Burrowing Crab Chasmagnathus Granulatus Research Articles

Cadmium and Zinc in Mar Chiquita Coastal Lagoon (Argentina): Salinity Effects on Lethal Toxicity in Juveniles of the Burrowing Crab Chasmagnathus granulatus

The large Argentine marine littoral zone is characterized by great number of wetlands and includes only one coastal lagoon, Mar Chiquita, which has been declared as a Biosphere Reserve by the Man and Biosphere Reserve Program from UNESCO. Its margins present populations of Chasmagnathus granulatus, a semiterrestrial crab distributed along wide salinity gradients that plays an important role as a key species within the corresponding trophic web. Dissolved cadmium (Cd) and zinc (Zn) concentrations present in this ecosystem were determined. Cadmium concentrations ranged between n.d. and 0.82 mug/L and zinc levels ranged between n.d. and 1224.38 mug/L within the mentioned coastal lagoon. Cd and Zn acute semistatic toxicity bioassays were carried out for 96 h on juvenile crabs of C. granulatus. LC(50) 96-h values were 2.24 mg Cd(2+)/L and 7.07 mg Zn(2+)/L at 5 psu, and 15.42 mg Cd(2+)/L and 11.41 mg Zn(2+)/L at 25 psu. Higher salinities resulted in lower metal toxicity. This effect was stronger for Cd than for Zn. C. granulatus juveniles LC(50) 96-h values determined for Cd were three to four orders of magnitude higher than the corresponding dissolved metal concentrations in the Mar Chiquita coastal lagoon; nevertheless, those Zn values determined were similar to several ones corresponding to natural water samples.

Indirect effects of the intertidal burrowing crabChasmagnathus granulatus in the habitat use of Argentina’s South West Atlantic salt marsh birds

The burrowing crab,Chasmagnathus granulatus, is the dominant benthic macroinvertebrate of southwest Atlantic salt marshes and tidal flats, having strong ecosystem engineering effects by direct and indirect effects on soil, vegetation, invertebrates, fishes, and birds. Vegetation structure is a main component for bird habitat selection, since greater habitat complexity generates higher niche diversity. This environmental complexity can be modified by species interactions or disturbance events (i.e., by crabs), in turn modifying the associated community. The bird species of salt marshes of the southwest Atlantic are highly dependent on these ecosystems. We assessed the effects of the burrowing crab on the structure of the cordgrass,Spartina densiflora, marshes, and how these changes affected the composition and diversity of birds. This study was conducted at the Mar Chiquita Coastal Lagoon, Argentina (37°40′S, 57°23′W).S. densiflora marshes were classified within three categories: marshes without burrowing crabs, marshes with low burrow densities, and marshes with high burrow densities. We made vegetation and bird surveys during spring beforeS. densiflora produce seeds and in summer when seeds are abundant. We also manipulated inflorescence (i.e., spike) density (a variable affected by crabs) to assess the effect on bird habitat use. The highest inflorescence density ofS. densiflora and highest bird diversity were recorded at low burrow densities. Cover and height ofS. densiflora were negatively correlated with burrow density. The number of bird species and the number of red-capped wren-spinetail,Spartonoica maluroides, were negatively related with cover and height ofS. densiflora. Experiments confirmed that granivorous species used areas with higher spike densities, a variable related to crab burrow density. Burrowing crabs directly and indirectly affect the cover, height, spike density, and morphologic characteristic of seeds ofS. densiflora marshes. These effects indirectly affect the bird community that uses these marshes, being negatively related toS. maluroides and total bird abundance, and positively related to bird diversity.

Changes in distribution and abundance of juvenile fishes in intertidal soft sediment areas dominated by the burrowing crab Chasmagnathus granulatus

Intertidal estuarine areas located between southern Brazil and the northern Argentinean Patagonia are characterised by extended beds of the burrowing crab Chasmagnathus granulatus. Their activity leads to profound changes in the structure, quality and dynamics of sediments, which affect the entire benthic community and create a patchy distribution of resources for other species. In this study, the differences in habitat use by juvenile fishes above soft sediments inhabited by C. granulatus were evaluated. Adjacent areas with (‘crab beds’) and without crab burrows in intertidal mudflats of the Mar Chiquita Coastal Lagoon (37°32'S, 57°19'W) were sampled seasonally from 2000 to 2002. There were more fish species in crab-bed areas. Abundances of benthivorous and planktivorous fishes were also higher in crab beds during flood tide; however, during ebb tide, planktivorous fishes were more abundant outside crab-bed areas, whereas benthivores did not vary between areas. The present study demonstrated that species composition and abundance of juvenile fishes can be affected by the presence of bioturbator species such as C. granulatus.

Effects of the dominant SW Atlantic intertidal burrowing crab Chasmagnathus granulatus on sediment chemistry and nutrient distribution

Through field experiments and chemical analysis of the sediment and pore water, we investigated the effect of the burrowing crab Chasmagnathus granulatus on the distribution and availability of electron acceptors in mudflat sediments. The results show that the presence and activ- ity of C. granulatus and its burrows determine the chemical characteristics of pore water and the redox state of mudflat sediments. Crabs enhance the transport of particulate material in the sediment column, completely mixing the upper 7 cm of sediment in a few days. Comparative analyses of pore water from areas (1) with crabs and burrows, (2) with unoccupied burrows, and (3) without burrows or crabs reveal a large increase in sediment oxygenation, modification of pore water salinity and the distribution of dissolved inorganic nitrogen and sulphate when crabs are present. Denitrification and organic matter (OM) degradation rates were estimated by a diagenetic model that searches for simultaneous agreement between measured and model-calculated depth profiles. OM degradation rates (kG) were found to be greater in bioturbated (kG = 10.6 ± 6.1 (SD) µM s -1 ) than in non-bio- turbated sediments (kG = 4.9 ± 17.8 µM s -1 if unoccupied burrows were present; kG = 0.02 ± 0.013 µM s -1 without burrows). Model-estimated denitrification rates revealed that OM degradation pathways are also affected by C. granulatus activities and their burrows. These changes in the sediment chemistry that are directly and indirectly produced by C. granulatus could comprise the force that drives the pathways of microbial processes and nutrient flows to neighbouring systems.

The effect of the SW Atlantic burrowing crab Chasmagnathus granulatus on the intertidal razor clam Tagelus plebeius

The deposit feeding burrowing crab Chasmagnathus granulatus and the suspension feeding burrowing clam Tagelus plebeius occur sympatrically in the intertidal flats of the south western Atlantic estuarine environments. Common ecological theory predicts negative effects of deposit feeder bioturbation on co-occurring suspension feeders up to complete exclusion of the latter. By means of descriptive and experimental approaches performed at the Mar Chiquita coastal lagoon (37°32′S–57°19′W; Argentina) we show that within beds of C. granulatus (i) frequency of clam shell damage is higher, (ii) shell geometry shows proportionally shorter shells, (iii) clam condition is lower and, (iv) shell growth rates are lower. The negative effect of crabs increases towards lower tidal level. Changes in shell geometry are only detectable in damaged shells, suggesting that this pattern results from direct physical effects by crabs. The negative effect of crabs on clam condition and growth is mediated by indirect mechanisms. Burrowing and migrating of crabs may disrupt clam feeding and/or may force clams to invest more energy into escape movements and burrow rebuilding, thus leaving less energy for growth. Our study indicates that the adaptation of T. plebeius to a highly dynamic intertidal environment enables the clam to coexist with this strong bioturbator, too. The additional stress inflicted by C. granulatus does affect the clam negatively, but is not sufficiently high to eliminate it from the system.

Effect of fish predation on intertidal benthic fauna is modified by crab bioturbation

The burrowing crab Chasmagnathus granulatus is an important bioturbator in SW Atlantic estuaries where they generate dense and extended intertidal beds. Its bioturbation leads to profound changes in the structure, quality and dynamics of sediments with concomitant impacts on the entire benthic community. In this study, we evaluate whether the presence of this crab affects the predator–prey interaction between juvenile fishes and their benthic prey. Gut content and benthic prey selection by juvenile fishes inside and outside crab beds were evaluated, and predation effect was experimentally contrasted between areas using fish exclosures. The results show that in crab beds the percentage of fish with empty guts was lower and the number of polychaetes consumed by fish higher than outside crab beds. The silverside Odontesthes argentinensis and the catfish Pimelodella laticeps fed on larger polychaetes outside than inside crab bed areas, while the white mouth croaker Micropogonias furnieri preyed upon larger polychaetes inside crab beds. In addition, field experiments shows that fish predation decreases polychaete abundances only in crab beds. These results suggest that crab bioturbation facilitate fish predation on benthic prey.

Small- and large-scale effect of the SW Atlantic burrowing crab Chasmagnathus granulatus on habitat use by migratory shorebirds

Here we address the question of whether the presence of the burrowing crabs Chasmagnathus granulatus affects small- and large-scale habitat use by migrant shorebirds. This crab is the dominant species in soft bare sediments and vegetated intertidal areas along the SW Atlantic estuaries (southern Brazil 28°S to the northern Argentinean Patagonia 42°S). They generate very extensive burrow beds in soft bottom intertidal areas. Our information shows that this burrowing crab affects the small-scale habitat use by shorebirds, given that shorebirds never walk through the funnel-shaped entrances of burrows. Given that crab burrow entrances occupy up to 40% of the intertidal area, there is a large decrease of available shorebird habitat in crab beds, restricting their activity to the spaces between the burrows. The southern migratory shorebird Charadrius falklandicus maximize the use of these areas by foraging closer to the burrows than the other bird species. Neotropical migrants, such as Calidris fuscicollis, Pluvialis squatarola and Tringa melanoleuca, used foraging paths that tended to maximize the distance from burrows, especially the distance to larger burrows. A field experiment showed that this was not necessarily due to a decrease in the availability of polychaetes near the crab burrows. A combination of landscape measurements and satellite images showed that crab beds covered up to 40% of the intertidal area of the Mar Chiquita coastal lagoon (37°40′S, Argentina), and nearly 100% of the intertidal area of the Bahia Blanca estuary (38°48′–39°25′S, Argentina). These two estuaries are located along the migratory flyway of Neotropical migratory shorebirds, but the Bahia Blanca estuary (area∼110,000 ha) shows a much lower shorebird diversity than Mar Chiquita (area∼4500 ha). The most common species in Bahia Blanca is the two-banded plover C. falklandicus, the species least affected by crabs at Mar Chiquita and which prefers to use high-density crab areas as foraging sites. The oystercatcher Haematopus palliatus was also most abundant in high-density crab areas, but they used these areas for resting. The abundances of preys varied during the study period and between the crab density areas, indicating that the use of these areas by birds is independent of crab density. However, burrowing crabs affect the depth distribution of polychaete and thus their availability to shorebirds. We suggest that this shorebirds–burrowing organism interaction could be generalized for other intertidal estuarine habitats.

Effects of the intertidal burrowing crabChasmagnathus granulatus on infaunal zonation patterns, tidal behavior, and risk of mortality

Rhythmic movements in response to tidal cycles are characteristic of infaunal inhabitant of intertidal soft-bottoms, allowing them to remain in the area with best living conditions. The effect of bioturbators as modifier of local environmental conditions and thus of gradients in intertidal habitats, has not been investigated yet. The Atlantic estuarine intertidal areas are dominated by the burrowing crabChasmagnathus granulatus that generates strong environmental heterogeneity by affecting the physical-chemical characteristics of the sediment. The comparison between intertidal areas with and without crab shows that sediments in the crab beds remain more humid, softer, and homogeneous across the intertidal and along the tidal cycle than areas without crabs. The densities of infauna were higher at high intertidal zones in crab beds than in similar areas without crabs. Infaunal organisms performed vertical movements into the sediment following the tidal cycle that were always of higher magnitude in habitats without crabs. Infaunal species tend to spend most of the time buried into the sediment in the crab bed. Migratory shorebirds use the Atlantic estuarine environments as stopover or wintering sites. They feed (mainly on polychaetes) in the low intertidal zones of both habitats (with and without crabs), but they also feed in the upper intertidal of the crab bed; polychaete per capita mortality rate is higher in the upper part of the crab bed. Environmental heterogeneity produced by crab disturbance has an effect on the infaunal behavior, risk of mortality, and the zonation pattern. This is another example of the ecosystem engineering ability of a burrowing intertidal species.

Evidences of habitat displacement between two common soft-bottom SW Atlantic intertidal crabs

The intertidal burrowing crab Chasmagnathus granulatus Dana is the dominant species in soft sediments and vegetated intertidal areas along the SW Atlantic estuaries (southern Brazil 28°S to the northern Argentinean Patagonia 41°S) where it produces dense and extensive burrowing beds. The mud crab Cyrtograpsus angulatus Dana coexists with Ch. granulatus in this area, but it also inhabits areas to the south (northern and central Argentinean Patagonia). A survey covering both areas showed that C. angulatus rarely live in burrows when coexisting with Ch. granulatus, but form large burrowing beds when not coexisting with Ch. granulatus. When both species coexisted, burrowing beds of C. angulatus are restricted to sandy–muddy areas. Only rarely are burrows of C. angulatus found within Ch. granulatus beds. However, when Ch. granulatus were experimentally excluded within their burrowing beds, new settlers of C. angulatus made burrows and maintained them until they reached large size. Paired (inside and outside Ch. granulatus burrowing bed) sampling during high tide using beach nets showed that C. angulatus rarely venture inside the Ch. granulatus crab beds. Other field experiments showed that adults Ch. granulatus always displace C. angulatus from burrows. Furthermore, in several sites located south of the limit of distribution of Ch. granulatus at the Patagonian coast, soft bare intertidals are dominated by burrowing beds of C. angulatus mixed with the congener C. altimanus Dana. Together, these evidences suggest that the mud crab C. angulatus is displaced from soft bottom areas by the burrowing crab Ch. granulatus. It is an example of competitive exclusion through aggressive interference in soft-bottom habitats when the shared resource is the access to sediment surface, a two-dimensional well-defined resource.

Changes in the feeding behavior of the deposit-feeding polychaete Laeonereis acuta on soft sediments inhabited by burrowing crabs

In this paper, we evaluate the influence of the burrowing crab Chasmagnathus granulatus on the feeding behavior, body condition, burrowing activity, and fecal production of the deposit-feeding polychaete Laeonereis acuta. Previous results and ours show that, due to crab activity, sediment organic matter decreases with depth outside a crab bed, but there were no differences inside. Also, particle sizes were smaller inside the crab bed. Polychaetes showed better body condition inside the crab bed, suggesting higher energy gain. They fed on the surface more frequently outside the crab bed than inside. However, feeding rate was higher inside the crab bed, which is consistent with the larger weight and volume of fecal pellets found inside crab bed. In both areas, the bedload sediment transport was not similar to the sediment ingested by this polychaete. A 1-month exclusion/inclusion experiment performed outside and inside the crab bed showed that the body condition of polychaetes changed between areas but that this was not directly due to crab manipulation. Thus, our results suggest that the higher sediment quality inside a crab bed can positively affect the feeding behavior of the deposit-feeding polychaete L. acuta, increasing its feeding rate and consequently enhancing its body condition.

The SW Atlantic burrowing crab Chasmagnathus granulatus Dana affects the distribution and survival of the fiddler crab Uca uruguayensis Nobili

In this paper, we address the question of whether the presence of the burrowing crab Chasmagnathus granulatus affects the habitat use of the fiddler crab Uca uruguayensis. Field samples showed that the species have a disjoint spatial distribution. Male fiddler crab density decreased in zones with C. granulatus, however, female density increased. Male fiddler crabs avoided feeding on sediment affected by C. granulatus and were more preyed. Predation was higher during the fiddler crab reproductive season and, probably due to predation risk, males showed lower reproductive display in shared zones. Field experiments shows that when C. granulatus were excluded, densities of U. uruguayensis increased mainly due to an increase in density of males. Habitat differentiation of these species may be because C. granulatus affects U. uruguayensis in several ways, including direct predation, disturbance and behavioural changes associated to predation risk. Males and females are affected differentially probably because of the extreme sexual dimorphism of this crab species. Coloration on enlarged claw and waving activities are all factors that increase predation risk for male and the presence of only one feeding claw may increase sediment-mediated effects.

Does the presence of the SW Atlantic burrowing crab Chasmagnathus granulatus Dana affect predator–prey interactions between shorebirds and polychaetes?

The burrowing crab Chasmagnathus granulatus is an important bioturbator that generates dense burrow assemblages (crab beds) characteristic of intertidal habitats of SW Atlantic estuaries. Crab bioturbation affects the topography and hydrodynamics of the sediment, increasing sediment water and organic matter content, decreasing sediment hardness and changing the grain size frequency distribution. In this study, we found that burrowing crabs can decrease the impact of predation by shorebirds on polychaetes. The polychaete Laeonereis acuta Treadwell has U-shaped burrows outside crab beds, which are associated with surface deposit-feeding while their burrows are mainly I-shaped inside which is associated with subsurface deposit feeding behavior. This pattern is likely the result of larger vertical sediment mixing inside crab beds due to crab burrowing. As a result of their feeding strategy, polychaetes appear on the surface more often outside crab beds, which increases their availability for shorebirds. In addition, shorebird species differentially use crab beds. The White-rumped Sandpiper, Calidris fuscicollis Vieillot, preferentially forage outside crab beds, meanwhile the Two-banded Plover Charadrius falklandicus Latham forage more frequently inside crab beds. However, experiments excluding shorebirds inside and outside crab beds showed negative effects of shorebirds only outside crab beds. Thus, our results show that the SW Atlantic burrowing crab C. granulatus affects the strength of the predator–prey interaction between shorebirds and polychaetes.

Elevated predation risk changes mating behaviour and courtship in a fiddler crab

The fiddler crab, Uca beebei, lives in individually defended burrows, in mixed-sex colonies on intertidal mud flats. Avian predation is common, especially of crabs unable to escape into burrows. Mating pairs form in two ways. Females either mate on the surface at their burrow entrance ('surface mating') or leave their own burrow and sequentially enter and leave ('sample') courting males' burrows, before staying in one to mate underground ('burrow mating'). We tested whether perceived predation risk affects the relative frequency of these mating modes. We first observed mating under natural levels of predation during one biweekly, semi-lunar cycle. We then experimentally increased the perceived predation risk by attracting grackles (Quiscalus mexicanus) to each half of the study site in two successive biweekly cycles. In each experimental cycle, crabs were significantly less likely to mate on the side with more birds. Moreover, on the side with elevated predation risk, the number of females leaving burrows to sample was greatly reduced relative to the number of females that surface-mated. Males waved less and built fewer mud pillars, which attract females, when birds were present. We discuss several plausible proximate explanations for these results and the effect of changes in predation regime on sexual selection.