Temperature-associated decreases in demographic rates of Afrotropical bird species over 30 years.

Tropical mountains harbor globally significant levels of biodiversity and endemism. Climate change threatens many tropical montane species, yet little research has assessed the effects of climate change on the demographic rates of tropical species, particularly in the Afrotropics. Here, we report on the demographic rates of 21 Afrotropical bird species over 30 years in montane forests in Tanzania. We used mark-recapture analyses to model rates of population growth, recruitment, and apparent survival as functions of annual mean temperature and annual precipitation. For over one-half of focal species, decreasing population growth rates were associated with increasing temperature. Due to the trend in temperature over time, we substituted a time covariate for the temperature covariate in top-ranked population growth rate models. Temperature was a better explanatory covariate than time for 6 of the 12 species, or 29% of all focal species. Population growth rates were also lower for species found further below their elevational midpoint and for smaller-bodied species. Changes in population growth rates were more closely tied to changes in recruitment than to changes in apparent survival. There were no consistent associations between demographic rates and precipitation. This study demonstrates temperature-associated demographic impacts for 6 (29%) of 21 focal species in an Afrotropical understory bird community and highlights the need to incorporate the impacts of climate change on demographic rates into conservation planning across the tropics.

Positive abundance-elevational range size relationship weakened from temperate to subtropical ecosystems.

Describing the patterns and revealing the underlying mechanisms responsible for variations in community structure remain a central focus in ecology. However, important gaps remain, including our understanding of species abundance. Most studies on abundance-based relationships are from either temperate ecosystems or tropical ecosystems, and few have explicitly tested abundance-based relationships across a temperate to tropical ecotone. Here, we use a comprehensive dataset of breeding birds across elevation spanning a temperate to subtropical gradient in the Himalayas-Hengduan Mountains of China to examine the relationship between species abundance and (a) elevational range size, (b) body size, (c) elevational range centre and (d) endemicity. We tested a priori predictions for abundance-elevational range size relationship, abundance-body size relationship and abundance-elevational range centre relationship, and explored how these relationships change along this temperate to subtropical mountain ecosystem. We found that species abundance was significantly positively correlated with elevational range size across the study sites, demonstrating the key importance of elevational range size towards species abundance. Body size and elevational range centre are weakly correlated with abundance. A novel finding of our study is that the abundance-elevational range size relationship gradually weakened from temperate to subtropical ecosystems, adding to a growing body of evidence suggesting that abundance-elevational range size tracks a temperate to tropical ecotone. Our study demonstrates that abundance range-size relationship can transition across ecotones where faunas of different evolutionary origins converge. Furthermore, measuring abundance relationships across different environmental variables at the same spatial scale with comparable biogeography is a key strategy that can reveal the underlying mechanisms behind abundance patterns.

Little genetic structure in a Bornean endemic small mammal across a steep ecological gradient.

Janzen's influential "mountain passes are higher in the tropics" hypothesis predicts restricted gene flow and genetic isolation among populations spanning elevational gradients in the tropics. Few studies have tested this prediction, and studies that focus on population genetic structure in Southeast Asia are particularly underrepresented in the literature. Here, we test the hypothesis that mountain treeshrews (Tupaia montana) exhibit limited dispersal across their broad elevational range which spans ~2,300 m on two peaks in Kinabalu National Park (KNP) in Borneo: Mt Tambuyukon (MT) and Mt Kinabalu (MK). We sampled 83 individuals across elevations on both peaks and performed population genomics analyses on mitogenomes and single nucleotide polymorphisms from 4,106 ultraconserved element loci. We detected weak genetic structure and infer gene flow both across elevations and between peaks. We found higher genetic differentiation on MT than MK despite its lower elevation and associated environmental variation. This implies that, contrary to our hypothesis, genetic structure in this system is not primarily shaped by elevation. We propose that this pattern may instead be the result of historical processes and limited upslope gene flow on MT. Importantly, our results serve as a foundational estimate of genetic diversity and population structure from which to track potential future effects of climate change on mountain treeshrews in KNP, an important conservation stronghold for the mountain treeshrew and other montane species.

Why are tropical mountain passes "low" for some species? Genetic and stable-isotope tests for differentiation, migration and expansion in elevational generalist songbirds.

Most tropical bird species have narrow elevational ranges, likely reflecting climatic specialization. This is consistent with Janzen's Rule, the tendency for mountain passes to be effectively "higher" in the tropics. Hence, those few tropical species that occur across broad elevational gradients (elevational generalists) represent a contradiction to Janzen's Rule. Here, we aim to address the following questions. Are elevational generalists being sundered by diversifying selection along the gradient? Does elevational movement cause these species to resist diversification or specialization? Have they recently expanded, suggesting that elevational generalism is short-lived in geological time? To answer these questions, we tested for differentiation, movement and expansion in four elevational generalist songbird species on the Andean west slope. We used morphology and mtDNA to test for genetic differentiation between high- and low-elevation populations. To test for elevational movements, we measured hydrogen isotope (δ2 H) values of metabolically inert feathers and metabolically active liver. Morphology differed for House Wren (Troglodytes aedon) and Hooded Siskin (Spinus magellanicus), but not for Cinereous Conebill (Conirostrum cinereum) and Rufous-collared Sparrow (Zonotrichia capensis) respectively. mtDNA was structured by elevation only in Z. capensis. δ2 H data indicated elevational movements by two tree- and shrub-foraging species with moderate-to-high vagility (C. cinereum and S. magellanicus), and sedentary behaviour by two terrestrial-foraging species with low-to-moderate vagility (T. aedon and Z. capensis). In S. magellanicus, elevational movements and lack of mtDNA structure contrast with striking morphological divergence, suggesting strong diversifying selection on body proportions across the c. 50 km gradient. All species except C. cinereum exhibited mtDNA haplotype variation consistent with recent population expansion across the elevational gradient, potentially concurrent with Holocene anthropogenic habitat conversion for agriculture. In different ways, each species defies the tendency for tropical birds to have long-term stable distributions and sedentary habits. We conclude that tropical elevational generalism is rare due to evolutionary instability.

Cryptic species diversity reveals biogeographic support for the 'mountain passes are higher in the tropics' hypothesis.

The 'mountain passes are higher in the tropics' (MPHT) hypothesis posits that reduced climate variability at low latitudes should select for narrower thermal tolerances, lower dispersal and smaller elevational ranges compared with higher latitudes. These latitudinal differences could increase species richness at low latitudes, but that increase may be largely cryptic, because physiological and dispersal traits isolating populations might not correspond to morphological differences. Yet previous tests of the MPHT hypothesis have not addressed cryptic diversity. We use integrative taxonomy, combining morphology (6136 specimens) and DNA barcoding (1832 specimens) to compare the species richness, cryptic diversity and elevational ranges of mayflies (Ephemeroptera) in the Rocky Mountains (Colorado; approx. 40°N) and the Andes (Ecuador; approx. 0°). We find higher species richness and smaller elevational ranges in Ecuador than Colorado, but only after quantifying and accounting for cryptic diversity. The opposite pattern is found when comparing diversity based on morphology alone, underscoring the importance of uncovering cryptic species to understand global biodiversity patterns.

Life stage, not climate change, explains observed tree range shifts.

Ongoing climate change is expected to shift tree species distribution and therefore affect forest biodiversity and ecosystem services. To assess and project tree distributional shifts, researchers may compare the distribution of juvenile and adult trees under the assumption that differences between tree life stages reflect distributional shifts triggered by climate change. However, the distribution of tree life stages could differ within the lifespan of trees, therefore, we hypothesize that currently observed distributional differences could represent shifts over ontogeny as opposed to climatically driven changes. Here, we test this hypothesis with data from 1435 plots resurveyed after more than three decades across the Western Carpathians. We compared seedling, sapling and adult distribution of 12 tree species along elevation, temperature and precipitation gradients. We analyzed (i) temporal shifts between the surveys and (ii) distributional differences between tree life stages within both surveys. Despite climate warming, tree species distribution of any life stage did not shift directionally upward along elevation between the surveys. Temporal elevational shifts were species specific and an order of magnitude lower than differences among tree life stages within the surveys. Our results show that the observed range shifts among tree life stages are more consistent with ontogenetic differences in the species' environmental requirements than with responses to recent climate change. The distribution of seedlings substantially differed from saplings and adults, while the distribution of saplings did not differ from adults, indicating a critical transition between seedling and sapling tree life stages. Future research has to take ontogenetic differences among life stages into account as we found that distributional differences recently observed worldwide may not reflect climate change but rather the different environmental requirements of tree life stages.

A test for community saturation along the Himalayan bird diversity gradient, based on within-species geographical variation.

The idea that ecological communities are unsaturated is central to many explanations for regional gradients in species diversity. We describe a test for differing degrees of saturation across a regional diversity gradient, based on within-species geographical variation in ecological attributes. If communities in species-poor regions are less saturated than communities in species-rich regions, species that straddle both regions should have broader niches in species-poor regions, exploiting resources that are consumed by other species in species-rich regions. We studied 10 species of Old World leaf warblers that range across the Himalayas. Elevational range and feeding method showed niche contractions in the species-poor north-west Himalayas with respect to the species-rich south-east Himalayas, whereas prey size did not vary geographically. Niche contractions are contrary to the expectation of character release in depauperate environments, as has been shown, for example in mainland-island comparisons. We show that arthropod abundances are likely a limiting resource, and that niche contractions are consistent with measurements of a narrowing of resource availability. Results suggest that north-western warbler communities are at least as saturated as the south-east and that lower resource diversity drives reduced species numbers.

Geographic determinants of gene flow in two sister species of tropical Andean frogs.

Complex interactions between topographic heterogeneity, climatic and environmental gradients, and thermal niche conservatism are commonly assumed to indicate the degree of biotic diversification in montane regions. Our aim was to investigate factors that disrupt gene flow between populations and to determine if there is evidence of downslope asymmetric migration in highland frogs with wide elevational ranges and thermal niches. We determined the role of putative impediments to gene flow (as measured by least-cost path (LCP) distances, topographic complexity, and elevational range) in promoting genetic divergence between populations of 2 tropical Andean frog sister species (Dendropsophus luddeckei, N = 114; Dendropsophus labialis, N = 74) using causal modeling and multiple matrix regression. Although the effect of geographic features was species specific, elevational range and LCP distances had the strongest effect on gene flow, with mean effect sizes (Mantel r and regression coefficients ß), between 5 and 10 times greater than topographic complexity. Even though causal modeling and multiple matrix regression produced congruent results, the latter provided more information on the contribution of each geographic variable. We found moderate support for downslope migration. We conclude that the climatic heterogeneity of the landscape, the elevational distance between populations, and the inability to colonize suboptimal habitats due to thermal niche conservatism influence the magnitude of gene flow. Asymmetric migration, however, seems to be influenced by life history traits.

Species richness of Orthoptera declines with elevation while elevational range of individual species peaks at mid elevation.

Species richness has been shown to decrease, and elevational range increase (the Rapoport effect), with elevation as a consequence of biotic and abiotic factors, but patterns are inconsistent across taxonomic groups. Despite being an important indicator taxon and a component of local communities, Orthoptera distributions at higher elevations in Europe remain unclear. We investigated the relationship of Orthoptera species richness and elevational range with elevation in the Pyrenees mountains, Europe. We conducted sweepnetting surveys supplemented by hand-sampling, at 28 sites stratified by elevation, across three study areas. Using generalised linear models, we found that species richness declined with elevation. Elevation was an important predictor of species richness, but sampling effort and vegetation structure (height and cover) also contributed to estimates of species richness. Using a nonlinear regression to model the elevational range of species over the elevational gradient, we did not observe a Rapoport effect, with elevational range peaking at mid-elevation instead. Smaller elevational ranges of species found at high elevations may be due to a combination of sampling over a restricted elevational range and the presence of specialist high-elevation species. We argue that our findings are useful for understanding species distributions with elevation at the interface between local and regional scales. Clarifying the biotic and abiotic predictors of species distribution is important for informing conservation efforts and predicting consequences of climate change.

Elevational range extension of the Puna Mouse, Punomys (Cricetidae), with the first record of the genus from Chile.

We report an elevational record for the Andean sigmodontine Puna Mouse Punomys, which is also the first record of the genus in Chile. The record is based on a mummified specimen that we discovered at an elevation of 5,461 m (17,917 feet) in the caldera of Volcán Acamarachi, Región de Antofagasta, Chile. Results of a morphological assessment suggest that the specimen can be provisionally referred to the species P. lemminus. This new record also extends the known geographic distribution of the genus by 700 km to the south and brings the known Chilean mammal richness to a total of 170 living species and 88 genera. This finding highlights the need for increased survey efforts in more remote, high-elevation regions and demonstrates that there is still much to be learned about the mammal fauna of the Andean Altiplano.

Se reporta un registro altitudinal para el roedor sigmodontino Punomys, el cual corresponde a su vez al primer hallazgo del género para Chile. Este se basa en un espécimen momificado encontrado a una elevación de 5,461 m en la caldera del Volcán Acamarachi, Región de Antofagasta, Chile. Los caracteres morfológicos sugieren que el espécimen puede ser referido provisionalmente a la especie P. lemminus. Este nuevo registro amplía la distribución geográfica conocida del género en 700 km al sur, y eleva la riqueza de mamíferos vivientes chilenos a un nuevo total de 170 especies y 88 géneros. Este hallazgo resalta la necesidad de aumentar los esfuerzos de prospección en las regiones más remotas y de mayor altitud y demuestra que aún queda mucho por aprender sobre el ensamble de los mamíferos del Altiplano andino.

Prioritized carbon allocation to storage of different functional types of species at the upper range limits is driven by different environmental drivers.

The upper elevational range limit of tree species (including treeline and non-treeline species) is generally considered to result from either carbon limitation or sink limitation. Some evidence also suggests that tree line might reflect preferential carbon allocation to NSC storage at the expense of growth. How might the importance of these potential mechanisms be determined? We used an elevational gradient to examine light-saturated photosynthesis (Asat) and NSC concentrations in plant tissues of three different functional types of tree species. We also examined the effects of consecutive 4 years of in situ defoliation on growth and NSCs at the upper elevational range limit. Declining temperature with increasing elevation did not reduce Asat in any of the species. We found NSC increased with elevation in major storage tissues (e.g., roots and twigs) but not in leaves. The defoliation showed that C storage took priority over growth. Such preferential carbon allocation, directly caused by growth decline, always existed in the deciduous tree species. In the evergreen tree species, however, growth decline resulted from preferential carbon allocation to storage was only detected in 2017 and then disappeared as the intensity of defoliation increased. Our results showed that trees prioritized sustaining stores of C more highly than allocation of growth, regardless of the trees' C or sink limitations. At the cold range limits, the prioritized carbon allocation to storage in deciduous tree species was in response to low temperature stress, while in evergreen tree species, the prioritization of carbon allocation was only a transient physiological response to defoliation disturbances.

Ecological Factors Generally Not Altitude Related Played Main Roles in Driving Potential Adaptive Evolution at Elevational Range Margin Populations of Taiwan Incense Cedar (Calocedrus formosana).

Population diversification can be shaped by a combination of environmental factors as well as geographic isolation interacting with gene flow. We surveyed genetic variation of 243 samples from 12 populations of Calocedrus formosana using amplified fragment length polymorphism (AFLP) and scored a total of 437 AFLP fragments using 11 selective amplification primer pairs. The AFLP variation was used to assess the role of gene flow on the pattern of genetic diversity and to test environments in driving population adaptive evolution. This study found the relatively lower level of genetic diversity and the higher level of population differentiation in C. formosana compared with those estimated in previous studies of conifers including Cunninghamia konishii, Keteleeria davidiana var. formosana, and Taiwania cryptomerioides occurring in Taiwan. BAYESCAN detected 26 F ST outlier loci that were found to be associated strongly with various environmental variables using multiple univariate logistic regression, latent factor mixed model, and Bayesian logistic regression. We found several environmentally dependent adaptive loci with high frequencies in low- or high-elevation populations, suggesting their involvement in local adaptation. Ecological factors, including relative humidity and sunshine hours, that are generally not altitude related could have been the most important selective drivers for population divergent evolution in C. formosana. The present study provides fundamental information in relation to adaptive evolution and can be useful for assisted migration program of C. formosana in the future conservation of this species.

A new species of Brachycephalus (Anura, Brachycephalidae) from the coast of Santa Catarina State, southern Atlantic Forest, Brazil.

We describe a new species of Brachycephalus from municipality of São Francisco do Sul and municipality of Itapoá, in the state of Santa Catarina, Brazil, southern Atlantic Forest. The new species is known from six localities from near sea level up to 250 meters and represents the first record of a "pumpkin-toadlet" occurring in the lowlands. Morphological traits and phylogenetic analysis of a fragment the 16S mtDNA gene place the new species in the Brachycephalus pernix group. The new species is supported by external morphology, osteology, advertisement call, and mtDNA divergence. It is characterized, among other traits by a dorsal body color dark green with a dark brown vertebral stripe, and an orange background; snout-vent length of 9.2-10.8 mm in males and 11.1-12.4 mm in females; and advertisement call short (0.02-0.03 seconds), composed of one high-frequency note (dominant frequency 6.6-7.3 kHz). We observed synchronized alternation in the emission of vocalizations among neighbor males, indicating that males of the new species are able to hear and use vocalizations to interact with each other. We provide descriptions of clutch, eggs, and juvenile and observations on parental care. The new species has not been recorded within any protected area and can be threatened by human-induced habitat loss and modification.

Support for the elevational Rapoport's rule among seed plants in Nepal depends on biogeographical affinities and boundary effects.

As one of the most important hypotheses on biogeographical distribution, Rapoport's rule has attracted attention around the world. However, it is unclear whether the applicability of the elevational Rapoport's Rule differs between organisms from different biogeographical regions. We used Stevens' method, which uses species diversity and the averaged range sizes of all species within each (100 m) elevational band to explore diversity-elevation, range-elevation, and diversity-range relationships. We compared support for the elevational Rapoport's rule between tropical and temperate species of seed plants in Nepal. Neither tropical nor temperate species supported the predictions of the elevational Rapoport's rule along the elevation gradient of 100-6,000 m a.s.l. for any of the studied relationships. However, along the smaller 1,000-5,000 m a.s.l. gradient (4,300 m a.s.l. for range-elevation relationships) which is thought to be less influenced by boundary effects, we observed consistent support for the rule by tropical species, although temperate species did not show consistent support. The degree of support for the elevational Rapoport's rule may not only be influenced by hard boundary effects, but also by the biogeographical affinities of the focal taxa. With ongoing global warming and increasing variability of temperature in high-elevation regions, tropical taxa may shift upward into higher elevations and expand their elevational ranges, causing the loss of temperate taxa diversity. Relevant studies on the elevational Rapoport's rule with regard to biogeographical affinities may be a promising avenue to further our understanding of this rule.