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AND PHYLOGENY, BIOGEOGRAPHY SYSTEMATICS, OF (MELASTOMATACEAE) CHAETOLEPIS Grimm and Almeda Daniel Frank Department of Botany Academy of Sciences California 55 Music Concourse Golden Gate Park Drive, dgrimmO @yahoo.com; [email protected] I RESUMEN INTRODUCTION montane shmbs and subshrubs genus of neotmpical mostly small Mi,. (Melastomalaceae) a is charac,enaed^4.merous,d.^^^ readily thatare species fatelyannuaU). consismofll As interpreted here, it welWeveloped pedoconnecuves^ lack stamens that «emonous isomVic inappendrcula.e mostly flowers, compre^flatened mostly or withatestathatconsists seeds MocuUr.aptcallyse, and cochleate use ovaries, Andean hasindudedspecresof South inthe past, circumscribed i»h"ligi.adngcefc. The genus,asithasbeen Cuba Recently the motrotypic genus and America, America, Central the Guayana Highlands, southern ''-ophlr.NaudimLmweltropicalAfricawas.ramferredtoCIwtdephO-que^Felix monophyly of the geographically dispa- Because the Study has attempted Chaetolepis or Nerophila. evaluate to seemed ripe for testing. been evaluated, it rate had never assemblage Chaetolepis of^taxa refelêd to 218 The primary goals of this study were to determine if Chaetolepis is monophyletic, assess the phylogenetic relationships of the c species based on morphology, and evaluate character evolution by using A cladistic techniques. L ttaa: xonomic revision is provided that includes detailed descriptions of all recognized n microscope images of seeds summary known for selected species, a of chromosome numoers, miormation on and habitat geographic distributions, flowering phenology, distribution maps, il- lustrations of all recognized species, citations of specimens examined, and a list of excluded taxa. This was supplemented by limited field study of four species in Costa Rica, Colombia, and Venezuela. — Tribal and Generic Affinities. Chaetolepis belongs to the tribe Melastomeae, the core mbersof^^ i are characterized by cochleate seeds with a testa that is tuberculate or papillate, a crown inenomes oi ur ap- pendages on the ovary apex, and the presence of staminal pedoconnectives (Renner and Meyer 2001; Freire- Fierro 2002; Michelangeli Based on et al. 2013). overall morphological Wurdack similarities, (1973) hypothesized that Chaetolepis most No is closely related to Tibouchina Aubl. cladistic analyses or molecular data, however, had been used to assess the possible sister relationships of Chaetolepis. Other neotropical relatives of Chaetolepis include Don, & Aciotis D. Arthrostemma Pav. ex D. Don, Heterocentron Hook Am., Microlicia D. Don, Microlepts (DC.) Monochaetum Miq., (DC.) Naudin, Nepsera Naudin, and Pterolepis (DC.) Miq., Rhexia based on L. a study using ndhF and cpDNA rpll6 sequences 24 members of of Melastomeae and outgroups (Renner and Meyer 2001). Based on an expanded DNA molecular sampling using sequence data, Bucquetia DC., Castratella Naudin, and Monochaetum now are considered the closest sister taxa of Chaetolepis (Peimeys etal. 2010; Michelangehetal. 2013). Like Chaetolepis, the majority of species in thesegenera are also restricted to higher elevation montane environments Andes in the of northwestern South America and have tetramer- all ousnêrs, an apicallysetulose ovary thatUfourlocuUrandfree tarn hypamhium, the and capsularfrulB. Morphology (Characters Taxonomic of Value).—The species of Chaetolepis are easily dis- tinguished from each other by characters involving the indument, anther connective, seed surface, petal coloration, and petal margin. The indument found on leaves, intemodes, and nodes, organs includes floral eglandular tnchomes, glandular trichomes, barbellate/dendritic trichomes, and The scales. distribution of the structures vanes greatly. ranges from It absent to densely covering the surfacesÎfTeeeLtS^lnTfl^^l organs. The anther connective 1^ prolongation ranges from absent to minutely nrolnno^ft t 1 i T dwTrwtb 1^^^^ prevailingly smooth with compressed/flattened imerdmatim'^ZttZ t Lk interdigitating The cells. petals vary in color fromyel- llooww iTalel ttoo pink oorr magenta anHd their margins are either glabrous or ciliate ainoris.ics.udi«(Su„dCl9MTribJuW^™'’'lTo^^^ Wurdack Almeda 1973; some ' Over of 2001). the past 80 years the soecies have been k a • .he placemen, of Leon and Alain MamnsT989) 1974; ciesa„d.h.eevade.ies. Because a, ^ identify symapomorphies and only cumorv w, aupmlon “ ' vegetative and ” reproductive llco^tunWiSr chamctem. monograph example of some Melastoma.aceae, of the °f for chatacters used to characieri varyingin color from rose .op„rple;4-Cw^t.ts“f"''“^^^^ smZr connectives; and ovoid cochleateLds With ofasmallersamplingofspecielTstiWbuX^^^^^^^ T and the seed surface appeam -smooth or dimta fit^ly and No any ^ ûriculate.” work mentioned sculptural prior details of the seed SFM testa because n ^ ,1 and consistency detail relating of to the seed and other mavlT'^ structures perhaps iolepis, biased by the four-merous flowers and ‘U greltirreduc^ greatly reduced some pedoconnective. Furthermore, 219 monophyly doubt about the of the genus has persisted because no cladistic analysis of the group has ever been and undertaken to evaluate character states their evolution. — Ingroup. Âll sixteen taxa thought to comprise Chaetolepis form the ingroup for this study. Chaetolepis alpina var. alpina, C. alpina var. latijolia, C. anisandra, C. cubensis, C. ciôdontisii, C. gentianoides, C. lindeniana, C. lori- carella, C. microphylla, C. perijensis var. perijensis, C. perijensis var. glandulosa, C. phelpsiae var. phelpsiae, C. C andC. phelpsiaevar. chimantensis, C. santamartensis, sessilis, thymifolia. Outgroup.—To monophyly among assess the of Chaetolepis, seven species the genera Tibouchina, Mi- and Monochaetum were used as the outgroup. The exemplars used were gleasoniana Wurdack, crolicia, T. T. Wurdack, Almeda and Naudin, M. Cogn. narinoensis wurdackii Todzia, Microlicia canastrensis obtusifolia T. ex Romero, Monochaetum amabile Almeda, and M. compactum Almeda. Tibouchina was chosen as part of R. common Wurdack most outgroup because hypothesized shares a recent ancestor the (1973) that likely it with Chaetolepis based on many shared characteristics, such as perigynous flowers, campanulate or urceolate and hypanthia, superior ovaries that are apically pubescent, capsular fruits, cochleate seeds. In addition, T. êasoniana, narinoensis, and wurdackii were chosen as part of the outgroup because the tuberculate seed T. T. The testa and foliar scales (see Todzia and Almeda 1991) are identical to those found in C. cubensis. species of Microlicia and Monochaetum were chosen other members of the outgroup because of their close phylogenetic as proximity to Tibouchina based on molecular studies by Renner and Meyer (2001). Material Examined.— Over 400 herbarium specimens, including types, were used in the phylogenetic analysis and taxonomic Specimens from the following herbaria with significant holdings of Central revision. DUKE, and on BM, BR, CAS, COL, DS, South American Melastomataceae were borrowed or studied site: F, UTMC images FMB, G, GH, HECASA, M, MO, NY, and (acronyms according to Thiers 2012). Internet K, US, P, on of type specimens OSTOR Plant Science) were examined for those taxa that were not readUy available loan. and The Characters.— and characters were used in the analysis (Tables 1 2). Forty-one vegetative floral was informative in this study. type of indument and structures particularly covering both vegetative floral Data Analysis.—The program PAUP* 4.0bl0 (Swofford 2002) was used to generate phylogenetic software matrix estimates. Maximum parsimony with branch and bound search was selected to evaluate the data that a Two were conducted. Analysis was generated in Maclade (Maddison and Maddison 2000). analyses 1 4.03 ingroup was provided insight into relation- performed and analysis 2 monophyly Chaetolepis to assess the of was prompted by the results of the ships cubensis) after the exclusion of one of the species of Chaetolepis (C. formed and 23 Sixteen species the taxa. first analysis. The analysis used 41 morphological characters first and formed Monochaetum, Microlica the presumed ingroup and seven species from the genera Tibouchina, was appUed backbone constraint that a outgroup. analysis Default options were used run the analysis. In 2, to unique and/or seed characters grouped because of their floral outgroup members together all plus C. cubensis & Mad- MacClade (Maddison grouped was done by constructing a tree in ingroup This all taxa together. loading the dison PAUP (Swofford 2002), derooting the tree, 2000) with opening in the data matrix, the file the search. constraint in the as a backbone and enforcing finally constraint in the analysis, generated, a bootstrap analysis phylogenies To various evaluate clades in the the strength of the different emp branch swapping was oyed; ot er- employing Stepwise-addition PAUP. a heuristic search was utilized in default options were used. . The among the lineages of Chaetolepis. “l MacClade was icters used to infer the evolutic ges,” and were chosen. “all possible changes” options mo conducted^ a™lys« were O Chartolepis, gain — biogeography of insight into the historical witnm taxa patterns of the diilerent peerrffoormr^mserdi j_ r^rioin aanndd ddiissopeerrssaall patl .. origin to provi; de insight into the MacClade, a data mai ^«nca. opir used Fitch Central America, and This analysis Africa. '°"P«ratingallChaetolepi glands sessile (2). 5 {1); and Almeda, Systematics of Chaetolepis Grimm PAUP with generated in in which they are found (Table and one of the 26 most parsimonious tree topologies 3), may how have Chaetolepis taxa was performed to infer polytomies resolved The second analysis arbitrarily. number only in the of area states (five) migrated within the Andes. This analysis differed from the first analysis were examined. (MPR’s) reconstructions used. In the analysis, equally most parsimonious all Chromosome H. Cytology Camoy’s (Bradley 1948) solution (4 chloro- modified Horal bud material was collected in the field, fixed in under 70% ethanol and stored refrig- form, 3 24 hours, transferred to ethanol, glacial aceitic acid, v/v/v) for 1 v Chaetolepis cufodontisii eration The new chromosome count reported here for until studied. andalOOxoil with phase contrast microscope acetocarmine smears mother using a Zeiss light of pollen cells The 2100x. magnification of made by camera lucida at a immersion were Drawings meiotic figures objective. of among specimens cited. voucher new marked with an asterisk collection count for this is on based dried her- characters and reproductive vegetative all :e an mature open flower in the center of one from either taken e numerous being recorded, were ot were characteristics r scored from matur^branches. To eLure that aberrant were specimens used. were examined and with typical traits those “actoljs morphobgy of . i A seed study the scanning efecmn (SEM) was used io microscope t«th gold-palladium and sputter-coated tape double-stick Seeds were aluminum stubs with affixed to laboramryatthe^^^^ microscope ^l»y.San,ples scanning electron were photographed the in SEM Voucher collectton data for the 20kV, microscope at “(Sciences using Leo 14M BP scanning electron a ‘“'“ges are provided legends. in the relevant figure RESULTS ^fogeny.-in 22 n the analysis, first cladistic was and the reter 0.66 224 which Cl, of nine are unique, three are homoplasious and one include ambiguous. The is unique characters ^te, appresse flattened scales, basally attached with a raised thickening near the point where the scale is at- ’ tached to the leal surface and an erose margin on the hypanthium base, hypanthium surface, apex, adaxial leaf abaxial primacy leaf surface veins, branch surface intemodes, branch surface nodes Lercalydne lobe sinus, alyxvesiuure excluding the midrib, and calyx midrib (characters 3, 6, 11, 17, 20 23 25 28 31 respectively). (char- m homoplasious ""oo* ‘>"= egUndular smooth glandular tri- trichomes, MUK/d S^mes or b Systematics of Chaetolepis Almeda, and Grimm 226 ffi