Thermoregulatory behavior and diel activity of yearling winter flounder,Pseudopleuronectes americanus (Walbaum)

Synopsis Sixteen yearling winter flounder, tested individually for 3-day periods in electronic shuttleboxes, voluntarily occupied an 8–27°C range of temperature, with a modal final thermal preferendum of 18.5°C (mean 18.7°C, median 19°C, midpoint 17.5°C, s.d. 1.9°C, S _k + 0.33). The locomotor activity pattern of the yearling fish in the laboratory was markedly nocturnal, with mean hourly nocturnal activity exceeding mean hourly diurnal activity by a factor of 3.4. Maximum activity occurred at 0300 EST, minimum activity at 1400 EST. While activity generally increased with temperature, a local activity minimum occurred at 18.7°C, coinciding with the 24-hour mean final thermal preferendum. Comparison of these laboratory data with previously published field data suggest that behavioral responses to temperature and light play significant roles in determining age- and size-specific differences in seasonal depth stratification and onshore-offshore distributions in this species.     

Thermoregulatory behavior of juvenile cromileptes altivelis (serranidae), a tropical marine fish

Ten juvenile Cromileptes altivelis were tested individually for 3-day periods in electronic shuttleboxes to measure their thermoregulatory behavior. The fish voluntarily occupied a 21–27°C span, out of a potentially available 0–50°C range. The mean final thermal preferendum was 24.5°C for all individuals. The thermoregulatory performance of this species is similar to that of cool temperate freshwater fishes.     

Integrating metabolic performance,thermal tolerance,and plasticity enables for more accurate predictions on species vulnerability to acute and chronic effects of global warming

Predicting species vulnerability to global warming requires a comprehensive, mechanistic understanding of sublethal and lethal thermal tolerances. To date, however, most studies investigating species physiological responses to increasing temperature have focused on the underlying physiological traits of either acute or chronic tolerance in isolation. Here we propose an integrative, synthetic approach including the investigation of multiple physiological traits (metabolic performance and thermal tolerance), and their plasticity, to provide more accurate and balanced predictions on species and assemblage vulnerability to both acute and chronic effects of global warming. We applied this approach to more accurately elucidate relative species vulnerability to warming within an assemblage of six caridean prawns occurring in the same geographic, hence macroclimatic, region, but living in different thermal habitats. Prawns were exposed to four incubation temperatures (10, 15, 20 and 25 °C) for 7 days, their metabolic rates and upper thermal limits were measured, and plasticity was calculated according to the concept of Reaction Norms, as well as Q₁₀ for metabolism. Compared to species occupying narrower/more stable thermal niches, species inhabiting broader/more variable thermal environments (including the invasive Palaemon macrodactylus) are likely to be less vulnerable to extreme acute thermal events as a result of their higher upper thermal limits. Nevertheless, they may be at greater risk from chronic exposure to warming due to the greater metabolic costs they incur. Indeed, a trade‐off between acute and chronic tolerance was apparent in the assemblage investigated. However, the invasive species P. macrodactylus represents an exception to this pattern, showing elevated thermal limits and plasticity of these limits, as well as a high metabolic control. In general, integrating multiple proxies for species physiological acute and chronic responses to increasing temperature helps providing more accurate predictions on species vulnerability to warming.     

Effect of temperature on locomotor activity in the goldfish (Carassius auratus) and the bluegill (Lepomis macrochirus): Presence of an ‘activity well’ in the region of the final preferendum

Goldfish (Carassius auratus) and bluegill sunfish (Lepomis macrochirus) were placed in aquaria where their locomotor activity was monitored by photocells, and tested at various acclimation temperatures over a range encompassing their final thermal preferenda. Activity was pooled over 24-hour periods to eliminate any circadian rhythm effects. Both species exhibited an activity well of reduced locomotor activity in the region of the final preferendum. Goldfish, tested either singly or in groups of 2–5 individuals, exhibited a social-interaction effect which became more pronounced at higher temperatures. These results are discussed in relation to a thermokinetic interpretation of thermo-regulatory behavior in fishes, and to the correspondence between thermal preferenda and thermal optima.     

Temperature selection and critical thermal maxima of the fantail darter,Etheostoma flabellare,and johnny darter,E. nigrum,related to habitat and season

Synopsis Riffle dwelling fantail darters (Etheostoma flabellare) selected lower temperatures in winter (19.3°C) compared to pool dwelling johnny darters (E. nigrum; 22.0°C. A similar trend was evident in summer tests (fantail darters, 20.3°C; johnny darters, 22.9°C). Summer tested animals selected higher temperatures than winter tested animals maintained at the same acclimation temperature and photoperiod. When tested together in the same gradient, both species appeared not to thermoregulate, but tended to avoid each other. Critical thermal maxima (CTMax) did not differ between seasons for either species (fantail darters, 31.1°C winter, 31.3°C summer; johnny darters, 30.9°C winter, 30.5°C summer). Differences in the thermal responses of these darters correlated with differences in their respective habitats.     

Temperature preferences and oxygen consumption of three species of sculpin (Cottus) from the Pit River drainage,California

Synopsis I examined the temperature preferences and routine metabolic rates of Pit sculpin, Cottius pitensis, marbled sculpin, C. klamathensis macrops, and rough sculpin, C. asperrimus, of the Pit River drainage of California to determine if the distributional patterns of these species can be explained on the basis of physiological or behavioral responses to temperature. The routine metabolic rates of these species did not increase significantly between 10 and 15°C, indicating an area of thermal compensation. Metabolic rates then rapidly increased between 15 and 20°C (Q₁₀ values>4.0) followed by little increase between 20 and 25°C (Q₁₀ values >2.0), indicating another area of thermal compensation. The final temperature preferenda of Pit, marbled and rough sculpin were 11.2, 12.1 and 13.5°C, respectively. Marbled and rough sculpin appear to be more stenothermal than Pit sculpin. At acclimation temperatures of 10, 15 and 20°C the acute preferred temperatures of marbled and rough sculpin ranged from 11.1 to 14.7° C and 13.3 to 14.4°C, respectively. Values for Pit sculpin ranged from 9.9 to 16.4°C at acclimation temperatures of 10, 15 and 20°C. The distributions of marbled and rough sculpin are consistent with their behavioral and metabolic responses to temperature. The widespread distribution of Pit sculpin is consistent with its greater tolerance of high temperatures and eurythermal behavior, but the absence of Pit sculpin from habitats dominated by marbled and rough sculpin is not consistent with a temperature related explanation.     

Thermal niche variation among individuals of the poison frog,Oophaga pumilio,in forest and converted habitats

The conversion of natural habitats to human land uses often increases local temperatures, creating novel thermal environments for species. The variable responses of ectotherms to habitat conversion, where some species decline while others persist, can partly be explained by variation among species in their thermal niches. However, few studies have examined thermal niche variation within species and across forest‐land use ecotones, information that could provide clues about the capacity of species to adapt to changing temperatures. Here, we quantify individual‐level variation in thermal traits of the tropical poison frog, Oophaga pumilio, in thermally contrasting habitats. Specifically, we examined local environmental temperatures, field body temperatures (T_b), preferred body temperatures (T_pref), critical thermal maxima (CT_max), and thermal safety margins (TSM) of individuals from warm, converted habitats and cool forests. We found that frogs from converted habitats exhibited greater mean T_b and T_pref than those from forests. In contrast, CT_max and TSM did not differ significantly between habitats. However, CT_max did increase moderately with increasing T_b, suggesting that changes in CT_max may be driven by microscale temperature exposure within habitats rather than by mean habitat conditions. Although O. pumilio exhibited moderate divergence in T_pref, CT_max appears to be less labile between habitats, possibly due to the ability of frogs in converted habitats to maintain their T_b below air temperatures that reach or exceed CT_max. Selective pressures on thermal tolerances may increase, however, with the loss of buffering microhabitats and increased frequency of extreme temperatures expected under future habitat degradation and climate warming. Abstract in Spanish is available with online material.     

Elucidating the functional evolution of heat sensors among Xenopus species adapted to different thermal niches by ancestral sequence reconstruction

Ambient temperature fluctuations are detected via the thermosensory system which allows animals to seek preferable thermal conditions or escape from harmful temperatures. Evolutionary changes in thermal perception have thus potentially played crucial roles in niche selection. The genus Xenopus (clawed frog) is suitable for investigating the relationship between thermal perception and niche selection due to their diverse latitudinal and altitudinal distributions. Here we performed comparative analyses of the neuronal heat sensors TRPV1 and TRPA1 among closely related Xenopus species (X. borealis, X. muelleri, X. laevis, and X. tropicalis) to elucidate their functional evolution and to assess whether their functional differences correlate with thermal niche selection among the species. Comparison of TRPV1 among four extant Xenopus species and reconstruction of the ancestral TRPV1 revealed that TRPV1 responses to repeated heat stimulation were specifically altered in the lineage leading to X. tropicalis which inhabits warmer niches. Moreover, the thermal sensitivity of TRPA1 was lower in X. tropicalis than the other species, although the thermal sensitivity of TRPV1 and TRPA1 was not always lower in species that inhabit warmer niches than the species inhabit cooler niches. However, a clear correlation was found in species differences in TRPA1 activity. Heat‐evoked activity of TRPA1 in X. borealis and X. laevis, which are adapted to cooler niches, was significantly higher than in X. tropicalis and X. muelleri which are adapted to warmer niches. These findings suggest that the functional properties of heat sensors changed during Xenopus evolution, potentially altering the preferred temperature ranges among species.     

Growth and thermal tolerance of a Tibet fish Schizopygopsis younghusbandi juveniles acclimated to three temperature levels

Schizopygopsis younghusbandi is an endemic fish of Tibet characterized by slow growth. Artificial stock enhancement was applied to rebuild the natural population of S. younghusbandi in recent years. However, the optimal growth temperature and thermal tolerance of S. younghusbandi has not been studied, which restricts the production of S. younghusbandi fingerling for stock enhancement. The purpose of this paper is to determine the growth, critical thermal maximum (CTMax), lethal thermal maximum (LTMax) and acclimation response ratio (ARR) of S. younghusbandi juveniles (body weight 5.7 ± 1.2 g) at three acclimation temperature levels (10, 15, 20°C). The results showed that acclimation temperature significantly affected the growth, CTMax, LTMax and ARR of the experimental fish. Largest final weight (7.5 ± 2.3 g) was recorded in 15°C group. At a heating rate of 1°C/30 min, CTMax ranged from 30.98 to 32.01°C and LTMax ranged from 31.76 to 32.31°C in the three acclimation temperatures. Schizopygopsis younghusbandi had lower ARR value (0.097) than most other fish species. Low ARR value indicates that S. younghusbandi may have narrower thermal tolerance range and weaker acclimation ability to global warming. For successful aquaculture of S. younghusbandi juveniles, temperature should be maintained around 15°C.     

Comparisons of behavioural and TRPA1 heat sensitivities in three sympatric Cuban Anolis lizards

Thermal tolerances of organisms play a role in defining geographic ranges and occurrence of species. In Cuba, three sympatric species of Anolis lizards (Anolis allogus, Anolis homolechis and Anolis sagrei) inhabit different thermal microhabitats. A previous study found that these species showed distinct gene expression patterns in response to temperature stimuli, suggesting the genetically distinct thermal physiology among species. To investigate whether the Anolis species inhabiting locally distinct thermal habitats diverge their thermal tolerances, we first conducted behavioural experiments to analyse the temperatures at which the three Anolis species escape from heat source. Then, for each of the three species, we isolated cDNA encoding a putative molecular heat sensor, transient receptor potential ion channel ankyrin 1 (TRPA1), which has been suggested to play a role on eliciting behavioural responses to heat stimuli. We performed electrophysiological analysis to quantify activation temperature of Anolis TRPA1 to see whether the pattern of divergence in TRPA1 responses is congruent with that of divergence in behavioural responses. We found that temperatures triggering behavioural and TRPA1 responses were significantly lower for shade‐dwelling species (A. allogus) than for sun‐dwelling species (A. homolechis and A. sagrei). The ambient temperature of shade habitats where A. allogus occurs stays relatively cool compared to that of open habitats where A. homolechis and A. sagrei occur and bask. The high temperature thresholds of A. homolechis and A. sagrei may reflect their heat tolerances that would benefit these species to inhabit the open habitats.     

Worker thermal tolerance in the thermophilic ant Cataglyphis cursor (Hymenoptera, Formicidae)

In the thermophilic ant genus Cataglyphis, species differing in their physical caste system have developed alternative mechanisms to face extreme heat by physiological and/or behavioural adaptations. In this study, we tested whether thermal tolerance is related to worker size in the ant Cataglyphis cursor that presents intermediate worker size compared with previously studied species (size range 3.5–10 mm). Thermal tolerance at two temperatures was tested in the laboratory on colonies originating from two habitats (seaside versus vineyard), known to differ in average worker size. As expected large workers were more resistant to high temperature than small workers, but the effect of worker size on thermal resistance was less pronounced under the more extreme temperatures. The pattern of thermal tolerance was similar in the two habitat types. After controlling for worker size, worker thermal tolerance significantly varied amongst colonies, but this variation was not related to colony size. Our results suggest that a higher thermal tolerance can confer an advantage to larger workers especially during foraging and are discussed in the context of the evolution of worker size in ants.     

A laboratory study on the thermal tolerance of four southeastern stream insect species (Trichoptera,Ephemeroptera)

The acute thermal tolerances of four southeastern stream insect species, Ephemerella invaria (Walker), Stenonema ithaca (Clemens and Leonard), Symphitopsyche morosa (Hagun), and Brachycentrus lateralis (Say) were determined using an artificial stream enclosure. All species were acclimated at 10°C for 72 hours prior to instantaneous immersion into heated water for 96 hours. Percent mortality was recorded and the temperature at which 50% mortality occurred determined (LT5o). Data were subjected to standard statistical analysis.Thermal tolerance values were compared between species tested and to results from previous investigations using similar methodologies. The evolution and life histories of these species were also discussed in relation to their thermal tolerance values.     

Larval consumption rates,interspecific predation,and local guild composition of egg-overwintering Agabus (Coleoptera,Dytiscidae) species in vernal ponds

Summary In vernal ponds in the boreal region, egg-over-wintering Agabus species form a guild that feeds mainly on larvae and pupae of aedine mosquitoes. The regular co-existence of very similar Agabus species indicates local communities not structured by interspecific competition. However, the lower number of species in local guilds than in the regional species pool poses a problem of limited membership. We suggest that the species of this guild display habitat differences mainly with respect to water temperature, pond size and prey density. In this view, habitat selection reflects body size and thermal growth response of the species, mainly in connection with larval development. We present field data from two northern Swedish vernal ponds. Based on these data, feeding experiments were performed to test the hypothesis outlined above. At a high prey density, larvae of all instars of the larger species A. erichsoni Gemm. & Har. had a significantly higher consumption rate than those of the smaller species A. opacus Aubé. At a low prey density the differences were smaller, and only the third instar larvae differed significantly. At 2° C, larvae of A. opacus had a significantly higher consumption rate than those of A. congener (Thunberg). At 15° C, no significant difference was observed. In studies of within-guild interspecific predation, always the larger larvae consumed the smaller ones. Field data show that egg hatching is spread out in time, and show interspecific differences. Consequently, the effects of unexpected droughts differ with species.     

THERMAL ECOPHYSIOLOGY OF LAURENCIA PACIFICA AND LAURENCIA NIDIFICA (CERAMIALES,RHODOPHYTA) FROM TROPICAL AND WARM‐TEMPERATE REGIONS1

Laurencia is a globally distributed genus with about 80 species (order Ceramiales) that inhabit tropical, subtropical, and warm‐temperate regions of both sides of the Atlantic and Indo‐Pacific oceans. This study investigated how two species of Laurencia distributed in different thermal environments (California and Hawaii) varied in their photosynthetic responses to temperature. The thermal ecophysiology of both species was investigated at different temporal scales (short‐term responses and seasonal acclimatization) using oxygen evolution and pulse‐amplitude‐modulated (PAM) fluorometry. Our results indicated that seasonal acclimatization of both species of Laurencia influenced the short‐term photosynthetic response at both locations. Greater seasonal differences in the photosynthetic performance were observed for L. pacifica Kylin, which reflects the ability of this species to acclimatize to local environmental conditions characterized by short‐term fluctuations and a broader annual temperature range. Photosynthetic performance of L. nidifica J. Agardh was consistent with the less variable local environment (no short‐term fluctuations and a narrower temperature range). These results suggest that acclimatization to temperature variability in the environment can influence the degree of flexibility of physiological responses of species in this genus.     

Behavioral thermoregulation in the spiny lobster Panulirus Argus (latreille)

Ten spiny lobsters (Panulirus argus) were allowed to thermoregulate individually for 3-day periods in an electronic thermoregulatory shuttlebox which allowed them to control water temperatures (and thereby their own body temperatures) by their movements. The range of preferred (voluntarily occupied) temperatures was 25–35°C (mean 29.9°C; mode 30.0°C; median 30.0°C; midpoint 30.0°C; S_k (skewness, Pearson's coefficient) –0.04; s.e.m. 0.19°C; S.D. 2.32°C). The final thermal preferendum (by the gravitation method) in this species is 30°C.     

Interspecific variation in temperature dependence of egg development of five congeneric stonefly species (Protonemura Kempny, 1898, Nemouridae,Plecoptera)

Embryonal development of the five congeners Protonemura auberti Illies, 1954, P. hrabei Rauser, 1956, P. meyeri (Pictet, 1841), P. nitida (Stephens, 1835), and P. praecox (Morton, 1894) was studied under various laboratory temperatures and different photoperiods.Mean number of eggs in field collected batches was between 470 (P. praecox) and 1211 (P. auberti). Spring species had smaller egg batches than autumn species (Table 1). Mean hatching success in the laboratory was 50–100% at 2–18 °C. In most species hatching success decreased slightly with increasing temperature (Figs. 1a-e). None of the eggs incubated at 24 °C developed. Hatching pattern followed an asymmetric frequency distribution. In general, the hatching periods were the shorter the higher the incubation temperature.Embryonic development of all five species was inversely temperature dependent (Figs. 2a-e), and well described by a power law relationship (Figs. 3a-e). Interspecific differences in incubation periods were notable at nearly all temperatures (Fig. 4). There was a distinct interspecific sequence in length of incubation period (with steps of about 4 days), which was the same as can be seen in the flight periods: The later the species flies the longer the incubation period. Temperature fluctuations and variations in photoperiod had no influence on incubation and hatching periods or hatching success.The thermal demand of the egg stage neither explains the recent geographical distribution of the Protonemura species, nor does it directly correspond to the field temperatures common during their egg development. However, it is optimal in respect to resource partitioning between the five species, with the consequence of temporal displacement of life cycles.Derived from Brittain's (in press) proposal to compare the two constants a and b of the regressions describing the temperature dependence of embryonal development, a new index (Integral Development Time, IDT) indicating the thermal demand was created for easier comparison of numerous species (Table 2). Evaluation of the IDT for various species of Plecoptera (Fig. 5) suggests that species belonging to the family group Systellognatha generally have higher thermal requirements in the egg stage than species belonging to the Euholognatha.     

Heat freezes niche evolution

Climate change is altering phenology and distributions of many species and further changes are projected. Can species physiologically adapt to climate warming? We analyse thermal tolerances of a large number of terrestrial ectotherm (n = 697), endotherm (n = 227) and plant (n = 1816) species worldwide, and show that tolerance to heat is largely conserved across lineages, while tolerance to cold varies between and within species. This pattern, previously documented for ectotherms, is apparent for this group and for endotherms and plants, challenging the longstanding view that physiological tolerances of species change continuously across climatic gradients. An alternative view is proposed in which the thermal component of climatic niches would overlap across species more than expected. We argue that hard physiological boundaries exist that constrain evolution of tolerances of terrestrial organisms to high temperatures. In contrast, evolution of tolerances to cold should be more frequent. One consequence of conservatism of upper thermal tolerances is that estimated niches for cold‐adapted species will tend to underestimate their upper thermal limits, thereby potentially inflating assessments of risk from climate change. In contrast, species whose climatic preferences are close to their upper thermal limits will unlikely evolve physiological tolerances to increased heat, thereby being predictably more affected by warming.     

How recruitment, intraspecific interactions, and predation control species borders in a tidal estuary

We examined the relative contribution of recruitment, intraspecific species interactions, and predation in controlling the upper intertidal border of the northern acorn barnacle, Semibalanusbalanoides, in a tidal estuary in Maine. We hypothesized that the contracted border at sites that experienced low tidal currents was due to flow-mediated recruitment that resulted in reduced survival due to the absence of neighbor buffering of thermal stress (i.e., positive intraspecific interactions). We tested this hypothesis by manipulating the density of recently settled barnacles and their thermal environment in a field experiment. Counter to our original hypothesis, barnacles with neighbors suffered severe mortality at low-flow sites. When density-dependent predation by the green crab (Carcinusmaenus) was experimentally eliminated, however, we did detect evidence for positive interactions at the low-flow sites but not at the high-flow sites. In spite of the close proximity of the sites, maximum daily rock temperatures at the low-flow sites were slightly, but consistently, greater than those at high-flow sites. Our findings suggest that the upper intertidal border of S. balanoides in the Damariscotta River is limited at low-flow sites by a combination of reduced recruitment, elevated mortality from thermal stress and enhanced predation by green crabs. More generally, our findings highlight how physical stress and predation interact to alter the nature of density-dependent species interactions in natural assemblages. Received: 6 August 1998 / Accepted: 11 October 1998     

Divergence of thermal physiological traits in terrestrial breeding frogs along a tropical elevational gradient

Critical thermal limits are thought to be correlated with the elevational distribution of species living in tropical montane regions, but with upper limits being relatively invariant compared to lower limits. To test this hypothesis, we examined the variation of thermal physiological traits in a group of terrestrial breeding frogs (Craugastoridae) distributed along a tropical elevational gradient. We measured the critical thermal maximum (CT_max; n = 22 species) and critical thermal minimum (CT_min; n = 14 species) of frogs captured between the Amazon floodplain (250 m asl) and the high Andes (3,800 m asl). After inferring a multilocus species tree, we conducted a phylogenetically informed test of whether body size, body mass, and elevation contributed to the observed variation in CT_max and CT_min along the gradient. We also tested whether CT_max and CT_min exhibit different rates of change given that critical thermal limits (and their plasticity) may have evolved differently in response to different temperature constraints along the gradient. Variation of critical thermal traits was significantly correlated with species’ elevational midpoint, their maximum and minimum elevations, as well as the maximum air temperature and the maximum operative temperature as measured across this gradient. Both thermal limits showed substantial variation, but CT_min exhibited relatively faster rates of change than CT_max, as observed in other taxa. Nonetheless, our findings call for caution in assuming inflexibility of upper thermal limits and underscore the value of collecting additional empirical data on species’ thermal physiology across elevational gradients.     

Critical thermal limits and their responses to acclimation in two sub-Antarctic spiders: <Emphasis Type="Italic">Myro kerguelenensis</Emphasis> and <Emphasis Type="Italic">Prinerigone vagans</Emphasis>

Despite the relative richness of spider species across the Southern Ocean islands remarkably little information is available on their biology. Here, the critical thermal limits of an indigenous (Myro kerguelenensis, Desidae) and an introduced (Prinerigone vagans, Linyphiidae) spider species from Marion Island were studied after 7–8 days acclimation to 0, 5, 10 and 15°C. Critical thermal minima (CT_Min) were low in these species by comparison with other spiders and insects measured to date, and ranged from −6 to −7°C in M. kerguelenensis and from −7 to −8°C in P. vagans. In contrast, critical thermal maxima (CT_Max) were similar to other insects on Marion Island (M. kerguelenensis: 35.0–35.6°C; P. vagans: 35.1–36.0°C), although significantly lower than those reported for other spider species in the literature. The magnitude of acclimation responses in CT_Max was lower than those in CT_Min for both species and this suggests decoupled responses to acclimation. Whilst not conclusive, the results raise several important considerations: that oxygen limitation of thermal tolerance needs to be more widely investigated in terrestrial species, that indigenous and alien species might differ in the nature and extent of their plasticity, and that upper and lower thermal tolerance limits might be decoupled in spiders as is the case in insects.