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Phylogenetic Relationships, Based on Allozyme Data, Between Six Cycad Taxa Indigenous to South Africa

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Phylogenetic Relationships, Based on Allozyme Data, Between Six Cycad Taxa Indigenous to South Africa 182 S. Afr. J. Bot. 1998.64(3) 182- 188 Phylogenetic relationships, based on allozyme data, between six cycad taxa indigenous to South Africa F.H. van der Bank', P. Vo rster' and M. van der Bank' 'Department of Zoology, 'Department of Botany, Rand Afrikaans University, P.O. Box 524, Auckland Park, 2006 Republic of South Africa 1 Department of Botany, University of Stellenbosch, Private Bag X1 , Matieland, 7602 Republic of South Africa Received 10 June 199i; revised 24 January /998 Phylogenetic relationships between Encephalartos altensteinil Lehmann, E. friderici-guifielmii Lehmann, E. /ehmannii Lehmann, E. nalalensis Dyer & Verdoorn, E. transvenosus Stapf & Burtt Davy and E. villosus Le maire were studied, using Cyeas revoluta Thunberg as outgro up. Th ree continuous and one discontinuous buffer systems were used and gene products of 21 enzyme coding loci were examined by horizontal starch gel-electrophoresis. A biochemical key, based on fixed allele differences, is presented. Fixed allele differences at one locus between E. altensteinii and E. nata lensis may confirm that these species do not share the same gene pool. However, the genetic distance is the least (O.042) between these two species, compared to the mean genetic distance value of 0.222 for the other ingroup taxa. The results are discussed with reference to affinities based on morphology and distribution. Keywords: Phylogenetic relationships, cycads, Encephalartos, allozyme . • To whom correspondence should be addressed. E-mail: [email protected] Introduction tris-EDTA-borate (MF; pH 8.6; Markert & Fau lhaber 1965), and The taxonomy of the Cycadales has never been easy. In most lithium-borate (electrode: pH 8.0) Iri s-citrat e (gel: pH 8.7) (RW; groups really distinctive morphological characteristics are lack­ Ridgway et aJ. 1970). Statisti cal analysis of allozyme data was exe ~ ing, and species are us uall y defined in terms of sets of character­ cuted using BIOSYS-I (SwotTord & Sel and er 1981) and DISPAN (Ota 1993). istics. When vegetative as well as reproductive structures are known, it is usually possible to identify a specimen; but there is Results reason to believe that th e characteristics used for identification are not of phy logenetic significance, due to a high degree of con­ Locus abbreviations, enzyme commission numbers, and mono­ vergence or divergence. morphic loci are li sted in Table 2. All ele products at the follow­ Due to the fact that cycads mature slowly, lack mechani sms ing loci were monomorphic: GPI-2, PE PB and PEP-LTI , and for long-distance dispersal of their seeds, and are generally con­ those of ACP, EST-2 (for the ingroup species) and PER-2 fi ned to specialised habitats which tend to be isolated within migrated cathodally. All ele frequencies for polymorphic loci are larger tracts of unsuitable habitat, geographical distribution prob­ presented in Tabl e 3, and allozyme phenotypes of putative heter­ ably provides a clue to phylogenetic relationships; and thi s is one ozygotes were congruent with those expected on the basis of the of the reasons why the phylogenetic significance of so many quaternary structure of the enzyme (Ward 1977). A zymogram diagnostic characters is in doubt. Vorster (1993) provided a list showing di stin ct differences between species in allele product of diagnostically useful characteristics for the South A fr ican spe­ mobilities at AK are presented in Figure I a. Fixed all ele mobility cies, and Vorster (i n press) attempted to do the same for some differences between species occurred at AK, AA T- 1, EST, tropical arborescent species which are very poor in di s tinguish~ GPI-2, LA P, MDH, MNR, PER, PEP-D, -LT2, PGM, PGDH and ing traits. In both papers it was stressed that the phylogeny SKDH. Fixed allele differences at SKDH between E. a/tensteinii remains a grey area. and E. nafalellsis were observed (Table 3). Genetic markers to The purpose of the present investigation, of which this paper identify the taxa studied are presented in a biochemical key in presents the first results, is to investigate similarities between Table 4. species of Eneepho/arlos on the molecular level, hopefully to The mean value of Wri ght's (1978) fixation index ofindividu­ gain insight into the phylogenetic relationships berween species. als relative to the total population, Fts, is 0.233; 0.80 1 fo r the total population and its subpopulations (FIT) and FST = 0.740 for Material and Methods the am ount of differentiation among subpopulations relative to Electrophoretic data fo r 30 individuals, from seven taxa (Table I), the limiting amount under complete fixation (Table 5). The loci were com pared. Of necessity, we had to use plants in cu lti vat ion, bu t that contributed least to inter- and intraspecifi c differences are \"e have used only specimens of undisputed identity . When sampling AAT-2 (0. 175), ACP (0.286), PGM-l (0.367) and PGM-2 diffe rent specimens of the same species, we used specimens fro m (0.437) because all ingroup taxa shared one of th e two alleles at diffe rent provenances to prevent sampling of the same clone or pop­ these loci (Table 3), ul at ion . Herbarium voucher speci mens of the plants exam ined were Nei's (1978) unbiased genetic di stance values between deposited at the Uni versity of Ste llenbosch, Botany Department populations and taxa are presented in Table 6. The mean genetic (STEU). Leaf tissue extracts were prepared and analysed by starch distance value between the ingroup taxa studied is 0.210, 0.042 gel electrophoresis (12% gels) using the extraction buffer, standard between E. a/tensteinii and E. natalensis, and 0.925 between the electrophoretic procedures, method of interpretation of ge l banding ingroup and outgroup taxa studied. These va lu es were 0.231, patterns and locus nomenclature referred to by Van der Bank eJ al. 0.058 and 0.935 respectively fo r Nei' s (1972) geneti c distance. (1995). The huffers used are: Iris-EDTA-borate (A; pH 8.6; Gon­ The latter values are a little higher compared to the values in charenko., al 1992); histidine ci trate (HC; pH 5.7: Kephart 1990); Table 6, but reflect similar groupings, and are included for s. Afr. 1. Bot. 1998,64(3) 183 Table 1 Locality description of the cycad individuals studied Sample Species locality (probable origin) Cycas revolUla Cultivated female (Japan) 2 Cycas revo(uta Cultivated male (Japan) 3 Cycas reva/uta Cultivated female (Japan) 4 Cycas revaluta Cultivated plant (China) 5 Encepha{artos transvenOSU$ Trichardtsdal 6 Encephalartos transve110SlIS Modjadje's Village, TZaneen 7 EncephaJartos transvenosus Cultivated 8 Encephalarlos lransvenosus Cultivated 9 Encephalartos villosus Cultivated 10 Encephalartos villosus East London II Encephalarlos villoslis Port SL lohna 12 EllcephalarfOs villosus Cultivated 13 EllcephalarlOs villoslI!)' Cultivated 14 Encephalartos fride ric i-q II if ie 1m i i Cathcart 15 Encepha!arlos fride rici-q uif ie/m i i Cultivated 16 Encepho!ar (Os fride r ic i-q II if ie 1m i i Cultivated 17 Encephaiarlos jriderici-quiliefmii Cultivated 18 Encephalartos nalalensis Howick 19 Encepha/artas naw/ensis Cultivated 20 Encepha/arlOS nala/ensis Vryheid 21 Encepha/anos n£lta/ellsis Melmoth Z2 Encepha/artos a/fellsteinii Grahamsto\"ll 23 £ncephalartos alfensfeinii Bathurst 24 Encephafarlos a/lens/eillii Buslunans River Mouth 25 Encephalarfos a/tens/einii Mkambati River 26 Encephalarlos ailens/einii Cultivated 27 Encephalartos lehmanni; Cultivated 28 Encephalartos lehmannii Cultivated 29 Encephalarlos iehmannii Steytlerville 30 Encepha!artos lehmannii Kirkwood comparisons with values listed in the literature, Phylogenetic N I and N2 is: P < (_1_)2 N2 . Fi xed allele mobi lity diffe rences relationships based on Nei's (1978) genetic di stance values are 2NI depicted in Figure lb. The cophenetic correlation value for the occurred at one locus for the two closest taxa studied (£. allellsleinii result in Figure Ib is 98.4% (using BIOSYS), with the onl y poor bootstrap value (42) between the branch linking E. villos1ts with and E. nalalensis) (Table 3). Therefore, the probability that these the other ingroup taxa (using D1SPAN). two species are fro m the same gene pool is extremely small (P < 0.39 x 10-11 ). Discussion Several other statistically based measures of genetic distance The taxonomic uses of allozyme electrophoretic data were reviewed are also available to reduce genetic differentiation between popu­ by many authors, including Avise (1974), Thorpe (1982) and lations over a range of enzyme loci to a single figure level, but Thorpe and Sole-Cava (1994). These reviews give details ofmeth­ Nei's (1978) measure is now used predominantly (Thorpe & ods for di stinguishing and identifying cryptic and sibling species. Sole-Cava 1994). Shaklee el 01 (1982), Thorpe ( 1982) and The criterion which must be applied if electrophoretic data are to be Thorpe and Sole-Cava (1994) showed the relationship between used, is to assess the level of differentiation found between taxa (a taxonomic divergence and genetic di stance, and concluded that test of whether or not they are from the same gene pool), Such a test the genetic distance (Nei 1972) average 0.05 (range: 0.002-(L07) is discussed by Thorpe and Sate-Cava (1994). The method to esti­ for conspecific populations; 0.30 (range: 0.03-0.6 1) for conge­ mate statistical probabilities for fixed allelic differences in samples neric species; and it ranged 0.58- 1. 2 1 between genera in the 184 s. Air. 1. 1301. 1998.64(3) a Origin II"l II"l V1 II"l 0 0 0 0 000 0 0 0 0 0 0 0 0 0 0 e-- e-- e-- e-- 00 00 00 00 00 0 0 0 0 0 0 0 0 ~ ~ ~ ~ '" '" '" '" ~ ~~ ~ ~ ~ V1 V1 V1 II"l ~ ~ ~ ~ 000 0 0 ~ ~ ~ ~ ~ ~ ~ ~ e-- e-- e-- e-- 0 0 0 0 00 e-- CXl CXl CXl 0 0 0 0 0 0 0 0 00000 0 0 0 0 0 0 0 0 -00000---- }- '" '" '" '" ~ .......
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