Thermal dynamics and climate variability

Thermal dynamics and climate variability: a review of crustacea from differing climates/ different latitudes
Sydney Henderson
Abstract
In a world dominated by ectotherms, organisms whose body temperature closely follows that of the environment, temperature can be considered the most pervasive abiotic factor. Temperature impacts all aspects of ectothermic life, from cellular processes to global distribution patterns. Temperature, and consequently climate is tightly linked to thermal dynamics such as locomotory activity, behavioural and physiological adaptations, life history stages and acclimation ability. Temperature, as well as temperature variability changes on a latitudinal gradient. Tropical climates exhibit high, relatively stable temperatures, and consequently inhabitant crustacea are adapted to rely on behavioural thermoregulation such as shade-seeking and burrowing. Furthermore tropical crustacea live close to their upper thermal range so have a relatively limited acclimation ability. Similarly, polar climates are characterized by low but stable temperatures. Polar crustacea, in a relatively unavoidable cold temperatures, rely on more physiochemical adaptations to prevent cellular freezing. Crustacea from polar and tropical climates experience stable temperatures and will have relatively low thermal tolerance and acclimation ability, given that they do not experience fluctuating seasonal temperature which would necessitate thermal plasticity and acclimation ability. In comparison, temperate climates have seasonal and diurnal temperature fluctuations, requiring wider thermal tolerance ranges and acclimation ability. Climate is intrinsically linked to crustacea thermal dynamics. Understanding the way organisms adapt and react to temperature change will be imperative for effective biodiversity management and conservation in the face of anthropogenic climate change.

Introduction
Temperature can be considered the major environmental factor, particularly for ectotherms, whose body temperature follows that of their environment to varying degrees (Angiletta, 2009). The extrinsic importance of temperature lies in that it dictates the rate of all physiological and biological reactions, and organisation. Extreme high or low tempertures can be lethal. Crustacea (phylum: Arthropoda) are poikilotherms, meaning their body temperature follows closely that of the environment (Hickman et al. 2007) and as their internal metabolic heat production is relatively low, they are also considered bradymetabolic. Temperature has a fundamental impact on all stages of crustacean life and physiology, impacting growth (Hartnoll, 2001), reproduction (Nagaraju, 2011), locomotion (Sudo, 2003) as well as larger scale effects such as distribution limits (Somero, 2004; Terblanche et al. 2011). Consequently, temperature impacts all levels of organisation from cellular to the whole ecosystem.
As ectotherms, crustaceans’ internal temperature, and therefore, a range of thermal-dependant metabolic processes, relies on that of their external environment (Zi-Ming et al. 2013). Crustacea are found all over the globe, inhabiting a diverse range of thermal regimes. They display a variety of thermal adaptations, both behavioural and physiological to adapt to the thermal dynamics of their environment, in order to maintain an efficient internal metabolic rate (Lagerspetz ; Vainio, 2006). The distribution of the sub phyla crustacea across a global latitudinal gradient allows comparative investigation of thermal dynamics under different environmental conditions. Trends in global thermal dynamics have previously been put forward, linking thermal range and acclimation ability to environmental temperatures and thermal stability of an organisms’ natural habitat (Addo-Bediako et al. 2000; Nguyen et al. 2011; Somero 1996; Stillman, 2003). In this review these proposed trends will be examined in the case of crustacea from tropical, temperate and polar latitudes. An understanding of how organisms react and adapt to temperature change acquires key relevance in the face of anthropogenic climate change, and will be essential for conservation and biodiversity management (Chevin et al. 2010).
While thermal variance has been studied in microhabitats such as intertidal areas (e.g. Nguyen et al. 2011; Somero & Stillman 2000), and globally in other groups such as bivalves (Compton et al. 2007) and tropical insects (Addo-Bediako et al. 2000), a global review of crustacean thermal dynamics is presented here. In this review, the major influence of temperature on crustacea will be investigated through five sections; 1) temperature and locomotory activity, 2) behavioural adaptation and thermal preference, 3) physiological adaptations, 4) temperature and life history stages, and 5) thermal acclimation. These five aspects of thermal dynamics will present a conclusive illustration of how climate variability impacts thermal dynamics in crustacea.
1. Temperature and locomotory activity
The Van’t Hoff rule describes how the rate of chemical reactions and biological functions increases approximately two-fold for every 10°C increase, within a biological temperature range. The rhythmical movement of the cirral legs of barnacles illustrates this rule. Sessile adult barnacles (crustacea, cirripedia) rely on beating and pumping of their cirral legs to filter feed and for respiratory ventilation. Movement rate depends on temperature, in accordance with the Van’t Hoff rule; there is a positive correlation between increased temperature and increased leg movement (Nishizaki & Carrington, 2014). Mobile crustaceans’ temperature-performance curves are more complicated given the added factor of behavioural thermoregulation. The majority of experiments in this area are laboratory based, and have been collated in Table 1. It is important to note, however, that laboratory experiments cause stress to the animal which may affect locomotion, and that in-situ behavioural thermoregulation is impossible to account for in a laboratory setting. Furthermore locomotory responses, as a physiological response, may also be affected by other environmental conditions (e.g. salinity) and again it would be near impossible to test for all environmental variables in a laboratory. That being said, focusing on the effects of temperature on locomotory activity may offer a good representation and serve as a prediction for understanding how organisms will be affected by climate change.
The information in Table 1 demonstrates the varying effects of increasing temperature on crustacea from tropical, temperate and polar habitats. Polar species were relatively unaffected by a temperature increase, even at temperatures above their ambient range (Young et al. 2006). Although Young et al. (2006) only increased temperatures by a very small margin in comparison with the other studies, this is still relevant in perspective of the very narrow thermal range experience in polar climates. The ability of polar ectotherms to withstand higher temperatures than their living temperatures was also evidenced in a 2014 study which found that crustacea living in the polar environments had a higher critical thermal maximum (CTmax) than their maximum habitat temperature, whereas temperate crustacea had a lower CTmax than their maximum habitat temperature (Faulkner et al. 2014). Indeed when the temperate species Cancer magister was subjected to temperatures above its ambient water temperatures (11-13°C) there was a complete failure of neuromuscular transmission (Florey ; Hoyle, 1976). The ability of polar species to withstand temperatures higher than their environmental range may be due to having a lower thermal dependence, most likely due to physiological adaptations such as the ability to regulate membrane structure, which permits activity at extreme temperatures. As seen in Table 1, both Antarctic species showed no breakdown of motor activity despite temperatures exceeding that of their ambient environmental range (-1.9 to + 1°C) (P?rtner 2002). Despite being able to withstand temperatures higher than that of their normal environmental range, Wells (1979) reported that the upper temperature at which G. antarcticus shows 33% mortality is 11°C. As Young et al. (2006) reported no change in behavioural ability was observed up to 3.5°C, it is expected that between 3.5°C and 11°C, locomotory rate of G. antarcticus would rapidly fall. It is important to take into account thermal range, not just ambient temperatures, when considering behavioural reactions to increasing temperatures. The polar species examined in this example have a narrower range of temperatures throughout which they can perform behaviours. Generally, species acclimatized to lower temperatures have a reduced range of temperatures at which they can perform normal behaviours (Barlow ; Kerr, 1969; McLeese ; Wilder, 1958), suggesting that a penalty of adaptation to low temperatures is a reduction in scope of behavioural ability (Somero et al. 1996; Wilson et al. 2001), evidencing the high cost of living in the polar regions.
The capacity of some crustacean species to survive polar temperatures has been linked to their ability to regulate magnesium levels in the haemolymph (Thatje ; Arntz, 2004; Thatje et al. 2005) – an ability present in the suborder Natantia, which includes isopods and amphipods, successful in polar waters (Frederich et al. 2001). The availability of oxygen to tissues relies on oxygen distribution via the haemolymph. High, unregulated levels of magnesium restrict circulation, particularly at low temperatures, restricting oxygen availability. Inability to regulate magnesium is a limiting factor for many crustacean species, explaining their absence in polar regions (Frederich et al. 2000).

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Temperate crustacea are able to fully take advantage of increasing temperatures to increase locomotory behaviour (Schmalenbach ; Buchhol, 2013; Lindst?m ; Fortelius, 2001). Temperate species experience the least stable environment, with high variability in temperate water temperatures daily and seasonally. As poikilotherms, crustaceans’ internal body temperature closely follows that of their environment, and so they would have a wide thermal range. Two of the three temperate crustacea referenced in the table above were able to increase locomotory activity up to the maximum temperature tested, evidencing a high upper thermal tolerance.

Tropical species were able to take advantage of higher temperatures to increase locomotion up to a point, after which there are detrimental effects (Florey & Hoyle, 1976; Weinstein & Full, 1994). Tropical crustacea live close to their upper thermal limits, and therefore increased temperatures can only increase locomotory activity up to a point, after which they reach their thermal maxima. Increased temperatures beyond their upper limit can have potentially lethal effects. Similar patterns are observed for reproduction and population growth, for example in population growth of the tropical calenoid copepod, Arcatia sinjiensis (Milione & Zheng, 2008). Population growth of A. sinjiensis was highest at 30°C before decreasing at 34°C. Tropical species living in warm stable climates, would already be living close to their upper thermal limits, and therefore are especially susceptible to increases in temperature (Nguyen et al. 2011).
It is clear that environmental temperatures set the limits for activity (Stillman & Somero, 2000), as seen in the above examples. Consequently, changing global temperatures due to climate change will lead to changing ranges and distribution patterns of species (Van Dijk et al. 1999), with knock on effects on food web stabilities, as species migrate to remain within a thermal range. Understanding how animal’s locomotory ability is affected by temperature change can be used as a means to study their metabolism and to calculate physiological performance curves, which are a useful predictive tool for anthropogenic climate change.

2. Behavioural adaptation and thermal preference.
Temperature is an important determinant of activity and, as ectotherms, crustacea must employ behavioural thermoregulation as they lack endogenous temperate control mechanisms. Mobile crustacea employ behavioural strategies to actively select preferred temperatures in a heterogeneous thermal environment (Re et al, 2012; Dillon et al. 2012). Behavioural thermoregulation allows organisms to remain at temperatures that are energetically favourable and avoid harmful temperatures, and has been observed in a wide range of crustacea (e.g. Crossin et al. 1998; Campos ; van der Veer, 2008). Behavioural thermoregulation and thermal selection varies not only between ectothermic species, but within a species due to factors such as sex (Podrabsky et al. 2008), salinity (Reiser et al. 2017) and season (Clark ; Green, 1991). Behavioural avoidance of harmful temperatures has been seen in all groups of crustacea (Lagerspetz ; Vainio, 2006), and varies between species and dependant on thermal history.
An example of a crustacean utilising behavioural adaptations for temperature avoidance was reported by Nguyen et al (2011). It is a well-studied trend that heat tolerance of marine intertidal animals is related to their vertical distribution on the intertidal shore. A study which investigated the thermal tolerance of 34 marine invertebrate species in 7 phyla, found a positive correlation between upper thermal limits (UTLs) and position on the shore, from subtidal to upper tidal. An anomaly in this study was the upper intertidal crab, Dotilla myctiroides, which had a low UTL, falling in the range of the sub tidal group (Nguyen et al. 2011). This discrepancy can be explained by the behavioural adaptations to thermal stress of this soldier crab, which burrows in the sand for thermal regulation, actively selecting temperatures that are low enough fall into the ‘subtidal’ range. This is an example of how behaviour can be harnessed to inhabit a microhabitat of more favourable temperatures, a key aspect of ectothermic thermal adaptations.
Most scientific studies on behavioural thermoregulation of invertebrates have focused on determining the preferred temperature of a species, and less focused on determining the behavioural thermoregulation used to allow the animal to actively select this temperature. One apparent trend is decreased orthokinesis (minimum motor activity) in the middle of an organisms’ preferred temperature range as observed in a variety of crustacea, from lobsters (McLeese & Wilder, 1958), to freshwater planktonic crustacean (McKenzie et al. 1992). Decreased movement when at the preferred temperature means the organism is most likely to remain at that preferred temperature.
Probably the most important behavioural adaptation of crustacea is simply the ability to avoid potentially harmful temperatures and to move in relation to temperature (Dillion et al. 2012). A 2003 study on the behavioural thermoregulation of the purple shore crab found that animals’ warmed and cooled more rapidly in water than air, and crabs’ behaviour and movement takes advantage of this to mediate internal temperatures (Mcgaw, 2003). Crabs were able to shuttle between air and water to precisely control body temperature within a narrow range, an indispensable behavioural adaptation in their rocky shore environment, which can have a thermal range of 20 °C.

Originally postulated by Fry in 1947, the final thermal preferendum paradigm (FTPP) states that a given species will have a species-specific thermal preference, and all individuals of that species will ultimately aggregate at this certain temperature, regardless of prior thermal history (Fry, 1947). This theory has largely been discredited due to a growing number of studies demonstrating how factors, such as seasonality (Reiser et al. 2016), ontogeny (Lafrance et al. 2005), nutritional state (Pulgar et al. 2003) and gender (Podrabsky et al. 2008) can effect thermal selection by ectotherms. Short-term laboratory experiments have shown various crustacean species to be affected by previous thermal acclimation (e.g. Diaz et al. 2002; Reiser et al. 2014).
Capacity to cope with a wide range of temperatures within a species, such as the shrimp Cangon Cangon, which has a distributional range over a large thermal range, from European coasts and throughout the Mediterranean (Campos & Van der veer, 2008), evidences the importance of acclimation on thermal preference and tolerance, and further discredits the FTPP. Members of this species do not aggregate to a specific temperature, and are able to exploit a wide range of temperatures, over a latitudinal gradient.

Behavioural thermoregulation across latitudinal gradients has been well studied in species of insects (MacLean et al. 2016; Samietz et al. 2005; Addo-Bediako et al. 2000) and reptiles (Artacho et al. 2017; Cadby et al. 2014). Lower thermal limits appear to be more strongly coupled with environmental temperature than upper thermal limits (Sunday et al. 2011; Addo-Bediako et al. 2000). Indeed upper thermal limits are often unrelated to lower limits. This may be due to evolutionary constraints on adaptations to higher temperature, or the use of behavioural thermoregulation to buffer the effects of high temperatures, through behaviours such as burrowing or shade seeking. Ability to cope with increasing temperatures may more strongly rely on behavioural adaptations, whereas lower thermal limits are set by physiological capacity, such as magnesium- haemolymph regulation described in section 1, and membrane mobility, described in section 3.
3. Physiological adaptations
Cellular membrane structure is an important aspect of physiological adaptations to temperature. High temperatures affect permeability of membranes, through increasing the fluidity and activity of integral membrane proteins. Extreme high temperatures can cause denaturation of proteins and the break down of membranes. Low temperatures increase rigidity of the membrane, detrimentally decreasing fluidity. In extremely low temperatures intercellular liquid can freeze and form crystals that are highly damaging to the membrane. Crustacea are capable of thermally-mediated regulation of membrane fluidity as an adaptation to changing external temperatures. This homeoviscous adaptation to maintain cellular structure was first described in E. coli by Sinenksy (1974).
Using a range of mechanisms, such as changes in saturated fatty acids and metabolism of phospholipids, crustacea are able to adjust membrane component compositions in relation to changing environmental temperatures (Pruitt, 1990). Phospholipids isolated from copepod species over a latitudinal range demonstrate that animals in temperate seas have more lateral mobility in their membrane components than those in tropical seas (Pruitt, 1990). Temperate sea species would experience higher variation in temperatures, both seasonally and daily, and need to have controlled membrane flexibility to adapt at a cellular level and cope with their dynamic environment. In regards to polar seas copepods, membrane adjustment ability is indispensable, given the extremely low temperatures and potential for fatal intracellular freezing. Indeed, Pruitt noted that the ability to adjust membrane fluidity is most evident in crustacea at extremely low temperatures (Pruitt, 1990). A reliance on physiological adaptations in colder environments is a consequence of the relatively unavoidable low temperatures of polar habitats. In comparison, crustacea in warmer habitats may rely more heavily on behavioural adaptations, such as shade seeking and burrowing.
Not only do species across a latitudinal thermal gradient demonstrate differences in membrane structure and regulatory ability, but also within the same species, in temperate waters experiencing seasonal thermal variation. In a study on the amphipod crustacean, Gammarus spp., there was a drastic difference in the fluidity of phospholipids vesicles – a key membrane component – obtained from animals in the spring and in the summer (Lahdes et al. 2000). Evidently temperature has a major impact on physiochechemical characteristics in this genus, who acclimatize on a cellular level to their environment.
Species in changing, dynamic habitats such as estuaries or the intertidal have developed physiological adaptations to survive in this fluctuating environment. The crab, Carcinus maenas regulates their respiratory metabolism to stabilise oxygen consumption by increasing the amount of oxygen bound to haemocyanin, delaying hypoxia and sustaining cardiac performance over a wide temperature range (Giomi & P?rtner, 2013). This mechanism has been linked to an increase of cardiac work at higher temperatures, allowing maintenance of metabolic scope below critical temperature in the giant tiger prawn, Penaeus monodon (Ern et al., 2014, 2015).

4. Temperature and life history stages
Survival and growth of decapod crustacea strongly depends on water temperatures. The effect of temperature on moulting increment and growth rate has been investigated in a variety of crustacea as listed in Table 2. These studies have important implications for aquaculture, as utilising temperature is commercially beneficial. Studies in this area have focused on commercially relevant species such as Acanthomysis robusta, a mysid which forms the basis of the diet of juvenile Japanese flounder and Paralichthys olivaceu; a target species for stock enhancement (Sudo, 2003). The crucifix crab, Charybds ferriatus is of economic importance in East Asian markets, although despite this commercial importance it is not cultured commercially (Parado-Estepa et al. 2002). If C. ferriatus was to be commercially cultured, research into optimum growth temperature would be invaluable. Also of commercial importance is the blue king crab, Paralithodes platypus, whose Alaskan fisheries population have crashed, due to overfishing and unfavourable environmental conditions effecting recruitment (Dew & McConnaughey, 2005; Orensanz et al. 1998). Stoner et al. (2013) aimed to determine how temperature might be harnessed to redevelop efficient blue king crab hatcheries.
As seen in Table 2, temperate species growth increases right up to the highest trialled temperature. This may be due to temperate species experiencing less stable temperatures in their environment seasonally (and therefore a wider range of temperatures, than species closer to the equator) and so are adapted to effectively take advantage of increased temperatures. This evidences the concept that animals living in thermally stable environments have reduced thermal acclamatory ability (Nguyen et al. 2011). Worryingly regions closer to the poles and the equator (i.e. relatively thermally stable environments) have faster warming rates than the global average (Collins et al. 2010; Liu et al. 2005), meaning animals in these environments, which already have weaker thermal acclamatory ability, are extremely vulnerable under anthropogenic current climate change conditions (Faulkner et al. 2014; Stillman, 2003).

Although temperate and tropical studies investigated different temperatures these are relative to the species’ natural environment and explore the range of possible ambient temperatures. Similar to the previously discussed behavioural effects of increased temperature (section 1), temperate species appear to benefit from temperatures outside their normal range, whereas tropical species already live close to the upper limit of their thermal range and so growth does not increase ‘exponentially’ with temperature. Tropical species appear to have a maximum, not identified in temperate studies. Interestingly the blue king crab does appear to have a maximum growth temperature (8°C) (Stoner et al. 2013), after which growth levels off. This is more similar to the tropical species, which may be to do with environmental thermal stability in P. platypus’ natural habitat, with populations centred in the colder, and seasonally more stable waters of the Gulf of Alaska and the Bering Sea (Armstrong, 1981).
The temperature-size rule (TSR), a well-established phenomenon, dictates that although higher temperatures may increase the speed of growth by decreasing the time between moults, individuals at lower temperatures will attain a larger final size upon maturity (Atkinson & Sibley, 1997; Hartnoll, 2001). Simply put, in colder temperatures, crustacea will live longer and reach a larger size. This rule is considered an important indicator for climate change, and variations in body size can be used as a gauge of even minor environmental thermal change (Daufresne et al. 2009). It is unclear why this is the case, although perhaps it can be attributed to disparities between the process which determines the onset of the next moult, and the process which determines the growth. These processes may respond differently to temperature, and independently of each other.
Ontogeny can also effect the impact of temperature on growth. Wyban et al. (1995) studied the effect of temperature on the Pacific white shrimp, Panaeus vannami, by investigating response to temperature in three different size classes; small, medium and large. Wyban et al. (1995) found that small or medium shrimp grow faster at higher water temperature, whereas larger shrimp do not. This is most likely due to planktonic P. vannami larvae habituating mangrove estuaries which are subject to relatively extreme diurnal temperature fluctuations. Thus, small postlarvae and juvenile P. vannami are adapted to highly variable thermal environments, and growth that responds directly to temperature. This ontogenic differentiation is a perfect illustration of a larger scale trend. temperate species generally have a wider thermal range as they experience higher fluctuations in seasonal temperature. Their tropical and polar counterparts have reduced physiological flexibility due to having evolved in more thermally stable environments (Gaston, 2009; P?rtner, 2006; Stevens, 1989). More on this matter will be discussed in section 5. This pattern can also be noted in the microhabitat of the intertidal shore, where heat tolerance is related to vertical distribution on the shore and the varying thermal microclimates due to tidal patterns in each of these areas (Nguyen et al. 2011.)
An important aspect of examining crustacean growth is the moult period, as growth is not continuous, as described in the commonly used von Bertalanffy model (1938) but discrete and biphasic. The biphasic nature of crustacean growth can be used as a foundation to model more appropriate models of crustacean growth, by combining the frequency of moulting with the amount of growth. Such models have been investigated in several crustacea including the Alaskan king crab, Paralithodes camtschatica (McCaughran & Powel 1977) and the Dungeness crab, Cancer magister, (Wainright & Armstrong 1993). While an important starting point for potential aquaculture, these laboratory based models however do not aceknowledge potential environmental factors, particularly temperature, which can greatly impact aspects of moulting and growth (Fowler et al. 1971).
Another important factor involved in the effects of temperature on various life history stages is mobility. Mobile life stages (typically adult stages) are able to rely on behavioural thermoregulation such as actively selecting a preferred temperature through klinokinesis. Immobile life stages (for example planktonic or sessile) may evolve different thermal sensitivities (Kingsolver et al. 2011) and rely more heavily on physiological adaptations. Furthermore, lower temperature can decrease the rate of growth in larval stages, extending the larval period in which they are vulnerable to predation (Criales & Angler, 1986), which can set the distribution limits for many species, as seen in the brown shrimp, Crangon crangon (Campos & van der Veer, 2008). C. crangon larvae’ distributional range reflects temperature conditions as egg incubation time is dependent on water temperature, with increasing water temperature decreasing egg development time (Wear 1974). Longer egg development time would leave C. crangon vulnerable to predation, therefore setting the colder water distribution limits.
In temperate species, seasonality sets the limits for life history stages with recruitment occurring during warmer summer months (Sprung 2001). In the North Sea, Carcinus maenas is the dominant planktonic species. In the southern limits of C. maenus’ distribution, larvae release and recruitment shift from summer to winter months, which brings with it different ecological conditions (Sprung, 2001), for example food availability and different predators. A match of physiological and ecological factors determine larvae survival and therefore sets the southern limits of this temperate species. An equilibrium must be met, between physiological factors, predominantly temperature, for successful larvae release, and efficient ecological factors, such as food availability for larvae survival.
5. Thermal acclimation
Crustacea can rely on thermal acclimation ability to survive temperature extremes. Acclimation plays an important role in the preferred temperature range of an individual, an aspect that has been examined in a multitude of crustacea such as the Jonah crab Cancer borealis (Lewis & Ayers, 2014), white shrimp Litopenaeus vannamei (Kumlu et al. 2010) and the common brown shrimp, Cangon cangon (Reiser et al. 2014). Ability for thermal acclimation is directly positively related to the preferred or living temperature of a species.
The trend between living temperature and acclimation ability has been noted within a species in laboratory experiments, which demonstrate that artificially induced acclimation temperature affects the preferred temperature of an individual. Lewis and Ayers (2014) found that acclimation temperature played an important role in the preferred temperature of individual Cancer borealis crabs – crab acclimated in the laboratory to higher temperatures had significantly higher preferred temperatures. Temperature gradient preference experiments may be influenced by positive thigmotaxis, whereby crustacea exhibit contact-seeking behaviour and aggregate in corners. Lewis and Ayers (2014) avoided this by partitioning off the corners of the chamber with plastic grating. Acclimation ability investigated in the laboratory however should not be considered categorically correct and definite, as it does not take into account the potential for evolutionary change or potential for behavioural thermal regulation such as thermal avoidance.
Studies have shown that increased acclimation temperature increases critical thermal maximum (CTmax). CTmax is a quantified measurement of an organism’s upper thermal limits, defined as the temperature at which an animal can no longer effectively organise locomotory activity (Cowles ; Bogert, 1944). In decapod crustacea it is commonly measured by the loss of righting response (LRR), which indicates environmental death. The red sea crab, Portunus pelagicus demonstrated increasing CTmax with increasing acclimation temperature (Qari ; Aljarari, 2014). Figure one illustrates six examples of increasing CTmax in correlation with increased acclimation temperature. Interestingly, the highest CTmax observed in temperate animals is similar to the lowest CTmax of acclimated tropical animals, evidencing well-defined temperature limits (Fig. 1). Of the examined studies, temperate species had an upper limit of 34.9°C (Cumilaf et al. 2016), whereas the lower range of tropical crustacea is 36°C (González et al. 2010). There are obviously well-defined temperature limits, dictated by environmental temperatures that prescribe an organisms’ acclimation ability to upper thermal limits.
Acclimation ability responds to environmental temperatures, and for temperate species such as the red sea crab Portunus pelagicus, seasonal temperature is directly linked to their CTmax. Crabs tested during the summer months had significantly higher CTmax and CTmin than in the winter months, demonstrating the high importance of ambient temperature on acclimation ability (Qari & Aljarari, 2014). As ambient seasonality temperatures effect acclimation ability so does thermal variation within crustaceans’ natural environment. Faulkner et al. (2014) investigated the warming responses of five species of crustacea from two distinct thermal regimes; the temperate and variable conditions of South Africa and the colder, but more stable environment of Marion Island in the Sub-Antarctic. Their results demonstrate the temperate species have a weaker ability to tolerate warming than populations from cooler regions. This complies with Janzen’s rule, which stipulates a positive relationship between thermal tolerance range and latitude (see Gaston et al. 2009). In an analysis of ectothermic animal’s thermal tolerance breadths, Sunday et al. (2011) also found that thermal tolerance breadths increase with latitude. This rather simplistic model, however, should not be relied on for climate change related predictions as other factors, such as rates of change, as well as behavioural thermoregulation may further complicate predictions. Indeed, in contradiction, Peck (2010) reports that Antarctic stenotherms – inhabiting high latitude – typically have limited tolerance and plasticity. An explanation of this lies in the climate variability hypothesis, which assumes that a primary determent of a poikilothermic species’ thermal tolerance is the thermal variation experienced in its natural environment. Poikilothermic animals living in temperate environments experience a wider range of temperature variation, whereas lower ranges of temperature variation at tropical latitudes generally leads to narrower tolerance windows (Stevens 1989). In this case invertebrates living in thermally buffered environments with minimal daily or seasonal temperature variation would be expected to exhibit narrow thermal tolerance breadths (Stevens, 1989).
A global analysis of ectothermic thermal tolerances in conjuncture with latitude showed a positive relationship between temperature variability and thermal tolerance breadth (Sunday et al. 2011). Highly stenothermal Antarctic invertebrates have thermal tolerance ranges that are two to four times smaller than those of lower latitude species. Antarctic species living at very stable temperatures of 5°C cannot tolerate a temperature increase of 2°C (Peck et al. 2004), whereas experiments on temperate species, such as the purple shore crab which experiences habitat thermal variation of up to 20°C have shown an ability to with stand temperatures up to 33.6°C (Todd and Dehnel, 1960). Indeed within a species, even small-scale latitudinal variation appears to have an impact on thermal range. Populations of the green shore crab, Carcinus maenas, on the west coast of America exhibited higher CTmax at the southern extent of their distribution than northerly populations, corresponding with sea temperature variation (a range of 12°C in the south and 8°C in the north) (Kelley et al. 2011). Clearly acclimation ability is closely linked to environmental temperatures.

Conclusion
Six conclusions can be drawn from this review. Firstly, and most generally, there are distinct disparities between the thermoregulatory abilities and adaptations of crustacea across a global latitudinal gradient. This is to be expected given the tight linkage between temperature and ectotherm adaptation. Impacting all levels of crustacea biology, temperature has a direct impact on species distribution patterns as well as organismal effects such as locomotory activity and life history stages.
Secondly environmental temperature is reflected in the way crustacea locomotory activity reacts to temperature. Polar crustacea, despite no acute breakdown of motor co-ordination at temperatures above ambient range due to lower thermal dependence, have a small thermal range, reflecting the thermal stability of their environment. Temperate species are able to take advantage of increasing temperatures to increase locomotory activity, reflecting the thermal variability of their environment and their subsequently wide thermal range. Tropical species are similar to polar species in that, they too experience stable temperatures and have a corresponding low thermal range. Furthermore tropical species, living in high temperatures, are already at the upper range of their thermal limits and therefore are unable to take advantage of increased temperatures for increased locomotory activity.
Thirdly, environmental temperature impacts mechanisms by which organisms thermoregulate. Behavioural thermoregulation is an important ability of mobile crustacea in order to avoid harmful temperatures and actively select a preferred temperature. Tropical organisms are more likely to reply on behavioural adaptations such as shade-seeking and burrowing, whereas polar species are more likely to reply on physiochemical adaptations. This pattern is also illustrated in temperate organisms, who have varying physiochemical modifications depending on the season, and corresponding temperatures.
Fourthly, temperature is a key consideration for aquaculture, given the prolific impact of temperature on all crustacea life stages. Temperate species life history are majorly affected by seasonality, in comparison with the stable tropical and polar climates, and seasonality sets the limits for life history stages in temperate crustacea.
Lastly, thermal acclimation ability is directly linked to climate, specifically climate variability. Ectotherms’ body temperature closely follows that of their external environment, and high environmental variability will mean subsequent wide thermal ranges and acclamatory ability. Contrastingly, organisms in stable environment, such as polar and tropical latitudes will have small thermal ranges and low acclamatory ability, which acquires special relevance given the current and threatening issue of climate change. Incidentally polar and temperate regions are the areas predicted to be the worst affected by climate change. Understanding the extent to which organisms, in this case crustacea, will be affected and are capable of adapting to such change is essential for conservation. In particular, understanding the disparity in warming and warming response from crustacea from different environments will be crucial for spatially appropriate conservation and biodiversity management. However, there is perhaps hope in the potential for further evolution, be it behavioural and/or physiological, which may allow crustacea to unexpectedly adapt and survive.
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Tables and Figures
Table 1: Studies on the effect of increasing temperature on locomotory behaviour in crustaceans.

Species Region Temperature change Effect on behaviour Reference
Horned ghost crab, Ocypode ceratophthalma Tropical
Increase up to 37°C Movement increased up to 36°C, but 37°C caused erratic movements and irreversible damage to the neuromuscular system. Florey & Hoyle, 1976
Atlantic ghost crab, Ocypode quadrata Tropical Increased to 24°C Walking speed increased in a linear relationship until 24°C, at which point it decreased. Weinstein & Full, 1994.

Dungeness crab, Cancer magister Temperate Increase to 17°C Increase in movement until complete failure of neuromuscular transmission at 17°C. Florey & Hoyle, 1961
Benthic amphipod, Monoporeia affinisTemperate Increased to 18°C Swim speed increased with temperature, and seemingly maximum swim speed was still not reached at 18°C. Lindst?m & Fortelius, 2001
Common lobster, Hommarus gammusTemperate Increased to 18°C Locomotory activity of lobsters increased with increasing temperature. Schmalenbach & Buchhol, 2013
Benthic isopod, Glyptonotus antarticusAntarctic Increase up to 3.5°C No acute breakdown of motor co-ordination. Young et al. 2006
Antarctic gammarid, Paraceradocus gibber Antarctic Increase up to 3.5°C No acute breakdown of motor co-ordination. Young et al. 2006
Table 2: Studies on the effect of temperature on growth and moult increment in crustaceans.

Species Region Temperature change Result Reference
Ornamental red cherry shrimp, Neocaridina heteropodaTropical Groups held at 24, 28 and 32°C, over a 90 day period Lower temperatures only delay growth. All females at 32°C lost their eggs, indicating negative effects of high temperature on ovarian maturation Tropea et al. 2015
Crucifix crab, Charybdis ferriatusTropical Groups held at temperatures of 20, 26 and 32°C until all larvae and juveniles had reached the next stage of development (around 5 days) Lower temperatures delay the start of development and prolong the moulting period. Baylon & Suzuki, 2007
Pacific white shrimp, Panaeus vannameiTropical Groups held at 23, 27 and 30°C Shrimp growth increased in correlation with temperature, but is size specific i.e. smaller shrimp have a higher preferred temperature than larger shrimp. Wyban et al. 1995
Copepod, Acartia sinjensisTropical Groups held at 10, 15, 20, 25, 28, 30, 34 and 38°C Population growth up to 30°C, before significantly dropping at 34°C, and high mortality at 38°C Milione & Zeng 2008
Epibenthic mysid, Acanthomysis robustaTemperate Groups held at 10, 15, 20 and 25°C Water temperature did not affect moult increment, but did shorten the intermoult period, leading to a faster growth rate. Sudo 2003
Amphipod, Eogammarus sinensisTemperate Groups held at 15, 18, 21, 24 and 27°C Higher temperatures decreased the duration of embryonic development. Xue et al. 2013
Atlantic Blue Crab, Callinectes sapidusTemperate Groups held at 16, 20, 24 and 28°C Intermoult period decreased as temperature increased, up to the maximum temperature Brylawski & Miller, 2006
Blue King crab, Paralithodes platypus Temperate to sub-polar Groups held at 1.5, 4.5, 8 and 12°C Growth rate increased rapidly up to 8°C and then levelled off. The effect of temperature on survival was relatively small. Stoner et al 2013

5429251333596202513335Cold
Temperate
Tropical
Cold
Temperate
Tropical

Figure 1: Acclimation temperatures with corresponding CTmax of tropical and temperate crustaceans. Litopenaeus stylirostris (Re et al. 2006); Litopenaeus vannamei (González et al. 2010); Portunus pelagicus (Qari and Aljarari, 2014); Cancer antennarius (Padilla-Ramírez et al. 2015); Hemigraspus nudus (McGaw, 2003) and Hemigraspus crenulatus (Cumilaf et al. 2016).

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