Intertidal burrows of the air-breathing eel goby, <Emphasis Type="Italic">Odontamblyopus lacepedii</Emphasis> (Gobiidae: Amblyopinae)

Odontamblyopus lacepedii inhabits burrows in mudflats and breathes air at the surface opening. Investigations of the intertidal burrows using resin casting demonstrated a highly branched burrow system. The burrows are composed primarily of branching patterns of interconnected tunnels and shafts that communicate into two to seven surface openings. Bulbous chambers (i.e., dilated portions of the burrow) at branching sections of the tunnels or shafts are common features of the burrow. The presence of these chambers accords the fish adequate space to maneuver inside the burrow, and thus constant access to the surface. The combination of all burrow characteristics and previously reported variability in air breathing patterns are ostensibly of selective value for aerial predator avoidance during air breathing in O. lacepedii.     

Burrow structure and use in the sand fiddler crab, Uca pugilator (Bosc)

Uca pugilator, the sand fiddler crab, constructs two kinds of burrows in protected, sandy upper-intertidal and supratidal substrates on the west coast of Florida. Temporary burrows are built and used as a refuge by non-breeding crabs during high tide periods and at night when crabs cease feeding in the intertidal zone. Breeding burrows are constructed and defended by courting males and are the site of mating, oviposition and the incubation of eggs by females. Up to three ovigerous females may be accommodated in a single breeding burrow, each female sequestered in a separate terminal chamber. The construction and defence of burrows specialized for breeding may be an adaptive response by males to the preferences females exhibit when selecting a breeding site.     

Burrow morphology and utilization of the goby (Parapocryptes serperaster) in the Mekong Delta,Vietnam

Some fish species living in mudflats construct burrows for dwelling and hiding. The goby Parapocryptes serperaster is a burrowing fish in mudflats of many estuaries in South East Asia. This study was carried out in the Mekong Delta, Vietnam, to examine burrow morphology and usage by this species. Morphology of the burrows constructed by P. serperaster was investigated by resin castings in situ to obtain the physical structure and configuration of each burrow. Fish from the burrows were caught and measured before burrow casts were made. Fish burrows comprised several openings, a few branching tunnels and multi-bulbous chambers. The surface openings were circular, and the shapes of branching tunnels were nearly round. The burrows had interconnected tunnels and various short cul-de-sac side branches. The burrow structure differed between fish sizes, but burrow dimensions were positively correlated with fish size, indicating that larger fish can make larger and more sophisticated burrow. The burrow structure and dimensions were not different between the dry and wet seasons. Laboratory observations showed that P. serperaster used body movements to dig burrows in the sediment. Burrows could provide a low-tide retreat and protection from predators, but were not used for spawning and feeding for this goby species. This study indicates that the burrowing activity of gobies is an important adaptation for living in shallow and muddy habitats.     

Depth of artificial Burrowing Owl burrows affects thermal suitability and occupancy

Many organizations have installed artificial burrows to help bolster local Burrowing Owl (Athene cunicularia) populations. However, occupancy probability and reproductive success in artificial burrows varies within and among burrow installations. We evaluated the possibility that depth below ground might explain differences in occupancy probability and reproductive success by affecting the temperature of artificial burrows. We measured burrow temperatures from March to July 2010 in 27 artificial burrows in southern California that were buried 15–76 cm below the surface (measured between the surface and the top of the burrow chamber). Burrow depth was one of several characteristics that affected burrow temperature. Burrow temperature decreased by 0.03°C per cm of soil on top of the burrow. The percentage of time that artificial burrows provided a thermal refuge from above‐ground temperature decreased with burrow depth and ranged between 50% and 58% among burrows. The percentage of time that burrow temperature was optimal for incubating females also decreased with burrow depth and ranged between 27% and 100% among burrows. However, the percentage of time that burrow temperature was optimal for unattended eggs increased with burrow depth and ranged between 11% and 95% among burrows. We found no effect of burrow depth on reproductive success across 21 nesting attempts. However, occupancy probability had a non‐linear relationship with burrow depth. The shallowest burrows (15 cm) had a moderate probability of being occupied (0.46), burrows between 28 and 40 cm had the highest probability of being occupied (>0.80), and burrows >53 cm had the lowest probability of being occupied (<0.43). Burrowing Owls may prefer burrows at moderate depths because these burrows provide a thermal refuge from above‐ground temperatures, and are often cool enough to allow females to leave eggs unattended before the onset of full‐time incubation, but not too cool for incubating females that spend most of their time in the burrow during incubation. Our results suggest that depth is an important consideration when installing artificial burrows for Burrowing Owls. However, additional study is needed to determine the possible effects of burrow depth on reproductive success and on possible tradeoffs between the effects of burrow depth on optimal temperature and other factors, such as minimizing the risk of nest predation.     

Semi-monthly reproductive cycles in male and female fiddler crabs, Uca pugnax

In the fiddler crab, Uca pugnax, numbers of displaying males, numbers of burrows with half-dome superstructures (hoods), numbers of females with decalcified vulvar opercula, and numbers of females hatching their eggs, all follow a semi-monthly cycle with peaks near the time of the spring tides. These are all aspects of reproduction in Uca. Decalcification of female opercula, although necessary for mating and egg deposition, is neither necessary nor sufficient for behavioural response to courting males. In fact, behavioural responsiveness to males probably precedes decalcification in most cases. The different aspects of the reproductive cycle are coordinated so that females and males are ready to mate at about the same time near one spring tide, and the resulting eggs will be ready to hatch on the next spring tide. Two decalcified females which were kept isolated from males produced fertile clutches, suggesting that stored sperm from a previous mating may fertilize eggs in a later cycle. This may be one reason for the relative rarity of observed matings or precopulatory behaviour in Uca.     

Lifestyle of Korean mudskipper <Emphasis Type="Italic">Periophthalmus magnuspinnatus</Emphasis> with reference to a congeneric species <Emphasis Type="Italic">Periophthalmus modestus</Emphasis>

To investigate the life history and ecology of the mudskipper Periophthalmus magnuspinnatus, observations and collection were made on coastal mudflats in southern Korea. Periophthalmus magnuspinnatus was active from May to September on the mudflats, exclusively occupying rough and elevated or sloped mudflats of the seashore or the stream mouth, usually vegetated with halophilous grasses. The congeneric species, P. modestus, mainly occurred on extensive low-elevation and level mudflats with no visible vegetation. An apparent alternation of habitat use by P. modestus took place on mudflats at the stream mouth in mid-October, when P. magnuspinnatus began wintering in its burrow and P. modestus came onto the vacated mudflats to construct burrows for wintering. The active season for P. magnuspinnatus at 17°C or higher air temperature was a little shorter than that of P. modestus. Periophthalmus magnuspinnatus constructed a burrow for the entire season in the highest area of the intertidal mudflat, where they hid themselves during high tide or when frightened, whereas P. modestus were likely to use any burrow constructed by other animals or sunken places to hide. The main stomach contents of P. magnuspinnatus were crabs and gammarids. During the reproductive season from May to July, P. magnuspinnatus performed mating behaviors and constructed a spawning burrow similar to the ones known for P. modestus, except their body color turned dark and quivering body movements were observed in the mature male instead of a pink or orange body color and wiggling body movements as in P. modestus. Eggs, measuring 1.56–1.69 mm in major axis and 0.94–1.0 mm in minor axis, were laid on the ceiling and the side wall of the “J”-shaped spawning room generally known for Periophthalmus species. Young of both species started to occur on the mudflat in June.     

Environmental factors affecting the spawning burrow selection by the gobiid Valenciennea longipinnis

Spawning burrow selection by the longfinned goby Valenciennea longipinnis was studied in the near-shore moat on coral reefs, Okinawa, Japan. The gobies make several burrows within their home range, and spawn in one of them. To examine the factors important for spawning burrow selection three characteristics were investigated: current strength, burrow length and effect of underground water on the burrow. Among the burrows, pairs tended to spawn in a larger burrow irrespective of their body sizes. Most of the other non-spawning burrows were too small for a pair to stay together, because hard substrata may prevent the fish from excavating and shaping the burrow as they like. Moreover, pairs preferred to spawn in burrows where the underground water was oozing out, probably because the male's parental burden will decrease due to the higher dissolved oxygen concentration in such burrows. Although current strength may affect a water-exchange in a V. longipinnis burrow in relation to water-exchange function of a mound, it did not affect the spawning burrow selection because of the smaller velocity difference among the burrows relative to the daily fluctuation of tidal current.     

Microhabitat and rhythmic behavior of tiger beetle Callytron yuasai okinawense larvae in a mangrove forest in Japan

Mangrove forests are regularly flooded by tides at intervals of approximately 12.4 h (tidal rhythm). Larvae of the tiger beetle Callytron yuasai okinawense in a mangrove forest made shallow burrows in mounds up to 1 m in height constructed by the mud lobster Thalassina anomala. No larval burrows were observed on the forest floor, which was very muddy even during low tide. Some larvae plugged the burrow openings before they were submerged at high tide. The mean interval between consecutive burrow plugging events was 12.37 h, which is similar to the period of tidal cycles. Nine out of 30 larvae plugged the burrow openings even when the burrows did not become submerged. Plugging behavior may be governed by an endogenous biological clock, or may be a response to exogenous information about tidal level (e.g. moisture seeping through the ground).     

HABITAT DIFFERENCES IN THE TIMING OF REPRODUCTION OF THE INVASIVE ALGA SARGASSUM MUTICUM (PHAEOPHYTA,SARGASSACEAE) OVER TIDAL AND LUNAR CYCLES1

Sargassum muticum (Yendo) Fensholt is an invasive species that is firmly established on intertidal and subtidal rocky shores of Europe and the Pacific coast of North America. Local success and spread of S. muticum is thought to rely on its reproductive potential that seems dependent on exogenous factors like tidal and lunar cycles. This study is the first to compare the reproductive patterns (periodicity of egg expulsion and embryo settlement) of this invader in two different habitats: the middle and low intertidal. The combination of monthly, daily, and tidal samples at triplicate sites within each habitat showed a semilunar periodicity of egg expulsion and embryo settlement coincident with increasing tidal amplitude just before full and new moons. In both habitats, duration of each egg expulsion event was ～1 week, and embryo settlement occurred during the first daily low tide and with the incoming high tide during spring tides. However, both expulsion and settlement started 1–2 d earlier, expulsion saturation was faster, and settlement was higher in the mid‐ compared to the low intertidal. Our results suggest that the exact timing of gamete expulsion and embryo release of S. muticum responds to local factors, including tidal cues, which result in differences between mid‐ and low‐intertidal habitats.     

Direct evidence for aerial egg deposition in the burrows of the Malaysian mudskipper, <Emphasis Type="Italic">Periophthalmodon schlosseri</Emphasis>

The presence of mudskipper eggs in an air-filled chamber was confirmed by direct endoscopic observation of intact burrows of Periophthalmodon schlosseri in a mudflat in Penang, Malaysia. For all five burrows from which video images of egg chambers were successfully obtained, the presence of air was unequivocally demonstrated by the existence of an air–water interface inside the chambers. Of these burrows, eggs were found in two, but not in the others. Eggs were laid uniformly in a monolayer on the inner top surface of the chamber. The much brighter color of the surface mud of the egg chambers than the surrounding mud, irrespective of the presence or absence of the eggs, suggested that the surface mud had been oxidized by deposited air.     

Homing in fiddler crabs (Uca lactea annulipes and Uca vomeris : Ocypodidae)

Fiddler crabs emerge from burrows on intertidal sand- and mudflats to feed during low tide. In the species studied here (Uca lactea annulipes, Uca vomeris) a crab normally wanders no more than about 1 m away from its burrow and, when frightened, dashes back along a straight line to take cover. Feeding crabs tend to move sideways, without changing orientation, along paths radiating from the burrow. When they move along circumferential paths they adjust their orientation so that one side continues to point towards the burrow. The crabs do not need to see the burrow in order to stay aligned with the home vector, and they are not misled by a dummy hole close to their own burrow unless they have come to within about 10 cm of it. The home runs of crabs end within a few centimeters of a burrow that is covered with a sheet of sandpaper and then give way to search runs, centred upon a position slightly short of the burrow location. Feeding crabs can be displaced on sandpapers and their subsequent home runs end at a position where the burrow would be, had there been no displacement. Landmarks close to the burrow do not influence the home runs of displaced crabs. Crabs that are rotated on a sheet of sandpaper, counter-turn to keep their original orientation constant. Fiddler crabs thus employ path integration with external compass information and close range visual guidance for homing. Accepted: 11 May 1998     

Circatidal rhythmic behaviour in the coastal tiger beetle Callytron inspecularis in Japan

Larvae of the coastal tiger beetle Callytron inspecularis (W. Horn) (Coleoptera: Cicindelidae) plug their burrow opening before submergence at high tide. Field observations showed that burrow plugging was a rhythmic behaviour that coincided with the tidal cycle (ca. 12.4 h). On average, larvae plugged their burrows 41.8 min before the tide covered the habitat. The mean interval between consecutive burrow-plugging events in the field was 12.40 h. In the laboratory, in the absence of tidal inundation, the mean interval between consecutive burrow-plugging events was 12.45 h. This suggests that the burrow-plugging rhythm of the coastal tiger beetle is governed by an endogenous circatidal rhythm.     

Nesting activity,breeding success and colony size for the Wedge-tailed Shearwater Puffinus pacificus on Heron Island

Abstract The Capricorn Group of islands in Australia's Great Barrier Reef Marine Park sustains one of the world's largest breeding populations of the Wedge-tailed Shearwater Puffinus pacificus. Heron Island, a 13.5 ha coral cay which supports tourist and research station leases as well as a national park, is the third largest nesting island in the group. Sample censuses of breeding burrows were conducted each year between 1985 and 1990 and a further survey was completed in 1993. These returned estimates of between 13 264±1387 and 16 337±1545 active burrows (Y±SE). Burrow densities within each of the habitats monitored showed no significant trends between years, although there were large differences in burrow density between habitats. There were roughly the same number of burrows in the developed (west) and national park (east) halves of the cay. A miniature video camera system (burrowscope), which allowed nesting chambers at the ends of burrows to be inspected, was used in 1989, 1990 and 1993. This demonstrated that around half the burrows were occupied by incubating birds. Variations were found in the distribution of incubating birds between habitats, although this did not remain constant between the years. In the 1993 season, breeding activity was traced from the burrow establishment to fledging stage. Fifty-one per cent of burrows were used for breeding (eggs laid), 77% of eggs hatched and 80% of chicks produced a fledgling. Overall breeding success for the island was estimated at 61%. In 1993 the area designated as Buildings was found to have significantly lower hatching success compared with natural habitats. Most mortality occurred at the egg stage; however, in the Fringe habitat, mortality was highest at the chick stage. Previous surveys have estimated the breeding population from burrow counts. It now appears that only about 30% of such burrows produce fledglings.     

A new species of Poly dor a (Polychaeta: Spionidae) from the Indo-West Pacific and first record of host hermit crab egg predation by a commensal polydorid worm

A new spionid polychaete, Polydora robi, is described from intertidal and shallow subtidal areas in the Philippine Islands and Bali, Indonesia. Polydora robi belongs to the Polydora ciliata/ websteri species group and is characterized by a rounded prostomium, triangular occipital tentacle, needlelike posterior notosetae, and a pygidium with digitiform composite cirri surrounding the anus. Adults burrow into empty gastropod shells inhabited by hermit crabs. The burrows of the worms typically extend from an external opening in the apex of the shells to an opening in the central body whorls along the columella. The species was found to ingest the fertilized eggs and developing embryos attached to the pleopods of host hermit crabs. The occurrence of egg predation and the symbiotic relationship between polydorids and hermit crabs is discussed. Known egg predators of hermit crabs are reviewed.     

Unusual nonmarine burrows from the Middle Jurassic of Scotland

The Kilmaluag Formation of the Great Estuarine Group (Middle Jurassic) of Scotland represents deposition of mixed carbonate and clastic sediments in a low‐salinity coastal lagoon to floodplain lake setting. Large, unusual trace fossils occur at two horizons within the formation. One type consists of platelike structures about 50 cm in diameter, which are found on wave‐rippled sandstone. These structures, strikingly similar to burrows produced by modern mudskippers, are assigned to fish that shallowly burrowed into the lagoon‐shore sediment.

The second type of burrows, found in brecciated, dolomitic limestones, are pipelike, about 4 to 7 cm in diameter and as much as 50 cm deep. One example has a chamber at the base of the pipe. Although most features of these structures appear similar to modern lungfish burrows, the chamber is most similar to structures produced by modern crayfish. The animal probably burrowed into the moist, mudflat sediment to escape desiccation during seasonal aridity.     

Reproductive ecology of three ocypodid crabs II. Incubation sites and egg mortality

Mortality of eggs during incubation was estimated for three ocypodid crabs,Scopimera globosa, Ilyoplax pusillus andMacrophthalmus japonicus, and the influence of incubation sites was discussed. These crabs all lived in isolated burrows and fed on sediments during day time low tide.S. globosa andI. pusillus inhabited the upper intertidal sandflats, whereasM. japonicus inhabited the lower intertidal mudflats. Females of bothS. globosa andI. pusillus remained in their plugged burrows without feeding throughout incubation and the mortality of eggs was low despite large broods relative to body size. On the other hand, females ofM. japonicus fed actively on surface mud during incubation and the mortality of eggs was high despiite small broods relative to body size. InS. globosa andI. pusillus, the ovaries of ovigerous females were small until egg-hatching, whereas inM. japonicus, the ovaries grew rapidly during incubation and females were able to produce consecutive broods. I conclude that incubation of eggs in burrows may be advantageous in species which inhabit the upper interidal sandflats, even though the crabs cannot forage during incubation, since otherwise their eggs would be exposed to strong heat stress and desication during the summer. Furthermore, such species may produce few large broods because of less frequent interruption of feeding than that associated with production of many small broods.     

The composition and density of fauna utilizing burrow microhabitats created by a non-native burrowing crustacean (<Emphasis Type="Italic">Sphaeroma quoianum</Emphasis>)

The non-native isopod, Sphaeroma quoianum, has invaded many estuaries of the Pacific coast of North America. It creates extensive burrow microhabitats in intertidal and subtidal substrata that provide habitat for estuarine organisms. We sampled burrows to determine the effects of substratum type on the community of inquilines (burrow inhabitants). The density of inquilines was higher in wood and sandstone than marsh banks. Inquilines, representing 58 species from seven phyla, were present in 86% of samples. Inquilines equaled or outnumbered S. quoianum in 49% of the samples. Non-native fauna comprised 29% of the species and 35% of the abundance of inquilines, which is higher than other estuarine habitats in Coos Bay. Sessile non-native species were found living within burrows at tidal heights higher than their typical range. Thus, the novel habitat provided by burrows of S. quoianum may alter the densities and intertidal distribution of both native and non-native estuarine fauna.     

ECOLOGY AND EVOLUTION OF MATING SYSTEMS OF FIDDLER CRABS (GENUS UCA)

1. General accounts of the natural history and behaviour of fiddler crabs suggest there exist two broad mating patterns in the genus. Most western and Indo-Pacific species mate on the surface of intertidal substrates near burrows females defend. The sexes associate only briefly during courtship and mating. In contrast, males of many American species court from and defend burrows to which females come for mating. Copulation occurs underground in burrows plugged at the surface; the sexes usually remain together for at least several hours. Here we summarize and contrast recent detailed field studies of the mating systems of U. pugilator, an American species, and U. vocans, a species widely distributed in the western and Indo-Pacific. We indicate how differences in the breeding ecology of these two species may account for basic differences in modes of sexual selection leading to the two broad mating patterns in the genus. 2. U. pugilator burrows in protected sandy substrates in the upper intertidal and supratidal zone. During ebb tide, nonbreeding crabs leave burrows they occupy during high tide to forage on food-rich substrates in the lower intertidal zone. Reproductively active males remain in the burrow zone where they fight for and defend burrows from which they court. Large males win most fights for burrows and tend to defend burrows high on the elevation gradient, especially during periods with relatively high tides. Females usually approach and descend the burrows of several males before choosing their mates by remaining in males' burrows. Males remain underground with their mates for 1–3 days until after they oviposit their eggs. Some males then emerge and leave their burrows while others sequester their mates in the chambers where mating and oviposition has occurred, dig new chambers and resume courtship, perhaps attracting additional females. In either case, females remain underground for approximately 2 weeks, finally emerging to release their planktonic larvae. Burrows that do not collapse due to tidal inundation or flooding by groundwater are best for breeding and usually are located relatively high on the elevation gradient. Females choose mates indirectly by preferring to breed in burrows that will remain intact while they oviposit and incubate their eggs. Large males mate more often than small males because they are better able to defend burrows at locations females prefer to breed. The mating system of U. pugilator may be classified as resource-defence polygyny. 3. U. vocans burrows in open muddy substrates in the mid- to lower intertidal zone. At a site near Chunda Bay, Australia, where the reproductive behaviour of this species has been studied in depth, both sexes feed near burrows they defend. Females tend to occupy their burrows for longer periods and move shorter distances than do males. Mating occurs on the surface near the burrows that females defend. Females accept both resident and wandering males as mates. They show no preference for mating with larger males. Female choice may be based on other male morphological or behavioural characteristics. Females oviposit their eggs either while on the surface or in their burrows. They produce relatively small clutches and are active on the surface throughout their breeding periods. Males fight both their neighbours and wandering males. Large males tend to win fights and defend burrows in areas where large females, which produce relatively many eggs, are most dense. Such areas may offer greater protection from predators than areas occupied by smaller females. Small males mate about as often as large males but may father fewer larvae. The mating system of U. vocans is resource-free and promiscuous. 4. The mating systems of U. pugilator and U. vocans differ fundamentally in that female U. pugilator require access to a specific microenvironment to breed successfully, while female U. vocans do not. We suggest this difference occurs because of contrasts in clutch sizes and the mobility and movement patterns of feeding females. Female U. pugilator produce relatively large clutches and probably experience more intense selection from factors that can cause egg loss and mortality than do U. oocans, which produce clutches of sufficiently small volume to be protected by their abdominal flaps. Hence, the range of suitable breeding environments for U. pugilator is small compared to that for U. vocans. In addition, U. pugilator burrows in areas that are relatively food-poor, leading to daily migrations to and from food-rich substrates in the lower intertidal zone, preventing female defence of an area suitable for both breeding and feeding. U. vocans, however, burrows in areas sufficiently rich to support feeding, leading to relatively low female mobility and defence of burrows that are also suitable breeding sites. 5. Adaptive radiation of the genus Uca in the Americas is manifest by trends toward smaller adult size, higher population densities, more frequent microgeographic sympatry and increased terrestriality, compared to species in the western and Indo-Pacific regions. We outline the general features of the selection mechanisms tying each of these trends to the evolution of resource—defence mating systems. Intraspecific variation in the courtship behaviour and site of mating in U. lactea and U. vocans supports our contention that resourse—defence behaviour tends to occur at high population densities. Additional data are needed to evaluate the other hypotheses critically.     

Solar incubation cuts down parental care in a burrow nesting tropical shorebird,the crab plover Dromas ardeola

We describe an unknown mode of solar‐assisted egg development in the crab plover Dromas ardeola, a shorebird that breeds in self‐excavated burrows. The insulating properties of the nest burrow and the intense solar radiation allowed egg development at near‐optimal temperature (35.2°C±0.2) and humidity (60.2%±4.4), allowing a very low incubation attendance by the parent birds (28.3% of time, with recesses lasting up to 58 h). Crab plovers did not abandon completely parental incubation, possibly because of the need to turn their egg, and because the slight warming provided by parents (0.8°C) may improve hatching. This is the first case of solar assisted incubation in a species unrelated to the Megapodiidae, the only birds known to develop their eggs without contact incubation.     

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.