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Toward a Phylogeny of Maxillariinae Orchids: Multidisciplinary Studies with Emphasis on Brazilian Species

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Toward a Phylogeny of Maxillariinae Orchids: Multidisciplinary Studies with Emphasis on Brazilian Species LANKESTERIANA 7: 57-60. 2003. TOWARD A PHYLOGENY OF MAXILLARIINAE ORCHIDS: MULTIDISCIPLINARY STUDIES WITH EMPHASIS ON BRAZILIAN SPECIES RODRIGO B. SINGER & SAMANTHA KOEHLER Departamento de Botânica, I.B., Universidade Estadual de Campinas Campinas, SP, Brazil, 13083-970. [email protected] It has been demonstrated that the neotropical sub- trnL-F, matK, atpb) of Brazilian Maxillariinae species tribe Maxillariinae comprises a well-supported mono- with emphasis on Brazilian endemic groups, in col- phyletic orchid group, according to cladistic analyses laboration with W.M. Whitten and N.H. Williams of nuclear and chloroplast DNA regions (Whitten et (Florida Museum of Natural History, University of al. 2000). The results presented by Whitten et al. Florida, U.S.A.). Here we present a brief account of (2000) brought to light significant changes concern- the results of morphological studies, with emphasis ing the delimitation of the subtribe Maxillariinae. on variability of flower features, especially pollinari- First, in order to avoid the description of an additional um morphology, among different alliances of subtribe within the Maxillarieae to place the genus Brazilian Maxillariinae species. Infrageneric classifi- Xylobium Lindl., orchids formerly assigned to the cation into alliances follows Pabst & Dungs (1977). subtribes Lycastinae and Bifrenariinae (Dressler Acquisition of specimens. The existence of two large 1993), including Xylobium, were merged into a living orchid collections in the state of São Paulo, broader Maxillariinae. This taxonomic decision is located at the "Instituto de Botânica de São Paulo" also supported by floral and vegetative characters. and at the "Escola Superior de Agronomia Luiz de Second, it has been demonstrated that Maxillaria Queiroz" has allowed the collection of samples for Ruiz & Pav., by far the largest genus in the subtribe, DNA studies as well as of fresh flowers for morpho- is polyphyletic, since species of Cryptocentrum logical studies. The cultivation of plants also makes Benth., Chrysocycnis Linden & Rchb.f., Mormolyca possible the observation and comparison of the vege- Fenzl and Trigonidium Lindl. appeared nested within tative architecture of different species. Another Maxillaria (M. Whitten and N. Williams, pers. advantage of such complete living collections is the comm.). Once again, this seems to be a very reason- possibility to conduct breeding system studies as well able finding, since Maxillaria as currently accepted as to observe eventual pollinators, as many bee comprises plants with extremely dissimilar vegetative species occur in the area of the collections. Voucher architectures. materials of the individuals sampled are being In order to contribute with the reevaluation of the deposited at the SP and UEC herbaria. generic boundaries within the subtribe Maxillariinae (sensu Whitten), a team of Brazilian researchers, Morphological features. Vegetative features are including molecular and field biologists as well as known to present large variation within the chemists has been brought together in a multidiscipli- Maxillariinae. Plants may or may not bear pseudob- nary project. Our goals are (1) to conduct morpholog- ulbs and the rhizome, when present, may be long and ical studies of vegetative and flower features of repre- apparent or extremely short. The number of leaves sentative species; (2) to determine the chemical struc- may vary from one to three in Brazilian species. ture of flower rewards and fragrances of representa- Flower features are quite conservative. The lip is tive species; (3) to perform pollination and breeding articulated (rarely fused) to the base of the column. system studies with representative species and; (4) to The anther is incumbent and holds a pollinarium with perform sequencing of multiple DNA regions (nrITS, four pollinia. The pollinarium always bears a well- 58 LANKESTERIANA Nº 7 Figure 1. Examples of studied taxa of Maxillariinae. A-C. Maxillaria picta Hook. A. Flower in lateral view. B. Column, with lip hinged at its base. C. Pollinarium. D-F. Maxillaria parviflora (Poepp. & Endl.) Garay. D. Flower in lat- eral view. E. Lip and column in lateral view. F. Pollinarium. G-I. Trigonidium obtusum Lindl. G. Flower in lateral view. H. Column with lip hinged at its base. I. Pollinarium. Mayo 2003 SINGER & KOEHLER - Phylogeny of Maxillariinaes 59 developed viscidium. Pollinium stalks (namely, tegu- Brazilian Maxillariinae. Flower, and particularly, the lar stipes) may or not be present. pollinarium structure are well suited for this kind of Some vegetative and flower features appeared to be pollinators. The well-developed semilunar viscidium highly conservative within the Maxillaria alliances embraces the scutellum of the Hymenopteran pollina- studied. Interestingly, these morphological conserva- tor when the insect retreats the flower. Such pollinari- tive alliances appeared as monophyletic groups um deposition is ecologically significant, since the according to preliminary molecular data (M. Whitten, scutellum is a difficult place for the insects to clean. pers. comm.). The 'Maxillaria picta' alliance compris- Noteworthy, Maxillaria species that present round, es species whose pseudobulbs are always bifoliate. pad-like viscidia, such as Maxillaria parviflora Besides, flowers of this orchid assemblage are gener- (Poepp. & Endl.) Garay, have pollinaria deposition on ally rewardless and present pollinaria devoid of the face of their pollinators. To date, we have record- stipes. The 'Maxillaria madida' alliance possesses a ed bumble-bee and Euglossine pollination in different and remarkable situation. Whereas vegeta- Bifrenaria harrisoniae (Hook.) Rchb.f., ant and bee tive features, such as the number and width of the (Meliponini) pollination in M. parviflora, and leaves and length of the rhizome, are quite variable, Meliponinae and wasp pollination in several species the flowers present an extremely similar morphology. of the 'Maxillaria picta' alliance. Pollination through Up to this moment, species from this alliance have pseudocopulation mediated by Plebeia (Meliponinae) shown to be rewardless, the flowers being yellowish, drones was recently confirmed in Trigonidium brownish or vinaceous and displaying a shining, obtusum Lindl. (Singer 2002). We also have found smooth labellum. Pollinarium structure is also very nectar in M. rigida Barb. Rodr. and M. parviflora. conservative within this alliance, with most species In the past, flower features have been widely used bearing a semilunar, often thickened viscidium, in orchid taxonomy. However, it seems that vegeta- attached to large tegular stipes. tive features are less subject to homoplasy in the Moreover, both the 'Maxillaria discolor' and the subtribe Maxillariinae and we should pay more 'Maxillaria valenzuelana' alliances present rewarding attention to these characters in phylogenetic studies. flowers (with trichomes, wax-like or resin-like com- A good example of this is the old concept of pounds) and very similar pollinarium structure, since Oncidium Sw. Until a few years ago, Oncidium was most species show a semilunar viscidium and large a vegetative diverse genus with more than 400 tegular stipes. Preliminary molecular data suggest species, composed of plants with more or less simi- these alliances are close related (M. Whitten, pers. lar flowers (predominantly yellow, with some brown comm.). They include species either with unifoliate spotting, quite often bearing complex elaiophores). pseudobulbs or without pseudobulbs, as Maxillaria Williams et al. (2001) have demonstrated that the valenzuelana (A. Rich.) Nash and M. equitans genus Oncidium, as formely circumscribed, repre- (Schltr.) Garay, respectively. Similar vegetative shifts sents a polyphyletic assemblage of plants defined by (unifoliate pseudobulbs to pseudobulbless plants) its morphologically similar flowers which are attrac- have already been verified in well-supported mono- tive to similar pollinators (mostly oil-gathering bees phyletic groups of Oncidiinae, namely the Notylia- of family Apidae). Instead, "Oncidium-like" flowers Macroclinium and Erycina-Psygmorchis complexes have arisen several times in subtribe Oncidiinae (Williams et al. 2001). The loss of the pseudobulb in (Williams et al. 2001). Maxillariinae may represent a heterochronic shift with retention of seedling morphological features ACKNOWLEDGEMENTS. Both authors acknowledge their (neoteny), as already suggested for Oncidiinae (Chase advisor, Maria do Carmo E. Amaral, from the Botany 1986). Department of Universidade Estadual de Campinas, Brazil. This contribution was made possible through grants con- Reproductive biology. Field observations by R.B. ferred by FAPESP (Fundação ao Amparo à Pesquisa do Singer indicate that Hymenoptera, especially Estado de São Paulo, processes 01/08958-1 and 02/02161- Meliponinae bees, are the main pollinators of 7) for both authors. 60 LANKESTERIANA Nº 7 LITERATURE CITED Whitten, W.M., N.H. Williams & M.W. Chase. 2000. Chase, M.W. 1986. A reappraisal of the Oncidioid Subtribal and generic relationship of Maxillarieae Orchids. Syst. Bot. 11 (3): 477-491. (Orchidaceae) with emphasis on Stanhopeinae: com- Dressler, R.L. 1993. Phylogeny and classification of the bined molecular evidence. Amer. J. Bot. 87 (12): 1842- orchid family. Dioscorides Press, Oregon. 1856. Pabst, G. & F. Dungs. 1977. Orchidaceae Brasilienses Vol. Williams, N.H., M.W. Chase, T. Fulcher & W.M. Whitten. 2. Hildesheim, Brucke. 2001. Molecular Systematics of the
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