Re-Evaluation of Species Limits and Taxonomy of the Endemic Genus Stemonoporus Thw
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Cey. J. Sci. (Bio. Sci.) 36 (1): 17 - 34, 2007 17 RE-EVALUATION OF SPECIES LIMITS AND TAXONOMY OF THE ENDEMIC GENUS STEMONOPORUS THW. (DIPTEROCARPACEAE) USING MORPHOLOGICAL DATA S.C.K. Rubasinghe1*, D.M.D. Yakandawala1 and D.S.A. Wijesundara2 1Department of Botany, University of Peradeniya, Peradeniya, Sri Lanka 2Royal Botanic Gardens, Peradeniya, Sri Lanka Accepted 14 May 2007 ABSTRACT Stemonoporus Thw. is the most species-rich endemic dipterocarp genus in Sri Lanka and all its members are categorized as highly threatened or threatened in the IUCN red data book. Species limits within this important taxon are ill-defined, with some authors recognizing only a few variable species, and others recognizing a number of separate morphologically circumscribed species. These controvasies regarding the number of species and the species limits of Stemonoporus Thw. are a hindrance to implementing management measures to conserve this taxon. The main aim of the present study was to evaluate the species limits of the endemic genus Stemonoporus using morphological characters. Numerical and cladistic analyses were performed based on morphological data obtained from specimens collected from different geographical locations and herbarium specimens. Cluster analysis and Cladistic analysis on 170 specimens, divided the genus into 27 different clusters. These clusters corresponded to the 26 species recognized by Kostermans (1992) and in additional the present study identified a new morphologically distinct species, making the species limit of the genus 27, further strengthening its position as the most species rich endemic dipterocarp genus in Sri Lanka. The examination of 73 variables by cluster analysis and cladistic analysis revealed a number of clear differences in morphological features between the currently recognized species. In addition to the previously recognized characters, many important characters of leaves, flowers and especially of the leaf venation were recognized to define species limits. The study has revealed the long standing ambiguity regarding the species limits of Stemonoporus and further has recognized a new species to the genus. Key words: Stemonoporus, Dipterocarpaceae, cladistics, phenetics, species limits, morphological data. INTRODUCTION genera and a monotypic South American genus in Stemonoporus Thw. is an endemic genus that the subfamily Monotoideae, and one species of belongs to the family Dipterocarpaceae whose one genus in the South American subfamily origin dates back to the Gondwana in the early Pakaraimoideae (Kajita et al., 1998). However, Cretaceous period and thought to have migrated the disjunct distribution of closely related taxa to present day South-east Asia through the both in Sri Lanka and Malaysia suggests that Deccan plate. Blume in 1825 recognized the dipterocarps must have already diverged to Dipterocarpaceae, which forms a very closely- generic or even infrageneric sections before they knit family, and considered it to be related to entered the Laurasian plate from the Deccan plate Tiliaceae and Clusiaceae (Kostermans, 1992). (Dayanandan et al., 1999). The family Dipterocarpaceae consists of three subfamilies distributed widely in the tropics. In Sri Lanka the family Dipterocarpaceae is According to the recent molecular phylogenetic represented by 9 genera with 58 species of which classification, the family is nested within the Doona and Stemonoporus are endemic. More Rosids, in the Eurosids II clade, under the order interestingly all of the Sri Lankan dipterocarp Malvales close to Thymelaeaceae and Cistaceae species are endemic (Kostermans, 1992). Sri (APG II, 2003). Based on the most recent Lankan dipterocarp studies, apart from a note on classification of the family (Ashton, 1982), Vateria by Linnaeus in his Flora Zeylanica approximately 470 species in 13 genera are (1747), start with Thwaites’s book Enumaratio recognized in the Asian subfamily Plantarum zeylaniae (1864). Thwaites Dipterocarpoideae, 39 species in two African recognized three endemic genera Doona, Stemonoporus and Monoporandra. *Corresponding author’s e-mail: [email protected] S.C.K. Rubasinghe, D.M.D. Yakandawala and D.S.A. Wijesundara 18 Stemonoporus is the most species-rich ‘growing especially as small or large gregarious endemic dipterocarp genus in Sri Lanka with up groups in the understorey or on river banks in the to 26 species (Kostermans, 1992). All its lowlands’, or as ‘frequently common to sub- members are placed under the highly threatened dominant trees in the mid-mountain forests at or threatened category in the IUCN red data book 1000-1600 m’. Further, ‘each species has its own (IUCN Sri Lanka, 2003). The entire genus is well-defined habit, geographical and ecological confined to the per-humid forests in the wet zone range’ (Ashton, 1980). Kostermans (1992), in his of the island, from the Knuckles region south and account of the genus describes the ecology as westwards, and eastwards to Rakwana, with the ‘always in the wettest parts, many along streams exception of S. acuminatus, which occurs in and streamlets in the wet zone’. ‘Occurrence is Badulla district in the intermediate zone (Ashton, independent of the depth of soil or kind of soil 1980). Several species reach a height of about and often they occur in small populations far 1800 m, the highest altitude recorded for any apart and only a few reach timber size’ dipterocarp species in Sri Lanka (Ashton, 1980). (Kostermans, 1992) (Figure 1). Ashton (1980) described their ecology as Figure 1. Ecology of Stemonoporus Thw. A- S. petiolaris, occurs near a large rock on a mountain top close to Kitulgala/Yatiyantota area. B - S. gracilis is only known from an area along a tributary of the Kelani River, in the Kitulgala region. C - S. scalarinervis at Gilimale Forest Reserve. D - S. oblongifolius at Wewaltalawa near Yatiyantota. Species limits and taxonomy of Stemonoporus Thw. 19 Stemonoporus species are small trees with currently been re-established as a proper genus. resinous wood, rarely reaching timber size. Thwaites originally recognized 11 species under Leaves simple, spirally arranged, usually elliptic Stemonoporus and 3 under the Monoporandra. to oblong, very variable in shape and size. Inflorescence axillary and extra-axillary, Considering the three main classification principally panicles, showing reduction to systems; Trimen (1893) in his treatment on the racemes and single flowers, but their paniculate Dipterocarpaceae in volume I (1893) of his origin is always indicated. Flowers bisexual, Handbook to the Flora of Ceylon, separated sepals and petals are five-merous. Stamens 5 or Stemonoporus from Vateria. He described 13 15 (10 -13) in number, in one or two whorls, 10 species in Stemonoporus and 2 in the reinstated external, 5 internal slightly shorter. The genus Monoporandra while Ashton (1980) in his distinctive feature of Stemonoporus is the treatment of the family in the Revised Handbook chrome-yellow stamens, which form a very to the Flora of Ceylon has recognized 15 species. conspicuous cone-like structure around the style The most recent treatment of the family in the centre. No connective appendage, anthers Dipterocarpaceae by Kostermans (1992) opening by a pseudo-pore. Two types of fruits are recognized 26 species of Stemonoporus, where recognized in Stemonoporus: globose and ovoid, the genus is broadly divided into two sub-genera pointed (Figure 2). (Monoporandra and Stemonoporus) based on the number of stamens. A summary of the different Taxonomic history taxonomic treatments is given in Table 1 and a Stemonoporus was first recognized by comparison of major taxonomic treatments of the Thwaites in 1854. Since its inception, it has genus is given in Table 2. moved through Vateria and Vatica and has Table 1. A summary of the taxonomic treatments of Stemonoporus Thw. (Adopted from Kostermans, 1992). Taxonomic treatment Number of species recognized 11 species of Stemonoporus Thw. and 3 of Monoporandra Thwaites (1854) Thw Referred Stemonoporus Thw.to Vateria L. and added 3 to Thwaites (1864) Stemonoporus Thw. Reduced Stemonoporus Thw.to Vatica kept Monoporandra de Candolle (1868) Thw.separate. Separated Stemonoporus Thw.from Vater and recognized Trimen (1893) 13 species of Stemonoporus Thw. and 2 of Monoporandra Thw. Added 2 new species of which one belonged to Vatica (16 Hooker (1900) proper Stemonoporus Thw.) Included Monoporandra Thw. in Stemonoporus Thw. (15 Alston (1931) proper Stemonoporus) Restored S. lancifolius(Thw) Ashton but excluded S. moonii Ashton (1980) Thw (15 proper Stemonoporus Thw.) Recognized two sub-genera while distinguishing 26 species Kostermans (1992) of Stemonoporus Thw. S.C.K. Rubasinghe, D.M.D. Yakandawala and D.S.A. Wijesundara 20 Figure 2. Flowers and fruits of Stemonoporus Thw. A and B - The characteristic cone-like structure around the ovary formed by stamens of Stemonoporus that are 5 (b) or 10-15 (a) in number. C - The characteristic anthers of Stemonoporus, the margins of the two valves touching in one place, where the margin bulges, forming a long oblique orifice apically and a straight one below the bulges. D - a and D - b - Two types of fruit found in Stemonoporus; globose to sub-globose thin-skinned (D -a), conical, pointed, thick-skinned (D - b). Species limits and taxonomy of Stemonoporus Thw. 21 Table 2. A comparison of major taxonomic treatments of the genus Stemonoporus Thw. Trimen (1893) Ashton (1980) Kostermans (1992) S. nitidus S. nitidus S. nitidus S. oblongifolius S. oblongifolius S .oblongifolius