Experimental effect of cold,La Niña temperatures on the survival of reef fishes from Gorgona Island (eastern Pacific Ocean)

The eastern tropical Pacific (ETP) reefs are affected at irregular times by extremely cold temperatures that occur principally during La Niña events. The effects of these low temperatures on the survival of reef fishes were experimentally assessed by determining the critical thermal minimum (CTM) of 15 reef fish species from Gorgona Island (ETP), and comparing these CTMs with the records of temperature during past La Niña events. Among species, mean CTMs ranged from 10.8°C to 16.3°C, which were lower than the coldest temperature recorded during the last La Niña event (18°C during La Niña 1998-1999). However, the observed ranges of CTM for two species (Thalassoma lucassanum and Eucinostomus gracilis) extended above 18°C. These results suggest that most of the reef fishes we studied are physiologically tolerant to the cold temperatures encountered during La Niña, though decreases in at least two populations may be expected as a result of the mortality of less tolerant individuals. Although tolerant to cold temperatures, reef fish populations may still experience negative changes during La Niña, because other determinants in population maintenance (e.g. reproduction and recruitment) are more temperature sensitive. The effects of other cold phenomena on reef fish survival are also discussed herein.     

Thermal tolerance of early development in tropical and temperate sea urchins: inferences for the tropicalization of eastern Australia

The thermal envelope of development to the larval stage of two echinoids from eastern Australia was characterized to determine whether they fill their potential latitudinal ranges as indicated by tolerance limits. The tropical sand dollar, Arachnoides placenta, a species that is not known to have shifted its range, was investigated in Townsville, northern Australia (19°20′S, 146°77′E), during its autumn spawning season (May 2012). The subtropical/temperate sea urchin, Centrostephanus rodgersii, a species that has undergone poleward range expansion, was investigated in Sydney, southern Australia (33°58′S, 151°14′E), during its winter spawning season (August 2012). The thermal tolerance of development was determined in embryos and larvae reared at twelve temperatures. For A. placenta, the ambient water temperature near Townsville and experimental control were 24 °C and treatments ranged from 14 to 37 °C. For C. rodgersii, ambient Sydney water temperature and experimental control were 17 °C, and the treatment range was 9–31 °C. A. placenta had a broader developmental thermal envelope (14 °C range 17–31 °C) than C. rodgersii (9 °C range 13–22 °C). Both species developed successfully at temperatures well below ambient, suggesting that cooler water is not a barrier to poleward migration for either species. Both species presently live near the upper thermal limits for larval development, and future ocean warming could lead to contractions of their northern range limits. This study provides insights into the factors influencing the realized and potential distribution of planktonic life stages and changes to adult distribution in response to global change.     

The 1998 bleaching event and its aftermath on a coral reef in Belize

Widespread thermal anomalies in 1997-1998, due primarily to regional effects of the El Niño-Southern Oscillation and possibly augmented by global warming, caused severe coral bleaching worldwide. Corals in all habitats along the Belizean barrier reef bleached as a result of elevated sea temperatures in the summer and fall of 1998, and in fore-reef habitats of the outer barrier reef and offshore platforms they showed signs of recovery in 1999. In contrast, coral populations on reefs in the central shelf lagoon died off catastrophically. Based on an analysis of reef cores, this was the first bleaching-induced mass coral mortality in the central lagoon in at least the last 3,000 years. Satellite data for the Channel Cay reef complex, the most intensively studied of the lagoonal reefs, revealed a prolonged period of elevated sea-surface temperatures (SSTs) in the late summer and early fall of 1998. From 18 September to 1 October 1998, anomalies around this reef averaged +2.2°C, peaking at 4.0°C above the local HotSpot threshold. In situ temperature records from a nearby site corroborated the observation that the late summer and early fall of 1998 were extraordinarily warm compared to other years. The lettuce coral, Agaricia tenuifolia, which was the dominant occupant of space on reef slopes in the central lagoon, was nearly eradicated at Channel Cay between October 1998 and January 1999. Although the loss of Ag. tenuifolia opened extensive areas of carbonate substrate for colonization, coral cover remained extremely low and coral recruitment was depressed through March 2001. High densities of the sea urchin Echinometra viridis kept the cover of fleshy and filamentous macroalgae to low levels, but the cover of an encrusting sponge, Chondrilla cf. nucula, increased. Further increases in sponge cover will impede the recovery of Ag. tenuifolia and other coral species by decreasing the availability of substrate for recruitment and growth. If coral populations are depressed on a long-term basis, the vertical accretion of skeletal carbonates at Channel Cay will slow or cease over the coming decades, a time during which global-warming scenarios predict accelerated sea-level rise.     

Viability Analysis of Reef Fish Populations Based on Limited Demographic Information

Abstract:  Marine protected areas (MPAs) that allow some degree of artisanal fishing have been proposed to control the overexploitation of marine resources while allowing extraction by local communities. Nevertheless, the management of MPAs is often impaired by the absence of data on the status of their resources. We devised a method to estimate population growth rates with the type of data that are usually available for reef fishes. We used 7 years of spatially explicit abundance data on the leopard grouper ( Mycteroperca rosacea ) in an MPA in the Gulf of California, Mexico, to construct a matrix population model that incorporated the effects of El Niño/La Niña Southern Oscillation on population dynamics. An environmental model that estimated different demographic estimates for El Niño and La Niña periods performed better than a single-environment model, and a single-habitat model performed better than a model that considered different depths as different habitats. Our results suggest that the population of the leopard grouper off the main island of the MPA is not viable under present conditions. Although the impact of fishing on leopard grouper populations in the MPA has not yet been established, fishing should be closed as a precautionary measure at this island if a priority of the MPA is to ensure the sustainability of its fish populations.     

The effects of an extraordinary El Niño / La Niña event on the size and growth of the squid<Emphasis Type="Italic"> Loligo opalescens</Emphasis> off Southern California

The population structure of the California market squid Loligo opalescens was studied for the Channel Islands region off Southern California between June 1998 and March 2000. During this time Californian waters were exposed to an extraordinary El Niño event that was possibly the most dramatic change in oceanographic conditions that occurred last century. There was then a rapid transition to record cool La Niña conditions. Statolith increments were used to determine age parameters and increment periodicity was validated for the first 54 days of life. Based on statolith increment counts, the oldest males and females were 257 and 225 days respectively and individuals matured as young as 129 and 137 days respectively. No distinct hatching period was detected. There was a general trend of increasing body size throughout the study period. Squid that hatched and grew through the El Niño were strikingly smaller and had slower growth rates compared to squid that grew through the La Niña. This was related to oceanography and associated productivity. There was a positive correlation between squid mantle length and upwelling index and a negative correlation between mantle length and sea temperature. The 'live-fast die-young' life history strategy of squid makes them ideal candidates for following the effects of the dramatic changes in oceanographic conditions off California. We propose that squid can serve as ecosystem recorders and productivity integrators over time and space and are useful organisms to tie oceanography to biology.     

Coral Bleaching and Global Climate Change: Scientific Findings and Policy Recommendations

Abstract: In 1998, tropical sea surface temperatures were the highest on record, topping off a 50-year trend for some tropical oceans. In the same year, coral reefs around the world suffered the most extensive and severe bleaching ( loss of symbiotic algae) and subsequent mortality on record. These events may not be attributable to local stressors or natural variability alone but were likely induced by an underlying global phenomenon. It is probable that anthropogenic global warming has contributed to the extensive coral bleaching that has occurred simultaneously throughout the reef regions of the world. The geographic extent, increasing frequency, and regional severity of mass bleaching events are an apparent result of a steadily rising baseline of marine temperatures, combined with regionally specific El Niño and La Niña events. The repercussions of the 1998 mass bleaching and mortality events will be far-reaching. Human populations dependent on reef services face losses of marine biodiversity, fisheries, and shoreline protection. Coral bleaching events may become more frequent and severe as the climate continues to warm, exposing coral reefs to an increasingly hostile environment. This global threat to corals compounds the effects of more localized anthropogenic factors that already place reefs at risk. Significant attention needs to be given to the monitoring of coral reef ecosystems, research on the projected and realized effects of global climate change, and measures to curtail greenhouse gas emissions. Even those reefs with well-enforced legal protection as marine sanctuaries, or those managed for sustainable use, are threatened by global climate change.     

Global effects of extreme temperatures on wild bumblebees

Climate plays a key role in shaping population trends and determining the geographic distribution of species because of limits in species’ thermal tolerance. An evaluation of species tolerance to temperature change can therefore help predict their potential spatial shifts and population trends triggered by ongoing global warming. We assessed inter- and intraspecific variations in heat resistance in relation to body mass, local mean temperatures, and evolutionary relationships in 39 bumblebee species, a major group of pollinators in temperate and cold ecosystems, across 3 continents, 6 biomes, and 20 regions (2386 male specimens). Based on experimental bioassays, we measured the time before heat stupor of bumblebee males at a heatwave temperature of 40 °C. Interspecific variability was significant, in contrast to interpopulational variability, which was consistent with heat resistance being a species-specific trait. Moreover, cold-adapted species are much more sensitive to heat stress than temperate and Mediterranean species. Relative to their sensitivity to extreme temperatures, our results help explain recent population declines and range shifts in bumblebees following climate change.     

Amazonian Tree Mortality during the 1997 El Niño Drought

Abstract: In 1997, the Amazon Basin experienced an exceptionally severe El Niño drought. We assessed effects of this rare event on mortality rates of trees in intact rain forest based on data from permanent plots. Long-term (5- to 13-year) mortality rates averaged only 1.12% per year prior to the drought. During the drought year, annual mortality jumped to 1.91% but abruptly fell back to 1.23% in the year following El Niño. Trees dying during the drought did not differ significantly in size or species composition from those that died previously, and there was no detectable effect of soil texture on mortality rates. These results suggest that intact Amazonian rainforests are relatively resistant to severe El Niño events.     

Temperature tolerance and latitudinal range of brown algae from temperate Pacific South America

Lower and upper survival temperatures of microthalli of 25 species of South American Phaeophyceae isolated from central Peru (14°S) to the Canal Beagle (55°S) were determined using 2-wk exposure for the upper and 4-wk exposure for the lower limit. All species survive 4 wks at -2°C. With respect to the upper limit, species reported only from southernmost South America tolerate 19.9 to 24.5°C (n=8), and species occurring from Cape Horn to central Chile 24.6 to 27.4°C (n=7). Three species that occurred as far north as northern Chile and Peru before the 1982–1983 El Niño event, and whose northern limit was dramatically shifted southwards in 1983, tolerate 20.8 to 25.3°C, whereas five species that have survived in Peru tolerate 25.6 to 28.5°C. Tinocladia falklandica which tolerates 27.8 to 28.1°C but lives only in southernmost South America and Striaria attenuata, which tolerates 31.6 to 31.9°C but occurs at ca. 42°S, are exceptional. Their high temperature tolerance may have no adaptive value in South America. They are restricted to the cold-temperate region due to low temperature requirements for reproduction or for reasons yet unknown. In general, the northern distributional limits of the Phaeophyceae studied along the temperate Pacific coast of South America are reproduction boundaries, except in El Niño years when they are redefined according to the species' upper suvival limits. Temperature tolerance of isolates from northern Chile and Peru agrees well with maximum temperatures reached during the 1983 El Niño.     

Effects of More Frequent and Prolonged El Niño Events on Life-History Parameters of the Degu, a Long-Lived and Slow-Reproducing Rodent

Abstract:  Global climate change (GCC) can have profound effects on species whose ecology is governed primarily by climatic factors. The ecology of small mammals inhabiting semiarid Chile is strongly affected by the El Niño Southern Oscillation (ENSO). During La Niña events in this area, dry conditions prevail and species may disappear from the thorn-scrub habitat. Conversely, El Niño events bring high rainfall, and associated pulses of food trigger small-mammal population increases. We used capture–mark–recapture to study responses of the degu ( Octodon degus ), a dominant small mammal, to variation in rainfall over 18 years. In response to a recent trend toward wetter conditions, degus reached record-high densities and maintained more stable numbers in the area. Underlying mechanisms involved variation in adult survival, juvenile persistence, and fecundity linked to rainfall changes during consecutive years (i.e., rainfall phases). During prolonged droughts, degus had low survival and produced fewer offspring, with low persistence. Following high rainfall, these parameters reversed; consecutive wet years resulted in further increases. Weak declines in fecundity and adult survival and high persistence of juveniles explained delayed responses to deteriorating conditions in initial dry years. If GCC leads to increased frequency of El Niño events, we anticipate greater numerical dominance of degus in semiarid Chile and possible range expansion. Furthermore, degus have strong impacts on other small mammal and some plant species, are important prey species, and are agricultural pests and disease reservoirs. Hence, GCC has the potential to dramatically influence their ecology in northern Chile and to have cascading effects on other components of this system.     

Behaviour moderates climate warming vulnerability in high-rocky-shore snails: interactions of habitat use, energy consumption and environmental temperature

High-rocky-shore intertidal animals are predicted to be generally more vulnerable to climate warming than lower-shore species, because their thermal tolerances lie closer to maximum environmental temperatures (T _e). However, this prediction is based on taxonomically and ecologically limited information. The present study investigated the effect of habitat use on climate warming vulnerability of the tropical high-shore snail, Echinolittorina malaccana (from Brunei Darussalam, 5°N), which aestivates in sun-exposed or shaded habitats. The thermal regimes of these habitats differed vastly, but snails showed similar daily energy consumption in either habitat, due to temperature-insensitive metabolism (TIM) between 35 and 46 °C in the sun-resting snails. However, maximum T _e values in the shade and the sun were 35 and 46 °C, respectively, suggesting that sun-resting snails, which presently experience temperatures near the incipient lethal temperature range (46–56 °C), should be more threatened by further warming than shade-resting snails, which have an 11 °C ‘safety margin’. Thus, vulnerability of high-shore species to climate warming could be moderated by availability of shaded habitat, making predictions for these species more complex than previously realized.     

Green abalone, <Emphasis Type="Italic">Haliotis fulgens</Emphasis> infected with the agent of withering syndrome do not express disease signs under a temperature regime permissive for red abalone, <Emphasis Type="Italic">Haliotis rufescens</Emphasis>

All California abalone species have been shown to be susceptible to infection with the bacterial agent of abalone withering syndrome (WS), although expression of signs of the disease may vary between species and with environmental conditions. We examined thermal modulation of WS expression in green abalone Haliotis fulgens at temperatures mimicking El Niño (18.0°C) and La Niña (14.2°C) events in southern California. In contrast to results obtained from previous experiments with red abalone, H. rufescens, the higher temperature did not result in higher infection intensities of the causative agent of the disease nor increase in clinical signs of disease. These results demonstrate clear differences in thermal regulation of disease expression between abalone species, and provide further data suggesting that green abalone should be a target species of recovery efforts in southern California, where WS is endemic.     

The effects of temperature and pressure acclimation on the temperature and pressure tolerance of the shallow-water shrimp Palaemonetes varians

Anthropogenic effects on marine ecosystems (e.g. hypoxia, warming) at and beyond continental margins are assumed to affect physiological and biochemical boundaries to species’ distributions, potentially leading to habitat contraction across depth. Whether or not shallow-water benthic invertebrates are capable of undergoing depth-related migrations in response to such perturbations remains largely unknown. The few studies available have focused solely on whether colonisation of deep waters may be ongoing and on the ability of shallow-water species to tolerate low temperatures and high hydrostatic pressures: two physical parameters, which are thought to limit the depth range of a species. Those studies did not consider the effects of acclimation to low temperature and, especially, acclimation to high hydrostatic pressure on pressure tolerance. We demonstrate that acclimation to both low temperature (5 °C) and to high hydrostatic pressure (10 MPa) increases the pressure tolerance within the shallow-water shrimp Palaemonetes varians. Previous studies have demonstrated the impressive temperature and pressure tolerance of this shallow-water shrimp. Here, we provide evidence that a shallow-water species may acclimate to low temperature and high pressure and show greater pressure tolerance, suggesting that shallow-water organisms may be able to rapidly—and potentially stepwise—acclimate to the low temperature and high pressure conditions typical of the deep sea. These findings are of importance for understanding phylogenetic development from shallow- to deep-water species and the processes behind past, present and future bathymetric range shifts in species.     

Temperature acclimation of upper and lower thermal limits and freeze resistance in the nonindigenous brown mussel, Perna perna (L.), from the Gulf of Mexico

Acute and chronic upper and lower thermal limits and freeze resistance were investigated in the nonindigenous brown mussel, Perna perna, from the Texas Gulf of Mexico coast in order to assess its potential distribution in North American coastal waters. This species' long-term, incipient lower and upper thermal limits were 7.5°C and 30°C, congruent with the seasonal ambient water temperature range of 10-30°C reported for other populations worldwide. Effects of temperature acclimation and individual size on survival time were most pronounced on chronic exposures to lethal temperatures approaching incipient lower or upper thermal limits. When exposed to temperature increasing at 0.1°C min^-1, the acute upper lethal limit was 44°C regardless of acclimation temperature or individual size. P. perna had a limited freeze resistance, being intolerant of emersion at -2.5°C. This species' narrow incipient thermal limits, limited capacity for temperature acclimation and poor freeze resistance may account for its restriction to subtidal and lower eulittoral zones of cooler subtropical rocky shores. Near extinction of P. perna from Texas Gulf of Mexico waters occurred in the summer of 1997 when mean surface-water temperatures approached its incipient upper limit of 30°C.     

Trophic cascades result in large-scale coralline algae loss through differential grazer effects

Removal of predators can have strong indirect effects on primary producers through trophic cascades. Crustose coralline algae (CCA) are major primary producers worldwide that may be influenced by predator removal through changes in grazer composition and biomass. CCA have been most widely studied in Caribbean and temperate reefs, where cover increases with increasing grazer biomass due to removal of competitive fleshy algae. However, each of these systems has one dominant grazer type, herbivorous fishes or sea urchins, which may not be functionally equivalent. Where fishes and sea urchins co-occur, fishing can result in a phase shift in the grazing community with subsequent effects on CCA and other substrata. Kenyan reefs have herbivorous fishes and sea urchins, providing an opportunity to determine the relative impacts of each grazer type and evaluate potential human-induced trophic cascades. We hypothesized that fish benefit CCA, abundant sea urchins erode CCA, and that fishing indirectly reduces CCA cover by removing sea urchin predators. We used closures and fished reefs as a large-scale, long-term natural experiment to assess how fishing and resultant changes in communities affect CCA abundance. We used a short-term caging experiment to directly test the effects of grazing on CCA. CCA cover declined with increasing fish and sea urchin abundance, but the negative impact of sea urchin grazing was much stronger than that of fishes. Abundant sea urchins reduced the CCA growth rate to almost zero and prevented CCA accumulation. A warming event (El Ni?o Southern Oscillation, ENSO) occurred during the 18-year study and had a strong but short-term positive effect on CCA cover. However, the effect of the ENSO on CCA was lower in magnitude than the effect of sea urchin grazing. We compare our results with worldwide literature on bioerosion by fishes and sea urchins. Grazer influence depends on whether benefits of fleshy algae removal outweigh costs of grazer-induced bioerosion. However, the cost-benefit ratio for CCA appears to change with grazer type, grazer abundance, and environment. In Kenya, predator removal leads to a trophic cascade that is expected to reduce net calcification of reefs and therefore reduce reef stability, growth, and resilience.     

Biodiversity in Littorina species (Mollusca: Gastropoda): a physiological approach using heat-coma

 The upper critical thermal limits were measured as heat-coma temperatures in Littorina species as a tool for examining physiological diversity. Thermal tolerance was found to be stable within a population, but extensive variation occurred between populations and species. All species examined displayed heat-coma values of ∼30 °C and did not show a positive correlation with shore height. Eulittoral-fringe species tended to have higher coma temperatures than eulittoral species. Coma temperatures varied both seasonally and geographically. Lethal thermal limits were also investigated; these were ∼10 C° higher than coma temperatures. Received: 16 August 1999 / Accepted: 13 June 2000     

The synergistic effects of increasing temperature and CO2 levels on activity capacity and acid–base balance in the spider crab, Hyas araneus

With global climate change, ocean warming and acidification occur concomitantly. In this study, we tested the hypothesis that increasing CO₂ levels affect the acid–base balance and reduce the activity capacity of the Arctic spider crab Hyas araneus, especially at the limits of thermal tolerance. Crabs were acclimated to projected oceanic CO₂ levels for 12 days (today: 380, towards the year 2100: 750 and 1,120 and beyond: 3,000 μatm) and at two temperatures (1 and 4 °C). Effects of these treatments on the righting response (RR) were determined (1) at acclimation temperatures followed by (2) righting when exposed to an additional acute (15 min) heat stress at 12 °C. Prior to (resting) and after the consecutive stresses of combined righting activity and heat exposure, acid–base status and lactate contents were measured in the haemolymph. Under resting conditions, CO₂ caused a decrease in haemolymph pH and an increase in oxygen partial pressure. Despite some buffering via an accumulation of bicarbonate, the extracellular acidosis remained uncompensated at 1 °C, a trend exacerbated when animals were acclimated to 4 °C. The additional combined exposure to activity and heat had only a slight effect on blood gas and acid–base status. Righting activity in all crabs incubated at 1 and 4 °C was unaffected by elevated CO₂ levels or acute heat stress but was significantly reduced when both stressors acted synergistically. This impact was much stronger in the group acclimated at 1 °C where some individuals acclimated to high CO₂ levels stopped responding. Lactate only accumulated in the haemolymph after combined righting and heat stress. In the group acclimated to 1 °C, lactate content was highest under normocapnia and lowest at the highest CO₂ level in line with the finding that RR was largely reduced. In crabs acclimated to 4 °C, the RR was less affected by CO₂ such that activity caused lactate to increase with rising CO₂ levels. In line with the concept of oxygen and capacity limited thermal tolerance, all animals exposed to temperature extremes displayed a reduction in scope for performance, a trend exacerbated by increasing CO₂ levels. Additionally, the differences seen between cold- and warm-acclimated H. araneus after heat stress indicate that a small shift to higher acclimation temperatures also alleviates the response to temperature extremes, indicating a shift in the thermal tolerance window which reduces susceptibility to additional CO₂ exposure.     

Activation energy for skeletal aragonite deposited by the hermatypic coral Platygyra spp.

Average radial growth rates of the hemispherical aragonite colonies deposited by the Indo-Pacific scleractinian reef coral Platygyra spp. were determined by measuring the thickness of density variations in the skeleton that are revealed by X-radiography. Ninety-one specimens from 21 localities were examined, but only 54 of these exhibited well-defined growth-banding. The apparent temperature dependence of growth rate is linear over the range 23.9° to 29.3°C, averaging 5.4±0.94 mm/year at 24°C, 8.0±0.42 mm/year at 27°C, and 9.7 ±0.58 mm/year at 29°C (90% confidence limits). Expression of the influence of temperature on growth rate in terms of the Arrhenius equation yields an activation energy of 20,680 cal/mole, which is comparable to values for typical biological reactions, but is only half that reported for skeletogenesis in another reef coral, Pocillopora damicornis, on the basis of controlled incubation studies involving ⁴⁵Ca uptake.     

Area-specific temporal changes of species composition and species-specific range shifts in rocky-shore mollusks associated with warming Kuroshio Current

Increases of low-latitude species in various sea areas and poleward shifts of ranges of many species have been reported and linked to warming environments. To examine the generality of these trends for mollusks, we conducted 7 quadrat surveys during 1978–2006 on Pacific rocky shores in Japan (26.6–45.0°N). Results showed that the dominance of southern species increased on 11 of the 15 shores in a southern, warming sea area but on only 1 of the 6 shores in a northern area with no warming trend. The latitudes of species ranges increased on average but showed large interspecific variations which showed a weak, positive correlation with interspecific taxonomic distance. The differences of these results from the previously reported trends are discussed in relation to the unique current patterns in our study area and the phylogenetic constraints of species-specific responses to a warming environment.     

Physiological and behavioral responses of temperate seahorses (Hippocampus guttulatus) to environmental warming

The aim of the present study was to evaluate, for the first time, the effect of environmental warming on the metabolic and behavioral ecology of a temperate seahorse, Hippocampus guttulatus. More specifically, we compared routine metabolic rates, thermal sensitivity, ventilation rates, food intake, and behavioral patterns at average spring temperature (18 °C), average summer temperature (26 °C), temperatures that they endure during summer heat wave events (28 °C), and in a near-future warming scenario (+2; 30 °C) in Sado estuary, Portugal. Both newborn juveniles and adults showed significant increases in metabolic rates with rising temperatures. However, newborns were more impacted by future warming via metabolic depression (i.e., heat-induced hipometabolism). In adult stages, ventilation rates also increased significantly with environmental warming, but food intake remained unchanged. Moreover, the frequency of swimming, foraging, swinging, and inactivity did not significantly change between the different thermal scenarios. Thus, we provide evidence that, while adult seahorses show great resilience to heat stress and are not expected to go through any physiological impairment and behavioral change with the projected near-future warming, the early stages display greater thermal sensitivity and may face greater metabolic challenges with potential cascading consequences for their growth and survival.