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Nord. J. Bot. - Section of tropical

Phylogeny and classification of the and Memecy laceae

Susanne S. Renner

Renner, S. S. 1993. Phylogeny and classification of the Melastomataceae and Memecy- laceae. - Nord. J. Bot. 13: 519-540. Copenhagen. ISSN 0107-055X.

A systematic analysis of the Melastomataceae, a pantropical family of about 4200- 4500 in c. 166 genera, and their traditional allies, the , with c. 430 species in six genera, suggests a phylogeny in which there are two major lineages in the Melastomataceae and a clearly distinct Memecylaceae. Melastomataceae have close affinities with and Lythraceae, while Memecylaceae seem closer to , all of which were considered as possible outgroups, but sister group relationships in this plexus could not be resolved. Based on an analysis of all morph- ological and anatomical characters useful for higher level grouping in the - taceae and Memecylaceae a cladistic analysis of the evolutionary relationships of the tribes of the Melastomataceae was performed, employing part of the ingroup as outgroup. Using 7 of the 21 characters scored for all genera, the maximum parsimony program PAUP in an exhaustive search found four 8-step with a consistency index of 0.86. Because of the limited number of characters used and the uncertain of some of the tribes, however, all presented phylogenetic hypotheses are weak. A synapomorphy of the Memecylaceae is the presence of a dorsal terpenoid-producing connective gland, a synapomorphy of the Melastomataceae is the perfectly acrodro- mous venation. Within the Melastomataceae, a basal monophyletic group consists of the Kibessioideae (Prernandra) characterized by fiber tracheids, radially and axially included phloem, and median-parietal placentation (placentas along the mid-veins of the locule walls). A second clade, the Melastomatoideae, with libriform fibers and lacking included phloem, and with axile-basal or axillary placentation, comprises the remainder of Melastomataceae and consists of eight reasonably well differentiated tribes, one of them paleotropical (Astronieae), three pantropical (Sonerileae, Melasto- meae, and Miconieae), and four neotropical (Merianieae, Microlicieae, Rhexieae, and Blakeeae). The Astronieae have several plesiomorphic characters, such as anthers opening by two slits and axile-basal placentation; all other clades have primarily poricidal anthers’ and consistently axillary placentas. A new subfamilial and tribal classification based on the analysis includes an annotated list of all currently recog- nized genera with numbers of species, distribution and important synonyms. Signi- ficant features of the classification include: the placement of Pternandra in a subfamily of its own and the dissolution of the Astronioideae; the merger of three neotropical and paleotropical tribal pairs; and the placement of in a tribe of its own. Biogeo- graphy, fossil history, and hypothesized phylogeny of the Melastomataceae imply an east Gondwanian origin of the family.

S. S. Renner, Dept of Systematic Botany, Llniv. of Aarhus, Nordlandsvej 68, DK-8240 Risskov, Denmark. Present address: Institute for Sysiemaric Boiany, Benizel-Weg 2, 55 099 Mainz, Germany.

Accepted 4-3-1993 @ NORDIC JOURNAL OF BOTANY NORD. J. BOT 13: 519-540

Nord. I. Bot. 13 (5) (1993) 519 groups can go in the Melastomataceae or Memecylaceae. Introduction and intraordinal Bremer’s (1988) suggestion that Melastomataceae are paraphyletic if the Crypteroniaceae and Memecylaceae relationships are excluded is not supported by my analysis because The Melastomataceae Jussieu are a large family of c. 166 there is no derived character which would support such a genera and 420&4500 species. Although distributed pan- clade and because in many attributes Memecylaceae ap- tropically, the family has a marked concentration of spe- pear more basal in the than Melastomataceae. cies in the New World where there are c. 2950 species. Hypotheses on Memecylaceae sister group relationships About 1275-1550 occur in the Old World, depending on are discussed below in the section on Memecylaceae DC. the estimates for the two largest genera, and Arguments for a close relationship between Cryptero- . Melastomataceae are easily recognized among niaceae, Lythraceae, and Melastomataceae are their cap- dicots by having with a characteristic acrodromous sular (replaced by in derived clades of venation ( Hickey 1973) and numerous small, exal- Melastomataceae and Lythraceae), morphology buminous seeds. Melastome are likewise charac- (Pate1 et al. 1984; Nowicke, pers. comm. 1991; S. Gra- teristic of the family. They have two locules at maturity ham, in litt. 1992), similar wood (van Tieghem (except in Rhexia), often a non-fibrous endothecium and 1891a, b; Vliet et al. 1981), and the often opposite leaves poricidal dehiscence, and their connectives lack dorsal with marginal nerves. These arguments do not appear glands. Excellent complete descriptions of the family strong, and I have therefore considered it best to develop may be found in Gleason (1932), Wurdack & Kral hypotheses on ancestral character states based on evi- (1982), Whiffin (1990), and Watson & Dallwitz (1991). dence from various sources (similar to the approach taken Such is the morphological distinctness of the Melasto- by Johnson & Briggs (1984: 709)) and to root the cla- mataceae that there has been very little controversy over dogram using part of the ingroup as outgroup, rather than their status and circumscription, except for the treatment base my analysis on an uncertain sister group. Neither the of Mernecylon and and their segregates and Axi- Crypteroniaceae nor the Lythraceae (as defined by Gra- nandra and Dactylocladus. The former two have vari- ham et al., in press) are distinguished from other myr- ously been regarded as a separate family, Memecylaceae, talean families by known autapomorphies, and recent or included as a subfamily in the Melastomataceae. Be- cladistic analyses of the Lythraceae based on morphology low I argue for the recognition of two families. (Graham et al., in press) and molecular data (Graham, Axinandra comprises four Asiatic species of trees with pers. comm.) moreover indicate that Onagraceae too are glabrous penninerved leaves and few-seeded capsules; very close to Lythraceae. Clearly, however, Crypteronia- closely related is Dactylocladus stenostachys, likewise ceae (and Alzateaceae and Rhynchocalycaceae), Melas- from southeast . Axinandra has been seen as lythra- tomataceae, Memecylaceae, and Myrtaceae will have to ceous (e.g., Bentham & Hooker 1867; Lourteig 1965), be appropriately broadly sampled before this plexus can melastomaceous (Triana 1871 ; Baillon 1877; Cogniaux be disentangled. 1891; Krasser 1893; Gilg 1897; Bakhuizen 1946/1947), intermediate (Meijer 1972), or as belonging in a separate family together with another Asiatic , Crypteronia (four species). Apparently Axinandra, Dactylocladus, History of the classification of the and Crypteronia share only plesiomorphic characters, such as brochidodromous venation, racemose inflores- Melastomataceae and Memecylaceae cences, and stamens with lengthwise dehiscent anthers, The history of the classification of the Melastomataceae and the heterogeneity of the group has been stressed by and Memecylaceae, compared to that of groups of com- Johnson & Briggs (1984) and Tobe & Raven (1987a, b). parable size, is uncomplicated (Tab. 1). However, since Leaves in Crypteronia, but not Axinandra and Dactylo- internal melastome classification has not been reviewed cladus, have stipules, and mainly on account of this but recently, a summary is necessary to understand more also because of its Crypteronia has rarely recent developments. been included in the Melastomataceae (Hallier 1918; Jussieu (1789) was the first to recognize the Melasto- Vliet 1981). More often it has been placed in Lythraceae mataceae as a natural unit, but it was David Don (1823) (Bentham & Hooker 1867; Hallier 1911; Lourteig 1965) who put structure into the family, using mainly placenta- or a separate family (A. De Candolle 1868), to which tion and seed characters. Don excluded and were sometimes added the monotypic South African Mouriri from the Melastomataceae because they differ and the small Central and South American from the rest of the family in precisely these characters. (A. De Candolle 1868; Beusekom-Osinga & Memecylon (placed in the Onagraceae by Jussieu!) was Beusekom 1975). More recently, Johnson & Briggs first included in the Melastomataceae by Du Petit- (1984) and Graham (1984) have argued for recognizing Thouars (1811), and Mouriri was added by Brown Rhynchocalyx and Alzatea at the family level and for ( 18 18). Shortly thereafter Blume (1826) again excluded excluding Axinandra, Dactylocladus, and Crypteronia Memecylon from the Melastomataceae, preferring to treat from the Lythraceae. Below it is shown that none of these it as a Myrtaceae, while De Candolle (1828a, b) created a

520 Nord. I. Bot. 13 (5) (1993) Tab. 1. Summary of the classifications of Melastomataceae and Memecylaceae. Triana’s subfamilies were formalized by Hooker in Bentham & Hooker (1 867). Cogniaux’s and Krasser’s Axinandreae comprise only Axinandra, Vliet et al.’s Crypteronieae comprise Axinandra, Dactylocladus, and Cypteronia. Cassebeerieae = Sonerileae, Osbeckieae = Melastomeae, Pleromeae = Tibouchineae, Pternandreae = Kibessieae, Pyxidantheae = Blakeeae, Tamoneae = Miconieae.

Triana 1871 Cogniaux 1891 Krasser 1893 Vliet et al. 1981 this paper

Melastomatoideae Melastomatoideae Melastomatoideae Melastomatoideae Melastomatoideae Microlicieae Microlicieae Tibouchineae Tibouchineae Astronieae Pleromeae Tibouchineae Osbeckieae Sonerileae Sonerileae Osbeckieae Osbeckieae Rhexieae Rhexieae Merianieae Rhexieae Rhexieae Microlicieae Osbeckieae Rhexieae Merianieae Merianieae Merianieae Microlicieae Melastomeae Ox ysporeae Oxy sporeae Oxy sporeae Miconieae Microlicieae Sonerileae Sonerileae Bertolonieae Merianieae Miconieae Bertolonieae Bertolonieae Cassebeerieae Dissochaeteae Blakeeae Dissochaeteae Dissochaeteae Dissocheteae Blakeeae Miconieae Miconieae Tamoneae Astronieae Py xidantheae Blakeeae Blakeeae Kibessioideae Kibessieae Astronioideae Astronioideae Astronioideae Astronieae Astronieae Astronieae Kibessieae

Memecyloideae Memecyloideae Memecyloideae Memecyloideae Memecylaceae Mouririeae Memecyleae Memecyleae Pternandreae Axinandreae Axinandreae Memecyleae Cypteronioideae Crypteroniaceae Cry pteronieae

family for the two. Chamisso (1836) once more included 1983, 1988; Takhtajan 1983; Cronquist 1988; Thorne them in the Melastomataceae, as did Lindley (1836, 1992). Triana, a native Colombian with extensive knowl- 1846), while others (Gardner 1840; Janssonius 1950; edge of the family in the field, worked in London and Favarger 1952, 1962; Lowry 1976; R. Dahlgren in R. Paris and thus had access to all important collections then Dahlgren & Thorne 1984 and in G. Dahlgren 1989; available. Geographic isolation played a paramount role Johnson and Briggs 1984; Renner 1989b; Huber 1991) in his understanding of the family; differing from his supported the recognition of a distinct family and still predecessors, he rigorously separated Old and New others returned to placing Mouriri and Memecylon in the World genera, recognizing the new groups thus formed at Myrtaceae (Richard 1845; Grisebach 1864). the tribal level. In order to key out the resulting new In his Quinarian arrangement of the Melastomataceae, tribes Triana (1865, and following him all later workers) De Candolle (1828b) grouped the 68 melastome genera had to rely on characters such as number of floral parts, he recognized according to seed type (cochleate with connective appendages, and geography (the Dissochae- curved vs. straight), degree of fusion between teae are distinguished from the Miconieae and the Ti- and and type, ovary summit (with bouchineae from the Osbeckieae by the first of each pair outgrowths vs. unadorned), and type of anther dehiscence being paleo-, the second neotropical). At the subfamilial (by one pore, two pores, or by longitudinal slits). From level, on the other hand, Triana merged Naudin’s Astro- later arrangements, De Candolle’s differs most strikingly nioideae and Kibessioideae, recognizing only three major in its complete disregard of geographic origin. taxa: Memecyloideae, Astronioideae, and Melastomatoi- The next major student of the family, Naudin (1849- deae (formalized by J. D. Hooker in Bentham & Hooker 1853), emphasized type of placentation and kind and 1867). number of seeds, which were used to distinguish five Severe discrepancies between the distribution of wood subfamilies (Melastomatoideae, Astronioideae, Kibes- anatomical characters and Triana’s classification were sioideae, Memecyloideae, and Mouririoideae). Within first pointed out by van Tieghem (1891a) who studied the the Melastomatoideae, Naudin recognized four tribes, wood of most of the genera recognized by Triana and Microlicieae, Lasiandreae (more or less todays Melasto- concluded (1) that wood anatomy did not support the meae), Rhexieae, and Miconieae (which include Dis- merging of Astronioideae and Kibessioideae, and (2) that sochaetinae, Blakeinae, Merianiinae, and an extremely wood anatomy did not support those of Triana’s tribes heterogeneous Sonerilinae). that were based solely on connective configuration. Triana’s system, published in 1865 and slightly mod- Groupings based on seed morphology, on the other hand, ified in 1871, is the one in use today (Cogniaux 1891; were supported by wood anatomy. Krasser 1893; Melchior 1964; Hutchinson 1973; Bremer A total of 489 genera have been proposed in the fam-

35 Nod I. Bot. 13 (5) (1993) 52 1 Tab. 2. Classification of the Melastomataceae and Memecylaceae, with the genera assigned to subfamily and tribe according to the new scheme. Generic synonymizations made in recent monographic and floristic literature are indicated but no new ones are suggested, except in the cases of Tigridiopulmu C. Chen, Tylunthera C. Hansen, and Vietseniu C. Hansen. See text for perceived generic relationships. Numbers in parentheses give the total species for a genus; species numbers and geographic range data are mainly from herbarium surveys and recent revisions and floras as cited; only the most general indication of the geographic range is given. Melastomataceae total: c. 166 genera, 4200-4500 species. Memecylaceae total: 6 genera, 430 species.

Memecylaceae DC. B. Sonerilu-Berroloniu-Gravesia Alliance Amahiblemmu Naudin (13)~, - trouical West Africa: Jacques- Lijndeniu 2011. & Moritzi (10) - W Malesia, Java, Ceylon (3), kix (1974a) Magascar (6),West Africa (I); Bremer (1982) Aschistanfheru C. Hansen (I) - Vietnam; Hansen (1987b), prob- Memecylon L. (incl. Kluineustrum Pierre ex A. Chev.) (300) - ably identical with Phylluguthis Asia and Oceania, Africa (70), Madagascar (78); Bremer Berroloniu Raddi (18) - SE Brazil, 1 sp. in ; Baum- (1979, 1983); Maxwell (1980b) gratz (1989-1990) Mouriri Aubl. (81) - tropical America, most diverse in Brazil; Boyuniu Wurdack (I) - Morley (1976, 1985, 1989) Brirteniu Cogniaux (1) - Borneo (Sarawak); Hansen (1985d) Sputhundru Guill. & Perr. (2) - W Africa; Jacques-Fklix (1978b, Culvoa J. D. Hooker (19) - tropical Africa; close to Gruwesia 1985) Cenrrudeniu G. Don (4) - Mexico to coastal ; placed Votomiru Aubl. (8) -eastern Amazonia (Brazil, French Guiana); here following Almeda (1977 and in press) Morlev<~ (1963. 1985. 1989) Cenrrudeniusrrum Cogniaux (2) - northern (Almeda Gilg (33) Africa (24), Madagascar (9); Jacques- 1977) Fklix (1978b, 1985) Cincinnobotrys Gilg (incl. Gruwesiellu A. & R. Fernandes, Ha- plophyllophorus A. & R. Fernandes, and Primuluriu Brenan) (6)- tropical Africa; Jacques-Fklix (1975) Melastomataceae Jussieu Cyanundrium Stapf (5) - Borneo Subfamily Kibessioideae Naudin (1849) Dicellundra J. D. Hooker (2) - tropical West Africa; Jacques- Fklix (1974b) Tribe Kibessieae Krasser (1893) Diplurpeu Triana (I) - , Colombia Pternundru W. Jack (incl. Kibessia DC., Ewyckiu BI., Macro- Enuulophyton Steenis (I) - plncis BI., Recromitria BI.) (15) - SE Asia, Malaysia; Max- Feliciudamia Bullock (I) - Africa (Guinea); nomenclatural sub- well (1981). stitute for Adumiu Jacques-Fklix (195 1) Fordiophyton Stapf (incl. Stapfophyron Li) (8) - , In- dochina Subfamily Melastomatoideae Naudin (1849) Gruwesia Naudin (incl. Veprecella Naud., Phornorhamnus Bak., Tribe Astronieae Triana (1865) (Irothecu Gilg, Peralonemu Gilg, Orthogoneuron Gilg, Neo- Maxwell & Veldkamp (1990a, b) peralonemu Brenan) (100) - Madagascar (95), Africa (5) Asfrocalyx Merrill (I) - Philippines Kerriothyrsus C. Hansen (1) - Laos; Hansen (1988b), prob. Astronia Blume (59) - Malay archipelago identical with Phyllugarhis A. Gray (incl. Nuudiniellu Krasser) (67) - Malay Macrocenrrurn J. D. Hooker (21) - N , reaching archipelago Brazil Beccuriunthus Cogniaux in Boerlage (incl. Bumlera Lauterb. & Muguireanrhus Wurdack ( 1 ) - Guyana; Wurdack (1964a) K. Schum. and Eweretriu Merr.) (22) - Malay archipelago Monolenu Triana (10) - Guatemala to Amazonia Opisrhocenrru J. D. Hooker (I) - Colombia, Venezuela to N Tribe Sonerileae Triana (1865) Brazil (Rio Negro) A. Oxysporu Alliance Phylluguthis Blume (incl. Tigridiopulmu C. Chen and Tq'lan- Anerincleistus P. W. Korthals (incl. Pomurostomu Stapf, them C. Hansen (1990b)) (30) - southern China to Thailand, Creughiellu Stapf, Oritrephes Ridley, Plugiopetulum Rehder Sumatra and Borneo (Hansen 1988a), Phuulunrhus Ridley, Perilimnustes Ridley, Preussiella Gilg (2) - tropical West Africa (Cameroun, Gabon); Krusseru 0. Schwartz) (30) - and Malesian region; Jacques-Fklix (l977b) Maxwell (1989) Sulpingu Martius ex DC. (incl. Saccolena Gleason) (8) - N Burtheu J. D. Hooker (I) - China, Taiwan, Vietnam; Hansen South America (1980b) Surcopyrumis Wallich (I) - Nepal, China, Taiwan, Thailand, Blustus Loureiro (12) - Assam, China to Vietnam, Malay archi- Malay archipelago; Hansen (1979) pelago; Hansen( 1982) Scorpiothyrsus Hui-Lin Li (6) - China; very close to Phylla- Blume (incl. Tashiroeu Matsum.) (13) - China, Japan, gurhis Formosa Sonerila Roxburgh (100-175) - tropical Asia; Lundin (1983) Driesseniu P. W. Korthals (incl. Triuruntheru Backh. and Vier- Tateunrhus Gleason (I) - Venezuela seniu C. Hansen (Hansen 1984)) (18) - Vietnam, Borneo, Triu/enu Naudin (incl. Diolena Naudin) (22) - Mexico to Ama- Sumatra, Java; Hansen (1985b) zonia Neodriesseniu Nayar (6)- Borneo; Hansen (1985a) Tryssophyton Wurdack (I) - Guyana; Wurdack (1964a) Ochthochuris Blume (incl. Phueoneuron Gilg) (7) - Asia (5); Africa (2); Hansen & Wickens (1981) Tribe Merianieae Triana (1865) Oxysporu DC. (incl. Allomorphia BI., Cumpimiu Ridley (Han- Adefobotrys DC. (25) - Mexico and Jamaica to Brazil and sen 1988c), Cyphotheca Diels (Hansen 1990a), Styrophyron Bolivia Hu, and Tuyloriophyton Nayar) (65) - India, China, Malay Axinaeu Ruiz & Pavon (30) - Costa Rica to Bolivia ; close to Peninsula, Sumatra, Melanesia (Fidschi Is.); Maxwell (1982) Meriuniu; Eves ( 1936) Poikilogyne E. G. Baker (incl. Dicerospennum Bakh. f. and Behuriu Chamisso (I 0) - SE Brazil Scrobiculuriu Mansf.) (21) - SE Asia Benevidesiu Saldanha & Cogniaux (2) - SE Brazil (Bahia) Poilunnammia C. Hansen (4) - Vietnam; Hansen (1987a) Bisglaziowiu Cogniaux (1) - SE Brazil (Bahia) Sporoxeiu W. W. Smith (4) - upper Burma, China Centroniu D. Don (1 5) - Panama to and Brazil, Guayana Stusseniu C. Hansen (based on membrunifolius Li = Dolichouru Brade (1) - SE Brazil (Espirito Santo) Neodriesseniu membrunifoliu (Nayar) C. Hansen, 1980) (I) - Graffenriedu A. P. DC. (incl. Culyptrella Naud. and Ptilanthus Vietnam; Hansen (I 980a, 1985~) Gleason) (44) - Central and S. America to Bolivia; West lndies Huberia A. P. DC. (6) - SE Brazil, I sp. in the Andes Acioiis D. Don (30) - Mexico, Lesser Antilles, Brazil and 0. Swartz (74) - Central and S. America to Bolivia; Bolivia West Indies Acisanihera P. Browne (17) - Venezuela, Guianas, Brazil to Merianihera Kuhlmann (3) - SE Brazil Paraguay Neblinanihera Wurdack (1) -Venezuela to NW Brazil (Cerro de A. P. DC. (I) - Guianas and northeastern Brazil la Neblina) Arihrosiemma Pavon ex D. Don (7) -from Mexico and the West Wurdack (I) - Venezuela Indies to Bolivia (absent from Brazil) Pachyloma A. P. DC. (incl. Urodesmium Naudin) (6) - Col- Brachyoium (A. P. DC.) Triana (58) - Colombia to Argentina; ombia, Venezuela, Guyana, N Brazil Wurdack (1953) Phainaniha Gleason (4) - Guayana Chaeiolepis (A. P. DC.) Miouel (10) - Colombia. Venezuela. I Markgraf (7) - Colombia, Ecuador, and Peru: West indies, N Brazil Wurdack (1989); Almeda (1989b, 1990b) A. P. DC. (22) - tropical South America, most diverse in Brazil Tribe Rhexieae DC. (1828) Cornoliopsis Wurdack (I) - Venezuela (Cerro de la Neblina) Rhexia L. (13) - North America, concentrated in SE coastal Desmoscelis Naudin (1) - Venezuela to Brazil and Bolivia plain; Kral and Bostick (1969) (savannas) Ernesria A. P. DC. (incl. Brachypremna Gleason and Pseu- Tribe Microlicieae Naudin (1849) doernesiia (Cogn.) Krasser) (16) - Andes and upper Amazon Bucqueiia A. P. DC. (3) - Northern Andes basin, Guianas, Brazil DC. (incl. Pyramia Cham.) (21) - south-central Friizschia Chamisso (I) - south-central Brazil Brazil; Martins (1984) Heierocentron W. J. Hooker & G. A. Walker Arnold (27) - Naudin (2) - Colombia, Venezuela: a high-altitude Mexico and ; Whiffin (1972) Lnriculepis Brade (I) - Brazil (upper Rio Negro) Chaetostoma DC. (12) - south-central Brazil; very close to A. P. DC. (incl. Siphaniheropsis Brade) (22) - most diverse in Guayana, also Colombia, Brazil, and Bolivia; Ren- Eriocnema Naudin (1) - south-central Brazil ner (1989a) DC. (46) - south-central Brazil Mallophyion Wurdack (I) - Venezuela (Chimanti Mt.); Wur- Lithobium Bongard (1) - south-central Brazil dack (1964a) Microlicia D. Don (100) - south-central Brazil, a few species in A. P. DC. (27) - Venezuela, Guayana, Colombia, Guayana, Peru, and Bolivia Brazil; Martins (1989) DC. (15) - most diverse in SE Brazil, but ex- Microlepis (A. P. DC.) Miquel (4) - south-central Brazil tending to Mexico; Renner ( 1990) Monochaerum (A. P. DC.) Naudin (45) - Mexico to Peru and Stenodon Naudin (1) - Brazil (Bahia) Venezuela; Almeda (1978) DC. (I 1) - south-central Brazil Naudin (I) - Central America, West Indies, to central Brazil & - Tribe Melastomeae (Osbeckieae DC. (1828))~ ,, Pilocosra Almeda Whiffin (4) Costa Rica to Ecuador; Paleotropical genera Almeda and Whiffin (1980); close to Tibouchina Amphorocalyx J. G. Baker (5) - Madagascar; close to Dionycha Poieranihera Bongard (2) - Brazil, Venezuela; Wurdack (1963) Antherotoma (Naudin) J. D. Hooker (2) - Africa (2). Mada- Pterogasira Naudin (2) - northern South American savannas; gascar (I) Renner (in press) Cailliella Jacques-Fklix (I) - Africa (Guinea) (A. P. DC.) Miquel (14) Mexico and West Indies to Dichaetanihera Endlicher (incl. Sakersia Hook. f. and Bar- Paraguay; Renner (in press) beyastrum Cogn.) (30) - Madagascar (23), Africa (7) Sandemania Gleason (I) - Venezuela, Brazil, Peru; Renner Dinophora Bentham (1) - tropical West Africa (Sierra Leone to (1987) Angola) Schrvackaea Cogniaux (I) - Mexico to Colombia; Renner (in Dionycha Naudin (3) - Madagascar; Perrier de la Bithie (1951) press) Dionychastrum A. Fernandes & R. Fernandes (1 ) - Africa (Tan- Siphanihera Pohl ex DC. (incl. Meisneria DC. and Farringionia zania) Gleason)( 16) - Colombia, Venezuela, Brazil, Bolivia Bentham (incl. Podocaelia (Bth.) A. & R. Fernandes) Svitrumia Chamisso (2) - Brazil (Bahia, Minas Gerais) (140) - Africa (120), Madagascar (20); close to Tibouchina Aublet (incl. Purpurella Naudin and Itatiaia Ule) Guyonia Naudin (1-2) - tropical West Africa; close to Osbeckia (240) - Mexico, West Indies to northern Argentina Heterotis Bentham (14) - tropical West Africa; close to Os- Tibouchinopsis Markgraf (2) - Brazil (Bahia) beckia; Jacques-Felix (1981b) Melastoma L. (50) - Asia; 1 sp. introduced in Madagascar Tribe Miconieae DC. (1828) Melastomastrum Naudin (6) - tropical West Africa; close to Paleotropical genera; see especially Maxwell ( 1980a. 1983) Osbeckia; Jacques-FLhx (1974) Boerlagea Cogniaux (1) - Borneo Nerophila Naudin ( I ) - Africa (Senegal); perhaps a Chaeiolepis Caianihera F. V. Mueller (incl. Hederella Stapf and Phylla- (Wurdack, pers. comm. 1985) pophysis Mansfeld) (16) - Malesian region (New Guinea, Osbeckia L. (incl. Rousseauxia DC.; Jacques-Felix (1973)) (50) Borneo, Sumatra) - Asia (+ 30); ; Africa (5),Madagascar (13); Hansen Creochirorl Blume (incl. Eisocreochiron Quisumb. & Merrill ( 1977) and Anplectrella Furtado) (9) - New Guinea, Java, Philip- Otanthera Blume (incl. Lachnopodium Blume) (3) - China to pines the Philippines and Australia; very close to Melastoma Dipleciria (Blume) H. G. L. Reichenbach (incl. Anplectrum A. Pseudosbeckia A. & R. Fernandes (1) - Africa (Mopmbique); Gray, Backeria Bakh.) (8) - Burma, Indo-China, Malesian close to OJbeckia region; Veldkamp et al. (1978); Diplecfria will be treated as A. L. Jussieu (incl. Tetraphyllaster Gilg) (14) - synonymous with Dissochaeia in the Flora of Thailand (Ren- Africa; 1 sp. in Madagascar and the Mascarene Islands; close ner, ms) to Osbeckia: Jacques-Fklix (1976) Dissochaeia Blume (incl. Omphalopus Naudin, Dalenia Korth., and Neodissochaeia Bakh. f.) (22) - India, Malasian region Neotropical genera Kendrickia J. D. Hooker (I) - South India, Ceylon Acanthella J. D. Hooker (2) -Venezuela, Colombia. Brazil (Rio Macrolenes Naudin ex Miquel (1 1) - Malay Peninsula, Philip- Negro) pines

3S* Nod J. Bot. 13 (S) (1993) 523 Medinilla Gaudichaud-BeauprC (incl. Hypenunthe BI., Curioniu trum Naud., Platycenrrum Naudin) (175) - Mexico to Argen- Naudin, Medinillopsis Cogn., Cephulomedinillu Merr.) (200- tina, absent in the West Indies, most diverse in southeastern 400) - Madagascar (70), Africa (2), SE Asia (85): Maxwell Brazil (1978), Regalado (1990), Whiffin (1990) Llewelyniu Pittier (1) - Venezuela (coastal mts.); should be incl. Pachycentria Blume (10) - SE Asia; Maxwell (1978) in Henrietteu; Judd (1989) Plethiandra J. D. Hooker (9) - Borneo, Malay Peninsula Loreya A. P. DC. ( incl. Heteroneuron Hook. f.) (13) - Central Pogonunthera Blume (1) - Malay Peninsula, Philippines, Malay America to Brazil and Bolivia; Renner (1989a) archipelago; Maxwell (1978) Aublet (3) - Amazon basin (esp. Colombia, Venezuela, Nayar (6) - Assam to Indo-China Brazil) J. D. Hooker (incl. Ekmaniocharis Urban) (23) - Neotropical genera Greater Antilles; Skean (1993) Pilger (7) - Andes Ruiz & Pavon (incl. Copedesma Gleason, Pachydesmia Allormieta Gleason (1) - Colombia Gleason, and [curia Macbr.) (1OOO) - throughout tropical Anaectocalyx Triana (3) - Venezuela (Miconieae) America Necker ex Rafinesque (7) - tropical America; Renner Myriusporu A. P. DC. (I) - Guyana and Amazon basin; incl. in (1989a); incl. in Loreyu by Judd (1989) Loreyu by Judd (1989) A. P. DC. (incl. Momrnseniu Urban & Ekman) Myrmidone C. F. P. Martius (incl. Horrnoculyx Gleason) (2) - (3 1) - Greater Antilles Colombia, Guyana, Venezuela, Brazil (Amazon basin); incl. Catocoryne J. D. Hooker (1) - Colombia in by Judd (1989) Naudin (1) - Colombia Necrumium Britton (1) - Trinidad and Venezuela D. Don (5) - West Indies, Jamaica (l), Lesser Ossuea A. P. DC. (incl. Octopleuru Griseb. and Diclemia Nau- Antilles; polyphyletic fide Judd (1989), see also Judd & din) (91) - West Indies and Mexico to Brazil; likely poly- Skean (1991) phyletic and dismembered by Judd (1989) into Pentossaea Clidemia D. Don (incl. Heterotrichum DC.) (120) - Mexico to Judd and Sugrueu DC.; see also Judd & Skean (1991) Peru; axillary-flowered species excl. as Sugrueu by Judd Puchyunthus A. Richard (16) - West Indies and Cuba ( 1989) Naudin ex A. Gray (7) - southeastern Brazil; D. Don (43) - Mexico to Peru, reaching the West scarcely distinguishable from Leandru (Wurdack 1962) and Indies and Brazil incl. in that genus by Judd & Skean (1991). Cyphosryla Gleason (2) - Colombia Tetrazygiu L. C. Richard ex A. P. DC. (incl. Tetrazygiopsis A. P. DC. (12) - Amazon basin, esp. Brazil Borhidi) (21) - Antilles and Guayana Henriettellu Naudin (5 1) -West Indies and Guatemala to Bolivia Tococu Aublet (incl. Microphyscu Naudin) (54) - Mexico to and Brazil; should probably be included in Henrietteu; Judd Brazil and Bolivia; West Indies (Tobago) ( 1989) Huilaea Wurdack (4) - Colombia; close to Chulybeu Tribe Blakeeae Hook. in Benth. & Hook. (1867) Killipia Gleason (4) - Colombia, Ecuador Blukeu P. Browne (incl. Pyxidunthys Naudin) (100) - Mexico Kirkbridea Wurdack (2) - Colombia: Wurdack (1976) and West Indies to Bolivia; Almeda (1990a) Raddi (incl. Tschudya DC., Oxymeris DC., Clidemius- Topobeu Aublet (62) - Mexico to Amazonia, absent in the West Indies ily, 90 of these since Cogniaux (1891); only about 166, Gondwanian distribution. Memecylon today comprises however, are currently recognized (Tab. 2). The Cypho- about 300 species in Asia (Maxwell 198Ob; Bremer 1979, styleae (Gleason 1929) and Feliciadamieae (Jacques-Fe- 1982, 1983), 70 in Africa (Jacques-Felix, in litt. 1991) lix, in litt. 1991) are the only tribes that have been added and 78 in Madagascar (Jacques-F6lix 1984, 1985). Very to the 13 recognized traditionally. On the other hand, close to Memecylon are Wurneckeu with 33 species in several workers have suggested merging the Sonerileae Africa and Madagascar (Jacques-Felix 1978a, b, 1985), and Bertolonieae (J. D. Hooker in Bentham & Hooker Lijndeniu with three species in southeast Asia, six in 1867; Gleason 1932; Bakhuizen 19461947; Wurdack Madagascar and one in West Africa (Bremer 1982; Jac- 1964a), and J. D. Hooker (in Bentham & Hooker 1867) ques-Felix 1985), and Spafhundru with two species in and Wurdack (in a letter to J. Hutchinson of 8 September Africa (Jacques-F6lix 1978a, b, 1985 and in litt. 1991). 1969) among others also doubted the distinctness of the The two neotropical genera, Mouriri with 81 species Tibouchineae and Melastomeae as traditionally con- (Morley 1953, 1976, 1985, 1989) and Votomitu with eight ceived. species (Morley 1963, 1985, 1989), are very close to each other and should perhaps be merged. Memecylaceae and Melastomataceae share no apo- morphic states and differ in all but a few basic myrtalean The status and relationships of the features such as intraxylary phloem, diplostemonous Memecylaceae DC. , and presence of a well developed floral tube. Characters that disagree in the two families are listed in The six genera of Memecylaeae form a morphologically Tab. 3, and the states found in Memecylaceae are dis- homogeneous group, and Naudin’s (1 849-1853) sugges- cussed below; the respective states in Melastomataceae tion to recognize two subfamilies, the paleotropical Me- are discussed in the next section and not repeated here. mecyloideae and the neotropical Mouririoideae, based on Memecylaceae have 1-locular ovaries with a free cen- differences in placentation and seed number was never tral placenta or 2-5-locular ovaries with central-basal, taken up. It is assumed that the present range of the basal, or in ten species of Mouriri and four of Votomita family represents the remnant of a once continuous axillary placentation (Morley 1953). Up to 18 ovules

524 Nord. J. Bot. 13 (5) (1993) ules in the species in which this has been investigated (Morley, in litt.) and often radially thickened endothe- cium cells (Eyde & Teen 1967), the plesiomorphic condi- tion in the order although not necessarily in the family (Morley 1953: 252). The anthers open by two longitudi- nal slits in Memecylon and its segregates and by two short, drop-shaped slits in most species of Mouriri. Func- tionally the latter are equivalent to pores: Mouriri but not Memecylon flowers are buzz-pollinated by pollen collect- ing bees. A few species of Mouriri and Votomita that do not have thickened endothecium cells or any detectable endothecium still have anthers that dehisce longitudinally A (Morley 1953, 1976). Assuming an ancestral condition c------( which combined a thickened endothecium with a longici- 0,5 mm -5mrn dal mode of dehiscence (as represented in Memecylon), the problem becomes the sequence of character build-up Fig. 1. A. Seed of Melastomataceae. - B. Seed of Memecyla- ceae. during the evolution of the family for the syndrome (i.e., poricidal dehiscence and loss of an endo- thecium). have been recorded in Memecylon and up to 16 per locule Dorsally the connective carries a concave gland cov- in Mouriri (M. trunciflora; Morley 1976), but usually ered by a viscuous liquid that at various times was only one or two of them develop into a large subglobose thought to be or floral oil, but probably is neither. seed (Fig. I), containing an with well developed The role of the clearly terpenoid glandular products in cotyledons that are either rolled or wrinkled (in Memecy- pollination is unclear (Renner 1989b). Ion; Jacques-Fdix 1977a; Bremer 1981) or thick and Mernecylon and Mouriri have large sclereids that are fleshy (Mouriri and Memecylon rivulare). Germination in associated with the terminations of the veins (Foster Memecylaceae is cryptocotylar, with the testa being 1946; Morley 1976; Rao et al. 1980, 1983; Rao & Jac- raised above the soil and enclosing the cotyledons (Duke ques-Fklix 1978); Memecylon stands out in the posses- 1969; Vogel 1980; Burtt 1991; pers. obs.). The seeds sion of globular silica bodies in the epidermis (Baas store starch. 1981). Stomata in Memecylaceae are paracytic (Mentink Memecylaceae flowers have antepetalous ovary loc- & Baas 1992). The wood has fibre-tracheids and sxially

Tab. 3. Differences between Memecylaceae and Melastomataceae.

Memecylaceae Melastomataceae

Anther connective with a terpenoid-producing gland lacking glands Endothecium fibrous not fibrous Anther dehiscence longicidal, slits sometimes short and poricidal, rarely longicidal; the pores functioning as pores (Mouriri) developing in an epidermis-free region Placentation axillary, axile-basal, basal or free central axillary, axile-basal, or parietal (Prernandru) (Memecylon) Locules opposite the opposite the except in Pternandru and Rhexia Seeds few, large and with thick or convolute numerous, small, and with short cotyledons cotyledons with fibrous exotegmen without fibrous exotegmen Germination cryptocotylar phanerocotylar Leaf venation brochidodromous acrodromous Stomata paracytic anomocytic, polocytic, cyclocytic Leaf sclereids large absent Indumentum usually glabrous usually with hairs and/or emergences and mite domatia absent common in Miconieae Wood with fibre-tracheids and interxylary phloem with libriform fibers and lacking interxylary phloem except in Pternandru Growth form trees and shrubs, treelets, herbs, climbers

Nord. J. Bot. 13 (5) (1993) 525 included (interxylary) phloem and vessel-ray pits that are high; the latter are concentrated in the Merianieae and small with vestured borders. Miconieae (Alloneuron, Loreya, Miconia, Tessmannian- Chemical differences between the Memecylaceae and thus). Root climbers are quite common in the Miconieae Melastomataceae include the frequent occurrence of acy- (Catanthera, Clidemia, s. l., Kendrickia, lated anthocyanins in the latter, whereas so far only Leandra (incl. Pleiochiton), Macrolenes, Medinilla, a non-acylated anthocyanins have been found in the Me- few Miconia) but occur also in the Astronieae (a few mecylaceae (Lowry 1976; but only two species of Meme- species of Astronia), and Merianieae (, Phai- cylon have been investigated), and Favarger (1952, 1962) nantha). Several non-climbing species of Blakeeae and noted a difference in the colour of the root tips between Miconieae (e.g., Medinilla spp., Pogonanthera, Pachy- the two families, Melastomataceae but not Memecyla- centria) are capable of growing as facultative epiphytes, ceae containing anthocyanin in the root tips. and many herbaceous species, especially of Sonerileae, Based on pentamery and the more primitive forms of grow low on trunks or on fallen logs (Renner 1986 for a placentation occurring only in the New World, Morley survey). (1953 and pers. comm.) suggested that Mouriri is closer Forty-four genera are largely or entirely herbaceous, to the ancestral condition than Memecylon. This is sup- but although this is about a quarter of the total number of ported by cotyledon morphology, which in Memecylon genera, only relatively few species are involved, the and appears more derived than in Mouriri. shrubby groups comprising by far the majority of all Anther dehiscence, on the other hand, appears plesio- species. Herbaceous species are concentrated in the Me- morphic in Memecylon and relatives. lastomeae and Sonerileae, of which 19 and 20 genera, The closest affinities of the Memecylaceae, in my respectively, include herbs. Many sonerilean herbs are opinion, lie with Myrtaceae, but I know of few potential acaulescent with basal rosettes of large, somewhat turges- synapomorphies uniting these groups. These are the ter- cent leaves. Tuber formation occurs in Cincinnobotrys, penoid-containing connective glands, in Myrtales found Heterotis, , and . only in Myrtaceae and Memecylaceae (but erroneously The question as to the ancestral growth habit in Melas- scored as absent in the latter by Johnson & Briggs tomataceae can be answered by looking at closely related (1984)), and the few large starch-containing seeds. Fur- families, such as Lythraceae, Crypteroniaceae, and Myr- ther light on the relationships of both families will prob- taceae all of which are basically ligneous. Although ably only be shed by new kinds of data. Other myrtalean growth form was among the characters surveyed for all families, such as the Crypteroniaceae and Lythraceae, genera, I have not included it among the characters used appear more distantly related to the Memecylaceae be- in the phylogenetic analysis because it proved too varia- cause of their capsular fruits, numerous small seeds with ble. minute embryos, and lack of axially included phloem.

Reproductive architecture Melastomataceae characters: variation The variety of inflorescences in the Melastomataceae and polarization corresponds to the size of the family but shows less diversity than, for example, the Myrtaceae (Johnson & The purpose of this section is to pull together all in- Briggs 1984). Inflorescences are determinate thyrsoid formation relevant to solving infrafamilial relationships ramification systems (Mora Osejo 1966; Cremers 1986; in the family and, for the subset of characters found Weberling 1988). Uniparous (scorpioid) cymes are par- phylogenetically informative, to discuss polarization. Un- ticularly frequent in the Sonerileae, but occur scattered in less otherwise indicated, the circumscription of the higher other groups. Single terminal flowers are rare, occurring taxa is that proposed in the Phylogenetic analysis below mainly in the fruticose and herbaceous groups (Soneri- and shown in Tab. 2. leae, Melastomeae); they are the result of reduction and suppression as has been shown in various genera (Wur- dack 1953; Almeda 1978; Cremers 1986). Cauli- and Germination ramiflory is most frequent in the Miconieae and is a Germination is phanerocotylar, the eophylls being decus- derived feature. sate with the cotyledons, but seedlings of only relstively The systematic usefulness of architecture few species have been studied (Ziegler 1925; Favarger in Melastomataceae appears limited to low taxonomic 1952, 1962; Gadella et al. 1969; Rao & Chin 1972). levels, but several generic realignments in Miconieae based on inflorescence position (terminal vs. lateral) have been proposed by Judd (1986a, 1989) and Judd & Skean Vegetative architecture (1991). A recent attempt at interpreting the distribution of different inflorescence types on the background of the Melastomataceae exhibit a diversity of growth habits, traditional system (Sell & Cremers 1987) suffers from a including herbs, shrubs, treelets, and trees to 3W5 m misrepresentation of that system. Thus it is said that the

526 Nord. I. Bot. 13 (5) (1993) Memecyleae, Blakeeae, and Miconieae are the most Anisophylly is common in the Sonerileae sensu lato primitive tribes in the family, certainly not the orthodox and Miconieae, less so in the Kibessieae, Astronieae, view (cf. History of the classification of Melastomataceae Merianieae, Microlicieae, Melastomeae, and Rhexieae. and Memecylaceae). (Sonerileae: Anerincleistus, Barthea, Blasrus, Bredia, Driessenia, Neodriessenia, Stussenia, , , Cincinnobotrys, Fordiophyton; Phyllagathis Wood anatomy (incl. Tylanthera), Surcopyramis, Sonerila; , , Diplurpeu, , Monolenu, Trio- The distribution of wood-anatomical characters in the lena; Merianieae: Phainantha; Miconieae: Allomaieta, family is very important (Tieghem 1891a, b; Janssonius Catanthera, Clidemia, Maieta, Medinilla, Myrmidone, 1914; Metcalfe & Chalk 1950; Vliet et al. 1981) and ; Blakeeae: , Topobeu). Sometimes alter- provides one of the major reasons for breaking up the nate leaves on a stem are reduced or caducous so that Astronioideae sensu Triana ( 1871) and Cogniaux ( 1891). phyllotaxy appears alternate (species of Blakea, Catan- Typical Melastomataceae wood has libriform fibers, of- thera, Centradenia, Cincinnobotrys, Clidemia, and Mac- ten large and simple vessel-ray pits, fiber dimorphism, rocentrum), and in Phainantha all leaves on one side of libriform septate and non-septate fibers, and grouped the stem are replaced by climbing roots. vessels. Most members of Triana’s Astronioideae (Astro- Ant domatia in melastomes are confined to the Mico- , Astronidium, Astrocalyx, and Beccarianthus) have nieae (Allomaieta, Clidemia, Conostegia, Henriettellu, this kind of wood; Pterizandra, however, has fiber trache- Maieta, Myrmidone, Ossaeu, Tococa) and Blakeeae and ids and axially and radially included phloem. are typically situated in leaf bases, though hollow stems I have used one wood anatomical character in the occupied by occur in Blakea, Miconia, Tococa, and phylogenetic analysis, namely fiber type, fiber tracheids Topobea. In the Old World, leaf domatia are rare (but are being generally considered plesiomorphic in the Myrtales found in Medinilla dispariifolia), their ecological func- and in the angiosperms as a whole, libriform fibers as tion being taken by root domatia (Pachycentria, Pogo- derived. nanthera). Mite domatia have so far been reported from Miconieae, Blakeeae, and Sonerileae (Schnell 1963; Al- meda 1989a, I990a; Baumgratz 1989-1 990). Leaf morphology and anatomy Extensive summaries of the pioneer studies of Pflaum Epidermal features (1 897), PalCzieu (1899), and Gottschall (1900) on melas- tome leaf morphology and anatomy may be found in Stomata1 types in Melastomataceae are extremely varied; Solereder (1899 & 1908) and Metcalfe and Chalk (1950; especially frequent are the following kinds: anomocytic the latter is difficult to use because of the numerous (neighbouring cells not different from guard cells), polo- synonymous generic names and the insufficient distinc- cytic (guard cells incompletely surrounded by a single tion in the summary statements of Memecylaceae and subsidiary cell), diacytic ( enclosed by a pair of Melastomataceae). A recent complete survey is that of subsidiary cells whose common wall is at right angles to Mentink & Baas (1992). the guard cells), and tetracytic (stomate surrounded by Melastomataceae leaves are entire or dentate and de- four subsidiary cells, two lateral and two polar), but other cussate and typically have a petiole. The primary lateral types and intermediate kinds also occur (Mentink & Baas vascular bundles run parallel to the leaf margin from the 1992). Anomocytic stomata are the basic type in the base to the apex. This type of venation is rare in the Myrtales (Baas 1981; R. Dahlgren & Thorne 1984: 636; angiosperms (Hickey 1973) and nowhere as persistent as Keating 1984: 816), but it is difficult to polarize stomata1 in the Melastomataceae. A few isolated genera and spe- types. The epidermis may be glabrous or possess an cies, notably Centradenia, Sonerila, and some Miconieae elaborate indumentum of glandular, stellate, or simple (e.g., Loreyu arborescens, Macairea rufescens; Renner hairs and emergences (Wurdack 1986; Mentink & Baas 1989a) have imperfectly acrodromous or penninerved 1992), and the cuticle may carry waxy Ornamentations leaves, and it would be interesting to know the devel- (illustrations: Wurdack 1986, Renner 1989a). With the opmental mechanisms behind such switches. exception of the conspicuous trichomes that characterize A feature of the ground tissue that may have taxonomic the leaves and floral tubes of many Melastomeae (the significance is the presence or absence of diverse calcium “tibouchinoid hairs” of Wurdack 1986), few major group- oxalate crystals (raphides, styloids, druses, and crystal ings are evident from epidermal features. An example are sand). Thus, Baas (1981) suggested that one or two paral- the Astronieae, which are characterized by non-glandular lel lines of specialization are exemplified in the Astro- peltate scales, mainly anomocytic stomata, a 1-3-layered nieae and Miconieae where the crystal complement in the hypodermis, and a complex vascular pattern (Mentink & leaves has come to consist exclusively of styloids, and Baas 1992). Some of these features also occur scattered Judd (198613) has shown that within the Miconieae, crys- in the Miconieae. The Kibessieae, according to Mentink tal type is sometimes useful at the generic level. and Baas, are clearly distinct from the Astronieae, and

Nord. J. Bot. 13 (S) (1993) 527 majority of genera, however, they are 1-pored, 2-pored or, rarely, 4-pored (some Miconia). A few Miconieae (some species of Charianthus, Mecranium, and Miconia sect. Chaenanthera) have anthers with rimose pores or slits, apparently a secondary development from poricidal opening (Wurdack, pers. comm.; Judd, in litt. 1992). The 2-pored condition occurs scattered in the Merianieae (Tessmannianthus), Sonerileae (Scorpiothyrsus, Sone- rila), Miconieae (Bellucia, Chalybea, Huilaea, Loreya, A B Mecranium, Medinilla, Miconia), and Blakeeae (Blakea, Topobea), and is variable within certain genera (e.g., Fig. 2. Melastomatoideae pore opening mechanism. - A. Cross Loreya) which comprise 1- and 2-pored species. The lack section through mature anther. - B. Longitudinal section. The pore develops in an epidermis-free region via the desiccation of of the usual fibrous thickenings of the radial endothecium exposed mesophyll. cell walls is correlated with a special mode of anther opening in Melastomataceae (Fig. 2): the pores develop in an epidermis-free region via the desiccation of exposed characterized by anomo-cyclocytic and paracytic stomata mesophyll (Ziegler 1925; Eyde & Teeri 1967). In other (stoma accompanied on either side by one or more sub- myrtalean families the irregular shrinkage of the endo- sidiary cells parallel to the long axis of the pore and guard thecium cells during anther dehydration leads to the rup- cells), druses, the absence of non-glandular peltate scales ture of the wall (e.g., in Memecylon: Eyde & Teeri 1967). and of a hypodermis, and the occurrence of a simple In the cladistic analysis, anthers that still have a func- vascular pattern in midrib and petiole. The remainder of tional endothecium and open along their entire length the family is heterogeneous in leaf anatomy (Mentink & were scored as the ancestral condition. Baas 1992). In Sonerileae s.1. polocytic stomata are gen- Melastome connectives are of striking plasticity, but erally present, and in the Blakeeae anomocytic ones. two major types may be discerned (Triana 187 1 : plates 1-5): one with dorsal connective spurs and basally un- prolonged connectives, the other with a basal-ventrally Floral morphology and anatomy prolonged connective for which the term pedoconnective was introduced by Jacques-F6lix (1953: 977; 1981a). The General considerations criterion I have used for scoring a taxon as having pedo- The floral features most useful for higher level system- connectives was that in the affirmative case the connec- atics in melastomes are placentation, configuration'of the tives of the largest stamens found are curved so that a ovary apex, and stamina1 morphology. These are dis- knee is formed between connective and filament. Be- cussed in turn below. Much has been made of the number cause other myrtalean families do not have such genic- of floral parts, for example, by Triana (1865, 1871), but ulate connectives or appendages, in the cladistic analysis this feature is usually too variable for generic grouping an unprolonged and dorsally unappendaged connective (Wurdack 1964a; Renner 1989a). aestivation is was assumed to be the ancestral condition. right-contorted and petal venation is imperfectly acrodro- mous. The gynoecium The androecium Ovary locules in Melastomataceae are antesepalous ex- Flowers in the family are actinomorphic, often with zygo- cept in Pfernandra and Rhexia where they are antepeta- morphic androecia, and are usually diplostemonous, with lous. An important character provided by the gynoecium the inner set of stamens usually smaller than the outer is its perigyny or epigyny. The degree of adnation be- one. The opposite is the case in (Almeda tween ovary wall and hypanthium, however, is not al- 1978) and Dissochaeta (incl. Diplectria) where due to ways easy to discern in herbarium material (Eyde & Teeri ontogenetic displacements the stamens in the outermost 1967) and is not correlated with either the hypanthium or ring are placed in front of the petals, the smaller in front ovary wall becoming fleshy in fruit (see below: fruit of the sepals (obdiplostemony). In some other species the characters). inner whorl is more or less rudimentary, with the antepe- The ovary apex in capsular-fruited melastomes is often talous stamens either staminodial or lacking entirely (ha- crowned by bristles, scales, or lobes around the style that plostemony). Immature anthers are tetrasporangiate and persist into the fruit stage and sometimes play a role in dithecal; mature anthers are generally bisporangiate with seed dispersal (Weber 1987; Stone & Weber 1988). Such the exception of Rhexia where they are unisporangiate, ovary outgrowths are especially common in the Soneri- one of numerous autapomorphies of that genus. In a few leae s. s. and Melastomeae and may characterize these genera the anthers open by two longitudinal slits (Pter- lineages but their homology is unclear. I consider ovaries nandra and the four genera of Astronieae). In the vast with unadorned tops plesiomorphic.

528 Nod J Bot. 13 (5)(1903) wall becomes fleshy and there is no pulp surrounding the seeds. Conflicting information on fruit type is often given by collectors on herbarium labels and in floras and mono- graphs (Bakhuizen 1946-1947; Maxwell 1981), and great care must be taken to ascertain the primitive condition in each case. Dry capsules from epigynous ovaries are found in the Merianieae (Merianthera, Tateanthus), So- nerileae (Feliciadamia), Melastomeae (Dinophora), Rhexieae (Rhexia), and Miconieae (Alloneuron, Allo- maieta, Cyphostyla), and fleshy capsules from perigy- nous ovaries have evolved repeatedly in the Melasto- 0,5mm meae. Further critical examination of fruit attributes in B the family is a matter of high priority, and may reveal the - phylogenetic affinities of some unusual genera, such as Fig. 3. Melastomeae seed. - A. The emerging radicle pushes the operculum from the micropyle. - B. Germination. Kendrickia or Alloneuron and its allies. Because in Crypteroniaceae and Lythraceae dry cap- sules are plesiomorphic and because completely inferior ovaries are generally considered more derived than peri- Placentation gynous ones, in the cladistic analysis fleshy fruits from There are axillary, axile-basal, and median-parietal pla- inferior ovaries and with seeds embedded in pulp were centas, and this is one of the most useful characters for scored as apomorphic. phylogenetic reconstruction because the three types ap- pear to have become fixed early in the phylogeny of the family. Median-parietal placentation is only found in Ovules, embryos, and seeds Pternandra (incl. Kibessia) and ontogenetically results from the strong expansion of the ovary bottom which Embryological studies in Melastomataceae, including pushes initially axillary placentae onto the walls of the ovule ontogeny, megagametogenesis, and embryogeny, ovary (Schumann 1887). Basal placentas are found in the have been conducted in 20 genera (Hofmeister 1858; Astronieae, where they are large and may ascend into the Ruys 1924; Ziegler 1925; Mauritzon 1939; Subrama- locules (Schumann 1887; Maxwell &Veldkamp 1990a. nyam 1942, 1944, 1946, 1948, 1951a, b; CrCtC 1956, b). Axillary placentas characterize the remainder of the 1957, 1960a, b; Iconomidis 1958; Etheridge & Herr family. Although there is some variation within this last 1968), representing all tribes except the Astronieae, Mer- type (the placentas may fill out the entire locule space, be ianieae, and Blakeeae. The functioning megaspore devel- restricted to the central part of the ovary column, or may ops into the Polygonum type megagametophyte. The be stalked), it appears that axillary placentation arose ovules are anatropous except in Rhexia where they are only once in ancestral Melastomataceae. Based on Schu- atropous (Etheridge & Herr 1968; Schmid 1984) and mann’s ontogenetic study and by comparison with bitegmic, with both integuments contributing to the mi- closely related groups, especially the Crypteroniaceae cropyle (resulting in a more or less Z-shaped micropyle). and Lythraceae, a plastic axile-basal or axile placentation A micropyle formed by both integuments is common in appears to be the plesiomorphic condition. the Myrtales, as are the mature exalbuminous seeds char- acteristic of all Melastomataceae. Seeds are of three major types: straight with a smooth or tubulate surface, straight with a foveolate surface, or The fruit cochleate (campylotropous; Fig. 3) with a tubulate sur- About 2150-2350 species in 41 genera have berries, face (Don 1823; Cogniaux 1891 ; SEMs: Whiffin & Tomb 2000-2200 in 125 capsules. Fleshy fruit walls arose sev- 1972; Renner 1989a; C. Hansen’s numerous unpublished eral times, correlated with dispersal by , mainly SEMs of the seeds of Asiatic species are available for birds, and even closely related species, such as Melas- study in the herbarium in Copenhagen). Campylotropy toma sanguineum and M. malabathricum or Pternandra becomes manifest rather late during the ontogeny of the echinata and €? coerulescens, may differ in seed dispersal ovules, and only the mature seeds are fully campylotro- and fruit morphology (further examples are found in pous (Bouman & Boesewinkel 1991). Such seeds with a Otanthera and ). While the capsule of the first distinctly convex antiraphal side and the micropyle sit- species in these pairs opens loculicidally to release wind- uated close to the chalaza are found in Melastomeae and dispersed seeds, the second species are dispersed. Rhexia. The adaptive advantage of campylotropy, as dis- In M. malabathricum this is achieved by the dry exocarp cussed by Bouman and Boesewinkel, may be that it tearing open irregularly to expose dark red, fleshy placen- enables seeds to contain embryos twice as long as the tae in which the seeds remain embedded until eaten by a seed itself, giving better opportunities for the early estab- bird; in €? coerulescens the hypanthium but not the ovary lishment of the seedling.

Nord. J. Bot. 13 (5) (1993) 529 Tab. 4. Character coding for Melastomataceae cladistic analy- ses; see text for discussion of additional characters and rationale Karyology for polarizations. Almeda & Chuang (1992) have recently summarized 1. Libriform fibers: 0 = absent; 1 = present karyological work in the family (mainly by Favarger 2. Anther dehiscence: 0 =by two slits; 1 =by one or two pores 1952, 1962, and Solt & Wurdack 1980) and based on this 3. Pedoconnective: 0 = absent; 1 =present and their own work have suggested a base-number of 4. Dorsal connective appendages: 0 = absent; 1 =present n=17 for the neotropical Miconieae. Polyploidy is fre- 5. Ovary crowned by outgrowths: 0 =absent; 1 =present 6. True bemes: 0 =absent; 1 =present quent in the family. 7. Consistently axillary placentation: 0 =absent; 1 =present

Chemistry Nectar production and pollination biology Relevant data may be found in Hegnauer (1990), and some references were provided above (The status and A summary of pollination in melastomes, including a relationships of Memecylaceae DC.) to chemical traits survey of the relevant older literature, is available (Ren- distinguishing Memecylaceae and Melastomataceae. ner 1989b). The vast majority of the total species offer Chemical characters that would underline the naturalness pollen as a reward and are pollinated by bees that use of certain intrafamilial groups have not yet been found. thoracic vibrations which are transmitted to the anthers to collect the pollen. Nectar production is rare in Melasto- mataceae, but occurs in 70-80 neotropical species be- longing to Blakea, , Meriania, , Brach- yoturn, Tibouchinu, Chalybea and Huilaea, Charianthus, Phylogenetic analysis Mecranium, and Miconia (references in Renner 1989b). The following 21 characters were analyzed in repre- Melastome nectaries lack histologically differentiated sentatives of all genera of Melastomataceae and Meme- nectariferous tissue and may be located on the filaments cylaceae using herbarium material (studied during visits near the insertion of the connective, at the filament in- to BK, BKF, BM, C, E, K, L, and US) and literature (cited sertion into the hypanthium, or at the petal tips (Medinilla above and in Tab. 2) and in a few cases fresh material: rnagnifca; Tococa guianensis; Tobe et al. 1989; Rut- growth form, leaf venation, presence and degree of ani- ishauser, pers. comm. 1991). According to Tobe and sophylly, presence and type of domatia, inflorescence collaborators (Stein & Tobe 1989; Tobe et al. 1989) they position and architecture, floral bracts (presence, shape, are unique in Myrtales and perhaps among angiosperms whether persistent), calyx type (whether persistent) and as a whole. Nectar secretion so far is known only from shape, sex (hermaphrodite or unisexual), mery, New World Melastomeae, Merianieae, Miconieae, and number, mode of anther dehiscence, presence and Blakeeae, but the status of knowledge in this area and the degree of stamen dimorphism, presence and type of con- unstructured nature of melastome nectaries make it im- nective appendages, presence of a pedoconnective, con- possible so far to use the presence of nectar secretion as a figuration of the ovary apex, ovary locule number, pla- systematic character. centation type, ovule number, fruit type, fruit cross sec- tion, and seed shape. Only seven of these (Tab. 4) were used in the cladistic analyses because they were known for each genus, sufficiently invariable at the level of Pollen morphology investigation, and of reasonably likely homology; all aut- Based on the limited number of studies, c. 110 species in apomorphies were deleted from the data matrix. The 30 genera (Patel et al. 1984; Renner 1989a; Skean, in analyses were performed using the program PAUP ver- press; Nowicke, unpubl.; Almeda, unpubl.; Hansen, un- publ.), there exists little variation in pollen morphology. The pollen small, tricolporate, radially symmetrical, Tab. 5. Data matrix for the Melastomataceae. The Kibessieae are is outgroup for the remainder of the Melastomataeae. and isopolar; occasionally grains may have 4, 5 or more colpi, and some grains are dicolporate or heteropolar. Melastomataceae character states Faint to distinct pseudocolpi (or subsidiary colpi, in the terminology of Patel et al. (1984)) are usually present. Kibessieae 00 00 000 The surface sculpture is smooth and striate or rugulate or Astronieae 10 00 000 rugulate-verrucate. Polyads and tetrads known from Sonerileae 11 00 101 are Merianieae 11 01 001 Tococa spadiciforu and Miconia melanotricha (Patel et Microlicieae 11 10 001 al. 1984; Skvarla, pers. comm.). Melastomeae 11 10 101 Contrary to earlier statements, mature pollen of Melas- Rhexieae 11 01 001 tomataceae contains two, not three cells (Tobe & Raven Miconieae II 00 011 Blakeeae 11 00 011 1984).

530 Nord. J. Bot. 13 (5) (1993) Kibessieae Kibessieae

Astronieae Astronieae

Sonerileae Sonerileae

Merianieae Melastorneae

Rhexieae Merianieae

Microlicieae R hexieae

Melastorneae Microlicieae

Miconieae Miconieae

Blakeeae Blakeeae

Kibessieae

Astronieae Astronieae // Sonerileae / ,I Sonerileae

Melastorneae Microlicieae

Merianieae

Rhexieae

Miconieae I Miconieae

Blakeeae '7 Blakeeae

Fig. 4. 14. Topologies of the four equally parsimonious trees generated by PAUP in an exhaustive search with Kibessieae as functional outgroup; length = 8; C.I. (excluding uninformative characters) = 0.86. Characters: 1. libriform fibers; 2. anther dehiscence by pores; 3. pedoconnective; 4. dorsal connective appendages; 5. ovary crowned; 6. true berries; 7. consistently axillary placentation; single lines = synapomorphies, double lines = homoplasies, x = losses (reversals). See Tab. 4 and text for character states and explanation of characters. I : preferred hypothesis.

Nord. 1. Bot. 13 (5) (1993) 53 1 sion 3.0s (Swofford 1991) on a Macintosh PC, and the calfe & Chalk 1983, quoting Solereder) and may be characters were polarized as discussed in the preceding present or absent even within single genera, viz. Strych- section and as shown in Tab. 4. The Kibessieae was used nos (Baas, in litt.) and the roots of Lythrum salicaria to root the cladogram because they retain the ancestral (Metcalfe & Chalk 1950). An independent origin of the condition in all characters employed in the analysis (cf. axially included phloem in Memecylaceae and Pternan- data matrix, Tab. 5). dra is supported by the fact that vessel ray pits in Pter- The units accepted as monophyletic and employed in nandra are simple (like those of other Melastomataceae) the analysis are those appearing as headings in the fol- whereas Memecylaceae have half-bordered pits. Radially lowing discussion; they are formally recognized at the included phloem, on the other hand, while not known tribal level to conform as much as possible with taxo- from Memecylaceae has been found in other Melastoma- nomic tradition in the family. Their composition may be taceae (Medinilla). found in Tab. 2 in which all genera currently recognized in the family are assigned to tribe. Since the discussion that follows incorporates much information presented in Astronieae Melastomataceae characters: variation and polarization, references to morphological studies will not be repeated This paleotropical group of four genera (Maxwell & but may be found above under the respective character Veldkamp 1990a, b) is macromorphologically, leaf ana- type. tomically, and biogeographically very homogeneous. As- An exhaustive search using the data matrix shown in tronia with 59 and Astronidium with 67 species differ Tab. 5 yielded four equally parsimonious 8-step trees from one another mainly in stigma and placenta shape with a consistency index of 0.86 (excluding uninforma- and the smaller Beccarianthus (22 species) is separated tive characters). The alternative topologies shown in Fig. on calyx and capsule morphology. The monotypic Astro- 4 assume homoplasy (including losses) for either pedo- calyx, finally, is simply a pleiostemonous Astronia. As- connectives, dorsal connective appendages, or ovary tronieae share several plesiomorphic character states, crowns. Tree 1, hypothesizing the independent evolution such as a woody habit, dry capsules with cuneate or linear of ovary crowns in Melastomeae and Sonerileae s. s., was seeds, basal or axile-basal placentation, longicidal anther selected as the preferred hypothesis because this seems opening, and a complex vascular pattern in the midrib intuitively likely (compare discussion of this character (Mentink & Baas 1992), but seem to lack an autapo- under Gynoecium). Less likely seem a parallel evolution morphy. A synapomorphic trait joining them with the of pedoconnectives in Melastomeae and Microlicieae Melastomatoideae is the possession of libriform fibers (tree 2) or a loss of ovary crowns in Microlicieae (tree 3). (van Tieghem 1891a; Vliet 1981). Tree 4, which assumes a loss of pedoconnectives in Sbne- rileae, is contradicted by the present distribution of Sone- rileae, Melastomeae, and Microlicieae (see below). Addi- Sonerileae tional evidence from characters not used in the formal analysis as well as distribution data will be considered in The Sonerileae s. 1. occur mainly in Southeast Asia and the further discussion of the preferred cladogram. Madagascar, whereas they have few species in the ne- otropics; they may comprise 560-600 species in 40 gen- era. The clade is heterogeneous, and the synapomorphic Kibessieae character accepted in the cladistic analysis as character- izing it, namely the apically crowned ovaries, is only The Kibessieae, comprising only Pternandra (incl. Ki- found in two-thirds of the genera. I have recognized two bessia) with 15 species in southeast Asia (Maxwell provisional alliances within the Sonerileae s. 1. (Tab. 2), 1981), are rich in autapomorphies, such as median-parie- the -Anerincleistus alliance and the Sonerila- tal placentation, antepetalous ovary locules, and radially Bertolonia-Gravesia group, but the distinction between included phloem, but share no derived traits with the the two is weak, and for example Brediu and Driesseniu remainder of the Melastomataceae other than the acrodro- could go in both (compare Tieghem 1891a; Cogniaux mous venation, the synapomorphy of the family. As is 1891; Diels 1932; Hansen 1985b). The first alliance com- apparent from the data matrix (Tab. 5) Kibessieae are prises mainly shrubs with terminal or axillary, umbellate, here regarded as the sister taxon to all other tribes and, fasciculate, or thyrsoid paniculate cymes (scorpioid in following Naudin (1 849-1953), formally recognized as a some Neodriessenia) and 8- or 10-ribbed, occasionally subfamily (Tab. 2). In wood anatomy, Pternandra stands tetrangular (Bredia) capsules. The second consists mainly out in having axially (in addition to radially) included of herbs with often uniparous (scorpioid) inflorescences, phloem and fiber tracheids, and it has therefore been more rarely with paniculate cymes or flower clusters and proposed as a link between the Melastomataceae and typically 3- or 4-angular capsules. The core of the first Memecylaceae (Thorne in R. Dahlgren & Thorne 1984: group is made up of Oxyspora, Anerincleistus, Poilan- 677). Interxylary phloem, however, is common in alli- nammia, Bredia, Barthea, Driessenia, Neodriessenia, ances with intraxylary phloem such as the Myrtales (Met- Blastus, and Stussenia. The second group in Asia com-

532 Nord. I. Bot. 13 (5) (1993) prises Sonerila, , and Phyllagathis (incl. mainly on account of their stamens and seeds. The seeds Tigridopalma) and the satellites Scorpiothyrsus, Aschis- of Pachyloma, however, are unlike those of all other tanthera, and Kerriothyrsus plus a few small or mono- Merianieae. In some of these genera the dorsal connec- typic genera (Tab. 2). tive appendages are present in the larger stamens only Most Sonerileae are herbaceous of shady hab- and moreover are bifid and curved so that they appear itats, often with a basal whorl of large, somewhat turges- like ventral appendages (compare illustrations of Tess- cent leaves, sometimes with tubers. As in many herb- mannianthus in Wurdack 1989, and Almeda 1989b, aceous melastomes, there is a pronounced tendency to 1990b). More difficult to place are the monotypic, prob- reductions in the androecium, with the inner set of sta- ably ancient Guayana groups Neblinanthera (Wurdack mens becoming staminodial. Old World Sonerileae usu- 1964a) and Ochthephilus (Wurdack 1972). Ochthephilus ally have isomerous flowers whereas New World ones differs from all other Merianieae in being herbaceous. typically are anisomerous (5-merous androecia with 3- locular ovaries). Exceptions are the neotropical Trysso- phyton, which is isomerous (Wurdack 1964a: 156) and the paleotropical Calvoa, which is anisomerous. Rhexieae Most Asiatic Sonerileae belong to the taxonomically In the preferred cladogram (Fig. 4.1 .) Rhexia appears as very poorly understood genus Sonerila, while the major- the sister group to Merianieae on the basis of shared ity of African and Madagascan Sonerileae belongs to stamen characters (dorsally thickened connectives, lack Gravesia and its close relatives Amphiblemma and Cal- of pedoconnectives), but the evidence is weak. Rhexia is voa. More isolated are the small or monotypic Cincinno- the only genus of Melastomataceae with an entirely North botrys, Dicellandra, Preussiella, and Feliciadamia, the American distribution and rich in autapomorphies, such last recently elevated to tribal status (Jacques-Fklix, in as atropous ovules, capsules from completely inferior litt. 1991) because of its capsular fruits that develop from ovaries, locules in front of the petals instead of alternat- an epigynous ovary. This condition, however, has ing with them, and mature unilocular anthers. The co- evolved repeatedly in the family (see Melastomataceae chleate seeds found in most but not all species of Rhexia characters: variation and polarization), and on its own develop from atropous ovules and therefore must have does not have much significance. The neotropical Soneri- originated independently from the cochleate seeds of Me- leae, finally, consist of a core made up by Bertolonia, lastomeae, which develop from anatropous ovules. Pa- Macrocentrum, Monolena, , , and Di- chyloma, placed next to Rhexia by Triana and Cogniaux, plarpea and some phenetically more isolated groups, shares no derived characters with that genus and is here such as Maguireanthus, Opisthocentra, Tateanthus, and regarded as merianiean. Boyania (Wurdack 1964a, Solt & Wurdack 1980). Al- meda (1977 and in press) has suggested that Centradenia, Centradeniastrum, and a new central American genus too Melastomeae are soneriloid, mainly because of their pyramidal seeds. Two relatively strong characters support a clade con- A huge well-defined clade (c. 850 species in 47 genera) sisting of the Sonerileae and all following tribes (Fig. 4), the members of which share the following synapomor- namely fixed axillary placentation and poricidal anthers. phies: cochleate seeds, stamens with pedoconnectives, By contrast, the Sonerileae themselves, whether in the and ovaries typically crowned by persistent trichomes. strict sense (excluding the Oxyspora-Anerincleistus alli- Melastomeae are mainly annual or perennial herbs, sub- ance) or in the broad sense accepted here, are a poorly shrubs, or shrubs, rarely treelets. Inflorescences are ter- defined group. Ontogenetic studies of sonerilean fruits minal paniculate or rarely glomerulate cymes; occasion- are planned. ally they are few-flowered and axillary or consist of solitary flowers. Flowers are usually diplostemonous with unequal stamens, or one whorl is staminodial or lacking (some species of Tibouchina, Monochaetum, Aci- Merianieae santhera, , and Poteranthera). The capsules This clade, the core of which is made up by Axinaea, of neotropical representatives are dry and have wind- Meriania, Centronia, , Adelobotrys, and dispersed seeds, those of some paleotropical species tear Phainantha, consists almost entirely of treelets, shrubs, open to expose fleshy placentae (cf. Melastomataceae or woody climbers with dry capsules, leathery leaves and characters: variation and polarization) or have true berries rather large flowers with dorsally thickened and variously (Melastoma orientale). The adaptive significance of this spurred stamens without pedoconnectives. It may com- and also of the campylotropous seeds has already been prise 16 genera and c. 220 species. A number of small discussed (Melastomataceae characters: variation and po- southeastern Brazilian shrubs (Behuria, Benevidesia, larization). Unusual are the flattened and broadly winged Merianthera, Dolichoura, Huberia, and Bisglaziovia), seeds of Acanthella, and the placement of this genus is the Orinoco-Rio Negro savanna genus Pachyloma, and therefore uncertain. Indumentum complexity in the fam- the Andean Tessmannianthus are added to this core ily reaches its pinacle in the Melastomeae (Wurdack

Nord. J. Bot. 13 (5) (1993) 533 1986), and there are striking parallelisms in hypanthium were proposed by Gleason (1929) to form the tribe Cy- indumentum in, for example, Osbeckia, Dissotis, and phostyleae, which he thought characterized by capsules Pterolepis from Asia, Africa, and South America, respec- developing from inferior ovaries, haplostemonous an- tively. droecia, and calyptrate calyces. However, and as pointed Monochaetum, traditionally placed in the Rhexieae out by Gleason, each of these features has developed (Triana 1871; Cogniaux 1891), belongs in the Melasto- repeatedly in the family (capsules from inferior ovaries in meae on account of its campylotropous seeds (Almeda Melastomeae, Merianieae, Rhexieae, and Sonerileae, ha- 1978), setose ovary apex, wood anatomy (ter Welle & plostemonous androecia in Melastomeae, Microlicieae, Koek-Noorman 198 l), and chromosome number (Al- and Miconieae, and calyptras in Astronieae, Blakeeae, meda & Chuang 1992). Kibessieae, Merianieae, and Miconieae), and it is partic- ularly in the Miconieae that these derived characters abound (viz. haplostemony in Dissochaeta p.p. (incl. Omphalopus), and Miconia p.p.; pleiostemony in some Microlicieae Clidemia, Conostegia, Llewelynia, Miconia, and Pleth- Microlicia, Chaetostoma, Lavoisiera, Rhynchanthera, iandra; calyptras in Bellucia, Centronia, Conostegia, Trembleya, and Carnbessedesia form a cohesive assem- Llewelynia, Mecranium, Myriaspora, and some Tococa; blage centered in south-central Brazilian savannas. Auta- for miconiean genera with ant and mite domatia, nectar pomorphic characters are the straight, sometimes slightly secretion, or woody dimbers compare the section on winged seeds with a usually foveolate surface (SEMs: Melastomataceae characters). Whiffin & Tomb 1972; Renner 1990) and shrubby, often microphyllous habit. A synapomorphy with Melastomeae is the presence of pedoconnectives (Fig. 4). Problematic is Bucquetia, an Andean group of three species placed variously in Microlicieae (Triana 187 1; Blakeeae Cogniaux 1891 ), Melastomeae-Tibouchineae (Baillon Blakea and Topobea, two extremely close taxa (Gleason 1877; van Tieghem 1891a; Krasser 1893), and Meria- 1945; Wurdack 1957; Almeda 1990a), are traditionally nieae (Wurdack 1980). Wood-anatomically it is anoma- separated from the Miconieae because of their persistent lous in any of these alliances because of the mainly floral bracts, axillary flowers, and conspicuously cross- procumbent ray cells (ter Welle & Koek-Noorman 1981: venulate leaves with often very numerous, strictly paral- 372). The seeds are pyramidal, the ovary apex glabrous or lel lateral veins. They share baccate fruits with seeds setose, and the stamens simple. Bucquetia is here placed embedded in pulp with Miconieae (Fig. 4), and are also in the Microlicieae because that is where it seems to closest to Miconieae in wood anatomy (Koek-Noorman disagree least. et al. 1979). Possibly therefore Miconieae become par- The monotypic genera Castratella, Eriocnema, and aphyletic by ranking Blakeeae at the tribal level. On the Lithobium differ from other Microlicieae in being herb- other hand, little is gained at present by including this aceous and Lithobium in addition by having 3-merous distinct clade in the already huge Miconieae. flowers. Indeed, Baillon (1877) and Krasser (1893) held Eriocnema to be soneriloid (bertolonioid).

Miconieae A new classification of the The Miconieae comprise c. 2200 species in 38 genera mostly of shrubs, trees, and treelets and are distinguished Melastomataceae by epigynous ovaries that usually develop into baccate The arrangement proposed in Tab. 2 and for which the fruits typically dispersed by birds. In some Old World rationale has been presented above differs from the tradi- Miconieae, such as Dissochaera, Medinilla, and Macro- tional one (Cogniaux 1891) as follows: (1) the Memecy- lenes, the fusion of ovary and hypanthium is incomplete laceae are removed from the Melastomataceae, (2) Pter- because there are deep pockets in the tissue between the nandra is placed in a subfamily of its own, (3) the ovary wall and receptacle, corresponding to the bud posi- Astronioideae are abolished, (4) three neotropical and tions of the stamens; and in some New World genera the paleotropical tribal pairs are merged (Bertolonieae with hypanthium and ovary wall do not become fleshy in fruit Sonerileae-Oxysporeae, Tibouchineae with Melasto- (Allomaieta, Alloneuron, Cyphostyla). Kendrickia, a meae, and Miconieae with Dissochaeteae), (5) the Cy- woody climber, traditionally placed in the Oxysporeae phostyleae are submerged in the Miconieae, and (6) because of its dry fruit, is here placed in the Miconieae Rhexia is placed in a tribe of its own. As pointed out in because it appears closely related to Medinilla in habit the historical review of melastome classification, most of and floral morphology. The Andean genera Allomaieta, these changes have also been suggested by other workers Alloneuron, and Cyphostylu (with together ten species) or present a return to pre-Triana classificatory schemes.

534 Nord. J. Bot. 13 (5) (1993) S. America: Blakeeae (UlSO) Melastomeae (301550) Merianieae (16/220) Miconieae (30/1700) Microlicieae (11/21 0) Sonerileae (13190)

Fig. 5. Preferred phylogenetic hypothesis and present day distribution of Melastornataceae superposed on a Late Cretaceous (80 m.y.b.p.) reconstruction to indicate continental locations during initial tribal diversification. Figures in brackets indicate the number of generdspecies in the different areas.

and Pliocene pollen has been reported from Costa Rica Geographical distribution and fossil and Panama (A. Graham 1987, 1991). Unfortunately me- lastome pollen is of a general type, similar to pollen of evidence several other myrtalean families, and may be difficult to The geographic distribution of all genera is given in Tab. assign with certainty (Pate1 et al. 1984). Macrofossils, 2, and these data will now be used, together with the however, clearly show that the Melastomataceae were fossil record and the hypothesized phylogeny (Fig. 4), to present by the early Tertiary. Thus, there are Eocene develop a broad picture of the evolutionary history of the leaves from Colombia (Huertas 1977), Brazil (Duarte family. 1956), North America (Hickey 1977), and Borneo The acrodromous venation emerges as the single most (Geyler 1887), and an Upper Miocene leaf has been important character of Melastomataceae, acquired before reported from Sumatra (Krausel 1929). The most in- the poricidal mode of anther dehiscence. Based on wood teresting record in terms of the biogeographic history of anatomy there are two major groups, Kibessioideae and the family is an Antarctic leaf from Tertiary deposits on Melastomatoideae, and they are supported by placenta- Seymour Island (DusCn 1908). Although a critical mod- tion characters: median-parietal placentas are typical of em investigation is lacking, DusCn’s illustration shows a Kibessioideae and axillary ones of Melastomatoideae in- leaf with typical melastome venation, and the identifica- cluding Astronieae (since the axile-basal placentation of tion appears reliable according to E. Zastawniak who has the latter ontogenetically derives from axillary placentas). studied the specimen (Pingen, in litt. 1992). The fossil record of the Melastomataceae has recently Combining all available evidence, the most likely been reviewed (Pingen & Collinson, in press). There is place of origin of the Melastomataceae appears to be East one tentatively acceptable pollen fossil from the Paleo- Gondwana (Antarctica + Australia + New Guinea; Fig. 5): cene of Colombia (Hammen & Garcia de Mutis 1966), because the family exhibits much greater fundamental which would be the oldest record of the family; Miocene diversity in the east-Gondwanan region; because the most

Nord. J. Bot. 13 (5) (1993) 535 plesiomorphic groups (Kibessieae and Astronieae) occur systematic significance in some Mexican Melastomataceae. - Svst. Bot. 17: 583-593. the region; there record in Malesian because is a fossil - & Whiffin, T. 1980. Pilocosta, a new genus of tropical from Antarctica; and because there are three tribes that American Melastomataceae. - Syst. Bot. 5: 294-311. span from Southeast Asia to South America. Africa has a Baas, P. 1981. A note on stomata1 types and crystals in the number of Melastomeae (13 genera) and Sonerileae (7 leaves of Melastomataceae. - Blumea 27: 475479. Baillon, H. E. 1877. Monographie des MClastomacCes. Histoire genera), but lacks all other tribes, most notably the Mico- des plantes 7: 1-65. - L. Hachette et Cie., Paris, London, nieae, except for two introduced species of Medinilla. Leipzig. This may indicate that the Miconieae originated at a time Bakhuizen van den Brink Jr., R. C. [1943] 1946-1947. A contri- when the distance between South Africa and East Gond- bution to the knowledge of the Melastomataceae occurring in the Malay Archipelago, especially in the Netherlands East wana was already too large for Africa to be entered from Indies. - Rec. Trav. Bot. NCerl. 40: 1-391. (Reprinted in the south, while it was still possible to reach South Meded. Bot. Mus. Herb. Rijks Univ. 91: 1-391. 1947.) America. The suggestion of Madagascar as a possible Baumgratz, J. F. A. 1989-1990. 0 g&nero Berfoloniu Raddi cradle (Femandez & Femandez 1954) is unlikely because (Melastomataceae): RevisZo taxon6mica e consideraGes ana- t6micas. - Arq. Jard. Bot. Rio de Janeiro 30: 69-213. Madagascar harbours none of the basal groups. It does Bentham, G. & Hooker, J. D. (eds). 1867. Genera plantarum. - have some Sonerileae (Gravesia) and Melastomeae (a Love11 Reeve & Co., London, pp. 725-773. (Melastomaceae few species of Osbeckia, an introduced species of Melas- by J. D. Hooker). toma, a few representatives of African genera (Dissotis, Beusekom-Osinga, R. J. van & Beusekom, C. F. van 1975. Delimitation and subdivision of the Crypteroniaceae (Myr- , Antherotoma, Tristemma) and two en- tales). - Blumea 22: 255-266. demics, Amphorocalyx and Dionycha), but lacks all Mi- Blume, C. L. 1826. Bijdragen tot de flora van Nederlandsch conieae except Medinilla (Perrier de la Bfithie 1951). Indie. - Batavia. Osbeckia and Medinilla probably arrived in Madagascar Bouman, F. & Boesewinkel, F. D. 1991. The campylotropous ovules and seeds. their structure and functions. - Bot. Jahrb. from Asia where they comprise numerous species, Syst. 113: 255-270. whereas they have no native representatives in Africa. Bremer, K. 1979. Taxonomy of Memecylon (Melastomataceae) I. India, suggested by Nayar (1972) as a place of origin for in Ceylon. - Opera Bo< 50: 1-32. the Melastomataceae, can be excluded because of the - 1981. Seeds and embryos in Skri Lanka (Ceylonese) species unbalanced representation of the family there, which con- of Memecylon, with notes on Spnfhandru (Melastomata- ceae). - Nord. J. Bot. 1: 6245. sists of weeds and relatively derived species, e.g., of - 1982. , a re-established paleotropical genus of the Sonerila (Lundin, pers. corn. 1991). Melastomataceae - Memecylaceae. - Nord. J. Bot. 2: 121- A west-Gondwanan origin, favoured by Raven & Ax- 124. elrod (1974), is contradicted by the present distribution of - 1983. Taxonomy of Memecylon (Melastomataceae) in Bor- neo. - Opera Bot. 69: 147. the tribes and the restriction of the Kibessieae and Astro- - 1988. Melastomataceae. - In: Dussanayake, M. D. and Fos- nieae to Malesia, while an east-Gondwanan origin not berg, F. R. (eds), A revised handbook to the Flora of Ceylon. only agrees with the fossil record but also with a possible - Faridabad, India. Vol. 6, pp. 157-240. ancestry from crypteronioid-lythroid stock. Brown, R. 1818. Observations ... on ... C. Smith’s collection of plants from the vicinity of the river Congo. Appendix 5, p. 434. - In: Tuckey, J. K., Narrative of an expedition to Acknowledgements - I thank P. Baas, S. Graham, W. Judd, J. explore the river Zaire, usually called the Congo, ... in 18 16 Kadereit, T. Morley, and J. Wurdack for helpful comments on ... . London. (Reprinted 1866 in: Bennett, J.J. (ed.), The the manuscript; P. Baas, H. Mentink, M. Pingen, and F. Almeda miscellaneous botanical works of Robert Brown. R. Hard- for letting me see at the time unpublished manuscripts; and K. wicke, London. Vol. 1, p. 18.) Larsen for an opportunity to see Old World Melastomataceae Burtt, B. L. 1991. On cryptocotylar germination in . during the 1992 Thai-Danish expedition. - Bot. Jahrb. Syst. 113: 429442. Candolle, A. P. de 1828a. Melastomaceae. - Prodr. 3: 99-102. - 1828b. MCmoires sur la famille des MClastomacCes. - Paris. Candolle, A. L. P. P. de 1868. 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