Ability of some gastropod egg capsules to protect against low-salinity stress

The ability of the egg capsules of three intertidal gastropod species to protect embryos against low-salinity stress was examined. Encapsulated embryos of Ilyanassa obsoleta (Say), Nucella lamellosa (Gmelin), and N. lima (Gmelin) were far more tolerant of transfer to water of reduced salinity than were embryos which had been prematurely removed from egg capsules and transferred to low-salinity sea water directly. However, the walls of N. lamellosa and N. lima capsules were found to be permeable to salts and at least to small carbohydrate molecules (glucose). Correspondingly, indirect evidence for all three species and direct evidence for N. lamellosa indicates that the osmotic concentration of intracapsular fluid declines to near ambient after transfer of egg capsules to dilute medium. Experiments conducted using embryos of N. lamellosa suggest that the egg capsules may protect embryos by reducing the rate at which the osmotic concentration of intracapsular fluid decreases rather than by reducing the magnitude of the decrease.     

Mechanical characterization of an unusual elastic biomaterial from the egg capsules of marine snails (Busycon spp.)

Egg capsule material serves as a putative protection mechanism for developing snail embryos facing the perils of the marine environment. We conducted the first quantitative study of this acellular structural protein with the goals of characterizing its chemical and mechanical properties and the relationship of these properties to its biological protective function. We have found that this protein polymer exhibits long-range elasticity with an interesting recoverable yield evidenced by an order of magnitude decrease in elastic modulus (apparent failure) that begins at 3%-5% strain. This material differs significantly from other common structural proteins such as collagen and elastin in mechanical response to strain. Qualitative similarities in stress/strain behavior to keratin, another common structural protein, are more than coincidental when composition and detailed mechanical quantification are considered. This suggests the possibility of alpha-helical structure and matrix organization that might be similar in these two proteins. Indeed, the egg capsule protein may be closely related to vertebrate keratins such as intermediate filaments. We conclude that while this material's bimodal tensile properties may serve as useful protection against the impact loading egg capsules encounter in the intertidal zone, the full biological importance of these capsules is not known.     

A physico-chemical investigation of the jelly capsules surrounding eggs of the Common frog (Rana temporaria temporaria)

The mucopolysaccharide capsules surrounding eggs of the Common frog, Rana temporaria temporaria L. were investigated. Frogs were obtained from ponds in northern England ranging in altitude from 46 to 838 m. Egg capsules acted as insulators, keeping the centre of an egg mass warmer on average than the surrounding water. The size of an egg capsule and its insulating efficiency varied in different pond waters. Egg capsules in water from highland ponds were larger and more efficient insulators than egg capsules in water from lowland ponds. In a series of laboratory experiments the factors controlling capsular swelling were investigated). The ionic concentration of the fluid surrounding the eggs was found to be the principal factor involved, but the temperature and pH of the fluid and the valency and identity of the ions in solution were also important. The water chemistry of ponds varied with altitude. This accounted for the differential swelling of egg jelly in highland and lowland ponds. The fact that egg capsules are larger in highland pond water may increase egg survival at low temperatures. This is probably fortuitous as there is no evidence to suggest that frogs select their breeding ponds because the water in them will promote capsular swelling.     

Capsule walls as barriers to oxygen availability: Implications for the development of brooded embryos by the estuarine gastropod Crepipatella dilatata (Calyptraeidae)

Encapsulation of developing embryos imposes potential restrictions, because the capsule wall must allow for adequate inward diffusion of oxygen and for increased diffusion of oxygen as metabolic demand increases with continued development. Samples of egg capsules from the gastropod Crepipatella dilatata were used to document surface characteristics, composition of the different capsule wall layers, and alterations in wall thickness during development. The diffusion coefficient and capsule wall permeability were determined experimentally for capsules containing embryos at different developmental stages. We also determined oxygen consumption rates for various embryonic stages and for nurse eggs, which provide food for embryos during development. The capsule wall of C. dilatata possesses 2 differentiated layers: the external capsular wall (ECW) and the internal capsular wall (ICW). The ECW is compact and fibrous, features that remain invariable during development, and lacks surface features that might make some portions of the capsule wall more permeable to oxygen than others. On the other hand, the ICW is initially spongy and thick, but significantly decreases in thickness over time, particularly before the embryos begin feeding on nurse eggs. Although the capsule wall is a serious barrier to diffusion, permeability to oxygen increases over time by 112% due to the dramatic thinning of the inner capsule wall layer. Nurse eggs consume oxygen but at very low rates, supporting the idea that they correspond to living embryonic cells that have stopped their development. Respiration measurements indicated that embryos are initially supplied with enough oxygen within the egg capsules to carry out the activities characteristic of embryogenesis, even though the capsular walls show their maximum thickness and lowest permeability at this time. However, as the embryo develops its velum and becomes more active, capsule wall thickness decreases and capsule permeability to oxygen increases. Correspondingly, the oxygen demands of metamorphosed but still encapsulated specimens are approximately 135% higher than those of pre-metamorphosed sibling embryos.     

Latitudinal variation in biophysical characteristics of avian eggshells to cope with differential effects of solar radiation

Solar radiation is an important driver of animal coloration, not only because of the effects of coloration on body temperature but also because coloration may protect from the deleterious effects of UV radiation. Indeed, dark coloration may protect from UV, but may increase the risk of overheating. In addition, the effect of coloration on thermoregulation should change with egg size, as smaller eggs have higher surface‐volume ratios and greater convective coefficients than larger eggs, so that small eggs can dissipate heat quickly. We tested whether the reflectance of eggshells, egg spottiness, and egg size of the ground‐nesting Kentish plover Charadrius alexandrinus is affected by maximum ambient temperature and solar radiation at breeding sites. We measured reflectance, both in the UV and human visible spectrum, spottiness, and egg size in photographs from a museum collection of plover eggshells. Eggshells of lower reflectance (darker) were found at higher latitudes. However, in southern localities where solar radiation is very high, eggshells are also of dark coloration. Eggshell coloration had no significant relationship with ambient temperature. Spotiness was site‐specific. Small eggs tended to be light‐colored. Thermal constraints may drive the observed spatial variation in eggshell coloration, which may be lighter in lower latitudes to diminish the risk of overheating as a result of higher levels of solar radiation. However, in southern localities with very high levels of UV radiation, eggshells are of dark coloration likely to protect embryos from more intense UV radiation. Egg size exhibited variation in relation to coloration, likely through the effect of surface area‐to‐volume ratios on overheating and cooling rates of eggs. Therefore, differential effects of solar radiation on functions of coloration and size of eggshells may shape latitudinal variations in egg appearance in the Kentish plover.     

Inshore spawning grounds of the squid Doryteuthis gahi suggest the consistent use of defoliated kelp Lessonia trabeculata in Central Chilean waters

Coastal spawning grounds of the squid Doryteuthis gahi were identified for the first time at three sites in Central Chile during 2014 and 2015. A total of 15 egg masses were collected from between 10 and 15?m depth and brought to the laboratory for evaluating capsular, embryonic and when possible, paralarval characters. Egg capsules from all spawning areas were similar in size (～25?mm in length) and number of embryos per capsule (～15). Egg-laying patterns, in addition to field observations, suggest that relatively small D. gahi may be using shallow waters in semi-protected environments to reproduce and spawn. Additionally, the differential use of healthy versus defoliated kelp Lessonia trabeculata suggests that egg-laying females selectively choose substrata that limit mechanical damage to the fragile egg capsules and that also allow for adequate water flow, which could be reduced in a mass of kelp fronds. These findings highlight the opportunistic behaviour of D. gahi which use overgrazed L. trabeculata, a condition that is widespread in areas with high herbivory pressure.     

EFFECTS OF UV RADIATION ON GROWTH AND PHLOROTANNINS IN FUCUS GARDNERI (PHAEOPHYCEAE) JUVENILES AND EMBRYOS1

Diminishing levels of atmospheric ozone are increasing UV stress on intertidal algae. Early developmental stages tend to be more susceptible to environmental stresses; however, little research has examined how these stages are protected from UV radiation (UVR). Many brown algae contain high levels of phlorotannins, which are thought to function in screening UVR. In this study, we tested the effects of ambient levels of UV‐B and UV‐A on growth and phlorotannin production in 1‐ to 2‐cm juvenile and microscopic postsettlement embryos of the intertidal alga Fucus gardneri Silva. Algae were grown in four light treatments: 1) ambient light; 2) under cellulose acetate, which lowered light quantity but did not affect light quality; 3) under Mylar^TM, which filtered UV‐B; and 4) under Plexiglas^TM, which blocked UV‐A and UV‐B. Over a 3‐week period, UV‐B inhibited and UV‐A enhanced the growth of F. gardneri embryos, whereas the growth of juveniles was not affected. Phlorotannin concentrations of both embryos and juveniles did not differ in any of the light treatments. Our results suggest that embryos of F. gardneri are susceptible to UV light but develop a tolerance to it as they mature. This tolerance may result from increases in phlorotannin concentrations that occur during maturation; however, phlorotannin production in embryonic or juvenile stages is either not induced by UV light or takes more than 3 weeks to occur.     

Development and the influence of nurse egg allotment on hatching size in Searlesia dira (Reeve, 1846) (Prosobranchia : Buccinidae)

The reproductive biology of the intertidal prosobranch Searlesia dira (Reeve, 1846) was examined with special attention given to variability in the nurse egg to embryo ratio among capsules, among clutches and among geographically isolated populations. Embryos and nurse eggs were distributed among the capsules in a manner consistent with the hypothesis that nurse eggs were genetically predetermined, that each female had a genetically defined nurse egg to embryo ratio, and that each capsule represented a random sample of that ratio. The binomial distribution of embryos and nurse eggs among the capsules resulted in some capsules receiving many more embryos per nurse egg than others. The number of nurse eggs an embryo succeeded in eating was proportional to the number of capsule-mates sharing a capsule. Embryos eating more nurse eggs hatched out at a larger size. Differences in the nurse egg to embryo ratios among capsules in the same clutch were much larger than that of the mean ratios among clutches. Among-site differences in the mean nurse egg to embryo ratios suggest that selection pressure for different mean hatching sizes may have acted on the mean nurse egg to embryo ratios.In contrast to the predictions of optimal hatching size theory, hatching size varied widely within clutches as a consequence of differences in nurse egg to embryo ratios among capsules. This variance may be adaptive for species that lay their eggs months before juveniles emerge into an unpredictable environment, or simply be a consequence of an imperfect mechanism for increasing hatching size.     

Embryonic development and reproductive seasonality of Buccinanops globulosus (Nassariidae) (Kiener, 1834) in Patagonia,Argentina

Buccinanops globulosus mated all year round, with higher frequency from May to September, prior to spawning months. Gravid females were found between October and March. Oviposition peaked during rising temperatures and longest daylength while hatching peaked with high water temperature and declining daylength. Gravid females measured between 20 and 41?mm in shell length. The spawn consisted on average of 31 egg capsules, each containing 1266 eggs. Embryos usually completed development within each egg capsule by ingesting small fragments of the uncleaved nurse eggs, which were not a limiting resource. Egg capsules with more than one embryo were not common; in those cases, the embryos had different sizes probably related to intracapsular competition for nutrients, and were on average smaller than solitary embryos in the other capsules. Embryos hatched as crawling juveniles with a mean hatchling shell length of ~3.4?mm. In a few cases, malformed embryos were found, but it was not a common phenomenon. The information recorded in this study, as the minimum reproductive size and spawning season, is valuable for fisheries management.     

ENCAPSULATION OF EGGS BY MARINE GASTROPODS: EFFECT OF VARIATION IN CAPSULE FORM ON THE VULNERABILITY OF EMBRYOS TO PREDATION

Representatives of many plant and animal taxa enclose their embryos within some form of protective structure. Inter- and intraspecific differences in the morphology of these egg coverings may have profound effects on the development and survival of encapsulated embryos, yet in many taxa little is known about the causes or potential consequences of this variation. Comparisons of capsule morphology among populations of the rocky shore gastropod, Nucella emarginata, revealed significant variation in the thickness of capsule walls, the only barrier separating developing embryos from the external environment. Laboratory experiments demonstrated that thicker-walled capsules were more resistant to predation by a co-occurring isopod, Gnorimosphaeroma oregonense, than were thinner-walled capsules. Control experiments confirmed that these differences in vulnerability were not caused by differences in the palatability of the capsule wall or attractiveness of the capsule contents. The actual mechanism by which thick-walled capsules differentially protect developing embryos remains unclear, although decreased vulnerability of thick-walled capsules to these isopods may simply result from increased handling time by predators. Subtle differences in capsule morphology thus appear to have substantial effects on the survival of encapsulated embryos. Hence, predators may have played an important role in selecting for the production of thick-walled capsules among populations of N. emarginata.     

Hatching plasticity in the tropical gastropod Nerita scabricosta

Hatching plasticity has been documented in diverse terrestrial and freshwater taxa, but in few marine invertebrates. Anecdotal observations over the last 80 years have suggested that intertidal neritid snails may produce encapsulated embryos able to significantly delay hatching. The cause for delays and the cues that trigger hatching are unknown, but temperature, salinity, and wave action have been suggested to play a role. We followed individual egg capsules of Nerita scabricosta in 16 tide pools to document the variation in natural time to hatching and to determine if large delays in hatching occur in the field. Hatching occurred after about 30 d and varied significantly among tide pools in the field. Average time to hatching in each pool was not correlated with presence of potential predators, temperature, salinity, or pool size. We also compared hatching time between egg capsules in the field to those kept in the laboratory at a constant temperature in motionless water, and to those kept in the laboratory with sudden daily water motion and temperature changes. There was no significant difference in the hatching rate between the two laboratory treatments, but capsules took, on average, twice as long to hatch in the laboratory as in the field. Observations of developing embryos showed that embryos in the field develop slowly and continuously until hatching, but embryos in the laboratory reach the hatching stage during the first month of development and remain in stasis after that. Instances of hatching plasticity in benthic marine invertebrates, like the one in N. scabricosta, could greatly enhance our ability to investigate the costs and benefits of benthic versus planktonic development, a long‐standing area of interest for invertebrate larval biologists.     

Chemical defense in the egg masses of benthic invertebrates: an assessment of antibacterial activity in 39 mollusks and 4 polychaetes.

Many marine invertebrates deposit benthic egg masses that are potentially vulnerable to microbial infection. To help counter this threat these species may have evolved some form of chemical protection for their encapsulated embryos. In this study the egg masses from 7 marine mollusks were tested for antibacterial activity against 4 marine pathogens: Enterococcus sericolicida, Vibrio anguillarum, Vibrio alginolyticus, and Vibrio harveyi. Extracts from all of these egg masses were found to inhibit the growth of at least 1 marine bacterium at concentrations that approximate the natural concentration of extract in the egg masses. The egg masses of 39 mollusks and 4 polychaetes were then tested for antibacterial activity against 3 human pathogenic bacteria; Escherichia coli, Staphylococcus aureus, and Pseudomonas aeruginosa. Activity was detected in the egg masses from 34 species, including 2 polychaetes and mollusks from two classes and 18 families. Antibacterial activity in molluskan egg masses was found to extend across the marine, estuarine, freshwater, and terrestrial environments. Both gelatinous egg masses and tough egg capsules were found to inhibit microbial growth, suggesting that physical protection alone may not be sufficient to protect the eggs. Antimicrobial activity was observed in the fresh egg masses but not in the well-developed egg masses of a subset of species. The results of this study indicate that a wide range of invertebrates use chemical defense to protect their early stage embryos against bacterial infection.     

Role of maternal provisioning in controlling interpopulation variation in hatching size in the marine snail Nucella ostrina

This study examined the role of maternal provisioning in controlling interpopulation variation in hatching size in nine isolated populations of the intertidal gastropod Nucella ostrina, in which development to the early juvenile stage takes place within an egg capsule. Variation among populations was almost entirely due to the ratio of nurse eggs to embryo, which explained 65% of the variation in hatching size. Egg size was not a significant predictor of hatching size. Differences among seven of these populations in the nurse egg/embryo ratio were entirely due to the number of nurse eggs allocated per capsule; these populations allocated different numbers of nurse eggs per capsule but allocated the same number of embryos. Intriguingly, the two most wave-sheltered populations allocated significantly more nurse eggs and more embryos to each capsule than did the seven other populations, but they maintained nurse egg/embryo ratios consistent with patterns observed in the other populations. Inter- and intrapopulation variation in hatching size appears to be controlled largely by different mechanisms: within-population variation being controlled mainly by differences in allocation of embryos per capsule, whereas most among-population variation being due to differences in allocation of nurse eggs per capsule.     

Potent phototoxicity of marine bunker oil to translucent herring embryos after prolonged weathering

Pacific herring embryos (Clupea pallasi) spawned three months following the Cosco Busan bunker oil spill in San Francisco Bay showed high rates of late embryonic mortality in the intertidal zone at oiled sites. Dead embryos developed to the hatching stage (e.g. fully pigmented eyes) before suffering extensive tissue deterioration. In contrast, embryos incubated subtidally at oiled sites showed evidence of sublethal oil exposure (petroleum-induced cardiac toxicity) with very low rates of mortality. These field findings suggested an enhancement of oil toxicity through an interaction between oil and another environmental stressor in the intertidal zone, such as higher levels of sunlight-derived ultraviolet (UV) radiation. We tested this hypothesis by exposing herring embryos to both trace levels of weathered Cosco Busan bunker oil and sunlight, with and without protection from UV radiation. Cosco Busan oil and UV co-exposure were both necessary and sufficient to induce an acutely lethal necrotic syndrome in hatching stage embryos that closely mimicked the condition of dead embryos sampled from oiled sites. Tissue levels of known phototoxic polycyclic aromatic compounds were too low to explain the observed degree of phototoxicity, indicating the presence of other unidentified or unmeasured phototoxic compounds derived from bunker oil. These findings provide a parsimonious explanation for the unexpectedly high losses of intertidal herring spawn following the Cosco Busan spill. The chemical composition and associated toxicity of bunker oils should be more thoroughly evaluated to better understand and anticipate the ecological impacts of vessel-derived spills associated with an expanding global transportation network.     

Chondrichthyan egg cases from the south-west Atlantic Ocean

Egg cases of 21 oviparous chondrichthyan species from the south-west Atlantic Ocean are described and compared. The catshark Schroederichthys bivius has a cigar-shaped egg case with curled tendrils only at the posterior end. Egg cases of the elephant fish Callorhinchus callorynchus are spindle-shaped with anterior and posterior tubular extensions and lateral flanges. The skate Amblyraja doellojuradoi presents medium-sized egg cases (71 mm in length) with a lateral keel extending to the first portion of the horns. The endemic skate species of the genus Atlantoraja have medium to large egg cases (69-104 mm in length) and present relatively large posterior horns. Egg cases of the genus Bathyraja have a medium size, 75-98 mm in length, and are characterized by a very similar morphology, a relatively smooth to rough surface case and posterior horns strongly curved inwards. Egg cases of the genera Dipturus and Zearaja are very large, 115-230 mm in length, and have a well-developed posterior apron. Despite the problematical identification of skates at species level, the egg capsules of the endemic genus Psammobatis are easily diagnosed; the capsules are small (25-53 mm in length), those of Psammobatis rutrum being the smallest known to date in the world. Egg cases of Rioraja agassizi have a medium size, 61-68 mm in length, relatively straight sides, a smooth surface and silky attachment fibres placed in the lateral keel next to each horn. Those of the genus Sympterygia are small to medium sized, 51-86 mm in length, and display the thickest lateral keel and the longest posterior horns among the skates of the world. Egg cases can be a useful tool for identifying species and egg-laying areas; therefore, a provisional key for the south-west Atlantic Ocean chondrichthyan capsules is presented.     

Seasonal differences in egg size in three species of crabs from a tropical upwelling zone

Egg size and offspring size are fundamentally important aspects of the life histories of all animals. However the impact of environmental conditions on intraspecific variation in egg size of marine invertebrates is poorly documented. Here we followed three species of intertidal crabs Xanthodius sternberghii, Petrolisthes armatus and Clibanarius albidigitus to understand how seasonal environmental variation in temperature and salinity associated with seasonal upwelling impacts egg size. Ovigerous females of both P. armatus and C. albidigitus were found year round, while X. sternberghii has a limited reproductive season, with ovigerous females found only between November and February. In all three species more than half of the variation in egg size was attributable to variation among broods from different females. Eggs collected during the dry, upwelling season were significantly larger than those collected during the wet, non-upwelling season. Multiple regression analysis showed that average egg size from each brood was significantly negatively correlated with temperature for all three species. Egg size was also negatively correlated with salinity in P. armatus when we controlled for temperature. Overall these results support the idea that changes in environmental temperature caused by seasonal upwelling play a significant role in generating seasonal differences in egg size.     

Effects of UV radiation on five marine microphytobenthic Wadden sea diatoms isolated from the Solthörn tidal flat (Lower Saxony,southern North Sea) – Part I: growth and antioxidative defence strategies

The unconsolidated sediment of intertidal mudflats constitutes a highly unstable environment, due to continuously changing water levels and currents as well as temporary exposure to the air. Therefore, diatoms inhabiting marine intertidal areas are subjected to strongly changing surface light and UV intensities due to exposure at low tide. Five marine intertidal diatoms (Achnanthes exigua, Cocconeis peltoides, Diploneis littoralis, Navicula digitoradiata and Amphora exigua) were isolated from the Solthörn tidal flat (Lower Saxony, southern North Sea). Semi-continuous cultures were used to determine the effect of UV radiation (photosynthetically active radiation only [PAR], PAR+UV-B, PAR+UV-A, PAR+UV-B+UV-A) during short- and long-term exposure (6 h or 30 days). Growth rates, chlorophyll a (chl a), antioxidant capacities, accumulation of phenolic compounds (e.g. flavonoids) and DMSP, and activities of antioxidant enzymes (superoxide dismutase, ascorbate peroxidase, monodehydroascorbate reductase and glutathione reductase) were assessed. UV-A had only minor effects on cells, while growth rate, chl a content and protein content were significantly reduced after long-term UV-B exposure. Achnanthes exigua extracts showed the highest antioxidant capacity. The highest activity of SOD, APX and MDHAR was found under long-term combined UV exposure (PAR+UV-B+UV-A). Overall, the antioxidative defence of the five isolates was stimulated during exposure to UV radiation, as may be found during emersion. Emersion induces oxidative stress and, as a result, growth of the five diatom taxa was inhibited to suit changing environmental conditions. All five taxa tested in the present study showed species-specific acclimatization potentials, providing possible explanations for variability in population, species composition and ecosystem structures in the face of climatic variations.     

Early development of the limpet Siphonaria lessonii Blainville, 1827 in populations affected by different physical stressors

The limpet Siphonaria lessonii is very common along Atlantic Patagonian intertidal rocky shores. We studied the early intracapsular embryonic development of this limpet in detail in two populations in north Patagonia, with different environmental conditions (i.e. wave exposure, wind, temperature). Early development in both populations was achieved at controlled and equal conditions (13°C). The spawn consisted of a series of enchained egg capsules embedded in a jelly mass. The development from egg to hatching veliger took 9–11 days in embryos from both populations. The developmental process at both sites was identical, differing only in the embryos' sizes at each stage. Larger adult individuals producing larger embryos were registered at the sheltered site. The differences in sizes of adult and embryos of S. lessonii could be attributed to distinct environmental stressful conditions between sites.     

Fossilized embryos are widespread but the record is temporally and taxonomically biased

We report new discoveries of embryos and egg capsules from the Lower Cambrian of Siberia, Middle Cambrian of Australia and Lower Ordovician of North America. Together with existing records, embryos have now been recorded from four of the seven continents. However, the new discoveries highlight secular and systematic biases in the fossil record of embryonic stages. The temporal window within which the embryos and egg capsules are found is of relatively short duration; it ends in the Early Ordovician and is roughly coincident with that of typical "Orsten"-type faunas. The reduced occurrence of such fossils has been attributed to reducing levels of phosphate in marine waters during the early Paleozoic, but may also be owing to the increasing depth of sediment mixing by infaunal metazoans. Furthermore, most records younger than the earliest Cambrian are of a single kind-large eggs and embryos of the priapulid-like scalidophoran Markuelia. We explore alternative explanations for the low taxonomic diversity of embryos recovered thus far, including sampling, size, anatomy, ecology, and environment, concluding that the preponderance of Markuelia embryos is due to its precocious development of cuticle at an embryonic stage, predisposing it to preservation through action as a substrate on which microbially mediated precipitation of authigenic calcium phosphate may occur. The fossil record of embryos may be limited to a late Neoproterozoic to early Ordovician snapshot that is subject to dramatic systematic bias. Together, these biases must be considered seriously in attempts to use the fossil record to arbitrate between hypotheses of developmental and life history evolution implicated in the origin of metazoan clades.     

Reproductive biology of the intertidal spider Desis marina (Araneae: Desidae) on a New Zealand rocky shore

Desis marina is an intertidal spider which lives within rock crevices and cavities in the holdfasts of the brown kelp Durvillaea antarctica , where it is submerged in the sea for long periods. Spiders live within silk-lined retreats which enclose an air bubble, and mate location is restricted to periods when the nest is exposed to the air. Eggs are laid from September to January and emergence is complete by May, with the major recruitment period being from March to April. During June to August, females are reproductively inactive. Egg development requires two months and the first two instars remain in the nest for a further two months. Number of egg sacs laid is independent of female size but number of eggs per sac increases with size. Egg size and number of eggs per sac is independent of brood sequence. Female reproductive investment is only 17–6% of body weight, but with 3 to 4 egg sacs/female, nest-guarding time is in excess of five months. Females reproduce only once per year and may reproduce again in the following summer. Desis marina has a much lower clutch size than comparable terrestrial spiders and is a 'bet-hedger', producing sequential broods which spread recruitment over time and reduce the risk of total loss due to storms.