Generic concepts in Australian Gardenieae (Rubiaceae): a cladistic approach (original) (raw)

Abstract

Cladistic analyses of a morphological database for 37 Australian species of Gardenia Ellis and Randia Houst. ex L. demonstrate the polyphyly of both genera. Five clades are identified, with Randia audasii C. T. White placed in an isolated position. Four of these clades are assigned to extant genera: Aidia Lour., Atractocarpus Schltr. et K. Krause, Gardenia s.s. and Kailarsenia Tirveng. The following genera are considered as falling within Atractocarpus: Neofranciella Guillaumin, Sukunia A. C. Sm., and Trukia Kaneh. Pelagodendron Seem. belongs with Aidia. Whereas Randia audasii falls within the current concept of Rothmannia Thunb., available data suggest that genus is paraphyletic, and should be confined to the African taxa. Introduction The Gardenieae and Coffeeae together include one-fifth of all rubiaceous genera, following revised delimitation of the tribes by Bremekamp (1934, 1966) and Verdcourt (1958). These tribes were initially associated by their contorted corolla lobes (Bremekamp 1934) and later studies (e.g. Robbrecht 1979) have reinforced this link over the earlier system based on number of ovules. While many genera with large flowers and fruits and numerous seeds immersed in pulp were referable to the Gardenieae, and others with small flowers and fruits, and 1-seeded locules with no pulp were referable to the Coffeeae, many intermediate taxa remained difficult to assign (Verdcourt 1981). Although many small, often monotypic, genera have been described (e.g. Griffithia Thwaites, Xeromphis Raf., Basanacantha Hook. f.), they have often been separated by rather artificial characters (Standley 1929) and few have been widely accepted (e.g. Rothmannia Thunb.). For almost two centuries Randia Houst. ex L. and Gardenia Ellis have served as reservoirs for difficult elements with contorted aestivation within the tropical woody Rubiaceae, species with two locules in the ovary being placed in the former, and those with one in the latter (Bentham 1861, 1867; Hooker 1873). Whereas Keay (1958), Bridson et al. (1980) and later workers have radically divided Randia and Gardenia in tropical Africa, these narrower generic concepts have generally been rejected in the neotropics (e.g. Standley and Williams 1975; Dwyer 1980; Rogers 1987). Most recent workers on the palaeotropical taxa, however, have accepted the conclusion of Fagerlind (1943), Taylor (1944), Bremekamp (1957) and Keay (1958), that Randia is restricted to the Americas; a recent cladistic analysis of Ixoroideae revealed no close relationship between Randia s.s. and the two representatives of Australian Randia s.l. included in the study (Andreasen and Bremer 1996). There has been a proliferation of new genera in the eastern palaeotropics (e.g. Tirvengadum and Sastre 1979; Tirvengadum 1983) but no testing of the concepts. Indeed, few taxa have been described or revised in such detail that one can be confident of their generic identity (e.g. Tirvengadum 1983). The aim of this study is to resolve the generic placement of the Australian species of Gardenia s.l., Randia s.l. and several new taxa in light of the narrower generic delimitations of both Gardenia and Randia.

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References (83)

  1. K. tentaculata (Hook. f.) Tirveng.
  2. R. kanisii Puttock 12 110000000001001000111022211110230210100110-000001010002101010221101000022000202211101100(12)100 R. macarthurii F. Muell. 3 9010091099901111199299930119002210912001011109102091901910990210990019922991111099999990 9 199 R. moorei F. Muell. ex Benth. 12 100100010001001000011022221111230200100100-000001010000001110221100000022000202211101100 2 100 R. ossicarpa 12 110100010001001000011121221111230210100100-099999999909999999291100000122909999999999909 9 999 R. sessilis F. Muell. 123 0010001000001111001100200010102210011000011100102011102210110210001011122101111000000000(12)110 R. stipularis F. Muell. 123 0010001000001011010211201211102210112001011103102011102210120210100099922901211000002000(01)110 R. tahitiensis Nadeaud 3 9010091199901100299199930019002210911001011109102091902910990210999099999991111099999990 9 199 R. tuberculosa F. M. Bail. 12 100000000001001000011022211111230210100100-000001010002201010221100000022000202211101100(01)100
  3. Coffea L. -Outgroup 12 190000000009999990999999991111230219900900-09-000010009902120222209099299990012319999990 9 900 relationship with Gardenia s.s is strongly supported (+5, 99%). On this basis it appears to merit generic recognition. The R1 (+6, 87%) and Aidia clades (+4, 95%) are both quite robust, and hence also merit generic status. The R2 clade is unsupported in this database, decaying at one extra step and being present in < 50% of parjack trees. The divergence of R. audasii above the Aidia clade also receives little support (+2, < 50%), although its placement below R1 is well supported (+6, 87%). Synapomorphies for the major clades are listed in Fig. 1. 190 C. F. Puttock and C. J. Quinn A. racemosa A. cowleyi R. stipularis R. fitzalanii R. sessilis G. merikin R. hirta R. chartacea R. benthamiana G. actinocarpa G. psidioides G. rupicola G. scabrella G. fucata G. megasperma G. pyriformis G. resinosa G. vilhelmii G. dacryoides G. ewartii G. faucicola G. jabiluka G. gardneri G. kakaduensis G. schwarzii G. sericea G. tessellaris G. ovularis K. jardinei K. ochreata K. suffruticosa R. audasii R. moorei R. 'ossicarpa' R. tuberculosa R. kanisii R. sp. (IronRa.) Coffea -Outgroup The type species of Gardenia s.s. was described from horticultural material originating in southern China. It is estimated that there are 120 species. Subgenus Bergkias, which is characterised by terminal thorns, is endemic in Africa (Verdcourt 1979). Little recent work has been done on subgenus Gardenia: there are recent floristic treatments for Fiji (Smith and Darwin 1988) and Hawaii (Wagner et al. 1990), a minor revision for Peninsula Malaysia (Wong 1982), and descriptions of three new species in Asia (Tirvengadum 1983). An examination of available material from New Guinea and the Pacific suggests there are almost twice as many names in current use as there are species. There are also several species in New Guinea awaiting description. At the species level, Gardenia is notoriously difficult (Verdcourt 1979; Bridson and Verdcourt 1988). The genus, however, as defined here, is very distinct, being supported by nine synapomorphies in this database (Fig. 1). Atractocarpus heterophyllus (Montr.) Guillaumin et Beauvis. and Neofranciella pterocarpon (Guillaumin) Guillaumin, Porterandia anisophylla (Jack ex Roxb.) Ridl., Sukunia pentagonioides (Seem.) A. C. Sm., Sulitia obscurinervia (Merr.) Ridsdale and Trukia carolinensis (Valeton) Kaneh. et Hatus. all cluster within R1 (Fig. 2). Whereas the entire clade is well 192 C. F. Puttock and C. J. Quinn A. cochinchinensis A. racemosa A. cowleyi Pelagodendron Atractocarpus G. merikin Neofranciella Porterandia anisophylla R. benthamiana R. chartacea R. fitzalanii R. sessilis R. stipularis Trukia carolinensis R. hirta Sukunia pentagonioides Sulitia obscurinervia G. augusta G. ovularis G. actinocarpa K. jardenii K. ochreata K. suffruticosa K. tentaculata Rothmannia globosa R. audasii R. moorei R. 'ossicarpa' R. tuberculosa R. kanisii R. sp. (IronRa.)
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