THE PHILIPPINE RUBIACEAE GENERA : UPDATED SYNOPSIS IN INTKEY DATABASES OF THE DELTA SYSTEM

Changes in generic concepts and delimitation resulted in 80 genera (6 of which are cultivated) of Philippine Rubiaceae. Generic descriptions of all genera are coded in DELTA format (Version 2000) and translated by the directive CONFOR into INTKEY database for interactive identification and information retrieval. The descriptions incorporate mostly compiled data of Philippine Rubiaceae species and observations on over 130 characters. Recently accepted classification (subfamily, tribes, and subtribes) and significant characters defined by Robbrecht (1988, 1993) are appended to the generic descriptions. The database is available on the World Wide Web at http://www.uni-bayreuth.de/ departments/planta2/wgl/delta_ru/index.html.


INTRODUCTION
The family Rubiaceae is the fourth largest in number among vascular flora comprising 659 genera and approximately 13,000 species (Robbrecht, 1988).In the Flora Malesiana region, it is estimated only second to the Orchidaceae with about 150 genera and 1,830 species.The most recent enumeration of Philippine Rubiaceae genera is nearly eighty years old (Merrill, 1923) and out of date in many respects.Furthermore, the dearth of generic descriptions essential to the understanding of Rubiaceae in the Philippines prompted us establish an automated databank for Philippine Rubiaceae genera under the DELTA system.
The present study endeavours to contribute to the understanding of Philippine Rubiaceae.It also ascertains the comparability of this study with other research efforts both on the Philippine flora and on the Taxonomy and Systematics of Rubiaceae worldwide.Moreover, it also validates the new names and combinations with the view to help scientists or foresters interested to get acquainted with the various nomenclatural and taxonomic changes that have occurred in the Rubiaceae of the Philippines.This up-to-date information on the genera of Rubiaceae occurring in the Philippines is an important first step towards a comprehensive treatment of the family for the island and an important contribution to the Flora of the Philippines project, since no recent survey has been done after Merrill's (1923) work.
The lack of comprehensive generic descriptions for Philippine Rubiaceae and the need for an interactive identification tool combined with information retrieval have initiated the use of a DELTA database.The DELTA format is a versatile tool for the storage and management of descriptive taxonomic data (Dallwitz, 1980) while the DELTA system is an integrated set of programs based on the use of the DELTA format.It is advantageous for the first stage in a project as large as the Philippine Rubiaceae, because it is easily accessible and versatile enough to allow corrections and additions throughout the whole project.

MATERIAL AND METHODS
The DELTA format is mainly composed of 3 files (Dallwitz et al., 1999): 1) a character file (CHARS); 2) a specification file (SPECS), describing character traits like number and type of characters; and 3) a taxon description file (ITEMS).The data are gathered against a character list which can be viewed also on the website.The characters and character states were compiled mostly by consulting literature sources, direct observation, and measurements on herbarium specimens borrowed from the Philippine National Herbarium (PNH), the National Herbarium Netherlands (L), and the National Botanical Garden of Belgium (BR).The list encompasses a total of 139 characters used to generate a natural language description of the Philippine Rubiaceae genera.Likewise, the character list gives information on questions of character definition, and includes other comments which do not appear in the printed taxon descriptions.
A sample of the genus descriptions, generated from the databank by CONFOR and typeset automatically, is appended to this article.However, the genus descriptions are continuously improved and revised in the database.Thus, new characters are added from time to time, and existing definitions are improved or rendered less inclusive as more precise data become available.Likewise, characters and character states which are still undesirably inclusive or clumsy will be improved in due course.Against this background, an internet-accessible database has obvious advantages.Moreover, only few images are presently included in the CIMAGES file, and new images are added continuously.
To be able to use the program INTKEY, for interactive identification and information retrieval generated by the directive CONFOR, a PC with Windows 95/NT or later version is needed and the program INTKEY, available at http://biodiversity.uno.edu/delta, must be installed.

Revisions in Philippine Rubiaceae species
During the extensive preliminary studies for the Rubiaceae of the Flora of the Philippines project as well as present revisionary work on Philippine Mussaenda Burm.ex L. and Villaria, a number of name and rank changes has become inevitable which will be discussed briefly in the following.
Generic revisions involving Philippine Rubiaceae were initialized by Bremekamp (1934Bremekamp ( , 1939Bremekamp ( , 1940aBremekamp ( -d, 1947a, b) , b) who published extensive Latin descriptions.Bremekamp (1934Bremekamp ( , 1939) ) published a monograph of the genus Pavetta L. with emphasis on delimitation against the closely related genus Ixora L. He added four new species of Pavetta to the Philippine record.Six years later, Bremekamp (1940b) published a preliminary account of the Philippine species of Urophyllum Jack ex Wall., Pleiocarpidia Schum.& Bremek., and Praravinia Korth.& Bremek.In his investigation of Urophyl lum, he noticed four uncommon features : 1) long-pedicellate flowers arising singly or in groups from leaf axils; 2) presence of acaridomatia on the leaves; 3) different kinds of hairs in the corolla; and 4) linear instead of oblong or ovate anther shape.Based on the unusual character of syngenesious anthers, Bremekamp (1940a) introduced a new endemic genus to the Philippines in the same year -Antherostele.
The four genera mentioned are placed in the same tribe, Urophylleae.Moreover, Bremekamp (1940b) revised the genus Williamsia Merr.and included some Urophyl lum species in the genus Praravinia.However, he stated that, compared to other genera, delimitation of Urophyllum presents more difficulties.Similarly, he encountered difficulties in the classification of the Philippine species which are imperfectly known to the present day.He refrained from describing several Urophyllum species since the herbarium specimens he studied lack reproductive parts (Bremekamp, 1940b).He also experienced the same difficulty in the treatment of the genus Praravinia and most of the descriptions were based on fruiting material only.In this regard, re-investigation of the closely related Philippine species of Urophyllum and Praravinia is needed.
Moreover, Bremekamp published a monograph on the genus Acranthera Arn.ex Meisn. (1947b) andStreblosa Korth. (1947a).He recognized a single species of Acranthera in the Philippines -A.philippinensis Merr.and added one new endemic species and five new varieties of Streblosa.Another early work was made on a systematic arrangement of the closely interrelated Malaysian species of Wendlandia Bartl.ex DC. (Cowan, 1936).Out of four series in Cowan's classification, he grouped the Malaysian Wendlandia in one series -Subinclusae (anthers almost sessile).This series is further subdivided into two subseries, ser.Tinctoriae and ser.Paniculatae, both represented by Philippine species.In 1964, Jayaweera published his treatment of the rubiaceous genus Mussaenda of the Philippine Islands.He added two more Mussaenda species from Merrill's list (1923) endemic to the Philippines and recognized a variety of M. macro phylla Wall.However, Bahadur (1968) remarked that Jayaweera recorded heterostyly in all the Mussaenda species he studied but did not mention dioecism.Heterostyly and dioecism are common on the tribe Mussaendeae (Bahadur, 1968).
The genus Ceriscoides (Hook.f.) Tirveng.has a particularly complex history discussed by Tirvengadum (1978).Tirvengadum (1978Tirvengadum ( , 1983) ) published studies in the tribe Gardenieae, mostly in the Randia-Gardenia assemblage for tropical Asia.Gardenia Ellis was formerly separated from Randia on the basis of ovary structure (one-celled in Gardenia and two-celled in Randia).Tirvengadum (1978) raised the section Ceris coides of Randia to generic status accommodating J.D. Hooker's section Ceriscoides of Gardenia.In the Philippines she transferred only one species from Gardenia to Ceriscoides (Tirvengadum, 1983).
Most of the revisions on Philippine Rubiaceae species were made by Ridsdale (1978aRidsdale ( -c, 1979Ridsdale ( , 1982Ridsdale ( , 1985Ridsdale ( , 1989Ridsdale ( , 1996)).In 1978 he published a worldwide revision on Mitragyna Korth., Uncaria Schreb., and the tribe Naucleeae (Ridsdale, 1978c) of which 6 genera (Ludekia, Metadina, Myrmeconauclea, Neonauclea, Petrusadina, and Nauclea) are represented in the Philippines.He revised the heterogeneous genus Adina and Philippine representatives were placed under Metadina and Pertusadina.His detailed revisions of Nauclea L. and Neonauclea Merr.(Ridsdale,1978b(Ridsdale, , c, 1989) ) have set understanding and delimitations of both genera.He made one combination from Neonauclea to Ludekia, a new genus record to the Philippines.In 1979, he recognized the taxonomic position of Sulitia Merr., an endemic monotypic genus in the Philippines.He transferred the genus from the tribe Cinchoneae to the Gardenieae with reference to the position and form of its inflorescence.Moreover, he published a revision of Badusa (Ridsdale, 1982) describing a new species from Palawan, B. palawensis.However, he recognized B. philippica as dubious and taxonomic re-investigation revealed that this species is a synonym of B. corymbifera (Soerjarto et al., 1996) found in Tonga, Fiji to Solomon Island, and New Guinea.Jansen & Ridsdale (1983) published a revision of the genus Dolicholobium A. Gray.They recognized the two Philippine representatives as conspecific.In 1985, Ridsdale described a new species of Fagerlindia Tirveng.(F.emanuelssoniana Ridsdale) and a new combination, F. microcarpa (Bartl.ex DC.) Ridsdale (Gardenia micropora).Fagerlindia had been segregated from Randia s.l. by Tirvengadum (1983).Ridsdale's (1996) recent published work deals with a review of Aidia in southeast Asia and Malesia.He described one new species and three new combinations for Philippine record.Ridsdale is currently making combinations in Greeniopsis Merr.and Hymenodictyon Wall.included in Table 1 which will be published in the near future.Jansen (1984) published a synopsis of Guettardella Champ.ex Benth., where all the Philippine species of Antirhea were transferred to Guettardella as new combinations and one new species (G.caudata) was described.He assigned all Old World Antirhea species to Guettardella except for two species from the Mascarene Islands.Meanwhile, Chaw & Darwin (1992) made a systematic study of the palaeotropical genus Antirhea including the segregate genus Guettardella.Their extensive study resulted to 36 species of Antirhea, eight out of 13 species in Table 1 are new records for the Philippines.
Ali & Robbrecht (1991) published remarks on Asian and Australian taxa included in Diplospora DC. or Tricalysia A. Rich.ex DC. and showed that the two genera form an artificial assemblage of species.The Philippine Tricalysia species were all transferred to Diplospora leaving Tricalysia as an exclusively African genus.Two new combinations in Diplospora were made but the identity of D. sessilis is yet to be determined.Discospermum Dalzell which is recognized as a synonym of Diplospora, was upheld at generic rank (Ali & Robbrecht, 1991).It is represented by only one species, which had been transferred from Gardenia by Ali & Robbrecht (1991).Huxley & Jebb (1991a, b, 1993) published a series of tuberous epiphytes recognizing five genera (Anthorrhiza, Hydnophytum, Myrmecodia, Myrmephytum, and Squamellaria) and united them in a new subtribe, the Hydnophytinae, on the base of tuber cavity development and form of inflorescences.Three genera of the tuberous epiphytes (Hydnophytum Jack., Myrme codia Jack., and Myrmephytum Becc.) are represented in the Philippines.Revisions are available for Myrmephytum (Huxley & Jebb, 1991b) and Myrmecodia (Huxley & Jebb, 1993).All previously described Philippine Myrmecodia species were subsumed under one species, M. tuberosa Jack.A revision of Philippine species of Hydnophytum is still wanting.In 1993, the tribal affiliation of Scyphiphora (Puff & Rohrhofer, 1993) was changed from tribe Knoxieae (subfam.Antirheoideae) to the tribe Gardenieae (subfam.Ixoroideae).Likewise, the tribe Schradereae was re-examined (Puff et al., 1993).Puff et al. (1993) concluded that Lucinaea DC. is a synonym of Schradera Vahl.In a follow-up study (Puff et al., 1998) all previously described Philippine Lucinaea species were transferred to Schradera.One new combination and one new endemic species (Schradera elmeri) were included for the Philippine record.Other generic revisions comprising Philippine genera are those of Xanthophytum Reinw.ex Blume (Axelius, 1990) and Paederia L. (Puff, 1991), but they did not result in name changes or additional records for the Philippines.Three other genera (Hedyotis L., Oldenlandia L., and Plectronia L.) included in Merrill's list (1923) are wanting revision.While revising the African Oldenlandia species of tribe Hedyotideae, Bremekamp (1952) split off the genus Exallage Bremek.He refrained from giving a full description of the genus because he could not include a study of the numerous Asian species in his study.In the Philippines, he transferred four species (E. auricularia, E. philippinensis, E. perhispida, and E. radicans) to Exallage out of the 49 Hedyotis species listed by Merrill (1923) and only E. auricularia was provided with a description.The African Exallage is a well-defined genus based on the indehiscent fruit, etc.A similar variation exists in Asian Hedyotis s.l., the identities of which require a meticulous and laborious work to investigate.
In 1985, Bridson reinstated the genus Psydrax after evaluating the heterogeneous African species of Canthium.She divided the genus Psydrax into two subgenera, Psy drax (palaeotropical) and Phallaria (African).She revised only the African species, resulting in 36 species of Psydrax which include new combinations from Canthium and Plectronia.Seven years later, Bridson (1992) published a revision of the genus Canthium in tropical Africa with a complete list of all African epithets of Canthium and Plectronia.Furthermore, she indicated the group affinities of Malesian Canthium which is a good basis for revising the Asian species.The genera Exallage and Psydrax will be included in the Philippine Rubiaceae database when revisions for the Philippine species become available.
Table 1 summarizes all the changes in Philippine Rubiaceae species since Merrill (1923).The old names are written beside currently used names to enable the reader to compare and cross check.Detailed information of all the genera can be retrieved using the INTKEY database.

Taxonomic position of the Philippine Rubiaceae genera
Classification of the family Rubiaceae has been a point of dispute among contending taxonomic views (Schumann, 1891;Verdcourt, 1958;Bremekamp, 1966;Robbrecht, 1988Robbrecht, , 1993) ) and for molecular data (Bremer et al., 1995;Bremer, 1996;Bremer & Thulin, 1998;Bremer & Manen, 2000;Andreasen & Bremer, 2000;Razafimandimbison & Bremer, 2002).This large family is still undertreated, many genera needing revision and the complexity of subfamilial and tribal classification remain unsettled (Robbrecht, 1993).The latest accepted classification of Robbrecht (1988Robbrecht ( , 1993) ) is basically adopted here but molecular phylogenetic results were also incorporated.Refer to Table 2 for quick identification of Philippine Rubiaceae genera placements which are also included in the generic descriptions of the INTKEY database.The four subfamilies (Cinchonoideae, Ixoroideae, Antirheoideae, and Rubioideae) recognized by Robbrecht are all represented in Philippine Rubiaceae genera, subfam.Rubioideae being the largest in genus numbers.Of 44 tribes (five of which are tribus incertae as to subfamiliar placement) in his list (Robbrecht, 1988), a total of 25 tribes are found in the Philippines.
In contrast to Robbrecht's (1988Robbrecht's ( , 1993) ) classification, molecular data of Bremer (1996) suggest only three subfamilies (Cinchonoideae, Ixoroideae, and Rubioideae).Phylogenetic results showed that the members of subfam.Antirheoideae are nested among the three other subfamilies.Likewise, she made some modifications of the tribal groupings particularly in the subfamily Rubioideae.However, Bremer (1996) stated that there is a need for more genera of Rubiaceae to be analyzed to obtain strong evidence and relevant data for subfamilial revisions of Rubiaceae classification.Indeed, the impact of phylogenetic analyses show that Rubiaceae is still systematically unsettled.
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