 0.9-1 cm long; umbels 4-flowered, one umbel per node, primary and secondary peduncles absent. Bracts 0.2—0.7 mm long, 0.2- 0.3 mm wide, narrowly to broadly triangular or semicircular, glabrous, some- times velutinous or tomentulose abaxially and/or on the margins; bracteoles 0.1—0.3 mm long, 0. 1—-0.3 mm wide, triangular, glabrous, sometimes the margins velutinous. Pedicels 17-24 (-31) mm long, velutinous or velutinous-tomentulose. Sepals 4.8-6 (-7) mm long, 2.5-3.5 mm wide, adaxially glabrous or the apex tomentulose or the proximal 4—%4 glabrous, the distal 4-4 sparsely velutinous, glands 1.8-2.6 mm long, I-1.5 mm wide. Claw of the lateral petals 2.5— 6.2 mm long, limb 11-14 mm long, 10.8-13.7 mm wide, orbicular or broadly obovate, base briefly attenuate, fimbriae up to 0.8 mm long, glandular. Claw of the posterior petal 3.7-4.5 mm long, limb (7-) 7.5-10.5 mm long, (5.2-) 5.8- 9.2 mm wide, obovate or broadly so, fimbriae up to 0.6 mm long, capitate- glandular, glands of the apex a little smaller. Filaments of the lateral stamens 4.5-5.2 mm long, posterior filament 4.2-4.5 mm long, always shorter than the lateral four, arced toward the posterior petal, anthers 1.2-1.6 mm long, glandu- lar connectives 0.5-0.9 mm long. Pollen grains 42-50 wm. Filaments of the staminodes (3.2-) 4.14.8 mm long, sometimes the anterior a little longer than the lateral four, posterior-lateral filaments inflexed between the posterior styles, glands 1.2-1.5 mm long, the anterior usually a little longer than the other four, anterior and anterior-lateral glands 1.3—1.4 mm wide, posterior-lateral glands -I-1.2 mm wide, all glands with the apex and abaxial face smooth. Anterior style (4.3-) 4.7-5 mm long, 0.4—-0.5 mm wide, arced toward the posterior petal, posterior styles (4.5-) 5-5.3 mm long, 0.3-0.4 mm wide, always exceeding the anterior style, divergent, arced toward the posterior-lateral petals, all styles velutinous in the proximal 1/5 of their lengths, stigmas 0.4—0.5 mm in diameter, capitate. Mature samara not seen. Type. Claussen s.n. Brazil. Minas Gerais. “In sylvis post incendia redivivis (vulgo capueiras) . . .” (holotype P-JUSS!, isotype P!). istribution. (Fig. 7). Central Minas Gerais and south-central Goias; wood- lands and gallery forests. AZIL. Goias: Serra dos Pirineus, ca 15 km (str. line) N of Corumba de Goias, 1250-1300 m, Anderson 10222 (MICH, NY, UB); Fazenda do Palmital, Glaziou 20749 (BR, C, G, L, P, R); Luziania, Rio Vermelho a 15 km de cidade, Heringer 17258 (MICH); Allegrio, Pohl 2946 (1563d) (F, K NY, W). Minas Gerais: Claussen 522 (C, F, GH, P, 8), Claussen 643 (G), Claussen s.n. (F 1540004, 1540005), Claussen s.n. (G); Mendonca 976 (NY); Schwacke s.n. (R 19578). 49 Peixotoa ae is readily recognized by the large stipules, which are often persistent on the stem and always so at the more distal nodes of the inflores- cence. P. cig of Mato Grosso also has large, persistent inflorescence stipules but differs in its dense, white pubescence that gives the plants a silvery- grey appearance. Its leaves are usually densely tomentose below. The stem and leaf pubescence of P. leptoclada is moderate. In P. cordistipula the staminode filaments are unequal: the anterior exceeds the anterior-lateral two, and these exceed the posterior-lateral two. In P. leptoclada the staminode filaments are equal or the anterior may exceed the lateral four. The anterior style is longer than the posterior two in P. cordistipula but shorter in P. leptoclada. Another species with unusually large stipules is P. hatschbachii from central Goias. It is easily distinguished by its glabrous stems and leaves and the stalked leaf glands. Some forms of P. goiana have large stipules but these are deciduous in the inflorescence. Also, the inflorescence leaves of P. goiana are sessile. Those of P. leptoclada are petiolate and often bear the glands slightly and obliquely stalked. Peixotoa paludosa of central Minas Gerais and southern Bahia may be a close relative. Its inflorescence stipules, though smaller, are often persistent. It also has petiolate inflorescence leaves, similar staminodes, and an anterior style that is shorter than or subequal to the posterior two. It differs from P. leptoclada in that the leaf glands are borne on the surface of the lamina away from the base and costa. Such glands do occur sometimes in large leaves of P. leptoclada but usually in addition to a basal pair. The flowers of P. leptoclada are larger than those of P. paludosa. No mature fruits are at hand, but immature samaras show that the dorsal and lateral wings are large and perhaps similar to those of P. magnifica and P. jussieuana, Samaras of P. paludosa also have wide lateral wings. hile the majority of collections seen are presumed to be sexual, Anderson 10222, Glaziou 20749 and Heringer 17258, all from Goias, are apomicts. The pollen, contained in closed anthers, is 81-83% non-staining in cotton blue in lactophenol. These specimens are noticeably more robust than the sexual plants; especially the inflorescence axes appear sturdier and less graceful than in the specimens that Jussieu saw. 19. Peixotoa magnifica C. Anderson, sp. nov. Fig. 19. Frutex scandens vel liana usque ad 8m. Stipulae vegetativae (10—-)12— 23mm _ longae, cordatae vel triangulari-cordatae, acutae, deciduae, in- florescentiae interdum bifidae. Petioli (6.5-)8-28 mm longi. Laminae 7- 28.2 cm longae, 4.7—14 cm latae, ovatae vel ellipticae vel anguste ellipticae inter- dum suborbiculares interdum obovatae, basi cordatae interdum parum cordatae raro obliquo-cordatae, supra aureo-velutinae, subtus ferentes pilos T-formes vel tomentosae, basi juxta costam biglandulosae interdum 3—4-glandulosae. Folia inflorescentiae petiolo brevi, distaliora anguste lanceolata vel triangularia vel linearia, Cr a vel caudata. Inflorescentia terminalis vel axillaris, umbellis singularibus. Bracteae bracteolaeque praesentes interdum absentes. Pedicelli (8-) 17-27 mm fe Sepala 5-8 mm longa. Limbus petalorum lateralium 15- 17.8 mm pOneES orbicularis, margine glanduloso-fimbriato. Limbus_ petali postict 9.5—12 mm longus, obtrapezoideus vel orbicularis vel late obovatus, margine glandulis capitatis ornato. Filamenta staminum antico-lateralium in- clinata versus petalum posticum, filamentum staminis postici multum arcuatum versus petalum posticum. Styli subaequales vel saepius anticus quam postici longior vel raro brevior, anticus 7.3-9.2 mm longus, arcuatus versus petalum posticum, stigmate oblongo abaxiali, postici 7.5-9.2 mm longi, divergentes, arcuati versus petala postico-lateralia praesertim 4 distali, stigmatibus capitatis. Ala dorsalis samarae 2.9-4.2 cm longa, 1.7-2.4 cm lata, alae laterales 7-10 (-12) mm latae, 10.5-20 mm altae, extensae usque ad 8 mm subter nucem. 50 Climbing or scandent shrub or vine to 8 m. Vegetative branches densely, usually golden, velutinous. Stem stipules (10-) 12-23 mm long, 8.8-22 mm wide, cordate or triangular-cordate, apex acute, velutinous adaxially, most densely so at the apex and base, velutinous-tomentose or velutinous abaxially, deciduous; inflorescence stipules gradually decreasing in size toward the distal nodes, the smallest not less than 5 mm long, 5 mm wide, cordate, concave, entire with the apex acute or apiculate, sometimes bifid, adaxially the proximal 4-4 densely velutinous, the distal %4—%4 glabrous or sometimes very sparsely velutinous, abaxially densely velutinous-tomentose or sometimes only velutinous, deciduous or persistent. Stem leaves with the petiole (6.5-) 8-28 mm long, densely, usually golden, tomentose-velutinous, laminas 7—28.2 cm long, 4.7—14 cm wide, ovate or elliptical or narrowly so, sometimes suborbicular or obovate, apex apiculate, base cordate or slightly so, rarely oblique-cordate, golden velutinous above, usually loosely, sparsely, sometimes densely, pubescent with T-shaped hairs or tomen- tose below, a pair of sessile glands at the base at the costa or halfway on the petiole, rarely on the petiole just below the lamina, rarely borne on the surface of the lamina up to 0.9 mm above the base, up to 1.5 mm from the costa, each gland (0.7-) 1-2 mm in diameter. Inflorescence leaves with the petiole at least | mm long, laminas abruptly smaller than the cauline, the smallest not less than 2.3 mm long, 0.6 mm wide, the more proximal elliptical or narrowly so or oblanceolate, the more distal narrowly lanceolate or triangular or linear, apex acuminate or caudate, sparsely velutinous or glabrous above, densely velutinous below, a pair of glands halfway on the petiole or sometimes at the base at the costa, each gland 0.5—1.7 mm in diameter, sometimes with a second, smaller pair (0.4-0.7 mm in diameter) above or below the larger pair. Inflorescence terminal or axillary, the axes usually densely golden velutinous or velutinous-tomentu- lose, internodes of the primary axis 1.6—11 cm long, 2° axes (0.5) 1.5-17 cm long, 2° subsidiary axes 1.5-5.5 cm long, 3° axes 0.5—-5.2 cm long, 4° axes 1.4-1.8 cm long; umbels 4-flowered, one umbel per node, primary and secondary peduncles absent. Bracts 0.3-2.4 mm long, 0.1-0.6 mm wide, triangular or linear, glabrous, sometimes sparsely tomentulose abaxially and/or on the mar- gins; bracteoles 0.1-0.6 mm long, 0.1-0.3 mm wide, triangular, glabrous; some- times bracts and bracteoles absent. Pedicels (8-) 17-27 mm long, densely velutinous or velutinous-tomentose, usually golden, becoming up to 43 again as long in fruit. Sepals 5-8 mm long, 2.8-4.2 mm wide, adaxially the proximal 1/2-7/8 glabrous, the distal 1/2-1/7 velutinous or tomentulose, glands 2.1- 3.7 mm long, I—1.8 mm wide. Claw of the lateral petals (3—-) 3.5-5.3 mm long, limb 15-17.8 mm long, 15-17 mm_ wide, base briefly attenuate or rarely truncate, fimbriae up to | mm long, glandular. Claw of the posterior petal 5— 7.3 mm long, limb 9.5-12 mm long, 8-12 mm wide, orbicular or broadly obovate or obtrapezoidal, fimbriae up to 0.7 mm long, capitate-glandular. Filaments of the lateral stamens 6.5-8.4 mm long, the distal 4% of the anterior-lateral two curved toward the posterior petal, posterior filament 5-5.6 (-6) mm long, strongly arced toward the posterior petal, anthers 1.4-2 mm long, glandular connectives 0.6-1.1 mm long. Pollen grains 60-63 wm. Filament of the anterior staminode (6—) 6.5-7.8 mm long, anterior-lateral filaments (4.8-) 5.5-6.5 mm long, posterior-lateral filaments (4—) 5-6.3 mm long, strongly inflexed between the posterior styles, glands (1-) 1.2-1.6 mm long, 1.2-1.5 mm wide, the poste- rior-lateral glands usually a little smaller than the anterior three, all glands with the apex and the abaxial surface smooth, rarely bearing | or 2 fertile locules, 0.7—l1 mm long. Anterior style 7.3-9.2 mm long, ca 0.5 mm wide, strongly arced with especially the distal 4 tapered and curved toward the posterior petal, the stigma oblong, 0.5-0.6 mm long, borne laterally at the apex on the abaxial sur- face, posterior styles 7.5—-9.2 mm long, 0.4—0.5 mm wide, usually a little shorter than the anterior style, divergent, gently arced, especially the distal 4, toward ~ 51 the posterior-lateral petals, sugmas 0.4—0.5 mm in diameter, capitate, all styles usually with scattered hairs in the proximal 4 or less of their lengths. Ca arpo- phore up to 2.6 mm long. Torus up to 8 mm high. Samara the dorsal wing 2.9-4.2 cm long, 1.7—2.4 cm wide, upper margin gently arced, lower margin usually falcate, entire or gently erose, lateral wings 7-10 (-12) mm wide, 10.5- 20 mm high, semicircular, extending up to ca 8 mm below the nut, areole 4.5— 7.8mm long, 4.4-7 mm wide, nut 5-8 mm long, 5-7 mm in diameter, mature seeds not seen. Type. Anderson 9831. Brazil. Mato Grosso: Ca 12 km ENE of Barra do Gargas, ca 840 m, 6 May 1973 (holotype UB, isotypes MICH, NY). Distribution. (Fig. 7). Southern Goias, including the Distrito Federal, central and southwestern Mato Grosso, and eastern Bolivia; cerrado and woodlands. BOLIVIA. Dept. Santa Cruz: Prov. Nuflo de Chavez, Estancia Novicia, ca 30 km S de Con- cepcion, 62°12'W, 16°29'S, 500 m, Krapovickas & Schinint 32117 (CTES, MICH); Prov. Velasco, San Ignacio, 60°58'W, 16°22'S, 370 m, Krapovickas & Schinini 32251 (CTES, MICH)); Prov. Velasco 27kmS a San fuera io, camino a San Miguel, 61°W, 16°36/S, ca 400 m, Krapovickas & Schinini 32379 (CTES, CH); Prov. Chiquitos, 3 km NW de Robore, 59°46'W, 18°20'S, 300 m, Krapovickas & Schinini sae (MIC H): Prov. Velasco, Kuntze s.n. (NY). BRAZIL. Distrito Federal: entre Brasilia e Neca Pires et al. 9709 (UB, US). Goias: ca 10 km by road W of Ipora, 600 m, Anderson 9359 eae ms , UB); Serra do Caiapo, ca 5 ee (str. line) S of Caiaponia, 850 m, enan 9474 (MICH, F B): ee ra Dourada, ca 17 km (str. line) S of Goias Velho, 6 km NE of Mossamedes, 700-780 m, ean 10162 (MICH, NY, UB); ca 5 km W of Niquelandia, ca 750 m, Anderson et al. 35028 (MICH, \ : ve | en Peixotoa - #& jussieuana @ leptoclada @ magnifica O megalantha @ paludosa FIG. 7. Distribution of P. jussteuana, P. leptoclada, P. magnifica, P. megalantha, and P. paludosa. b2 NY); prope urbem Goyaz oe) Burchell 7133-2, 7084-2 (K); Serra Dourada, along affluent of the Rio Canna Brava, 28 km E of Formoso, 48°50'W, 13°45’S, Dawson 15005 (LAM); Chapada dos Veadeiros, Pouso Alto, 1800 m, Ferreira 215, 216, 217 (UB); Serra da Santa Brida Teen Gardner 3623 (K); Caiaponia, Eaahban 37700 (MICH); drainage of the upper Rio Araguaia, ca 70-87 km SE of Aragargas, 700 m, /rwin et al. 17494 (MICH, NY, UB), 17554, 176317 (MICH, UB); Barra Pillar, Villa Boa [Gotas], Pohl 1739, 3555 (1559d) p.p. (K, W); vicinity of Estrela do Norte, Belém-Brasilia, Prance & Sue 1a 58433 (MICH, NY); Natividade, Rizzo 9819 (MICH); Salinas, Weddell 2024 (P). Mato Grosso: Serra do Roncador, mun. Barra do Gargas, 250 km along new road NNE of Xavantina, ca 450 m, Eiten & Eiten 9014 (NY, US); Vale do Sonho, Xavantina, pg gee da pou Brasil Central, Fonseca 276 (NY, UB); Xavantina, Harley & Souza 11037 (K, MO, NY, P, UB); Xavantina- Cache road, 85 km fon Xavantina, Hunt 5700 (NY); neighborhood oh Navantina Hund 5868 NY); ee erra do Roncador, ca 60 km N of ee 550 m, Irwin et al. 15997 (MIC , UB); Navant 400 m, Jrwin et al. 16792 (MICH, , UB); C ' yaba ee Maine ae ‘Nioaque, Eppes 618 (NY): ca3 km Sof senna oe W, 14°44’S, Ratte 12 p.p. (K, UB); 1 km S of Xavantina, 52°20’W, 144 4'S, Ratter & Ramos 370 (K, NY, P, RB, ie Pantanal, Rio Paraguai, Fazenda Bela Vista, Schaller 89 (NY); Urucum, Corumba, 600 ft, Smith 74 (K). The usually large inflorescences of P. magnifica bear showy flowers most of which are open at about the same time. The unusual anterior style bears an oblong stigma at the apex but laterally on the abaxial surface. P. jussieuana from Ceara, Piauy and Maranhao is the only other species in the genus with such a stigma and is probably a close relative. It differs in that its lateral petals have obovate limbs while those of P. magnifica are orbicular. The samaras of P. magnifica have large, semicircular lateral wings that are recurved and give the fruit an inflated appearance. In P. jussieuana the lateral wings are rectangular and smaller. The cauline stipules of P. magnifica are 12-23 mm long while those of P. jussieuana are 6-8 mm long. Peixotoa magnifica has often been assigned to P. cordistipula of central Mato Grosso and eastern Bolivia. They are readily separated on the basis of their styles. Those of P. cordistipula all bear the usual capitate sugma and are less than 5.5 mm long. The styles of P. magnifica are 7-9.2 mm long. Most striking in P. cordistipula are the large inflorescence stipules which persist even after the pedicels have been shed. While P. cordisipula also produces large inflorescences, only a few flowers are open at any one time. Its dense white pubescence gives the plants a silvery-grey appearance that is not evident in P. magnifica. A structure commonly tound in P. cordistipula but absent in P. magnifica is a central, glandu- lar mass in the umbel. The staminode glands of P. magnifica rarely bear one or even two small locules that remain closed and contain largely aborted pollen. Collected in flower and fruit from April through December. 20. Peixotoa megalantha C. Anderson, sp. nov. Fig. 21 Frutex caulibus gracilibus usque ad 1.5m. Stipulae vegetativae 16-23 mm longae, acutae vel bifidae, persistentes, interdum deciduae. Petioli 5-9.5 mm longi. Laminae 10.5-20 cm longae, 7-17.2 cm latae, late ellipticae vel ovatae, rugosae, supra juniores velutinae maturae glabrae, subtus tomentosae, basi juxta costam biglandulosae. Folia inflorescentiae petiolo brevi distaliora sessilia, linearia vel subulata. Inflorescentia terminalis, umbellis singularibus. Bracteae bracteolaeque praesentes. Pedicelli 22-27 mm longi. Sepala 6.5-7.6 mm longa. Limbus petalorum lateralium 16-16.6 mm longus, orbicularis, margine glandu- loso-fimbriato. Limbus petali postici 9.5-10 mm longus, late obovatus, margine glandulis capitatis ornato. Filamentum staminis postici erectum. Stylus anticus ca -2mm longus, parum inclinatus versus petalum posticum, styli posticl 5.4— 5.5mm longi, divergentes, arcuati versus petala postico-lateralia; stigmata capitata. Samara ignota. Shrub with slender stems to 1.5 m. Vegetative branches tomentose, pubes- cence abraded from older parts. rae supules 16-23 mm long, 14-20 mm wide, S) 53 cordate, entire with the apex acute, or bifid, glabrate or tomentulose adaxially, tomentulose-velutinous abaxially, persistent, sometimes deciduous; inflores- cence stipules like the cauline, entire with the apex acute or notched, usually more densely pubescent, persistent. Stem leaves with the petioles 5—9.5 mm long, tomentulose-velutinous, laminas 10.5-10 cm long, 7-17.2 cm wide, broadly elliptical or ovate, apex apiculate, base cordate, rugose, the younger leaves velutinous above, the older leaves glabrous above, all tomentose below, a pair of sessile glands at the base at the costa or more commonly halfway on the petiole, each gland 1.6—3 mm in diameter. Inflorescence leaves with a small petiole at the more proximal nodes, the more distal sessile, laminas abruptly smaller than the cauline or gradually decreasing in size toward the distal nodes, the smallest not less than 4.5 mm long, 0.6 mm wide, shape of the more proximal like the cauline, the more distal linear or subulate, apex acuminate, finely velutinous above, tomentose-velutinous below, a pair of glands at the base at the costa, each gland ca 1-2 mm in diameter, sometimes with a second, smaller pair (ca 0.6 mm in diameter) above the larger pair. Inflorescence terminal, the axes tomentulose, internodes of the primary axis 3.8-9.8 cm long, 2° axes 3.6-6.3 cm long, mature 3° axes not seen, subsidiary axes absent; wines 4-flowered, one umbel per node, primary and secondary peduncles absent. Bracts 0.7—-0.8 mm long, ca 0.7 mm wide, triangular, glabrous, sometimes tomentulose on the margins; bracteoles 0.4—0.5 mm long, 0.3-0.4 mm wide, triangular, glabrous. Pedicels 22-27 mm long, tomentose. Sepals 6.5-7.6 mm _ long, 3.5-4mm_ wide, adaxially tomentulose-velutinous, glands 2.3-3.5 mm long, 1.5-1.7 mm wide. Claw of the lateral petals 2.5-2.8 mm long, limb 16-16.6 mm long, 15-16 mm wide, orbicu- lar, base attenuate, fimbriae up to 1 mm, glandular. Claw of the posterior petal ca 5mm long, limb 9.5-10 mm long, ca 9-9.5 mm wide, broadly obovate or suborbicular, fimbriae up to 0.9 mm long, capitate-glandular, glands of the dis- tal 4 much smaller. Filaments of the lateral stamens 5—5.3 mm long, posterior filament 4.5-4.7 mm long, erect, anthers 1.6-1.8 mm long, glandular connec- tives 0.8-0.9 mm long. Pollen grains 63-74 wm. Filaments of the anterior and anterior-lateral staminodes 4.8-5 mm long, posterior-lateral filaments 4.4— 4.5 mm long, inclined toward the posterior styles, anterior and anterior-lateral glands 1.4-1.5 mm long, ca 1.4 mm wide, posterior-lateral glands ca 1.2 mm long, ca | mm wide, all glands with the apex and abaxial face smooth. Anterior style ca 5.2 mm long, ca 0.4 mm wide, the distal Ys slightly inclined toward the posterior petal, posterior styles 5.4—5.5 mm long, ca 0.3 mm wide, divergent, the distal ¥3 arced toward the posterior-lateral petals, all styles with eoattered. hairs in the proximal ’—% of their lengths, stigmas ca 0.5 mm in diameter, ¢ capitate. eae not seen. e. Anderson et al. 36870. Brazil. Bahia: Espigao Mestre, ca 100 km WSW of ee 750-800 m, 8 Mar 1972 (holotype UB, isotype NY Peixotoa megalantha is known only from the type (Fig. 7). It is a coarse, low shrub which ts Foharactenized by being larger in nearly all its parts than other species. The persistent stem and inflorescence stipules are 16-23 mm long. The orbicular limbs of the lateral petals are ca 16 mm in diameter. Such large flowers are found in only three other species. P. hispidula, which grows ee a coast from Bahia to Sao Paulo, has petiolate inflorescence leaves; those of megalantha are sessile. P. magnifica of Goias, Mato Grosso and eastern Bolivia has styles 7-9.2 mm long; the anterior style bears an oblong stigma at the apex but laterally on the abaxial surface. The styles of P. megalantha are less than 6 mm long and all bear capitate stigmas. In P. reticulata, an apomict with usually closed Se s containing mostly aborted pollen, the more distal inflorescence stipules e less than 15 mm long, deciduous, and smaller than the stem stipules. P. Caine has not been reported from Bahia. 54 21. Peixotoa octoflora C. Anderson, sp. nov. fig. 21, Suffrutex usque ad 2m. Stipulae vegetativae deciduae, ignotae, inflores- centiae 7.5-8.2 mm longae, acutae, deciduae. Petioli 10-17 mm longi. Laminae 7.8-11.6cm longae, 6.6-10.4cm = latae, ovatae vel suborbiculares, supra velutinae, subtus ferentes pilos T-formes interdum laxe tomentosae, basi justa costam biglandulosae. Folia inflorescentiae petiolo brevi, distaliora linearia, supra glabra, subtus dense velutina. Inflorescentia terminalis, umbellis singulari- bus, 8-floribus; pedicelli 13-26 mm longi, 4 sessiles, 4 pedunculati, pedunculis 1-2.5 mm longis. Bracteae bracteolaeque praesentes. Sepala 7 7.2-7.8 mm longa. Limbus petalorum lateralium 14.5-15.2 mm longus, orbicularis, margine glanduloso-fimbriato, interdum eglanduloso. Limbus petali postici 10-12 mm longus, orbicularis, margine 4 proximalibus glandulis capitatis ornato, “4 distali glanduloso-fimbriato. Filamenta staminum subaequalia. Filamenta staminodi- orum anticorum subaequalia postica superantia. Styli subaequales, 4.8-5.3 mm longi, anticus erectus vel parum inclinatus versus petalum posticum, postici divergentes, arcuati versus petala postico-lateralia; stigmata capitata. Samara 1gnota. Subshrub to 2 m. Vegetative branches golden tomentulose-velutinous. Stem stipules deciduous, not seen; inflorescence stipules 7,5-8.2 mm long, 8.6—10 mm wide, cordate, apex acute, adaxially the proximal 4 densely velutinous, the distal % glabrous, abaxially densely velutinous, deciduous. Stem leaves with the petiole 10-17 mm long, densely golden velutinous, laminas 7.8—11.6 cm long, 6.6— 10.4 cm wide, ovate or suborbicular, apex apiculate, base cordate, velutinous above, pubescent with T-shaped hairs or sometimes loosely tomentose below, a pair of sessile glands halfway on the petiole, or sometimes at the base at the costa, each gland 1.2-1.4 mm in diameter. Inflorescence leaves with the petiole at least 2 mm long, laminas abruptly smaller than the cauline, the smallest not less than 2.8 mm long, ca | mm wide, the more proximal shaped like the cauline but more densely pubescent, the more distal linear, apex acuminate, glabrous above, densely velutinous below, a pair of glands at the base at the costa or halfway on the petiole, each gland I-1.7 mm in diameter. Inflorescence terminal, the axes densely velutinous, internodes of the primary axis 3.1-7.5 cm long, 2° axes 2.4— 14.7 cm long, 3° axes 2.1-5.5 cm long, subsidiary axes absent; umbels 8- flowered, of four pairs, in each pair one pedicel pedunculate, one pedicel sessile, primary peduncles 1-2.5 mm long, secondary peduncles absent. Bracts 0.6— 1.2 mm long, 0.8—-I1 mm wide, triangular, glabrous adaxially, tomentulose abax- ially; bracteoles 0.4—0.5 mm long, 0.3-0.5 mm wide, oblong, glabrous adaxially, glabrous or sparsely tomentulose abaxially. Pedicels 13-26 mm long, densely velutinous. Sepals 7.2—-7.8 mm long, 0.6-0.8 mm wide, adaxially the proximal /% glabrous, the distal tomentulose, glands 1.3-1.7 mm long, 0.9-1.2 mm wide. Claw of the lateral petals 3.2-3.5 mm long, limb 14.5-15.2 mm long, [4-15 mm wide, orbicular, base briefly attenuate, fimbriae up to | mm long, glandular or sometimes eglandular. Claw of the posterior petal 5.5-6 mm long, limb 10- 12 mm long, 10-11 mm wide, orbicular, fimbriae up to 1 mm long, those of the proximal %4 capitate-glandular, the distal 4 fimbriate-glandular. Lateral stamen filaments 5-6 mm long, posterior filament 4.8—5 mm long, usually a little shorter than the lateral four, arced toward the posterior petal, anthers 1.6-1.7 mm long, glandular connectives 0.7-0.8 mm long. Pollen grains 57-70 yum. Filament of the anterior staminode 5.5-6 mm long, anterior-lateral filaments 5—5.5 mm long, equal to or a little shorter than the anterior filament, posterior-lateral filaments 4.24.5 (-5) mm long, inflexed between the posterior styles, anterior and ante- rior-lateral glands 1.4—1.5 mm long, ca 1.3 mm wide, posterior-lateral glands ca 1.1 mm long, ca 0.9 mm wide, all glands with the apex and abaxial face smooth. Styles 4.8-5.3 mm long, 0.4-0.5 mm wide, anterior stvle usually a little stouter 55 than the posterior two, erect or slightly inclined toward the posterior petal, posterior styles divergent, arced toward the posterior-lateral petals, all styles glabrous or with a few scattered hairs near the base, sugmas 0.4—0.6 mm in diameter, capitate. Samara not seen. / e. Irwin etal. 16681a. Brazil. Mato Grosso: Serra do Roncador, ca 60 km N of Navantina, 550 m, 6 Jun 1966 (holotype UB, isotypes MICH, NY). Peixotoa octoflora is known only form the type (Fig. 8). It is the only species in the genus which bears 8-flowered umbels. Each umbel consists of four pairs of flowers. Each pair has the pedicel of one flower sessile, the other pedicel on a short (1-2.5 mm long) primary peduncle. The type is apparently an apomict. The closed anthers contain pollen that is mostly aborted and 95% non-staining in cotton blue in lactophenol. 22. Peixotoa paludosa Turczaninow, Bull. Soc. Nat. Moscow 31(1): 393. 1858. itera ba) P. parviflora var. B primaeva Niedenzu, Verz. Vorles. Akad. Braunsberg W-S 1912-1913: 34. 1912. Type. Glaziow 13598, pro parte holotype B, destroyed, isotypes, C! K!). — Vine over 5 m long. Vegetative branches velutinous or tomentulose, pubes- cence abraded from older parts. Stem stipules ca 10 mm long, ca 9 mm wide, triangular, apex acute, finely velutinous adaxially, velutinous or tomentulose- velutinous abaxially, deciduous; inflorescence stipules gradually decreasing in size toward the distal nodes, the smallest not less than 2.5 mm long, 3 mm wide, densely velutinous in the proximal 4 and sparsely so or glabrate in the distal % adaxially, tomentulose-velutinous abaxially, very densely white tomentulose along the margins, persistent or eventually deciduous. Stem leaves with the petiole (4—) 5-17 mm long, densely velutinous or densely tomentose, laminas 4.8—-14.3 cm long, 3-9.2 cm wide, elliptical or broadly elliptical or obovate, apex apiculate, base slightly cordate or almost truncate, velutinous or densely so above, loosely, sometimes sparsely, pubescent with T-shaped hairs or tomentose below, a pair of sessile glands borne on the surface of the blade 3-29 mm above the base, 1.5-—17 mm from the costa, each gland 0.4—1.2 mm in diameter, or one or both glands absent. Inflorescence leaves with the petiole 1.2 mm or more long, laminas abruptly smaller than the cauline or gradually decreasing in size toward the distal nodes, the smallest not less than 1.8 mm long, 0.5 mm wide, shape of the more proximal like the cauline, the more distal narrowly lanceolate or linear, apex acuminate and sometimes with a tiny gland, sparsely velutinous or glabrous adaxially, densely velutinous or sericeous-velutinous abaxially, a pair of glands near or at the base or at the costa or sometimes halfway on the petiole, each gland 0.3—0.8 mm in diameter, rarely the larger laminas with a second pair near the base; rarely the most distal leaves reduced to sessile, linear or subulate, eglandular bracts, 0.7-+ mm long, 0.3-1.6 m wide. Inflorescence terminal or axillary, the axes velutinous or velutinous-tomentulose, internodes of the pri- I mary axes 2.3-10 cm long, 2° axes 0.7-12.5 cm long, 2° subsidiary axes 0.9-8 cm long, 3° axes 0.5-6.2 cm long, 3° subsidiary axes 0.5-20 mm long, 4° axes ().4— 12mm long; umbels 4-flowered, one umbel per node, primary peduncles absent, secondary peduncles rare, up to 0.1 mm long. Bracts 0.2-0.4 (-1.2) mm long, 0.3-0.9 (-1.2) mm wide, triangular or semicircular, glabrous, sometimes tomentulose on the margins; bracteoles 0.1 mm long and wide, tri- angular, glabrous; sometimes bracts and bracteoles absent. Pedicels 11-19 mm long, velutinous or velutinous-tomentulose, often a little longer in fruit. Sepals 3.2-4.8 mm long, |.8-2.5 mm wide, adaxially finely velutinous, or the proximal J. % glabrous, the distal /3 velutinous, glands 1.4-2.4 mm long, 0.7—1.3 mm wide. 56 Claw of the lateral petals 2-2.7 mm long, limb 9-11.5 mm long, 9.3—10 mm wide, broadly obovate or broadly elliptical or suborbicular, base briefly attenu- ate, fimbriae up to 0.9 mm long, usually glandular, sometimes a few eglandular. Claw of the posterior petal 3.3—4 mm long, limb 5.3—7 mm long, 5—6.3 mm wide, obovate, fimbriae up to | mm long, capitate-glandular, or sometimes the proxi- mal 2 capitate-glandular, the distal 2 fimbriate-glandular. Stamen filaments 3.6—4.5 mm long, posterior filament equal to or shorter than the lateral four, strongly arced toward the posterior petal, anthers |.1-1.4 mm long, glandular connectives 0.6-0.7 mm long. Pollen grains ca 50 wm. Anterior staminode fila- ment 3.7-4.3 mm long, always exceeding the lateral four, anterior-lateral and posterior-lateral filaments 3.5—4.1 mm long, posterior-lateral filaments strongly inflexed between the posterior styles, anterior gland 1.4-1.6 mm long, 1.3- 1.5 mm wide, anterior-lateral and posterior-lateral glands 1.1-1.4 mm_ long, anterior-lateral glands 1.1-l.4 mm wide, posterior-lateral glands 0.8-1.3 mm wide, usually narrower than the anterior-lateral two, all glands with the apex and abaxial face smooth. Styles 4—5.5 mm long, 0.3-0.4 mm wide, the anterior style sometimes a little longer and usually a little stouter than the posterior two, strongly arced toward the posterior petal, posterior styles strongly divergent, inclined toward the posterior-lateral petals, stigmas 0.3-0.5 mm in diameter, capitate. Carpophore up to 3mm long. Vorus up to ca 3.5 mm high. Samara with the dorsal wing 2.8—3.1 cm long, 1.5-1.7 cm wide, upper margin arced or straight, lower margin arced or slightly so, erose, lateral wings 6-8.5 mm wide, 12-I4mm_ high, rectangular or semicircular, entire, erose, or irregularly dentate, areole 3.2-3.5 mm long, 2.7-3.5 mm wide, nut ca 4.5 mm long, 3.5— +mm in diameter, mature seeds not seen. Type. Blanchet 2867. Brazil. Bahia: “...in paludibus d’Ihabira etal prope urbem di Barro.” (holotype CW?, isotypes BM! BR! F! G! K! LE! NY W)). Distribution. (Fig. 7). North-central Minas Gerais, especially the Serro do Espinhaco, and Bahia; cerrado, campo sujo, caatinga, and at forest margins. BRAZIL. Bahia: type q.v.; ca 35 km N . ee Jesus da Lapa, on main road to ge ages ca 450 m, 43°19'W, 12°58'S, Harley et al. 21561 CH); o hapadao Ocidental da Bahia, 37 km SE of Covrentng: on road to Jaborandr, ca 600 m, ca 28 35'S, Harley et al. 21964 (MICH). ee Gerais: 22 km by road W of Januaria on road to Serra ‘das Araras, 610 m, Anderson 9197 (MICH, NY); Belo ue Barreto 7545 (UB), 8681 (R ‘ Claussen 68A (BR, F, W); mun. Santa Luzia, Lagoa Santa, da Costa s.n. (R 72044); entre Diamantina, Mendanha e Morro ae Ramao, Glaziou 13598 p.p. Ki type of PF; a var. ae hely entre Taquarugu e Serrinha, Glaziou 18929 p.p. (C, K, LE, R); mun. Caimopolis, rod. Fernao Dias, Hatschbach 41427 (MICH); Serra do ‘Vaquaril, Nova Lima, Roth a (RB); Warming 238 (O Contagem, Rio Paraopeba, Warming s.n. (C); Warming s.n. GOET US); mun. Belo Horizonte, Morro das Pedras, Williams J Assis 6490 (GH); mun. Bile ian ehe Colonia Afonsa Pena, Williams © Assis 7231 (GH). ~ = — In P. paludosa the leaf glands are borne on the surface of the lamina away from the base and costa. Within the range of P. paludosa only P. spinensis also bears the glands in this manner. It differs in that its leaves are densely tomentose to woolly below, and in that the anterior and anterior-lateral eae glands have an indentation across the apex and decurrent on the abaxial face. The leaves of P. paludosa are more moderately pubescent, and the staminode glands are smooth. Also, the flowers of P. paludosa are slightly smaller than those of P. spinensis. In P. leptoclada, of Minas Gerais and Goias, the larger leaves sometimes bear the glands on the surface away from the base and costa. This species, which may be a close relative, has similar but larger flowers, but differs most strikingly in its large stipules. Those of the stem are ca 11-20 mm long and wide and are usually persistent. Those of the inflorescence are at least 7 mm long and are persistent at the more distal nodes. In P. paludosa the stem stipules are ca 9- 10 mm long and wide, and deciduous. ‘hose of the inflorescence are usually less ay than 7 mm long and wide (only rarely more), and, while persistent for a while, are eventually deciduous. Roth 1587 is unusual in that the specimens are apparently apomictic. The anthers are closed and contain aborted pollen that is 99% non-staining in cotton blue in lactophenol. e type locality is unknown. According to Urban (1895), during the years 1833-1847 Blanchet sent an assistant on collecting trips along the southern coast of Bahia and as far inland as the Rio Sao Francisco. Many of the exact localities for the collections made during these expeditions have not been ascertained. Only two additional collections of P. paludosa are known from Bahia. These were made by Harley in 1980. The specific epithet is doubtlessly a misnomer. P. paludosa has never been collected in wet areas. More likely the type was found in a dry area near the “marais d’Itabira.” Niedenzu (1912) described P. parviflora var. primaeva based on Glaziou 13598. He considered this variety an ancestral type, because in his material the staminodes had tiny locules (*Androecium fere Banisteriae sectionis Pachyzeug- matis. ..”). However, Glaziou 13598 is a mixed collection of P. paludosa and of Banisteriopsis malifolia (Nees & Martius) Gates var. malifolia. The eae a the holotype (at B, now destroyed) shows a specimen of P. paludosa in bud. isotypes at C and K are also in bud, but flowers of B. malifolia are contained in i. packets mounted with the specimens. Niedenzu doubtlessly based his description on such flowers supplied with the Berlin specimen Collected in flower from January through September, in fruit from April through August. Figs. 2 ye P. discolor Grisebach, Linnaea 13: 216. 1839. Type. Sellow s.n. [Sellow ie c1313 (28 May 1819), fide Niedenzu, 1912] (holotype, B en isotype K!). 23. Peixotoa parviflora Adr. Jussieu in St. Hilaire, Fl. Bras. Mer. 3: 62. 1832 [1833]. Scandent shrub or vine to 3m. Vegetative branches finely or loosely, often golden, velutinous, pubescence abraded from older parts. Stem stipules 5.5— 17 mm one 5-17 mm wide, cordate, apex acute, velutinous or tomentulose adaxially, velutinous-sericeous or velutinous-tomentulose abaxially, deciduous; inflorescence stipules gradually decreasing in size toward the distal nodes, the smallest not less than 3mm long, 4 mm wide, entire with the apex acute or notched, or rarely bifid, finely velutinous or tomentulose adaxially, sericeous- eee or tomentulose-velutinous abaxially, deciduous or persistent. Stem leaves with the es 4-32 mm long, (often eo) velutinous or velutinous- tomentulose, lamina 5.5-17.5 cm long, 3-10.6 cm wide, ovate or elliptical, some- times obovate or cee apex apiculate, ee cordate or slightly so or rarely truncate, velutinous to densely so above, loosely pubescent with T-shaped hairs or loosely tomentose below, with a pair of sessile glands at the base at the costa or halfway on the petiole or rarely on the petiole just below the lamina or rarely on the lamina up to | mm above the base at the costa, each gland 0.7-2.3 (-4.3) mm in diameter, rarely one or both glands absent. Inflorescence leaves with the petiole (0.6-) 1mm or more long, laminas usually abruptly smaller than the cauline, sometimes gradually decreasing in size toward the distal nodes, the smallest not less than 3 mm long, 0.6 mm wide, shape of the more proximal like the cauline, the more distal narrowly elliptical or lanceolate or oblanceolate or linear, apex acuminate or sometimes caudate, velutinous or sparsely so above, sericeous or sericeous-velutinous below, a pair of glands at the base at the costa or half-way on the petiole or rarely on the petiole just below the lamina, each gland (0.2—) 0.3-1.5 mm in diameter, rarely with a second, smaller pair (ca 58 0.4 mm in diameter) borne just above the pair of larger glands; rarely some of the inflorescence leaves reduced to sessile, linear, eglandular bracts, 2.5-3.5 mm long, ca 0.5 mm wide. Inflorescence terminal, the axes velutinous or velutinous- tomentulose, internodes of the primary axis 1-8.3 cm long, 2° axes 0.9-12.5 cm long, 2° subsidiary axes 0.8—4.2 cm long, 3° axes 1-10.8 cm long, 3° subsidiary axes 0.7—2.2 cm long, mature 4° axes not seen, usually at the more distal nodes the 2° and 3° axes suppressed, with | or 2 umbels borne on secondary peduncles in the leaf axil, very rarely subsessile, always with 3 umbels at the terminal nodes, each umbel 4-flowered, primary peduncles absent, secondary peduncles 0.5- 17 mm long. Bracts 0.3-2.2 (-3) mm long, 0.3-1 mm wide, triangular or linear or rarely semicircular, glabrous or the margin velutinous or tomentulose; bracteoles 0.1-0.7 mm long, 0.1-0.5 mm wide, triangular or linear or oblong glabrous or the margin velutinous or tomentulose. Pedicels 7-19 mm long, velutinous or sometimes velutinous-tomentulose, often a little longer in fruit. Sepals 2.2-4.7 mm long, 1.2-2 mm wide, adaxially glabrous or with the apex tomentulose or the oe ¥3—Yo glabrous, the distal 24-4 tomentulose, glands 0.7—2 mm long, 0.4—1.1 mm wide. Claw of the lateral petals 1.5-2.2 mm long, limb (4.3-) 5.2-9.5 mm ane (4—) 5-8. 5 mm wide, orbicular or broadly obovate, base briefly attenuate, fimbriae up to 0.5 mm one usually glandular, Tee a few eglandular. Claw of the posterior ae @2—) 2, 7-4 mm long, limb 3.5— (—6.8) mm long, 3-6.2 mm wide, obovate or broadly so or suborbicular, bees up to 0.5mm _ long, capitate-glandular or sometimes the apex fimbriate- glandular. Lateral stamen filaments 2.2-4 mm long, always exceeding the poste- rior filament, posterior filament (1.9-) 2.1-3 mm long, arced toward the poste- rior petal, anthers 0.8-1 mm long, glandular connectives 0.6-0.7 mm long. Pol- len grain ca 42 um. Anterior staminode filament 2.1—3.3 (-4) mm long, always exceeding the lateral 4, lateral staminode filaments 1.8—3.2 (3.8) mm long, the anterior-lateral filaments usually longer, sometimes shorter, than the posterior- lateral two, or sometimes the four lateral filaments subequal, posterior-lateral filaments inflexed or only slightly so between the posterior styles, anterior and anterior-lateral glands (1-) 1.1-1.5 mm long, 1.3-1.4 mm wide, posterior- oe glands 0.8-1.2 mm long, 0.9-1 mm wide, usually a little shorter but always - rower than the anterior three, all with the apex x and abaxial face smooth. aie 2.3-3.7 mm long, 0.3-0.4 mm wide, the anterior style often a little longer, some- times a little shorter, than the posterior two, anterior style arced toward the posterior petal, posterior styles divergent, arced (not as strongly as the anterior style) toward the anterior-lateral petals, all styles usually with scattered hairs in the proximal '2 of their lengths, stigmas 0.3-0.4 mm in diameter, capitate. Carpophore up to 0.5 mm long. Torus up to 3.5 mm high. Samara with dorsal wing 1.6-1.9 cm long, 0.8—0.9 cm wide, upper margin straight or gently sigmoid, ee margin arced or slightly sigmoid, entire or slightly erose, lateral wings 2.5-3.6 mm wide, 6.7-8.8 mm high, rectangular or semicircular, sognuy erose to sometimes dentate, areole 3-3.3 mm long, 3-3.5 mm wide, nut 4.5-5 mm long, 2.8—3.5 mm in diameter, seed ca 6 mm long, outer cotyledon ca 6.5 mm long, ca 2.5mm wide, folded at ca % of its length, inner cotyledon ca 4.2 mm long, ca 2.2mm wide, folded at ca % of its length. ype. St. Hilaire sn. Brazil. Minas Gerais: [Conceicio} . ” (holotype P!, isotype P!). Dice puGon: (Fig. 8). poi central and southern Minas Gerais and Rio de Janeiro to eastern Sao Paulo and eastern Parana; cerrado, campo, woodlands, and gallery forests. ee . in pascuis juxta Conceao RAZIL. Minas Gerais: Ackermann s.n. (BR); Morro do ae Black & Magathaes 51-12228 (UB); Serra do Taquaral, Ouro Preto, de Brito, Jr. s.n. (SP 18835); 4 km W of Morro do se 20 km from road between Conceigao and Be lo ee ene a (MIC H); Serra de Biribiri, 7 km ve “ Diamantina, 1200 m, Glassman & Gomes, jr. 8tl Vaquarugu a Serrinha, Glaziou 13929 p.( — 7 — if ~ Peixotoa > & octoflora I _\ | @ parviflora “| f \ A | a i. cee 1 | ee ae ee ae, pion > Fes FIG. 8. Distribution of P. octoflora and P. parviflora. Diamantina a Tombador, Glaziouw 18930 (C, P, R); Presidente Soares, Heringer 10295 (UB); Santa Barbara, Hoehne s.n. (SP 5052); Serro Frio, Martius s.n. (M); Serra dos Pinheiros, Pohl 3556 (d. 1557) (K, W); Caldas, Regnell-IIT 118 (US), 11-318 (F, P, S, US); [1-3 18x (S), [11-1652 (F, 8, US, W); Santa Luzia, Riedel & Luschnath I (LE, S); Serra do Taquaral, ae Preto, Silva Novaes s.n. (SP 18839): Ouro Preto, Texeira s.n (SP 18826); Galdae! Widgren in 1845 (BR, F, K, LE, S, US); Williams & Assis 6619 (GH). Parana: 2 km W of moss of PR-11 and the eee prea at ihe border of Parana and Sao Paulo, 620 m, Davidse et al. 11371 (MO); Sengés, Dusén Gan H, S); Jaguariahyva Jaguariaiva], 740 m, Dusén 340a (GH, MO, a 9225 (A, BM, F, G, GH, K, MO, NY, S, US), 9619 (S), 9690 (F, S), 11685 (A, BM, G, GH, K, MO, NY, S, US), 13032 (S), ine (S); mun. Tibagi, Balsa para Pirai, 696 m, Hatschbach 6028 (US), mun. Tibagi, Fazenda Monte Alegre, Harmonia, Pee 283 1 MICH, RB); mun. Sengés, Fazenda Morungava, Rio do Funil, Hatschbach 5347 (MICH, US); Jaguariaiva, Hoehne s.n. (SP 23414); 4km N of Wenceslau Braz, Lindeman & Haas 3131 (K | MBM. NY); Tibagi, Reiss 65a (F); mun. Sengés, Fazenda Morungava, Rio Funil, ca 24°08'S, 49°29" WwW, 800 m, Smith et al. 14834 (US). Rio de Janeiro: Rio de Janeiro, Ackermann 232 (P-JUSS), ee s.n. (BR), Burchell A882 (K); Peirdpolis a Santo Antonio, cae 5785 (C, P), 8581 (C, P), 10371 (B C, G, K, LE, P, US); pr. Sumidero, na rodovia BR-3 entr Itaipava e Pedro do Rio, ca 600 m, ae 10321 (RB); Serra dos Orgaos, Riedel IIT (LE, S); SelbuT: it. [a 1814] (K?, NY); Petropolis, mata do Judio, ca 700 m, Sucre ee (MICH); mun. Araras, 850-980 m, Sucre 10614 (MICH). Sao Paulo: SP); Campinas, eee Novaes 651 (SP eae Cotia, Constantino s.n. (RB 45430); Jaragua, Poche. s.n. (SP 5479); estrada de Pirapora a C lara a, Hoehne s.n. (F, GH, NY, SP 12905); Sao Paulo, 800 m, Holway & Holway 1496 (US); Amparo, Kuhn 1192 (SP); [garatd, Kuhlmann 2758 (SP); ca ae km SW de Jundar. Serra do Japr, Leitao Filho et al. 3164 (NY, US); Lojgren 2 C); mun. Itapeva, Itangua, ca 1000 m, Vidal s.n. (R 72157, R 72400); Sorocabat Wacket s.n. (W); circa Parnahyba ee ad fl. Tieté, ca 700 m, Wettsten & Schiffner s.n. (W). — ey ist) oq e oy & ot a aq iss) cay ‘oS w —_~ ad = — Peixotoa parviflora is readily recognized by its inflorescence structure. The terminal nodes bear three umbels and thus are 12-flowered. The lateral umbels and often also the terminal umbels are borne on secondary peduncles. Com- monly at the more distal nodes the secondary and tertiary axes are suppressed and one or two umbels are borne on secondary peduncles in each leaf axil. The inflorescence leaves are always petiolate. The flowers are unusual in that the posterior styles are curved toward the anterior-lateral petals instead of toward the posterior-lateral petals. Only three other species share the inflorescence 60 structure but not the curvature of the styles. P. adenopoda from east-central Bahia has stalked leaf glands and staminode glands that have an indentation across the apex and decurrent on the abaxial face. P. catarinensis from coastal Santa Catarina has tiny leaf glands (0.2—0.7 mm in diameter) which are borne on the surface of the lamina away from the base and costa. In P. parviflora the stami- node glands are smooth. The leaf glands are sessile and borne at the base at the costa or halfway on the petiole or rarely entirely on the petiole just below the lamina. They are 0.7—2.3 (-4.3) mm in diameter. P. tomentosa is probably closely related and the species most likely to be confused with P. parviflora. Its range, the central region of the Serra do Espinhago of Minas Gerais, falls within that of P. parviflora. Its flowers are larger than those of P. parviflora. Sessile inflorescence leaves and sessile umbels at the more distal nodes are common. The pubescence of the inflorescence axes and of the lower leaf surface also usually help to separate the two species. The young branches and inflorescence axes of P. tomentosa are very densely golden velutinous. In P. parviflora the hairs are com- monly white and only sparsely to moderately abundant. The leaves of P. tomen- tosa are usually densely tomentose or woolly below while those of P. parviflora are sparsely to moderately pubescent with T-shaped hairs or moderately tomentose but never woolly below One of the few species that have a relatively wide range, P. parviflora shows some geographical variation in flower size. Specimens from Rio de Janeiro tend to have the smallest flowers, those from Parana the largest. Collected in flower throughout the year, in fruit in January, June, July, and August. 24. Peixotoa psilophylla C. Anderson, sp. nov. Fig. 2 Suffrutex usque ad 2 m. Stipulae vegetativae 8.6—15 mm longae, acutae vel bifidae, interdum persistentes. Petioli 3.3-6.5(-8.8) mm longi. Laminae (3.7-) 6.3-20.7 cm longae, (3.6—-)5-9.5 cm latae, rugosae, ellipticae vel ovatae vel lanceolatae, supra glabrae vel sparsim velutinae, subtus glabrae vel in venis sparsim velutinae, basi juxta costam biglandulosae. Folia inflorescentiae petiolo brevi vel sessilia, distaliora lineari-lanceolata vel subulata. Inflorescentia termi- nalis vel axillaris, umbellis singularibus. Bracteae bracteolaeque praesentes. Pedicelli 16-34 mm longi. Sepala 4.5-6.5(-7) mm longa. Limbus petalorum ateralium 11.5-15 mm longus, orbicularis, margine denticulato, glanduloso vel eglanduloso. Limbus petali postici 8.5—9 mm longus, late obovatus, margine 74— ¥%4 proximalibus glandulis capitatis ornato, 43-4 distali denticulato, glanduloso vel eglanduloso. Filamenta staminum subaequalis. Glandulae 3 staminodiorum anticorum apice non profunde indentatae, plerumque indentatione in pagina abaxiali decurrenti. Stylus anticus 3.2-4 mm longus, saltem parum inclinatus versus petalum posticum, styli postici 3.44.3 mm longi, interdum inaequales, divergentes, arcuati versus petala postico-lateralia; stigmata capitata. Samara ignota Gubshnus to 2m. Vegetative branches purplish-brown, sparsely tomentu- mse when young, older parts glabrous. Stem stipules 8.6-12 mm long, 9- 5mm wide, cordate or triangular-cordate, entire with the apex acute or Sore or bifid, finely tomentulose or velutinous adaxially, velutinous- tomentulose abaxially, deciduous, sometimes persistent; inflorescence stipules gradually decreasing in size toward the distal nodes, the smallest not less than 6.3 mm long, 6.2 mm wide, entire with the apex acute or sometimes notched, more densely pubescent than the cauline, eventually deciduous. Stem leaves with the petiole 3.3—6.5 (-8.8) mm long, tomentulose-velutinous or sparsely so, some- times glabrous, laminas (3.7—) 6.3-20.7 cm long, (3.6—) 5-9.5 cm wide, elliptical or ovate or lanceolate, apex apiculate, base cor date: rugose, glabrous or sparsely — 61 velutinous above, especially along the veins, glabrous or with scattered hairs on the major veins below, especially near the base, a pair of sessile glands at or up to 0.7 mm above the base at the costa or halfway on the petiole, each gland I- 2.1 mm in diameter. Inflorescence leaves petiolate or sometimes sessile at the distal nodes, laminas abruptly smaller than the cauline, linear-lanceolate to linear to subulate, the smallest not less than 3 mm long, 0.6 mm wide, apex acuminate, tomentulose-velutinous above, densely velutinous below, a pair of glands at the base at the costa or halfway on the petiole, each gland 0.8—1.6 mm in diameter, sometimes with a second, smaller pair (0.5-0.6 mm in diameter) below the larger pair, rarely one or both glands absent. Inflorescence terminal or axillary, the axes velutinous-tomentulose, internodes of the primary axis 2.7— 15 cm long, 2° axes 2.8-11.1 cm long, 2° subsidiary axes |.8-2.7 cm long, 3° axes 0.9-3.3 cm long; umbels 4-flowered, one umbel per node, primary and second- ary peduncles absent. Bracts 0.4-1.3 mm long, 0.5-1 mm _ wide, triangular, glabrous or tomentulose, or tomentulose only abaxially and/or on the margins; bracteoles 0.2-0.5 mm long, 0.2—0.4 mm wide, triangular, sometimes linear or semicircular, glabrous or tomentulose, or sometimes tomentulose only abaxially and/or on the margins. Pedicels 116-34 mm long, velutinous, Sepals 4.5-6.5 (-7) mm long, 2.3-2.7 (-3) mm wide, adaxially the proximal 4-4 glabrous, the distal *%4—'4 velutinous-tomentulose, glands 1.6—2.4 (-2.7) mm long, 0.8-1.2 mm wide. Claw of the lateral petals 2.3-2.6 mm long, limb 11.5-15 mm long, | I- 14.5 mm wide, orbicular, base briefly attenuate, denticulate, teeth up to 0.6 mm long, glandular or eglandular. Claw of the posterior petal 4—4.5 mm long, limb 8.5-9 mm long, 8.5—10 mm wide, broadly obovate, margin of the proximal %4—%4 capitate-glandular, fimbriae up to 0.3mm long, margin of the distal 4-% denticulate, teeth up to 0.1 mm long, glandular or eglandular. Lateral stamen filaments 3.5-4.1 mm long, posterior filament 3.3-3.7 mm long, always at least slightly shorter than the lateral four, slightly arced toward the posterior petal, anthers 1.4—1.5 mm long, glandular connectives 0.6—-0.8 (-1) mm long. Pollen grains 50-57 wm. Filament of the anterior staminode 3.8—5 mm long, anterior- lateral filaments 3.64.5 mm long, posterior-lateral filaments 3.4—3.8 mm long, always shorter than the anterior three, inflexed between the posterior styles, anterior and anterior-lateral glands 1.3-1.4 mm long and wide, posterior-lateral gland 1.2—-1.3 mm long, 1.1-1.2 mm wide, anterior and anterior-lateral glands usually with a shallow indentation across the apex and decurrent on the abaxial face, posterior-lateral glands smooth. Anterior style 3.24 mm long, ca 0.4 mm wide, slightly inclined toward the posterior petal, posterior styles 3.4—4.3 mm long, ca 0.3 mm wide, sometimes unequal, always exceeding the anterior style, divergent, arced toward the posterior-lateral petals, all styles usually glabrous, sometimes with scattered hairs in the proximal /% of their lengths, stigma of the anterior style 0.4—-0.5 mm in diameter, sues of the posterior styles 0.3-0.4 mm in oe all capitate. Samara not se Type. Irwin et al. 16860. Brazil. ae Grosso: Ca 15 km S of Xavantina, 400 m, 10 vie 1966 (holotype UB, isotypes MICH, NY). Distribution. (Fig. 10). Known only from the vicinity of Xavantina of east- central Mato Grosso; cerrado and margins of gallery forests. RAZIL. Mato Grosso: Xavantina, 400 m, Irwin et oe 16778 (NY, UB); ca 70 km : of Xavantina, 500 m, /rwin et al. 17439 (MICH, NY, UB); ca 3 km S of Xavantina, Ratter R112 p.p. (NY); Xavantina, Richards 6429 (K, NY, P); Chavantina SCS Gnay, Sick B481 (RB). Peixotoa psilophylla is the only species in Mato Grosso with the stem leaves glabrous or very sparsely pubescent when mature. ‘The internodes of the pri- mary inflorescence axis and the slender secondary axes are commonly 5-15 cm long, and give the inflorescence an open appearance. The hairs of the inflores- cence and any other part of the plant are white. The anterior and anterior- 62 lateral staminode glands usually have an indentation across the apex and decur- rent on the abaxial face. The margin of the limbs of the lateral petals and of the distal section of the limb of the posterior petal is denticulate. The geographically nearest glabrous species is P. hatschbachi of south-central Goias, which is distin- guished by its stalked leaf glands. In P. psilopylla the leaf glands are sessile. P hatschbachu also differs in its smaller flowers whose petals are fimbriate, its larger stem stipules (over 17 mm long and wide), and its smooth staminode glands. ‘ollected in flower in June and July. oe 25. Peixotoa reticulata Grisebach, Linnaea 13: 213. 1839. Plgsie? Aegon P. aa ee Grisebach, Linnaea 13: 214. 1839. Type. Sellow s.n. ie t 5505, fide Niedenzu, 1912] (Lectotype NY!). P. lateritia Adv. Jussieu, Ann. Sci. Nat. Sér. II, Bot. 13: 279. 1840. Type. Claussen s.n. (holotype P-JUSS!). P. hirta £. 1. minor poses Verz. Vorles. Akad. Braunsberg W-S 1912 1913: 35. 1912. Type. Warming s.n. (23 Sep 1864) (lectotype C!, Coon EF] LE) US|). Peron: eel eee (Grisebach) Niedenzu, Verz. Vorles. Akad. Brauns- berg W-S 1912-1913: 35. 1912. P. macrophylla £. L. minor (Niedenzu) Niedenzu, Das Pflanzenreich IV. 141: Doe 9g? ae bid goles £ Il. typica (Grisebach) Niedenzu, Das Pflanzenreich IV. 141: 28, = Pee ie Ferreira, An. Soc. Bot. Brazil, XX Cong. Nac. Bot. Goiana: 281. 1969. Type. Ferreira 2b (UB 37618) (holotype UB!). Shrub or subshrub, sometimes scandent, to 3 m. Vegetative branches finely velutinous or tomentulose, older parts usually glabrate. Stem stipules 8-25 mm long, 9.6-25 mm wide, entire with the apex acute, or bifid, velutinous or tomen- tulose or glabrate adaxially, tomentose-velutinous or sericeous-velutinous abaxially, deciduous; inflorescence stipules gradually decreasing in size toward the distal nodes, the smallest not less than 5.5 mm long and wide, entire with apex acute or notched, or bifid, sometimes the most distal fused at the base, deciduous. Stem leaves with the petiole 3-14 (-15.5) mm long, (sometimes golden) tomentose or velutinous-tomentose, laminae 3.5-17.5 cm __ long, 2.2-12 cm wide, elliptical, ovate, suborbicular, rarely lanceolate, the smaller sometimes obovate or rhombic, apex apiculate, base cordate or sometimes truncate, rugose, velutinous A tie: loosely pubescent with ‘T-shaped hairs or loosely tomentose or tomentulose below, rarely the pubescence restricted to the major veins below, a pair of sessile glands at the base at the costa or, more commonly, halfway on the petiole, rarely borne on the surface of the blade up to lim above the base, up to 1.5mm from the costa, each gland (0.7—) 1-2.2 (-3.2) mm in diameter, rarely with a second smaller pair (0.4—0.8 mm in di- ameter) immediately above the larger pair. Inflorescence leaves with the more proximal petiolate, usually the more distal sessile, laminas abruptly smaller than the cauline, the smallest not less than 1.4 mm long, 0.6 mm wide, shape of the more proximal like the cauline, the more distal lanceolate or oblanceolate or linear, the most distal usually subulate, apex acuminate, sparsely velutinous or glabrous above, densely velutinous or velutinous-tomentose below, a pair of elands at the base or halfway on the petiole, each gland 0.3—2.3 mm in diameter, rarely with a second, smaller pair (0.6—1 mm in diameter) immediately above or adjacent to the larger pair, rarely one or both glands absent. Inflorescence terminal or axillary, the axes s densely velutinous-tomentulose, internodes of the primary axis 1-12.5 cm long, 2° axes 0.7-11.8 (- Is 5.2) cm long, 2° subsidiary axes ~o rare, 2.5-3 cm long, 3° axes 0.5-6.5 cm long, 4° axes 0.4-3.8 cm long, 5° axes 63 el Sie cm long; umbels 4-flowered, one umbel per node, primary peduncles rare, up to 1.5 mm ou ee peduncles up to 6 mm long or absent. Bracts 0.3-3. 8 mm long, 0.2-1.3 (-2.3) mm wide, triangular or subulate, glabrous or velutinous or romeniiloc or sparsely so on both surfaces or only abaxially or only on the margins; bracteoles 0.1—-0.9 mm long, 0.1-0.6 mm wide, triangular or oblong or subulate, usually glabrous, sometimes tomentulose abaxially; bracts and bracteoles rarely absent. Pedicels 8-32 mm long, (sometimes golden) dense- ly tomentulose-velutinous, usually at least slightly longer in fruit. Sepals 4— 8.5 mm long, 24.3 mm wide, adaxially finely tomentulose or velutinous or pubescent with T- shaped hairs or sparsely so, or the proximal W424 glabrous, the distal ¥4—-¥3 pubescent, glands 1.4—3 mm long, 0.7-1.5 mm wide. Claw of the lateral petals 2.2—2.5 (-3.5) mm long, limb (11.2—) 12-15 (-18) mm long, 12.5-15 (—18) mm wide, orbicular or broadly obovate, base briefly attenuate, fimbriae up to 1.2 mm long, glandular or eglandular. Claw of the posterior petal 3.5—5 mm long, limb 8-10 (-12) mm long, 7—10 (-12) mm wide, orbicular or broadly obo- vate or broadly elliptical, fimbriae up to 0.8 (-10) mm long, those of the proxi- mal 1/2—4/5 capitate-glandular, the distal 1/2-1/5 fimbriate-glandular or some- times all capitate-glandular. Filaments of the lateral stamens 3.5-4.7 (—5.3) mm long, posterior filament 3.3-4.7 mm long, usually shorter, sometimes subequal to the lateral four, slightly or strongly arced toward the posterior petals, anthers 1.3-1.7 mm long, glandular connectives 0.6—0.9 mm long. Pollen grains 57-63 (-70) win. Filaments of the anterior and anterior-lateral staminodes 3.54.6 (-4.9) mm long, posterior-lateral filaments 3.2-4.3 mm long, usually a little shorter, sometimes equal to the anterior three, inflexed between the posterior styles, sometimes the anterior-lateral filaments exceeding the other three, eee 1.2-1.6 (-2) mm long, 1.1-1.5 mm wide, the posterior-lateral glands usually little smaller than the anterior three, all glands with the apex and abaxial ne smooth. Styles 3.6—5 mm long, 0.2-0.4 mm wide, equal or the anterior longer or shorter than the posterior two, the anterior style always a little stouter than the posterior two, anterior style arced or at least slightly inclined toward the poste- rior petal, posterior styles divergent along their entire length or only in the distal ¥3, arced toward the posterior-lateral petals, all styles usually glabrous, some- times velutinous in the proximal % of their lengths, stigmas 0.4—-0.5 mm in diameter, capitate. Samaras variable in size, the smaller usually with the dorsal wing 1|.3-2 cm long, 0.9-1.3 cm wide, upper margin straight or gently sigmoid, lower margin arced or nearly so, erose or slightly so, lateral wings 1.3— 2.5 mm wide, 6-8 mm high, rectangular, erose or sometimes lacerate, the larger samaras usually with the dorsal wing 2.3-4 cm long, 1.3-2.3 cm wide, upper margin str aight, arced or sigmoid, lower margin sigmoid or slightly so or arced, lateral wings (2.7—) 3-7.5 mm wide, 9-15 mm high, rectangular or semicirculat erose or slightly o Bel samaras witht the areole 3.5-6.5 mm long, 3—6.8 mm wide, nut ca 4-6.5 (-7.5) mm long, 3—5.5 mm in diameter, seed 6.2—7.4 mm el outer cotyledons 7.4-11.5 mm long, 1.5-2.2 mm wide, folded at ca 4—-%4 of Its ength, inner cotyledon 4.2-6 mm long, 1-2 mm wide, folded at ca 4/5—5/6 of its length or straight. ype. Sellow sm |Selow ii MinG 13 105(2 NN 1819), tide Niedenzu, 1912) Brazil. Minas Gerais: “... pr. Juesshuy [Suacui].” (lectotype NY!). Distribution. (Fig. 9). From central Minas Gerais to Parana, southern Goias to central and southern Mato Grosso, eastern Paraguay and eastern Bolivia; cerrado and campo. — BOLIVIA. Prov. Velasco, Kuntze s.n. (NY). BRAZIL. Distrito Federal: Brasilia, Ferreira 3, 9, 13 (UB), Heringer 8410 (UB, US), 11670, 11811 (UB), Irwin et al. 11103 (MICH, NY), fan oF Re 5067 (MICH, NY), 6717 (MICH), Martin 407 (GH), Pires et al. 9086 (UB, US), Sucre 274 (NY,R JB); Braclancict Ferreira 4 (UB); Lagoa Paranoa, 1000 m, Irwin et al. 15340, 26641 (MICH, on Beaciondin: Lima et al. 1 (NY, UB); entre Brasilia e Fercal, Pires 38097 (MICH, NY). Goias: . e : . eee. Ee Peixotoa reticulata yee fen , f \ 4 a phates | FIG. 9. Distribution of P. reticulata. Serra do Caiapo, ca 25-30 km S os Caiaponia, 800-1200 m, Anderson 9391, 9589 (MICH, NY); Serra i N of Corumba de Goias, 1250-1300 m, Anderson 10260 (MICH, NY); Goias, Burchell 7037 te Che ilcante, "Burchell 7596 (K); eee ze Veadeiros, Pouso Ali 1800 m, Ferreira 12 (UB); Goiania, Guimaraes et al. 292 (MICH, RB); mun. Ves BR-60, Hatschbach 34639 (ICH); mun. Luzania, Heringer 14691 (UB); Rio da Prata, ca 6 km S of Posse, 14°S, 46°W, Irwin et al. 14340a (NY); Serra do Catapo, ca 50 km S$ of Caiaponia, 1000 m, abi etal. 17830 (MICH); Serra dos Pirineus, 20 km N of Corumba Ss Golas on road to Niguelindia, 1150 m, [rwin et al. 18762 (NY); Serra dos Pirineus, ca 18 km E of Pirenopolis, oe m, /rwin al. 34248 (MICH, NY); ey ee cachoeira do Abade, Onishi et al. 74 (UB); ad S. Pedro, Pohl 472- Rea (K, W). Mato Gro ca 5 km N of Barra do Gargas, 500 m, Haas 9871 (MICH, NY); Campo Grande, Archer & can 140 (SP, roe yeas Caneiro, Hartmann 113 (SP); Rio Brilhante, Hatchbach & Guimaraes 21770 (C, M, ie P); Terenos, Hatschbach 24625 (MBM, MICH); Casa da Pedra, Hoehne s.n. (SP 302! 56); ee Grande: Hoehne &F Gehrt s.n. (SP 35760); Serra do Roncador, ca 210 km N of Xavantina, 500 m, Irwin et al. 16087 (MICH, NY, UB), ca 87 km N of Xavantina, 550 m, Trwin etal. 16484 (MICH, NY, UB), ca 60 km N of Sor annna. 550 m, Irwin et al. 16681b (MICH, NY); mun. Ha 42 km de Parts de 15 de Novembro, a 14392 (C, ne: 14396 (CTES); 23-40 km E de Aquiduana, Krapovickas & 7 Schinini 32918, 32893 (MICH); 4 km e Ribas do Rio Pardo, Krapovickas oo ae 34372 (MICH); 6 km de Campo Grande, Krapovic oa oe oO ristobal 34542 (MICH); Campo Grande, Lagoa Rica, 600-700 m, Lutz s.n. (R 72142); Cha apada, Moore 153 (BM); Campo Grande, Occhioni ae (MICH); mun. Dourados, ca 20 km E of Itaum, Pedersen 11116 (MICH); Fazenda Sta. F oo 56°40'W, 19°] 3" S, Prance & Schaller 26269 (NY); 8 km NE of Base Camp (12°54'S, 51°52'W), ca 270 km N of Xavantina, Ratter et al. 1990 (K); ca 200 km N of Navantina, Xavantina-Sao k -elix So de Santos & aan 1637 (K, MO, NY, P, RB, UB): Buriti, NE of 7 ee 22 50 ft, Smeth ads hae mun. Campo Grande, Fazenda das Varzeas, Sucre 10416 (RB); ca Fk of Campo Grande, Swalen 9606 (US). Minas Gerais: 6km N of Gouvéia on road to Diamantina 1250 m, yisrienee 8564 (MICH, NY); mun. Belo Horizonte, Serra do Taquaril, Barreto J7 AF, US), 7578 pp. (F); Aaa antina, Brade 13776 (RB); Belo Horizonte, Brade 14819 & Barreto ly (RB): Bau, Brade 17802 & Altamira (RB); Diamantina, Brito 100 (R); Claussen s.n. (P-JUSS, lectotype of P. lateritia); Claussen s.n. (BR, G); Carandai-Crespo, Duarte 45] (RB); Patos de Minas, 800 m, Duarte 2843 (RB); Cachambu, ca 1000 m, Duarte 3787 (RB); rodovia Uberaba-Uberlandia, a 40 km de Uberaba, Ferreira 2b (UB 37618, holotype on: gli) rodovia Goiania-Brasilia, entre Uberaba e Uberlandia, ilk 43 (NY, UB); Gaudichaud 584 (P); Pouso Alegre, Hoehne s.n. (SP 19391); Serra do Espinhacgo ca 25 km S . of Diamantina, on i. to ae 1300 m, [rwin et al. 22091 (NY), ca 20 km S W of Diamantina, 1200 m, /rwin et al. 22294 (MICH, NY); Ituiutaba, Macédo 38 (MO, S, SP), 2428 (BM, MO, S, US), 4962 (SP); Capao d’Antas, Nova ae Magathaes 241 (UB); bles _ i ~ ~ n lol ia S —_ Laon | 4 =) 65 40 km 0 de eben eee 19041 (NY); Diamantina, 1250 m, Mexia 5798 (A, BM, G, GH, MO, NY, US); Belo Horizonte, Porto G@ Fagundo 2200 (RB); Baependi, Rabello 893 (R); han [Uberaba], Regnell I- 28 (US), IT- 28a (S); Baguart, Regnell [1-28d (F, S, US); Caldas, Regnell I1-28e (C, R, S); Passos, Serra de Ventania, Regnell [/-28x (S); Serra do Curral, Belo Horizonte, Roth 1591 (RB); Schwacke s.n. (R 72455); Serra da Mantiqueira, Vidal 2054 (R); mun. Thermopolis, C hapadao e Jacuhy, Vidal 1-780, 1-800, 1-836 (R); José Alves, Warming s.n. (C); Lagoa Santa, Warming s.n. (C LE, US); outskirts of Belo Horizonte, 1000 m, Williams & Assis 5827 (GH); mun. Belo See Morro das Pedras, 1100 m, tae cf Assis 6938 (GH). Parana: Fazenda Lagéa, ca 20 km N of Cianorte, Anderson 11155 (MICH, NY, UB); mun. Campo Mourao, Hatschbach 8819 (MICH, US); Fazenda Lagoa, Cianorte, See 14243 (F, MBM, MICH, NY, P, US), 21572 (MICH, P); Campo Mourao, Labouriau 60 (SP). Sao Paulo: Mogi-Mirim, Burchell 5090 (K), 5220 (GOET, K); Araras, de Andrade s.n. (SP 24514); Itirapina, de Paula 11] (SP); Morro Pellado, Edwall s.n. (SP 12109); mun. Moji-Guacgu, Fazenda Campininha, ca 4 km NNW of Padua Sales, 575-650 m, Eiten & Eiten 1901 (NY, SP, US), ca 10 km NNE of Padua Sales, Eiten & Eiten 2280 (SP); mun. Sao Carlos, 3.5 km NNW of center of Sao Carlos, 21°59'S, 57°55'W, 825 m, Eiten et al. 2914 (K, NY, SP, US); mun. Ibaté, oe 8 km NW of center of city of Sao Carlos, ca 1 km NE of Ibate, Evten et al. 2928 (MO, NY, SP, US); un. Sao Carlos, 13-15 km NW of Santa Eudoxia, Eiten & Eiten 3291 (NY, SP, UB); mun. Sao Gabe near boundary with mun. Araraquara, 19.9 km NW of railroad st item at Santa Eudoxia, 625 m, Evten & Campos 3449 (NY, SP); mun. Moji-Guacu, eee Campininha, 6.2 km NNW of Padua Sales, 575-650 m, Fiten & Eiten 5616 (NY), 10 km E of ome Sales, Eiten & Eiten 5701 (NY, UB, US); Itirapina, Gehrt s.n. (SP 8328); mun. MeL 18 km N of Botucatu, 14 re E of Sao Miguel, ca 550 bees ot 6571 (NY); caminho para Sete-Que nee Heiner 269 (S); Mogi-Mirim, Hoehne s.n. (SP 20525); ae Hoehne s.n. (SP 20569); Sao Simao, Kuhlmann 4144 (SP), Lima 6268 (SP); Rio Claro, ee 587 (C); Sorocaba, Lund s.n. (C, UB); [tirapina, Matos 92 (SP); mun. Moji- Guagu, Fazenda Campininha, 10 km de Padua Sales, Mattos 125 & Rizzini (RB); mun. Moji-Guacu, Fazenda Campininha, 3,2-3.4 km NNW de Padua Sales, Mattos & Mattos 8305 (SP, US); Sao Joao de Boa Vista, Mosén 4050 (P, S); Votuporanga, Pires 57925 (NY, UB); inter Casa Branca et Sao Simao, Regnell [1-28b, H-28g (S); Minas, Riedel 584 (R); Sorocaba et Ypanama [Ipanema], Riedel & Luschnath 2007 (A, GH, LE, NY, P, S, US); zwischen Rio Pirituba und anes Sellow IV. ut. 5505 (NY fragment, BS onne of P. macrophylla); prope Franca, Wacket s.n. (W). PARAGUAY: Cerro Cora, 36 km Pedro Juan Caballero, 300 m, Anderson 11782 (MICH); 3 km < of Rio Ypané on road to Pedro nee Cabellero, 175 m, Anderson 11790 (MICH); Est. Santa Maria, Cerro Noaga, Anisits 2372 (S); dept. Kanendiju, col. Fortuna, 8 km de Kuruguatz, Arenas 629 (CTES, MICH); dept. Paraguar, Salto de Pirareta, AGhe etal. 1750 (MICH); Cordillera de Altos, Cerro Chocha, eee 390 (A, BM, F, , K, M); Centurion, ce Rio Apa und Rio Aquidaban, Fiebrg 4017 (BM, G, GH, K), 4590 (GH); San Luts, zwischen Rio Apa und Rio Aquidaban, 22-— 23°S, Fiebrig 5181 (G, M); camino Bernal- ae Hassler ye (G); Cordillera de Altos, Hassler 2976 (BM, G, NY); Bernal-Cué, Hassler 4055 ay G, NY, P); e fl. Corrientes, Hassler 4509 (BM, ae K, P); Ipé-hu, Hassler 5054 (BM, G, K, P); a Capi, ee 5927 (A, BM, F, G, K, MO, , P, S, W); prope Chololo, in valle Ae Y-aca, Hassler 6774 (A, BM, F, G, K, NY, P, S, W); Rio ee Hassler 8166 (A, BM, F, G, K, MO, NY, P, S, W), 8/66a (BM, F, co K, NY); Sierra de Amambay, Hassler 9881 (A, G, K, NY, P, W), 988/a (GH K), Conrados Estrella, epee 10056 (G); dept. San Pedro, 43 km de San Estanislao Norte, Krapo- vickas et al. 14265 (CTES, MO); dept. Paraguarr, Pirareta, Schinini 4348 (F, MICH, P); dept. San Pedro, Ruta 3, 2km S de Tacuara, Schinini 8809 (MICH), 34km N de Tacuara, Schinini 8887 (MICH); dept. Amambay, camino entre Ruta 5 y Bella Vista, 9km N de Ruta 5, Schinini 8956 (MICH). Peixotoa reticulata is a widespread taxon of which almost all of the known collections are apomictic. The anthers remain closed and contain mostly aborted pollen that 1s 80— 97% non-staining in cotton blue in lactophenol. Ina few excep- tions this percentage is much lower (see also the section Apomixis). Rarely a few flowers of a specimen (e.g. Fiebrig 4590) will have a few or all of the anthers open. Pollen fertility may be higher than or the same as in samples from closed anthers. Only in two collections, Mexia 53798 and Occhioni 5775, do all flowers have all anthers open. The pollen of Mexia 5798 is 45-98% non-staining, while that of Occhioni 5775 is almost 100% staining. The Occhioni collection is perhaps a rare, sexual individual. Chromosome counts of the pollen mother cells of a collection from Para- guay (Anderson 11790) show that while 2n=30, the separation of anaphase I/ telophase I is highly irregular. Most frequently 17 and 13 or 16 and 14 pairs would move to the opposite poles, but figures with 15 and 15 were also seen (W. R. Anderson, pers. comm.). In the only sexual species counted, P. glabra (Ander- son 11549), n=10(W. R. Anderson, pers. comm.). Since Gates (1981) found that 66 in Banisteriopsis, the most likely ancestor of Peixotoa, n=10 in nearly every count, this is probably also the base number for Peixotoa. P. reticulata may be the product of a cross between a diploid and a tetraploid. The resulting sterile triploid was then propagated apomictically. Such crosses may have happened several times. Also, as noted in the section on apomixis, it appears that hybridization between P. reticulata and sexual species does take place. Thus, P. reticulata as circum- scribed here is probably polyphyletic. Such a history would account for its con- siderable morphological variation, which is reflected by a number of regional types. Their distinctions are summarized in Table I. These groupings are not abolutes, but indicate the combination of characters most likely to be encoun- tered in each region. The extremes are quite distinct, yet are linked by many intermediate forms. The samaras are extremely variable in the shape and the size of the wings. Those from the eastern part of the range tend to have the dorsal wing with the upper and lower margin straight or sigmoid and often parallel to each other. The lateral wings are 3-5 mm wide. Those from the more western part tend to have an arced or sigmoid upper margin and usually a semicircular or arced lower margin; their lateral wings are 4-7.5 mm wide. Perhaps these reflect hybridization with the large-winged P. cordistipula and P. magnifica. The varia- lion in size is striking. In Hatschbach 2465 from Terenos in southern Mato Grosso the dorsal wing ts ca 3.5-4 cm long and ca 2-2.5 cm wide. The lateral wings are ca 1.6 cm high and ca 6-7 mm wide. In Hatschbach 21770 from Rio Brilhante, south of Terenos, the dorsal wing is ca I-1.5 cm long and ca 1 cm wide. The lateral wings are ca 6-6.5 mm high and ca 2 mm wide. These fruits appear immature or aborted, yet contain mature, apparently normal embryos. n the past plants of P. reticulata from the western part of the range have often been assigned to P. cordistipula of central Mato Grosso and eastern Bolivia. This species differs from P. reticulata in its larger and persistent inflorescence stipules, longer staminode filaments, and the commonly present glandular mass in the center of the umbel. Also, the dense, white pubescence gives P. cordistipula a silvery-grey appearance. Grisebach based his P. reticulata on two Sellow collections from east-central Minas Gerais. These were deposited at Berlin, but are no longer extant. A fragment of Sellow I, it. c. 1310, consisting of a leaf and flowers, is at NY and is here designated lectotype. Photographs of the Berlin specimen are at F, MICH, and NY. Mexia 5798 is an excellent match for this collection. It is noteworthy that the flowers of the NY fragment have open anthers. Collected in flower and fruit throughout the year. 26. Peixotoa sericea C. Anderson, sp. nov. Bigs) 2. Liana (?). Stipulae vegetativae deciduae, inflorescentiae 6—10 mm longae, acutae, persistentes. Petioli 8-10 mm longi. Laminae 7.1—10.3 cm longae, 3.3- 9 cm latae, ellipticae vel anguste ellipticae, supra sparsim appresso-sericeae vel glabratae, subtus appresso-sericeae, biglandulosae 4.5-9 mm supra basim juxta vel prope costam. Folia inflorescentiae petiolo brevi, distaliora sessilia, linearia. Inflorescentia terminalis vel axillaris, umbellis singularibus. Bracteae bracteo- laeque absentes. Pedicelli 13-15 mm longi. Sepala 3.8-4.5 mm longa. Limbus petalorum lateralitum ca 10mm longus, ellipticus, margine glanduloso- fimbriato. Limbus_ petali posuct 6-6.5 mm longus, ellipticus vel anguste obo- vatus, margine glandulis capitatis ornato. Styli subaequales, 44.5 mm longi, anticus arcuatus versus petalum posticum, postici divergentes, inclinati versus petala postico-lateralia; stigmata capitata. Samara ignota. Viner Vegetative branches finely sericeous-tomentulose when very young, soon becoming glabrous. Stem stipules deciduous, not seen; inflorescence ‘Table I. Summary of the regional morphological types of P. reticulata. central Minas Gerais estern & southern Minas Gerais, & Sao Paulo il centrz Mato Grosso southern Mato Grosso Paraguay shape of stem laminas —_ abundance o hairs on lower surface of stem lamina presence of petioles in inflorescence leaves presence of secondary peduncle shape of limb of posterior etal margin of limb of posterior petal curvature of anterior style — ovate, elliptical, suborbicular, rhombic Sparse to moderate rare obovate, elliptical capitate- or fimbriate- glandular arced toward posterior petal ovate, elliptical, suborbicular dense common obovate, orbicular capitate- glandular arced toward posterior petal Parana Goias ovate, ovate, elliptical elliptical, suborbicular moderate to dense moderate common rare orbicular obovate capitate- glandular capitate- or fimbriate- glandular arced toward arced toward posterior posterior petal petal ovate, elliptical, lanceolate sparse to dense rare elliptical capitate- glandular slightly inclined > ovate, elliptical sparse to dense rare obovate capitate- glandular ovate, elliptical moderate almost always present orbicular, oblate capitate- or fimbriate- glandular erect or nD 68 stipules 6-10 mm long, 6-8 mm wide, cordate or triangular-cordate, apex acute, sparsely and finely velutinous or sericeous adaxially, sericeous abaxially, per- sistent. Stem leaves with the peuole 8-10 mm long, densely velutinous, laminas 7.1-10.3 cm long, 3.3-5 cm wide, elliptical or narrowly so, apex apiculate, base attenuate, sparsely appressed-sericeous to glabrous above, appressed-sericeous below, a pair of sessile glands borne 4.5-9 mm above the base at the costa or on the surface of the lamina up to 0.4 mm from the costa. Inflorescence leaves of the proximal nodes petiolate, the more distal sessile, laminas abruptly smaller than the cauline, the smallest not less than 2.2 mm long, 0.6 mm wide, the proximal narrowly elliptical, the distal linear, apex acuminate, glabrous above, densely sericeous-velutinous below, a pair of glands at the base at the costa or borne on the surface of the lamina up to 1mm above the base, up to 0.4mm from the costa, each gland 0.3-0.9 mm in diameter. Inflorescence terminal or axillary, the axes finely and densely tomentulose-velutinous, inter- nodes of the primary axis 2-4 cm long, 2° axes 1.7-6.8 cm long, subsidiary axes absent; umbels 4-flowered, one umbel per node, primary and_ secondary peduncles absent. Bracts and bracteoles absent. Pedicels 13-15 mm long, densely golden velutinous. Sepals 3.8-4.5 mm long, 1.7—1.8 mm wide, velutinous adaxi- ally, sparsely so near the base, glands 1.1-1.2 mm long, 0.7-0.8 mm wide. Claw of the lateral petals 2-2.2 mm long, limb ca 10 mm long, ca 7.5 mm wide, ellipti- cal, base attenuate, fimbriae up to 0.4 mm long, glandular. Claw of the posterior petal ca 4 mm long, limb 6-6.5 mm long, 4.5-5 mm wide, elliptical or narrowly obovate, fimbriae up to 0.5 mm long, capitate-glandular. Filaments of the lateral stamens 4.5—-4.8mm long, posterior filament ca 3mm long, strongly arced toward the posterior petal, anthers 1.3—1.4 mm long, glandular connectives 0.6— 0.7 mm long. Pollen grains 42-50 wm. Anterior staminode filament 4.5—5 mm long, anterior-lateral filaments 3.3-4 mm long, posterior-lateral filaments 3- 3.3 mm long, always shorter than the anterior lateral two, inflexed between the posterior styles, anterior and anterior-lateral glands ca 1.2-1.3 mm long, ca L.1- .2.mm wide, posterior-lateral glands ca | mm long, ca 0.8 mm wide, all glands with the apex and abaxial face smooth. Styles 44.5 mm long, ca 0.3 mm wide, anterior style arced toward the posterior petal, posterior styles divergent, in- clined toward the posterior-lateral petals, all styles densely pubescent with basi- fixed hairs in the proximal 4 of their lengths, stigmas ca 0.3-0.4 mm in di- ameter, capitate. Samara not seen. Type. Eupunino 120. Brazil. Bahia: Santa Cruz de Cabrdalia, Reserva Biologica do Pau-brasil, 5 Jan 1972 (holotype CEPEC, isotype MICH). Peixotoa sericea is known only from the type (Fig. 10). It is one of two species in the genus with sericeous leaves. The hairs are sessile or subsessile and have a straight, stiff trabecula. The lamina is very sparsely pubescent above but more densely so below. Yet, to the naked eye the leaves may appear glabrous. The leaves are also atypical for the genus in that the base is cuneate instead of cordate or subtruncate. The glands of the stem leaves are borne above the base but at the costa, usually at different levels from each other. P. adenopoda, the other species with sericeous leaves, differs in that the leaf glands are stalked and borne on the surface of the lamina away from the base and costa. Also, the terminal inflores- cence nodes bear three umbels, i.e., are 12-flowered. Those of P. sericea bear a single umbel and thus are 4-flowered. Unfortunately, the collector of the type did not note the habitat. According to Mori and Silva (1979), the reserve at Santa Catarina Cabralia consists largely of mata higrofila (tropical wet forest). Perhaps P. sericea grows at the forest edge or in the adjacent restingas. Cerrado, the vegetation type in which Peixotoa is most commonly collected, is not found in this area. Peixotoa hispidula ts the only other coastal species in Bahia. It has the typically 69 Peixotoa 4 psilophylla ‘~. MM sericea \| @ spinensis FIG. 10. Distribution of P. psilophylla, P. sericea, and P. spinensis. cordate leaves which are usually glabrous or nearly so or may be variously pubescent but never sericeous. Its large flowers are borne in a solitary umbel or in only slightly more elaborate inflorescences. 27. Peixotoa spinensis C. Anderson, sp. nov. Fig. 16. Frutex vel suffrutex caulibus usque ad 2 m longis, prostratis, ascendentibus vel volubilibus. Stipulae vegetativae 3—10.5 mm longae, cordatae vel triangu- lares, acutae vel bifidae, deciduae interdum persistentes. Petioli 2-8 mm longi. Laminae 3.6-9.3 cm longae, 1.8—6.5 cm latae, ellipticae vel ovatae vel lanceo- latae, supra velutinae raro tomentoso-velu utinae, subtus tomentosae vel lanatae, ferentes 2 (interdum 3-4) glandulas (2.2-)3-20 mm supra basim 0-14 mm a costa. Folia inflorescentiae petiolo brevi raro sessilia, linearia vel subulata. In- florescentia terminalis vel axillaris, umbellis singularibus. Bracteae bracteolae- que praesentes. Pedicelli 8-28 mm longi. Sepala 3—-6.1 mm longa. Limbus petalorum lateraltum 9.5-13.5 mm longus, orbicularis, margine glanduloso- fimbriato vel glanduloso- denticulato, interdum praesertim apice eglanduloso. Limbus petali postici 6—8.3 mm longus, ellipticus vel late ellipticus, raro sub- orbicularis, margine glandulis capitatis ornato vel apice glanduloso-fimbriato. Glandulae staminodiorum apice profunde indentatae, indentatione in pagina abaxiali decurrenti, interdum postico-lateralibus laevibus. Stylus anticus 3. 4.2 mm longus, erectus vel parum inclinatus versus petalum posticum, styli postici 3.4—4.5 mm longi, semper anticum excedentes, divergentes, inclinati versus petala postico-lateralia; stigmata uaa Ala dorsalis samarae ca 2 cm longa, ca 1.4 cm lata, alae laterales 2.3-3.5 mm latae, 7.3-8 mm altae. Shrub or subshrub with ascending, ca or prostrate stems to 2 m. Vegetative branches densely tomentulose- VS MUMS, pubescence abraded from older parts. Stem stipules 3-10.5 mm long, 3.5-10 mm wide, cordate or tri- angular, entire with the apex acute, or bifid, finely velutinous-tomentulose, 70 sparsely so adaxially, densely so abaxially, deciduous or sometimes persistent; inflorescence stipules like the cauline, sometimes more densel pubescent, deciduous. Stem leaves with the petioles 2-8 mm long, densely tomentose- velutinous, laminas 3.6—9.3 ¢m long, 1.8-6.5 cm wide, elliptucal or ovate to lanceolate, apex apiculate or in narrow leaves acuminate, base cordate, velu- tinous or rarely velutinous-tomentose above, tomentose or woolly below, a pair of sessile glands borne on the surface of the lamina (2.2-) 3-20 mm above the base, up to 14 mm from the costa, sometimes with one or two additional glands, each gland 0.5-1.5 (-2.3) mm in diameter. Inflorescence leaves with the petiole .2 mm or more long, rarely sessile, laminas abruptly smaller than the cauline, the smallest not less than 1.4 mm long, 0.7 mm wide, shape of the more proxi- mal like the cauline, the more distal lanceolate to linear to subulate, apex acumi- nate, velutinous or sparsely so adaxially, densely velutinous abaxially, a pair of glands borne on the surface of the lamina in the larger leaves, at the base at the costa or halfway on the petiole in the smaller leaves, each gland 0.3-1.3 mm in diameter, sometimes with one or two smaller glands (0.4—0.6 mm in diameter) above, below or adjacent to the larger pair. Inflorescence terminal or axillary, the axes tomentulose-velutinous, internodes of the primary axis 2-8.5 cm long, 2° axes 1.9-8.7 cm long, 2° subsidiary axes 1.2—-2 cm long, 3° axes 0.7-3.3 cm long; umbels 4-flowered, one umbel per node, primary and secondary pedun- cles absent. Bracts 0.3—1.6 mm long, 0.2-0.7 mm wide, linear, triangular or sometumes semicircular, glabrous, sometimes velutinous on the margins; bracteoles 0.1—0.5 (—0.7) mm long, 0. 1-0.4 mm wide, triangular or semicircular, glabrous. Pedicels 8-28 mm long, densely tomentulose-velutinous, the smaller ones up to twice again as long in fruit. Sepals 3-6.1 mm long, 1.6-3.5 mm wide, adaxially the proximal %—% glabrous, the distal 4-24 tomentulose, glands |- 2.1 mm long, 0.7—1.2 mm wide. Claw of the lateral petals 2.5-3.5 mm long, limb 9.5-13.5 mm long, 10-13 mm wide, orbicular, base attenuate, fimbriate and/or denticulate, fimbriae and teeth up to 0.4mm long, glandular, sometimes eglandular, especially at the apex. Claw of the posterior petal 3.2-4.3 mm long, limb 6-8.3 mm long, 5.2-7.2 mm wide, elliptical or broadly so, rarely sub- orbicular, fimbriae up to 0.3 (-0.5) mm long, capitate-glandular, sometimes the apex fimbriate-glandular. Filaments of the lateral stamens 3.5—4.6 mm long, posterior filament 3.24 mm long, always shorter than the lateral four, arced toward the posterior petal, anthers 1.1-1.4 mm long, glandular connectives 0.6— 0.8 mm long. Pollen grains ca 50 wm. Filament of the anterior staminode 3.5— + mm long, always exceeding the anterior-lateral two, anterior-lateral filaments 3.2-3.8 mm long, always exceeding the posterior-lateral two, posterior-lateral filaments 3-3.5 mm long, inflexed between the posterior styles, anterior and anterior-lateral glands 1.2-1.4 mm long, 1.2-1.5 mm wide, the anterior gland often a little larger than the anterior-lateral two, posterior-lateral glands 0.8— I.2 mm long, ca 1 mm wide, all glands with an indentation across the apex and abaxial face or sometimes the posterior-lateral two smooth. Anterior style 3.1- 4.2 mm long, 0.3-0.4 mm wide, always shorter and a little stouter than the poste- rior two, erect or very slightly inclined toward the posterior petal, posterior styles 3.44.5 mm long, ca 0.3 mm wide, divergent, inclined toward the posterior- lateral petals, all styles glabrous or sometimes with scattered hairs in the proxi- mal '3 of their lengths, stigmas 0.4—0.5 mm in diameter, capitate. Carpophore up to 1.7 mm long. Torus up to ca 4.5 mm high. Samara with the dorsal wing ca 2cm long, ca 1.4.cm wide, upper margin arced, lower margin straight, slightly erose, lateral wings 2.3-3.5 mm wide, 7.3-8 mm high, rectangular, margin erose to lacerate, areole 4.2-4.5 mm long and wide, nut ca 5.6 mm long, ca 3.5 mm in diameter, mature seeds not seen. Type. Anderson 8730, Brazil. Minas Gerais: Serra do Espinhago, 25 km by ~ 71 road NE of Diamantina, ca 1.5 km from Rio Jequitr, 790-900 m, 12 Apr 1973 (holotype UB, isotypes MICH, NY). Distribution. (Fig. 10). Known only form the northern and central part of the Serra do Espinhacgo of Minas Gerais; cerrado. RAZIL. Minas Gerais: 25 km by road E of Diamantina, 2 km W of SeReuaee 790 m, Ander- son 8364 (MIC H, NY, UB); estrada para Grao Mogul, Graziela [Barroso] s.n. (RB 171697); mun. Grao Mogul, Rio eens eeu Hatschbach 41240 (MICH); mun. Grao Mogul, rod. Cristalia, Hatschbach 41394 (MICH); ca 18 km W of Grao Mogul, 950 m, /rwin et al. 23662 (MICH, NY, UB); ca3 km N of Sao Joao da Chapada, 1200 m, /rwin et al. 28236c (NY); slopes N of Grao Mogul, Maguire 49285 (MICH, NY); entre Montes Claro e Riacho dos Machados, Santos s.n. (NY). Peixotoa spinensis is easily recognized by the leaf glands that are borne on the surface of the lamina away from the base and costa. The laminas are densely tomentose or woolly below. All or only the anterior and anterior-lateral stami- node glands have an indentation across the apex and decurrent on the abaxial face. P. paludosa is the only other species of the Serra do Espinhago in which the leaf glands are borne on the surface of the lamina. It differs in that its leaves are only moderately pubescent and in that its staminode glands are smooth. Also, its inflorescence stipules are usually persistent at the more distal nodes. The in- florescence stipules and also commonly the inflorescence leaves of P. spinensis are deciduous. The larger leaves of P. leptoclada sometimes have the glands borne on the lamina but usually in addition to a basal pair. P. leptoclada is distinguished by its large (at least 7 mm long), persistent inflorescence stipules Collected in flower in Febr uary through April, also in Naar and Septem- ber, in fruit in February. 28. Peixotoa tomentosa Adr. Jussieu in St. Hilaire, Fl. Bras. Mer. 3: 61. 1832 (kS33)]. Fig. 18. P. tomentosa B oligothrica Adr. Jussieu in St. Hilaire, Fl. Bras. Mer. 3: 62. 1832 [1833]. Type. St. Hilaire 392 (lectotype P!). P. tomentosa crys Grisebach, Linnaea 13: 212. 1839. Type. Sellow s.n. Caee TH. it. BIS40 c1306 (26 Dec 1818), fide Niedenzu, 1912] (holo- e B, eed fragment of holotype, NY!). ie finn Adr. Jussieu, Ann. Sci. Nat. Sér. II, Bot. 13: 280. 1840. Type. Martius s.n. (holotype M!). P. tomentosa f£. 2. cuneata Niedenzu, Verz. Vorles. Akad. Braunsberg W-S LO se 38. 1912. Type. Sellow IV. it. 5504 (lectotype NY!). P. tomentosa f. 3. gracilior Niedenzu, Verz. Vorles. Akad. Braunsberg W-S 2-1913: 38. 1912. Type. Glaziou 16731 (lectotype C!, isotypes BR! KLE! Pi RY): Erect, ascending or prostrate shrub or subshrub to 2 m. Vegetative branches loosely or densely golden velutinous or velutinous-tomentose when young, pubescence faded to white and often abraded from older parts. Stem stipules (2.5-) 3.1-9.5 mm, 3.2-10 mm wide, cordate or cordate-triangular or triangular, entire with the apex acute or notched, or bifid, glabrous or finely velutinous or tomentulose adaxially, densely velutinous-tomentulose abaxially, deciduous; inflorescence stipules like the cauline, sometimes a little larger, usually more densely pubescent, often persistent. Stem leaves with the petiole (1-) 2-9 mm long, densely golden velutinous or velutinous-tomentose, lamina 4.5—10.8 cm long, 2.6-8.7 cm wide, ovate or elliptical or suborbicular or sometimes narrowly elliptical or lanceolate or oblanceolate or rhombic, apex apiculate, base cordate or slightly cordate or almost truncate, densely golden velutinous above, tomen- tose or sometimes also pubescent with T-shaped hairs or woolly below, a pair of 72 sessile glands at the base at the costa or halfway on the petiole or sometimes on the petiole just below the lamina, each gland 0.8—2 (—2.5) mm in diameter. In- florescence leaves with the petiole 0.5 mm or more long or the most distal sub- sessile or sometimes sessile, laminas abruptly smaller than the cauline, the small- est not less than 3.8 mm long, | mm wide, shape of the more proximal like the cauline, the more distal lanceolate or linear, the most distal sometimes subulate, apex acuminate or caudate, velutinous or sparsely so above, densely velutinous or velutinous-tomentulose below, a pair of glands at the base at the costa or halfway on the petiole or rarely on the petiole just below the lamina, each gland Q.8—1.8 mm in diameter, sometimes with a second, usually smaller, pair (0.3— 1.3 mm in diameter) immediately above the pair of larger glands. Inflorescence terminal, the axes densely golden velutinous or velutinous-tomentulose, inter- nodes of the primary axes 1.5-8.8 cm long, 2° axes 1.4—-18.3 cm long, 3° axes 2.4—3.5 cm long, subsidiary axes absent, usually at the more distal nodes the 2° and 3° axes suppressed, with | or 2 umbels sessile or borne on secondary peduncles in the leaf axil, always with 3 umbels at the terminal nodes, each umbel 4-flowered, primary peduncles absent, secondary peduncles up to 12.5 cm long or absent. Bracts 0.6-1.8 mm long, 0.3-0.8 mm wide, triangular or narrowly triangular, usually glabrous, sometimes tomentulose abaxially or on the margins; bracteoles 0.1-1 mm long, 0.1-0.3 mm wide, triangular or nar- rowly triangular or semi-circular, usually glabrous, sometimes sparsely tomen- tulose on the margins; bracts and/or bracteoles sometimes absent. Pedicels 8—2 | (—29) mm long, densely golden velutinous or velutinous-tomentulose, sometimes a little longer in fruit. Sepals 3.3-6 mm long, 1.9-2.8 mm wide, adaxially velutinous or tomentulose or pubescent only in the distal 1/8-1/2, the proximal 1/2-7/8 glabrous, glands 1.2—2.2 mm long, 0.8—1.3 mm wide. Claw of the lateral petals 1.5-2.7 mm long, limb 9.5-12.3 mm long, 10-11 mm wide, orbicular or broadly elliptical, base briefly attenuate, fimbriae up to 0.5 mm long, glandular or eglandular. Claw of the posterior petal 3-3.5 mm _ long, limb 6.6-9.5 (—10.2) mm long, 5.9-9 (— 9.6) mm wide, orbicular or broadly obovate, fimbriae up to 0.7 (-1) mm long, capitate-glandular in the proximal 4—%4, the distal 43—% fimbriate-glandular. Lateral stamen filaments 3.2-4.5 mm long, usually sub- equal, sometimes the posterior- lateral filaments longer or shorter than the ante- rior-lateral two, posterior filament 2.8-4 mm long, usually shorter than the lateral four, arced toward the posterior petal, anthers I-1.2 (-1.4) mm long, glandular connectives 0.6-0.9 mm long. Pollen grains ca 50 ym. Anterior staminode filament 3.7-4.7 mm long, exceeding the anterior-lateral two at least slightly, anterior-lateral filaments 3.24.3 mm long, usually longer than or some- times subequal to the posterior-lateral two, posterior- -lateral filaments 3.1- + mm long, inflexed between the posterior styles, anterior and anterior-lateral glands |.1-1.4 mm long, I.1-1.4 m wide, the anterior gland often a little larger than the anterior-lateral two, posterior glands 1.1-1.2 mm long, 0.8-1.2 mm wide, always a little smaller than the anterior three, all glands with the apex and abaxial face smooth. Anterior style 3.44.2 mm long, 0.3—-0.4 mm wide, equal to or exceeding the posterior two, arced toward the posterior petal, posterior styles 3.2-4 mm long, 0.2-0.3 mm wide, divergent, inclined toward the posterior- lateral petals, stigmas 0. aes mm in diameter, capitate. Carpophore up to 2.8 mm Ong. ‘Torus up to ca 5 mm high. Samara with the dorsal wing 1.5-2.2 cm long, 1.1-1.5 cm wide, upper margin straight, sigmoid or arced, lower margin arced, erose, lateral wings 2.3-5.2 mm wide, 7.2-14 mm high, rectangular or semicircular, slightly erose, areole 2.2—5 mm long, 2.5-4mm_ wide, nut 3.5-6 mm long, 2.8-4.5 mm in diameter, seed 5.8-7 mm long, outer cotyledon 4.9-7.9 mm long, 1.8-2.3 mm wide, folded at 24—% of its length, inner cotyledon 4.5-6.1 mm long, 1.6—2.2 mm wide, folded at 2/3-4/5 of its length FIG. 1. Distribution of P. tomentosa. Type. St. Hilaire s.n. Brazil. Minas Gerais: “In montibus ... Serra da Caraga et Serra d’Ouro Branco necnon pagum Nossa Snra da Penha (Minas Novas) . (lectotype P!). Distribution. (Fig. 11). Known mostly from the central section of the Serra do Espinhaco of Minas Gerais; cerrado, campo, and along gallery forests. AZIL. Minas Gerais: 12 km by road W of Diamantina on road to Curvelo, 1450 m, Anderson 8430 (MICH, NY); 7 km W of Diamantina, KM 299 on MG-259, a m, Anderson 35155 (MICH, NY); mun. Belo Horizonte, Serra do Taquaril, Barreto 7578 p.p. (RB), 7622 (F); Sao Sebastiao do Paraiso, Morro do Liso, Beta 97 (R); Belo Horizonte, Brade 11822 (R); Sao Sebastiao a Paraiso, Bau, Brade 17810 & Altamira (RB); Ouro Preto, Campos s.n. (SP 18825); circa Cachoeira do Campo, Casaretto 2711 (G); Ouro Branco, Castellanos 25616 et al. (NY); Claussen 14 (P), 35 G, P). 75A (BR), 6A (BR), 131 (BM, BR, GH, MO, S, W), 13/c(W), 141 (BM), 157 (BM), 197 (BR, G, P, W), 414 (BR, G, W); de Rodrigo Silvio, anes 922 (G); ca 50 km N of Congonhas along highway 135 to Belo LeKOve eo: 1270 m, Davidse S Ramamoorthy 10795 (MICH, MO); mun. Ouro Preto, between eee and Belo Horizonte, KM 15, ca 1200 m, Davis & Shepherd D59601 (E); Serra do Se ar M 132, 1330 m , Duarte 2206 (RB, S); Serra do Cips, mun. Jaboticatubas, along road at KM 111, 19°19’S, 43°36'W, 1000-1050 m, Fiten & Fiten 6837 (K, NY, UB, US), along road at KM 116-117 from Almeida to Conceigao do Mato D entro, 1150 m, 11072 (MICH); mun. Caeté, Saou ae Felippe 31 (SP, US); Serra da Pinheiro, pres de Sao Joao del Rei, Glaziow 16731 (BR, C, LE, R); Serra ‘Ouro Branco, Sao Juliao, Glaziou s.n. (BM, P); Serra do Cipo, mun. Santa Ana ae Riche: Hatsch- bach 35286 (MICH); Serra Sao Joao del Rei, Itacolumit, Helmenreichen s.n. (W); Miguel Burnier, Hoehne s.n. (SP 5117); Serra do Helos inlay ca 45 km SE of Belo Horizonte, ca 1500 m, Irwin et al. 19514 (MICH, NY); Serra do Cipo, c a KM 112, ca 135 km N of Belo Horizonte, 1200 m, frwin et al. 20436 (MICH, NY); ca 18 km N EOF Diamantina on road to aera 1300 m, Irwin et al. 22676 Cine H, NY); ca 7 km NE of Diamantina on road to Mendanha, 1300 m, Irwin et al. 22833 (MICH, Y); 8km E of Diamantina, 1175 m, Jrwin et al. Hee (MICH, et ca 3km N of Sao Ieee rsh Cees 1200 m, /rwen et al. 28236a (MICH, NY); Serra da Caraga, i km W of Barao de Co 400 m, Irwin et i 28891 (MICH, NY), 28976 (NY), 29081 (MICH, NY); Serra do Cipo, mun. Jaboticatubas, KM 123 rodovia Lagoa Santa-Conceicao do Mato meee Diamantina, Joly & Semir 3592 ee te KM 112 rodovia Lagoa Santa-Conceigao do Mato Pee Diamantina we etal. 1009, 1401 . Ouro Preto, 1150 m, Krapovickas & Cristobal 33538 (MICH); Serra do Cipo, Kuhlmann & is 8 (RB); Belo Horizonte, Labouriau 1013 (RB); Langsdorff s.n. (LE, P); Serra da Caraga, Santa Barbara, Leithao Filho et al. 9568 (BM); Serra do Sao José, Sao Joao del Rei, Lutz 45 (R); Serra do 1% Itatiaia, Ouro Preto, Magalhaes 1124 (RB); Serra Rola Moga, Betim, eee 17015 (MICH); Serra do Curral e Mutuca, Nova Lima, Ba 19496 (UB); Serra Rola Moga, 8 km from Belo Hori- zonte, 4300 ft, sole le 44619 (MICH, NY); 3 mi from Diamantina on roa ral to Curvelo, 4300 ft, Maguire 44770 (MICH, NY); Serra do Mutuca, Belo Horizonte, 1200 m, Markgvaf etal. 35 41 (F, RB); Serra do Taquaril, — Horizonte, Markgraf et al. 3585 (RB); Congonhas do Campo, Martius s.n. (M, rae ge of i. phlomoides): Serro Frio, Sepes, Martius s.n. (M); Cachoeira do Campo, Martius 1168 R, G, LE, M, NY); prope Sabara, Netto 80 (R); Serra da Moeda, 26 km de Belo Horizonte, 1300 m, Pubs 7417 ni Pico de Itabirito, 1400-1600 m, Palacios et al. 3853 (R); Serra do Curraly cae 2462 F Pabst 3298 (RB); Serra da Moeda, BR-3, Peretra 7525 (M); ad Oliveira, Pohl 358 (F, ",W); Ouro ee Porto 1259 (RB): Tejuco {Diamantina}, Riedel 1258 (LE, S, US); Riedel XIII ae S); Cachoeira do Ca impo, Riedel s.n. ae Penha, S¢. Hilaire s.n. (P, syntype of P. tomentosa); Serva da Carassa [Caraga], St. Hilaire s.n. (P, type of P. tomentosa); Serra d’Ouro Branco, St. Hilaire s.n. (P, syntype of P. tomentosa); St. Hilaire ns 1 1399] ( P, le as e of P. tomentosa B ladle: Sao Juliao, Schwacke s.n. (R 1957 9); Pico d’Itabira, Schuache yn. (R 72391); Serra do ee , mun. Jaboticatubas, 110, rodovia Lagoa Santa-Conceicao do Mato Dentro: Diamantina, Sazima & Semir 3843 (EF); Serra da Moeda, Sellow B1840 c1306 (NY, fragment of holotype of P. ere Ns 'B rotundifolia); Serra do Cipo, mun. Jaboucatubas, KM 112, rodovia Lagoa Santa- Ke pas do Mato Dentro-Diamantina, Semir & Sazima 564 (E); Serra do Pinheiro, nalts 21 (R, RB); Serra do el del Rei, 3500-5500 ft, Stephan s.n. (K); Belo Horizonte, Texeira s.n. (SP 18829); ae do Curral, mun. Belo Horizonte, Vidal s.n. oS 129614, R 129617); Serra do Ci rere Chapeéu de Sol, 1060 m, Vidal 11-6302, V-6 (R); Serra do Cipo, Fazenda do Cipo, Vidal V-75, V-91 (R); Weddell 1208 p.p. (G, P); vicinity of Lagoa Seca, 20 km 5S of Belo Horizonte, Williams 5458 (GH); mun. Betim, Serra da Cee. 1600 m, Willians wo Assis 6238 " GH); Serra do Mutuca, mun. Belo Horizonte, ca 8 km beyond Lagoa Seca, 1100 m, Willams & Assis 628 1 (GH ); Serra do Curral, mun. Nova Lima, 1300 m, Williams & Assis 6394 (GH); a Nova Lima, Lagoa Grande, 1300-1500 m, Meheay, Assis 6536 (GH, SP, US); mun. Nova ima, Pico de Belo Horizonte, 1300 m, Welliams & Assis 71 a car mun. Betim, Serra da Caveira, ta m, Williams & Assis 7494 (F, GH, MO, NY, S). = — — ¥ Peixotoa tomentosa is readily recognized by its inflorescence structure. Each terminal node bears three, commonly sessile, umbels and thus is 12-flowered. At the more distal nodes the secondary and tertiary axes are usually suppressed and one or two umbels are borne sessile or on secondary peduncles in each leaf axil. The young branches, the petioles, and the inflorescence axes are densely golden velutinous or velutinous-tomentulose. The leaves are golden velutinous above and usually densely white or greyish tomentose or woolly below. The hairs of the lower surface are often so abundant that the epider mis is obscured and only the larger veins and costa are visible. The leaves vary in shape from ovate to broadly or narrowly elliptical to obovate to suborbicular to rhombic or less commonly to lanceolate or oblanceolate. Plants from the Serra do Cipo, especially, tend to be more robust than others and are more likely to have suborbicular or broadly ovate laminas. However, this variation extends throughout the range and does not justly recognition of infraspecific taxa. Niedenzu also recognized a form, gracilior, tor plants with particularly long and slender pedicels, which are indeed striking. However, the size of the pedicel is highly variable in this and all other species. Three other species have a similar inflorescence structure, but none are as densely pubescent as P. tomentosa. P. adenopoda of east-central Bahia differs in that its leaf glands are stalked and borne on the surface of the lamina away from the base and costa, and in that its staminode glands have an indentation across the apex and decurrent on the abaxial face. In P. catarinensis, from coastal Santa Catarina, the leaf glands are also borne on the lamina but are very small, 0.2- 0.7 mm in diameter. In P. tomentosa the staminode glands are smooth. Che leaf glands are sessile, 0.8—2.5 mm in diameter, and borne at the base at the costa or halfway on the petiole or sometimes on the petiole just below the lamina. The northern range of P. parviflora coincides with the range of P. tomentosa. P. parviflora is probably a close relative and the species most likely to be confused with P. tomentosa. Its flowers, which are smaller than those of P. tomentosa, are el in that the posterior styles curve toward the anterior-lateral petals in- stead of toward the posterior- -lateral petals. The pubescence of the inflorescence 75 branches is commonly white, though sometimes golden, and much sparser. The inflorescence leaves are always petiolate, never sessile as is common in P. tomentosa, and the umbels are always borne on secondary peduncles. Collected in flower throughout the year, in fruit from November through April and in July. EXCLUDED AND DOUBTFUL SPECIES P. cordobensis Kuntze, Rev. Gen. 3(2): 28. 1898. Cordobia al (Grisebach) Niedenzu, Verz. Vorles. Ak. Braunsberg 1912-1913: 41. 1912. See section Relationships within the Family, P. microphylla Turczaninow, Bull. Soc. Nat. Moscow 36(1): 584. 1863. Accor ding to Turczaninow, the type of this species was collected by “Hoker” or “Loker” and was seen by him in Robert Brown’s herbarium. All attempts to locate this type have ae unsuccessful. The description is too vague to allow this name to be assigned to any known species of Petxotoa. Perhaps the type does not belong to this genus at all. The author does not mention such structures as the cordate stipules, the four-flowered umbels or the unique androecium. More- over, in his brief discussion, he does speak of pedicels that are bibracteolate in the middle, a condition unknown in Peixotoa. LITERATURE CITED Sate W. R., 1978. Byrsonimoideae, a new subfamily of the Malpighiaceae. Leandra 7: 5-18. 197, — , 1979. Floral conservatism in neotropical Malpighiaceae. Biotropica 11: 219-223, , L981. Malpighiaceae, Botany of the Guayana Highland. Mem. N.Y. Bot. Gard. 32: 21-305. Cuatrecasas, J., 1958. Prima Flora Colombiana. 2. Malpighiaceae. Webbia 13(2): 343-664. Gates, B., 1982. A monograph of Banisteriopsis and Diplopterys, Malpighiaceae. Flora Neotropica }_9s Monogt -aph 30: 37 Jussieu, A. de, 1832. Welpianeecre In Saint- le A. de, Flora Brasiliae Meridionalis 3: 5-86 (1832 , text) and 161-180 (1833, plate ae 1S. , 1840. Malpighiacearum synops isa 1 Sc . Nat. Sér. I, Bot. 13: gies and 321-3: , 1843. Monographie des el enneees Arch. Mus. Hist. Nat. Paris 3: 5-151 and ae 616 Mori, S. A, & L. A. Mattos Silva. 1979. : ae Herbarium of the “Centro de Pesquisas do Cacau” at Itabuna, Brill Brittonia 31: 177 Morton, C. V., 1968. A typification on some e subfamily, sectional, and subsectional names in the family Malpighiace ae. Taxon 17: 314-324. Cee, Bey LO. eas in Malpighiaceae americanae II. Verz. Vorles. Akad. Braunsberg W-S 912-1913: 334 —_—_—______. ae eee In Engler, A. Das Pflanzenreich IV. Vol. 141: 1-870. ee W. T., M. T. A. da Silva, & A. ie So ice 1981. Contribuigao ao estudo citologico de I Mpighincese Arq. Jard. Bot. 25: 169-17 Pohl, eae equipment ae naee for the study of minute plant structures. ne 67. 95-G Urban, [., 1895. Biographische Se III. 4. Jacques Samuel Blanchet (1807-1875). Engl. Botan. ahrb, 21, Beiblatt 52: Mase a 1974. Olblumen en siaaaenee Bienen. Tropische und subtropische Pflanzenwelt, Nr. Viesbaden. ACKNOWLEDGMENTS m grateful to W. R. Anderson. for his generous advice, his numerous valuable suggestions and comments, and for providing the chromosome counts. I also wish to thank Stuart R. Lowrie, who shared his insights pene thet nature of malpighiaceous pollen, and who supplied the SEM micrographs. aes Gates kindly permitted me to consult her then} fe ae monograph of Banisteriopsis. The handsome qigicuon were drawn by Karin Dot am also indebted to the curators of the following herbari ain provided access to their collections and arranged for loans: A and GH (Harvard), BM foneon BR (Brussels), C (Copen- hagen), CEPEC ae CTES (Corrientes), E (Edinburgh), F (Chicago), G (Geneva), GOET (Gottingen), IAN (Belem), K (Kew), LAM (Los Angeles), LE ees M (Munich), (Curitiba), MG (Belem), MO (St. Louis), NY (New York), P (Paris), R, RB, and RFA (Rio de Janeiro), S (Stockholm), SP (Sao Paulo), U (Utrecht), UB (Brasilia), US fe a W (Vienna). This work was supported by grant G-DEB-79-04946 from the National Science Foundation to the University of Michigan. The distribution maps in Figs. 3-11 are based on Goode Base Map No. 203, copyright the University of Chicago. > > INDEX TO NUMBERED COLLECTIONS CITED Archer, , & Gehrt, A., 140 P. hispidula. ae 232 P. parviflora. Allemao & oe 214 P. jussieuana Almeida, J., & Santos, T. S., 136 P. hispidula. ae pee 56, 57 P. hispidula. Anderson, W. R., 6201, 8090 P. goiana; 8364 P. spinensis; 8430 P. tomentosa; 8511 P. glabra; 8564 P. eeu 8623 P. barnebyi; 8730 P. spinensis; 9197 P. paludosa; 9359 P. magnifica; 9391 P. reticulata; 9474 P. magnifica; 9589 P. reticulata; 9831 P. magnifica; 9871 P. reticulata; 10024 P. golana; 10169 P. magnifica; 10222 P. leptoclada; 10260 P. reticulata; 10282 P. goiana; 11155 P. ea 11196 P. hispidula; 11537 P. barnebyi; 11549 P. glabra; 11591 P. andersonii; 11635, 650 P. eee, 11782, 11790 P. reticulata gee W. et al. 35028 P. magnifica; 35155 P. tomentosa; 35189 P. g abra; 36870 P. Andrade, A. G., 318 & Emmerich, M., 310 P. goiana. Araujo, D., 119. 124, 140 P. hispidu iG Araujo, D., 311 & Peixoto, A. L., 181 P. hispidula. Araujo, D., 690 & Peixoto, A. L., 492 P. hispidula. Arbo, M. M.. et al. 1750 P. retic nlaiae ata Atala, F., 2. Se P. cipoana; 80, 90 P. hispidula. Barclay, G. , 60 P. catarinensis Barreto, M., z 545 P, paludosa; 7 577, 7578 p.p. P. reticulata; 7578 p.p., 7622 P. tomentosa; 8681 P. paludos Belem, R. P., “3456 Ee seas Belem, R. P., & Mendes, J. M., 290 P. hispidula. Berg 243 P. hispidula. Black, G. A., & Magalhaes, M., 51-12228 P. parviflora. Blanchet, J. S 116, 1908 P. hispidula; 2867 P. parviflora. Bowie & C eA ae 214 P. parviflora. Brade, - ae 5949 P. parviflora; 11003, 11396 P. hispidula; 11822 P. tomentosa; 12014 P. hispidula; 13 » 13775 P. glabra; 13776 P. reticulata; 13777, 13781 P. glabra. Brade, A. C., & Altamiro 17802 P. reticulata; 17810 P. tomentosa. Brade, A. a 14819 & Barreto, M., 1335 P. reticulata. Brito, S., 100 P. reticulata. mea W. J., A882 P. parviflora; 5090, 5220, 7037 P. reticulata; 7084-2, 7133-2 P. magnifica; 7596 reticulata. en on P. hispidula. Campos P., SP18825 P. tomento Campos Novaes, J., SP12072 P. aes. Casaretto, G., 2711 P. tomentosa. Castellanos, A., et al. 25616 P. tomentosa. 77 C eae PB: 14.35. P. matinee 68A P. paludosa; 75A, 76A, 131, 131C, I41, 157, 197, 414 P. entosa; 522 p.p., >. leptoclada; 522 p.p. P. reticulata. Constantino, D., RB. 15430 a ee ae Cuatrecasas, J., & Duarte, A. P., 26636 P. hispidula. Damazio, L.. 929 Pp, tomentosa; 2054 P. cipoana. da Costa, B. M., R72044 P. paludosa. da Silva Manso, P., (L howky 99, 117) P. ae (Lhotzky 257) P. hispidula. Davidse, e ,eta i 37 Sa pauU aor 12208 P. bahiana. Davi se, » & Ramanoorhy . P., 10795 P. tomentosa. Pace E. es oe BR nae, de Santos, R. R., & Souza, R., 1637 P. reticulata. D’Orbigny es P. cordistipula. Duarte, A. 451 P. reticulata; 2206 P. pines 2843 P. reticulata; 3339 P. catarinensis; 3655 P. ae 3787 P. reticulata; 4173, 5966 P. hispidula. Dusén, P., 340a, 9225, 9619, 9690, 9949, 11685. 13032, 13082 P. parviflora. Edwall, Ge SPI ee P. reticulat Eiten, fete ,& Eit T., 1901, 9980, 2914, 2928, 3291 P. reticulata; 4835, 4917 P.jussieuana; 5616, P. ee 6837 P. tomentosa; 9014 P. magnifica; 11072 P. tomentosa. ° en, vs & Campos, J. B., 3449 P. reticulata. see G. F., 8 P. goiana. sien A,, 120 P. sericea. . M., 31 P. tomentosa. ae reira, M. B., 2a(UB37281 P. goiana; 2b(UB37618), 3, 4, 5 P. reticulata; 6 P. goiana; 7 P. reticu- lata; 8 P. goiana; 9 P. reticulata; 10, 11 P. goiana; 12, 13 P. reticulata; 32 P. goiana; 43 P. reticulata; 46, 47, 118, 150, 152 P. goiana; 215, 216, 217 P. magnifica. Fiebrig, K., 390, 4017, 4590, 5181 P. reticulata. Flaster 35 P. hispidula. Fonseca, S. G., 276 P. magnifica. Frazao "RB 11 ies P. hispidula. Fromm, E. R., et al. 1334 P. hispidu Gardner, G., 3067, 2498 P. jussieuana; coe P. gardneri; 3069 p.p. P. goiana; 3623 P. jussieuana. oe B. E., 348 P. parviflora. Gates, B. E., & Estabrook, G. F., 103 P. goia ere C., 584 P. reticulata aan 964 5 Taney yeh Glassman, S.F., & Gomes, Jr., J. C., 81 19 P. ees 8064 P. cipoana. Glaziou, A., 1328, 3891 P. hispidula; 5785, 8581 P. parviflora; 9680 P. jussieuana; 10371 P. parvi- flora; 11815, 11819 P. hispidula; 13598 P P. paludosa; 16731 P. tomentosa; 18929 p.p. P. beac 18929 p.p., 18930 P. parviflora; 18937 P. glabra; 20749 P. leptoclada; 20749a P. olan Gomes 1056 P. goiana. Gottsberger, I. S., 6571 P. reticulata Graziela [B Ar ocdl a P. ey RBI71697 P. spinensis. Guimaraes, a F., et al. 292 ue Harley, R. ee Sowa ae 1037 magnifica; 21561, 21964 P. paludosa. Harley, R. ‘c al. ae Bea 11432 P. goiana; 21561, 21964 P. paludosa. Hartmann, T. nee a reticulata Hassler, E., 1141, 2976, 4055, 4509, 5054, ! ae 6774, 8166, 8166a, 9881, 988 la, 10056 P. reticulata. Hatschbach, G., 1413 P. jussieuana; 2831, 5347, 6028 P. parviflora; 8819, 14243, 21572, 21770 P. reticulata; 28921 P. ahione "346 639 P. reticulata; 34713 Bi rdistipula; 35286 P. tomentosa; 36675 s Baa ee 36904 P. anadenanthera; 37700 P. SERCE 41240, 41394 P. spinensis; 4142 aludosa. Heringer, E. P., 8394 _ fe 8410 P. reticulata; 10295 P. parviflora; 11670, LI811, 14691 P. reticulata; 17258 P. leptoclada. Hoehne, F. C., 158 P. cordistipula; SP5052 P. parviflora; SP5117 P. tomentosa; SP5479, SP12905 P. parviflora; SPI9391, SP20525 P. reticulata; SP20569, SP23414 P. parviflora; SP30256 P. Hoehne, F. C., = Gehrt, A., SP35760 P. reticulata. Holway, E. W. D., & Holway, M. M., 1496 P. parviflora. Hunt, D. A Oo 5868 P. magnifica. 78 ae FH. §., et al. 8221, 10291, 106838, 10773 P. Boone 11103 P. reticulata; 11705, 12351, 12855, 5139 2: peers 14340a P. reticulata; 14889 P. bahiana; 15340 P. reticulata; 15997 P. magni- fica 16087 P. reticulata; 1668 La P. sre ea 1668 1b, 16484 P. reticulata; 16778 P. psilophylla; 16792 P. magnifica; 16860 P. Scio gaa 17176 P. cordistipula; 17439 P. psilophylla; 17494, 17554. 176517; pg nian 17562 P. goiana; 17830 P. reticulata; 18616 P. goiana; 18762 P reticulata; 19271 P. goiana; 19514, ae 36 P. tomentosa; 22091, 22294 P. reticulata; 2676. 22833 P. tomentosa; 23662 P. spinensis; 24985 P. goiana; 26641 P. reticulata; 27661, 28236a P. tomentosa; 28236b P. irwinii; 28236¢ P. spinensis; 28371, 28412 P. glabra; 28891, 28976, as P. tomentosa; 32799, 34061, 34096 P. golana; 34248 P. reticulata; 34743, 34744 P. gola Irwin, H. os ie Soderstrom, T. R., 5067 P. reticulata; 6083a P. goiana; 6117 P. reticulata; 7443 P. axillat Jesus,: J: a 92 P. adenopoda. Jobert 1045, Bie: P. jussieuana. Jobert & Schwacke 1053 P. jussieuana. Joly, A. B., & Semir, J., 3592 P. tomentosa. oly, A. B., et al. 1009, 1401 P. tomentosa. , 34372, 34542 P. reticulata Krapovickas, A., & Schinini, A., 32117, "3995 |, 32379 P. magnifica; ‘ 39918, 32893 P. reticulata; 36399 in <3] — 1) 4 — a = ~ = t4 % wn - Ym. 1 ms nH 7 = o ss i) oo —_ or 9 we Krapovickas, A., et al. 5 P. reticulata; 23197 P. hispidula. Kuhlmann, i; G., RBo6o7: 9 P. hispidula; RB150098 P. tomentosa; 6221 P. hispidula. Kuhlmann, J. G., & Edmundo 8 P. tomentosa. Kuhlmann, M., 1192, 2758 P. parviflora; 4144 P. os Labourtiau, L., 60 P. oe 1013 P. tomento Leitao Filho, H. F., et al. 3164 P. pi einige Q! 568. 9673 P. tomentosa. Lima. As etal. dP; reticulata: 5 P. gol Lima, A. S., 6268 P. reticulata Lindman, C. A. 1 Bi eo i cordistipula. Lindeman, J. C., 6327 P. hispidt ula. Lindeman, J. C., & Haas, J. H., 3131 P. parviflora. Lisboa, A., 2! 506 P. jussieuana. Lofgren, a 272 P. parviflora; 587 P. reticulata. Lutz, B., R72142 P. reticulata; R129613 P. catarinensis; 45 P. tomentosa; 653 P. hispidula. Pen. P. von, 1599, 1721 P. jussieuana. Maas, P. J. M., & Carauta, P., 3129 P. hispidula. Macédo, A., 23 P. goiana; 38, 2428 P. reticulata; 3671 P. goiana; 4962 P. reticulata. aes ene B., 44619, 44770 P. tomentosa; 49144 IP. ap ere 49285 P. spine nsis Malme, G. O. A., 1610 : pall ts eae IG 10x P. cake 1919 P. cordistipula. Markgraf, | ee Brade, A. C., 3047, 3788 P. hispidula. Markgraf, F., et al. 3541, ee PB: tomentosa. Martin, D., 407 P. reticulata. Martin, R., 476 P. goiana. Martius, C. F. P., 656 P. cordistipula; 1168 P. tomentosa; 1169 P. hispidula; 1343 P. glabra; 1756 P. hirta. Matos, M. E. R., 92 P. reticulata. Mattos, A., & Rizzini, C. S., a P. reuculata, Mattos, J. R., & Mattos, N. F., 8305 P. reticulata. Medeiros, D., 15 P. sre Mello, O., RB67422 P. jussieua Mello Filho, L. E., 830, 980, OBi "1199 P. hispidula. Mendonga, F. R., 976 P. leptoclada. Mexia, Y., 5798 P. reticulata; 5804 P. glabra. Miquel, J., el al. 1 P. hispidula, Moreira, A. X., & Angosto, fe R72 184 P. hispidula. Moreira, H., & Moreira, I., 495 P. catarinensis. Moseén, C. W. H., 4050 P. reticulata. Netto, L., 80 P. tomentosa; 88 P. hispidula. Netto, L., et al. R19558 P. hispidula. 79 Occhioni, P., RB26440 P. hispidula; 23 P. goiana; 650, 1153, 3554, 3675, 4313 P. hispidula; 5009, 5054 P. cipoana; ee P. reticulata; 6389, 7226 P. hispidula. Onishi, E., et al. 74 P. culata Pabst, G. F. J., 7117 P. tomentosa; 10321 P. parviflora. Pedersen, T. M., 11116 P. reticulata. Pereira, E., 55, 536, 576 P. hispidula; 1535 P. glabra; 3569 P. hispidula; 7525 P. tomentosa; 10525 P. his ae dula. Pereira, E., & Pabst, G. F. J., 3298 P. tomentosa. Philcox, D, & Freeman, B., 4696 P. cordistipula. Pimentel, T., & Barbosa, E. V., 15 P. hispicula. Pinheiro, . S., 1870 P. Le KOE e El Pires, J. M., 57925 58097 P. reticulat Pires, J. M., et al. oe P. reticulata; ‘9709 P. magnifica. Plowman, T., 9943 P. goiana. Pohl, J. E., 358 P. tomentosa; 472-2598(1498d) P. reticulata; 1739.3555(1559d) P. magnifica; 2946(1563d) P. leptoclada; 3556(d 1557) P. parviflora. Porto, C., 1259 P. tomentosa. Porto, C., ie aera 2200 P. reticulata. Prance, G. T., et al. 19239 P. oa Prance, G. i fe Schaller, = ee P. reticulat Prance, G. , & Silva, N OO P. goiana; ree P. magnifica. Puttemans, ie SPI2118 P. ioe: Saag C. 893 P. reticulata. 3 P. catarinens Ramamoorthy T. P., 618 P. aeninees Ramariz, D., 114 P. tomentosa. Rangel R197 e puspieula. Ratter, J. A., 2 p.p. P. psilophylla; R112 p.p. P. magnifica. aes aes & ee, J., 370 P. magnifica Ratter, J. A., et al. 1990 P. reticulata; 2806, 2899, 3129 P. goianz RC encl A. F., [-29 P. parviflora; [[-28, [1-28a, H-28b, II-28d, II- 98e, I[-28g, I1-28x P. reticulata; III-118, Te 318, HI-318x, 111-1652 P. parviflora. Reiss, R., 65a P. parviflora. Reitz, R., 3240, 5083 P. catarinensis. Reitz, R., & Klein, R. M., 2752, 6779 P. catarinensis. Rente 2, 238 P. hispidulz Richards, P. W., 6429 P. aalopaa Riedel, L., III, XII p.p. P. parviflora; XII p.p. Bees XUI P. glabra; 584 P. reticulata; 617 P. hispiduta 1013 P. See cs 1156 P. glabra; 1258 P. tomentosa. Riedel, L., & Luschnath, B., I P. parviflora; VII, on P. hispidula; 2007 P. reticulata. Rizzo, ‘~ "3997, 4128, 4442, 4494, 4597, 4638 P. goiana; 9819 P. magnifica. Rodrigues 92 P. eae: Rosa, F. R., 67 P. goiana. Roth, L., . ee "i ace 1591 P. reticulata. Salzmann, P., 93 I idula. 5 stl 34 = aaa Ae Cre 5086, 5087, 5088 P. hispidul Sazima, M., & Semir, J., 3843 P. tomentosa. Schaller, G., 89 P. magnifica. Schinini, A., 4348, 8809, 8887, 8956 P. reticulata. Schott 3677 P. ee =} ay o 5 oe Schwacke, C. A. 7130 P. hispidula; 8370 P. glabra; R19578 P. leptoclada; R19579, R72391 P. tome ee RED) 155 P. reticulata. si a t. [a. 1814] P. parviflora; III. it. B1840 c1306 P. tomentosa; IIT. it. 1310, TV. it. 5505 P. ie sei L. & Saz M., 564 P. tomentosa. Sick, K., B481 P. ‘silo Silva, M. G., & Maria, J., 3193 P. ea a Silva Novaes SP 18839 P. parviflorz Silveira, A., 21 P. tomentosa. Smith, L. a 6367, 6570 P. ees Smith, L. B., & Reitz, R., 6085, 12276 P. catarinensis. 80 Smith, L. B., et al., 14834 P. parviflora Sucre, D 274 P. reticulata; 3853, 4291 Pp. hispidula; 7560 P. parviflora; 10416 P. reticulata; 106 14 P. parviflora. Swallen, J. R., 9606 P. reticulat Teixeira, A. R., SP18826 P. parvittors SP18829 P. tomentosa. Trinta, Z. A., & Fromm, E. 1618, 2042, 2042.4 P. hispidula. Trinta, Z. A., et al. 1386 P. roa. ie Ule, E. H. G., 711 P. catarinensis; 4706 P. hispidula; 7446 P. jussieuana. Vauthier 459 P. glabra. Veloso, H., 1235, 1 1 P. cordistipula. Vianna, M. OF 15, 35, 537 P. hispidula. Vidal, ie 2054 P. rare R3929 1 P. reticulata; R72121 P. glabra; see R72400 P. parviflora; R108249 P. ere R129614, R129617 P. tomentosa; I-780, I-800, 1-836 P. reticulata; II- 6302, V-6, V-75, V-91 P. tomentosa; V-100 P. hispidula. Wawra & Maly 142 P. hispidula. Weddell, H. A., 1208 Williams, L. O., 5458 P. tomentosa. b.p. P. leptoclada; 1208 p.p. P. tomentosa. O., & Assis, V., 5827 P. reticulata; 6238, 6281, 6394 P. tomentosa; 6490 P. paludosa; 536 P. tomentosa; 6619 P. parviflora; 6938 P. reticulata; 7147 P. tomentosa; 7231 P. paludosa; ore P. tomentosa. INDEX TO SCIENTIFIC NAMES italics. Bz anisteriopsis Rob. in Small 2-3, 6, 9 Banisteriopsis malifolia (Nees & Martius) Gates var. malifolia 57 C. argentea (Griseb.) Nied. ae Bea Griseb, 2-3 Adr. Juss. 11-12 P, fete C. Anderson 3, 5, 6, 9, 11, 13 17-19, 25, 28, 29, 60, 68, 74, . P. amadenanthera G; eee pare 7, 12, 14, 18, , 82 ’ —20, 33, : Pe sndenann . ae 3, 7, 11, 14, 16, 18, 20-21, 83 P. axillaris C. Anderson 6, 10, LL, 14, 18, 21- , 44, 84 Pe Balan C. Anderson 15, 23-25, 83 P. barnebyi C. Anderson 15, 16, 17, O4, 25-26, 85 P. catarinensis C. Anderson 5, 6, 9, 13, 18, 24, 8, 60, 74, 86 P. cipoana C. Anderson 5, 11, 15, 24, 26, 28— 30, P. opel Adr. Juss. 5, 10, 16, 30-32, 33, 49, 52, 66, 88 P. cordobensis O. el 1, 75 P. discolor Griseb. P. gardneri C. ne 3, 15, 20, 32-33, 41, 82 . glabra Adr. Juss. 3, 9, 13, 34-35, — 65, 86 . golana 7 Pani 8, 10, 13, 15, 17, 33, 35-38, 4 8 ie grandiftora See 62 P. achiu C. Anderson 3, 6, 11, 12, 13, 19, 38-40, 49, 62, 84 P. a Adr. Juss. 3, 5, 15, 20, 33, 40, 41-42, EG ee oe f. 1. minor Nied. P. hirta f. 2 macrophylla eas Nied. P. hispidula Adr. Juss. 6, 9, 11, 13, 16, 03, 28, 35, 40, 42-44, 53, 68, 86 P. hispidula f. 1. eiionenios Nied. 42 Accepted taxa are in Roman type; the main entry for each is in bold face. Synonyms are in P. hispidula : 2 subglabrata Mae P. haspidula { 3. micrantha Nied. P. irwini C., An deresn 8, 12, 13. 35, a 4445, 83 r; jusvstate: Adr. Juss. 5, 8, 9, 12, 16, 46-47, a0, 81 51, 52, P. Jussteuana var. O pect ee 47 P. jussieuana var. B velutina Nied. 46 P. lateritia Adr. Juss. 62 ie pare nn Juss. 5, 10, 14, 16, 47-49, P. naropt ne 62 P. macrophylla f. 1. minor (Nied.) Nied. 62 P: macrophylla f. II. Diind | (Griseb.) ie 62 P. aoe lifica C. our 8, 9, 10, 12, 16, 32 49-52, , 88 Pp. Re peice a Anderson 15, 51, 52-53, 90 P. microphylla Vure P. octoflora C. eee 6, 10, L1, 13, 54455, 59, 90 P. paludosa Turez. 5, 9, 10, 14, 16, 25, 49, 51, , 84 , or P. parviflora Adie ae - 8, 9, 12 18, 21, 26, 28, 57-60, 74, P. parviflora var. B prince Nied, 55, 57 P. phlomoides Adr. Juss. 71 P. a C. Anderson 3, 13, 40, 60-62, , 90 P. ee Griseb. 6, 9, 10, 12, 13, 14, 32, 35, 38, 53, 62-67, 91, 92 P. sericea C. Anderson 3, 5, 9, 11, 14, 25, 29, 66, 68-69, 81 P. spinensis C. Anderson 5, 16, 26, 29, 56, 69- 71, 85 P. tomentosa Adr: Juss. 6, 14,18, 21, 26, 28, 29, 60, 71-75, 87 ; 6 le f. cuneata Nied. 71 P. tomentosa £. 3. gracilior Nied. 71 pine ap roe ca Adr. Juss. 71 >. tomentosa 8 ie ee Griseb. 71 Cas Stigmaphyllon Adr. Jus a eS ute c Dp 5 “ft “, x RON Jac: hay, > or pans ye x 2 SS a PAN loon mane’ ae PINs Teen” t yor DET IQ : “pea OS . EN J FIG, a P. eae P. sericea and P. di lee a—e, P. adenopoda: a) ELS Rae au stem leaf, x 0.5, detail x 2 ) leaf gland, x 5; c) base of a distal umbel, x 2.5; d) flower, x 2.5 posterior- er (left) ane enon (right) s ee nee x 5. £ ] i 0.5, detail X 2.5; g) base of a distal umbel, x 2.5; S flower, x 2.5. i-l, P P. jussieuana: 1) and stem leaf, x a detail x 2.5; j) flower, x 2.5; k) anterior (left) and posterior- lateral (right) staminode glands, x 10; 1) samara, x 1. a—e from Jesus 392, fh from Rane 120, i-k om Eitten & Eiten 4835, | from Lutzelburg 2 82 . 13. P. a alee and P: leaf, x ie 5, detail 2 gardnert. a-e, P. pe aaa a) flowering branch and stem e of a distal umbel, * 1.5; c) f ; 1D: ees view of anthe x 7.5; e)anterior (lett) Bete te lateral (right) ae gla f-j, P. gardnert: f) stem leat. x 0.5, detail x 2.5; g) base ¢ Ifa on il umbel, * 1.5; h) axes, all pedicels shed, ~ 25 te dearpe ae ont bases of secondary x 1.5; ]) an 93 1) flov j) anterior (left) and posterior-lateral (right) stami- node glands, x 7.5. a-e from Aces. 36904 fj from Gardner 3069 p.p. FIG. 14. P. bahiana, P. andersonti and P. irwinit. a—-e, P. bahiana: a) flowering branch, x 0.5; b) base of a distal umbel, 5: a ee (above) and _ anterior- ne eral (betow w) petals, x 3; d) posterior- -lateral (left) and anterio , x 5;e) gyn i f-i, ee andersoni: f) flowering fea and stem leaf, x 0. 5; g) base of a distal umbel, x 1.5; h) flower, x i ; 1) posterior-lateral (left) — Sate (right) staminodes, x 5. j+m, P. irwinuw: j) stem leaf. x 0.5; k) base of a iy tal umbel, 5; 1) flower, x 1.5; m) anterior (above) and posterior-lateral (below) renee 5. a-e aa ee et al. 14889, f-i from Anderson 11591, j-m from Irwin et al. 82366. 20° IG. 15. P. » paludosa, P. hatschbachu ee P. axillaris. a-e, a ae soe a) yeas: branch and m leaf, x 0.5; b) base of a distal umbel, .5; c) flower, x d) anterior (left) and posterior- ee ight) staminode glands, x 5; e) samara, X 0.75. f-k, A suas f) flowering branch and stem leaf, x 0.5, detail x 1.5; g) base or stem leaf, x 2.5; h) fl x o i) Se lateral (above) and anterior acing) staminodes: x 7.5; j) samara, X 1; she embryo, x ca. 5. ep, P. axillaris: |) flower- ing shoot and stem ei x Bute m) flower, < 1; n) ant vr stamin ode land, x 75: 0) posterior- lateral staminode gland, 5; p) samara, X l.a ee branch from Williams & Assis 6940, leaf from Barreto 8681, b-d an Williams & Assis 6940, e from Warming s.n., f-k from Hatschbach 36675, l-p from Irwin & Soderstrom 7443 = 85 FIG. 16. P. spinensis, P. ee and P. pane a-d, P. spinensis: a) flowering branch and stem leaf, x 0.5; b) Base of a distal umbel, x 2.5; c) flower, x 1; d) posterior- coe (left) and anterior (right) staminode Cee x ie 5. e-h, P. leploclad a: e) flowering branch and stem leaf, x 0.5; f) base of a distal umbel, = 2.5; g) flower, x 1; h) posterior-lateral ee, and anterior caene staminode glands, x 7.5. i-l, P. barnebyi: i i) flowering branch and stem leaf, x 0.5; j) base of a distal umbel, x 2.5; k) flower, X 1; 1) posterior-lateral (left) and anterior oe yee eras x 7.5. a-d from Anderson 0, e-h from Pohl 2946, i-l from Anderson 862 86 way os ah oe P. glabra and P. hispidula. a—-e, P. catarinensis: a) flowering aes and joe stem oe x 0.5, - 2.5; b) base of a distal umbel, x 1.5; c) androecium and gynoecium, claw o compu abe tal, 3B: d) anterior (left) and posenon lateral (right) staminodes, x 7.5; e) samara, ; a Pla p) flowering branch, x 0.5, detail x 2.5: g) base of a distal ie x 1.5; h) flower, ysterior-lateral ene! ae AU SISE rare staminode glands, x 7.5; j) samara, X 1. k-o, 2 oe k) stem leaf, x 0.5, detail x 2.5: 1) flow ng bran x 0.5: m) ae of posterior petal, androecium anes out, WU I se lor- ie a 1 sienan ee stamen and | anterior-lateral staminode removed, x 3.5: n) anterior (above) and posterior-lateral (below) staminode glands, 7.5: 0) Samara, X ee abi wan from Smith & Bee 12276, leaf from Reitz 32: Reitz 12276, f-1 from Irwin et al. 28371, j from Irwin et al. 28412, k-n from Harley etal. 18510, 0 from Araujo 140. WAMU see Muy a $ ter . 18. P. cipoana, P. parviflora and P. tomentosa. a—c, P. cipoana: a) flowering branch and stem ee x 0. 5; b) flower, x 1.5; c) anterior (above) and poster Ne -lateral (below) staminode glands, x 10. rela He 7 povenns pee and stem leaf, x 0.5; e) base of a distal umbel, x 1.5; ee petal, 5; g) androecium and gynoecium, with posterior styles to left, x 7.5; h) anterior (above) and poser Steral (below) one glands, x 10; 1) samara, X 0.75. j-n, P. tomentosa: }) flowering branch, x 0.5; k) base of a distal umbel, x 25 5: 1) Ronen x 1.5; m) So ea neaity (left) and anterior (1 ight) penioode glands, x 10; n) samara, x 0.75. a from Atala 62, b, c from Vidal R1I08249, d flowering branch from Dusén 9225, leaf from Widgren s.n., e from ee 10371, f-h from Sucre 7560, i trom Widgren s.n., j pe Langsdorff s.n., k from Irwin eval 28236a, 1,m, from iten & Eiten 11072, n from Maguire 4477 tty oy, “! fote f, es P. magnifica and P. cordistipula. a-t, P. ee a) flowering branch and stem leaf, x 0.5 on ie e of a distal umbel, x 1.5; c) posterior — .5; d) androecium and gynoecium, claw of posterior 05 x 3; : acted of anterior style, x 15; f) samara, X 0.75. g—k, P. cordistipula: g) flow ae branch and s eaf, x 0.5; h) base of a distal umbel, x 1.5; 1) posterior petal, ye oD androecium ea an nee claw of posterior petal, x 3; k) samara, x 0.75. a, b from fre 9831, c—-e from Anderson 9474, f from sails etal. hee g branch from Lindman A3533, leaf from ene, 1919, h-j from Lindman A3533, k from Kuntze s. IG. 20. P. ee and P. hirta. a-h, P. goiana: a) flowering branch and stem leaf, * 0.5; b) base ofa Aietall umbel, 9; c) flower, X 1.5; d) posterior petal, x 1.5; e) anterior (above) and posterior- lateral (below) staminode glands x 10; fy Caer anterior style to left, x 5; g, Z samara, ile i-m, P. hirta: 1) stem leaf, x 0.5; j) base of stem leaf, x 2.5; k) base of a distal nae 2.5; 1) flower, x 1.5; m) ae a ibov e han yeaa ern (below) aaa x 10. a branch coe Irwin et al. Ge leaf from Irwin et al. , bec, e-f from /rwin et al. 18616, d from ee 6201, g from Lima et al. 5, h from /[rwin et a aan i-m ae Nie sn, Rs Whur 7) > & NMA gy y & ¥ UA, = Su hE WODLOTAOL | TF ~~ ry ae / Sn RPM, x t spare by ah 3 | 7 / d Z, ty ae mn & BdAyrhi c ° ys ayaa AR rm 7 4 R ayupLuny si f ™ a Wud, . 44 oly : Ka ya OM axe ostaylon, P. eee and oe psilophylla. a-d, P. octoflora: a) flowering branch and stem ven x 0.5; b) base of dist ul umbel (with 2 ae removed), X 2; c) flower, x 1 (left) and posterior- eel (right) eee ae x 7.5. e-h, P. megalantha: e) flowering bran a and stem leaf, ib: ase ef a distal umbel, eo g) flower, X A; h) anterior staminode gland, x 7.5. i-l, < PCED 1) flowering branch and stem leaf, x 0.5 base of a distal umbel, x 2; k) ee. aes Oe anteric : — ower, 5; 1) anterior (above) and sis mats (below) seminede glands, x from Irwin et A 16681a, e-h from oo et pee J, 1 gee from Irwin et al. 16778, leaf from Sick B+ 6860. SI, Jj trom Irwin et al. 16778, k, | from ie win et oa I G pevintegare w WA ws vA a a ald UAL yo Adiga y Gn eink ee v4 FIG. 22. P. reticulata. a-d, from east-central Mato Grosso: a) stem leaf, x 0.5; b) wee = a distal umbel, x 2; c) flower, x 1; d) anterior (left) and posterior-lateral (right) ee e—h, from Minas Gerais: e) flowering branch and stem leaf, x 0.5; f) flower, x g) posterior- ee (left) and anterior (right) staminode glands, x 5; h) samara, x 0.75; i-m, ee Sao Paulo: 1) flower- ing branch and stem leaf, x 0.5; j) flower x 1.5; k) anterior (left) and posterior-lateral (right) staminode glands, x 5; 1) samara on torus, 0.75; m) embryo, x 3.5. a-d from /rwin et al. 16681b, e branch from Mexia 5798, leaf from Brade 14819, f-h from Mexia 5798, i from Eiten & Eiten 3291, j,k from Eiten & EFiten 2914, 1m, from Fiten & Eiten 2928. 92 Why \t ose vat De Ms, vy FIG. 23. P. reticulata. a-f, from Paraguay: a) flowering branch and stem leaf, x 0.5; b) base of a distal umbel, x 2.5; ¢) posterior Riera x 2 35 a) androecium (posterior- -lateral stamen and anterior- le wteral staminode and stamen removed), x 5; €) gynoecium, anterior style to right, x 5; f) samara, US: ae Bet sous Mee: Can ee owen ing shoot and stem leaf, x 0.5; h) base of a distal cael 1) ie x 1; j) posterior-lateral (left) and anterior (right) scimnode glands, x 5; k) anterior cz fee 1s samara, X 0.75. a branch from Hassler 8166, leaf from Hassler 4055, b-e from Hassler 8166, c aes Hassler 988 1a, g-l from Hatschbach 24625 Contr. Univ. Mich. Herb. 15:93-136. 1982. NOTES ON NEOTROPICAL MALPIGHIACEAE—I William R. Anderson Herbarium The University of Michigan Ann Arbor, Michigan 48109 My current studies of American Malpighiaceae include both monographic and floristic projects, as well as routine identification of many specimens sent as gifts. In the course of this work I regularly encounter taxa new to science, situations in need of nomenclatural adjustment, and material that supplements our previous knowledge, such as flowering material of a species hitherto known only in fruit. It will be many years before I can complete and publish revisions for large difficult genera like Byrsonima and Heteropterys. These notes will serve as a vehicle for the earlier publication of miscellaneous descriptions and other observations as they accumulate. BYRSONIMA The species treated below are presented in approximately the order in which I believe they would have appeared in Niedenzu’s monograph of the family (1928). This is done in order to facilitate their comparison and integration with that treatment. Byrsonima hatschbachii W. R. Anderson, sp. nov. Fig. 1. Suffrutex caulibus 23-40 cm altis plerumque non ramosis ex xylopodio enascentibus. Lamina foliorum majorum 11.5—16.5 cm longa, 1.4—1.6 cm lata, lineari-elliptica; petiolus 2-3 mm longus; stipulae 2—2.7 mm longae, basi con- natae distaliter liberae. Inflorescentia bracteis bracteolisque pemtennbus, Petala lutea; petalum posticum ungue eglanduloso. Antherae 2-2.5 mm longae, sparsim sericeae praecipue inter loculos, connectivo loculos + aequanti. Ovari- um glabrun oa subshrub, with stems 23-40 cm tall, faely sericeous to glabrate, unbranched or nearly so, all arising from a xylopodium. Lamina of the larger leaves 11.5-16.5cm long, 1.4-1.6cm wide, linear-elliptical, very gradually tapered to an acute or acuminate apex and attenuate base, thinly sericeous to glabrate, with the short white hairs longest persistent on the adaxial midrib, the lateral veins prominulous on both sides; petiole 2-3 mm long, not clearly distin- guishable from the lamina, sericeous to glabrate; stipules 2—2.7 mm long, con- nate only in the proximal third, the free portions triangular or acuminate, abaxially sericeous, adaxially glabrous. Inflorescence 9-11 cm long, finely sericeous with the hairs brown to gray, the flowers borne | per bract or (especi- ally proximally) in compact cincinni of 2—3 flowers; bracts 2-4 mm long, 0.6— 1 mm wide, subulate or narrowly triangular, thinly sericeous to glabrate, stiff; primary floriferous peduncle (from bract to joint) 0-3 mm long, developed only at lower nodes; bracteoles like the bracts but shorter; bracts and bracteoles 93 94. FIG, eel oe a affinis. a-f, B. hatschbachu: a) habit, x 0.5; b) ee es supules, adaxial y c) 2-flowered cincinnus, X 2.5; d) posterior petal, x 3.5; e) stamens 7,93 f) facet aen cal ve ata away from posterior petal, x 7.5. gl, B. affinis: @) flowering branch, x 0.5 ie abaxial lamina x 2.5 in as h) stipules, x 1.5 be eaelets Spa x 1.5; )) stamens, X 5; yhnoecium, with styles bending away from posterior petal, x 5; 1) fruit, x 0.5. Drawn by De Douthit, a—f from Pieeabnch 37593, gk from. Neen 11274, oe Anderson 9518. 95 persistent at least past flowering. Pedicel 4.5-6 mm long in flower, sericeous, circinate in bud, decurved in old flowers. Flower 12-13 mm in diameter. Sepals all biglandular, ca 1.5-2 mm long beyond the glands, 1.5 mm wide, obtuse, abaxially sericeous in center, ciliate on margin, adaxially glabrous, recurved at the apex in anthesis; glands 1-2.5 mm long. Petals yellow, glabrous; lateral petals reflexed, with the claw ca 2.5 mm long, the limb 4.5-5 mm long, 5-6 mm wide; posterior petal with the claw 2.5 mm long, erect, eglandular, the limb 4 mm long and wide, reflexed. Filaments 1.9 mm long, abaxially glabrous, adaxially hirsute at the base; anthers 2—2.5 mm long, sparsely sericeous, especially between the locules, the locules unwinged, often free at the x, the connective not or hardly exceeding the locules. Ovary 1.3 mm high, anes all 3 locules fertile; styles 3 mm long, distally slightly bent away from the posterior petal. Fruit un- known. TYPE: BRAZIL. Mato Grosso: Mun. Chapada dos Guimaraes, Capao Seco, campo cerrado, borda das depressoes umidas, 15 Nov 1975 fl, Hatschbach 37593 (MBM, holotype; MICH, isotype). Byrsonima hatschbachu is named in honor of Gert Hatschbach, a fine botanist whose “olho vivo” seldom misses a novel plant. The species seems to be most closely related to B. intermedia Adr. Juss., from which it differs in its short, nearly unbranched stems, long narrow leaves, nearly distinct stipules, and persistent bracts and bracteoles. B. intermedia is rather variable and as presently defined may well comprise more than one species. However, even in its broadest inter- pretation it could not accommodate Hatschbach 37 593, nor have I seen any speci- mens intermediate between the two species. Byrsonima morii W. R. Anderson, sp. nov. Fig. 7 Frutex 1.5-2 m altus. Lamina foliorum majorum (5—) 6-8.2 cm longa, (3-) 4—6.1 cm lata, late elliptica vel parum ovata obovatave vel fere orbicularis, coriacea, subtus pertinaciter sericea vel appresso-tomentosa vel glabrescens, reticulo plerumque stramineo et prominenti, praecipue in pagina adaxiali; petiolus (3—-) 5-12 (-15) mm longus; stipulae (1.5-) 2-3 mm longae, omnino connatae. Inflorescentia (4—) 6-9 (-13) cm longa, floribus in dimidio distali congests, bracteis bracteolisque demum deciduis. Petala lateralia alba, petalum posticum flavum. Antherae 2—2.9 mm longae, glabrae vel eeeeae sericeae inter loculos, connectivo loculos + aequanti. Ovarium glabru Shrub 1.5—2 m tall, the stems sericeous, the short ae brown and persistent in the first season of growth, turning white and eventually deciduous in later seasons. Lamina of the larger leaves (5—-) 6-8.2 cm long, (3—-) 4-6.1 cm wide, broadly elliptical or slightly ovate or obovate to nearly orbicular, obtuse, rounded, or emarginate at the apex, slightly revolute at the margin, cuneate or truncate and shortly decurrent at the base, coriaceous, often glaucous above, sericeous or appressed-tomentose, persistently so or early or eventually glabrescent, the hairs above rather twisted and soon turning white, the hairs below nearly straight to serpentine or moderately twisted, reddish or sometimes eventually turning white, the lateral veins and reticulum usually yellowish and prominent, especially above; petiole (3—) 5-12 (-15) mm long, subsericeous to glabrescent; stipules (1.5-) 2-3 mm long, rounded, smoothly and completely connate, abaxially sericeous to glabrate, adaxially glabrous. Inflorescence (4-) 6-9 (—13) cm long, with flowers mostly densely clustered in the distal half, sub- sericeous proximally to nearly velutinous distally, the flowers borne mostly | per bract, occasionally in 2 _flowered cincinni; bracts 1.5-3 mm long, 0.7—1.4 mm wide, triangular, stiff or reflexed apically, abaxially hairy, adaxially glabrous; peduncle 0-1 (—3) mm long; bracteoles like the bracts but smaller; bracts and 96 bracteoles epee through anthesis, mostly deciduous before maturation of the fruit. Pe 7-12 mm long (—15 mm in fruit), reddish-tomentose, circinate in bud, pence and ype twisted in fruit. Flower 12-15 mm in diameter. Sepals all biglandular, 1.5-2.5 mm long beyond the glands, 1.8-2.5 mm wide, obtuse or rounded at the apex, abaxially loosely but densely sericeous, adaxially glabrous or sparsely tomentose distally, appressed but recurved at the apex in anthesis, accrescent (to 2-3 times as large) in fruit; glands 2-3 mm long. Petals glabrous, the lateral 4 white, the posterior bright yellow turning red-orange in age; lateral petals strongly reflexed, with the claw 3-3.5 mm long, the limb 4.3-6.5 mm long, 5.5-8 mm wide; posterior petal erect, with the claw 3-3.5 mm long, the limb 3.5 mm long, 4 mm wide, corrugated. Filaments 2—2.5 mm long, abaxially glabrous, adaxially hirsute at the base; anthers 2-2.9 mm _ long, glabrous or with a few long straight hairs between the locules, the locules un- winged, sometimes free at the apex, the connective not or hardly exceeding the locules (up to 0.3 mm). Ovary 1.4-1.7 mm tall, glabrous, all 3 locules fertile; styles 3-4 mm long, straight or bent at the apex. Mature fruit not seen. YPE: BRAZIL. Bahia: Mun. Mucuge (=Mucuje), 3km S de Mucuje, campo rupestre, 1000 m, 22 Dec 1979 fl, Mort & Benton 13158 (CEPEC, holo- type; MICH, isotype). PARATYPES: BRAZIL: Bahia: 10-15 km N of Mucuge on road to Andarat, sandstone hillside with shallow soils and large bluffs of exposed rock, 1100 m, Feb fl, Harley 18864 (MICH); Mun. Mucuje, 3 km S de Mucujé, campo rupestre, 1000 m, Jul fl, Mori et al. 12609 (MICH, US); Mun. Mucujeé, 10-12 km W de Mucuje, campo rupestre, 1000 m, Jul fl, Mori et al. 12667 (MICH, US); Mun. Lengois, 15 km NW of Lencois, campo rupestre, 900 m, Jun fl, Mort & Boom 14282 (MICH), Jun fr, Mort & Boom 14283 (MICH); Mun. Lengois, 5 km N of Lengois, campo rupestre, Jun fl, Mori & Boom 14402 (MICH). cs When I first saw this species, I identified it (Harley 18864) as “Byrsonima bumelufolia Adr. Juss. vel aff.” However, since seeing the type of B. bumeliifolia in Paris | have changed my opinion. That type has very dense, tightly twisted, persistent hairs on the abaxial side of the lamina, as in B. variabilis Adr. Juss., and it seems likely that those two species are closely related, perhaps even con- specific. B. mori is more probably related to B. brachybotrya Nied., a species of Parana and Santa Catarina. They have similar flowers crowded in short pseudo- racemes, although these are longer and bear more flowers in B. morii. B. morii has the lamina much larger, especially wider (two or three times as wide) and borne on a longer petiole. The two also differ dramatically in their habit. B. morii is a shrub 1.5-2 m tall, while B. brachybotrya is a subshrub with slender branches 15-50 cm high from a woody, more or less subterranean, base. Byrsonima moru honors Scott A. Mori, a fine botanist whose collections from southern Bahia comprise a major contribution to our knowledge of that flora. Byrsonima onishiana W. R. Anderson, sp. nov. Fig. 2 Suffrutex caulibus erectis usque ad 30 cm altis. Lamina foliorum majorum 10-17 cm longa, 1.5-3.5 cm lata, anguste oblonga vel obovata, apice acuta vel obtusa, basi attenuata; petiolus 5-12 mm longus; stipulae 2-3 mm_ longae, liberae vel usque ad % connatae. Inflorescentia bracteis bracteolisque persistenti- bus. Petala pallide rosea. Antherae 2.1-2.7 mm longae, glabrae vel sparsim sericeae inter loculos, connectivo loculos + aequanti. Ovarium glabrum. Fructus —11 mm diametro. Subshrub, the leafy stems erect, slender, up to 30cm tall, arising from a thicker prostrate (7) underground (7) stem, loosely sericeous to glabrate. Lamina of the larger leaves 10-17 cm long, 1.5-3.5 cm wide, narrowly oblong or obo- 97 G. 2. OC aee onishiana and B. spinensis. a-f, B. onishiana: a) habit, including underground base, x 0.5; b >) stipu ules, x ; ae c) flower, x 2.5; d) stamens, x 7.5; e) gynoecium, w an styles bending away oe eee petal, x 5; f) fruit, drawn from behind to show enlarged sepals and persistent petals, x 2. gl, B. spinensis: g) pope branch, x 5h) stipules, X 5; 1) cincinnus, xX 2.5; j) stamens, X 7.5; k) gynoecium, x 5; 1) ee: fruit, x 2 pene by Karin Douthit, a-f from Jrwin et al. 25824, g-l from Anderson et 7 3533 98 vate, acute or obtuse at the apex, very gradually attenuate at the base, thinly appressed-tomentose to glabrescent, the lateral veins and reticulum prominu- lous on both sides but more strongly so above; petiole 5-12 mm long, distin- guishable only with difficulty from the decurrent lamina, appressed-tomentose to glabrate, swollen at the base; stipules 2-3 mm long, triangular or ovate, dis- tinct or connate for up to % of their length, abaxially loosely sericeous, adaxially glabrous. Inflorescence 3-13 cm long, loosely sericeous, the flowers borne 1 (-2) per bract; bracts 2-3 mm long (-8 mm at lowest flowers), I-1.3 mm wide, nar- rowly triangular to subulate, stiff and ascending or spreading or somewhat re- curved; peduncle none; bracteoles like the bracts but smaller; bracts and bracteoles persistent past maturity of the fruit. Pedicel 5-10 mm long or longer in fruit, loosely sericeous, circinate in bud, decurved and eventually twisted in fruit. Flower ca 14 mm in diameter. Sepals all biglandular, 1.5-2.5 mm long beyond the glands, 1.7-2.3 mm wide, obtuse at the apex, abaxially loosely sericeous, adaxially glabrous, recurved at the apex in anthesis, accrescent (to ca 3 times as large) in fruit; glands 1.7-2.3mm long. Petals “very pale pink,” glabrous; lateral petals reflexed, with the claw 2-3 mm long, the limb 5-5.5 mm long, 5.5-7 mm wide; posterior petal erect, with the claw 2.5 mm long, the limb 3.7mm long, 4.5 mm wide, corrugated. Filaments 2-2.5 mm long, abaxially glabrous, adaxially hirsute at the base; anthers 2.1-2.7 mm long, glabrous or with a few long, straight hairs on inner faces of locules, the locules unwinged, often free at the gig the connective not or hardly exceeding the locules (up to 0.2 mm). Ovary 1.2-1.6 mm high, glabrous, all 3 locules fertile; styles 3.6— 3.8 mm long, straight or bent away from posterior petal at apex. Fruit 9-11 mm in diameter, glabrous; petals sometimes persistent nearly to maturity of fruit. TYPE: BR AZIL. Minas Gerais: Serra dos Oculos, campo, ca 60 km NE of Patrocinio, 1050 m, | Feb 1970 fl/fr, Irwin et al. 25824 (UB, holotype; MICH, NY, isotypes). PARATYPE: BRAZIL. Verusa (without further locality data), Oct fl, Heringer LO7O1 (UB). Byrsonima onishiana is named in honor of Eunice Onishi, Brazilian botanist and one of the collectors of the type. It is most closely related to B. oxyphylla Adr. Juss., which has a similar habit and generally similar flowers. In B. oxyphylla the leaves are smaller (seldom over 6 cm long and 1.5 cm wide) and more numerous on each stem; its inflorescence is shorter and more congested, often almost umbellate; its petals are white (the lateral four) and yellow (the posterior one, turning red in age); and the drupe is smaller, 5-6 mm in diameter. Byrsonima cacaophila W. R. Anderson, sp. nov. Pig. Arbor 10-20 m alta. Lamina foliorum majorum 13-24 cm longa, 5.5-10 cm lata, glabrata vel in costa pertinaciter sericea; petiolus 12-32 mm longus; stipulae 3.5-6.5 mm longae, omnino connatae vel distaliter distinctae. Inflorescentia cincinnis 1—3-floris, bracteis plerumque 3-5.5 mm longis, subulatis, reflexis, pedunculo primario (2-) 4-10 mm one bracteolis 1-2 mm _ longis, ovatis, bracteis bracteolisque eee any Sepala adaxialiter glabra. Petala alba, aetate rubescentia. Antherae 2.2-2.9 mm longae, sericeae praecipue inter loculos pilis fusiformibus 0.4-0.5 mm longis, connectivo loculos + aequanti. Ovarium glabrum. Fructus (siccus) 1] mm longus, 15-18 mm diametro, depressus, nuce costata. Tree 10-20 m tall, the trunk 25-35 cm in diameter, the stems sericeous, eventually glabrescent. Lamina of the larger leaves 13-24 cm long, 5.5-10 cm wide, acute or obtuse to rounded and often abruptly short-acuminate at the apex, slightly revolute at the margin, cuneate at the base, glabrate at maturity or 99 FIG. 3. eNOS ed ne Be B. bahiana. a-f, B. cacaophila: a) flowering branch, x 0.5; b) stipules 2; c) cincinnus, : stamens, X 7.5; e) gynoecium, with < mete (ied) from above, x I. x; 1) bract and br acteoles on inflorescence axis, X 5; j) immature Fruit with enlarged sepals and Baoan petals, x 2; ruit from above, ai 3. Drawn by Karin Douthit, a-e from Belem & Pinheiro 2881, £ from en Magathaes 637, ae ae Harley 18475 100 persistently sericeous on the midrib or rarely on the lateral veins below, the lateral veins prominent, especially below, the reticulum prominulous on both sides; petiole 12-32 mm long, sericeous to glabrate; stipules 3.5-6.5 mm long, triangular, completely connate or free distally, smooth or abaxially sulcate, abaxially sericeous to glabrescent, adaxially glabrous. Inflorescence (8-) 11— 19cm long, sericeous proximally to loosely sericeous to velutinous distally: cincinni with 1-3 flowers; bracts mostly 3-5.5 mm long, shorter at apex of in- florescence, 1-1.5 mm wide, subulate, mostly revolute at apex or reflexed or twisted, abaxially sericeous, adaxially glabrous; primary floriferous peduncle (from bract to first joint) (2-) 4-10 mm long; bracteoles 1-2 mm long, I-1.5 mm wide, ovate, abaxially sericeous, adaxially glabrous, borne at or up to 2mm below apex of peduncle; bracts and bracteoles persistent past maturity of the fruit. Pedicel 5-7 mm long, loosely sericeous to velutinous, slightly circinate in bud, apparently straight in fruit. Flower 17 mm in diameter. Sepals all biglandu- lar, 2 mm long beyond the glands, 2-3 mm wide, acute or obtuse at the apex and revolute in anthesis, abaxially sericeous, adaxially glabrous; glands 3-4 mm long. Petals white, turning reddish in age, glabrous; lateral petals reflexed, with the claw 2.5-3 mm long, the limb 5-6 mm long, 6-7 mm wide, concave; posterior petal erect, the claw 3-3.8 mm long, eglandular, the limb 44.5 mm long, 5- 6 mm wide, corrugated. Filaments 2.5—-3 mm long, abaxially glabrous or sparsely hirsute at base, adaxially hirsute at base; anthers yellow, 2.2-2.9 mm long, the locules linear, unwinged, sericeous, especially on inner faces, with the fusiform hairs only 0.3-0.5 mm long, the connective equalling the locules or occasionally exceeding them by up to 0.2 mm. Ovary 1.5 mm high, conical, sulcate, glabrous, all 3 locules fertile; styles 3.5-4 mm long. Fruit (dried) 11mm long, 15- 18 mm in diameter, glabrous, depressed, the nut with prominent bony ribs. TYPE: BRAZIL. Bahia: Rod. Urucuca a Taboquinha, plantagao de cacau, 22 Nov 1971 fl, T. S. Santos 2200 (CEPEC, holotype; MICH, isotype). PARATYPES: BRAZIL. Bahia: Centro de Pesquisas do Cacau, CEPLAC, Ilheus, Apr fr, Belém & Magalhaes 637 (CEPEC, IAN, NY, UB): Itapebr plantagao de cacau, Nov fl, Belém & Pinheiro 2881 (CEPEC, NY, UB); Itajuipe, plantagao de cacau, Nov fl, Belém & Pinheiro 2918 (CEPEC, NY, UB); Itabuna, Estrada da Burarema, mata de cacau, Nov fl, Mello Filho 2902/Emmerich 3440 (CEPEC, R); Ilhéus, CEPEC, plantagao de cacau, Dec fl, Santos 530 (CEPEC, MICH). [am not sure where this interesting species fits best in the taxonomy of the genus. [t resembles B. maguirei Anderson in its stipules, inflorescence, anther- shape, and sulcate ovary, but differs in having reflexed bracts, white petals, hairy anthers, and all three carpels fertile. The reflexed subulate bracts suggest the complex that includes B. crispa Adr. Juss., but B. cacaophila differs from those species in its long peduncles, white petals, persistent bracts and bracteoles, and short fusiform anther-hairs. The epithet cacaophila draws attention to the fact that all kown collections of this species come from cacao plantations, including those from the CEPEC re- serve, which was founded on an old plantation. When the native forests were cut many years ago to plant Theobroma, some native trees were left to shade the cacao trees. Byrsonima cacaophila seems to be one of those regularly spared; the species may owe its persistence to this practice. Many other undescribed species of those wonderful forests probably disappeared into extinction at that ume. According to the label with Santos 2200, this species has the common name “eis 101 Byrsonima schunkei W. R. Anderson, sp. nov. Fig. 4. eas eee Adr. Jussieu var. velutina Niedenzu in Engler, Pflanzenr. IV. 715. 1928. Lectotype: Tessmann 3450 (NY!). Frutex vel arbor 2-10 m alta. Lamina foliorum majorum 20-29 cm longa, 8-14 cm lata, apice acuminata, supra mox glabrata praeter costam plerumque velutinam, subtus dense et pertinaciter velutina; petiolus (16— 2023 lenam longus, velutinus; stipulae 4-8 (—10) mm longae, connatae, apice rotundatae. Inflorescentia bracteis 3-6 (-10) mm longis, 1.5-2.5 mm latis, cincinnis 1—4- floris, pedunculis primariis 3-15 mm longis, bracteolis 1.5—4 (—7) mm longis, 5-3 (-5) mm latis, sub apice pedunculi portatis. Sepala eee appresso- tomentosa. Petala lutea. Antherae 2.0-3.5 mm longae, laxe sericeae praecipue inter loculos, connectivo loculos aequanti vel usque 0.5 mm superanti et apice ut videtur glanduloso. Ovarium sericeum Shrub or tree 2-10 m tall; stems velutinous, eventually glabrescent. Lamina of the larger leaves 20-29 cm long, 8-14 cm wide, ellipical or broadly elliptical or somewhat obovate, cuneate at the base, slightly revolute at the margin, acumi- nate (sometimes aruptly so) at the apex, soon glabrate above except usually persistently velutinous on the midrib and sometimes on the lateral veins, densely and persistently velutinous below, the hairs 0.25—0.50 mm long, erect, bifurcate, the branches as long as or shorter than the stalk; midrib and lateral veins prominent below; petiole (16—) 20-31 mm long, velutinous like the lamina with a proximal admixture of some longer, sub-basifixed hairs; stipules 4—8 (—10) mm long, connate, the pair rounded at the apex, abaxially loosely sericeous, adaxially glabrous. Inflorescence 6-24 cm long, velutinous; cincinni comprising 1-4 flowers, often better developed proximally than distally in the same inflores- cence; bracts 3-6 (-10) mm long, 1.5-2.5 mm wide, subulate, lanceolate, or ovate, stiffly spreading or somewhat flexuous distally, abaxially sericeous, adaxially glabrous; primary floriferous peduncle (from bract to first joint) 3— 15 mm long; bracteoles 1.5—4 (-7) mm long, 1.5—3 (—5) mm wide, ovate, borne at various heights on peduncle from just above base to somewhat below apex; bracts and bracteoles deciduous during or after anthesis, all gone before matu- ration of the fruit. Pedicel 4-8 mm long, velutinous, circinate in bud, decurved or twisted in fruit. Flower ca 13 mm in diameter. Sepals all biglandular, 2-3 mm long beyond the glands, rounded at the apex, revolute, appressed-tomentose, goualy so abaxially, densely to sparsely so adaxially, accrescent in fruit; glands 54.5 mm long. Petals yellow, glabrous, the outermost covering all others in me lateral petals reflexed, with the claw 2.5-3 mm es the limb 5.5- 6.5 mm long, 6-8 mm wide, concave, especially in the anterior pair; posteror petal erect, the claw 3-4 mm long and thicker, eglandular, ae limb 3-4 mm long, 3.5-4.5mm _ wide, corrugated. Filaments 2.0-2.5 mm _ long, adaxially hirsute at very base with hairs only ca 0.5 mm long; anthers 2.0-3.5 mm long, loosely sericeous, especially between the locules, the connective equalling the locules or exceeding them by up to 0.5 mm and apparently glandular at the apex, the locules narrowly linear, separated on the connective, free at the apex. Ovary |.3-1.6 mm high, conical, sericeous, at least on the distal half, all 3 locules fertile; styles 3-3.5 mm long, straight or slightly bent at the apex. Fruit yellow, 13-15 mm long, 11-13 mm in diameter (dried), ovoid with an attenuate apex, glabrate or distally thinly sericeous. TYPE: PERU. San Martin: Prov. Mariscal Caceres, Dtto. Tocache Nuevo, Apr fl, Schunke V. 3929 (MICH, holotype). PARATYPES: PERU. San Martin: Mariscal Caceres, Tocache Nuevo, Mar fr, Gentry et al. 25569 (MICH), Jan fr, Schunke V. 3699 (MICH), Nov fl, Schunke V. 5589 (MICH), Aug tr, Schunke V. 6859 (MICH, MO), Jul fl, Schunke V. 7332 102 hIG. es alee schunkei and B. surinamensis. a-f, B. schunkei: a) flowering branch and large leaf, x 0.5, pr ur from abaxial side of lamina x fics b) supules, x 1.5; c) cincinnus, x 2.5; d) BRAGS, with ee ane short % anthers from same flower, x 5: e) gynoecium, with styles bent away from posterior petal, * 5; f) fruit, x 1. g-l, B. surinamensis: 8) foverine branch, x 0.5, with leaf-tip 5; h) supules, x 3.5; 1) flower bud with bracts ee Bracteo les, x 2.5; j) stamens, with long and short anthers from same ae er, X 7.5; k) gynoecium, * 7.5; 1) aes sae from behind to show enlarged sepals and twisted pedicel, x 1.5. Drawn by Karin Dorie a-e from Schunke 7332, f from Maas et al. P12974, g—k from Lindeman et a 597, | trom Mort & Bolten 8437 103 (MO, US); Uchiza, Pampa Yacu, alt. 450 m, Jan fl/imm. fr, Schunke V. 5750 (F, LA). Loreto: Yarina cocha, middle Ucayali, Dec fl, Tessmann 3450 (NY). Psa Villa Isabel, rio Cuchara, Sep fr, Schunke V. 5676 (F, LA). BRAZIL. Acre: Mpio. Rio Branco, highway to Porto Velho, Albuquerque et al. 1279 (MICH): ); Porangaba, Rio Jurua-mirim, May fr, Maas et al. P12974 (MICH, NY); km 45 Rod. Rio Branco-Porto Velho, Feb imm. fr, Santos et al. 53 (MICH). Amazonas: Antimary, Rio Acre, Mar fr, Huber s.n. (MG 4251). BOLIVIA. Pando: Cobya, Dec fl, Ule 9488 (MG). Byrsonima schunke? is named in honor of Jose Schunke Vigo, who has made several beautiful collections of it. The species grows in tall forest or, more often, in secondary forest or thickets, always on terra firma. Schunke has reported the following common names for the species in Peru: “Indana,” “Indano,” “Indano Colorado,” and “Sacha Uvilla.” He also records the following uses in folk- medicine: “La corteza la utilizan los nativos para curarse de tuberculosis, bebiendo en cocimiento” (3929). “A quarter of a glass of a maceration of the bark is drunk for dysentery” (5750). Niedenzu described this species as B. poeppigiana var. velutina, but his con- cept of the taxon was vague and his syntypes included at least one collection referable to a third species, B. linguifera. | am selecting a lectotype for Niedenzu’s name here, but nevertheless I feel it would be best to give the taxon a distinct epithet at the level of species. Byrsonima schunkei belongs to a complex of four species native to western Amazonia, which are closely related but easily distin- guished. The following key will facilitate their recognition: 1. Leaves sericeous to glabrate below, the hairs sessile, straight, appressed. B. arthropoda Adr. Juss. 1. Leaves thinly to densely xe eee al the persistent hairs Y-shaped with a straight, erect stalk, the branches mostly s r than the stalk 2. Leaves strongly revolute pe peer ete velutinous above over eae surface o lamina. inguin Cuatr. Leaves flat or only slightly revolute at the margin, soon glabrate above or pe velutinous on the midrib. 3. Floriferous pecunele 0.3—2 mm long (—3 mm in fruit); bracteoles borne at apex of peduncle, 0.8—1.5 (-2) mm long, | mm wide; larger leaves with the lamina 10- me cm long and 4.5—9 cm wide, the petiole 6-15 (-18) mm long; sepals adaxially glab B. Pee Adr. Juss. Floriferous eran 3-15 mm long; bracteoles borne below apex of peduncle, 1.5—4 (-7) mm long, 1.5-3 (-5) mm wide; larger leaves with the lamina 20-29 cm long a and —14 cm wide, a petiole (16—) 20-31 mm long; sepals adaxially aa to der appressed-tomentose. : can eee Anderson No oO Byrsonima affinis W. R. Anderson, sp. nov. Bigeel: Arbor (3—) 4-8 (—10) m alta, trunco 30 cm diametro, torto, suberoso. Lamina follorum undulata, margine revoluta, basi auriculata, sessilis, subtus arcte ne reticulo subtus prominent. Inflorescentia bracteis 3-6 mm longis, 2.5-3.5 mm latis, anguste tr iangularibus, revolutis et/vel reflexis, bracteolis 1-2.5 mm iis bracteis bracteolisque deciduis. Petala lutea; petalum posticum ungue eglanduloso. Antherae 2.5-3.5 mm longae, loculis glabris, connectivo extus plerumque laxe sericeo, inter loculos semper sericeo, loculos non vel usque ad 0.4 mm superanti. Ovarium lanatum. Fructus usque ad 3.7 cm diametro, luteus, nuce 12 mm alta, 18 mm lata, trilobata, basi excavata. Tree (3—) 4-8 (-10) m tall; trunk up to 30 cm (or more?) in diameter, often twisted, with thick, fissured, corky bark; young stems tomentose with admixture of spreading hairs, soon glabrescent. Lamina of the larger leaves | 1-20 cm long, 104 4—9 (—10) cm wide, elliptical or slighty obovate, acute, obtuse, or rounded at the apex, undulate, revolute at the margin, shallowly to deeply auriculate at the base, sessile, glabrate above except tomentose on the midrib, glaucous and smooth or somewhat rugose above, tightly and persistently woolly below, the hairs light brown or white, the lateral veins and reticulum very prominent below; petiole none; stipules 4—6 mm long, smoothly and completely connate or distinct at the apex, acute, abaxially tomentose or loosely sericeous, adaxially glabrous. In- florescence 12-24 cm long, erect in flower and pendent in fruit, densely and persistently tomentose, the flowers borne | per bract or in clusters (compact cincinni) of 2 (—37) flowers; bracts 3-6 mm long, 2.5-3.5 mm wide, narrowly triangular, abaxially tomentose, adaxially glabrous, mostly revolute and/or re- flexed; peduncle 0; bracteoles smaller than the bracts, especially shorter (1- 2.5mm long), revolute or not; bracts and bracteoles deciduous, sometimes during anthesis, often later. Pedicel 8-15 mm long, thickened and slightly elongated in fruit, densely spreading-tomentose or velutinous, straight or slightly circinate in bud, often somewhat decurved after anthesis but apparently + straight in mature fruit. Flower 18-22 mm in diameter. Sepals all biglandular, 2-3.5 mm long beyond the glands, 2.5—-3.5 mm wide, obtuse or acute at the apex, abaxially densely appressed-tomentose, adaxially glabrous or distally tomentose, recurved at the apex in anthesis, accrescent (to ca twice as large) in fruit; glands 2-3.5 mm long, yellow. Petals yellow, turning red-orange in age, glabrous; lateral petals reflexed, with the claw 2.5-3.5 (+4) mm long, the limb 6-9 mm long, 7-Ll mm wide; posterior petal with the claw 3-4.3 mm long, erect, eglandular, the limb 4—5.5 mm long, 5.5-7 mm wide, corrugated, reflexed. Fila- ments 2-3 mm long, abaxially glabrous, adaxially hirsute at the base; anthers 2.5-3.5 mm long, longer opposite sepals than petals, the locules glabrous, un- winged, often slightly free at the apex, the connective usually loosely sericeous outside locules and always sericeous between locules, often somewhat enlarged and flattened at the apex but not or hardly exceeding the locules (up to 0.4 mm). Ovary 1.8—2 mm high, densely woolly, all 3 locules fertile; styles 2.5-3.9 mm long, straight or bent away from the posterior petal. Fruit globose, to 3.7 cm in diameter, glabrate at maturity or persistently tomentose at the apex, yellow at maturity with thick juicy flesh, the nut 12 mm high and 18 mm wide, 3-lobed, deeply excavated at the base, leaving behind a rounded torus which fits into the excavation. TYPE: BRAZIL. Mato Grosso: Mun. Rio Verde, Serra da Pimenteira, 25— 35 km SW of town of Rio Verde, cerrado on sandy soil, 8 Feb 1975 fl/imm. fr, Anderson 11274 (MBM, holotype; MICH, isotype). ARATYPES: BRAZIL. Mato Grosso: Mun. Alto Araguaia, vic. of Riberao Claro, NW of Alto Araguaia, 800 m, cerrado, Feb fl, Anderson 11404 (MICH): Cuiaba-Vilhena, Km 206, 56°28’ W, 14°11’ S, 550 m, campo cerrado, Mar fr, Bamps 5498 (MICH); 12°54’ S, 51°52’ W, Xavantina-Sao Felix road, cerrado on red sandy soil, R. R. de Santos et al. R.1244 (MO). Goias: 12 km S of Caiaponia, 840 m, cerrado, Apr fl/fr, Anderson 9518 (MICH, NY, UB); Mun. Rio Verde, open woodland: Jan fl/imm. fr, Pedersen 12136 (MICH). Minas Gerais: 70 km $ of Uberlandia, cerrado, Feb fl, Anderson 12394 (MBM, MICH); Mun. Prata, cerrado, May fr (MICH); 9km S of Frutal, cerrado, Mar fl, Hatschbach 38223 (MICH); 4km N of Patrocinio, 1000 m, cerrado, Jan fl, Irwin et al. 25736 (MICH, NY); Mun. Frutal, sandy grassland with scattered trees, Jan fl, Pedersen 12072 (MICH); Uberaba, Regnell IIT 302 (MO). Byrsonima affinis is closely related to B. basiloba Adr. Juss., a shrubby species which it resembles in herbarium specimens, which undoubtedly explains why it has not been named previously. The two species can be separated by the char- acters in the following couplet: 105 Tree (3-) 4-8 (-10) m tall, the trunk with thick, corky bark; lamina not only revolute at the margin but He undulate; anthers with the connective usually loosely sericeous outside locules and always sericeous between them; inflorescence mien in fruit; fruit oe 3 cm in diameter. 3. affinis. Shrub 0.3-1.5 (-2.5) m tall, much-branched from base, the stems not corky; lamina ee al the margin but otherwise flat; anthers glabrous; inflorescence erect in fruit; fruit up 2 cm in diamete1 B, beer In addition to these differences, B. basiloba differs from B. affinis in the following tendencies: Its bracts are seldom revolute and mostly earlier decidu- ous; the anthers are generally shorter, probably seldom exceeding 2.5 mm in length; and the ovary is usually loosely sericeous instead of woolly, with the hairs straighter and more nearly parallel. The distribution of B. affins and B. basiloba is shown in Figure 5. For the most part the two species occur in different areas, but they seem to come very close together in the Triangulo Mineiro, south of Uberlandia. They should be studied in that area, to see whether they hybridize; perhaps they differ in eco- logical requirements that effectively separate them even there. The few collec- tions of B. basiloba from the area of Vilhena, Rondonia, are disjunct from the rest of the species. They are somewhat atypical, especially in having smaller leaves ae flowers, but they could not be assigned to any other species, certainly not to affinis. Further perce may reveal more populations of B. basiloba in ee across north-central Mato Grosso. Byrsonima fonsecae W. R. Anderson, sp. nov. Fig. 6 Arbor 5 m alta. Lamina foliorum majorum 10.5—15.5 cm longa, 5.5-7.5 cm lata, obovata vel paene elliptica, apice plerumque rotundata, subtus parce FIG. 5. Distribution of ie caffe squares) and B. basiloba (circles). Goode Base Map 203, C pede The University of Chicag 106 appresso-tomentosa vel subsericea; petiolus 14-20 mm longus; stipulae 4-6 mm longae, omnino connatae vel apice distinctae. Inflorescentia bracteis 1.5 2.5 mm longis, 1.5-2 mm latis, persistentibus, pedunculis 1.5—-6 (-10) mm longis, bracteolis bracteis similaribus vel minoribus. Pedicellus circinatus in alabastro, decurvatus in fructu. Sepala utrinque tomentosa, valde appressa per anthesin. Petala alba, demum rosea. Antherae glabrae, loculis cylindricis, 1.4—1.6 mm longis, connectivo 0.3-0.9 mm superatis. Fructus 12-13 mm diametro (siccus). Tree 5 m tall, the stems thinly sericeous to glabrate. Lamina of the larger leaves 10.5-15.5 cm long, 5.5-7.5 cm wide, obovate or nearly elliptical, cuneate at the base, revolute at the margin, mostly rounded at the apex, sometimes emarginate, sometimes broadly obtuse, coriaceous and somewhat rugose above, soon glabrate above or loosely sericeous on the midrib and lateral veins, thinly appressed-tomentose or subsericeous below with some hairs straight and ap- pressed and others twisted and spreading, eventually glabrescent, the lateral veins and reticulum visible above, prominent below; petiole 14-20 mm long, loosely sericeous to glabrate; stipules 4-6 mm long, completely connate or free just at the apex, abaxially sericeous to glabrate, adaxially glabrous. Inflorescence 4-12 cm long, loosely sericeous or tomentose to subvelutinous, the cincinni mostly |-flowered, a few 2- or 3-flowered; bracts 1.5-2.5 mm wide, lanceolate or ovate, abaxially appressed-tomentose, adaxially glabrous, all or most per- sistent past maturity of the fruit; peduncle 1.5—6 (-10) mm long; bracteoles like the bracts or somewhat smaller, apical or | slightly subapical. Pedicel 7-8.5 mm long (somewhat longer in fruit), appressed-tomentose, circinate in bud, de- curved and eventually twisted in fruit. Flower 13 mm in diameter. Sepals all biglandular, 1.5-2 mm long beyond the glands, 2-2.5 mm wide, obtuse or rounded, reddish-tomentose on both sides, strongly appressed in anthesis, some- what accrescent in fruit (to ca 4mm long and 5 mm wide); glands 2.5-3 mm long, pink. Petals glabrous, white, their claws becoming red or pink in age, the posterior petal with a yellow claw, the whole petal becoming red in age; lateral petals spreading, with the claw 3-3.5 mm long, the limb 4.5-5.5 mm long, 6— 7 mm wide, concave, especially in the anterior pair; posterior petal erect, the claw 3 mm long and thicker, the limb 3.5—4.5 mm long, 4 mm wide, corrugated. Filaments 2-2.7 mm long, abaxially nearly glabrous, adaxially long-hirsute, especially on proximal '; anthers 1.7-2.4 mm long, glabrous, the locules 1.4— 1.5 mm long, cylindrical, separated on the connective, rounded and sometimes slightly detached at the apex, the connective yellow, extended 0.3-0.9 mm be- yond the locules, the extension rounded or obtuse and somewhat recurved. Ovary ca 1.5 mm high, ovoid, glabrous, all 3 locules fertile; styles 3-3.5 mm long, white, all bent distally away from the posterior petal. Fruit 12-14 mm in di- ameter (dried), green, depressed-globose, short-beaked, glabrous, the nut prominently ribbed 7 — PE: BRAZIL. Minas Gerais: Serra do Cipo, ca 1225 m elev, in woods near stream, 18 Feb 1972 fl/fr, Anderson et al. 36183 (UB, holotype; MICH, NY, isotypes). Byrsonima fonsecae is named in honor of Sidney Geraldo da Fonséca, plant collector, mechanic, and friend, whose diverse talents were critical to the success of many expeditions in central Brazil led by Howard Irwin, myself, and others. It seems to be most similar to Byrsonima vernicosa Niedenzu, a rare plant known only from near Petropolis outside Rio de Janeiro, and B. nervosa DC., a species common in the Planalto of Brazil. All three have similar anthers, large fruits, pedunculate flowers, and large, petiolate, obovate, coriaceous leaves. However, B. vernicosa and B. nervosa have the leaves very densely and persistently tomen- tose below, whereas in B. fonsecae the lamina is only thinly hairy below at matu- 107 rity. Glaziou stated that B. vernicosa has yellow petals, but this requires confirma- tion, because all other species with anthers of this type have the petals white/ ink pin Byrsonima souzae W. R. Anderson, sp. nov. Fig. 6. Arbor 15 m alta. Lamina foliorum majorum 16-23.5 cm longa, 10.5-14 cm lata; petiolus 32-60 mm longus; stipulae 3-6 mm longae, omnino connatae. In- florescentia floribus in cymulis 3-floris dispositis, bracteis 1-2 mm_longis latisque, post maturitate fructus persistentibus, pedunculo 2-10 mm_ longo, bracteolis bracteis similibus vel minoribus, persistentibus. Petala rubra, glabra. Antherae glabrae, connectivo glanduloso loculos 1.2 mm superanti, conoideo, Ovarium sericeum; styli 2 mm longi, recti. Fructus 21-25 mm diametro, |0- 13 mm altus; nux disciformis. Tree 15 m tall, the trunk 13 cm in diameter; stems densely sericeous or appressed-tomentose with short, somewhat twisted hairs, eventually glabrescent. Lamina of the larger leaves 16—23.5 cm long, 10.5-14 cm wide, obovate or broadly elliptical, cuneate or rounded and tten slightly decurrent at the base, rounded at the apex and often abruptly short-acumuinate with the acumen up to 1 cm long, glabrate at maturity or sparsely appressed-tomentose on the midrib, especially below, the veins and reticulum prominent on both sides; petiole 32- 60 mm long, sericeous like the stems to glabrate; stipules 3-6 mm long, smoothly and completely connate, rounded at the apex, abaxially appressed-tomentose to glabrate, adaxially glabrous. Inflorescence 15-18 cm long, appressed-tomentose like the stem, composed of short cymules of at least 3 flowers, possibly more; bracts 1-2 mm long and wide, broadly ovate, abaxially tomentose, adaxially glabrous; primary floriferous peduncle (from bract to joint) 2-10 mm long, longest at base of inflorescence and diminishing distally; peduncles of lateral branches of cymule apparently not developing; bracteoles like the bracts or smaller, the first bracteoles often borne below apex of peduncle; bracts and bracteoles persistent past maturity of fruit. Pedicel ca 5 mm long in flower, probably longer in fruit, tomentose, somewhat circinate in bud, perhaps straight in fruit. Flower ca 16mm in diameter. Sepals all biglandular, ca 2 mm long beyond the glands, 2.5mm wide, obtuse or rounded, abaxially appressed- tomentose, adaxially glabrous, strongly appressed in anthesis, wider but not or hardly longer in fruit: glands 3-4 mm long, white. Petals red, glabrous; lateral petals reflexed, with the claw 2-3 mm long, the limb 4—6 mm long, 5-6 mm wide; posterior petal erect or somewhat reflexed, the claw 2.5 mm long, the limb 3 mm long and wide. Filaments 2 mm long, abaxially glabrous, adaxially hirsute at the base; anthers 2.1-2.4 mm long, glabrous, the locules 0.9-1.2 mm long, unwinged, tapered distally to an acute apex, the connective enlarged and glandular, 1.7-1.8 mm long, extending ca 1.2 mm beyond the locules, tapered distally to a point, straight or somewhat recurved. Ovary 1.5 mm high, densely sericeous, all 3 locules fertile; styles 2 mm long, straight. Fruit 21-25 mm in diameter and 10-13 mm high, transversely depressed-obtrulloid, with the thin orange flesh following the contours of a disc-shaped nut, glabrate or sparsely sericeous distally. RAZIL. TYPE: Amazonas: Manaus, Km 45 da BR 17, t. firme, solo argiloso, mata virgem, 3 May 1961 fl, Rodrigues & Coelho 2462 (NPA. holotype; NY, SP, isotypes). PARATYPE: Amazonas: Manaus, Estr. Torquato Tapajos, Km 105, lado direito, solo argiloso, Sep fr, Rodrigues & Loureiro 7168 (INPA, NY). his species is named in honor of my late friend, Raimundo Souza, a fine field botanist whose participation was a great asset to scores of expeditions led by . 6. Byrsonima fonsecae and B. souzae. a-h, B. ee a) flowering branch, x 0.5, with nla mar portion of abaxial side of lamina x 2; ae supu x 3.5; c) portion of inflorescence with Sea ate flower-bud, x 2.5 35 aa) flower, « 2.5; e, f) ene x 10: g) Ryne ne styles bent away from posterior pet tal, 9; h) fruit, * 1.5.1 B. souzae: 1) (sue es and axis of in nflore escence, al oo x 09%*)) stipules, x 1.5; are FAA OStE ucted ee x 2.5; 1) apex of peduncle to show bracteoles of oe x 7.5;m,n) stamens, X 10; 0) gynoecium, x oop) ne uit, with flesh and base in longitudinal section to expose the nut, x 0.75. Drawn by Karin Douthit, a-h from Anderson et al. 36183, i-o from Rodrigues & Coelho 2462, p from Rodrigues & Loureiro 7168. 109 both Brazilians and foreigners in all parts of Brazil. B. souzae is probably most closely related to B. christianeae Anderson, to which it is quite similar in most features, including the anthers and gynoecium. They differ in the inflorescence and fruit, both of which are unique in B. souzae. Its flowers are borne in stalked, dichasial cymules, with both bracteoles fertile, instead of one-sided, helicoid cymules (cincinni) as in other Malpighiaceae with compound inflorescences. | cannot tell from the material at hand whether more than three flowers develop in older cymules. B. christianeae has the flowers borne 1-3 in a cluster, actually a sessile cincinnus. The bizarre fruit of B. souzae, with a wide, depressed, disc- shaped nut, is quite different from the globose fruit of B. christianeae and all other species of Byrsonima. Byrsonima spinensis W. R. Anderson, sp. nov. Fig. 2 Arbor 4-6 m alta. Lamina foliorum eae 4—8 (-9) cm longa, 1.8-3.2 (-3.6) cm lata, plerumque obovata, apice obtusa vel rotundata, glabrata vel paene glabrata; petiolus 6-9 (-10) mm orcas stupulae 1.6—-2.3 mm longae omnino connatae vel apice distinctae. Inflorescentia bracteis ee ee persistentibus, cincinnis 1—2-floris, pedunculo plerumque 1-5 mm _ longi Pedicellus circinatus in alabastro, decurvatus in- fructu. Sparsissime sericea, valde appressa per anthesin. Petala alba, demum rosea. Antherae glabrae, loculis cylindricis, 1-1.6 mm longis, connectivo 0.7—1.1 mm sane Tree 4-6 m tall, the stems a a equccons (2), soon glabrate. Lamina of the larger leaves 4-8 (—9) cm long, 1.8-3.2 (-3.6) cm wide, obovate or sometimes elliptical, obtuse or rounded at the apex (sometimes abruptly short-acuminate), slightly revolute at the margin, tapered or cuneate at the base, soon glabrate or very sparsely sericeous below on the midrib, the lateral veins and reticulum visible but hardly gcarertens petiole 6—9 (-10) mm long, thinly sericeous to glabrate; stipules 1.6-2.3 mm long, completely connate or distinct just at the apex, smooth or Cee in the middle, abaxially sericeous to glabrate, adaxially glabrous. Inflorescence 3-8 cm long, thinly sericeous to glabrate, the cincinni |-2-flowered; bracts 1-2 mm long, ca 1mm wide, triangular, glabrous or abaxially thinly sericeous, stiff; primary floriferous peduncle (from bract to joint) mostly I—-5 mm long, occasionally undeveloped; bracteoles like the bracts but smaller; bracts and bracteoles persistent at least past flowering. Pedicel (3.5—) 5-8 (-10) mm long, sericeous, moderately circinate in bud, decurved in old flowers and enlarging fruits. Flower 12-16 mm in diameter. Sepals all biglandu- lar, 1.5-2.3 mm long beyond the glands, 2-2.5 mm wide, broadly obtuse or rounded, very sparsely sericeous on both sides and ciliate on the margin, strong- ly appressed in anthesis; glands white in bud, pink in anthesis, 1.6-2.5 mm long. Petals white, becoming pink in age, glabrous; lateral petals spreading to re- flexed, the claw 2.5-3.5 mm long, the limb 4—5.5 mm long, 4.5-6 mm wide; posterior petal erect, the claw 2.7-3 mm long, the limb 3-3.5 mm long, 4+ 4.5 mm wide, corrugated. Filaments 2-2.5 mm long, sparsely hirsute abaxially at base, hirsute adaxially on proximal half; anthers 1.8-2.4 mm long, glabrous, the locules 1-1.6 mm long, cylindrical, rounded or minutely apiculate and some- times slightly detached at the apex, the connective yellow, extended 0.7—1.1 mm beyond the locules, the extension conoid, bluntly pointed, nearly straight to recurved. Ovary 1-1.5 mm high, glabrous, all 3 locules fertile; styles 3.1-3.7 mm long, straight or bent away from the posterior petal. Fully enlarged fruit not seen. TYPE: BRAZIL. Minas Gerais: Serra do Espinhago, valley ca 5 km SSE of Pico de Itambe, 1140 m, by stream, 14 Feb 1972 fl, Anderson et al. 35969 (UB, holotype; MICH, NY, isotypes). 110 PARATYPES: BRAZIL. Minas Gerais: Serra do Espinhago, ca 8 km N of Gouveia on road to Diamantina, 1220 m, wet place at edge of gallery forest, Feb fl, Anderson et al. 35339 (MICH, NY); same locality and date as type, Anderson et al. 35970 (MICH, NY, UB). ‘This species is closely related to Byrsonima myricifolia Grisebach, which is also a much-branched tree with similar clusters of small, nearly glabrate leaves. ‘The both have the petals all white, turning pink in age, the bracts and bracteoles persistent, and the floriferous peduncle usually well developed. Byrsonima spinensis differs from B. myricifolia in having the pedicel circinate in bud and in having many of the cincinni 2-flowered; in B. myricifolia the pedicels are straight, even in very young buds, and the cincinni seem never to bear a second flower. Also, the lamina ts usually obtuse or rounded in B. spinensis, acuminate or acute in B. myricifolia. The epithet spinensis refers to the elongated Serra do Espinhago; espinhaco is a Portuguese word meaning backbone or spine. Byrsonima bahiana W. R. Anderson, sp. nov. Vigo: Arbor 8m alta. Lamina foliorum majorum 8-10.5 cm longa, 4.3-5.6 cm lata, obovata, apice rotundata vel obtusa, coriacea, fere glabra; petiolus 14- 16 mm longus; stipulae 1.5-2.5 mm longae, distinctae, rotundatae. Inflores- centia fere glabra, bracteis bracteolisque 1.2-1.5 mm longis, usque ad 1.9 mm latis, rotundatis, persistentibus. Pedicellus decurvatus in fructu. Sepala glandulas 2.5mm superantia, apice rotundata, fere glabra, in fructu carnosa et accres- centia, minimum 10 mm longa et 9 mm lata, basi auriculata. Petala alba et rosea. Antherae 2—2.5 mm longae, glabrae, loculis ut videtur nec complanatis nec alatis, connectivo vix superatis (0.1 mm). Ovarium glabrum, tantum 2 loculis fertilibus. Fructus immaturus ca 15 mm diametro. Tree to 8m tall, the trunk 20 cm in diameter; stems initially very thinly sericeous, soon quite glabrate except short-hirsute in axils of stipules. Lamina of larger leaves 8-10.5 cm long, 4.3-5.6 cm wide, obovate to almost elliptical, rounded or obtuse at the apex, revolute at the margin, cuneate at the base, coriaceous, tinged with red above, especially on margin and midrib, glabrous or bearing a few straight appressed hairs on the midrib below, not glaucous below, not or slightly glaucous above, the principal lateral veins ca 10, prominulous below; petiole 14-16 mm long, glabrous or bearing a few straight appressed hairs; stipules 1.5-2.5 mm long, distinct, broadly rounded, inserted so that one stands partially inside the other, glabrous. Inflorescence 10.5-11 cm long, sericeous at the very base, distally very sparsely sericeous to glabrate, the flowers borne | per bract; bracts 1.2—1.5 mm long, 1.6-1.9 mm wide, broadly rounded, glabrous or bearing a few hairs; peduncle none; bracteoles about as long as the bracts but narrower, usually not as wide as long; bracts and bracteoles persistent past maturity of the fruit. Pedicel in fruit 10-13 mm long, bearing scattered appressed hairs, decurved. Sepals all biglandular, 2.5 mm long beyond the glands, 3 mm wide, rounded, glabrous except for tiny hairs on the margin, revolute in anthesis, appressed and greatly enlarged in fruit; glands 2.5-3 mm long. Petals “strongly reflexed with reddish claw and small concave white limb, often tinged with pink,” persistent in immature fruit. Filaments 2-3 mm long, abaxially glabrous, adaxially hirsute at base; anthers 2-2.5 mm long, glabrous, the locules apparently not flattened or winged, exceeded only slightly (0.1 mm) by the connective. Ovary 1.5 mm high, glabrous, with only 2 locules developed and fertile. Immature fruit ca 15 mm in diameter, with thick flesh around a small nut ca 5-6 mm in diameter, glabrous; accrescent sepals fleshy, appressed, at least 10 mm long and 9 mm wide, somewhat auriculate at the base. 111 TYPE: BRAZIL. Bahia: 5 km SE of Marau, 39°00' W, 14°08’ S, low restinga on sand, elev. 0-50 m, 2 Feb 1977 fr, Harley 18475 (CEPEC, holotype; MICH, isotype). This species is readily placed in Niedenzu’s series Platylepis on the basis of its white and pink petals, distinct stipules, and small, subequal, persistent bracts and bracteoles. It is most similar to Byrsonima laevis Niedenzu and B. amoena Cuatrecasas. The most dramatic characteristic distinguishing B. bahiana is its fleshy sepals that are large in flower and enlarge greatly in fruit. In contrast to the dimensions given above for this species, the sepals of B. laevis and B. amoena are only 1.5 mm long or less in flower and enlarge to no more than 4 mm long in fruit. In B. bahiana the calyx glands, filaments, and anthers are longer than in B. laevis and B. amoena; also, the anther locules in B. bahiana do not appear to be flattened and winged as in the other two species, and the ovary has only two locules fertile instead of three. Additional differences may become apparent when good flowering material of B. bahiana is available for study. ‘The type bears immature fruits and my only flowers were old, degraded, and incomplete. Byrsonima laevis and B. amoena are species of northwestern Amazonia, in the drainage area of the Rio Negro in Brazil, Venezuela, and Colombia. B. bahiana is known only from the type, which was collected 2500 km from Manaus, where B. amoena grows, and even farther from the nearest populations of B. laevis. The mazonian affinities of many species of coastal Bahia have been abundantly documented, but this is still an impressive disjunction. Also, it is worth noting that this is a species of white-sand restinga, not wet forest as is true of many species or species-pairs disjunct between Bahia and Amazonia. Byrsonima laevis and B. amoena are also species of open, white-sand vegetation, the Rio Negro analogue of the restingas of Bahia. This suggests that when the Amazonian flora extended down the coast into Bahia, it probably comprised not only forests, but white-sand “campinas,” just as it does today. Byrsonima alvimii W. R. Anderson, sp. nov. Fig. 7 Arbor usque ad 20 m alta. Lamina foliorum majorum 12-18 cm longa, 5.5- 9.3 cm lata, omnino vel fere glabra; petiolus 18-25 mm longus; stipulae 5-7 mm longae, omnino connatae, glabrae. Inflorescentia bracteis 4-7 mm longis, 2— 3.3 mm latis, reflexis, demum deciduis (?), cincinnis 1—3-floris, pedunculo (0-) 1-4.5 mm longo, bracteolis bracteis similibus, 2.5-4 mm longis, plerumque non reflexis. Pedicellus circinatus in alabastro. Sepala utrinque velutina vel sericea. Petala rosea. Antherae loculis sericeis vel hirsutis praecipue apice, cylindricis, 1.5-2.1 mm longis, apice rotundatis, connectivo (0.4—) 0.7—1.1 mm_ superatis. Ovarium glabrum; styli 4.5-5 mm longi. Tree to 20 m tall, the trunk 35 cm in diameter; stems nearly glabrous ae hirsute in axils of stipules. Lamina of the larger leaves 12-18 cm long, 5.5- 9.3 cm wide, elliptical, acute or obtuse and apparently abruptly falcate at a apex, revolute at the margin, cuneate at the base, coriaceous, glabrous or with a few appressed hairs on the midrib below, the principal lateral veins 8-10, moder- ately prominent below; petiole 18-25 mm long, glabrous or very sparsely sericeous; stipules 5-7 mm long, smoothly and completely connate, the pair broadly triangular, wider than the petiole, glabrous. Inflorescence 13-16 cm long, sericeous proximally to velutinous distally; cincinni 1—3-flowered, the sec- ond and third flowers more likely to develop in proximal cincinni; bracts 4— 7 mm long, 2—3.3 mm wide, flat, lanceolate, mostly strongly reflexed, loosely and sparsely sericeous to glabr ate; primary floriferous peduncle (0-) 1-4.5 mm long, longest in eae cincinni, velutinous; bracteoles like the bracts but smaller (2.5-4 mm long, 1.2-—2.5 mm wide), mostly not reflexed; bracts and bracteoles t it wy ay Se ? Eas Ny x 3 , ) Se aa * 7. Bytsonima alvimu and B. mori. a—e, B. alvimii: a) Aa ae x 0.5; b) SPE: gC) peat with | posterior-lateral petal removed ange ape 5; d) stamens, X 7.5; e) gynoecium, with tips of styles bent away from posterior petal, . fy, B. mori: £) pope) branch, x 0.5, with reticulum of upper surface of lamina x :) ae x 2.5 1) 2a a fond stamens, X /. L2r 8 5; J) gynoecium, with anterior style to left, x 5. Dr rawn by Karin Deathit fro om Ae types. 113 apparently deciduous late in anthesis or afterwards. Pedicel 9-11 long, veluti- nous, circinate in bud. Flower 18 mm in diameter. Sepals all biglandular, 1.5-— 2mm long beyond the glands, 2.5 mm wide, obtuse, densely velutinous or sericeous on both sides, erect or revolute at the apex in anthesis; glands 2.5— 3.5 mm long. Petals pink, glabrous; lateral petals reflexed, the claw 3.5-4 mm long, the limb 6-8 mm long, 7-9 mm wide; posterior petal erect, the claw 4 mm long, the limb 5 mm long and wide, corrugated. Filaments 3-3.5 mm long, abaxially glabrous or sparsely hirsute at base, adaxially densely red-hirsute on proximal % and sometimes sparsely hirsute distally; anthers 1.9-3.2 mm long, longest opposite 3 anterior sepals and shortest opposite 3 posterior petals, the locules 1.5—2.1 mm long, cylindrical, sericeous or hirsute, especially at the apex, rounded and sometimes detached at the apex, the connective yellow, massive, extended ().7—1.1 mm beyond the locules (only 0.4 mm in anther opposite poste- rior petal), the extension truncated or bluntly pointed, straight or slightly re- curved. Ovary 1.8 mm high, glabrous, all 3 locules fertile; styles 4.5-5 mm long, bent in the distal 4-3. Fruit not seen. E: BRAZIL. Bahia: Una, Faz. Sao Rafael, mata, 10 Nov 1969 f1, 7. S. dos Santos 479 (CEPEC, holotype; MICH, isotype). PARATYPE: BRAZIL. Bahia: Ubaitaba-Marau, Dec fl, Vinha 53 (CEPEC, MICH). Another collection that may well represent this species is Pinheiro 1691 (or 16012) (MICH). It was a tree 15 m tall, collected in forest on the road from Santa Luzia to Canavieiras, Bahia. The material is in very young bud, so that color of the petals was not known, but the anthers do seem to be hairy and the connective exceeds the locules. It resembles B. alvimii in the size and shape of the leaves, bracts, and bracteoles, but all parts of the plant are much hairier and the stipules are smaller. Byrsonima alvimi is named in honor of Dr. Paulo de T. Alvim, who as Scien- tific- Technical Director at CEPLAC has played an important role in encouraging collection and study of the flora of Bahia. This species falls into Niedenzu’s series Psilonemis, due to its pink petals and hairy anthers with an extended connective. Within that group, it seems to have no close relatives. It is distinguished by its stature, large coriaceous petiolate leaves, 1—3-flowered cincinni with the peduncle often developed, very large bracts and bracteoles, and sepals densely hairy on both sides. It seems to be most similar to B. schomburgkiana, but I am not convinced that they are really very closely related. According to the label on the type, the common name of this species Is “mucambo roxo.” Byrsonima surinamensis W. R. Anderson, sp. nov. Fig. 4. Arbor 6-17 m alta, ramis sericeis. Lamina foliorum majorum 5-8.5 (—10) cm longa, 2.4-3.7 cm lata, anguste obovata, apice plerumque rotundata vel emarginata, mucronata; petiolus 2-4 (-5) mm longus, laxe sericeus vel demum glabrescens. Inflorescentia bracteis bracteolisque caducis, pedunculo nullo, floribus singulis. Pedicellus 4—5.5 mm longus (-10 mm in fructu), rectus vel aulo circinatus in alabastro, decurvatus vel tortus in fructu. Petala rosea. Antherae 1.4—2.8 mm longae, loculis 0.8-1.0 mm longis, cylindricis, distaliter laxe sericeis, apice rotundatis vel acutis, connectivo loculos 0.6—1.8 mm super- anti. Ovarium glabrum; styli 2.5 mm longi. Fructus sepalis accrescentibus, usque 6-8 mm longis, distaliter 3-4 mm latis, lingulatis, proximaliter auriculatis, membranacelis. 114 Tree 6-17 m tall, the trunk 15-30 cm in diameter; stems densely sericeous, glabrescent only belatedly; internodes mostly very short, the leaves borne in dense apical clusters. Lamina of the larger leaves 5-8.5 (-10) cm long, 2.4— 3.7 cm wide, obovate or narrowly obovate, gradually narrowed to cuneate at the base, broady obtuse or (usually) rounded or emarginate and mucronate at the apex, glaucous below, glabrate at maturity except loosely sericeous on the midrib, especially below, and on the apical mucro, with scattered appressed hairs persistent elsewhere on the lamina, the lateral veins prominulous below or on both sides; petiole 2-4 (-5) mm long, loosely sericeous or eventually glabrescent; stipules 2-3.5 mm long, smoothly connate but the pair usually emarginate at the apex, abaxially sericeous, adaxially glabrous. Inflorescence 6-12 cm long, veluti- nous-tomentose; flowers borne | per bract; bracts 2.5-3.5 mm long, ca | mm wide, narrowly lingulate, spreading and + straight, abaxially loosely sericeous, adaxially glabrous or sparsely sericeous in the center; peduncle none; bracteoles like the bracts but shorter; bracts and bracteoles deciduous before anthesis. Pedicel 4—5.5 mm long (-10 mm in fruit), appressed-tomentose, straight or slightly circinate in bud, decurved or sigmoid-twisted in fruit. Sepals all bi- glandular, 2 mm long beyond the glands, ca 2 mm wide, ovate-lingulate, strong- ly revolute in anthesis, abaxially appressed-tomentose, especially in the center, and ciliate on the margin, adaxially loosely sericeous proximally, accrescent (especially elongated) in fruit; glands 1.8-2.4 mm long, reddish. Petals pink, glabrous, the outermost covering all others in bud; lateral petals reflexed, with the claw 2.5-3 mm long, the limb 5-6.5 mm long, 5.7-7.3 mm wide, concave, especially in the anterior pair; posterior petal erect, the claw 3 mm long and thicker, the limb 4 mm long, 4.5 mm wide, corrugated. Filaments 1.5-2 mm long, longer opposite petals than sepals, abaxially glabrous, adaxially hirsute on the proximal 4; anthers 1.4—2.8 mm long (in the same flower), loosely sericeous on the locules, especially distally, the locules 0.8—1.0 mm long, rounded or acute and slightly detached at the apex, cylindrical, the connective extended 0.6— 1.8mm _ beyond the locules, the extension cylindrical, blunt, straight or the longer ones recurved, longest on anterior stamens and shortest on posterior. Ovary 1.5 mm high, ovoid, glabrous, all 3 locules fertile; styles 2.5 mm long, straight. Fruit ca 9-10 mm in diameter, glabrous; accrescent sepals 6-8 mm long, distally 3—4 mm wide, lingulate, basally somewhat auriculate, membranous. YPE: SURINAME. Lely Mts., 175 km SSE of Paramaribo, east slope of plateau no. 1, 500-700 m, 10 Oct 1976 fr, Mori & Bolten 8437 (MICH, holotype; NY and 7 undistributed duplicates, IsOLy pes). PARATYPES: SURINAME. Nassau Mts., savanna forest near km | 1.6, 16 Mar sterile, Lanjouw & Lindeman 2737 (LAN); Lely Mts., SW plateaus covered by ferrobauxite between 550 and 710 m alt., along base line on plateau 5 in low savanna-like forest on ferrite, 30 Sep fl, Lindeman et al. 597 (MO, N Byrsonima surinamensis is similar to B. schomburgkiana Bentham, a species known from southern Guyana and adjacent Brazil and Venezuela. B. schom- burgkiana differs in the following characteristics: More twisted hairs of stems and leaves, producing a vesture that I would call tomentose; larger and especially wider leaves that are more gradually tapered distally, with a longer petiole; persistent bracts and bracteoles; longer pedicels; anthers with longer locules (1.3-1.9 mm) and the connective extended only 0.5—-1.1 mm beyond the locules; longer styles. For a full description, see p. 98 of my Guayana Highland paper (1981). 115 Byrsonima aubletii Kostermans, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 25: 10. 1936, nom. nov. for B. altissima (Aublet) DC. NEESER altissima Aublet, Hist. Pl. Guiane Frang. 1: 455. 1775, not M. altissima in, 1764. Type: Aublet, French Guiana, “in sylvis Sinemariensibus” (BM!). Byrsonima altissima (Aublet) DC. Prodr. 1: 579. 1824. Byrsonima discolor Pilger, Repert. Spec. Nov. Regni Veg. 42: 179. 1937. Type: Ducke, Brazil, Para, ilhas altas do Macujubinsinho (RB 20950!). all tree, 18-30 m tall. Lamina of the larger leaves 10-15 cm long, 5.5-8 cm wide, elliptical or obovate, obtuse or rounded and often abruptly short- acuminate at the apex, sericeous to glabrate at maturity above, densely and persistently rufous-sericeous below, the reticulum prominent above; petiole 20— 33 mm long, sericeous; stipules 3-4 mm long, completely connate, abaxially sericeous, adaxially glabrous. Inflorescence with the flowers borne 1 per bract; bracts and bracteoles alike, very small, ca 0.5-1 mm long and wide, broadly triangular or rounded, persistent past maturity of the fruit; peduncle 0-0.5 mm long. Pedicel 8-10 mm long (—16 mm in fruit), straight in bud, nodding in fruit. Sepals all biglandular or all eglandular, abaxially densely sericeous, adaxially sparsely sericeous, especially proximally. Petals white or pink (probably white turning pink in age), abaxially densely sericeous. Anthers densely sericeous, with some hairs on the locules and many more on the connective between the locules and outside beside them; locules extended at the apex into long, tapered, distally sterile horns about equalling the connective; connective enlarged at the apex into a swollen, globular or elongated mass, larger in some anthers than others in the same flower. Ovary densely sericeous; all 3 locules fertile. Immature fruit ovoid, with an apical beak, thinly sericeous. In 1935, Sandwith pointed out that the correct name for Byrsonima altissima sensu Niedenzu (1928) is B. aerugo Sagot. Not only is the epithet alézssima illegiti- mate; its type was not the species for which Niedenzu used the name. However, Sandwith did not know which species was actually represented by Aublet’s type. Kostermans did not understand the taxonomy of Byrsomima even as well as Sandwith, but he went ahead anyway and published the substitute name B. aubletii. When I was able to study Aublet’s type in 1981, I discovered that the species is the one described in 1937 as B. discolor Pilger. The only difference between Aublet’s collection and the others I have seen, which have all come from Para or Maranhao, is that the latter have eglandular sepals while Aublet’s speci- men has all the sepals biglandular. The occurrence of eglandular forms is very common in Byrsonima species and does not merit taxonomic recognition, Byrsonima aubletii is quite distinctive. Its anthers are those of Niedenzu’s Section Acrotheca Subsection Uroceras. In that group this is the only species with the leaves densely and persistently sericeous below and the petals sericeous. In fact, this is the only species in the genus with densely hairy petals. In a few others (e.g. B. maguirei) the petals bear a few hairs, but in most the petals are glabrous. The brief description given above is intended to facilitate recognition of this rare and interesting plant, which is not described in Niedenzu’s 1928 monograph. HETEROPTERYS These species, like those of Byrsonima, are presented here in 5 as ea the order of their closest relatives in Niedenzu’s monograph (192 116 Heteropterys actinoctenia W. R. Anderson, sp. nov. Fig, &; Liana lignosa. Lamina foliorum majorum 7—10.5 cm longa, 3.5—4.2 cm lata, falcata, elliptica, apice acuminata, glabrata, subtus inter costam et marginem aliquot glandulis parvis munita; peuiolus 6—9 mm longus, eglandulosus. In- florescentia paniculata, 3-4 cm longa, ex 3-5 umbellis vel corymbis 3—-8-floris constans; pedunculus 4-6 mm longus, apice vel saepius infra apicem bibracteo- latus. Calycis glandulae 3.24 mm longae, obovatae, decurrentes. Nux samarae utrinque 4-8 alulis parallelis ex areola ventrali radiantibus munita. Woody vine; stems sericeous to glabrate, with many punctiform lenticels. Lamina of the larger leaves 7-10.5 cm long, 3.5-4.2 cm wide, falcate, elliptical, acuminate at the apex, thin and + flat at the margin, rounded at the base, initially sericeous? quite glabrate at maturity, bearing on each side below a row of small (0.5 mm in diameter) crateriform glands approximately midway between midrib and margin, shiny above, the lateral veins and reticulum visible below, barely so above; petiole 6-9 mm long, glabrate or bearing scattered appressed hairs at maturity, eglandular; stipules not found. Inflorescence axillary and terminal, 3-4 cm long, a raceme of 3-5 short-stalked or subsessile umbels or corymbs of 3-8 flowers each, tomentose-sericeous; floriferous bracts 1—1.5 mm long, triangular, concave, appressed, eglandular, persistent; peduncle 4-6 mm long; bracteoles like the bracts or smaller, one or both usually borne up to 1mm _ below apex of peduncle. Pedicel 8 mm long (in fruit), tomentose- sericeous. Sepals (in fruit) triangular, flat (not revolute), 1.5-2 mm long, ca 2mm wide, hardly exceeding the glands, abaxially tomentose, adaxially glabrous, the anterior eglandular, the lateral 4 bearing 8 large glands 3.2-4 mm long, obovate, compressed, somewhat decurrent onto pedicel. Petals and stamens unknown. Style stigmatic on the inner angle of the apex, briefly ex- tended dorsally and rounded or obtuse. Samaras 30-40 mm long, brown- sericeous, especially proximally and toward abaxial edge of wing; dorsal wing 25-32 mm long, 12-16 mm wide, often with a rounded projection on adaxial edge near nut, the abaxial margin straight or slightly curved upward; nut ca 6 mm long and high, bearing on each side a series of 4-8 short parallel winglets oriented at right angles to margin of ventral areole, irregular in size and shape, the largest 2.5 mm wide, often alternating with much smaller outgrowths; ventral aerole ovate, 3-4 mm high, 2-2.5 mm wide; pyramidal torus ca 2 mm high. TYPE: PERU. Loreto: Prov. Maynas, upland forest remnant and adjacent scrub on white sand just past Quistococha, ca 200 m, 27 May 1978 fr, A. Gentry 22302 ES N. Jaramillo (MICH, holotype). Heteropterys actinoctenia resembles H. cristata Bentham and H . floridana Cuatr. in its leaves and inflorescence, so I assume it will prove to have pink carinate or winged petals when it is found with flowers. It differs from both of them, and from all other species in the genus, in its samara, which bears on the nut a series of parallel winglets oriented at right angles to the margin of the ventral aerole. The epithet actinoctenia refers to this “radiating comb”; it is the name Niedenzu gave to the section of Banisteria (Banisteriopsis) in which this kind of samara occurs. In other species of Heteropterys the nut of the samara is smooth-sided or bears on each side a single transverse winglet oriented parallel to the margin of the ventral areole. The samara of H. cristata and H. floridana is of the latter type. Heteropterys actinoctenia is also distinguished by its somewhat decurrent calyx glands and its relatively short peduncles and pedicels, which produce an in- florescence denser than that of its close relatives. 117 IG. ; b) rete cones old flower after loss of samaras, with pedicel and part of pedu Uncles 9<, 275) d) ti f style, x he . base of samara, x 2. fH, H. marleneae: f) fruiting branch, x 0.5; g) pseudo- 2.5; h) tip of style, x 25; 1) immature samara, x 10. yew by Karin 8. Het ee actinoctenia and H. marleneae. a—e, H. actinoctenia: a) fruiting branch, x 0.5; c) ¢ raceme ae 2 old flowers, x 2 Douthit from the types. 118 Heteropterys Series Rhodopetalis Niedenzu Subseries Stenostigma Niedenzu In his treatment of Heteropterys for Das Pflanzenreich, Niedenzu divided the pink-flowered species into two principal subseries, Bradystigma and Stenostigma, plus a third subseries for the Antillean species, H. purpurea. The most important difference between the first two lay in the stigmas, which are dorsiventrally compressed in two or all three styles in Bradystigma, laterally compressed in Stenostigma. Study of the species assigned to Stenostigma by Niedenzu has led me to remove the following species, for the reasons given: Hf. rubiginosa Adr. Juss.: The type proves to have the styles of Bradystigma. Hf. trichanthera Ady. Juss.: The petals are yellow. H. machaerophora Nied.: Two of the three styles are dorsiventrally compressed as in Bradystigma. HM. martiana Adr. Juss.: The petals are yellow; this species is probably related to . syringifolia Grisebach. H, dusenu Nied.: The petals are yellow. This leaves only Heteropterys pteropetala Ady. Juss. in Stenostigma. H. ptero- petala is a widespread, polymorphic species that merits careful study, which will probably result in its division into several entities. [am not prepared to address that problem at this time, but there are two additional species of Stenostigma in eastern Brazil that seem to be distinct from H. pleropetala. They are decribed below. Heteropterys caducibracteata W. R. Anderson, sp. nov. Fig. 9. Frutex usque ad 2 mm altus vel liana lignosa. Lamina foliorum majorum 3-4.8 cm longa, 1.6-2.5 cm lata, utrinque pertinaciter tomentosa, subtus 0, 1, vel 2 glandulas prope basim ferens; petiolus 3-4 mm longus, eglandulosus. Bracteae 1.8-2.5mm_ longae, 0.4-0.5 mm latae, anguste triangulares vel ellipticae, eglandulosae, deciduae; bracteolae bracteis similares sed persistentes. Petala rosea et alba; petala lateralia dorsaliter alata, ala alba, pilosa, 0.6-1 mm lata: petalum) posticum dorsaliter carinatum. Filamenta inaequalia, illa sepalis opposita plerumque longiora, illa petalis postico-lateralibus Opposita crassissima, 3 postica inter stylos posticos inflexa; antherae 0.9—-1.1 mm longae. Shrub to 2 m tall or woody vine climbing to 4 m; stems to 3 cm in diameter, tomentose-sericeous, belatedly glabrescent. Lamina of the larger leaves 3-4.8 cm long, 1.6-2.5 cm wide, elliptical or ovate, acute or obtuse at the apex, flat at the margin, cuneate at the base, persistently tomentose or subvelutinous above with stalked, mostly T-shaped sinuous hairs, persistently densely tomentose below, bearing 0, I, or 2 slightly peltate glands below near base between midrib and margin, the lateral veins prominent below, obscure above; petiole 3-4 mm long, appressed-tomentose, eglandular; stipules 0.2 mm long, triangular, borne on petiole above its base. Inflorescence paniculate, densely tomentose, the flowers borne in corymbs or dense racemes of 10-20 (or moreyr); bracts 1.8-2.5 mm long, 0.4—0.5 mm wide, narrowly triangular or elliptical, thin, abaxially densely tomentose, adaxially glabrous or sparsely tomentose, eglandular, deciduous late in anthesis; peduncle 5-7.5 mm long, densely tomentose; bracteoles like the bracts or slightly shorter, persistent (at least through anthesis; older stages not seen), borne at or slightly below apex of peduncle. Pedicel 2.5-4.5 mm long, densely tomentose. Flowers ca 12 mm in diameter. Sepals 1.5-2 mm long, I- 1.3 mm wide, triangular, obtuse at the apex, appressed in anthesis, abaxially densely and uniformly tomentose, adaxially glabrous, the anterior eglandular or bearing | or 2 glands, the lateral 4 all biglandular, the glands 1.2—1.6 mm long, pink. Petals pink with white wings, pilose especially on the wings; lateral petals — 119 FIG. 9. Heteroplerys ae a) flowering branch, x 0.75; b) lamina below, x 7.5; c) lane) awe x 7.5; d) flower bud, with pedicel and peduncle, x 3; e) flower bud from ee 3.75; f) bracts, adaxial ae left, a oe right, x 10; g) flower, with pedicel, peduncle, and falling eee x 4.5; h) lateral petal, x 1) parual androecium, flattened out, with stamen to right Opposite posterior petal, x 10; }) style ne eel view left, adaxial view right, x 30. Drawn by Karin Douthit from the type 120 spreading to reflexed, the winged claw 1.5-2 mm long, the limb 3.5-4.5 mm long, 2—2.3 mm wide, elliptical, entire or slightly erose, eglandular, bearing a prominent decurrent dorsal wing 0.6—1 mm wide; posterior petal erect with the limb reflexed, the claw 2.8-3.5 mm long, | mm wide, the limb 3.3—4 mm long, 2.3-2.7 mm wide, oblong, erose, eglandular, dorsally carinate with the keel 0.2 mm wide, decurrent. Filaments red in age, 2.2-3.5 mm long, connate in the basal 0.8—1 mm, glabrous, unequal, generally longer opposite sepals than petals, thickest Opposite posterior-lateral petals, the posterior 3 inflexed; anthers glabrous, 0.9-1.1 mm long, subequal, largest opposite posterior-lateral petals. Ovary 1.5 mm high, densely sericeous; styles 1.7—2.2 mm long, equal or the anterior slightly shorter, + straight, somewhat divergent, flattened laterally, truncate dorsally at the apex, the stigma internal and elliptical. Fruit unknown. ZIL. Bahia: Caatinga between Breijao da Caatinga and Delfino (4 1°0-20' W, 10°25' S), sandy soil, 600 m, 9km W of Breijao, 7 Mar 1976 f1, Anderson 11742 (MBM, holotype; MICH, isotype). Heteropterys caducibracteata is named for its deciduous bracts, which fall late in anthesis, in contrast to the persistent bracts of the other species of Stenostigma. The bracts and bracteoles are also unusual in their narrowly elliptical shape and thin texture. The leaves in this species are smaller than in H. perplexa and H. pteropetala sens. lat., in which the larger laminas are at least 6 cm wide, and usually quite a bit larger. Heteropterys perplexa W. R. Anderson, sp. nov. Fig. Liana lignosa. Lamina foliorum majorum 6.5-13.5 cm longa, 3.8-8 cm lata, ovata vel elliptica, subtus pertinaciter subvelutina et 2 magnis glandulis prope basim munita; petiolus 5-6 mm longus, eglandulosus. Bracteae bracteolaeque persistentes; pedunculus 5-9 mm longus. Pedicellus 3-5.5 mm longus. Sepala abaxialiter sericea. Petala limbo albo et roseo, ungue rubro; petala lateralia dorsaliter alata, ala 1.2—2.5 mm lata; petalum posticum dorsaliter carinatum, carina 0.1-0.3 mm lata. Filamenta rubra, valde inaequalia, illa petalis antico- lateralibus opposita brevissima, illa petalis postico-lateralibus opposita crassis- sima, 3 postica inter stylos posticos inflexa. Joody vine climbing to 10 m; stems densely appressed-tomentose or sub- sericeous, eventually glabrescent. Lamina of the larger leaves 6.5-13.5 cm long, 3.8-8 cm wide, ovate or elliptical, acute or slighty acuminate at the apex, flat at the margin, broadly cuneate to rounded at the base, appressed-tomentose to glabrate above, persistently subvelutinous below with T-shaped hairs with the trabec ula much longer than the stalk and straight or sinuous, bearing 2 large, often swollen or slightly peltate Suahie abaxially above the base, on the midrib or on a lateral vein between midrib and margin, the lateral veins and coarse reticu- lum prominent below, impressed above; petiole 5-6 mm long, appressed- tomentose to glabrescent, eglandular; stipules 0.2 mm long, borne on the petiole just above its base. Inflorescence paniculate with widely spreading branches, densely tomentose, containing reduced leaves with 2 large prot rant and some- times short-stalked glands, the flowers borne in umbels or tight corymbs of 4-15; bracts 1.3-1.8 mm long, 0.6-1 mm wide, triangular or ovate, abaxially densely and uniformly tomentose, adaxially glabrous, appressed, eglandular, persistent (at last while peduncle persists); peduncle 5-9 mm long, densely tomentose; bracteoles 1-1.5 mm_ long, 0.8-lmm_ wide, triangular or ovate, abaxially tomentose in center and glabrate and membranous toward mar gin, adaxially glabrous, spreading, usually bearing 2 small marginal glands or glandular spots near the base, persistent, borne at or just below apex of peduncle. Pedicel 3— 121 FIG. 10. dee BES a, g) aus branches, x 0.5, yea circles aa flower buds with pedicel and peduncle, x 2; c, i) flower buds from above, = d, }) tower ee above, x 3; e,k) partial androecia, flattened out, with stamen to left opposite Een petal, x 7.5; f, 1) style ups, lateral view left, adaxial view right, x 20. Drawn by Karin Douthit, a—-f from . Gone 11749, g-l from Irwin et al. 23203 War 5.5 mm long, densely tomentose. Flowers 10-16 mm in diameter. Sepals 1.5— 2 mm long, 1-2 mm wide, triangular, acute or obtuse at the apex, appressed in anthesis, abaxially uniformly sericeous or glabrous at the margin, adaxially glabrous, the anterior eglandular, the lateral 4 bearing 8 glands 1.3—2 mm long. Petals white on the margins, pink i in the center, and red in the claw, glabrous or sparsely pilose on edge of wing; lateral petals spreading, the winged claw 1. 1— 2 mm long, the limb 3—5.5 mm long, 2—3.3 mm wide, elliptical, entire, eglandular, bearing a prominent dorsal wing |.2—2.5 mm wide; posterior petal erect with the limb somewhat reflexed, the claw 2-3 mm long, 1-1.4 mm wide, the limb 2.3- 3.5 mm long and wide, irregularly rectangular, revolute, slightly erose, eglandu- lar, dorsally carinate with the keel 0.1—0.3 mm wide, decurrent. Filaments red, 1.5-3.5 mm long, connate in the basal 0.5-1 mm, glabrous, strongly unequal, shortest opposite anterior-lateral petals, thickest opposite posterior-lateral petals, the posterior 3 bent inward between the 2 posterior styles; anthers glabrous, unequal, 0.9-1.8 mm long, shortest opposite the 4 lateral sepals and the posterior petal. Ovary I-1.5 mm high, densely sericeous; styles 2-3 mm long, glabrous, straight and erect or divergent or leaning toward posterior petal, flattened later- ally, rounded or truncate or apiculate dorsally at the apex, the stigma internal and elliptical. Fruit unknown. TYPE: BRAZIL. Bahia: Caatinga 10-20 km W of Delfino on ae trom Lagoinha to Campo Largo, 41°20' W, 10°27'S, 650-750 m, 8 Mar 1976 f1, Anderson 11749 (MBM, holotype; K MICH, NY, RB, US, ge) PARATYPES: BRAZIL. Bahia: 8km NW of Lagoinha, 5.5 km SW of Delfino, on road to Minas do Mimoso, at ee om frequently burned and cut over, ca 850 m, ca 41°17’ W, 10°24’ S, Mar fl, Harley 16775 (MICH). Minas Gerais: Serra do eles wooded valley slopes, ca 5 km NE of Francisco Sa, road to Salinas, 950 m, Feb fl, Irwin et al. 23203 (MICH, NY, UB). I originally intended to describe this as two species, one based on Anderson 11749, the other on /rwin et al. 23203. They have flowers that are dramatically different in size, with petal- wings 2-2.5 mm wide vs. 1.2-1.5 mm wide, respec- tively, and longest filaments 3-3.5 mm long vs. 2.5 mm. Figure 10 was drawn to illustrate those and other differences. Then I realized that Harley 16775 1s gen- erally similar to my //749 and came from the same area, but has flowers that are somewhat intermediate in size, especially in the petal- wings, which are 1.5- 1.7 mm wide. Since the three collections are similar in most vegetative features, it seems best to treat them as one species, at least for the present. Perhaps when more collections are available it will become possible to achieve a finer resolution of this perplexing situation. The only character that reliably separates this species from H. pteropetala Adr. Juss. sensu lato is habit. Heteropterys perplexa is a woody vine, while H. pteropetala is an erect subshrub, shrub, or small tree. The common form of H. pteropetala differs further in having larger suborbicular leaves with denser hairs below. As noted above, H. pteropetala requires thorough study. Heteropterys krapovickasii W. R. Anderson, sp. nov. Fig. 11: Liana lignosa. Lamina foliorum majorum 7.5-10.5 cm longa, 3.5-5.1 em lata, eglandulosa, subtus pertinaciter metallo-sericea; petiolus 7-13 mm longus, 2 glandulas magnas inter medium et apicem ferens. Inflorescentia paniculata, folis minoribus petiolo vel basi laminae biglandulosis et saepe margine laminae aliquot glandulis ace gees munitis; flores in umbellis vel corymbis (2—) 4—8- floris portati, saepe I-2 paribus proximalibus adjectis; bracteolae plerumque prope medium pedunculi portatae. Petala flava, ungue 1.3-2.4 mm longo, limbo 3-4 mm longo, 2.3-3 lato. Filamenta 1.7-2.5 mm longa, recta; antherae |— 123 1.1 mm longae, glabrae. Styli recti. Samara 23-30 mm longa; ala dorsalis 13- 15 mm lata, flabelliformis. Woody vine, the stems puberulent to glabrate. Lamina of the larger leaves 7.5-10.5 cm long, 3.5—5.1 cm wide, elliptical or slightly obovate, obtuse, round- ed, or emarginate and often apiculate at the apex, flat or slightly recurved at the margin, cuneate or obtuse at the base, eglandular (but see inflorescence leaves), initially sericeous above but soon glabrate or persistently sericeous on the midrib, very densely and persistently metallic-sericeous below, the hairs both brown and white, sometimes abraded from oldest leaves, the lateral veins prominent below, the reticulum visible above; petiole 7-13 mm long, sericeous to glabrate, bearing 2 large glands between middle and apex; stipules not found. Inflorescence FIG. le Heteropterys krapovickasu. a) fruiting branch, x 0.55, circle x is 2.8: b) base 5 leaf, abaxial side, x 1.65; c) poruon of inflorescence in fruit, x 2.2; d) samara, = |. ) flower, X 3.85; f) tals, aaa eee petal left, lateral petal pee x 5.5; g) portion of ees opened out, abaxial ‘lew, stamen to left opposite posterior petal, x 8.2 35 5; h) up of style, side view, x 22. Drawn by Karin Deut hit, a-d from Pires & Santos 16372, e-~h from Krapovickas & oF Schinini 36259. 124 paniculate, subvelutinous or appressed-tomentose, containing abruptly reduced leaves with large glands on petiole or base of lamina and often several smaller glands distally on margin of lamina; flowers borne ultimately in umbels or corymbs of (2—) 4-8 flowers, often with | or 2 additional pairs of flowers borne more proximally; bracts 0.9-1.2 mm long, 0.5-0.8 mm wide, ovate, appressed, abaxially sericeous, adaxially glabrous, eglandular, persistent; peduncle (1.5-) 2.5-4 mm long; bracteoles like the bracts but slightly shorter (0.7—1 mm long), borne mostly near middle of peduncle, sometimes somewhat above or below. Pedicel 3-5.5 mm long, appressed-tomentose. Flowers 10-11 mm in diameter. Sepals 2—2.5 mm long, 1.5—-1.7 mm wide, triangular or ovate, obtuse or rounded at the apex, appressed in anthesis, abaxially sericeous, adaxially glabrous, all eglandular or the lateral 4 bearing 7-8 glands 1.1-1.4 mm long, elliptical, sepa- rated on the sepals. Petals yellow, glabrous, eglandular, nearly entire (minutely denticulate or erose); lateral petals horizontal, + spatulate, with the claw 1.3— 1.7 mm long, the limb 3-4 mm long, 2.3-3 mm wide, ovate or broadly elliptical, somewhat concave; posterior petal apparently erect, hardly different in size and shape from the lateral 4, the claw 2-2.4 mm long, the limb 3 mm long, 2.3— 2.5 mm wide, truncate at the base. Filaments 1.7—2.5 mm long, mostly shorter and stouter opposite petals than sepals, straight, glabrous, basally connate; anthers I-1.1 mm long, alike, glabrous. Ovary | mm high, densely sericeous; styles 2.2 mm long, straight and + erect, equalling or slightly exceeding the anthers, truncate and acute or slightly apiculate dorsally at the apex. Samaras 23-30 mm long, sparsely sericeous; dorsal wing 23-30 mm long, 13-15 mm wide, extending over the adaxial edge of the nut, much flared distally to produce a flabellate shape, the abaxial margin nearly straight; nut 6 mm long, 5 mm high, with prominent parallel longitudinal wrinkles (dried) but no lateral crests or wings; ventral areole circular, 1.5—-2 mm in diameter. TYPE: BOLIVIA. Santa Cruz: Prov. Sandoval, San Matias (58°26' W, 16°21' S), 5km SW del Aeropuerto, 170m, 19 Apr 1980 fl, Krapovickas & Schinint 36259 (MICH, holotype; CTES, NY, isotypes). PARATYPE: BRAZIL. Mato Grosso: 57°50’ W, 14°48" S, capoeira de mata seca, Aug fr, Pires & Santos 16372 (MICH, MG). This species is named in honor of my friend, Antonio Krapovickas, a fine botanist who specializes in the systematics of the Malvaceae. Heteropterys krapovickasii is closely related to H. hassleriana Nied., from which it differs chiefly in having the hairs on the abaxial side of the lamina very dense, straight, sessile, appressed, and_ persistent, producing a metallic-sericeous vesture. In H. hassleriana the hairs are sparser and less appressed and the leaf is usually soon glabrate. Also, in H. hassleriana the samara wing is not or hardly flared distally, so that the two edges of the wing are nearly parallel. Both of these species are probably closely related to the larger-flowered H. tomentosa, although Niedenzu placed H. tomentosa and H. hassleriana in different subsections. Heteropterys sincorensis W. R. Anderson, sp. nov. Figs il. Frutex vel arbor parva usque ad 3 (-4) m alta. Lamina foliorum majorum (5-) 6-10 cm longa, 4.5-6.5 cm lata, coriacea, late ovata vel elliptica vel sub- orbicularis, apice rotundata et interdum apiculata vel emarginata, basi rotundata vel cordata, mox glabrata, basi margine 2 glandulas magnas impressas et inter- dum distaliter aliquot glandulas minores marginales ferens; petiolus 2-4 mm longus, eglandulosus. Pedunculus (1.5-) 4-7 mm longus, apice bibracteolatus. Sepala appressa, plerumque omnia biglandulifera, glandulis sericeis. Petala flava, abaxialiter rubra in centro, denticulata; petalum posticum limbo. basi glanduloso. Androecium glabrum. Styli apice truncati vel interdum apiculati. 125 all, Heteropterys sincorens!s. a) flowering branch, x 0.5 left and upper surface in circle to right, both Xx ca 2.5; b) base of leaf, abaxial view, X 3; c) umbel, ae): flower, X 3.5; e) ae abaxial view, anterior- ae (left) and posterior ae x 3.5; f) anthers, ae view (left) and adaxial view (right), 0; g) style-tip, x 20; h) samara pope on receptable, . Drawn by Karin Douthit, a-g from fae 18821, h Hone Mori et al. 12635. , with leaf below enlar ae in circle to 126 Samarae ala dorsalis 23-27 mm longa, 11-14 mm lata, nux sine cristis laterali- JUS. Shrub or small tree up to 3 (4) m tall; stems loosely sericeous, soon glabrescent. Lamina of the larger leaves (5—) 6-10 cm long, 4.5-6.5 cm wide, broadly ovate or elliptical to suborbicular, rounded and sometimes apiculate or emarginate at the apex, slightly revolute at the margin, rounded or cordate at the base, coriaceous, at first sparsely sericeous but soon glabrate, bearing 2 large impressed marginal glands at base of blade or halfway onto petiole and often additional smaller glands distally in marginal sinuses, the lateral veins and coarse reticulum prominent below, the fine reticulum prominent above; petiole 2—4 mm long, thick, loosely sericeous to glabrate, eglandular except for the often decurrent glands at base of lamina; stipules often present on petioles of small inflorescence-leaves as minute (0.1—0.2 mm long) protuberances borne on peuole above base, apparently absent from larger leaves. Inflorescence a much- branched, many-flowered panicle containing abruptly reduced leaves, the ulti- mate branches sometimes racemose but usually corymbose or umbellate com- prising 4-10 or more flowers, loosely sericeous; bracts and bracteoles ]-2 mm long, 0.5-1 mm wide, elliptical or obovate, rounded at the apex, appressed, persistent, the bracteoles borne at apex of peduncle, usually bearing 2 tiny marginal glands at the base; peduncle (1.5-) 4-7 mm long. Pedicel 5-10 mm long, loosely sericeous. Flowers 11-13 mm in diameter, “very sickly scented” (Harley 15981). Sepals 1.6-2 mm long, 1.2-1.5 mm wide, ovate or triangular, acute or obtuse at the apex, appressed in anthesis, abaxially sericeous, adaxially glabrous, all biglandular or (in | collection) the anterior eglandular, the glands 1.5-2.5 mm long, obovate, densely sericeous, attached below free portion of sepal. Petals yellow with red abaxial streak, turning orange-red in age, glabrous; lateral petals spreading to eventually strongly reflexed, with the claw I— 1.8 mm long, winged, the limb 3.5-4.5 mm long, 2-3 wide, oblong or obovate, dentuculate and eglandular at the margin, thickened and slightly carinate in the middle; posterior petal with the claw 1.5-2.3 mm long, thick and winged, erect, the limb 3-3.5 mm long, 2.3-3 mm wide, reflexed, similar to lateral petals but with glandular zones or discrete glands at the base. Filaments 1.5—2.5 mm long, longer Opposite sepals than petals, erect and straight, connate in the proximal 0.7-1.3 mm, glabrous; anthers I-1.4 mm long, glabrous, subequal. Ovary ca 1.5 mm high, densely sericeous; styles 1.5-2 mm long, glabrous, alike, erect to eventually divergent with the anterior bending back toward the anterior sepal and the posterior 2 bending toward the posterior petal, equalling or slightly exceeding the anthers, truncate at the apex, sometimes minutely apiculate dorsally. Samaras 30-35 mm long, borne quite erect when single, suberect when 2 develop, reddish purple (dried), sparsely and loosely sericeous; dorsal wing 23-27 mm long, 11-14 mm wide, the abaxial margin nearly straight; nut ey Saghesl long, 4—5 mm high, with prominent veins but no lateral crests or wings; ventral areole ovate or circular, ca 3 mm in diameter; torus ca 2.5 mm high. TYPE: BRAZIL. Bahia: Serra do Sincora, by Rio Cumbuca ca 3 km S of Mucuge, near site of small dam on road to Cascavel, approx. 41°21’ W, 13°01’ S, 850 m, 4 Feb 1974 f1, Harley 15981 (CEPEC, holotype; MICH, isotype). PARATYPES: BRAZIL. Bahia, Serra do Sincora: 5 km S of Andarar on road to Mucuge, by bridge of Rio Paraguagu, 41°19’ W, 12°50’ S, ca 400 m, Feb fl, Harley 18587 (MICH); 2-3 km SW of Mucugé on road to Cascavel, open scrub on white sand, 41°24’ W, 13°01'S, ca 950 m, Feb fi, Harley 18821 (MICH); 6.5 km SW of Mucugé on Cascavel road, sandstone rock and sandy soils, 41°25’ W, 13°01’ S, ca 1000 m, Mar fl, Harley 21019 (MICH); Mpio de Mucuje, na estrada para Jussiape, 41°24’ W, 13°00'S, 1000 m, campo rupestre, Jul fr, Mori et al. 12635 (MICH, US). Ag This distinctive species is notable for its arborescent habit, glabrate coriace- ous leaves with marginal glands and short petioles, umbels or corymbs of pedunculate flowers, hairy calyx glands, oblong yellow petals with a red abaxial streak, and the flag petal with glands at the base of the limb. It is probably most similar to Heteropterys bahiensis Nied., known only from the type, which I have not seen. The photograph of the specimen at G (F neg. 24258) shows it to have relatively long, narrowly elliptical leaves, cuneate at the base, with slender, well developed petioles. When H. bahiensis is collected with good flowers and fruits, further differences from H. sincorensis will surely be found. Heteropterys sincorensis is known only from the Serra do Sincora, for which it is named; the place of origin of Blanchet’s type of H. bahiensis is not known. Other species in this complex are H. megaptera Adr. Juss., from Rio de Janeiro, and H. lasseri Anderson, from Venezuela. Heteropterys conformis W. R. Anderson, sp. nov. Bide 13: Liana lignosa, ramis mox glabratis. Lamina foliorum major um 5.5-9.5 (-12.5) cm longa, 3-4.5 (—5.7) cm lata, margine tenuis, primo sericea permox omnino glabrata, reticulo utrinque + prominenti; petiolus 7-15 mm longus, basi biglandulosus. Inflorescentiae axillares et terminales, 1.54 cm longae, floribus congestis. Pedicellus 4-7 mm longus, 0.3-0.4 mm diametro, sessilis. Sepala appressa per anthesin, eglandulosa. Petala flava, valde reflexa, limbo 2-2.5 mm longo, 1.8-2.3 mm lato. Styli angulo interno stigmatici. Woody vine; stems initially sericeous, soon glabrate. Lamina of the larger leaves 5.5-9.5 (-12.5) cm long, 34.5 (—5.7) cm wide, elliptical, rounded, obtuse, or acute at the apex, sometimes emarginate or abruptly short-acuminate, thin and slightly revolute at the margin, cuneate to rounded at the base, initially sericeous but very soon quite glabrate, eglandular or (Glaziou 13608) bearing several small glands on or just within the margin, the lateral veins and reticulum prominent on both sides; petiole 7-15 mm long, sericeous to soon glabrate, biglandular at or just above the base, the glands circular, slightly raised, 0.7— 1.3 mm in diameter; stipules not found. Inflorescence a series of dense axillary and terminal clusters, borne among current leaves, each unit 1.5—4 cm long, sericeous, a raceme of sessile and stalked umbels of 2-6 flowers with some flowers borne singly on the axis; bracts and bracteoles 0.5—1 mm long, narrowly ovate or triangular, eglandular, persistent; floriferous peduncle none. Pedicel 4—7 mm long, 0.3-0.4 mm in diameter, sericeous. Flowers 6-8 mm in diameter. Sepals 2—2.5 mm long, 0.9-1.5 mm wide, ovate or elliptical, rounded at the apex, appressed in anthesis, abaxially loosely sericeous, adaxially glabrous, all eglandu- ar. Petals yellow, glabrous; lateral petals soon strongly reflexed, the claw 1.2- 1.8mm long, the limb 2-2.5 mm long, 1.8-2.3 mm _ wide, ovate, concave, denticulate, eglandular; posterior petal hardly different from the lateral 4, the limb or limb and claw strongly reflexed, the claw 1.5-2.5 mm long, the limb 2—2.5 mm long, 2 mm wide, ovate, bearing tiny glandular-thickened zones at the base or tiny sessile glands around the margin. Filaments 1.5—2 mm long, connate in the proximal 0.5-0.7 mm, strongly unequal, stoutest and straightest ec the 2 posterior-lateral petals, the posterior 3 bent inward, glabrous; anthe1 0.8-1.2 mm long, largest opposite ree a lateral petals, glabrous Ue con- nective dark red to black. Ovary I-1.3 mm high, tomentose; styles 1.7—2.2 mm long, equalling or slightly exceeding ae stamens, glabrous, pone eae to somewhat arcuate, laterally flattened, truncate at the apex or dorsally acute, stigmatic on the inner angle. Fruit unknown. TYPE: BRAZIL. Bahia: Itaju do Colonia a 25 km da estrada Itaju—Pau Brasil, solo pedregoso em capim, 23 Jan 1969 fl, T. S. Santos 346 (CEPEC, holotype; MICH, isotype). — 128 FIG. 13. de dea RON arrnas and Hf. sanctorum. a-f, H. nS a) Des: branch, x 0.5 circles X 2.5; b) petiole, 2.5: c) portion of inflorescence, 5: d) flower, x 5; e) at aceniae stretched out, sea il view, with stamen to left Opposite anterior aes 10; £) gynoecium, x 7.5 g—m, H. sanctorum: g) leat, abaxial view, x ae ae x 2.5: h) wena on old stem, * 0.5; 1) portion of inflorescence, * 2.5; }) lowe »; k) portion of eee oecium, x ! Peat anterior style to left, x 7.5; m) style-tip, x 25. ee by Karin Douthit an the ae 129 PARATYPES: BRAZIL. Minas Gerais: Biribiry, pres de Diamantina, 1882, Glaziou 12491 (C, G, P); Rio Manso, Glaziou 13608 p-p. (P). I doubt that either of the paratypes actually came from Minas Gerais. For some reason, Glaziou falsified the label data for many of his collections, ascribing to Rio de Janeiro or Minas Gerais plants that very probably came from Bahia or farther north. Glaziou 18925a, the type of Byrsonima glazoviana Nied., is B. crispa Adr. Juss. and surely came from Bahia or north, not Minas Canis: Another example is Glaziou 9680, which is Peixotoa jussieuana Adr. Juss.; Glaziou said it came from Rio de Janeiro, whereas it probably came from Maranhao, Piaur, or Ceara (C. Anderson, pers. comm.). This species, and the next two, belong to the series Holopetalum, which | would define as comprising species with appressed sepals, yellow petals, sessile pedicels, and the lamina not persistently metallic-sericeous below. Heteropterys conformis is quite similar to H. ternstroemiifolia, and the epithet chosen denotes that similarity. The two species can be distinguished by the characteristics sum- marized in the following couplet: Inflorescence axillary and terminal, 1.5—!cm long, the flowers densely crowded; calyx eglandular; lamina initially sericeous ee ae soon quite glabrate, the margin thin, the reticulum + prominent below; limb of petals 2-2.3 mm long, 1.8—2.3 mm w fide: stigma ongly internal. H. cfr eae terminal, 2-14 cm long, open; calyx bearing 8 glands on the 4 lateral sepals amina initially densely tomentose, usually glabrate at maturity, the margin thic ened and cordlike, the reticulum invisible below; limb of petals 3-4.5 mm long, 2.5-3 mm wide; stigmas terminal or subterminal. H. ternstroemitfolia. — seer ternstroemiifolia Adr. Juss., Ann. Sci. Nat. 2° Sér. Bot. 13: 274. 1840. Type: Martius 116, “in sylvis caeduis ad latera montium prope Sebasti- anopolim ei. Mandiocca,” Rio de Janeiro, Brazil (M! holotype; M! probable Isotype, P! isotype). Banisteria fischeriana Regel & See Ind. sem. horti Petrop. 48. 1858. Type: Riedel 173, Corcovado, Rio de Janeiro, Brazil, Jan 1832 fl (LE! lectotype; GH! LE! U! Ww! Haleenisos or isosyntypes Banisteria ternstroemifolia (Adr. Juss.) Nied., Verz. Vorles. Ak. Braunsberg 1912- 1913: 14. 1912. Woody vine, the stems sericeous to glabrate, longitudinally fissured and bearing punctiform lenticels in age, occasionally rooting at the nodes. Lamina of the larger leaves 6.5-13 cm long, 2-6 cm wide, elliptical or obovate, cuneate or occasionally rounded at the base, thickened and cordlike at the margin and sometimes corrugated, acute to rounded and often apiculate and retuse at the apex, initially densely rusty-tomentose on both sides, soon glabrescent, usually nearly or quite glabrate at maturity and often shining above, eglandular, the lateral veins and reticulum obscure or prominulous above, obscure below; petiole 8—15 (—20) mm long, Be oe tomentose to glabrate, biglandular at the base, the glands circular, 0.5-1.5 mm in diameter; stipules subulate, ca 0.5 mm long, borne at base of petiole just below glands. Inflorescence 2-14 cm long, open, loosely sericeous to glabrate, terminal, simple or occasionally basally ternate, a raceme of sessile or stalked umbels or corymbs of 2-6 flowers each, the umbels often subtended by biglandular bracts; floriferous bracts 0.7-1 (-1.5) mm long, triangular, eglandular, persistent; peduncle none; bracteoles like the bracts but smaller, usually ca 0.5 mm long. Pedicel 4-10 mm long, loosely sericeous, tumid at the apex, thickening in fruit, especially toward the apex. Sepals 2-3.5 mm long, 1.5-2 mm wide, ovate, acute or rounded at the 130 apex, appressed in anthesis, abaxially loosely sericeous, adaxially glabrous or sericeous at the margin, the anterior eglandular, the lateral 4 bearing 8 elliptical glands, the glands |.4—2.5 mm long, free and sometimes revolute at the apex, not compressed. Petals yellow, ee all 5 very strongly reflexed [flipped completely back] in anthesis, the claw 1.5—1.7 mm long, the limb 34.5 mm long, 2.5-3 mm wide, obovate, convex in center, revolute at margin, the margin eglandular, erose or denticulate; posterior petal hardly different from the lateral 4, the limb slightly longer. Filaments 2-3 mm long, alike, straight, connate for about 2 their length, glabrous on both sides; anthers 1—1.6 mm long, alike, Ae ous, erect to reflexed. Ovary 1.5 mm high, reddish-tomentose, 3-lobed, all 3 locules fertile; styles 1.5-1.8 mm long, equalling or slightly exceeding the stamens, glabrous, straight, diverging from each other, truncate at the apex, stigmatic on the apex or on the oblique inner angle. Samara 38-45 mm long, sparsely tomentose on the nut at maturity, otherwise glabrate; dorsal wing 25— 31mm long, 15-18 mm wide, extending over the adaxial side of the nut, the abaxial margin bent abruptly upward beyond the nut; nut 12-15 mm long, 7-10 mm high, the sides bulging and without winglets or crests, the veins prominent, the ventral aerole ovate to circular, 5 mm high, 4 mm wide. BRAZIL. Rio de Janeiro: 1832 fl, Ackermann (BR); 1851 fl, Andersson (S); 1847 fl, Forssell 111 (S); Copacabana, 29 Oct 1867, Glaziou 2116 (BR, C, P); Corcovado loco umbroso, 15 Sep 1874 fr, Mosén 2441 (S); Copacabana, Aug 1862 fr, Nadeaud (P). This species is endemic to the immediate area of Rio de Janeiro. I have described it in full here because Niedenzu’s description, under Banuisteria, is inadequate. I have seen no collection made more recently than 1874, which makes me fear that the species is extinct. However, perhaps it is hiding in some “loco umbroso” at the foot of Corcovado. Heteropterys sanctorum W. R. Anderson, sp. nov. Fig. I3. Liana lignosa. Lamina foliorum majorum 18-25 cm longa, 8-11 cm lata, subtus pertinaciter sparsim sericea, margine aliquot glandulis parvis munita; petiolus 15-22 mm longus, basi biglandulosus. Inflorescentia in ramis vetustiori- bus portata, 4-8 cm longa. Pedicellus 6.5-9 mm longus, 0.5—-0.8 mm diametro, sessilis. Sepala appressa per anthesin, 4 lateralia 6-8 glandulas ferentia. Petala flava, valde reflexa, inter se similaria. Woody vine, the stems sericeous to glabrate. Lamina of the larger leaves 18-25 cm long, 9-11 cm wide, ovate, elliptical, or obovate, mostly acuminate at the apex, slightly revolute at the magin, cuneate, obtuse, or rounded at the base, initially sericeous but soon glabrescent, with persistent scattered hairs below at maturity, bearing a series of small sessile marginal or very slightly (to 1.5 mm) submarginal glands, often in sinuses, the lateral veins visible above, lateral veins and reticulum prominent or at least visible below; petiole 15-22 mm long, sericeous to glabrate, biglandular at the base, the glands 2 mm in diameter; stipules not found. Inflorescence borne on old, leafless stems, axillary to fallen leaves, 4—8 cm long, sericeous, an open unbranched raceme of stalked or sessile umbels of 2-4 flowers, with some flowers borne singly on the axis, the small bracts (2 mm long) bearing 2 large glands; floriferous bracts and bracteoles ca 0.5 mm long, ovate, sericeous, eglandular, persistent; floriferous peduncle none. Pedicel 6.5-9 mm long, 0.5—-0.8 mm in diameter, sericeous. Flower ca 12 mm in diameter. Sepals 1.5 mm long beyond the glands, I-1.5 mm wide, triangular, appressed in anthesis, abaxially sericeous, adaxially glabrous, the anterior eglandular, the lateral 4 bearing 6-8 glands (4 on the posterior-lateral 2, 1-2 each on the anterior-lateral 2), the glands 1.4-1.7 mm long, circular or elliptical. 13] Petals yellow, glabrous, all completely reflexed, the lateral 4 with the claw 1.5- 1.8 mm long, the limb 3.5-4 mm long, 2.7 mm wide, obovate, erose or denti- culate, eglandular, the posterior like the others or the limb slightly wider. Fila- ments 2.2-2.5 mm long, subequal, connate in the proximal Imm, glabrous, those opposite the Sa ecli pressed inward, those opposite the petals leaning out- ward; anthers 1-1.5 mm long, slightly longer opposite sepals than petals, glabrous, reflexed. Ovary 1.5 mm high, loosely sericeous; styles 2-2.5 mm long, glabrous, strongly divergent, truncate at the apex, stigmatic on the upper- internal abe Fruit unknown TYPE: BRAZIL. Bahia: Sao Jose, estrada a Una, Pedra branca, mata, 6 May 1971 fl, 7. S. dos Santos 1603 (CEPEC, holotype; MICH, isotype). PARATYPE: BRAZIL. Bahia: Uruguca/Ubaitaba, Apr imm fl, Santos 761 (CEPEC). This species is named in honor of Talmon Soares dos Santos, who collected both the type and the paratype. Mr. Santos has made many fine collections in a previously neglected region and thereby contributed much to Brazilian botany. Heteropterys sanctorum is most similar to H. transtens Nied., which seems to be endemic to the area of Rio de Janeiro. They are compared in the characters known for both in the following couplet: Inflorescence 4-8 cm long, without long lateral branches, borne on old, leafless stems; larger laminas 8-11 cm wide, with all glands nearly or quite marginal; petiole 15-22 mm long. H. sanctorium. Inflorescence 15-30 cm eae compound, axillary and terminal on currently leafy stems larger laminas 11-15 cm wide, bearing a row of glands 5-15 mm inside the margin: petiole 30-50 mm one . dransiens. Heteropterys marleneae W. R. Anderson, sp. nov. Bign38. Liana lignosa. Lamina foliorum majorum 15-17 cm longa, 7.3-8.5 cm lata; petiolus 13-16 mm longus, 3-4 glandulis immersis munitus. Inflorescentia ae paniculata, 10-20 cm longa, floribus in Reeog or acem 2-6 cm longis, 6-26-floris, tomentoso-velutinis portatis; bracteae I-1.5 mm longae lataeque, 2 glandulis prominentibus munitae, persistentes; pedunculus nullus vel interdum usque ad 0.5 mm longus; bracteolae bracteis similares sed minores et eglandu- losae. Pedicellus 5 mm longus. Sepala apice revoluta. Woody vine. Lamina of the larger leaves 15-17 cm long, 7.3-8.5 cm wide, elliptical or obovate, obtuse or abruptly short-acuminate at the apex, flat at the margin, rounded at the base, glabrous (at maturity), bearing below in a row on each side 3-4 impressed gle ands 3—9 mm from the margin, the lateral veins and ee cross-veinlets prominent below, obscure above; petiole 13-16 mm long, tomentose to glabrate, bearing 3-4 glands 1.5-2 mm in diameter, partially immersed in the thick petiole; stipules not seen. Inflorescence an axillary panicle 10-20 cm long, dark brown tomentose-velutinous, simple or compound, with the flowers ultimately borne in pseudoracemes 2-6 cm long containing 6-26 decussate flowers; floriferous bracts 1-1.5 mm long and wide, broadly tri- angular, bearing 2 prominent glands at the base, abaxially tomentose, persistent; peduncle none or occasionally up to 0.5 mm long; bracteoles like the bracts but smaller and eglandular. Pedicel (in fruit) ca 5 mm long, velutinous with a mix- ture of short T- and Y-shaped hairs, distally enlarged. Sepals (in fruit) 2.5 mm long, 2mm wide, triangular, revolute at the apex, abaxially appressed-tomen- tose, adaxially glabrous, the anterior eglandular, the lateral 4 bearing 8 glands 1.8—2 mm long, elliptical, separated on the sepals. Petals and stamens unknown. Style bearing a short pedaliform apical-dorsal extension. Immature samaras up 132 to 36 mm long, widely spreading but the wings bent upward beyond the nut, scurfy-tomentose proximally; dorsal wing ca 30 mm long, 10 mm wide, straight or curved; nut 8-10 mm long, 4—5 mm high, without lateral crests or wings. TYPE: BRAZIL. Amazonas: Lago do Castanho-Mirim, estrada da Petrobras, mata virgem de terra firme, solo argiloso, humoso, meio umido, 23 Jun 1973 imm fr, Albuquerque, Coelho, & Mello 839 (INPA 39180, holotype; MICH, isotype). Heteropterys marleneae belongs to the natural but difficult subgenus Para- banisteria, in which the sepals are always revolute at the apex. It will undoubtedly prove to have yellow petals like the rest of the subgenus. In that group it is unusual in having the short, distally enlarged pedicel sessile or subsessile. It is also notable for its compound inflorescence with dark brown, scurfy, tomentose- velutinous axes, and for the short persistent bracts and bracteoles, the bracts bearing two large glands. This distinctive and interesting species is named in honor of Dr. Marlene Freitas da Silva, Curator of the herbarium at INPA in Manaus. Under her careful supervision that collection has become one of the most important research herbaria in South America. JANUSIA All students of the Malpighiaceae have agreed that Janusia Adr. Jussieu and Schwannia Endlicher are closely related genera. They really differ only in these characters: sae 6 (5 opposite the sepals plus | opposite the posterior petal); anthers densely tomen- tose; stout vines; cleistogamous flowers absent. Schwannia. Seamens “5 (op posite the cee or fewer; anthers glabrous; slender vines; cleistogamous owers present in | specie fanusia. I have now encountered a new species that is perfectly intermediate between the two groups. It is generally very similar to /. guaranitica in habit and in possessing cleistogamous flowers, but it has six fertile stamens. Its anthers are sparsely hairy. It is therefore necessary to unite the two genera. The two generic names seem to have been published essentially simultaneously, i.e. both in April 1840 (Stafleu & Cowan, 1976; 1979), and they have never before been con- sidered synonyms, so I must select one of the two names for the genus. I here choose Janusia Ady. Juss., for reasons that I shall discuss in some detail in my monograph of the genus. It is also necessary to select a lectotype for Janusia, and [ here choose Janusta guaranitica (St. Hil.) Adr. Juss. Below are published several new combinations that are necessary when Janusia and Schwannia are combined, and the new intermediate species is described. Janusia janusioides (Adr. Juss.) W. R. ee comb. nov, Fimbriaria janustoides Ady. Juss., Ann. Sci. Nat. 2° Sér. Bot. 13: 250. 1840. Janusia lindmanii (Skottsberg) W. R. Anderson, comb. nov. Schwannia lindmani Skottsberg, Kongl. Svenska Vetenskapsakad. Handl. 35(6): 29 Janusia malmeana (Nied.) W. R. Anderson, comb. nov. Schwannia malmeana Nied. in Engler, Pflanzenr. IV. 141: 540. 1928. Janusia mediterranea (Vell.) W. R. Anderson, comb. nov. Banisterta mediterranea Vell., Fl. Flum. 191 [text]. 1825 [1829, fide Carauta, 1973]; 4 pl. 162 [atlas]. 18: 133 This is surely an older name for the species widely known as Schwannia elegans (Adr. Juss.) Adr. Juss. Vellozo’s description and figure leave little room for doubt as to its identity; note especially these features: six stamens, one style, red ciliate petals with white hairs, elongated white calyx glands, growing in inland thickets. [ know of no other member of the Malpighiaceae that this could be. Janusia schwannioides W. R. Anderson, sp. nov. Fig. 14. Planta volubilis, caulibus gracilibus. Lamina foliorum majorum 3—4.4 cm longa, 1.3-2.2 cm lata, elliptica, basi plerumque rotundata, apice rotundata et apiculata, subtus prope basim 2 glandulis disciformibus stipitatis munita; petiolus 6-10 mm longus, eglandulosus. Flores chasmogami in umbellis axillari- bus (2—) 3-4-floris portati; pedunculus 8-15 (-21) mm longus, apice vel paulo infra apicem bibracteolatus. Pedicellus 7-16 mm longus. Petala glabra, profunde laciniata, limbo 6—-12.5 mm longo, 6-13 mm lato. Stamina 6, subaequalia; fila- menta 4.5-6.5 mm longa, 3 sepalis anticis opposita libera, 3 sepalis posticis et petalo postico opposita ’%—%4 connata; antherae 0.8-1.2 mm longae, paucipili- ferae. Flores cleistogami 3 carpellis muniti. Samara basi rudimento alarum lateralium munita. Vine with slender twining stems, forming thickets in shrubs to 3 m high; stems loosely sericeous to velutinous, glabrate in age, with an understory of thinner, shorter white hairs and an overstory of stiffer, longer golden hairs, these varying from strongly appressed to strongly spreading. Lamina of the larger leaves 3-5.2 cm long, 1.3-3 cm wide, elliptical or narrowly elliptical, mostly rounded and apiculate at the apex, slightly revolute at the margin, broad- ly cuneate or rounded at the base, sericeous above at first but soon glabrescent with some hairs persistent on the midrib, persistently sericeous, appressed- tomentose, or tomentose below, or belatedly glabrescent, bearing at or somewhat above the base below 2 stalked disciform glands 0.25—0.5 mm in diameter with stalks ca 0.3 mm long, the lateral veins and reticulum obscure on both sides; petiole 6-15 mm long, loosely sericeous to glabrate, eglandular, 0.4—0.5 mm in diameter; stipules 0.5-0.7 mm long, triangular, borne on stem at base of petiole. Chasmogamous and cleistogamous flowers both borne in axillary umbels or terminating short lateral shoots, produced simultaneously but mostly not in the same umbels. Chasmogamous flowers borne in umbels of (2—) 3-4 flowers, witha common stalk 5-25 mm long usually bearing 2 bracts or small leaves; floriferous bracts [subtending floriferous peduncles] I-1.8 mm long, 0.3-0.5 mm _ wide, narrowly elliptical, membranous, glabrous or bearing a few hairs, eglandular; peduncle 8-15 (-21) mm long, 0.3-0.4 mm in diameter, loosely sericeous to glabrate; bracteoles like the bracts, appressed, usually borne at apex of peduncle, ee aie slightly below. Pedicel 7-16 mm long, loosely sericeous to glabrate except persistently sericeous at the apex, proximally slender like the peduncle but distally enlarged to 1 mm or more in diameter at the apex. Sepals 4-6 mm long, 1.5-2.7 mm wide, elliptical or obovate, rounded at the apex, thin, strongly incurved or inrolled in anthesis, bearing a few straight sessile hairs abaxially, ciliate on the margin at the apex, adaxially glabrous, the anterior eglandular and narrower, the lateral 4 biglandular, the glands green , 1.5—-3 mm long, narrowly elliptical. Petals carrot-yellow, glabrous, deeply fimbriate-laciniate (the divisions at least | mm long), eglandular; 4 lateral petals widely spreading, with the claw 2.5-3.5 mm long, the limb flat and nearly circular or broadly cuneate at the base, 6-12.5 mm long, 6-13 mm wide; posterior petal with the thick claw erect, 3.3- 4.5 5 mm long, the limb strongly reflexed, circular or obovate, 6-11 mm long, 5-12 mm wide. Stamens 6 (5 opposite the sepals plus | opposite the posterior Bee, all fertile, subequal; filaments 5—6.5 mm long opposite sepals, 4.5-5.3 mm 134 ane | SS ; wie ai re & . 14. Janusta schwannioides. a) habit glands, x 10; ¢) stip ee > 1) samara, x 1.5 t, X 0.5; b) base of comes abaxial side, to show hairs and pules, x 10; d) bud of chasmogamous | flow 3; e) cha ciara flower, 5; a ndroecium and gynoecium of chasmogamous ee eee anterior sepal to left and claw of posterior petal to right, | anterior-lateral sepal removed, 5; g) anthers of casa flower, ee view (left) and adaxial view (right), x 15; h) eae flower, * 10; ee cane flower with samaras enlarging and sepals pulled ‘back to show aan: pet ae 2 stam style, x 10; j) infruc aed with fruits all from ercetany! flowers, X fen eee neces x type. 2: k) fruit froma sin ae >: m) embryo, X 5. Drawn by Karin Douthit from the 135 long opposite posterior petal, glabrous, the anterior 3 free, the posterior 3 one- half to three-fourths connate, all crowded around the style and curved down- ward toward the anterior sepal and then upward; anthers 0.8-1.2 mm long, alike, often reflexed, bearing a few appressed hairs on sides of locules and sometimes a tuft of longer, twisted hairs at apex or base. Carpels 1—-1.5 mm high, borne on a low common receptacle, densely sericeous; style 1, borne on the anterior carpel, 5-6.5 mm long, glabrous, stout, lying in plane of symmetry of flower, curved downward toward anterior sepal and then abruptly upward at the apex, the terminal stigma often slightly oblique, extending just beyond the anthers. Cleistogamous flowers borne singly or, usually, in umbels of 2-4 flowers, with a common stalk 1-5 mm long; bracts and bracteoles like those of chasmogamous flowers but mostly shorter; peduncle 2-8 mm long; cleistoga- mous flowers occasionally replacing 2 of the 4 flowers in a chasmogamous umbel. Pedicel 3-8 mm long. Sepals 1.5 mm long, 0.7 mm wide, eglandular, appressed, narrowly elliptical, abaxially sparsely sericeous, adaxially glabrous. Rudimentary petals 1-5, 0.1-0.4 mm long, non-vascularized, concealed be- tween bases of sepals. Stamens 2, opposite posterior-lateral sepals, with fila- ments 0.15—-0.2 mm long and anthers 0.25-0.3 mm long, indehiscent. Gynoe- cium with all 3 carpels developed and fertile; style 1, borne on the anterior carpel, ca | mm long, hook-shaped, bent over between posterior carpels to bring stigma into contact with anthers. Fruit, whether derived from chasmogamous or cleistogamous flowers, comprising 3 samaras, or fewer through failure of | or 2 carpels to mature; samaras separating from a low, rounded or obscurely 3-sided torus | mm or less high; samara a 20 mm long, loosely sericeous proximally; dorsal wing 12-15 mm long, 5.5-7 mm wide, with a rounded projection 1-2 mm high at the adaxial base; nut ca 7 mm high, 4 mm long, laterally flattened; lateral wings restricted to base of nut, partially to completely connate, forming an apparent flattened extension of the nut 2-3 mm long and wide at right angles to dorsal wing; cartilaginous carpophore well developed and functional, extending from receptacle down line of fusion of lateral wings. Embryo with the cotyledons subequal, flattened, folded back in the distal third. : BRAZIL. Bahia: Trail from Lagoinha (which is 5 km SW of Delfino) to Minas do Mimoso (41°16' W, 10°27’ S), elev 640 m, caatinga 1.5 km NW of Lagoinha, 7 Mar 1976 fl/fr, Anderson 11748 (MBM, holotype; K, MICH, NY, RB, isotypes). PARATYPES: BRAZIL. Bahia, Mun. Pocoes: 2.5 km W of Pocoes on road to Bom Jesus da Serra, dense roadside brush, Mar fl/fr, Anderson 12514 (MICH, RFA); Km 10 da estrada que liga Pocoes (BR-116) ao povoado de Bom Jesus da Serra (ao W de Pogoes), transigao entre Mata de Ci upo e Caatinga, 750 m, Mar fl/imm fr, Mor? et al. 9517 (CEPEC, MICH). Janusia schwannioides resembles J. guaranitica in its habit, inflorescence, samaras, and production of cleistogamous flowers, but in the latter species the petals are subentire, there are only five stamens, the stamens and style are + erect and straight, and the cleistogamous flowers have only two carpels. The other species that resembles J. schwannioides is J. janusioides sensu stricto. It has shorter petioles, floriferous peduncles very short or absent, smaller flowers with the less deeply laciniate petals hairy adaxially on the claw, shorter, less curved androecium and gynoecium, and hairier anthers; it does not produce cleisto- gamous flowers. MASCAGNIA In the last volume of these Contributions (Anderson, 1980), I pointed out that Banisteria riedeliana Regel was an older name for Mascagnia metallicolor 136 Niedenzu, and made the appropriate new combination. Recently, while studying in oe the types of species described by Adrien de Jussieu, I discovered that there are two stll older names for this species. The new combination and a full eer for this species as I now understand it are given below. See the 1980 paper for a description and illustration of the species. Mascagnia ocr aha (Adr. Jussieu) W. R. Anderson, comb. nov. Hiraea renidens Adr. Jussieu, Ann. Sci. Nat. 2° Ser. Bot. 13: 26. 1840. Hiraea herpes Adr. Jussieu, Arch. Mus. Hist. Nat. 3: oe L435. Banisteria riedeliana Regel, Ind. Sem. Hort. Bot. Petropol. 17. 1855 Tetrapterys renidens (Adr. Jussieu) Grisebach in Martius, Fl. Bras. 121), 88. 1858. sa ya heteropetala (Adr. Jussieu) Grisebach in Martius, Fl. Bras. 12(1): 88 858. wean metallicolor Niedenzu, Arb. Bot. Inst. Lyc. 3: 20. 1908. Mascagnia riedeliana (Regel) W. R. Anderson, Contr. Univ. Mich. Herb. 14: 21. 1980 ACKNOWLEDGMENTS My studies of the Malpighiaceae are currently supported by a grant from the National Science Foundation to The University of Michigan (DEB 81- 03522); that grant is also helping to pay for this publication. I am most grateful to Dr. Alicia Lourteig, whose hospitality during the summer of 1981 made my lengthy visit at the Muséum National in Paris both pleasant and profitable. I extend my thanks also to Drs. Carmen Cristobal, Antonio Krapovickas, T. Myndel Pedersen, Gert Hatschbach, and Paulo Occhioni for their friendly as- sistance during my recent visits to Argentina and Brazil, and to the many cura- tors who sent specimens for my study as gifts or loans; their ee are repre- sented by the following acronyms: BM, BR, C, CEPEC, CTES, F, G, GH, IAN, INPA, K, LA, LE, M, MBM, MG, MO, NY, P, R, RB, S, SP, U, ie US, W. Karin Douthit drew the beautiful illustrations, with her noun combination a skill and attention to detail. LITERATURE CITED dency William R. 1980. Notes on Mascagnia in South America. Contr. Univ. Mich. Herb. |4: 17-23. 1981. ig poe m ‘The botany of the Guayana Highland—Part XI. Mem. New ork Bot, Gard. 32: 21-305 ence pd carl etl be io The text of Vellozo’s Flora Fluminensis and its effective date of publication. ‘Taxon 22: 281-28 Niedenzu, ee 1928. eee in A. Engler. Das Pflanzenreich IV. 41: 1-870. eaneNlle N. Y. I ne . Review of the species of Byrsonima occurring in British Guiana. Kew Bull. 311- Stafleu, ee A, : "Ri chard S. Cowan. 1976. Taxonomic Ae sceOnd, edition. Vol. I: A-G. ——————.. 1979. Tone literature, second edition. Vol. Contr. Univ. Mich. Herb. 15:137-140. 1982. ON MEIOTHECIUM (SEMATOPHYLLACEAE) William R. Buck New York Botanical Garden Bronx, NY 10458 The Sematophyllaceae are unusual among the pleurocarpous mosses in having many genera, most of which are mono- or oligotypic. Of approximately 53 genera, about 35 have four or fewer species. Often the genera are based on small peristomial differences, such as endostomial development. Some genera are separated on leaf papillosity differences, whereas other genera, such as Acroporium, permit this variation within a single species. The degree of alar development, which runs a full spectrum, is often the difference between two genera. The Sematophyllaceae are a relatively recent family, as are most of the pleurocarps, probably derived from a Northern Hemisphere Hypnobryalean stock (Vitt, 1982). Thus, it is assumed that the Sematophyllaceae, a predomi- nantly Southern Hemisphere family, migr ated and radiated after continental drifting, which may explain the generic proliferation. Although there can be little doubt that some of the monotypic genera truly represent isolated endpoints of evolution, it is difficult to believe that two-thirds of the genera in the family fall into that category. The discrepancy is probably a result of both poor obser- vation and a lack of understanding the values of morphological variation. The fact that most of the taxa occur in the tropics complicates the situation further because many tropical species are known only from type collections, which are often scanty or lacking mature sporophytes. The confusion is additionally com- pounded by some of the larger genera of Sematophyllaceae which are surely polyphyletic. Mevothecitum combines many of the problems outlined above. The genus is characterized by usually more or less ovate leaves with shortish cells; the alar cells are differentiated and somewhat inflated and colored, but not as extremely so as in many of the Sematophyllaceae. The capsules are erect to suberect, and the peristome is described as single with the exostome variously spaced and orna- ented. Brotherus (1908) recognized and described three sections: Eu- Mevothecium, Pterogonidiopsis and Pseudo-Meiothecium. Uhe three sections are sepa- rated mainly on the basis of sporophyuc characters, although there is some gametophytic correlation. Section Metothectwm, containing the majority of spe- cles, is recognized by widely spaced, papillose exostome teeth. The calyptra is decidedly roughened at its apex. The section Pterogonidiopsis, with only a few South American species and a single African one, has the exostome teeth closely placed and + smooth on the front surface except for prominent bars separating the individual plates. The calyptra is smooth. Pseudo-Meiothecium has plants more robust and an exostome whose teeth are closely placed. The ornamentation is described as papillose (Brotherus, 1908), but it is not like that of sect. Meio- thectum. Rather, in sect. Pseudo-Meiothecium the papillae are arranged in tight horizontal rows, the type of sculpturing often seen in reduced hypnoid peristomes (Buck, 1980). The four species (all of northern South America) h lave smooth calyptrae. et 137 138 Soa first appeared in the literature as a Schimper name “in herb. Mus. Par.” (Jaeger, 1877) in a catalogue with no description but with 33 species listed under it. In 1894 Brotherus described two new African species in the genus, P. usambarica and P. stuhlmannu, even though the genus had sull not been described nor had any indication been given in the literature as to how Schimper himself circumscribed it. In 1897 Brotherus described another African species, P. chloroclada. \t is obvious that at this time Brotherus had no intention of de- scribing Plerogoniella as a new genus. However, both Wik et al. (1967) and Farr et al. (1979) recognized P. chloroclada as a descriptio generico-specifica. In 1908 Brotherus placed Pterogoniella in synonymy under Meiothecium. Both P. usambarica and P. chloroclada were transferred to en P. stuhlmannii was transferred to Rhaphidostegium (= Sematophyllum). Very similar to Mevothecium is Mevotheciopsis. Metotheciopsis supposedly differs from Meiothecium by a double peristome and from the sect. Piero, gonidiopsis by the exostome teeth more strongly barred on the front surface. An examination of almost all South American taxa of Meiothecium, as well as a representative selection of Old World species, indicates that the genus 1S polyphyletic. Meiothecitum, described from Samoa (Mitten, 1868), is easily circum- scribed sporophytically. Che peristome of widely spaced, narrow, non-bordered, papillose exostome teeth and no endostome is surprisingly constant across the broad range of gametophytic variation. However, careful dissection of mature capsules indicates that the species of sections Plerogonidiopsis and Pseudo- Meitothectum not only have very different exostomes, but also have reduced endostomes. At times the segments can even be seen with a dissecting micro- scope projecting straight up between the recurved teeth. In the New World there seem to be only two true species of Meiothecium: M. boryanum (C. Mull.) Mitt. (syn. M. scabriusculum Besch. and M. wrightii (C. Mull.) Broth.) from the West Indies and M. revolubile Mitt. of Amazonian campinas. As noted above, the section Plerogonidiopsis is very close to Mevotheciopsis. Vhe supposed differences in the peristomial development in Meiothectopsis (including the presence of an endostome and the more strongly barred exostome) are based ona poor understanding of sect. Pterogonidiopsis. A comparison of the peristomes of Meiotheciopsis lageniformis (C. Mull.) Broth. and Meiothectum commutatum (C. Mull.) Broth., reveals that they are very similar, with the endostome of M. commutatum only somewhat more reduced. Gametophyucally the plants are also similar. Because of the resemblances a congeneric alliance is appropriate. — Meiotheciopsis commutata (C. Mull.) Buck, comb. nov. Neckera commutata C. Mull., Bot. Zeit. (Berlin) 15: 385. 1857. Pterogoniella commutata (C. Mull.) Jaeg., Ber. Vat. St. Gall. Naturw. Ges. 1875-76: 21 877, comb. inval. Weiihrann lenerum Mitt., J. Linn. Soc. London, Bot. 12: 470. 1869, syn. nov. Meiotheciopsis usambarica (Broth.) Buck, comb. nov. Pterogoniella usambarica Broth., Bot. Jahrb. Syst. 20: 208. 1894, nom. inval. Meiothectum usambaricum Broth., Nat. Pfl. 1(3): L101. 1908. Meiotheciopsis laxiuscula (Card.) Buck, comb. nov. Pterogoniella laxiuscula Card., Rev. Bryol. 36: 51. 1909, Meiotheciopsis longidens ( (Angstr. ) Buck, comb. nov. Potamium longidens Angstr., Ofv. Forh. K. Sv. Vet.-Akad. 33(4) ~ 1876: Pterogonium longidens C. Mull. ex Angstr., loc. cit., nom. nud. in syn. Plerogoniella longidens ( (Angstr.) ) Jaeg., Ber. Tat. St. Gall. Naturw. Ges. 1875- 76: 211. 1877, comb. inval. Metothecium longidens (Angstr.) Broth., Nat. Pfl. 1(3): LLO1. 1908. 139 Meiotheciopsis lagenifera (Mitt.) Buck, comb. nov. Meiothecium lageniferum Mitt., J. Linn. Soc. London, Bot. 12: 470. 1869. Pterogoniella lagenifera (Mitt.) Jaeg., Ber. ‘Vat. St. Gall. Naturw. Ges. 1875- 76: 207. 1877, comb. inval. Mevotheciopsis is probably most closely related to Schraderella of Venezuela. Type material of the sole species, S. pungens C. Mull., at NY has no peristome, but based on the illustration in Brotherus (1908) it (presumably) differs in the absence of an endostome and exostome teeth paired. Meiothecium sect. Pseudo-Meiothecium has a reduced hypnoid exostome and a fairly conspicuous endostome. The leaf cells are often short, but in some species they are linear. e species assigned to this section are not related to either Meiothecium s.str. or Metotheciopsis. Rather, they seem most closely related to Sematophyllum caespitosum (Hedw.) Mitt. and its allies. The group is characterized by mostly ovate, short- pointed leaves with relatively short leaf cells. The setae are short, and the capsules are mostly erect. The peristome is scarcely reduced. Although widespread, S. caespitosum should not be interpreted as widely as Dixon (1920) suggested. Leaf shape and areolation appear constant. The following combinations are thus proposed: Sematophyllum aptychoides (C. Mull.) Buck, comb. nov. Neckera aptychoides Schlieph. ex. C. Mull., Bot. Zeit. (Berlin) 15: 382. 1857. Fs aptychoides (C. Mull.) Mitt., J. Linn. Soc. London, Bot. 12: 471. 86 e terogoniella a (C. Mull.) Jaeg., Ber. Vat. St. Gall. Naturw. Ges. 1875-76: 9. 1877, comb. inval. Meiothecitum sulcatum Mitt., J. Linn. Soc. London, Bot. 12: 470. 1869, syn. nov. Sematophyllum fabronia (Besch.) Buck, comb. Mevothecium fabronia Besch., J. Bot. Mesa res 1891. Pterogoniella fabronia (Besch.) Par., Ind. Seal 1047. 1898, comb. inval. Sematophyllum negrense (Mitt.) Buck, comb. nov. Mevothecium negrense Spruce ex Mitt., J. Linn. Soc. London, Bot. 12: 471. 9. Pterogoniella negy ensis (Mitt.) Jaeg., Ber. Tat. St. Gall. Naturw. Ges. 1875-76: 209. 1877, comb. inval. Sematophyllum mittenianum Buck, nom. nov. Mevothecium andinum Mitt., J. Linn. Soc. Bot. 12: 470. 1869 non Semato- phyllum andinum Mitt., op. cit. 483. It may be possible to segregate Sematophyllum caespitosum and closely related ecies generically. The short leaf cells and short setae with erect capsules are eee in Sematophyllum. However, until this large, widespread genus is better known it seems best to retain it in a catch-all genus. LITERATURE CITED Brotherus, V. F. 1894. Musci ea . Bot. Jahrb. Syst. 20: 176-218. —————.. 1897. Musci africani. II. Bot. Jahrb. Syst. 24: 232-284 8. Seinatophllaceae 7 A. Engler & K. Prantl, Die naturlichen Pflanzenfamilien 1(3): 1098- 112 Buck, W. R. 1980. A ge eneric revision of the SHMOCO NEES 2: J. Hattori Bot. Lab. 48: 71-159. Dixon, H. N. 1920. Rhaphidostegium caespitosum (Sw.) and its affinities. J. Bot. 58: 81-89. 140 Farr, E. R., J. A. Leussink & F. Staffleu. 1979. Index Nominum cg jail (Plantarum). Volume IIL. Paria Innes Regnum Veg. 102: 1277-18 Jaeee A. 1877. Genera et species muscorum Via ae at disposi seu. Adumbratio florae uscorum totius orbis terrarum. VII. Ber. ‘Tat. yall. 1875-76: 201-371. Mitten, ue 1868. A list of the Musci collected a the Rev. Thom Powell in the Samoa or Navigator's ds. J. Linn. Soc. London, Bot. 10: 166-195, tabs. 5, 6. Vitt, D uf 1982. Bryopsida. /n S. P. Parker (ed.), Synopsis - classificauon of living organisms 1: 07-336. McGraw-Hill, New York. Wik, ay van der, W. D. a el P. A. Florschutz. 1967. Index Muscorum. Volume IV (P-S). Regnum Veg. 48: 1-604 Contr. Univ. Mich. Herb. 15:141-162. 1982. THE NORTH AMERICAN DISTRIBUTION OF THE GENUS DRYOPTERIS Thomas M. Carlson Division of Biological Sciences W. H. Wagner, Jr. Herbarium and Division of Biological Sciences The University of Michigan Ann Arbor, Michigan 48109 INTRODUCTION The Woodterns, Dryopteris, are conspicuous, well collected ferns of mainly forests and swamps that occur over most of North America except the Great Plains. They become rare and sporadic in southwestern United States and in southern Florida. The 15 sexual taxa display a diverse array of mainly temper- ate, boreal, and arctic distribution patterns that appear to have been greatly influenced by Pleistocene glaciations. These patterns contribute toward an understanding of the complex phylogenetic relationships of these plants. There are nine diploid taxa and six polyploid taxa. Comparisons of possibly related Old World species provides insight into past distributions and migration patterns. Five of the polyploids are perhaps the most interesting in terms of glacial his- tory; three of these gTOW exclusively in eastern North America. Glaciation may have played a major role in the origins of these allopolyploids by creating condi- tions conducive to hybridization. Two other allopolyploid Woodferns have amphi-Atlantic distributions, and Wagner (1971) proposed that they may have had a common parent (SS), now presumably extinct. As will be seen, similarities between the ranges of these two ferns support this suggestion. The first comprehensive treatment of the overall geography of ferns was by Christ (1910). However, there are few in-depth studies (Smith, 1972; Tyron, 1969, 1970, 1971; Wagner & Wagner, 1965). MATERIALS AND METHODS The ranges are plotted of 13 of the 15 sexual taxa of Dryopteris occurring in North America, north of Mexico (including the two primarily Mexican taxa whose ranges barely extend into the southwest United States). Some of the distribution maps include localities in Iceland, a land mass often considered to be part of the Old World. The square symbols on the maps (Figs. 1, 3, 6, 8-19) represent herbarium specimens that we both examined and annotated. The dots represent localities learned from reliable literature or through correspondence with other pteridologists. Many of the specimens represented by dots were also seen by Wagner. RESULTS Dryopteris fragrans (L.) Schott (Fragrant fern) is a diploid that prefers shaded or exposed and usually north facing cliffs (Wherry, 1961; Carlson, 1979). Its 141 142 =) Ve oo fa 7a vee vr ‘ goo Ne , wre it CK, 4 *. ory f om by “ * ¢ Ven a4 x aa, Uh Se. % 2 ie ie ae ‘ 4 ‘ey “ iv, my y. Ce < A x he & -s nee sy | ile ae re 5 a: : Ars "ea wy 4 / ~ = nw eC BY f A ( ey a a x I | poe (\ ee e D. FRAGRANS | hay By FIG. 1. Dryopterts fragrans range in North America —- = AP NO. COPYRIGHT | 80 | AMERICAN MAP CO, INC. | | j | co ay wet s | re | aes al y) 4 ) = iS - 23 [ / 170 é r aN ¥ U en Py 1 [ ‘ = 5 | iat | Lt / a : 268 FIG. “a. Dryopteris fragrans approximate global range 143 circumboreal-arctic range extends in northern latitudes through North America (Fig. 1) and across northern Asia and Siberia and southward into China, Korea, and Japan (Fig. 2), with disjunct populations in northern Finland and northwest Russia. The circumboreal Dryopteris filix-mas (L.) Schott (male fern) is a tetrapoloid. Its taxonomy in North America is poorly understood. It may be an autopoly- D. FILIX-MAS FIG. 3. Dryopteris filix-mas range in North America. 144 ploid (Wagner, 1971) or an allopolyploid (Britton, pers. comm.). It occurs in mountainous regions or in areas with exposed bedrock and/or talus slopes (Carlson, 1979) in most parts of its North American range (Fig. 3). In the west D. filix-mas extends through the Rocky Mountains from British Columbia to north- ern Mexico. Male fern also has a disjunct population in the San Bernadino Mountains in southern California. In eastern North America D. filix-mas is scat- tered from the Great Lakes to Greenland and Iceland. Dryopteris arguta (Kaulf.) Watt (Coastal Woodfern), a diploid of rocky ravines and partially shaded slopes (Grillos, 1966), has a Cordilleron range extending along the west coast of North America from northern Baja Calfornia to southern British Columbia. It also has populations in the High Sierras of California as well as disjunct populations in the mountains of south-central Arizona in Pimal and Gila counties. Jermy (2n (itt., 1979) stated that when he examined specimens of this species from the University of California (Berkeley) herbarium, what appeared to be two distinct spore sizes were found. He suggested that the western American D. arguta May contain a tetraploid cytotype as well as the known diploid. Another variable character is broad and narrow frond forms. There are three frond types (narrow, normal and broad) (Fig.’s 4A, 4B, 4C). As illustrated (Fig. 5), the range of the narrow form is restricted to the High Sierras of California, while the broad form occurs on Santa Catalina and Santa Cruz Islands and along with one locality in Humboldt Co., California. The normal form grows predominantly along the coast but its range also overlaps with those of the narrow and broad forms. ‘The two extreme forms may only be environmentally induced variations of D. arguta produced by local conditions. Dryopteris expansa (Presl) Frazer-Jenkins (Spreading Woodfern), best known in previous botanical literature as D. dilatata (Hoffm.) Gray or D. assimilis S. Walker is a diploid of cool talus slopes and swamps, mainly in woodlands (Carlson, 1979). It exhibits an amphioceanic distribution pattern (Fig. 7) with populations in north temperate and sub-boreal eastern and western North America (Fig. 6), Europe, and eastern Asia. Much confusion has existed regarding the North American range of Spread- ing Woodfern because it has been lumped with other species in the D. carthusiana FIG. 4. Dryopteris arguta: silhouettes of herbarium specimens. 145 complex, which have collectively been called D. austriaca Jacq.) Woynar (often also referred to as D. dilatata). For example, Hultén (1968) illustrated the range of D. austriaca as continuous between the Pacific and Atlantic oceans across Canada. owever, careful examination and revision of western Canadian Dryopteris her- H f barium specimens an the subsequent mapping of their localities results in a pattern (Fig. 6) that supports Britton’s (1972) suggestion that in western Canada a st as western the range of the Spreading Woodfern extends only as far ea Alberta. D. expansa has also been confused with the allotetraploid D. campyloptera Clarkson (Mountain Woodfern) of eastern North America. ae x D. ARGUTA. COMPLEX 30 @ NORMAL FORM O NARROW FORM % BROAD / . 5. Dryopteris arguta range. 146 oe of Cabrcors ats = = PA Rt 4 Shee #6 ‘ en : ee EO) , yt H 4 . : ae ' ( ; Ei “ee i of i . \ ‘ Ps oe iM) 7 a 4 ‘| ey, > 0 EXBANSA os rs expansa range in North America. , | | SCHOOL SERIES { Outline Map RLD MAP NO.1723 COPYRIGHT Ri AMERICAN MAP CO. INC. FIG. 7. Dryopteris expansa approximate global range, 147 Four diploid Woodferns have ranges restricted to eastern North America. The two most common are Dryopteris intermedia (Muhl.) Gray (Glandular Wood- fern, Florist Fern) and D. marginalis (L.) Gray (Marginal Woodfern). D. intermedia grows predominantly in rich woods while D. marginalis prefers drier more up- land forests, particularly ones with rocky slopes (Wherry, 1961; Carlson, 1979). D. intermedia extends (Fig. 8) southward through the Appalachian Mountains into northern Georgia and Alabama and westward into eastern Missouri, central Iowa and northern Minnesota. In Canada it grows along the Great Lakes and the St. Lawrence Seaway, in Newfoundland, Nova Scotia, New Brunswick, southern Quebec and southeastern Ontario. The northern part of the range of D. marginalis (Fig. 9) is very similar to that of D. intermedia but it does not extend into Minnesota. However, the south Marginal Woodfern extends much farther west through the Plains states of Missouri, Arkansas, Nebraska and Oklahoma, per- haps because it tolerates drier habitats than does the Grandular Woodfern. Dryopteris goldiana (Hooker) Gray (Goldie’s Woodfern), a fern of extremely rich mesic woodland habitats (Wherry, 1961; Carlson, 1979), is another diploid Woodfern that is restricted to eastern North America, but it is considerably less common and its range (Fig. 10) is smaller than that of D. intermedia or D. margin- alis, presumably because its habitat is much more specific. The southern bound- 1 ' ' ! 4 f ' t FIG. 8. Dryopteris intermedia range. e qe sere 9% ij : i ort ; oe. 3 aoe oe Oe eon ~"ee fie ee : Sh - re “eee “OU 2°, e eeee “ Seceee” He ee ee . eote, e | ieee? ~. fe y i@ 0” | a ee D. MARGINALIS | } an \ ; | \ ae ‘ . ) | fs \ FIG. 9. Dryopteris marginalis range. aries of its range are similar to those of D. intermedia but it does not extend so far north; it reaches only southeast Ontario and southwest Quebec. D. goldiana does, however, extend farther west into Missouri than D. intermedia. Dryopteris ludoviciana (Kze.) Small (Florida Woodfern), is a diploid whose distribution (Fig. 11) is restricted to damp woods and swamps and rocky ledges in hammocks (Wherry, 1972) on the coastal plain and the lower piedmont of the United States from North Carolina to Florida and west to Louisiana. It does however, extend up into the Cumberland plateau in Alabama. Dryopleris celsa (Palmer) Small (D. goldiana x ludoviciana) (Log Fern) is an endemic allotetraploid which grows on rich hummocks and on rotting logs in swamps (Carlson, 1979). The present distribution of the Log Fern (Fig. 12 extends south from New Jersey to northern South Carolina and northern Alabama and west to northern Louisiana and western Arkansas. The range has two interesting disjunct areas, namely the vicinity of Rochester, New York region and southwestern Michigan. Disjunct populations of Woodwardia areolata (L.) Moore (Netted Chain-fern) and Lygodium palmatum (Bernh.) Sw. (Climbing Fern) are also known from southwestern Michigan. Several new localities for D. celsa have been found in recent years including additional localities in four south- western Michigan counties, and a specimen from Missouri originally thought to be D. clintoniana (Eaton) Underwood (Clinton’s Fern) was determined by us to be D. LUDOVICIANA FIG. 11. Dryopterts ludoviciana range. D. celsa. Single localities were also recently verified in both Illinois and Indiana, firsts for both states. Due to the morphological similarity between D. celsa and D. clintonianal, many D. celsa collections have been misidentified as D. clintoniana. As these mistakes are corrected, the known records of D. celsa may increase, particularly in the northern part of its range between its presently known dis- junct populations and the major area of its assumed range. FIG. 1 CELSA D. GOLDIANA HHMI ' Dp. LUDOVICIANA = | I | Z | J eee Gee | | ; ae) aT 3. Dryopteris celsa range with ranges of its parents (D. goldiana & D, ludoviciana). FIG. 15. Dryopteris clintoniana range with ranges of its parents (D. cristata & D. goldiana \ Pe | NN 7, a . b.c CLINTONIANA a CRISTATA hee GOLDIANA bays These misidentifications have also led people to believe that the range of Dryopteris clintoniana (D. cristana x goldiana), a North American allohexaploid, extends much farther south than it actually does. Montgomery (1976) showed that in New Jersey D. clintoniana does not reach below the glacial border. The southern boundary of its range (Fig. 14) in Pennsylvania and New York also follows the southern boundary of the ice sheet advance. [It has populations in northeast Ohio, northwest Indiana and much of the Lower Peninsula of Michi- gan. [ts range in Canada is similar to that of one of its parents (D. goldiana) (Fig. 10) since it grows in [southern] southeast Ontario, [southern] southwest Quebec, in addition to a single known locality on the Gaspe Peninsula. FIG. 16. Dryopteris campyloptera range. 153 Dryopteris campyloptera 1s an endemic allotetraploid whose parents are D. expansa and D, intermedia (Wagner and Hagenah, 1962; Wagner, 1963; Wagner, 1971; Gibby, 1977). Its distribution (Fig. 16) extends into southeastern Canada as far north as Labrador and southward down the Appalachians through the United States to Tennessee and North Carolina. The two allotetraploid species, Dryopteris cristata (L.) Gray (Crested Wood- fern) and D. carthusiana (Vill.) H. P. Fuchs (syn. D. spinulosa (Muell.) Watt) (Spinulose Woodfern) have amphi-Atlantic ranges. Because of their similar dis- tributions, their ranges are described together. Both D. cristata and D. carthusiana are common throughout the Great Lakes region and in New England. Their ranges (Figs. 18 & 19) extend as far west as eastern North Dakota, and both have disjunct poulations in the same region in western Nebraska. The southern boundary of the range of D. cristata runs through southern Iowa, northern Illinois, southern Indiana and southern Ohio. In the Appalachians it occurs further south, being known from West Virginia, Virginia and North Carolina. The area of D. carthusiana extends south to northern Arkansas, southeastern Missouri, southern Illinois, Kentucky and northern South Carolina. Herbarium i D. CAMPYLOPTERA -&- A _| D. EXPANSA = : 35, ‘) D. INTERMEDIA _Iillll ra) N ia aN 4 \ < ! NU \, 4} ar D. CARTHUSIANA Gs i AX a | | +. SCHOOL SERIES Oudine Map ORL R 3690 200 E at 090 ae va | MAP NO. 1723 2 COPYRIGHT | AMERICAN MAP CO., INC 4 — 2 cy « & i 10, io To, ise in FIG. 21. Dryopteris carthusiana approximate global range. 157 and (or) in Europe. A. C. Jermy (¢n litt, 1979) suggests that D. arguta may be closely related to the Old World D. villaru ida)-D. pallida complex of Europe and Asia. Such vicarious affinities suggest that this western North American range may be a relic of a once widespread arctotertiary distribution. Other eastern North American Woodferns that may have Old World vicarious affinities will be discussed below. Many of these may have had ancestral arctotertiary ranges oO which only isolated relicts at lower latitudes exist today. During the Pleistocene epoch fluctuating climatic and geologic conditions altered ranges in northern latitudes. The distributions of the five North American allopolyploids will be analyzed in respect to the ranges of their respective parents. Since the present ranges of the parents of D. celsa (D. goldiana, and D. ludoviciana) (Figs. 12, 11, 10) do not overlap (Wagner & Wagner, 1965), it seems likely that during the advances and retreats of the ice sheet the two ranges conjoined and hybrids were formed which went through polyploidization to form the fertile allotetraploid, D. celsa. When the ice sheet finally retreated D. goldiana migrated northward separating from D. lucoviciana, and the newly formed D. celsa established a range (Fig. 12, 13) that is latitudinally and altitudinally intermediate between those of its parents (Wagner, 1971). Dryopteris clintoniana (Fig. 14) probably resulted from the many disturbed habitats left by the glacier. These habitats allowed for its parents (D. cristata and D. goldiana) to become established in close proximity as their ranges followed the retreating ice sheet northward (Wagner, 1971). Hybrids formed which under- went polyploidization to form D. clintoniana. Its distribution is latitudinally inter- mediate between and narrower than its parents (Figs. 10, 14, 15, 18), restricted to land covered by deposits of glacial till. The southern border of its range faithfully follows the southern boundary of the ice sheet advance. The fern’s localization on glacial till may be a major factor limiting the expansion of its range to the south or north. The parents of D. campyloptera, D. expansa and D. intermedia, overlap in the northern Great Lakes, southwest Quebec, in the Gaspe Peninsula, Anticosti Island, north of Anticosti Island on the southern coast of eastern Quebec, and west of the Gaspe Peninsula on the northwest shore of the St. Lawrence River, and in the Saguenay River Valley. However, the primarily Appalachian distri- bution of D. campyloptera (Fig. 14) does not directly reflect these areas of over lap (Fig. 15). We suspect that several times during advances of the ice sheet the two parents mingled to a much greater extent as a result of the southward extension of their ranges. Hybrids between them went through chromosome doubling to produce D. campyloptera which then followed the retreating glacier northward and developed its present range, the boundaries of which are more northern than those of D. intermedia but much more southern than those of D. expansa. While D. campyloptera may occur in low woodlands in the north, it grows south- ward almost exclusively in high mountain forests. This supports findings (e.g., Beaman et al., 1962; Reese, 196 1a) that suggest that some polyploids are ee equipped than their diploid ancestors to inhabit extreme environments at higher altitudes. The two circumboreal (more specifically amphioceanic) allotetraploid spe- cies, Dryopteris cristata and D. carthusiana form a natural pair. Wagner (1971) suggested that these tetraploids have a common parent (“D. semicristata’) that possessed a hypothetical SS genome and is now presumably extinct. Among the similarities between these two ferns are leaves partially deciduous, triangular basal pinnae, and preference for moist, acidic lowland habitats. Perhaps the greatest similarity is in their ranges (Figs. 18-21), although a noticeable dif- ference 1s the more northerly range of D. carthusiana. This is not surprising since 158 the extant putative parent (D. intermedia, Fig. 8) of the former has a greater northern distribution than that of D. ludoviciana (Fig. 11) the parent of the latter. Since the range of each allopolyploid and one of its parents is known, the ap- proximate range of the hypothetical missing parent, “D. semicristata” can be estimated. Stebbins (1950) stated that there is a tendency for angiosperm polyploids to be more widely distributed than their diploid ancestors. This assumpuon. Is evidently not true in several primary temperate fern polyploid complexes. Hirabayashi (1974) found that distribution of the tetraploid Japanese Dryopteris are narrower than those of their diploid ancestors. Of temperate North Ameri- can pteridophyte complexes, twelve examples were taken from the literature (Wagner, 1965, 1966; Wagner and Wagner, 1965; Smith, 1971) and from the Gee presented above in the present paper. Of them, most have narrower ranges than either of their parents while the rest have a distribution similar to one parent but narrower than the other. While these patterns hold true for most allopolyploid pteridophytes that have been studied, some do have ranges larger than one or both parents. For example, the areas of D. cristata (Figs. 18, 20) and D. carthusiana (Figs. 19, 21) are both larger than the existing ranges of the presumed extant parents (igs 6. Ti). but mainly in an east-west direction. If these allopolypoids are very old the original relationship between their range and those of their parents may have become distorted and the patterns no longer evident. Genetic divergence of old polyploids probably slows down considerably and thus acts to conserve primitive ecological and distributional features while related diploids have undergone change (Ehrendorfer, 1980). Since the known parent of each, D. intermedia and D. ludoviciana, are diploids, their ranges may have shifted more than those of the allopolyploids. Their parental diploids may even have had world-wide distributions in the past. To understand the original relationships between the ranges of allopolyploids and those of their parents, the most revealing studies may be on what we assume to be fairly recently developed allopolyploids (e.g., D. campyloptera, D. celsa, and D, clintoniana). The generalized relationship between the ranges of allopolyploid pteridophytes and those of their parents has been summarized as follows (Wagner 1969): “Of the hybrid pteridophytes . .. those which are not entirely sympatric with the parents show less east-west correlation with the parental ranges than north-south. The north- south distribution is intermediate as we might expect.” No matter what size the range of an allopolyploid pteridophyte is in relation to those of its parents, latitudinal intermediacy between them can usually be anticipated. Therefore, we expect that if “D. semicristata” were present today it would grow approximately within 50°-80° N latitude and possibly have a circumpolar distribution. If we assume that Dryopterts ludoviciana is the other parent of D. cristata in addition to “D. semicristata,” then the origin of D. cristata must have occurred in the New World. However, a native Asian diploid fern, D. tokyoensts (Mak.) C. Chr., with the same chromosome number as D. ludoviciana (2N = 82) grows in Japan, Korea, and parts of China in only slightly higher latitudes than D. ludoviciana and in a similar climate. (A similar vicariant relationship exists be- tween D. goldiana of eastern North America and D. monticola C. Chr. of Japan which are morphologically similar.) D. ludoviciana (Fig. 22A) and D. tokyoensis (Fig. 22B) are similar morphologically, but the former has fronds which are slightly more dissected and pinnae which are more dimorphic than the latter. Perhaps they have a common ancestor that had a worldwide range that through time became fragmented, and geographical separation of the populations di- verged into vicarious species. Where the ancestral range overlapped with that of “D. semicristata” in the northwestern and north central Old World, hybrids may have formed that underwent chromosome doubling to produce D. cristata, which eventually developed a circumboreal distribution. When the Pleistocene con- 159 FIG. 22. Frond A is Dryopteris ludoviciana. Frond B is Dryopteris tokyoensis. tinental ice sheet extended southward, all the populations of the circumpolar “D. semicristata” were presumably eradicated by the advancing ice. The present eastern Asian range of D. tokyoensis may be a fragment of a former broad Old World range that survived the glaciation. The ability of D. cristata to survive the extreme conditions produced by glaciauion more successfully than its diploid parents may be explained by suggestions (e.g., Stebbins, 1950; Reese, 1958, 196 1a) that polyploids are better equipped than their diploid ancestors to adapt to extreme conditions since they have more alleles and therefore greater genetic flexibility. Hence the or igin of D. cristata may have been in either the Old or the New World or possibly in both hemispheres Assuming that Dryopteris intermedia is ike other parent of D. carthusiana in addition to “D. semicristata,” the origin of D. carthusiana may also have been in the New World. However, the Old World species, D. madierensis Alston and D. azorica (Christ) Alston, apparently possesses the same basic genome as the D. intermedia (Gibby, 1977; Gibby and Walker, 1977) and the three comprise a species aggre- gate. These findings were determined by chemical (Widen & Britton, 1969; Widen et al., 1970; Widén, Britton et al., 1975; Widén, Lounasmaa et al., 1975), cytological (Gibby & Walker, 1977) and morphological (Walker, 1961) compari- sons. One difference that exists between them is that the Old World representa- tives do not possess the epidermal glands so characteristic of the New World D. intermedia. Vhis lack of glands is important since D. carthusiana is glabrous or nearly so. If the glandular New World D. intermedia were a parent of D. carthusiana one oie expect to find more glands on D. carthusiana since they are present on other North American Woodfern hybrids involving D. intermedia. Since D. carthusiana 1s glabrous or nearly so we suspect the existing glandular New World D. ienmedta is not a likely immediate parent, although a glabrous New World ancestor may have been a parent of D. carthusiana. Presently there exist nearly glabrous populations of D. intermedia in the southern part of its range in the Appalachian mountains of North Carolina (Wagner, unpublished). 160 ‘These “glabrous” populations are probably primitive judging from outgroup comparison with its relatives and indicate that the ancestor of the glandular D. intermedia was glabrous. Under these circumstances D. carthusiana could have originated in the New World. An Old World origin of D. cathusiana is, however, equally possible, even though presently D. madierensis is only known on the continental island, Madeira Island (450 miles from Morocco) while D. azorica is only known on the oceanic islands, the Azores (900 miles from Portugal). Most taxa on island floras are the same as, or related to, widely distributed continental species. The dominance of widely distributed species in island fern floras clearly reflects an ability to migrate from continents. Their proportion on islands are much higher than their proportions in source areas (Tryon, 1970). Page (1973) concluded that the Canary Island (50 miles from Cape Yubi, Africa) fern flora is largely made up of ancient species that existed in Tertiary Europe, and have survived, little changed, as relics in the island mountains. ae on Madeira Island (1. Manton, J. D. Lovis and G. Vida, unpublished; I. Manton et al., unpublished; as cited in Lovis, 1977) have also yielded ee that these islands may have provided refugia for forms that are no longer capable of surviving elsewhere. Accordingly the insular D. azorica and D. madierensis may actually be remnants of an ancestral range which extended through nor oon Africa into western and central Europe in mid-Tertiary times. Where the ancestor overlapped that of “D. semicristata,” hybrids formed that ener to produce D. carthusiana which eventually developed its circumboreal distribution. Gibby (1977) suggested that D. madierensis or D. azorica may have occurred to- gether with D. expansa in the past in Europe to give rise to the allotetraploid D. dilatata (s. str.), which has a similar genome, and may be conspecific with the North American D. campyloptera. When the Pleistocene ice sheet extended south- ward, the entire continental range(s) of D. madierensis and D. azorica or their ancestor may have been destroyed along with that of “D. semicristata. island populations survived the Pleistocene see because of a buffered island habitat. The continental range of D. carthusiana was able to survive glacia- tion for the same reason mentioned above for D. cristata. Hence, the origin of D. carthusiana may also have been either in the Old World or the New World or possibly in both hemispheres independently. In the modern arctic climate, habitat instability is still an outstanding feature, and it is likely that new polyploid races are being produced where opportunities for hybr idization and successful establishment of new genotypes obtain (Johnson, Packer & eese, 1965). The frequency of polyploids in areas that were altered by climatic conditions of the Pleistocene epoch suggests that these environments were outside the tolerance limits of many of the existing diploid species and that they were in- vaded by older polyploids (e.g., D. cristata and D. carthusiana) as well as by the proliferation of new variable hybrid polyploids (e.g., D. campyloptera, D. celsa and D. clintoniana.) SUMMARY The 15 sexual taxa of Woodferns (Dryopteris) living in North America north of Mexico, display a diverse array of geographical ranges that correlate with topography, climate, past glacial activity, habitat availability. The ranges bear upon possible phylogenetic origins. wo diploids, D. fragrans and D, expansa and the tetraploid, D. filax-mas all have circumboreal distributions. The western D. arguta extends mainly through Pacific coastal ranges while D. cinnamonea and D. patula var. rossi reach their northern limits in southern United States. Four diploid species (D. goldiana, D. intermedia, D. marginalis and D. ludoviciana) grow exclusively in eastern North America. All of these occur from Canada south 161 through the Appalachians to the upper fringes of the southeastern United States coastal plains except D. ludoviciana, which is restricted wholly to the southeastern United States coastal plain. The other Woodferns that grow exclusively in east- ern North America are fertile allopolyploids: D. campyloptera, mainly of northern and middle latitudes in the mountains; D. clintoniana, of middle latitudes; and D. celsa, of middle and southern latitudes. Pleistocene glaciation probaby influ- enced the origins of these allopolyploids by forcing co-habitation of their re- spective parents in response to advancement and retreat of the ice sheet. The final two North American Woodfern species (D. cristata and D. carthusiana) are both circumboreal allopolyploids. It has been suggested that they may have a common parent (SS genome) now presumably extinct. Strong similarities be- tween the ranges of these two allotetraploids support this suggestion. Their origin may have been either in the Old World or the New World or possibly in both hemispheres. ACKNOWLEDGMENTS We thank Donald M. Britton, Michael G. Price, and A. A. Reznicek for reviewing the final manuscript. We also are grateful for help in various ways to P. Adams, D. J. Bay, J. M. Beitel, W. S. Benninghoff, J. G. Bruce, R. B. Cranfill, . D. Freeman, C. H. Haufler, R. Haynes, M. J. Huft, A. C. Jermy, D. M. Johnson, D. B. Lellinger, R. Kral, M. R. Mesler, J. T. Mickel, R. S. Mitchell, R. H. Mohlenbrock, J. D. a ee a B. Ownbey, J. H. Peck, T. Reeves, J. Senko, . W. Short, A. R. Smith, W. C. Taylor, J. W. Thieret, R. M. Tryon, G. R. Upchurch, E. G. Voss, F. S. Wagner, K. S. Walter, and J. R. Wells. We also acknowledge the curators of several herbaria for supplying herbarium speci- mens for study. These herbaria are BLH, CAN, CI, DAO, GH, IND, KY, MICH, MIL, MIN, MPPD, MT, MTMG, NYS, TKB, UC, UNA, UWL, WIS, and WU. This research was supported by NSF grant DEB-03550 AOL. LITERATURE CITED Beaman, J. H., 1. D. De Jong, and W. P. Stoutamire. 1962. Chromosome studies in the alpine and alpine Mora os ens co and Guatemala: Am. J. Bot., 49: 41-50. Britton Donald M. 19 ee Woodferns Dryopteris, in western North America. The Can. Field-Nat. 86: 24 194 Carlson, Thomas J. 1979. T e ue pie and frequencies of interspecific hybridization in Michigan Woodferns. Mich. , 18(2): 47-56. Christ, H. 1910. Die Geographie ve ame. fees Gustav Fischer. 35 Ehrendorfer, Friedrich. 1980. Polyploidy and distribution. pp. 15-60 in, “Lewis, Walter H. (Ed.). 1980. Polyploidy: Biological relevance. Basic Life Sciences—Volume 13. ander Hollaen- der, General Editer. Plenum Press, New York and London. xu + 583 Gibby, M. 1977. The origin of Dryopteris campyloplera. Can. J. Bot. 55: 1419- i oe -————_____ S. Walker. 1977. Further cytogenetic studies and a fy dae of the diploid an- ry in the Dryopterts carthusiana complex. Fern Gaz. 11(5): 315-3 Grillos, Steve J. 1966. Ferns and fern allies of Calfornia. University of C ae nia Press, Berkeley. 104 Beobarcetiel . 1974. Cytogeographical Studies on Dryopteris in Japan. Harashobo Co., Ltd., Tokyo, apan. 176 pp: Hulten, Eric. 1968. Flora of Alaska and ne ighboring territories: A manual of vascular plants. Stan- ford University Press, Stanford, Calif. xxii + L0( eee A. W., J. C. Packer, and G. Reese. 1965. Polyplocy distribution and environment. pp. 49 ioe in Wright, H. E. Jr. and D. G. ars 965. The quaternary of the United States. 5 Pris eton University Press. Princeton, N.J. vi + 565 pp. ae ae as H. and Robert H. Peebles. 195 1. Arizona Flora. University of California Press, ‘keley. vill + 1032 pp Lovis, ID. 197 . Evolutionary patterns and processes in ferns. Adv. in Bot. Res. 4: 229-440. Montgomery, Tames D. 1976. The distribution and abundance of Dryopteris in New Jersey. Am. Fern J. 66(2): 53-59. 162 bee C. N. 1973. Monographiae biol. Canariensis. 4: ae Reese, Gerd. 1958. Polyploidie und Verbreitung: Z. Botanik. 46: 339-354, ——.. 196 la. Goebotanische Bedeutung de OO und Chromosomenstruktur. Naturwiss. Rdsch. 14: 140-145. Reeves, Sea 1978. Notes of North American Lower vascular plants. Am. Fern J. 68(2): 61-62. Smuch, DG 1971. Systematics of the neotropical species of Thelypteris sect. Cyclosorus. Univ. Calif. -ubl. ne 59: 1-136 LO? Comparison of fern and ees plant distributions with some evolutionary erpretations for ferns. Biotropica 4(1 9. eye G. L. 1950. Variation and evolution in ie) Columbia Univ. Press, New York. xix + 643 —~ — avon Rol 1969. Taxonomic problems in the geography of North American ferns. Bioscience 19(9): 790-795. . 1970. Development and evolution of fern floras of oceanic islands. Biotropics 2(2 76-84. —————.. 1971. The process of evolutionary migration in species of Selaginella. Brittonia 23 89-100. Wagner, W.H., Jr. 1963. Pteridophytes of the Mountain Lake Area, Giles County, Virginia, Includ- ing notes ae Whitetop mountain. Castanea 28: 113-150. 196: hee ee in North Carolina and the question of its origin. J. Elisha Mitchell = SI: 966. oe new § a of ferns from the United States. Am. Fern J. 56(1): 45-46. 971. Evolution of us in relation to the Appalachians. In Holt, Perry C. (Ed.), The Distribu sets History of the of the southern Appalachians. Part 2: Flora. Virginia Polytechnic Inst. & ae e-Un — ch Div. Monogr. 2: 2. a oF cee Hagenah 1962. Dryopteris in the Huron Mountain Club Area of Michigan. Eee o 90-100 and F. S. W ee 1965. Rochester Area Log Ferns (Dryopteris celsa) and their hybrids. Proc. Rochester Acad. 11(2): 57-104. Walker, S. - G1: mee eerie in the Dryopteris spinulosa complex. II. Am. J. Bot. 48(7): 607-6 1 Wherry, Edgar T. 1961. The fern guide. Doubleday & Co., Inc., Garden City, N. : ee pp. ——_———— we The les fern guide. Doubleday & Co., Garden Cit ity, N.Y pp: Widen, F. a 1D. M. Britton. 1969. A chromatographic and cytological study of ee dilatata in eastern eee ere Can. J. Bot. 47: 1337-1344. . Britton, W. H. Wagner, Jr., and F. S. Wagner. 1975. Chemotaxonomic studies on hybrids ae Dryopteris i in eastern North America. Can. J. Bot. 53: 1554-1567. , M. Lounasmaa, G. Vida, and T. Reichstein. 1975. Die phloroglucide von drei Dryopteris— Arten von as Azoren sowie zwei Arten von Madeira und den Kanarischen Inseln zum vergleich. H elv. Chiam. Acta. 58: 904, ikko Sora, and Jaakko Sarvela. 1970. Dryopteris dilatata S. Lat. in Europe and a Island of Madeira: A chromatographic and cytological study. Acta Bot. Fennica 91: 1 —_~ _ _ ~I Contr. Univ. Mich. Herb. 15:163-169. 1982. THE ITINERARY OF MARY Se rami IN QUEENSLAND, AUSTRAL A.M. Cartei University of California Herbarium erkeley, California 9472( The names of Joseph and Mary Strong Clemens will long remain legend in the Philippines, in Borneo, and in New Guinea. Their botanical specimens housed in herbaria throughout the world will stand as a testimony of their many vears of collecting in those areas and as a valued contribution to the science of botany. Forced by the Japanese invasion to flee from New Guinea (where she had remained following the death of her husband in 1936), Mrs. Clemens ar- rived in Brisbane, Australia, on 31 December 1941 aboard a special train for refugees, and there she made her home. At an age when most of us would be content to take life a bit easier, she commenced field work in Queensland. Her journal, whose daily entries commence with a notation about the weather and a quotation from the Scriptures, shows her celebrating her 71st birthday (3 Janu- ary 1944) by a collecting trip to the summit of Mount Mistake near Brisbane. During the war years most of her collecting was confined to short trips in the vicinity of Brisbane. After the war, and until early in 1950, she made at least one or two longer collecting trips annually. Mrs. C lemens died in Brisbane on April 13, 1968, at the age of 95. Her collections have served as the basis for a number of the new species included by C. T. White in his “Contributions to the Flora of Queensland.” These have appeared in various numbers of the Proceedings of the Royal Society of Queensland. Sets of most of the collections made by Mrs. Clemens during the period from 1943 through 1950 were sent to Professor H. H. Bartlett, Director of the Botanical Garden of the University of Michigan, Ann Arbor, Michigan, U.S.A., who undertook the distribution and sale of duplicates in Mrs. Clemens’ behalf. Professor Bartlett's extensive correspondence with Mrs. Clemens is on file at the University of Michigan Herbarium. [t was in connection with the distribution of these duplicates that preparation of an itinerary of Mrs. Clemens’ Queensland field work seemed advisable in order to clarify some inconsistencies that ap- peared on the labels. The major part of the itinerary was compiled from the collections during 1957-1958 while the author was in residence at the University of Michigan processing the herbarium collections amassed by H. H. Bar tlett. Later, through the courtesy of Dr. S. L. Everist of the Brisbane Botanic Garden, Mrs. Clemens’ journals were made available to the writer. She consulted them again in Brisbane in 1976. The cooperation of Dr. Rogers McV augh and of Mrs. Robert Dieterle, both of the Herbarium of the University of Michigan, is much appreciated, The collections which Mrs. Clemens made during the period from February 1943 thr ough October 1944 (in part) were assigned consecutive field numbers in continuation of the series begun by Mr. and Mrs. Clemens. Assignment of these field numbers was made at the University of Michigan, rather than in Queens- land by Mrs. Clemens. These numbers, assigned under the direction of Pro- fessor Bartlett, ran from 41,964 to 44,767. Consequently, duplicates of type specimens may bear a collection number whereas the holotype housed in the 163 164 Brisbane Botanic Garden does not bear a number. Subsequent collections, through January 1950, were not assigned field numbers. Including duplicates, approximately 18,380 sheets of Mrs. Clemens’ Queensland collection were processed at the Herbarium of the University of Michigan. In addition to the set at the University of Michigan, the following institutions received sets of these duplicates: Botanic Museum and Herbarium, Brisbane; British Museum; Chicago Natural History Museum; Conservatoire and Jardin Botaniques, Geneve; Gray Herbarium of Harvard; Naturhistoriska Riksmuseet, Stockholm, New York Botanical Garden; Rijksherbarium, Leiden; Royal Botanic Gardens, Kew; State University of lowa; United States National Herbarium; University of California Herbarium, Berkele In the following itinerary, dates for specimens collected in the immediate vicinity of Brisbane have been omitted. When Mrs. Clemens does not use the term “station” in the usual Australian sense of “homestead” or huge shee ranch, she specifies that the locality is at a railroad station. Figures 1—3 illustrate Mrs. Clemens’ collecting localities. An illuminating account of Mrs. Clemens, as a person and a botanical col- lector, written by R. F. Langdon, is included in People and Plants in Australia (edited by D. J. and S. G. M. Carr, published by Academic Press, Sydney, New York, London, et al., 1981). It injects life into the itinerary! 1942 February to December: some collections in the vicinity of Brisbane. 1943 January: some collections in the vicinity of Brisbane. February 9, 24, 29, Mt. Coot-tha. March 2, 4, Cooper's Plains [SW of Sunnybank, $ of Brisbane]; 9, near Clermont; 11, Mt. Gravatt [S of Brisbane]; 22, Ashgrove; 29, Stafford. April 5, Ferny Grove; 19, Phornside Station, Birkdale; 26, Lawnton, Petrie. May 3, Kalinga, Clayfield; 11, Lockyer Valley; 18, Kenmore; 22, Lone Pine, Kenmore: 28, Ngun- gun, Glasshouse Mountain, near summit. June 7, near Mt. Gravatt; 11, Landsborough: 12, Beerwah; 22, Sunnybank; 29, Pine Mountain. July 3, Klimbah BY of Brisbane]; 6, Mt. Elimbah, summit saddleback, Elimbah; 15, Mt. Coot-tha, Taylor Range; 20), Nugee, Virginia, Banyo; 26-27, Landsborough, Caloundra, Mooloolah. August 5, Sunnybank; 6, Sherwood: 10, Sherwood Arboretum; 16-17, Cooper’ s Plains; 23, Mt. Gravatt; 30, Mt. Gravatt, Sunny- bank. September 7-9, ‘Tugun; 20, Chermside; 21-22, Glasshouse Mountain; 22, Caloundra; 30, Chermside, Woolscour. October 5, Caloundra; 12, Ferny Grove, Sanford Range; 19, Brisbane to Springbrook via Mudgeeraba; 20-28, Spring- brook National Park; 29, returned to Brisbane via Mudgeeraba. November 10, from Brisbane to Stanthorpe; 11-12, Stanthorpe; 13, Stanthorpe to Liston (N.S.W.); 14-18, Liston; 19, Liston to Stanthorpe to Warwick to Toowoomba; 20, Toowoomba to Ipswich to Brisbane; 26-27, Springbrook. December 21, Brisbane to Imbil, Lutton’s Saw-mill; 22-30, imi: 30, Imbil, Gympie; 31, return to Brisbane. 19444 January 3, Mt. Mistake, summit; 18, Rosewood, Phagoona railroad station, Redwood; 25, Brisbane to Laidley, Currumbin; 25-31, Laidley (Laidley hills, Forest Hill, Grandchester). February I-t, Mt. Mistake; 4, Mt. Mistake, from summit to Crosby's to Laidley; 5, Laidley to Brisbane; 17, Brisbane to Crows Nest; 18, Crows Nest, Peechey; 19-24, Crows Nest; 25, Crows Nest via 165 eee ee, 52 Tully Miskoxise: = Ingham elownsville fo) =i See 20 Mt. Dalyrymple y< 2?7~—m | | QUEENSLAND e) Mackay | Ogmore ® | Tropic_of Capricore _ _ _| __* Springvale _Ngckhompion — —_— ee | ..*Lancevale i Yalleroi@ | Scarnesse Maryboroughe ® Charleville Brisbane a Stanthorpe been Guin: ef ea yas 140° 145° I50° FIG. 1. Principal collecting localities on Mrs. Clemens’ extended trips in Queensland. ‘Toowoomba to Brisbane. March 15, Brisbane to Kingaroy, “152 miles via bus”; 16, Kingaroy; 1-7, Kingaroy to logging camp in Bunya Mountains; 18-24, Bunya Mountains, sawmill near summit; 25, Bunya Mountains to Toowoomba via Dolby; 26, return to Brisbane. April 18, Brisbane to Bribie Island; 20-30, Bribie Island. May 1, Bribie Island; 2, Bribie Island to Brisbane; 23, Brisbane to Stradbroke Island; 26, Stradbroke Island: pumping station to Amity Camp from Dunwich; 27, Amity to One Mile, Dunwich; 28-31, Dunwich. June I-6, Strad- broke Island; 6, Dunwich to Brisbane. July: Brisbane. August 1, Brisbane to Yarraman; 18, Yarraman to Brisbane. September [no definite dates]: Sanford, Sunnybank, Tambourine Mountain. October 5, Brisbane to Wallangarra; 6-19, 152° 152°30 1o3? 153°30 3 ss = ad ack no) Cc Oo 4 Cc ° 2 3 3 Crows Nest Sie waren Mt Glorious 3- «Nudgee e Banya ; ° *Stafford ~ | amity fe Ferny Grove "Chermside e Ashgrove ® BR A 27230 ne Mr Coot-thase | 8 Lockyer Valley Kenmore * “ + AIth rough one K specimen, that of Gymnogramma calomelanos var. denudata Harr., bears a locality not on the MICH sheet. 5For a sample of Harrington’s early writing, see McVaugh (1970, p. 220). OWhile in China, Harrington himself collected a few plants, including at least one fern, Cheilanthes argentea (Gmel.) Kunze, which came to MICH in 1928 with the Allmendinger hebarium. 200 structure and the form of the receptacle show an unmistakable affinity to Mecodium. Hymenophyllum fraternum Harr., J. Linn. Soc. Bot. 16: 26. 1877 (non Pres 1843) H. steerer C.Chr., Ind. Fil. 368. 1905. Lectotype: Philippines, Panay, Hoilo Prov., mountains above Maasin, Jan. 1875, Steere s.n. (MICH). Currently accepted names: Hymenophyllum fimbriatum J.Sm. ex Hook.; Mecodium fimbriatum (J.Sm. ex Hook.) Copel. According to Harrington this was found growing with the preceding, and there was only a single plant. The lectotype (or holotype if no other separate specimen was originally made) has a length of rhizome with only a single frond, from which a single pinna that had been detached for study by C opeland is now mounted on a separate sheet. Davallia philippinensis Harr., J. Linn. Soc. Bot. 16: 27. 1877. Microlepia philippinensis (Harr.) Copel., Polyp. Philip. 56. 1905. Tapeimidium philippinense (Harr.) C.Chr., Ind. Fil. Suppl. 3: 176. 1934. Lectotype: Philippines, Luzon, Mt Mahayhay (= Mt Banahaw), June—July 1874, Steere s.n. (MICH). Currently accepted name: Re al luzonicum (Hook.) Kramer. Having not seen the sheet at MICH, which had been misfiled, Kramer (1968, p. 552) was only able to cite the specimen at K, although not explicitly as lectotype. Lomaria areolaris Harr., J. Linn. Soc. Bot. 16: 28. 1877. Blechnum areolare (Harr.) Copel., Polyp. Philip. 90. 1905. Stenochlaena areolaris (Harr.) Copel., Philip. J. Sci. 2C: 406. 1908. Lectotype: Philippines, Luzon, Mt Mahayhay (= Mt Banahaw), growing in the leaves of a Pandanus, June-July 1874, Steere s.n. (MICH Currently acepted name: Stenochlaena areolarts. ‘This is the most unusual of the ferns collected by Steere, and it is also rather strange that his collection of it was the first, since the same area had been previously combed for ferns by, among others, Cuming and Brackenridge. The fern is a scandent epiphytic aquatic, rooting in and clambering between the water-filled leaf bases of Pandanus sumplex Merrill, which is common in the area and even cultivated. The pandan is the mainstay of an important cottage in- dustry, its long leathery leaves being woven into mats, baskets, hats and other articles. = leaf bases contain a substantial amount of water and harbor an interesting fauna, including small crabs. Plant-held waters were termed phyto- telmata by Maguire (1971), although Pandanus was not mentioned. My observa- tions are that prothallia only develop in leaf pockets not already inhabited by the Stenochlaena, possibly indicative of allelopathy. he tips of the rhizoids of prothallia develop into branched and expanded holdfasts anchored to the sides of the pocket that may function to help prevent the gametophytes from being flooded out in the frequent heavy rainstorms such as characterize the vicinity of Mt Banahaw. Asplenium steerei Harr., J. Linn. Soc. Bot. 16: 28. 1877. Lectotype: Philippines, Luzon, Mt } Malashay (= Mt Banahaw), epiphytic on trunks, June-July 1874, Steere s.n (MICH). Currently accepted name: Asplenium steeret. While reducing Asplenium benguetense Hieron. to A. steeret, Price (1974a) suggested that A. steerer might in turn have to be reduced to one of three earlier 201 names: A. acutiusculum BL., A. anisodontum Presl, or A. insitictum Brack. That idea is now withdrawn. Asplenium acutiusculum Bl. Enum. Pl. Jav. 178. 1828 (Holotype: Java, Blume s.n., L sheet no. 908,311-892) is a smaller individual of the same species as A. longissimum Bl. published on the same page (Holotype: Borneo, collector un- known, L sheet no. 922, 167-1020). I here designate A. longzssimum the accepted name for the species when the two names are combined. For a modern descrip- tion of the species, see Holttum (1955). The Philippine Asplenium anisodontum Presl based on Cuming 128 is a mem- ber of the complex of A. caudatum Forst.; the Hawaiian A. insiticium Brack. seems to be a large elaborate form of A. contiguum Kaulf. None of these are closely related to A. steeret. Nephrodium luerssenii Harr., J. Linn. Soc. Bot. 16: 29. 1877. eae luerssenu (Harr.) Christ, Bull. Herb. Boiss. 6: 193. 1898 (non Doerfler, ee ene (Harr.) C.Chr., Ind. Fil. 276. 1905. Lectotype: Philippines, “Bulukai Island,” 1874, Steere s.n. (MICH). Currently accepted names: Thelypteris ligulata (Presl) Ching; Pneumatopteris ligulata (Presl) Holtt. . Harrington localized this as from “Bulukai Island,” and the labels on sheets I have seen read “Bulukai, Luzon,” and “Buluka Island, Luzon,” but there was and is no such place name. I am convinced that Bulukai [sland is a misrendering of Balabac Island from Steere’s original field label, long discarded. There is also another possibility. On 23 June 1874, on the first day of his journey on foot from Mahayhay to Mauban, Luzon, Steere reached the “village of Bulukon” and could have found this plant in that vicinity. I agree with the supposition of Hubbell (1964) that “Bulukon” in that case must have been the town of Lucban. Unable to examine the MICH specimen, which had been misfiled, Holttum (1974a; 1982) cited only a K specimen, but not specifically as lectotype. Nephrodium eatonii var. formosanum Harr., J. Linn. Soc. Bot. 16: 29. 1877. Lectotype: Taiwan, Keelung, March 1874, Steere s.n. (MICH). Currently accepted name: Ctenitis eatoni (Bak.) Ching. Nephrodium bakeri Harr., J. Linn. Soc. Bot. 16: 29. 1877; Bak. in J. D. Hook. Icon. Plant. 17: pl. 1664. 1886. Dryopteris bakeri (Harr.) Copel., Philip. J. Sci. 2C: 405. 1907. Haplodictyum bakeri (Harr.) Ching, Sunyatsenia 5: 251. 1940; Holtt. Kali- kasan 2: 63. 1974. Cyclosorus bakeri (Harr.) Copel., Gen. Fil. 142. 1947. Thelypteris bakeri (Harr.) Reed, Phytologia 17: 262. 1968. Pronephrium bakeri (Harr.) Holtt., Fl. Males. II, 1: 516. 1982. Lectotype: ee Panay, Iloilo Prov., mountains above Maasin, Jan. 1875, Steere s.n. (MICH Currently accepted names: Thelypteris bakeri; Haplodictyum bakeri; Prone- phrium baker. Haplodictyum, when recognized as a genus, consists of four species, all endemic to the Philippines (Holttum, 1974b). Nephrodium subpedatum Harr., J. Linn. Soc. Bot. 16: 30. 1877. Aspidium subpedatum (Harr.) Diels, Natur. Pflanzenfam. 1(4): 186. 1899. eee subpedata ee ) Ching, Sinensia 2: 23, t.4. 1931. Lectotype: Taiwan, E. of Takao (= Kaohsiung), March 1874, Sleere s.n. (MICH) Currently accepted name: Tectaria polymorpha (Wall. ex Hook.) Copel. 202 Ching (1931, p. 23) published a photo of the “type” but, instead of a Steere specimen, it is the type of a name that is based on a later Morse collection from the China mainland and that Ching assumed to be synonymous. In Flora of Taiwan (DeVol et al. 1975), this species will key out to Tectaria trifolia, a synonym of T. angulata (Willd.) Copel. However, I have seen no specimens referrable to T. angulata from Taiwan and I believe that all so named are T. polymorpha; for the distinctions, see Price (1974b, p. 45). Harrington noted that only one of the two fronds collected was fertile, and it is the fertile frond that is present at MICH, thus supporting the choice of the MICH specimens as lectotypes of the Harrington names, rather than those of K. Polypodium aoristisorum Harr., J« Lann, Soc: Bot. 16: 40137 7 Nephrodium aoristisorum (Harr.) Copel, Polyp. Philip. 28. 1905. Dryopteris aoristisora (Harr.) C.Chr., Ind. Fil. 252. 1905. Cyclosorus aoristisorus (Harr.) Copel., Gen. Fil. 142. 1947. Thelypteris aoristisora (Harr.) Reed, Phytologia 17: 260. 1968. Nannothelypteris aoristisora (Harr.) Holtt., Blumea 19: 38. 1971; Kalikasan 2: 1974; Fl. Males. II, 1: 538. 19892. Lectotype: Philippines, Panay, Hoilo Prov., mountains above Maasin, Jan. 1875, Steere s.n. (MICH). Currently accepted names: Thelypteris aoristisora; Nannothelypteris aoristisora. This species provides the type of the generic name Nannothe lypteris, which, like Haplodictyum (see Nephrodium bakeri above), applies to several diminutive species endemic to the Philippines (Holttum, 1974b). Kt 6 Polypodium schenkii Harr., J. Linn. Soc. Bot. 16: 31. 1877. Lectotype: Philippines, Panay, Iloilo Proy., mountains above Maasin, Jan. 1875, Steere s.n. (MICH). Currently accepted names: Clenopteris obliquata (Bl.) Copel.; Prosaptia obliquata (Bl.) Price, infra. The types of Ctenopteris, Cryptosorus, and Prosaptia all belong to a natural species-group of Grammitidaceae characterized by pinnatifid or just pinnate fronds, with sori impressed or sunken into the lamina, and with rhizome paleae clathrate and ciliate. The earliest generic name for these is Prosuptia, and thus Clenopteris as a genus name cannot stand despite its current wide use. The proper combination for this species. is: Prosaptia obliquata (Bl.) Price, comb. nov. (basionym: Polypodium obliquatum BI. Enum. Pl. Javae 129. 1828). Polypodium craterisorum Harr., jinn. Soc Bott. 16) 31.1877. Lectotype: Philippines, Luzon, Mt Mahayhay (= Mt Banahaw), June-July 1874, Steere s.n. (MICH). Currently accepted names: Clenopteris celebica (Bl.) Copel.; Prosaptia celebica (BL) Tag. & Iwats. This species is rather unusual among Grammitidaceae by having the stipes conspicuously articulate; they join short, broad phyllopodia. Polypodium hammatisorum Harr., J. Linn. Soc. Bot. 16: 32. 1877. Lectotype: Philippines, Luzon, Mt Mahayhay (= Mt Banahaw), June-July 1874, Steere s.n. (MICH). Currently accepted name: Crypsinus pyrolifolius (Goldm.) Copel. Although collected by Steere “on fallen timber,” the usual habitat of this fern is high on small branches and twigs in mid-mountain and mossy forest. Polypodium steerei Harr., J. Linn. Soc. Bot. 16: 32. 1877. Microsorium steeret (Harr.) Ching, Bull. Fan Mem. Inst. Biol. Bot. 4: 306. 1933. 203 Lectotype: Taiwan, ‘Vakow (MICH). Currently accepted name: Microsorium punctatum (L.) Copel. Although the name M. steerei is accepted in Flora of Taiwan (DeVol et al. 1975), this plant is only an inconsequential form of the wide-ranging and vari- able M. punctatum. — = Kaohsiung), Ape’s Hill, Oct. 1873, Steere s.n. Drymoglossum carnosum var. obovatum Harr., J. Linn. Soc. Bot. 16: 33. 187 D. subcordatum var. Chena (Harr.) Bak., J. Bot. 28: 267. 1890 D. obovatum (Harr.) Christ, J. de Bot. 19: 73. 1905. Lemmaphyllum Fe var. obovatum (Harr.) C.Chr., Dansk Bot. Arkiv 6(3): 47. 1929 Lectotype: Taiwan, Vamsui (This was wrongly published as Posia by Harr- ington. It is evident from the herbarium labels that he interchanged, by error, the localities of var. obovatum and var. subcordatum, next on the same page), Dec. 1873, Steere s.n. (MICH) Currently accepted name: Lemmaphyllum microphyllum Presl. Christensen (1929, p. 47) cited the type as Steere 487 (US), but this must be a misinterpretation of a stray number, as none of the ferns bore collection num- ders. Adiantum steerei Harr., J. Linn. Soc. Bot. 16: 34. 1877. Lectotype: Peru, Dep. Amazonas, Poma Cocha, 1872, Steere s.n. (IK, chosen by Tryon, Cont. Gray Herb. 194: 176. 1964) Currently accepted name: Adiantum ruizianum Klotzsch. This is the only case where one cannot really argue for making a MICH specimen the lectotype. Harrington cited two different collections in the original publication, both from northern Peru, one from Poma Cocha and the other from between Moyobama & Chachopoyas. From between these syntypes, Tryon (1964) chose the Poma Cocha specimen at K and, as the first to choose, he must be followed. ‘There is no Poma Cocha specimen now at MICH, it apparently having been sent off for exchange (Tryon cited isotypes at GH, MO, and US). Three sheets remain at MICH, two from between Moyobamba & Chachapoyas, and one labeled Castia, Peru. Polypodium xantholepis Harr., J. Linn. Soc. Bot. 16: 36. 1877. Lectotype: Peru, between San Bartolomeo & San Mateo, Oroya-Linia R.R., March 1873, Steere s.n. (MICH). Currently accepted name: Polypodium xantholepis. This is close to P. thyssanolepis A.Br. ex Klotzsch, but is somewhat smaller; the paleae of the lamina below are darker and more strongly clathrate, and the upper surface glabrous. Polypodium laevigatum war. rigidum Harr., J. Linn. Soc. Bot. 16: 36. 1877. Lectotype: Peru, Dep. Amazonas, Rio Utcubamba, July 1872, Steere s.n. (MICH C urrently accepted name: Campyloneuron phyllitidis (L.) Presl. The lectotype bears the following unpublished habitat data: “Dry places on rocks and branches.” Gymnogramma calomelanos var. denudata Harr., J. Linn. Soc. Bot. 16: 37. 1877 Lectotype: Peru, 1872, Steere s.n. (MICH). Currently accepted name: ale calomelanos (L..) Link. ‘Tryon (1962, p. 60) said the “isotype” at was ied as Pebas, Peru, which Steere visited in Jan. 1872, nat that name is not mentioned in the original 204 publication nor does it appear on the two sheets at MICH. Tryon (1964, p. 68) listed the “type” at K but did not explicitly make it the lectotype. ACKNOWLEDGMENTS I am appreciative of the help of Howard Crum (MICH), R. E. Holttum (K), Kenneth Lester Jones (MICH), A. A. Reznicek (MICH), Mary Steere (Ann Arbor), William C. Steere (NY), and Edward G. Voss (MICH), LITERATURE CITED Baker-Blocker, Anita. ee Mark Harrington, Michigan’s First Meteorologist. Chronicle, Mag. Hist. Soc. eee ape Ching, R. C. ‘The ce of Chinese Ferns VII. A Revision of the Genus Tectaria from China and eee ee Sinensia 2: 9-36 Christensen, C. 1929. Taxonomic Fern-Studies ‘2 Il. Dansk Bot. Arkiv 6(3): 1-99, index, pl. 1 Copeland, E. me 1908. Notes on the Steere Collection of Philippine Ferns. Philip. J. Sci. 2C: ane ————.. 1936. The Philippine Ferns Collected by J. B. Steere. Philip. J. Sci. 60: 19-25. Crum, Tee 1977. William Campbell Steere: An Account of His Life and Work. The Bryologist —694 DeVol, C. E.etal. ‘1975. Flora of Taiwan, Vol. [. Pteridophyta and Gymnospermae. Taipei, 562 pp. aoe Brederick 1932. Joseph Beal Steere—Naturalist, Explorer, Educator. Mich. Alumnus 38: Harrington. M. W. 1875. Se ay the President of the University. Proc. Board Regents Univ. M pes meeting 187 6-482. : Che ‘Tropical oe Collected by Professor Steere in the Years 1870-75, J. Linn. o rt. oe 25-37 aoa R. a 1955: Ferns of Malaya. Singapore, 643 pp. —————.. 1974a. ee in the Family ‘Thelypteridaceae V. The genus Pneumatopteris Nakai. a As 293-32! 974b. Studies in the Family Thelypteridaceae. VI. Haplodictyum and Nannothel- ypteris. Kalikasan 2: 58-68. ——__—.. 1982. me aa eee Fl. Males. TH, 1: 331-560. Hooke W. J. & J. G. Baker. 1868. Synopsis Filicum. London, 482 pp., Hubbell T. H. 1964. ‘The University of Michigan Beal- Steere Expedition, secne a Unpub. MS, PP: Jones, Kenneth Lester. 1978. Mark hoe : ae (1848-1926): Botanist, Astronomer, feteorologist ... Mich. Academ. Kramer, K. U. 1968. The Lindsaeoid Ferns of ee on World II. A Revision of Tapeinidium. Blumea 15: 545-556. Maguire, sen tt, Jr. 1971. Phytotelmata: Biota and ae Structure Determination in Plant- held Waters. Ann. Rev. Ecol. Syst. 2: 439-46< Mains, E. B 1956. The Herbarium. The University a Michigan. An Encyclopedic Survey 8: 1442— 145 Melaven R . 1970. Botanical Results of the Mic higan yaa Survey under the Direction of Douglass eerie 1837-1840. Mich. Bot. 9: 213-2 Morton, C. V. 1968. The Genera, Subgenera, and re . the Hymenophyllaceae. Cont. U.S. Natl. Herb. 38: 153-214. Price, M. G. 1974a. Overlooked Names of Philippine Ferns. Kalikasan 2: 109-113. —————.. 1974b. The Pteridophytes Described from Mount Makiling, Luzon. Philip. Agric. 57: 37-4 per ‘Kruseman, M.J. van. 1950. Malaysian Plant Collectors and Collections. Fl. Males I. Djakarta, PP- eee : B. 1879. (Bill of sale) to the pe of the University of Michigan ... Proc. Board Regents Univ. dere June Meeting 1879: 938a. A Naturalist ae in South America. Excerpts from Correspondence. . . . Mich. Alums he Rev. 44: 331-344. Bimenican Naturalist in the Far East. Excerpts from Letters. ... Mich. Alum 1us ee v. 45: 47-61 Tryon, Rolla 1962. Taxonomic Fern Not tes. II. Cont. Gray Herb. 189. 52-76. ——————.. 1964. The Ferns of Peru. Polypodiaceae (Dennstaedtieae to Oleandreae). Cont. Gray Herb. 194: 1-253. Voss, alae G. 1978. Botanical Beachcombers and ee Pioneers of the 19th Century in the Upper Great Lakes. Cont. Univ. Mich. Herb. 13: 1-100. Contr. Univ. Mich. Herb. 15:205-207. 1982. LINUM MCVAUGHI, A NEW SPECIES FROM MEXICO C. NM ers Department of Biological Sciences Vayne State University Detroit, Michigan 48202 A specimen of Linum, collected in the state of Jalisco, Mexico, was recently sent to me for examination by Rogers McVaugh, well known for his studies of the flora of Mexico. His surmise that it might represent a new species was correct, In my opinion, and I take considerable pleasure in describing it and naming it in his honor. The following description is based on the single collec- tion. Linum mevaughii C. M. Rogers, sp. nov. . mexicano affine, sed inflorescentia ramis patentibus, caule porcis sub- erosis, foliis comparate parvis et numerosis, stylis longioribus et fere omnino connatis. Coarse, glabrous perennial herb or semishrub, ca 90 cm tall; stem up to 5 mm in diameter and somewhat irregularly corky-rigid below; leaves simple, entire, subcoriaceous, mostly opposite, occasionally whorled below, the upper- most and bracts frequently alternate, all sessile or subsessile, ovate to subcordate, the largest 15 mm long, 7 mm wide, gradually reduced in the inflorescence, also smaller and more dense at the bases of flowering branches and on non-flowering branches; stipular glands present; inflorescence many-flowered, paniculate, more or less leafy, with spreading branches; sepals five, ca 3 mm long, ovate, entire or with a few delicate marginal teeth; petals five, yellow, oblanceolate, ca 10 mm long; stamens five, ca 10 mm long, very closely appressed to the style, with very slender intervening staminodia less than | mm long; anthers narrowly elliptic, 1.5-2 mm long; styles five, ca 10 mm long, united nearly to the summit; stigmas Capitate; mature fruits and seeds not seen. Holotype: R. Gonzalez T. 466, 30 Sept., 1971. Alt. 1520 m, bosque de encino y pino en ladera de cerro, entre Cuale y Minas de Oro, Municipio de Talpa, Jalisco, Mexico (MICH The new species is illustrated in Fig. 1. It is clearly closely related to a group of three known species, L. orizabae Planch., L. guatemalense Benth. and L. mexicanum H.B.K. (Rogers, Brittonia 20: 107-135, 1968). Of the three it most closely resembles the last. Linwm mexicanum is also the only one of the three which is so far known from Jalisco. The locality for L. mevaughi lies outside the known range of L. mexicanum, however. Fig. 2 shows the distribution of the two species. Linum mevaughu differs from L. mexicanum in having corky-ridged stems, more spreading inflorescence branches, smaller, more numerous leaves and longer stamens and styles, the former more closely appressed to the style and the latter united to a greater degree, nearly to the summit. 206 sap “A_/ ae 4 aed { \ ft Karin Douthit. a, flowering branch HG: 1. Linum miCURUBTE, drawn from the type by oS loeb, stlOWET ye 2.5: 2c 1 flower with petals fallen x 6; d, androecium and gynoecium x 5; e, anther ace stigm: is x 25 CHIHUAHUA YUCATAN STD ue CAMPECHE a Apka $ 2] i cH @ras ij =S \, GUATEMALA i) —— FIG. 2. Distribution of Linum mexicanum (circles) and L. mcvaughii (triangle). Be == == == _— os Contr. Univ. Mich. Herb. 15:209-217. 1982. COMMENTS ON THE MIELICHHOFERIOIDEAE OF CENTRAL AMERICA, WITH THE DESCRIPTION OF A NEW SPECIES OF SYNTHETODONTIUM Jonathan Shaw and Howard Crum Herbarium and Division of Biological Sciences The University of Michigan Ann Arbor, Michigan 48109 The Bryaceae are fairly well defined, yet the subfamilial and even the generic limits are difficult to fix. Brotherus (1924) recognized three subfamilies, the Orthodontioideae (with basal membrane none and segments linear), the Bryoideae (with both exostome and endostome well-developed), and the Mielichhoferioideae (with lateral inflorescences and exostome, endostome, or both, reduced or lacking). Andrews (1935, 1940) and Ochi (1959) divided the Bryoideae further into the Pohlioideae and the Bryoideae in a narrow sense. The distinctions between them are not absolute. The Pohlioideae (defined here in a narrow, though informal sense to include Pohlia, Pseudopohlia, and Epiptery- gium) have narrow, lanceolate leaves with long, narrow cells and costa ending near the apex. The Bryoideae, generally considered the most primitive sub- family, retain such features as long-tapered, bordered, and trabeculate exostome teeth and well-developed endostomes, with broad, perforate segments and ap- pendiculate cilia rising from a high basal membrane. Of course, many of the Bryoideae show varying degrees of reduction from the well-developed primitive peristome. The Pohlioideae share some features of significance with the Mielichho- ferioideae (including Haplodontium, Mielichhoferia, and Synthetodontium). ‘Vhese include lanceolate leaves with long cells and costa not or only shortly excurrent. The Mielichhoferioideae are especially defined in terms of peristome reduction, but there is no single feature that separates that subfamily from the Pohlioideae. Synthetodontium costaricense, described here as a new species, could be justifiably placed in Pohlia instead. The criterion most commonly used to separate the Mielichhoferioideae is pleurocarpy. However, inflorescences that appear to be lateral are in some cases terminal on short stems that become overtopped by innovation and displaced to a pseudolateral position. Meusel (1935) suggested that timing determines the degree of lateral displacement. In taxa where the innovation develops early in the season and therefore has time for considerable growth, the inflorescence appears to be borne on a short side branch. But in most species of Bryum and Pohlia the subfloral innovations develop late and thus have the appearance of secondary branches. There is considerable diversity of branching patterns in the Mielichho- ferioideae. In M. macrocarpa (Drumm.) Bruch & Schimp., for example, the plants are freely branched, mainly by innovation. Inflorescences terminate the main stem as well as innovations and fork branches (Fig. 2). In M. mielichhoferr (Hook.) Loeske, stems are even more branched, often repeatedly forked (Fig. 1). Male plants tend to branch more freely than female, and perigonia are consid- erably differentiated as reddish buds at the ends of short lateral branches. 209 210 Generally the main stem is identifiable among several finer grades of lateral branches. Innovations occur infrequently, the branches bearing perigonia are scarcely strong enough to give rise to further innovative growth. They are highly specialized reproductive branches with scarcely any vegetative leaves subtending the perigonia. Compared with M. macrocarpa and M. mielichhoferi, M. graciliseta (Hampe) Mitt. is weakly branched (Fig. 3). Although apparently lateral, the paroicous inflorescences terminate main stems or more rarely lateral branches. The stems fork but not abundantly. Synthetodontium costaricense has inflorescences consistently terminal, on short, simple or sparsely forked stems, but appear to be lateral because of multiple innovation (Fig. 4), In Pohlia innovations are common, but forking of the stem is rare. In a few species, such as P. atropurpurea (Wahl.) Lindb., the tiny plants are unbranched. Pohlia drummondii (C. M.) Andr. normally has innovations below the perigonia but not the perichaetia (Fig. 6); and sterile plants are simple. Pohlia papillosa (Jaeg.) Broth., on the other hand, often innovates below female inflorescences, but male plants are simple (Fig. 7). Both male and female plants of Pohlia wahlenbergu (Web. & Mohr) Andr. innovate consistently, and unusually robust sterile plants sometimes branch by forking (Fig. 5). Such examples show that the 7. Branching patterns. 1. Mielichhoferia muelichhofert. 2. M. macrocarpa. 3. M. graciliseta. 6. P FIGS, 1- 4. Synthetodontium costaricense. 5. Pohlia wahlenbergii. 6. P. drummondii. 7. P. papillosa. (Hollow circles represent perigonia, solid circles perichaetia.) ce) ala difference between highly branched Mielichhoferias and simple or sparsely branched Pohlias is a matter of degree, albeit with vastly different extremes. The Mielichhoferioideae are characterized by peristome reduction, with exostome, endostome or both variously reduced or lacking. Yet Brotherus (1924) listed 13 species of Meelichhoferia with double peristomes and both species of Synthetodontium have an exostome as well as a fairly well-developed endo- stome It is evident that the primitive Mielichhoferioideae are not entirely distinct from some of the more advanced Pohlioideae. Such continuities are to be ex- pected, of course, if the two families are derived from a common ancestry. We discuss below several taxa of the Mielichhoferioideae with a relatively primitive position. THE MEXICAN AND CENTRAL AMERICAN MIELICHHOFERIOIDEAE Synthetodonttum has a double, fairly well-developed peristome, with the endostome partly or almost entirely adherent to the exostome. The genus Con- sists of two species: S. costaricense Shaw & Crum (described below) and S. pringlei Card., known from two collections in Mexico (Jalisco). Mielichhoferia, consisting of several species enumerated below, has a more reduced peristome—lacking, single and consisting of endostome only, or double. Haplodontium has a reduced, single peristome consisting of exostome only. Haplodontium argentifolium (Mitt.) Jaeg. & Sauerb. grows on soil of banks and in rock crevices at altitudes of about 12,500 ft. in Mexico (Distrito Federal and Puebla). It should be noted that Mielichhoferia lacks definition. In at least one species of more northern distribu- tion, M. mielichhoferi (Hook.) Loeske, the single peristome is actually exostome. Since M. mielichhofer: provides the type of the genus, this observation carries significant nomenclatural implications. However, a much needed re- -evaluation of the subfamily might lead to combining Mielichhoferia and Haplodontium to allow for varying kinds and degrees of peristomial reduction. We thus make no transfers pending results from studies in progress. KEY TO THE SPECIES OF MIELICHHOFERIA IN MEXICO (AND ELSEWHERE IN CENTRAL AMERICA) Peristome lacking (or sometimes present but very poorly developed and shed with th operculum). 3. M. ire Bensloiie os ssent P eristo ne sin le. donne Soe papillose, irregular-margined; leaves distinctly serrulate. 2. M. serrata actos segments smooth, not particularly irregular- margined. 1. M. campylocarpa re Peristome cae le. 4. Capsules broadly pyriform; exostome teeth not or scarcely exceeding the aR . M. landu +. Capsules cylindric; exostome teeth %—%4 the length of the exostome. 5. graciliseta _ ned M. campylocarpa (Hook. & W.—Arn. ex Hook.) Mitt. On turfy and rocky soil at 9750—15,000 ft. alt. on Mexico’s high volcanic peaks; Jalisco (Nevado de Colima), México (Nevado de Toluca), Puebla (Ixtaccrhuatl), and Veracruz (Orizaba; also between Perote and El Cofre).—Mexico and Guatemala; Ecuador, Bolivia, and Peru; recently reported from a wide range in South se Rhodesia, Kerguelen, Australia, New Zealand, Marion Island, and Hawai 2. M. serata Card. & Herz. ex Card. On rocky soil, 9200-13,000 ft. ne Mexico (Nevado de Toluca), Puebla (La Venta), and Veracruz (nba Endemic. 212 3. M. schiedeana C. M. On soil among rocks and in crevices at 10,500— 15,000 ft. on Mexico’s high volcanoes (Toluca and nearby Zinantecatl, Popo- catepetl, Ixtaccrhuatl, and Orizaba).—Endemic. 4. M. landiui Card. On soil at 7000-11,000 ft. alt., in pine zones or in the transition to open terrain; Distrito Federal, Hidalgo, Mexico, Oaxaca, and Puebla.—Endemic. 5. Mielichhoferia graciliseta (Hampe) Mitt., Jour. Linn. Soc. London, Bot. 12: 324. 1869. Leptochlaena graciliseta Hampe, Linnaea 32: 130. 1863. Small or medium-sized, whitish or reddish plants. Stems 5-10 mm high, branched by innovation, sometimes forked. Rhizoids restricted to the base of the stem, orange, papillose. Leaves of innovations erect to erect-spreading, 0.8- 1.2 mm long and 0.4—0.6 mm wide, lanceolate; margins plane, serrate near the apex, serrulate to the midleaf or more: costa ending a few cells below the apex; cells linear-rhomboidal and + vermicular, thin-walled, 45—75(130) x 7-10 um, not differentiated at the margins. Paroicous; inflorescences terminal on main stems or short to elongate side branches, bulbiform; perichaetial bracts concave, broadly lanceolate, reddish. Setae 8-13 mm long; capsules inclined at 45-1009, 1.5-3 mm long, narrowly pyriform; annulus broad (of 2-3 rows of cells): operculum convex to conic, blunt or apiculate; exothecial cells rectangular, 50— 75 X 20-35 wm, with evenly thickened, + sinuose walls; stomata superficial, with guard cells 40-50 ym long; exostome teeth irregularly lanceolate, blunt, 45-34 as long as the endostome, varying from 150-280 um long, sometimes on the same capsule, hyaline or very pale yellow, smooth or finely roughened, not or scarcely bordered, moderately trabeculate, with trabeculae irregularly projecting at the sides; endostome hyaline or pale-yellow, consisting of a distinct, though low basal membrane (scarcely exceeding the annulus), linear segments, scarcely keeled or perforate, often joined at the tips, papillose in the upper half, nearly smooth below, and cilia lacking. Spore 15-19 ym, very finely roughened. Figs. 8-16. On soil or rock ledges and in crevices (at least sometimes on limestone), sometimes on banks of streams, at 2900-3600 m. alt.; Mexico (Oaxaca, Ixtlan, Hermann 269]2a: Sierra Juarez, Richards, Sharp & Sharp 2769, Sharp 98506); Colombia, Venezuela, and (as M. castanea Herz.) probably Bolivia). Bartram (1951) erroneously reported M. graciliseta from Costa Rica. The specimen is re- corded below as the type of Synthetodontium costaricense. The double peristome, with the endostome considerable longer than the exostome, is the most significant feature of this species, not previously recorded from Mexico. Hampe described the species in the genus Leptochlaena which he distinguished from Mielichhoferia on the basis of the double peristome. The generic name is, however, a later homonym of Leptochlaena Spreng. Mitten (1869) used the name at the sectional level, where it is legal. Brotherus (1924) established the subgenus Mielichhoferiopsis for double-peristome species of Mielichhoferia. Use of the suffix -opsis is, however, contrary to article 2 of the international code of botanical nomenclature. Brotherus’ treatment (1924, pp. 353-354) is not entirely trustworthy. For example, M. lindigii Hampe is characterized as having a double peristome. Mitten (1869) placed that species in a group with “peristomium simplex internum.” Hampe did not clearly indicate whether the peristome is single or double, but his statement that Leptochlaena graciliseta has the stature of a Mielich- hoferia but a double peristome suggests that he would have placed M. lindigii in Mielichhoferia because of its single peristome. A more confusing situation is met with in Brotherus’ treatment of M. graciliseta. He illustrated it (p. 354) as having an endostome about twice the length of the exostome, but his key characterizes the peristome as double with both exostome and endostome similar in length or the endostome shorter. A specimen in Hampe’s herbarium with data matching that of the type (Bogota, La Penna, 2800 m, Lindig 2014) has teeth about half as 7a 3) 38. 8-16. Mielichhoferia graciliseta, 8. Peristome, X 115. 9. Exothecial cells, x490. 10. Upper —s FIC leaf cells, x490. 11. Alar cells, x490. 12. Capsules, x11. 13. Leaves, x30. 14. Annulus, x490. 15. Costa in section, X490. 16. Stoma, x490. long as the segments, just as in Brotherus’ illustration. Mitten cited, in addition to that collection one also from the Andes Bogotenses, Wier 181, represented at BM and in triplicate at NY. One of the specimens at NY is accompanied by Mitten’s sketches showing endostome and exostome of equal length. Examina- tion of the material, however, shows that Mitten was in error as the peristome is exactly as Brotherus illustrated it. It may be that Brotherus’ confused text was influenced by Mitten’s sketches. i veristome of M. castanea Broth. ex Herz., as illustrated by Herzog (1916), closely matches that of M. graciliseta. In both species the segments of the endostome are commonly joined at the tips. Very likely the names are synony- mous. Pohlia pseudobarbula (Ther.) Shaw (of Mexico) has a peristome scarcely more complete than that of M. graciliseta. The exostome teeth, though longer than the endostome, are blunt, hyaline and nearly smooth below, and scarcely trabeculate or bordered, as in M. graciliseta. It is tempting to think of the double-peristome 214 Mielichhoferia as an advanced Pohlia (judged in terms of peristome reduction in more advanced members of the genus Pohlia). Herzog (1926), in fact, suggested such a relationship between Mielichhoferia and Pohlia. The only other Mexican Mielichhoferia with a double peristome is M. landii. It has the peristome even more reduced. The exostome is very short, no more than 100 ym high and usually completely hidden by the annulus. Mielichhoferta macrocarpa (Hook.) Bruch & Schimp. has been reported from Mexico on the basis of a sterile collection made by Hahn on the Nevado de Toluca. The plants do not compare at all well with M. macrocarpa as the leaf margins are less clearly recurved and more strongly toothed, and the leaf cells are shorter and thicker-walled. Because of a well-developed, almost Pohlia-like peristome the following new species of Synthetodontium is particularly significant in indicating a phylogenetic trend from the Pohlioideae into the Mielichhoferioideae: Synthetodontium costaricense Shaw & Crum, sp. nov. Plantae graciles, 5-9 mm altae, rubrae vel aureae, + nitidae, caulibus Innovando ramosis, fructificationibus simulate lateralibus. Folia 0.8—1.2 mm longa, lanceolata, acuta, haud vel + limbata; costa subpercurrens; cellulae rhomboideales. Dioicum. Seta usque ad 10 mm altitudine; capsula + erecta, anguste pyriformis, 2-3.5 mm longa; annulus latus; operculum convexum: peristomium duplex, exostomii dentes ca 3000 pm longi, lanceolato-acuminati, papillosi, inferne aurantiaci; endostomium membrana basilari + alta, processi- bus angustis, carinatis, + perforatis. Sporae 12-17 uw, laeves vel subtiliter papil- losae. Very small, reddish to golden, + glossy plants. Stems 5-9 mm high, freely branched by subfloral innovation, occasionally forked, with a poorly developed central strand surrounded by gradually larger, thin-walled cells and 2-4 periph- eral layers of abruptly differentiated thick-walled cells. Rhizoids restricted to the base of the stems, orange, densely papillose. Leaves of innovations erect to erect-spreading, + concave, 0.8—-1.2 mm long and 0.3-0.6 mm wide, lanceolate, acute; margins plane or slightly reflexed; costa ending a few cells below the apex; upper cells rhomboidal, varying from 45-75(125) x 9-15 wm, rather thin- walled, undifferentiated at the margins or + abruptly elongate in 3-5 rows forming a distinct, sometimes yellowish border; basal cells gradually to abruptly enlarged and rectangular, or sometimes scarcely differentiated. Dioicous: perichaetia terminal but appearing to be lateral because of well-developed in- novations, perichaetial bracts reddish, concave, lanceolate to broadly triangular, with margins + recurved; perigonia bulbiform, at the ends of short, inconspicu- ous stems, sometimes appearing lateral because of innovation, the perigonial bracts erect, concave, ovate, shortly and broadly acute; paraphyses abundant, filiform, pink to orange. Setae up to about 10 mm; capsules erect or nearly so, 2-3.5 mm long, narrowly pyriform, with the neck about as long as the urn; annulus broad consisting of 2-3 rows of cells and about 100 wm high; operculum convex, blunt; exothecial cells 45-65 x 12-26 ym, rectangular, with straight, evenly thickened walls; stomata superficial or slightly sunken, the guard cells ca. +0 fm long; exostome teeth ca. 300 in long, lanceolate, slenderly acuminate, orange and very finely papillose below, gradually becoming pale to hyaline and finely papillose, above, strongly bordered, trabeculate; endostome vellow, some- tumes + adherent to the exostome, with a basal membrane about 4 the length of the exostome, narrow but keeled and £ perforate segments, smooth or finely papillose below, and subhyaline above, cilia none. Spores 12-17 wm, smooth or finely roughened. Figs. 17-26. Cloud forest, ca. 10,000 ft., Roberts’ Farm, S. slope of Mt. Irazu, prov. Cartago, Costa Rica, Ruth D. Svihla (holotype, FH, a Mielichhoferia graciliseta). ES 23 “es SAL ce CC eo Merk FIGS. 17-26. Synthetodontium costaricense. 17. Leaves, X30. 18. Peristome, x 115. 19, 20. Upper leaf cells, 490. 21. Annulus, *490. 22. Exothecial cells, x490. 23. Costa sections, 490. 24. Stoma, «490. 25. Exothecial cells near the capsule mouth, x490. 26. Capsules, x 11. The genus is characterized by a double peristome with perforate segments arising from a basal membrane about one-third the length of the exostome which consists of long-tapered teeth, orange below, hyaline above. The only other species of the genus, S. pringle: Card., is known from two localities in the state of Jalisco, Mexico. It has endostome segments adherent to the exostome teeth giving a chambered appearance rather like that of Brywm algovicum Sendtn. 216 ex GC. M. According to Cardot each segment becomes split into two parts each adhering to an adjacent tooth and appearing to form a narrow, sinuose border. Actually, as we have seen them and as Georg Roth showed them in manuscript drawing, they are very delicate and irregular in outline but fully adherent and not bifid. In S. costaricense the exostome and endostome are mostly free (though some- times adherent below). It has red-brown, acute leaves and spores 12-17 wm Oe ae pringlei has pale-green, broadly acute to obtuse leaves and spores 25-30 us he cells at the leaf margins are sometimes differentiated as a border in S, costaricense: the border may vary from fairly distinct and often yellowish to none at all even on the same plants. The genus OU ee has some phylogenetic significance since its peristome, being relatively well developed, is decidedly primitive among the more reduced peristomes generally representative of the Mielichhoferioideae. The exostome teeth are regular in development and long-tapered, showing scarcely any resemblance to those of the double-peristome Mielichhoferias such as iM. eraciliseta. There is, in fact, little to distinguish the exostome from that of such an otherwise dissimilar species as Bryum algovicum. Furthermore, the endo- stome segments are keeled and more or less perforate, unlike those of other Mielichhoferioideae, though like those commonly found in Pohlia. We must admit that with no prior knowledge of Synthetodontium as a genus we might have described S. costaricense asa Pohlia, though per haps in its own subgenus. In any case, Synthetodontium may be close to the ancestral origins of the Mielichho- ferivideac and the Pohlioideae. he rather well-differentiated bud-like inflorescences seem to link Syntheto- dontium to the Mielichhoferioideae, at least as they are now constituted. How- ever, such a linkage presents a dilemma. Both species of Synthetodontium and the double-peristome Miclichhoferias suggest relationships to Pohlia, but to unre- lated taxa. Synthetodontium pringtei Card. and S. costaricense have exostome teeth resembling those of primitive Pohlias (or even many species of Brywm). Mielichho- feria graciliseta (and to a lesser degree M. landii), on the other hand, show re- semblances to such advanced Pohlias as P. pseudobarbula. Kither the Mielichhoferioideae are polyphyletic, with only distant phylo- genetic linkage between Mielichhoferia and Synthetodontium, or there is evidence for convergence, by one or the other genus, toward Pohlia. These alternatives are shown tn figures 27-28: Figure 28 implies that apparently lateral bud-like inflorescences evolved independently, with Mielichhoferia evolving from the ancestors of present-day Pohlias with similar peristomes. Figure 27 implicates a Mielichhoferia Mielichhoferia Pohlia Synthetodontium Pohlia Synthetodontium 27 28 FIGS. 27-28. Alternative hypotheses of phylogenetic relationships of Pohlia, Synthetodontium, and en ZY) common ancestor of Pohlia and Synthetodontium with long-tapered exostome teeth and well-developed endostomes. The similarity in the exostomes of such eae as aaa graciliseta and Pohla pseudobarbula seems due to con- vergenc It may be significant that both Synthetodontium and Mielichhoferia graciliseta (at least) have anatomically similar costae. In Pohlia there are usually median guide cells and stereids moderately differentiated or none. In the Mielichho- ferioideae examined, stereids fill most of the costa, and guide cells are absent. ETE ERATURE CITED pe - L. 1935, 40. Bryaceae. /n A. J. Hens Moss Flora of North America North of Mexico. . 2, pp. 184-242, 261-262. Keene aan E. B. 1951. Additional Costa Rican mosses. IV. Bryol. 54: Brotherus, V. F. 1924. Musci. /n A. Engler & K. P1 -antl, Die eae PRerenearnilien: (Ed. 2). ol. 10. Leipzig. Herzog, T. 1916. Die Bryophyten meiner zweiten Reise durch Bolivia.—Biblio. Bot. 87, pp. I-172. ————.. 1926. Geographie der Moose. 439 pp. Jena. ae H. a As uchsformen und Wuchstypen der europaischen Laubmoose.—Nova Acta d0ld., 3: 123-277. sive, W. 186 a ae austro-americani: enumeratio Muscorum Omnium austro-americanorum uctori Speke cognitorum—Jour. Linn. Soc. London, Bot. 12: 1-64( Ochi, HL 1959. A revision of the Bryaceae in Japan and the adjacent regions. Publ. Biol. Inst. Lib. pon Tottori Univ., 124 pp. —_ “ac. ye =, ~~ = _s == -— ee = — = =. a a a s ia xe 7 Se Sh Se ited «Sok en eee ae ak Contr. Univ. Mich. Herb. 15:219-295. 1982. POHLIA HEDW. (MUSCI) IN NORTH AND CENTRAL AMERICA AND THE WEST INDIES Jonathan Shaw University of Michigan Herbarium and Division of Biological Sciences Ann Arbor, Michigan 48109 Within the Bryaceae, basic features defining Pohlia are lanceolate, often serrate leaves, costae ending before or in the apex, and long, generally narrow cells. The peristome is double and the endostome is well-developed although cilia, when present, generally lack appendicula. Groups of taxa within the genus are more or less uniform with respect to such attributes as exostome and endo- stome form and ornamentation, exothecial cell shape, the presence or absence of annuli and the position of stomates relative to their guard cells. Within such groups, species differ mainly with respect to gametophytic details. Axillary gemmae have evolved at least twice in unrelated lines. Rhizoidal tubers are present in P. ante P. pacifica and P. integra, and are of some use in char- acterizing these les The group of species segregated by some as the genus Mniobrywm are in- cluded here in Pohlia, although it is fully acknowledged that Mniobryum consti- tutes a distinctive assemblage. The presence of such species as P. lescuriana, which is a Pohlia by any definition but approaches the Mniobryum group in several significant respects, makes including the latter as an infra-generic taxon in Pohlia at least defensible. Crum and Anderson (1981), following Andrews (1935), placed Bryum apiculatum Schwaegr. (= P. cruegert Hampe ex C. Mull.) in Pohlia. However, I agree with Ochi (e.g. 1981) that it is more at home in Bryum. It is the long cells that might indicate a relationship to Pohlia, but this trait is to be found rather frequently elsewhere in Bryum. In addition, the costa is shortly excurrent, and the leaf margins are entire (or vaguely serrulate), features un- common in Pohlia. Although appendiculate cilia are found in a few Pohlia spe- cies, they are uncommon, and their presence in B. apiculatum casts further doubt on a closer relationship to Pohlia than to Bryum. Finally, the lowland circum- tropical distribution of B. apiculatum would be unique in Pohlia, though not rare in Bryum. Pohlia peracuminata Bartr. from Guatemala is a similar case where generic placement is uncertain, but according to Ochi (in litt. 1981) it can be Soren to Bryum microchaeton Hampe. I also follow Ochi in interpreting Anomobryum as more allied to Bryum than Pohlia, although generic status seems justified. The long-narrow cells, which are not even characteristic of all Anomobryum species, is the only feature that relates Anomobryum to Pohlia. With about 110 species, Pohlia is represented on every continent. The cir- cumboreal taxa are by far the best understood, and of the thirty-two Pohlias in North America north of Mexico, twenty-five occur on at least one other north- ern continent. There have been no revisions of tropical or Southern Hemisphere species. pe) of the American Pohlia species are rare throughout their range and t nus is poorly represented in temperate eastern North America. Pohlia 7 ee and P. lescuriana are restricted to eastern North America and are fairly AAG) 220 widespread, though relatively uncommon (P. lescuriana is also known from two localities in Alaska). Pohlia wahlenbergii and P. nutans are common in the north- eastern states and extend southward to a considerable extent. Otherwise, species reaching ‘Tennessee and North Carolina : so by following the Appalachian mountain chain. These include P. cruda, P. elongata, and P. longicollis. Pohlia annotina is fairly widespread in eastern North America, in the mountains and elsewhere. In the Rocky Mountains and westward, Pohlia is one of the most diversified of moss genera. Collecting on the slopes of Mt. Rainier (Washington) for ex- ample, can yield up to eight species in a day. Relatively moist areas with acidic soils in the mountains of southern British Columbia are also uncommonly di- verse in Pohlia species. Several species extend southward in the Rockies to Colorado, but only P. elongata, P. nutans, P. wahlenbergii and P. tundrae are con- firmed in Arizona and (or) New Mexicc Of the North American species, p. elongata, P. cruda, P. nutans and P. wahlenbergu reach Mexico, but only P. elongata is common there. Other aes 4 fexico and Central America are either endemic, for example, P. integra, oerstediana and P. pseudo-barbula, or have phytogeographic affinities a ae plants. The latter includes P. richardsii and P. polycarpa. Of the Pohlia taxa occurring north of Mexico, six (19%) are endemic. Three of these are found in the Pacific coastal region near or at sea level (P. longi- bracteata, P. columbica and P. pacifica), whereas the other three are alpine-high montane in the Cascade and (or) Rocky Mountains. Endemism in Mexico and Central America is more difficult to evaluate since collecting there has been relatively scarce. Pohlia integra and P. pseudo-barbula are confined to mountains near Mexico City, and P. oerstediana extends from central Mexico to Costa Rica. There are no West Indian Pohlia endemics and only P. elongata, P. wahlenbergii, P. richardsu and P. papillosa are known from the islands. As a whole, Pohlia covers a significant diversity of habitats. Most taxa occur on soil, although the substrates of particular species are usually well-defined in terms of pH, texture, organic matter content, and the levels of various ions. The Mniobryum group (species 29-36) is found almost exclusively on soils of high pH, .9 or above, and low organic content (less than 2.5%). Of those species only P. longibracteata occurs frequently on more acidic soil. Species 14-17, including the group of weedy propaguliferous species, also prefer soil with relativ ely low levels of organic matter (3.5% or less), but they usually occur on mildly acidic soil. The remaining species are almost entirely restricted to acidic soils (pH 4-6). y far the most common habitat in which to find Pohlias are roadbanks and other disturbed sites. Some such species as P. annotina are rarely collected in undisturbed habitats, although most other, less weedy species are found also on streambanks or in oil-filled crevices. Several species reach the alpine zone, and P. crudoides, P. longicollis and P. andrews are characteristic of crevices among siliceous rocks in tundra. Most other species, when they occur in the alpine zone, grow in more exposed sites. Pohlia obtusifolia and P. bolanderi var. seriata prefer exceptionally lush snowmelt areas. Pohlia sphagnicola is unique in being restricted to Sphagnum hummocks lacking mineral soil The main purpose of this paper is to provide an aid in identification. The infra-generic evolution and classification of Pohlia will be dealt with elsewhere, and the genus is left undivided here. However, the arrangement of species is intended to represent relationships, insofar as a linear sequence can. Species I-13 have long-rectangular, nonsinuose exothecial cells, whereas in species 14— 35 they are short-rectangular to isodiametric, usually with strongly sinuose walls. Species 10-13 have whitish exostome teeth densely covered with papillae (less so in P. pseudo-barbula) and lack cilia altogether. I have dispensed with tradition in, for example, placing P. elongata next to P. nutans rather than with other species 221 having narrowly split endostome segments. It is open to question whether endo- stome simplification has occurred only once in those species, and the present arrangement is intended as noncommital, reflecting gametophytic as well as sporophytic similarity. Of the species with sinuose exothecial cells, 14-27 have annuli, while they are lacking in 28-34. In species with sinuose exothecial cells, they become di, straight when thoroughly moistened. Axillary gemmae characterize species 11-13 and 17-26. Many a ohlcae relating to nomenclature and typification are not discussed for the sake of brevity. The location of lectotypes are indicated, and further details on specimens are available from the author. The problem of in versus ex in some author citations was not resolved completely. In such cases as Webera columbica Kindb. in Kindb. & Macoun, the in allows the name to be typified from Kindberg’s herbarium, as most bryologists would agree it should. It should be admitted that in such other cases as P. longibracteata Broth. in Roell., the choice of in is to facilitate typification since there ts little evidence from the or Bue publi- cation to make a decision as to whether im or ex is more appropria A revision of species 18-26 was published previously (Shaw, 198 La), and only brief descriptions and illustrations are given here. Pohlia beringiensis (17) 1s described for the first time. Although sterile, similarities in gemmae structure relate it to P. drummondu (18). Gemmae similar to those of species 17-26 are produced by P. papillosa, P. richardsu and P. pseudo-barbula, but other characters, especially sporophytic, indicate that the two groups are only distantly related. eys are provided for both fertile and non-fruiting plants, although identi- fication may require at least a determination of sexuality. e key to non- fruiting plants is offered somewhat apologetically since gametophytic distinc- tions, though often reliable, are difficult to define and consequently to use. ept where noted, geographic distributions given are based on specimens examined. I did not attempt to exhaustively otal Lute the local distributions of such common species as P. wahlenbergii, P. cruda and P. nutans, and so maps should be considered synoptic. Greenland Aemntion: shown on maps are given incidentally. Pohlia Hedw., Spec. Musc. 171. 1801. Webera Hedw., Spec. Musc. 168. 1801 (hom. illeg., non Webera Schreb.). Lamprophyllum Lindb., Oefv. K. Vet. Ak. foedne 20: 12. 1863 (hom. illeg., non Lamprophyllum Miers. Mniobryum Schimp. ex Limpr., Laubm. Deutsch. 2: 272. 1892. 7 Small to robust, glossy or dull plants generally forming populations of restricted size, occasionally mixed with other mosses. Stems erect, short to very long, simple or branching at the base or less commonly above when sterile, simple or often innovating when bearing gametangia, in transverse section round to pentagonal, with a + distinct central cylinder of small, thin-walled cells, 4—7 rows of gradually larger thin- to thick-walled cells and 1-4 (—6) rows of small, often pigmented thick-walled peripheral cells. Rhizoids pale orange-red to brown, papillose, sometimes bearing tubers. Leaves little altered when dry, symmetric or + asymmetric, lanceolate to ovate-lanceolate (linear in the S. Hemisphere P : tenuifolia), acute or rarely rounded at the apex, generally + decurent; margins plane to recurved (especially the perichaetial bracts), ser- rate to serrulate or rarely entire near the apices, entire to serrulate below; costa single, ending above mid-leaf to percurrent, rarely shortly excurrent, occasion- ally abruptly widened below, in transverse section rounded to plane on the ventral (upper) surface, + strongly convex on the dorsal surface, with 2—4 median guide cells, a + well-developed dorsal stereid band, sometimes a few ventral stereids, and 1 row of enlarged firm-walled cells on the dorsal and Zee ventral surfaces; upper cells hexagonal to linear, sometimes large and lax, smooth, with thin to thickened walls, lower cells hexagonal, often quadrate across the insertion, marginal cells undifferentiated or in some species longer and narrower than the median. Propagula sometimes present in the upper leaf axils, arising singly or in clusters of 3-many, consisting of a multicellular body and (0-)I-4 apical or subapical leaf primordia. Dioicous, paroicous or rarely autoicous; perichaetia terminating short to elongate main stems, the bracts scarcely to strongly differentiated, with recurved margins; perigonia terminal on main stems or sometimes innovations, the bracts shortly acute to longly acumi- nate from an ovate, concave, + red-brown base, erect to spreading or sometimes squarrose from the erect base; paraphyses filiform, orange to pink. Sporophytes rare tO Common; setae erect, In transverse section similar to the stem but with a more strongly developed central cylinder; capsules | or rarely more per in- florescence, inclined to erect, ovate-pyriform to linear-cylindric, sometimes urceolate, with a well-developed neck, very small to large; exothecial cells isodiametric to elongate and rectangular, sometimes collenchymatous, the walls straight to sinuose, smaller, thicker-walled and less sinuose near the TIM; 1s0- diametric with thin to firm, straight walls in the neck, gradually more elongate with thickened, straight walls just above the seta; stomata abundant in the neck, with 2 guard cells, superficial, sunken, or deeply immersed; annuli well devel- oped to absent, (1-)2-3 rows of cells, revoluble or separating irregularly; opercula conic to convex, acute, apiculate, umbonate or rarely rostrate; calyptrae cucullate, smooth; peristome, double; exostome teeth 16, lanceolate, acute to acuminate and 300-550 wm long, or rarely + blunt and about 150-300 um long, strongly to obscurely bordered, with dorsal plates wider but ventral plates thicker; endostome well-developed or incomplete, hyaline or sometimes yellow; basal membrane low to high; segments generally elongate and keeled, broadly to narrowly (or not at all) perforate; cilia long and nodulose to short or absent. Spores very finely to coarsely roughened. KEY TO STERILE PLANTS I. Plants bearing axillary gemmae. 2. Gemmae arising singly in the leaf axils. 3. Plants conspicuously whitish; leaves erect. 17. P. beringiensis 3. Plants green, reddish or black; leaves erect or spreading. Gemmae translucent orange-red; leaf primordia the same color as the body. 13. P. pseudo-barbula 4. Gemmae opaque, reddish to orange or green, sometimes black; primordia green or pale. 5. Gemmae cylindrical to oblong, reddish when moist; leaf primordia green and conspicuous, scattered on the body and at the apex. 18. P. drummondii 5. Gemmae elliptic to ovoid, orange when moist (often black when dried); primordia pale, stiffly triangular, mostly restricted to the apex. 19. P. filam 2. Gemmae arising in clusters. ; 6. Gemmae spheric to short-oblong, hardly longer than wide. Primordia of gemmae triangular-laminate, forming a dome over the apex. 22. P. bulbifera 7. Primordia of some or all gemmae peglike (just 1-2 cells broad at the base), erect or + incurved over the apex. 8. Gemmae yellow or brown, isodiametric or nearly so. 26. P. camptotrachela 3 > red, a little longer than wide. NO 8. Gemmae orange o1 ‘ : 23. P. andrewsii 9. Plants dull (tropical). 12. P. nichardsii 6. Gemmae oblong, obconic or + elongate and vermicular. 10. Gemmae small (about 110 zm) and orange-red, with the primordia peglike and incurved. go to 8 10. Gemmae larger, oblong, obconic or linear-vermicular. Il. Plants glossy. 223 12. Leaf gen tgs laminate, at least 3-4 cells broad at the base. 13. emmae oblong or obconic. 20ear apse 3. Gemmae tae aaa 21. P. tundrae 12. Leaf primordia peglike, 1-2 cells broad at the base Se ae vermicular). aes bi vend IA vee dull. . Gemmae linear-vermicular with one primordium, less commonl oblong with 24 primordia (tr eee eve subtropical). LL. P. ap 14. Gemmae oblong or ce uc to linear-obconic. 15. Stems 1 ara simple; gemmae coe obconic or oblong, green, ean yaline or orange (temperate and = subarctic North Amer ae) 25. P. annotina sere atten forking, gemmae broadly oblong or ee reddish- tooo (Mexico). . P. pseudo-barbula ot 1]. Plants without axillary gemmz 16. Rhizoidal tubers Sonar pre 17. Plants shiny yellow-green; eae laxly and broadly rhomboidal; tubers red (Mexico). 36. P. integra 17. Plants Soa or somewhat glossy green; cells linear rhomboidal, tubers aa orange or orange-brown (temperate North America). 28. P. De uriana ers lac hig 18. Leaf margins strongly recurved. Stems red, leaves longly lanceolate; cells linear-hexagonal, thin-walled. ] n a ae a . P. crudoides 19. Stems green, leaves lanceolate; cells rhomboidal, rather thick-walled, a umes porose. . P. cardotu Leaf margins plane (except the perichaetial bracts). 20. Cells broad, 13 4m or more wide. 21. Plants dull whitish, in calcareous seepy or wet areas. 29. P. wahlenbergi 21. Plants yellow-green, reddish or blackish, at least below, in calcareous or acidic, usually wet sites 22. Plants oes robust, to | cm or more high, often blackish mae a 9 ee pale green just near the tips. eee Brean 22. Plants ae 3-8 mm high, yellowish or Rees. 23. Plants slender, yellowish, ee hat glossy 30. P. melanodon 23): ie tiny, populations ae consisting of more ae a few . P. atropurpurea a a ems, re 20. one narrow, es 10(-12) em wide. 24. Cells sabe sopemet thick-walled. re Plants dioicous; leaves nearly entire. 1. P. sphagnicola 25. ae paroicous; leaves + serrate near the apex. 2. P. nutans or 3. P. elongata 24. Cells with thin to firm walls. 26. Plants whitish, with the leaves often erect and seriate. 27. Plants slender, to | cm ee 32. P. brevinervis Qk eee medinnl size to ro Leaves erect and often seriate, lanceolate. 62 bolanderi var. seriata + to elliptic. 4. P. cruda 26. Plants shiny or dull green, the leaves sometr rect but not whitish. 29. Plants paroicous. 30. Plants dull or tutes cae pale green, 1.5-3.5 cm high Mexico, Central America). 5. P. oerstediana 30. Plants, aes glossy, aes or pale green, 0.5-2.0 mm ee 31. Leaves long-laneolate, erect, coarsely serrate near he ape ele longi 31. Leaves broadly lanceolate, spreading to erect- re - 4. ae: 28. Leaves + spreading, not seriate, broadly eee , serrate or serrulate near the apex. P. cruda 29. Plants see 32. Leaves widespreading, narrowly lanceolate; ec ial bracts very longacunina ate, to6mm long. — 35 P.lonibrarteat 32. Leaves erec + spreading, ee ee: or broadly lanc late; peri es bracts shorter, to 3-4 n 33. Plants robust; leaves broadly acute to » rounded, long- decurrent, ee low on the stem often pin iat P ludwigit 226 os oe Plants slender to medium size; leaves acute, shortly or not at all decurrent, ae er leaves green (all leaves + reddish ers alan ca). 34. Plants high in the ann or in the north. 35. Stems red. 36. Plants slender with erect-appressed leaves. 33. P. vexans 36. Plants medium size ae erect-spreading, soft-appearing leave 16. P. erecta 5. Stems green, orange or a own 37. Plants of South and Central ane pale Cae Bh, Plants of the western United Be and anada. 6. P. bolanderi Plants at or near sea level. 38. Plants very glossy light green. . P. pacifica 38. Plants dull or slightly glossy, green or ees Plants dull or slightly glossy, Jae peri- gonial bracts erect or erect-spreading. 200F: a 39. Plant dull reddish; perigonial Bee wi spreading. apes ee oD OS ae KEY TO PLANTS BEARING SPOROPHY TES 1. Exothecial cells plainly bulgin 2. Gemmae red or orange, shortly oblong to isodiametric; leaf primordia mies P. richardsit 2. Gemmae green, yellow, pink or hyaline, linear-vermicular or less commonly oblong: primordia erect, rarely poorly developed and incurved. . P. papillosa 1. Exothecial cells not bulging. 3. Annulus absent; exothecial cells little “ be longer than wide. Cells laxly and broadly rhomboidal, 12 um or more wide. 5. Plants tiny (2-6 mm), often seria stomates superficial or somewhat imme cells of perichaetial eae ts narrow (6-10 um). 30. P. cope D: are small to robust (5-35 mm), yellowish or whitish- ee most stomz deeply immersed; cells or perichaetial oe broad, 12-20 um broad s Plants pale, whitish-green, 1-5 cm 28. P. wahlenbergu Plants yellowish glossy-green, 0.5-1. i cm n high. 29. P. melanodon — C a linear-hexagonal, 6-10 um wide. 7. Plants robust; leaves wide-spreading; perigonial bracts long- inser to 6 long. os ingbrceat 7. Plants slender; leaves erect to erect-spreading; perigonial bracts ee to : 4mm long. 8. elas superficial or slightly sunken. . Stems elongate, to 1 cm or more high; leaves erect, whitish; cells en al leaves linear, 6-9 ps m wide. 31. P. brevinervis 9. Stems short, 2-6 mm; leaves few, spreading (except pobre ae bac cells of leaves on sterile plants or ie stem leaves broadly hexag 10-11 pm or more wide. UD ca cane 8. Stomates + deeply immersed. 10. Stems red, leaves small and erect, glossy gre 32. P. vexans 10. Stems green to orange; leaves larger, erodes dull, often sev ee oo columbica 3. eee present; exothecial cells elongate-rectangular. I Capsule erect; basal membrane low; segments narrow or rudimentary, not keeled. 12. Endostome segments long, + perforate; leaf margins recurved; cells rhombic, thick-walled. 5. P. cardotii 12. Endostome segments ae reduced to absent; leaf margins plane; cells linear-rhomboidal:; thin-w 16. Capsule ee inclined; fel membrane low to high; segments broad and keeled or reduc 13. ee S aie long-rectangular, 60-90 2m long, with straight or slightly sinuose walls. 4. a aae segments narrowly or not at all split along the keel; cilia rudi- mentary or absent, rarely long . erecta ia) on 225 15. Exostome teeth yellowish-white, coarsely papillose from apex to base. . P. crudoides 15. Exostome teeth brown to yellow-brown or hyaline, more coarsely Babee in the upper 4—% than below . Some or all plants with more than « one seta per Base . P. polycarpa 16. Plants with one seta per inflorescence. 17. Plants with gemmae; exostome teeth + blunt, eae in the upper half, nearly smooth below. P. pseudo-barbula 17. Plants without gemmae; exostome teeth acute or acuminate, coarsely papillose in the upper 3, gradually less papillose below. 18. Plants a oe oe thick-walled. 19 4-7 mm long; basal membrane 12 a exostome meee segments keeled. P. elongata 19. Capsule 2-4 mm long; basal membrane to Ys the exostome length; segments eae: keeled, often ular . P. elongata var. greenii = 18. Plants eG glossy; cells thin-walled. 20. fee dioicous; leaves serrulate to serrate a the . P. bolandert 20. Plants paroicous; leaves coarsely serrate - he apex Plants very glossy (Canada and the -P. longicollis 21. Plants slightly glossy, pale green (Mexico and Central America). 5. P. oerstediana 14. Endostome Oe broadly split along the keel; cilia long. 22. Plan . dio 23. Plants growing with Sphagnum; leaves not decurrent; oe with thickened wall P. sphagnicola 23. Plants on ae or subalpine soil; leaves longly decurent cells with see to firm walls. _P. ludwigit 22. Plants paroic 24. Basal eo low; segments narrow, scarcely keeled Gaise roadly perforate); cells lax and thin-walled, 12-20 4m obtusifolia 24. pee membrane 4 the exostome length; segments broad and keeled; s linear, 6-10 wm wide. 25. Leaves dull; cells with thickened walls. 2. P. nutans 25. Leaves slightly to strongly glossy; cells with thin walls. 26. Leaves pa, lanceolate to elliptical, serrulate to serrate 4. P. cruda 1S tt 26. Leaves noes aaneeann coarsely serrate at the apex. 5. P. oerstediana : me cells short siesta 25-60 ym long; with strongly sinuose walls Plants with axillary ger Species 18-26 27, Beats without axillary ee mae Cuecnas tubers sometimes present) 28. Plants oe ae eee urrent; costa of peric Secamse nee u ened at the base. wer ludwigt 28. Plants ee ie Vile or not at ae ae costa of per Sie bracts not much broadened at the 29. Plants very glossy; leaves ne serrate at the apex; cells 11 ise) ho ~I 180 wm lon 2H 5 pacifia 29. Plants not or slighty eee leaves serrulate to serrate at saa cells 65-120 pin | 28. eee l. ons ca ae (B. & S.) Lindb. & Arn., K. Svensk. Vet. Ak. Handl. 23: Oo: ee Pi B. & S. in B.S. G., Bryol Eur. 4: 158. 1846. (Lectotype: BM-Schimp. Webera sphagnicola (B. & S.) Schimp., Coroll. 6. 1856. Slender, dull-green plants generally scattered among Sphagnum. Stems 10- 30 mm, simple or forking when sterile, simple or rarely innovating below the 226 inflorescence when fertile; leaves erect-spreading, distant, lanceolate, 1.0- 1.6 mm long, entire to serrulae at the apices; upper cells hexagonal to rhom- boidal, 25-60 wm long, 6-9 um wide, thick-walled. Dioicous; perichaetial bracts to 3.5 mm long; perigonial bracts + long-acuminate, erect to erect-spreading. Capsules inclined 90-145° from vertical, cylindric-pyriform, 3-5 mm. long; exothecial cells elongate-rectangular, 50-90 wm long, with straight’ walls; stomates superficial; annuli of 2-3 cell rows, revoluble; opercula conic, acute to apiculate; exostome teeth orange-brown, acute-triangular, most coarsely papil- lose near the apices, bordered, trabeculate; endostome hyaline; basal membrane 72 the exostome length; segments keeled, broadly perforate; cilia long, nodulose. Spores 10-15 wm, almost smooth. Fig. Restricted to Sphagnum hummocks in bogs and fens; near sea level or montane. Northern Asia, Europe and North America; Oregon, Washington, Alberta, Manitoba, Quebec, Nova Scotia, Prince Edward Island, Newfoundland: Greenland. Map. |. Pohha sphagnicola is circumboreal, but apparently rare throughout its range. It is restricted to Sphagnum hummocks but even there it is far less common than P. nutans. Habitat alone is not diagnostic for P. sphagnicola; in fact, Hedwig’s type of P. nutans is mixed with a bit of Sphagnum. Bruch, Schimper and Giimbel described a variety sphagnetorum for P. nutans to cover forms growing with Sphagnum since they often have long stems and distant leaves. Pohlia sphagnicola differs from P. nutans in having entire or weakly serrulate leaves, shorter cells, and smaller, more nearly smooth spores, in addition to dioicous inflorescences. The two are no doubt closely related, but the differences are, for the most part, consistent (Fig. 2). In an unpublished study at the Uni- versity of Alberta, R. Belland made the same conclusions using different quanti- tative methods. 2. Pohlia nutans (Hedw.) Lindb., Musci Scand. 18. 1879. Webera nutans Hedw., Spec. Musc. 168. 1801. (Lectotype: G: Hedw.— Schwaegr.). Bryum trichodes Hedw., Spec. Mus. 168. 1801 (nom. nud. in synon.—W, nutans Hedw.). B. interruptum Dicks., Pl. Crypt. Brit. fasc. 4. 13. 11f. 6. 1801. (Lectotype: BM-—Dicks.). B. nigricans Dicks., Pl. Crypt. Brit. fasc. 4. 15. 11f. 10. 1801. (Lectotype: BM-—Dicks.). B. sericeum Huds. ex With., Syst. Arr. Brit. Pl. ed. 4, 3: 822. 1801 (nom. illeg. incl. spec. prior—w. nutans). Hypnum nutans (Hedw.) Web. & Mohr, Ind. Mus. Pl. Crypt. 3. 1803. Bryum nutans (Hedw.) Turn., Musc. Hib. 117. 1804. Webera bicolor Hoppe & Hornsch. ex Hornsch. 2: 102. 1819. (Isotype: G: Hedw.—Schwaegr.). Bryum insulare Houtt. ex Steud.,! Nomencl. Bot. 2: 88. 1924 (nom. nud. in synon.—W. nutans Hedw.). Bryum subdenticulatum Brid., Bryol. Univ. 1: 634. 1826. (Lectotype: B-Brid.; Isotype: G: Hedw.—Schwaegr.). Webera strangulata Nees ex Hueb., Musc. Germ. 428. 1833. (2lsotype: BM— Ha — mpe). Lamprophyllum nutans (Hedw.) Lindb., Act. Soc. Sc. Fenn. 10: 25. 187 | Article 46c of the international code of botanical nomenclature states that ex is appropriate in author citations only where the name was validly published. However, ex is used in this and similar citations for bibliographic completeness. FIG. 1. Pohkhia sphagnicola. A. Annulus, X290. B. Capsules, x 15. C. Exothecial cells, 290. D. Upper leaf cells, x 290. E. Stomate, x 290. F. Peristome, x80. G. Male plant, x 16. H. Leaves. x22. Webera canaliculata C. Mull. & Kindb. in Macoun, Cat. Canad. Pl. 6: 113. 1892. (Lectotype: S—Kindb.; Isotype: BH). Bryum cucullatiforme Kindb., Eur. N. Am. Bryin. 2: 387. S—Kindb.: Isotype: BH). Bryum nutans ssp. macounu Kindb., Eur. N. Am. Bryin. 2: 385. 1897. (Lecto- type: S—Kindb.; Isotype: BH). B. nutans ssp. canaliculatum (C. Mull. & Kindb. in Macoun) Kindb., Eur. N. Am. Bryin 2: 385. 1897. Bryum canaliculatum (C. Mull. & Kindb. in Macoun) C. Mull., Gen. Muse. Fr. 221. 1901. 1897. (Lectotype: 20 Fo) fo) fe) 72 fe) 18r e fe) 9 fe) ro) E F cine. O — o e) ‘16+ 2 ® N a sd O hed 14+ ‘a ® o e a . e 7,) Ce 12} ee 1 1 l l n n 1 r L 1 L 12+ Oo r) e * . a 0 10+ = CP o e c O O o. 6° Oo O fe) — 8 e .° mes A an 3 6 ° = F e eo 4} e ee E e © @ ot e l 1 1 1 Ll L 1 L l 1 1 1 L of es 20 30 40 50 60 70 cell length--pm FIG, 2. Scatter diagrams comparing Pohha nutans 7 os sphagnicola with respect to leaf cell length, length of teeth on leaf margins, and spore diamete Bryum pertenellum Bryhn. & Ryan in Bryhn, Rep. Sec. Norw. Arctic Exp. Fram no. 11. 1906. (Lectotype: O). Pohlia polygama Kindb., Rev. Bryol. 34: 92. 1907. (Lectotype: S—Kindb.). Bryum graniticum Warnst. & Geh. ex Podp., Mag. Bot. Lapok. 18: 21. 1919. (nom, inval. in synon.—P. nutans). Pohlia subdenticulata (Brid.) Podp. in Gyoerffy, Mag. Bot. Lapok. 18: 21. 1919. Webera polygama (Kindb.) Broth. in Engler & Prantl, Nat. Pfl. ed. 2, 10: 360. 1924. Bryum implexum Sw. ex Podp., Act. Ac. Sc. Nat. Moray. 17: 91. 1945 (nom. nud. in synon.—P. nutans). Bryum compactum Aust., Bot. Gaz. 2: LLL. 1877. (Lectotype: NY). 2g) Slender to medium-size, dull-green plants scattered singly or forming ex- tensive turfs. Stems 4-30 mm, simple or forking when sterile, simple or rarely innovating below the inflorescence when fertile. Leaves erect to spreading, crowded to distant, lanceolate, 0.8-1.6 mm long, serrate to serrulate near the apices; upper cells linear-rhomboidal, 50-90 ym long, thick-walled. Paroicous; perichaetial bracts to 4.0(—5.0) mm long. Capsules inclined 90-—145° from verti- cal, cylindric-pyriform, 3-5 mm long; exothecial cells elongate-rectangular, 50- 90 wm long, with straight walls; stomates superficial; annuli of 2-3 cell rows, revoluble; opercula conic, acute to apiculate; exostome teeth yellow to orange- brown, acute-triangular, most coarsely papillose near the apices, bordered, trabeculate; endostome hyaline, rarely yellow-brown; basal membrane 2 exo- stome length; segments keeled, ae perforate; cilia long, nodulose. Spores 16-22 ym, distinctly roughened. I In rock crevices, on mineral oe rotting logs, tree bases, roadbanks and mixed with Sphagnum; sea level to alpine. Across Canada and the northern United States, Europe and Asia; widespread at high latitudes of the Southern Hemisphere; south in North America to Maryland, Virginia, Missouri, Arizona, New Mexico and California (reported from Georgia and Arkansas); known in Mexico from only two localities (Potosr and Nevado de Toluca); apparently absent from the New World tropics. Map 2 Exsiccati: Austin, Musci Appal. 510 as B. elongatwm (UBC); Bauer, Musci Eur. Exs. 318-320 as P. nutans (ALTA), 909-910 as P. nutans (WTU), 911 as P. nutans f. umbrosa (WTU), 1095, 1096a & bas P. nutans v. caespitosa (WTU), 1097 as P. nutans v. subglobosa (WTU); Bauer, Bryoth. Bohem. 23 sphagnetorum (WTU); Boros, Plantae Hungar. Exs. 878 (CAS); Brotherus, Bryoth. Fenn. 335 as P. nutans v. longwseta (WTU); Drummond, Musci Amer. 262 as B. ae (MICH); Grout, N. Am. Musci Perf. 23 as P. nutans (UC, CAS); ee Musci Acro. Bor.-Am. 143a & b as W. commutata (MICH, NY), 166 as W. nutans (MICH, UC, WTU), 167 as W. nutans v. Se (MICH, UC, WTU), 524-525 as W. nutans (MICH, UC, met ); Kobsch, Bryoth. Sax. 50 as P. nutans (MICH), 421 as B. nutans f. elata (MICH), 526 as P. nutans v. caespitosa (MICH); WMaccun: Canad. Musci 160 as W. nutans (UC); Noguchi, eee: 840 as P. nutans (MICH); Rabenhorst, Bryoth. Eur. 329b as B. nutans (UC), 584 as a nutans (UC); Sull. & Lesq., Musci Bor.-Am. (ed. 1) 179 as B. nutans (MICH), 178 as B. nutans M ,ed. 2, 267 as B. nutans (UC, MICH), 268 as B. nutans (MICH); Warnstorf, Mark. Laubm. 64 as W. aa (C AS), 175 as W. nutans v. sphagnitorum (CAS). ~ Pohlia nutans is characterized by its thick-walled cells giving the plants a dull appearance, and paroicous inflorescence. The endostome is well-developed with keeled, broadly perforate segments and nodulose cilia. When sterile, plants are difficult to distinguish from P. elongata, except by habitat and range. Even some fruiting specimens approach P. elongata in having more narrowly perforate endostome segments, and (or) short cilia, making a definite determination impossible. One poorly developed specimen from central Mexico (Nevado de Toluca, Rzedowski 21676, US) is unusual in having a yellow-brown endostome with rather narrowly split segments. The type of Webera luisieri Dix. from Madeira also appears to be decrepit P. nutans with a yellow endostome. I do not consider such forms worthy of nomenclatural recognition, at least on the basis of the rather poor material available. Although rare, a few collections otherwise referable to P. nutans appear to be dioicous. Crum and Anderson (1981) reported one such New York collection (Smith & Glider 43837, as questionable P. sphagnicola. Pohlia schimperi (C. Mull.) Andr. is said to differ from P. nutans in its lush habit, red color, and dioicous inflorescence. North American specimens determined by Andrews as P. schimperi are all rather unexceptional P. nutans, and I have seen only one Alaskan collection (Peters Lake, Steere 650725-21) that agrees with the type of P. schimperi. The differences between the two species are not convincing. I have excluded P. schimperi from the North American flora, but have not placed it in the synonomy of P. nutans pending study of more collections. he many infra- FIG. 3. Pohlia nutans. A. Exothecial cells, x290. B. Capsules, X15. C. Peristome, x80. D. Annulus, 290. E. Female plant, x5. F. Stomate, x290. G. Exostome tooth, lateral view showing relative thickness of dorsal and ventral plates and trabecula, x 250. H. Leaves, x22. I. U per leaf cells, x 290. specific taxa proposed under P. nutans should be evaluated in the light of a world-wide study, 3. Pohlia elongata Hedw., Spec. Musc. 171. 1801. Bryum cylindricum Dicks., Pl. Crypt. Brit. Fasc. 4, 12. 11f. 4. 1801. (Lectotype: B M—Dicks.). Bryum elongatum (Hedw.) With., Syst. Arr. Brit. Pl. ed. 4, 3: 815. 1801. Leskea elongata (Hedw.) Web. & Mohr, Ind. Muse. Pl. Crypt. 3. 1803. Mnium elongatum (Hedw.) P. Beauv., Prodr. 75. 1805. ~ M. pohlia Hoff. ex P. Beauv., Prodr. 75. 1805. (nom. elongatum). Pohlia arcuata Hoppe & Hornsch. ex Hornsch., Flora 2: 98. 1819. (Lectotype: GZU-Hoppe). Bs ete tale us & Hornsch. ex Hornsch., Flora 2: 96. 1819. (Lecto- type: GZU-Hoppe) P. tenella Hoppe & Hornsch. ex Hornsch., Flora 2: 95. 1819. (Lectotype: GZU-Hoppe). Webera caespitosa Hoppe & Hornsch. ex Hornsch., Flora 2: 102. 1819. (Lecto- type: GZU-Hoppe). Pohlia acuminata Hoppe & Hornsch. ex Hornsch., Flora 2: 94. 1819. (Lecto- type: GZU: Hoppe; Isotype: G: Hedw.—Schwaegr.). P. polymorpha Hoppe & Hornsch. ex Hornsch., Flora 1: 100. 1819. (Lecto- Roe GZU: Hoppe). poe ee Hoppe & Hornsch. ex Hornsch., Flora 2: 97. 1819. (Lectotype: GZU: Hoppe). P. pulchella Hoppe & Hornsch. ex Hornsch., Flora 2: 95. 1819. (Lectotype: GZU: Hoppe; Isotype: G: Hedw.-Schwaegr.). P. xanthocarpa Hoppe & Hornsch. ex Hornsch., Flora 2: 97. 1819. (Lecto- type: GZU: Hoppe). Webera macrocarpa Hoppe & Hornsch. ex Hornsch., Flora 2: 104. 1819. (Iso- type: G: Hedw.—Schwaegr.) Pohlia cylindrica (Dicks.) Hornsch., Flora 2: 93. 1819 fey recognita Reichenb., Flora 5: 540. 1822 (nom. nud. in synon.—P. elongata). P. minor var. affins (Hoppe & Hornsch. ex Hornsch.) Brid., Bryol. Univ. 7: 612. cone P. elongata var. cylindrica (Dicks.) Brid., Bryol. Univ. 7: 610. 1826. eee als var. brachycarpa (Hoppe & Hornsch. ex Homi h.) Brid., Bryol. Univ. 7: 627. 1826. Webera elongata var. acuminata (Hoppe & Hornsch. ex Hornsch.) Schwaegr., Suppl. Spec. Pl. ed. 4, 5 P. elongata var. acuminata (Hoppe & Hornsch. ex Hornsch.) Hueb., Muse. Germ. 473. 1833. le fee var. affinis (Hoppe & Hornsch. ex Hornsch.) Hueb., Muse. Germ. 470. je, Iohmorpa var brachycarpa (Hoppe & Hornsch. ex Hornsch.) Hueb., Musc. Germ. 470. les ee ca var. arcuala (Hoppe & Hornsch. ex Hornsch.) Schwaegr. ex Hueb., Musc. Germ. . 18: Bryum boissiert D. Not., Syll. 134. 1838. (Lectotype: R: De Not.). a po iare Hoppe & Hornsch. ex Hornsch.) Hartm., Handb. Skand. Fl. ed. : 296. 18 : Oi ee var. macrocarpum (Hoppe & Hornsch. ex Hornsch.) B. & S. in B.S. G., Bryol. Eur. 4: 102. 1839. B. acuminatum (Hoppe & Hornsch. ex Hornsch.) B. & S., Bryol. Eur. 4: 91. 1839. Bryum alpinum var. Wepre ets & Hornsch. ex Hornsch.) Lesq., Mem. Soc. Sc. Neuchatel. 3: 37. °B. integridens C. Mull., Syn. 1: aon 1848 (type not seen). B. spectabile C. Mull., Syn. 2: 583. 1851. (Lectotype: H-Br.). Webera acuminata (Hoppe & Hornsch. ex Hornsch.) Schimp., Coroll. 64. 1856. W. acuminata var arcuata (Hoppe & Hornsch. ex Hornsch.) Schimp., Coroll. 64. 1856 W. acuminata var. tenella (Hoppe & Hornsch. ex Hornsch.) Schimp., Coroll. 64. 1856 ed W. elongata var. macrocarpa (Hoppe & Hornsch. ex Hornsch.) Schimp., Coroll. 65. 1856. Webera polymorpha (Hoppe & Hornsch. ex Hornsch.) Schimp., Coroll. 65. 1856. vi ns var. affinis (Hoppe & Hornsch. ex Hornsch.) Schimp., Coroll. 65. 1856 ee pobimnp var. brachycarpa (Hoppe & Hornsch. ex Hornsch.) Schimp., Coroll. 65. 185 ee ie elongatum (Hedw.) Lindb., Act. Soc. Sc. Fenn. 10: 27. 1871. W. spectabilis (C. Mull.) Besch., Mém. Soc. Sc. Nat. Cherbourg 16: 196. 1872. 2W. integridens (C. Mull.) Schimp. ex Besch., Mem. Soc. Sc. Nat. Cherbourg 16: 196. 1872. W. cylindrica Schimp. ex Besch., Mém. Soc. Sc. Nat. Cherbourg 16: 196. 13/2. (Isotype: NY; ?Isotype: BM). Bryum cylindricum Mont. ex Besch., Mém. Soc. Sc. Nat. Cher bourg 16: 196. 1872 (nom. inval. in synon.—W. cylindrica). Webera muelleriana Schimp. ex Besch., Mem. Soc. Sc. Nat. Cherbourg 16: 196. 1872. (Isotype: BM: Schimp.). W. gracilicarpa Hampe, Vid. Medd. Naturh. For. Kjoebenh. ser. 3, 6: 149. 1875. W. ambigua Limpr., Laubm. Deutschl. 2: 237. 1891. (Lectotype: BP-Limpr.). B. elongatum ssp. ambiguum (Limpr.) Kindb., Eur. N. Am. Bryin. 2: 390. 1890. B. elongatum ssp. acuminatum (Hoppe & Hornsch. ex Hornsch.) Kindb., Eur. N. Am. Bryin. 2: 390. 1890. Bryum muellerianum (Schimp. ex Besch.) C. Mull., Gen. Musc. Fr. 220. 1901. Bryum ambiguum (Limpr.) Kindb. ex Par., Ind. Bryol. Suppl. 56. 1900 (nom. inval. in synon.—B. elongatum spp. ambiguum). Pohlia ambigua (Limpr.) Broth. in Engler & Prantl, Nat. Pfl. 1(3): 548. 1903. P. muelleriana (Schimp. ex Besch.) Broth. in Engler & Prantl, Nat. Pfl. 1(3): 547. Ae P. spectabilis (C. Mull.) Broth. in Engler & Prantl, Nat. Pfl. 1(3): 547. 1903. ?P. integridens (C. Mull.) Broth. in Engler & Prantl, Nat. Pfl. 1(3): 547. 1903. P. elongata var. EES (Hoppe & Hornsch. ex Hornsch.) Herz., Bot. Jahrb. 47: 482. 1912 P. acuminata var. . nella (Hoppe & Hornsch. ex Hornsch.) Amann, Fl. Mouss. Suisse 2: 178. 1919. P. acuminata var. arcuata (Hoppe & Hornsch. ex Hornsch.) Amann, FI. Mouss. Suisse 2: 178. 1919. P. polymorpha var. acuminata (Hoppe & Hornsch. ex Hornsch.) C. Jens. Danm. Moss. 2: 392. 1923. Pohlia zacatecana Williams, Bryol. 26: 33. 4. 1923. (Isotype: a ). Webera zacatecana (Williams) Ther., Smiths. Misc. Coll. 81: 1928. Webera polymorpha var. acuminata (Hoppe & Hornsch. ex Hor Her C. Jens., Skand. Bladmfl. 166. 1939. Pohlia polymorpha ssp. acuminata (Hoppe & Hornsch. ex Hornsch.) Podp., Consp. 331. | : ie a var. arcuata (Hoppe & Hornsch. ex Hornsch.) Podp., Consp. S51, 19 P. minor var. tenella (Hoppe & Hornsch. ex Hornsch.) Wyk & Marg., Taxon 8: 74. 1959. P. minor var. arcuata (Hoppe & Hornsch. ex Hornsch.) Wijk & Marg., Taxon 8: 74. 1959. P. minor var. brachycarpa (Hoppe & Hornsch. ex Hornsch.) Wijk & Marg., Taxon 8: 74. 1959. 233 P. minor ssp. acuminata (Hoppe & Hornsch. ex Hornsch.) Wijk & Marg., Taxon 8: 74. 1959 P. minor var. pulchella eee & Hornsch. ex Hornsch.) Dem. & Wilz., Bull. Jard. Bot. Nat. Belg. 48: 449. 1978. Slender to medium size, dull-green plants forming small populations, often in rock crevices. Stems 5-25 mm, simple or forking when sterile, frequently innovating below the inflorescence when fertile. Leaves erect to erect-spreading, crowded to distant, lanceolate, 0.9-1.7 mm long, serrulate to serrate near the apices; upper cells rhomboidal to linear-hexagonal, 45-85 um long, thick-walled. Paroicous or rarely autoicous; perichaetial bracts little to strongly differentiated, to 3.5mm, perigonia in autoicous plants terminal on short to elongate side branches, the perigonial bracts shortly acute. Capsules inclined 10°-90° from vertical, shortly to longly cylindric-pyriform, 3.5-7.0 mm long; exothecial cells elongate-rectangular, 50-110 4m long, with straight walls; stomates superficial; annuli of 2-3 cell rows, revoluble; opercula conic, acute; exostome teeth yellow to brown, acute-triangular, most coarsely papillose near the apex, bordered, trabeculate; endostome hyaline; basal membrane 4—% the exostome length; segments keeled but narrowly perforate; cilia short, absent, rarely long and + nodulose. Spores 16—23 um; distinctly roughened. Fig. Roadbanks and streambanks; frequent in rock crevices in arctic or alpine regions; reportedly epiphytic in New Guinea. Widespread at high latitudes across the three northern continents. In North America, apparently absent from the Cascade-Sierra Nevada system of the western United States, but extending southward in the Appalachians to Tennessee and North Carolina. Ranging in the Rockies from Alaska to Arizona, and extending in the mountains through Mexico, Central America and, through scattered records, to southern South America; Africa, China, Nepal, S. E. Asia, Phillipines, Japan, and New Guinea. Map 3 Exsiccati: Austin, Musci pees (Suppl. 1) 510 as a fee (CANM, NY); Bauer, Musci Eur. Exs. 903 as P. acuminata ae TU); Bauer, Bryoth. Bohem. 233 as W. elongata v. oder f. facets Sar (BM); Erbar, Crittogam. Ital. 609 as ie ee (UC, NY): Fleischer. Musci Frond. A ee Indici 285 as W. brachydonted (NY); ca ut, N. . Musci Perf. 398 as P. elongata (CANM, MICE , UC); Hartman, cand. 376 as W. eu (BH): Ha a Musci Acro. Bor.—-Am. 145 as i Gara DANM, MICH NY, UC); Husnot, Musci Gall. 461 as W. elongata (BH, BM, NY); Macoun, Canad. Musci 521 s W. Paes (MICH), 806 as W. ee (CANM); Noguchi & Hattori, Musci Jap. 386 as P. to bees (ALTA, BH, MICH, NY); Pringle, Plant. Mex. 10623 & 10648 as W. cylindrica (MICH, MO); a orst, Bryoth. Eur. 275 as W. elongata (1 Nee ae 360 as W. ee (NY, UC); Sc hultz, Flora t Germ. Exs. 1185 as B. elongatum (BM); Sull. & Lesq., Musci Bor. Amer. (ed. 1) 1 rongann (BH, MICH, NY), (ed. 3) 266 as B. eee (BH, MICH, NY, UC), 270 as B. luduigii (BH). — Pohha elongata is distinguished from related species by its (usually) paroicous inflorescence, longly cylindric capsules (4-7 mm), and especially the narrowly o scarcely perforate endostome segments. Unlike P. longicollis and P. bolanderi, which have similar peristomes, the leaves are dull with cell walls incrassate. The sexuality of P. elongata var. elongata is almost uniformly paroicous, although a few specimens, including the type of Webera ambigua Limpr., are autoicous. Capsule size and shape are extremely variable. Mexican and Central Amer- ican plants generally known as P. spectabilis are characterized by exceptionally long, narrow capsules. However, other than in exothecial cell length which is obviously correlated in development with capsule length, such plants can scarce- ly be distinguished consistently from more typical P. elongata. Although a some- what coherent distribution in Mexico and Central and northern South America suggests at least limited genetic distinctiveness for P. spectabilis, frequent inter- gradation makes recognition of two taxa impractical. A specimen from Virginia [*\ las anal (zal i \p aa RSE FO ect a ——— ( SS OSS.» Se li FIG. 4. Pohlia elongata. A. Stomate, X290. B. Peristome, X80. C. leaves, X22. D. Upper cells of branch leaf, x 290. E. Habit, x7. F. Lateral view of annular cells, X290. G. Annulus, x 290. H. Upper cells of perichaetial bract, x 290. (Brown, 1890, UBC) also approaches P. spectabilis. In Mexico, as both the “specta- bilis” and typical expressions, P. elongata is more common then anywhere else in North America. At the other extreme, scattered collections of P. elongata from arctic, alpine, or other harsh habitats are characterized by a suite of distinctive attributes. These plants can be distinguished as the var. greenii. 3b. Pohlia elongata var. greenii (Brid.) Shaw, comb. nov. Pohlia minor Schwaegr., Spec. Musc. Suppl. 1: 70. 64. 1816. (Lectotype: G: Hedw.—Schwaegr.; Isotype: B—Brid.). 235 ee tes mene Hornsch. ex Hornsch., Flora 2: 100. 1819. (Lectotype: G: Hedw.—Schw Tee ae eee & Hornsch. ex Hornsch., Flora 2: 99. 1819. (?Isotype: M) P. curviseta Hoppe & Hornsch. ex Hornsch., Flora 2: 98. 1819. (Lectotype: GZU: Hoppe; Isotype: G: Hedw.-Schwaegr.; ?Isotype: NY). P. green Brid., Mant. Musc. 115. 1819. (Lectotype: B: Brid.). P. minor Schwaegr. var. greenii (Brid.) Brid., Bryol. Univ. 1: 612. 1826. P. elongata var. minor (Schwaegr.) Hartm., Handb. Skand. FI. ed. 2L 336. 1832. P. polymorpha var. minor (Schwaegr.) Hueb., Musc. Germ. 470. 1833. P. ees var. polyseta (Hoppe & Hornsch. ex Hornsch.) Hueb., Muse. Germ. 470. 18 mae oe var. minus (Schwaegr.) Hartm., Handb. Skand. FI. ed. 3: 296. 1838. Webera gracilis (Hoppe & Hornsch. ex Hornsch.) Schimp., Coroll. 65. 1856. W. acuminata var. minor (Schwaegr.) Schimp., Coroll. 64. 1856 W. acuminata var. polyseta (Hoppe & Hornsch. ex Hornsch.) Schimp., Coroll. 64. 1856. Bryum nudicaule Lesq., Mem. Calif. Ac. Sc. 1. 21. 1868. Webera nudicaulis (Lesq.) Lesq. & James, Man. N. Am. Moss. 220. 1884. Pohlia EU i es (Hoppe & Hornsch. ex Hornsch.) Braithw., Brit. Moss Fl. 2 18 Pohha ee eee aBronh ex Engler & Prantl, Nat. Pfl. 1(3): 548. 1903. P. brachycarpa var. polyseta (Hoppe & Hornsch. ex Hornsch.) Schwaegr. ex Par., Ind. Bryol. ed. 2, 5: 105. 1906. P. acuminata var. minor (Schwaegr.) Amann, Fl. Mouss. Suisse 3: 178. 1919. P. acuminata var. ia (Hoppe & Hornsch. ex Hornsch.) Amann, FI. Mouss. Suisse 3: 178. 1919. . minor var. gracilis (Hoppe & Hornsch. ex Hornsch.) Wijk et Marg., Taxon 8: 74. 1959. Tiny, dull-green plants scattered singly or in small populations. Stems very short, with 5-10 leaves. Leaves scarcely 1 mm long, broadly lanceolate; upper cells rhombic, 35-65 um long, thick-walled. Paroicous, autoicous or rarely dioicous; perichaetial bracts undifferentiated, broadly lanceolate, to 1.5 mm long; perigonia in dioicous plants terminal on short, simple stems, the bracts shortly and broadly acute, erect. Capsules inclined 145°-190° from vertical seg- ments broadly pyriform, 2-3.5 mm long; basal membrane low, up to % the exostome length, segments often ir peter linear, scarcely keeled, narrowly perforate to entire; cilia absent. Fig. 5 Rock crevices in tundra areas, less frequently on exposed soil. Alaska, Wash- ington, California, Colorado, Utah, Newfoundland; Greenland. Reported from northern Europe. Ma Although no single character is diagnostic, several morphological attributes distinguish the var. greeni from so-called typical expressions: smaller overall size; shorter, broader leaves, more strongly inclined, broader, shorter capsules, low basal membrane, irregular segments, and complete lack of cilia. Ironically, it is Hoppe and Hornschuch’s extensive list of “new species” that provides the most persuasive argument against recognizing the var. greenii at the specific level. With over ten taxa, they (Hoppe and Hornschuch ex Hornschuch 1819) typified practically every conceivable intermediate between the long and short capsule forms. However, sporophytic and gametophytic differences be- tween the varieties, coupled with geographic differences, warrant separation, at least on the basis of North American plants. Unlike P. spectabilis, the var. greenii Tai N B —_L N L \ | : K ye FIG. 5. A-D. Pohlia elongata var. elongata. A. Capsules, Mexican plants commonly called P. spectabilis, X 13. B. Exothecial cells from capsules as in A, x 230. C. Capsules, typical North American lants, X13. D. Exothecial cells from capsules as in C, x230. E-J. Pohlia elongata var. greenwi. E. Exothecial cells from capsules as in F, x 230. F. Capsules, x 13. G. Leaves, x 18. H. Peristome, X58. I. Stomate, X230. J. Upper leaf cells, x 230. differs from the var. elongata in more than just capsule shape, and notably so in endostome morphology. Webera carneoides Dix & Badw. of the Himalayas repre- sents the var. greenii at its extreme. ‘The plants are minute, with capsules scarcely longer than wide. It is difficult to typify the var. greenii because so many names are available representing the extensive intergradation that occurs between plants with long and short capsules. Most of the old types are fragmentary and could scarcely endure yet one more peristome dissection. My concept of the var. greenii is the 23d short capsule- reduced endostome form at or near its extreme (Fig. 5). Extensive variation in capsule shape is allowed in the var. elongata. The names P. polymorpha and P. acuminata have been used for plants with relatively short capsules. Speci- mens in Hoppe’s herbarium with these names have intermediate to elongate eee (e.g. Fig. 5c), falling within the range of the var. elongata. 4. Pohlia cruda (Hedw.) Lindb., Musci Scand. 18. 1879. Mnium crudum Hedw., Spec. Musc. 189. 1801. (Lectotype: G: Hedw.- Schwaegr.). Hypnum crudum (Hedw.) Web. & Mohr, Ind. Mus. Pl. pee 3: 1803. Webera cruda (Hedw.) Fuernr., Flora 12(Ergbl. 2): 35. 1829 Lamprophyllum crudum (Hedw.) Lindb., Act. Soc. Sc. Fenn. 10: C7187 1. Webera cruda var. nivalis Podp. ex Herz., Wien. Bot. Zeitschr. 93: 45. 1944. (Lectotype: BRNM Robust to slender, shiny, pale whitish-green plants generally forming small populations. Stems 5-20 mm, simple or forking when sterile, usually simple when fertile. Leaves wide- spreading to erect-spreading, distant to crowded, lanceolate to elliptic, sometimes + asymmetric, 1.0-2.0 mm long, serrate to serrulate at the apices; upper cells linear-rhomboidal, 70-120 ym long, 6— 94m wide, thin-walled. Paroicous or dioicous; perichaetial bracts variable, lanceolate to linear, to 4.0-5.0 mm long; perigonial bracts longly acuminate, to 5.0 mm long. Capsules inclined 10°-135° from vertical, narrowly or rarely broadly cylindric-pyriform, 4—6 mm long; exothecial cells elongate-rectangular, 40-80 44m long, with straight walls; stomates superficial; annuli of 2-3 cell rows, revoluble; opercula conic; exostome yellow-brown to red-brown; acute-triangu- lar, bordered, trabeculate; endostome hyaline; basal membrane 4 the exostome length; segments keeled, broadly Bena cilia long, nodulose. Spores 18— 26 um, rather coarsely papillose. Fig. 6 Soil of roadbanks, streambanks, andes often in crevices under rocks or exposed roots; rarely on rotting wood. Circumboreal; across northern North America south to California, Arizona, Tennessee and North Carolina; Mexico (Puebla, Tlaxcala, Mexico); Chile; Antarctica; New Zealand; Auckland Islands; Hawau; India; Japan; reported from eastern Asia. Map 5 Exsiccati: Allen, Mosses of the Cascade Mts. 58 as B. crudum (MICH); Bauer, Musci Eur. Exs. 311 as P. cruda (ALTA); Drummond, Musci Amer. 269 as B. crudum (MICH, UC); Grout, N. Am. Musci Perf. 398a as P. elongata (MICH), 443a as P. cruda (CAS, MICH, UC), 443b as P. cruda (MICH); Holmen, Bryoph. Dan. Exs. 155 as W. cruda (ALTA, UC); Holzinger, es Acro. Bor.-Am. 144 as W. cruda v. minor (MIC a re Kobsch, Bryoth. Sax. 336 as W. cruda (MICH); Macoun, Canad. Musci 161 as W. cruda (UC), 4 s W. cruda (MICH), ae as W. micro- plata (MICH); Noguchi, Musci Jap. 1134 as P. cruda (} (ice; Rabenhorst, Bryoth. Eur. 1024 as W. cruda (CAS); Sull. & Lesq., Musci Bor. Am. (ed. 1) 179b as B. cruduwm (MICH); nee Mark. oe 270 as W. cruda (CAS). ~ Readily distinguished by its pale, whitish-green color and strong gloss, P. cruda is not to be confused with any other Podlia. It is further characterized by the longly cylindric capsules and well-developed endostome with broadly perforate segments and long cilia. I have seen a few sterile Pohlia collections with elongate stems (to 2 cm) and glossy pale-green leaves. These are distinctive, with the leaves lanceolate, erect, and overlapping. One such gathering (N.Y., Westchester Co., Shaw, 26 Dec. 1981, MICH) grew on seepy granitic rock in beech-maple forest. The habitat, as well as the aspect of the plants, brought Mielichhoferia to mind. In the absence of sporophytes, these are interpreted as P. cruda, mostly by default. Ecologically and morphologically this expression is well characterized. ad | <= FIG. 6. Pohlia cruda. A. Exothecial cells, 290. B. Leaves, X22. C, Capsules, X15. D. Habit, «4.5. E. Stomate, X290. F. Peristome, x80. G. Annulus, 290. H. Upper leaf cells, x 290. 5. Pohlia oerstediana (C.. Mull.) Shaw, comb. nov. Bryum oerstedianum C. Mull., Syn. 2: 583. 1851. (Lectotype: H-Br; Isotype: BM-—Hampe). Webera oerstediana (C. Mull.) Jaeg., Ber. S. Gall. Naturw. Ges. 1873-74: 128. 13795. Bryum seleri C. Mull., Bull. Herb. Boiss. 5: 181. 1897. (Lectotype: H-Br). Pohlia seleri (C. Mull.) Broth. in Engler & Prantl, Nat. Pfl. 1(3): 548. 1903. Webera orizabensis Card., Rev. Bryol. 40: 38. 1913. (Holotype: PC). Robust, shiny to dull, pale whitish-green plants often in deep turfs. Stems (5-) 10-30 mm, simple or forking when sterile, generally simple when fertile. 239 Leaves erect to erect-spreading, crowded and imbricate, lanceolate, 1.5-2.5 mm long, coarsely serrate near the apices; upper cells linear-rhomboidal, 70-135 wm long, 6-11 wm wide, thin-walled. Paroicous; perichaetial bracts gradually en- larged, to 3.5 mm long. Capsules inclined 10°-100° from vertical, longly cylin- dric, to ca. 6 mm long; exothecial cells elongate-rectangular, 40-80 um long, with straight walls; stomates superficial; annuli of 2-3 cell rows; opercula conic; exostome yellow to red-brown, acute-triangular, bordered, trabeculate; endo- stome hyaline; basal membrane 2 the height of exostome teeth; segments keeled, broadly to narrowly perforate; cilia long to short, sometimes nodulose. Spores 19-25 um, rather coarsely papillose. Fig. 7. cS we PO SI = Q oy = Od ) iw) iv) SS S ey =) SS) \ s RB) a) a) > =) en) a SS } a 2, FIG. 7. Pohlia oerstediana. A. Peristome, X80. B. Capsules, x 15. C. Leaves, X22. D. Exothecial cells, X290. E. Stomate, x290. F. Habit, x4. G. Annulus, x290. H. Upper leaf cells, x 290. 240 Soil, moist rocks, tree bark, rotting logs, at high elevations in Oak-Pine forests. Mexico (Hidalgo, Mexico, Oaxaca, Puebla, Tlaxcala, Veracruz). Guatemala (San Marcos, Huehuetenango) and Costa Rica (Volcan Irazu). Map 6. Exsiccati: Pringle, Plant. Mex. 10548 as W. cylindrica (FH). Pohlia oerstediana is distinguished from P. cruda by its larger size, and less glossy leaves that are coarsely serrate near the apex. It is exceptionally robust in microscopic details as well as overall habit (Fig. 7). Vhis species seems to be more common than P. cruda in Mexico and Central America, with P. cruda more or less restricted to alpine habitats. Most or all records of P. cruda given by Bartram (1949) should be referred to P. verstediana. I have seen few capsules of P. verstediana with intact peristomes, but the segments seem to be variable with respect to perforation. The Asian Webera laticuspis Broth. is very similar to P. verstediana gametophytically and has nar- rowly perforate segments. Such a disjunction, though interesting, is not unheard of among mosses (e.g. Avengstroemia orientalis Mitt.). 6. Pohlia bolanderi (Sull.) Broth. in Engler & Prantl, Nat. Pfl. 1(3): 548. 1903. Bryum bolandert Sull., Mem. Calif. Acad. Sei. 1. 22: 1868.(Lectotype: NY). Slender to medium-size, glossy green plants in compact turfs. Stems 3—10 (—20) mm, simple or forking when sterile, simple when fertile. Leaves erect to erect-spreading, crowded and imbricate, lanceolate, 1.0—1.6 mm long, serrate to serrulate at the apices; upper cells linear-rhomboidal, 55-100 zm long, 6- 9 wm wide, thin-walled. Dioicous; perichaetial bracts scarcely differentiated, to 2 mim long; perigonial bracts shortly acute, erect. Capsules inclined 35°-90° from vertical, elongate-cylindric, 3-5 mm long; exothecial cells elongate-rectangular, 40-85 pm long, with straight walls; stomates superficial; annuli of 2-3 cell rows, revoluble; opercula conic, acute; exostome teeth yellow to yellow-brown, acute- triangular, most coarsely papillose near the apices, bordered, trabeculate; endo- stome hyaline; basal membrane 3—' the exostome length; segments keeled but narrowly perforate; cilia short, rarely long. Spores 16-26 wm, distinctly rough- ened. lig. 3. Rather dry alpine or subalpine soil, less commonly along streams, high montane to alpine. Apparently endemic to western North America: British Columbia; Washington, Oregon, California, Nevada. Map 7. In addition to the dioicous inflorescence, P. bolanderi is distinguished from its close relative, P. longicollis, by the smaller overall size and less coarsely serrate leaves. The distinctive geographic range of P. bolanderi gives meaning to these relatively slight morphological differences. The distributions of the two species overlap only in southern British Columbia. Reports of P. longicollis from Cali- fornia, Oregon, and Washington are referable to P. bolanderi. 6a. Pohlia bolanderi var. seriata, Shaw, var. nov. Plantae erectae, strictae, dense confertae, plerumque steriles. Folia pallida, dense seriata, imbricata, lanceolata, carinata. Flores dioici. Perichaetia parva, inconspicua, bracteis paucis. Dull, whitish plants often in deep, luxuriant turfs. Stems erect, up to 5.0 cm high. Leaves + distinctly arranged in 4—5 vertical rows, pale green to whitish, erect to erect-spreading. Dioicous; perichaetia terminal on simple stems, incon- spicuous, the bracts not differentiated from lower leaves. Perigonia and sporo- phytes as in the var. bolanderi. Big, 9, i i ( m = + = ; SS" S| FES FF ea ee SEZ iN SS SS E On 1, =a | Bai cae FIG. 8. Pohlia bolanderi. A. Capsules, X15. B. Annulus, X290. C. Exothecial cells, x290. D. Leaves, x22. E. Habit, x9. F. Stomate, 290. G. Peristome, X80. H. Upper cells, x290. TYPE: U.S.A. Colorado, Gilpin Co.: 5 mi. E of Corona Pass on Corona Pass Rd., Shaw 2628 (Holotype: MICH; Isotype: J. Shaw). Paratypes: Colorado, Garfield Co: 30 mi. N of Glenwood Springs, Hermann 24272 (COLO). Washing- ton, Pierce Co.: Mt. Rainier Nat. P., above Paradise Pt., Shaw 2453 JJ. Shaw). Moist snowmelt areas in the alpine zone. Colorado, Utah, Wyoming, Wash- ington. Map 7. e var. seriata differs from the var. bolanderi in its luxuriant growth and extremely pale leaves often arranged more or less distinctly in vertical rows. At their extremes, the two varieties are very distinct, although several collections FIG, 9. Pohha bolanderi var. seriata. A. Habit, X35. B. Costae, transverse se base (above) and apex (below), 290. C. Upper leaf cells, x290. D. Leaves, x3 ctions near the leaf 35: show intergradation (e.g. Shaw 2448, ALTA: Lawton 2833, WTU). When well developed, the var. seriata looks much like Conostomwm ltetragonum (Hedw.) Lindb. Whereas the var. bolanderi is often fertile, | have seen only one specimen of the var. sertata with sporophytes (Utah, Cache Co., Behling 288, NY). One other collection (Shaw 2678, ALTA) has female inflorescences, but all other specimens seen are completely sterile. Geographic ranges of the two varieties are different, though overlapping (Map 7). The var. seriata is characteristic of lush snowmelt areas whereas the var. bolanderi grows most frequently on rather dry tundra soil. 7. Pohlia longicollis (Hedw.) Lindb., Musci Scand. 18. 1879. Webera longicollis Hedw., Spec. Musc. 169. 41. f. 1-5. 1801. (Lectotype: G: Hedw.—Schwaeer.). Bryum longicolle (Hedw.) Sw., Monthl. Rev. 34: 538. 1801. Hypnum longicolle (Hedw.) Web. & Mohr, Ind. Mus. Pl. Crypt. 3: 1803, Webera alpina Hoppe & Hornsch. ex Hornsch., Flora 2: 104. 1819. (Lecto- type: GZU—Hoppe; Isotype: G: Hedw-Schwaegr.). Bryum grimsulanum Brid., Bryol. Univ. 1: 628. 1826. (Lectotype: B-Brid.; Isotype: G: Hedw.-Schwaegr.). Mnyrium crudum var. longicolle (Hedw.) Wahlenb., Fl. Suec, 2: 725. 1826. Webera grimsulana (Brid.) Brid., Bryol. Univ. 2: 829. 1827. Bryum hornschuchii Spreng., Syst. Veg. 4: 212. 1827 (nom. illeg. incl. sp. prior.—W., alpina, W. grimsulana). 243 VV ae ee ae var. alpina (Hoppe & Hornsch. ex Hornsch.) Fuernr., Flora 12(Ergbl. 29, W. eee var. grimsulana (Brid.) Fuernr., Flora 12(Ergbl. 2): 36. 1829. ye eee var. alpinum (Hoppe & Hornsch. ex Hornsch.) B. & S. in B.S. G., Bryol. Eur. 4: 102. 1839. Sis clavata Sendtn., Flora 23: 61. 1840 (hom. illeg.; Lectotype: M). Bryum lucidum Jay in Durand, J. Ac. Nat. Sc. Philadelphia ser. 2, 3: 202. 1856. (Lectotype: Lamprophyllum eee (Hedw.) Lindb., Act. Soc. Sc. Fenn. 10: 26. 1871. Bryum nitens Kindb., Bih. K. Svensk. Vet. Ak. 7: 66. 1883 (hom. illeg.; Lecto- type: S—Kindb.). B. elongatum ssp. longicolle (Hedw.) Kindb., Bih. K. Svensk Vet. Ak. Hand. 7 66. 1883 B. nitescens Kindb., Bot. Not. 196 (based on B. nitens). Webera pennina De Not. ex Par., Ind. Bryol. 1353. 1898 (nom. nud. on synon.— W. ee V. elongata var. alpina eye ppe & Hornsch. ex Hornsch.) Herib., Mem. Ac. 310, ‘1 — Sc. C See Ferrand ser. 2, Pohlia longicollis vee alpine (Hoppe & Hornsch. ex Hornsch.) Williams, Bull. New York Bot Gard. 364. 1902. Webera nitescens ae Kindb. ex Macoun, Cat. Canad. Pl. 7: 246. 1902. Pohlia longicollis var. grimsulana (Brid.) Amann, Fl. Mousse. Suisse 2: 179. ( Robust, shiny green plants frequently forming small populations in rock crevices. Stems 8-30 mm, generally simple though sometimes forking. Leaves erect to erect-spreading, crowded and imbricate, lanceolate, acute to acuminate, 1.5—2.6 mm long, coarsely serrate near the apices; upper cells linear rhomboidal, 80-140 pm long, 6-9 ym wide, thin-walled. Paroicous; perichaetial bracts gradu- ally differentiated from lower leaves, to 3.8 mm. Capsules inclined 10°—90° from vertical, elongate-cylindric, 4-6 mm long; exothecial cells elongate-rectangular, 50-90 wm long, with straight walls; stomates superficial; annuli of 2—3 cell rows, revoluble; opercula conic, blunt to acute; exostome teeth yellow to brown, acute- triangular, most coarsely papillose near the apices, bordered, trabeculate; endo- stome hyaline; basal membrane 43— the exostome length; segments keeled but eagle) perforate; cilia short, rarely long. Spores 16-23 ym, distinctly rough- ened. yy Sail of roadbanks and streambanks, frequently in rock crevices with P. crudoides at high latitudes. Circumboreal; across northern North America south in the Rockies to Colorado and in the Appalachians to Tennessee and North Carolina; absent from the Cascade-Sierra Nevada systems. Japan; India; re- cently reported from East Africa (Mt. Kenya). Map ; ati: Bauer, Musci Eur. Exs. 908 as W. ocoe cae (BH, COLO); Eee: Musci Amer. 270 as ene (C ANN M, MICH, NY); Grout, N. Am. Musci Perf. 398 as P. elongata (CAS, COLO), Hartmann, Bryac. Scand. Exs. 164 as W. longicolla (BH, COLO), viene & Nestler, Surpes Crypt. 1409 as W. ele (BH, CANM); Rabenhorst, Bryoth. Eur. 359 as W. longicolla (NY, UC), 873 as W. longicolla (NY Characterized by its large size, glossy, crowded leaves, and paroicous in- florescences, P. longicollis is strikingly beautiful. Like P. elongata and P. bolanderi, P. longicollis has the endostome segments narrowly (or scarcely) perforate. 8. Pohlia polycarpa (Mitt.) Broth. in Engler & Prantl, Nat. Pfl. 1(3): 548. 19038. Bryum polycarpum Mitt., Linn. Soc. Bot. 12: 293. 1869. (Lectotype: NY) Webera polycarpa (Mitt.) Jaeg., Ber. S. Gall. Naturw. Ges. 1873-74: 134. 1875. Bryum aggregatum C. Mull., Bull. Herb. Boiss. 5: 181. 1897. (Lectotype: BM-—Hampe). — = Ese. OE \..-{ ae Al a == Cy UV —=Fp= (oh | \ {| 4 i FIG. 10. Pohlia longicollis, A. Exothecial cells, x 290. B. Capsules, X15. C. Annulus, x290. D. Leaves, X22. E. Stomate, x290. F. Peristome, x80. G. Upper leaf cells, «290. Webera aggregata (C. Mull.) Par., Ind. Bryol. 1343. 1898. Pohha aggregata (C. Mull.) Broth. in Engler & Prantl, Nat. Pfl. 1(3): 547. 1903. P. pluriseta Herz., Beth. Bot. Centralbl. 26: 71. 14. 1910. (Lectotype: JE). Slender to tiny, glossy green plants forming low turfs. Stems 3-10 mm high, generally simple when sterile, branching by I-3 innovations below the peri- chaetia, simple or with a subfloral innovation when bearing perigonia. Leaves erect to erect-spreading, crowded and imbricate, lanceolate, 0.7—1.4 mm long, serrate to serrulate at the apices; upper cells linear-rhomboidal, 50-95 wm long, 245 thin-walled. Dioicous; perichaetial leaves lanceolate to linear-lanceolate, scarcely differentiated or gradually longer than the lower leaves, to 2.5 mm, perigonial bracts shortly acute, erect to erect-spreading. Capsules inclined [0°-95° from vertical, narrowly cylindric, 2.5—4 mm long, |-4 per perichaetium; exothecial cells elongate-rectangular, 50-90 wm long, with straight walls; stomates super- ficial; annuli of 2-3 cell rows, revoluble; opercula convex to conic, apiculate; exostome teeth yellow-brown to brown, acute-triangular, most coarsely papillose near the apices, bordered, trabeculate; endostome hyaline to very pale yellow; basal membrane ca. 43 the exostome length; segments weakly to strongly keeled, not or scarcely perforate; cilia rudimentary to absent. Spores 16-20 pm, very finely roughened. Fig. 11. DOPO CG, 00AD cS OD (a) o) 6! Jeno nH FIG. 11. Pohlia polycarpa. A. Lateral view of annular cells, «290. B. Leaves, including a long perichaetial bract with recurved margins, X22. C. Stomate, x290. D. Upper leaf cells, X290. E. Peristome, X80. F. Habit, x8. G. Exothecial cells, x 290. H. Capsule, x15. I. Annulus, 290. 246 On alpine soil; Guatemala (Volcan del Fuego—type, Tajumulco); Bolivia; Peru, Chile. Map 9. An Andean species, P. polycarpa is known in Central America from only two localities in Guatemala. Distinguished at once from related species by the poly- setous condition, P. polycarpa is otherwise very similar to the northern P. bolandert. Webera costesti Card. & Ther. is synonymous with P. polycarpa according to a note in the New York Botanical Garden written by ‘Theriot. 9. Pohlia obtusifolia (Brid.) L.. Koch, Leafl. West. Bot. 6: 20. 1950. Bryum obtusifolum Brid., Musc. Rec. 2(3): 52. 1803 (type not seen). Bryum cucullatum Schwaegr., Spec. Musc. Suppl. 1(2): 94. 68. 1816. Pohlia cucullata (Schwaegr.) Bruch ex Hochstetter, Flora 1: 274. 1826. Bryum vogesiacum Dub. in Cand., Bot. Gall. 2: 552. 1830. (Lectotype: RO). Webera cucullata (Schwaegr.) Schimp., Coroll. 66. 1856. W. cucullata var. hausmannii De Not., Atti Univ. Genova 1: 430. 1869. (Lecto- tvpe: RO). Lamprophyllum cucullatm (Schwaegr.) Lindb., Act. Soc. Sc. Fenn. 10:87. 1871. Pohlia cucullata (Schwaegr.) Lindb., Musci Scand. 19. 187 Webera microcaulon C. Mull. & Kindb. in Macoun, Cat. Canad. Pl. 6: 114. 1892. (Lectotype: S—Kindb.). W. polymorphoides Kindb. in Macoun, Cat. Canad. Pl. 6: I 11. 1892. tvpe: S—Kindb.). Pohlia polymorphoides (Kindb. in Macoun) Broth. in Engler & Prantl, Nat. Pfl. 3): 348. 1903. Bryum obrutum Stirt. ex Dix., J. Bot. 61: 71. 1923 (nom. nud. in synon.—P. cucullata). Brywm oxoniense Dix., Bryol, a2: 2. 1929. (Lectotype BM—Dix: Isotvpe: BH). Pohlia serrulata Bruch ex Podp., Consp. 337. 1954 (nom. nud. in synon.—P. cucullata). Lecto- — l — Very small to elongate, pale-green to blackish, rather dull plants in low to deep turfs. Stems 3—8(—25) mm high, simple or rarely forking when sterile, simple or rarely with I—4 innovations when fertile. Leaves spreading to erect, crowded to distant, broadly lanceolate, broadly acute, subtly cucullate, 0.7- I.fmm long, serrulate to serrate at the apices: upper cells broadly rhombic, 30-55(-70) wm long, 12-16(-20) wm wide, lax and thin-walled. Paroicous: perichaetial bracts scarcely to moderately differentiated, to 2.8 mm long. Cap- sules I160°-180° from vertical, broadly pyriform, 2.0-3.5 mm long; exothecial cells elongate-rectangular, 40-75 win long, with straight walls; stomates super- ficial; annuli of 2-3 cell rows, revoluble; opercula blunt or umbonate, rarely acute-conic; exostome. teeth slenderly triangular, most coarsely papillose near the apices, bordered, trabeculate; endostome hyaline; basal membrane low, up to ¥3 the exostome length; segments + keeled but narrow and delicate, rather broadly perforate; cilia rudimentary to absent. Spores 17-25 ym, distinctly roughened. Fig. 12. . On soil, typically in the alpine zone, frequent in moist, rich snowmelt areas. Circumboreal; in North America known from Alaska, Yukon Territory, North- west’ Lerritories, British Columbia, Alberta, Washington, California, Montana, Wyoming, Colorado, Utah, Labrador, Newfoundland, Quebec, Vermont: India: Japan. Map 10. Exsiccati: Bauer, Musci Eur. et Amer. Exs. 1925 as P. cucullata (CANM), Musci Eur. exs. 312 as P. cucullata (BH); Erbar, Crittogam Ital. O12 as W. cucullata (NY, UC); Husnot, Musci Gall. 462 as W. ludwigtt (BH, NY); Lisowski, Bryoth. Polon. 613 as P. cucullata (CANM): Macoun, Can. Musci 468 as W. cucullata (NY), 636 a W. polymorphoides (NY); Mougeot & Nestler, Stirpes Cryptog. 830 as P. cucullata (CANM, NY); Rabenhorst, Bryoth. Eur. 442 as W. cucullata (NY); 1215 as W. ludwigt (NY); Annulus, x290. D. 12. Pohlia obtusifolia. A. Upper leaf cells, x290. B. Capsules, x 15. ves, X22. F : G: Stomate, x290. E. Lea fF. Peristome, x80. G. Exothecial cells, x 290. Sull. & Lesq., Musci Bor. Am. (ed. 1) 176 as B. cucullatum (MICH, NY), (ed. 2) 265 as B. cucullatum (BH, MICH, NY). Pohla obtusifolia is readily distinguished from other species by a combination of gametophytic and sporophytic attributes: paroicous inflorescence, lax, thin- walled leaf cells, and the delicate endostome segments which, although poorly developed, are broadly perforate. Fruiting plants are typically very small (3-6 mm high), but sterile forms growing in unusually moist sites can reach 2—4cm. Such forms rarely even produce gametangia, and are most reliably identified by the subtly cucullate 248 leaves and broad cells (12-13 wm or more wide). It is characteristic of elongate expressions to be almost black except for the apical few leaves which are pale- reen. Pohlia obtusifolia is not obviously related to any other North or Central American Pohlia. Vhe non-sinuose exothecial cells and revoluble annuli indicate a possible relationship to the previous eight species, but the endostome is more similar to that of P. cardoti. In spite of extensive inquiries to European and North American herbaria, I have been unable to typify P. obtusifolia. However, | accept the name rather than using the slightly later P. cucullata primarily because after Sc hwaegrichen de- scribed Brywm cucullatum in 1816, he noted (1830) that his species was synony- mous with Bridel’s Bryum obtusifolium, Untortunately, there is no specimen of B. obtusifolium in Schwaegr ichen’s herbarium. 10. eon e Serres (Sull. & Lesq.) Broth. in Engler & Prantl, Nat Pfl. 1(3): aa a Sull. & Lesq., Proc. Am. Ac. Arts & Sc. 4: 278. 1859. (Iso- type: NY-Mit c. Webera crudoides (Sull. & Lesq.) Jaeg., Ber. S. Gall. Naturw. Ges. 1877-78: 440. 1879. Pohlia crassinervis Lindb., Bot. Centralb. 6: 363. 1881. (Lectotype: H-SOL). P. crassidens Lindb., Rev. Bryol. 10: 5. 1883. (Lectotype: H-SOL). Webera trachydontea San., Bot. Centralbl. 13: 247. 1883. (Lectotype: H- ae F Bryum crassidens (Lindb.) Kindb., Bih. K. Svensk. Vet. Ak. Handl. 7 : 66. Webera crassidens (Lindb.) Kindb., Forh. Vid. Selsk. Christiania 1886; 22. 1888. W. crassinervis (Lindb.) Par., Ind. Bryol. 1343. 1898. Bryum trachydonteum (San.) C. Mull., Gen. Muse. Fr. 220. 1901. Medium size to robust, somewhat glossy plants frequently forming small populations in rock crevices. Stems 4—10(-25) mm, deep cherry-red, generally simple but sometimes forking near the base when sterile. Leaves erect- -spreading to erect, rather distant, narrowly or rarely broadly lanceolate, acute, |.0-2.2 (—2.6) mm long, serrate to serrulate at the apices; with strongly recurved mar- gins; upper cells linear-hexagonal, 35-60(-85) wm long, with thin walls. Dioicous; perichaetial bracts lanceolate to linear lanceolate, to 3 mm long; perigonial bracts shortly to longly acuminate, erect to erect-spreading. Capsules inclined 30°—90° from vertical, ovoid-cylindrical, 2.5—4 mm long; exothecial cells shortly to longly rectangular, 30-80 ae ae with straight to weakly sinuose walls; stomates superficial; annuli of 2-3 cell rows, revoluble or separating irregularly; opercula convex to conic, ee to bluntly apiculate; exostome teeth whitish to yellow, slenderly triangular, rounded at the apex, coarsely papillose throughout, scarcely bordered, weakly trabeculate; endostome whitish to hyaline; segments narrow and scarcely keeled, very narrowly perforate to entire; basal membrane low, to 43 the exostome length: cilia absent or rarely rudimentary. Spores 15-25 ym, finely roughened. Fig. | On soil especially in rock crevices in tundra paras according to Ochi (1959), on soil also in the deciduous zone of Japan (as var. revolvens). Alaska, Yukon Territory, Northwest ‘Territories and British Columbia; Greenland; northern Europe and Asia; Japan; India. Map ee Noguchi, Musci Jap. 936 as « crudoudes (CANM, NY); Savicz-Lubitzkaja, Hep. et Musci U.R.S.S. Exs. 96 as P.crudoides (CANM, CAS). 2658 ; eek 20 oo8 aa af ue Sen eD ie WR RE eed 9208 yy 13. Pohlia crudoides. A. Upper leaf cells, x290. B. Capsules, x15. C. Annulus, x 290. D. Habit, . E. Exothecial cells, x 290. F. Peristome, x80. G. Lateral view of exostome tooth showing weakly cee trabecula, x80. H. Leaves, perichaetial bracts on the leaf, x22. I. Stomate, x 290. Pohlha crudoides can be recognized by cherry-red stems and obviously re- curved leaf margins. The leaves are generally long, narrow and somewhat suff, although I have seen a few sterile plants from the Yukon Territory with broader, less stiff leaves and hardly recurved margins. Pohlia crassinervis Lindb. typifies such expressions, which do not seem to justify separation. Pohlia crassidens Lindb. and Webera trachydontea San. are based on the same type specimen (collected in Opdal by Kaurin). There is no specimen represent- ing W. trachydontea in the Sanio herbarium (Rusinka in litt, 1979). To avoid future ambiguity, I have chosen a single specimen in Lindberg’s herbarium to 250 lectotypify both names. The Himalayan type of Webera timmioides Broth. is close tou. ee ee solely by its very robust stature. Both syntypes lack sporophyte 11. Pohlia papillosa (C. Mull. ex Jaeg.) Broth. in Engler & Prantl, Nat. Pfl. 1(3): 552. 1903 Bryum papillosum C. Mull., Syn. 1: 326. 1848. (Lectotype: NY; Isotype BM-— Besch.). hom. illeg.—non B. papillosum Dicks. B. tenellum Schimp. ex C. Mull., Syn. 2: 572. 1851. (Lectotype: BM—Schimp.; Isotype: NY). Brachymenium tenellum Cari ex C. Mull.) Schimp. ex Besch., Mém. Soc. Sc. Nat. Cherbourg | Or2: ~ 3. papillosum (C. Mall ex Jaeg., Ber. S. Gall. Naturw. Ges. 1873-74: 122. ee Schimp. ex Besch., Ann. Sci. Nat. Bot. ser. 6, 3: 204. 1876. ee BM-—Besch.; Isosyntype: NY, BM—Hampe). Bryum ay ae C. Mull, Bull. Herks Boiss. 5: 550. 1897. (Lectotype: Brachymenium mammillosum (C. Mull.) Par., Ind. Bryol. Suppl. 39. 1900. Pohlia mammillosa (C. Mull.) Broth. in Engler & Prantl, Nat. Pfl. 1(3): 552. Pali subglobosa (Schimp. ex Besch.) Broth. in Engler & Prantl, Nat. Pfl. 1(3): TOU: re papillosa (C. Mull. ex Jaeg.) ) Herz., Biblioth. Bot. 87: 79. 1916. W. mammillosa (C. Mull.) Broth. in Engler & Prantl, Nat. Pfl. ed. 2, 10: 362. 1924. W. ee (Schimp. ex Besch.) Broth. in Engler & Prantl, Nat. Pfl. ed. 2 10: 362. 19 Medium-sized dull green plants sometimes forming extensive turfs. Stems 9-25 mm, simple or commonly forking when sterile, with 1-4 innovations below the perichaetia, simple when bearing perigonia. Leaves erect-spreading, fre- quently distant on sterile stems, crowded when fertile, lanceolate, 0.7—1.4 mm long, serrate to serrulate at the apices; upper cells rhomboidal, 60-90(-1 10) zm long, 6-9 «um wide, with firm walls. Gemmae numerous in the upper leaf axils, arising in clusters in each axil, linear-vermicular, oblong, or obconic, 100-250 ym long but apparently indeterminate in growth, brown or whitish, pink or reddish, with I-4 peglike, erect, leaf primordia at the apex, the primordia sometimes 2-3 cells broad at the base. Dioicous; perichaetial bracts linear-lanceolate, to 3.0 mm long; perigonial bracts erect-spreading. Capsules inclined 10°-100° from verti- cal, broadly to narrowly pyriform, narrowed at the mouth, sometimes curved and slightly asymmetric, 2.5—4.0 mm long; exothecial cells isodiametric to shortly rectangular, 35-60 wm long, with straight or rarely sinuose, often strongly bulging walls; stomates superficial to somewhat sunken; annuli of 1(2) cell rows, separating irregularly; opercula convex and apiculate or shortly and bluntly rostrate (and sometimes curved upward); exostome teeth whitish to yellow, + fused near the base; slenderly triangular, + blunt at the apices, coarsely papil- lose throughout, scarcely bordered, weakly trabeculate; endostome whitish to hyaline, papillose; segments narrow and scarcely keeled, very narrowly perfor- ate to entire; basal membrane low, to 3 the exostome ee cilia absent or rarely rudimentary. Spores 17-21 um, finely roughened. Fig. 14-15 On soil of roadbanks or in other disturbed sites, most frequent at moderate altitudes in cloud forests; Mexico (Chiapas, Chihuahua, Hidalgo, Mexico, Michoacan, Nayarit, Oaxaca, Puebla, Querétaro, Veracruz); Cuba; Dominican Republic; Haiti: Jamaica; Guatemala; Costa Rica; Panama: Columbia, Venezu- ela; Peru; Ecuador; Brazil. Map — 2 \ > aes FIG. 14. Pohlia papillosa. A. Stomates, 290. B. Upper leaf cells, x 290. C. Perichaetial bracts, x22. D. Capsule, x15. E. Peristome, x80. F. Leaves, X22. G. Exothecial cells, x290. H. Annulus, x 290. MO). — Exsicatti: Pringle, Plantae Mex. 10635 as W. didymodontia Ecologically, P. papillosa replaces P. annotina and related species in the Neotropics. It reproduces by axillary gemmae, and is common in disturbed areas. The gemmae are variable in shape, encompassing most of the forms characteristic of P. proligera and P. annotina combined. They are usually very small (ca. 100 wm) and linear-vermicular with one primordium, but are some- times obconic with 2—4 peglike (or rarely more or less laminate) primordia. Many specimens have only dense clusters of small vermicular gemmiae. Several Mexican collections have very small ovoid gemmae with poorly developed leaf primordia gay BV ANSE a FIG. 15. Gemmae of Pohlia and Bryum species. A. Pohlia richardsit. B. Pohlia papillosa. C. Bryum sp., Fife 2827. D. Bryum gemmiluscens Wilz. & Dem., Dehle, 1824 (BM). (e.g. Bowers et al. 5385; Sharp et al. 1325; Iwatsuki & Sharp 53396; all in TENN). These may prove to be poor examples of P. papillosa, but require further study. Contrary to Andrews (1950), I find P. papillosa to be consistently distinct from the Old World, P. flexuosa Hook. The two are quite similar, but P. flexuosa has longly acuminate exostoine teeth (370-600 um vs 240-370 um in P. papillosa), completely separate to the base and segments distinctly keeled and narrowly perforate. The exothecial cells are often only weakly bulging whereas they are strongly so in P. papillosa. Gemmae of P. flexuosa are often scarce, but are generally linear-obconic to ovoid with 3-5 peglike primordia. I have seen P. flexuosa trom Japan, the Philippines, China, India, Nepal and Africa (Rwanda), and it is apparently endemic to the Old World. 253 A number of species have been segregated from P. flexuosa on the basis of differences in gemmae form. Andrews (1950) referred Pohlia delicatula (Mitt.) Broth. to the synonymy of P. flexuosa, but it is a clearly distinct species distin- guished by leaf shape and aereolation as well as gemmae morphology. Like the European P. le aca Wilz. & Dem., P. delicatula has two kinds of gemmae: linear with poorly developed primordia and tiny, isodiametric to ovoid types. Unlike P. eae the ovoid gemmae do not have protruding walls. The Hawaiian P. leucostomoides (Broth.) Bartr. has larger, translucent ovoid gemmae devoid of primordia in the leaf axils. 12. Pohlia richardsii Shaw, sp. nov. Plantae caespitosae vel gregariae. Caules fertiles ramosissimi, breves, steriles plerumque simplices, gemmifert duplo et triplo longiores, uniformiter foliosi, erecti. Folia lanceolata vel lineali-lanceolata, apice irregulariter serrata, inferne margine reflexa, areolis cellulis superioribus linearibus. Capsulae late pyri- formes. Gemmae abundantes, axillis foliorum summorum, intense rubrae, ovoideae. Dull green plants. Stems 5-15 mm, simple when sterile or bearing anther- idia, innovating rather abundantly when bearing archegonia. Leaves erect- spreading to ple when moist, scarcely altered on drying; 0.7—1.4 mm long, 0.3-0.6 mm wide, lanceolate, acute, somewhat decurrent by 3-4 rows of rectangular ee margins plane to + reflexed, serrate to serrulate near the apex or halfway to the base, entire below; costa ending 1-3 cells below the apex; upper cells linear-rhomboidal, (40—)50-80(-95) ym long, 6-9 wm wide, thin- to firm- walled, not or rarely vermicular, undifferentiated along the margin, rectangular near the base. Gemmae numerous to few in upper leaf axils, arising in clusters in each axil, rarely appearing single on older shoots, spheroidal, ovoid, obovoid, or obconic, about 110-160 ym long but apparently indeterminate in growth, deep red to orange-red, with 1-4 short, inconspicuous leaf primordia at the apex, the primordia rarely rudimentary, sometimes peglike, but mostly laminate (3 cells or more broad at the base), incurved over the apex, paler than the body but not green. Dioicous; perigonia bulbiform; outer bracts longly acute to acuminate, erect to erect-spreading from an erect base; perichaetia terminal, rather incon- spicuous; outer bracts to ca. 2.2 mm long, triangular- to linear-lanceolate; mar- gins recurved from base to apex. Sporophytes uncommon; setae 10-15 mm high, orange to orange-red; capsules inclined about 90°, 2.5-3.5 mm long, tan to brown, broadly pyriform, abruptly contracted to a well-developed neck; exo- thecial cells isodiametric to short-rectangular, 35-55 wm long, 18-40 ym aoe with non sinuose but bulging walls; stomates superficial to + sunken, 35-55 ps long; annuli of 1(2) cell row(s), separating irregularly; opercula bluntly convex to hemispheric; exostome teeth white to yellowish-white, narrowly lanceolate, + blunt, (200-)220-330(-370) um long, weakly bordered, hardly trabeculate, densely and rather coarsely papillose from apex to base dorsally; endostome white to hyaline; basal membrane ca. 4 the exostome length; segments linear to linear-laneolate, not or slightly keeled, not perforate; cilia none. Spores 10- 20 wm, finely papillose to nearly smooth. Fig. 15-16. TYPE: Costa Rica, Volcan Poas, on banks of volcanic ash near edge of water c. 2,500 m, 20 Feb. 1965, Richards R 6010 (Holotype: MICH). Paratypes: Colombia, Dept. Santander: Vicinity of California, 3000 m, Kilip & Smith (NY). Dept. Cundinamarca: ca. 1 km S of Usme, ca. 9000 ft., King et al. C-6000 (US). Ecuador, Prov. Napo: Paramo de Guamami, N of Quito-Baeza Rd., 4200 m, Steere E-250 (NY). Venezuela, Nova Granada, Linding (NY). n soil. Costa Rica (De Alajuela, San José); Colombia; Venezuela; Ecuador. Recents collected in the Dominican Republic (La Vega, Shaw 5621). Map 13. MIG. 16. Pohlia richards. A. Stomate, X290. B. Exothecial cells, x 290. C. Upper leaf cells, x290. D. Capsules, x15. E. Leaves, x22. Pohlia richardsu was collected by Paul Richards in 1965 and was subsequently sent to H. Crum, who recognized it as a new species. I am grateful to them for permission to describe it here. This is apparently a species of the nothern Andes and Central America, penetrating northward only to Costa Rica, though disjunct in the Dominican Republic. The only fruiting plants of P. richardsti available, all collected by D. Griffin and collaborators in Costa Rica, are unfortunatly mixed with P. papillosa. I was able to convince myself that at least some of the sporophytes belong to P. richards: gametophytes, although there is nothing to distinguish the capsules from those of P. papillosa. Vhis similarity is comparable to the uniformity among capsules of P. annotina and related species. Pohlia richardsii is distinguished from P. papillosa by the deep red to orange, spherical to short-oblong gemmae. The only time that P. papillosa approaches P. richardsu in gemmae form is in very poorly-developed expressions of the former that have few, small gemmae, with rudimentary primordia. In such cases, the gemmae are pale, rather than dark and translucent. Even in mixed specimens of P. papillosa and P. richardsii, it is possible to separate stems of the two. Pohlia richardsi can also be distinguished from such other propaguliferous species as P. pseudo-barbula and Pseudopohlia didymodontia by smaller gemmae. Gemmae like those of P. richardsii are found in several unrelated species of Bryaceae. For example, Bryuwm gemmilucens Wilz. & Dem. has gemmae practically indistinguishable from those of P. richardsii (Fig. 15). Similar forms occur on stems of a sterile and unidentified Brywmn collected by A. Fife in Newfoundland (Fig. 15), and those of P. andrewsii are also similar (Bigs 25): 13. Pohlia pseudo-barbula (Ther.) Crum ex Shaw, comb. nov. Webera pseudo-barbula Vheér., Smiths. Misc. Coll. 85: 28. 17. 1931. (Lectotype: PC; Isosyntype: F). 255 Slender dull-green plants. Stems 5-15 mm, generally unbranched when sterile, with I—4 innovations below the perichaetia, simple when bearing perigonia. Leaves erect to erect-spreading, crowded on the stem, lanceolate, 0.7-1.4mm_ long, serrate or serrulate to midleaf or below; upper cells rhomboidal, 50—75(-90) wm long (longer on perichaetial leaves) 6—9 um wide, with firm walls. Propagula present in clusters in upper leaf axils, ovoid to spherical or obconic, abruptly narrowed to a uniseriate stalk, red to orange, (140—) 160-230(-300) ym long, with 2-5 apical and subapical, erect or incurved leaf primordia, the primordia laminate in form (2-5 cells broad at the base). Dioicous; perichaetial bracts lanceolate to linear-lanceolate, to 3.0 mm long; perigonial bracts shortly to longly acuminate, erect-spreading. Capsules inclined 10°-90° from vertical, narrowly pyriform to cylindrical, 2.5-4.0 mm long, slightly curved and asymmetric; exothecial cells longly to shortly rectangular, 35-65 ym long, with somewhat sinuose walls; stomates superficial; annuli of 2 cell rows, separating irregularly; opercula conic, acute or apiculate; exostome hyaline to whitish, narrowly triangular, + blunt, coarsely papillose in the upper ¥%, more finely roughened below, scarcely bordered, weakly trabeculate; endo- stome hyaline; basal membrane low, to 13 the exostome length; segments linear but slightly keeled, not or very narrowly perforate; cilia absent or rarely rudi- mentary. Spores 13-20 ym, nearly smooth or somewhat roughened. Fig. 17 On soil, especially along roadsides in montane Oak-Pine forests. Appar a endemic to Mexico, where it is known from the volcanic region around Mexico City (Distrito Federal, Hidalgo, Michoacan, Puebla). Map This species is related to P. papillosa, as indicated by he weakly bordered and trabeculate, rather short exostome teeth, linear scarcely keeled, narrow ly split segments, and low basal membrane, lacking cilia altogether. Like P. papillosa, it reproduces by axillary gemmae. It is distinguished by narrowly cylindrical capsules with rectangular, nonbulging exothecial cells, and far less coarsely papillose exostome teeth that are almost smooth below. have examined all four of the syntypes cited by Theriot. The lectotype is an abundant specimen bearing both sporophytes and axillary gemmae. An addi- tional specimen collected by Brother Amable in Cuautzin (no. 1827) was re- ported by Theriot (1932). A duplicate in the Mitten collection (NY) agrees pre- cisely with the syntypes. Crum (1951) made the combination Pohlia pseudo- barbula in his thesis, but since that is not considered effective publication, the combination is validated here. The author citation should therefore be P. pseudo- barbula (Theér.) Crum ex Shaw. 14. Pohlia ludwigii (Schwaeer.) Broth., Act. Soc. Sc. Fenn. 19: 27. 1892. Leskea ludwign [Spreng. ex] Web. & Mohr, Ind. Musc. Pl. Crypt. 3: 1803. (nom. nud.). Bryum ludwigu Schwaegr., Spec. Musc. Suppl. 1(2): 95. 68. 1816. (Lectotype: G: Hedw.—Schwaegr.). Webera ludwigit (Schwaegr.) Fuernr., Flora pee 2) eoOs Loe: Bryum bigeloviu Sull., Rep. Expl. Sway. Rout. Railr. 4: 187. 5. 1856. (Lecto- type: FH). Webera breidler’ Jur., Verh. Zool. Bot. Ges. Wien 20: 167. 3b. 1870. (Lecto- type: : Bryum breidleri (Jur.) Hobk., Grevillea 1: 157. 1873. Webera bigelovu (Sull.) Lesq. & James, Man. N. Am. Moss. 223. 1884. Pohha bigelovii (Sull.) Broth. in Engler & Prantl, Nat. Pfl. 1(3): 50. 1903. Webera ludwigit var. breidlert (Jur.) Roth, Eur. Laubm. 2: 26. 1904. Bryum grande (Kindb.) Par., Ind. Bryol. ed. 2, 1: 217. 1904 (nom. inval. synon.—B, ie wigil). Mniobryum bigelovi (Sull.) Loeske, Stud. Morph. Syst. Laubm. 124. 1910. CO CUA CCAD, A / SEER Ay Ss, FIG. 17. Pohlia pseudo-barbula. A. Peristome, X80. B. Exothecial cells, x 290. C. Upper leaf cells, x 290. D. Annulus, 290. FE. Lateral view of annular cells, x 290. F. Capsule, x 15. G. Stomate, x 290. H. Leaves, x22. I. Gemmae, x 180. M. ludwigu (Schwaegr.) Loeske, Stud. Morph. Syst. Laubm. 124. 1910. Pohlia sudetica Podp. ex Gyoerfty, Mag. Bot. Lapok 18: 21. 1919 (nom. illeg. incl. spec. prior.—B. ludwigii). P. ludwigtt var. breidleri Jur.) Amann. FI. Mouss. Suisse 2: 182. 1819. Webera sudetica Giac., Atti Ist. Bot. Univ. Lab. Critt. Pavia Ser. 5, 4: 231. 1947 (nom. illeg. incl. spec. prior.—B. ludwigit). Mostly robust, dull green plants in deep turfs. Stems 8-35 mm, simple or frequently forking when sterile, simple when bearing perichaetia, simple or with . i) . © . . . = . . I-3 innovations when bearing perigonia. Leaves spreading to erect-spreading, 257 distant to crowded, ovate-lanceolate, acute to rounded at the apices, 1.5-2.0 (—2.6) mm long, serrate near the apices, + longly decurrent; upper cells rhombic to rhomboidal, 40-65(-85) um long, 7- “19 m wide, with firm to rather thin walls. Dioicous; perichaetial bracts lanceolate, not strongly differentiated in shape but with the costa very broad at the base, to 3.2 mm long; perigonial bracts shortly to longly acuminate, erect-spreading. Capsules 90°-180° from vertical, broadly pyriform, 2-3.5 mm long; exothecial cells shortly rectangular, 25-65 ym long, with sinuose ee stomata superficial; annuli of 2 9-3 cell rows, separating irregularly; opercula conic, blunt to apiculate; exostome yellow to brown, acute- triangular, most coarsely papillose at the apex, bordered, trabeculate; endo- stome hyaline; basal membrane '2 the exostome length; segments keeled and broadly perforate; cilia long, nodulose. Spores 14-21 ym, finely roughened. Fig. 18 Rare; on soil, especially in arctic or alpine tundra, often in lush snowmelt areas. North America (Alaska, British Columbia, Washington, Montana); north- ern and central Europe; Japan. Map 15. Exsiccati: Bauer, Musci Eur. Am. Exs. 2027 as W. ludwigit m ANM), Musci Eur. Exs. 1624 as P ludwigit Coens NY), 1625 as P. ludwigi f. ea (ALTA, NY), 1626 as P. ludwigit f. fragilis (ALTA, NY), 1627a as P. ludwigit f. aquatica (ALTA, NY), 162 7b as P. ludwigi f. erecta (NY); Husnot, Musci cai 520 as W. ludwigit v. major (BH); een Bryoth. Polon. 286 as P. ludwigit (CANM). Pohha ludwigit is uncommon in North America. It is characteristic of moist, rich snowmelt areas in tundra sites, where it is usually sterile but forms rather deep cushions. Schofield and Tan collected beautiful fruiting plants in British Columbia (74439, UBC). According to Schofield (in litt., 1980), capsules are produced late in the season, thus rarely collected. The species is recognized when sterile by the pink color of the lower leaves, and broadly lanceolate, longly decurrent leaves with the apices more or less cucullate (when moist). Plants collected by Schnooberger (/382) and Moxley (1936, 1941) from Bruce Co., Ontario and distributed as P. ludwigit are Bryum gemmiparum De Not. Gollecuons made by Bartram in Arizona and determined by Andrews as P. commitata (ez, drummondii) OL. ludwigit are also B. ere ne type of Bryum bigelovu Sull. is, by default, an odd expression of P. ludwigii. Sporophytically it is more or less typical, but the leaves are not very decurrent and have rather lax, thin-walled cells. It represents the only record of P. ludwigii in California (although it is not surprising). The type of Brywm ludwigii ssp. grande Kindb. is a sterile Philonotis. A specimen of P. ludwigit in Schwaegri- chen’s herbarium closely matches his illustration, providing one of few instances where typification is unambiguous! 15. yen reer (Ren. 7m Ren. & Card.) Broth.in Engler & Prantl, Nat. Pfl. 1(3): 5 903. oe aes Ren. im Ren. & Card., Rev. Bryol. 15: 71. 1888. (Type not seen). Bryum commutatum ssp. cardotu (Ren. in Ren. & Card.) Kindb., Eur. N. Am. Bryin. 2: 388. 1897. Pohlia porosa H. Lindb., Bull. Torr. Bot. Club 27: 318. 21. 1900. (Lectotype: H; Isotypes: COLO, DUKE, MICH, NY, UBC). Bryum cardotu (Ren. in Ren. & Card.) Kindb. ex Par., Ind. Bryol. Suppl. 1900 (nom. inval. in synon.—W. cardotii). Webera porosa (H. Lindb.) Par., Ind., Bryol. ed. 2, 5: 120. 1906. Medium-sized to robust, dull green plants often forming deep turfs. Stems 3—10(—25) mm high, simple or forking when sterile, simple or rarely innovating below perichaetia, simple when bearing perigonia. Leaves erect to erect- eT —— = tal ub ii “i fy FIG. 18. Pohlia ludwigi. A. Upper cells of branch leaf, x290. B. Upper cells of perichaetial A ‘Ee 9 5 3 bract, x 290. C. Exothecial cells, x 290. D. Peristome, X80. E. Annulus, X290. F. Capsules, x 15. ‘Transverse section of perichaetial bract near the base, 290. H. Leaves, x22. spreading, crowded, lanceolate, 0.7—1.5 mm long, serrulate to entire at the apex; costa strong, broad; upper cells rhombic, 18-35(—55) um long, 6-9 um wide, with thickened, often porose walls. Dioicous; perichaetial bracts somewhat differenti- ated, lanceolate to triangular, to 2.6 mm long; perigonial bracts shortly acute, erect. Capsules erect or nearly so, symmetric, narrowly pyriform, 2.0-3.0 mm long; exothecial cells shortly rectangular, 35-55 pm long, with + strongly sinuose walls; stomates superficial; annuli of 2—4 cell rows, separating irregular- ly; opercula conic, acute; exostome teeth yellow to brown, + narrowly triangu- lar, obtuse or + acute, most coarsely papillose near the apices, bordered, eae nd CH IOSOT SOS oS ecoe 1 O. Sg eo SOs KORY if OS (al¢) 2009569 fa) FIG. 19. Pohlia cardotii. A. Leaves, X22. B. Upper leaf cells, x 290. C. Exothecial cells, x 290. D. Stomates, x290. E. Habit, x3. F. Peristome, x80. G. Capsules, x15. H. Annulus, 290. trabeculate; endostome hyaline; basal membrane very low, less than 13 the exo- stome length; segments narrow but somewhat keeled and broadly perforate; cilia rudimentary to lacking. Spores 14-19 ym, finely roughened. Fig. 19. Uncommon; on soil in alpine or subalpine sites. Apparently endemic to western North America; Alaska, British Columbia, Washington and California. Map 16. | Exsiccati: Allen, Mosses of the Cascade Mts. 56 as P. porosa (isotypes, see above for representative herbaria). A relatively rare alpine species, P. cardotii is easily recognized by the erect capsules, very low basal membrane, and especially when sterile, strongly re- curved leaf margins and rather thick-walled cells. Pohlia crudoides is the only other American species with strongly recurved margins (except that the perichaetial bracts of most are recurved). When sterile, P. crudoides is distin- guished by the red stems, narrower leaves and longer, thinner-walled cells. Except for one Alaskan specimen of P. cardoti their ranges do not overlap. 260 A type for P. cardotit could not be located in Paris. However, the distinctive nature of this species leaves little question of the application of the name. In the future, if an isotype is not found, a neotype should be selected. 16. Pohlia erecta Lindb., Bot. Not. 1882: 195. 1889. (Lectotype: H-SOL; Iso- types: EO. WIL), Mielichhoferia defecta Sanio, Bot. Centralbl. 13: 248. 1883. (Lectotype: H- Webera defecta Sanio, Bot. Centralbl. 13: 248. 1883 (nom. nud. in synon. defecta). Mielichhoferia erecta (Lindb.) Kindb., Forh. Vid. Selsk. Christiana 1888(6): 1888. M. oo. Webera erecta (Lindb.) Limpr., Laubm. Deutschl. 2: 239. 1891. Pohlia defecta (Sanio) Andr. in Grout, Moss Fl. N. Am. 2: 189. 71. 1935. Slender, rather glossy plants. Stems 3-8 mm, generally simple when sterile or bearing perichaetia, with I-2 innovations when bearing perigonia. Leaves erect-spreading to erect, more or less crowded, lanceolate; 0.7—1.2 mm long, serrate to serrulate near the apex; upper cells rhomboidal, 35-60 ym long, 6— 9 um wide, thin-walled. Dioicous: perichaetial bracts slightly differentiated, to ¢.2 mm long; perigonial bracts shortly acuminate, erect. Capsules erect or nearly so, symmetric, slenderly pyriform, 2.0-3.0 mm long; exothecial cells shortly rectangular, 25-60 pm long, with sinuose walls; stomates superficial; annuli of I(2) cell row(s), separating irregularly; exostome teeth yellow to light brown, narrowly triangular, somewhat irregular, obtuse, papillose (but not coarsely so) near the apices, bordered, + trabeculate; endostome hyaline; basal membrane very low, less than 3 the exostome length; segments rudimentary and irregular or + elongate and perforate; cilia absent. Spores 15-21 um, finely roughened. Hise 20; Very rare, on soil in the alpine zone. Western North America (Yukon Ter- ritory, British Columbia and Washington) and Fennoscandia. Ma The erect capsules resemble those of P. cardotii, but the endostome of P: erecta 1s even more reduced. Sanio (1883) described this species as a Mielichho- feria. Although P. erecta seems at home in Pohlia, the distinctions between plants of the so-called Mielichhoferioideae (Mielichhoferia, Synthetodontium, Haplo- dontium) and Pohlia are sometimes ill-defined. Pohha erecta differs from P. cardotii in having soft-textured leaves with plane margins, thinner-walled cells and narrower costae. When sterile, P. erecta resembles a small version of P. drummondu, especially in having reddish stems. Further similarities between P. erecta and P. drummondii include the aspect and branching of male plants (Fig. 20) and strongly sinuose exothecial cells. These combined probably indicate close phylogenetic affinity. 17. Pohlia beringiensis Shaw, sp. nov. Plantae erectae, graciles, plerumque steriles. Caulis simplex, 4-15 mm altus. Folia pallida, erecta, imbricata, lanceolata, ad apicem serrulata, cellulis lineari- dus. Gemmae frequenter adsunt, oblongae, cerasinae. Slender, pale-whitish plants diffusely scattered in small populations. Stems +—15 mm, simple. Leaves erect, imbricate (moist or dry), 0.9-1.3 mm long, 0.3- 0.6 mm wide, narrowly lanceolate, gradually acute, slightly decurrent; margins plane, serrulate near the apex, entire below; costa ending 3-5 cells below the apex; upper cells linear-rhomboidal, + vermicular, thin-walled, 60-90(-100) um long, 6-10 gm wide, somewhat longer and narrower along the margin. Gemmae single in Id upper leaf axils, oblong-bulbiform, about 500—-650(—1000) um long, deep cherry-red when moist, red or black when dried, with 3-6 large, broadly 261 \ Vy ) i ae. tEeieveses LE ~¢, BOSS ree CoO Obes; YI CO? e, BOGO IO CSS BOGS erooos 0290800 FIG. 20. Pohlia erecta. A. Habit, female plant, x5. B. Exothecial cells, x 290. C. Upper leaf cells, x 290. D. Stomate, x 290. E. Annuli, x 290. F. Habit, male plant showing subfloral innovations, 5. G. Leaves, x22. H. Peristome, x 250. laminate leaf primordia at the apex and frequently lower, the primordia green. Inflorescences and sporophytes unknown. Fig. 21. TYPE: Alaska, Paxson area, 11.7 km W of Paxson on Hwy. 8., 22 July 1979, Shaw 3794 (Holotype: MICH). Paratypes: Alaska, Honolulu area, 57.5 km S on Hwy. 3 from junction with Hwy. 8 at Cantwell, 24 July 1979, Shaw 3889 (ALTA). Yukon Territory, Ogilvie Mts., Tombstone Mt. area, 71.3 km N of Hwy. 3 on the Dempster Hwy., 16 July 1979, Shaw 3466 (MICH). n organic-poor soil in mountainous areas, especially in roadside ditches. Central Asia (Altai Mts.) and northwestern North America (Alaska and Yukon Territory). Map 18. Ne YA ae iy) ) (] ay ZINN DN fy enn VAR mi iM Mi WN} HY FIG. 21. Pohha beringiensis. A. Upper leaf cells, x290. B. Habit, x 10. C. Gemmae, x 100. D. Leaves, X22. Pohlia beringiensis was mentioned previously (Shaw 1981, p. 33) as a variant possibly related to P. drummondii. Continued study has convinced me that it should be segregated as a new species. Compared to P. drummondii, plants of P. beringiensis are very small, with leaves whitish, erect and slenderly lanceolate. The gemmae are of a similar form to those of P. drummondii, but are even more deeply cherry red (at least when fresh). Red gemmae on plants with such pale, whitish leaves gives P. beringiensis an aspect different from any expression of P. drummondi. The micro-habitat in which P. beringiensis occurs also differs from that of P. drummondi. In western North America, P. drummondii grows on relatively organic-rich soil banks or frequently in alpine tundra (Shaw 1982), whereas P. beringiensis prefers sandy or muddy, organic-poor sites such as roadside drainage ditches. Even allowing for some variation in the microhabitats occupied by P. drummondu, there does seem to be ecological differences between the two. Especially since sporophytes of P. beringiensis are not known, differences between it and P. drwmmondii are mainly with respect to aspect, color, size and habitat of plants. The gemmae, in terms of size and development, are similar to those of P. drummondu, differing only in color. Varietal status for P. beringiensis (under P. drummondi) could be defended, and there is no correct answer to the problem of appropriate rank. Other pairs of Pohlia species are distinguished by such ill-defined features as habit and color. For example, few collectors confuse P. melanodon and P. wahlenbergu, although they differ only in size and sheen. The distribution of P. beringiensis, seemingly disjunct from central Asia (Altai Mts.) to Alaska and the Yukon Territory, falls more or less within the circumboreal range of P. drummondii. In northwestern North America, P. drummondii is far 263 more common than P. beringiensis. In addition to my own collections of P. beringiensis (seven) it has otherwise been collected only twice: Alaska, Liberator Lake, Steere 610723-—39 (NY), and U.S.S.R. (or China), Altai Mts., Barzduna, 1966 (NICH). POHLIA ANNOTINA AND RELATED SPECIES Species of this group, characterized by axillary gemmae, were revised by Shaw (198 la). Brief descriptions, and illustrations of gemmae are provided here but the reader is referred to Shaw (198la & b) for a more detailed treatment. All nine species are virtually uniform sporophytically, characterized especi- ally by rectangular, sinuose exothecial cells and well-developed peristomes with endostome segments hyaline, broadly keeled and perforate. 18. Pohlia drummondii (C. Mull.) Andr. 7» Grout, Moss Fl. N. Am. 2. 196. 1935. Medium-sized, glossy plants with red stems (when fresh). Gemmae arising singly in 1—-3(-5) upper leaf axils (sometimes lacking in alpine plants), oblong- bulbiform, 350-600 zm long, green when young, red when older, with 3-6 lanceolate-laminate leaf primordia scattered at the apex and below, the pri- mordia generally green. Fig. 22A. Roadbanks, streambanks and gullies in montane areas, frequent on non calcareous tundra soil. Circumboreal, in North America south to California, Colorado and New England. Map 19 19. Pohlia filum (Schimp.) Mart., Svensk Vet.-Ak. Arh. Natur. 14: 149. 1956. Slender, rather glossy plants frequently black below when dry. Leaves erect, imbricate to remote. Gemmae arising singly in I—3(—5) upper leaf axils, ege- shaped, 300-600 um long, green when young, orange to orange-brown when older, frequently becoming black when dry, with 3-6 rather stiff, triangular- ae leaf primordia at the apex and sometimes lower, the primordia green to pale. Fig. 22B Common on glacial outwash in northwestern North America, rarely along roadsides, restricted to nearly organic-free soils. Circumboreal; in North Amer- ica south to Oregon, Alberta, Quebec and Prince Edward Island. Map 20. Pohlia filum is distinguished from P. drummondu by its slender habit, erect leaves, and egg-shaped gemmae with stiffly appearing primordia often re- stricted to the gemmae apex. 20. Pohlia andalusica (Hohn.) Broth. in Engler and Prantl, Nat. Pfl. 1(3): 551. 1903. Slender, glossy plants with erect, + imbricate leaves. Gemmae in clusters of 3-8 in 1-4 upper leaf axils, obconic to oblong, 250-500 um long, green when young, brown to red-brown when older, with 3-6 lanceolate-laminate leaf pri- Benes more or less restricted to the apex, the primordia green. Fig. 22C. 1 soil in disturbed sites; rare in western North America, from Alaska and the aie south to Washington and Colorado; more eaueU in northern New York, New England and the Maritime provinces. Map 21. Pohlia andalusica is distinguished from related species by its slender, glossy appearance and gemmae with laminate primordia arising in clusters in a few upper leaf axils. IN \ i a Ae WW a Vy) i) aS « IN A AUN LAAN h eK) Max?) Pe) \) LUNN FIG. 22. Gemmae, 150. A. Pohlia drummondii. B. Pohlia filum, C. Pohlia andalusica. 21. Pohlia tundrae Shaw, The Bryologist 84: 65. 1981. Medium-sized, glossy plants commonly forming compact turfs on tundra soil, more lax and elongate in protected sites. Gemmae in clusters of 3-many in the upper leaf axils, cylindrical to linear-oblong, 275-500 pm long, hyaline, green or red-brown, with 3-6 flexuose, lanceolate-laminate leaf primordia at the apex and frequently lower on the body, the primordia green. Fig. 23A. Common on alpine and subalpine soil, less frequent along roadbanks and in other disturbed sites below treeline. Endemic to western North America: Wash- 265 KY eS Maan Alinabas ss ESP ngeas ie FIG. 23. Gemmae, x 150. A. Pohlia tundrae. B. Pohlia bulbifera. C. Pohlia andrewsit. ington and Montana south to California and Arizona; northern British Columbia and southern Alaska. Map 21. The narrowly cylindrical gemmae distinguish P. tundrae from other species with laminate leaf primordia. It differs from P. annotina in having glossy leaves in addition to the gemmae characters given in the key. 22. Pohlia bulbifera (Warnst.) Warnst., Krypt. Fl. Brandenburg 2: 429. 1904. Slender, glossy plants forming compact turfs. Leaves wide-spreading. Gemmiae in clusters of 3-8 in upper leaf axils, spherical to obovate, 200-400 um in the longest dimension, green to yellow, yellow-orange, or orange-red, with 4—5 broadly triangular-laminate leaf primordia in a tight spiral (almost whorled) 266 at ne eh, the primordia green, incurved over the apex and forming a dome. Hig. 2 oe Soll in disturbed places, often where the ground is compacted and hard. Circumboreal; rare in western North America (Alaska, Yukon ‘Territory, Alberta and Colorado), more common in northern New oe New England, and the Maritimes, west to Michigan and Wisconsin. Map 23 Pohlia bulbifera is known at once by glossy leaves and more or less spheroidal gemmae with the laminate leaf primordia forming a dome over the apex. 23. Pohlia andrewsii Shaw, The Bryologist 84: 70. 1981. Medium size to robust shiny plants forming small populations in rock crev- ices. Gemmae in clusters of 4-many in the upper leaf axils, oblong to linear- oblong or spheroidal, 50-170 wm long, pale green when young, red to orange- red when older, strongly translucent, with I-4(-6) short, incurved, peglike leaf primordia at the apex or rarely lower, the primordia of some gemmae becoming laminate with age. Fig. 23C. Soil in rock crevices in tundra habitats, less commonly along roadbanks. Arctic and subarctic North America, Greenland, Spitzbergen, Norway, Sweden and Czechoslovakia. Map 22. Pohla andrews may be recognized by glossy leaves and orange to red, trans- lucent gemmae with the primordia, whether peglike or laminate, incurved over the apex. 24. Pohlia proligera (Lindb. ex Breidl.) Lindb. ex Arn., Bot. Not. 1894: 54. 1894. Medium-sized to robust, glossy plants forming small to extensive popula- tions. Gemmae in dense clusters in upper leaf axils, linear-vermicular to nar- rowly oblong, 65-110 zm long, green when young, yellow to golden, orange, pink or sometimes red when older, strongly translucent, with I-2 peglike apical leaf primordia, the primordia (when single) often bent at an angle to the propagulum body. Fig. 24/ On soil, especially along the sides of roads, paths and streams; sometimes on disturbed tundra soil in the far North but rarely reaching the alpine zone in more southerly mountains; rarely on rotting wood. Circumboreal: our most common propaguliferous species across northern North America, south to California, C olorado, Wisconsin, Michigan and southern New England. Map 25. Pohlia proligera is well characterized by glossy leaves and linear-vermicular gemmae forming a felty mass in upper leaf axils. The gemmae are uniform in shape and have ‘only one or two apical primordia. The type specimens of P. proligera are located in GJO-—Breidler rather than GZU as previously stated (Shaw 1981b, p. 21) 25. Pohlia annotina (Hedw.) Lindb., Musci Scand. 17. 1879. Dull, slender plants often forming + extensive turfs. Gemmae in clusters of 5-many in the upper leaf axils, extremely variable in shape, oblong, obconic or + elongate and linear, although almost always tapered to the base, rarely oblong-bulbiform when old, 150-300 ym long, occasionally longer, hyaline, yel- low, green, orange or red when young, generally orange-red when large and pnb oni strongly translucent, with (2—-)3-5 peglike, erect, apical leaf pri- mordia, the primordia occasionally becoming 2—4 cells broad at the base on old gemmae. Fig. 24B Common in disturbed habitats; sandy soil in gardens, on roadbanks and in gullies, rarely found in undisturbed sites. Possibly circumtemperate; specimens Ky KK FIG. 24. Gemmae, x 150. A. Pohlia proligera. B. Pohlia annotina. C. Pohlia camptotrachela. confirmed from northern and central Europe and North America. Bicentric in North America and having a suboceanic distribution: Newfoundland to Georgia and Arkansas, Wisconsin, Missourl and lowa; Alaska, British Columbia, Montana, Washington, Oregon and California. Map 26. Characterized by dull leaves and oblong to linear-obconic gemmae with 268 peglike leaf primordia, P. annotina can be confused only with P. camptotrachela. From P. proligera it is distinguished by dull leaves and gemmae having three to as many as five primordia. 26. Pohlia camptotrachela (Ren. & Card.) Broth. in Engler & Prantl, Nat. Pfl. [yep 2> LOS: Slender, dull plants forming small isolated populations. Gemmae in clusters of 5-many in upper leaf axils, uniform in shape, spheroidal or slightly longer than wide, 70-120(—150) ym in the longest dimension, yellow or occasionally brown, strongly translucent, with 1-4 short, peglike leaf primordia at the apex, the primordia incurved over the apex or less commonly erect. Fig. 24C. Not uncommon on soil of roadbanks, pathbanks and stream margins in montane areas along the Sierra Nevada—Cascade systems; rather poor alpine soil in the central Rockies. Western North America (British Columbia south to Cali- fornia, Nevada and Colorado); the British Isles, central and northern Europe. Map 27. Unlike P. annotina, the gemmae of P. camptotrachela are uniform in shape and size: roughly spheroidal and less than 150 um in diameter. Among the North American propaguliferous species, only P. annotina and P. camptotrachela have notably dull leaves. Virtually identical male plants of the two species give further indication of a close relationship. 27. Pohlia pacifica Shaw, sp. nov. Plantae erectae, graciles, valde nigrescentes. Caulis simplex, e basi innovans et ramulosus, erectus, 1.0-8.0 mm altus. Folia inferiora lanceolata, comalia elongato-lanceolata, ad apicem valde serrata, cellulis linearibus, angustls. Peristomia duplices, exostomii dentibus 16, luteo-bruneis. Annulus latus, duplex vel triplex, Slender, pale green, shiny plants usually in small populations. Stems 2-8 mm, simple when sterile or bearing gametangia. Leaves erect-spreading to spreading when moist, scarcely altered on drying, 1.1-1.8(-2) mm long, 0.5- Q.8 mm wide, lanceolate, acute to shortly acuminate, not or slightly keeled, not to slightly decurrent; margins plane, coarsely serrate near the apex, entire to subentire below; costa ending 3-5 cells below the apex; upper cells linear- rhomboidal, vermicular, thin-walled, (100-)115-135(-150) ym long, 6-9 um wide, somewhat longer and narrower along the margin (to ca. 180-200 pm). Dioicous: perigonia bulbiform; outer bracts + longly acuminate, spreading to erect-spreading from an erect base, serrate near the apex; perichaetia rather conspicuous, outer bracts to 3.2 mm long, narrowly and longly lanceolate. Sporophytes frequent in spring; setae 5-10 mm long; capsules inclined 120° 180° from vertical, 1.5-2.5 mm long, orange to tan, pyriform, the neck shorter and narrower than the urn; exothecial cells shortly rectangular, 25-55 um long, 12-25 um wide, with evenly thickened, strongly sinuose walls; stomata super- ficial, 35-55 pm long; annuli of 2(3) rows of cells, separating irregularly; opercula convex-conic, obtuse or rarely acute; exostome teeth vellow-brown, narrowly acute, bordered, trabeculate, coarsely papillose near the apex dorsally, more finely roughened below; endostome hyaline; segments keeled and broadly perforate; basal membrane about 4 the height of the exostome teeth; cilia short to long, usually nodulose. Spores 14-20 um, nearly smooth. Fig. 25, TYPE: U.S.A. California, Del Norte Co.: Del Norte Redwoods State Park, southern boundary, clay roadbank, 15 April 1980, Shaw 4173 (Holotype: MICH; Isotype: J. Shaw). Paratypes: Oregon, Tillamook Co.: 2.7 mi N of Arcada Beach on Hwy. 1OL, muddy roadbanks and drainage ditches, 14 April 1980, Shaw 4110 1S, DLW eS [> ASO y ex. y ‘s) 9 i aa SS FIG. 25. Pohha pacifica. A. Habit, x3. B. Stomate, x 230. C. Capsules, x 12. D. Peristome, x64. E. Annulus, 232. F. Exothecial cells, x232. G. Costae, transverse sections near the apex (left) and base (right), ¥ 232. H. Leaves, x17. I. Habit, female plant, x3. J. Habit, sterile plant x3. K. Upper leaf cells, «232. (CANM), 4127 (NY). Washington, Clallam Co.: Sol Duc Hot Springs, Lawton 4298 (WTU). Acidic, sandy or clay soil, especially in wet, disturbed places at sea level; drainage ditches, stream banks. Endemic to the Pacific Coast of North America; Queen Charotte Islands of British Columbia (Schofield 14172a, UBC), Washing- ton, Oregon, California (south to Santa Cruz County). Map 28 Pohlia pacifica generally grows in the same habitats with Epipterygiwm tozeri (Grev.) Lindb. and P. longibracteata. It is frequently mixed with the Epipterygium, 270 oS = AY \ 5a | ( a | IG. 26. Polygraph comparing poulations of Pohlia lescuriana and Pohlia pacifica. A. Perichaetial leaf length. B. Perichaetial leaf width. C. Branch leaf cell length. D. Branch leaf length. E. Branch leaf width. F. Perichaetial leaf cell length. Lines connect values for different measurements from one specimen. whereas P. longibracteata prefers wetter sites (sometimes a few feet away) that are periodically submerged. Pohlia longibracteata is much larger (8-15 mm), lacks an annulus and has immersed stomates. Epipterygium tozeri is the same size as P. pacifica but has dull, bluish leaves, with broad cells (14-25 yum wide), and strongly collenchymatous exothecial cells. Pohlia pacifica is closely related to P. lescuriana but is distinguished by the more robust habit, very glossy leaves and longer cells. Figure 26 compares popu- lations of the two species with respect to six morphological characters; P. pacifica is more robust in all respects. Although many populations of P. lescuriana have rhizoidal gemmae, | have seen one or two apparently attached to P. pacifica only once, as pointed out to me by A. C. Crundwell ina specimen sent for his opinion. ‘Tan, Shaw and Ireland (1981) reported P. lescuriana from western North America, but several of the specimens have proved to be misidentified. I can confirm P. lescuriana in the west on the basis of only two collections from Alaska: Curry, Schofield 1892 (BH); Kenai Peninsula, Shaw 4016 (MICH). 28. Pohlia lescuriana (Sull.) Grout, Moss. Handlens Microsp. 210. 1906. Bryum pulchellum Hedw., Spec. Musc. 180. 1801. Hypnum pulchellum (Hedw.) Web. & Mohr, Ind. Musc. Crypt. 3. 1803. Bryum carneum var. pulchellum (Hedw.) Turn., Musc. Hib. 119. 1804. Mnium pulchellum (Hedw.) P. Beauv., Prodr. 76. 1805. Bryum tenellum Brid., Bryol. Univ. 1: 653. 1826 (nom. nud. in synon.—B, pulchellum). Webera pulchella (Hedw.) Fuernr., Flora 12(Ergbl. 2): 35. 1829. W. carnea var. pulchella (Hedw.) Hampe, Flora 20: 277. 1837. Bryum lescurianum Sull., Mem. Am. Ac. Arts. Sci. n. ser. 4: 171, 1849. Webera lescuriana (Sull.) Jaeg., Ber. S. Gall. Naturw. Ges. 1873-74: 1875. Pohha pulchella (Hedw.) Lindb., Musci Scand. 17. 1879. 271 Webera iwozanica Besch., Ann. Sci. Nat. Bot. ser. 7. 17: 342. 1893. Mniobryum pulchellum (Hedw.) Loeske, Stud. Morph. Syst. Laubm. 124. 910. Tiny, dull or somewhat glossy plants. Stems 2—8 mm, simple when sterile or bearing gametangia. Leaves erect-spreading on sterile plants, few, erect, and overlapping on fertile plants, 0.8—1.4 mm long, serrulate near the apices; upper cells linear-rhomboidal, 65-110 um long, 6-9 um wide, with thin walls. Tubers sometimes present on rhizoids, orange, orange-brown or yellow, 70-120 um long. Dioicous; Bovidueteue bracts linear-lanceolate to lanceolate, to 2.5 mm long; perigonial bracts + longly acuminate, erect to erect-spreading. Capsules inclined 95°—180° from vertical, shortly and broadly pyriform, 1.5—3.0 mm long; exothecial cells short-rectangular, 12-40 um long, with strongly sinuose walls; stomates superficial or slightly sunken; annuli of 1-3 cell rows, separating irregularly; opercula low to high conic; exostome teeth light yellow-brown, acute-triangular, most coarsely papillose near the apices, bordered, trabeculate; endostome hyaline; basal membrane 4 the exostome length; segments keeled, broadly pertorale, ae long, nodulose, rarely short. Spores 18-26 um, distinctly roughened. Fig. 2 Uncommon; on esi: often in weedy places, roadbanks, fields, stream banks, near sea level. Fennoscandia; British Isles; Central Europe; Japan; North America; Maritime provinces and New England south to Virginia and Tennes- see, west to Michigan and Iowa; disjunct in Alaska, where it is known from the Kenai Peninsula and near Curry on the southern edge of the Alaska Range. Map 28. Exsiccati: Austin, Musci Appal. 190 as B. lescurianum (CANM, CAS, MICH, UBC); Brotherus, Bryoth. Fenn. 45 as P. eae (BH); Grout, N. Am. Musci Perf. as P. lescuriana (CANM, CAS COLO, MICH, NY, UBC, UC); Hartman, Bryac Seana: Exs. 404 as W. pulchella (BH); Holzinger, Musci Acro. Bor.—Amer. Sie is W. lescuriana (GAND {, MICH, UC; =P. wahlenbergu in COLO), Husnot, Musci Gall. 572 as W. pulchella (BH); Macoun, Canad. Musci 163 as W. lescuriana (CANM: =P. annotina in UC); Renauld and Cardot, Musci Am. Sept. Exs. 222 as W. lescurtana (BH, CANM); Sull., Musci Allegh. 101 as B. eg en (MICH); Sull. & Lesq., Musci Bor. Am. (ed. 1) 181 as B. leeanianum (BH, MICH); (ed. 2) 2 s B. lescurianum (BH, CANM, MICH). One of the tiniest species, P. lescuriana looks much like a miniature P. annotina without gemmae. It is characterized by the slight stature, well- developed peristome, sinuose exothecial cells, separating annuli and super ficial (or slightly sunken) stomates. I find that North American plants sometimes, but not always, have gemmae on their rhizoids. These range from scarce to abun- dant and are orange to yellow, round to ovoid, 70-120 um in diameter, and without bulging superficial cells. The European P. lutescens (Limpr.) H. Lindb. is ee from P. lescuriana by its stellate rhizoidal gemmae (Whitehouse 1973, 1978), although very similar in most other respects. Ecological and phytogeographic information needs to be accumulated with which to evaluate P. lutescens as a biological entity. 29. Pohlia wahlenbergii (Web. & Mohr) Andr. 7x Grout, Moss FI. N. Am. 2: 203. 1935 Hypnum albicans [Wahlenb. ex] Web. & Mohr, Ind. Muse. Pl. Crypt. 3. 1803. (nom. d.). cae wahlenbergu Web. & Mohr, Bot. Taschenb. 280, 475. 1807 (type not seen). Mnium albicans Wahlenb., Fl. Lapp. 353. 1812 (nom. illeg. incl. spec. prior.—H. wahlenbergir). Bryum wahlenbergit (Web. & Mohr) Schwaegr., Spec. Musc. Suppl., 1(2): 92. 70. 1916. FIG. 27. Pohlia lescuriana. A. U pper leaf cells, x290. B. Capsules, 15. C. Leaves, x22. D. Peristome, 80. E. Stomate, x 290. F. Rhizoidal tubers, x270. G. Exothecial cells, 290. H. Annulus, «x 290). B. nitens Sw. ex Schwaeer., Spec. Muse. Suppl. 1(2): 92. 1816 (nom. nud. in synon.—B. wahlenbergi). B. glaciale Brid., Bryol. Univ. 1: 852. 1826. (Lectotype: B-Brid.). Webera wahlenbergu (Web. & Mohr) Fuernr., Flora L2(Erepl. 2): 35. 1829. Bryum wahlenbergii var. glaciale (Brid.) Hueb., Musc. Germ. 456. 1833. Webera wahlenbergii var. glacialis (Brid.) Hampe, Flora 20: 277. 1837. Webera albicans Schimp., Coroll. 67. 1856 (nom. illeg. incl. spec. prior.—H. wahlenbergi). W. albicans var. glacialis (Brid.) Schimp., Coroll. 67. 1856. Zu Lamprophyllum albicans Lindb., Oefv. K. Vet. Ak. Foerh. 23: 75. 1886 (nom. illeg. incl. spec. prior.—H. wahlenbergi). Pohtia albicans Lindb., Musci Scand. 17. 1879 (nom. illeg. incl. spec. prior.—H., } (ee ahle nbe jer ee var. glacialis (Brid.) Broth. & Sael., Act. Soc. F. Fl. Fenn. 6(4): 59. 1890. Mniobryum albicans nee Laubm. Deutschl. 2: 277. 1892 (nom. illeg. incl. spec. prior.—H. wahlenbergi) Pohha obtusata rales Ree Bryol. 32: 37. 1905. (Lectotype: S—Kindb.). Mniobryum wahlenbergi (Web. & Mohr) Jenn., Man. Moss. W. Penn. 146. 18. 3 Mniobryum wahlenbergu var. glaciale (Brid.) Wijk & Marg., Taxon 8: 74. 1959. Very robust to slender, dull, whitish plants forming small populations or covering | arge areas. Stems 4—15(-50) mm, simple when slender and sterile, forking when robust and fertile, with 1-3 innovations below perichaetia or perigonia; leaves erect-spreading, crowded to remote, 0.9-1.8 mm long, toothed to serrulate in the upper 3, acute or sometimes + rounded at the apex, + longly decurrent; upper cells broadly oblong-hexagonal, 65-110 um long, 12-20(-30) um wide, gradually longer and narrower near the margin, with thin to firm walls. Dioicous; perichaetial bracts scarcely differentiated, to 2.0 mm long; perigonial bracts shortly acuminate, wide- spreading. Capsules inclined 140°-240° from. vertical, shortly and_ broadly pyriform or urceolate, 1|.5— 2.5 mm long; exothecial cells + isodiametric, 22-55 um, with strongly sinuose walls; stomates + deeply immersed; annuli lacking; opercula low-convex to conic, apiculate; exostome teeth light brown to red-brown, acute-tr langular, most coarsely papillose near the apices, bordered, trabeculate; endostome hyaline to yellow; basal membrane slightly more than 4 the exostome length; segments keeled, broadly perforate; cilia long, nodulose. Spores 13-18(- 24) um, very finely roughened. Fig. 28. On calcareous rocks or soil in wet seepy areas, ditches, stream margins, springs, glacial outwash; sea level to the alpine zone. Widespread in North America (across Canada and south to California, Arizona, Arkansas and North Carolina), Europe, and northern Asia; Iceland; Greenland; Mexico (Baja Cali- fornia, Distrito Federal, Michoacan, Oaxaca, Puebla, Sonora); Guatemala (Quezaltenango, San Marcos); Dominican Republic (Shaw 5656); Chile; Argentina; Australia (also reported from New Zealand); Japan. Map 29. Exsiccati: Allen, Moss. Cascade Mts. 53 as P. albicans (DUKE, MICH, NFLD); Bauer, Musci Eur. Exs. 915 as M. albicans (WTU); Drummond, Musci Amer. 260 as B. carneum (C oe MICH, NY); Grout, N. Am. Musci Perf. 260 as M. carneum (UC); Holmen, Bryoph. Dan. Exs. 151] as W. ae (ALTA, NFLD, UC); Holzinger, Musci Acro. nage 168 as M. albicans (MICH, UC): 220 as W. lescuriana (COLO); 393 as M. albicans (MICH, UC); 437 as M. albicans (MICH, UC, WTU); ee Bryoth. Sax. 52 as M. albicans ae 1); 244 as M. albicans (MICH); 420 as P. commutata (MICH); Macoun, Canad. Musci 165 as W. albicans (MICH); 208 as W. carnea ae figula, Krypt. Germ., Aust. el a = 228 as M. than (MICH); Rabenhorst, Bryoth. Eur. 1215 as W. ie eae! Ren. & Card., Musci Am. Sept. Exs. 303 as W. albicans (MICH); ee Ljubitzkaja, . Musci URSS Exs. 126 as i wahlenbergr (WT ti): 127 as M. seeds Vv. Lidia (WTU); Sc eal Bryoph. Can. 9 as P. wahlenbergii (ALTA, CANM, NFLD)); Sull. & Lesq. Musci Bor.-Am. (ed. 1) 100 as B. wahlenbergii (MICH); 182 as B. wahlenbergu (MICH); ed. 2, oe as B. eee (MICH); Warnst. Mark. Laubm. 172 as W. albicans (CAS). Distinguished by rather dull, whitish-green leaves, P. wahlenbergiu is one of the most common Pohlias of the northern United States and Canada. It is most luxuriant in seepy calcareous areas at high altitudes and latitudes, where the stems may attain a height of several centimeters and the leaves are darker green on deep red stems. In ecologically or geographically marginal sites, P. wahlen- WrENY, { D 7 ho d j | So oS. COCCaBP2? Sosne. F0056890 20899700 9C 0000 pose ODDO0O DOO Genco 7OL00ges OS OM OnPOCO SEL OOS IG. 28. Pohlia wahlenbergi. A. Capsules, x 15. B. Exothecial cells, x290. C. Stomate, x 290. D. Peristome, X80. E. Habit, male plant, x6. F, Leaves, x22. G. Upper leaf cells, «290. ‘ee — i 2 ae tS = Sa. - bergu may be very slender, with scarcely any superficial! resemblance to well- developed forms. The lax cells are helpful in identifying such plants. Pohlia wahlenbergii, like the following species, produces sporophytes in the spring, and they generally disappear by mid summer. In some areas at least, capsules are abundant in suitably moist sites. All Mexican and Central American collections are sterile. Small forms may be confused with P. melanodon, which is always small but has somewhat glossy, yellow-green leaves that are more acuminate than those of P. wahlenbergu. The European P. wahlenbergii var. calcarea (Warnst.) Warb. is the size of typically well-developed P. wahlenbergii but has the gloss and yellowish ras leaves of P. melanodon. The differences between the var. calcarea and typical P. wahlenbergi are impressive and warrant further study. Plants collected by A. Radlow in Baja California (50 mi SW of Mexicali, 8504, 8505, TENN) are less than one centimeter high, green to reddish-brown, somewhat glossy, and have rhomboidal cells (Fig. 29). They are apparently dioicous, but perigonia or sporophytes are lacking. Although in all liklihood distinct, they are included under P. wahlenbergi pending further collections. Pohha wahlenbergu has variably shaped cells but they are almost always broader. Among the Mniobryums, P. longibracteata and P. columbica have cells as narrow as those of the Baja California plants, although in the absence of sporophytes the affinities of the latter is a moot question. The habitat on clay soil along a stream 1s suggestive of the Mniobryum group. FIG. 29. Plants collected by A. Radlow in Baja California A. Habit, 30, B. Uppper leaf cells, x 100. C. Costae, transverse sections, X 100. D. Leaves, 35. 276 30. Pohlia melanodon (Brid.) Shaw, Uhe Bryologist 84: 506. 1982. Bryum delicatulum Hedw., epecs Muse. 179. 1801 (type not seen). Bryum carnewm [L. ex] With., Syst. Arr. Brit. Pl ed. 4, 3: 820. 1801 (nom. illeg. incl. spec. prior.—B. delicatulum). Hypnum carneum Web. & Mohr, Ind. Muse. PI. Crypt. 3. 1803 (nom. leg. incl. spec. prior.—B. delicatulum). Mnium carneum P. Beauy., Prodr. 74. 1805 (nom. illeg. incl. spec. prior.—B. delicatulum). Bryum melanodon Brid., Bryol. Univ. 1: 845. 1827. (Lectotype: B-Brid.). Webera carnea Schimp., Coroll. Bryol. Eur. 67. 1855. Pohlia carnea (Schimp.) Lindb., Musci Scand. 17. 1879. Webera delicatula (Hedw.) C. Jens., Skand. Bladmfl. 154. 1939, Pohlia delicatula (Hedw.) Grout, Moss Fl. N. Am. 2: 266. 1940 (hom. illeg.—P. delicatula [Mitt.] Broth.). Slender, glossy, vellow-green_ plants forming small populations. Stems 2— 6 mm, simple when sterile or fertile or rarely with an innovation below peri- gonia; leaves spreading to erect-spreading, somewhat distant, lanceolate, acute or shortly acuminate, 0.7—1.3 mm long, serrulate near the apex; upper cell laxly and broadly rhomboidal, 60-100 um long, 10-22 wm wide, somewhat narrower near the margin, with thin walls. Dioicous; perichaetial bracts not strongly dif- ferentiated, to 1.9mm long; perigonial bracts short-acuminate, spreading. Capsules inclined + 180° from vertical, shortly and broadly pyriform, some- times urceolate when dry and empty, 1.0-3.0 mm long; exothecial cells iso- diametric, + collenchymatous, 20-15 pm long, with strongly sinuose walls: stomates + deeply immersed; annuli absent; opercula conic, blunt to acute; exostome teeth dark brown or red-brown, acute-triangular, most coarsely papillose near the apices, bordered, trabeculate; endostome yellow to yellow- brown; basal membrane slightly more than '% the exostome length; segments keeled, broadly perforate; cilia long, nodulose. Spores 15-22 um, finely roughened. Fig. 3 On. calcareous clay soil in disturbed, often wet places (ditches, gullies, streambanks, field margins); mostly near sea level. Eastern North America; Ontario and Nova Scotia, New York, New Jersey, Tennessee, Michigan, Ohio and lowa; Europe; reported from northern Asia; Japan. Sterile specimens from Illinois, Arkansas and Maine probably belong to P. melanodon. Map 30. Exsiccati: Bauer, Musci Eur. Exs. 326 as M. carneum (ALTA, BH), 327 as M. carneum (ALTA, BH), 1103 as M. carneum (ALVA); Erbar, Crittogam Ital. 816 as W. carnea (UC); Gyrorffy, Flora Hung. Exs. 929 as M. carneum (ALTA, BH, MICH, UC); Harman, Bryac. Scand. Exs. 15 as W. carnea (BH) Holmen, Bryoph. Dan. Exs. 152 as W. delicatula (ALVA, CAS, UC); Husnot, Musci Gall. 234 as W. carnea (BH); Ren. & Card., Musci Fur. Exs. 283 as W. carnea (BH); Warnstorf, Mark. Laubm. 173 as W. carnea (CAS). , ~~ The delicate little plants of P. melanodon are best recognized by their glossy yellowish leaves and small capsules with dark-brown exostome teeth, yellow endostomes and deeply sunken stomates. Other than in size and sheen, there is little to distinguish this species from P. wahlenbergii, yet the two are rarely con- fused. Pohlia melanodon is similar to P. atropurpurea and P. brevinervis, but the three differ in leaf areolation. In P. atropurpurea, the cells of leaves on sterile plants are broad and lax (11-20 um wide), whereas those of perichaeual bracts are narrow and hexagonal (7-I1l um wide). Pohlia brevinervis has narrow, linear cells throughout and P. melanodon has lax cells throughout. These differences, though not absolute, help distinguish collections lacking sporophytes (Fig. 32). several Mniobryum species have exothecial cells just below the rim lax, thin- S 0, 0.0 7590500 COURT IONUAO? FIG. 30. Pohlia melanodon. A. Exothecial cells near the rim, 290. B. Habit, female plant and capsules, x8. C. Habit, sterile plant, x 10. D. Exothecial cells, x 290. E. Peristome, x80. F. Leaves, x22. G. Stomate, 290. H. Upper leaf cells showing gradation in shape toward the margin, 290. walled and somewhat overlapping (Fig. 30). ‘This is visible under a dissecting microscope as a pale, narrow strip between the urn and rim. It is particularly visible in P. melanodon and P. atropurpurea, but is also present in such other (non-Mniobryum) species as P. lescurtana. 31. Pohlia atropurpurea (Wahlenb.) H. Lindb., Acta Soc. F. Fl. Fenn. 16: 14. 1-7. 1899. Bryum atropurpureum Wahlenb. ex Web. & Mohr., Ind. Pl. Crypt. 4. 1803. Bryum pulchellum var. atropurpureum Wahlenb., Fl. Lapp. 360. 1812. (Lecto- type: S 278 B. carneum var. atropurpureum (Wahlenb.) Wahlenb., Fl. Lapp. Suppl. 1(2): Ol 1616. B. atropurpureum (Wahlenb.) Wahlenb. ex Fuernr., Flora 12(Ergbl.): 56: 1829. B. carneum var. minus De Not., Syll. 120. 1838 (nom. illeg. incl. var. prior.—B. carneum Vv. atropurpureum). Pohha borealis Arn. ex H. Lindb., Act. Soc. F. Fl. Fenn. 16: 14. 1899 (nom. nud, in synon.—P. atropurpurea). P. arnelli H. Lindb., Act. Soc. F. Fl. Fenn. 16: 14. 1899 (nom. nud. in synon.— P. atropurpurea). P. bardoensis Arn. ex H. Lindb., Act. Soc. F. Fl. Fenn. 16: 16:14. 1899 (nom. nud. in synon.—P. atropurpurea). Webera atropurpurea (Wahlenb.) Warnst., Verh. Bot. Ver. Brandenburg 42: 202, 1901. Mniobryum atropurpureum (Wahlenb.) Hag., Musc. Norv. Bor. 114. 1901. Webera atropurpurea (Wahlenb.) C. Jens., Skand. Bladmfl. 155. 1939. Tiny green or reddish plants generally scattered singly with other mosses. Stems 2—6 mm, simple when sterile or fertile. Leaves erect to erect-spreading, distant on sterile stems, overlapping on fertile stems, lanceolate, 0.6—1.3 mm long, serrulate near the apices; upper cells of leaves on sterile plants broadly rhomboidal, 65-110 um long, 10-15 um wide, thin-walled, those of the peri- chaetial bracts narrower (6-11 ~m wide). Dioicous; perichaetial bracts + dif- ferentiated, linear-lanceolate, to 3 mm long; perigonial bracts short-acuminate, erect-spreading to spreading. Capsules inclined + 180° from vertical, shortly pyriform when mature, urceolate when dry and empty, 1.0-2.5 mm long; exo- thecial cells isodiametric, 15-50 um long, with strongly sinuose walls; stomates superficial, slightly sunken or + immersed; annuli lacking; opercula conic, acute; exostome teeth dark brown, acute-triangular, most coarsely papillose near the apices, bordered, trabeculate; endostome yellow to yellow-brown; basal membrane slightly more than '4 the exostome length; segments keeled, broadly perforate; cilia long, nodulose. Spores 15-21 wm, finely roughened. Fig. 31. On calcareous, clay soils in wet places (roadbanks, ditches, stream margins), frequently mixed with P. vexans and Bryobrittonia longipes; low-montane boreal and subarctic habitats, sometimes higher in the mountains. Northern North America; Alaska, Yukon Territory, British Columbia, Washington, Minnesota, Manitoba, Ontario, lowa, Idaho, and New York: Europe and Asia. Map 30 Pohla atropurpurea is characterized by its tiny size, shortly urceolate capsules, almost superficial stomates, dark-brown exostome teeth and yellow endostomes. Pohlia vexans is easily separated by its lighter exostome teeth and hyaline endo- stome. Although P. atropurpurea can be distinguished from P. melanodon by several morphological features, the two are obviously closely related. Pohlia atropurpurea is a smaller plant and may represent a northern sister species to the more temperate P. melanodon. Vhey occupy virtually identical habitats on wet clay soil and when either species grows in the same site with P. wahlenbergii, they prefer slightly elevated, presumably drier microhabitats. 32. Pohlia brevinervis Lindb. & Arn., Musci Asiae Bor. 51. 1890. (Lectotype: H-SOL). Webera brevinervis (Lindb. & Arn.) Kindb., Enum. Bryin. Exot. 108. 1891. Small to very small, whitish, somewhat glossy plants. Stems 2-15 mm, generally simple when sterile or bearing perichaetial, sometimes innovating when bearing perigonia. Leaves erect to erect-spreading, + crowded and FIG. 31. Pohla atropurpurea. A. Habit, x12. B. Leaves, x22. C. Exothecial cells, x290. D. Stomate, X290. E. Peristome imbricate, lanceolate, 0.7—1.4 mm long, serrulate near the apices; upper cells linear-rhomboidal, 60-110 zm long, 6-9 ym wide, with thin walls. Dioicous; perichaetial bracts slightly differentiated, to 1.9 mm long; perigonial bracts short- to long-acuminate, erect or somewhat spreading. Capsules 150°-180° from vertical, shortly pyriform, 1.0-3.0 mm long; exothecial cells isodiametric, somewhat collenchymatous, 20-40 um in diameter, with strongly sinuose walls; stomates superficial or somewhat sunken; annuli lacking; opercula conic; exo- stome teeth dark to light brown or red-brown, acute-triangular, most coarsely papillose near the apices, bordered, trabeculate; endostome light yellow; basal membrane slightly more than 4 the exostome length, segments keeled, bros dly perforate; cilia long, nodulose. Spores 16-21 ym, finely roughened. Fig. 33. On clay soil along streams, outwash and other wet places in subarctic and arctic, montane habitats. Western Siberia and northwestern North America (Alaska, Yukon Territory, and Alberta). Map 31. This very rare species has been collected fewer than ten times. It is known from six localities in western North America, all close to or within areas thought to be ice-free during the last glacial episode. On the basis of the few specimens available, P. brevinervis is distinguished by relatively elongate stems with erect, whitish leaves, dark exostome teeth, vellow endostomes and slightly sunken stomates. The habit of sterile plants resembles P. vexans because of the erect, somewhat shiny leaves, but in sporophytic features P. brevinervis resembles P. atropurpurea. a — IG. 32. Upper cells from branch leaves of sterile plants versus perichaetial bracts in “Mnio- yum” species, X 260. A—B. Pohlia melanodon. A. Branch leaf cells. B. Perichaetial bract. C-D. Pohla atropurpurea. C. Branch leaf. D. Perichaetial bract. E-F. Pohlia brevinervis. E. Branch leat. Perichaetial bract. 33. Pohlia vexans (Limpr.) H. Lindb., Acta Soc. F. Fl. Fenn. 16: 20. 1899. Mniobryum vexans Limpr., Laubm. Deutschl. 2: 273. 1892. (Lectotype: BP). Webera vexans (Limpr.) Par., Ind. Bryol. 1361. 1898. Slender, glossy plants scattered singly or gregarious, sometimes forming extensive turfs. Stems 5—15 mm, generally simple when fertile or sterile. Leaves erect, Imbricate to remote, lanceolate, 0.6-1.3 mm long, serrulate near the apex; upper cells linear-rhomboidal, 70-110 um long, 6-9 ym wide, with thin walls. Dioicous; perichaetial bracts somewhat differentiated, to 1.8 mm long; peri- gonial bracts short to long-acuminate, spreading to erect-spreading. Capsules + 180° from vertical, shortly pyriform, 1.0-3.0 mm long; exothecial cells irregularly isodiametric, 15-45 wm in diameter, with slightly to strongly sinuose, evenly thickened walls; stomates deeply immersed, rarely almost superficial; annul lacking; opercula conic, apiculate; exostome teeth yellow or light brown; acute- triangular, most coarsely papillose near the apices, bordered, trabeculate; endo- stome hyaline; basal membrane slightly more than % the exostome length; seg- ments keeled, broadly perforate; cilia long, nodulose. Spores 17-23 ym, finely roughened. Fig. 34. Fairly common on calcareous clay soils in wet or moist places (streambanks, roadbanks and other disturbed sites). North-western North America (Alaska, Yukon Territory, Northwest Territories, British Columbia, Alberta, Montana, I+ FIG. 33. Pohlia brevinervis. (From the Bryologist: in press, 1982) A. Peristome, X 13. B. Stomate, x490. C. Capsules, x15. D. Stomate, x490. E. Leaves, x33. F. Exothecial cells, x 490 Manitoba); northern and central Europe; reported from northern Asia by Crundwell and Nyholm (1977). The dot map showing the distribution of this species obviously represents collection localities along the Alaska Hwy. rather than the actual distribution of P. vexans populations. Map 32. Exsicatti: Bauer, Musci Eur. Exs. 328 as M. vexans (BH), 1628 as M. vexans (WTU); Husnot, Musci Gall. 826 as W. lutescens (BH); Macoun, Flora Canadensis 628a as B. pulchellum (CANM). Pohha vexans is best recognized by erect, glossy leaves on a red stem (at least when moist). It is more common than P. atropurpurea and P. brevinervis, occur- ring along many of the sizable streams with clay banks in Alberta, British Columbia and the Yukon. It is restricted to soils of high pH, 6.5 or above. 34. Pohlia columbica (Kindb. in Macoun & Kindb.) Andr. in Grout, Moss. Fl. N. Am. 2: 202. 1935. Webera columbica Kindb. in Macoun & Kindb., Cat. Canad. Pl. 6: 115. 1892. G. 34. Pohlia vexans. A. Exothecial cells, x 290. B. Habit, male plant, x7. C. Upper leaf cells, 5. | 10. G 80. FIG. «290. D. Capsules, x 15. E-F. Stomates, «290. G. Leaves, x22. H. Peristome, x (Lectotype S—Kindb.). Bryum columbicum (Kindb. in Macoun & Kindb.) Kindb., Eur. N. Am. Bryin. 2: 386. 1897. Pohlia decurrens H. Lindb., Act. Soc. F. Fl. Fenn. 16: 12. 1899. (nom. illee. incl. spec. prior.—W. columbica). Mniobryum columbicum (Kindb. in Macoun & Kindb.) Loeske, Stud. Morph. Syst. Laubm. 124. 1910. Slender, green or reddish, rather dull plants. Stems 4-8 mm, simple when sterile or fertile. Leaves erect-spreading or spreading, rather distant, narrowly 283 lanceolate to lanceolate, 1.4-2.5 mm long, serrate near the apex or to midleaf; upper cells linear-hexagonal, 85-150 ym long, 6-10 um wide, with thin walls. Dioicous; perichaetial bracts + differentiated, linear-lancelate, to 3.0 mm long; perigonial bracts rather long-acuminate, wide-spreading. Capsules inclined 100°-180° from vertical, broadly pyriform or urceolate, 1.5-3.5 mm long; exo- thecial cells isodiametric, somewhat collenchymatous, 20-40 ym in diameter, with strongly sinuose walls; stomates immersed; annuli lacking; opercula conic, acute; exostome teeth yellow or light brown, acuminate, most coarsely papillose near the apices, bordered, trabeculate; endostome hyaline or very pale yellow; basal membrane slightly more than '2 the exostome length; segments keeled, broadly perforate; cilia long, nodulose. Spores 13-18 pm, nearly smooth. Fig. 35. Very rare; on soil in disturbed places and along streams; mostly near sea level along the Pacific Coast. eee endemic to coastal Alaska, British Columbia and Washington. Map 3 ‘his species is close to P. re but is distinguished by its duller, often reddish leaves, and less conspicuously long perigonial bracts. Whereas P. longibracteata is common along the Pacific coast, P. columbica is exceedingly rare. I have had no success finding it at the type locality or elsewhere near the coast, although W. B. Schofield has collected it in several places in British Columbia. The Japanese P. fauriei (Card.) Ochi, including Webera densiretis Broth., is a rather puzzling, variable intermediate between P. columbica and P. longibracteata, but according to Ochi (1959) distinct from both. Shiny plants of P. faurie? that look like compact expressions of P. longibracteata lack the diagnostic very long perigonial bracts. This small complex of the three closely related species is of some phytogeographic interest since they are endemic to the North Pacific region. 35. Pohlia longibracteata Broth. in Roell, Bot. Centralbl. 44: 419. 1890. (Lecto- type: H-SOL Webera longibracteata (Broth. im Roell) Ren. & Card., Rev. Bryol. 19: 66. 92 Bryum longibracteatum (Broth. in Roell) Kindb., Eur. N. Am. Bryin. 2: 384. Q7. Webera lachenaudu Card. & Ther. in Bailey, Bryologist 7: 66. 1904. (Isotype: BH, CANM, UC). Robust, shiny plants forming small populations. Stems 5-25 mm, generally simple when sterile or fertile. Leaves wide-spreading, narrowly and longly lanceolate, distant to crowded, 23.5 mm long, rather coarsely serrate near the apices; upper cells linear-hexagonal, 80-160 um long, with thin walls. Dioicous; perichaetia terminal, or on short, axillary branches; perichaetial bracts + dif- ferentiated, to 4 mm long; perigonia terminal or on short, axillary branches; perigonial bracts extremely long-acuminate, to 6mm long, wide-spreading. Capsules inclined + 180° from vertical, broadly pyriform to urceolate, 1.5-3 mm long; exothecial cells isodiametric, 20-40 ym in diameter, with straight to sinuose walls; stomates immersed; annuli lacking; opercula shortly to longly and obliquely rostrate; exostome teeth light-brown to yellow, acuminate, most coarsely papillose near the apices, bordered, trabeculate; endostome hyaline; basal membrane slightly more than 2 the exostome height; segments keeled, eau perforate; cilia long, nodulose. Spores 14-20 um, almost smooth. Fig. 36. Fairly common on calcareous, neutral or mildly acidic soils in wet, often disturbed places at sea level (ditches, gullies, roadbanks, stream margins). noges bc SST HAG Vadeo (LCG gS oie i PON IerEADOA FIG. 35. Pohhta columbica. A. Exothecial cells, x 290. B. Peristome, x80. C. Stomate, x290. D. Habit, x9. E. Capsules, x 15. F. Upper leaf cells, x 290. G. Leaves, x22. H. Habit, sterile plant, <9. Endemic to western North America; British Columbia, Washington, Oregon and California (south to Santa Clara Co.). Map 34. Exsiccau: Grout, N. Am. Musci Perf. 135 as M. albicans (MICH); Holzinger, Musci Acro. Bor.-Am. 95 as W. lachenaudii (BH, CANM, UC). Our only common Podlia along the coast of California, Oregon and Wash- ington, P. longibracteata is striking because of its large size and very glossy leaves. It is most common along roadsides in wet ditches, but can also be found in undisturbed sites such as muddy streambanks in Redwood forests. Che name refers to the perigonial bracts, which are very conspicuous, reaching six milli- meters in length. 285 a ae. LHS ROS DoCUeaZ eGo nOCaSC B2E PFC 0& 059% RO ROBES C098 IGONGO GL FIG. 36. Pohlia longibracteata. A. Exothecial cells, x 290. B. Stomate, x 290. C. Upper leaf cells, x 290. D. Habit, female plant with perichaetia terminal and on short axillary branches, x4. E. Habit, male plant with terminal and axillary perigonia, x4. F. Peristome, x80. G. Leaves, X22. H. Capsules, 5 x 228 x15. I. Perichaetial bracts, It is an interesting feature of P. longibracteata that although the inflores- cences are generally terminal, it is not uncommon to find them also on short, axillary branches (Fig. 36). Such lateral inflorescences are of the same sex as those that are terminal on the main stem. I have not found any autoicous plants. 36. Pohlia integra (Card.) Shaw, comb. nov. Webera integra Card., Rev. Bryol. 38: 4. 1911. (Isotypes: BH, BM, MO, US). Mniobryum integrum (Card.) Broth. in Engler & Prantl, Nat. Pfl. ed. 2, 10: 362. 1924. it ik | FIG. 37. Pohlia integra. A. Upper leaf cells, x 290. B. Rhizoidal tubers, x 200. C. Leaves, X22. 7 ; Very small, pale green to yellow-green, glossy plants. Stem up to 28 mm high, simple. Leaves delicate, flexuose when dry, rather distant, 0.6—-1.3 mm long, entire to crenulate near the apices; upper cells laxly elongate-hexagonal, 70-140 pm long, 13-20 um wide, with thin walls. Gemmae abundant on matted rhizoides at the stem base, also present but infrequent on single rhizoids in the leaf axils, round or more commonly oblong, red-brown, 140-240 um in the longest dimension; consisting of about 10-14 + isodiametric cells, each about 90 pm in diameter, the superficial cells not bulging. Inflorescences and sporo- phytes unknown. Fig. 37. On calcareous clay in rock crevices; endemic to central Mexico (Ixtaccthuatl). Map 35. Although P. integra is known only from the type locality, the original speci- men must have been copious for isotypes are present in a number of herbaria. I have not seen a specimen from the Cardot herbarium in Paris, but since the type material seems to be uniform it would be surprising if such a specimen differs significantly from others. Although sterile, a relationship to the previous “Mniobryum” species is indicated by the lax leaf cells. Pohlia integra is distinguished most readily by abundant, deep red, rhizoidal tubers. Compared to P. wahlenbergii, which also occurs in central Mexico, P. integra has more yellowish, glossy leaves. DOUBTFUL TAXA Type specimens for a relatively extensive list of species described by Hoppe and Hornschuch and other early authors were not available for study. Such early names could prove important if types are found in the future. Webera micro- apiculata C. Mull. & Kindb. in Macoun, Cat. Canad. PI. 6: 115. 1892: A type for this species is in the Kindberg herbarium. It is dioicous, lacks sporophytes, and has erect, somewhat glossy leaves. Although it does not agree with any of the species included here, it hardly seems distinctive enough to recognize at present. Only additional collections will help distinguish it as a good species rather than an aberrant form of something else, perhaps P. cruda. oy os Ns ry . 1 Ly Vy Se Yh Ly Yi — , // : ae > so pp Ly Yi ay ad MAPS 1+. I. Pohlia sphagnicola, 2. Pohlia nutans. 3. Pohlia elongata var. elongata. 4. Pohlia Hate var. greeni MAPS 5-8. 5. Pohlia cruda. 6. Pohlia oerstediana. 7. Pohlia bolanderi var. bolanderi (solid circles) and var. seriata (hollow circles). Half black circles on Mt. Rainier, Washington marks an area of sympatry. 8. Pohlia longicollis. 289 eS iS AAS SS fl \ 9. Pohha polycarpa. 10. Pohlia obtusifolia. 11. Pohlia crudoides. APS 9-11. M 290 \ 2 5 ' . Tropic of Capricorn i ay! : . MAPS 12-15. 12. Pohlia papillosa. 13. Pohlia richardsii. 14. Pohlia pseudo-barbula, 15. Pohlia ludwigit. 29] )} ye A Tye EA f bh iw i, a if 21. 16. 6 MAPS 1 0. Pohlia filum. Pohla cardotii. 17. Pohlia erecta. 18. Pohlia beringiensis. 19. Pohlia drummondii. 21. Pohha andalusica. 2 292 c 5. Pohlia proligera. 26. 23. Pohlia bulbifera. 24. Pohlia andrewsii. 2 APS 22-27, 22. Pohlia tundrae. 27. Pohlia camptotrachela. Pohlia annotina. M 293 ete Contr. Univ. Mich. Herb. 15:297-309. 1982. SELECTED SPECIES OF LEUCOCOPRINUS FROM THE SOUTHEASTERN UNITED STATES Helen V. Smith Herbarium and the Matthaei Botanical Gardens The Cer) of Michigan and Nancy ‘Ss Weber Herbarium The University of Michigan Ann Arbor, Michigan 48109 The present contribution on Leucocoprinus is part of a larger project on the macrofungi of the southeastern United States by Nancy S. Weber, Alexander H. Smith, and Dan Guravich. Among the numerous problems encountered were collections belonging to the genus Leucocoprinus, some of which are here re- ported upon. This report covers those members of Leucocoprinus sensu stricto examined to date, but is not a comprehensive treatment of the genus for the Southeast. The species discussed here are gilled mushrooms with thin-fleshed, typically fragile fruiting bodies whose buttons often resemble those of certain species o Coprinus. The pileus is striate to plicate-striate by maturity at least near the margin, and the lamellae are separated from the stipe by a collarium. The spores are smooth and white or nearly so in deposit. Brachybasidioles are present and usually inflated in the mature hymenium. Dried material was studied in the usual manner for dried mushrooms. The pilear cuticle was studied using sections and squash mounts of the disc and about midway between the disc and the margin. Except as noted, all collections studied are deposited in the Herbarium of the University of Michigan (MICH). Color notations in quotation marks are taken from Ridgway (1912). Of particular importance to this project were the collections and excellent notes on them made by Ervin Hillhouse of Liverpool, ‘Texas. KEY TO SPECIES — eee body white, dingy white, beige, tan, vinaceous, or lilac brown; lacking yellow lor I. entre body yellow or yellowish, with or without a brown disc. 5 Fruiting body white to dingy white, ee or light tan; lamellae white or nearly so; spores hyaline to light ochraceous in KOH. 3 2. Fruiting body vinaceous brown to fh ac brown on the disc and scales of the pileus, similarly colored to paler elsew ate ue and ee (in KOH) becoming pale peach color to pale flesh color at least 1 eo be iain 3. Striae on pileus long, crowded, eens nearly to the disc; surface finely eee ee finely fibrillose-scaly. oe mgr 3. Striae on pileus more widely spaced, shorter; surf: red with floccose sc Beco 4 4. Fruiting body pure white and remaining so (sometimes otk a tinge of buff on ae ce and surface of the stipe when bruised); soft warts present over the disc; typically fruit- e singly or in small clusters. 3. L. breviramus 4. Fruiting body off-white to dirty white, sometimes brownish or grayish; disc lacking warts, glabrous; typically fruiting in large clusters 4. L. Bere 297 298 5. Fruiting body exceedingly fragile; pileas membranous and soon collapsing. 6 5. Fruiting body + fleshy and not collapsing when picked. fd 6. Pileus bright yellow except for the brown disc; large cystidia (35-105 x 16-25 jum) present on young lamellae. 5. L. magnicystidiosus 6. Pileus pale yellow, disc dull olive yellow; large cystidia absent from lamellae. 6. L. fragilissimus 7. Pileus pale greenish yellow with a contrasting brown disc; spores less than7 wm long. 7. L. flavescens ae 8 Pileus (including disc) bright or deep yellow; spores over 7 wm long. . Pileus radially lacerate-fibrillose, scarcely striate until partially dried; spores lacking a conspicuous apical pore. 8. L. flavus 8. Pileus at first covered with small floccose scales, distinctly striate when fresh; spores with a conspicuous apical pore. 9. L. luteus |. Leucocoprinus lilacinogranulosus (Henn.) Locq., Bull. Soc. Linn. Lyon 12: 95. 1943. Lepiota lilacino-granulosa Henn., Verh. Bot. Vereins Prov. Brandenburg 40: 145. 1889. Illustrations: Pl. 1, Fig. 4, a—c; Pl. 2, Pig. -4,a, b:; Pileus 1.5-4.5 cm broad, 2.0-3.3 em high at maturity, columnar and truncate at first, becoming broadly conic at maturity, plicate-striate nearly to the disc; cuticle over disc continuous, elsewhere breaking into fine scales which are more widely spaced toward the margin; buttons “deep brownish vinaceous” to “light grayish vinaceous,” darkest over disc, mature specimens “benzo brown” over the disc and scales, otherwise “tilleul buff.” Context thin. Odor and taste mild. Lamellae off-white to very pale pinkish buff in age, moderately crowded, becoming wrinkled in age. Stipe 4-5 cm long, 2.5-4.0 mm thick at the apex, 7.5-10.0 mm broad at the widest part of the somewhat bulbous base; white with a “light grayish vinaceous” cast over the base, unchanging when injured. Annulus thin, located just above the bulb. Spores 7.5-11.0 x 5.0-7.5 sm, elliptic to ovate in face view, inequilateral in profile, moderately to strongly dextrinoid, hyaline to pale peach color in KOH; apical pore prominent, with a hyaline, domelike cap. Basidia from young pileus 37-42 x 7.5-9.0 in, 4-spored, apex truncate, base tapered. Brachybasidioles not readily demonstrated in young pilei where basidia are elongating, prominent in pilei with mature spores and then 12-16 wm in diam. Pleurocystidia not observed. Cheilocystidia 30-66 x 8-17 wm, clavate to irregularly elongate-inflated, hyaline, abundant on young specimens, apparently collapsed by maturity. Clamp connections not seen. Pilear cuticle over disc a zone several cells thick of inflated, mutually compressed, + ellipsoid cells whose long axis is perpendicular to the surface of the cap; end cells staggered, not forming a palisade, contents pale rust color as viewed in KOH: toward the edge of the disc, cells become decumbent and longer; transition to radially-arranged hyphae evident, tips of hyphae in fascicles forming the scales. Cells in radially-arranged hyphae with peglike branches, slightly inflated, some- times disarticulating under pressure, versiform cells increasingly abundant toward the margin and between scales, these cells often with stubby branches; cells in scales ellipsoid, subglobose, boomerang-shaped, or elongate-inflated, often with a dull purplish tan content. Material studied. Guravich 1280, gregarious to cespitose in a longleaf pine nursery at the Harrison Experimental Forest of the De Soto National Forest, Harrison Co., MS: 10 July 1981. leg. Weber, Guravich, Cibula, and Izral. This species was described from specimens that fruited in the orchid beds in the greenhouses of the Berlin Botanical Gardens, and so far as we know has not been previously reported from North America. Except that our specimens and Zo9 their spores are slightly larger than indicated for European material, they match the European description and illustrations well. Moser (1978) gives the spores as (7) 8.5-10.5 (12) * 6-6.5 (7.5) wm which approximates that observed in our material; however Hennings gave ae as 10-13 x 7-9 wm and more recently Bon (1981) indicated they are 8-9.5 9-7 (8) wm. We are unable at present to account for these variations. 2. Leucocoprinus longistriatus (Peck) Smith & Weber, comb. nov. Lepiota longistriata Peck, Bull. Torrey Bot. Club 25: 368. 1898. Illustrations: Pl. 1, Fig. 2, ac. Pileus 4.3-8.0 cm broad, convex to plane or the margin uplifted in age, umbo distinct or not at maturity; surface plicate-striate nearly to the disc at maturity, striae fine, close, smooth; cuticle probably entire at first, remaining smooth over the disc but breaking up into fine fibrillose scales and fibrils especi- ally over the ridges of the striae, scales easily brushed off; cuticle smoky brown to “vinaceous buff” over the disc, light buff becoming “ivory yellow” elsewhere. Lamellae “cartridge buff’ to duller in age, edges very finely fimbriate, even, concolor. Stipe 4.0-8.3 cm long, 3-6 mm at the apex, 6-12 mm at the base when mature; shiny, glabrous, “cartridge buff” to “ivory yellow” or white, staining dull ochraceous when handled when young. Annulus a small, tight, ascending collar with a pale brown margin; usually located well below the midpoint of the stipe, often just above the small bulb, deciduous or persistent. Spores 6-8 (—9) x 4.5—-5.0 wm, ovate in face view, elliptic In profile, lacking a distinct apical pore but under oil immersion there is a ee of a thin spot at the apex, not dextrinoid. Basidia 4—spored, 16-21 x 9-11 wm, scarcely projecting beyond the brachybasidioles. Pleurocystidia not seen. Cheilocystidia 91-33 x 8-12 um, clavate or fusoid-ventricose, some mucronate or with a finger-like projection, hyaline. Pilear cuticle over the disc of radially arranged, anastomosing hyphae; the cells not inflated, many disarticulating at the septa and with a short branch just below the septum that forms a false clamp; cuticle over striate portion of fascicles of radially arranged hyphae 6-19 wm broad, end cells usually not erect, their apex rounded, slightly tapered, or clavate; content hyaline to light ochraceous brown in KOH. Clamp connections not seen. Material studied. Guravich ee cespitose to scattered on chipped-bark mulch and on =k under sweet gum and live oak, Davis Bayou picnic area, Gulf Islands National Seashore, Oceat Springs, Nees Mississippi; 13 July 1981, leg. Weber and Guravich; F. S. Earle, Alabama, Ta (TYPE, NYS). This species was described by Peck from specimens collected by F. S. Earle in Alabama and was later reported by Murrill (1911, 1972) from lawns and woods in Cuba and Jamaica as well as Florida. The distinguishing features are the numerous, fine, long striae on the cap, light bown disc and paler margin, and easily removed fibrils and scales. Jur notes on the microscopic features of the holotype follow: Spores 6.5— 7.5 X 4.5-5.0 wm, ovate to elliptic in face view, elliptic in profile, lacking an obvious apical pore, faintly yellow as revived in KOH, not or only faintly dextrinoid. Basidia reviving poorly, about 10.5-13.5 wm broad, apparently embedded in a pavement of branchybasidioles, probably 4-spored as there are numerous tetrads of spores. Pleurocystidia not seen. Cheilocystidia 27-37 x 8-15 wm (length estimated only), subcylindric to clavate, abundant. Pilear aa over striate portion of + radially arranged hyphae; the cells elongate, not 1 flated, seldom disarticulating, not distinctly pigmented; terminal cells dee at the apex; hyphae seldom branched. 300 3. Leucocoprinus breviramus Smith & Weber, sp. nov. Pileus 3.5-9.0 cm latus, conicus demum campanulatus vel late convexus, ad marginem brevistriatus vel non-striatus, candidus vel ad apicem sordide hraceus, squamulosus demum glaber. Lamellae librae, albidae. Stipes 5.5— 8.0 cm longus, 3-5 mm ad apicem, deorsum 10-12 mm crassus, annulatus, sursum glaber, albidus, tactu + ochraceus. Annulus parvus, evanescens. Sporae 7.5-9.0 (10) x 5.5-6.5 wm, dextrinoideae, ad apicem pori praeditus. Pleuro- cystidia nulla. Cheilocystidia versiformia, 34-67 x 9-12 um. Specimen typicum in Herb. Univ. Mich. conservatum est: legit E. Hillhouse 264, Liverpool, Brazoria — Co, Texas, 6 Aug, 1971; Illustrations: Pl. 1, Fig. 3, a, b; Pl. 2, Fig. 1, a, b. Pileus 3.5—9.0 cm broad when fully mature, at first cylindric or finger-like, then conic to campanulate and finally plane or somewhat concave with the margin uplifted; striate near the margin only when mature, at times scarcely striate; snow white and remaining so except for a tinge of buff on the disc; covered with small rounded floccules which are smaller and denser on the disc, scattered toward the margin, easily rubbed off and sticking to the fingers; margin usually incurved until near maturity, frequently appendiculate with white floccules. Context white, 3-4 mm thick near the stipe, tapering and thin at the margin. Odor slightly musky, taste when raw mild. Lamellae white, unchang- ing, ventricose, of two lengths, about 15 per cm near the margin, edges even. Stipe 5.5-8.0 cm long, 3-5 mm thick at the apex, 10-12 mm thick at the elongate bulb; clothed with small floccules like those of the pileus below the annulus, glabrous above; white but at times staining slightly yellow when handled; hollow, stuffed, or solid. Annulus superior, rather small, membranous but mealy below, usually disappearing and leaving only small particles on the stipe, or remaining as a skirtlike ring, white. Spores 7.5-9.0 (10) x 5.5-6.5 wm, broadly ovate to elliptic in face view, elliptic in profile; the distinct small apical pore covered by a hyaline dome; dextrinoid. Basidia 4-spored, stubby. Brachybasidioles inflated. Pleurocystidia not seen. Cheilocystidia 34-67 x 9-12 wm; subcylindric to faintly ventricose or clavate; abundant; hyaline. Pilear cuticle over the disc of inter- woven hyphae of variable diameter (3.5—12.0 wm broad), predominantly narrow at the surface, septate, not inflated, branched, the cells mostly irregular in shape, scattered, + erect clavate cells present also especially in the scales; toward the margin of narrow hyphae that are tangled, not inflated and divided into short, branched cells suggestive of pieces of a 3-dimensional puzzle. Oleiferous hyphae present in context. Clamp connections not observed, but “false clamps” present. Material studied. Hillhouse 264 (TYPE, MICH), Liverpool, Brazoria Co., Texas, 6 Aug. 1971; also Guravich 867, Greenville, Washington Co., Mississippi, 29 Aug. 1977. Hillhouse reported that it usually fruits singly but sometimes in small clusters, on lawns, in gardens, and on wet hay in the summer and fall. Dried fruiting bodies are warm ivory on the pileus, the lamellae are + pinkish buff, and there are no signs of discoloration. According to Hillhouse it is “some years plentiful and other years not to be found at all. Seems to be best during dry summers and comes on two or three days following a good shower. The summer of 1970 was a very wet year and I found none at all. . . the cespitose habit is not very prominent in this area with five in one tuft the most I have ever found: most are found as single... . Slight musky odor, raw taste mild, cooked taste of young buttons very good, no upsets from any amount eaten which at times have been large servings.” L. breviramus differs from L. cepaestipes by its persistent clean white appear- 301 ance, pileus that is striate only near the margin with prominent and persistent floccules on the disc, firm flesh, and non-cepsitose habit of fruiting. The specific epithet refers to the short, stubby branches of the cuticular cel L. breviramus is closely related to Lepiota cretata Locquin (in Haller, 1950) of Europe. However, Locquin proposed Lepiota cretata as a new name for Agaricus cretaceus Bulliard which is presumably the type for the species. An excellent description and discussion of L. cretata is presented by Josserand (1955). He described the species as a beautiful white Lepiota whose pilei are covered with snow white floccose scales as in the case with our species and he pointed out that his species could not be Buillard’s species. In Builliard’s illustration the covering of the pileus and stipe is clearly shown as consisting of slender fibrils intead of soft, almost globose to granular floccules. Josserand rejected the Builliard plate as representing Josserand’s taxon as we reject it for ours. Consequently we also reject the name Lepiota cretata for our species. Although there is considerable similarity between our species and that cur- rently called Leucocoprinus cretatus in Europe, there are differences. The Euro- pean taxon is seemingly more robust; fruits in large clusters; has larger spores, a bitter taste, and firm to hard flesh when young. 4. Leucocoprinus cepaestipes (Sow.: Fr.) Pat., Tab. Analyt. Fung. 7: 45. 1889. Agaricus cepaestipes Sow.: Fr., Syst. Mycol. 1: 280. 182 Illustrations: Pl. 1, Fig. 1, a, b; Pl. 2, Fig. 3, a, b. Pileus 2-8 cm broad, fragile, very thin, cylindric to ovoid, becoming broadly convex to plano-convex; white to whitish at first, becoming dingy white to buff and the disc often pale tan to yellowish brown; surface continuous over the disc and glabrous to minutely tomentose, toward margin decorated with fine mealy to fibrillose scales; disc truncate when young, sometimes umbonate at matur ity; margin entire at first, becoming striate, striate-rimose, or plicate-striate to sulcate. Context thin, fragile, white or staining straw yellow to light pinkish cinnamon when bruised. Odor and taste not distinctive. Lamellae remote from stipe, crowded, thin, white becoming dingy white; margin even, finely fimbriate under a lens. Stipe 3-13 cm long, 2-6 mm at apex, enlarged downward and somewhat fusiform-ventricose to distinctly clavate toward the base, sometimes terminating in a short point in age; glabrous to faintly pruinose above, tomen- tose below; white at first, becoming delicately tinged pinkish tan to pale brown, sometimes straw yellow then light brown after handling. Annulus rather large, flaring, membranous with a lacerate eae puperOl, white to whitish, sub- persistent to evanescent. Spores 9-11 x ~7.5 am, ovate to subelliptic in face view, somewhat inequilateral in profile, nae truncate from a small apical pore, thick-walled, moderately to strongly dextrinoid, hyaline or in age dull a eae in KOH. Basidia 35-40 xX 11-13 wm (estimated). Bra Sy ae esent, reviving poorly. Pleurocystidia not seen. Cheilocystidia 37-6 _ 16 wm (or longer, base hard to locate), clavate to bottle- -shaped with a ane sometimes forked at apex, hyaline, abundant on young lamellae, collapsing by maturity. Pilear cuticle over disc a zone several cells thick of inflated, versiform, globose, or ellipsoid cells 11-40 x 9-22 um, these usually mutually nes giving rise to a turf of elongate erect or nearly so pileocystidia 45-99 x 4-9 ww toward the margin of fascicles of radially arranged hyphae which give rise to sausage-shaped to oblong or irregular cells with stubby branches at the surface. Material studied. Hoseney 2/15, on lawn extension. 620 Church Street, Ann Arbor, Washtenaw Co., Michigan, 14 July 1972 Observations. L. cepaestipes and its close relatives are considered “southern” species by most American mycologists and it is consequently included although 302 we have not found it there to date. It is, however, another common species which “everyone knows” but for which, when published descriptions are compared, there is no consensus on its characters. ‘The concept presented here is adapted from H. V. Smith (1981) for the macroscopic characters and Hoseney 2115 for the microscopic features. This collection is the one illustrated in A. H. Smith and N.S. Weber (1980, pl. 163). In summary, although L. cepaestipes has been known to mycologists for about two centuries, the limits of the species are still in need of clarification. 5. Leucocoprinus magnicystidiosus Smith & Weber, sp. nov. Pileus 4-8 cm latus, conicus, campanulatus vel planus, ad marginem valde striatus, laete luteus vel ad centrum brunneolus; fragilis; gustus leviter amarus. Lamellae luteae vel pallidior, 2-3 mm latae. Stipes 7-15 mm longus, 2-4 mm crassus, bulbosus, luteus, annulatus. Annulus superne albidus, inferus luteus, evanescens. Sporae 9.0-13.5 x 7.5-9.0 um, valde dextrinoideae, ad apicem porl praeditus. Pleurocystidia 35-105 x 16-25 (30) um, tenui-tunicata, ovoidea ellipsoidea vel subfusoidea. Cheilocystidia 18-25 x 10-17 ym, vesiculosa, clavata vel versiformia. Cuticula pileorum cellulae globosae vel subglobosae praeditus. Specimen typicum in Herb. Univ. Mich. conservatum est: legit F. Hillhouse 266A, Liverpool, Texas, AUG 197. 1, Illustrations: Pl. 3, Fig. 1, a-g. Pileus 4-8 cm broad when fully expanded, when young cylindric and up to 1.5 cm wide and 3.2 cm high, soon elliptic to narrowly conic becoming campan- ulate to plane or concave due to an uplifted margin; striate to plicate nearly to the disc; bright lemon yellow except for the brown disc when young, at maturity the striae bright yellow, grooves white; extremely fragile at all stages, sticking to the fingers and tearing readily; context white, less than 1 mm thick except at the disc. Odor not distinct, taste faintly bitter. Lamellae yellow to white, paler than the surface of the pileus, ventricose, 2-3 mm broad: lamellulae few. Stipe 7- 15 cm long, 2-4 mm broad, equal or nearly so down to the somewhat bulbous base; yellow like the pileus or darker, sometimes reddish brown: extremely fragile, sometimes larger specimens break from their own weight. Annulus yel- low below, white above; soon evanescent except for a flimsy ring on the stipe and a few small fragments appendiculate on the margin of the pileus. Spores 9.0-— 13.5 X 7.5-9.0 «um, ovate to + lemon-shaped in face view, ovate to slightly inequilateral in profile, drawn out into a snout-like apex with a distinct pore covered with a domelike cap; moderately to strongly dextrinoid, the dome over the pore remaining hyaline; spore wall thick, complex. Basidia 4-spored, 20-39 x 10-12 wm, becoming greatly elongated at sporulation. Brachybasidioles in- flated in mature specimens, 13.5-16.5 (-21) um broad, forming a pavement. Pleurocystidia 35-105 x 16-25 (-30) ym, thin-walled, hyaline, elongate-elliptic tapering to a rounded or pointed apex, base narrow and pedicel-like; cystidia extend well beyond hymenium, often longer than lamellae are thick, most abundant near the edge of the lamellae, common in young specimens, pre- sumably collapsed in old ones. Cheilocystidia of two types: 1) those similar in size and shape to the pleurocystidia, and 2) those 18-25 x 10-17 fm, abundant, variable in shape (vesiculose, broadly obovate, or pyriform), and persistent in old specimens. Pilear cuticle over disc a thick zone of inflated, mutually compressed cells terminating in a pavement of rounded cells 9-13 am in diam: context below the cuticle of uninflated hyphae many with false clamp connection, numerous oleiferous hyphae also present; cuticle in striate portion a mixture of globose, subglobose, and inflated ellipsoid cells 15-72 x 13-54 wm, and narrow angular cells 3-8 wm broad with stubby branches. 303 Material ec Hilthouse 266A (TYPE, MICH), 7 Aug. 1971, Liverpool, Brazoria Co., Texas; A. H. Smith 14871, 17 Aug. 1939, Cades Cove, Tennessee. Hillhouse gave the following information on 266A and two ae collections not discussed here: “Summer and fall, scattered and gregarious on St. Augustine lawn and in pine and mixed woods growing from the soil and in layers of pine needles and leaves among grass, most aoe at times. Found only during and shortly after rains or in early morning when dews are he L. magnicystidosus differs from L. fragilissimus by its brighter colors and presence of immense pleurocystidia. L. thoenit Heinem. from Zaire is similar but has larger spores (11.4—16.0 x 8.4-10.0 wm) and lacks pleurocystidia. L. magnicystidosus has extremely fragile fruting bodies, Hillhouse reported that the stipe was “extremely fragile and even a slight breeze will cause it to break once the fruit has reached maturity. [ have had the stem break on specimens up to 8 cm across just from my breath while trying to get a picture. The larger specimens seem to sometimes break from their own weight. ... Hot sun causes these to quickly deteriorate and disappear and none will be found except early in the morning on hot days.” Dried specimens have a distinct vinaceous cast over the disc and the striae. 6. Leucocoprinus fragilissimus (Rav. in Berk. & Curt.) Pat., Essai Taxon. p. 171. 1900. FAiatula fragilissima Ray. in Berk. & Curt. Ann. Mag. Nat. Hist., ser. 2, 12: 429. 1853 Illustrations: Pl. 3, Fig. 1, a-. Pileus 2—5(—5) cm broad, becoming plane with a depressed umbo or the margin uplifted, striate nearly to the umbo, pulverulent with small yellow squamules except over the disc; disc “ecru olive” (dull olive yellow), to “dee colonial buff,” elsewhere “pinard yellow” to “straw yellow;” fragile. Lamellae white, narrow, subdistant to crowded. Stipe 6-10 cm long, 1-2 mm thick at apex, 5-6 mm at base; fragile, pale yellow, powdery then glabrescent. Annulus small, superior. Spores 9-14 x 7.0-8.5 wm, broadly elliptic in face view, broadly inequilateral in profile; apex drawn out and almost snout-like, capped with a hyaline dome over the apical pore; apiculus inconspicuous; strongly dextrinoid. Basidia 21-25 x 9.0-10.5 um, stout. Brachybasidioles 9-11 x 10-15 wm, abun- dant. Pleurocystidia not seen. Cheilocystidia 13-25 (-36) x 9-15 (-20) um, broadly clavate, sometimes incrusted, single or in fascicles. Pilear cuticle over the disc a zone several cells thick of inflated, mutually compressed cells terminating in a pavement of rounded cells; transition to rest of pileus has mounds of in- flated, more or less globose cells 12-54 x 12-41 wm in diam mixed with narrow (3.5-7.0 wm wide) hyphae whose cells are branched and irregular in shape, the whole apparently formed by the ends of radially aranged hyphae and continuing in this manner to the margin, a mixture of big cells and narrow ones, many of the inflated cells have diffuse eae gray contents. iterial studied. Guravich 1346, scattered in mixed woods, Gulf Islands National Seashore, os a ee Jackson Co., Vastshas oar 1981; leg. Weber and Guravich; also Guravich 1268, scattered under Se ae Harrison Experimental Forest of the De Soto National Forest, Harrison Co., Mississippi; 18 July 1981. Sections mounted in KOH often release a yellow pigment; the disc and striae dry light gray to brownish gray. These fruiting bodies are almost unbeliev- ably fragile and difficult to collect. L. fragilissimus was described from South Carolina and is widely distributed in tropical and subtropical regions: Pegler (1972) reported it from Ceylon (now Sri Lanka), and Heinemann (1977) re- ported it from Zaire. 304 7. Leucocoprinus flavescens (Morg.) H. V. Smith, Michigan Bot. 20: 50. 1981. Lepiota flavescens Morgan, J. Mycol. 13: 5. 1907 ‘This species was described from southern Ohio and has been reported from greenhouses in Massachusetts and California as well. It fruits in warm humid habitats and we expect it will be found in the southeastern United States. ‘The description is from H. V. Smith (1981). Pileus (1.7) 2.5-4.0 cm broad, cylindric-ovate becoming expanded to plane with an ones or flattened umbo, plicate-striate almost to the disc, pale green- ish-yellow (near “napthalene yellow”), the disc brown and glabrous and contrast- ing strongly with the rest of the surface, ridges floccose to powdery, the grooves glabrous and white. Flesh thin, white. Lamellae yellowish, remote from the stipe, close to subdistant, narrow, subventricose. Stipe 3-6.5(-8) cm long, 2 mm thick at apex, enlarging to 3—5 mm thick at base, pale sulphur-yellow and covered wuh fine Bocenlee like the pileus. Annulus superior, pale yellow, delicate and often evanescent. Spores 4.8-6.6(-7.2) x (3.5-)4.9-5.5 wm, broadly ellipsoid, sub- globose or globose, pale oe brown to rusty-brown in Melzer’s sol., apical pore lacking. Basidia 15-18 x 7-8 wm, clavate. Brachybasidioles 14-18 x 10- 15 wm, saccate and thin-walled. Cheilocystidia 17-43 x 8-15 wm, abundant, clavate, subcylindric, fusoid-ventricose with an obtuse apex, or cylindric with a short neck and a head. Lamellar trama loosely interwoven, oleiferous hyphae present. Pileal trama loosely interwoven, oleiferous hyphae present. Cuticle of pileus a compact layer of enlarged, clavate to pyriform, vesiculose, broadly fusoid-ventricose, or globose, thin-walled cells 10-50 wm broad which may also be found in clusters along the ridges of the striae, but generally not free- floating in the mount. Clamp connections not found. 8. Leucocoprinus flavus moe Heinem. Bull. Jard. Bot. Nat. Belg., Bull. Nat. Plant, Bele. 47: 84. 1977 Lepiota flava Beeli, Bull. - Roy. Bot. Belgique 64: 215. 1932. Illustrations: Pl. 1, Fig. 1, a-c. Pileus 1.8—2.7 cm broad, conic to broadly conic, radially lacerate-fibrillose, scarcely striate until dried, brilliant yellow with the umbo deeper in color and drying brown, umbo slight, disc lubricous. Context thin. Taste mild. Lamellae distant sues the stipe. Stipe 5-6 cm long, about 2 mm at apex, nearly equal to a bulb 3-7 mm thick at the base, pale yellow, base sometimes with a small mycelial mass. ae flaring upward, membranous, persistent, pale yellow. Spores s (6— ) 7.5-9.0 X 3.8-5.3 wm, narrowly ovate to narrowly almond- shaped i in face view, ee ovate to elliptic in profile: apical pore absent; wall not conspicuously thickened; spores moderately dextrinoid. Basidia 4—-spored, approximately 25 x Yum, not reviving well. Brachybasidioles not conspicuously inflated at maturity. Pleurocystidia not seen. Cheilocystidia 33-49 x 7-11 wm, abundant, hyaline, subcylindric to somewhat fusoid-ventricose. Lamellar trama interwoven, of large cells. Minute refractive granules present in many cells in the hymenium. Pilear cuticle near the disc on mature pilei bearing mounds of short, curved (sausage-like) cells 4.4—10.5 wm broad, slightly inflated, sometimes bearing one or more branches, end cells broadly S-shaped or narrowly clavate; toward the margin of fascicles of radially arranged to loosely interwoven hyphae, hyphae unbranched, not disarticulating, 4.5-12.0 (-20) wm broad. Material studied. Weber 4583, under mixed hardwoods, McCormick’s Creek State Park, Owen Co., Indiana, 15 Aug. 1981 Unlike most species in the genus, the margin of the cap in L. flavus does not become striate until the carpophore is at least partially dried. The brachy- 305 basidioles do not become inflated; Beeli (1932) described them as slenderly cylindric, and we had difficulty finding them. Our specimens agree well with the descriptions and illustrations given by Beeli (1932, 1936) and Heinemann (1977). This represents the first report of L. flavus from the United States. It seems likely that it will be found in the South as it was described from Zaire where warm, humid weather is also present. 9. Leucocoprinus luteus (Bolt.) Locq., Bull. Mens. Soc. Linn., Soc. Bot. Lyon 14: Agaricus luteus Bolton, An History of Fungusses growing about Halifax 2: 50. 1788. Leucocoprinus birnbaumii (Corda) Sing., Sydowia 15: 67. 1962. ¢ Agaricus birnbaumu Corda, Icon. Fung. 3: 48. 182 ihistrationss Rly 1; Fig-6, a, b; Pl: 2, Fig. 2; a, b. Pileus 3.0—-7.5 cm broad when mature; finger-like to conical at first, becom- ing broadly expanded to plane with a slight umbo; long striate to almost sulcate; covered with powdery to mealy, soft, floccose warts that are easily rubbed off; evenly lemon yellow to sulphur yellow. Context thin, at most 2—3 mm thick near the stipe, rather brittle when young, pliable when mature. Odor not distinctive; taste mild and pleasant but leaving a bitter aftertaste. Lamellae not crowded, pale yellow, soft and readily collapsing, of 3-4 lengths. Supe 6-9 cm long, 3- 4 mm thick at apex, up to 11 mm thick at the clavate base; minutely floccose- pulverulent near the base; brilliant yellow to pale greenish yellow; solid to pithy, usually hollow with age; white mycelioid covering sometimes present at the base. Annulus small, yellow, collar-like, often evanescent. Spores (8—) 9.0-10.5 (-12) x 6.0-7.5 wm, ovate in face view, inequilateral in profile; apex truncate or not, apical pore with a hyaline dome; wall thick; dextrinoid. Basidia 33-53 X 6-8 wm, elongate at the time of sporulation and projecting well beyond the surrounding brachybasidioles. Brachybasidioles broad, inflated to subglobose, some with weakly dextrinoid content. Pleurocystidia not seen. Cheilocystidia 40-66 x 10- 15 wm; clavate, elongate, tapered, + ventricose, or irregular, some with finger- like apical projection; abndant; hyaline; collapsing at maturity. Pilear cuticle over the disc of hyphae with a mixture of short, branched, uninflated cells and predominantly of inflated, sausage-shaped, ellipsoid, pyriform to subglobose or globose cells; toward the margin of loosely interwoven hyphae forming mounds of narrow intact hyphae and short, disarticulated cells with stubby branches. Material studied. Hillhouse 147, Liverpool, Brazoria, Co., Texas; 13 June 1970; Guravich 1373, Davis Bayou picnic area, Gulf Islands National Seashore, Ocean Springs, Jackson Co., Mississippi; 17 July 1981, leg. Weber and Guravich. Hillhouse reports that this species occurs on decaying plant material, particularly decaying hay, in rich garden soil, on lawns or rarely on logs or stumps or in shaded lawns under live oak from late spring to fall in Texas. This species is often found in greenhouses and similar environments such as shopping malls in cooler regions but it grows outdoors in the warm regions of the world. It has been reported to be poisonous, but Hillhouse recorded the following observations on collection 147: “Cooked taste about like the com- mercial variety of Agaricus but with a slight bitter aftertaste. | have eaten fairly large amounts of these on several occasions without upset of any kind. For the past three or four years I have had about a hundred bales of hay stacked near my garden where I use it for mulch. I have had large amounts of this mushroom and another I believe to be Lepiota cepaestipes come up in the hay and in my garden. I eat both without upset.” e use of the pre-Freisian epithet /uteus instead of the post-Friesian epithet birnbaumii is mandated by the rule changes promulgated at the International Botanical Congress in Sidney, Australia in 1981. PLATE 1. FIG. 1, L. flavus (Weber 4583), a, cheilocystidia; b, basidium; c, ee FIG. 2, L. longistriatus (Guravich aa a, basidium and genkey b, spores; c, eee stidia. FIG. 3, L. breviramus (Hillhouse 264), a, cheilocystidra; ee oe res. FIG. 4, L. hlacinogranulosus a ra a, asidium and brachybasidiole; b, cheiloc c, Sp _ FIG. 5, L. cepaestipes (Hoseney 2115), a, cheilocystidia, b, spores. FIG. 6, L. aan ie 147), a, spores; b, cheilocystidia. All x 1,000. — 4b PLATE 2. FIG. 1, L. breviramus (Hillhouse 264), a, cells from plicate portion; b, cells from. disc. FIG, 2, L. birnbaumiu (Hillhouse 147), a, cells from plicate portion; b, cells over disc. FIG. £ cepaestipes (Hoseney 2115), a, cells from plicate portion; b, section of cuticle over disc. FIG. 4, L. lilacinogranulosus (Guravich 1280), a, hyphae over plicate portion; b, cells from disc. Fig. 1 x 1,000, all others x800. Pree yeviias PLATE 3. FIG. 1, L. fragilissimus (Guravich 1346), a, cells from striate poruon of pileus; b, cheilocystidia; ¢, spores. All x800. FIG. 2, L. magnicystidiosus (Hillhouse 266A), a, section of pilear cuticle over disc; b, large cysudia, both pleuro- and c eilocystidia; c, cells from cuticle in striate portion; d, persistent cheilocystidia; e, spores; f, basidium; g, brachybasidioles. All x 1,000 except 0 a &c which are X800. — 309 ACKNOWLEDGMENTS The directors and staff of the Matthaei Botanical Gardens and the Herbarium generously and the Latin diagnoses. The junior author’s field work was aided by Dr. Harold Howse who made the facilities of the Gulf Coast Research Laboratory, Ocean Springs, Mississippi available. Dr. William Cibula helped her become acquainted ae Mississippi and its mushrooms. ‘The courtesy and cooper- ation of the staffs of the Harrison Exper Feel Forest of the DeSoto National Forest and the Gulf Islands National Seashore in allowing A to collect in their respective areas is much appreciated. LITERATURE CITED Beeli, M. mere Fungi Goossensiani. IX. Genre Lepiota. Bull. Soc. Roy. Bot. Belg. 64: 206-222. ————.. 1936. Lepiota, Annularia. oe hii beet des C Tete du Sone II: 2948. ; eat Cle monogrta aphique des “Lépiotes” de’Europe. Doc. Mycol. 11(43): 1-77 ee R. 1950. Beitrag zur Kennt tnis ae aargausichen Pilzflora. Lepiota lutea (Withering) Godfrin Lepiota cretata. Mitth. Aargauischen Naturf. Ges. 23: 82.(complete reference not ere Heinemann, P. 1977. Leucoprinus (Agaricaceae). Flore illustrée des Champignons d’ Afrique 5: 87— 101. ears M. ee ee critiques sur quelques Champignons de la Région Lyonnaise. Bull. Soc. Mycol. Fr. 71 125 races ae 1978. ie Ril ace a Blatterpilze. 4th ed. Bnd. IIb/2 of H. Gams’ Kleine Krypto- gamenflora. Gustav Fischer, Stuttgart. 532 p Murrill, W. A. 1911. The Agaricaceae of tropical North America—II. Mycologia 3: 79-91. —__——.. 1972. Keys to the fleshy Basidiomycetes of Florida (compiled and edited by James Kimbrough, Department of Botany, Univ. of Florida). Inst. Food and eas a 199 pp. Pegler, D. N. 2. A revision of the genus Lepiota from Ceylon. Kew Bull. 27: Ridgway, R. 1912. Color staydards and color nomenclature. Washington, D.C. ie >. a can A. H., and N. S. Weber. 1980. The Mushroom Hunter’s Field Guide. The Wnierity of fichigan Press, Ann Arbor. 31 Smith, H. V. 1981. Some species of Leucocoprinus which grow in greenhouses. Mich. Bot. 20: 45— a a , : _ - - = a —— >- ——s ean a - —_— = s~- a —e — eye : 7 7 7 7 1; 7 —_— sy * as Contr. Univ. Mich. Herb. 15:311-313. 1982. A NEW SPECIES OF RHYNCHOSPORA (CYPERACEAE) FROM VENEZUELA Wm. Wayt Thomas! Division of Biological Sciences and Herbarium The University of Michigan Ann Arbor, Michigan 48109 According to Koyama’s treatment of the Rhynchosporeae for the Botany of the Guyana Highland (1972), over 86 species of Rhynchospora are known from Venezuela. Despite Koyama’s important contribution to our knowledge of Venezuelan Rhynchospora there still exist species to be described. Because of its capitate inflorescence, white spikelet scales, and lack of hypogynous bristles, Rhynchospora chimantensis at first appears to be a new species of section Dichromena. However, it has several features uncharacteristic of section Dichromena which are found in section Pseudocapitatae (Kukenthal, 1949): achenes |-2 per spikelet, basal spikelet scales smaller than those more distal, relatively loosely aggregated spikelets in fascicles, and achene surfaces which are only lightly rugulose. For these reasons, I did not include this species in my recent revision of Rhynchospora section Dichromena (Thomas, 1982) but consider it allied to section Pseudocapitatae. Rhynchospora chimantensis Thomas, sp. nov. Fig 21, Cespitose perennial; culms erect to arching, to 60 cm tall, 1.1-1.3 mm wide basally, 0.8-0.9 mm wide distally, obtusely triquetrous, glabrous, green; leaves —5, I-2 cauline, green to greyish-green; leaf blades 5-30 cm long, 1.5-3 mm wide, flat to conduplicate, lateral veins + prominent, sparsely villous or glabrous adaxially except for the villous midrib, villous abaxially; margins villous; leaf sheath 3—5 cm long, villous; inner band whitish, chartaceous, densely short- villous, its summit truncate to concave; inflorescence bracts 1-3, green to greyish-green, flat to conduplicate, midrib prominent, base whitish; basal bract longest, 4.2-6 cm long, 1.6-1.9 mm wide, linear, white along basal 3.5-6 mm and an extension along midrib, sparsely pilose, more densely so along adaxial midrib; inflorescence congested, globose, head-like, with 24-38 fascicled spike- lets with basal peduncles occasionally to 3 mm long; spiklets ovoid to lanceoloid, 4—7 mm long, 0.8-1.3 mm wide, with ca. 9-10 scales; scales ovate, ca. 5 mm long, ca. 1.8 mm wide, smaller towards base of spikelet, cymbiform, distal “3 carinate, white to pinkish, lightly and finely streaked with reddish brown basally; stamens 3: filaments 2.9-3.3 mm long; anthers 2—2.1 mm long; stigma and style 3.1— 3.6 mm long; stigma branches 2.3-2.6 mm long; achene lenticular, usually bi- laterally symmetrical, ellipsoid, yellowish or beige darkening to light reddish- brown at base, |.1-1.3 mm long, 0.7-0.9 mm wide; edges of style base confluent with margins of achene; style base persistent, triangular, 0.6—1 mm long, 0.6- 0.7 mm wide, brown; summit of achene horizontal; achene surface transversely rugulose, often indistinctly so; epidermal cells narrowly elliptic, often indistinct. Ipresent Address: Carnegie Museum of Natural History, Pittsburgh, Pennsylvania 15213. 311 312 FIG. 1. Rhynchospora chimantensis. A. Whole plant, culm 60 cm tall; B. Spikelet, 5 mm long; € Spikelet scale, 4mm long; D. Achene, 1.8 mm long (Steyermark & Wurdach 362 (NY)). Planta caespitosa, perennis; culmi erectiusculi vel arcuati, usque ad 60 cm alti; fohorum laminae 5-30 cm longae, adaxialiter glabrae vel villosae, abaxialiter villosae; laminarum costae villosae; inflorescentia laxe congesta, globosa; spiculae 24-38, ovate vel lanceolate, 4-7 mm longae, 0.8-1.3 mm latae, 1-2 achaeniis; achaenium lenticulare, ellipsoideum, 1.1- y 3mm longum, 0.7— 0.9 mm latum; achaenii testa leviter transverse rugulos VENEZUELA: BOLIVAR: Chimanta Massif, oa Section, scrub forest near Summit Camp, rare on dry sandy banks bordering river, heads gray-white, 1925 m, 2 Feb. 1955, Steyermark & Wurdack 362 (F, holotype; MO, NY isotypes). Although this species is known from only one collection, it is quite possible that it occurs on other tepui mountains as well as on Chimanta. R. chimantensis is characterized by its height, narrow leaves, green bracts with pilose midribs, the loosely congested head of spikelets, the ellipsoid achenes, and the long style 313 bases. Despite having seen only one collection (3 sheets), I feel that there is little possibility of this being simply an extreme variant of a previously described taxon. ACKNOWLEDGMENTS I would like to thank the curators of F, MO, fa NY for the loan of their specimens and Ms. Kathy eee for the line drawing she prepare LITERATURE CITED Koyama, T. 1972. Cyperaceae—Rhynchosporeae and Pia in ia Mz astute ed. The Botany of the Guyana Highland—Part 9. Mem. New York Bot. Gard. 23-89 Kukenthal, G. 1949. Vorarbeiten zu einer eRe der eave Me Bot. Jahrb. Syst. 75: 290-3 14. Thomas, W. W. 1982. The Systematics and Pollination of ee secuon Dichromena. Ph.D. Dissertation. University of Michigan. Ann Arbor, Michigan - wo la 9 @- 4¢ a ee a ee —_— a co — * aa _ - eee =? - Contr. Univ. Mich. Herb. 15:315-324. 1982. BOTRYCHIUM RUGULOSUM eaigabntris ned ee A NEWLY RECOGNIZED SPECIES OF EVERGREEN GRAPEFERN IN THE GREAT LAKES AREA OF NO AMERICA f lagner, Jr. and F. S. Wa Herbarium a Division of Biological cee ‘he University of Michigan Ann Arbor, Michigan 48109 The name Botrychium ternatum (Thunb.) Sw. was long applied to the ever- green grapeferns (Botrychium subg. Sceptridium) of North America. Not until Underwood's pioneering revision of 1898 did there arise any question as to whether true B. ternatum, an Asiatic species described originally in 1784 from Nagasaki, Japan, really occurred in North America at all. In his opinion, nothing known to him from the New World could be identified as B. ternatum. However, in the late 1950’s, Wagner (1959) encountered grapeferns in Michigan that seemed at the tme to be the same as, or closely related to, the Japanese species, and its name was once again taken up for certain American collections by \ various authors. Wherry (1961) referred to the plants in question as the ternate grapefern” and described it as “a relative of the Asiatic taxon.” Other authors, including ourselves, subsequently simply referred to it as B. fernatum without qualification. In connection with monographic research on Botrychiwm during the past several years, it has become evident that typical B. éernatum does not occur in North America, and that the plants that we referred to are distinct from that species in a number of characters. They constitute a new species, closely related to two much more common species that usually occur with it, the endemic American B. dissectum Spreng. and the circumboreal B. multifidum (Gmel.) Rupr. The purpose of this paper is to describe this new species and to compare it with species of near relationship. Botrychium rugulosum W. H. Wagner, sp. nov. Frons deltoidea, 3 (—4)-pinnata, pinnis usque ad apicem dlivisis, pinnulis ultimis angulatis, oblique rhomboideis, lanceolatis vel oblongis, 0.2—-0.5 cm latis, vivis convexis et rugulosis, marginibus grosse dentatis (raro subintegris). In habit resembling B. dissectum and B. multifidum with which it usually grows, its fronds emerging from the ground before the former and after the latter. Sterile blades deltoid (Fig. 1, 2) the stalk more or less the same length as the blade (shorter in sun forms, longer in shade forms), the stalk and blade together of mature, fertile plants averaging 8—16 cm (3 cm in sun to 30 in shade), the blade itself averaging 4-8 (2-16) cm long. Sterile blade 3-(2—4)-pinnate, divided to the pinna tips with regular reduction in symmetry. Lateral and basal pinnae ovate-deltoid, the pinnules rhomboidal, ovate, or oblong, usually strongly angled, 0.2—0.5 cm wide, the laminar surface in the living state convex above and more or less coarsely rugulose (Fig. 1, Wagner 1962, Pl. 1). Pinnule margins with nearly regular, somewhat rounded, wide teeth (except in rare 315 516 IG. 1. Frond of Botrychium rugulosum (Michigan: Gratiot Co., w. of Meridian Rd., Fall, 1981). Note rugulose and convex segments, indicated by arr« subentire forms) ue. 2). Lateral veins mainly somewhat spreading rather than ne: rly oe (Fig. 3). Chromosomes n = 45. SoA MIC HIGAN, Midland Co., Mills Twp., south side of Baker Road, sect. 29, R. On.’ I’. 16N. Common in more or less open, dry, sandy field with scattered trees and shrubs, east of oil well. 9 September 1959. W. H. Wagner 9099, M. Sargent, and R. L. McAdams. rae Botrychium multifidum (Gmel.) Rupr. f. dentatum R. Tryon (1939) is a syno- nym, based upon compact forms of B. ee from northern Wisconsin. COLLECTIONS EXAMINED CANADA: ONTARIO: CARLETON CO.: Merivale, Minshall s.n., no oe (DAO). MIDDLE- SEX CO.: Near London, Balkwill, J.A. s.n., 10 Oct 1893 (MICH.) MUSKOKA CO.: Georgian Be Ie Beausoleil 1, Starrett 1212, no date (DAO). NORTHUMBERLAND CO.: C seton, Macoun, J. s Aug 1860 (NY). COUNTY UNKNOWN: Lake region and Ontario, Macoun, J. s. 15 Aug | 1874 MTMG). JEBEC: CHAMBLY CO.: Chambly, no rae 25 Sep 1927 (MTMG). GATINEAU CO.: Ironside, Hull, Rodland- silo a no date (MTMG). RICHELIEU CO.: St-Joseph-de-Sorel, Marie- Victorin, F 28082, 23 Sep 1928 (GH). ST ea CO.: Hatley, Mousley, H. s.n., 1919 (US). Hi Sapo nae CO. poreree Terrill s.n., 23 Oct 1927 (MTMG). VAUDREIL CO.: Robert, C.S.U. no date (DAQ). UNIT ED ST ATES: CONNECTICUT: WINDHAM CO.:: Brooklyn, Merritt, A.O. s.n., Oct 1903 (MICH). MICHIGAN: BENZIE CO.: Sect 1, NW Shorter Lake, Hagenah, D.J. 6585, 12 May 1967 (BLH). CHIPPEWA CO.: Sugar I, Sect 33, Hiltunen, ].K. 2695, 4 Sep 1960 (MICH.) CLARE CoO.: Sect 16, 17, S side Rte 115, Wagner, W.H. et al. 63193, 14 Nov 1963 (MICH). GRAND TRAVERSE CO.: Sand Lake, Ate W.H. et al. 9389, 22 May 1961 (US). SE Traverse City, Sect 28, Wagner, WH. & FS. 9451, 5 Jul 1961 (MICH). SE Traverse C ity, unnamed lake, le See W.H. et al. 9316, 9319, 9322, | Me 1960 (all MICH). Sect 29, T 27 N, R9 W, Hagenah, DJ. 4516, oe ie ie 4614, 3 Jun 1961 (both BLH). ue mi ESE ‘Traverse City, beach small lake, Dieterle, J. é oe 1960 (MICH). GRATIOT CC ect 25, % mi S M-46, W of Meridian Rd, Wagner, W.H. oo 442, aaae 1961, Wagner, W.H. ye bee 12 Apr 1959, 9108, 20 Sep 1959 (all MICH). TOSCO CO.: oe l 23 Rigaud, FIG. 2. Sterile frond silhouettes of eastern North American Botrychium subg. eae in MICH). R = B. ee Cas (1) Monroe Co., 8670, (2) St. Clair Co., Hagenah H-S-T, (3 yeas nee 5594, (4) Mo e Co., 8598, (5) Coarse form, Monroe Co., 8594, (6) Sana Co, 70503; (7) Jus bea form, Moe Co., 8598, (8) finely cut form, Muskegon Co., Vos 9180. cine (Michigan): (1) hae Co., mea ee 11146, (2) Wexford 5 - 9314. D = B. ia (Indiana): (1) Jefferson Co., mith SDM-5-1, (2) the same, SDM-2C-1, (3) eae extreme the same, SDM-3A-B. (All fel fe nos, erie entence designated are W. Ww agner). N,R5 E, Wagner, WH. et al. 9329, 2 Oct 1960 (MIC H). JACKSON CO.: 4 mi ENE as & Seymour R Wagner, W.H. 73460, 13 Oct 1973 (MICH). KALKASKA CO.: Sect 14, 4 mi W Kalkaska, Hagenah, D.J. 4 4606, sun 1961 (BLH). KENT CO.: Lake Shore Dr | miS SontelGiy Ww agner, W.H. et al. 9205a, 23 Apr 1960 (BLH, MICH). LEELANAU CO.: NW Cook Lake, Sect 35, Hagenah, D.J. 6582, 12 May 1967 (BL H). LIVINGSTON CoO.:: ee Lime Lake, peer W.H. Z al. 62358, 26 Oct 1962 nee CH). S side S ee Lake, Wagner, W. H.8 27 Jun 1957 (MICH). MIDLAND CO.: ; bv Jefferson Rd near Shearer ee WHS a 2 F. 8999, 3 Jun 1959 (MICH). Sect 29, E of oil well, Wagner, W. H. et al. 9029, 11 Jun 1959, 9099, 9 Sep 1959,9258, 14 Jul 1960 (all Nate a WP oi) WSS IG. 3. Segment shapes and venation patterns of eastern North American Botrychium subg. Sceptridium (all MICH). R = B. rugulosum (Michigan): (1) Monroe Co., 8603, (2) Monroe Co., 9123, (3) Gratiot Co., 9/08, (4) Midland Co., 9099, (5) same as (1), (6) same as (4). M = B. multifidum (Michi- gan): (1) Midland Co., 9097, (2) same as (1), (3) Mecosta Co., 91/6, (4) St. Clair Co., 9053. I dissectum: (Michigan): (1) Monroe Co., 8596a, (2) Cheboygan Co., 8063a, (3) Midland Co., 9096, (4) (Canada, Ontario): Kent Co., 91/58. (All field nos. are W. Wagner.) W side Sect 15, Kawkawlin R, Hagenah, D.J. 4623, 10 Jun 1961 (BLH). MISSAUKEE CO.: Sect 22, T 23 N, R 7 W, Hagenah, DJ. 5027, 11 Aug 1962 (BLH). MONROE CO.: Dixon Rd, 2 mi NE Petersburg, Wagner, W.H. 8396b, 11 Jun 1957, 9067, 12 Jul 1959, Wagner, W.H. & BLASDELL, R. k. 8594, 16 Sep 1958 (all MICH). Grames Rd near ‘Tuttle Hill Rd, Wagner, W.H. 8598, 16 Sep 1958, Wagner, W.H. & McAdams 9126, 9 Oct 1959 (both MICH). Ida ‘Twp, Wagner, W.H. & Hauke, R.&S K. 8632, 2 Oct 1958, Wagner, W.H. & McAdams 9123, 9 Oct 1959 (both MICH). Ida Twp, E of US-23 near irrigauon ditch, Wagner, W.H. 8670, 14 Nov 1958, Wagner, W.H. & Blasdell, RF. 8649, 18 Oct 1958, Wagner, W.H. & Hagenah, Df. 8635, 4 Oct 1958, Wagner, W.H. & McAdams 9123, 9 Oct 1959, Wagner, W.H. etal. 8670, 14 Nov 1958, 9261, 4 Aug 1960, s.n., 2 Nov 1961 (all MICH). Plank Rd near Ostrander Rd, Wagner, W.H. & White, R.A. 9100-3, 16 Sep 1959 (MICH). Rutherford’s, Dixon Rd, Wagner, W.H. & Blasdell, RF. 8594, 16 Sep 1958, Wagner, W.H. & Sargent, M 9066, 5 Jul 1959 (both ICH). Sect 5, Hitchingham Rd, Blasdell, RF. 300, 12 Oct 1958 (MICH). W of Plank Rd, N of Ostrander Rd, Wagner, W.H. & Shetler, §. 8603, 19 Sep 1958 (MICH). MONTCALM CO.: Eof 131,58 of Mecosta Co, Wagner, W.H. et al. 9114, 26 Sep 1959 (MICH). MUSKEGON CO.: Gerber Scout Reservation, Wagner, W.H. 9249, 5 Jul 1960 (MICH). NW of Holton, Voss, E.G. 9180, 13 Sep 1959 (MICH). OGEMAW CO.: Sect 29, T 23.N, R23 E, Wagner, W.H. et al. 9326, 2 Oct 1960 (MICH). OSCODA CO.: Wagner Lake, Wagner, W.H. 70163, 30 Apr 1970 (MICH). Wagner Lake, Sect 13, Hagenah, D.J. 6039, 21 Apr 1963 (BLH). SAGINAW CO.: Fordney Rd, Wagner, W.H. et al. 9048, 21 Jun 1959 (MICH). Fordney Rd, N of Brady Rd, Wagner, W.H. et al. 8589, 15 Sep 1958, 9045-5, 21 Jun 1959 (both MICH). SAGINAW CO.: M-46, S of Hess Rd, Wagner, W.H, 70503, 4 Nov 1970 (MICH). Nehmer’s Woods, W of Saginaw, Case, F. s.n., 25 Sep 1958 (MICH). Rte 85, 5 mi S 319 Frankenmuth, Churchill, JA. s.n., 20 Aug 1956 (MSC). 1% mi E Burt, Wagner, W.H. et al. 8586, 15 Sep 1958 (MICH). 1/5 miS Gratiot Co line, Wagner, W.H. et al. 8683, 12 Apr 1959 (MICH). 234 mi N Chapin, Wagner, W.H. et al. 9104, 20 Sep 1959 (MICH). 6/10 mi N Birch Run Rd, W. agner, W.H, 8520, 22 Mar 1958 (MICH). ST CLAIR CO.: Algonac, Farwell, a 3552-5, 12 Oct 1913, 7260, 15 Oct 1924 (both BLH). E side Wales Rd, Wagner, W.H. et al. 9305, 23 Aug 1960 (MICH). N side Nolan Rd, Wagner, W.H. et al. 62010, 31 Mar 1962 (MICH). N side Rte 21, Wagner, W.H., 9055, 26 Jun 1959, 9311, 23 Aug 1960 (MICH). NE edge Algonac, Wagner, W.H. et al. 9163, 20 Dec 1959 (MICH). Nolan Rd, Hagenab, D.J. H-S-T, 11 Nov 1961 (MICH). TUSCOLA CO.: Swaffer & Bray Rds, Wagner, W.H. & Barnes 8616, 23 Sep 1958 (MICH). WASHTENAW CO.: Waterloo Rec Area, N of Cedar Lake, Wa agner, F.S. 81026, 12 May 1981 ee 1] mi NE Willis McKean Rd, Grether, D.F.s.n., 6 Moy fe (MICH). WAYNE CO.: Center Sect 30, Sumpter Twp, W ie W.H. & Blasdell, R.F. 8614, 22 Sep 1958, Wagner, W.H. et al. 9194a, 9194b, +30 Mar 1960 (all MICH). eee R Dr, junct Flat R Rock, I-94, Wagner, W.H. & Morzenti 9220, 20 May 1960, Wagner, W H. et al. 8640-5, 11 Oct 1958 (both MICH). Merriman Rd, near Pennsylvania Rd, W agner, W.H. 9347, 12 Mar 1961 (MICH). N side Oakville-Wally Rd, Pious W.H. 62095, Jun 1962 (MICH). Sect IL. off Willis Rd, Blasdell, R.F. ee oe 1958 (MICH). MI ESOTA: AIT KIN CO.: N side Rte 200, 5 mi E Cass Co, Wagner, W.H. & F.S. 793 16-1, 28 a 1978 (MICH V YORK: HE RKIMER CO.: Dutch Hill, 6 mi E Utica, Haberer, JV. 1901, Nov 1905 (US). MAISON CO.: Lenox, Oneida Lake, Haberer, JV. s.n., Oct 1906 (US). ST LAWRENCE CoO.: Parishville, St John, E.P. s.n., 11,21 Sep 1942 (US). 3 mi SW Parishville, St John, E.P. s.n., 28 Jul 1943 (US). 3 mi W ae St John, E.P. s.n., 2 Aug 1943 (MICH). VERMO A DOISON CO.: Base Cae Mt, Middlebury, Brainerd, EF s.n., 8 Sep 1899 (NY). Bristol, ae) me .n., 26 Sep 1878, Pringle, C.G. s.n., 26 Sep 1878 (both NY). Brot! Cobble Hill, Brainerd, E s.n., 12 Sep 1900 (MO). Mt Philo, Wagner, W.H. & Lellinger, D.B. 9084, 19 Aug 1959 (MICH). CAL EDONL A CO.: St Johnsbury, Harris Hill, Rooney, B.M. s.n., 99 Oct 1903 (US). St Johnsbury, Padduck Village Cemetery, ees B.M. s.n., 9 Oct 1902 (US). CHITTENDEN CO.: Hinesburgh, Hosford, F.H. s.n., 22 Sep 1880 (NY,US). FR. ee ESUIN CO.: Fletcher, Tryon, R.M. 4642, 14 Oct 1939 (MQ). I+ mi W Fletcher intersect, Wagner,W.H. & Lellinger, D.B. 9087, 19 Aug 1959 (MICH). ORANGE CO.:: eae Coney Denslow, H.M. s.n. “9 Sep 1910 (NY). ORLE: ANE CO.: 1%4mS eae Seymour re Wdorner. fd. 51.2 31 ed 1939 (NY). RUTLAND CO.: Rutland, Eccleston, W.W. 2 Sa0 18 3 (NY). WE eOn eo: Woodstock, Billings Hill, Kittredge, E.M. s.n. 4-23 Sep 1930 (MICH WISCONSIN: BAY FIELD CO.: Barnes Twp, Sandbar Lake, Peck, J.H. s.n., 21 Sep 1979 (UWL). Sand lake near Barnes, Tryon, R.M. 4100, 14 Jul 1938 (WIS). Sect 31. NW 4, T 45 N,R&8W Moran, R.C, 125752, 3 Sep 1980 (MIL). DOUGLAS CO.: Black Fox Lake, Somerville, M.F. 71, 20 Sep yes (WIS). C heney Lake, Somerville, M.F. 47, 10 Sep 1931 (WIS). Deer Print Lake, Somrerilles M.F. 20 Sep 19! 30 (WIS). alee Somerville, M.F. s.n., 1 Sep 1929 (MO). Gordon, 2nd beach sand lake, a eee M.F. s.n., 1 Sep 1929 (MICH). Lower eae Claire Lakes, Somerville, M.F. 55, 5 Sep 1932 (WIS). N of sand lake, Peck, A fe s.n., 17 Jul 1979 (UWL). Sand lake, Conklin & Somer 1180-83, 6 Sep 1938 (WIS), Fisher, M 79-81, 17 Jul 1979 (Pigeon Lake Field Station), Fuller, . 3926, 6 Aug 1930 (MIL), Somerville, MF. s.n., 25 Jul 1931 (WIS). Sand lake near pee Tryon, mail 4100, 14 Jul 1938 (WIS). Sand lake on 2nd beach, Somerville, M.F. s.n., Sep 1938 (MIN). Sand lake, Ist & 2nd beaches, Conklin & Somerville 109595a, 6 Sep 1938 (MIL), 1182 al 1187 (US). Smith Lake, bees M.F. 49, 21 Aug 1931 (WIS). 12 ai E Solon Springs, sand lake on county H, Tryon, R.M. & 4 Jul 1940 (MO). MARINETTE CO.: Near Peshtigo, Grassl, C.O. 7393, 2 Sep 1936 canteen, Pours GE CO.: N of Almond, ee JW. s.n., 24 Aug 1938 (WIS). WAUSHARA CO.: Sand lake, Sorenson 2660, 17 Jul 1962 (WIS). Various aspects of the systematic biology of B. ruguloswm (identified as B. ternatum) are dealt with by Wagner (1959, 1960a,b, 1961, 1962a). This is a very rare fern, in spite of the many localities recorded in the range map, Fig. 4. It seems to es most numerous in Vermont and Michigan, but this may be due in part to the intensive collecting in these two states. At the majority of places there are only one or a few individuals, occurring with related species which are much more common. It grows associated with other members of subgenus Sceptridium: in its northern occurrences mainly with B. multifidum and in its southern occur- rences with B. dissectum. In Michigan we deliberately set out to discover sites, and invariably we found either B. multifidum or B. dissectum first and then searched for B, rugulosum. We suspect that there are many localities yet to be discovered, especially in southern Ontario, the Upper Peninsula of Michigan, and northern Wisconsin. Figure 5 presents the results of thorough collecting in 24 places in south- 10 5 FIG. 5. Bar graph of incidence of 4 plants of B. rugulosum in 24 localties in - southeastern Michigan. Y axis = number of localities; X axis = no. of plants. (Single lo- 2 cality with 130 plants not shown). eastern Michigan from Midland County south to Monroe County, and including eight counties altogether. The bar graph gives the number of localities that produced progressively 1-5, 6—L0, and so on, plants. The best spot (Monroe Co., 2 mi. n. e. of Petersburg, W. Wagner 8594, (MICH)) yielded 130 specimens; it is not shown on the graph. As will be seen, however, most of the habitats gave far less. Indeed, 15 of them had ten or less individuals, 62.5 percent of the total, and 6 (25 percent) (shown in black in first bar) had only one! The survey, which was made over a three-year period (1957-1959), gave results which have since been repeatedly confirmed. This means that if a large stand of evergreen grapeferns of more common species is discovered, it should be carefully examined, for there 321 may be one or a few B. rugulosum mixed in. The median number per locality in our survey was seven, and the mean 17. Botrychium dissectum was found in all 24 localities, B. onezdense (Gilb.) House in 12, and B. multefidwm in 11. If our survey had been made in northern Michigan or Wisconsin, B. dissectum and B. oneidense would probably have been uncommon or absent, and B. multifidum would be invariably present The substratum in all localities is sandy or silty with more or less included black organic matter. The soil evidently ranges from circumneutral to acidic, more commonly the latter. In the southern portion of the range, B. Saint 1S found mostly in low swampy areas, especially in old second-growth grassy places and along paths. Past grazing evidently stimulates the formation of populations. Old apple orchards abandoned for twenty or more years provide productive sites, as do brushy old fields and second-growth upland woods. In Vermont and New York actively pastured open fields and meadows have proved to be ideal and in these places we have frequently been interrupted in our studies by the visits of curious cows. The grazing does not seem to have any serious effects on the plants, although some of them appear trampled and many are dwarfed. The most dwarfed and compact sun forms of B. rugulosum and other sceptridiums occur In open, exposed grassy areas around and near borders of sandy lakes as in northern Michigan (especially Grand Traverse Co.) and northern Wisconsin ee Co.), where its most common associate is B. multifidum. In the more northern localities for B. rugulosum the haircap moss, Poly- trichum, is usually abundant, associated with low shrubs such as Gaultheria pro- cumbens and Rubus hispidus. Other woody plants found with it are generally scattered or clumped, and include Acer rubrum, Betula papyrifera, Hamamelis virginiana, Populus tremuloides, Prunus serotina, Salix spp., Spiraea alba, anc Vaccinium angustifolium. Also found are such herbaceous genera as Antennaria, Fragaria, Hieracium, Lycopodium, Osmunda, Pteridium, and Solidago. In southern localities (as illustrated in Michigan in the region from Saginaw and St. Clair counties to Monroe Co.), grapeferns grow in richer sites as a rule, and the plants of B. rugulosum reach the largest size known, some of the sterile fronds attaining 25 or more cm in length (Wagner, 1959, pl. 9). Associated with the fern in these richer sites are such woody taxa as Acer rubrum, Cornus drummondu, C. racemosa, C. stolonifera, Corylus americana, Populus tremuloides, Sassafras albidum, Ulmus americana, and Vitis riparia. Among the herbs are species of Anemone, Aster, Desmodium, Equisetum, Fragaria, Solidago, and Viola. The spleenwort fern, Asplenium platyneuron, has in recent years become one of its most frequent associates. The actively grazed pastures that we studied in Franklin Co., Vermont, and in St. Lawrence Co., New York, contain much more freshly disturbed conditions than the foregoing. The botrychiums tend to be most common in the most open areas, where the cattle trample some and perhaps nibble some. Trees occur mainly at the edges in the less disturbed parts of the pastures. Where grazing is most intense, the plants of B. rugulosum may occur in almost lawn-like habitats. Clumped vegetation is produced by Dennstaedtia punctilobula, Pteridium aquilinum, Rubus idaeus, Comptonia peregrina, Spiraea alba, S. tomentosa, and J uniperus virginiana. Colonies of these species overtop the other vegetation which is made up of such genera as Achillea, Antennaria, Danthonia, Fragaria, Gnaphalium, Fieracium, ee Panicum, Plantago, Poa, Polygonum, Polytrichum, Prunella, Rumex, and V In eles oe B. rugulosum we often encounter not only one to three other species of evergreen grapeferns, but the moonworts, B. matricarufolium and B. simplex, the rattlesnake grapefern, B. virginianum, and the northern adder’s- tongue, Ophioglossum vulgatum var. pseudopodum. At the type locality for B. rugulosum there were seven species of Botrychium plus the Ophioglossum. The 22 seasonal sequence of leaf development in the four species of Great Lakes ever- green grapeferns is in the order (1) B. multifidum, (2) B. oneidense, (3) B. rugulosum, and (4) B. dissectum (Wagner 1960, 1961). Leaf development in all of these ferns is exceptionally slow, taking 3—4 months from the time of appearance of the young frond above ground in May or June and the maturation of the leaf and sporangia in September and October. The contrasts are striking in mixed populations in late June and July when the species differ widely in the extent of leaf development. Botrychium rugulosum new leaves average |.3-1.8 times as developed as those of B. dissectum during June and July. During the same period the young leaves of B. multifidum are approximately 2-4 times further developed than those of B. rugulosum. The same phenological differ- ences exist in respect to sporangial maturation in the fall, as measured by the changes in sporangial color from green to yellow to brown, although these changes are more variable than the early vernation changes. Once a field botanist becomes familiar with the characters of B. ruguloswm it is unlikely that he will confuse it with any of the three other members of subg. Sceptridium that occur in the Great Lakes area. A key to these is given below. Of all the species, B. rugulosum has the most slender appearance and a somewhat thinner texture. Although over-all frond size is extremely variable both within and between populations, B. rugulosum can be told immediately from B. dissectum because it lacks the long undivided terminal segment of the pinna tips (Figs. 2, 3). Allof the pinnules of B. ruguloswm tend to be smaller than those of B. dissectum and are usually only one- to two-thirds as large. In the wild the distinctive rugulose and convex pinnule surfaces of B. rugulosum are especially valuable in distinguishing it from B. dissectum. The bizarre dissected form of B. dissectum is connected by intermediates to the “normal” form, f. obliquum. Some of the inter- mediates can occasionally be confused with B. ruguloswm, unless the pinnule shapes are carefully examined. Even though they may be rather finely cut, the forms of B. dissectwm always show the outline of the long terminal segments. Those of B. rugulosum are not only evenly divided to the tip, but the tips tend to be blunter. Botrychium rugulosum is readily separated from B. multifidum by its smaller segments; their strongly angular, rather than rounded, outlines (Fig. 3); their rugulose, rather than smooth, surfaces (Fig. 1); and the marked differences in timing of vernation (Wagner, 1960a). Unfortunately the spores of all of these grapeferns are variable, and we have not yet been able to find differences de- pendable enough to use in routine identification. The true B. ternatum of Japan is a plant of wooded hillsides, pathways, and grassy terraces. It is more finely divided than B. rugulosum, and the ultimate divisions tend to be linear, linear-lanceolate, or lanceolate-ovate, with their sides more parallel. The segments are not only narrower, but their margins are beset with delicate, very narrow, pointed teeth, these of very different shape, asa rule, from those of B. rugulosum. The texture of B. ternatwm is generally thinner and more herbaceous than that of B. ruguloswm. Smith (1967) found in an analysis of their phenolic compounds that B. ternatum is more similar to B. japonicum (Prantl) Underw., and that B. ruguloswm is more similar to B. dissectum. B. ternatum differs from B. rugulosum, according to Smith, in presence or absence of six phenolic compounds. Of other New World sceptridiums that might be confused with B. ruguloswn only B. schaffneri Underw. of Mexico southward is likely to cause any problems. It is a plant of pine woods and pastures at high altitudes with pinna outlines more narrow and more sharply deltoid than in B. ruguloswm. The primary pinnules are more contracted basally, and the median and submedian pinna bases are more inequilateral, the basiscopic first pinnule becoming nearly sessile on the rachis, the acroscopic first pinnule remote from the rachis. The sterile segments of B. schaffneri are slightly more divided, the ultimate pinnules more distant, smaller, and commonly unequally bifid with the acroscopic lobe larger. 323 Chemical evidence found by Smith (1967) indicates closer relationship of B. schaffneri to other southern and tropical American taxa (B. lunarioides (Michx.) Sw. and B. jenmanii Underw.) than to any other botrychiums. DISCUSSION Botrychium rugulosum is unquestionably a distinct species, and the combina- tion of its geographical range, periodicity, blade cutting, segment shapes, laminar contours and marginal teeth set it apart from all other species of subg, Sceptridium. The populations of B. rugulosum are consistent in repeating the specific characters uniformly in over 70 localities in an area of the Great Lakes region that extends approximately 1000 miles (1600 km) from Minnesota in the west to Connecticut in the east. Most importantly, B. rugulosum grows intermixed with other sceptridiums in the same habitats. It exists side-by-side with B. dissectum and B. multifidum and sometimes B. oneidense as well. The distinctive characters of B. rugulosum are therefore not merely environmental modifica- tions. It is intriguing to speculate on the origin of this species. How is it possible for a species to be limited to an area that 15,000 years ago was completely covered by Wisconsin glaciation? We might also ask the same question about the rare and local little goblin fern, B. mormo W. Wagner, of northern Michigan, Wisconsin, and Minnesota (Wagner & Wagner, 1980). There are a number of possible hypotheses to explain these ranges. Among them we can propose ex- planations invoking (1) long-distance dispersal, (2) allopatric speciation, (3) relicts, and (4) migration. As to long-distance dispersal by spores, our original idea that B. rugulosum was actually the same as the Asiatic B. ternatum is now abandoned and an assump- tion that we had received this species in the New World from spores carried by prevailing winds from the west is now without support. We know of no place in the world today, other than the Great Lakes region, where there are plants that conform to B. ruguloswm. The second hypothesis involving de novo evolution of a new species following glaciation also seems questionable. It is difficult for us to imagine a new species of Botrychium based on a number of characters (Wagner 1960b) evolving as a result of temporarily localized allopatry independent of its relatives, and then quickly spread over such a wide range. The hypothesis that B. rugulosum may represent a relict that survived in non-glaciated areas does not seem unreasonable, but there is as yet scanty evi- dence of the role of refugia. The classical idea of nunataks, for example, has usually been contradicted by substantial geological data. On the other hand, B. rugulosum may have survived glaciation in, for example, the Driftless Area of Wisconsin, and when the glacier receded and the climate changed, the fern returned eastward and northward to its present range. To us, the hypothesis of migration southward with the glacial front is at present the most appealing. Recent phytogeographers are inclined to believe that our northern flora was in general shifted southward by glaciation and that, ideed, many if not most species were forced into the middle and southern states. Support for this hypothesis comes largely from pollen core data. If B. rugulosum had actually been an ancient northern species that migrated southward at the leading edge of the glacier and then later returned, this picture would not be different from that of numerous well known species, especially of trees It is interesting to speculate that B. rugulosum may have had something to do with the variation of the enigmatic grapefern, B. dissectum, and its origin. The northern Botrychiwm rugulosum might have been forced, during glaciation, into contact with the southern B. biternatum that today occurs from the Ohio Valley southward to the Gulf Coast. In some ways B. dissectum appears to be intermedi- ate between B. rugulosum and B. biternatum. Perhaps B. dissectum constitutes a 324 more or less stabilized hybrid swarm at the diploid level. Toward the north it 1s more similar to B. rugiwlosum and toward the south B. biternatum. However, even without f. dissectum, B. dissectum is considerably more variable than either of them. B. dissectum ft. obliquum would represent the “normal” form of the hybrid and the striking and unique f. dissectum with its curiously lacerate pinnules might represent the result of morphogenetic upsets due to genetic conflicts (cf. Wagner, 1962b). KEY TO GREAT LAKES BOTRYCHIUM SUBG. SCEPTRIDIUM I. Sterile pinnules deeply and conspicuously lacerate, the ultimate Sl linear or nar- rowly oblong, with mostly 1-2 veinlets. ge f. dissectum (in ee to B. d. f. abliquum) 1. Sterile pinnules dentate to entire, not lacerate, the ultimate segments wider, aay ovate, spatulate, or lanceolate, with numerous veinlets Terminal pinna divisions more or ne undivided, linear lanceolate to ovate; plants peouile more frequent south of 43° N. Plants of various habitats; aes pinna blades elongate, linear-lanceolate to lanceolate, acute; onions pl ae in winter tending to become bronzy; roots wan nantly dark and corky, av. 2.5 mm thick (dried) 1 cm from stem; new leaves a ing in early Ju . dissectum £. liga Plants of low, shaded forests and swamps; aise pinna blades ovate-lanceolate ovate, rounded; lamina exposed in winter remaining dull green; roots dominant ivory- to tan- colored, smooth, av. 2.0 mm thick (dried) 1 cm from stem; new leave appearing in middle May. B. ee ‘Terminal ae divisions regularly divided to the tips, plants becoming more frequent ta 5, a north oe 4. Pinnules rounded, larger, mostly 4-8 mm wide; the living lamina flat; pinnule eee ee finely crenate to entire; new leaves appearing in early May. B. multifidum 4. Pinnules angular, smaller, mostly 2-5 mm wide; the living lamina convex a rugulose; pinnule margins mostly coarsely dentate; new leaves appearing in lat ay. B ; nee ACKNOWLEDGMENTS We are grateful to the many individuals who helped us in the field, the herbarium curators who lent us specimens, and those persons who provided us relevant information about where these plants might occur. We thank David J. Bay and Joseph M. Beitel for their help with the illustrations, and we give special thanks to Kerry S. Walter for his aid in prov ee computer programs for collection data. W. R. Anderson translated the diagnosis into Latin. This research was carried out in large part under NSF Grant DEB 8005536 for Evolution and Rea: of Botrychiwm. LITERATURE CITED Smith, D. C. 1967. Chemical characters as a guide to the taxonomy of Botrychium. Doctoral thesis, University of Michigan, Ann Arbor. ‘Tryon, R. M., Us 1939. Notes on the ferns of Wisconsin. Amer. Fern Jour. 29: |-9, 1898. American ferns—1; the ternate species of Botrychium. Bull. Torrey Bot. Club 25: 521-54 1. Wagner, W. H., Jr. 1959. American iS aa ns resembling Botrychium ternatum: a preliminary aber Amer. Fern Jour. 49: 97-10 —————.. 1960a. Periodicity and Se eas in Botrychium subg. Sceptridium in the north- eastern United States. Bull. Torrey Bot. Club. 87: 303-325. 1960b. Evergreen grapeferns and the ee a eee categories as used in North els pteridophytes. Amer. Fern Jour. 50 me new on on the vernation ee “OF Botrychium dissectum and B, eet “Amer. Fern Jour. 51: 31-33. 32a. Plant cinpactnes st leaf produc tion in Botrychium multifidum “ssp. typicum” and “forma dentatum. mer. Fern Jour. 1-18. —————.. 196 2b. Irregular eee development i in hybrid ferns. Phytomorphology 12: 87-100. —————- and F Wagner. 1981. New species of moonworts, ae subg. Botrychium (€ »phioglossaceae) from North America. Amer. Fern 1: 20- Wherry, E. VT. 1961. The Fern Guide. Doubleday and Co., New York. Contr. Univ. Mich. Herb. 15:325-337. 1982. OBSERVATIONS ON FOUR SPECIES OF DELESSERIACEAE (RHODOPHYTA) FROM THE SOUTH SANDWICH ISLANDS, THE ANTARCTIC Michael J. Wynne Herbarium The University of Michigan Ann Arbor, Michigan 48109 A collection of sublittoral benthic marine algae from ae South Sandwich Islands in the South Atlantic was recently sent to me by Dr. M. A. Hoban of Texas A & M University. The collection was made by trawl on 30 Sept., 1978, at Zavodovski I (56°19.5 'S; 27°34.5'W), the northernmost island of the archi- pelago, aaa is the only active volcanic arc in Antarctic regions. These islands occupy an intermediate location between sub-Antarctic South Georgia and the maritime Antarctic region of the South Orkney Islands, and the terrestrial vege- tation is essentially Antarctic in character (Longton & Holdgate, 1979). The South Sandwich Islands lie in the Low Antarctic province of Wace (1965) and are situated south of the Antarctic Convergence. Although Kemp & Nelson (1931) reported that some brown and red algae were collected by trawling by a British expedition on board R.R.S. ‘Discovery’ and that the kelp Macrocystis pyrifera was conspicuously absent, the only other prior marine algal collection from these islands was that reported by Zinova (1966), who listed 8 species, none the same as those reported here Numerous specimens of 4 species of Delesseriaceae were contained in the present collections, and they are known from only a few records in the litera- ture.! These species, Delesseria salicifolia Reinsch, Microrhinus carnosus (Reinsch) Skottsb., Neuroglossum ligulatum (Reinsch) Skottsb., and Phycodrys austro- bee Skottsb., were all originally described from South Georgia, 470 km to the N Two of them, D. salicifolia and N. ligulatum, have been reported only from ny island. Moe and DeLaca (1976) reported Microrhinus carnosus from the South Shetland Is. and Phycodrys austro-georgica from the South Shetland Is. and the Antarctic Peninsula. The present paper provides new information about the vegetative and re- productive morphology of these species and discusses their taxonomic place- ment within the family Delesseriaceae. Evidence is presented demonstrating that Delesseria stephanocarpa (A. & E. S. Gepp) Skottsb., a species that has been re- corded north to 47° S latitude on the Argentine coast, should be included with D. salicifolia. Representative specimens of the four species have been deposited in the University of Michigan Herbarium (MICH). should be inserted that the records of all these species from the “Argentine littoral” sts es and ye coaatiaee (1972) are not from the Argentine mainland proper but from i Antarctic and sub-Antarctic islands. Thus, they do not represent new records except for that of Delesseria stephanocarpa 5 Kuhnemann (1969). 325 o20 Delesseria salicifolia Reinsch (Figs. 1-9). Syn.: Delesseria stephanocarpa (A. & KE. S. Gepp) Skottsb. Reinsch (1888) described D. salicifolia from S. Georgia and later (1890) provided illustrations to show a much branched Delesseria composed of blades 4—11 cm long and 11-16 mm wide bearing well developed opposite nerves and with special tetrasporophylls only 2-3 mm long that develop from the midrib. Reinsch pointed out that outwardly it resembled D. lyallii [= Cladodonta lyallii (Hook. f. & Harv.) Skottsb.] except in the character of the margin, but in regard to its production of special tetrasporophylls from the midrib it agreed with D. sanguinea. . Kylin and Skottsberg (1919) later re-recorded this species from S. Georgia and included a photograph of a small tetrasporophyll with serrate margins. They also stated that cystocarpic plants were not known in this species and called attention to the possibility that D. stephanocarpa could very well represent the female plant of this species. Subsequent authors (Skottsberg, 1923, 1941, 1953; Papentfuss, 1964) did not refer again to the possible connection. Delesseria stephanocarpa was described (as Hydrolapathum stephanocarpum) by Gepp & Gepp (1905, 1912) from the South Okney Is. The Gepps stated that their species was “most nearly related” to D. sanguinea but differed in having cystocarps decorated with simple outgrowths, forming a “wreath of appendages.” They referred to the blades of their alga as having conspicuous veins and having a width of 3—4 mm. Skottsberg (1941), who transferred it to Delesseria, said that it was re- lated to D. sanguinea and D. fuegiensis. Its distribution was extended to western Antarctica and the South Shetland Is. by Skottsberg (1941) and Moe and DeLaca (1976). It had also been recorded from the Argentine mainland at Puerto Deseado, at 47°45’ S lat. (KUhnemann, 1969). The material from Zavodovski I. includes both tetrasporic and cystocarpic specimens. Individual blades range from 4 mm to 8 or more mm in width; one older blade measured up to 18 mm in width. This range overlaps the values given for D. salicifolia and D. stephanocarpa. The margins of the blades are entire and slightly undulating. A well developed midrib is present. Lateral veins are simple and extend almost to the margins in mature blades. Blades are densely branched from the midrib. Wings (= alae) are monostromatic, except for the veins. Sections of midribs of mature blades showed rhizoidal cells intermingled with large cells (cf also Skottsberg, 1923), a structural arrangment distinguishing Delesseria from Apoglossum, in which a sheath of rhizoidal cells encircles a layer of large cells (Kylin, 1956). Tetrasporangia and procarps are produced on small special leaflets arising from the midrib. The tetrasporophylls are 1000-1200 wm long and 600-700 xm wide and have serrate margins (Figs. 7-9), agreeing with the description of D. salicifolia by Skottsberg (in Kylin & Skottsberg, 1919). Female leaflets are 800- 1000 wm long and 400-500 xm wide at the time that early post-fertilization stages are discernible. These female leaflets also have serrate margins, and small spine-like processes develop on the young pericarp (Figs. 1-4). The mature cystocarpic proliferations are 2-4 mm long, and numerous elaborate append- ages or spines cover the pericarp (Figs. 5 & 6). Comparably adorned cystocarps occur in two other delesseriacean species, namely, Polyneura hilliae (Grev.) Kyl., in which a mass of spines cover the pericarp (Kylin, 1924), and Zinovaea acantho- carpa Wynne, in which spiny membranous extensions course over the pericarp (Wynne, 1970). The most significant point concerning the close similarity between the female and tetrasporic bladelets is that they complement the vegetative similari- ties existing between D. salicifolia and D. stephanocarpa and necessitate the con- clusion that a single species is involved, a possibility first suggested by Kylin and — B20 Skottsberg (1919). Delesserta salicifolia appears to be distinct from the 9 other species of Delesseria listed by Papenfuss (1964) from the Antarctic and Sub- Antarctic. Most of these species are not well known, but two, D. fuegiensis Skottsb. and D. macloviana Skottsb., were recently studied in some detail (Mendoza, 1974). Cystocarps in both species are unornamented, and venation is less pro- nounced than in D. salicifolia. Cystocarps are also unornamented in D. lancifolia (J. D. Hook.) J. Ag. (Skottsberg, 1923) and in D. belayevi Zinova (Zinova, 1963). Microrhinus carnosus (Reinsch) Skottsb. Figs. 10-18. Microrhinus is a monotypic genus that was established by Skottsberg (1923) on the basis of Delesseria carnosa Reinsch (1888) described from S. Georgia. Skottsberg (in Kylin & Skottsberg, 1919) initially transferred Reinsch’s alga to Chauvinia Harv. (presently Chauviniella Papenfuss, 1956) but subsequently recognized it as a distinct genus, since its apex did not con form to that pattern typical of the Hypoglossum group of Kylin (1924), to which Chauviniella 1s as- signed. Other characteristics of Microrhinus include a polystromatic blade with pronounced midrib, production of female and tetrasporangial reproductive organs on small special bladelets, and the occurrence of these bladelets in clusters arising not only from the region of the midrib but also from the blade margin and randomly over the blade surface. As Skottsberg remarked, the manner in which both vegetative and reproductive proliferations sprout in clusters from the midrib, margins, and surface of the blades makes M. carnosus an easily recognizable alga. Simple lateral veins, extending almost to the margins, are visible in younger blades, but these veins are obscured in the thickened older blades. Reinsch (1890 Pl. VU, fig. 6, Pl. VIII) did not observe reproductive stages, and Kylin & Skottsberg (1919) did not discuss reproduction other than to indi- cate that female and tetrasporic thalli were collected, but later (1923) described female and tetrasporic stages upon establishing the genus. His illustrations in- cluded early stages of cystocarpic development. The several specimens in the present collection consist of female/cystocarpic and tetrasporic thalli. Examination of the apex of very small bladelets revealed a pattern of the Delesseria type (Figs. 16-18), in which intercalary cell divisions occur in the second-order cell rows rather than that of the Membranoptera type, in which intercalary cell divisions are absent. Skottsberg (1923) had referred to the apex as being the Delesseria type, although his figures showed no intercalary divisions in the second-order cell rows. He stressed its resemblance to another poly- stromatic alga, Pantoneura, and Kylin (1924) assigned Microrhinus to the Membranoptera group because of its resemblance to Pantoneura, one of the members of that group. The consequent transfer of Microrhinus from the Membranoptera group to the Delesseria group necessitates a re-examination of the genera within that group. It currently contains the following genera: Delesseria, Apoglossum, Marionella, Laingia, Pseudolaingia, Yamadaphycus,2, and Odontolaingia, which was i recently described by Mendoza (1976). Of these 7 genera only Yamadaphycus 2Whether Okamurina (Zinova, 1972) or Yamadaphycus (Mikami, 1973) is the correct name of this genus has become confused. Recently Mikami and Yoshida (in Yoshida, 1980) adopted the name Okamurina. However, because Zinova (1972) based her genus Okamurina on Laingia pacifica Yamada (Yamada, 1932) [actually Pseudophycodrys pacifica Yamada, 1930}, which had already been used as the type of Mikami’s genus Congregatocarpus (Mikami, 1971b), on the basis of Article 63 of the Inter- national Code of Biological Nomenclature (Stafleu et al., 1978) Okamurina must be regarded as nomenclaturally superflous and thus illegitimate. Zinova’s (1976) later move to rectify the situation is inadmissable. Therefore, the correct name of this genus is Yamadaphycus. mm 5,6,10-15 I-15. Figs. 1-9 Delesseria salicifolia; Figs. 10-15 Microrhinus carnosus. Figs. 1-4, youn ea ‘igs. 5 & 6 FIGS. 8 stages of developing cystocarps on special leaflets; F igs. 5 & 6, mature cystocarps on special leaflets; seems similar enough to warrant comparison. There are strong similarities be- tween Yamadaphycus and Microrhinus beyond the coincidental sharing of the epithet ‘carnosus.’ Both have coarse, polystromatic blades, and cystocarps and tetrasporangia are produced on special proliferations, usually in clusters that are scattered over the entire blade surface. New blades arise both from the midrib 329 and also from the blade margins. Midribs are present in both genera. Lateral venation is evident in both genera, although in Yamadaphycus it is “often incon- spicuous” (Mikami, 1973) and in Microrhinus not visible in older blades. One apparent difference is the arrangement of tetrasporangia on the special pro- liferations. Zinova (1972) described the tetrasporangia as occurring in rows on the sporophylls. Such a pattern, however, is not evident in Mikami's (1977, fig. 8) figure of a mature tetrasporophyll nor in a specimen of this species deposited in the University of Michigan Herbarium and determined by H. Mikami. Tetra- sporangia are randomly arranged on the tetrasporophylls of Microrhinus (Figs. 14 & 15). Skottsberg (1923) stated that in Microrhinus new sporophylls arise from the stipes of older ones and that this process can be repeated, resulting in a dense tuft of leaflets. In Yamadaphycus the tetrasporic proliferations radiate from a central core or base (Mikami, 197la; Zinova, 1972). The tetrasporophylls of Yamadaphycus measure 1.5-2.5 mm in length and 0.5-0.1 mm in width (Mikami, 1977), whereas the average size of those of Microrhinus was reported by Skotts- berg (1923) to be 4-5 by 3 mm. Much variation in size of tetrasporophylls (and cystocarpic proliferations), however, was observed in the Zavodovski I. material. Many tetrasporophylls are 1.5-2.5 mm long and about 1.2 mm wide, as Figs. 14 & 15 represent; but some tetrasporophylls are up to 5 mm long and 3 mm wide, agreeing with the average values given by Skottsberg. The overlap in size of tetrasporophylls between these two taxa is obvious. Cystocarpic proliferations demonstrate some differences. Although dimen- sions of the bladelets are similar, the apices are attenuate or mucronate in Yamadaphycus but rounded in Microrhinus. Furthermore, ostioles of Microrhinus are provided with the prominent curved beak (Figs. 10-12; Skottsberg, 1923), whereas those of Yamadaphycus are simple. A single cystocarp is borne per bladelet in Yamadaphycus; usually one but less commonly two cystocarps are borne per bladelet in Microrhinus (Fig. 12). Carposporangia are produced in chains in both genera. n summary, the present observations call for the transfer of Microrhinus from the Membranoptera group to the Delesseria group and point out the strong resemblance to Yamadaphycus carnosus. My interpretation is that these two taxa (one occurring in the South Atlantic and the other occurring in the northwestern North Pacific) are closely related but can be distinguished on features of the cystocarpic proliferations and the arrangement of tetrasporangia, 1.€., allegedly in distinct rows in Yamadaphycus but irregularly arranged in Microrhinus. In my opinion there is a strong likelihood that the differences cited warrant separating these taxa at only the specific level rather than at the generic level. Phycodrys austro-georgica Skottsb. Figs. 19-29. This distinctive species was established by Skottsberg (1923), who described and illustrated sori of tetrasporangia as developing in the swollen lateral nerves of blades and also in special proliferations that arise from the midrib. Skottsberg described but did not illustrate cystocarps. Justification for placing this species in Phycodrys came from its Phycodrys-type apical growth (Fig. 19; cf. fig. 5c in Skottsberg, 1923), in which intercalary divisions occur in the primary cell row and opposite secondary cell rows are developed equally rather than alternately, the monostromatic nature of the blades (except for the midrib and nerves), and the structure of the midrib. The blades of P. austro-georgica have a conspicuous midrib. The lateral veins are macroscopic and become branched in older blades. Blades have marginal dentations, which are outgrowths of the second-order cell rows. The unusual feature of this species is the dense proliferation of vegetative FIGS. 16-19. Figs. 16-18 Microrhinus carnosus, camera-lucida drawings of apical region of blades; 7 = cells derived from intercalary cell divisions in second-order cell rows; Fig. 19 Phycodrys austro-georgica, camera-lucida drawing of apical region, showing the result of an intercalary division 1) in the primary cell row (i branches from the midrib, or from the remaining rachis of denuded blades, resulting in a shaggy appearance of this alga. The only other species in the genus which has secondary vegetative blades arising from the primary blade surface is P. amchithensis (Wynne, 1970), but the degree of such branching is minimal in that species compared to that in P. austro-georgica. It is known that in older specimens of P. rubens (L.) Batt., the type of the genus, fertile leaflets (tetrasporic and cystocarpic) can develop from the midrib, lateral nerves, and lamina (Turner, 1808, pl 35; Kylin, 1923, fig. 43d), although vegetative blades do not arise from these areas. The production of vegetative blades would not seem to be such a departure as to exclude P. austro-georgica from this genus. Also, a counter- part situation can be cited in the genus Membranoptera, another delesseriacean genus with typically only marginal branching. Membranoptera dimorpha Gardn. 331 (Gardner, 1926; Abbott & Hollenberg, 1976) has abundant branching from both the blade margins and the midrib regions, and it continues to be accepted as a Membranoptera ( (Widdowson, 1974; Phinney, 1977; Hawkes et al., 1978). I con- clude that it is most reasonable to include this species, P. austro-georgica, within Phycodrys. Procarps are randomly scattered over the blade surface (Fig. 20), as is ty pical of the subfamily Nitophylloideae, and many cystocarps can develop on a given blade. The procarp conforms to the Phycodrys- -type (Kylin, 1924), which is char- acterized by a single carpogonial branch associated with two sterile-cell groups, one consisting of usually 4 to 6 cells and the other of 2 or 3 cells. The mature, unfertilized procarp of P. austro-georgica commonly has one sterile-cell group comesed of 4 cells and the other sterile-cell group composed of two cells (Figs. Y) mas 25). Tetrasporic plants produce tetrasporangia both in discrete swollen sori located at the ends of lateral nerves or blades (Fig. 27) or in small leaflets ar ising from the sides of the midrib (Fig. 26). Male plants produce spermatangia in leaflets also arising from the midrib (Fig. 28) and completely covering the sur- faces except for the margins. Cortical cells produce spermatangial mother cells, the latter cutting off the spermatangia (Fig. 29). The current status of species within the genus Phycodrys deserves some at- tention. Dawson (1962) compiled a “tentative working key” to the species of Phycodrys, which contained I4 spec ies, including + new ones. That list should be modified by the following points. Phycodrys bullata Gardn. is the basionym of Tokidadendron bullata (Wynne, 1970). A second species, P. ambigua Gardn., was also transferred to that genus but should be regarded as a synonym of the former species in that it represents a first-year blade of this perennial algae (unpubl. obs.). Phycodrys elegans Setch. & Gardn. is a member of the subfamily Delesserioideae and thus should be excluded from Phycodrys (unpubl. obs.). The correct name for P. sinwosa (Huds.) Kuetz. is P. gic (Dixon, 1964). Taxa that have been described subsequent to Dawson's key are P. cerrateae (Dawson ef al., 1964), P. isabellae (Norris & Wynne, 1968), P. Cee (Wynne, 1970), and P. polycarpa and P. pumila (Zinova, 1972). Taxa omitted from Dawson’s account are P. radicosa (Okam.) Yamada & Inagaki and P. rossica (Sinova) Zinova (Zinova, 1955). Although Dawson included P. fimbriata (de la Pyl.) Kyl. in his tally, Atlantic records of that species are commonly treated as P. rubens (Vaylor, 1962), while records of P. lags from the Pacific Ocean (such as Nagai, 1941; Zinova, 1955; Mikami, 1972) have more recently been recorded as P. riggti Gardn. (Perestenko, 1980). Likewise, Perestenko (1980) has pl aced P. serratiloba (Rupr.) Zinova (Zinova, 1965) within the synonymy of P. riggti. Therefore, a total of 17 species of Phycodrys are presently recognized. Neuroglossum ligulatum (Reinsch) Skottsb. Figs. 30-37. This species is placed in a Southern Hemisphere genus containing 3 species: N. binderianum Kuetz., the type, occurring in South Africa; N. multilobum, re- stricted to Tristan de Cunha in the South Atlantic (Baardseth, 1941), and N. ligulatum, hitherto known only from South Georgia. Wagner (1954) studied the type species in some detail. Neuroglossum ligulatum, on the other hand, is known only from the original description (Reinsch, 1888, 1890), from a few figures of details contributed by Skottsberg (1923), and from Zinova’s (1966) report of a cystocar ‘pic specimen from S. Georgia. A photograph now in MICH taken by Dr. W. Taylor of a specimen in Skottsberg’s collection at the University of Uppsala shows an alga collected by K. A. Andersson at Possession Bay (Stat. 17), S. Georgia, and identified by Skottsberg as N. ligulatum. The photograph shows many ligulate blades arising from the midribs of two eroded axes. Lateral nerves 25 stcy CAR tea cb4 cb2 Cc 2 a -——————__ 400ym 21-25,29 3mm 400pm 20 32-35 26-28 30,31 cb3—first, cp—c porting cell; stc}, stco—first and second group Of sterile cells, respectively; Figs. 26 & 27, tetra- dortion of a blade bearing scattered procarps an young cystocarps; Figs. 21-25, procarps: cb) ,cbo I P go p g Pp p l the midrib; Fig. 29, surface view of male bladelet, spermatangia being budded from spermatangial mother cells; Fig. 30, transection of a portion of a blade; Fig. midrib; Figs. 32-35, procarps (see above for explanation of symbols). are absent, as stated by Skottsberg (in Kylin & Skottsberg, 1919). Skottsberg had been able to compare his material with a collection of this species in the Ham- burg Museum. The material from Zavodoyski I. fully agrees with the photo- graph of the specimen identified as N. ligulatum by Skotutsberg and cited in Kylin and Skottsberg (1919). When Skottsberg (in Kylin & Skottsberg, 1919) transferred Reinsch’s Delesseria ligulata to this genus, he also asserted that Choreocolax delesseriae Reinsch (1890, pl. 15, figs. 4-6) was not a separate organism but in reality represented the early stages in the production of lateral blades from the region of the midrib of N. ligulatum. Skottsberg also disagreed with Reinsch’s statement that midrib proliferations are lacking in this alga, pointing out that Reinsch’s (1890, pl. VI) figure illustrated such proliferations. Zinova (1966) also referred to clusters of small leaves emerging from the midrib of older axes and larger blades. | concur with Skottsberg’s and Zinova’s characterization of this alga as having lateral blades commonly produced, either singly or in clusters, from the region of the midrib (Figs. 36 & 37). In fact, the occurrence of lateral blades as marginal outgrowths is relatively rare in the specimens at hand. Individual blades become lobed and proliferous along the margins, but the more significant branching system is based on the origin of blades from the midrib rather than from the margins of blades. Reinsch and Skottsberg referred to occasional blades becom- ing furcate, and I also observed some blades forking. he very different type of branching seen in N. ligulatwm from that observed in the type species (N. binderianum) (cf. Kiitzing, 1843; Wagner, 1954; Simons, 1976) raises the question whether these two species can be placed in the same genus. Specimens of N. ligulatum from Zavodovski I. are very robust, some reach- ing 50cm in length. Skottsberg (1923) referred to a specimen that was 36 fy 5 cm FIGS. 36 & 37. Neuroglossum ligulatum. Fig. 36, portion of a cystocarpic specimen; Fig. 37, portion of a tetrasporic specimen. 334 60 cm in length. Blades can be 10-12 (occasionally to 14) cm long and are usually l-2 cm wide but occasionally reach 3.5 cm in width. Lateral venation is absent. Skottsberg (1923) remarked that a single apical cell could be detected in the “almost microscopic proliferations” and that the leaflets had assumed marginal growth by the time they were a few mm long. Present observations revealed a single apical cell was present in proliferations (in the tufts arising from the midrib) less than 1 mm in length. But by the time they are | mm long marginal growth has become active, and the single apical cell is no longer evident by the time the proliferation is 2 mm long. Male, female/cystocarpic, and tetrasporangial specimens are represented in the collections from Zavodavski I. Examination of male blades showed that close to the margin primary cells cut off a cortical cell to either blade surface by periclinal divisions. Uhese cortical cells then undergo a series of anticlinal divi- sions such that each produces about 16 cells (a range of 8 to 20 cells) that function as spermatangial mother cells. One sees in sectioned, reproductively mature blades a tristromatic structure: a central layer of relatively large cells and numerous smaller flask-shaped spermatangial mother cells on either surface. The spermatangial mother cells cut off single spermatangia outwardly. On female blades the procarps are randomly scattered over the blade sur- faces. Each procarp consists of a single four-celled carpogonial branch and two sterile-cell groups, each consisting of one ov two cells (Figs. 32-35). This procarp pattern is the Myriogramme-type (Kylin, 1924). The scattered cystocarps are |.5— 2.0 mm in diam. Skottsberg (in Kylin & Skottsberg, 1919) referred to the cysto- carp of N. ligulatum as having a thick papilla or crown perforated by an ostiolar canal, which he indicated as characteristic of the genus. The cystocarp of this species, when observed in section, seems to be inter- mediate between the Schizoseris-type and the Gonimocolax-ty pe (Kylin, 1924), even though that of N. binderianum conforms to the Schizoseris-type (Kylin, 1924; Wagner, 1954). The carposporangia are ovoid to tear-shaped, reach 90 wm in length, and are arranged in chains. They are produced on elaborately branched gonimoblast filaments that arise from a much dissected fusion cell (of about 225-285 wm diam.). The fusion cell is connected by processes to numerous content-rich central cells on the floor of the cystocarp, which ts similar to the Gonimocolax-type of cystocarp. But the discrepancy with that type is the presence of a fusion cell in the cystocarp of N, ligulatm. The discrepancy with the Schizoseris-type cystocarp is the presence of the numerous content-rich primary cells and the connections of processes from the fusion cell with those primary cells as well as the fact that the gonimoblast filaments are not all completely converted into carposporangia, in contrast to the situation in N. binderianwm (Wagner, 1954). This difference in the organization of the cystocarp may be further evidence that N. ligulatum should not be placed in Neuwroglossum. Skottsberg (in Kylin & Skottsberg, 1919) described the tetrasporangial sori as being sharply delimited, oval, a few mm in diam., and developing between the midrib and the margin. I observed tetrasporangial sori also to be oval, about 0.8 by 1.2 mm, and first appearing near the margins of the lower half of the blade. In later stages the initially discrete sori often coalesce into irregularly linear extensions running lengthwide on the blade, to 1 cm long or more but only | mm broad. These sori generally cover the entire blade but leave a central clear zone, corresponding to the midrib, in the lower half of the blade. Neuroglossum, based upon N. binderianum, is most closely related to Platy- clinia, another genus of the Myriogramme group of Kylin with a Southern Hemisphere distribution. Both genera have polystromatic blades, which show a similar appearance in section: a gradual reduction in size from the largest medullary cells seen as a median row to the smaller medullary cells on either side of the central row finally to the smallest cells of the cortical layers, which ts also 335 true of the anatomy of N. ligulatum (Figs. 30 & 31; cf. also Reinsch, 1890, pl. VII, figs. 7 & 8; Skottsberg, 1923, fig. 19c). The two genera were separated by Kylin (1956) on the basis that the midrib is detectable only near the base in Platyc linia but continues into more distal portions of the blades in Neuroglossum. SUMMARY Four species of Delesseriaceae are recorded from the South Sandwich Islands in the Antarctic. Delessaria salicifolia is shown to include D. stephanocarpa on the basis of the morphological similarity of their vegetative habit and of their special female and tetrasporic leaflets. Microrhinus carnosus is transferred to the Delesseria group, and its strong resemblance to Yamadaphycus carnosus 1s pointed out. Procarps of Phycodrys austro-georgica and Neuroglossum ligulatum are figured. The former species conforms to the genus Phycodrys despite the unusual branch- ing pattern, whereas the latter species seems less closely related to the genus Neuroglossum. The production of clusters of vegetative leaflets from the midrib in N. ligulatum, which then develop into the indeterminate branch system, is unlike the branching pattern present in N. binderianum, the type of the genus. ACKNOWLEDGMENTS I wish to thank Dr. Michael A. Hoban for providing me with these collections made on the search vessel /slas Orcadas, while he was a Dauicipant on a joint cruise (no. 17-78, Stat. 29) of the Neurell Science Foundation and the Argentine Navy. Dr. Hoban’s participation in the cruise was made possible by NSF grants (DPP 76-80738 and DPP 78- 23463) to Dr. Greta Fryxell, and her assistance is also gratefully acknowledged. Iam also most grateful to Mr. Richard L. Moe and Dr. Max . Hommersand for their careful reading of the manuscript and offering much useful criticism. EVEBRATURE CILeD Abbott, I. A., & G. J. Hollenberg. ee Marine Algae of California. xii + 827 pp., 701 figs. Stanford University Press, Stanford, Ca aad oa E. 1941. The marine = of Tristan da Cunha. Results of the Norwegian Scientific xpedition to nace da Cunha, 1937-1938. No. 9. 174 pp., 75 figs. ee E. Y. 1962. Marine red algae of Pacific Mexico, Part 7. Ceramiales: Ceramiaceae, Delesseriaceae. Allan Hancock Pac. 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'T - masculine cae of Delesseria aCe Skottsberg and the occurrence of he genera Delesseria L amouroux and oe yserts Kylin as hodophyta) in Tierra del Fuego and Isla cre os Fie tados. Phy PS cec, iN 33: —504, 15 | 6, Odontlingi, ee tea fs la familia Duane (Rhodophyta). Bol. Soc. Ar pelle oe 17: 190-198, 9 fig Mikami, 197 1a. A new member of ee (Rhodophyta) from Hokkaido (preliminary secon. Bull. Jap. Soc. Phycol. 19: 5-8, 9 figs. 97 Ib. Congregatocarpus, a new genus of the Delesseriaceae (Rhodophyta). Bot. Mag., Tokyo 243-246, 9 figs. es On Phycodrys fimbriata (De la Pyl.) Kylin. Bull. Jap. Soc. Phycol. 20: 77-82, 11 figs. ———_.. 1973. ear us, a new genus of the Delesseriaceae (Rhodophyta). Phycologia 12: 139- 143, he 1977. on fully mature plants of aie carnosus (Rhodophyceae, Delesseriaceae) from Japan. Bull. aap, Soc. ae 25: 7-11, Moe, ‘i | be _ KE. DeLaca. 1976. Occurrence of pee ee algae along the Antarctic Peninsula. arctic i of US. 11: 20-24, figs. Nica M. 1941. 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Univ. of Michigan ae ou Arbor, Michigan. Turner, D. 1808. ol. 1, London, [2] + 164 + [2] pp., pls. Wace, N. M. 1965. ene ere In: Biogeo graphy and cre in en ae (J. van Mieghem & Re ee Oye, eds.), pp. 201-266, 13 figs., 8 tables. W. Junk, the Hague. Wagner, F. S. 1954. Contributions to the morphology of the Delesemerea Univ. Calif. Publ. Bot. 27: ae 346, 290 figs. Ww oe T. B. 1974. The marine algae of British Columbia and northern ep es revised list and keys. Part Il. Rhodophyceae (red algae). Syesis 7: 143-186, 8 figs., 2 tables Wynne, M. us 970. Marine algae of Amchitka Island (Aleutians Islands). I. Delesseriaceae. Syesis a 95-144, 92 figs. Yamada, Y. 1930. Notes on some Japanese algae I. J. Fac. Sci. Hokkaido Univ., ser. 5, 1: 27-36, 2 figs., pls. II-V ——————.. 1932. Nowe on some Japanese algae IIL. J. Fac. Sci. Hokkaido Univ., ser. 5, 1: 109-123, »» pls. I-XXV. +4 Sree - 1980. ae ae notes on some Japanese marine algae (3). J. Jap. Bot. 55: 310-314, fi m Zinova, D. 1955. [Classification of red hae of the northern seas of the U.S.S.R.}| Komarov Bot. Inst., Akad. Nauk SSSR, 220 pp., 169 figs. 337 Delesseriaceae anus ee pyaar et Macquarie. Trudy Bot. Inst. Akad. Nauk 3 one 65. Species fami liae melee eriacearum neg ae in parte septentrionali Oceani Pacifici. Novit. Sys tem. Plant. non V: ascul. 1965: 78-97, s. 1966. Novitates ae algis marinis Antarcticis ac iis ae Novit. System. Plant. non . 1963. Vascul. 1966: 103-1 97 ae familiae Delesseriaceae rear a iar in parte septentrionali Oceani Pacific. 2 ete Sys Plant. non Vascul. ¢ 976. Species \caane Deles saenta (Rhowtophya) in parte septentrionali Oceani Pacific. 3: Nout System. Plant. non Vascul. ed ee eee _—_ = _ > oe a os = i oe oe > a wee a A = a @e 1s CONTRIBUTIONS FROM THE UNIVERSITY OF MICHIGAN HERBARUM Vol. 9, No. I. 30 September 1966 La Vegetacion de Nueva Galicia Jerzy Rzedowski & Rogers McVaugh pp. 1-123, map Vol. 9, No. 2. 1 December 1969 Notes on the Distribution of West Indian Marine Algae Particularly in the Lesser Antilles Wm. Randolph Taylor pp. 127-203 ol. 9, Nos. 3-7. 30 March 1972 sence = eae in Neuva Galicia, Mexico from 1790 to the Presen ee McVaugh pp. 207-357, map Tae aa Mexicanarum Pugillus Rogers McVaugh pp. 361-484 North American Sy pee of Rie orientalis (Compositae) Rogers McVaugh & Christiane Ander pp. 487-493 The Genus Trigonospermum fess Cannone, Heliantheae) Rogers McVaugh & Chester W. Laskowski pp. 497-506 The Oaks (Quercus) Described by Nee (1801), and De ee & Bonpland (1809), with Comments on Related Spec h Cornelius H. Muller & Rogers McVaug pp- 509-522 Vol. 9, No. 8. 31 August 1972 Preliminary Studies on the Dothideales in Temperate North America Margaret E. Barr pp. 525-638 Index Volume 9, numbers 1-8. Errata. Title page. pp. 639-664 ol. 10 18 September 1973; corrected printing 10 January 1974 Mosses of the ae Lakes Forest Howard Cru pp. 1-404 [N.B.: A pies edition was ince perently siblshed’ in 1976.] Vol. 11, No. lI. 7 November 1973 New and Re eee Mexican Umbelliferae Mildred E. Mathias & Lincoln Constance pp. 1-24 Vol. 11, No. 2. 1 August 1975 Marasmius Section ee in the Northeastern United States Martina S. Gillian pp. 2540 The Taxonomy of ee (Malmrehiaeese) William R. Anders pp. 41-50 Notes on Bunsiorbact hein South-central Brazil William R. Anderson & Bronwen Gates pp. 51-55 Mexican Species on Pedicularis (Scrophulariaceae) Hitherto Confused we P. tripinnata Mart.& Gal. ers McVaugh & T. Lawrence Mellichamp pp. 57-63 Rene avery of Lobelia dielsiana Wim mee and a Related Species New to Science Rogers McVaugh & Michael J. Hu pp. 65-68 eee parry: (Boraginaceae) See with Heliotropium limbatum rs McVaugh & Audrey S. Delcourt pp. 69-71 A eee Sargassum from the Western Atlantic Wm. Randolph Taylor pp. 73-75 A Noteworthy Variant Caulerpa m. Randolph Taylor pp. 77-79 A New Species of Halimeda from Malaysia Wm. Randolph Taylor pp. 81-83 Notes on the DibuGen of Sphagnum tenellum Howard Crum pp. 85-87 Comments on Sphagnum capillaceum Howard Crum pp. 89-93 Belonia americana, ae ee and Robergea Richard C, Har pp. 95-96 Vol. 11 Sno: 1 August 1977 ener Results of the Sessé & Mocino Expedition (1787-1803) I. Summary of Excursions and Travels Rogers McVaugh pp. 97-195 Vol. 11, 4. 2 June 1978 ro Taxonomic ene of the Luzulae Group of Cyperus inda F. Den pp. 197-271 Vol. 11, No. 5. 21 November 1978 Two New Species of anaes from Western Mexico William R. Anderso pp. 273-276 Kosteletekya tubiflora (Malvaceae): A New Name based on a essé & Mocino Paintin Orland J. Blanchard, Jr., & Rogers McVau ugh pp. 277-280 Sphagnum richardsianum, a New Species from Mexico Howard Crum pp. 281-283 Calceolaria mexicana and_C. tripartita in Mexico Leslie R. Landrum & Rogers McVaugh pp. 284-288 Asclepias zacatecana, a New Species from Zacatecas, Mexico Rogers McVaugh pp. 289-290 Galeotti’s Botanical Work in Mexico: The Numbering of his Collections and a Brief Itinerary Rogers McVaugh pp. 291-297 A New Species of Pedicularis fon Jalisco, Mexico Rogers McVaugh & Suzanne Koptur pp. 298-300 A New Species of Arbutus (Ericaceae) from Western Mexico Rogers McVaugh & Thomas J. Rosatti pp. 301-304 Three New Species of Triwmfetta from Western Mexico . Wayt Thomas & Rogers McVaugh pp. 305-309 Vol. 12, Part I, No. 3. 12 March 1974 Flora Novo- Galiciana [Fagaceae] Bo McVaugh pp. 1-93 Vol. 31 May 1978 Beni Beachcombers and Explorers: Pioneers of the 19th Century in the as ere Lakes Edward G pp. 1-100 Vol. 14. 5 August 1980 The ae ae Researches of Charles Darwin and Joseph Hooker James Affolte pp. 1-10 Was a New Genus of Malpighiaceae from Colombia and Peru erson pp. 11-16 Notes on Mascagnia in ae America William R. Anders pp. 17-24 A Cara to ae Identiication of Mexican Sphagna How pp. 25-52 Comments on pian henryense Howard Crum pp. 53-56 Carowrightia mcvaughu: A Polymorphic Species from Southwestern Mexico Thomas F. Daniel pp. 57-60 The Genus Justicia (Acanthaceae) in the Chihuahuan Desert Thomas F. Daniel pp. 61-68 Two New Cucurbitaceae from Mexico Dieterle pp. 69-74 Additions to the Hypogeous Mycoflora of the Canary Islands and Madeira Robert Fogel pp. 75-82 A Stryphnodendron New to the Amazonian Flora Elena Maria Occhioni Martins pp. 83-84 Eight New Species of Ipomoea and Quamoclit from Mexico Gordon D. McPherson pp. 85-98 Botanical Results of the Sessé & Mocino Expedition (1787-1803) Is Icones Florae Mexicanae Rogers McVaugh pp. 99-140 Karwinski’s Itineraries in Mexico, 1827-1832 and 1841-1843 Rogers McVaugh pp. 141-152 The Taxonomy of Carex Section Stellulatae in North America North of Mexico A. A. Reznicek & P. W. Ball pp. 153-204 Notes on ene Algae ae the Tropical Atlantic Ocean—VIII Wm ph Taylo pp. 205-208 Boat adbie (Coates Ceramiales), a New Red Algal Genus from Alaska J pp. 209-220 ichael J. Wynne Beringiella (Rhodomelaceae, Ceramiales), a New Red Algal Genus from Al Michael J. Wynne pp. 221-229