Size-related selection of two intertidal gastropods by the reef crab Ozius truncatus

Reef crabs, Ozius truncatus H. Milne-Edwards 1834, and gastropods, Bembicium nanum (Lamarck, 1882) and Nerita atramentosa Reeve, 1855, were collected from a South Australian rocky intertidal platform in April 1981. Laboratory experiments were conducted to determine (1) the size of prey chosen, and (2) the energy gained by the reef crabs feeding on B. nanum and N. atramentosa. The time taken to break open and consume prey increased exponentially with prey shell height. Crabs with larger master claws had shorter handling times for a given prey size. When offered a range of prey sizes, crabs selected prey of a certain size, well below the maximum prey size that they could handle. The mean prey size selected by crabs did not concide with the size predicted to yield the maximum rate of energy gain. For example, one reef crab preferred B. nanum 5 to 8 mm in size, but the size class predicted to yield the greatest energy gain was 4 to 4.9 mm. The results of the laboratory prey-choice experiments are discussed in relation to the shore-level size gradients of B. nanum and N. atramentosa.     

Biotic resistance to green crab, <Emphasis Type="Italic">Carcinus maenas</Emphasis>, in California bays

The distribution of the introduced European green crab, Carcinus maenas, was investigated in the central California embayments of Bodega Bay Harbor (BBH), Tomales Bay, and Bolinas Lagoon using baited traps and snorkel surveys. Adult green crabs were very spatially limited in all three embayments and occurred primarily in warm, shallow areas that lacked large native Cancer spp. crabs. The green crabs that were found in closest proximity to populations of Cancer spp. exhibited high levels of limb damage and loss; damage was strongly correlated with low ratios of intertidal area: edge, indicative of narrow areas of intertidal that are more easily accessed by large Cancer spp. moving up to forage during periods of tidal inundation. Up to 70% of the green crabs tethered in areas of BBH that are utilized by Cancer spp. experienced limb loss, while those tethered in the marsh, where there are no Cancer spp., were undamaged. The results suggest that the potential distribution of green crabs in the northeastern Pacific will be far less than has been predicted, and that their impacts may be largely attenuated through predation by and competition with native crab species.     

Influence of occupant microhabitat on the composition of encrusting communities on gastropod shells

The exploitation of microhabitats is widely considered to increase biodiversity in marine ecosystems. Although intertidal hermit crabs and gastropods may inhabit the same shell type and shore level, their microhabitat may differ depending on the state of the tide. On the south coast of Wales the hermit crab Pagurus bernhardus mainly inhabits the shells of Nucella lapillus (84%). Hermit crab shells had a significantly different encrusting community compared with live N. lapillus shells. At low tide the live gastropods were found on exposed rock surfaces whereas hermit crabs were restricted to tidal pools. Communities encrusting live gastropod shells were characterised by lower species richness and abundance compared with shells inhabited by hermit crabs (12 species found in total). A greater abundance and richness of epibionts was recorded from both shell types during the summer compared with winter. Differences in community composition between shell occupant types were attributed to microhabitats used by gastropods and hermit crabs and the associated desiccation pressures, rather than competitive interactions or shell characteristics. This contradicts earlier studies of subtidal shells where biological processes were considered more important than physical factors in controlling species abundance and richness patterns. The use of rockpool microhabitats by hermit crabs increases the biodiversity of rocky shores, as some species commonly found on hermit-crab-inhabited shells were rare in other local habitats.Communicated by J.P. Thorpe, Port Erin     

Abundance,population structure and claw morphology of the semi-terrestrial crab <Emphasis Type="Italic">Pachygrapsus marmoratus</Emphasis> (Fabricius, 1787) on shores of differing wave exposure

Wave action is known to influence the abundance and distribution of intertidal organisms. Wave action will also determine the duration and suitability of various foraging windows (high-tide and low-tide, day and night) for predation and can also affect predator behaviour, both directly by impeding prey handling and indirectly by influencing prey abundance. It remains uncertain whether semi-terrestrial mobile predators such as crabs which can access intertidal prey during emersion when the effects of wave action are minimal, are influenced by exposure. Here, we assessed the effect of wave action on the abundance and population structure (size and gender) of the semi-terrestrial intertidal crab Pachygrapsus marmoratus on rocky shores in Portugal. The activity of P. marmoratus with the tidal cycle on sheltered and exposed shores was established using baited pots at high-tide to examine whether there was activity during intertidal immersion and by low-tide searches. Because prey abundance varies along a wave exposure gradient on most Portuguese shores and because morphology of crab chelipeds are known to be related to diet composition, we further tested the hypothesis that predator stomach contents reflected differences in prey abundance along the horizontal gradient in wave exposure and that this would be correlated with the crab cheliped morphology. Thus, we examined phenotypic variation in P. marmoratus chelipeds across shores of differing exposure to wave action. P. marmoratus was only active during low-tide. Patterns of abundance and population structure of crabs did not vary with exposure to wave action. Stomach contents, however, varied significantly between shores of differing exposure with a higher consumption of hard-shelled prey (mussels) on exposed locations, where this type of prey is more abundant, and a higher consumption of barnacles on sheltered shores. Multivariate geometric analysis of crab claws showed that claws were significantly larger on exposed shores. There was a significant correlation between animals with larger claws and the abundance of mussels in their stomach. Variation in cheliped size may have resulted from differing food availability on sheltered and exposed shores.     

Shell choice in<Emphasis Type="Italic"> Pagurus longicarpus</Emphasis> hermit crabs: does predation threat influence shell selection behavior?

Pagurus longicarpus hermit crabs depend on empty gastropod shells for protection against predation. Hermit crabs avoid gastropod shells in which holes have been drilled by naticid gastropods, and hermit crabs forced to occupy drilled shells are more vulnerable to predation by green crabs, Carcinus maenas. In this study, we examined the effect of predator cues on P. longicarpus shell investigation behavior and shell choice. In paired laboratory shell choice trials, we examined hermit crab response to green crab chemical cues. We compared hermit crabs from two sites differing in the percentage of Littorina littorea shells with drill holes. The percentage of time hermit crabs spent occupying intact shells increased significantly in the presence of predator cues. The effect of predator cues on the amount of time hermit crabs spent investigating shells differed between individuals from the two sites. Predator effluent had a marginal effect on the proportion of hermit crabs initially choosing intact shells and within 15 min most hermit crabs in both treatments occupied intact shells due to shell switching. These results indicate that predation cues alter P. longicarpus shell choice behavior favoring intact shells, which provide greater protection. In summary, predation appears to be a key factor influencing hermit crab shell selection behavior.Communicated by T. Czeschlik     

Temporal and spatial settlement patterns of sympatric hermit crabs and the influence of shell resource availability

Larvae of marine organisms often need specific resources or environments at settlement, and their success at settlement might be strongly influenced by the abundance and distribution of such specific resources. The larvae of hermit crabs need small shells to settle, so it is thought that the distribution and abundance of small shells influence the settlement pattern of hermit crabs. To investigate the influence of small shell distribution on the settlement of pagurid hermit crab larvae, we conducted a field experiment at an intertidal rocky shore in Hakodate Bay, Japan. From the line-transect sampling in the field, we found that Pagurus middendorffii settled extensively in the offshore side of the intertidal zone while P. nigrofascia settled in the uppermost area of the intertidal zone. Small shells were most abundant in a narrow shallow trough, slightly offshore from the uppermost area of the intertidal zone. For both species, settler abundance was high where adults were abundant, but settler abundance did not appear to be related to shells abundance. An experiment to clarify settlement patterns showed that larval recruits tended to be similar to those in the line-transect sampling of settlers. Thus shells may not be a primary factor affecting settlement patterns at relatively large scale within the intertidal flat. However, when we analyzed the relationship of settlers and shells separately within each transect, the distribution of settlers was well explained by shell resource availability. Therefore on a smaller scale, shell availability may influence the number of settlers. Settlement periods of P. middendorffii and P. nigrofascia fully overlapped, so their larvae probably were affected by similar transport factors, such as current and tidal movement. Nevertheless they showed different spatial patterns of settlement.Communicated by T. Ikeda, Hakodate     

Analysis of lipid components for determining the condition of anchovy larvae,Engraulis mordax

It has been presumed that intertidal spawning by Limulus polyphemus minimizes the loss of egges to subtidal predators; however, this strategy involves considerable risks. Massive beach strandings of adults accompany seasonal spawning migrations of crabs along Cape May in Delaware Bay, (USA). At least 190000 horseshoe crabs, approximating 10% of the adult population, died from beach stranding along the New Jersey shore of Delaware Bay during the 1986 (May to June) spawning season. Abnormalities of the telson (which is used in righting behavior) were significantly more common among stranded crabs than among individuals actively spawning on the intertidal beach. The number of stranded crabs per day was not correlated with tidal height or environmental variables (wind speed, wave height) which characterized the conditions at spawning. A complex suite of factors, including the size of the available spawning population, tidal and weather conditions, and beach slope, influence the number stranded during the breeding season. Horseshoe crab stranding results in a large loss of gravid females from the population, and may represent a major input of organic matter to intertidal sandy beaches in certain regions of Delaware Bay.     

Negative interactions between two SW Atlantic intertidal crabs in soft-bottom habitats

The intertidal crabs Chasmagnathus granulatus and Cyrtograpsus angulatus coexist across the SW Atlantic intertidal. Previous studies in this region suggest that C. granulatus displace C. angulatus in soft sediment, where C. granulatus build burrows (“burrowing beds”). We examined variation in abundance, size-frequency distribution, sex ratio, incidence of autotomies, and diet of both species in C. granulatus crab beds and adjacent areas without burrows. We also experimentally tested the hypothesis that in the absence of C. granulatus, C. angulatus will build burrows. Only large sized individuals of C. angulatus venture into C. granulatus crab beds. The sex ratio of C. angulatus was always biased towards females, with higher bias outside crab beds (1:8 outside, 1:2 inside). Although the items consumed in the crab beds did not differ from those consumed outside, the males of C. angulatus had a higher frequency of empty stomachs in crab beds. The incidence of limb autotomies of C. angulatus was higher outside C. granulatus crab bed areas. After a long rainy period in which C. granulatus was absent from these areas, the pattern of habitat use of C. angulatus changed. During this period C. angulatus showed higher abundance in the areas, where C. granulatus previously constructed burrows, and there were no differences between areas in any of the measured parameters. In the absence of C. granulatus, C. angulatus built their own burrows and never used C. granulatus burrows. The interaction between C. granulatus and C. angulatus may be a good example of competitive exclusion, when the shared resource is the access to surface soft-sediment.     

Shell-size selection by intertidal sympatric hermit crabs

This study evaluated selection for shell size by three species of tropical intertidal hermit crabs, Clibanarius antillensis, C. sclopetarius, and C. vittatus, from species of shells which are frequently used in nature. Crab size and weight were strongly and significantly related to all measured parameters of the selected shells. The strength of these relationships (r² values) depended neither on the crab nor on the shell variables taken into account. The relationships between crab size and the dimensions of the selected shells showed higher r² values than the corresponding relationships with the shells that the crabs had occupied when they were collected (0.482–0.903 in comparison to 0.091–0.652, respectively), indicating that the crabs were occupying sub-optimal shells in nature. Negative allometry was frequently found in the relationships between crab and shell variables, indicating that large crabs select and use proportionally lighter shells than do small crabs. This negative allometry was stronger for the shells used in nature (except for C. antillensis), i.e. larger crabs tended to select heavier shells in the laboratory than in nature. Different allometric relationships were also recorded among the dimensions of shells used in nature and those selected by the hermit crabs in free-access experiments: as shell length increased, the selected shells were heavier and had larger apertures than the shells used in nature. The relationships between crab size and the length and weight of the selected shells did not depend on the species of crab or species of shell, but only on crab size. Therefore, analyses using these variables can be performed without taking the species of crab or shell into account, i.e. data from different crab or shell species can be pooled. The influence of crab and/or shell species was recorded only in the models fitted for aperture length and width, variables which were more related to shell architecture than did shell length or weight. In contrast, if crab weight is used as an independent variable, different crab or shell species can be analyzed together independently of the particular shell parameter. This indicates that crab weight may be less susceptible than crab shield length to shell morphological constraints. Finally, the results indicate that the preferred size of a given shell type chosen by a given hermit crab will depend more on crab size or weight, than on the crab or shell species under consideration, i.e. crab shell-size relationships are not species specific.Communicated by P.W. Sammarco, Chauvin     

Utilization of partially predated snail shells by the hermit crab Pagurus longicarpus Say, 1817

The hermit crab Pagurus longicarpus was shown to inhabit shells that were partially predated from intertidal areas of Long Island, New York. Among field collections of P. longicarpus, 2.13% of the hermit crabs (46 of 2155) were found with shells with snail tissue present. Over 90% of these partially predated snail shells were occupied by male hermit crabs. Although hermit crabs were in 8 species of snail shells, only Littorina littorea and Nassarius obsoletus were found occupied by hermit crabs and containing snail tissue. In the laboratory, we found that specimens of the spider crab Libinia emarginata were able to pull off the operculum of snails, leaving damage as found in field collections. In contrast, specimens of P. longicarpus were not able to prey on live, healthy snails. When specimens of P. longicarpus were placed in communal tanks, hermit crabs preferred partially predated snail shells to empty and original shells. However, original shells and empty shells were occupied with more frequency than partially predated shells when crabs were isolated. These findings indicate P. longicarpus actively seeks shells soon after attack and abandonment by snail predators, especially in the presence of competitors.     

Feeding habits of the dungeness crab Cancer magister as determined by the index of relative importance

Specimens of the Dungeness crab Cancer magister, were collected subtidally and intertidally from an estuary in Washington State, USA in June and September 1980, and January, April, May and July, 1981. Gut contens of freshly collected crabs were analyzed by the Index of Relative Importance; for each prey taxon, this method measured frequency of occurrence, percentage of total biomass, and percentage of total numbers consumed. The most important higher taxon eaten was fish; however, the most important prey genus was the shrimp Crangon spp. There was greater predation on Crangon spp. at night at the intertidal site, and during winter and spring when the shrimp were most abundant there. Feeding activity, as indicated by a weight-specific gut-fullness index, showed no consistent diel pattern. There were significant ontogenetic changes in feeding patterns: first-year crabs preyed primarily on very small bivalves or small crustaceans including their conspecifics; second-year crabs preferred Crangon spp. and fish, and third-year crabs preyed less on Crangon spp. and more on fish. Such changes in feeding habits with ege could be purely due to mechanics of food handling, but might also reduce competition among age groups of crabs, possibly partitioning resources within the estuary. Findings are discussed in terms of optimal foraging and compared to other similar studies.Contribution No. 599, School of Fisheries, University of Washington, Seattle, Washington 98195, USA     

The role of individual behavior type in mediating indirect interactions

Trait-mediated indirect interactions (TMII) play an important role in structuring natural communities, and numerous studies have experimentally demonstrated their presence in a variety of systems. However, these studies have largely examined the presence or absence of traits that are responsible for these interactions, without considering natural variation between individuals in the extent to which these traits are manifested. We used a well-documented TMII to investigate the importance of individual behavior type for determining the strength of the TMII. The toadfish Opsanus tau has an indirect positive influence on bivalve survival because the mud crab Panopeus herbstii, a consumer of bivalves, reduces foraging effort in the presence of toadfish. We quantified variation in the strength of persistent individual mud crab responses to toadfish and resulting variation in the strength of TMII. We demonstrate that the strength of this TMII is strongly influenced by mud crab size and behavior type, strengthening with the intensity of response of individual mud crabs to toadfish predator cues. Further, we demonstrate that the spatial distribution within intertidal oyster reefs of crabs with different behavior types is not random; mud crabs inhabiting subtidal areas, where predator cues are more persistent, are significantly less responsive to toadfish cues than mud crabs from intertidal areas. This spatial behavioral structure should lead to spatial variation in the strength of TMII. Given the widespread importance of TMII and the broad occurrence of individual personality or behavior types across numerous taxa, these results should be generally applicable. The distribution of behavior types within a population may therefore be a useful metric for improving our ability to predict the strength of TMII.     

Factors limiting distribution and activity patterns of the soldier crab Dotilla myctiroides in Phuket, South Thailand

This study was carried out between January and March 1995 on the intertidal sand flats of Tang Khen Bay, Phuket, South Thailand, where the soldier crab Dotilla myctiroides (H. Milne-Edwards) occurs in densities of up to 120 m⁻². In this bay, long, ribbon-like sand waves (wavelength 40 m, height 0.4 m) are interspersed with shallow pools, running approximately parallel to the shore. During daylight low-tides, exposure of the sand waves is followed 15 to 20 min later by the emergence of the crabs which have been buried under the sediment surface during high tide. Their subsequent burrowing and feeding activity results in the production of large numbers of sand pellets on the sediment surface. Most crabs retreat down their burrows, and some also plug the burrow entrance, prior to being covered by the incoming tide. The crab burrows have a distinct distribution on the sand waves. Burrows are most dense at the top of each sand wave, and a band of unburrowed sediment adjoins the adjacent tidal pools. Crabs are most abundant between mean high-water neap-tide level and mean low-water neap-tide level, where the median particle size of the surface sediment is ∼2 . Measurements of water-table depth below the sand waves and the exposure time of the sediment indicated that, where sediment size is suitable, the main factor controlling crab distribution is the duration of daytime exposure. This observation is in contrast to those of many previous studies, which have suggested that water-table height and sediment drainage are the main factors controlling the distribution of D. myctiroides. Received: 14 January 1998 / Accepted: 6 May 1999     

Catch rate and emergence of male and female spanner crabs (Ranina ranina) in Australia

Catches of the spanner crab, Ranina ranina (Linnaeus, 1788), in southern Queensland, Australia in 1982 and 1983 showed a seasonal cycle, with peak catches from August to September. In most catches, males outnumbered females. Catches of both male and female spanner crabs declined in November and December, when a high proportion of females caught were ovigerous. Fish, echinoderms and polychaetes were the main food items. Closed-circuit time-lapse video recording was used to study the emergence behaviour of R. ranina in a 6 m-diameter tank. Spanner crabs in this tank were buried most of the time, emerging on average for only 1.7 h d^-1. They emerged mainly between 16.00 and 24.00 hrs and, less often, between 01.00 and 15.00 hrs. Males remained emerged significantly longer than females. The duration of emergence of the females peaked before the spawning season, becoming shorter during the period when they were ovigerous. Temperature and emergence were negatively correlated. It is concluded that reproductive behaviour has a strong influence on catch composition of spanner crabs.     

Life-history patterns in two forms of Macrophthalmus japonicus (Crustacea: Brachyura)

Two behaviorally distinct forms — courtship signal — (Form L and V) of Macrophthalmus japonicus (De Haan) are known. Monthly samples of each form were collected from two areas in Japan: Form L from Amakusa, western Kyushu, between April 1985 and February 1986; Form V from Fukuoka, northern Kyushu, between November 1982 and December 1983. Life-history traits were compared between two populations and discussed. Individuals from the Amakusa population grew more slowly and matured at a smaller size than those from the Fukuoka population. Maturation period and longevity did not differ significantly. In the first year of reproduction, females in the Fukuoka population produced more, and larger broods than those in the Amakusa population and crab mortality was higher in every age group. Egg size did not differ significantly. The major environmental factors — air-temperature, nitrogen content of the surface mud and period of aerial exposure were similar in both areas, and it is suggested that the differences in life-history exhibited by these crabs are genotypic, and perhaps related to their different form.     

Body and resource size at the land–sea interface

Body size in animals varies with many parameters, amongst them taxonomic affiliation, lifestyle and ambient environment oxygen levels. Size has considerable implication to possibilities for animals; for example, parasites need to be small and top predators large. Body size and resource requirements (shell size) were investigated across the land–sea interface in hermit crabs (Crustacea: Malacostraca: Decapoda) and snails (Mollusca: Gastropoda: Prosobranchia). These are two of the few taxa to occur in the sea, on the shore and on land as residents. Both taxa are also appropriate for such an analysis as they are abundant, speciose, cohabit the same environments and are linked—gastropod shells are a critical resource to hermit crabs. Both the maximum and mean sizes of hermit crab species showed parabolic relationships with shore height, decreasing from the sublittoral and supralittoral to the eulittoral. Average maximum size of gastropods exhibited a similar intertidal minimum although variability was high. It is suggested that this pattern is robust: not only did two distantly related taxa show the same pattern, but neither region nor site contributed significantly to total variability. The mass of resources (gastropod shells) used by hermit crabs, however, showed a converse pattern. The smallest shells (relative to hermit crab body size) were used in the sublittoral and supralittoral. Response to environmental stress and predation pressure are offered as two alternate theories to explain the observed body dwarfism and resource gigantism in the intertidal zone.Communicated by J.P. Thorpe, Port Erin     

Annual fluctuation of life-history traits in the mud crab Macrophthalmus japonicus

Interannual fluctuation of life-history traits in the mud crab Macrophthalmus japonicus (De Haan) was studied for a 7-yr period (1981 to 1987) in an intertidal mudflat in the estuary of the Tatara-Umi River, Fukuoka, Japan. Density of juveniles fluctuated greatly, and maximum annual density was low in 1981 to 1984 (22.0 to 44.0 m^-2) and very high in 1985 to 1987 (98.5 to 188.6 m^-2). In 1980, recruits may have been very few, though there is no data on or before 1980, because no crab of the 1980 cohort was found during the study period. Density of adult crabs was almost stable (11.0 to 32.6 m^-2), perhaps because of density-dependent mortality. Under the high density conditions prevailing from 1985 to 1987 crabs grew slowly and longevity was shorter. Male chela and female abdomen were dwarfish, and females produced few and small broods. The changes of life-history traits under high density conditions were thought to result from strong intraspecific competition for space.     

Resource utilization for decorating in three intertidal majid crabs (Brachyura: Majidae)

 For three spider crabs (Tiarinia cornigera, Micippaplatipes and Pugettia quadridens quadridens), patterns of algal utilization for decorating were compared with the dynamics of algae on an intertidal rocky shore reef where the crabs co-occurred. T. cornigera and P. quadridens quadridens were most abundant from autumn to spring when the dominant algae (Boodleacoacta, Sargassum hemiphyllum, S. thunbergii and Corallina pilulifera) occurred in high coverage, while M. platipes was most abundant from spring to autumn. Monthly change of algae used for decorating was not correlated with algae growing in the crab habitat for T. cornigera, but for M. platipes, it was positively correlated for two algal species, and for P. quadridens quadridens, negatively correlated for one algal species. Each species of the spider crabs used some algal species preferentially for decoration. Decorating preference experiments conducted in the laboratory showed that M. platipes and P. quadridens quadridens exhibited similar preference to their algal utilization in the field, whereas for T. cornigera, algal preference in the experiment differed from utilization in the field. Comparisons between materials used for decoration and gut contents revealed that T. cornigera and M. platipes used algal species differently for decorating and feeding, while P. quadridens quadridens used the same algal species for both decorating and feeding. Different tactics for camouflage are discussed in terms of algal utilizations by the three majid species. Received: 25 September 1999 / Accepted: 22 June 2000     

Past predation risk induces an intertidal whelk (Nucella lamellosa) to respond to more dilute concentrations of its predator’s scent

Unless sensory control is coupled with developmental flexibility, organisms are unable to produce adaptive phenotypes in response to environmental risk cues. However, empirical examples of adaptive phenotypic plasticity focus largely on development and rarely on sensory control. Here, I ask whether past exposure to predation cues affects an intertidal whelk’s (Nucella lamellosa) sensitivity to the scent of a known predatory crab (Cancer productus). I pre-conditioned whelks for 30 days with the smell of crabs eating other whelks and then, after a period of rest (14 days in fresh seawater free of predator effluent), re-introduced them to different concentrations of waterborne crab scent. Relative to controls, pre-conditioned whelks expressed crab-defences at significantly lower concentrations of crab effluent, suggesting the pre-conditioning reduced their risk–response threshold. Results remind us that the full scope of factors controlling adaptive phenotypic plasticity in natural populations remains underappreciated.     

Dispersal,population structure and burrow shape of Ocypode cursor

The distribution of the sand crab Ocypode cursor (L.), as indicated by the number of burrows, was studied for 2 years in a 50×50 m sand beach area in northern Israel. A definite relationship was established between the distribution pattern from the seashore inwards towards the sand dunes, and the degree of sand moisture as it changed seasonally. During autumn, more crabs were found at a distance of 15 to 25 m from the sea where sand moisture was about 14%. At the beginning of winter crabs dispersed evely, disappearing with advancing winter. Crabs reappeared in spring, although in smaller numbers, dispersing in a pattern similar to that in autumn. At the beginning of summer and later on, more crabs appeared and concentrated closer to the sea (5 to 10 m). The population structure was analysed directly by measuring the crab's dimensions, and indirectly by counting burrows and measuring the diameter of their openings. Direct analysis revealed two distinct sizeage groups: smaller crabs 0.5 to 3 cm long, and larger ones over 4 cm long. The smaller burrows were inhabited by the first group and were mostly found closer to the sea; the second group was found more landwards. Three main types of burrow shapes are described.