The Phytogeography of the Chamaecytisus Proliferus (L. Fil.) Link (Fabaceae: Genisteae) Complex in the Canary Islands: a Multivariate Analysis
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Vegetatio 110: 1-17, 1994. @ 1994 Kluwer Academicifublishers. Printed in Belgium. 1 The phytogeography of the Chamaecytisus proliferus (L. fil.) Link (Fabaceae: Genisteae) complex in the Canary Islands: a multivariate analysis J. Francisco-Ortega 1, M. T. Jackson 1*, A. Santos-Guerra 2, M. Fernandez-Galvan 2 & B. V. Ford-Lloyd 1** 1School of Biological Sciences,The University of Birmingham, P.O. Box 363, Birmingham, B15 2TT, ~ UK 2Jardin de Aclimatacion de La Orotava, Calle Retama Num. 2, 38400, Puerto de La Cruz, Tenerife Canary Islands, Spain * Present address: International Rice Research Institute, P.O. Box 993, 1099 Manila, Philippines; ** Author for correspondence 7.9.1992 Keywords:Ecogeography, Fodder-legumes, In-situ-conservation, Biodiversity, Germplasm,Tagasaste Abstract Chamaecytisusproliferus(L.til.) Link (Fabaceae:Genisteae) represents a speciescomplex in the Canary Islands. Floristic data from 147releves from the whole complexwere collected and analysedby clas- sification (TWINSP AN) and ordination (DECORANA) methods.Results indicate that white escobon of Tenerife, escobonof El Hierro, white escobonof Gran Canaria and typical tagasastein La Palma are associatedwith those plant communitiesfrom the north of theseislands which are under the influ- ence of the north-easterntrade winds. Narrow-leavedescobon in Tenerifeand La Gomera, escobonof southernGran Canaria and white tagasasteof La Palma are found in those areaswhich are not under the direct influenceof thesewinds. Morphological forms from the more easterlyislands (Gran Canaria and Tenerife-La Gomera) have the broadestecological range and they have played an important role in the floristic changeswhich have taken place after the destructionof the forests in theseislands. The highestpriorities for in situ conservationshould be givento wild populations of typical tagasaste,white escobonof Tenerife and escobonof El Hierro. Abbreviations:International Board for Plant Genetic Resources(IBPGR), Detrended Correspondence Analysis (DECORANA), Operational Taxonomic Unit (OTU), Two Way Indicator SpeciesAnalysis (TWINSP AN) Nomenclature:Hansen, A. & Sunding,P. 1985.Flora of Macaronesia.Checklist of vascularplants. 3rd ed. Sommerfeltia 1: 1-167; Acebes-Ginoves,J.R., Del Arco, M. & Wildpret, W. 1991. Revisiontaxo- nomica del genera Chamaecytisus(t. fil.) Link en Canarias.Vieraea 20: 191-202. ~Accepted Introduction truly domesticated species (Harlan 1983). No clear morphological and agronomic differences The genus Chamaecytisus Link (Fabaceae: exist between typical tagasaste plants from wild Genisteae) comprises approximately 28 species. and from cultivated populations. This means that The centre of highest diversity occurs in the Bal- unlike many other more domesticated crops, ta- kans where more than 15 species can be found gasastegermplasm collected in the wild is of value (Cristofolini 1991). The number of species de- as in many instances it can be used without hav- creasestowards the rest of Europe, the Near East, ing to resort to complex plant breeding proce- North Africa and the Canary Islands. Chamae- dures. The importance of germplasm from wild cytisus proliferus (L.ftl.) Link forms a taxonomic populations of this fodder speciesdetermines that species complex in the Canary Islands (Fig. 1). ecogeographical studies within its distribution are Both Acebes-Ginoves (1990) and Francisco- extremely useful in the establishment of strategies Ortega (1992) reported seven morphological for subsequent evaluation and utilisation of its forms within this complex namely, white escobon plant genetic resources. Furthermore as the cen- of Tenerife (C. proliferus ssp. proliferus sensu tre of origin and diversity of tagasaste is found stricto), narrow-leaved escobon (C. proliferus ssp. within an archipelago, it is essential to consider angustifolius (Kuntze) Kunkel), typical tagasaste the general patterns of island biogeography prior (C. proliferus ssp. proliferus var. palmensis (Christ) to any study on the phytogeography and plant Hansen & Sunding), white tagasaste (C. proliferus genetic resources of this species. ssp. proliferus var. calderaeJ .R. Acebes), escobon Studies concerning the biogeography of the Ca- of southern Gran Canaria (C. proliferus ssp. me- nary Islands have been reported extensively else- ridionalis J.R. Acebes), white escobon of Gran where (e.g. Webb & Berthelot 1836-1850; Kunkel Canaria (C. proliferus ssp. proliferus var. canariae 1976; Bramwell 1979; Santos-Guerra 1983a; (Christ) Kunkel) and escobon of El Hierro (C. Rivas- Martinez 1987). Due to the oceanic posi- proliferus ssp. proliferus var. hierrensis (Pitard) tion of the archipelago and the influence of the J.R. Acebes). cold north-eastern and the hot north-western Both kinds of tagasaste are endemic to the is- trade winds there is a stratification of different land of La Palma. Escobon ofEl Hierro, escobon climates in the archipelago both in terms of alti- of southern Gran Canaria, white escobon of Gran tude and orientation. This stratification is also Canaria and white escobon of Tenerife are only reflected by the patterns of distribution of the found in their respective islands whereas narrow- vegetation of the Canary Islands in five different leaved escobon occurs in Tenerife and La Go- life-zones namely, infra-canarian zone (semi- mera (Fig. 1). desert scrub), thermo-canarian zone (arid scrub, Although originally from La Palma, typical ta- Laurus azorica wood and heath belt), meso- gasaste is also cultivated in El Hierro, La Go- canarian zone (Pinus canariensis forest), supra- mera, Tenerife and Gran Canaria. The other six canarian zone (high altitude scrub) and oro- morphological forms are not cultivated but heavily canarian zone (only Viola cheiranthifolia). Within pruned and grazed in their wild habitats (Perez de this ecological framework the other two factors Paz et al. 1986; Francisco-Ortega et al. 1990). which determine the patterns of biogeographical White escobon of Gran Canaria has beenreported variation found in the Canary Islands are the ob- to be semi-cultivated in some areas of north- vious geographical isolation which exists between western Gran Canaria (Hernandez-Gonzalez each island and the abrupt topography. 1987; Francisco-Ortega et al. 1990) where it is Habitat stratification in life-zones has pro- used to feed livestock. duced several examples of adaptive radiation in Tagasaste appears to follow the same pattern the genera Aeonium (Lems 1960), Argyranthemum of domestication of other fodder species and al- (Humphries 1976) and Sonchus (Alridge 1980). though it is cultivated it cannot be regarded as a On the other hand geographical isolation due to 3 ,J ,10 20, km iW ~~ ~m EI Hierro ~; !2 ~m20 Fig. 1. The distribution of the seven morphological forms of C. proliferus (based on Acebes-Ginoves et al. (1991) and Francisco- Ortega (1992». Taxa are coded as follows: 0 C. proliferus ssp. proliferus var. proliferus; .C. proliferus ssp. angustifolius; f,. C. proliferus ssp. proliferus var. canariae; ..C. proliferus ssp. meridionalis;.C. proliferus ssp. proliferus var. palmensis; 0 C. proliferus ssp. proliferus var. calderae and .C. proliferus ssp. proliferus var. hierrensis. island topographyand/or insular isolationhas led Previousreports on the phytogeographyof C. to vicariance,examples of this evolutionarypro- proliferns (Ceballos & Ortufio 1951; Esteve- cessbeing the patterns of morphologicalvariation Chueca 1969; Sunding 1972; Rivas-Martinez and ecologyin the genera Cheirolophus,Crambe 1987)were imprecise,as firstly they did not in- and Limonium (Bramwell 1972). clude the whole distribution range of the species, 4 and secondly they were more concerned with the and in zones with low and high rainfall of 175 and ecology of the life-zones of the archipelago rather 1200 mm respectively. than with the ecological characteristics and For each locality cover values of vascular plant taxonomy of C. proliferns itself. Only recently has species were estimated using the Braun-Blanquet it been proposed (Acebes-Ginoves 1990) that scale. Special emphasis was placed upon the typical tagasaste, white escobon of Gran Canaria, compilation of Canarian and Macaronesian en- white escobon of Tenerife and escobon of El demics and of those species reported in previous Hierro were linked to the laurel (Laurns azorica) works (e.g. Santos-Guerra 1983b; Rivas-Mar- wood and the heath (Erica arborea) forma- tinez 1987) as characteristic of each life-zone, tions from the north of these islands, whereas the since they are clearly related with both the ecol- other three morphological forms were in associ- ogy and historical biogeography of the Canary ation with the plant communities of P. canarien- Islands. Numerical analyses were carried out on sis. a matrix in which each quadrat was regarded as No multivariate analyseson the phytogeogra- an OTU. Data from the Braun-Blanquet scale phy of this specieshave beenreported previously, were transformed following Feoli-Chiapella & and from our study both herbarium specimens Feoli (1977). A classification of releves was ob- and germplasmwere collected from most of the tained by the polythetic divisive method of localities of C. proliferus in the Canary Islands TWINSPAN (Hill 1979a). Ordination of reIeves with the objective that an understandingof the was accomplished using DECORANA (Hill phytogeographyof this specieswould lead to a 1979b) which avoids the curvilinear distortion better interpretation of its patterns of morpho- known as "arch effect" which is produced with