Patterns of endemism in African birds: how much ...

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Patterns of endemism in African birds: how much does taxonomy matter? Jon Fjeldså To cite this article: Jon Fjeldså (2003) Patterns of endemism in African birds: how much does taxonomy matter?, Ostrich, 74:1-2, 30-38, DOI: 10.2989/00306520309485367 To link to this article: http://dx.doi.org/10.2989/00306520309485367

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Patterns of endemism in African birds: how much does taxonomy matter? Jon Fjeldså

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Centre for Tropical Biodiversity, Zoological Museum, Universitetsparken 15, DK-2100 Copenhagen, Denmark e-mail: [email protected]

It has been suggested that the ‘right’ biogeographic patterns will only arise with a species concept reflecting the finest splitting of terminal phylogenetic branches. The significance of taxonomic resolution was assessed using distributional data for all resident African land-birds, held at the Zoological Museum of Copenhagen. The primary database (in a one-degree geographical grid) served as a template for creating two new databases: one, where the distributional records were allocated to species according to the ‘Speciation Atlases’ for African birds (Hall and Moreau 1970, Snow 1978) and two, according to the finest taxonomic splitting that has since then been suggested in the literature. With this spatial resolution, the species richness pattern is almost exactly the same whether old or new taxonomy is used. The endemism follows another pattern with marked local aggregates. The location of principal aggregates are quite robust to recent splitting, although a slightly more complex structure appears. Also some small new areas of endemism emerge, each with 2–3 narrow endemics. However, large portions of African savannah and lowland rainforest are still characterised by a total lack of narrow endemism. Based on the current understanding of diversification processes and adaptive re-distribution it is suggested that the uneven distribution of narrowly endemic and widespread species will persist even if it were possible, in the future, to define terminal taxa in a consistent way.

Introduction In recent years, approximately one genuinely new bird species is discovered per year in Africa, but a much higher number of ‘new’ species is generated by taxonomic revision. Many birds, referred to with binominals in the 19th and early 20th centuries, were lumped (as subspecies under a broad biological species concept) in various volumes of Peters’ Checklist and in the ‘Speciation Atlases’ of African birds (Hall and Moreau 1970, Snow 1978). It was primarily explorernaturalists who emphasised the equivalency of allo- and parapatric forms, thus fostering the concept of broad biospecies. However, the next generation of field biologists, who paid attention to vocalisations and interaction between populations, found that many forms that replace each other in adjacent geographical areas effectively maintain their integrity in spite of possibilities for contact, and therefore qualify as species. Such insights were supplemented with those gained using molecular data, which repeatedly demonstrate that broadly defined bio-species did not reflect the historical relationships of populations (e.g. Klein et al. 1993, Leisler et al. 1997, Roy et al. 1998, Ryan and Bloomer 1999, Beresford 2002, Bowie 2003, Bowie et al. in press). Traditional interpretations of speciation were replaced by new ones (Moritz et al. 2000), and also the perceptions of what constitutes a species changed (Zink 1997). The biological species concept (Mayr 1942, 1962), which was part of a framework for allopatric speciation, gave way to ideas that are more in agreement with phylogenetic systematics. A consequence of this was to define species as the smallest ‘diagnosable clusters of individuals within which there is a parental pattern of ancestry and descent’ (Eldredge and Cracraft 1980, p 92). Since historical descent and reproduc-

tive isolation jointly illuminate biotic discontinuity, the discrepancy between the concepts is in fact smaller than what the heated debate might suggest. It is sometimes claimed that, with a consistent mapping of the smallest diagnosable evolutionary units, complex new patterns of endemism would emerge (e.g. Daugherty and Triggs 1990, Crowe and Brooke 1993 reviewing ICBP 1992, Cracraft 1983, Beresford and Cracraft 1999). The widespread use of a broad (Mayrian) biospecies concept admittedly translates into lack of attention to many distinctive local forms with their own unique adaptations (Barrowclough 1992, Hazevoet 1996). This view is particularly relevant in conservation biology, where management units are often defined using molecular markers (see Avise 1989, Arctander et al. 1996, 1999). However, conservation work is constrained, in most nations, by the fact that the legislation deals with formally accepted species. The dispute about what is the right species level classification continues, and some taxonomists argue that enormous efforts are still needed before true biogeographic patterns can be described and conservation targets defined (examples in Cracraft and Grifo 1999). Rather than accepting this defeatist view I ask, in this paper, how much the lack of consensus matters. Thus I examine how much species diversity patterns in Africa are affected by the taxonomic research that took place since the publication of the abovementioned ‘Speciation Atlases’. Since nobody tried to revise the classification of African birds in a consistent way (for instance, identifying the terminal branches) it is too early to evaluate the consequences of specific taxonomic philosophies. My aim was therefore to create two new databases

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that will allow me to test how sensitive the biogeographic pattern is to the taxonomic splitting that has been proposed. Methods


Computer software and distribution data Computer programs have been developed to accommodate distribution data for large numbers of species and provide gridded maps showing various taxic diversity measures. I used the WORLDMAP software (Williams 1992), a PCbased graphical tool designed for fast, interactive handling of distributional data for large numbers of taxa. Distribution data Distributional data for the resident birds of sub-Saharan Africa were compiled over several years (as part of a research programme for describing and explaining biogeographic patterns and develop efficient proactive conservation plans; see Brooks et al. 2001, and www.zmuc.dk/commonweb/research/blueprint-africa.htm for data sources). The data were entered in a geographical projection grid of one geographical degree (12.000km 2 at the equator), a compromise between sampling unevenness and the need for a fine resolution in geologically complex parts of Africa. This is probably the finest geographical resolution permissible today for such a large geographical area. Sampling bias is reduced by conservative interpolation: considering the dynamic nature of species distributions (the neutral dispersal assembly perspective of Hubbell 2001) it was assumed that species are at least temporarily present between collecting points, provided that the grid-cells include appropriate macrohabitat. This was determined using numerous topographic and ecological maps and satellite images. However, the interpolation was conservative, and takes into account published evidence indicating distribution gaps. For species that are considered to be genuinely rare or local, only confirmed records were used. A rarity-based heuristic algorithm (Williams 1999) was used to select priority sets of complementary conservation targets. Complementarity explicitly describes the degree to which a target area contributes otherwise un-represented taxa to a set of areas (Vane-Wright et al. 1991, Humphries et al. 1996). The rarity-based algorithm produces results very close to optimal (Csuti et al. 1997), and unlike ‘optimal’ linear-programming branch-and-bound methods it allows the flexibility of the result to be assessed. Taxonomies Species concepts Two new databases were created by copying the original database, defining new ‘taxon identification cards’, and then moving the original records between these ‘cards’ as appropriate. Thus the distribution input is the same between the datasets; it is only the taxonomic allocation of records that is changed (except in cells representing a suture between two ‘new’ species, and a 0.017% increase of data input for 28 genuinely new-discovered species). The new databases were: (1) Speciation Atlas taxonomy: Here I combined all distributional data for taxa assembled under binominals by Hall and

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Moreau (1970) and Snow (1979), notwithstanding that these authors suggested that some of these might comprise two or more incipient species. 1 595 species were recognised in this way (resident species, excluding seabirds and some palaearctic species that are very marginally present in the study area, and neglecting all migrant visitors). (2) Current taxonomy: Since there is no consistent classification of terminal taxa I used the finest species splitting that has been suggested in publications since the appearance of the Speciation Atlases. For this I reviewed taxonomic and regional handbooks, group revisions and recent comprehensive molecular studies. Various people have disputed the validity of some splits, but I will consider this personal opionions, except when concrete evidence is presented documenting that a split was not justified. Genuinely new-discovered species were included, except Hirundo perdita, whose breeding area is unknown. The number of species was raised by 215 to a total of 1 810 species (Appendix 1). Results Taxon richness Since the majority of new species represent geographical replacements, recent splitting does not lead to changes in species richness that are visible with this kind of graphical presentation. The difference in individual grid cells amount mainly to 0–3 species (maximum 6, or 1.3%) in zones with many species replacements. I therefore show only one species richness map (Figure 1). The highest species density is in areas with great topographic complexity (maximum 611 ‘current’ species in the Semliki/Rwenzori grid-cell on the Congo/Uganda border) and in East African habitat mosaics and in the savannah woodlands of Zambia and Zimbwabwe. Endemism Patterns of endemism were assessed using two kinds of cutoff limits: (1) the lower quartile (25%) of range-sizes in the two datasets (see Gaston 1994 for rationale), in which case different area thresholds mean that the resulting maps

Figure 1: Species richness of resident birds in sub-Saharan Africa. This map, made from the ‘Speciation Atlas’ dataset, is virtually indistinguishable from that based on the ‘Current’ taxonomy


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(Figures 2A and B) are not fully comparable; and (2) fixed area thresholds, namely species inhabiting maximum 20, and maximum 10 grid-cells (Figures 3 and 4). For these latter comparisons I excluded marginal species (along the northern range boundary) that would exceed the threshold if their extra-limital distributions were taken into account. Considering that grid-cell areas are larger than areas of occupied habitat, the 10-cells threshold may in fact be close to the 50.000km 2 threshold used by BirdLife International to describe global patterns of endemism (Stattersfield et al. 1998). Recent systematic splitting leads to an increase in number of range-restricted species. Thus the lower quartile is 446 species distributed in maximum 31 grid-cells (versus 398 species in maximum 45 cells when following the ‘Speciation Atlas’ data). Using the 20-cell threshold gives 336 (versus 215) species, the 10-cell threshold 195 (versus 117) species. The endemism is quite locally aggregated (Figures 2–4) with the principal centres located in the Cameroon Mts, along the Albertine Rift, in the Kenya highlands and Eastern Arc Mts of Tanzania, along the Angola Scarp, at the Cape and in the Ethiopia highland. Only in some parts of the continent do these aggregates correspond to areas of high species richness (De Klerk et al. 2002). Using the ‘lower quartile’ criterion, the ‘Speciation Atlas’ data give a rather ‘smooth’ picture with extensive areas with moderately high endemism (Figure 2A), reflecting the large area threshold (45 cells, see above). The most localised pattern appears when using the 10-cell threshold, which means that the Cape and much of the Ethiopia Highland do no longer figure as areas of endemism (Figure 4). Although the 216 ‘new’ species contribute significantly to the total number of endemic species, the geographic pattern that they describe did not change much. Among the main centres of endemism, only the Eastern Arc area became clearly more conspicuous using ‘Current’ data, and the narrow endemism also increased on the north side of the African Horn, between Danakil/East Ethiopia Highland and

the Ahl Madow escarpment. Small changes are also visible outside the principal centres: a slightly more complex pattern of endemism appears around the Bight of Biafra, with scattered new endemics in the rugged Ogodué-Sangha district (see especially Figure 3B). Small new areas, each with 2–3 narrowly endemic species (more or less neglected by BirdLife International, Stattersfield et al. 1998) appear in the northern Savannahs (most evident at Jos, Kano and south of Lake Tchad), in the lowlands of Uganda and interior of Tanzania (south of Lake Victoria), on Mt Kulal in northern Kenya and Taita Hills in the south (see Brooks et al. 1998), in the coastal forests of Lindi, south-eastern Tanzania, scattered in the uplands of southern Congo/northern Zambia (corresponding to the Mwinilunga centre of plant endemism), in southern Zambia, in the high grasslands of the Drakensberg Massif and in the Namib Desert. High endemism in Damaraland/Namib desert and in Katanga in Figure 2A is reduced in Figure 2B because of the different threshold range-size, but small new peaks appear in Ovambo-Etosha and southern Namibia.

A

B

Near-minimum set A near-minimum set based on ‘Speciation Atlas’ taxonomy comprises 59 areas of which six are irreplaceable, 21 conditionally flexible (in which case selection of alternative areas may require a higher total number of areas) and 32 fully flexible. The minimum set using ‘Current’ taxonomy is 87 areas, of which 25 are irreplaceable, 16 conditionally flexible and 46 fully flexible. The increased number of irreplaceable cells is due to the increased number of single-cell endemics, mainly in the montane regions of eastern Africa (Ethiopia to Tanzania) and in the adjacent Congo/Zambia border-zone. Because of the flexibility of such minimum sets, it is not meaningful to focus on the amount of mis-match between them, but more appropriate to see how well the ‘Speciation Atlas’ minimum set covers the ‘Current’ species. This is done by pre-selecting the 59 areas identified from the first database in the other database, to identify the residuum of ‘new’ species not covered by these 59 cells.

Figure 2: Endemism of resident African birds, expressed as richness of species in the lower quartile (25%) of range-sizes. A: based on the ‘Speciation Atlas’ data (398 species with 44-cells cut-off range-size). B: based on the ‘Current’ data (443 species and 31-cells cut-off rangesize)


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A

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B

Figure 3: Endemism of resident African birds, expressed as richness of species with a range of maximum 20 grid-cells. A: 215 species from the ‘Speciation Atlas’ dataset. B: 333 species from the ‘Current’ data

A

B

Figure 4: Endemism of resident African birds, expressed as richness of species with a range of maximum 10 grid-cells. A: 117 species from the ‘Speciation Atlas’ dataset. B: 193 species from the ‘Current’ data

The result (Figure 5) is that 52 ‘new’ species (out of 215) would not be covered, and that 39 extra cells would be needed (on top of the 59 cells) to cover all 1 810 ‘Current’ species. These 52 ‘new’ species are found locally in the northern savannas (Cisticola dorsti, Prinia fluviatilis, Lagonosticta virata and sanguinodorsalis, Vidua mariae), Upper Guinea forest (Phyllastrephus leucolepis, Prinia leontica), Sangha area (Stiphrornis sanghensis), Congo-Sudan boundary (Psalidoprocne mangbettorum and olaeginia), Ethiopia (Caprimulgus solala, Psalidoprocne oleaginia, Mirafra degodiensis, Heteromirafra sidamoensis, Calandrella blanfordi, Lagonosticta larvata), Somalia (Laniarius liberatus), northern Kenya (Zosterops kulalensis), Uganda (Sylvietta chapini, Ploceus ugandae), along the Eastern Arc Mts (Xenoperdix udzungwensis, Turdus helleri, Zosterops silvanus and Z. winnifredae, unnamed Sheppardia sp., Apalis fuscigularis, Euplectes psammocromicus, and in the adjacent Kilombero swamps Ploceus burnieri and two unnamed Cisticola spp.), in south-

ern coastal Tanzania (Stactolaema woodwardi, Batis reichenowi), along the Albertine Rift (Glaucidium albertinum, Muscicapa itombwensis, Apalis kaboboensis), along the Malawi Rift (Agapornis lilianae, Euplectes psammocromicus, Apalis flavigularis and lynesi), in Katanga/Zambia (Agapornis lilianae, Stactolaema sowerbyi, Cisticola melanurus, Apalis kaboboensis, Sylvietta chapini, Ploceus ruweti and katangae), western Angola (Laniarius brauni, Certhilauda benguelensis), Namibia (Certhilauda barlowi and burra), the Karoo (Certhilauda subcoronata, Anthus longicaudatus and pseudosimilis) and Drakenberg highland (Heteromirafra ruddi). Some of these species could be covered, though, by taking into account the flexibility in the ‘Speciation Atlas’ minimum-set. Discussion The species richness pattern is not affected by taxonomic change, as the taxonomic changes consist mainly of splitting


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Figure 5: Target areas for conservation of resident African birds. Dots mark a near-minimum set of complementary areas based on ‘Speciation Atlas’ data, and the grey shades represent the distribution of the 52 new species (in the ‘Current’ dataset) that are not covered in these former cells. A total of 39 extra cells are needed to cover these 52 species

of populations that inhabit complementary geographic areas. The richness pattern is dominated by the widespread species, since the 25% of most-widespread species contribute 70.5% of the total data input; the density of such species being effectively explained by current environmental factors, most importantly Net Primary Productivity (Jetz and Rahbek 2002). The few genetic studies of widespread species suggest considerable phylogenetic age and genetic variation but often an unclear historical population structure (e.g. Simonsen et al. 1998, Bowie 2003 for Nectarinia olivacea), suggesting considerable population movements and gene flow. Apparently, the distribution of such species is dynamic, as they respond to the climatic instability that characterises large parts of Africa. According to biogeographic null models, endemism is assumed to be nested within areas with high species richness (Gaston 1994). This holds true in some parts of Africa (notably in the Guinea-Congolean rainforest region, Fjeldså et al. in press) but not in others. Some parts of Africa are species-rich but have very few endemics, and some peaks of endemism are somewhat displaced compared with richness peaks or positioned near the margins of the continent in moderately species-rich areas (Jetz and Rahbek 2001, De Klerk et al. 2002). Overall, narrow endemism correlates poorly with environmental variables other then altitudinal range. Since the lower quartile of range-sizes contributes only with 1.3% of the total data input (Jetz and Rahbek 2002) the endemics contribute little to the overall species richness pattern, but it is important to note that it is mainly among these species that we find Africa’s most threatened birds (BirdLife International2000 and Figure 7b in Brooks et al. 2002). The principal centres of endemism changed only slightly because of recent splitting (Figures 2–4), reflecting a high spatial congruence between distributions of very distinctive geographical relics (many of them monotypic genera) and

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weakly differentiated neoendemics, many of which were recently assigned species rank. Fjeldså and Lovett (1997) interpret this congruence as a result of accumulation of relict taxa over long time spans in places with predictable ecological conditions. Stability may relate to local orographic moderation of moisture-giving winds and mist formation, and steepness of the terrain, which makes it possible for endemic species to adapt to climatic change by moving a few hundred metres up, or down, the mountain slopes (Fjeldså et al. 1999). It is sometimes suggested, although rarely in the literature, that a large number of new areas of endemism would appear if a phylogenetic species concept were adopted, recognising all terminal phylogenetic branches. According to Figures 2–4, taxonomic splitting adds some complexity to known zones of endemism, and it gives rise to a few new areas with 1–3 endemic species each. Undoubtedly, further studies will strengthen some of these areas. However, it is worth noting that large parts of Africa are still characterised by widespread species with subtle and intergrading subspecies and a total absence of narrow endemics. Fjeldså (2000) analysed biogeographic patterns for Andean birds, using a similar approach as in this paper. Also in this case, the main peaks of endemism were fairly robust to taxonomic change, as a consequence of how endemic species are aggregated in certain montane areas. The aggregated nature of endemism is less evident in tropical lowlands. The majority of savannah birds are widespread, with subtle geographical variation, and most ‘new’ species in the woodland savannas are patchily distributed over a moderately wide area. Molecular studies of Guinea-Congolian forest birds suggest much more phylogeographic structure than believed before (Beresford 2002). Although the endemism will probably rise in the Upper Guinea Rainforest, in Cameroon-Gabon and near the eastern margin of the Congo Basin, it remains to be seen how many definite geneflow breaks will be found in the widespread species, and how many ‘new’ species will qualify as narrow endemics. Roy et al. (1997) point out a clear distinction between ecoregions characterised by well-marked phylogeographic structure, and others (such as lowland floodplains and inherently unstable savannah regions) that are dominated by widespread and geographically-overlapping species, mostly of considerable phylogenetic age. Such differences lead to complex and scale-dependent variations in richness on different taxonomic levels. Overall, in situ evolution of new species may be restricted to rather few areas with wellmarked topography, and to some parts of the ‘Arid Corridor’, mainly in the Namib/Karoo and Horn of Africa areas (see Arctander et al. 1996 and 1999 for diversification of savannah mammals). Based on the current understanding of the processes of speciation and adaptive re-distribution, I believe that the uneven distribution of narrowly endemic and widespread species will persist even if it were possible, in the future, to define terminal taxa in a consistent way. Recent efforts to make global conservation priorities have used endemic bird areas (Stattersfield et al. 1998), ecoregions (Olsen and Dinerstein 1998) and ‘hotspot’ approaches (Mittermeier et al. 1998). Especially the two latter kinds of priority areas are substantial in size, so the area-


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efficiency and precision for conservation planning is low (Burgess et al. 1998). Conservation actions need to be precisely targeted, and this requires effective and flexible analytic algorithms (Pressey and Nicholls 1989, Williams 1999) and precise distributional data, which can identify true evolutionary units and areas that are important for the diversification process. Focusing on larger regions where high biodiversity is maintained by high ecoclimatic heterogeneity makes it difficult to define precise targets and is a failure when it comes to targeting the threatened species. Although recent systematic revision did not alter the geographic patterns of avian diversity much, it allows us to more precisely target unique populations of conservation concern. For instance, Figure 5 identifies needs for new conservation initiatives in Katanga and adjacent parts of Zambia, and in the Namib and Karoo areas (South Africa). If we want conservation plans that cover the full avian diversity, the efforts to map and analyse diversification patterns must continue, and this requires continued collecting effort and detailed phylogeographic analysis. Acknowledgements — The development of the databases would not have been possible without long-term support from the Danish Research Council and the Danish Council of Development Research. Pamela Beresford and Rauri Bowie are thanked for keeping me updated about their ongoing molecular studies. I wish to thank Mr Louis A Hansen for assistance with the graphic presentations and literature search, C Rahbek for stimulating discussions, and P Beresford and RJ Dowsett for comments on an earlier manuscript.

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Ornithologists’ Club 116: 225–229 Pressey RL and Nicholls AO 1989. Efficiency in conservation evaluation: scoring versus iterative approaches. Biological Conservation 50: 199–218 Prigogine A 1975. The status of Estrilda kandti and Estrila atricapilla graueri. Bulletin of British Ornithologists’ Club 95: 1518 Prigogine A 1983. Un noveau Glaucidium de l’Afrique centrale (Aves, Strigidae). Revue Zoologie de Afrique 97: 886–895 Prigogine A 1985. Recently recognized bird species in the Afrotropical Region — a critical review. Proceedings International Symposium on African Vertebrates: 91–114 Roy MS, Arctander P and Fjeldså J 1998. Speciation and taxonomy of montane greenbuls of the genus Andropadus (Aves: Pycnonotidae). Steenstrupia 24: 51–66 Roy MS, da Silva JC, Arctander P, García-Moreno J and Fjeldså J 1997. The role of montane regions for the speciation of South American and African birds. In: Mindell DP (ed) Avian Molecular Evolution and Systematics. pp 325–343. Academic Press, San Diego Ryan PG and Bloomer P 1999. The long-billed lark complex: a species mosaic in southwestern Africa. Auk 116: 194–208 Ryan PG, Hood I, Bloomer P, Komen J and Crowe TM 1998. Barlow’s Lark: a new species in the Karoo Lark Certhilauda albescens complex of southwest Africa. Ibis 140: 605–619 Safford RJ, Asj JS, Duckworth JW, Tefler MG and Zewdie C 1995. A new species of nightjar from Ethiopia. Ibis 137: 301–307 Sibley GC and Monroe BL 1990. Distribution and Taxonomy of the Birds of the World. Yale University Press, New Haven Simonsen BT, Siegismund HR and Arctander P 1998. Population structure of African buffalo inferred from mtDNA sequences and microsatellite loci: high variation but low differentiation. Molecular Ecology 7: 225–237 Smith FG, Arctander P, Fjeldså J and Amir OA 1991. A new species of shrike (Laniarius) from Somalia, verified by DNA sequence data from the only known individual. Ibis 133: 227–235 Snow NW 1978. An Atlas of Speciation in African Non-passerine Birds. Trustees of the British Museum (Natural History), London Stattersfield AJ, Crosby MJ, Long AJ and Wege DC 1998. Endemic Bird Areas of the World. Priorities for Biodiversity Conservation. BirdLife International, Cambridge

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Received August 2002, accepted October 2002 Editor: WRJ Dean

Appendix 1: Taxonomic changes since the ‘Speciation Atlases’ (Hall and Moreau 1970, Snow 1978) Lumping and splitting of African birds have been discussed in a large number of papers but when possible I refer to the taxonomy used in ‘Birds of Africa’, where references can often be found to primary sources. It should be noted that many of these and other splits have been questioned, especially by Dowsett and Dowsett-Lemaire (1993). Since the aim of this paper was to examine how splitting affects patterns of endemism, and not to provide a ‘correct’ taxonomy, I ignored such opinions, except when actual information was presented that clearly documented that suggested splits were not justified. Genuinely new species are: Xenoperdix udzungwensis (Dinesen et al. 1994); Melignomon eisentrauti (Louette 1981); Glaucidium albertinum (Prigogine 1983), Caprimulgus prigoginei (Louette 1990) and C. solala (Safford et al. 1995); Mirafra ashi (Colston 1982), M. degodiensis and Heteromirafra sidamoensis (Erard 1975); Laniarius liberatus (Smith et al. 1991); Sheppardia sp. nov. (Beresford et al. submitted); Stiphrornis sanghensis (Beresford and Cracraft 1999); Melaenornis annamarulae (Forbes-Watson 1970); Chlorocichla prigoginei (De Roo 1967), Phyllastrephus leucolepis (Gatter 1985); Acrocephalus avicenniae (Ash et al. 1989, for species rank see Leisler et al. 1997); Cisticola dorsti (Chappuis and Erard 1991) and two unnamed Cisticola spp. (Stevenson and Fanshawe 2002; JF, own observations); Nectarinia rufipennis (Jensen 1983); Anthus longicaudatus (Liversidge 1996) and A. pseudosimilis (Liversidge and Voelker 2002); Ploceus ruweti (Louette and Benson 1982, but see Dowsett and Dowsett-Lemaire 1993), P. burnieri (Baker and Baker 1990) and P. victoriae (Ash 1986, but see Dowsett and Dowsett-Lemaire 1993); Malimbus ballmanni (Wolters 1974); Lagonosticta sanguinodorsalis (Payne 1998); Vidua maryae (Payne 1982); Serinus ankoberensis (Ash 1979, 1998, but see Van den Elzen 1985). Splits accepted in ‘Birds of Africa’ (Brown et al. 1982, Urban et al. 1986, 1997, Fry et al. 1988, 2000, Keith et al. 1992) are: Gymnobucco sladeni (split from G. peli),Trachyphonus usambiro (split from T. darnaudi); Campethera scriptoricauda (from C. bennettii) and C. mombasica (from C. abingoni); Dendropicos lugubris (from D. gabonensis) and D. spodocephalus (from D. griseocephalus); Rhinomomastus aterrimus (from R. cyanomelas), Tockus leucomelas (from T. flavirostris) and T. jacksoni (from T. deckeni); Ceratogymna albotibialis (from C. cylindri-


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cus); Centropus neumanni (from C. leucogaster) and C. burchelli (from C. superciliosus); Bubo ascalophus (from B. bubo) and B. vosseleri (from B. poensis); Caprimulgus ruwenzorii (from C. poliocephalus), C. nigriscapularis (from C. pectoralis) and C. clarus (from C. fossi); Balearica regulorum (from pavonina); Eupodotis gindiana and savilei (from E. ruficrista) and E. afroides (from E. afra); Accipiter erythropus (from A. minulus); Buteo archeri (from B. augur). Lanius marwitzi (split from L. collaris); Oriolus pervicali (from O. larvatus); Dicrurus modestus (from D. adsimilis), Tchagra anchietae (from T. minuta); Laniarius brauni and amboimensis (from L. luhderi); Prionops rufiventris (from P. caniceps); Tersiphone bedfordi (from T. rufiventer); Batis occulta (from B. poensis) and B. ituriensis (from B. minima); Platysteira jamesoni (from P. blisetti); Chaetops aurantius (from C. frenatus); Zoothera tanganjicae (from Z. piaggiae), Z. crossleyi (from Z. gurneyi) and Z. guttata (from Z. fischeri); Diaptrornis brunnea and fischeri (split from D. chocolatina); Parus cinerascens and thruppi (from P. afer); Hirundo rufigula (from H. preussi); Pycnonotus somaliensis, dodsoni and tricolor (from P. barbatus); Phyllastrephus cabanisi and placidus (from P. fischeri) and P. hypochloris (from P. baumanni); Criniger chloronotus (from C. barbatus); Cisticola nigriloris and discolor (from C. chubbi), Cisticola lepe (from C. erythrops), C. angusticaudus (from C. fulvicauda) (but C. restrictus was accepted by Hall and Moreau as ‘New species’) and C. angolensis and aberdare (from C. robustus); Prinia fluviatilis (from P. subflava); Schistolais (Prinia) leontica (from S. leucopogon); Apalis personata (from binotata); A. bamendae and goslingi (from A. sharpii) and A. kobobensis and chapini (from A. porphyrolaema); Bradypterus lopezi and mariae (from B. barratti); Turdoides hartlaubi (from T. leucopygius); Hyliota usambarae (from H. australis); Anthrepres axillaris (from A. fraseri), Nectarinia nanoensis, ludovicensis, stuhlmanni and prigoginei (from N. afra) and N. obscura (from N. olivacea); Mirafra sharpei (from M. hypermetra), M. archeri (from M. ruddi), Calandrella blanfordi and erlangeri (from C. cinerea), C. athensis (from C. somalica); Anthus crenatus (from A. lineiventris) and A. hoeschi (from A. cinnamomeus). Other, mainly more recent splits are: Struthio molybdophanes (split from S. camelus; Freitag and Robinson 1993); Stactolaema woodwardi (from S. olivacea, Clancey 1979) and S. sowerbyi (from S. whytii; Clancey 1995a); Agapornis fischeri, lilianae and nigrigenis (from A. personata; Dowsett and Dowsett-Lemaire 1993); Tauraco livingstonii (from T. schalowi; Dowsett-Lemaire and Dowsett 1988); Otus granti (from O. leucotis; König et al. 1999), Bubo cinerascens (from B. africanus; Holt et al. 1999) and B. mackinderi (from B. capensis; König et al. 1999), Glaucidium ngamiense, castaneum and scheffleri (from G. capense; Sibley and Monroe 1990); Melierax poliopterus (from M. canorus; Sibley and Monroe 1990); Accipiter tousenelii (from A. tachiro; Sibley and Monroe 1990); Cursorius somalensis (from C. cursor, Pearson and Ash 1996). Batis reichenowi, dimorpha and margaritae (from B. mixta/capensis); Turdus olivaceus to be split into olivaceus (basal branch), smithi, swynnertoni, roehli, helleri, abyssinicus and ludoviciae (Bowie 2002); Monticola pretoriae (from M. brevipes; Sibley and Monroe 1990); Muscicapa itombwensis (from M. lendu; Sibley and Monroe 1990); Alethe castanea (from A. diademata; Sibley and Monroe 1990, Beresford 2002); Sheppardia poensis (from S. bocagei; Wolters 1982, Beresford 2002); Stiphrornis gabonensis and xanthogaster (from S. erythrothorax; Beresford and Cracraft 1999); Onychognathus neumanni (from O. morio; Feare and Craig 1998); Psalidoprocne petiti, chalybea, mangbettorum, oleaginea, antinorii, orientalis and holomelas (from P. pristoptera; Prigogine 1985); Hirundo domicella (from H. daurica, Sibley and Monroe 1990); Parus carpi (from P. niger; Clancey 1995b), P. pallidiventris (from P. rufiventris, Harrap and Quinn 1996) and P. guineensis (from P. leucomelas; Sibley and Monroe 1990), Anthoscopus sylviella (from A. caroli; Sibley and Monroe 1990, Zimmerman et al. 1996); Andropadus tephrolaemus divided in A. tephrolaemus, nigriceps, neumanni, chlorigula and fusciceps, A. olivaceiceps split from milanjensis and A. kakamegae from A. masukuensis (Roy et al. 1998); Phyllastrephus poliocephalus (from flavostriatus; Stuart and Jensen 1986), Phyllastrephus alfredi (from P. flavostriatus; Sibley and Monroe 1990); Bleda notata (from B. eximia; Chappuis and Erard 1993), the eastern ugandae then split from notata and woosnami from B. syndactyla (Beresford 2002); Criniger ndussumensis (from C. olivaceus; Sibley and Monroe, Beresford 2002); Cisticola emini (from C. aberrans; Sibley and Monroe 1990) and C. bodessa (from C. chinianus; Sibley and Monroe 1990) and C. distinctus (from C. lais; Sibley and Monroe 1990); Prinia melanops (from P. bairdii, Sibley and Monroe 1990); Apalis viridiceps (from A. flavida, Sibley and Monroe 1990); Pseudalcippe atriceps (from P. abyssinica, Sibley and Monroe 1990); Sylvietta chapini (from S. leucophrys; Sibley and Monroe 1990); Nectarinia stuhlmanni and prigoginei re-defined and N. fuelleborni and usambaricus split from N. mediocris (Bowie et al. in press); N. tsavoensis (from bifasciata, Cheke and Mann 2001); Mirafra alopex and sharpei (from M. africanoides and africana; Wolters 1982), M. naevia (from M. sabota; Sibley and Monroe 1990); Certhilauda subcoronata, brevirostris, semitorquata, benguelensis and damarensis (from C. curvirostris, Ryan and Bloomer 1999) and C. barlowi (from C. albescens, Ryan et al. 1998); Anthus camaroonensi (from ‘A. novaeseelandiae’, Wolters 1982) and A. banneremanni (from A. similis; Sibley and Monroe 1990); Passer motitensis and ruficinctus (from P. iagoensis; Summers-Smith 1984, Dowsett and Dowsett-Lemaire 1993); Ploceus vitellinus (from P. velatus, Wolters 1982), P. katangae (from P. reichardi, Louette and Benson 1982), P. nicolli (from P. olivaceiceps; Wolters 1982), Euplectes psammocromius (from hartlaubi, Dowsett and Dowsett-Lemaire 1980), Pytilia lineata (from P. phoenicoptera, Sibley and Monroe 1990), Lagonosticta virata (from L. rhodopareia, Nikolai 1982), L. virata and landanae (from L. rubricata, Clement 1993) and L. vinacea (from L. larvata; Wolters 1982, Sibley and Monroe 1990); Estrilda nigriloris (from astrild; Wolters 1982, Clement 1993), E. kandti (from E. atricapilla, Prigogine 1975, Sibley and Monroe 1990) and E. charmosyne (from E. erythronotus; Sibley and Monroe 1990); Lonchura nigriceps (from L. bicolor; Sibley and Monroe 1990). Vidua is difficult to compare, V. nigeriae and maryae were in funerea, but Hall and Moreau (1970) may have referred some specimen records to V. raricola and larvaticola, and V. camerunensis is of very uncertain status (Klein et al. 1993, Payne and Payne 1994); Serinus xantholaema (from S. flavigula; Erard 1974), S. xanthopygius (from S. atrogularis, Erard 1974), S. frontalis and hypostictus (from S. citrinelloides, Van den Elzen 1985), S. canicapillus and reichardi (from S. gularis; Wolters 1982), S. buchanani (from S. donaldsoni; Wolters 1982, Sibley and Monroe 1990), S. leucolaema (from S. alario; Sibley and Monroe 1990), S. whytii (from S. striolatus, Wolters 1982) and S. melanochrous (from S. burtoni, Jensen and Brøgger-Jensen 1992). BirdLife International (2000) treats as separate species Apalis flavigularis, fuscogularis and lynesi (from A. thoracicus) and Zosterops kulalensis, winifredae and silvanus (from Z. poliogaster). The following suggested species were not taken into account, because of concrete published evidence (mainly in ‘Birds of Africa’ and Dowsett and Dowsett-Lemaire 1993) that they are not justified: Centropus epomidis, Apus toulsoni, Tersiphone tricolor, Lamprotornis elisabeth, Zoothera kibalensis, Bradornis pumilus, Andropadus hallae, Cisticola mongalla and taciturnus, Eremomela salvadorii and canescens, Anthus latistriatus, Estrilda quartina and ochrogaster, Lagonosticta umbrinodorsalis and rufopicta, and Vidua lorenzi and incognita. Many accepted species with highly disjunct ranges, such as some francolins (Peliperdix coqui/albogularis group), may require further splitting, but this needs to be properly evaluated using molecular markers. Species accepted in the ‘Speciation Atlases’ but later synonymised are Pogoniulus leucolaima (now in P. bilineatus) and Phoeniculus damarensis (now in P. purpureus; Cooper et al. 2001). Also Pogoniulus makawai (described from one specimen) has been neglected but new material has now become available for scrutiny (J Erritzøe).