Ecological responses to variation in seasonal snow cover.

Seasonal snow is among the most important factors governing the ecology of many terrestrial ecosystems, but rising global temperatures are changing snow regimes and driving widespread declines in the depth and duration of snow cover. Loss of the insulating snow layer will fundamentally change the environment. Understanding how individuals, populations, and communities respond to different snow conditions is thus essential for predicting and managing future ecosystem change. We synthesized 365 studies that examined ecological responses to variation in winter snow conditions. This research encompasses a broad range of methods (experimental manipulations, measurement of natural snow gradients, and long-term monitoring), locations (35 countries), study organisms (plants, mammals, arthropods, birds, fish, lichen, and fungi), and response measures. Earlier snowmelt was consistently associated with advanced spring phenology in plants, mammals, and arthropods. Reduced snow depth often increased mortality or physical injury in plants, although there were few clear effects on animals. Neither snow depth nor snowmelt timing had clear or consistent directional effects on body size of animals or biomass of plants. However, because 96% of studies were from the northern hemisphere, the generality of these trends across ecosystems and localities is also unclear. We identified substantial research gaps for several taxonomic groups and response types; research on wintertime responses was notably scarce. Future research should prioritize examination of the mechanisms underlying responses to changing snow conditions and the consequences of those responses for seasonally snow-covered ecosystems.

Respuestas Ecológicas a la Variación de la Cobertura Estacional de Nieve Resumen La nieve estacional se encuentra entre los factores más importantes que determinan la ecología de muchos ecosistemas terrestres, pero las crecientes temperaturas mundiales están cambiando los sistemas de nieve y causando declinaciones generalizadas en la profundidad y la duración de la capa de nieve. La pérdida de la capa de nieve aislante cambiará fundamentalmente el ambiente. El entendimiento de cómo los individuos, las poblaciones y las comunidades responden a las diferentes condiciones de nieve es esencial para predecir y manejar los cambios del ecosistema en el futuro. Sintetizamos 365 estudios que examinaron las respuestas ecológicas a la variación en las condiciones invernales de nieve. Esta investigación engloba una gama amplia de métodos (manipulaciones experimentales, medida de los gradientes naturales de nieve y monitoreo a largo plazo), localidades (35 países), organismos de estudio (plantas, mamíferos, artrópodos, aves, peces, líquenes y hongos) y medidas de respuesta. El deshielo temprano estuvo asociado continuamente con el adelanto de la fenología de plantas, mamíferos y artrópodos en primavera. La reducción de la profundidad de la nieve con frecuencia incrementó la mortalidad o las lesiones físicas en las plantas, aunque tuvo pocos efectos visibles sobre los animales. Ni la profundidad de la nieve ni la temporalidad del deshielo tuvieron efectos direccionales claros o consistentes sobre el tamaño corporal de los animales o la biomasa de las plantas. Sin embargo, ya que el 96% de los estudios se realizó en el hemisferio norte, la generalidad de estas tendencias en todos los ecosistemas y localidades tampoco está clara. Identificamos vacíos importantes en la investigación en torno a varios grupos taxonómicos y los tipos de respuesta; la información sobre las respuestas invernales estaba particularmente reducida. Las futuras investigaciones deberían priorizar el análisis de los mecanismos subyacentes a las respuestas ante las condiciones cambiantes de nieve y las consecuencias de aquellas respuestas para los ecosistemas cubiertos de nieve.

Taxonomic notes on the Australian skyhopper genus Kosciuscola Sjstedt (Orthoptera: Acrididae: Oxyinae).

The Australian skyhopper genus Kosciuscola Sjstedt consists of brachypterous species that inhabit the Australian alpine and subalpine region. The genus used to include 5 species and 1 subspecies, but according to a recent phylogenomic study, there could be as many as 14 species in the genus, that are genetically and geographically isolated from each other. This study represents the first step in describing and documenting the diversity of this interesting genus. In this study, we redefine the type species K. tristis, and elevate its subspecies K. tristis restrictus as a valid species on the basis of distinct morphological traits, geographical isolation, and phylogenomic evidence.

How is adaptive potential distributed within species ranges?

Quantitative genetic variation (QGV) represents a major component of adaptive potential and, if reduced toward range-edge populations, could prevent a species' expansion or adaptive response to rapid ecological change. It has been hypothesized that QGV will be lower at the range edge due to small populations-often the result of poor habitat quality-and potentially decreased gene flow. However, whether central populations are higher in QGV is unknown. We used a meta-analytic approach to test for a general QGV-range position relationship, including geographic and climatic distance from range centers. We identified 35 studies meeting our criteria, yielding nearly 1000 estimates of QGV (including broad-sense heritability, narrow-sense heritability, and evolvability) from 34 species. The relationship between QGV and distance from the geographic range or climatic niche center depended on the focal trait and how QGV was estimated. We found some evidence that QGV declines from geographic centers but that it increases toward niche edges; niche and geographic distances were uncorrelated. Nevertheless, few studies have compared QGV in both central and marginal regions or environments within the same species. We call for more research in this area and discuss potential research avenues related to adaptive potential in the context of global change.

Predicting species and community responses to global change using structured expert judgement: An Australian mountain ecosystems case study.

Conservation managers are under increasing pressure to make decisions about the allocation of finite resources to protect biodiversity under a changing climate. However, the impacts of climate and global change drivers on species are outpacing our capacity to collect the empirical data necessary to inform these decisions. This is particularly the case in the Australian Alps which have already undergone recent changes in climate and experienced more frequent large-scale bushfires. In lieu of empirical data, we use a structured expert elicitation method (the IDEA protocol) to estimate the change in abundance and distribution of nine vegetation groups and 89 Australian alpine and subalpine species by the year 2050. Experts predicted that most alpine vegetation communities would decline in extent by 2050; only woodlands and heathlands are predicted to increase in extent. Predicted species-level responses for alpine plants and animals were highly variable and uncertain. In general, alpine plants spanned the range of possible responses, with some expected to increase, decrease or not change in cover. By contrast, almost all animal species are predicted to decline or not change in abundance or elevation range; more species with water-centric life-cycles are expected to decline in abundance than other species. While long-term ecological data will always be the gold standard for informing the future of biodiversity, the method and outcomes outlined here provide a pragmatic and coherent basis upon which to start informing conservation policy and management in the face of rapid change and a paucity of data.

Do different rates of gene flow underlie variation in phenotypic and phenological clines in a montane grasshopper community?

Species responses to environmental change are likely to depend on existing genetic and phenotypic variation, as well as evolutionary potential. A key challenge is to determine whether gene flow might facilitate or impede genomic divergence among populations responding to environmental change, and if emergent phenotypic variation is dependent on gene flow rates. A general expectation is that patterns of genetic differentiation in a set of codistributed species reflect differences in dispersal ability. In less dispersive species, we predict greater genetic divergence and reduced gene flow. This could lead to covariation in life-history traits due to local adaptation, although plasticity or drift could mirror these patterns. We compare genome-wide patterns of genetic structure in four phenotypically variable grasshopper species along a steep elevation gradient near Boulder, Colorado, and test the hypothesis that genomic differentiation is greater in short-winged grasshopper species, and statistically associated with variation in growth, reproductive, and physiological traits along this gradient. In addition, we estimate rates of gene flow under competing demographic models, as well as potential gene flow through surveys of phenological overlap among populations within a species. All species exhibit genetic structure along the elevation gradient and limited gene flow. The most pronounced genetic divergence appears in short-winged (less dispersive) species, which also exhibit less phenological overlap among populations. A high-elevation population of the most widespread species, Melanoplus sanguinipes, appears to be a sink population derived from low elevation populations. While dispersal ability has a clear connection to the genetic structure in different species, genetic distance does not predict growth, reproductive, or physiological trait variation in any species, requiring further investigation to clearly link phenotypic divergence to local adaptation.

Physiological Limits along an Elevational Gradient in a Radiation of Montane Ground Beetles.

A central challenge in ecology and biogeography is to determine the extent to which physiological constraints govern the geographic ranges of species along environmental gradients. This study tests the hypothesis that temperature and desiccation tolerance are associated with the elevational ranges of 12 ground beetle species (genus Nebria) occurring on Mt. Rainier, Washington, U.S.A. Species from higher elevations did not have greater cold tolerance limits than lower-elevation species (all species ranged from -3.5 to -4.1°C), despite a steep decline in minimum temperature with elevation. Although heat tolerance limits varied among species (from 32.0 to 37.0°C), this variation was not generally associated with the relative elevational range of a species. Temperature gradients and acute thermal tolerance do not support the hypothesis that physiological constraints drive species turnover with elevation. Measurements of intraspecific variation in thermal tolerance limits were not significant for individuals taken at different elevations on Mt. Rainier, or from other mountains in Washington and Oregon. Desiccation resistance was also not associated with a species' elevational distribution. Our combined results contrast with previously-detected latitudinal gradients in acute physiological limits among insects and suggest that other processes such as chronic thermal stress or biotic interactions might be more important in constraining elevational distributions in this system.

Conserved and narrow temperature limits in alpine insects: Thermal tolerance and supercooling points of the ice-crawlers, Grylloblatta (Insecta: Grylloblattodea: Grylloblattidae).

For many terrestrial species, habitat associations and range size are dependent on physiological limits, which in turn may influence large-scale patterns of species diversity. The temperature range experienced by individuals is considered to shape the breadth of the thermal niche, with species occupying temporally and/or geographically stable climates tolerating a narrow temperature range. High-elevation environments experience large temperature fluctuations, with frequent periods below 0 °C, but Grylloblatta (Grylloblattodea: Grylloblattidae) occupy climatically stable microhabitats within this region. Here we test critical thermal limits and supercooling points for five Grylloblatta populations from across a large geographic area, to examine whether the stable microhabitats of this group are associated with a narrow thermal niche and assess their capacity to tolerate cold conditions. Thermal limits are highly conserved in Grylloblatta, despite substantial genetic divergence among populations spanning 1500 m elevation and being separated by over 500 km. Further, Grylloblatta show exceptionally narrow thermal limits compared to other insect taxa with little capacity to improve cold tolerance via plasticity. In contrast, upper thermal limits were significantly depressed by cold acclimation. Grylloblatta maintain coordinated movement until they freeze, and they die upon freezing. Convergence of the critical thermal minima, supercooling point and lower lethal limits point to adaptation to a cold but, importantly, constant thermal environment. These physiological data provide an explanation for the high endemism and patchy distribution of Grylloblatta, which relies on subterranean retreats to accommodate narrow thermal limits. These retreats are currently buffered from temperature fluctuations by snow cover, and a declining snowpack thus places Grylloblatta at risk of exposure to temperatures beyond its tolerance capacity.

Strong genetic structure corresponds to small-scale geographic breaks in the Australian alpine grasshopper Kosciuscola tristis.

BACKGROUND: Mountain landscapes are topographically complex, creating discontinuous 'islands' of alpine and sub-alpine habitat with a dynamic history. Changing climatic conditions drive their expansion and contraction, leaving signatures on the genetic structure of their flora and fauna. Australia's high country covers a small, highly fragmented area. Although the area is thought to have experienced periods of relative continuity during Pleistocene glacial periods, small-scale studies suggest deep lineage divergence across low-elevation gaps. Using both DNA sequence data and microsatellite markers, we tested the hypothesis that genetic partitioning reflects observable geographic structuring across Australia's mainland high country, in the widespread alpine grasshopper Kosciuscola tristis (Sjösted). RESULTS: We found broadly congruent patterns of regional structure between the DNA sequence and microsatellite datasets, corresponding to strong divergence among isolated mountain regions. Small and isolated mountains in the south of the range were particularly distinct, with well-supported divergence corresponding to climate cycles during the late Pliocene and Pleistocene. We found mixed support, however, for divergence among other mountain regions. Interestingly, within areas of largely contiguous alpine and sub-alpine habitat around Mt Kosciuszko, microsatellite data suggested significant population structure, accompanied by a strong signature of isolation-by-distance. CONCLUSIONS: Consistent patterns of strong lineage divergence among different molecular datasets indicate genetic breaks between populations inhabiting geographically distinct mountain regions. Three primary phylogeographic groups were evident in the highly fragmented Victorian high country, while within-region structure detected with microsatellites may reflect more recent population isolation. Despite the small area of Australia's alpine and sub-alpine habitats, their low topographic relief and lack of extensive glaciation, divergence among populations was on the same scale as that detected in much more extensive Northern hemisphere mountain systems. The processes driving divergence in the Australian mountains might therefore differ from their Northern hemisphere counterparts.

Niche breadth predicts geographical range size: a general ecological pattern.

The range of resources that a species uses (i.e. its niche breadth) might determine the geographical area it can occupy, but consensus on whether a niche breadth-range size relationship generally exists among species has been slow to emerge. The validity of this hypothesis is a key question in ecology in that it proposes a mechanism for commonness and rarity, and if true, may help predict species' vulnerability to extinction. We identified 64 studies that measured niche breadth and range size, and we used a meta-analytic approach to test for the presence of a niche breadth-range size relationship. We found a significant positive relationship between range size and environmental tolerance breadth (z = 0.49), habitat breadth (z = 0.45), and diet breadth (z = 0.28). The overall positive effect persisted even when incorporating sampling effects. Despite significant variability in the strength of the relationship among studies, the general positive relationship suggests that specialist species might be disproportionately vulnerable to habitat loss and climate change due to synergistic effects of a narrow niche and small range size. An understanding of the ecological and evolutionary mechanisms that drive and cause deviations from this niche breadth-range size pattern is an important future research goal.

Estimating genetic benefits of polyandry from experimental studies: a meta-analysis.

The consequences of polyandry for female fitness are controversial. Sexual conflict studies and a meta-analysis of mating rates in insects suggest that there is a longevity cost when females mate repeatedly. Even so, compensatory material benefits can elevate egg production and fertility, partly because polyandry ensures an adequate sperm supply. Polyandry can therefore confer direct benefits. The main controversy surrounds genetic benefits. The argument is analogous to that surrounding the evolution of conventional female mate choice, except that with polyandry it is post-copulatory mechanisms that might bias paternity towards males with higher breeding values for fitness. Recent meta-analyses of extra-pair copulations in birds have cast doubt on whether detectable genetic benefits exist. By contrast, another meta-analysis showed that polyandry elevates egg hatching success (possibly due to a fertilization bias towards sperm with paternal genes that elevate embryo survival) in insects. A detailed summary of whether polyandry elevates other components of offspring performance is lacking. Here we present a comprehensive meta-analysis of 232 effect sizes from 46 experimental studies. These experiments were specifically designed to try to quantify the potential genetic benefits of polyandry by controlling fully for the number of matings by females assigned to monandry and polyandry treatments. The bias-corrected 95% confidence intervals for egg hatching success (d = -0.01 to 0.61), clutch production (d = 0.07 to 0.45) and fertility (d = 0.04 to 0.40) all suggest that polyandry has a beneficial effect (although P values from parametric tests were marginally non-significant at P = 0.075, 0.052 and 0.058, respectively). Polyandry was not significantly beneficial for any single offspring performance trait (e.g. growth rate, survival, adult size), but the test power was low due to small sample sizes (suggesting that many more studies are still needed). We then calculated a composite effect size that provides an index of general offspring performance. Depending on the model assumptions, the mean effect of polyandry was either significantly positive or marginally non-significant. A possible role for publication bias is discussed. The magnitude of the reported potential genetic benefits (d = 0.07 to 0.19) are larger than those from two recent meta-analyses comparing offspring sired by social and extra-pair mates in birds (d = 0.02 to 0.04). This difference raises the intriguing possibility that cryptic, post-copulatory female choice might be more likely to generate 'good gene' or 'compatible gene' benefits than female choice of mates based on the expression of secondary sexual traits.