Gas exchange characteristics, metabolic rate and water loss of the Heelwalker, Karoophasma biedouwensis (Mantophasmatodea: Austrophasmatidae)

This study presents the first physiological information for a member of the wingless Mantophasmatodea, or Heelwalkers. This species shows cyclic gas exchange with no evidence of a Flutter period (more typical of discontinuous gas exchange in insects) and no indication that the spiracles are fully occluded during quiescent metabolism. Standard metabolic rate at 20 degrees C was 21.32+/-2.73 microl CO(2)h(-1) (mean+/-S.E.), with a Q(10) (10-25 degrees C) of 1.7. Increases in V()CO(2) associated with variation in mass and with trial temperature were modulated by an increase in burst period volume and a decline in cycle frequency. Total water loss rate, determined by infrared gas analysis, was 0.876+/-0.08 mg H(2)Oh(-1) (range 0.602-1.577, n=11) whilst cuticular water loss rate, estimated by linear regression of total water loss rate and metabolic rate, was 0.618+/-0.09 mg H(2)Oh(-1) (range 0.341-1.363, n=11). Respiratory water loss rate was therefore no more than 29% of the total rate of water loss. Both total water loss rate and estimated cuticular water loss rate were significantly repeatable, with intraclass correlation coefficients of 0.745 and 0.553, respectively.     

Water detection in the desert sand scorpion,Paruroctonus mesaensis (Scorpionida,Vaejovidae)

Summary For the sand scorpion, Paruroctonus mesaensis, substrate moisture is a powerful and fast-acting stimulus of discrete behaviors related to localization and imbibitory uptake of water. These behaviors are readily observed in the field and quantified in the laboratory when free-roaming animals encounter sand substrates dampened by small amounts of water. Of 10 behaviors we monitored in laboratory tests, 5 (pedipalp-pull, rototiller-digging, prolonged stops, headstand, and backingup) occurred only after contact with a moistened substrate. These water-stimulated behaviors were selectively blocked when all 8 tarsal leg segments were coated with wax; coverings of the chemosensory pectine appendages had little to no effect. Electrophysiological recordings from chemoreceptor organs on the tarsi showed that neurons innervating the dorsal tarsal organ, were highly sensitive to humid air stimuli while the numerous, poretipped hairs on the ventral surface were responsive to aqueous solutions applied directly to their tips. Selective blocking of the 8 tarsal organs had no effect on water sensitive behavior indicating that the chemosensory hairs mediate detection of substrate moisture. Such localized, sensory triggering of a robust and directed behavior presents a useful model for further neuroethological studies.Abbreviations H headstand - PP pedipalp-pull - RD rototillerdigging - B backing-up - C wall-climbing - P pause - W walk-through - S stop - T turn - R rest - PST pacific standard time - SEM scanning electron microscopy     

Rapid changes in desiccation resistance in Drosophila melanogaster are facilitated by changes in cuticular permeability

Insects can improve their desiccation resistance by one or more of (1) increasing their water content; (2) decreasing water loss rate; or (3) increasing the amount of water able to be lost before death. Female Drosophila melanogaster have previously been reported to increase their resistance to desiccation after a desiccation pre-treatment and recovery, but the mechanism of this increased desiccation resistance has not been explored. We show that female, but not male adult D. melanogaster increased their resistance to desiccation after 1 h of recovery from a 3 to 4.5 h pre-treatment that depletes them of 10% of their water content. The pre-treatment did not result in an increase in water content after recovery, and there is a slight increase in water content at death in pre-treated females (but no change in males), suggesting that the amount of water loss tolerated is not improved. Metabolic rate, measured on individual flies with flow-through respirometry, did not change with pre-treatment. However, a desiccation pre-treatment did result in a reduction in water loss rate, and further investigation indicated that a change in cuticular water loss rate accounted for this decrease. Thus, the observed increase in desiccation resistance appears to be based on a change in cuticular permeability. However, physiological changes in response to the desiccation pre-treatment were similar in male and female, which therefore does not account for the difference in rapid desiccation hardening between the sexes. We speculate that sex differences in fuel use during desiccation may account for the discrepancy.     

Respiratory and cuticular water loss in insects with continuous gas exchange: comparison across five ant species

Respiratory water loss (RWL) in insects showing continuous emission of CO(2) is poorly studied because few methodologies can measure it. Comparisons of RWL between insects showing continuous and discontinuous gas exchange cycles (DGC) are therefore difficult. We used two recently developed methodologies (the hyperoxic switch and correlation between water-loss and CO(2) emission rates) to compare cuticular permeabilities and rates of RWL in five species of ants, the Argentine ant (Linepithema humile) and four common native ant competitors. Our results showed that RWL in groups of ants with moderate levels of activity and continuous gas exchange were similar across the two measurement methods, and were similar to published values on insects showing the DGC. Furthermore, ants exposed to anoxia increased their total water loss rates by 50-150%. These results suggest that spiracular control under continuous gas exchange can be as effective as the DGC in reducing RWL. Finally, the mesic-adapted Argentine ant showed significantly higher rates of water loss and cuticular permeability compared to four ant species native to dry environments. Physiological limitations may therefore be responsible for restricting the distribution of this invasive species in seasonally dry environments.     

Water loss and gas exchange by eggs of Manduca sexta: Trading off costs and benefits

Like all terrestrial organisms, insect eggs face a tradeoff between exchanging metabolic gases (O₂ and CO₂) and conserving water. Here I summarize the physiology underlying this tradeoff and the ecological contexts in which it may be important. The ideas are illustrated primarily by work from my laboratory on eggs of the sphingid moth Manduca sexta. In particular, I discuss: (1) dynamic changes in metabolic demand and water loss during development; and (2) how the eggshell layers and embryonic tracheal system control the traffic of gases between the embryo and its environment. Subsequently, I identify three areas with interesting but unresolved issues: (1) what eggs actually experience in their microclimates, focusing particularly on the leaf microclimates relevant to eggs of M. sexta; (2) how egg experience influences whether or not hatchling larvae succeed in establishing feeding sites on host plants; and (3) whether Hetz and Bradley's [Hetz, S.K., Bradley, T.J., 2005. Insects breathe discontinuously to avoid oxygen toxicity. Nature 433, 516−519] oxygen toxicity hypothesis for discontinuous gas-exchange cycles applies to insect eggs.     

Respiratory evaporative water loss during hovering and forward flight in hummingbirds

Hummingbirds represent an end point for small body size and water flux in vertebrates. We explored the role evaporative water loss (EWL) plays in management of their large water pool and its use in dissipating metabolic heat. We measured respiratory evaporative water loss (REWL) in hovering hummingbirds in the field (6 species) and over a range of speeds in a wind tunnel (1 species) using an open-circuit mask respirometry system. Hovering REWL during the active period was positively correlated with operative temperature (T_e) likely due to some combination of an increase in the vapor-pressure deficit, increase in lung ventilation rate, and reduced importance of dry heat transfer at higher T_e. In rufous hummingbirds (Selasphorus rufus; 3.3 g) REWL during forward flight at 6 and 10 m/s was less than half the value for hovering. The proportion of total dissipated heat (TDH) accounted for by REWL during hovering at T_e > 40 °C was < 40% in most species. During forward flight in S. rufus the proportion of TDH accounted for by REWL was ~ 35% less than for hovering. REWL in hummingbirds is a relatively small component of the water budget compared with other bird species (< 20%) so cutaneous evaporative water loss and dry heat transfer must contribute significantly to thermal balance in hummingbirds.     

The ultrastructure of hemocytopoietic organs in the desert scorpion, Paruroctonus

The light and electron microscopes were used to examine possible hemocytopoietic tissue in the desert scorpion, Paruroctonus mesaensis. Results agree with earlier light microscopic studies that cells are released into the blood from the two lateral lymphoid organs and the supraneural gland. The former are sacciform structures attached by their anterior ends to the diaphragm. The supraneural gland forms the thickened wall of the supraneural artery in the mesosoma from the first to the third abdominal ganglia. The lateral lymphoid glands have an acellular stroma in which are embedded granular and agranular cells. The stroma is apparently formed by specialized cells which release membranous cell fragments that become the matrix of the gland. Cells are released into the body cavity from the periphery of the two organs. The supraneural gland has a fibrous stroma in which are embedded a variety of cell types. The cells appear to be released in greatest abundance into the blood in the lumen of the gland. The gland has cells with opaque granules (0.9-1.4 micron diameter) and agranular cells of variable shape. The most abundant cell, possibly the stem-cell for the others, is about 10 micron diameter and often has processes of variable length. In addition, muscle cells at various stages of differentiation are found at the inner margin of the gland. These cells have thick and thin myofilaments (24-32 and 5-8 nm diameter) and dense bodies which sometimes become organized into sarcomeres with Z-bands before the cells are released into the gland lumen. The function of these muscle cells is unknown, but possibly they contribute to the maintenance of blood pressure and the release of cells into the blood from the inner margin of the gland.     

Respiratory water loss during rest and flight in European Starlings (Sturnus vulgaris)

Respiratory water loss in Starlings (Sturnus vulgaris) at rest and during flight at ambient temperatures (T(amb)) between 6 and 25 degrees C was calculated from respiratory airflow and exhaled air temperature. At rest, breathing frequency f (1.4+/-0.3 Hz) and tidal volume Vt (1.9+/-0.4 ml) were independent of T(amb), but negatively correlated with each other. Mean ventilation at rest was 156+/-28 ml min(-1) at all T(amb). Exhaled air temperature (T(exh)) at rest increased with T(amb) (T(exh) = 0.92.T(amb)+12.45). Respiratory water loss at rest averaged 0.18+/-0.09 ml h(-1) irrespective of T(amb). In flying Starlings f was 4.0+/-0.4 Hz and independent of T(amb). Vt during flight averaged 3.6+/-0.4 ml and increased with T(amb) (Vt = 0.06.T(amb)+2.83) as, correspondingly, did ventilation. T(exh) during flight increased with T(amb) (T(exh) = 0.85.T(amb)+17.29). Respiratory water loss during flight (average REWL(f) = 0.74+/-0.22 ml h(-1)) was significantly higher than at rest and increased with T(amb). Our measurements suggest that respiratory evaporation accounts for most water loss in flying Starlings and increases more than cutaneous evaporation with rising ambient temperature.     

Thermogenic characteristics and evaporative water loss in the tree shrew (Tupaia belangeri)

Thermogenic characteristics and evaporative water loss were measured at different temperatures in Tupaia belangeri. The thermal neutral zone (TNZ) of T. belangeri was 30–35 °C. Mean body temperature was 39.76±0.27 °C and mean body mass was 100.86±9.09 g. Basal metabolic rate (BMR) was 1.38±0.03 ml O₂/g h. Average minimum thermal conductance (C_m) was 0.13±0.01 ml O₂/g h °C. Evaporative water loss in T. belangeri increased when the temperature rose; the maximal evaporative water loss was 3.88±0.41 mg H₂O/g h at 37.5 °C. The results may reflect features of small mammals in the sub-tropical plateau region: T. belangeri had high basal metabolic rate and high total thermal conductance, compared with the predicted values based on their body mass whilst their body temperatures are relatively high; T. belangeri has high levels of evaporative water loss and poor water-retention capacity. Evaporative water loss plays an important role in temperature regulation.     

Breathe softly, beetle: continuous gas exchange, water loss and the role of the subelytral space in the tenebrionid beetle, Eleodes obscura

Flightless, diurnal tenebrionid beetles are commonly found in deserts. They possess a curious morphological adaptation, the subelytral cavity (an air space beneath the fused elytra) the function of which is not completely understood. In the tenebrionid beetle Eleodes obscura, we measured abdominal movements within the subelytral cavity, and the activity of the pygidial cleft (which seals or unseals the subelytral cavity), simultaneously with total CO2 release rate and water loss rate. First, we found that E. obscura has the lowest cuticular permeability measured in flow-through respirometry in an insect (0.90 microg H2O cm(-2) Torr(-1) h(-1)). Second, it does not exhibit a discontinuous gas exchange cycle. Third, we describe the temporal coupling between gas exchange, water loss, subelytral space volume, and the capacity of the subelytral space to exchange gases with its surroundings as indicated by pygidial cleft state. Fourth, we suggest possible mechanisms that may reduce respiratory water loss rates in E. obscura. Finally, we suggest that E. obscura cannot exchange respiratory gases discontinuously because of a morphological constraint (small tracheal or spiracular conductance). This "conductance constraint hypothesis" may help to explain the otherwise puzzling phylogenetic patterns of continuous vs. discontinuous gas exchange observed in tracheate arthropods.     

Sexual dimorphism in desiccation responses of the sand scorpion Smeringurus mesaensis (Vaejovidae)

The osmoregulatory and respiratory responses of male and female Smeringurus mesaensis (Vaejovidae) to prolonged desiccation were measured. No significant effect of sex on mass-loss rates (MLRs) was found. Still, females maintained their haemolymph osmolality when desiccated to 10% mass loss, whereas that of males increased significantly after loss of as little as 5% of initial mass. Females had a 3-fold larger hepatopancreas, significantly higher hepatopancreas water content and higher metabolic rates when adjusted to hepatopancreas-free dry mass. Thus, females not only store more water in the hepatopancreas but also mobilise it to the haemolymph at a higher rate during desiccation, thus maintaining haemolymph osmolality. Gas exchange rates of both males and females decrease as desiccation progresses. An initial respiratory exchange ratio (RER) of approximately 0.9 is followed by a significant increase at mass loss levels of 7.5% and higher. RER values greater than 1.0 may result from partial shift to anaerobic catabolism, which allows closure of the book lung spiracles for longer duration, thus minimising respiratory water loss. The effects of gas exchange rates on rates of water mobilisation between body compartments and water loss to the environment suggest a trade-off between maintaining osmotic stability and conserving body water stores under stressful conditions.     

A new species of scorpion of the genus Neoscorpiops Vachon, 1980 (Scorpiones: Euscorpiidae) from India

A new species of scorpion of the genus Neoscorpiops Vachon, 1980 is described from Northern Maharashtra. Neoscorpiops maharashtraensis sp. nov. is a medium-sized scorpion species ranging from 36 to 42 mm. Carapace, mesosoma and metasoma brown, legs and telson yellow; pedipalp in a shade of brown with carinae dark drown to black. Granulation on carapace coarse, with a few smooth patches; on mesosoma, which is restricted to the posterior half. Anterior margin of carapace with a moderately deep “U”-shaped emargination medially lacking elevated edges. Pectines well developed 7/7. Trichobothria on patella ventral 13–16. Pedipalp chela narrow and elongate in males, length-to-width ratio in males is 4.86–5.05.     

Metabolism,thermoregulation and evaporative water loss in the Chaotung Vole (Eothenomys olitor) in Yunnan-Kweichow Plateau in summer

Proper adjustment of thermoregulatory mechanisms ensures the survival of mammals when they are subjected to seasonal changes in their natural environment. To understand the physiological and ecological adaptations of Eothenomys olitor, we measured their metabolic rate, thermal conductance, body temperature (T_b) and evaporative water loss at a temperature range of 5–30 °C in summer. The thermal neutral zone (TNZ) of E. olitor was 20–27.5 °C, and the mean body temperature was 35.81±0.15 °C. Basal metabolic rate (BMR) was 2.81±0.11 ml O₂/g h and mean minimum thermal conductance (C_m) was 0.18±0.01 ml O₂/g h °C. Evaporative water loss (EWL) in E. olitor increased when the ambient temperature increased. The maximal evaporative water loss was 6.74±0.19 mg H₂O/g h at 30 °C. These results indicated that E. olitor have relatively high BMR, low body temperature, low lower critical temperature, and normal thermal conductance. EWL plays an inportant role in temperature regulation. These characteristics are closely related to the living habitat of the species, and represent its adaptive strategy to the climate of the Yunnan-Kweichow Plateau, a low-latitude, high-altitude region where annual temperature fluctuations are small, but daily temperature fluctuations are greater.     

A new species of the rare buthid scorpion genus Lissothus Vachon, 1948 from Central Algeria (Scorpiones,Buthidae)

Taxonomic considerations are given for the genus Lissothus Vachon, 1948 (Scorpiones, Buthidae). Two species are currently known, Lissothus bernardi Vachon, 1948 from Libya and Lissothus occidentalis Vachon, 1950 from Mauritania. In this contribution, a new species, Lissothus chaambi sp. n., is described from the desert of Central Algeria. The new species is most closely related to L. bernardi. The geographical distribution of the genus is discussed.     

A model of pupal water loss in Glossina

The results of a long-established investigation into pupal transpiration are used as a rudimentary data set. These data are then generalised to all temperatures and humidities by invoking the property of multiplicative separability, as well as by converting established relationships in terms of constant humidity at fixed temperature, to alternatives in terms of a calculated water loss. In this way a formulation which is a series of very simple, first order, ordinary differential equations is devised. The model is extended to include a variety of Glossina species using their relative surface areas, their relative pupal and puparial loss rates and their different 4th instar excretions. The resulting computational model calculates total, pupal water loss, consequent mortality and emergence. Remaining fat reserves are a more tenuous result.The model suggests that, while conventional wisdom is often correct in dismissing variability in transpiration-related pupal mortality as insignificant, the effects of transpiration can be profound under adverse conditions and for some species, in general. The model demonstrates how two gender effects, the more significant one at the drier extremes of tsetse fly habitat, might arise. The agreement between calculated and measured critical water losses suggests very little difference in the behaviour of the different species.     

Living in a shallow burrow under a rock: Gas exchange and water loss in an Australian scorpion

The scorpion, Urodacus manicatus (Scorpionida: Urodacidae), inhabits temperate sclerophyll woodland in south-eastern Australia and excavates a shallow burrow into soil beneath a rock. Mean minimum and maximum temperatures within a burrow were respectively higher and lower than outside measurements, and relative humidity within a burrow remained higher than outside the burrow at all times. Using flow-through respirometry, VCO₂ and water loss were measured at three ecologically relevant temperatures (10, 20 and 30 °C) to assess patterns of gas exchange and the temperature dependence of VCO₂ and water loss rates. Daytime VCO₂ corresponds to a lower metabolic rate than reported for the closely related, but more arid region inhabiting species Urodacus armatus and Urodacus yaschenkoi. CO₂ output is continuous at 10 °C; however, at 20 °C scattered sharp depressions are observed and these become more frequent and periodic at 30 °C. Total water loss in U. manicatus is nearly double that in U. armatus, indicative of lessened selective pressure for water conservation within a mesic rather than xeric environment.     

Participation of book lungs in evaporative water loss in Paraphysa parvula, a migalomorph spider from Chilean Andes

Small animals need efficient water conservation mechanisms for survival and reproduction, which is relevant for the spiders that have large book lungs with large respiratory surface. If lung evaporation is relevant to limit water loss, adjustments of the spiracle opening to metabolic demands should be expected. In this study, we measured the metabolic rate and total evaporative water loss mediated by the opening of the spiracles in the migalomorph spider Paraphysa parvula, a resident of fluctuating Mediterranean environments of the mountains of central Chile. We found that the metabolism of P. parvula was similar to other Theraphosidae and low compared to other arthropods. Carbon dioxide production and evaporative water loss increased with temperature, particularly at 40 °C. The total evaporative water loss at 40 °C increased dramatically to about 10 times that found with the lower temperatures. Thus, 40 °C will be the limit temperature for this species after which evaporative water loss starts to become damaging, so it has to avoid it. The exposition to hypercapnic environments had as a consequence an increase in evaporative water loss and the involvement of the book lungs in this loss was about 60%. The possibility of losing water could condition this species to seek temperate and oxygenated shelters under rocks.     

A new high-elevation scorpion species of the genus Scorpiops Peters, 1861 (Scorpiones: Euscorpiidae: Scorpiopinae) from the Himalayas,India

A new high-elevation scorpion species of the genus Scorpiops is described from the Indian state of Himachal Pradesh. Scorpiops spitiensis sp. nov. is the second highest-elevation scorpion species in Asia and the first one from India occurring at elevations above 4200 m. The new species closely resembles Scorpiops petersii, but it can be distinguished from it based on a suit of characters, one of which is the presence of 16 trichobothria on the external aspect of the patella, which is unique to the new species.     

Evaporative water loss and energy metabolic in two small mammals,voles (Eothenomys miletus) and mice (Apodemus chevrieri), in Hengduan mountains region

Evaporative water loss (EWL) and energy metabolism were measured at different temperatures in Eothenomys miletus and Apodemus chevrieri in dry air. The thermal neutral zone (TNZ) of E. miletus was 22.5–30 °C and that of A. chevrieri was 20–27.5 °C. Mean body temperatures of the two species were 35.75±0.5 and 36.54±0.61 °C. Basal metabolic rates (BMR) were 1.92±0.17 and 2.7±0.5 ml O₂/g h, respectively. Average minimum thermal conductance (C_m) were 0.23±0.08 and 0.25±0.06 ml O₂/g h °C. EWL in E. miletus and A. chevrieri increased with the increase in temperature; the maximal EWL at 35 °C was 4.78±0.6 mg H₂O/g h in E. miletus, and 5.92±0.43 mg H₂O/g h in A. chevrieri. Percentage of evaporative heat loss to total heat production (EHL/HP) increased with the increase in temperature; the maximal EHL/HP was 22.45% at 30 °C in E. miletus, and in A. chevrieri it was 19.96% at 27.5 °C. The results may reflect features of small rodents in the Hengduan mountains region: both E. miletus and A. chevrieri have high levels of BMR and high levels of total thermal conductance, compared with the predicted values based on their body masses, while their body temperatures are relatively low. EWL plays an important role in temperature regulation.