Low-latitude Populations Research Articles

Transcriptomic responses and physiological changes to cold stress among natural populations provide insights into local adaptation of weeping forsythia

Genetic mechanisms of species local adaptation are an emerging topic of great interest in evolutionary biology and molecular ecology. In this study, we compared the changes of physiological and phenotypic indexes and gene expression of four weeping forsythia populations under cold stress through a common garden experiment. Physiological and phenotypic results showed that there were differences in cold tolerance among populations. cold tolerance of high the latitude population (HBWZ) was the strongest, followed by the middle latitude population (SXWL), while the low latitude populations (SXHM) and (SXLJ) expressed the weakest cold tolerance. We identified significant differences in gene expression of cold tolerance related pathways and ontologies, including genes of oxylipin and isoquinoline alkaloid biosynthetic process, galactose, tyrosine and unsaturated fatty acids metabolism, among these populations under the same experimental temperature treatments. Even under the same degree of stress, there were notable differences in gene expression among natural populations. In this study, we present a working model of weeping forsythia populations which evolved in the context of different intensities of cold stress. Our study provides new insights for comprehending the genetic mechanisms of local adaptation for non-model species.

Diminished warming tolerance and plasticity in low-latitude populations of a marine gastropod.

Models of species response to climate change often assume that physiological traits are invariant across populations. Neglecting potential intraspecific variation may overlook the possibility that some populations are more resilient or susceptible than others, creating inaccurate predictions of climate impacts. In addition, phenotypic plasticity can contribute to trait variation and may mediate sensitivity to climate. Quantifying such forms of intraspecific variation can improve our understanding of how climate can affect ecologically important species, such as invasive predators. Here, we quantified thermal performance (tolerance, acclimation capacity, developmental traits) across seven populations of the predatory marine snail (Urosalpinx cinerea) from native Atlantic and non-native Pacific coast populations in the USA. Using common garden experiments, we assessed the effects of source population and developmental acclimation on thermal tolerance and developmental traits of F1 snails. We then estimated climate sensitivity by calculating warming tolerance (thermal tolerance - habitat temperature), using field environmental data. We report that low-latitude populations had greater thermal tolerance than their high latitude counterparts. However, these same low-latitude populations exhibited decreased thermal tolerance when exposed to environmentally realistic higher acclimation temperatures. Low-latitude native populations had the greatest climate sensitivity (habitat temperatures near thermal limits). In contrast, invasive Pacific snails had the lowest climate sensitivity, suggesting that these populations are likely to persist and drive negative impacts on native biodiversity. Developmental rate significantly increased in embryos sourced from populations with greater habitat temperature but had variable effects on clutch size and hatching success. Thus, warming can produce widely divergent responses within the same species, resulting in enhanced impacts in the non-native range and extirpation in the native range. Broadly, our results highlight how intraspecific variation can alter management decisions, as this may clarify whether management efforts should be focused on many or only a few populations.

Diversity Patterns of Bermuda Grass along Latitudinal Gradients at Different Temperatures in Southeastern China.

Cynodon dactylon (L.) Pers. (common Bermuda grass) has a limited capacity to grow at low temperatures, which limits its geographical range. Exploring its evolutionary relationship across different environmental gradients is necessary to understand the effects of temperature change on the genetics of common Bermuda grass. In this study, high-throughput transcriptome sequencing was performed on 137 samples of C. dactylon from 16 latitudinal gradients to explore the differential molecular markers and analyze genetic diversity and structure along latitudinal gradients at different temperatures. We primarily sampled more high-quality single nucleotide polymorphisms (SNPs) from populations at lower and middle latitudes. Greater intraspecific genetic variation at each level of temperature treatment could be due to factors such as wind pollination and asexual breeding. Populations of C. dactylon at high latitudes differed from populations at middle and low latitudes, which was supported by a principal component analysis (PCA) and genetic structure analysis, performed at different temperatures. We observed more genetic variation for low-latitude populations at 5 °C, according to an analysis of three phylogenetic trees at different temperature levels, suggesting that low temperatures affected samples with low cold resistance. Based on the results of phylogenetic analysis, we found that samples from high latitudes evolved earlier than most samples at low latitudes. The results provide a comprehensive understanding of the evolutionary phenomenon of landscape genetics, laying the groundwork for future structural and comparative genomic studies of C. dactylon.

D-lactate production in the acorn barnacleBalanus glandula(Darwin, 1854) (Cirripedia: Balanidae) under emersion stress

AbstractAnaerobic metabolism is an important response to stress in many organisms. Intertidal species often face heat stress during low tide. Balanus glandula (Darwin, 1854) is a high-shore intertidal barnacle common to the Pacific that experiences prolonged low-tide air exposure. It is not known whether B. glandula uses anaerobic metabolism during emersion, or if its use varies by latitude. We measured low tide D-lactate production in two US west coast populations of B. glandula separated by 14 degrees of latitude. We exposed barnacles to seven low-tide air temperatures (10, 15, 20, 25, 30, 35, and 38 °C) for which aerobic respiration has been previously measured. Our northern population of B. glandula increased D-lactate production at high air temperatures where aerobic metabolic depression is known to occur, indicating sublethal stress. In contrast, our southern population showed little increase in D-lactate over the same temperature range, coincident with high aerobic respiration across those temperatures. In a second experiment, we quantified D-lactate at 1, 2, and 6 hours post-emersion for northern B. glandula exposed to either a 10 or 38 °C low tide, to measure their potential lactate usage. While D-lactate was elevated at 38 °C compared to the 10 °C control immediately following low tide exposure, it dropped to control levels, and was likely excreted, within 1 hour of re-immersion. Our results suggest that the low latitude population of B. glandula may be more resilient to climate change than its high latitude counterpart in the absence of adaptation, which has strong implications for species distribution.

Two experimental designs generate contrasting patterns of behavioral differentiation along a latitudinal gradient in Lestes sponsa-Common-garden not so common after all?

Understanding why and how behavioral profiles differ across latitudes can help predict behavioral responses to environmental change. The first response to environmental change that an organism exhibits is commonly a behavioral response. Change in one behavior usually results in shifts in other correlated behaviors, which may adaptively or maladaptively vary across environments and/or time. However, one important aspect that is often neglected when studying behavioral expressions among populations is if/how the experimental design might affect the results. This is unfortunate since animals often plastically modify their behavior to the environment, for example, rearing conditions. We studied behavioral traits and trait correlations in larvae of a univoltine damselfly, Lestes sponsa, along its latitudinal distribution, spreading over 3,300 km. We compared behavioral profiles among larvae grown in two conditions: (a) native temperatures and photoperiods or (b) averaged constant temperatures and photoperiods (common‐garden). We hypothesized latitudinal differences in behavioral traits regardless of the conditions in which larvae were grown, with northern populations expressing higher activity, boldness, and foraging efficiency. When grown in native conditions, northern larvae were bolder, more active and more effective in prey capture than central and low latitude populations, respectively, as well as showed the strongest behavioral correlations. In contrast, larvae reared in common‐garden conditions showed no differences between regions in both individual traits and trait correlations. The results suggest different selective pressures acting on the studied traits across populations, with environment as a central determinant of the observed trait values. Common‐garden designed experiments may evoke population‐dependent levels of plastic response to the artificial conditions and, hence, generate results that lack ecological relevance when studying multi‐population differences in behavior.

Behavioral constraints on local adaptation and counter-gradient variation: Implications for climate change.

Resource allocation to growth, reproduction, and body maintenance varies within species along latitudinal gradients. Two hypotheses explaining this variation are local adaptation and counter‐gradient variation. The local adaptation hypothesis proposes that populations are adapted to local environmental conditions and are therefore less adapted to environmental conditions at other locations. The counter‐gradient variation hypothesis proposes that one population out performs others across an environmental gradient because its source location has greater selective pressure than other locations. Our study had two goals. First, we tested the local adaptation and counter‐gradient variation hypotheses by measuring effects of environmental temperature on phenotypic expression of reproductive traits in the burying beetle, Nicrophorus orbicollis Say, from three populations along a latitudinal gradient in a common garden experimental design. Second, we compared patterns of variation to evaluate whether traits covary or whether local adaptation of traits precludes adaptive responses by others. Across a latitudinal range, N. orbicollis exhibits variation in initiating reproduction and brood sizes. Consistent with local adaptation: (a) beetles were less likely to initiate breeding at extreme temperatures, especially when that temperature represents their source range; (b) once beetles initiate reproduction, source populations produce relatively larger broods at temperatures consistent with their local environment. Consistent with counter‐gradient variation, lower latitude populations were more successful at producing offspring at lower temperatures. We found no evidence for adaptive variation in other adult or offspring performance traits. This suite of traits does not appear to coevolve along the latitudinal gradient. Rather, response to selection to breed within a narrow temperature range may preclude selection on other traits. Our study highlights that N. orbicollis uses temperature as an environmental cue to determine whether to initiate reproduction, providing insight into how behavior is modified to avoid costly reproductive attempts. Furthermore, our results suggest a temperature constraint that shapes reproductive behavior.

Identifying global hotspots of avian trailing-edge population diversity

Climate change is causing the ranges of many species to shift poleward and to higher elevations. Trailing-edge populations near the low-latitude edge of a shifting range are predicted to be at high risk of climate-induced extinction, but conservation efforts are hindered by a lack of information about the global distribution of trailing-edge populations. We used a large spatial dataset on the ranges of nearly all extant avian species to identify potential hotspots where trailing-edge populations represent a large proportion of the total avifauna. We identified potential trailing-edge population hotspots by isolating and overlaying low latitude regions of species’ ranges, and computing the proportion of total species richness in a location comprised of low-latitude populations. We identified potential hotspots on all continents other than Antarctica. Potential trailing-edge population diversity was highest near the equator, low-latitude margins of mountain ranges, desert edges, and along coastlines. Because a potential trailing-edge population hotspot might not be an actual trailing-edge population hotspot if the low-latitude populations are not declining, information on population trends is necessary for confirmation. As a case study, we focused on one of the identified hotspots, the Southern Appalachian Mountains, where our analysis indicated that 30 bird species have potential trailing-edge populations. Even though more population studies have been conducted in the Appalachian Mountains than in most of the other potential hotspots that we identified, there was insufficient information available from the high elevations where these species occur to make strong inferences about population declines. Our research highlights the need for a concerted effort to gather more information about population trends in the regions we identified as potential hotspots of trailing-edge population diversity.

Investigating reindeer pastoralism and exploitation of high mountain zones in northern Mongolia through ice patch archaeology.

In interior Eurasia, high mountain zones are crucial to pastoral subsistence, providing seasonally productive pastures and abundant wild resources. In some areas of northern Mongolia, mountainous tundra zones also support a low-latitude population of domestic reindeer herders–a lifestyle whose origins are poorly characterized in the archaeological record of early Mongolia. Traditionally, reindeer pastoralists make significant seasonal use of munkh mus (eternal ice) for their domestic herds, using these features to cool heat-stressed animals and provide respite from insect harassment. In recent years, many of these features have begun to melt entirely for the first time, producing urgent threats to traditional management techniques, the viability of summer pastures, and reindeer health. The melting ice is also exposing fragile organic archaeological materials that had previously been contained in the patch. We present the results of horseback survey of ice patches in Baruun Taiga special protected area, providing the first archaeological insights from the region. Results reveal new evidence of historic tool production and wild resource use for fishing or other activities, and indicate that ice patches are likely to contain one of the few material records of premodern domestic reindeer use in Mongolia and lower Central Asia. The area’s ancient ice appears to be rapidly melting due to changing climate and warming summer temperatures, putting both cultural heritage and traditional reindeer herding at extreme risk in the years to come.

Metabolic rates from Bluntnose minnow (Pimephales notatus) populations at lower latitudes are more sensitive to changes in temperature than populations at higher latitudes

AbstractPredicting the potential effects of changes in climate on freshwater species requires an understanding of the relationships between physiological traits and environmental conditions among populations. While water temperature is a primary factor regulating metabolic rates in freshwater ectotherms, how metabolic rates vary across the species range is unclear. In addition, photoperiod has also been hypothesised to influence metabolic rates in freshwater taxa based on seasonal changes in activity rates. Using an experimental approach, we investigated whether variation in routine metabolic rate (RMR) and sensitivity of RMR to changes in temperature are correlated with local thermal regimes, photoperiods and body mass among ten populations across the geographic range of the Bluntnose minnow (Pimephales notatus), a North American freshwater fish species. Routine metabolic rate data were collected from populations acclimatised to three temperature treatments (9, 18 and 27°C) and correlated with water temperature and photoperiod estimates at collection locations for each population. Routine metabolic rate was negatively correlated with minimum photoperiod at 9°C, negatively correlated with weekly high temperature at 18°C and positively correlated with weekly high temperature at 27°C. Body mass was also a predictor of RMR at each temperature treatment. Thermal sensitivity of RMR was positively correlated with weekly high temperature, indicating that individuals from warmer low latitude populations experienced greater sensitivity of RMR to changes in temperature than individuals from cooler high latitude populations. These results indicate differential responses among populations to variation in temperature and suggest the importance of recognising this variation when characterising responses of freshwater taxa to increases in water temperature.

Integrating patterns of thermal tolerance and phenotypic plasticity with population genetics to improve understanding of vulnerability to warming in a widespread copepod.

Differences in population vulnerability to warming are defined by spatial patterns in thermal adaptation. These patterns may be driven by natural selection over spatial environmental gradients, but can also be shaped by gene flow, especially in marine taxa with high dispersal potential. Understanding and predicting organismal responses to warming requires disentangling the opposing effects of selection and gene flow. We begin by documenting genetic divergence of thermal tolerance and developmental phenotypic plasticity. Ten populations of the widespread copepod Acartia tonsa were collected from sites across a large thermal gradient, ranging from the Florida Keys to Northern New Brunswick, Canada (spanning over 20° latitude). Thermal performance curves (TPCs) from common garden experiments revealed local adaptation at the sampling range extremes, with thermal tolerance increasing at low latitudes and decreasing at high latitudes. The opposite pattern was observed in phenotypic plasticity, which was strongest at high latitudes. No relationship was observed between phenotypic plasticity and environmental variables. Instead, the results are consistent with the hypothesis of a trade-off between thermal tolerance and the strength of phenotypic plasticity. Over a large portion of the sampled range, however, we observed a remarkable lack of differentiation of TPCs. To examine whether this lack of divergence is the result of selection for a generalist performance curve or constraint by gene flow, we analyzed cytochrome oxidase I mtDNA sequences, which revealed four distinct genetic clades, abundant genetic diversity, and widely distributed haplotypes. Strong divergence in thermal performance within genetic clades, however, suggests that the pace of thermal adaptation can be relatively rapid. The combined insight from the laboratory physiological experiments and genetic data indicate that gene flow constrains differentiation of TPCs. This balance between gene flow and selection has implications for patterns of vulnerability to warming. Taking both genetic differentiation and phenotypic plasticity into account, our results suggest that local adaptation does not increase vulnerability to warming, and that low-latitude populations in general may be more vulnerable to predicted temperature change over the next century.

Latitude-associated evolution and drivers of thermal response curves in body stoichiometry.

Trait-based studies are needed to understand the plastic and genetic responses of organisms to warming. A neglected organismal trait is elemental composition, despite its potential to cascade into effects on the ecosystem level. Warming is predicted to shape elemental composition through shifts in storage molecules associated with responses in growth, body size and metabolic rate. Our goals were to quantify thermal response patterns in body composition and to obtain insights into their underlying drivers and their evolution across latitudes. We reconstructed the thermal response curves (TRCs) for body elemental composition [C (carbon), N (nitrogen) and the C:N ratio] of damselfly larvae from high- and low-latitude populations. Additionally, we quantified the TRCs for survival, growth and development rates and body size to assess local thermal adaptation, as well as the TRCs for metabolic rate and key macromolecules (proteins, fat, sugars and cuticular melanin and chitin) as these may underlie the elemental TRCs. All larvae died at 36°C. Up to 32°C, low-latitude larvae increased growth and development rates and did not suffer increased mortality. Instead, growth and development rates of high-latitude larvae were lower and levelled off at 24°C, and mortality increased at 32°C. This latitude-associated thermal adaptation pattern matched the 'hotter-is-better' hypothesis. With increasing temperatures, low-latitude larvae decreased C:N, while high-latitude larvae increased C:N. These patterns were driven by associated changes in N contents, while C contents did not respond to temperature. Consistent with the temperature-size rule and the thermal melanism hypothesis, body size and melanin levels decreased with warming. While all traits and associated macromolecules (except for metabolic rate that showed thermal compensation) assumed to underlie thermal responses in elemental composition showed thermal plasticity, these were largely independent and none could explain the stoichiometric TRCs. Our results highlight that thermal responses in elemental composition cannot be explained by traditionally assumed drivers, asking for a broader perspective including the thermal dependence of elemental fluxes. Another key implication is that thermal evolution can reverse the plastic stoichiometric thermal responses and hence reverse how warming may shape food web dynamics through changes in body composition at different latitudes.

Metabolomic and Lipidomic Signatures of Cold Tolerance in Bumble Bees

Thermal tolerance is a key physiological trait that often varies among populations and species, and can be a key determinant of current and future geographic distributions. The cellular basis for organismal differences in thermal tolerance is an active area of research, with chemical constituents of the cytosol and cell membrane likely playing important roles in maintaining cell function at low temperatures. For freeze‐avoiding organisms, the abundance and makeup of sugars and related molecules depress the freezing point, reducing the risk of ice formation. At less extreme temperatures, remodeling of membrane lipids can help maintain fluidity (and function) at cold temperatures. We have found striking differences in critical thermal minima (CTmin) of worker bumble bees reared in common‐garden conditions from queens collected at low and high latitudes. We compared metabolomic and lipidomic signatures of these bees to identify key molecules that likely underlie differences in cold tolerance. PCA analyses of GC‐MS data for aqueous abdominal extracts revealed clear separation of low and high latitude populations and reveal higher levels of several key molecules, including L‐proline, in the high latitude population. Lipid extractions and subsequent GC‐MS reveals variation in fatty acid composition of nutrients (triacylglycerols) and of cellular membranes, for which phospholipid composition is also critical for function at low temperatures. This broad spectrum approach to metabolomics and lipidomics allows us to compare a wide range of metabolites between bees differing in cold tolerance, facilitating ongoing research on the mechanistic basis for this physiological trait linked to the ecology of diverse organisms.Support or Funding InformationThis material is based upon work supported by the National Science Foundation under Grant Nos. DEB‐1457659 and RII Track 2‐1826834. Any opinions, findings, and conclusions or recommendations expressed in this material are those of the author(s) and do not necessarily reflect the views of the National Science Foundation.This abstract is from the Experimental Biology 2019 Meeting. There is no full text article associated with this abstract published in The FASEB Journal.

Complex interactions between local adaptation, phenotypic plasticity and sex affect vulnerability to warming in a widespread marine copepod.

Predicting the response of populations to climate change requires an understanding of how various factors affect thermal performance. Genetic differentiation is well known to affect thermal performance, but the effects of sex and developmental phenotypic plasticity often go uncharacterized. We used common garden experiments to test for effects of local adaptation, developmental phenotypic plasticity and individual sex on thermal performance of the ubiquitous copepod, Acartia tonsa (Calanoida, Crustacea) from two populations strongly differing in thermal regimes (Florida and Connecticut, USA). Females had higher thermal tolerance than males in both populations, while the Florida population had higher thermal tolerance compared with the Connecticut population. An effect of developmental phenotypic plasticity on thermal tolerance was observed only in the Connecticut population. Our results show clearly that thermal performance is affected by complex interactions of the three tested variables. Ignoring sex-specific differences in thermal performance may result in a severe underestimation of population-level impacts of warming because of population decline due to sperm limitation. Furthermore, despite having a higher thermal tolerance, low-latitude populations may be more vulnerable to warming as they lack the ability to respond to increases in temperature through phenotypic plasticity.

Temperature variation makes an ectotherm more sensitive to global warming unless thermal evolution occurs.

To assess long-term impacts of global warming on species, there is growing interest in latitudinal intraspecific patterns in thermal adaptation. Yet, while both mean temperatures and daily temperature fluctuations (DTFs) are expected to increase under global warming, latitudinal differences in the effects of DTFs have not been documented. We tested whether low-latitude populations of an ectotherm deal better with greater DTF than high-latitude populations, especially at a high mean temperature close to the optimal temperature for growth where DTF causes exposure to extreme high temperatures. We evaluated the impact of DTFs when assessing the effect of gradual thermal evolution at the high latitude with a space-for-time substitution. We compared effects of both mean temperatures (20 and 24°C) and DTFs (constant=0°C, low=5°C and high=10°C) on growth rates between low-latitude and high-latitude populations of the damselfly Ischnura elegans in a common-garden experiment. DTFs, if anything, reduced growth and were generally stressful as indicated by reductions in body condition, antioxidant defence and metabolic rate, and increases in oxidative damage. Most negative effects of DTFs were only present at a mean of 24°C when too high temperatures were reached during a daily cycle. Notably, while 4°C warming was beneficial in terms of growth rate at both latitudes at a constant temperature regime, this changed in a negative effect at high DTF. Moreover, this modulating effect of the mean temperature by DTF differed between latitudes indicating local thermal adaptation. While 4°C warming at low DTF still caused faster growth in low-latitude larvae, it already slowed growth in high-latitude larvae. This supports the emerging insight that warming would increase growth in high-latitude larvae in the absence of DTF, yet would decrease growth in the more realistic scenarios with DTF. In contrast, a space-for-time substitution approach suggested that under gradual thermal evolution, the evolved high-latitude larvae would no longer suffer a growth reduction in the presence of DTF. Our study provided important proof-of-principle that jointly integrating gradual thermal evolution and the expected increase in DTF generates opposing predictions of effects of global warming on this ectotherm.

Variation in developmental temperature alters adulthood plasticity of thermal tolerance in Tigriopus californicus.

In response to environmental change, organisms rely on both genetic adaptation and phenotypic plasticity to adjust key traits that are necessary for survival and reproduction. Given the accelerating rate of climate change, plasticity may be particularly important. For organisms in warming aquatic habitats, upper thermal tolerance is likely to be a key trait, and many organisms express plasticity in this trait in response to developmental or adulthood temperatures. Although plasticity at one life stage may influence plasticity at another life stage, relatively little is known about this possibility for thermal tolerance. Here, we used locally adapted populations of the copepod Tigriopus californicus to investigate these potential effects in an intertidal ectotherm. We found that low latitude populations had greater critical thermal maxima (CTmax) than high latitude populations, and variation in developmental temperature altered CTmax plasticity in adults. After development at 25°C, CTmax was plastic in adults, whereas no adulthood plasticity in this trait was observed after development at 20°C. This pattern was identical across four populations, suggesting that local thermal adaptation has not shaped this effect among these populations. Differences in the capacities to maintain ATP synthesis rates and to induce heat shock proteins at high temperatures, two likely mechanisms of local adaptation in this species, were consistent with changes in CTmax owing to phenotypic plasticity, which suggests that there is likely mechanistic overlap between the effects of plasticity and adaptation. Together, these results indicate that developmental effects may have substantial impacts on upper thermal tolerance plasticity in adult ectotherms.

Impacts of increased ocean temperatures on a low-latitude coral reef fish – Processes related to oxygen uptake and delivery

Increasing temperatures are expected to significantly affect the physiological performance of ectotherms, particularly in tropical locations. The shape of an organism's thermal reaction norm can provide important information on its capacity to persist under climate change scenarios; however, difficulty lies in choosing a measurable trait that best depicts physiological performance. This study investigated the effects of elevated temperatures on processes related to oxygen uptake and delivery, including oxygen consumption, haematology, and tissue health for a low-latitude population of coral reef damselfish. Acanthochromis polyacanthus were collected from the Torres Strait (10°31–46′S, 142°20–35′E) and maintained at current average ocean temperatures (+0 °C; seasonally cycling), + 1.5 °C and + 3 °C higher than present day temperatures for 10 months. Aerobic performance indicated a limit to metabolic function at + 3 °C (33 °C), following an increase in aerobic capacity at + 1.5 °C (31.5 °C). Neither haematological parameters nor gill morphology showed the same improvement in performance at + 1.5 °C. Gill histopathology provided the first indicator of a decline in organism health, which corresponded with mortality observations from previous research. Findings from this study suggest thermal specialisation in this low-latitude population as well as variation in thermal sensitivity, depending on the physiological trait.

In hot water: sustained ocean warming reduces survival of a low-latitude coral reef fish

Tropical species are predicted to be particularly vulnerable to the impacts of climate change given the relatively narrow thermal range they naturally experience. Within the tropics, average temperature and thermal variation can differ among populations and consequently low-latitude populations may respond differently to increased temperatures than higher latitude tropical populations. In this study, we investigate the long-term effects of climate change relevant temperature increases on commonly measured condition metrics for a low-latitude population of damselfish (Acanthochromis polyacanthus). Adult fish were randomly assigned to one of the three seasonally cycling treatments: (1) current average ocean temperatures for the collection locations, (2) 1.5 °C, or (3) 3 °C higher than current average temperatures. Treatments were maintained for approximately 10 months. At the end of the experimental period, Fulton’s K and hepatosomatic index were calculated for fish from each treatment group and critical thermal limit (CTMax) was measured for a subset of fish at control temperatures. Fish mortality was recorded throughout the experimental period, as well as at the end of the experimental period after the introduction of a secondary exercise stressor. No significant effect of temperature was observed on fish condition (Fulton’s K and hepatosomatic index); however, significant mortality was observed for fish maintained at 3 °C higher than current average temperatures. When a secondary exercise stressor was introduced, significant mortality was also observed at 1.5 °C higher than current average temperatures. Acute exposure to higher temperatures (CTMax) suggested a much higher thermal tolerance for this population than long-term mortality, producing a thermal limit of 37.1 °C compared with a chronic thermal limit of 33 °C. Our results show that some basic measures of fish condition may not be capable of detecting lethal and sub-lethal effects of increased temperature. The results of this study are consistent with the hypothesis that low-latitude species are already living close to their thermal maximum.

Voltinism-associated differences in winter survival across latitudes: integrating growth, physiology, and food intake.

Species that span large latitudinal gradients face strong differences in voltinism and in winter conditions within their range. Latitudinal gradients in winter survival and especially their underlying mechanisms and association with voltinism patterns are poorly studied. We tested in the damselfly Enallagma cyathigerum whether high-latitude populations were better in dealing with the longer winters compared to central- and low-latitude populations and whether this was associated with changes in voltinism. We thereby evaluated whether higher initial levels and/or lower reductions during winter of energy storage (measured as fat content) and investment in immune function [measured as the activity of phenoloxidase (PO)], and/or stronger compensatory responses in food intake contributed to the higher winter survival in high-latitude populations. To this end, we simulated a long high-latitude winter at 4°C under manipulated food conditions. Across food levels, winter survival was highest in Swedish larvae, intermediate in Belgian larvae, and lowest in Spanish larvae, indicating latitude-specific thermal adaptation that could be partly linked to differences in voltinism. The semi-voltine Swedish larvae were growing slower before winter and as a result accumulated the highest fat content and PO activity when the winter started compared to the univoltine, faster growing Belgian, and Spanish larvae. Fat content and PO activity declined during the winter, yet equally across latitudes, and were not buffered by compensatory food intake. Our data identified possible underlying physiological mechanisms of winter survival and support the hypothesis that widespread latitude-associated voltinism shifts may be a selective factor contributing to latitudinal shifts in winter survival.

Demographic compensation does not rescue populations at a trailing range edge

Species' geographic ranges and climatic niches are likely to be increasingly mismatched due to rapid climate change. If a species' range and niche are out of equilibrium, then population performance should decrease from high-latitude "leading" range edges, where populations are expanding into recently ameliorated habitats, to low-latitude "trailing" range edges, where populations are contracting from newly unsuitable areas. Demographic compensation is a phenomenon whereby declines in some vital rates are offset by increases in others across time or space. In theory, demographic compensation could increase the range of environments over which populations can succeed and forestall range contraction at trailing edges. An outstanding question is whether range limits and range contractions reflect inadequate demographic compensation across environmental gradients, causing population declines at range edges. We collected demographic data from 32 populations of the scarlet monkeyflower (Erythranthe cardinalis) spanning 11° of latitude in western North America and used integral projection models to evaluate population dynamics and assess demographic compensation across the species' range. During the 5-y study period, which included multiple years of severe drought and warming, population growth rates decreased from north to south, consistent with leading-trailing dynamics. Southern populations at the trailing range edge declined due to reduced survival, growth, and recruitment, despite compensatory increases in reproduction and faster life-history characteristics. These results suggest that demographic compensation may only delay population collapse without the return of more favorable conditions or the contribution of other buffering mechanisms such as evolutionary rescue.

The Late Pleistocene Canis lupus (Canidae, Mammalia) from Avetrana (Apulia, Italy): reappraisal and new insights on the European glacial wolves

The modern wolf, Canis lupus Linnaeus, 1758, has one of the largest ranges amongst carnivorans, and for this reason it shows local and regional differences for adaptation to a great variety of habitats, ranging from the arctic tundra to the Arabian desert. These differences are particularly evident as wolves follow the Bergmann's ecogeographical rule, with low latitude populations being generally smaller than those living at high latitudes. The fossil record of the modern wolf dates back to the second half of the Middle Pleistocene. The earliest records come from the French site Lunel-Viel 1 and are ascribed to the subspecies Canis lupus lunellensis Bonifay, 1971, and from the Italian Polledrara di Cecanibbio. These Middle Pleistocene forms were generally small in size, slightly larger than the last representative of Canis mosbachensis Soergel, 1925. During the last 400 kyr, European continental environments were profoundly affected by the glacial/interglacial cycles and much evidence suggests a trend toward a size increase for C. lupus. In particular, a number of these European large-sized wolves seem to be typical of cold phases, for example, Canis lupus maximus Boudadi-Maligne, 2012 from the OIS 3-2. The aim of the present study was to describe the morphological and morphometric variability of C. lupus from Avetrana bed 8 in comparison to other populations from northern and southern Italy, as well as from other localities in Europe, to obtain a better understanding of the biochronology, palaeobiogeography and evolution of this large carnivore in the last 125 kyr. The morphological and morphometric analyses pointed out the difference of C. lupus of Avetrana compared to the Apulian and to the Northern Italian wolves but also identified a marked affinity with the C. lupus of the glacial site Cardamone (Apulia), referred to OIS 2. This similarity support the idea of the dispersal of large morphotypes (glacial wolves) during cold phases.