A clear challenge for ecological niche modelling is determining how to best mitigate the effects of sampling bias from commonly collected biodiversity data. Recent approaches have focused on filtering occurrences in overrepresented regions based on geographic or environmental proximity. We tested the efficacy of filtering in geographic and environmental space using occurrence data from four species. Our evaluation strategies examined 14 distance measures in geographic and environmental spaces and eight combinations of environmental variables and their ordinations. This resulted in 78 datasets for each species, which we evaluated using area under the curve (AUC), the difference between training and testing AUC, omission rate, the true skill statistic, and Schoener's D to examine the effects of different filtering schemes. The degree of change produced by filtering on predicted suitability and evaluation statistics increased with increasing range size. Environmental filtering resulted in higher model fit at larger extents and retained more occurrences than geographic filtering. Our results indicate that models should be evaluated using multiple evaluation statistics at multiple thresholds. The use of bin sizes when filtering in environmental space allows for simple comparison between species and filter types and makes for an easily reportable and repeatable distance metric. We specifically recommend that ecological niche models using natural history collection data filter in environmental space with variables derived from permutation importance or the first few axes of a principal components ordination.
The Riverine Barriers Hypothesis (RBH) posits that tropical rivers can be effective barriers to gene flow, based on observations that range boundaries often coincide with river barriers. Over the last 160 years, the RBH has received attention from various perspectives, with a particular focus on vertebrates in the Amazon Basin. To our knowledge, no molecular assessment of the RBH has been conducted on birds in the Afrotropics, despite its rich avifauna and many Afrotropical bird species being widely distributed across numerous watersheds and basins. Here, we provide the first genetic evidence that an Afrotropical river has served as a barrier for birds and for their lice, based on four understory bird species collected from sites north and south of the Congo River. Our results indicate near-contemporaneous, Pleistocene lineage diversification across the Congo River in these species. Our results further indicate differing levels of genetic variation in bird lice; the extent of this variation appears linked to the life-history of both the host and the louse. Extensive cryptic diversity likely is being harbored in Afrotropical forests, in both understory birds and their lice. Therefore, these forests may not be “museums” of old lineages. Rather, substantial evolutionary diversification may have occurred in Afrotropical forests throughout the Pleistocene, supporting the Pleistocene Forest Refuge Hypothesis. Strong genetic variation in birds and their lice within a small part of the Congo Basin forest indicates that we may have grossly underestimated diversity in the Afrotropics, making these forests home of substantial biodiversity in need of conservation.
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. 1.8Ma-11,700Ka). 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. 7.5Ma), while the speciation events for the rest of the genus are dated to the Pliocene (c. 5-1.8Ma). 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 diversit...
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