The control of vascular resistance in the southern rock lobster,Jasus edwardsii (Decapoda: Palinuridae)

In Jasus edwardsii (Hutton) the vascular resistance of each of the seven major arterial systems leaving the heart was increased in response to several of the following neurotransmitters and neurohormones: acetylcholine, adrenalin, serotonin, dopamine, octopamine and peptides proctolin and FLRFamide-related peptide F(1). The resistance to flow through the infrabranchial sinus (IBS), part of the venous system, was also sensitive to these drugs. Unexpectedly, the responses of the IBS continued after removal of the gills. Differences in the profiles of responses of the arteries to individual hormones and in the magnitudes and the time courses of back pressure changes, eliminate a common downstream location such as the venous sinuses or gills, as the source of the arterial responses. Vasoactive drugs were effective when applied either via the lumen or, with longer delay, to the basal side of an artery via the IBS. It is concluded that the resistance of each of these sections of the vascular system is independently controllable by hormones.     

The role of olfaction during mating in the southern temperate spiny lobster Jasus edwardsii

Chemosensory communication may be crucial during mate choice and mating in the southern temperate spiny lobster Jasus edwardsii to ensure that females mate with large males capable of supplying adequate numbers of sperm during the short mating window. To clarify the role of pheromones during this process, three laboratory experiments were carried out. In an experiment where the output of urine, which contains sex-specific pheromones, from large and small catheterized males was switched, large post-molt females did not make a clear choice of mate. This indicates that while females distinguish among females, males, and juveniles using their chemosensory sense, they distinguish among males using visual and tactile senses in combination with olfaction. Further, two antennule-ablation experiments were carried out to determine if detection of pheromones by the antennules of females or males was critical for mate selection, courting, or mating. In both cases, we observed a (nonsignificant) trend of slightly delayed mating of treatment females. We found that disruption of female olfaction causes less impact on courtship or mating than ablation of male antennules which increased the variance in the length of the period between molting and mating and resulted in a systematic reduction in clutch size. This lesser impact of female ablation may be because females can still respond to their own internal cues about egg ripeness whereas males cannot. In J. edwardsii, unlike the American clawed lobster, Homarus americanus, one fully functional partner of either sex appears sufficient to initiate mating.     

Integumental organs of the phyllosoma larva of the rock lobster Jasus edwardsii (Hutton)

The external morphology and distribution of the integumentalorgans of the final-stage phyllosoma of the rock lobster Jasusedwardsii were examined. Seven types of organs were recognizedon the integument of the body trunk, antennules and antennae:plumose setae, simple setae, porous setae, aesthetasc setae,simple pores, dorsal cuticular organs and dome-shaped structures.The plumose setae and simple pores were abundant on the dorsalsurface of the cephalosome, abdomen and telson. The dorsal cuticularorgans were present only on the dorsal surface of the cephalosome,and the aesthetasc setae were restricted to the antennule. A comparison of the morphology of these organs with the senseorgans of other decapods with known function suggests that thedorsal surface of the body trunk is one of the major sites ofreception of near-field water movement in the Jasus phyllosoma.     

Onset of maturity in male southern rock lobsters Jasus edwardsii in Tasmania,Australia

Summary

The reproductive contribution of male southern rock lobsters Jasus edwardsii has recently become of interest for managing this fishery as changes to the length of seasonal closures have shifted fishing effort towards males. To assess size at onset of maturity, a total of 387 male southern rock lobsters Jasus edwardsii were collected using baited traps from four of the eight fishery regions around the Tasmanian coast during July and August 2000. Sizes at onset of maturity in these samples were estimated by examining the vasa deferentia for evidence of the commencement of spermatid production. In addition, the total combined weight of the vasa deferentia and testis was compared with body size (as a gonad somatic index, GSI). Estimates of size at onset of maturity from the two methods were similar, although estimates from GSI appeared to have broader confidence limits. Although these physiological indices of maturity do not necessarily translate to functional maturity, they indicate that maturity is well below that of the legal minimum size of 110 mm carapace length used for managing the fishery (all estimates by either method for length at 50% sample maturity ≤70 mm carapace length). Production of sperm by males occurs below the size at onset of maturity of females from the same region and spatial patterns in maturity estimated by onset of spermatid production were similar to those of females.     

In situ video monitoring of by-catch interactions within commercial rock lobster (Jasus edwardsii) fishing traps

The incidental capture of non-target and undersized target species during commercial fishing operations has been identified as a major issue and ongoing challenge for fisheries management. In the Northern Zone rock lobster (Jasus edwardsii) fishery (NZRLF) of South Australia, escape gaps have been mandatory in all commercial fishing traps since 2002 to reduce non-target species (by-catch) catch rates and juvenile lobster mortality. This study used in situ video monitoring of by-catch species in traps with and without escape gaps to understand how by-catch species interact within commercial fishing traps. Twenty-two different by-catch species were observed entering commercial rock lobster traps, 20 of which were temperate reef finfish. Overall, on-board by-catch rates were significantly higher in hauled traps without escape gaps and were dominated by reef-dwelling finfish species, particularly Meuschenia hippocrepis and Notolabrus tetricus. However, in situ video monitoring within traps showed no difference in the maximum number of finfish by-catch (MaxN) in traps with or without escape gaps. We suggest that the difference in by-catch rates between hauled traps with and without escape gaps may be driven by species-specific behavioural interactions within traps prior to hauling. Overall, this study shows that the implementation of escape gaps in the NZRLF has considerably reduced fishery by-catch rates. Furthermore, the use of in situ video monitoring has provided a better understanding of by-catch behavioural interactions within traps, not previously identified in on-board sampling research, which may partly explain the mechanisms underpinning escape gap effectiveness.     

Temperature effects on metabolic rate and cardiorespiratory physiology of the spiny rock lobster (Jasus edwardsii) during rest,emersion and recovery

Although spiny rock lobster (Jasus edwardsii) is a wholly sub-littoral species, they show a considerable ability to survive prolonged emersion, a fact exploited during the commercial export of this species. Yet, despite this remarkable hardiness, basic information on how this species responds physiologically to emersion is somewhat lacking. Using flow-through respirometry and electrophysiological techniques, we identified that J. edwardsii undergoes marked physiological changes during rest, emersion and recovery over a broad range of temperatures (3.7–17.8 °C). Under resting conditions, routine metabolic rates (RMR) were 22.57 ± 2.39, 9.69 ± 0.55 and 8.09 ± 0.27 mL O₂ h^?1, average heart rates (Hr) were 54.72 ± 4.46, 37.68 ± 2.86 and 29.67 ± 0.59 BPM, and ventilation frequencies were 83.71 ± 5.86, 45.34 ± 2.91 and 41.62 ± 0.65 BPM at 15.0, 7.5 and 3.7 °C, respectively. Notably, the surgical implantation of electrodes elevated RMR compared with non-surgical treatments. In surgery and non-surgery groups, Q ₁₀ was calculated to be ca. 3.0. Upon emersion, rate of oxygen consumption and Hr decreased below resting rates in a temperature-dependent manner, but, along with rate of CO₂ production, increased steadily during 24-h emersion. Ventilation frequencies upon emersion showed a contrasting response and increased significantly above resting rates. When returned to flow-through sea water for recovery, elevated respiration rates provided clear evidence of an O₂ debt, and near-complete recovery was observed after 17 h at both 15.0 and 7.5 °C, but close to no debt was recovered at 3.7 °C. In addition, J. edwardsii was observed to undergo marked diurnal and periodic ventilation cycles, characterised by synchronous changes in RMR, Hr and ventilation frequency.     

Neural development in the planktonic and early benthic stages of the palinurid lobster Jasus edwardsii

Jasus edwardsii has eleven phyllosoma larvae that are morphologically adapted for a plank‐tonic existence. These are followed by a puerulus stage which, although it has the general body form of the adult, is also adapted for swimming. The next moult is the first of the truly benthic juvenile stages. Using transmission electron microscopy we compared transverse sections of connectives in the ventral nerve cord of the first phyllosoma, the puerulus and the first post‐puerulus stage at a number of points. We found that the anterior cord of the phyllosoma has approximately the same number of neurons as the puerulus. This has implications for our understanding of crustacean larval sensory systems because previous work shows that most fibres in the adult nerve cord are sensory. Fibre counts in the post‐puerulus increase in the expected manner. The cord structure of the puerulus and post‐puerulus is the same as that found in the adult whereas that of the early phyllosoma is not.     

Ontogenetic changes in enzyme activities associated with energy production in the spiny lobster, Jasus edwardsii

The larval life of the spiny lobster Jasus edwardsii is one of the longest and most complex of any marine organism and is poorly understood due to the difficulty of studying cryptic, pelagic organisms. Hence, the capacity for active swimming in the phyllosoma, puerulus and juvenile stages and the use of possible metabolic fuel reserves was inferred from a number of enzyme activities, including citrate synthase, lactate dehydrogenase, and HOAD. High activities of CS and LDH in abdominal tissues of Stage 11 phyllosoma and pueruli are consistent with a capacity to commence active on-shore movement. The activities of LDH and HOAD showed positive allometry while CS was independent of body mass. The body mass dependence of LDH activity may reflect the developing ability of the lobster to initiate brief escape manoeuvres, and the scaling of HOAD reflects an increased use of lipid fuel reserves. Aerobic enzyme activities were higher in abdominal tissues than in cephalic tissues of pelagic pueruli, but high activities appear in the cephalic tissues of juveniles. These changes mirror a developmental shift in activity from pelagic oceanic swimming to a benthic existence on the seabed of the near shore. The low LDH activity in pueruli confirmed previous findings that they have limited feeding capacity, with carbohydrate contributing little towards the major energy reserves. The highest LDH activities occur in the abdominal muscles of juveniles and correlate with rapid tail-flicking escape behaviour. The activities of HOAD increased throughout development, and in the abdominal tissues of juveniles, may reflect lipid transformation and accumulation as an energy reserve. Enzyme activities, therefore, provide useful information concerning migratory behaviour that is presently unavailable from ecological studies.     

Haemolymph glucose concentrations of juvenile rock lobsters, Jasus edwardsii, feeding on different carbohydrate diets

Postprandial changes in haemolymph glucose concentration ([Glc]H) were measured in 4-day-fasted juvenile intermoult spiny lobsters, Jasus edwardsii, provided with meals composed of glycogen, maltose, sucrose, glucose, or fructose in a gelatine base, or with gels of the algal glycans agar, alginate and carrageenan. Baseline [Glc]H was 0.61+/-0.02 mmol L(-1). After consumption of glycogen, maltose or sucrose, [Glc]H approximately doubled, peaked after 3 h and returned to baseline between 12 and 24 h. Glucose and fructose meals were followed by periods of sustained hyperglycaemia lasting more than 24 h (peaking at approximately 2.5 times baseline at 6 and 3 h respectively). Suggested explanations for augmented hyperglycaemic responses to glucose and fructose are: 1) these monosaccharides by-passed contact digestion and absorption in the R-cells of the digestive gland, directing them away from storage and toward transepithelial scavenging routes; or 2) glucose and fructose directly elicited release of crustacean hyperglycaemic hormone via a chemosensory reflex. Agar and alginate induced significant postprandial glycaemic responses, consistent with reports of carbohydrases in this species and indicating their potential for inclusion in artificial diets as both binders and energy sources. Carrageenan, a highly sulphated galactan, did not produce a glycaemic response. The measurement of glycaemic responses is a quick method of obtaining nutritional information on carbohydrates considered for inclusion in formulated diets prior to lengthy growth trials.     

Identifying potential prey of the pelagic larvae of the spiny lobster Jasus edwardsii using signature lipids

Signature lipid analyses were used to identify the natural prey of the pelagic phyllosoma larvae of the spiny lobster Jasus edwardsii. Lipid class, fatty acid and sterol composition were determined for associated potential prey items and for phyllosomes captured between 50 and 90 km offshore from the northeastern coast of New Zealand. Phospholipid was the dominant lipid class in all potential prey items and the transparent phyllosomes. The levels of other lipid classes varied between potential prey items, with the next most abundant classes being triacylglycerols, free fatty acids and sterols. A limited number of the potential prey items also contained wax ester. Major fatty acids in all potential prey items were generally 22:6ω3 (docosahexaenoic acid), 16:0, 18:1ω9c, and 20:5ω3 (eicosapentaenoic acid). Multivariate analyses of fatty acid and sterol content of the samples grouped the phyllosoma samples together regardless of their developmental stage or their collection location. However, the phyllosomes were not associated with any of the general groupings of pelagic ascidians, amphipods, chaetognaths, pteropods, euphausiids, fish, copepods or particulate matter that were formed by the statistical analyses. Although the sterol profiles of the potential prey items showed considerable variation, the phyllosomes contained predominately cholesterol, suggesting that other dietary sterols are converted to cholesterol or metabolised by the phyllosomes. Therefore, the use of sterols for tracing the prey of J. edwardsii phyllosoma appears to be limited. Our results suggest that phyllosomes are opportunistic predators that feed on a variety of prey and are preferentially retaining specific diet-derived fatty acids.     

Hydrogen ion titration and amino acid analysis of hemocyanin from the spiny lobster Jasus edwardsii.

Potentiometric and spectrophotometric titrations, isoelectric focusing and amino acid analyses, have been made on the hemocyanin from Jasus edwardsii. Counts of acidic neutral and alkaline groups were made from the titrations, enabling comparisons to be made with the amino acid analysis. Thermodynamic analysis of the data indicated that changes in the native protein structure took place at pH 4.0, 8.4 and 10.7. These observations are discussed in terms of dissociation, shifts in pK and conformational changes in the protein.     

The ontogeny of physiological response to temperature in early stage spiny lobster (Jasus edwardsii) larvae

The physiological response to temperature, in terms of oxygen consumption, nitrogen excretion and feed intake was examined in Jasus edwardsii larvae at mid-stages I-III. From stage I to stage III, the mass-specific oxygen consumption increased in a sigmoid pattern over the temperature range of 10-22 degrees C. The Q(10) value declined significantly from 14-18 to 18-22 degrees C range, indicating a reduced temperature dependence of larval metabolism at higher temperatures. At all stages, feed intake increased with increasing temperature but reached a plateau at the higher temperatures for stages I and II larvae. In contrast, nitrogen excretion increased linearly over this temperature range for all larval stages. Therefore, higher temperatures ( approximately 22 degrees C) may cause an energetic imbalance and reduce growth potential in early stage larvae. While the convection requirement index (quotient of feed intake and oxygen consumption) indicated an equivalent metabolic feeding efficiency from 14 to 22 degrees C, a consistent decline of the O/N ratio above 16-18 degrees C from stage I to stage III suggested that exposure to elevated temperatures may result in an increase in the amount of protein being diverted from growth to catabolic processes. Based on these results, a temperature of 18 degrees C is recommended for the culture of early stage J. edwardsii larvae.     

Metabolic characteristics of muscles in the spiny lobster, Jasus edwardsii, and responses to emersion during simulated live transport

The metabolic characteristics of five muscle groups in the spiny lobster Jasus edwardsii were examined in order to compare their anaerobic and oxidative capacities. Enzyme activities of phosphorylase, phosphofructokinase, pyruvate kinase, and lactate dehydrogenase were highest in abdominal muscles supporting anaerobic burst activity. Hexokinase, citrate synthase, and HOAD activities in the leg and antennal muscles indicated higher aerobic potential. Arginine kinase activities were high in all muscle groups indicating that muscle phosphagens are an important energy reserve. Arginine phosphate concentrations in 4th periopod and abdominal flexor muscle from lobsters sampled in the field were higher than any values from captive animals, and approximately five times those for ATP. Muscle lactates were high in captive animals. Responses to emersion during simulated live transport appear to exploit the capacity for functional anaerobiosis and further differentiated the muscle groups. Abdominal muscles were especially sensitive and after 24 h showed significant increases in lactate, glucose, ADP, and AMP. ATP levels appeared to be maintained by muscle phosphagens and raised doubts about the efficacy of the adenylate energy charge in evaluating the emersion response. Haemolymph glucose, lactic acid, and ammonia peaked after 24 h emersion and were largely restored following re-immersion. We propose that arginine phosphate concentrations in the 4th periopod are an appropriate index of metabolic stress, and could lead to improved commercial handling protocols.     

Temporal variation in the specific dynamic action of juvenile New Zealand rock lobsters, Jasus edwardsii

To enhance the on-growing of Jasus edwardsii in culture, it is important to understand the feeding physiology of juveniles. In crustaceans, there is a loss of energy and an increase in oxygen consumption (specific dynamic action or SDA) associated with feeding. The present research measured the SDA of juvenile J. edwardsii fed either in the morning or at night held at 15 degrees C. Closed box respirometry was used to measure oxygen consumption (MO(2)) and ammonia excretion in juvenile lobsters. Juveniles exhibited a nocturnal rhythm in both MO(2) and ammonia excretion. The factorial rise in MO(2) (1.58+/-0.03 times) for lobsters fed in the morning was significantly less than lobsters fed at night (1.80+/-0.01 times). Lobsters fed in the morning had a significantly shorter SDA (30+/-1.2 h) response compared to lobsters fed at night (36+/-1 h). Energy loss as a result of digestion was less for lobsters fed in the morning. Therefore, if juvenile J. edwardsii are fed in the morning, they could optimise the energy content of the meal and this could result in increased growth.     

The ontogeny of physiological response to light intensity in early stage spiny lobster (Jasus edwardsii) larvae

Early stage phyllosoma larvae of the spiny rock lobster Jasus edwardsii were examined for swimming speed, feeding, oxygen consumption and nitrogen excretion as instantaneous performance indicators when exposed to different irradiance levels. Swimming speed was measured in recently hatched Stage I larvae while all other parameters were measured in larvae from hatch to mid-Stage V. The swimming speed of recently hatched Stage I phyllosoma increased logarithmically between light intensities of 2.9 x 10(14) and 1.8 x 10(16) quanta s(-1) cm(-2) indicating that, within this range, swimming activity was only suppressed at the lowest irradiance level. Larvae examined under dark (no light) conditions showed lower feed intake, oxygen consumption and nitrogen excretion than larvae under low (7.7 x 10(12) q s(-1) cm(-2)) and high (3.9 x 10(14) q s(-1) cm(-2)) light intensities, and this was a consistent pattern observed throughout development from hatch to Stage V. There was no difference in feeding, oxygen consumption and nitrogen excretion between larvae exposed to low and high light intensities. However, from mid-Stage I to mid-Stage V, the metabolic feeding efficiency (feed intake:oxygen consumption ratio) was consistently higher in larvae exposed to low light intensity than in phyllosoma assessed in the dark and under high irradiance. A light intensity of about 7.7 x 10(12) quanta s(-1) cm(-2) and no higher than 3.9 x 10(14) quanta s(-1) cm(-2) is recommended to stimulate feeding and optimise metabolic feeding efficiency in early larval stages of J. edwardsii.     

Lipid reserves used by pueruli of the spiny lobster Jasus edwardsii in crossing the continental shelf of New Zealand.

Lipid is known to fuel the movement of the nektonic puerulus stage of the spiny lobster Jasus edwardsii across the continental shelf of New Zealand. Lipid class analysis of pueruli caught from two locations across the continental shelf showed that phospholipid predominated (86-96% of total lipid), with only smaller proportions of sterol (0.9-8.7%) and diacylglycerol (1.2-7.6%). Only traces of triacylglycerol, hydrocarbon and wax ester were present (<0.1% of total lipid). Comparison of the lipid class content of pueruli caught onshore and offshore showed that phospholipid reserves are primarily utilised during this important phase in the lifecycle and that diacylglycerol plays a less significant secondary role. Histology identified concentrations of phospholipid in fat bodies located in the haemocoel. The use of phospholipid as the dominant storage medium in the puerulus stage is unlike many other marine taxa, including crustacea, which tend to use triacylglycerol and wax ester. The use of phospholipid as a storage medium may well be related to its characteristic transparency, an important feature of this nektonic stage of lobster development that is highly vulnerable to pelagic visual predators.     

Agonistic behaviour in juvenile southern rock lobster,Jasus edwardsii (Decapoda,Palinuridae): implications for developing aquaculture

The Southern rock lobster, Jasus edwardsii, is a temperate species of spiny lobster with established well managed fisheries in Australia and New Zealand. It has also been under consideration as a species with aquaculture potential. Agonistic behaviour has important consequences under aquaculture conditions that encompass direct effects, such as damage or death of protagonists, and indirect effects on growth that relate to resource access, principally food and refuge. This study aimed to identify and characterize behaviours and to make a preliminary investigation of their occurrence under tank culture. Juvenile Jasus edwardsii were examined in a flow-through seawater system using a remote video camera system. Twenty-nine behaviours were divided into three sub-groups: aggressive (11), avoidance (6) and others (12). Aggressive behaviours included attacks, pushing, lifting, clasping and carrying an opponent. Avoidance behaviours included moving away in a backwards-, forwards- or side-stepping motion as well as with more vigorous tail flips. These behaviours were components of twelve behavioural groups that described contact, attack and displacement between individuals. Activity was crepuscular with two clear peaks, one in the morning and the other in the evening. The occurrence of behavioural groups was not different between the morning and evening. The frequency of aggressive behaviours was not affected by changes made to stocking density or access to food. The implications of agonistic behaviours are discussed further in relation to developing aquaculture.     

Estimating historical commercial rock lobster (Jasus edwardsii) catch inside Australian State territorial waters for marine protected area assessment: the binomial likelihood method

The rock lobster (Jasus edwardsii) fishery of South Australia is the State’s most valuable fisheries resource with an export value exceeding ~AU$100 million. The fishery operates primarily inshore (<60 m), and overlaps with a series of marine protected areas (MPAs) currently proposed for State territorial waters. As a result, the need to quantify the impact of proposed MPAs on commercial landings of rock lobster within territorial waters is an integral part of the MPA assessment process. Removing fishing effort displaced by MPAs prevents a corresponding increase in exploitation outside protected zones. We describe a binomial likelihood method that utilises historical commercial catch data to estimate catch totals of rock lobster inside South Australian State waters. Lobster catches per km² showed a high level of spatial variation with estimated historical lobster catch in State waters spanning approximately three orders of magnitude. The method identified key areas where high lobster catch (up to 500 kg/km²) overlapped with State waters. Binomial likelihood outputs have particular application to the estimation of net catch loss in situations where fishery buy-back or financial compensation are a considered option as part of the MPA implementation process.     

Biochemical composition during growth and starvation of early larval stages of cultured spiny lobster (Jasus edwardsii) phyllosoma

We examined biochemical changes accompanying feeding and starvation from hatch to Stage VI (day 74 after hatch) in spiny lobster, Jasus edwardsii, phyllosoma larvae. Larval dry weights (dw) increased 17-fold from hatch (80+/-1 microg) to Stage VI (1415+/-44 microg). Larvae starved for 6-11 days at Stages II, IV and VI were 14-40% lighter than their fed counterparts fed enriched Artemia. The increases and losses in total dry weight during feeding and starvation were associated with changes in the content of protein (constituting 31.4-41.7% of dw) and carbohydrate (constituting 2.6-5.3% of dw), while larger changes in lipid content indicated its greater importance as an energy substrate. Lipid content increased from 7.9% of dw at hatch to its highest of 12.5% at Stage IV, but declined by 50% or more during starvation. This suggests that protein, carbohydrate and lipid are all important energy stores, although lipids are catabolized at a greater rate during food deprivation. The principal lipid class was polar lipid (PL; 79-92% of total lipid), followed by sterol (ST; 6-20%), with triacylglycerol and other lipid classes at <2%. PL were catabolized and ST were conserved during starvation. Changes in the fatty acid (FA) profile had mostly occurred before the first moult at day 8 after hatch, with gradual changes thereafter to Stage VI, reflecting their abundance in the Artemia diet. There was some conservation of the major essential FAs, 20:4n-6, 20:5n-3, 22:6n-3, and the FA profile showed large gains in the C(18) polyunsaturated FA, 18:1n-9, 18:2n-6. Ascorbic acid content increased 10-fold from hatch to the end of Stage I (36 and 333 microgg(-1) dw, respectively), while the content at the end of Stage II was higher in fed than that in starved larvae (439 and 174 microgg(-1) dw, respectively). Our study will assist in the development of alternatives to nutritionally incomplete diets, such as live ongrown Artemia, to meet the requirements of phyllosoma in culture.     

Carabid fecundity as affected by extrinsic and intrinsic factors

Summary Eight species of carabids (six of them represented by two subpopulations) were sampled. It appeared that the mean number of ripe eggs in the ovaries could be regressed on the specific body weight (ranging from 20 to 285 mg dry wt.): the mean decreased in heavier species, and was higher in the autumn breeders than in the spring breeders. The rate of disappearance of ripe eggs from ovaries, estimated at the end of the reproductive season, was regarded as the egg deposition rate. It turned out to be inversely correlated with the actual female mobility. The calculated fecundities in the separate species and subpopulations with the same type of development could be satisfactorily regressed on the specific body weight. Thus, the developmental type, the specific body weight, and actual female mobility are involved in egg production and deposition.