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eISSN: 2357-044X Taeckholmia 38 (2018): 84-106 Numerical re-assessment of the phenetic relationship between Apocynaceae and Asclepiadaceae A. El-Gazzar*1, A. El-Ghamery2, A. Khattab3, A. El-Saied2, A.A. El-Kady2 1Department of Botany and Mcrobiology, Faculty of Science, El-Arish University, N. Sinai. 2Botany and Microbiology Department, Faculty of Science (Male), Al-Azhar University, Cairo. 3Botany and Microbiology Department, Faculty of Science, Cairo University, Cairo, Egypt. *Corresponding author: [email protected] Abstract Recognition of the Apocynaceae and Asclepiadaceae as one family or as two separate but related families with five subfamilies was for long highly controversial. Raising some of the subfamilies to the rank of family was also disbutable. To resolve this issue, 55 morphological and palynological characters were recorded in a data matrix comprising a sample of 41 species belonging to 24 genera of Apocynaceae and 75 species from 31 genera of the Asclepiadaceae. The data matrix was subjected to cluster analysis using the Sørensen’s measure of dissimilarity and Ward's method of clustering in the PC-ORD version 5. Out of three possible interpretations of the result, we opted for that which suggestd considering the five subfamilies Apocynaceae-Rauwolfioideae, Apocynaceae- Apocynoideae, Asclepiadaceae-Asclepiadoideae, Asclepiadaceae-Periplocoideae and Asclepiadaceae- Secamonoideae as five distinct, neatly defined and homogeneous families of equal rank based on a unique set of diagnostic structural and functional characters for each of them. Key words: Apocynaceae, Asclepiadaceae, classification, cluster analysis, morphology, pollinia, tetrads. Introduction The Apocynaceae are one of the largest and not stigmatic. Fruit a berry, drupe, capsule, families of angiosperms with 375 genera and or follicle. Seeds with or without coma. The over 5100 species (Endress et al., 2007). Apocynaceae are cosmopolitan, except in the According to Watson and Dallwitz (1992), Li arctic regions. Numerous species are grown for Ping-tao et al., (1995) and APG IV (2016) their medicinal and ornamental values, and the members of the Apocynaceae are mostly trees, wood of trees such as Alstonia and Dyera shrubs, or vines, rarely subshrubs or herbs, species is a source of lightweight hardwood containing milky latex or rarely watery juice. (Alan and Wilkes, 1964; Chin et al., 2006). A Leaves simple, opposite, sometimes whorled similar account of the Asclepiadaceae s.l. was or alternate, pinnately veined; stipules absent recently given by El-Gazzar et al. (2018). or rarely present. Inflorescences cymose, The Apocynaceae were established by axillary or terminal, with bracteoles. Flowers Jussieu (1789) as (“Apocineae”) to include 24 bisexual, 5- [or 4]-merous, actinomorphic. genera subdivided into three groups essentially Calyx 5- or rarely 4-partite, quincuncial, basal defined by characteristics of fruits and seeds. glands usually present. Corolla 5- or rarely 4- Brown (1810a and b) separated seven genera lobed, funnelform, salverform, urceolate, or (Ceropegia, Pergularia, Stapelia, Periploca, rarely rotate. Stamens 5 or rarely 4; filaments Cynanchum and Asclepias) from Jussieu’s short; anthers mostly sagittate, free or “Apocineae” and combined them with 46 connivent into a cone adherent to the stigma, newly described genera to establish a separate dehiscing longitudinally, base rounded, family, the Asclepiadaceae. Brown also cordate, sagittate, or prolonged into an empty recognized three groups within the spur; pollen in monads or tetrads. Ovaries Asclepiadaceae: the "Asclepiadeae verae" (= superior, rarely half-inferior, connate or Asclepiadoideae), the Periploceae, and an distinct, 1- or 2-locular; ovules 2–numerous unnamed mono-generic group containing only per locule. Style 1; stigma capitate, conical, or Secamone. In 1838, Endlicher divided the lampshade-shaped, base stigmatic, apex 2-cleft latter genus into three genera (Secamone, Received on 28/06/2018 and accepted on 18/08/2018 Numerical re-assessment of the phenetic relationship Toxocarpus and Goniostemma) and assigned Subsequent phylogenetic studies them the rank of “subordo” ‘Secamoneae’. supported the reunion of the Apocynaceae s.s Numerous new genera were added to the and Asclepiadaceae into the Apocynaceae s.l., Apocynaceae and Asclepiadaceae during the with the same five subfamilies (e.g. Sennblad succeeding decades and Schumann (1895a and and Bremer, 1996; Potgieter and Albert, 2001; b) proposed comprehensive classifications of Endress and Bruyns 2000; Endress et al., 2007; the two families. Schumann (1895a) divided Livshultz, 2010). Furthermore, Potgieter and the Apocynaceae sensu stricto into two Albert (2001) showed that the subfamilies: the Plumerioideae (with three Asclepiadoideae, Secamonoideae and tribes and six subtribes), and the Echitoideae Periplocoideae are well-supported and (with two tribes). Similarly, he (1895b) monophyletic, but no support could be found classified the Asclepiadaceae into two for the monophyly of the Apocynoideae and subfamilies: the Periplocoideae (with a single Rauvolfioideae as delimited previously by tribe) and the Cynanchoideae (with five tribes Endress and Bruyns (2000). Similar support and seven subtribes). The name for the three subfamilies was evident from the ‘Cynanchoideae’ in Schumann’s (1895b) cluster analysis of a comprehensive set of scheme had to be changed to Asclepiadoideae characters of vegetative and floral morphology R. Br. ex Burnett, as it included the type genus combined with types and features of pollen of the family (Asclepias), while the name aggregation (El-Gazzar et al., 2018). Plumerioideae K. Schum. was replaced with The apparent contradiction between Rauvolfioideae Kostel. In subsequent considering the Apocynaceae and classifications of the Asclepiadaceae, Asclepiadaceae as two distinct families or as Schumann’s tribe ‘Asclepiadaceae- one family prompted us to initiate the present Secamoneae’ was elevated to the rank of study with a view to put the currently accepted subfamily Secamonoideae Endl. (e.g. classifications of this closely knit assemblage Klackenberg, 2001; Livshultz, 2010). of genera and species to a rigorous practical test. A controversy over the taxonomic relationship between the Apocynaceae and Material and methods Asclepiadaceae persisted ever since. Some Herbarium specimens of a cosmopolitan authors (e.g. Judd et al., 1994; Struwe et al., sample of 41 species from 24 genera of the 1994; Sennblad and Bremer, 1996; APG IV, Apocynaceae sensu stricto and 75 species from 2016) considered the Apocynaceae and 31 genera of the Asclepiadaceae were examined in the Cairo University herbarium Asclepiadaceae as one large family the (CAI; acronym according to Holmgren et al., Apocynaceae s.l. Other authors (e.g. 1990). The number of specimens representing Takhtajan, 1987; Cronquist, 1981 and 1988; each taxon ranged between one and 30. Dahlgren, 1983; Rosatti 1989a and 1989b) Identity of taxa was confirmed using treated them as two distinct families. Whether appropriate local floras and nomenclature of this assemblage of genera and species are the species was updated according to the two treated as one family or as two families, their websites (http://www.theplantlist.org/), and subdivision into subfamilies remained almost (http://www.tropicos.org), where full lists of unaltered: the Apocynaceae are divided into synonyms can be found. Full names with the Rauvolfioideae and Apocynoideae, while author citations of taxa and collection data of the specimens examined are given in Appendix the Asclepiadaceae comprise the 1. Asclepiadoideae, Periplocoideae and Variation in vegetative, floral and pollen Secamonoideae (Endress and Bruyns, 2000 morphology observed in the available and Endress et al., 2007). Schlechter (1905) specimens was recorded comparatively in a raised the Asclepiadaceae-Periplocoideae to data matrix. The outcome was subjected to family Periplocaceae, which was later cluster analysis under the combination of accepted by other authors (e.g. Bullock, 1956; Sørensen’s distance measure and Ward's Hutchinson, 1959; Dyer, 1975). clustering method and the dendrogram method 85 El-Gazzar et al. was selected to express the result. All methods representing the 55 genera of the Apocynaceae are available in PC-ord version 5 (McCune, and Asclepiadaceae were defined accurately to 1997). express the variation in the plants’ vegetative and floral morphology as well as their pollen Results characteristics. Of the 55 characters in Table 1, States of the 55 characters observed in 12 are multistate and the rest are binary. the available specimens of 116 species Table 1: List of the 55 characters and character-states recorded comparatively for the 116 taxa belonging to 55 genera representing the Apocynaceae and Asclepiadaceae in the present study. 1. Plant herb 1/ shrub 0 2. Stem herbaceous 1/ woody 0 3. Stem cylindrical 1/ angled 0 4. Stem erect 1/ climbing-twinning 2/ procumbent 3 5. Stem obese 1/ not obese 0 6. Stem articulated 1/ continuous 0 7. Stipules present 1/ absent 0 8. Leaves petioled 1/ sessile–subsessile 2/ absent 3 9. Leaves fleshy 1/ not fleshy 0 10. Calcium oxalate prismatics present 1/ absent 0 11. Calcium oxalate druses present 1/ absent 0 12. Leaf arrangement alternate 1/ opposite 2/ whorled 3 13. Leaves simple 1/ pinnate 2/ absent 3 14. Leaf shape ovate-elliptic 1/ oblong 2/ orbicular 3/ linear-lanceolate 4/ cordate 5/ absent 6 15. Leaf blade hairy 1/ glabrous 0 16. Glandular hairs on leaves