Taxonomic and Phylogenetic Census of the Celastrales: a Synthetic Review Shisode S.B.1 and D.A
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Curr. Bot. 2(4): 36-43, 2011 REVIEW ARTICLE Taxonomic and phylogenetic census of the Celastrales: A synthetic review Shisode S.B.1 and D.A. Patil2 1 Department of Botany L. V. H. College, Panchavati, Nashik–422003 (M.S.) India 2 Post-Graduate Department of Botany S.S.V.P. Sanstha’s L.K.Dr.P.R.Ghogrey Science College, Dhule – 424 005, India K EYWORDS A BSTRACT Taxonomy, Phylogeny, Celastrales A comprehensive assessment of the taxonomic and phylogenetic status of the celeastralean plexus is presented. An attempt has been made to review synthetically C ORRESPONDENCE based on the data from different disciplines divulged by earlier authors and from present author’s study on the alliance. The taxonomic literature indicated that the D.A. Patil, Post-Graduate Department of Botany Celeastrales (sensu lato) are a loose-knit assemblage. The tribal, subfamilial, familial S.S.V.P. Sanstha’s L.K.Dr.P.R.Ghogrey Science and even ordinal boundaries are uncertain and even criss-cross each other. It College , Dhule – 424 005, India appeared that the alliance can be grouped under two taxonomic entities viz., the Celastrales and the Rhamnales which appear evolved convergently. E-mail: [email protected] E DITOR Datta Dhale CB Volume 2, Year 2011, Pages 36-43 Introduction Hipporcrateaceae are accorded an independent familial status. The order Celastrales (sensu lato) is a loose - knit The family Rhamnaceae is included under the Rhamnales assemblage. The taxonomic history clearly reflected that this alongwith the Vitaceae only. In the latest Engler's syllabus, alliance is not restricted to any taxonomic entity. The boundaries Melchior (1964), included the Celastraceae and between the various taxonomic entities based on exomorphic Hipporcrateaceae as two independent families within his order characteristics are uncertain. The alliance has been investigated Celastrales alongwith other 11 families. The family fairly in different domains of endomorphology. However, the data Celastraceae in his system is divided into five subfamilies viz., so accrued is not assessed critically to date. If at all assessed, Celastroideae, Tripterygioideae, Cassinoideae, Goupioideae they are assessed using information from a single discipline. The and Siphonodontoideae. The families have been divided again present authors studied vegetative anatomy of some into different tribes. However, he is silent about further Celastraceae, Hippocrateaceae and Rhamnaceae. They are also categorization of his family Hippocrateaceae. His order borrowed evidence from past literature and reviewed Rhamnales contains the families Rhamnaceae, Vitaceae and synthetically. The result of our in - depth study in this alliance Leeaceae. The family Rhamnaceae has been divided into five are being presented in this communication. tribes viz., Rhamneae, Zizypheae, Ventilagineae, Colletieae and Gouanineae. Treatment by Wettstein (1935) is essentially Taxonomic history similar in regard to the placement of the three families-the Celastraceae, Hipporcrateaceae and Rhamnaceae. Only In the treatment of Bentham and Hooker (1862-1883), Melchior (1964) included Leeaceae under the family Vitaceae the genera of the family Celastraceae and Hipporcrateaceae and accorded to it a subfamilial status. constitute a composite family, the Celastraceae (sensu lato). They Rendle (1959) included the family Celastraceae in the categorised these genera into two tribes - Celastreae and order Celastrales, alongwith Staphyleaceae, Aquifoliaceae and Hippocrateae. The tribe Hipporcrateae included only four genera Empetraceae, whereas the family Rhamnaceae is kept under viz., Hippocratea, Salacia, Siphonodon and Llavea, whereas the the order Rhamnales alongwith the family Vitaceae. He others are kept under the tribe Celastreae. The family included the Hipporcrateaceae (sensu stricto) under the Celastraceae (sensu lato), in their scheme, is kept under the Celastraceae (sensu lato) the Leeaceae under the Vitaceae. order Celastrales alongwith the families such as Rhamnaceae, Benson's (1957) order Sapindales contains the Celastraceae Vitaceae and Stackhousiaceae. The family Rhamnaceae is and Hipporcrateaceae as separate families alongwith many divided into four tribes viz., Ventilagineae, Zizypheae, Rhamneae others, few of which he considered of uncertain position. He and Gouanieae. Bessey (1915) included the families Rhamnaceae, also included the families Rhamnaceae and Vitaceae (sensu Vitaceae, Celastraceae and an independent family lato) under his order Rhamnales. Hipporcrateaceae in the same order Celastrales alongwith some Hutchinson (1959, 1969, 1973) in his revised system others. placed the Celastraceae and Hipporcrateaceae as distinct In the Englerian treatment, Engler and Diels (1936), families alongwith few others under the order Celastrales, the three families- Celastraceae, Hipporcrateaceae and whereas his Rhamnales contained the families viz., Rhamnaceae are treated under two independent orders viz., Rhamnaceae and Vitaceae alongwith Heteropyxidaceae and the Sapindales and Rhamnales. The former two families are Elaeagnaceae. Takhtajan (1969, 1980) recognised the included in the Sapindales alongwith other 22 families. The www.currentbotany.org ISSN: 2220-4822 Shisode S.B. and D.A. Patil Curr. Bot. 2(4): 36-43, 2011 Celastraceae and Hipporcrateaceae as independent families alliance in the different systems of classification. Engler and and included them under his Celastrales alongwith some Diels (1936) stressed simple leaves, perigynous flowers, others; few of which, in his opinion, are of doubtful position. He antipetalous stamens and basal ovules in the Rhamnaceae to includes the Rhamnaceae and Vitaceae under a single order distinguish it from the family Vitaceae which show Rhamnales; however, the latter has been divided into two predominantly climbing habit, presence of tendril, antipetalous separate families- the Vitaceae and Leeaceae. Cronquist (1968, stamens, axile placentation and berried fruit. Benson (1957) 1981, 1988) recognised the order Celastrales as containing the exploits the characteristics of sepals, fruits and presence of separate families Celastraceae and Hipporcrateaceae tendril to delimit these families. Rendle (1959) characterizes the alongwith few others. His order Rhamnales includes the order Celastrales by flowers bisexual or unisexual by abortion, families Rhamnaceae and Vitaceae; the Leeaceae has been regular hypogynous cyclic, four to five numerous petals free or given a separate familial status. sometimes connate at base, stamens alternate to petals and few Soo (1975) in his review of the new classification other ovarian and ovular features. He distinguishes the order systems of flowering plants included only two families within Rhamnales from the Celastrales giving emphasis on antipetalous the Celastrales-the Celastraceae and Aquifoliaceae. Likewise, stamens, the other features are overlapping in the Celastrales. his order Rhamnales contains the Rhamnaceae and Vitaceae Melchior (1964) and Wettstein (1935) emphasized similarity. only. He appears to have included Hipporcrateaceae under the Hutchinson (1959, 1973) stressed more on position of Celastraceae (sensu lato) and Leeaceae under the Vitaceae nectariferous disc in the flowers, position of stamens, aestivation (sensu lato). of petals, presence or absence of endosperm and nature of leaves Thorne (1992) in his recent treatment included while defining these orders. Hutchinson (1959) used pellucid, Celastraceae under his order Celastrales but divided it into five punctate leaves, inflorescence leaf opposed, presence of tendril subfamilies viz., Celastroideae, Tripterygioideae, Cassinoideae, and fruit baccate in Vitaceae, whereas these are not so in the Hippocrateoideae and Siphonodontoideae. The genera Goupia Rhamnaceae. Cronquist (1981) in his synthetic assessment of and Lophopyxis are kept under their independent families - the this alliance used large number of exomorphic as well as Goupiaceae and Lophopyxidaceae respectively. Hipporcrateaceae endomorphic features. Takhtajan (1969, 1980) also distinguishes (sensu stricto) is accorded a subfamilial rank under the family the order Rhamnales by the antitepalous stamens from that of Celastraceae (sensu lato). He included the family Rhamnaceae the Celastrales wherein the stamens are alternate to petals. and Elaeagnaceae in his order Rhamnales. It is interesting to Bentham and Hooker (1862-1883) placed the families note that the family Vitaceae has been kept under the order Celastraceae and Rhamnaceae under the same order Celastrales Cornales. The family Vitaceae has been divided into two and differentiated on the basis of stamens alternate to petals in subfamilies viz., Vitoideae and Leeoideae. the Celastraceae and opposite petals in the Rhamnaceae. These families also find place in different orders, the Celastraceae in Exomorphological significance and taxonomy the order Celastrales and the Rhamnaceae under the order Some systematists (Bentham and Hooker, 1862-1883; Rhamnales. Cronquist (1968, 1981, 1988) employed the position Rendle, 1959; Soo, 1975; Thorne, 1992), submerged the genera of of stamens in relation to the petals for the delimitation of the Hipporcrateaceae into the Celastraceae (sensu lato), while others orders Celastrales and Rhamnales. They are similarly treated by (Engler and Diels, 1936; Melchior, 1964; Westteein, 1935; Takhtajan (1969, 1980). Benson, 1957; Hutchinson, 1959, 1967, I973; Takhtajan, 1969, 1980; Cronquist, 1968, 1981, 1988), and split the latter into the