Speciation generally involves a three-step process--range expansion, range fragmentation and the development of reproductive isolation between spatially separated populations. Speciation relies on cycling through these three steps and each may limit the rate at which new species form. We estimate phylogenetic relationships among all Himalayan songbirds to ask whether the development of reproductive isolation and ecological competition, both factors that limit range expansions, set an ultimate limit on speciation. Based on a phylogeny for all 358 species distributed along the eastern elevational gradient, here we show that body size and shape differences evolved early in the radiation, with the elevational band occupied by a species evolving later. These results are consistent with competition for niche space limiting species accumulation. Even the elevation dimension seems to be approaching ecological saturation, because the closest relatives both inside the assemblage and elsewhere in the Himalayas are on average separated by more than five million years, which is longer than it generally takes for reproductive isolation to be completed; also, elevational distributions are well explained by resource availability, notably the abundance of arthropods, and not by differences in diversification rates in different elevational zones. Our results imply that speciation rate is ultimately set by niche filling (that is, ecological competition for resources), rather than by the rate of acquisition of reproductive isolation.
The primary explanation for the latitudinal gradient in species diversity must lie in why species fail to expand ranges across different climatic regimes. Theories of species gradients based in niche conservatism assume that whole clades are confined to particular climatic regimes because the traits they share limit adaptation to alternative regimes. We assess these theories in an analysis of the twofold decline in bird species richness along the Himalayas from the southeast to the northwest. The presence of fewer species in the northwest is entirely due to a steep decline in the number of forest species; species occupying more open habitats show a reversed gradient. Forest species numbers are exceptionally high at midelevations (1,000-2,000 m) in the southeast, which experience a warm, wet climate not present in the northwest, and a high proportion of these species fail to expand their range to the northwest. Despite this, many species do have populations or close relatives that straddle different climatic regimes along altitudinal gradients and/or the regional gradient, implying that climate-based niche conservatism per se does not strongly constrain range limits. We argue that climate- and competition-mediated resource distributions are important in setting northerly range limits and show that one measure of forest resources (foliage density) is lower in the northwest.
Many models to explain the differences in the flora and fauna of tropical and temperate regions assume that whole clades are restricted to the tropics. We develop methods to assess the extent to which biotas are geographically discrete, and find that transition zones between regions occupied by tropical-associated or temperate-associated biotas are often narrow, suggesting a role for freezing temperatures in partitioning global biotas. Across the steepest tropical-temperate gradient in the world, that of the Himalaya, bird communities below and above the freezing line are largely populated by different tropical and temperate biotas with links to India and Southeast Asia, or to China respectively. The importance of the freezing line is retained when clades rather than species are considered, reflecting confinement of different clades to one or another climate zone. The reality of the sharp tropical-temperate boundary adds credence to the argument that exceptional species richness in the tropics reflects species accumulation over time, with limited transgressions of species and clades into the temperate.
The swamp deerRucervus duvauceliiis the largest grassland-dwelling endemic cervid of India and Nepal. With a declining population trend across its range, this species is found in fragmented habitats of northern, north-eastern and central India and south-western Nepal. The northern swamp deer subspeciesRucervus duvaucelii duvauceliioccurs in small wetland patches across the states of Uttarakhand and Uttar Pradesh in India and has lost most of its habitat in the last century. Information about the distribution of the swamp deer in the upper Gangetic plains is limited, except in the Jhilmil Jheel Conservation Reserve in Uttarakhand and around the Bijnor barrage area of Hastinapur Wildlife Sanctuary in Uttar Pradesh. We surveyed the upper Gangetic plains between the Reserve and the Sanctuary, including some adjoining areas and three tributaries of the Ganges, to assess the status of these habitats, current swamp deer distribution and the threats faced by the species. We found several areas harbouring swamp deer within non-protected wetlands along the entire surveyed stretch of the upper Ganges and a previously unreported population in Uttar Pradesh. We documented major threats including habitat conversion, livestock grazing, poaching, conflict and other anthropogenic disturbances. We recommend community driven conservation and management ofRucervus duvaucelii duvauceliiin this fragmented landscape to ensure survival of this species and other threatened fauna of these wetlands and grasslands.
Analysis of one of the most comprehensive datasets to date of the largest passerine bird clade, Passerida, identified 10 primary well-supported lineages corresponding to Sylvioidea, Muscicapoidea, Certhioidea, Passeroidea, the 'bombycillids' (here proposed to be recognized as Bombycilloidea), Paridae/ Remizidae (proposed to be recognized as Paroidea), Stenostiridae, Hyliotidae, Regulidae (proposed to be recognized as Reguloidea) and spotted wrenbabbler Spelaeornis formosus. The latter was found on a single branch in a strongly supported clade with Muscicapoidea, Certhioidea and Bombycilloidea, although the relationships among these were unresolved. We conclude that the spotted wren-babbler represents a relict basal lineage within Passerida with no close extant relatives, and we support the already used name Elachura formosa and propose the new family name Elachuridae for this single species.
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