Article

Speciation in African Forest Robins (Stiphrornis): Species Limits, Phylogenetic Relationships, and Molecular Biogeography

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Abstract

The monotypic genus Stiphrornis (Aves: Turdidae) is revised under a phylogenetic species concept to include four species, one of which, from the southwest Central African Republic, is new. Mitochondrial DNA sequence data are analyzed to explore the phylogenetic relation- ships within Stiphrornis. These data indicate relatively high levels of sequence divergence among the species and corroborate their recognition as diagnosable taxa, a conclusion also supported by morphological evidence. These findings, along with the allopatric distributions of the species, compel attention to their phylogenetic and spatial history, which was not ex- plored when this group was ascribed to a single ''biological'' species. Data reviewed here also suggest that the northwest Congo Basin forest, where the new species was discovered, is more zoogeographically complex than has been previously sus- pected. In addition, application of a phylogenetic species concept emphasizes the narrow en- demism of S. gabonensis and S. sanghensis, along with its implications for conserving their threatened habitats. The findings of this paper also reinforce the notion that patterns of geographic variation in the lowland forests of West and Central Africa are still incompletely understood and that the impact of environmental and geological history on the diversification of the forest avifauna has not yet been fully explored.

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... Ours is the first such analysis for two species, Yellow-whiskered Greenbul Eurillas latirostris and Western Bluebill Spermophaga haematina, whereas for three others that have been the subject of previous genetic analyses -Olive Sunbird Cyanomitra olivacea, Green Hylia Hylia prasina and Forest Robin Stiphrornis erythrothorax -we supplement currently available data with additional sampling on both sides of the Dahomey Gap. Clements et al. (2018) recognise each of the five taxa above as a single species-level taxon, though several recent studies on Forest Robin (Beresford and Cracraft 1999;Schmidt et al. 2008;Voelker et al. 2013;Boana et al. 2015;Voelker et al. 2017) show that it comprises a number of genetically divergent lineages, several of which may merit species status. Marks (2010) found similarly divergent lineages across the range of Hylia prasina. ...
... For all five focal species, we amplified and sequenced the entire NADH dehydrogenase subunit 2 gene (ND2), which has been widely used in avian phylogenetics. For Forest Robin samples only, most of the mitochondrial cytochrome b gene (cyt-b) together with a portion of the flanking tRNA gene (1 066 bp in total) were sequenced to allow combined analysis of new samples from this study with previously published cyt-b data for this species (Beresford and Cracraft 1999;Schmidt et al. 2008;Voelker et al. 2013Voelker et al. , 2017. Both ND2 and cyt-b were amplified and sequenced in two overlapping fragments using avian-specific primers (Supplementary Table S2). ...
... Roy 1997; Sefc et al. 2003;Sorenson et al. 2004;Johansson et al. 2007;Sangster et al. 2010;Zuccon and Ericson 2010;Moyle et al. 2011). For Stiphrornis only, we also downloaded available cyt-b sequences to allow comparison with earlier studies on this genus (Beresford and Cracraft 1999;Schmidt et al. 2008;Voelker et al. 2017). ...
Article
This study evaluates phylogeographic structure in selected forest bird species distributed on either side of the Dahomey Gap in West Africa to extend analyses of avian diversification across the Guineo-Congolean forest. Mitochondrial DNA sequence data were collected for five species, including four with a break in distribution across the Dahomey Gap and one with a more continuous distribution. Data for the five focal species were included in a single analysis along with appropriate outgroup taxa, allowing estimates of divergence time on a common timescale. Both phylogeographic patterns and estimated divergence times were highly variable. In two cases (Green Hylia Hylia prasina and Yellow-whiskered Greenbul Eurillas latirostris), samples from east and west of the Dahomey Gap belong to divergent mitochondrial lineages, but with substantially older divergence in Eurillas (∼4.1 MY). In Western Bluebill Spermophaga haematina, relatively divergent eastern and western lineages (∼1.1 MY) were sampled in the same locality in eastern Ghana, suggesting east to west gene flow across the gap following earlier vicariance. In Forest Robin Stiphrornis erythrothorax, birds sampled within Ghana were from two highly divergent lineages (∼1.9 MY), one of which is more closely related to birds from Cameroon and Equatorial Guinea, such that divergence across the Dahomey Gap was comparatively recent (∼0.36 MY). For both Hylia and Stiphrornis, we corroborate recent studies documenting multiple divergent lineages within each genus, some of which may represent distinct species. Consistent with previous studies, our analysis shows that the Dahomey Gap can be a significant barrier to gene flow for species restricted to forest habitats. At the same time, individual species exhibit a broad range of divergence times and phylogeographic histories of variable complexity. We discuss the taxonomic implications of our results for each focal species.
... This dearth of genetic analyses (as compared to the betterstudied avifauna of the Eastern Arc Mountains of Africa) has been exacerbated by a general lack of well-preserved DNA for genetic analysis. While work in Liberia, Sierra Leone and Ghana in the last few decades has produced a limited number of voucher specimens with associated preserved tissue samples, most avian collections from the Congo Basin are relatively old (1970s and earlier) and lack associated tissue samples (but see Beresford & Cracraft, 1999;Schmidt, Foster, Angehr, Durrant, & Fleischer, 2008). ...
... The general lack of collecting in Afrotropical lowland forests is likely inhibiting the potential discovery of new species, although several have been described in recent years. For example, there have been two Forest Robin (Stiphrornis) species discovered in the last 15 years (Beresford & Cracraft, 1999;Schmidt et al., 2008). While the five described Forest Robin species (erythrothorax, gabonensis, pyrrholaemus, sanghensis, xanthogaster) are phenotypically rather similar, even the recently described taxa should not be considered cryptic, as all taxa show clear differences in genetics, song and plumage colour. ...
... In our analyses, we included representatives from all five currently described Stiphrornis lineages (erythrothorax, gabonensis, pyrrholaemus, sanghensis, and xanthogaster); these lineages are varyingly recognized as races of erythrothorax (e.g., Collar, 2005;Sinclair & Ryan, 2010) or as species (e.g., Beresford & Cracraft, 1999;Boano, Vinals, Durante, & Pavia, 2015;Schmidt et al., 2008). Whole genomic DNA was extracted from tissue using the DNeasy tissue extraction kit (Qiagen), from a total of 70 Stiphrornis. ...
Article
We describe three new species of forest robin in the genus Stiphrornis; two from West Africa and one from the Congo Basin. Each species represents a distinct phylogenetic lineage based on genetic analysis. In addition to genetic differentiation, each new species is diagnosable from other Stiphrornis lineages by morphology, and by plumage. One of the new species appears to be restricted to the Central and Brong-Ahafo Regions of Ghana, and another is restricted to Benin and the Central Region of Ghana. In Ghana, these two new species presumably come into contact with Stiphrornis erythrothorax (Western Region of Ghana and westward), and there is evidence that one of the new species has a distinguishably different song from erythrothorax. The distribution of the third new species is primarily on the south bank of the Congo River, near the city of Kisangani. Recognition of these species provides additional evidence that Afrotropical forests are harbouring substantial cryptic diversity, and that our knowledge of the drivers of this diversity remains poorly documented across the region. http://www.zoobank.org/urn:lsid:zoobank.org:pub:BF2A0BE6-1140-4EFF-9035-380D61AB03AE
... More recently, Beresford & Cracraft (1999), on the basis of relatively high among-taxon mitochondrial DNA divergences in the genus, and supported by morphology, recognized four phylogenetic species within Stiphrornis: S. erythrothorax, from Sierra Leone to the Niger Delta, primarily in lowland forest but also in riverine and savannah habitats within the Upper Guinea region (Keith et al. 1992); S. gabonensis, limited to evergreen coastal forest, ranging from just east of the Niger River Delta south to Gabon and east to Malen, Cameroon, and the island of Bioko, Equatorial Guinea; S. xanthogaster, the most widespread member of the genus, ranging from the River Dja in Cameroon south through NE Gabon to Lukolela, Democratic Republic of Congo east through southern Central African Republic, Sudan, the Ituri forest, and Uganda to W Kenya, mainly in lowland primary forest, but in eastern Democratic Republic of Congo S. xanthogaster occurs in transitional forest up to 1400 m (Keith et al. 1992); and a new taxon, Sangha Forest-robin S. sanghensis Beresford & Cracraft, 1999, known only from the Dzanga-Sangha Dense Forest Reserve, Central African Republic, in primary forest, old second-growth forest, and moderately inundated forest (Beresford & Cracraft 1999). ...
... More recently, Beresford & Cracraft (1999), on the basis of relatively high among-taxon mitochondrial DNA divergences in the genus, and supported by morphology, recognized four phylogenetic species within Stiphrornis: S. erythrothorax, from Sierra Leone to the Niger Delta, primarily in lowland forest but also in riverine and savannah habitats within the Upper Guinea region (Keith et al. 1992); S. gabonensis, limited to evergreen coastal forest, ranging from just east of the Niger River Delta south to Gabon and east to Malen, Cameroon, and the island of Bioko, Equatorial Guinea; S. xanthogaster, the most widespread member of the genus, ranging from the River Dja in Cameroon south through NE Gabon to Lukolela, Democratic Republic of Congo east through southern Central African Republic, Sudan, the Ituri forest, and Uganda to W Kenya, mainly in lowland primary forest, but in eastern Democratic Republic of Congo S. xanthogaster occurs in transitional forest up to 1400 m (Keith et al. 1992); and a new taxon, Sangha Forest-robin S. sanghensis Beresford & Cracraft, 1999, known only from the Dzanga-Sangha Dense Forest Reserve, Central African Republic, in primary forest, old second-growth forest, and moderately inundated forest (Beresford & Cracraft 1999). ...
... More recently, Beresford & Cracraft (1999), on the basis of relatively high among-taxon mitochondrial DNA divergences in the genus, and supported by morphology, recognized four phylogenetic species within Stiphrornis: S. erythrothorax, from Sierra Leone to the Niger Delta, primarily in lowland forest but also in riverine and savannah habitats within the Upper Guinea region (Keith et al. 1992); S. gabonensis, limited to evergreen coastal forest, ranging from just east of the Niger River Delta south to Gabon and east to Malen, Cameroon, and the island of Bioko, Equatorial Guinea; S. xanthogaster, the most widespread member of the genus, ranging from the River Dja in Cameroon south through NE Gabon to Lukolela, Democratic Republic of Congo east through southern Central African Republic, Sudan, the Ituri forest, and Uganda to W Kenya, mainly in lowland primary forest, but in eastern Democratic Republic of Congo S. xanthogaster occurs in transitional forest up to 1400 m (Keith et al. 1992); and a new taxon, Sangha Forest-robin S. sanghensis Beresford & Cracraft, 1999, known only from the Dzanga-Sangha Dense Forest Reserve, Central African Republic, in primary forest, old second-growth forest, and moderately inundated forest (Beresford & Cracraft 1999). ...
Article
We report the occurrence of the recently-described Stiphrornis pyrrholaemus in the Makokou area, northeastern Gabon,more than 600 kilometers NE from its type locality, in areas covered by dense primary to secondary forest. The previous records of Stiphrornis from the same area were referred to S. xanthogaster. We confirm this attribution on the basis of museum specimens. Although several recent sources treat all Stiphrornis as a single species, our findings strongly suggest the sympatric coexistence of two Stiphrornis species and thus that they should be treated as separate species under the Biological Species Concept.
... As part of a larger study of the avian family Muscicapidae, we assembled samples from the African forest robin assemblage (FRA). This group has had a taxonomically turbulent history (see, e.g., Irwin & Clancey, 1974; Beresford, 2003), and has increased in species number due to several recent discoveries (e.g. Beresford & Cracraft, 1999; Beresford et al., 2004; Schmidt et al., 2008). Included in our sampling of the FRA were most species ascribed to the genera Cossypha (12/15), Sheppardia (9/9), Pseudalethe (4/4; see Beresford, 2003), Erithacus (2/3), Pogonocichla (1/1), Stiphrornis (1/1), Swynnertonia (1/1) and Xenocopsychus (1/1) by Sibley & Monroe (1990). ...
... Included in our sampling of the FRA were most species ascribed to the genera Cossypha (12/15), Sheppardia (9/9), Pseudalethe (4/4; see Beresford, 2003), Erithacus (2/3), Pogonocichla (1/1), Stiphrornis (1/1), Swynnertonia (1/1) and Xenocopsychus (1/1) by Sibley & Monroe (1990). We also included Sheppardia aurantiithorax (Beresford et al., 2004), Sheppardia usambarae (often separated from Sheppardia sharpei), and Stiphrornis sanghensis (Beresford & Cracraft, 1999). Both Beresford & Cracraft (1999) and Schmidt et al. (2008) have addressed systematic relationships within Stiphrornis (now four species) and we do not reassess them here. ...
... We also included Sheppardia aurantiithorax (Beresford et al., 2004), Sheppardia usambarae (often separated from Sheppardia sharpei), and Stiphrornis sanghensis (Beresford & Cracraft, 1999). Both Beresford & Cracraft (1999) and Schmidt et al. (2008) have addressed systematic relationships within Stiphrornis (now four species) and we do not reassess them here. ...
Article
Aim Montane tropics are areas of high endemism, and mechanisms driving this endemism have been receiving increasing attention at a global scale. A general trend is that climatic factors do not explain the species richness of species with small to medium‐sized geographic ranges, suggesting that geological and evolutionary processes must be considered. On the African continent, several hypotheses including both refugial and geographic uplift models have been advanced to explain avian speciation and diversity in the lowland forest and montane regions of central and eastern Africa; montane regions in particular are recognized as hotspots of vertebrate endemism. Here, we examine the possible role of these models in driving speciation in a clade of African forest robins. Location Africa. Methods We constructed the first robustly supported molecular phylogenetic hypothesis of forest robins. On this phylogeny, we reconstructed habitat‐based distributions and geographic distributions relative to the Albertine Rift. We also estimated the timing of lineage divergences via a molecular clock. Results Robust estimates of phylogenetic relationships and clock‐based divergences reject Miocene tectonic uplift and Pleistocene forest refugia as primary drivers of speciation in forest robins. Instead, our data suggest that most forest robin speciation took place in the Late Pliocene, from 3.2 to 2.2 Ma. Distributional patterns are complex, with the Albertine Rift region serving as a general east–west break across the group. Montane distributions are inferred to have evolved four times. Main conclusions Phylogenetic divergence dates coincide with a single period of lowland forest retraction in the late Pliocene, suggesting that most montane speciation resulted from the rapid isolation of populations in montane areas, rather than montane areas themselves being drivers of speciation. This conclusion provides additional evidence that Pliocene climate change was a major driver of speciation in broadly distributed African animal lineages. We further show that lowland forest robins are no older than their montane relatives, suggesting that lowland areas are not museums which house ‘ancient’ taxa; rather, for forest robins, montane areas should be viewed as living museums of a late Pliocene diversification event. A forest refugial pattern is operating in Africa, but it is not constrained to the Pleistocene.
... The arid DG has been considered historically important in driving biodiversity patterns (Booth, 1958;Endler, 1982;Mayr & O'Hara, 1986) (Figure 1a). Genetic diversification patterns of a small number of bird and mammal taxa support this contention, where species do not share haplotypes across the gap (Beresford & Cracraft, 1999;Fuchs & Bowie, 2015;Fuchs et al., 2017;Hassanin et al., 2014;Marks, 2010;Nesi et al., 2013;Schmidt, Foster, Angehr, Durrant, & Fleischer, 2008;Voelker et al. 2017). ...
... For example, the sole genus of Afro-tropical birds (Stiphrornis) that has seen multiple, rigorous investigations (i.e., genetics, morphology, behaviour) was originally described as one widespread species lacking substantial plumage variation and included in Mayr and O'Hara's (1986) list of biogeographically uninformative taxa. Subsequently, the genus has been shown to contain no fewer than eight species (Beresford & Cracraft, 1999;Schmidt et al., 2008;Voelker et al. 2017). In addition, recent studies of the genera Bleda (Huntley & Voelker, 2016) and Criniger revealed higher than expected levels of diversification across the GCF than expected, especially given their inclusion on Mayr and O'Hara's (1986) list. ...
Article
Aim Vertebrate diversity in the Guineo‐Congolian forests (GCF) of Africa is high, yet mechanisms responsible for generating that diversity remain remarkably understudied. These forests have alternatively been viewed as centres of diversification (“cradles”) or more recently, as the opposite (“museums”). Here, we use a comparative dataset of avian and mammalian species to examine genetic diversification patterns across these forests and use these results to explain observed patterns in light of Plio‐Pleistocene climatic change and life‐history. Location Africa. Methods We analysed patterns of diversification across the GCF using a dataset composed of 629 and 1,048 mitochondrial sequences from 60 avian and 36 mammalian species, respectively. Uncorrected pairwise genetic distances were compared at three distinct geographical levels: west versus east of the Dahomey Gap, among the three major forest blocks, and among seven historical refugial areas. The timing of diversification was assessed for passerines, Rodentia, and Chiroptera. Results We found substantial signatures of diversification in all three levels of our geographical comparisons. We recovered substantial disparity in the amount and depth of structure of diversification patterns between low dispersers (understorey birds and rodents) and more capable dispersers (other bird species and bats). Additionally, our chronogram recovered recent speciation and intraspecific diversification across songbird and mammalian lineages. Main conclusions The discrete, and often deep, structuring of genetic diversification for both birds and mammals across the GCF revealed strong correlations between historic landscape fragmentation and dispersal ability. Our results revealed a striking amount of unrecognized genetic diversity, which may be suggestive of cryptic species. Given the rate at which these forests are being negatively impacted by human intervention, our general lack of knowledge concerning vertebrate diversity across these forests may very well impact our ability to identify evolutionary processes underlying diversity and enact meaningful conservation efforts in the future.
... Beyond these first two divergences, the speciation patterns and timing recovered within the genus can be explained through a combination of isolation in lowland forest refugia and the breakup of the Guineo-Congolian forests into western and eastern blocks, separated by the Dahomey Gap. Indeed, several investigations have recovered similar evidence for east-west genetic diversification of avian and mammalian populations distributed across the Dahomey Gap (birds: Beresford & Cracraft, 1999;Schmidt et al., 2008;Marks, 2010;Fuchs & Bowie, 2015;Huntley & Voelker, 2016;mammals: Gonder et al., 2011;Nesi et al., 2013;reptiles: Leaché et al., 2014). The patterns and timing of speciation we recovered in Criniger also support the involvement of the Dahomey Gap, in conjunction with more regional refugial scenarios, as a driver of two speciation events. ...
... prasina: Marks, 2010; Illadopsis rufipennis: Nguembock et al., 2009;N. olivacea: Bowie et al., 2004;Platysteira cyanea: Njabo et al., 2008;Sylvietta denti: Huntley & Voelker, 2017; Stiphrornis erythrothorax: Beresford & Cracraft, 1999, Schmidt et al., 2008Voelker et al., 2016b). The results from these ten 'uninformative' taxa in conjunction with investigations of several taxa with restricted ranges indicate the substantial complexity of biogeographic patterns within Guineo-Congolian lowland forests. ...
Article
Full-text available
The biogeographical history of Afro-tropical lowland forests during the Plio-Pleistocene is characterized by pervasive fragmentation-coalescence cycling due to global climatic oscillations. Vicariance scenarios driven by forest fragmentation have long been hypothesized as major mechanisms for the creation and maintenance of Afro-tropical avian diversity. However, the timing and centre of diversification events remains unclear. We undertook the first molecular phylogenetic and biogeographic investigation of the avian genus Criniger, a group of understory birds endemic to the lowland forests of West and Central Africa. Utilizing DNA from 43 specimens and a combination of molecular and biogeographic methods, we constructed time-calibrated phylogenies and ancestral area estimations. We estimated a widespread origin for the genus, with a basal divergence dating to the late Miocene. All other speciation events were dated to the Pliocene. However, we recovered substantial geographic structuring of genetic diversity, dating to the Pleistocene, within both Western and Central Africa for three species. The biogeographic patterns observed in the genus Criniger are likely the result of allopatric diversification driven by forest fragmentation during the Plio-Pleistocene. The results of this study indicate that Afro-tropical forests harbour substantially higher levels of cryptic diversity and greater genetic complexity than previously hypothesized.
... Throughout its fluctuating geological and climatic history, the region could have constituted, depending on the ecological tolerances of the plant or animal group concerned, either a corridor permitting, or a barrier preventing, east-west exchange of biotas (e.g. Beresford & Cracraft 1999). ...
... Throughout geological features and climatic history, the region could have constituted, depending on the ecological tolerances of the plant or animal group concerned, either a corridor or a barrier to east-west exchange of biotas (e.g. Beresford & Cracraft 1999). For P. misonnei, high genetic diversity was observed in the region of the Bakossi highlands (Mt Kupe, Mt Manenguba, Mt Bakossi), since specimens belonging to clades I1, I2 and I3 were recorded in this region. ...
Article
We studied the phylogeographical structure of Praomys misonnei (Rodentia: Muridae) in west-Central Africa based on 113 individuals from 31 localities and the sequencing of two mitochondrial genes (Cytb and CO1). Two clades can be identified through maximum likelihood, bayesian and network analyses: the first one is limited to the Korup National Park, while the second one has a broad geographical distribution in west-Central Africa. The second clade can be further divided into three sub-clades. The results of neutrality tests and mismatch distributions indicate that a population expansion occurred in these sub-clades. During the middle or upper Pleistocene, the west-Central African populations of P. misonnei may have been isolated in four different forest refugia, followed by population expansion and secondary contact during more humid periods. High genetic diversity was observed in the region of the Bakossi highlands, and the Cameroon Volcanic Line limits the geographical distribution of clades. Thus it seems likely that this volcanic line has in the past constituted a barrier that prevented the east–west exchange between P. misonnei populations through the expansion of the montane forest habitat. Our results also highlight the role of the Sanaga and Ogooué-Ivindo rivers in shaping P. misonnei genetic structure.
... For example, the widespread, monotypic Green Hylia (Hylia prasina) was shown to possess highly discrete geographic structure and deep genetic divergences between major lowland forest blocks (Marks, 2010). The systematics and phylogeography of the forest robin genus Stiphrornis has been analyzed no less than three times, with each investigation revealing deep-rooted cryptic diversity and new species (Beresford and Cracraft, 1999; Schmidt et al., 2008). In addition, the timing of diversification leading to geographic structuring within several recently studied lowland taxa has proven to be relatively recent (within the last 1.8 my), a result that seems to suggest the possibility of the Pleistocene Forest Refuge Hypothesis as a potential explanation for at least some diversification patterns (for birds: Voelker et al., 2013; Fuchs and Bowie, 2015; for mammals: Quérouil et al., 2003; Nicolas et al., 2008). ...
... In addition, and similar to the Congo River, the Ogooue River in Gabon could be acting as a barrier to gene flow between these subclades. Some species patterns have demonstrated the possibility of this river as a possible barrier to secondary contact after refugial expansion in the avian genus Stiphrornis (Beresford and Cracraft, 1999; Schmidt et al., 2008) and in several mammalian taxa (Quérouil et al., 2003; Tefler et al., 2003; Anthony et al., 2007; Nicolas et al., 2011). Greater sampling is needed in Gabon and Equatorial Guinea at finer scale to determine with certainty if this is the case in B. syndactylus. ...
Article
Recent investigations of distributional patterns of Afro- tropical lowland forest species have demonstrated to some degree our overall lack of understanding involving historical diversification patterns. Traditionally, researchers have relied upon two hypotheses, each of which views the lowland forest of Africa in differing roles. The Pleistocene Forest Refuge Hypothesis (PFRH) posits that biogeographic patterns of avian lowland species are explained via allopatric speciation during forest fragmentation cycles in the Pleistocene epoch (c. 1.8 Ma - 11, 700 Ka). The Montane Speciation Hypothesis (MSH) countered by suggesting that lowland forests are "evolutionary museums" where species, which originally evolved in montane forest refuge centers, remained without further diversification. Furthermore, investigations have largely regarded widespread, avian species which lack phenotypic variability as biogeographically "uninformative", with regards to historical biogeographic patterns. To test the tenets of these ideas, we investigated the systematics and biogeography of the genus Bleda, whose constituent species are restricted to lowland forest and are lacking in phenotypic variation. Using extracted DNA from 179 individuals, we amplified two mitochondrial genes and three nuclear loci and utilized Bayesian phylogenetic methods and molecular clock dating to develop a time-calibrated phylogeny of Bleda. We used LaGrange to develop an ancestral area reconstruction for the genus. Haplotype networks for three species were generated using Network. We recovered the four currently recognized species of Bleda, plus a monophyletic B. ugandae, a current sub-species which may warrant full species status. We found that the origins of the genus Bleda are estimated to be in the Upper Guinean forests of West Africa, dating to the Miocene (c. 7.5 Ma), while the speciation events for the rest of the genus are dated to the Pliocene (c. 5 Ma - 1.8 Ma). Our analyses recovered discrete and highly differentiated geographic structuring of genetic diversity in West and Central Africa in three of five species, with many of the diversification events dating to the Pleistocene. The biogeographic patterns observed in Bleda can be explained through a combination of isolation via forest refuges during the Plio-Pleistocene and riverine barriers limiting secondary contact after forest expansion. We find evidence for the PFRH as a driver of intra-specific diversity, but conclude that it does not facilitate an explanation for speciation in the genus Bleda. The "evolutionary museum" concept furnished by the MSH is countered by our evidence of in situ diversification in the lowland forests of Africa. Additionally, our results provide strong evidence of the value of seemingly "uninformative" widespread avian taxa for revealing complex patterns of forest diversity. Overall, our study highlights that past researchers have both underestimated the amount of diversity found in lowland forests and failed to understand the complexity of historical forces shaping that diversity. Gaining a better understanding of lowland forest diversity and the historical factors which have shaped it will crucial in determining conservation tactics in the near future.
... Nesi et al. 2013, Leaché et al. 2014. For example, no haplotypes were shared between morphologically similar populations sampled across the Dahomey gap for two songbird species, Stiphrornis erythrothorax (Beresford and Cracraft 1999;Schmidt et al. 2008) and Hylia prasina (Marks 2010). The biogeographic pattern is likely even more complex as the populations from the Upper Guinea Forest block were paraphyletic in Hylia (Marks 2010), suggesting that further phylogeographic structure is apparent within the Upper Guinea Forest block, a result in conflict with Mayr and O'Hara's (1986) hypothesis of only one refugium west of the Dahomey Gap. ...
... Previous studies on African lowland rainforest birds suggest that haplotypes are rarely shared between populations sampled across the Dahomey gap (Beresford and Cracraft 1999;Schmidt et al. 2008, Marks 2010) and our analyses are consistent with this result. The phylogeographic relationships of the populations inhabiting the forests west of the Lower Niger River and east of the Dahomey Gap (Western Nigeria, Benin, hereafter referred to as wLNR) ...
Article
Full-text available
The lowland forests of western and central tropical Africa are separated by several potential biogeographic barriers to dispersal for forest adapted vertebrates. The two primary barriers are 1) the Dahomey Gap, a savanna corridor that reaches the coast of southern Ghana, Togo and Benin, and separates the West African rainforest into the Upper (Ghana west to Guinea) and Lower Guinea (Nigeria to Uganda and Angola) forest blocks, and 2) the Lower Niger River, a large delta that separates Western and Eastern Nigeria. Previous studies on terrestrial vertebrates (lizards, mammals and birds) have highlighted a genetic break in the Dahomey Gap/Lower Niger River area although the relative importance of each barrier has not been assessed due to limitations in geographic sampling or properties of the markers used. We compared the phylogeographic history of two co-distributed sister-species of woodpeckers (Campethera caroli and C. nivosa) using data from three loci representing all inheritance modes. Our analyses revealed that both the Dahomey Gap and possibly the Lower Niger River acted as strong biogeographic barriers for the two woodpecker species, with the Lower Niger River being the first barrier to have formed, leading to three distinct populations of C. nivosa. Our divergence time analyses revealed that both these biogeographic barriers formed during the Pleistocene, supporting the Pleistocene refuge hypothesis, with the Dahomey Gap likely appearing about 0.5 myr BP. No genetic structure was recovered among sampled populations in either the Upper or the Lower Guinea Forest Block for both species, despite the considerable geographic area covered. Copyright © 2015 Elsevier Inc. All rights reserved.
... Throughout its fluctuating geological and climatic history, the region could have constituted, depending on the ecological tolerances of the plant or animal group concerned, either a corridor permitting, or a barrier preventing, east-west exchange of biotas (e.g. Beresford & Cracraft 1999). ...
... Throughout geological features and climatic history, the region could have constituted, depending on the ecological tolerances of the plant or animal group concerned, either a corridor or a barrier to east-west exchange of biotas (e.g. Beresford & Cracraft 1999). For P. misonnei, high genetic diversity was observed in the region of the Bakossi highlands (Mt Kupe, Mt Manenguba, Mt Bakossi), since specimens belonging to clades I1, I2 and I3 were recorded in this region. ...
Article
Full-text available
We studied the phylogeographical structure of Praomys misonnei (Rodentia: Muridae) in west-Central Africa based on 113 individuals from 31 localities and the sequencing of two mitochondrial genes (Cytb and CO1). Two clades can be identified through maximum likelihood, bayesian and network analyses: the first one is limited to the Korup National Park, while the second one has a broad geographical distribution in west-Central Africa. The second clade can be further divided into three sub-clades. The results of neutrality tests and mismatch distributions indicate that a population expansion occurred in these sub-clades. During the middle or upper Pleistocene, the west-Central African populations of P. misonnei may have been isolated in four different forest refugia, followed by population expansion and secondary contact during more humid periods. High genetic diversity was observed in the region of the Bakossi highlands, and the Cameroon Volcanic Line limits the geographical distribution of clades. Thus it seems likely that this volcanic line has in the past constituted a barrier that prevented the east—west exchange between P. misonnei populations through the expansion of the montane forest habitat. Our results also highlight the role of the Sanaga and Ogooué-Ivindo rivers in shaping P. misonnei genetic structure.
... In contrast, Beresford and Cracraft (1999) examined a lowland radiation of forest robins (Stiphrornis) and found geographic structure consistent with lowland Pleistocene refugia. Theirs was the only study to sample an African forest bird extensively across its range. ...
... Even with limited sampling, the tinkerbird data agree with the finding of Beresford and Cracraft (1999) and indicate that there is diversity within species that must be accounted for in speciation models. Populations of P. subsulphureus are 3.5% divergent between Ghana and Central African Republic (Figure 4.4), localities hypothesized to be in or near two of the Pleistocene refugia (Diamond andHamilton 1980, Crowe andCrowe 1982). ...
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ACKNOWLEDGMENTS Many people and institutions helped make thisdissertation possible. I thank my advisor, Fred Sheldon, for all of his help and support over the last six years. His advisory style combines helpful guidance with a hands-off approach to students that suited me particularly well. My graduate committee, of Van Remsen, Mark Hafner, Jim McGuire, Mike Hellberg, and Al Afton, was always available for advice or helpful discussions. At LSU I benefited from a wonderful group of fellow grad students. They helped me in a variety of ways from laboratory advice to reading manuscripts toinviting me over for dinner. In no particular order, they include: Ben Marks, Jason Weckstein, Chris Witt, Alex Aleixo, Alison Styring, Tom Devitt, Adam Leache, Jessica Light, Mark McRae, Dan Christian, Rob Faucett, Kwai Hin Han, Josie Babin, Kalan Ickes, Mario Cohn-Haft, David Reed, and Kazuya Naoki. In particular, Jason Weckstein took the time to show me around the lab and get me started on sequencing. Also, I owe Steve Cardiff and Donna Dittmann thanks for teaching me how to prepare bird specimens. I especially want to acknowledge,all of the work that Nannette Crochet does in the lab. She juggles requests from several labs and always seems to accommodate everyone. Tissue samples used in this dissertation were obtained from the following institutions: Field Museum of Natural History (John Bates,Dave Willard, and Raurie Bowie), American
... In addition to the six newly generated sequences, these included 61 sequences reported by Dietzen et al. (2003), one from Sangster et al. (2010), and 24 previously unreported cyt b sequences of E. rubecula from Germany (N = 4), Spain (N = 3), Morocco (N = 8), Madeira (N = 4), La Palma (N = 1) and the Azores (N = 4) which were obtained from GenBank (AY624077-AY624100). An additional member of Cossyphinae (Stiphrornis sanghensis, AF136731; Beresford & Cracraft, 1999), a member of Niltavinae (Niltava sundara, EF081354; Lei et al., 2007) and Turdus philomelos (AY495411; van der Meij et al., 2005) were used as outgroups. ...
Article
The European Robin Erithacus rubecula is currently treated as a single species with eight subspecies. A previous molecular study and new molecular, morphometric and bioacoustic data reported here strongly support the recognition of three species in this complex: E. rubecula (Europe, North Africa and Macaronesia except the central Canary Islands), E. superbus (Tenerife) and a recently described subspecies on Gran Canaria which we raise to species rank as E. marionae. The taxa on Tenerife and Gran Canaria have previously been lumped as a single taxon but differ from each other and from E. rubecula in territorial songs, tic calls, seep calls and wing length. All three species are characterised by moderate to high levels of interspecific mitochondrial DNA sequence divergence (mean 4.2–4.8%). Phylogenetic analysis indicates that E. marionae is sister to E. superbus + E. rubecula. Recognition of Gran Canaria and Tenerife Robins as separate species adds two single-island endemics to the Canary Islands avifauna.
... B.P., have also been found in West African populations of chimpanzees (3,25,26), woodpeckers (27), and frogs (28). Studies on other African lowland rainforest birds (29)(30)(31), forest-dwelling rodents (32), and African bushbucks suggest that haplotypes are rarely shared between populations sampled across the Dahomey Gap, although the relationships between clades are not well resolved. Finally, genome-wide data in lizards, snakes, and frogs revealed different levels of genetic divergence across taxa (33). ...
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Significance Although today the rainforest is continuous in Central Africa, the scarce fossil record suggests that arid conditions during the ice ages might have reduced tree density. However, the vast majority of the fossil pollen cores preserved in Tropical Africa are too young to inform about this period. Investigating whether the climate change left a genetic signature on trees can thus be useful to date past forest fragmentation. We use DNA technology to study five legume trees. Our results show significant differentiation of the populations of each species at a date compatible with forest fragmentation driven by ice age climatic oscillations. Contrasted timescales were obtained for each species, in agreement with different recolonization abilities in an expanding forest biome after fragmentation.
... B.P., have also been found in West African populations of chimpanzees (3,25,26), woodpeckers (27), and frogs (28). Studies on other African lowland rainforest birds (29)(30)(31), forest-dwelling rodents (32), and African bushbucks suggest that haplotypes are rarely shared between populations sampled across the Dahomey Gap, although the relationships between clades are not well resolved. Finally, genome-wide data in lizards, snakes, and frogs revealed different levels of genetic divergence across taxa (33). ...
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The rainforests of Tropical Africa have fluctuated over time. Although today the forest cover is continuous in Central Africa this may have not always been the case, as the scarce fossil record in this region suggests that more arid conditions might have significantly reduced the density of trees during the Ice Ages. Our aim was to investigate whether the dry ice-age periods left a genetic signature on tree species that can be used to date the past fragmentation of the rainforest. We sequenced reduced representation libraries of 182 samples representing five Legume tree species that are widespread in African rainforests and seven outgroups. Phylogenetic analyses identified an early divergent lineage for all species in West Africa (Upper Guinea), and two clades in Central Africa: Lower Guinea-North and Lower Guinea-South. As the structure separating the Northern and Southern clades cannot be explained by geographic barriers, we tested other hypotheses using demographic model testing. The best estimates recovered using ∂a∂I indicate that the two clades split between the Upper Pliocene and the Pleistocene, a date compatible with forest fragmentation driven by ice-age climatic oscillations. Furthermore, we found remarkably older split dates for the shade-tolerant tree species with non-assisted seed dispersal than for light-demanding long-distance wind-dispersed trees. We also show that the genetic diversity significantly declines with the distance from ice-age refugia in the two long-distance dispersed species only. Different recolonisation abilities after recurrent cycles of forest fragmentation seem to explain why we observe congruent genetic spatial structures across species with contrasted timescales. SIGNIFICANCE STATEMENT Although today the rainforest cover is continuous in Central Africa, the scarce fossil record suggests that arid conditions during the Ice Ages might have reduced the density of trees during the Ice Ages. However, the vast majority of the fossil pollen records preserved in Tropical Africa is too young to inform about this period. Investigating whether the past climate change left a genetic signature on trees can thus be useful to date past forest fragmentation. However, most genetic studies available to date lack resolution as they use limited numbers of loci. In this study we use modern DNA technology to study five Legume trees. Our results show significant differentiation of the populations of each species at a date compatible with forest fragmentation driven by ice-age climatic oscillations. Contrasted timescales were obtained for each species, which probably reflects their different recolonisation abilities after forest fragmentation.
... The recognition of new taxa in sub-Saharan Africa is uncommon relative to other areas around the globe (e.g. South America, Indonesia; Fjeldså 2013) and most new African taxa have come from the Lower Guinean Forest Block (Beresford et al. 1999;Schmidt et al. 2008) or the mountains of eastern and central Africa (Jensen 1983;Dinesen et al. 1994;Fjeldså & Kiure 2003;Beresford et al. 2004;Bowie & Fjeldså 2005;Fjeldså et al. 2006;Bowie et al. 2009Bowie et al. , 2016Voelker et al. 2010Voelker et al. , 2017. All these cases underscore the need for continued collecting of voucher specimens in sub-Saharan Africa (Bates et al. 2004). ...
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We describe a new species of drongo in the Square-tailed Drongo (Dicrurus ludwigii) complex using a combination of biometric and genetic data. The new species differs from previously described taxa in the Square-tailed Drongo complex by possessing a significantly heavier bill and via substantial genetic divergence (6.7%) from its sister-species D. sharpei. The new species is distributed across the gallery forests of coastal Guinea, extending to the Niger and Benue Rivers of Nigeria. We suspect that this taxon was overlooked by previous avian systematists because they either lacked comparative material from western Africa or because the key diagnostic morphological character (bill characteristics) was not measured. We provide an updated taxonomy of the Square-tailed Drongo species complex.
... Spatial analyses provide support for the influence of major geological features, rather than species ecology, as the primary mechanism genotypes, but did exhibit south to north and west to east clines detected by sPCA ( Figure 1). Osteolaemus is highly terrestrial and can be considered a true forest species; as such, the detected spatial structure likely reflects spatial expansion from Pleistocene forest refugia (Beresford & Cracraft, 1999;Leal, 2001;Maley, 1987), as seen in other forest species in this region (Anthony et al., 2007;Clifford et al., 2004;Marks, 2010). (Elenga, Schwartz, & Vincens, 1994;Jahns, 1996;Wirrman, Bertaux, & Kossoni, 2001). ...
... Most exciting, of course, is the fact that the continent undoubtedly holds further species as yet wholly un-known to science. Witness to that are the 26 new African species that have been described in the last two decades (van Rootselaar 1999); the subsequent description of Stiphrornis sanghensis (Beresford and Cracraft 1999) adds another species to this total. Astoundingly, these include two new genera: Xenoperdix (Dinesen et al. 1994) and Cryptosylvicola (Goodman et al. 1996). ...
... For example, European Robin Erithacus rubecula use a complex, melodious song (made up of prolonged notes, short warbles and trills) for territory defence, almost throughout the year (Hoelzel 1986(Hoelzel , 1989Scriba & Goymann 2010). African Forest Robins (Stiphrornis) have been reported to use two types of vocalisations (Beresford & Cracraft 1999). However, functions of these categories remain unclear. ...
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The physical characteristics and biological significance of song in the endemic Indian Robin Saxicoloides fulicata are described. Songs are discrete and composed of strophes (structural units) with frequency ranging from 1.03 to 8.00 kHz, preceded and followed by temporal intervals from 0.21 to 21.25 sec. Occasional, monosyllabic whistles are also identified. In a song bout usually the same type of strophe is repeated several times in a stereotyped manner with minor structural variations of elements before switching to another type of strophe. Most strophes are composed of two to five elements, having both simple and complex structure. Two categories of songs have been identified on the basis of their acoustical features and context of production. Type-A songs are simple, stereotyped, spontaneous and common, while type-B songs are rare, female-oriented and more complex than type-A. Song is used in both inter- and intra-sexual contexts. It seems that type-A songs are driven by male-male competition for territory and mates. Males also shorten the length of strophes and reduce gaps between strophes (in type-B songs) on the arrival of females in the vicinity, most probably to increase the song rate, suggesting it to be an indicator of male quality.
... We inferred a monophyletic Muscicapidae with strong support (Fig. 1 ). Evolutionary relationships within these subfamilies were largely congruent with recently published molecular phylogenies of various groups within Muscicapidae (Beresford & Cracraft 1999, Roy 2001, Dietzen et al. 2003, Cibois & Cracraft 2004, Voelker & Spellman 2004, Ertan 2006, Qiao-Wa et al. 2006, Seki 2006, Outlaw et al. 2007, 2010, Xin et al. 2007, Illera et al. 2008, Schmidt et al. 2008, Treplin et al. 2008, Sheldon et al. 2009, Zink et al. 2009, Sangster et al. 2010, Voelker 2010, Zuccon & Ericson 2010, Aliabadian et al. 2012, Oliveros et al. 2012). A comprehensive analysis of the topology inferred here is outside the scope of the current study; however, we provide a brief discussion of confirmed relationships and novel insights that we have generated from our increased taxonomic and molecular sampling (Supporting Information). ...
Article
The ecology of cavity nesting in passerine birds has been studied extensively, yet there are no phylogenetic comparative studies that quantify differences in life history traits between cavity and open-nesting birds within a passerine family. We test existing hypotheses regarding the evolutionary significance of cavity nesting in the Old World flycatchers (Muscicapidae). We used a multi-locus phylogeny of 252 species to reconstruct the evolutionary history of cavity nesting and to quantify correlations between nest types and life history traits. Within a phylogenetic generalized linear model framework, we found that cavity-nesting species are larger than open nesting species and that maximum clutch sizes are larger in cavity-nesting lineages. In addition to differences in life history traits between nest types, species that breed at higher latitudes have larger average and maximum clutch sizes and begin to breed later in the year. Gains and losses of migratory behaviour have occurred far more often in cavity-nesting lineages than in open-nesting taxa, suggesting that cavity nesting may have played a crucial role in the evolution of migratory behaviour. These findings identify important macro-evolutionary links between the evolution of cavity nesting, clutch size, interspecific competition and migratory behaviour in a large clade of Old World songbirds.This article is protected by copyright. All rights reserved.
... Similarly, supporters of the Phylogenetic Species Concept have identified species using not only diagnosability but also exclusive coalescence (Zink et al., 2002;Pavlova et al., 2005) or both (e.g. Beresford & Cracraft, 1999). Hybridzone dynamics indicating reproductive isolation have also been used in support of a proposal based on diagnosability (Sangster et al., 1999). ...
Article
The debate over species concepts has produced a huge body of literature on how species can, may or should be delimited. By contrast, very few studies have documented how species taxa are delimited in practice. The aims of the present study were to (i) quantify the use of species criteria in taxonomy, (ii) discuss its implications for the debate over species concepts and (iii) assess recent claims about the impact of different species concepts on taxonomic stability and the ‘nature’ of species. The application of six species criteria was examined in taxonomic studies of birds published between 1950 and 2009. Three types of taxonomic studies were included: descriptions of new species (N=329), proposals to change the taxonomic rank of species and subspecies (N=808) and the taxonomic recommendations of the American Ornithologists’ Union Committee on Classification and Nomenclature (N=176). In all three datasets, diagnosability was the most frequently applied criterion, followed by reproductive isolation and degree of difference. This result is inconsistent with the popular notion that the Biological Species Concept is the dominant species concept in avian taxonomy. Since the 1950s, avian species-level taxonomy has become increasingly pluralistic and eclectic. This suggests that taxonomists consider different criteria as complementary rather than as rival approaches to species delimitation. Application of diagnosability more frequently led to the elevation of subspecies to species rank than application of reproductive isolation, although the difference was small. Hypotheses based on diagnosability and reproductive isolation were equally likely to be accepted in a mainstream checklist. These findings contradict recent claims that application of the Phylogenetic Species Concept causes instability and that broader application of the Biological Species Concept can stabilise taxonomy. The criteria diagnosability and monophyly, which are commonly associated with Phylogenetic Species Concepts, were used throughout the study period. Finally, no support was found for the idea that Phylogenetic Species Concepts have caused a change in the ‘nature’ of species taxa. This study demonstrates that there is a discrepancy between widely held perceptions of how species are delimited and the way species are actually delimited by taxonomists. Theoretically oriented debates over species concepts thus may benefit from empirical data on taxonomic practice.
... Recent molecular phylogenetic studies have addressed relationships among African forest robins (Beresford, 2003; Voelker et al., 2009), chats and thrushes (Pan et al., 2006a ,b) and Asian flycatchers (Lei et al., 2007a,b). A number of molecular phylogenetic studies, mostly based on mitochondrial DNA sequences, involved limited taxon sampling to investigate particular genera, including Stiphrornis (Beresford and Cracraft, 1999; Schmidt et al., 2008), Sheppardia (Roy et al., 2001), Phoenicurus (Ertan, 2002Ertan, , 2006), Pseudocossyphus (Goodman and Weigt, 2002), Erithacus (Dietzen et al., 2003; Seki, 2006), Ficedula (Outlaw and Voelker, 2006), Oenanthe (Aliabadian et al., 2007), Monticola (Outlaw et al., 2007; Zuccon and Ericson, 2010a), Saxicola (Illera et al., 2008; Woog et al., 2008; Zink et al., 2009), Copsychus (Sheldon et al., 2009), and Oenanthe and Cercomela (Outlaw et al., 2010). The objectives of this study are to assess the phylogenetic relationships of representatives of 34 genera of Muscicapidae using DNA sequences from one mitochondrial and three unlinked nuclear loci, and to identify monophyletic groups for use in taxonomic revision. ...
... Most exciting, of course, is the fact that the continent undoubtedly holds further species as yet wholly un-known to science. Witness to that are the 26 new African species that have been described in the last two decades (van Rootselaar 1999); the subsequent description of Stiphrornis sanghensis (Beresford and Cracraft 1999) adds another species to this total. Astoundingly, these include two new genera: Xenoperdix (Dinesen et al. 1994) andCryptosylvicola (Goodman et al. 1996). ...
... Birds of New Guinea have a strong tendency to be locally distributed or geographically variable (Diamond, 1985;Beehler et al. , 1986;Cracraft, 1992). Few data exist for the Congo Basin, but distinct populations have been documented here as well (Beresford & Cracraft, 1999;Roy et al. , 2001). ...
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Attempts to determine global priorities for conservation and studies aimed at highlighting the conservation value of one continental region over another often devalue specific regions. Amazonia has been a particular recipient of this treatment in a number of recent studies. We suggest that this type of approach is not necessary at this scale, and we argue that the consequences could be devastating for the largest forests of the world. Among our concerns about the treatment of these forests (Amazonia, the Congo Basin and New Guinea) is that there seems to be a lack of appreciation for, or sufficient study of, important biogeographic subdivisions within these regions. In Amazonia, the south-eastern portion of the basin (the Belém/Pará region) has not been considered a global conservation priority, despite the fact that it is experiencing by far the highest deforestation rates. Parsimony Analyses of Endemicity and genetic data suggest that many Amazonian forest taxa are comprised of numerous regionally distinct units, and this may also be true in other large tropical forests. Such patterns need to be documented for adequate conservation of tropical biodiversity, but this might not happen if these regions are not recognized as priorities for conservation at a global scale.
... While Dickinson (2003) followed their decision, Rasmussen & Anderton (2005) and Collar (2005) retained both genera among chats. Although no comprehensive cladistic analysis for the family exists, several molecular studies addressed the relationships within selected genera: Erithacus (Seki 2006), Ficedula (Outlaw & Voelker 2006 ), Monticola and Pseudocossyphus (Goodman & Weigt 2002; Outlaw et al. 2007; Zuccon & Ericson in press), Oenanthe and Cercomela (Aliabadian et al. 2007; in press), Saxicola (Illera et al. 2008), Sheppardia (Roy et al. 2001; Beresford et al. 2004), Stiphrornis (Beresford & Cracraft 1999; Schmidt et al. 2008). A taxonomically more inclusive analysis (Beresford 2003) investigated the relationships among the African forest robins, a group of forest-dwelling chats dominated by the genera Cossypha, Sheppardia and Alethe first proposed by Irwin & Clancey (1974) . ...
Article
Zuccon, D. & Ericson, P. G. P. (2010). A multi-gene phylogeny disentangles the chat-flycatcher complex (Aves: Muscicapidae).—Zoologica Scripta, 39, 213–224. We reconstructed the first well-sampled phylogenetic hypothesis in the chat-flycatcher complex combining nuclear and mitochondrial sequences. The dichotomy between chats-terrestrial feeders and flycatchers-aerial feeders does not reflect monophyletic groups. The flycatching behaviour and morphological adaptations to aerial feeding (short tarsi, broad bill, rictal bristles) evolved independently from chat ancestors in three different lineages. The genera Alethe, Brachypteryx, and Myiophonus are nested within the Muscicapidae radiation and their morphological and behavioural similarities with the true thrushes Turdidae are presumably the result of convergence. The postulated close relationships among Erithacus, Luscinia and Tarsiger cannot be confirmed. Erithacus is part of the African forest robin assemblage (Cichladusa, Cossypha, Pogonocichla, Pseudalethe, Sheppardia, Stiphrornis), while Luscinia and Tarsiger belong to a large, mainly Asian radiation. Enicurus belongs to the same Asian clade and it does not deserve the recognition as a distinct subfamily or tribe. We found good support also for an assemblage of chats adapted to arid habitats (Monticola, Oenanthe, Thamnolaea, Myrmecocichla, Pentholaea, Cercomela, Saxicola, Campicoloides, Pinarochroa) and a redstart clade (Phoenicurus, Chaimarrornis and Rhyacornis). Five genera (Muscicapa, Copsychus, Thamnolaea, Luscinia and Ficedula) are polyphyletic and in need of taxonomic revision.
... Recent molecular phylogenetic studies have addressed relationships among African forest robins (Beresford, 2003;Voelker et al., 2009), chats and thrushes (Pan et al., 2006a,b) and Asian flycatchers (Lei et al., 2007a,b). A number of molecular phylogenetic studies, mostly based on mitochondrial DNA sequences, involved limited taxon sampling to investigate particular genera, including Stiphrornis (Beresford and Cracraft, 1999;Schmidt et al., 2008), Sheppardia (Roy et al., 2001), Phoenicurus (Ertan, 2002(Ertan, , 2006, Pseudocossyphus (Goodman and Weigt, 2002), Erithacus (Dietzen et al., 2003;Seki, 2006), Ficedula (Outlaw and Voelker, 2006), Oenanthe (Aliabadian et al., 2007), Monticola (Outlaw et al., 2007;Zuccon and Ericson, 2010a), Saxicola (Illera et al., 2008;Woog et al., 2008;Zink et al., 2009), Copsychus (Sheldon et al., 2009), and Oenanthe and Cercomela (Outlaw et al., 2010). ...
Article
The chats and flycatchers (Muscicapidae) represent an assemblage of 275 species in 48 genera. Defining natural groups within this assemblage has been challenging because of its high diversity and a paucity of phylogenetically informative morphological characters. We assessed the phylogenetic relationships of 124 species and 34 genera of Muscicapidae, and 20 species of Turdidae, using molecular sequence data from one mitochondrial gene and three nuclear loci, in total 3240bp. Bayesian and maximum likelihood analyses yielded a well-resolved tree in which nearly all basal nodes were strongly supported. The traditionally defined Muscicapidae, Muscicapinae and Saxicolinae were paraphyletic. Four major clades are recognized in Muscicapidae: Muscicapinae, Niltavinae (new family-group name), Erithacinae and Saxicolinae. Interesting relationships recovered by this analysis include: (i) a clade comprising the 'blue' flycatcher genera Niltava, Cyornis, Cyanoptila and Eumyias and some species of Rhinomyias; (ii) the position of Erithacus rubecula in a clade of otherwise exclusively African species; (iii) a close relationship between the shortwing Heinrichia calligyna and the flycatcher Rhinomyias insignis; (iv) a sister-relationship between forktails Enicurus and whistling thrushes Myophonus; and (v) a sister relationship of Ficedula and the 'chats'Monticola, Phoenicurus, Saxicola and Oenanthe. A high number of traditionally defined genera was found to be paraphyletic or polyphyletic.
... conserved speciation events) in tropical rainforest predate the Pleistocene, suggesting that the most recent climatic oscillations played a minor role in their evolution (Fjeldså & Lovett 1997;Moritz et al. 2000;Nicolas et al. 2006). Radiations of mountain and savanna birds took place in the Plio-Pleistocene (Fjeldså 1994;Fjeldså & Lovett 1997;Roy 1997;Roy et al. 1998), but many groups in putative Guineo-Congolian refuges diverged in the Mio-Pliocene, well before Pleistocene vicissitudes (Beresford 2003;Beresford & Cracraft 1999;Bowie et al. 2004a;2004b;Fjeldså 1994;Roy et al. 2001). Eight species of Hylomyscus woodmice diversified rapidly 2-6 Ma, mostly in the early Pliocene (Nicolas et al. 2006). ...
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Proefschrift Universiteit Leiden. Met samenvatting in het Nederlands.
... However, only a few studies have attempted to apply this approach to tropical African faunas. These studies focused on mountain and lowland forest birds (Beresford and Cracraft, 1999; Roy, 1997) and savanna-dwelling bovids (Arctander et al., 1999). We investigated the potential biogeographic scenarios proposed for the Central African lowland forest fauna by testing the congruence between phylogeographic patterns obtained for two sympatric species of forest shrews and the faunal regions previously recognised for mammals (Colyn, 1999; Deleporte and Colyn, 1999;Fig. ...
Article
We analysed the phylogeographic patterns of two congeneric and syntopic species of forest shrews and compared them with biogeographical scenarios proposed for the Central African tropical forest. Our results, based on 82 partial 16s rRNA mitochondrial sequences, suggest that both species must have originated in the Plio-Pleistocene and that their haplotype distributions could reflect the effect of forest fragmentation and expansion associated with paleoclimatic fluctuations during the Pleistocene. However, it seems that the two species responded very differently to environmental changes. While Sylvisorex johnstoni populations exhibit ancient haplotype segregation that may even represent currently unrecognised allopatric species, Sylvisorex ollula haplotypes are much less differentiated and suggest that this taxon has undergone a recent range expansion. The observed differences between these taxa may be explained by their presumably different ecological requirements and colonisation abilities, which in turn may be the result of a significant difference in body size between the two species. In conclusion, our results suggest that it is necessary to incorporate several ecologically well-documented species in studies that attempt to infer evolutionary processes from phylogeographic patterns
... Currently, at least 25 species concepts have been advanced (Coyne and Orr, 2004). Each one has certain limitations, and adhering to one particular concept may aVect the assessment of biological diversity (Avise and Walker, 1999; Beresford and Cracraft, 1999). On the other hand, gathering evidence from diVerent sources allows approaching entities that are acceptable as species regardless of which species concept is adopted (Avise and Walker, 1999 Walker, , 2000). ...
Article
Diversity of ants of the Tetramorium caespitum/impurum complex was investigated in a multidisciplinary study. Focusing on morphologically hardly distinguishable Western Palearctic samples, we demonstrate the genetic and phenotypic diversity, demarcate phylogenetic entities, and discuss the clades in terms of biogeography. Sequences of 1113bp of the mitochondrial COI gene revealed 13 lineages. COII data, worker morphometry and male genitalia morphology corroborated the COI results for seven lineages; the remaining six were disregarded because of small sample size. A comparison with published data on cuticular hydrocarbons showed correspondence. The seven entities show different distribution patterns, though some ranges overlap in Central Europe. Since no major discrepancy between the results of the different disciplines became apparent, we conclude that the seven entities within the T. caespitum/impurum complex represent seven species. Geographical evidence allows the identification of T. caespitum and T. impurum, and we therefore designate neotypes and redescribe the two species in terms of morphology and mtDNA. As the revision of about 50 taxon names would go beyond the scope of this study, we refer to the remaining five species under code names. We discuss our findings in terms of plesiomorphy and convergent evolution by visualizing the mtDNA phylogeny in morphological space.
... Another possibility is that the difference in the level of genetic divergence within the two species is demographically driven; for example by differences in population size. On a continental scale, the level of genetic differentiation between populations of these African forest taxa is similar to that found in some other species of lowland African rainforest birds (Beresford and Cracraft, 1999;Beresford, 2003; but see Roy, 1997;Bowie et al., 2004). The lone Asian taxon for which we have multiple individuals is Merops viridis, an inhabitant of open country from southeast China through the Indonesian and Philippine archipelagos to Java. ...
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The bee-eaters (family Meropidae) comprise a group of brightly colored, but morphologically homogeneous, birds with a wide variety of life history characteristics. A phylogeny of bee-eaters was reconstructed using nuclear and mitochondrial DNA sequence data from 23 of the 25 named bee-eater species. Analysis of the combined data set provided a well-supported phylogenetic hypothesis for the family. Nyctiornis is the sister taxon to all other bee-eaters. Within the genus Merops, we recovered two well-supported clades that can be broadly separated into two groups along geographic and ecological lines, one clade with mostly African resident species and the other clade containing a mixture of African and Asian taxa that are mostly migratory species. The clade containing resident African species can be further split into two groups along ecological lines by habitat preference into lowland forest specialists and montane forest and forest edge species. Intraspecific sampling in several of the taxa revealed moderate to high (3.7-6.5%, ND2) levels of divergence in the resident taxa, whereas the lone migratory taxon showed negligible levels of intraspecific divergence. This robust molecular phylogeny provides the phylogenetic framework for future comparative tests of hypotheses about the evolution of plumage patterns, sociality, migration, and delayed breeding strategies.
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The group of small forest robins, or akalats, that dwell in lowland and montane forests in Africa have complicated parapatric or partly overlapping distributions, the details of which are still being discovered. Here, we use external morphology and mitochondrial and nuclear DNA sequence data to determine the reciprocal monophyly of several populations, including one form that is related to Sheppardia lowei and S. montana. Those data corroborate the recognition of a new species, Sheppardia aurantiithorax, and show relatively high levels of sequence divergence among populations of the different species. The discovery of this new species, narrowly endemic in the Eastern Arc montane forests, emphasizes the complex biodiversity of the region and underscores the need for prompt and effective conservation measures.
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This report is the forth one of a series; Part 1 presents the results of a comprehensive literature screening in search for new bird taxa described in 2008, namely new genera, species and subspecies worldwide. We tracked four new genera, eight new species, 17 subspecies new to science which according to the International Code of Zoological Nomenclature were correctly described. New genera were erected for species or species groups, respectively, of the Rhinocryptidae, Cisticolidae and Parulidae. Six of the new species refer to Passeriformes and two to Non-Passeres, a storm-petrel and a parrot. The distributional areas of the new species often are minute, restricted to remote areas and were hitherto overlooked. In several cases the populations in question were known since long but their remarkable acoustical and genetic properties which led to description of new species were unexplored. In a zoogeographic context most of the new taxa originate from the Neotropics, followed by Palaerctic and Indomalayan Realms. The remainder of taxa are scattered over Australasia, the Afrotropics, and the southern Atlantic islands. In a taxon sequence by genunus/species/subspecies there is the following distribution: Neotropis and Caribbean (1/2/8), Palaerctic (-/2/4), Indo-Malaya (-/2/2), Nearctic (2/-/1), Afrotropics (1/1/-), Australasia (-/1/1), and southern Atlantic islands (-/-/1). Replacement names were proposed for two neotropical genera and four subspecies. – In Part 2 (this issue) a number of splits - namely those of known species into allospecies, which in most cases result in eographic representatives of a superspecies - are also addressed. But we restrict the treatment of these splits to the Palearctic and Indomalayan Realms. We suggest possible flaws in new descriptions and certain splits, regardless of the species concept addressed. However, in general this report should be taken as a documentation of new taxa, not as a critical review of recent changes in bird taxonomy and bird descriptions.
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The three missions of Systematics Agenda 2000 (SA2000) - to inventory Earth's species, to understand their relationships, and to use the latter to create predictive information systems-define an agenda of research for systematic biology. The recognition that systematic knowledge underpins biological knowledge in general, and applied biology in particular, has resulted in an amazing growth in systematics over the past decade. Increasingly, systematics is being used to solve, societal problems. This paper describes seven great questions within systematics and discusses their relevance for, and contribution to, conserving and sustainably using biodiversity. These questions fall into four broad categories: Questions about diversity: What is a species? and How many species are there? Questions about phylogeny: What is the Tree of Life? and What has been the history of character transformation? Questions about biogeography: Where are Earth's species distributed? and How have species' distributions changed over time? and Questions about phyloinformatics: How is phylogenetic history predictive?
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Our knowledge of the avian tree of life remains uncertain, particularly at deeper levels due to the rapid diversification early in their evolutionary history. They are the most abundant land vertebrate on the planet and have been of great historical interest to systematists. Birds are also economically and ecologically important and as a result are intensively studied, yet despite their importance and interest to humans around 13% of taxa currently on the endangered species list perhaps as a result of human activity. Despite all this no comprehensive phylogeny that includes both extinct and extant species currently exists. Here we present a species-level supertree, constructed using the Matrix Representation with Parsimony method, of Aves containing approximately two thirds of all species from nearly 1000 source phylogenies with a broad taxonomic coverage. The source data for the tree were collected and processed according to a strict protocol to ensure robust and accurate data handling. The resulting tree topology is largely consistent with molecular hypotheses of avian phylogeny. We identify areas that are in broad agreement with current views on avian systematics and also those that require further work. We also highlight the need for leaf-based support measures to enable the identification of rogue taxa in supertrees. This is a first attempt at a supertree of both extinct and extant birds, it is not intended to be utilised in an overhaul of avian systematics or as a basis for taxonomic re-classification but provides a strong basis on which to base further studies on macroevolution, conservation, biodiversity, comparative biology and character evolution, in particular the inclusion of fossils will allow the study of bird evolution and diversification throughout deep time.
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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.
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Phylogenetic relationships between species of akalats (Sheppardia) and alethes (Alethe) were derived from mitochondrial and nuclear sequence data, and phylogeographic pattern was also investigated in three widespread species (two Alethe and one Sheppardia) endemic to the African lowland rainforest biome. Co-distributed species of Alethe and Sheppardia showed similar spatial relationships between the Guineo-Congolian biome and the Western Rift mountains, and preliminary area relationships are shown for species of Sheppardia and Alethe distributed in the Cameroon highlands, the woodland areas north and south of the lowland forest block, and east African montane and lowland forest. Within widespread species, phylogeographic pattern was generally similar, with long branches between Upper and Lower Guinea populations, and between eastern and western Congolian populations. At the interspecific level, the nuclear intron used gave more resolution than the mitochondrial data, which gave poor resolution at basal nodes. While the affinities of some species groups were weakly supported, no support was suggested for the monophyly of Alethe and little for Sheppardia as those genera are currently circumscribed, and taxonomic changes are suggested. As found by previous workers, monophyletic clades corresponding to Muscicapini and Saxicolini of Sibley and Ahlquist (1990) were not recovered, and the membership of any monophyletic 'African forest robin assemblage' needs to be resolved.
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Los zorzales pequeños ("akalats") que se encuentran en los bosques montanos y de tierras bajas de África tienen complejas distribuciones que son parapátricas o se superponen parcialmente, y cuyos detalles están aún siendo descubiertos. En este estudio utilizamos la morfología externa y secuencias de ADN mitocondrial y nuclear para determinar la monofilia recíproca de varias poblaciones, incluyendo una forma que está relacionada con Sheppardia lowei y S. montana. Estos datos corroboran el reconocimiento de una nueva especie, Sheppardia aurantiithorax, y muestran niveles relativamente altos de divergencia en secuencias entre poblaciones de las diferentes especies. El descubrimiento de esta nueva especie, que es estrechamente endémica a los bosques montanos del arco oriental, enfatiza la compleja biodiversidad de la región y recalca la necesidad de implementar medidas efectivas de conservación con prontitud.
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This sixth part of “Type Specimens of Birds in the American Museum of Natural History” corresponds to taxa covered in Volume 10 of Peters' Check-list of Birds of the World. The original description of each taxon has been consulted unless otherwise noted, coordinates given for type localities when possible, currently accepted names for the taxa included, and comments on taxonomic history provided. Three hundred sixty-four published names are treated. Types of six of these are not in AMNH, and one was discussed in an earlier list.
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We describe Laniarius willardi, a new species of boubou shrike (Malaconotidae) from the Albertine Rift of Africa. The most conspicuous, distinguishing morphological feature of the species is a gray to blue-gray iris. This and external morphometric data indicate that L. willardi is diagnosable from other black or sooty boubous. Further, L. willardi is genetically diagnosable, and its closest relative is the Mountain Sooty Boubou (L. poensis camerunensis) from Cameroon. The Crimson-breasted Bush-shrike (L. atrococcineus) and the Lowland Sooty Boubou (L. leucorhynchus) are together the sister clade to L. willardi–L. p. camerunensis. Laniarius willardi and the geographically codistributed L. p. holomelas differ by .% in uncorrected sequence divergence, and elevational data taken from museum specimens suggest the possibility of elevational segregation of the species at ~, m, with L. willardi occurring at lower elevations. Our broad sampling of black and sooty boubou taxa indicate that () races of Mountain Sooty Boubou (L. poensis) do not form a monophyletic clade; () L. p. camerunensis may represent multiple, nonsister lineages; and () at least one race of Fülleborn’s Black Boubou (L. fuelleborni usambaricus) is genetically distinct from other races of that species. Received  June , accepted  December .
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It has been suggested that switching from the widely used Biological Species Concept to a Phylogenetic Species Concept, would result in the appearance of hitherto neglected patterns of endemism. The problem has mainly been analyzed with respect to endemic taxa and for rather limited geographical regions, but will here be analysed for the entire resident avifauna of sub-Saharan Africa. A database of African bird distributions was re-edited to create two new datasets representing 1572 biological species and 2098 phylogenetic species. Species richness patterns were virtually identical with the two taxonomies, and only subtle changes were found in the geographical variation in range-size rarity sum. However, there were some differences in the most range-restricted species, with increased complexity of long-recognized centres of endemism. Overall, then, the large-scale biogeographic patterns are robust to changes in species concepts. This reflects the aggregated nature of endemism, with certain areas acting as “species pumps” and large intervening areas being characterised by a predominance of widespread species which distribute themselves in accordance with contemporary environmental conditions. The percentages of phylogenetic and threatened species captured in a BSC near-minimum set of 64 grid-cells and a PSC near-maximum set, with the same number of grid-cells, are very similar.
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We describe a new species of forest robin from the Gamba Complex in southwest Gabon. This common bird, Stiphrornis pyrrholaemus sp. nov., inhabits primary lowland forest and forages on or near the ground like the other members of the genus Stiphrornis of central and western Africa. Unique phenotypic features of the new species include the male's bright orange chin, throat, and breast, creamy yellow belly, olive green back and rump, and gray flanks. Mitochondrial sequence divergence corroborates our assessment based on its distinct physical characteristics that this is a new species, and suggest that Stiphrornis erythrothorax is likely the most closely related congener.
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Oscine passerine birds make up almost half of all avian diversity. Relationships within the group, and its classification, have long been controversial. Over the last 10 years numerous molecular phylogenies have been published. We compiled source phylogenies from 99 published studies to construct an oscine supertree. We aimed to illustrate weak and strong parts of the phylogeny and set targets for future phylogenetic work and therefore preferred a heuristic approach where we judged the adequacy of taxon sampling and molecular method of each source tree instead of using matrices and automated tree-building programs. We present an estimate of the phylogenetic relationships of 1723 extant and one extinct species of oscine passerine birds (Aves: Passeri) — more than 37% of the total. We included 34/35 (97%) families, 38/39 (97%) subfamilies and 40/43 (93%) tribes. Overall resolution is 83% of a fully bifurcating tree. The basal lineages are all distributed in the Australo-Papuan region, but several more distal lineages dispersed out of this region and radiated in other parts of the world. However, taxa of the Australian region suffer from larger evolutionary gaps and the deep branches of the Sylvioidea and nine South American primaried oscines are still poorly resolved.
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A recent trend in the literature highlights the special role that tropical montane regions and habitat transitions peripheral to large blocks of lowland rainforest play in the diversification process. The emerging view is one of lowland rainforests as evolutionary 'museums'; where biotic diversity is maintained over evolutionary time, and additional diversity is accrued from peripheral areas, but where there has been little recent diversification. This leads to the prediction of genetic diversity without geographic structure in widespread taxa. Here, I assess the notion of the lowland rainforest 'museum' with a phylogeographic study of the green hylia (Aves: Sylviidae: Hylia prasina) using 1132 bp of mtDNA sequence data. The distribution of genetic diversity within the mainland subspecies of Hylia reveals five highly divergent haplotype groups distributed in accordance with broad-scale areas of endemism in the Afrotropics. This pattern of genetic diversity within a currently described subspecies refutes the characterization of lowland forests as evolutionary museums. If the pattern of geographic variation in Hylia occurs broadly in widespread rainforest species, conservation policy makers may need to rethink their priorities for conservation in the Afrotropics.
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The African forest robins (akalats) of the genus Sheppardia are represented by eight forest-dwelling species with restricted distributions in tropical Africa. All but three are strictly montane, inhabiting isolated highland areas in eastern and western Africa. Due to their subtle plumage variation and almost wholly allopatric distribution, determining systematic relationships based upon morphology has proven difficult. However, akalats, due to their distribution, offer an ideal opportunity to test models of speciation within tropical forests. We therefore investigated the phylogeny of species of this genus from presently sampled regions of their distribution using mtDNA sequence analysis. We found that the monophyly of described species, even from disjunct populations, is well supported. However, relationships among species is generally poorly resolved, with support given only to the paired relationships S. montana/S. lowei, S. bocagei/S. gunningi, and S. aequatorialis/S. cyornithopsis. This dataset lent support to S. montana and S. lowei representing a superspecies. All species appear to have evolved rapidly from a common ancestor around the Miocene/Pliocene transition, a time of the last uplift of East Africa's montane region. Surprisingly, intraspecific radiations suggest a far more recent population expansion in the upper Pleistocene, concordant with major climatic variation and vegetational changes. We discuss the implications of our results in the light of previous discussions of the montane speciation model.
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Species are defined using a variety of different operational techniques. While discussion of the various methodologies has previously been restricted mostly to taxonomists, the demarcation of species is also crucial for conservation biology. Unfortunately, different methods of diagnosing species can arrive at different entities. Most prominently, it is widely thought that use of a phylogenetic species concept may lead to recognition of a far greater number of much less inclusive units. As a result, studies of the same group of organisms can produce not only different species identities but also different species range and number of individuals. To assess the impact of different definitions on conservation issues, we collected instances from the literature where a group of organisms was categorized both under phylogenetic and nonphylogenetic concepts. Our results show a marked difference, with surveys based on a phylogenetic species concept showing more species (48%) and an associated decrease in population size and range. We discuss the serious consequences of this trend for conservation, including an apparent change in the number of endangered species, potential political fallout, and the difficulty of deciding what should be conserved.
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This study constitutes the first investigation of the phylogeographic structure of a forest bird distributed throughout the montane and lowland forest biomes of Africa. The key objective was to investigate the importance of Pleistocene climatic cycles on avian diversification across Africa. The Olive Sunbird is a relatively large polytypic sunbird widely distributed throughout evergreen, montane and coastal forests in Africa. Recently, it was split into two species, the Eastern Olive Sunbird (Nectarinia olivacea) and the Western Olive Sunbird (Nectarinia obscura), based on morphological grounds. Analyses of a 395bp fragment of the mtDNA NADH subunit 3 gene with flanking tRNA sequences, from 196 individuals of N. olivacea and 86 from N. obscura indicate that genetic divergence levels are low (1.0-2.4%) across some 9000km, from Ghana in the northwest of Africa to KwaZulu-Natal in eastern South Africa. Neither currently recognized Olive Sunbird species were monophyletic using either parsimony or likelihood tree-building methods. Phi(ST) values suggested that there was less variation partitioned among species than between most neighboring regions. Genetic diversity within the N. olivacea/obscura complex was dominated by three star-like phylogenies linked to each other by a single mutational step and two subnetworks (IV and V) separated from the core star-like phylogenies (subnetworks I, II, and III) by five to six mutational steps. The dominant evolutionary mechanism shaping genetic variation within the N. olivacea/obscura complex as identified by nested-clade analyses, appears to be one of range expansion possibly out of East Africa associated with a period of forest expansion during the mid-Pleistocene, some 1.1-0.7 million years ago. Mismatch profiles suggested that secondary contact has occurred between eastern and western lineages within the Ufipa Escarpment and possibly Zimbabwe, as well as between eastern lineages in the Kenyan Highlands and northern Eastern Arc Mts.
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This paper gives a historical overview of the African rain forest from its origins, towards, the end of the Cretaceous period. The areas around the Gulf of Guinea, in particular from Ivory Coast to Nigeria and especially Cameroon, Gabon and Congo, appear to have been already occupied at this time by wet tropical forest formations mainly composed of Angiosperms. In the course of the Tertiary period the combined effect of the equator being situated further north than now and the development of the Antarctic ice cap favoured the development of wet tropical conditions over a large part of North Africa. Towards the end of the Tertiary, the equator reached its present position and the northern hemisphere ice caps appeared, and these phenomena resulted in the disappearance of the forest formations spread across the north of Africa, and the concentration of these formations near the equatorial zone around the Gulf of Guinea and in the Congo-Zaire basin. From 800 000 years ago onwards the marked glacial variations at middle and high latitudes in both hemispheres, lowered temperatures in equatorial areas and brought arid climates at times of maximum glacial extension. -Author
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Small carnicore scats collected over a two-year period in the southwestern Central African republic revealed an unusually high occurrence of shrews (23.3%). Analysis of skull fragments yielded 311 individuals of 16 species. Shrew occurrence varied significantly with season (highest in the dry season), but not with proximity to water. The unusually high diversity of the shrew community in comparison to others in west-central Africa is attributed to a thorough and efficient sampling effort, and to a wide variety of available microhabitats maintained through disturbance factors such as a high density of elephants and selective logging activities. In addition to the scat remains, four shrew species were collected live: one is new to science and is described here as Sylvisorex konganensis. The species belongs to a group in which a basal portion of the tail is covered by long bristles. It is most similar to a yet unnamed species of Sylvisorex from Cameroon, with which it apparently occurs in sympatry. Both share critical characters with S. howelli and S. usambarensis, species endemic to the Uluguru and Usambara Mts. respectively.
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Late Pleistocene glaciations have been ascribed a dominant role in sculpting present-day diversity and distributions of North American vertebrates. Molecular comparisons of recently diverged sister species now permit a test of this assertion. The Late Pleistocene Origins model predicts a mitochondrial DNA divergence value of less than 0.5 percent for avian sister species of Late Pleistocene origin. Instead, the average mitochondrial DNA sequence divergence for 35 such songbird species pairs is 5.1 percent, which exceeds the predicted value by a factor of 10. Molecular data suggest a relatively protracted history of speciation events among North American songbirds over the past 5 million years.
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The prevailing explanation for the observed distributional patterns and areas of endemism of tropical forest organisms is the Pleistocene refuge hypothesis, which proposes that wide-ranging ancestral taxa were isolated into forest refuges during certain glacial periods, and that this isolation provided them with the opportunity to speciate. John Endler has recently argued that two predictions of the refuge hypothesis-that contact zones between vicars should be between refuges and that contact zones of rapidly reproducing butterflies should be wider than those of more slowly reproducing birds-are not borne out by the evidence. Endler therefore rejects the refuge hypothesis. We show that the data available are far too imprecise to permit any conclusions regarding contact zone widths and that, according to our reanalysis of the African bird data used by Endler, all the contact zones between vicars do indeed occur between refuges, exactly where they are expected. Additional strong support for the refuge hypothesis comes from the existence of many taxa endemic to the particular forest areas which have been postulated as refuges and from fragmented taxa which are still allopatric, never having come into secondary contact.
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Patterns of distribution, diversity and endemism for 1595 species and well-marked subspecies of Afrotropical (excluding marine islands) passerine and non-aquatic non-passerine birds are analysed. Patterns of distribution are well predicted by a hierarchical model of African avifaunal zones based on patterns of distribution and evolution in guineafowl (Family Numididae). The boundaries of avifaunal zones recognized for passerines and non-passerines are similar to one another, and largely parallel those of broad contemporary vegetation types along a rainfall gradient. However, there are interesting discrepancies between the distributions of contemporary vegetation types and inferred avifaunal zones, most notably the disjunct similarity of the avifaunas of three sections of lowland forest, much of which presently is an apparently continuous belt. This disjunct avifaunal similarity is interpreted in the light of palaeoecological and geomorphological data as evidence in favour of the hypothesis of forest refugia: that presently continuous forests were fragmented during arid conditions in the geological past, and that birds which prefer forested biotopes were confined to 'island' forest refugia. Patterns of diversity and endemism, when considered together with patterns of distribution, also support the refugium hypothesis, and suggest the existence of 2 additional refugia, one in the western Africa forest, the other in the eastern African equatorial savanna.-Authors
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The lake Barombi Mbo pollen record goes back to about 28,000 yr B.P. The pollen diagram based on 82 samples is subdivided into four main pollen zones. Zone I (ca. 28,000 to 20,000 yr B.P.) is characterized by relatively high frequencies of Caesalpiniaceae and also by a montane element with Olea capensis. The climate was cool and relatively wet. Zone II (ca. 20,000 to 10,000 yr B.P.). A sharp increase in Gramineae, the main non-arboreal land pollen taxon, began around 20,000 yr B.P. and lasted until 10,000 yr B.P. During this period the forest receded, giving way to a more open vegetation, but significant patches of forest (refuges) persisted in the area. This is confirmed by isotopic analyses (δ13C of sedimentary detritic organic matter from the catchment. Until ca. 13,000 yr B.P. Olea capensis was well represented indicating a relatively cool climate. Between 13,000 and 12,000 yr B.P. a warming trend associated with a strong increase in precipitation occurred. After this an abrupt reduction in precipitation linked to an increase in seasonality, but without temperature lowering, intervened between ca. 11,500 and 10,400 yr B.P. This last phase corresponds partly to the Younger Dryas time period. Zone III (ca. 10,000 to 2800 yr B.P.). After ca. 10,400 yr B.P. the climate became very wet until ca. 3000 yr B.P. A sharp decrease in the Gramineae intervened at ca. 10,000 yr B.P.; from ca. 9500 to 3000 yr B.P. they remained very low, between 0 and 3%, and the forest trees reached their maximum extension. Most of the trees exhibited large variations with quasi-periods of around 1000 to mainly 2000 yr (ca. 2200 calendar years), which could be related to large sylvigenetic or successional cycles. In this zone the Caesalpiniaceae were relatively well represented, with a maximum extension between 4500 and 3000 yr B.P. Podocarpus, a typical tree of the montane stratiform cloud forests, exhibited very low frequencies before 10,000 yr B.P. but their relative increase during the early and middle Holocene can only be explained by its growth on distant mountains. Its maximum extension phase was roughly synchronous with that of Caesalpiniaceae. The climate was warm and wet, but cooler on the mountains. Zone IV (ca. 2800 yr B.P. to present time). Around 2800 yr B.P. a sharp increase in the Gramineae, peaking at 30 to 40% of total pollen between ca. 2500 and 2000 yr B.P., indicates a sudden phase of vegetation opening and forest retreat, accompanied by severe erosion. Alchornea, a typical pioneer taxon, increased rapidly at the same time to large frequencies because it develops abundantly in all the openings. Elaeis guineensis, originally a pioneer palm tree, follows the same pattern. The climate was warm, relatively dry, and linked to an increase of seasonality. After 2000 yr B.P. the Gramineae returned to low frequencies, around 10%, associated with a strong increase in trees, indicating that the forest expanded again but not to the same extent as in the early and middle Holocene. The climate was warm and relatively wet, rather similar to the present-day climate.
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The composition of the forest avifauna of Southern Nigeria is analysed in the light of the most recent work in the region. It is found that east of the Niger the affinities are with Lower Guinea: at Lagos with Upper Guinea; in the Ondo-Benin area not definitely with either. A discussion of the barriers between the two sub-regions follows, and it is claimed that, whereas the Lagos-Accra savanna gap and the Cameroon montane area can each account for about one quarter of the total change, factors within Nigeria were almost twice as important in causing differentiation. It is suggested that these factors were a combination of climatic conditions and northward position of the coastline in late or post-Pliocene times, giving rise to a condition similar to that we find today between Lagos and Accra.
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Flightless insects give a clearer view of former distribution of montane habitat in Africa compared with highly mobile animals as birds and butterflies because passive long distance transport and long distance dispersal can be discounted. Only a few species in the twenty-one genera under study are shared between neighbouring mountains which can be explained in all cases by a Pleistocene lowering of the montane habitat by 850 m. Therefore a montane forest cover connecting the mountains at colder times as suggested by the pluvial theory can be refuted which is in correspondence with palynological findings suggesting a dry corridor between the mountains. No montane refuge for flightless insects can be identified, because the most species-rich mountain of a genus differs among the genera under study. Instead, each mountain served as a species refuge with a stable habitat. The requirement of a smaller habitat compared to vertebrates is indicated by endemic species on each single mountain suggesting pre-Pleistocene speciation which results even in endemic genera to one mountain. Different small patches of suitable habitat on one mountain could also explain the radiations found in some genera of flightless insects. In some genera species are lacking on Mt Kenya which indicates—with the findings of no endemic passerine bird on the mountain—a probably very dry condition during colder times. On Mt Cameroon no species of the flightless insect genera occurring on the Eastern mountains are found. This contrasts with the existing patterns of birds, grasses and butterflies.
Article
Using raw geographic distributions for Neotropical species and subspecies of the perching birds (Order Passeriformes), we present an hypothesis of area-relationships for twelve avian areas of endemism in the lowland Neotropics. With 1717 characters (distributions of species and subspecies) we find a single most parsimonious tree of area-relationships. This topology does not match area-relationships determined from phylogenetic studies of morphologic characters in some Neotropical birds. Analyses of data partitions representing major taxonomic subdivisions within Passerine birds provide many different hypotheses of area-relationships. These results suggest that a single set of Neotropical area-relationships is not likely. In the future, we suggest greater emphasis be placed on research to document patterns in the Neotropics, particularly phylogenetic patterns, than on speculation about what processes have been important for diversification.
Article
A total of 48 new species of birds were described in the ten year period from 1966 to 1975. Of these 28 can be considered good species but nine of the eleven doubtful species might also turn out to be valid (6, 7, 8, 13, 18, 27, 29, 30, 31). The 28 species can be tabulated as follows:Aa New species in new genera (2): 4, 40Ab New species not clearly members of a superspecies (14): 5, 9, 11, 14, 16, 19, 20, 24, 34, 39, 43, 45, 47, 48Ac Allospecies (members of superspecies) (12): 2, 3, 10, 15, 17, 21, 25, 32, 35, 36, 42, 46An additional 20 names cannot now be assigned the status of valid full species:Ba Species inquirendae (12): 6, 7, 8, 12, 13, 18, 26, 27, 28, 29, 30, 31Bb Subspecies (2): 1, 23Bc Synonyms (6): 22, 33, 37, 38, 41, 44In den 10 Jahren 1966–1975 wurden 48 Arten neu benannt, von denen 28 als gute Arten betrachtet werden knnen. Sie lassen sich wie folgt gruppieren:2 neue Arten in neuen Gattungen: 4, 4014 neue Arten, die nicht offensichtlich Mitglieder einer Superspezies sind: 5, 9, 11, 14, 16, 19, 20, 24, 34, 39, 43, 45, 47, 4812 neue Allospezies, die Mitglieder einer Superspezies sind: 2, 3, 10, 15, 17, 21, 25, 32, 35, 36, 42, 46Weitere 20 Namen sind entweder unsicher (6, 7, 8, 12, 13, 18, 26, 27, 28, 29, 30, 31), oder Subspezies (1, 23), oder einwandfrei Synonyme (22, 33, 37, 41, 44). Jhrlich werden im Durchschnitt immer noch 2.8 gute neue Arten beschrieben. Mehr neue Arten sind in der letzten Zehnjahresperiode aus der Neotropischen Region beschrieben worden als aus allen andern Teilen der Welt zusammen.
Article
A total of 18 species of birds was described as new in the five year period from 1976 to 1980. Of these, 12 can be considered good species (but three doubtful species [2, 6, 17] might well turn out to be valid):Aa New species in new genera (2): 7, 18 Ab New species not clearly members of a superspecies (3): 9, 13, 16 Ac Allospecies (members of a superspecies) (7): 3, 4, 8, 10, ?11, 12, 15 An additional 6 names cannot now be assigned the status of valid full species:Ba Species inquirendae (3): 2, 6, 17 Bb Subspecies: none Bc Synonyms (1): 1 Bd Invalid names (2): 5, 14 In den 5 Jahren 1976–1980 wurden 18 Arten neu benannt, wovon 12 als gute Arten betrachtet werden knnen. Diese lassen sich wie folgt gruppieren:Aa Neue Arten in neuen Gattungen (2): 7, 18 Ab Neue Arten, die nicht offensichtlich Mitglieder einer Superspecies sind (3): 9, 13, 16 Ac Neue Allospezies, die Mitglieder einer Superspezies sind (7): 3, 4, 8, 10, ?11, 12, 15 Weitere 6 Namen sind entweder unsicher (2, 6, 17) oder Synonyme (1) oder einwandfrei unerlaubt (5, 14).Jhrlich wurden 1976–1980 im Durchschnitt 2,4 gute neue Arten beschrieben (ungefhr 3,0 1981–1985; 3,1 1966–1975; 3,5 1956–1965; 2,6 1941–1955; 6,0 1938–1941; im Durchschnitt 3,3 1938–1985).Zwei der 12 guten Arten wurden in zwei neuen Gattungen beschrieben (Xenoglaux loweryi undNephelornis oneilli). Eine gute Allospezies vonSitta (S. [krueperi]ledanti) wurde aus der Palarktis beschrieben; nur eine neue gute Art (4) stammt aus Afrika. Alle anderen (10) sind aus Sdamerika beschrieben worden. Sechs bis sieben gute neue Arten (50–60 %) sind Allospezies in Superspezies.
Article
A widely accepted paradigm for speciation in tropical forests, the refuge theory, requires periodic habitat fragmentation driven by global climatic fluctuations to provide conditions for allopatric speciation. This implies that comparative species richness in refugia is due to loss of diverse communities in areas affected by climatic cycles. In this study we compare distribution patterns of bird and plant taxa which we consider to be of either deep phylogenetic lineages or recent radiations. It is demonstrated that lowland areas which have been postulated as Pleistocene refugia are dominated by species which represent lineages of pre-Pleistocene age. Since variations in species richness within these forest tracts reflect currently apparent environmental variables which might be considered to determine carrying capacity, we do not need to postulate that richness is the result of changes in forest cover in the past. Recently diversified taxa of plants and birds are found mainly at the periphery of the main rain forest blocks and in habitat islands outside them. Here, peak concentrations of young restricted-range species are often congruent with clusters of old and biogeographically relictual species. It is suggested that this reflects special intrinsic environmental properties of these areas, in the form of long-term environmental stability caused mainly by persistent orographic rain or mist. In this case, richness is not necessarily due to extinction outside these areas. Stability not only enables survival of relictual taxa, but also promotes morphological differentiation of radiating taxa, leading to aggregates of taxa of restricted distribution.
Article
"Reprinted from the Bulletin of the American Museum of Natural History, vol. 65." Vita. "Scientific results of the Congo Expedition. Ornithology, no. 12"--P. 3. Thesis (Ph. D.)--Columbia University, 1932. Includes bibliographical references.
Article
It is generally accepted that accentuated global climatic cycles since the Plio-Pleistocene (2.8 Ma ago) have caused the intermittent fragmentation of forest regions into isolated refugia thereby providing a mechanism for speciation of tropical forest biota contained within them. However, it has been assumed that this mechanism had its greatest effect in the species rich lowland regions. Contrary evidence from molecular studies of African and South American forest birds suggests that areas of recent intensive speciation, where mostly new lineages are clustered, occur in discrete tropical montane regions, while lowland regions contain mostly old species. Two predictions arise from this finding. First, a species phylogeny of an avian group, represented in both lowland and montane habitats, should be ordered such that montane forms are represented by the most derived characters. Second, montane speciation events should predominate within the past 2.8 Ma. In order to test this model I have investigated the evolutionary history of the recently radiated African greenbuls (genus Andropadus), using a molecular approach. This analysis finds that montane species are a derived monophyletic group when compared to lowland species of the same genus and recent speciation events (within the Plio-Pleistocene) have exclusively occurred in montane regions. These data support the view that montane regions have acted as centres of speciation during recent climatic instability.
Article
Ramphocelus tanagers are distributed throughout the Neotropical lowlands. In this paper, mitochondrial DNA (mtDNA) sequence data from cytochrome b and ND2 genes are used to estimate relationships among seven of nine species of the avian genus Ramphocelus. Genetic differentiation is high between Ramphocelus passerinii passerinii and Ramphocelus passerinii costaricensis, and the two subspecies are diagnosable and distinct from one another both morphologically and genetically. Thus, elevation to species status is recommended. Three clades are supported by both gene sequences; one clade contains R. passerinii, R. costaricensis, and R. icteronotus; the second clade contains Ramphocelus carbo, Ramphocelus bresilius, and Ramphocelus nigrogularis; the third clade contains Ramphocelus sanguinolentus. The degree of saturation was assessed for both genes and saturation of third position of codons occurs by 10-12% uncorrected pairwise sequence divergence. The general area cladogram suggests the following area relationships: Pacific and Caribbean Central America are sister areas, Chocó is the sister to the Central American area, and Amazonia/southeastern Brazil is the outgroup area to the Chocó/Central American clade.
Article
Pleistocene biogeographic events have traditionally been ascribed a major role in promoting speciations and in sculpting the present-day diversity and distributions of vertebrate taxa. However, this paradigm has recently come under challenge from a review of interspecific mtDNA genetic distances in birds: most sister-species separations dated to the Pliocene. Here we summarize the literature on intraspecific mtDNA phylogeographic patterns in birds and reinterpret the molecular evidence bearing on Pleistocene influences. At least 37 of the 63 avian species surveyed (59%) are sundered into recognizable phylogeographic units, and 28 of these separations (76%) trace to the Pleistocene. Furthermore, use of phylogroup separation times within species as minimum estimates of 'speciation durations' also indicates that many protracted speciations, considered individually, probably extended through time from Pliocene origins to Pleistocene completions. When avian speciation is viewed properly as an extended temporal process rather than as a point event, Pleistocene conditions appear to have played an active role both in initiating major phylogeographic separations within species, and in completing speciations that had been inaugurated earlier. Whether the Pleistocene was exceptional in these regards compared with other geological times remains to be determined.
Species limits in the genus Bleda
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Chappuis, C., and C. Erard 1993. Species limits in the genus Bleda Bon-NO. 3270
  • American
AMERICAN MUSEUM NOVITATES aparte, 1857. Z. zool. Syst. Evolutionsforsch. 31: 280-299.
Canary 1.2 User's manual
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Charif, R. A., S. Mitchell, and C. W. Clark 1995. Canary 1.2 User's manual. Ithaca, NY: Cornell Lab. Ornith.
Speciation and its ontology: the empirical consequences of alternative species concepts for understanding patterns and processes of differentiation
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Cracraft, J. 1989. Speciation and its ontology: the empirical consequences of alternative species concepts for understanding patterns and processes of differentiation, pp. 28-59.
Flore et faune du Parc National d'Odzala
Dowsett, R., and F. Dowsett-Lemaire (eds.) 1997. Flore et faune du Parc National d'Odzala, Congo. Turaco Res. Rep. 6.
Pleistocene forest refuges: fact or fancy?
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Endler, J. A. 1982. Pleistocene forest refuges: fact or fancy? In G. Prance (ed.) Biological diversification in the tropics: 641-656. New York: Columbia Univ. Press. Fay, J. M. 1988. Forest monkey populations in the Central African Republic: the northern limits. A census in Manovo-Gounda-St.
The avifauna of Dzanga-Ndoki National Park and Dzanga-Sangha Rainforest Reserve
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Green, A. A., and R. W. Carroll 1991. The avifauna of Dzanga-Ndoki National Park and Dzanga-Sangha Rainforest Reserve, Central African Republic. Malimbus 13: 49-66.
An Atlas of Speciation in African Passerine Birds
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Hall, B. P., and R. E. Moreau 1970. An Atlas of Speciation in African Passerine Birds. London: British Museum (Natural History).
A review of some genera of African chats (Aves: Muscicapidae, Erithacini)
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  • F P Jensen
Jackson, F. 1910. Bull. Br. Ornithol. Club 25: 85. Jensen, F. P. 1989. A review of some genera of African chats (Aves: Muscicapidae, Erithacini). Steenstrupia 15(6): 161-175.
The herpetofauna of the Central African Republic, with a description of a new species of Rhinotyphlops (Serpentes: Typhlopidae)
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Joger, U. 1990. The herpetofauna of the Central African Republic, with a description of a new species of Rhinotyphlops (Serpentes: Typhlopidae). In G. Peters and R. Hutterer (eds.), Vertebrates in the tropics: 85-102. Bonn: Mus. Alexander Koenig.
Birds of the African rain forests. Sounds of nature 9
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Keith, G. S., and W.W.H. Gunn 1971. Birds of the African rain forests. Sounds of nature 9. Fed. of Ontario Nat. and Am. Mus. Nat. Hist.
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Mayr, E., and R. A. Paynter, Jr. (eds.) 1964. Check-list of birds of the world, vol. 10. Cambridge, MA.: Mus. Comp. Zool.
Speciation patterns of birds in the central African forest refugia and their relationship with other refugia
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Prigogine, A. 1988. Speciation patterns of birds in the central African forest refugia and their relationship with other refugia. Acta XIX Congr. Int. Ornithol. 2: 144-157. Ottawa: Univ. Ottawa Press.
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Reichenow, A. 1874. J. Ornithol.:103.
An Atlas of Speciation in African Nonpasserine Birds
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Snow, D. 1978. An Atlas of Speciation in African Nonpasserine Birds. London: British Museum (Natural History).
A revised check-list of African shrikes, orioles, drongos, starlings, crows, waxwings, cuckoo-shrikes, bulbuls, accentors, thrushes and babblers
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White, C.M.N. 1962. A revised check-list of African shrikes, orioles, drongos, starlings, crows, waxwings, cuckoo-shrikes, bulbuls, accentors, thrushes and babblers. Lusaka: Govt. Printer.