Distributional Edge Research Articles

Westward expansion of the Evening Bat (Nycticeius humeralis) into Dawson and Lincoln counties of central Nebraska

The Evening Bat (Nycticeius humeralis) is a small, migratory species of temperate woodlands in the eastern United States. Along its western distributional edge in the Great Plains, this species resides in wooded riverine corridors. In recent decades, the distribution of Evening Bats has expanded westward throughout the Great Plains. Herein, we report on two county records that further document the distributional expansion of Evening Bats into central Nebraska. In 2019 and 2022, Evening Bats were collected in Lincoln and Dawson counties, respectively, in woodlands adjacent to the Platte River. Increases in wooded habitats along prairie waterways likely has resulted in their westward movements in the Great Plains. With the continued westward expansion, this species should be looked for farther west beyond known distributional limits along prairie waterways throughout the Great Plains.

Preliminary Investigation of Schmalhausen's Law in a Directly Transmitted Pathogen Outbreak System.

The past few decades have been marked by drastic modifications to the landscape by anthropogenic processes, leading to increased variability in the environment. For populations that thrive at their distributional boundaries, these changes can affect them drastically, as Schmalhausen's law predicts that their dynamics are more likely to be susceptible to environmental variation. Recently, this evolutionary theory has been put to the test in vector-borne disease emergences systems, and has been demonstrated effective in predicting emergence patterns. However, it has yet to be tested in a directly transmitted pathogen. Here, we provide a preliminary test of Schmalhausen's law using data on Marburg virus outbreaks originating from spillover events. By combining the two important aspects of Schmalhausen's law, namely climatic anomalies and distance to species distributional edges, we show that Marburgvirus outbreaks may support an aspect of this evolutionary theory, with distance to species distributional edge having a weak influence on outbreak size. However, we failed to demonstrate any effect of climatic anomalies on Marburgvirus outbreaks, arguably related to the lack of importance of these variables in directly transmitted pathogen outbreaks. With increasing zoonotic spillover events occurring from wild species, we highlight the importance of considering ecological variability to better predict emergence patterns.

High thermal plasticity, and vulnerability, in extreme environments at the warm distributional edge: The case of a tidepool shrimp

Climate change threatens the resilience of species, especially at their warm distributional edge in extreme environments. We investigated the thermal vulnerability of the tidepool shrimp, Palaemon elegans in the fast-warming southeastern Mediterranean Sea, its warm distributional edge. Tidepool organisms experience strong and fast thermal fluctuations. This might make them more resilient to change, but also bring them closer to their thermal limits during extreme conditions. To examine the shrimp's resilience, we tested three hypotheses: (1) P. elegance in the southeast Mediterranean has higher critical thermal maximum (CTMax) than in cooler regions, (2) the shrimp possess seasonal acclimatization, but (3) long exposure to extreme summer temperatures might erode its thermal performance making it vulnerable to future climate change. In a series of field observations and laboratory experiments we characterized the shrimp's thermal environment and population dynamics, determined its CTMax and tested diverse physiological performance attributes (respiration, digestion, activity, growth) under a wide range of temperatures during winter and summer. The thermal performance curves show that peak respiration rates occur at higher temperatures in the summer compared to winter, indicating seasonal acclimatization. The shrimp's CTMax was 38.1 °C; higher than its Atlantic counterparts, and an indication of biogeographic adaptation. However, its warming tolerance (the thermal safety margin: distance between maximum pool temperatures and CTMax) was only 0.3 °C, indicating low capacity for dealing with further warming in mostly isolated pools, compared to northeast Atlantic populations that have a wider safety margin. Performance also eroded with increase in exposure duration to high temperatures, and the optimum temperature reduces with time. We predict that prolonged exposure to current (~ 32 °C) and future mean summer values in open water would significantly reduce its performance and increase its mortality. Further, juvenile P. elegans were found to be more vulnerable to extreme (cold and hot) temperatures than adults. Together, these findings indicate that the population viability of this robust coastal water shrimp would eventually be reduced under continuous regional warming and the intensification of extreme events.

Roost Use and Movements of Northern Long-Eared Bats in a Southeast Nebraska Agricultural Landscape

Bats are important bio-indicators of ecosystem health and provide a number of ecosystem services. White-nose Syndrome and habitat loss have led to the decline of many bat species in eastern North America, including the federally threatened northern long-eared bat, Myotis septentrionalis. White-nose Syndrome was only recently found in Nebraska, which lies on the western extent of this species geographic range. To better understand how this forest-dependent species persists in an agriculturally dominated landscape amid a growing number of pressures, we investigated the roosting habits of this bat at the Homestead National Monument of America, located in southeast Nebraska. We mist-netted bats on eight nights in 2019 (16 August–26 August) and caught 55 bats across five species, including five juvenile northern long-eared bats. We located five unique roosts between two juvenile radio-tracked bats; most of the female roosts were in anthropogenic structures and tree cavities within 0.23 km of capture, while most of the male roosts were in snags and tree cavities as far as 2.73 km from the capture site. Fence cavities were also used by other undocumented northern long-eared bats. We recorded three radio-tagged bats that commuted between roosting sites and capture sites within hours after sunset. Our results provide evidence that at the distributional edge for this species, wooded areas, riparian zones, and human-built structures in an intensively managed agricultural landscape are used by this imperiled species.

Supercooling Points of Adult Dermacentor variabilis (Acari: Ixodidae) From a Population Near the Northern Distribution Limit.

The northern distributional limit of Dermacentor variabilis Say, the American dog tick, is expanding in Saskatchewan and Manitoba (western Canada). The ability of D. variabilis to continue to expand its range northwards will depend upon the ability of individuals within populations at the species distributional edge to withstand very low temperatures during winter. One component of cold hardiness is the supercooling point (SCP), the temperature below 0°C at which an individual freezes. In this study, the SCP was determined for 94 questing D. variabilis adults (44 females and 50 males) from an established population near Blackstrap Provincial Park in Saskatchewan. SCP values ranged from -18.2 to -6.7°C, with a median of -13.3°C. This suggests that host-seeking D. variabilis adults differ in their ability to survive exposure to subzero temperatures, for at least a short period of time, without freezing. The distribution of SCPs was bimodal, but there was no significant difference in SCP values between female and male ticks, and no relationship between SCP and tick body weight. It remains to be determined what factors contribute to the variation in SCP values among questing D. variabilis adults.

Local Scale Thermal Environment and Limited Gene Flow Indicates Vulnerability of Warm Edge Populations in a Habitat Forming Macroalga

Species inhabiting warm-edge populations of their distribution are suggested to be at the forefront of global warming due to reduced fitness, limited gene flow and living close to their physiological thermal limits. Determining the scale that governs thermal niche and the functional responses of habitat-forming species to environmental stressors is critical for successful conservation efforts, particularly as coastal ecosystems are impacted by global change. Here, we examine the susceptibility of warm-edge populations to warming, in the habitat-forming macroalga, Hormosira banksii, from south-eastern Australia. We use a quantitative breeding design to quantify intraspecific variation in thermal performance (growth, ontogenic development and photosynthetic efficiency) of different genotypes sourced from sites at the equatorward distributional edge (warm-edge) and those towards the center of its distribution (non-edge). The genetic diversity and structure of H. banksii was also examined using microsatellite markers amongst the same sites. Our results found variable responses in thermal performance for growth and development. Warm-edge germlings grew optimally in lower temperatures tested and had narrower thermal breadth compared to non-edge germlings which grew in higher and more broader temperatures. Warm-edge germlings however, showed greater plasticity to tolerate high light indicated by a greater proportion of energy being dissipated as regulated nonphotochemical quenching (Y(NPQ)) than nonregulated nonphotochemical quenching (Y(NO)). Overall genetic diversity was lower at the warm-edge location with evidence of increased structuring and reduced gene flow in comparison to the non-edge location. Evidence of genetic structuring was not found locally between high and low shore within sites. Together, these data suggest that non-edge populations may be “thermally buffered” from increased temperatures associated with ocean warming. Warm-edge populations of H. banksii, however, may be vulnerable to warming, due to narrower thermal breadth and sensitivity to higher temperatures, with genetic impoverishment through loss of individuals likely to further reduce population viability.

Latitudinal limit not a cold limit: Cold temperatures do not constrain an endangered tree species at its northern edge

AbstractAimA strong influence of climate on species’ range is often assumed, and forms the basis for projecting many future range shifts with changing climate. Particularly at poleward latitudinal or elevational edges, abiotic conditions are thought to play a major role in limiting distributions. We estimated the roles of climate and landscape features in shaping habitat at the northern distributional edge of a critically threatened mountain tree species.LocationBritish Columbia and Alberta, Canada.TaxonPinus albicaulis (Engelm.)MethodsWe used a hierarchical Bayesian model (HBM) to combine multiple scales, sources, qualities and types of species data. We jointly examined the climate influence on occupancy across scales, and on juvenile and adult abundance to quantify habitat quality at two life history stages.ResultsWe found that cold temperature was the strongest predictor of whitebark pine occurrence at regional scales, with colder areas being better (i.e. the sign was negative). Occupancy at local scales was best predicted by low growing degree‐days and declining precipitation as snow. These relationships with occupancy across scales indicate that suitable climatic and topographic habitats currently exist beyond the northern edge of whitebark pine's current range. We found high adult abundance was predicted in sunny, cool habitats with little climatic drought, whereas high juvenile densities were associated with higher precipitation as snow and more climatic drought.Main conclusionsThe negative relationship to temperature and the ample suitable habitats predicted to exist poleward of the current species’ range limit indicates whitebark pine is not limited by cold temperatures. We suggest that not all species’ ranges are cold limited at high latitudes or elevations. For whitebark pine this means warming temperatures may not directly result in a northern range expansion as a result of warming habitat.

Shaping up for stress: Physiological flexibility is key to survivorship in a habitat-forming macroalga

Organisms from all domains of life can have highly variable morphologies, with this plasticity suggested to increase fitness and survivability under stressful conditions. Predicting how organisms will adapt to environmental change requires an understanding of how variable morphologies perform under environmental stress. Morphological plasticity has been documented within marine macroalgae inhabiting environmental gradients, however the functional consequences of this variation has been rarely tested. In this study, form-function was assessed in the habitat-forming, intertidal macroalga Hormosira banksii. Morphological variation was quantified on two spatial scales (tidal gradient versus latitudinal gradient) and the performance tested (relative water content and photosynthetic efficiency) of morphological variants during heat and desiccation stress. At regional scales, individuals at the warm distributional edge were overall smaller in size, and had smaller vesicles (higher surface area to volume ratio; SA:VOL) than those from central populations. At local scales, individuals high on the shore were generally shorter and had larger vesicles than those low on the shore. Vesicle morphology (SA:VOL) was found to predict relative water content and photosynthetic performance during desiccation and rehydration. Differences in SA:VOL of vesicles between heights on the shore may reflect water requirements needed to maintain tissue hydration for photosynthesis during low tide. Warm-edge populations showed increased thermal sensitivity as indicated by decreased photosynthetic yield of PSII and delays in recovery after desiccation. Sensitivities to higher temperatures amongst warm-edge populations are potentially due to smaller fluctuations in regional temperatures as well as their morphology. This study provides a mechanistic understanding of the morphological variation among H. banksii populations. It suggests that H. banksii has a high degree of morphological plasticity reflecting local climate, topography and environmental conditions, with this morphological variation having functional consequences. Morphological variation across local and regional scales will be important for resilience of this species to future climate warming.

Evidence of hybrid origin for Tachyphonus nattereri Pelzeln, 1870 (Aves: Thraupidae).

Tachyphonus nattereri is one of the three "mystery birds" collected almost 200 years ago by J. Natterer in Brazil whose validity has been disputed in many publications over the last century. Known only from the male holotype and perhaps a female specimen without type status, it is currently treated as either a valid species, a subspecies or a doubtful taxon (an extreme of variation or an aberrant-colored individual). We reviewed the taxonomy of T. nattereri based on fieldwork near the type locality and a careful examination of the holotype and related museum specimens. Our extensive fieldwork revealed that no natural population found in the type locality matches the phenotype of T. nattereri. We found that the holotype is intermediate in plumage, morphometry and body shape between Tachyphonus cristatus and T. luctuosus, two sympatric species locally scarce and at their distributional edge around the type locality. We, therefore, suggest that T. nattereri is a hybrid. On the other hand, the purported female falls within the variation found in Tachyphonus c. madeirae, of which it probably represents an extreme phenotype.

Facilitative Effect of a Generalist Herbivore on the Recovery of a Perennial Alga: Consequences for Persistence at the Edge of Their Geographic Range.

Understanding the impacts of consumers on the abundance, growth rate, recovery and persistence of their resources across their distributional range can shed light on the role of trophic interactions in determining species range shifts. Here, we examined if consumptive effects of the intertidal grazer Scurria viridula positively influences the abundance and recovery from disturbances of the alga Mazzaella laminarioides at the edge of its geographic distributions in northern-central Chilean rocky shores. Through field experiments conducted at a site in the region where M. laminarioides overlaps with the polar range edge of S. viridula, we estimated the effects of grazing on different life stages of M. laminarioides. We also used long-term abundance surveys conducted across ~700 km of the shore to evaluate co-occurrence patterns of the study species across their range overlap. We found that S. viridula had positive net effects on M. laminarioides by increasing its cover and re-growth from perennial basal crusts. Probability of occurrence of M. laminarioides increased significantly with increasing density of S. viridula across the range overlap. The negative effect of S. viridula on the percentage cover of opportunistic green algae—shown to compete for space with corticated algae—suggests that competitive release may be part of the mechanism driving the positive effect of the limpet on the abundance and recovery from disturbance of M. laminarioides. We suggest that grazer populations contribute to enhance the abundance of M. laminarioides, facilitating its recolonization and persistence at its distributional range edge. Our study highlights that indirect facilitation can determine the recovery and persistence of a resource at the limit of its distribution, and may well contribute to the ecological mechanisms governing species distributions and range shifts.

Collapse of the echinoid Paracentrotus lividus populations in the Eastern Mediterranean--result of climate change?

The European purple sea urchin (Paracentrotus lividus) is considered to be a key herbivore throughout its distribution range—North-East Atlantic and Mediterranean Sea. It was also abundant in its eastern distributional edge, on rocky habitats of the coastline of Israel, but its populations have recently collapsed, and today it is an extremely rare species in the region. Field and laboratory experiments, that were carried out in order to examine the impact of the recent sea surface temperature rise in the Eastern Mediterranean, showed massive urchin mortality when temperatures crossed 30.5 °C before reaching peak summer values. These results suggest that elevated seawater temperatures in recent years may be a main cause for the disappearance of P. lividus from the southeast Mediterranean Sea, which may indicate distributional range contraction in this region.

Metabolic cold adaptation and aerobic performance of blue mussels (Mytilus edulis) along a temperature gradient into the High Arctic region

The blue mussel (Mytilus edulis) has recently expanded its northern distribution in the Arctic and is therefore considered to be a sensitive indicator of climate changes in this region. In this study, we compared aerobic performance of blue mussels from High Arctic, Subarctic and temperate populations at different temperatures. Standard metabolic rates (SMR) and active metabolic rates (AMR) were measured for each population, and absolute (AMR − SMR) and factorial ( $$\frac{AMR}{SMR}$$ ) scopes were calculated. Blue mussels from the temperate population had the lowest Q10 (= 1.8) and the largest thermal window (−1 to 25 °C), whereas Q10 values in the Arctic populations were 1.9 (Subarctic) and 2.3 (High Arctic), with a thermal window of −1 to 21 °C. Aerobic scope increased with rising temperatures, reaching a maximum at 14 °C (temperate) and 7 °C (Subarctic and High Arctic, respectively), after which a decrease was observed at temperatures exceeding 14 °C. At low temperatures (−1 °C), the average SMR of the High Arctic population was 93 % higher than that of the temperate population and 22 % higher than that of the Subarctic population. Combined, our results demonstrate physiological adaptation and plasticity of blue mussels across latitudes spanning from 56 to 77oN. This indicates that low ocean temperature per se does not constrain metabolic activity of Mytilus in the Arctic; rather, we speculate that maturation of reproductive tissues, larval supply and annual energy budgets are the most relevant factors influencing Mytilus populations near their northern distributional edge in the Arctic.

Thermal stress resistance of the brown alga Fucus serratus along the North-Atlantic coast: Acclimatization potential to climate change

Seaweed-dominated communities are predicted to disappear south of 45° latitude on North-Atlantic rocky shores by 2200 because of climate change. The extent of predicted habitat loss, however, could be mitigated if the seaweeds' physiology is sufficiently plastic to rapidly acclimatize to the warmer temperatures. The main objectives of this study were to identify whether the thermal tolerance of the canopy-forming seaweed Fucus serratus is population-specific and where temperatures are likely to exceed its tolerance limits in the next 200 years. We measured the stress response of seaweed samples from four populations (Norway, Denmark, Brittany and Spain) to common-garden heat stress (20°C–36°C) in both photosynthetic performance and transcriptomic upregulation of heat shock protein genes. The two stress indicators did not correlate and likely measured different cellular components of the stress response, but both indicators revealed population-specific differences, suggesting ecotypic differentiation. Our results confirmed that thermal extremes will regularly reach physiologically stressful levels in Brittany (France) and further south by the end of the 22nd century. Although heat stress resilience in photosynthetic performance was higher at the species' southern distributional edge in Spain, the hsp expression pattern suggested that this edge-population experienced reduced fitness and limited responsiveness to further stressors. Thus, F. serratus may be unable to mitigate its predicted northward shift and may be at high risk to lose its center of genetic diversity and adaptability in Brittany (France). As it is an important intertidal key species, the disappearance of this seaweed will likely trigger major ecological changes in the entire associated ecosystem.

Morphological, phytochemical and genetic variation in mixed stands and a hybrid swarm of Senecio germanicus and S. ovatus (Compositae, Senecioneae)

Mixed stands of Senecio ovatus subsp. ovatus and S. germanicus subsp. germanicus occur in the colline belt of central and eastern Europe. The latter species is adapted to more continental climate conditions and shows a later flowering time (August–September) than the widespread S. ovatus (July–August) that grows in more oceanic climates. We have surveyed 253 plants from 15 populations north of Regensburg (south-eastern Germany) using 16 qualitative and quantitative morphological characters and molecular markers [amplified fragment length polymorphisms (AFLP)] to detect introgressive hybridisation between these two species. Both multivariate statistical analyses based on morphological characters and the Bayesian clustering based on AFLP fingerprint data show that in most populations under study the two species form distinct entities and do not hybridise with each other. However, in one population from the Upper Palatine Forest a high number of intermediate individuals were found. A more detailed genetic (AFLP) and phytochemical (pyrrolizidine alkaloid, PA) analysis based on 125 individuals from this hybrid swarm indicated that these intermediate individuals are backcrosses towards S. germanicus. It is shown that the two species differ considerably concerning the qualitative and quantitative PA patterns and that backcrossed individuals either show an additive PA pattern or a PA pattern similar to S. germanicus, while in quantitative respects all of these individuals are approaching S. germanicus. These findings are discussed in terms of differential selection regimes influencing the fitness of pure and hybrid plants in an area which is an eco-climatological optimum for the more oceanic S. ovatus but which forms a distributional edge for the more continental S. germanicus.

Seasonal source–sink dynamics at the edge of a species' range

The roles of dispersal and population dynamics in determining species' range boundaries recently have received theoretical attention but little empirical work. Here we provide data on survival, reproduction, and movement for a Virginia opossum (Didelphis virginiana) population at a local distributional edge in central Massachusetts (USA). Most juvenile females that apparently exploited anthropogenic resources survived their first winter, whereas those using adjacent natural resources died of starvation. In spring, adult females recolonized natural areas. A life-table model suggests that a population exploiting anthropogenic resources may grow, acting as source to a geographically interlaced sink of opossums using only natural resources, and also providing emigrants for further range expansion to new human-dominated landscapes. In a geographical model, this source-sink dynamic is consistent with the local distribution identified through road-kill surveys. The Virginia opossum's exploitation of human resources likely ameliorates energetically restrictive winters and may explain both their local distribution and their northward expansion in unsuitable natural climatic regimes. Landscape heterogeneity, such as created by urbanization, may result in source-sink dynamics at highly localized scales. Differential fitness and individual dispersal movements within local populations are key to generating regional distributions, and thus species ranges, that exceed expectations.

Effects of Vegetated Field Borders on Arthropods in Cotton Fields in Eastern North Carolina

The influence, if any, of 5m wide, feral, herbaceous field borders on pest and beneficial arthropods in commercial cotton, Gossypium hirsutum (L.) (Malvales: Malvaceae), fields was measured through a variety of sampling techniques over three years. In each year, 5 fields with managed, feral vegetation borders and five fields without such borders were examined. Sampling was stratified from the field border or edge in each field in an attempt to elucidate any edge effects that might have occurred. Early season thrips populations appeared to be unaffected by the presence of a border. Pitfall sampling disclosed no differences in ground-dwelling predaceous arthropods but did detect increased populations of crickets around fields with borders. Cotton aphid (Aphis gossypii Glover) (Hemiptera: Aphididae) populations were too low during the study to adequately assess border effects. Heliothines, Heliothis virescens (F.) and Helicoverpa zea (Boddie) (Lepidoptera: Noctuidae), egg numbers and damage rates were largely unaffected by the presence or absence of a border, although in one instance egg numbers were significantly lower in fields with borders. Overall, foliage-dwelling predaceous arthropods were somewhat more abundant in fields with borders than in fields without borders. Tarnished plant bugs, Lygus lineolaris (Palisot de Beauvois) (Heteroptera: Miridae) were significantly more abundant in fields with borders, but stink bugs, Acrosternum hilare (Say), and Euschistus servus (Say) (Hemiptera: Pentatomidae) numbers appeared to be largely unaffected by border treatment. Few taxa clearly exhibited distributional edge effects relative to the presence or absence of border vegetation. Field borders like those examined in this study likely will have little impact on insect pest management in cotton under current insect management regimens.

Demographic responses of Virginia opossums to limitation at their northern boundary

The precise response of a population at its distributional edge to the limiting extrinsic factor should be mediated by the demography of the species. We applied this principle to understanding the northern distributional potential of the Virginia opossum (Didelphis virginiana Kerr, 1792). We reviewed the literature for demographic data that we then used to build model populations. Juvenile over-winter survival was adjusted to determine the survival necessary for a stable population. To put the results in the context of life-history strategy and ecological niche, we built models for two other medium-sized mammals with similar distributions, the raccoon (Procyon lotor (L., 1758)) and the muskrat (Ondatra zibethicus (L., 1766)). Northern raccoon populations may sustain juvenile winter survival rates of <0.50 because adult females live to reproduce in multiple years. Muskrat juveniles may need a winter survival rate of only 0.40 in average years because reproduction is very high. In contrast, young northern opossums need a survival rate of 0.81 over winter to compensate for low prewinter survival. Raccoons and muskrats, through different life-history strategies, should be able to expand their northern distribution to the winter-induced physiological limit. However, opossum populations should fail before the average individual physiological limit is reached.

Microsatellite heterozygosity, fitness and demography in natterjack toads Bufo calamita

AbstractRelationships between mean individual fitness and genetic heterozygosity remain controversial because multiple studies over many years have yielded inconsistent results. However, molecular measurements of genetic variation have mostly been based on allozyme markers, which may have substantial limitations due to their relative insensitivity and occasionally also to the effects of natural selection. We investigated variation in the number of tandem dinucleotide repeats at eight polymorphic microsatellite loci in 38 natterjack toad, Bufo calamita, populations that varied in census size, degree of isolation and distance from the distributional range edge. We also measured fitness attributes, notably larval survival and growth rates, under standardized conditions using samples from six of these populations. The results indicated that larval growth rates were positively and strongly correlated with expected heterozygosity (He) across the microsatellite loci. He was in turn highest in large populations with minimal isolation, but low near the biogeographical range edge irrespective of population size. Larvae from the smallest and most isolated natterjack population exhibited particularly low fitness and heterozygosity. Our results suggest that microsatellite loci may provide a valuable new approach in studies of links between fitness and heterozygosity.