Phylogeny of Willdenowia (Restionacaea): Implications For
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- - - Phylogeny of Willdenowia (Restionaceae): Implications for speciation mechanisms and evolution of regeneration strategies in the Fynbos. - - - - - A.J. Swift Systematics Project - Botany Honours 1994 Supervisor: Dr. H.P.Linder - - - - - - - - - - The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgement of the source. The thesis is to be used for private study or non- commercial research purposes only. Published by the University of Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University of Cape Town 80LUS LI8RARY - C24 0004 7836 I 111 I - ABSTRACT - The phylogeny of Willdenowia is investigated using cladistic analysis, by studying culm anatomy, reproductive morphology and nut morphology. Two major clades are resolved, - consisting of three and seven species respectively. Important characters being tepal shape, elaiosome and nut characteristics. Species groups within the genus are, in general, seperated by ecological factors, including altitudinal range, regeneration strategy (i.e. seeder/sprouter) - and to a lesser extent phenology. This suggests that sympatric speciation, caused by steep ecological gradients present in the south-western Cape, may be an important process in the Fynbos. The Willdenowia phylogeny also shows that the ability to resprout after fire is - ancestral, with sprouter-species giving rise to seeder-species in all cases. This suggests that sprouting is ancestral within the Restionaceae. This is discussed in relation to previous studies on regeneration strategies. This pattern also has implications for conservation; - sprouter species possibly deserve a higher conservation status, as a result of their increased speciation potential. Because of their resiliance to the variable fire regime of the Fynbos, sprouter species are more likely to persist in the environment and give rise to new species, - than are seeder species, which are more susceptible to local extinction. - - - - - - - - - - - - 1 - 1. INTRODUCTION - Elucidating the mechanisms behind the unusually high floral diversity of the Cape Flora - remains as one of the most challenging biological problems of this unique area (Linder and - Vlok, 1991). Two hypotheses have been advanced to explain the high species diversity of the Fynbos flora. Cowling (1987) invokes the effects of frequent, irregular fires which result in - an absence of competitive exclusion as the dominant cause, while Linder (1985a) (Linder and Vlok, 1991), proposes that the steep ecological gradients present in the Cape region are the primary force behind increased speciation events. - One method of approaching this problem is to uncover processes operating at the generic - level. Understanding processes at a generic level provides a "window" through which we can gain insight into processes occurring at a familial and landscape level. Through cladistic - analysis, a putative phylogeny of a genus can be calculated. From a phylogeny, sister species - can be isolated allowing the identification of mechanisms resulting in speciation events. This will, in tum, provide clues relating to the mechanisms behind the high species diversity of - the Cape flora. The genus Willdenowia Thun. was considered a suitable genus for such a - study, considering its size (i.e. eleven species) and its morphological variation. To date the study of evolution and speciation within the Restionaceae has been limited to - work on the genus Rhodocoma Nees (Linder and Vlok, 1991), a paper which provided valuable clues regarding speciation in the Cape Flora. - - 2 A second major aim of the study is to investigate the evolution of regeneration mechanisms - within the genus Willdenowia. By assessing the distribution of seeders and sprouters on the - phylogeny of Willdenowia, one can also gain insight into the processes behind the evolution of regeneration mechanisms. Pressing questions include the following: (i) Which state (i.e. - seeder or sprouter) is ancestral ? (ii) Does the distribution of seeders and sprouters show any - pattern, and if so, what are the implications of this pattern. - Willdenowia Thun. is a genus in the family Restionaceae, which includes eleven species. - Thunberg, who originally erected the genus, included three species in his description (Linder, - 1984). Pillans (1928), in his work on the African Restionaceae, expanded the genus to include twelve species, which was later increased to thirteen with the discovery and description of Willdenowia stokoei Pillans (Pillans, 1942). In his review of the African - Restionaceae, Linder ( 1984) transferred W. argentea, W. .fistulosa and W. .fimbriata to Ceratocyrum. The description of Willdenowia rugosa Esterhuysen (Linder, 1985), brought - the number of species in the genus to its current number of eleven. - The most important characteristic used to circumscribe Willdenowia is the form of the male - inflorescences, which are thyrsoid (Pillans, 1928, Linder, 1984, 1985). Other important - characteristics include linear male perianth segments, branching culms and the presence of an eliasome (the latter two characters are not consistent throughout the entire genus). The distribution range of the genus is centred in the south-western Cape, with individual taxa extending on a north-south axis (as far as Kamiesberg), or a west-east axis (as far as - Knysna). Species are found across a wide altitudinal range, and are usually found to occupy occasionally damp habitats in the landscape. - - - 3 - 2. MATERIALS and METIIODS - All species of Willdenowia, excluding Willdenowia a.ffinis Pillans, were included in the study. - This species is known only from the type specimen collected on Table Mountain. - 2.1 Reproductive structures. Morphological data were collected from herbarium specimens. The youngest female - inflorescence was selected for dissection. Material was reconstituted and softened in boiling - soapy water, facilitating easier dissection, which was carried out under a compound microscope at lOX. To prevent dehydration while dissecting, material was placed in water - in a petri dish. Dissected parts of the female inflorescence were flattened on a microslide - which was coated with Prestik®, a pliable putty. This prevents the margins of bracts and tepals from becoming inrolled, which results in difficulty when inspecting dry material. - - 2.2 Stem anatomy. - Unblemished lengths of intemode culm were cut and boiled in soapy water until soft. Specimens were sectioned using a sledge microtome, cutting at a thickness of between 20 µm - and 30 µm. Sectioned material was stained in Safrinin-Alcian blue (Tolivia and Tolivia, - 1987), dehydrated through an alcohol run, and then placed in xylene. Sections were mounted in Canada fixative on microslides, which were placed on a heating tray overnight. Microslides were dried in an oven for three days. Material was viewed under a light microscope between 1OX and 200X. - - 4 2.3 Nuts. - Nuts were obtained from herbarium collections, and viewed under a dissecting microscope. - Material was boiled to reconstitute eliasome tissue. - 2.4 General. - Herbarium specimens were inspected for ecological information, including regeneration status and phenology. The ability to sprout after fire can easily be identified from herbarium - material, by the charred remains of pre-fire culm growth combined with new post-fire vegetative growth. Inferring phenology from herbarium material can be misleading. Male flowers are retained - on the plant for differential periods after flowering, up to a few months in some cases. - Willdenowia species also commonly retain stigmas until the nut is ripe, giving a false impression of recent flowering. An example of this includes Willdenowia rugosa; on closer - inspection, apparently young female inflorescences reveal fully developed seeds. If one is aware of these difficulties, however, valuable phenological data can be obtained from herbarium specimens. Other information such as habitat type and geographical distribution - were also obtained from herbarium material. A number of field trips were undertaken to acquire missing data relating to regeneration status and phenology. The distribution of species are given in fig. 1 - fig. 5. Species generally occur across a wide geographical range, with individual ranges overlapping considerably. - 2.5 Phylogeny. Morphological and anatomical characters were scored onto a matrix (see Appendix 1 [Table 5 - - - - D l(lDI lOO IIEIIH - O CE AN D ]Ot-lotll(IIU IND I AN ~ 1ot-120o ,mu (a) sou, 0 IO 100 llllJll(UU - D uo,1.12ot ,mu .. ,,. ... ... 11" 11' - - - - - CAP - ,. - \ D m011ot1111m O CE AN D lOMot 11(11(1 IND I AN e:::zl 100-1100 1111111 (b) 100 mo,rnn D uovt 1200 •Elm !CAI.Es 0 lO ... 11" Fig. 1. Distribution of (a) Willdenowia purpurea and (b) Willdenowia rugosa. 6 - - - - - D 1n111 ioo ,um - O CE AN E] lOO-lft 111111(1 IN O I AN fZ:3 IOt-1 !Vt II! IMS (a) i 100 (1~1111(11(1 D 111M 12tt 111111(1 scau, o IO - .. ,r ... ... ,r - - - - - c::J 1£11113" ll!IIH O CE AN D lOO-lftll!IIEI 1 NO I AN fZ:3 IOt-1 ltf 11! 11(1 0 1111'1( !lot IUl(I !ICILl1 I IO 100 lllllll(IIH .. ,r ... ,,. ,r - Fig. 2. Distribution of (a) Willdenowia hwnilis and (b) Wil/denowia sulcata. 7 - - - - - - - D 1n1111 lOl ll£ffll ocEAN D lOI-IOt IIIJH 1NDIAH rn:I IOt-1 !0t 11(11(1 (a) scau,1.. ______ _.,ot llLIIIIEl1H D uo,,i J2Mll(1l(I ,,. ,,. - a• ,r - - - - - - D in1111 ,.. 11(11(1 OCEAN - D lO... Nll(lll(I 1NOIAN rn:I IOt-1 !Ot II( 111(1 (b)' scau, , IO lot IILOIIElll(S D •- 110t 11(11(1 - .. ... ... ... ,,. ,r - Fig. 3. Distribution of (a) Willdenowia bolusii and (b) Willdenowia I stokoei 8 - - - 30• - u• - - u• ,.. ... (a) ...... - JI" II' ... 20' 22' 2,• Fig. 4 .. Distribution of (a) Willdenowia glomerata and (b) Willdenowia teres · - - - - , 21• - I JO' - ·- J2° J2' - 34• 34• (a) ... !' .. (b) '" ,. .... - 35• 11• ... 20• 22' 2,• JI' II' 11• 20• 22° 2,• - Fig. 5. Distribution of (a) Willdenowia arescens and (b) Willdenowia incurvata. - - 9 1] for a list and description of characters used). Character polarity was assessed using - outgroup comparison (Maddison et al., 1984, as cited by Linder and Vlok, 1991).