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Revision of Stenostephanus (Acanthaceae) in Mexico PDF

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Preview Revision of Stenostephanus (Acanthaceae) in Mexico

Contr. Univ. Michigan Herb. 22: 47-93. 1999 OF STENOSTEPHANUS (ACANTHACEAE) REVISION MEXICO IN Thomas Daniel F. Department Botany of Academy California of Sciences Golden Gate Park San 94118 Francisco, California INTRODUCTION Stenostephanus Nees a neotropical genus of perennial herbs and shrubs is from comprising about 75 species that occur primarily at relatively high elevations Mexico The names apphed genus and current to Bolivia. history of to this its Wood were addressed by under Habracanthus) and Daniel delimitation (1988, (1995a). Most of the previously described Mexican species have been treated in Mexican Habracanthus or Hansteinia. In study, species previously either this all were and referred to Habracanthus, Hansteinia, and Stenostephanus studied are treated in the last genus. Combinations in Stenostephanus for most of the Central American and South American would belong genus have species that also in this be made. yet to This study based on examination of 460 herbarium specimens representing is 167 collections from 32 herbaria, field observations, and laboratory studies of Mexico. pollen and chromosomes. Fifteen species are recognized as occurring in In the only previous account of Mexican species oi Stenostephanus, Daniel (1995b) repeated herein treated nine species in Chiapas. Descriptions of those species are because, in most cases, they have been augmented substantially with additional from Chiapan graciUs) or collections data either additional collections (e.g., S, Mexico Four newly described from species are other states of silvaticus). (e.g., S. new proposed and two combinations Stenostephanus are for species herein in genera here congeneric with described in treated as it. INFRAFAMILIAL AFFINITIES Lindau treated most of the genera here recognized as constituting (1895) subsequent Lindau's Stenostephanus Isoglosseae subtribe Isoglossinae. in tribe placement of Syringidium in subtribe Porphyrocominae was hkely a mis- (1922) take on his part; the genus as described by him possesses the diagnostic characters of his Isoglossinae. Other genera of Isoglossinae treated by Lindau (1895) with World two stamens and monothecous anthers consisted of the Old genera Brachy- Bremekamp Nees and Oreacanthus Benth. retained Lindau's stephanus (1965) much-expanded Isoglossinae, renaming Rhytiglossinae, as a subtribe of his tribe it Bremekamp intended Stenostephanus (and doubtful that to retain Justicieae. It is however; he those taxa here considered congeneric with within his subtribe, it) noted Rhytiglossinae had bithecous anthers and consisted of genera confined that his World 47 CONTR. UNIVERSITY OF MICHIGAN HERBAI^IUM VOLUME 48 22 "lenticular with a pore in the centre of each of the flattened and marginal sides, a zone from He differing in structure the rest of the wall." suggested that genera (e.g., Brachystephanus) with pollen differing from those types he described for the would subtribes of Justicieae probably need be to referred to subtribes of their own. Subsequent observations of additional pollen by various workers reveal grains Bremckamp hke those described above by for his Rhytiglossinae in species of Steiiostephamis (see below) and Oreacanthiis (Scotland 1990), as well as diporate some pollen like that of Stenostephaiius in species of Brachystephanus (Figueiredo & Keith-Lucas Thus, would appear Brcmekamp's 1996). that circum- (1965) it scription of pollen types was too narrow and and that Stenostephaniis relatives its accommodated can be readily in subtribc Isoglossinae. Perhaps more relevant a question whether Stenostephaniis can be distinguished from is its relatives in the Other American subtribc. genera that are obviously related to Stenostephaniis, based on macromorphological and their palynological characteristics, include Razisca Oerst., Kalbreyeriella Lindau, and Cylmdrosolenhim Jndau. Lindau described 1 two subsequent the last to his infrafamilial classification of 1895. Without knowl- edge of palynological Lindau its characteristics, (1895) treated Razisea in tribe Odontonemeae Odontoneminae. The subtribe genus comprises four and species known from Central America and Colombia. androecium two monothe- is Its of cous stamens and diporate pollen its that similar to that of Stenostephaniis (see is below) confirm between The a close relationship these genera. genus usually is from distinguished those included here on in Stenostephaniis the basis of such characters as "spicate" (racemose defined below) its as inflorescence, gradually expanded corolla tube, and abruptly bent upper of the corolla (Gibson lip 1974; Wood Leonard These among 1958; 1988). features are encountered Mexican all species of Stenostephaniis, and doubtful that Razisea represents it is a distinct genus. Kalbreyeriella, which Lindau (1924) treated subtribe Porphyrocominae, in consists of three species occurring Colombia and Panama from in that differ Razisea primarily by their corollas with a rostrate tip in bud and a relatively long, narrow upper hp (Leonard Gibson 1958; 1974). Cylindrosokniiun a unispecific genus is from Ecuador and Peru. Lindau included and (1897) subtribe Isoglossinae in it concluded from that differed Stenostephaniis by having and looser fewer-flow- it ered inflorescences and corollas with a cylindrical tube and lingulate lobes. Addi- tional studies will be necessary to determine whether these three genera should continue to be treated as distinct from Stenostephaniis. Of the two paleotropical relatives of Stenostephaniis noted above, Brachystephanus was purported to diffcr by pollen and Oreacanthus was noted by and broad its to differ short corolla its tube (Lindau 1895). If these genera are to be maintained as distinct from Steno- stephaniis, other characters need be found will to to distinguish them. M Mexican stamens monothecous with which anthers, contain diporate pollen. MACROMORPHOLOGY Wood Me Stenostephaniis exhibit considerable morphological variation, particularly pubes- in may from cence, locality to This be due, locality. in part, to their rather isolated occurrences at high elevations (see below). Considerable variation pubescence in STENOSTEPHANUS DANIEL: 49 1999 of the rachis was observed in several Mexican species (e.g., S. glaber, S. haematodes, Wood monolophus, oaxacanus, and tacanensis). (1988) noted similar varia- S. S. S. among some Colombian chahtopes (Leonard) pubescence H. species tion in (e.g., Wood). R. J. I. Some appear be and aspects of inflorescence structure to relatively stable among Mexican most taxonomically useful characters Stenostephaniis. In species, reduced and from other leaves the distalmost pair of leaves in size differs in is which shape and attachment. In the following account, a terminal inflorescence (in leaves are replaced by highly reduced bracts) one that arises distal to this pair is proximal of leaves and axillary inflorescences arise in the axils of leaves to this many Most inflorescences are terminal and of those that superficially appear pair. These borne on to be axiUary are, in fact, also terminal. latter inflorescences are and subtended by reduced short branches in the leaf axils are a pair of leaves. Truly inflorescences are not subtended by a pair of reduced leaves. axillary Because of the presence of both bracts and bracteoles, the basic unit of the most dichasium considered be dichasium. Tn species the inflorescence to a is is several- to many-flowered; however, in some, comprises only a single flower. it Terms denoting the arrangement of dichasia are primarily dependent on the pres- and ence of peduncles stalks between the bract and the bracteoles) pedicels (i.e., known between the bracteoles and the flower). Tn dichasiate spikes (not stalks (i.e., among Mexican neither peduncles nor pedicels are present the species), (i.e., dichasia are sessile along the rachis). In dichasiate racemes, peduncles are absent and but pedicels are present. In dichasiate thyrses, peduncles are present pedicels may be present or absent. Panicles comprise branched inflorescences where the branches are indeterminate and consist of dichasiate spikes, racemes, or thyrses. from node Branching that results in a panicle often exclusively the basal of the is inflorescence in one or both axils of the distalmost pair of leaves). Inflores- (i.e., cences are usually pedunculate; however, either an indeterminate inflorescence if axis or a dichasium occurs in the axil of one or both of the distalmost pair of considered be leaves, then the inflorescence to sessile. is and pubescence from those Sterile inflorescences with calyces differing in size Wood Colombian noted by several species, of inflorescences, as (1988) in fertile were observed among Mexican However, such inflorescences might be not taxa. somewhat from which were sometimes difficult to distinguish juvenile ones, evi- dent proximal to the fertile inflorescences. more Various structures of Stenostephaniis tend to exhibit color than those of many other Mexican Acanthaceae; however, colors are not always preserved on herbarium specimens nor noted on Thus their usefulness for distinguishing labels. remains unknown. Color of the corolla generally taxa or assessing relationships is known and often useful taxonomic character. For example, appears to be a is it means monolophiis from Colors such the primary for distinguishing S. S. glaber. among maroon, and purple were noted trichomes (primarily the sep- as red, also calyces, and rachises of the inflorescence. Considerable variation in colora- tae), among was noted calyces of herbarium specimens. Tips of the calyx lobes are tion Sometimes often darker dried material than the proximal portion of the lobes. in darker coloration appears to be green and other times dark reddish or purplish. this The direction of dehiscence of the anthers was observed in most species to be some in the direction of the lower In species (e.g., S. latilabris) the direction of lip. dehiscence could not be determined with certainty from the herbarium specimens many appear available. In haematodes, collections have flowers with anthers that to S. CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 50 22 dehisce toward the upper and other collections have flowers with anthers that lip appear to dehisce toward the lower hp. Whether such variation the exists in species (or in the genus) or whether determination of the direction of dehiscence unreliable as viewed on herbarium specimens remains to be determined is Based on macromorphological characters, primarily features of the corolla, One there appear be two groups of Mexico. to at least species in consists of S. silvaticiis and latilabris, which both have relatively short, bluish to white corol- .S'. las with a well-developed and relatively long lower a short, cylindrical corolla lip; tube that hicks a distinct throat; a recurved to recoiled upper lip of the corolla; and stamens The that are inserted in the distal 1/3 of the corolla tube. remaining 13 species comprise a group characterized by having generally longer, reddish (at and known) least in part v/here corollas v/ith a poorly developed (or absent) to relatively short lower a mostly longer corolla tube that gradually or abruptly lip; is expanded distally into a distinct (usually saccate) throat; a straightforward to erect upper of the corolla; and stamens that are inserted the proximal lip in 1/3 to 1/2 of the corolla tube. This latter group readily divisible into those species with is racemose and known inflorescences those with thyrsoid inflorescences. not It is whether this inflorescence character actually segregates species into natural groups. For example, chiapensis and monolophiis, which greatly resemble each other S, S, in most features (including floral color and form), are distinguishable primarily by The this difference in inflorescence structure. ten species with thyrsoid inflores- cences are not easily distinguishable by large suites of mutually exclusive charac- ters. Their circumscriptions are sometimes confounded by lack of knowledge con- some cerning of their character states corolla color, capsule pubescence, (e.g., seed surface ornamentation). POLLEN Pollen of Mexican Stenostephanus was described by Daniel Examina- (1998). tion of additional pollen for the present study confirms and elaborates on those observations. Pollen of Mexican species was studied with a scanning electron all microscope. from which was examined Appendix Collections pollen are listed in 1. The palynological terminology used herein follows Walker and Doyle with (1975) modifications of Daniel (1998). Pollen that illustrates most of the variation encoun- among Mexican shown tered the species 1-3. in Figs. is Pollen of Steiiostephamis 2-porate and varies in shape from globose-elliptic is to globose-elongate (longer equatorial axis:shorter equatorial axis varies from The 1.00-3.33). longer equatorial axis the diameter of the apertural face in is all species except and haematodes, which diameter S. gracilis S. in the of the interap- sometimes ertural face longer. Outline of grains apertural view subcircular in is is and view from to circular, in interapertural varies narrowly elliptic to circular to it Using Walker and Doyle Mexican oblate. the size classes of pollen of (1975), Stenostephanus from medium-sized varies Stenostephanus to large. silvaticus (Fig. has the smallest grains ^im diameter apertural view) and 2f, (31 in gracilis g) S. ^m (Fig. le) has the largest (57 diameter in apertural view). The pores vary from 3-10 diameter and from gemmate jam in subpsilate to verrucate to to echinate in Each surface sculpturing. pore surrounded by a ± circular region 17-52 ^im is in The diameter. surface of the ± circular regions varies from subpsilate microver- to gemmae, rucate to microrugulate and, additionally, covered with baculae, and/ is The two ± or echinae. circular regions of a grain are separated from one another STENOSTEPHANUS DANIEL: 1999 51 FIG. 1. Pollen of Mexican Stenostephanus. a. S. ahishii {Ton 5340), interapertural view. b. S. breed- & Burns lovei {Breedlove 49644), interapertural view. chiapensis (Breedlove 72688), interapertural c. S. view. glaber {Daniel al 8405), interapertural view. gracilis {Croat 47536), interapertural d. S. et e. S. view. guerrerensis {Breedlove 61946), apertural view. guerrerensis {Breedlove 36060), intera- S. g. S. f. pertural view. h. S. harleyi {Reveal e( al. 4239), interapertural view. S. harleyi {Reveal et al. 4239), i. = apertural view. Scale bar 10 [im. by peripheral band 2-17 jim width. The band continuous and encircles the a in is most and some haematodes grain in species; in S, alushii (Fig. la) pollen of S. (Fig. and tacanensis either broken into segments of varying 3e, purpiisii, f), S. S, it is The lengths or does not completely encircle the grain. surface sculpturing of the A among band and sometimes peripheral varies species within a species. central row of gemmae, baculae, and/or echinae running the length of the band was observed in six species. among Mexican Daniel noted two forms of pollen the species of Steno- (1998) and Study stephanus, globose-elongate (e.g., Fig. Ig) globose-elhptic (e.g., Fig. la). of additional pollen reveals that these two forms based on shape are linked by globose-oblong pollen Fig. lb) found in several species. Because palynologi- (e.g., cal variation was observed in the widespread and morphologically diverse species « ' , 4 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 52 22 ') ^^^^^^^^R^y^^ ^b 1 JP o M^^^ Ap- A^j- ^^^^^^^^^^^^^^^^^^^k ^^^^^v^^ 1 ' \ # •'• ^^1 ^ g^^' ' ' ^' . . - f • • • >5 1 K^ \ L 1 1 '* ' b > * 1 * i ^ 1 .*' '* * ^ 'V • • • ^^^^^H^ m ^^_ . 'm %\ ^^^^H ^^^^^^^^pL%^^ - 'I r ^dJQ \^^ A ^ jnn A . QtoMrfHRiL*^'"'^vt^ai w.wfts,^ 1 ^P^_n \•- ^^F^EJ^v?J.^^^i^^^BC3V^VJ^ Wj«a^ssr >t.vt^.B^A"]Wn^|SIK^jF'!rhfMHc?•^*,^^^E,SS^t'^V.V.'..^'1'^^i'B-^i^>:V\ ^^^^^^^^^^^^H L^HB .^^^'' ^'^ W: fWT ^^H^KgIrfb^f^f^^^'^^-jTs^;v^tve^^^^^;';'-?** *^ M[ Slr^ 1 - V- w/Kr ^ ^ "-^^w !^^» il I^^^^^^^^^H ^I^E^ ^^^^^^H^^^K 1 ^^V .^ .^^^^^^^^^1 , 1>^^.^^^^^H ^^RqP^ ^Lj^ ^ ^n ^^mBh jB^^ « ^ ^ J^S^ r^^^^l^B ^^Bv^^^^^^^^E^^^^^H^^^^k_£^B~^Bi^K^^^^^H^^^^^^^^^^^^^H^HH^BIHvTv_ ^^^^^^^^^^^^^^^^^^^^^^^^^^1 f!^ ^^H |ifl|P l# ^^' r j^^^^^^^" "^^ ^fc "^^^^ ^^ \S?" ^^ ^ Ij^ ^E-- gK ^^\ !^fflA ^V^^in^'^ t^ J^iJI.r^ Y~i^I?^^ ^bC^ ''''# ^ U^^L . ' ; O ^^H^^^^P ^f^^ ^" ^^^"'^'^^a . ^F3 M^ &" ^" ^mc' * ^^^-i^^J- m^^K^.^ ^^a ^k]^ ^"W ii^ -^Ei^-"HiQj-A^^ai ^ ^^rj''-^J5 •^h Jj^^'.-^S^ -^B - ^^ Wi'-^-'^&^r ^''f'^'^W ^^^IK- ^* ^^i^H W^ l^^^fV '^f^t^^^K- _-"^^*fe^t,f-^'SJit»:^ lvfe!)_^'- ' ^^S' • Jr W--^L m . J :^^ M^^£ '1 ''" ^ T^^S^^^^^^'»^^ "^^SL'^r .JV^^ .Vi^^^^ 1 ^^^^^^^^^i^^^^^^^^SV^EkW^Ct^-wn^^^fH%c^•^r^B>^"mS<^TIp^t<^^^^^rJ^^''Wi^^Vc^^>^^B^^^^^t^^v~ivk^i^^k^Ui-^?r<^ni,^^^HH^^^*rJh^^T1^"^ja^SX^>r^SVv^kV [^ri-^-.^*-.K^^a-#SMfprL*_'.>k". '_.\-../:^r*'-a«^V- . ^y^m^^,^B>^m^c.*r^JB'Vd4t-^^-r^[B'4H-c'^SJi-ixfiir^f>J^^'^"^lL^->v^E.\^B¥^-K4^It»IrHu^^^^^^^^^^^^^^^^^^^^^^^^^^^l^B11 V}^ '^ ^^^^E^v~^i^^^k ^^J^P^HH^^UB^^£^-^&->;:, "'^^^SN^)^^^1^5Lbfij winT'+:'*\j"\rJ-x=. ^" ^ \ " tf^J ''^^JF:_r^^:v^^-^?/aF-^^ 1 r1i ^^^^^^^-- ^^^^^^BDCv^^^P^^^^^l^i L,:k-^3'j^ifxKll*WV'w-v].:^^:P^^^^k^^^ Tfc^^ML^ ^^ht^g *" ^ _w ^x-* KPnl^^WiBfcw^SL^,\^flwrJ .Vjr-.->LaBP^gj^ ^S:,; 1 ^^fe ^^^S^i^ ^^ !)^ B^^*^ i^S ^ ^V^ ^^» J ( FIG. Pollen of Mexican Sienoslephanus and Central American Razisea. madrensis (Cam- 2. a. 5. pos V. 4736), interaperlural view, fnonolophus {Matuda 3969)^ apcrtural view. monolophus b. 5. c. .S". {Matilda 3969), inlerapcrtural view. oaxacanus {Reko 3724), apcrtural view. oaxacanus d. S. e. S. & (Carlson 4022), inlerapertural view. f. S. sUvatinis (Breedlove Bartholomew 66948). g. S. silvaticus (Mexia 9273), interapertural view. h. S. tacaneusis (Nelson 3794), inlerapertural view. Razisea i. spicata (Daniel e( al. 6231), interapertural view. Scale: a-h, bar = 10 |im; bar = 12.5 i, Jj.m. haematodes, pollen of numerous individuals of was Some S. this species studied. of the palynological diversity observed within haematodes 5^. illustrated in Fig. 3 is and under discussed that species. is Mexican Pollen of Stenostephanus resembles described and/or that illustrated by Wasshausen (1985a, as Kalbreyemcanthus; 1984, 1985b, 1987a, 1987b, as Habra- Wood cantlius; 1985c, as Hansteinla), (1988, as Habracanthus), and Scotland (1990, as Among Habracanthus, Hansteinia, and Stenostephanus) South American for species. New other genera from the World, Razisea (Fig. 2i) and Kalbreyeriella (Scotland 1990) both have similar pollen. Pollen of the related genus Cylindrosolenium was described by Lindau (1897) as typical of Isoglossinae. Several genera from the Old World, including Brachystephanus and Oreacanthus, have also similar pollen Daniel (see 1998). , STENOSTEPHANUS DANIEL: 1999 53 T f ^ 1 . »^^ g^-'l^-^ m ii % % ^ h'^ 1 raMt^ jj M ' -s 4^ f^^^-^ > w ^^^^^^^^^^^s^ J w--- ^:^J^J^KtEm m;:;-I ^ ' l^^.pr* i^ 1* * '^ ^^^^^^l^ i#*^ ^^^^^^^f^ > J F I [ r -^ J 4- ^iyfi';^IO ^i ^^M-"'^ % w-^ f ' ^^^^^^^^^^^^ ^^^^^^^^^^^^^^^^^^^^^1 FIG. Pollen of Stenostephanus haematodes. Ventura A. 20491, interapertural view. Daniel 3. a. b. & Acosta C. 8359, interapertural view. Hinton al 10758, interapertural view. Ventura A. 3425, c. et d. apertural view. Ventura A. 4670, interapertural view. Ventura A. 4670, detail of surface showing e. f. sculpturing and terminus of peripheral band. Scale: a-e, bar = 10 ^m; bar = 2.2 \im. f, While additional sampling of Mexican species that are known from few col- lections necessary, several palynological characters appear to be useful for is still characterizing species and will likely be useful in phylogenetic studies of the genus. Various palynological attributes of each species are provided in both the species and Table descriptions 1. CHROMOSOME NUMBERS No chromosome counts have been reported previously for species here treated in Stenostephanus, During study, using methods noted by Daniel and Chuang this counts were obtained and haematodes (1993), for glaber as well as for the S. S. American known Central species currently Hansteinia blepharorhachis (Lindau) as Chromosomes Durkee. several were counted each taxon and counts in cells for all were n - The known chromosome number 18 (Fig. only in a related genus n 4). is - From which was reported for Razisea (Daniel the preceding 18, et 1990). al. discussion of intergeneric relationships, evident that Razisea might not be it is A common from chromosome number two distinct Stenostephanus, single to these chromosome taxa further supports their close relationship. Additional counts in Stenostephanus and relatives, especially the Old World genera Braehystepha- its nus and Oreacanthus, are desirable. AND FLOWERING DISTRIBUTION, HABITATS, TIMES Stenostephanus occurs from east-central Mexico (Hidalgo, Zf'OO'N) ca. lat. southward through Central America and northwestern South America Bolivia to The Andean 17°07'S). genus most species-rich South America, and (ca. lat. is in Colombia where The number distribution centered in 34 species occur. of its is species drops off dramatically in other countries: Mexico Venezuela (15), (8), 3 r HERBARIUM VOLUME CONTR. UNIVERSITY OF MICHIGAN 54 22 Mexico Tablc Palynological characteristics of species of Stenostcphaniis occurring in 1. (E = equatorial axis). Diameter of Diameter of OutHne of Width of Longer E: apertural face inleraperlural interapertural peripheral E shorter band (pm) face (|im) face (urn) broadly 41-48 40-44 3-5 alushii 1.06-1.20 elliptic to S. subcircular L66-L81 38-46 23-24 9-11 hreedlovei elliptic S. narrowly 46-56 20-24 13-17 2.11-2.77 chiapensis w/ S, elliptic broadly 46-50 38-42 globe 1.16-1.21 9 S. elliptic suboblate to 50-54 50-57 1.00-1.05 5. i^nicilis 11 circular narrowly 46-55 giicrrerensis 2.60 20 13 S. elliptic broadly elliptic to 35^6 39-47 2-10 Iiacnuitodes 1.03-1.23 S. subcircular to oblate 47-48 Iwrlcyi 1.46 32 elliptic 7 S. 36 31 subcircular 6 lalilabris 1.15 S. 1.10 44 40 subcircular 8 S. iihulrciisis C elliptic to L43-1.92 40-50 26-31 broadly 15-17 niofiolopliiis S. elliptic narrowly oaxaauuis 1.48-2.27 38-50 22-27 2-1 S. elliptic to 1 elliptic piirpusii 1.04 38 36 subcircular 7 S. narrowly 31-39 10-17 10-13 1-S5-3.33 S. silvciticus elliptic 5-7 50-52 44-49 1.03-1.12 subcircular tacanetisis 5. Costa Rica and Ecuador Guatemala Peru Panama and Bolivia (7), (6), (4), (3), Brazil and Nicaragua based on Glockeria glandiilosa Oerst., the identity (L?, (2), of which remains to hi^ confirmed). Species hkely occur in other mountainous countries of Central America Honduras) but have yet to be collected there. (i.e., Stenostephaniis represented in Central America by at least 13 species, most of is which Habracanthus are currently treated either or Hanstcinia. in DANIEL: STENOSTEPHANUS 1999 55 # # % 4 t i % % FIG. Camera chromosome lucida drawings of meiotic preparations, Hansteinia blepharorha- 4. a. & Almeda {Daniel ^ chis 6366), telophase I, /7 18. b. Stenostephanus glaber {Daniel e( ai 8405), & (chromosomes = telophase from only of shown), n haematodes {Daniel Acosta 1/2 cell 18. c. S. C. I - - 8359), late diakinesis n 18. Scale bar 10 j.im. 1, Most known from Species are only a single country; this distribution pattern reflects the high degree of local endemism characteristic of many species. Of the Mexican 15 species of Stenostephanus, ten are endemic to Mexico, three occur only Mexico and Guatemala, and two occur Mexico and Amer- in in other Central ican countries. Within Mexico, four species are endemic to Chiapas, two species known are only from Oaxaca, and two species occur only Guerrero. in Wood noted (1988) that species of Stenostephanus are largely restricted to wet cloud forests between 1700 and 3000 meters elevation. This generally true is Mexican which found from for species, are at elevations (400) 1000 to 2875 meters. Mexican All species occur at or above 1000 meters, and the presence of oaxaeanus S, known at 2875 meters elevation represents the highest occurrence of Acanthaceae in — —— — — VOLUME CONTR. UNIVERSITY OF MICHIGAN HERBARIUM 22 56 Mexico. Plants generally occur in vegetation types that vary from mesic to wet: temperate deciduous oak pine-oak pine-oak Liquidaiuhar forests, forests, forests, mesophytic montane montane and cloud Only forests, rain forests, forests. forests, haematodes, and can be found eleva- three species, S. gracilis, S. S, silvaticus, at tions below 1000 meters where they occur in tropical deciduous forests (apparently mon- rarely so and then always near water in these otherwise dry forests), lower montane The and mesophytic propensity of the majority tane rain forests, forests. of Mexican species to occur at relatively high elevations, particularly in relatively abundance isolated, cloud forest habitats, likely contributes to the of small-scale Andean endemism noted above and undoubtedly helps explain the largely charac- of the distribution of the genus. ter Wood swampy (1988) also noted that most species occur along streams or in ground and that plants found apart from such situations are sterile. Indeed, most Mexican collections of Stoiostephanns w^ere noted to have been collected from Of two near streams or springs. the species that studied in their native habitats, I one hacmatodcs) occurred along a stream and the other glaber) occurred {S, {S. on moist, level ground along a cloudswept ridge. Both were fertile. Although flowering among Mexican species of Stenostephanus has been recorded during every month of the year, most species have been collected in flower between October and March. This corresponds to the end of the wet season and a major Wood portion of the dry season Mexico. (1988) noted that species flower only in and propagated at irregular intervals for a short time. Plants oi S. chiapensis that I in a garden from seed of the type have failed to flower over a six-year period. Further studies of Stenostephanus might consider the possibility of gregarious New known among Acanthaceae from World) and flowering (not the the rela- between microhabitat and phenology. tionship floral TAXONOMY Stenostephanus Nees Martius, Fl (Jun) 1847. Type: StenostepJianus in bras, 91. 9: Nees. lobeliiformis DC, Habracanthiis Nees in A. Prodr. 11: 312. (Nov) 1847. Type: Habracan- Nees silvaticus [-Stenostephanus silvaticus (Nees) T. F. Daniel]. thiis DC, Galeottia Nees in A. Prodr. 11; 311. 1847, non Galeottia Rupr., 1842, nee DC, Nees Galeottia A. Rich., 1845. Glockeria in A. Prodr. 11: 728, 1847, non Nees Glockeria Gopp., 1836. Type: Galeottia gracilis [-Stenostepha- nus haematodes (Schltdl.) T. F. Daniel]. Dansk Kj0benhavn Hansteinia Oerst., Vidensk. Meddel. Naturhist. Foren. 1854: 142. 1855. Type: Hansteinia gracilis Oerst. {^Stenostephanus gracilis (Oerst.) T. F. Daniel]. & Gastranthus Moritz ex Benth. Hook, Gen. 1107. 1876, non pi. 2(2): f.. Gastranthus MuelL, 1868. Type: Gastranthus schlechtendahlii Moritz F. & Hook. Lindau ex Benth. [^Stenostephanus lasiostachyus Nees, fide f. The and author place of publication of G. schlechtendahlii are 1895]. Beatham Hooker, 1877)" by given as ''Moritz ex (in Ic. PI. 1210, t, Bremekamp The in Farr et (1979). generic description followed by a al. name Genera citation of the of the sole species in plantaruni fulfills the Nomen- Code requirements of Article 42 of the I/iternadofud of Botanical clature (Greuter et 1994) for a descriptio generico-specifica. al.

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