Basking Activity is Modulated by Health State but is Constrained by Conspicuousness to Predators in Male Spanish Terrapins

Aerial basking may have several benefits for freshwater turtles in addition to thermoregulation such as removing parasites from the skin, which would improve health state. However, basking outside of water may be risky because it may expose freshwater turtles to terrestrial predators. Here, we monitored the basking activity of male Spanish terrapins (Mauremys leprosa) in a wild population of the south‐western of the Iberian Peninsula. We also measured body size, health state parameters, parasite prevalence, and limb coloration of these individuals. We aimed to examine whether basking may improve health state of turtles. The results showed that male turtles with higher basking activity were those parasitized by Hepatozoon and that had lower total white blood cell (WBC). This might indicate that turtles in worse health condition increase their time spent basking to improve their immune system. In addition, because basking might be risky, we also expected that turtles with more conspicuous coloration should reduce their basking activity to avoid being detected by potential predators. We found that infected turtles, but not uninfected ones, that spent more time basking also had less bright coloration in limb stripes. Our study provides evidence that basking activity may improve health state of terrapins, but that color conspicuity may increase costs of basking, especially for parasitized individuals. Understanding the balance between the benefits and the costs of basking might be essential for the conservation of freshwater turtle populations.     

Are viviparous lizards from cool climates ever exclusively nocturnal? Evidence for extensive basking in a New Zealand gecko

Viviparity has evolved numerous times among squamate reptiles; however, the combination of viviparity and nocturnality is apparently rare among lizards. We used time‐lapse photography to examine evidence for diurnal activity in a viviparous lizard often described as nocturnal, the gecko Woodworthia ‘Otago/Southland’ from southern New Zealand (family Diplodactylidae). Evidence for diurnal emergence was extensive. Females have a higher incidence of basking compared to males, although no difference was detected between females in different reproductive conditions. Temperature loggers inserted into calibrated copper models were used to compare the body temperatures available to geckos in two basking positions and in two retreat types. Models in basking positions reached higher mean temperatures than models in retreats, although there was no significant effect of basking position or retreat type on model temperatures. Collectively, our results indicate that pregnant geckos that bask consistently could reduce gestation length by at least 14 days compared with females that remain in retreats. Extensive basking in this species adds to the growing evidence of diurno‐nocturnality in many New Zealand lepidosaurs, including other viviparous geckos. Our results lead us to question whether viviparity in lizards is ever compatible with ‘pure’ nocturnality in a cool climate. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●● , ●●–●●.     

Sea snake skin: Permeable to water but not to sodium

Summary Fasting yellow-bellied sea snakes (Pelamis) have a very low rate of exchange of Na with sea water. Influx and efflux are balanced at a value near 8 moles/100 g h. This is only a fraction of the rate of exchange found in marine fish. Na influx is due to uptake in the head region; dermal and cloacal influx are minimal. The impermeability of the skin to Na has been confirmed in isolated preparations. The outer keratin layer seems to be the primary barrier, since the shed skin alone is also impermeable. Na efflux can be increased to 140 m/100 g h by salt injections, and secretion by the sublingual salt gland can account for all of this loss. Fasting snakes are not in water balance in sea water. There is a net loss of water amounting to about 0.4% body wt/day that probably occurs mainly through the skin. The major osmotic problem ofPelamis in sea water seems to be water balance, not salt balance. Differences in salt gland size among sea snakes might be related to differences in skin permeability to water associated with dermal respiration. The importance of the skin as a permeability barrier suggests that the frequent skin shedding of sea snakes may be related to maintenance of low water permeability as well as to prevention of growth by marine fouling organisms.     

Behavioral and Physiological Thermoregulation of Crocodilians

Crocodilians, like other reptiles, regulate their body temperaturesby a combination of behavioral and physiological mechanisms.Behaviorally, they seek warm surface water or bask when cooland avoid overheating by the evaporation of water from theirdorsum, evaporation of water by gaping or by retreating to deep,cool water. Physiologically, crocodilians increase cutaneousthermal conductance by increasing blood flow to the skin (andsubdermal musculature) during warming. This hastens the warmingprocess. Cutaneous blood flow is reduced during general coolingand locally if the body temperature exceeds skin temperature.This enables crocodilians to increase body temperature significantlywhile basking in cool shallow water. Large crocodilians appearto be able to alter their rates of heat exchange to a largerextent than small ones and they can do so with less cardiovascularinvolvement. Large crocodilians, with their lower surface/volumeratio, are capable of producing sufficient metabolic heat toelevate their body temperature above water temperature.     

The energetics of basking behaviour and torpor in a small marsupial exposed to simulated natural conditions

Limited information is available on basking behaviour in torpid mammals and its energetic consequences. We investigated the effects of physiological and behavioural strategies on the energetics of the fat-tailed dunnart (Sminthopsis crassicaudata). Metabolic rate and body temperature during torpor, basking and rest were measured over 24 h in response to simulated environmental conditions: (a) constant ambient temperature (T _a) of 15°C, (b) constant T _a of 15°C with access to a radiant heat lamp, (c) a T _a cycle (range 15–31°C), and (d) a T _a cycle with access to a radiant heat lamp. When a radiant heat source was provided, all dunnarts (n = 16) basked during all measurements, which resulted in energy savings of up to 74% during rest. Overall, torpor was used on 59% of measurements with a maximum duration of 16.2 h and reductions in metabolic rate of 90% compared to normothermic values. Torpid dunnarts actively moved from a shaded area to position themselves under the heat lamp with body temperatures as low as 17.5°C and thereby reduced rewarming costs by 66%. We demonstrated, for the first time in the laboratory, that torpid animals actively move to a heat source to bask, and that this behaviour results in considerable energy savings. Our finding supports the view that basking during normothermia and rewarming from torpor substantially reduces energetic requirements, which may be important for the survival of small dasyurids living on limited resources in the Australian arid zone.     

Seasonal,environmental and anthropogenic influences on nocturnal basking in turtles and crocodiles from North-Eastern Australia

Many ectotherms bask in the sun as a behavioural mechanism to increase body temperature and facilitate metabolism, digestion or gamete production, among other functions. Such behaviours are common during the day, but some nocturnal species are also known to thermoregulate at night, in the absence of solar radiation, through shifts in body posture or microhabitat selection. Additionally, recent work has documented nocturnal basking in freshwater turtles in tropical Australia, though the purpose of the behaviour remains unknown. Here, we have built upon that work to test: 1. seasonal differences, 2. the influence of environmental factors and 3. the influence of anthropogenic development (e.g. river-front houses) on nocturnal basking behaviour. We visually surveyed transects repeatedly at night on the Ross River, Townsville, QLD, Australia from March to November 2020 and documented nocturnal basking in both freshwater turtles (Emydura macquarii krefftii) and freshwater crocodiles (Crocodylus johnstoni). For both taxa, we found significantly more nocturnal basking activity during the hotter months. Likewise, water surface temperature significantly influenced nocturnal basking in both taxa, especially when water temperatures were both high and warmer than air temperatures. We propose that nocturnal basking provides a mechanism for thermoregulatory cooling when water temperatures are high (e.g. 30°C) and above-preferred temperatures. After accounting for availability in basking habitat, both turtles and crocodiles basked more frequently on the undeveloped side of the river, suggesting avoidance of human activity or disturbance. This study is the first to document nocturnal basking activity temporally throughout the year as well as the first to identify the influences of environmental factors. Nocturnal thermoregulation has been documented in many reptiles, however, thermoregulatory cooling in tropical systems is less well-known.     

Too cold is better than too hot: Preferred temperatures and basking behaviour in a tropical freshwater turtle

Thermoregulation is critical to the survival of animals. Tropical environments can be particularly thermally challenging as they reach very high, even lethal, temperatures. The thermoregulatory responses of tropical freshwater turtles to these challenges are poorly known. One common thermoregulatory behaviour is diurnal basking, which, for many species, facilitates heat gain. Recently, however, a north-eastern Australian population of Krefft's river turtles (Emydura macquarii krefftii) has been observed basking nocturnally, possibly to allow cooling. To test this, we determined the thermal preference (central 50% of temperatures selected) of E. m. krefftii in an aquatic thermal gradient in the laboratory. We then conducted a manipulative experiment to test the effects of water temperatures, both lower and higher than preferred temperature, on diurnal and nocturnal basking. The preferred temperature range fell between 25.3°C (±SD: 1.5) and 27.6°C (±1.4) during the day, and 25.3°C (±2.4) and 26.8°C (±2.5) at night. Based on this, we exposed turtles to three 24 h water temperature treatments (‘cool’ [23°C], ‘preferred’ [26°C] and ‘warm’ [29°C]) while air temperature remained constant at 26°C. Turtles basked more frequently and for longer periods during both the day and night when water temperatures were above their preferred range (the ‘warm’ treatment). This population frequently encounters aquatic temperatures above the preferred thermal range, and our results support the hypothesis that nocturnal basking is a mechanism for escaping unfavourably warm water. Targeted field studies would be a valuable next step in understanding the seasonal scope of this behaviour in a natural environment.     

Basking and torpor in a rock-dwelling desert marsupial: survival strategies in a resource-poor environment

Australian deserts are characterized by unpredictability, low primary productivity, and high temperature fluctuations. Despite these adverse conditions the diversity of small insectivorous marsupials of the family Dasyuridae is surprisingly high. We quantified the thermal biology of the dasyurid Pseudantechinus madonnellensis (body mass ∼30 g) in the wild to gain some understanding of whether the success of dasyurids in the arid zone may be related to some extent to their use of energy conservation strategies. In winter, most free-ranging Pseudantechinus frequently (58.3% of 131 animal days) entered daily torpor after midnight (mean 0157 hours) in rock crevices when outside ambient temperatures (T _a) were low. Most animals remained torpid until the next morning when they moved while still torpid from rock crevices to sun-exposed basking sites. We visually observed basking during rewarming from torpor (mean commencement at 0943 hours) at body temperatures (T _b) as low as 19.3°C when radiant heat was high and T _a was rising. Basking continued for the rest of the day. Torpor use was not strongly correlated with T _a, but the temporal organization of daily torpor and activity were apparently linked to the thermal characteristics of basking sites. Our study suggests that by frequently employing daily torpor and basking and by appropriately coordinating their thermal biology with that of specific locations in their environment, Pseudantechinus can reduce daily energy expenditure and thus can live and reproduce in a challenging environment. It is likely that the success of other small dasyurids and perhaps many other small mammals living in deserts is linked to employment of torpor and basking for energy conservation.     

Thermal ecology of Pieris butterflies (Lepidoptera: Pieridae): a new mechanism of behavioral thermoregulation

Summary I document a new mechanism for behavioral thermoregulation, not previously described in animals, called reflectance basking. This behavior, described here for Pieris butterflies, involves the use of the wings as solar reflectors that reflect solar radiation onto the body to increase body temperature. Results show that Pieris require thoracic (body) temperature. between 29° and 40° C in order to take off and fly, and achieve these elevated temperatures by basking. Diurnal patterns of population flight activity are closely correlated with patterns of body temperature during basking. Behavioral studies indicate that 1) Pieris orient to solar radiation, 2) they use thermoregulatory postures consistent with reflectance basking, and 3) they do not use the basking postures found in other Pierid butterflies (i.e., the Coliadinae). There are consistent differences in wing angles used in reflectance basking between Pieris in different subgenera. Results are discussed with respect to thermoregulation and wing color in other Pierid butterflies, and suggest that a re-evaluation of the functional significance of melanization in Pieris is needed.     

Thermal physiological ecology of Colias butterflies in flight

Summary As a comparison to the many studies of larger flying insects, we carried out an initial study of heat balance and thermal dependence of flight of a small butterfly (Colias) in a wind tunnel and in the wild.Unlike many larger, or facultatively endothermic insects, Colias do not regulate heat loss by altering hemolymph circulation between thorax and abdomen as a function of body temperature. During flight, thermal excess of the abdomen above ambient temperature is weakly but consistently coupled to that of the thorax. Total heat loss is best expressed as the sum of heat loss from the head and thorex combined plus heat loss from the abdomen because the whole body is not isothermal. Convective cooling is a simple linear function of the square root of air speed from 0.2 to 2.0 m/s in the wind tunnel. Solar heat flux is the main source of heat gain in flight, just as it is the exclusive source for warmup at rest. The balance of heat gain from sunlight versus heat loss from convection and radiation does not appear to change by more than a few percent between the wings-closed basking posture and the variable opening of wings in flight, although several aspects require further study. Heat generation by action of the flight muscles is small (on the order of 100 m W/g tissue) compared to values reported for other strongly flying insects. Colias appears to have only very limited capacity to modulate flight performance. Wing beat frequency varies from 12–19 Hz depending on body mass, air speed, and thoracic temperature. At suboptimal flight temperatures, wing beat frequency increases significantly with thoracic temperature and body mass but is independent of air speed. Within the reported thermal optimum of 35–39°C, wing beat frequency is negatively dependent on air speed at values above 1.5 m/s, but independent of mass and body temperature. Flight preference of butterflies in the wind tunnel is for air speeds of 0.5–1.5 m/s, and no flight occurs at or above 2.5 m/s. Voluntary flight initiation in the wild occurs only at air speeds 1.4 m/s.In the field, Colias fly just above the vegetation at body temperatures of 1–2°C greater than when basking at the top of the vegetation. These measurements are consistent with our findings on low heat gain from muscular activity during flight. Basking temperatures of butterflies sheltered from the wind within the vegetation were 1–2°C greater than flight temperatures at vegetation height.     

Highly‐enhanced larval growth during the cold season mediated by the basking behavior of the butterfly Parnassius citrinarius (Lepidoptera: Papilionidae)

Some insect species are thought to grow quickly, even in low temperatures under natural conditions, presumably by conducting basking behaviors to use sunlight. However, whether basking behavior in fact enhances developmental speed and shortens the larval period in the field has not been determined. Moreover, few studies have examined whether basking is behavioral thermoregulation or simply the result of highly‐heterogeneous heat environments in the field. To examine these issues, we conducted field observations and laboratory experiments using larvae of Parnassius citrinarius Motschulsky, which mature within a short period after the thaw in early spring. First, body temperatures of larvae were measured under sunny and cloudy conditions. Second, larval preference for warmer locations was examined. Finally, we compared the developmental speed of larvae when they basked under field conditions and when did not bask in laboratory conditions under different air temperature regimes. Under sunny conditions, larval body temperature was substantially higher than either the temperature of the host plant or the air temperature, and was equivalent to the temperature of dead leaves, which the larvae used as basking sites. In contrast, no such tendency was observed under cloudy conditions. Larvae exhibited an exclusive preference for warmer locations. Moreover, in the field, despite the low ambient temperature, larvae grew much faster than those reared in the laboratory. These results imply that the basking behavior of P. citrinarius larvae is active thermoregulation to maintain high body temperatures in the cold season.     

Intraspecific alkaloid variation in ladybird eggs and its effects on con- and hetero-specific intraguild predators

Sexual size dimorphism (SSD) is a common phenomenon in animals. In many species females are substantially larger than males. Because body size plays a central role in modulating the body temperature (T _b) of ectotherms, intersexual differences in body size may lead to important intersexual differences in thermoregulation. In addition, because SSD is realized by differences in growth rate and because growth rate is strongly temperature dependent in ectotherms, a conflict between male reproductive behaviour and thermoregulation may affect the expression of SSD. In this study, we investigated the thermal implications of SSD in a reptile exhibiting spectacular female-biased SSD: the northern map turtle (Graptemys geographica). Over three seasons, we collected >150,000 measurements of T _b in free-ranging adult and juvenile northern map turtles using surgically implanted miniature temperature loggers. Northern map turtles exhibited seasonal patterns of thermoregulation typical of reptiles in northern latitudes, but we found that large adult females experienced a lower daily maximum T _b and a narrower daily range of T _b than adult males and small juvenile females. In addition, despite more time spent basking, large adult females were not able to thermoregulate as accurately as small turtles. Our findings strongly suggest that body size limits the ability to thermoregulate accurately in large females. By comparing thermoregulatory patterns between adult males and juvenile females of similar body size, we found no evidence that male reproductive behaviours are an impediment to thermoregulation. We also quantified the thermal significance of basking behaviour. We found, contrary to previous findings, that aerial basking allows northern map turtles to raise their T _b substantially above water temperature, indicating that basking behaviour likely plays an important role in thermoregulation.     

Energy and Mass Exchange Over Incomplete Vegetation Cover

Competition for fresh water between agriculture and domestic and industrial uses is increasing worldwide. This is forcing subsistence and commercial agriculture to produce more with less water. Consequently, it is crucial to properly and efficiently manage water resources. This requires accurate determination of crop water loss into the atmosphere, which is greatly influenced by the exchange of energy and mass between the surface and the atmosphere. Measurement of these exchange processes can best be accomplished by micrometeorological methods. However, most micrometeorological methods are very expensive, difficult to set up, require extensive post-data collection corrections and/or involve a high degree of empiricism. This review discusses estimation of evapotranspiration using relatively inexpensive micrometeorological methods in temperature-variance (TV), surface renewal (SR) and mathematical models. The TV and SR methods use high frequency air temperature measurements above a surface to estimate sensible heat flux (H). The latent heat flux (λE), and hence evapotranspiration, is calculated as a residual of the shortened surface energy balance using measured or estimated net radiation and soil heat flux, assuming surface energy balance closure is met. For crops with incomplete cover, the disadvantage of these methods is that they do not allow separation of evapotranspiration into soil evaporation and plant transpiration. The mathematical models (single- and dual-source) involve a combination of radiation and resistance equations to determine evapotranspiration from inputs of automatic weather station observations. Single-source models (Penman-Monteith type equations) are used to determine evapotranspiration over homogeneous surfaces. The dual-source models, basically an extension of single-source models, determine soil evaporation and plant transpiration separately over heterogeneous or sparse vegetation. These mathematical models have also been modified to accommodate inputs of remotely-sensed radiometric surface temperatures that enable estimation of evapotranspiration on a regional and global scale.     

Do the ornamented osteoderms influence the heat conduction through the skin? A finite element analysis in Crocodylomorpha

In order to assess the implication of the crocodylomorph ornamented osteoderms on the skin conduction during basking, we have performed three dimensional modeling and finite element analyses on a sample which includes both extant dry bones and well-preserved fossils tracing back to the Early Jurassic. In purpose to reveal the possible implication of the superficial ornamentation on the osteoderm heat conduction, we repeated the simulation on an equivalent set of smoothed 3D-modeled osteoderms. The comparison of the results evidenced that the presence of the apical sculpture has no significant impact on the osteoderm global conduction. Furthermore, as we also aimed to assess the influence of the inner bone porosity on the osteoderm conduction, we modified the heat equation parameters so that the 3D-modeled osteoderms successively score the compact and the cancellous bone properties (i.e. mass density, heat capacity, thermal conductivity and thermal diffusivity). Finally, we repeated the analyses using the soft-dermis properties which lead to outline that neither the degree of porosity nor the presence of the osteoderms (in itself) significantly modifies the heat conduction through the crocodylomorph skin. Consequently, as hypothesized by previous authors, if the dermal shield happens to be involved into heat capture during basking for crocodylians, this process must mainly rely on a convective effect based on the osteoderm relative degree of vascularization. This last assumption could thus explain why the crocodylians which produce little metabolic heat would carry an entire vascularized osteoderm shield.     

Thermoregulation and the determinants of heat transfer in Colias butterflies

Summary As a means of exploring behavioral and morphological adaptations for thermoregulation in Colias butterflies, convective heat transfer coefficients of real and model butterflies were measured in a wind tunnel as a function of wind speed and body orientation (yaw angle). Results are reported in terms of a dimensionless heat transfer coefficient (Nusselt number, Nu) and a dimensionless wind speed (Reynolds number, Re), for a wind speed range typical of that experienced by basking Colias in the field. The resultant Nusselt-Reynolds (Nu-Re) plots thus indicate the rates of heat transfer by forced convection as a function of wind speed for particular model geometries.For Reynolds numbers throughout the measured range, Nusselt numbers for C. eurytheme butterflies are consistently lower than those for long cylinders, and are independent of yaw angle. There is significant variation among individual butterflies in heat transfer coefficients throughout the Re range. Model butterflies without artificial fur have Nu-Re relations similar to those for cylinders. Heat transfer in these models depends upon yaw angle, with higher heat transfer at intermediate yaw angles (30–60°); these yaw effects increase with increasing Reynolds number. Models with artificial fur, like real Colias, have Nusselt numbers which are consistently lower than those for models without fur at given Reynolds numbers throughout the Re range. Unlike real Colias, however, the models with fur do show yaw angle effects similar to those for models without fur.The independence of heat loss from yaw angle for real Colias is consistent with field observations indicating no behavioral orientation to wind direction. The presence of fur on the models reduces heat loss but does not affect yaw dependence. The large individual variation in heat transfer coefficients among butterflies is probably due to differences in fur characteristics rather than to differences in wing morphology.Finally, a physical model of a butterfly was constructed which accurately simulates the body temperatures of basking Colias in the field for a variety of radiation and wind velocity conditions. The success of the butterfly simulator in mimicking Colias thermal characteristics confirms our preliminary understanding of the physical bases for and heat transfer mechanisms underlying thermoregulatory adaptations in these butterflies.     

Changes in body core temperatures and heat balance after an abrupt release of lower body negative pressure in humans

Changes in body core temperature (T _cor) and heat balance after an abrupt release of lower body negative pressure (LBNP) were investigated in 5 volunteers under the following conditions: (1) an ambient temperature (T _a) of 20 °C or (2) 35 °C, and (3)T _a of 25 °C with a leg skin temperature of 30°C or (4) 35°C. The leg skin temperature was controlled with water perfusion devices wound around the legs. Rectal (T _re), tympanic (T _ty) and esophageal (T _es) temperatures, skin temperatures (7 sites) and oxygen consumption were measured. The intensity of LBNP was adjusted so that the amount of blood pooled in the legs was the same under all conditions. When a thermal balance was attained during LBNP, application of LBNP was suddenly halted. The skin temperatures increased significantly after the release of LBNP under all conditions, while oxygen consumption hardly changed. The release of LBNP caused significant falls inT _cor s under conditions (1) and (3), but loweredT _cor s very slightly under conditions (2) and (4). The changes inT _es were always more rapid and greater than those ofT _ty andT _re. The falls inT _ty andT _re appeared to be explained by changes in heat balance, whereas the sharp drop ofT _es could not be explained especially during the first 8 min after the release of LBNP. The results suggest that a fall inT _cor after a release of LBNP is attributed to an increase in heat loss due to reflexive skin vasodilation and is dependent on the temperature of venous blood returning from the lower body. It is presumed thatT _es may not be an appropriate indicator forT _cor when venous return changes rapidly.     

Maternal thermoregulation influences offspring viability in a viviparous lizard

Gravid females of the viviparous scincid lizard Eulamprus heatwolei were maintained in the laboratory, with some females allowed to bask for 8 h/day and others for only 2 h/day. Maternal basking regimes influenced the gestation period, and significantly affected the body shapes, activity levels and running speeds of the offspring born to these females. Neonates from females with lesser basking opportunities were relatively short and fat, were very active, and were relatively fast runners. Effects of the embryos' thermal regime on the young lizards' morphology and running speed were still detectable two months after birth. Thus, much of the morphological and behavioral variation among neonatal reptiles may arise from phenotypically plastic responses to the thermal environments experienced during embryonic development, rather than from heritable genetic differences among individuals. Hence, selection on maternal thermoregulatory behavior may be an important avenue for adaptive modifications to neonatal phenotypes in reptiles.     

Functional genomics and sexual differentiation in amphibians

To succeed on land rather than in water, crabs require a suite of physiological and morphological changes, and ultimately the ability to reproduce without access open water. Some species have modified gills to assist in gas exchange but accessory gas exchange organs, usually lungs, occur in many species. In accomplished air-breathers the lung becomes larger and more vascularised with pulmonary vessels directing oxygenated haemolymph to the heart. The relative abundance of O₂ in air promotes relative hypoventilation and thus an internal hypercapnia to drive CO₂ excretion. Land crabs have a dual circulation via either lungs or gills and shunting between the two may depend on respiratory media or exercise state. During their breeding migration on Christmas Island Gecarcoidea natalis maintained arterial Po₂ by branchial O₂ uptake, while pulmonary O₂ pressure was reduced; partly because exercise doubled relative haemolymph flow through the gills. Related species rely on elevated haemocyanin concentration and affinity for O₂ to assist uptake but this compromises unloading at the tissues and thus the aerobic scope of tissues. Aquatic crabs exchange salt and ammonia with water via the gills but in land crabs this is not possible. Birgus latro has adopted uricotelism but other species excrete ammonia in either the urine or as gas. Land crabs minimise urinary salt loss using a filtration-reabsorption system analogous to the kidney. Urine is redirected across the gills where salt reabsorption occurs in systems under hormonal control, although in G. natalis this is stimulatory and in B. latro inhibitory. While crabs occupy a range of habitats from aquatic to terrestrial, these species do not comprise a physiological continuum but across the crab taxa individual species possess appropriate and specific physiological features to survive in their individual habitat.