Population structure of the South American Estuarine crab, Chasmagnathus granulatus (Brachyura: Varunidae) near the southern limit of its geographical distribution: comparison with northern populations

Crabs are among the most conspicuous and ecologically important invertebrates of the large intertidal zones that characterize estuarine and protected coastal areas in temperate regions. The habitat, population structure and breeding cycle of Chasmagnathus granulatus (Brachyura: Varunidae), a semiterrestrial burrowing crab endemic to the warm temperate coasts of the Southwestern Atlantic, were studied in San Antonio Bay (Argentina), near the southern limit of its range. San Antonio Bay has no freshwater input, winter is relatively colder, and summer warmer, than northern habitats of this species. Crabs lived both in vegetated and unvegetated zones, but density and sex ratio varied among dates and zones. The maximum observed density was 136 crabs/m² , the maximum carapace width (CW) was 32 mm (males) and 29.8 mm (females), ovigerous females were found only in November and January, and the smallest ovigerous female measured 17 mm CW. The population structure, spatial distribution, and recruitment pattern of C. granulatusdid not differ between San Antonio Bay and northern habitats. The higher density, smaller maximum size and shorter reproductive cycle observed in San Antonio cannot be atributed to changes associated with a latitudinal cline and other factors, such as thermal amplitude and food availability, need to be studied.     

Abundance,distribution and habitat characteristics of winter skate Leucoraja ocellata in the southern Gulf of St Lawrence: a population on the brink of extirpation?

Abundance indices of southern Gulf Leucoraja ocellata residing in Northumberland Strait, Canada, declined from 2001 to 2005, but increased during 2006 to 2008. Catch rates in 2009 were the lowest in the time series. The size of the area occupied in Northumberland Strait varied with changes in the abundance indices. Leucoraja ocellata were primarily caught in the north‐west half of the strait over sandy to gravelly seabed; a near total absence was documented from the eastern half despite suitable habitat and a previously established presence documented during research surveys conducted before 2000. In Northumberland Strait, L. ocellata occupied shallow (50% cumulative occurrence = 12 m) and warm (50% cumulative occurrence = 16·5° C) coastal waters and were not present in the cold (<1° C, >35 m) intermediate layer. Abundance and distribution data highlight the need for continuing studies to monitor the status of this population, which is classified as endangered by the IUCN and Committee on the Status of Endangered Wildlife in Canada.     

Effects of juvenile non-indigenous Carcinus maenas on the growth and condition of juvenile Cancer irroratus

The Atlantic rock crab, Cancer irroratus, is a commercially fished species and a critical prey item for the American lobster, Homarus americanus, in Atlantic Canada. The recent invasion of European green crab, Carcinus maenas, may have significant effects on the growth and condition of native C. irroratus, because both species overlap spatially and temporally and have similar habitat and dietary requirements. To examine such potential effects, we measured the growth of juvenile C. irroratus in the presence of juvenile C. maenas over a period of 4 months (growing season), under the following species combinations: (1) one C. irroratus (10-25 mm CW); (2) two C. irroratus (10-25 mm CW); (3) one C. irroratus (10-25 mm CW) and one C. maenas (10-15 mm CW). Morphological measurements included pre- and post-molt carapace width, chela height, abdomen width (mm), weight (g), and estimates of molt increment (%) and intermolt duration (days). Analysis of the hepatopancreas for % lipid content at the end of the experiment provided an estimate of physiological condition. The effect of the presence of C. maenas on the growth of C. irroratus shifted from negative to positive, when C. irroratus reached CW of 19-22 mm and gained a presumably significant size advantage over C. maenas. The positive effect resulted from increased energy intake through crab consumption. In the absence of crab consumption, the presence of a second crab (conspecific or C. maenas) had no effect on growth. C. irroratus consumed crabs more frequently when the second individual was a green crab than a conspecific. Consumption of C. maenas had a pronounced effect on the growth rate of C. irroratus, resulting in shorter intermolt periods and larger percent molt increments than in the presence of a conspecific. Therefore, the presence of juvenile C. maenas does not appear to have a prolonged negative effect on the growth of C. irroratus; rather, it may provide an additional food item as rock crabs grow, as long as encounters between the two species occur at high enough rates.     

Quantifying factors influencing American lobster (Homarus americanus) predation on the invasive green crab (Carcinus maenas) in Newfoundland,Canada

The European green crab (Carcinus maenas) is a newly invasive species in Newfoundland, where it has likely been present for ≤15 years. The green crab has been found in stomach contents of American lobster (Homarus americanus) in New England and Nova Scotia, Canada, but predation on this species has not yet been quantified in Newfoundland. We conducted feeding experiments to determine whether lobsters from Newfoundland were as likely as those from Nova Scotia (which have coexisted with green crabs for >60 years) to recognize and prey upon this new species. We also performed experiments to determine whether green crabs reach a size refuge from predation and whether factors including starvation, availability of alternate food sources, or habitat complexity would influence the probability of lobster attacking or feeding on green crabs. In our trials, lobster origin had no significant effect on crab predation; lobsters, irrespective of origin, were more likely to consume small (<40 mm carapace width [CW]) and medium (40–65 mm CW) crabs than larger (>65 mm CW) ones. Nevertheless, even small lobsters (73–76 mm carapace length, 300 g) were able to kill and consume the largest green crabs (78 mm CW, 100 g). Green crabs were less likely to be attacked or eaten when an alternative food source was present, suggesting that the lobsters were preying on the crabs, rather than simply killing them in a dispute over territory. The addition of a shelter provided a refuge for the green crabs; however, the crabs were only able to avoid being injured or eaten if this shelter was structurally complex. The green crab is slowly spreading westward around the island of Newfoundland, and so its long‐term effects, interactions with other organisms, and contribution to the diet of Newfoundland lobsters remain to be seen.     

Some aspects of the biology of the amphipods <Emphasis Type="Italic">Ischyrocerus anguipes</Emphasis> associated with the red king crab, <Emphasis Type="Italic">Paralithodes camtschaticus,</Emphasis> in the Barents Sea

This paper describes size composition, morphometric, and reproductive parameters of the summer generation of the amphipod Ischyrocerus anguipes (Krǿyer, 1838) associated with the red king crab Paralithodes camtschaticus (Tilesius, 1815) in Dalnezelenetskaya Bay (southern Barents Sea, Russia). In summer 2004–2007, prevalence and mean number of I. anguipes per host were 15.0% and 6.9 ± 1.1 specimens, respectively. The majority of amphipods were found on the host carapace (47.4%) and the limbs (43.9%). High infestation indices and absence of negative impacts for the host indicate that I. anguipes is a facultative commensal of the red king crab. The amphipods sex ratio was significantly biased toward females (F:M = 2:1). Females had a greater size than males. The size at 50% maturity of I. anguipes females was estimated to be 3.37 mm. The number of eggs laid is linearly correlated with the size of a female. Some differences in biological features of symbiotic and free-living amphipods (data obtained in 1940–1950) may be explained by climatic changes in the Barents Sea or advantages of living on the crab.     

Feeding Habits of Percnon gibbesi (H. Milne Edwards, 1853) in the Sicily Strait

The subtropical grapsid crab Percnon gibbesi (H. Milne Edwards, 1853) is one of the most recent alien decapods found in the Mediterranean, where it was discovered at Linosa (Pelagie Islands, Sicily Strait) in summer 1999. At present, the invasion of this species has been recorded in several other insular localities. We studied the feeding habits of the crab in an attempt to understand its success in the Mediterranean. The morphological characteristics of the chela, the feeding adaptations of the gastric mill and the analysis of stomach contents indicate that P. gibbesi is a strictly herbivorous species, a characteristic not shared with any other large-sized infralittoral Mediterranean crab.     

Effect of crab shell size on bio-demineralization with lactic acid-producing bacterium, <Emphasis Type="Italic">Lactobacillus paracasei</Emphasis> subsp. <Emphasis Type="Italic">tolerans</Emphasis> KCTC-3074

Demineralization (DM) from crab shell (CS) waste was carried out using a lactic acid-producing bacterium, Lactobacillus paracasei subsp. tolerans KCTC-3074 for 7 days at 25, 30, and 35°C. DM rates were 89∼92% and slightly affected by temperature. DM was also performed for four particle-sized shell samples (0.84∼3.35, 3.35∼10, 10∼20, and 20∼35 mm) with 10% inoculum, 5% shell, and 10% glucose at 30°C and 180 rpm for 7 days. It was found out that the shell size had a slight effect on the rate of DM. Negative relationships were found between DM and residual dry weight (r² = 0.960), and between DM and pH (r² = 0.906). Conversely, positive relationships were found between DM and medium protein (r² = 0.696), and between DM and total titratable acidity (r² = 0.630).     

Zooplankton response to a warmer northern Wadden Sea

Weekly measurements of mesozooplankton (>76 μm) and hydrographic parameters have been carried out since 1984 in the List Tidal Basin (northern Wadden Sea). Monthly water temperature significantly increased by 0.04°C year⁻¹. The largest increase by 3°C in 22 years occurred in September, implying an extension of the warm summer period. Mean annual copepod abundance and length of copepod season correlated significantly with mean temperature from January to May. Except for an increasing Acartia sp. abundance during spring (April–May), no long-term trends in copepod abundance were observed. The percentage of carnivorous zooplankton increased significantly since 1984 mainly due to a sudden increase in the cyclopoid copepod Oithona similis in 1997. We expect that global warming will lead to a longer copepod season and higher copepod abundances in the northern Wadden Sea.     

The effect of urban ground cover on microclimate,growth and leaf gas exchange of oleander in Phoenix,Arizona

We assessed how small patches of contrasting urban ground cover [mesiscape (turf), xeriscape (gravel), concrete, and asphalt] altered the microclimate and performance of adjacent oleander (Nerium oleander L.) plants in Phoenix, Arizona during fall/winter (September–February) and spring/summer (March–September). Ground-cover and oleander canopy surface temperatures, canopy air temperatures and pot soil temperatures tended to be lowest in the mesiscape and highest in the asphalt and concrete. Canopy air vapor pressure deficits were lowest in the mesiscape and highest in the asphalt plot. Rates of net photosynthesis of all oleander plants were highest in October and May, and declined through mid-summer (June–July), when rates tended to be highest in the cooler mesiscape, particularly when water was limiting. During fall/winter, oleanders in the mesiscape produced 20% less biomass, 13% less leaf area, and had 12% lower relative growth rates (R_G) than those in the other ground covers. Lower nighttime temperatures in the mesiscape in December led to oleander frost damage. During spring/summer, oleanders in the mesiscape produced 11% more biomass, 16% more leaf area, and had 3% higher R_G than those in the other cover types. The effects of urban ground cover on oleander performance were season-specific; while oleander growth was greatest in the mesiscape during spring/summer, it was lowest during fall/winter and these plants experienced frost damage. Because all oleander plants produced >10 times as much biomass during the spring/summer, on an annual basis oleanders in the mesiscape produced 5–11% more biomass than plants in the warmer ground covers.     

Evidence for a permanent establishment of the snow crab (Chionoecetes opilio) in the Barents Sea

In the Atlantic the snow crab (Chionoecetes opilio) is naturally distributed on the northwestern side, i.e. eastern Canada and west Greenland. Until recently, there have been no observations of snow crab in eastern Atlantic. However, in 1990s single and occasional reports were made of crabs captured in the eastern part of the Barents Sea, presumably introduced through ballast water. Special attention during the annual bottom-trawl surveys in the Barents Sea during February 2004–2006 were given to include recordings of snow crab to evaluate if the introduced species has succeeded to establish a self-sustaining population in this region. Recordings of snow crabs were systematically noted and biological measurements carried out. The results confirm previous Russian observations of snow crabs in the northern region of Gåsebanken. In addition, a significant number of crabs were also found in the central region of the Barents Sea, mainly in deeper waters from 180 to 350 m depth. The sizes ranged from 14 to 136 mm carapace width. All females above 70 mm were berried with fertilised eggs. A major fraction (31% in 2005; 76% in 2006) of the crabs consisted of juveniles below 50 mm CW, providing evidence for successful recruitment. The small-sized crabs were exclusively found in Gåsebanken, identifying the main recruiting area at present for snow crab in the Barents Sea. The results obtained show that the snow crab is now adapted to the northeast Atlantic.     

Use of Chitosan Membrane from the Carapace of the Soldier Crab <Emphasis Type="Italic">Mictyris brevidactylus</Emphasis> for Biosensor Construction

Glucose oxidase (EC 1.3.4.3) was immobilized on chitosan membrane (<0.1 mm in thickness) prepared from the carapace of the soldier crab Mictyris brevidactylus. A glucose electrode was constructed by covering a platinum electrode (2.0 mm in diameter) with the enzyme membrane. The enzyme electrode sensed glucose amperometrically (1.0 µA/mM, with linear range up to 0.5 mM, r = 0.999) when positively imposed with 0.6 V against an Ag/AgCl reference electrode. The glucose biosensor was sensitive (<0.1 µM, S/N > 3), reproducible (CV for 55 µM glucose <3%, n = 5), reagentless, and durable for months.     

The Effect of Tripalmitin Crystallization on the Thermomechanical Properties of Candelilla Wax Organogels

The effect of tripalmitin (TP) crystallization on the thermomechanical properties of organogels developed with candelilla wax (CW) was investigated using safflower oil high in triolein (HOSFO) as the liquid phase. Factorial combinations of CW (i.e., 0–3%) and TP (i.e., 0–1%) in the HOSFO were used to develop organogels at three different temperatures (T _set). The onset of crystallization (T _g) during the cooling stage (10 °C/min), the melting temperature (T _M), and the corresponding heat of melting (ΔH _M) of the organogels were determined by differential scanning calorimetry. Results showed that, without CW, the crystallization of TP in the HOSFO at the concentrations and T _set investigated (i.e., −10 °C to 25 °C) did not develop a three-dimensional network that provided significant viscoelasticity (i.e., solid-like behavior) to the HOSFO. The CW developed organogels in the HOSFO with T _M’s that increased from ≈30.5 °C up to ≈42.5 °C as a function of CW concentration. In contrast, in the CW–1% TP system, the co-crystallization of TP and CW resulted in organogels with T_M’s that varied just between 36 °C and 38 °C, independent of the CW concentration. Higher elastic modulus (G′) and yield stress (σ*) were obtained with 3% CW–1.0% TP organogels than with organogels developed just by CW, particularly at T _set’s of −5 °C and 15 °C. This research showed that co-crystallization of TP and CW, occurring at different extent as a function of T _set, resulted in organogels with thermomechanical properties different from the ones showed by CW organogels. The results showed that co-crystallization of triacylglycerides with CW might be a useful alternative to tailor particular physicochemical properties associated to a specific functionality (i.e., melting profile and texture). Organogelation of vegetable oil might be used to develop trans-free vegetable-oil-based spreads and coatings and also novel food products with new textural perceptions for the consumers.     

Summer and fall microhabitat utilization of juvenile bull trout and cutthroat trout in a wilderness stream, Idaho

Microhabitat use and availability were evaluated and compared between different size classes of juvenile resident bull trout (Salvelinus confluentus) and cutthroat trout (Oncorhynchus clarki) in a small wilderness stream within the South Fork Clearwater River basin, Idaho. The objective was to determine if utilization of measured habitat characteristics changed from summer to late fall. Sampling of fish was conducted with night snorkeling. During the summer, smaller juvenile bull trout (<66 mm) total length (TL) were associated with shallow stream margins over coarse substrates. In the fall, they moved to significantly deeper, lower velocity water, and closer to cover (p<0.05), but maintained their association with coarse substrates. During the summer, larger juvenile bull trout and larger juvenile cutthroat trout (66–130 mm TL) occupied significantly deeper water than smaller juvenile bull trout (p<0.05). Generally, larger juvenile bull trout were found closer to the bottom and in lower velocity water than larger juvenile cutthroat trout (p<0.05). In the fall, larger juvenile bull trout and larger juvenile cutthroat trout were associated with significantly deeper, lower velocity water located closer to cover than in summer (p<0.05). However, cutthroat trout occupied slightly deeper water over finer substrates than bull trout. Deep water with low velocities evidently provide important rearing areas for large bull trout and large cutthroat trout in the fall. Land management practices that maintain such environments will benefit these species.     

Survival of the mosquito predator,Notonecta unifasciata[Hemiptera: Notonectidae] embryos at low thermal gradients

Notonecta unifasciata Guerin eggs maintained at different stages of embryonic development in water at variable temperatures (2.2–25.6 °C) and for periods of 4–12 weeks revealed maximum viability (>80 %) at the highest temperature. However, optimum nondevelopmental viability was at 14.4 °C with eight-day-old embryos (>35 %). Short term (4 weeks) storage at 14.4 °C significantly increased egg viability. Survival was poor (<20 %) at the 2 lowest temperatures. Eggs held at 14.4 °C for 12 weeks and sustainingca. 50 % mortality, may be a practical procedure for biological control.      

Identifying patterns in the diet of mackerel icefish (<Emphasis Type="Italic">Champsocephalus gunnari)</Emphasis> at South Georgia using bootstrapped confidence intervals of a dietary index

Ontogenetic, inter-annual and regional variations in diet were investigated for mackerel icefish, Champsocephalus gunnari, in three successive summer seasons around South Georgia. Stomach contents from 2239 C. gunnari (130–560 mm total length) were examined. A bootstrapping technique was used to calculate confidence intervals for an index of relative importance of prey categories (% IRI_DC). Diet varied significantly between years and age classes but there was little regional difference in diet. In general, diet was dominated by krill, Euphausia superba and by the amphipod Themisto gaudichaudii. Smaller (younger) fish tended to prey on a higher proportion of T. gaudichaudii and small euphausiids such as Thysanoessa sp. and took smaller quantities of E. superba. In a season of poor krill availability (summer of 2003–2004) the proportion of krill in the diet, stomach fullness and fish condition (indicated by length–weight relationships) were significantly lower than in the other summer seasons. A large reduction (>80%) in the estimated annual (2005) biomass of the C. gunnari stock directly followed the season of poor krill availability. This decline was largely because of mortality of 2+ and 3+ fish, which were more krill dependent than 1+ fish. Younger fish appear to have survived, leading to an increase in the estimated population biomass in 2006.     

Occurrence of symbiotic turbellarians in the oyster,Crassostrea virginica

Crassostrea virginica was collected from several locations where it is cultured, both along the Northumberland Strait of New Brunswick and Malpeque Bay on the coast of Prince Edward Island. The oysters were found with two turbellarians on their gills. Urastoma cyprinae (Graff) was found in the oysters mostly during the warmer months of the year in numbers averaging as high as 50 worms per host (N = 50) and with as much as 78% of the host population infected (N = 100). Paravortex gemellipara (Linton) was also found during warmer months, but much less frequently or abundantly.Both male and female oysters were found to have U. cyprinae. No eggs or recent young of U. cyprinae were found in hosts; female-mature individuals of P. gemellipara with young were found from June through August.     

Diet,food preference,and algal availability for fishes and crabs on intertidal reef communities in southern California

Synopsis Herbivory by wide-ranging fishes is common over tropical reefs, but rare in temperate latitudes where the effects of herbivorous fishes are thought to be minimal. Along the west coast of North America, herbivory by fishes on nearshore reefs is largely restricted to a few members of the Kyphosidae, distributed south of Pt. Conception. This paper presents information on natural diets and results from feeding choice experiments for two abundant kyphosids from intertidal habitats in San Diego, California —Girella nigricans andHermosilla azurea, and similar data for the lined shore crab,Pachygrapsus crassipes, which also forages over intertidal reefs. These results are compared with the availability of algae in intertidal habitats measured during summer and winter, on both disturbed and undisturbed habitats. The diets of juveniles ofG. nigricans andH. azurea collected from nearshore habitats were dominated by animal prey (mainly amphipods), but adults of these fishes, andP. crassipes, consumed algae nearly exclusively, with 26, 10, and 14 taxa of algae identified fromG. nigricans, H. azurea, andP. crassipes, respectively. Algae with sheet-like morphologies (e.g.Ulva sp.,Enteromorpha sp., members of the Delesseriaceae) were the principal algae in the diets of the fishes, and calcareous algae (e.g.Corallina sp.,Lithothrix aspergillum) and sheet-like algae (Enteromorpha sp.) comprised the greatest identifiable portion of the shore crab's diet. Feeding choice experiments indicated that the fishes preferred filamentous algae (e.g.Centroceras clavulatum, Polysiphonia sp.,Chondria californica) and sheet-like algae (e.g.Enteromorpha sp.,Ulva sp.,Cryptopleura crispa) over other algal morphologies, whereas the shore crab chose jointed calcareous algae (e.g.Lithothrix aspergillum, Corallina vancouveriensis, Jania sp.) most frequently. The diets and preferences for algae by the fishes were generally most similar to the assemblage of algae available in early successional (disturbed) habitats during summer when sheet-like and filamentous algae are abundant. The shore crab exhibited the opposite trend with a diet more similar to late successional (undisturbed) habitats.     

Summer and spring trophic niche of larval and juvenile <Emphasis Type="Italic">Pleuragramma antarcticum</Emphasis> in the Western Ross Sea,Antarctica

The aim of this paper is to contribute to the knowledge on the feeding habits of larvae and juvenile Pleuragramma antarcticum in the western Ross Sea. In summer, the diet of P. antarcticum postlarvae (8–17 mm) was dominated by calanoid eggs (35.5%), Limacina (32.1%) and tintinnids (17.6%), while the principal food of juveniles consisted mainly of copepods (98.2%), with Oncaea curvata being the most abundant (85.1%) and the most frequently consumed prey. The food composition of P. antarcticum postlarvae (24–29 mm), collected in spring, suggest that they fed actively under the sea ice. Stephos longipes, Harpacticus furcifer and Paralabidocera antarctica sea ice copepods represent, in all their different developmental stages, the most abundant biomass food in Terra Nova Bay in this period. Our results therefore suggest that the diet of younger Pleuragramma specimens shifted in prey composition from the first summer to the following spring. This study draws attention to the key role of the copepod, P. antarctica, in the food web of Terra Nova Bay. This article belongs to a special topic: Five articles coordinated by L. Guglielmo and V. Saggiomo appear in this issue of Polar Biology and are a result of a workshop on Sea-ice communities in Terra Nova Bay (Ross Sea) held in August 2007 in Capo Calavà, Messina, Italy.     

An invasive crab alters interaction webs in a marine community

Over the last decade, the non-native, filter-feeding crab Petrolisthes armatus invaded oyster reefs of the South Atlantic Bight at densities of thousands m⁻². Mesocosm and field experiments demonstrated that P. armatus at ∼10–75% of mean summer densities: (1) suppressed growth of small oysters, biomass of benthic microalgae, and recruitment of native mud crabs, (2) enhanced oyster, mussel, and total bivalve recruitment, macroalgal cover, and survivorship of predatory oyster drills, but (3) did not affect native taxonomic richness. Laboratory feeding assays, field tethering experiments, and population changes in field and mesocosm experiments suggest that P. armatus is a preferred prey for native mud crabs and other consumers, thus relieving predation on native species and enhancing recruitment or survival of bivalves and oyster drills. In contrast, the invasive crab can consume crustacean larvae and via this feeding may suppress recruitment of native mud crabs. Our findings should be conservative given the low densities of P. armatus seeded into experimental plots and our inability to run longer-term experiments due to controls rapidly being colonized by non-native crabs recruiting from the plankton. Invasive crabs commonly impact native communities via predation, but community impacts of this invasive crab may be as much due to its role as a preferred prey of native consumers as to its predation on native prey. Given that oysters are foundation species for shallow reefs in the South Atlantic Bight, the long-term effects of this invasion could be considerable.     

Burrow Architecture of the Ghost Crab Ocypode ceratophthalma on a Sandy Shore in Hong Kong

The ghost crab Ocypode ceratophthalma (Pallas) creates burrows of variety shapes at different ages. Juveniles (mean carapace length 11 mm) produced shallow J-shaped burrows, which incline vertically into the substratum (mean depth 160 mm). Larger crabs (17–25 mm carapace length) have Y-shaped and spiral burrows (mean depth 361 mm). These Y-shaped burrows have a primary arm, which extends to the surface forming the opening, and a secondary arm which terminates in a blind spherical ending. The two arms join in a single shaft and end with a chamber at the base. The secondary arms and chambers are believed to be used for mating or as a refuge from predation. The spiral burrows have spiral single channel ending in a chamber. Older crabs (mean carapace length 32.6 mm) had simple, straight single tube burrows, which inclined into the substratum at mean of 73° and had a mean depth of 320 mm. During summer daytime periods, the burrows shelter the crabs from heat and desiccation stress. The sand surface temperature at the burrow opening was ~48 °C but temperatures inside the burrows can drop to 32 °C at a depth of 250 mm. Variation in the burrow architecture with crab age appears to be related to the crab’s behaviour. Juvenile crabs have smaller gill areas and move out of the burrows regularly to renew their respiratory water and, as a result, they do not need a deep burrow. Larger crabs, in contrast, can tolerate prolonged periods without renewing their respiratory water and therefore create deeper and more complex burrows for mating and refuges.