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Diversity and Distribution of Ferns in Sub-Saharan Africa, Madagascar

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Diversity and Distribution of Ferns in Sub-Saharan Africa, Madagascar Journal of Biogeography (J. Biogeogr.) (2004) 31, 1579–1604 ORIGINAL Diversity and distribution of ferns in ARTICLE sub-Saharan Africa, Madagascar and some islands of the South Atlantic J. J. Aldasoro*, F. Cabezas and C. Aedo Real Jardı´n Bota´nico, Consejo Superior ABSTRACT de Investigaciones Cientı´ficas, Madrid, Spain Aim This paper reports the diversity and endemism patterns of African ferns, and explores the potential role of diversity refuges and environmental and historical factors in the shaping of these patterns. Material and locations The extant fern taxa occupying Africa south of the Sahara, Madagascar and some islands of the South Atlantic. Methods The number of taxa in each area or operational geographical unit (OGU) was scored, and the correlation between this number and physical and climatic variables analysed by standard pairwise and stepwise multiple regression analysis (SPR and SMR). The effects of biological factors such as dispersal capacity, reproductive biology, genetic features and certain physiological adaptations were evaluated by comparing the number of species in each OGU. Floral affinities among OGUs were analysed using non-metric multi-dimensional scaling (NMS) and parsimonic analysis of dispersion (PAD), and compared with b-turnover and inter-OGU distances. Results OGU area, elevation and the distance between refuges determined the composition of local floras, but only greater OGU area and the existence of higher maximum elevations increased species richness. The distance between refuges also affected the number of endemic species, especially on islands. The biological features studied only slightly influenced fern distribution. The main climatic predictor of species number was humidity. SPR and SMR revealed three main groups of ferns with different ecological trends. NMS and PAD analyses separated the four areas of highest diversity in Africa, three of which are inhabited by ferns with distinct ecological requirements. The fourth area was Madagascar, which shows an accumulation of endemic and relict diversity that is not easy to explain. Main conclusions The distribution of ferns in Africa has been influenced by refuges. These probably allowed many species to recolonize the neighbouring areas after the extinctions of the Pleistocene. Three major components were detected in the African flora: Guinea-Congolian thermophilous, cold-tolerant Afro-montane, and Southern drought-tolerant elements. These are related to the three main refuge areas, i.e. the Gulf of Guinea area, the eastern tropical region, and the Cape region. Endemicity in ferns was found to be lower than that of seed plants due to the higher dispersability of fern spores. The distance between OGUs seems to be the main predictor of the number of endemic fern species these areas contain. ´ *Correspondence: J. J. Aldasoro, Real Jardın Keywords Bota´nico, Consejo Superior de Investigaciones Cientı´ficas, Plaza de Murillo 2, 28014 Madrid, Pteridophytes, biogeography, habitat diversity, centres of diversity, extinction, Spain. E-mail: [email protected] climatic variables, dispersal capability, isolation, c-diversity, d-diversity. ª 2004 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi 1579 J. J. Aldasoro et al. artefacts and that the pattern of African flora is well defined INTRODUCTION (Linder, 2001). Africa is quite poor in pteridophytes compared with other The main aim of this work was to document the patterns of tropical areas (Tryon, 1986). It is even poor compared with diversity and endemism of African ferns and to analyse the Madagascar, which is 40 times smaller: of a global pteridophyte potential role of environmental and historical factors in flora of c. 12,000 species, c. 627 grow in Africa and c. 557 in shaping them. Seven complementary analyses were undertaken Madagascar (Kornas´, 1993). that together provide the most complete picture yet of the According to some authors, the low pteridophyte diversity biogeography of African fern flora: of Africa is mainly the result of the paucity of the continent’s 1. The number of taxa in 29 operational geographical units rain forest flora, which is frequently explained by extinction (OGUs) was determined in order to evaluate c-diversity, and due to Tertiary droughts (Kornas´, 1993). Other authors the ratios between species and genera richness (and other taxa) indicate the absence of extensive mountain ranges in Africa compared to determine whether any OGUs do not follow the as the main reason for low fern diversity (Moran, 1995). In general pattern. contrast, the Malgache region is extraordinarily rich – a 2. The correlation between species number and latitude or discrepancy still unexplained. longitude was analysed to study diversity gradients. Standard Pteridophytes are generally distributed along a latitudinal pairwise and stepwise multiple regression analyses (SPR and gradient, with the highest diversity in the tropics (mainly in SMR) were used to determine the explanatory power of mountainous areas) (Jacobsen & Jacobsen, 1989; Kornas´, climatic and physical variables. 1993; Linder, 2001). However, in Africa, ferns are highly 3. Independent SMR analyses were performed for large genera diversified in the eastern mountain arc, and somewhat less so to show their climatic preferences, and in order to explain their in the western mountains (Schelpe, 1983). Three main distribution patterns. These analyses were performed using the latitude groupings – tropical, Mediterranean and Cape – are relative contributions of species to each OGU and environ- usually used to describe the continental pteridophyte flora of mental variables. Africa (Kornas´, 1993). Island floras include the Macaronesian 4. The influence of dispersal and reproduction methods on the flora, the flora of the Atlantic islands, and the flora of the colonization of isolated marine or ‘habitat’ islands and on the Malgache region. The Macaronesian flora is mainly related to evolution of endemism was assessed, relating the proportion of Mediterranean or Palaeomediterranean floras. Madagascar species with trilete spores and gemmiferous reproduction, etc., has an extraordinarily rich tropical flora with a very high to environmental factors. number of endemic species, probably influenced by a very 5. The floristic affinities among OGUs were analysed using conservative climate. The flora of the South Atlantic islands is non-metric multi-dimensional scaling (NMS) and compared most closely related to that of South America, the sub- with b-turnover and inter-OGU distances. Antarctic islands, and to a lesser degree to the Cape flora. 6. Patterns of endemicity were documented by performing A feature of this flora is their adaptation for long-distance several tests: (i) the determination of correlations between the dispersal (Smith, 1972). number of endemic species and latitude to look for gradients Habitat diversity, elevation, cloud cover, rainfall, tempera- in endemicity; (ii) the determination of correlations between ture, seasonality, exposure and soils have all been suggested to the number of endemic species and the distances between explain pteridophyte richness and distribution (Moran, 2002), OGUs, in order to study their isolation; and (iii) SMR analyses but biological factors such as dispersal capacity, reproductive of the relative contribution of endemic species and climatic biology, genetic features and certain physiological adaptations and physical variables, in order to determine these variables’ have also been proposed (Barrington, 1993; Kornas´, 1993). The explanatory power. role of these features, however, has not been examined in 7. The consistency of floral affinities deduced from NMS Africa and its related islands. analyses was tested. To phenetically classify biotas and to Refuges, places that have shown climatic stability over search for possible historical patterns, parsimonic analysis of geological time (Fjedsa˚ & Lovett, 1997), are areas from which dispersion (PAD) was performed. species spread out when climatic conditions became more favourable. Diversity and local endemism should therefore METHODS increase towards refuges because of their isolation and the maintenance of favourable climatic conditions. The existence Choice of taxa of refuges has been questioned by several authors. The main arguments against them are Neotropical (Colinvaux et al., Data on ferns south of the Saharan desert, Madagascar and 2000), while those in favour are based on evidence from Africa some islands of the South Atlantic (1001 species) were (Linder, 2001). Based on the distribution of several groups of obtained from published floral surveys and taxonomic mon- plants (Linder, 2001) and animals (Mayr & O’Hara, 1986; ographs. The main floral studies used were the following: Cape Harcourt, 2000), Africa is thought to harbour several of these Verde (Hansen & Sunding, 1993), Mt Loma, the Kwahu refuges. There is remarkable congruence between centres of Plateau and Togo Ranges (Alston, 1959), Mt Cameroon endemism and refuges, suggesting that the latter are not (Tardieu-Blot, 1964a; Edwards, 1998), Mt Oku (Edwards 1580 Journal of Biogeography 31, 1579–1604, ª 2004 Blackwell Publishing Ltd Diversity and distribution of African ferns et al., 2000), Annobo´n, Bioko and Rı´o Muni (Velayos et al., Atlantic islands as they have been described as refuges for 2002), Porto Principe and S. Tome´ (Figueiredo, 1998, 2001), animal and plant species (Fig. 1, Table 1). These are reported Mt Moukande´ (Tardieu-Blot, 1964b), Angola (Schelpe, 1977), to be particularly rich in
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