Chapter 4 Floristic and faunal Cape biochoria: do they exist?
Jonathan F. Colville, Alastair J. Potts, Peter L. Bradshaw, G. John Measey, Dee Snijman, Mike D. Picker, Şerban ProcheŞ, Rauri C. K. Bowie, and John C. Manning
4.1 Introduction factors, such as geology, altitude, latitude, and longi- tude can operate’. As such, proponents of a GCFR argue ‘There is no dispute that in the south of the continent for a shared evolutionary and ecological history, largely there are found numerous species, genera and even attributed to a cool-season moisture regime. families . . . that do not occur north. The question that It is not clear, however, whether a Greater Cape we have to answer is whether these forms constitute a Region is sensible only in the context of floristic data, separate . . . unit which is distinct, as a whole, from the other complexes of forms on the African continent?’ or whether the cool-season moisture zone shows bio- (Balinsky 1962). geographic coherence across a broader sampling of taxonomic groups (i.e. a Greater Cape Biochorion Many authors, over a long period, have remarked on (sensu Werger 1978)). A phytochorion or zoochorion the biotic distinctiveness of the southwestern corner of is defined geographically as an unranked area with Africa, both in terms of its flora (Bolus 1886; White 1976; a uniform composition of plant or animal species, Goldblatt 1978) and its fauna (Moreau 1952; Stuckenberg respectively (Linder et al. 2005; Olivero et al. 2012). Sev- 1962; Carcasson 1964; Poynton 1964; Holt et al. 2013). eral studies have explored the chorological divisions Climatically the region is defined by predominantly across southern Africa using individual taxonomic cool-season (autumn to spring) rainfall and mild tem- groups (e.g. plants (Born et al. 2007), frogs (Poynton peratures (Chapter 2), and its plant species richness is 1964; Crowe 1990), insects (Carcasson 1964; Endrödy- unmatched in the rest of Africa (Manning and Goldblatt Younga 1978), and birds (Crowe 1990; De Klerk et al. 2012; Snijman 2013). The Cape Floristic Region (CFR; 2002)). While these studies have generally highlighted or core Cape flora of Manning and Goldblatt 2012) is the distinctiveness of the predominantly mesic winter a distinctive phytogeographic feature (Goldblatt and rainfall Cape (see also Stuckenberg 1962), some have Manning 2000), previously recognized as one of six glob- demonstrated strong faunal links between the Cape al floral kingdoms on account of its high species richness and neighbouring succulent karoo areas (e.g. Endrödy- and endemicity (Marloth 1908; Good 1974; Takhtajan Younga 1978; Vernon 1999), while others have shown 1986; but see Cox 2001 who considered this ranking a stronger arid link, between areas of succulent karoo untenable). More recently, the concept of a Greater Cape and Nama karoo (e.g. Carcasson 1964; De Klerk et al. Floristic Region (GFCR), incorporating both the CFR 2002). Synthesizing the results of these studies to test and the succulent karoo region, has found favour as a the hypothesis of shared biotic regions has, however, more coherent biogeographical unit (Bayer 1984; Jür- been complicated by methodological inconsistencies gens 1991, 1997; Born et al. 2007). Bayer (1984) contends across these plant and animal studies. that once a larger, cool-season rainfall zone is consid- Establishing whether both fauna and flora are ered, then similarities in floristic composition and their aligned within core regions of a cool-season moisture patterns can be better understood ‘within which other zone, and whether the floristic distinctness of the GCFR
Fynbos: Ecology, Evolution, and Conservation of a Megadiverse Region. Edited by Nicky Allsopp, Jonathan F. Colville and G. Anthony Verboom. © Oxford University Press 2014. Published 2014 by Oxford University Press.
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is reflected in the biogeographic patterns of other lin- of regionalization for a Cape Biochorion, we briefly in- eages, is of great importance when seeking to under- troduce the biotic elements that make the broader Cape stand the historical and ecological development of the region so different from the rest of southern Africa. biota of the CFR and the adjacent succulent karoo re- gion (Rueda et al. 2010). Delineating and mapping of biogeographic regions utilizing a standardized method 4.2.1 Plants provides ‘units of area’ (Hausdorf 2002) to work with The GCFR, comprising fynbos, renosterveld, thicket, when searching for shared historical and evolutionary succulent karoo, and enclaves of afrotemperate forest patterns across taxa—the ‘biogeographic homology’ of vegetation (Chapter 1), has a documented vascular plant Morrone (2001). Biogeographic regions are likely to rep- flora of 11 423 species, with species endemism estimated resent areas of unique evolutionary history and ecolog- at 77.9%, and generic endemism at 22.2% (248 endem- ical process (Morrone 2009) and, as such, may be areas ic genera in 1119 genera; Snijman 2013). The region of high conservation importance (Kreft and Jetz 2010; includes four endemic or near-endemic plant fami- Olivero et al. 2012). These areas can be interpreted in lies, all confined to fynbos. These are dicots ofdiverse terms of historical events, such as geomorphic change affinity: Penaeaceae (Myrtales; 23 species), Grubbiaceae and climate stability (Chapter 8), or ecological pattern (Cornales; three species), Roridulaceae (Ericales; two (Wiens and Donoghue 2004; Beck et al. 2012), and they species), and Geissolomataceae (Crossomatales; one represent appropriate areas within which to investigate species; ordinal classification following APG3 2009). drivers of diversification and clade origin (Chapters 5, Asteraceae, typically the largest family in floras of arid 7, Rosen 1988; Jetz et al. 2004). Furthermore, by high- and semi-arid regions, is the most richly represented in lighting the biotic diversity and uniqueness of differ- the region, but the extraordinarily high contributions ent areas, they can be used to inform conservation and of Iridaceae, Ericaceae, and Aizoaceae is unique to this management strategies (Chapter 14), and to assess region, and consequently in southern Africa as a whole the threats posed to biodiversity by invasive species (Manning and Goldblatt 2012; Snijman 2013). Ericaceae, (Chapter 12) and climate change (Chapter 13). Proteaceae, Restionaceae, and Rutaceae have diversi- In this chapter we provide a systematic and consist- fied greatly on the impoverished sandstone soils of the ent biogeographic evaluation, at the species level, of Cape mountain ranges (Manning and Goldblatt 2012). five very different taxonomic groups (plants, birds, Geophytes comprise an exceptionally high proportion butterflies, reptiles, and frogs). For each taxonomic of the flora c.( 20%), possibly higher than in any other group, we group units of area together, based on their flora globally. Most geophytes are monocots(notably species similarity, and map these chorological patterns. Iridaceae, Hyacinthaceae, and Amaryllidaceae) but sev- We use these patterns to test the hypotheses of: (a) a eral dicot genera contribute large numbers of geophytes primary (biotic) break dividing South Africa into west- as well, namely Oxalis (Oxalidaceae), Pelargonium ern (predominantly cool-season rainfall) and eastern (Geraniaceae), and Othonna (Asteraceae; Manning and (predominately summer rainfall) regions; (b) an ex- Goldblatt 2012; Snijman 2013). tended Greater Cape Biochorion, grouping the CFR and the succulent karoo region; and (c) a more restrict- 4.2.2 Animals ed Cape Biochorion, centred on the CFR. Although less well appreciated, the GCFR harbours 4.2 The biotic uniqueness of the diverse and endemic-rich vertebrate and invertebrate southern tip of Africa faunas. Snakes and lizards represent the largest group of endemic vertebrates in southern Africa (Bauer The biogeographic regionalization of an area is deter- 1999). The GCFR has 191 reptile species (16 fami- mined by its species composition and how different this lies, 60 genera), of which 45 are considered endemic composition is to that of other regions (Croizat et al. or near endemic to the region. Amongst these, three 1974). The GCFR is particularly rich in both species groups stand out as having large numbers of endemic and endemics (Picker and Samways 1996; Kuhlmann taxa: geckos, cordylids, and chameleons. All three 2009; Linder et al. 2010; Manning and Goldblatt 2012; appear to be substrate specific, with the geckos and Snijman 2013). The elevated levels of endemism seen for cordylids having distributions that are strongly corre- many plant and animal groups suggest narrow ranges lated with particular rocky substrates and the distribu- for many taxa, and imply a strongly regionalized biota tions of chameleons being aligned to vegetation types (Rueda et al. 2010). Before we explore common patterns (Chapter 7, Tolley et al. 2006; Herrel et al. 2011).
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As has been highlighted repeatedly, the mesic winter the region is considered as a global centre of diver- rainfall area of the GCFR is globally unique in terms of sity and adaptive radiation for several insect groups its amphibian fauna. It has over 57 species, 31 of which (see Box 4.1). Moreover, the invertebrate fauna has are endemic, and two endemic genera. Poynton’s (1964) various unusual components, including a large pal- seminal study laid the foundation for future work, aeoendemic element (Stuckenberg 1962; Bowden 1978; emphasizing the high incidence of endemics, and the Endrödy-Younga 1978, 1988; Picker and Samways 1996; Gondwanan connections of the amphibian fauna in the Stuckenberg 2000; Day 2005; Kirk-Sprigs and McGregor mesic winter rainfall area. This area has continued to 2009; van Noort and Shaw 2009) and a diversity of be of interest to a number of faunal researchers (see neoendemics, associated with the adaptive radiations Poynton and Broadley 1978; Poynton 1980, 1987, 1992, of various pollinator groups (e.g. bees (Kuhlmann 1994; Drinkrow and Cherry 1995; Seymour et al. 2001; 2009), wasps (Gess 1992), flies (Hesse 1969; Usher Alexander et al. 2004; Poynton 2013; Schreiner et al. 1972; Greathead and Evenhuis 2001; Sinclair 2003; 2013), and most recently, the mesic winter rainfall area Barraclough 2006; Stuckenberg and Kirk-Sprigs 2009), has again been singled out as especially significant nemopterid lacewings (Tjeder 1967; Sole et al. 2013), for amphibians (Linder et al. 2012; Holt et al. 2013), Hopliini beetles (Peringuey 1902; Colville 2009), and with high levels of phylogenetic turnover between jewel beetles (Holm 1978; Holm and Gussmann 2004)). the mesic winter rainfall area and much of the rest of Stuckenberg (1998, 2000) and others (Usher 1972; sub-Saharan Africa. Barraclough 2006; Karolyi et al. 2012) have highlighted Although the CFR is relatively depauperate in terms the adaptive responses in mouthpart morphology of of its bird diversity and endemism (seven endemic spe- anthophilic genera found in at least seven fly fami- cies), the endemics that do occur are remarkable with lies. The high frequency of these adaptations in spe- respect to their evolutionary affinities. Two of Africa’s cies from the winter rainfall zone is significant because oldest songbird lineages (40–55 Ma; Barker et al. 2004; it indicates that the diversification of the GCFR flora Beresford et al. 2005; Fjeldså and Bowie 2008; Johansson has exerted strong selection on diverse anthophilic in- et al. 2008; Jetz et al. 2012), namely the sugarbirds sect groups. Other insect and invertebrate taxa, which (Promerops: Promeropidae) and rockjumpers (Chae- are either directly (e.g. phytophagous) or indirectly tops: Chaetopidae), each have one of their two species (e.g. predators) linked to plants, also show high bio- restricted to fynbos (Hockey et al. 2005). Additionally, geographic distinctiveness and/or signatures of adap- the orange-breasted sunbird (Anthobaphes violacea) is one tive radiation; for example Orthoptera (Dirsch 1965; of Africa’s most divergent sunbird lineages (Fjeldså and Naskrecki and Bazelet 2009), Mantophasmatodea Bowie 2008) and Victorin’s warbler (Cryptillas victorini), (Klass et al. 2003), Coleoptera (Sole et al. 2004), weevils once thought to be a warbler in the genus Bradypterus, (see Box 4.1), and scorpions (Prendini 2005). is now convincingly placed as a divergent monotypic The rich insect diversity of the GCFR is yet to be lineage in the Macrosphenidae (Alström, Olsson et al. comprehensively explored (Hesse 1969; Bowden 1978; 2013). The succulent karoo has few endemics, but shares Kuhlmann 2009); recent research, however, has yield- many bird species with the CFR and the Nama karoo, ed the discovery of a new insect order whose global and, together with the latter, supports one of the greatest centre of diversity is the winter rainfall zone (Picker regional concentrations of endemic birds in Africa (Allan et al. 2002), as well as the documentation of species et al. 1997; Hockey et al. 2005). These arid areas have sev- radiations (Naskrecki and Bazelet 2009; Sole et al. eral endemic larks (Alaudidae), chats (Muscicapidae), 2013) and unique adaptive forms (Picker et al. 2012). and cisticolid warblers (Cisticolidae). They also support three monotypic genera, comprising the cinnamon- breasted warbler (Euryptila subcinnamomea), the Namaq- 4.3 Assessment of the validity ua warbler (Phragmacia substriata), and the rufous-eared of a Cape Biochorion warbler (Malcorus pectoralis). The larks and chats com- ‘The first step toward . . . generalization is to determine prise some important in situ radiations (e.g. in Cercomela, what major types of coincident distributions (general- Certhilauda, and Calendulauda; Chapter 7, Outlaw et al. ized tracks) recur’. (Croizat et al. 1974) 2010; Alström, Barnes, et al. 2013), and show strong link- ages with species restricted to the north east arid zone of Early historical biogeographic studies were largely Africa (van Zinderen Bakker 1969; Verdcourt 1969). based on the intuitive, often subjective, and non- The invertebrates, including insects, show excep- replicable approach of examining the distribution maps tional richness and endemism within the GCFR, and of selected species (Drège 1843; Bolus 1875; Marloth
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Box 4.1 Insect elements unique to the Greater Cape Biochorion
Monkey beetles (Scarabaeoidea: Hopliini) collection and taxonomic work of the entomofauna of the Approximately 1040 described species (51 genera) of mon- Greater Cape region in general. key beetles are currently known from South Africa, the cen- Jumping cockroaches tre of adaptive radiation for the world’s Hopliini. Roughly 63% of the world’s species and 38% of the genera are con- The world’s only jumping cockroach (Saltoblattella mon- centrated here. Remarkably, >50% of the world’s species tistabularis; Box 4.1 Fig 1d) occurs in fynbos (Picker et al. occur in the Greater Cape. Two dominant genera, Hetero- 2012). This remarkable cockroach shows parallel evolution chelus (c.300 South African species, 70% of which occur in with grasshoppers, in terms of body plan and biomechanics fynbos vegetation types and 40% in succulent karoo veg- of jumping. In spite of its very unusual appearance, many of etation types) and Peritrichia (c.90 South African species, the features that make it unique amongst cockroaches are 77% of which occur in fynbos vegetation types and 64% locomotory adaptations, and the insect is probably a close in succulent karoo vegetation types; Box 4.1 Fig 1a), make relative of the cosmopolitan and modern subfamily Blatel- up approximately 35% of the South African monkey beetle linae (Djernæs et al. 2011). This evolutionary novelty is likely fauna. A remarkably high percentage of species (98%) and to be an adaptation for improved locomotion in a vertically genera (80%) are national endemics. Of equal significance stratified habitat such as stands of restios. is the huge (and globally unique) functional diversity seen in the South African monkey beetles. They pollinate a wide Weevils diversity of plants in the winter rainfall zones, with pollina- The largest insect superfamily, Curculionoidea, has an tion guild structure based on flower colour, shape, and floral estimated 12 000 species in southern Africa, and a large resources (Picker and Midgley 1996). The extreme sexual number of interesting groups in the GCFR. Radiations have dimorphism in both colour (c.82% of species) and hind-leg occurred in multiple lineages, some mirroring those of their size and shape (c.76%) suggests that sexual selection may host plants. Examples are: Sibinia (Tychiini) and Urodonti- be a strong driver explaining the huge diversity of species in dae (both on Aizoaceae); Apioninae (Apionidae; Box 4.1 Fig the Greater Cape (Colville 2009). 1e) on Aspalathus and Indigofera (Fabaceae); Gymnetrini on Scrophulariaceae; and Ceutorhynchini on Bruniaceae, Lacewings (Nemopteridae) Leucodendron (Proteaceae), and Heliophila (Brassicaceae). The 72 South African species of spoon- and thread-wing These are mostly small weevils (1–3 mm) which associ- lacewings (Box 4.1 Fig 1b) represent 57% of the world’s ate with the plants’ reproductive structures. Although host species, 79% of which occur in the Greater Cape and Nama specificity may still play an important role, radiation also ap- karoo regions. This represents almost half of the world’s spe- pears to be driven by abiotic factors such as geology. This cies (Sole et al. 2013), an indicator of the extensive radiation is the case in the large (mostly 1–3 cm), ground-inhabiting of the family in the western parts of southern Africa. Ap- weevils Brachycerus (Brachycerinae) and Episus (Microceri- proximately 94% of the South African species are endemic nae). There are also ancient lineages of Gondwanan affinity, (Tjeder 1967). As adults feed on pollen and nectar, they are mostly associated with Proteaceae (Tanaonini, several line- considered important pollinators in the arid and semi-arid ages in Entiminae sensu lato). regions. Palaeorelictual insect groups Heelwalkers (Mantophasmatodea) Of great evolutionary and biogeographical interest is the This is the most recently discovered order of insects (Klass presence of numerous palaeorelictual insect groups in et al. 2002; Picker et al. 2002), with all 20-odd species the winter rainfall Cape (e.g. Leptonyma sericea; Box 4.1 restricted to Africa. The majority (12 species) occur in the Fig 1f). These survivors of ancient Gondwanan lineages winter rainfall zone of South Africa (Eberhard et al. 2011). allow insights into the palaeohistory of the region, helping The Austrophasmatidae (Box 4.1 Fig 1c) is endemic to the to explain current speciation patterns of both the fauna and Greater Cape, and contains nine species (Klass et al. 2003). flora of the GCFR. Palaeorelictual insects are mostly strong The recent discovery of these fairly large nocturnal predators endemics, typically being confined to temperate habitats, in southern Africa underlines the need for more intensive especially forest, high-altitude fynbos, mountain streams
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Box 4.1 Continued
(e.g. Aphanicerca capensis; Box 4.1 Fig 1g), and caves. Due secondary hotspots in the Hottentots Holland Mountains to these attributes, they are considered of high conservation (80 species per quarter degree square) and the south- importance. They are best represented on the Table Moun- ern Cape forests (36 species per quarter degree square; tain chain (147 species per quarter degree square), but have Day 2005).
Bladder grasshoppers (Pneumoridae) This is an ancient family largely endemic to South Africa; all 17 species and nine genera (Dirsch 1965) occur in the region, with two forest species extending into East Africa (Dirsch 1965). The Greater Cape has a distinctive and rich fauna of bladder grasshoppers (Box 4.1 Fig 1h), with at least 12 species found here, all of these endemic to this region (Dirsch 1965).
Box 4.1 Figure 1 (a) Peritrichia monkey beetles belong to a guild of monkey beetles that feed almost exclusively on white, pink, and blue flowers (Photo: Sarah-Leigh Hutchinson); (b) the spoon-wing lacewing (Sicyoptera dilatata) was only known from a single specimen captured in 1836, and thought extinct until its recent rediscovery in the Riviersonderend mountain range (Photo: Mike Picker); (c) Austrophasma caledonense is a fynbos member of the insect order Mantophasmatodea, an order entirely restricted to Africa (Photo: Mike Picker); (d) the jumping cockroach (Saltoblattella montistabularis) shows convergence in body plan with grasshoppers (Photo: Mike Picker); (e) adaptive radiation of fynbos apionid weevils, with two species co-occurring on one Aspalathus species (Fabaceae) (Photo: Şerban Procheş); (f) the wormlion fly Leptonyma( sericea), showing elongated mouthparts for nectar feeding, is considered to belong to a relictual family with extensive radiation in the Greater Cape (Photo: Mike Picker); (g) the stonefly A( phanicerca capensis) is a representative of South Africa’s Gondwanan fauna, occurring only in temperate mountain streams (Photo: Mike Picker); (h) a male Bullacris showing the unique resonating bladder characteristic of this basal grasshopper family (Photo: Mike Picker).
1908; Nordenstam 1969; Hengeveld 1992). In the Cape, only relatively recently that the essentially comprehen- these early studies mostly used limited datasets, and sive lists of the Cape flora have been synthesized into retrieved broad-scale regional patterns (e.g. Bolus 1905; meaningful phytogeographical patterns (Goldblatt and Marloth 1908). As the flora of the Cape became better Manning 2000). The combination of modern analytical known, the numbers of taxa analysed and the resolu- techniques, increased computational power, and large tion of biogeographic detail improved (Weimarck 1941; multi-species distributional datasets has also given rise Dahlgren 1963), as did the resolution of subcontinental to a more objective means to identify biogeographic studies for both floristic (e.g. White 1976) and faunal patterns (e.g. Kreft and Jetz 2010; Linder et al. 2012; datasets (e.g. Carcasson 1964; De Klerk et al. 2002). It is Holt et al. 2013; Bradshaw et al. submitted).
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Table 4.1 Summary of the numbers of records and species richness of the South African plant and animal datasets analysed in this study.
Group Dataset(s) Approximate number Number of species Reference of records
Plants PRECIS including Protea 1.79 million 20 657 Germishuizen et al. (2006) South African Atlas Project Botanical Diversity Network Report No. 41;
1 Only terrestrial bird species considered; aquatic bird species (sensu Crowe 1990) removed on the basis of habitat usage taken from Hockey et al. (2005).
South Africa is fortunate to have well-surveyed dis- Cape Biochorion, and where this is situated. Since the tributional datasets generated from national atlasing analysed datasets are national, our results are unfortu- projects for a range of taxa, notably plants, birds, rep- nately restricted to South Africa, but this is problemat- tiles, frogs, and butterflies (Table 4.1). It is therefore ic only in the northernmost part of our area of interest, feasible to interrogate regional biogeographic patterns namely the arid winter rainfall zone of southwestern across taxa. The progression of biogeographic tech- Namibia. niques used for South African taxa has evolved from Except for frogs, data were analysed at the quarter purely descriptive accounts based on intuition and degree square (QDS) level of resolution (Edwards and expert knowledge (e.g. Weimarck 1941; Goldblatt and Leistner 1971) as historically these grid cells have been Manning 2000), through to high spatial resolution em- most widely employed for capturing biological distri- pirical analyses using point locality data and environ- bution data in South Africa. Frogs were analysed at mental themes (Gess 1992; Kuhlmann 2009; Prendini the half degree square level (HDS) of resolution due to 2005), with the utilization of multivariate statistics and the relatively low numbers of species found in South advanced algorithms for calculations of resemblance Africa, which translates into low species richness per matrices and clustering based on species-by-site matri- grid cell, therefore diminishing the discrimination abil- ces (Crowe 1990; Born et al. 2007; De Klerk et al. 2002; ity of clustering techniques. QDS and HDS are limited van Rensburg et al. 2004). These comprehensive spe- by their coarseness of resolution (a c.25 × 25 km block cies datasets and more robust quantitative analyses for QDS, and a c.50 × 50 km block for HDS), which ren- enabled us to undertake a comparative biogeographic ders them insufficiently sensitive to retrieve finer-scale study of different South African taxa, and assess their biogeographical patterns, or features with low taxon biogeographic patterns. representation (Moline and Linder 2006; Bradshaw Point locality data from collection-based records et al. submitted). In addition, sampling effort (number and recently collated national atlasing projects were of records per grid cell) varies dramatically between used to delineate biogeographic divisions based on a QDS or HDS, which may mislead biogeographic uniform, repeatable, and modern analytical approach analyses where the sampling effort for a given cell lies that clusters grid cells hierarchically according to the well below its potential species richness (i.e. species shared presence of species. In order to investigate the are omitted due to a lack of sampling). To minimize hypothesis of a Cape Biochorion, we analysed national this bias, we used conservative sampling effort thresh- species datasets in order to retrieve the major biogeo- olds to rid the datasets of poorly collected grid cells graphical patterns within South Africa. This allowed (Table 4.2). Unfortunately, this resulted in several grid us to determine whether the taxa analysed suggest a cells not being assigned to biochoria, with the result
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Table 4.2 Species richness and grid cell numbers per taxonomic group for each of the original and modified distributional datasets analysed, and the richness-per-grid sampling cut-off used to discard grid cells (deemed to be under-sampled; see text for details) for biogeographic analysis. While the sampling cut-off could result in a dramatic reduction in the area analysed (reptiles by 82%), species’ representation in the datasets dropped much less markedly (reptiles by 4%), if at all.
Original dataset Min. Modified dataset Maximum richness per grid cell1 cut-off richness No. spp. No. grid cells No. spp. No. grid cells Grid cells per taxon1
Plants 20 657 2009 (99%) 200 20 314 997 (49%) 2499
Reptiles 409 1469 (72%) 15 393 361 (18%) 73
Frogs 110 508 (94%) 6 110 318 (59%) 46
Birds 627 2009 (99%) 120 626 1361 (67%) 355
Butter- 656 1410 (69%) 40 650 394 (19%) 278 flies
1 Histograms were produced using the modified datasets. In the ‘grid cells per taxon’ histograms, species distribution sizes were separated into 20 bins, with the proportion of species of the given distribution sizes displayed for each bin. In the richness-per-grid histograms, grid cell species richness was divided into 20 bins, with the proportion of species richness displayed for each bin.
that the mapped results of cluster analyses typically technique was used to cluster grid cells using the func- contain gaps (e.g. Linder et al. 2005). In order to set tion hclust in the R statistical environment (R Devel- thresholds, the number of records for each grid cell opment Core Team 2012). UPGMA has generally been was plotted against its species richness. For under- found to outperform most other available hierarchical sampled grid cells, the species richness is usually equal agglomerative clustering methods (Shi 1993; Linder or near equal to the number of records (i.e. a strong 2001; Kreft and Jetz 2010). linear relationship between the two). We determined Instead of applying the commonly used, and argu- the threshold minimum number of records as the point ably arbitrary, phenon line (Rosen 1988; Rueda et al. at which this linear relationship started to break down 2010), we employed a branch ranking technique de- (i.e. where the number of records is no longer a strong rived from Strahler stream order assignment (Strahler predictor of species richness in a given grid cell). This 1957; Borchert and Slade 1981), which focusses to a threshold varies across datasets due to differences in greater extent on dendrogram structure than a visu- overall species richness. ally determined similarity cut-off (Bradshaw et al. Using a species presence matrix, a dissimilarity ma- submitted). The premise is that branches are scored trix of the relationships between cells was calculated from tip to root and upon merging, the ‘new branch’ using Kulczinski’s second (K2) similarity equation (see receives a higher branch order number when two or Jurasinski 2012), an equation commonly employed in more branches of equal score intersect (e.g. 1 + 1 = 2; biogeographic studies (Shi 1993; Moline and Linder 2 + 2 = 3; 3 + 3 = 4). Where branches are unequally 2006; Born et al. 2007). Importantly, K2 does not take scored, however, the highest branch order is retained shared absences into account (Shi 1993; Linder 2001), (e.g. 2 + 1 = 2; 3 + 2 = 3 and 3 + 1 = 3; 4 + 3 = 4 and and is not unduly influenced by differences in rich- 4 + 2 = 4 and 4 + 1 = 4). Branch order numbers were ness between cells (Born et al. 2007). The unweighted assigned to all branches in the dendrogram using the pair group method with arithmetic means (UPGMA; phytools library for R (Revell 2012). Using this branch Sokal and Michener 1958) agglomerative clustering order assignment, we were able to identify hierarchical
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biogeographical units (comprising grid cells) based on We defined our regional climate zones in terms species similarity. Being intrinsically hierarchical, the of their rainfall regimes, distinguishing winter and resultant dendrograms allowed us to determine the re- aseasonal (rain falling throughout the year, but pre- lationships between biogeographical areas, thus allow- dominantly between autumn and spring) rainfall ing us to test the hypotheses outlined above. Since we areas from those receiving most of their rainfall in were dealing with highly disparate taxonomic groups, summer. Within the winter/aseasonal rainfall zone biogeographic patterns retrieved at different hierar- (Fig 4.1), and concomitant with our stated hypoth- chical levels were interpreted against regional climate eses, we identify mesic (‘winter mesic’; total annual zones and gradients (e.g. whether biogeographic inter- precipitation: 275–900 mm; >65% rainfall in winter) vals are coincident with rainfall seasonality bounda- and arid winter (‘winter arid’; total annual precipi- ries, or recognized mesic/semi-arid intervals (Fig 4.1, tation: 100–275 mm; >65% rainfall in winter) rainfall Rutherford and Westfall 1986)). Also, given the com- zones, and mesic (‘aseasonal mesic’; total annual parative nature of this study and the disparate taxa precipitation: 275–900 mm; 35–65% rainfall in win- analysed, we restricted our investigation to the broad ter) and arid aseasonal (‘aseasonal arid’; total annual patterns reflected in the highest levels of the dendro- precipitation: 100–275 mm; 35–65% rainfall in winter) gram (i.e. primary and secondary breaks; Fig 4.2). rainfall zones.
Figure 4.1 Climatic zones used to describe biogeographic patterns for plant and animal groups within the aseasonal/winter rainfall zone. Seasonal rainfall zonation is as defined by Chase and Meadows (2007) andA ckerly et al. (Chapter 16). The winter rainfall zone is divided into winter-mesic (total annual precipitation: 275–900 mm; >65% rainfall in winter) and winter-arid (total annual precipitation: 100–275 mm; >65% rainfall in winter) zones, as is the aseasonal rainfall zone (mesic: total annual precipitation: 275–900 mm; 35–65% rainfall in winter; arid: total annual precipitation: 100–275 mm; 35–65% rainfall in winter). With the exception of the extreme northwestern corner of South Africa, which receives predominantly (>65%; but <100 mm) winter rainfall, the areas lying outside the rainfall zones indicated receive predominantly summer rainfall.
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4.4 Floristic and faunal biogeographic 4.4.2 Fauna patterns Regional patterns 4.4.1 Flora The animal groups showed considerable variation in the number of primary breaks (regional clusters) ob- Analysis of the plant dataset revealed a clear primary tained, ranging from two for reptiles and butterflies, split between the summer rainfall area and the rest four for birds, and six for frogs (Fig 4.2b–e, upper and of the country, with the winter and aseasonal rainfall lower panels). Although there were some similari- areas forming a single biogeographical unit, indicative ties in biogeographic boundaries across groups, none of a GCFR (Fig 4.2a, upper and lower panels). Nested of the clustering patterns were identical. For reptiles, within the GCFR, as retrieved here, were three subre- the primary division suggested a broad split between gions (Fig 4.2a, middle and lower panels), comprising: temperate and afrotropical reptiles, as retrieved by (a) the winter-mesic zone; (b) the aseasonal-mesic zone; Crowe’s (1990) biogeographic zones for snakes and and (c) the winter-arid zone. The winter-mesic subre- lizards. The precise location of the boundary between gion essentially incorporates Goldblatt and Manning’s these two major groups is somewhat obscured by the (2000) Northwest and Southwest phytogeographic cen- low level of sampling in central South Africa, but ap- tres and generally coincides with the mediterranean- pears to cut directly through the Highveld region, from type climate portion of the broadly winter rainfall zone the Kei River towards the northwest, with the western (Chapter 2). The aseasonal-mesic subregion incorpo- division incorporating the CFR, succulent karoo, thick- rates Goldblatt and Manning’s (2000) Karoo Mountain, et, Nama karoo, and the drier grassland and savanna Langeberg, and Southeast phytogeographic centres. regions. Similarly for birds, a western division, encom- The winter-arid subregion is essentially coincident passing the CFR, succulent karoo, and Nama karoo, with the western succulent karoo region (Chapter 1, was retrieved, but its eastern border shows more lim- see also Born et al. 2007; Snijman 2013). ited eastward and northward extension into savanna The winter-mesic subregion clusters most closely and grassland. Both Chapin (1932) and De Klerk et al. with the aseasonal-mesic subregion, forming an en- (2002) retrieved southwestern divisions for birds tity essentially equivalent to the CFR (sensu Manning comparable to our western division, but with deeper and Goldblatt 2012). These two subregions share a intrusion into savanna and grassland. The southeast- similar geology, namely the Cape Supergroup rocks ern coastal extent of our western division for birds is (Chapter 2). The eastern limit of the aseasonal-mesic essentially limited to the CFR. Chapin (1932) retrieved subregion, however, extends well beyond the eastern a very similar pattern with southern intrusions of afro- boundary of the CFR, stretching as far as East Lon- tropical bird fauna along coastal areas reaching well don (see also Weimarck 1941 and Bradshaw 2009). into the aseasonal-mesic zone. While extensive tracts of fynbos and renosterveld Butterflies showed a similar primary biogeographic vegetation in this area are restricted to the Cape Su- division to plants, centred on the boundary of the pergroup substrata of the Zuurberg range east of summer rainfall zone; however, this division was less Port Elizabeth, Cape floristic elements are com- distinctive than for plants with some summer rain- monly found in coastal grassland and thicket mosa- fall grid cells included within the winter/aseasonal ics, especially on calcareous substrata, well beyond rainfall zone. The primary division retrieved here for the Fish River (Cowling 1983). In contrast to the butterflies does not match either Carcasson’s (1964) winter-mesic and aseasonal-mesic subregions, the zoochorological divisions for butterflies nor Endrödy- winter-arid subregion is associated largely with shales Younga’s (1978) for beetles. Carcasson’s (1964) west- of the Karoo Supergroup and gneiss and granite of the ern ‘Cape Subregion’ division included the CFR, the Namaqua–Natal Metamorphic Belt (Chapter 2). The succulent karoo, and large areas of Nama karoo and southwestern extent of this subregion penetrated fur- grassland. The southeastern coastal extent of our west- ther south along the west coast than the Olifants River, ern division for butterflies, however, matched that of the boundary employed by Goldblatt and Manning Carcasson (1964), who retrieved a strong afrotemper- (2000) to delineate the CFR. This pattern is consistent ate forest element which extends deep into the asea- with Marloth’s (1908) West Littoral area and Acocks’ sonal-mesic CFR, having possibly displaced or limited (1953) vegetation map. Succulent karoo components, the northeastern reaches of fynbos species. By contrast, therefore, reach further south into the CFR than origi- Endrödy-Younga’s (1978) southeastern coastal extent nally thought (Chapter 1, Bradshaw et al. submitted). of Cape faunal elements extended northwards along
9780199679584-Allsopp.indb 81 22/08/14 10:30 AM 82 Fynbos: Ecology, Evolution, and Conservation of a Megadiverse Region Top panel: Geographical extent of the primary (regional) biogeographic clusters resolved by the plant and animal groups included in this study. The eastern boundaries of the aseasonal The clusters resolved by the plant and animal groups included in this study. extent of the primary (regional) biogeographic Geographical panel: Top
or frogs, the secondary decomposition is provided for the two westernmost regions, whereas for all other groups the decomposition is provided for the single westernmost region. Lower whereas for all other groups the decomposition is provided single westernmost region. the secondary decomposition is provided for two westernmost regions, or frogs, F Figure 4.2 Figure extent of the secondary (subregional) clusters identified within westernmost primary Geographical Middle panel: zones are indicated. (dashed lines) and winter (solid rainfall (regions). clusters representing the areas falling outside western primary cluster(s) are showing the relationships of secondary clusters falling within western regions; Simplified dendrograms panel: grid cells probably lack the These Linder 2001). these being a common feature of this type analysis (e.g. Grid cells not assigned to any clusters are shown as dashed lines, indicated by open boxes. instead having only ubiquitous species. taxa linking them to other clusters,
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the east coast into subtropical savanna. Endrödy- biomes (Chapter 1, Cowling 1983) and two rainfall Younga (1978) explained these southeastern extensions zones (Fig 4.1), it supports a diversity of reptile species through the presence of Cape relictual taxa found in drawn from each of these areas, resulting in an unusual the temperate coastal forests and Drakensberg Moun- assemblage of overlapping faunas (see also Chapter 7). tains and the radiation of these (Gondwanan) elements Subregion (b) incorporates succulent karoo, the north- into younger more adaptive groups. western portion of the CFR, Nama karoo, and desert. It Overall, the primary divisions revealed by our anal- offers a wide array of habitats and has many endemic ysis for reptiles, birds, and butterflies indicated broad- habitat specialist species (e.g. several limbless skink ly similar divisions between western temperate and sand specialists along the west coastal dune areas and eastern afrotropical faunas. Frogs, however, showed rupicolous (rock-dwelling) species found on exposed a distinct pattern from the rest of the animal groups, Table Mountain Group sandstone in the Cederberg with six regional biogeographic divisions at the pri- (Vernon 1999; Edwards et al. 2012)). The northeastern mary break. These were not aligned into western and portion of this subregion extends along the Orange eastern areas, as retrieved by (Crowe 1990; see also River to the east, including areas on both sides of Seymour et al. 2001; Alexander et al. 2004). Only two the Orange River. The reptile fauna is dominated by of the six regional clusters, the winter-mesic and pos- rupicolous species, but also has a high proportion of sibly the winter-arid, can be considered to constitute a sand species, since there are large deposits of alluvial Cape faunal division (Seymour et al. 2001; Alexander sand in this arid area. The continuity of sand along the et al. 2004; Linder et al. 2012) which is distinct from Orange River acts as a corridor for many species into the rest of South Africa, the latter being essentially this area which are otherwise associated with rather defined by afrotropical elements. Similar to birds and different habitats (e.g. Varanus niloticus). Subregions insects, these eastern tropical elements penetrate deep (a) and (b) cluster together, linking the CFR, succulent into fynbos along the coast, reaching as far south as the karoo, and Nama karoo. winter-mesic’s southeastern border. For birds, the western division also comprised three subregions (Fig 4.2d, middle and lower panels), Subregional patterns within the western comprising: (a) the winter-mesic and aseasonal-mesic cool-season rainfall zone zones; (b) the winter-arid and aseasonal-arid zones; Focussing within the western divisions for animals and (c) the strongly winter-arid, aseasonal-arid, and (Fig 4.2b–e, middle and lower panels), some similar- northern extent of the summer rainfall zones. Sub- ities were apparent in the delimitation of subregion- region (a) falls predominantly within the CFR. The al clusters, but, as for the primary regional divisions, southeastern coastal border of subregion (a), however, none of the biogeographic patterns were identical. does not reach the CFR’s southeastern limit, reaching Analysis of the reptile data retrieved three subre- just east of Knysna. Terrestrial bird studies by Crowe gions, comprising: (a) the southern winter-mesic and (1990) and De Klerk et al. (2002) both retrieved CFR- aseasonal-mesic zones; (b) the winter-arid and asea- coincident clusters (see also Werger 1978). Crowe’s sonal-arid zones; and (c) the aseasonal-arid zone plus (1990) southeastern coastal (see also Chapin 1932) the northern extent of the summer rainfall zone (this and northwestern borders were similar to ours. In northern subregion weakly defined). Crowe (1990) contrast, the southeastern coastal border of De Klerk grouped areas falling within our northern subregion et al.’s (2002) ‘Fynbos District’ extended beyond the with winter-arid areas. Our northern subregion is, CFR, including thicket, but their northwestern bor- however, considered too obscure to describe in detail, der was similar to ours. Although subregion (a) has likely extending beyond the northern study bounda- a relatively depauperate bird fauna, with only seven ries, and, for this reason, is not discussed further. Fynbos Biome-endemic species, several of these rep- Subregion (a) for reptiles incorporates mostly the resent some of the most ancient African bird lineages, southern and southeastern parts of the CFR (its north- highlighting the unique evolutionary affinities of this western extent did not reach beyond the Cederberg) winter-mesic and aseasonal-mesic subregion. Subre- and incorporated areas of southern Nama karoo, gion (b) for birds incorporates large areas of succulent thicket, and grassland. This subregion shows excep- karoo and the lower and upper parts of the Nama- tional reptile richness and endemism, and is also an Karoo Biome. In the north west it reaches the southern area of high zoogeographical complexity. Since the boundary of the Richtersveld. Both Crowe (1990) and southeastern sector of this subregion, in the vicinity of De Klerk et al. (2002) retrieved similar, but far broad- Port Elizabeth, represents a meeting point of several er, arid subregions, linking the succulent karoo with
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Nama karoo, savanna and grassland areas. They did of butterfly endemism (Mecenero et al. 2013). Many of not, however, separate out the Bushmanland, Rich- the winter rainfall endemic species show distributions tersveld and other desert areas which correspond to spanning the CFR and succulent karoo. For example, subregion (c). The Succulent Karoo and Nama-Karoo within the genus Chrysoritis (Lycaenidae) roughly 18 Biomes share many bird species, including eight spe- taxa with restricted winter-mesic ranges occur in both cies restricted to these two biomes (Vernon 1999). the CFR and succulent karoo. The clustering together Additionally, at least 30 grassland bird species extend of the CFR and succulent karoo for butterflies matches into the Nama karoo across the extensive contact area the pattern retrieved for plants. Congruence between between the borders of the Nama-Karoo and Grass- these two groups was, however, expected, as almost land Biomes (essentially equivalent to the eastern all species of butterfly are phytophagous and associ- border of subregion (b)). Many of these grassland spe- ate strongly with plants. It remains to be established cies also extend westwards into succulent karoo. Sub- whether this pattern will hold for other insect and in- region (c), which is somewhat similar to the Gariep vertebrate groups that are less dependent on plants. centre of endemism of van Wyk and Smith (2001), has Subregion (b) for butterflies is difficult to place bio- a handful of arid-adapted bird species, and which geographically. It resembles Endrödy-Younga’s (1978) extend into succulent karoo and Nama karoo (e.g. southeastern part of his ‘Cape-bilateral extension’ dis- Sclater’s lark (Spizocorys sclateri), which is found on persal pattern of Cape beetle taxa, which extends into the gravel plains of the Richtersveld). There are also thicket and grassland. He considered these extensions savanna bird species (e.g. fawn-coloured lark (Calen- to represent relic areas of Cape taxa. Subregion (b) dulauda africanoides) and sociable weaver (Philetairus clustered with subregion (a) indicating its strong links socius)) that extend across the savanna–Nama karoo with Cape elements. A number of putative Cape insect interface, into the northern areas of the Nama-Karoo groups (richness and endemism concentrated in the and Succulent Karoo Biomes. These arid-adapted winter rainfall zone) extend into aseasonal-mesic and and savanna elements separate subregion (c) from aseasonal-arid areas (e.g. grasshoppers: Dirsch 1965; subregion (b). As for reptiles, the bird data cluster flies: Hesse 1969; Usher 1972; Bowden 1978; Stucken- subregions (a) and (b) together, linking fynbos with berg 1997; Stuckenberg 2000; wasps: Gess 1992; beetles: core areas of succulent karoo and with the central and Holm 1978; Colville 2009). However, the area falling lower areas of Nama karoo. within subregion (b) is also known to include a strong Butterflies revealed two subregions within their representation of East African elements which extend western division (Fig 4.2e, middle and lower panels), their ranges southwestwards along the coast, intrud- comprising: (a) the winter-mesic, winter-arid, and ase- ing into the CFR. Carcasson (1964) considered thicket asonal-mesic zones; and (b) the southeastern extents to be characterized more by forest butterfly elements of the aseasonal-mesic, aseasonal-arid, and summer (e.g. Charaxes (Nymphalidae) and Neptis (Nymphali- rainfall zones (this southeastern subregion poorly de- dae)). The zoogeographical links between Nama ka- fined). Neither the earlier studies on butterflies (Car- roo, thicket, and surrounding winter rainfall clusters casson 1964) nor those on beetles (Endrödy-Younga highlights the interesting and complex faunal com- 1978), the two most detailed insect biogeographic position of subregion (b) with the occurrence of CFR, studies for South Africa to date, clustered the CFR and succulent karoo, Nama karoo, grassland, and savanna succulent karoo together in their zoochorological divi- insect elements being represented. sions. Carcasson (1964) considered the winter-mesic Cape as a separate subregion, whereas Endrödy-Youn- ga (1978), while recognizing that winter-mesic Cape 4.5 Cape biochoria: consensus elements extended into winter-arid areas, nonetheless and differences across taxa considered the northwestern parts of the succulent karoo to be part of a coastal Namib division. Both of Broad congruence across the butterfly, reptile, and bird these early biogeographers delineated the northwest- datasets supports the recognition of western and east- ern winter-arid areas of South Africa as part of an in- ern faunal divisions within South Africa (Table 4.3). dependent zoogeographic region that extended along Plants and butterflies showed the greatest similarity, a narrow band along the coast into northern Angola. A with their western divisions being mostly restricted to recent conservation assessment of South African but- winter and aseasonal rainfall areas. Birds and reptiles, terflies using comprehensive and detailed distribution on the other hand, retrieved a western division incor- maps highlights the winter rainfall zone as a hotspot porating both the winter rainfall and large components
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Table 4.3 Assessment of consensus of hierarchical clustering of biogeographical patterns across taxon groups to evaluate the hypothesis of a Cape Biochorion or Greater Cape Biochorion. Hierarchical clustering proceeds from primary breaks, subdividing South Africa into western and eastern regional divisions, through to lower level subregional delimitation. Nested Biogeographic consensus Biotic group levels plants reptiles frogs birds butterflies
I Primary split between broad ü ü û ü1 ü western versus eastern divisions
II Winter-mesic (Core Cape ü û ü ü û Biochorion)
II Winter-mesic + aseasonal-mesic ü û û ü2 û (Cape Biochorion)
II Winter-mesic + aseasonal-mesic ü ü3 û û û + southeastern extensions (Cape Biochorion with extensions) Primary division and chorology
III Winter-mesic + winter-arid + ü û û û ü4 aseasonal-mesic + aseasonal-arid + southeastern extensions (Greater Cape Biochorion)
III Winter-mesic + winter-arid + û ü û ü û aseasonal-mesic + aseasonal-arid + southeastern summer + northern summer (Greater Cape Biochorion with extensions)
1 There is a strong single primary division in the west, but multiple eastern divisions (see Fig 4.2d). 2 Incomplete inclusion of aseasonal-mesic zone. 3 Truncated northwestern winter-mesic zone. 4 Some inclusions of aseasonal-arid and summer rainfall zone areas.
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of the aseasonal-arid Nama karoo region and some between plants and animals is of importance when summer rainfall regional areas (Table 4.3). In contrast attempting to understand the development of the biota to the other groups, frogs did not display a broad pri- of both the CFR and the adjacent succulent karoo. The mary division, but rather revealed six clusters split at unequivocal support for the validity for a Greater Cape the primary level, with their western temperate faunal Floristic Region (Bayer 1984; Jürgens 1991; Born et al. division essentially reduced to a narrow, winter rain- 2007), using a comprehensive national species-level fall area consisting of two separate regions. plant dataset, is a significant finding. Plants showed Clear evidence of a Greater Cape Biochorion was a clear primary split between the summer rainfall area only found in plants and butterflies, in which winter- and the remainder of the country. Floristically, this in- mesic, winter-arid, and aseasonal-mesic zones grouped dicates that the floras of the CFR and succulent karoo together into a nested biogeographic unit (Table 4.3). are more similar to each than to any of the floras in In contrast to plants and butterflies, the winter-arid ar- the rest of South Africa. Plant biogeography therefore eas for reptiles and birds appeared to show greater fau- appears best considered within a GCFR concept (see nal affinity with aseasonal-arid Nama karoo areas to Manning and Goldblatt 2012; Snijman 2013). Faunisti- the east, rather than to winter-mesic areas to the south. cally, however, patterns retrieved in this study suggest For birds, the retrieval of two winter-arid subregions, that efforts to understand the ‘Cape’ biogeography linked to the southern and northern parts of Nama of animals will require a broader focus, extending karoo, highlights a deeper biogeographic complexity beyond the confines of the CFR and succulent karoo, between these two arid areas. Therefore, South African to incorporate areas of eastern neighbouring biomes. vertebrate groups appear not to conform to a Greater Additionally, the essentially floristically defined Cape Biochorion (Table 4.3, sensu lato Bayer 1984, Born biomes of South Africa appear to show limited similar- et al. 2007). For reptiles and birds, the arid subregions ity with faunal biogeographic patterns (van Rensburg (winter-arid plus aseasonal-arid) do, however, cluster et al. 2004; Procheş and Cowling 2007). with the winter-mesic and aseasonal-mesic subre- The environmental factors limiting the distribu- gions. The Greater Cape Biochorion, as retrieved for tions of plants and different animal groups, therefore, plants and butterflies, forms a core area of this group- appear to be quite variable. Several faunally defined ing. We therefore consider a Greater Cape Biochorion divisions, for example, reach beyond the confines of with extensions (of eastern aseasonal-arid/summer the rock types (Cape Supergroup) which are so criti- rainfall areas) as an appropriate biogeographic unit for cal in limiting the fynbos flora, as well as beyond the reptiles and birds (Table 4.3). winter-dominant rainfall area, well into and sometimes Within the plant- and animal-based Greater Cape beyond the aseasonal-arid zone. This can be seen in Biochorion divisions, the winter-mesic and aseasonal- reptiles, with species shared between the succulent ka- mesic rainfall areas retrieved differed substantially roo and Nama karoo with areas of thicket (Chapter 7, between groups in terms of both their bounds and ex- Vernon 1999; Meyer et al. 2010). Likewise, the distri- tents (Table 4.3). Broad support for a Cape Biochorion butions of several bird species appear to be climati- was evident in three of the five groups (plants, frogs, cally defined, resulting in distributions that cut across and birds), each of which retrieved a core winter-mesic vegetationally defined habitats which may be more area, with a similar northwestern border. The extent of closely linked to soil properties. Consequently, some the aseasonal-mesic area incorporated by each of these bird species are shared between the Cape, succulent groups was variable, however, as was the location of karoo, Nama karoo, and grassland regions (see also the coastal southeastern border. For plants, frogs, and Allan et al. 1997). birds, therefore, a Cape Biochorion is considered as Another pattern of biogeographic interest is the an appropriate biogeographic unit, but with variable variable position of the southeastern coastal border southeastern, aseasonal-mesic extensions (Table 4.3). of the aseasonal-mesic cluster across plants and ani- mals. Plant data identify the southeastern border of 4.6 Interpretation of biogeographic the aseasonal-mesic cluster as extending further east similarities and differences than the current CFR border. This easterly extension may reflect the underlying eastern extent of Cape The regional biogeographic patterns retrieved here for Supergroup rocks (Chapter 2), which may account plants and animals within the GCFR highlight several for the presence of Cape floristic elements as far as interesting and significant new findings. The retrieval the Great Fish River. This area receives some winter of both contrasting and similar biogeographic patterns rainfall (Fig 4.1) and is thus capable of supporting a
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flora that is adapted for winter growth. The crudeness we should not expect these taxa to respond to these of our grid cell units may, however, account for at least drivers in a uniform way. Frogs, for example, are ec- part of this pattern. Analysis of taxonomic datasets tothermic and mostly require water for survival and partitioned into biotic elements (high versus low alti- reproduction. They are also less vagile than birds and tude, mesic versus xeric, fynbos versus non-fynbos), butterflies. These physiological and life-history traits or at finer spatial scales, would be required to assess most probably make frogs more susceptible to habitat more accurately the eastern extent of the GCFR. The idiosyncrasies which can lead to narrow ranges and aseasonal-mesic and southeastern aseasonal-arid areas more detailed regional patterns (Rueda et al. 2010). are faunistically interesting in that they represent an The greater vagility of birds, by contrast, enables them area of transition or overlap between Cape clades and to track habitats (e.g. wooded river courses) and to fol- afrotropical elements. This limits the consensus for a low pulses of resource availability related to seasonal- Cape Biochorion, again emphasizing that chorological ity, as seen between areas of succulent karoo (winter divisions depend heavily on the taxa examined. arid) and Nama karoo (aseasonal arid; Allan et al. A lack of biogeographic congruence across groups is 1997). In this context, it is unsurprising that the fauna not unusual (see Rueda et al. 2010 for regionalization of and flora of the GCFR show different histories of as- European biota). One may expect that where primary sembly (Chapters 5, 7) and patterns of biogeographic producers show strong biogeographic pattern (i.e. the association. Understanding biogeographic structure unique GCFR vegetation), this will have a direct influ- and the formation of Cape biochoria requires consid- ence on the patterns shown by consumers. However, eration of a complex array of historical and ecological animal groups often show minimal fidelity to veg- factors, each of which will interact with a taxonomic etation types (Liversidge 1962; Allan et al. 1997; Cox group in an idiosyncratic manner. 2001; Jenkins et al. 2013), being aligned instead with other habitat variables (e.g. soil structure for scorpions 4.7 Conclusion (Prendini 2001a), insects (Endrödy-Younga 1978; Irish 1990; Sole et al. 2004; Botes et al. 2007), and reptiles Our analysis for plant and animal biogeographic pat- (Bauer 1999); aquatic habitats for frogs (Mokhatla et al. terns has the advantage of covering all of South Africa, 2012) and insects (Samways and Niba 2010); and nest- thereby providing a rigorous test of the relationships ing and roosting sites for bats (Monadjem et al. 2010)). between cool-season rainfall areas and summer rainfall Modern phylogenetic studies of Cape faunal groups areas. Only in plants and butterflies did the winter rain- provide support for this idea, linking areas not neces- fall and aseasonal-mesic areas cluster to form a Greater sarily along floristic lines, but rather along geographic Cape Biochorion in the traditional sense. For birds or altitudinal gradients, such as mountain ranges and and reptiles, winter-arid areas showed greater affinity lowland areas (e.g.Chapter 7, Prendini 2001b; Measey to the aseasonal-arid Nama karoo and arid summer and Channing 2003; Sole et al. 2004; Pitzalis and rainfall areas, than the winter-mesic areas to the south. Bologna 2010; Tolley et al. 2010; Predel et al. 2012). Frogs, on the other hand, retrieved the winter-mesic Several interrelated factors have undoubtedly been and winter-arid areas as distinct regions. Thus, limited influential in generating the differences outlined consensus was found for a Greater Cape Biochorion us- above. The winter–summer rainfall regime interface ing the distribution data of plant and different animal is an important boundary, and therefore driver, of taxa. For birds and reptiles, however, the CFR clustered biogeographic pattern (Linder and Mann 1998; van with succulent karoo and the aseasonal-arid parts of Wyk and Smith 2001; Cowling and Ojeda 2005). In ad- Nama karoo indicating that the faunal affinities of suc- dition, the Greater Cape region is an area of discrete culent karoo and Nama karoo appear closest with the geological boundaries (Chapter 2, Cowling et al. 2009), Cape fauna. Therefore, we consider a Greater Cape Bi- contrasting fire regimes (Chapter 3), habitat heteroge- ochorion with extensions as a concept for investigating neity (topography, climate, and soils; Chapter 2), and Cape faunal biogeography. The Nama-Karoo Biome concomitant ecological specialization (Chapters 6, 10, has, however, been poorly surveyed for most taxo- 11). These factors, explored elsewhere in the book, nomic groups (Gibbs Russell et al. 1984; Vernon 1999). have undoubtedly played a central role in structur- Increased collection and exploration of the Nama karoo ing the unique biota of the GCFR. Given the strongly may either strengthen its faunal relationship with the contrasting biologies and life-history requirements of winter rainfall areas, or highlight its faunistic distinct- plant and animal lineages, however, as well as their ness. A Cape Biochorion centred on the winter-mesic contrasting dispersal capabilities (Croteau 2010), CFR is a core area for plants, frogs (regional) and
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birds. However, the retrieval of a CFR-centred Cape Alström, P., Olsson, U., and Lei, F. (2013). A review of the Biochorion, incorporating both the winter-mesic and recent advances in the systematics of the avian superfam- aseasonal-mesic zones, requires consideration of dif- ily Sylvioidea. Chinese Birds, 4, 99–131. fering southeastern coastal borders between plants and APG3 (2009). An update of the Angiosperm Phylogeny Group classification for the orders and families of flower- animals related to the falling-off or extension of Cape ing plants: APG III. Botanical Journal of the Linnean Society, clades. The floristic coherence of the GCFR and CFR are 161, 105–21. not, therefore, necessarily reflected in the biogeograph- Balinsky, B.I. (1962). Patterns of animal distribution on the ic patterns of other taxonomic groups, as the ecological African continent. 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