Spatial Heterogeneity Created by Burrowing Crabs Affects Human Impact on Migratory Shorebirds

We show that, as previously described, there are conflicts between areas used by people and shorebirds. However, we propose that burrowing organisms add complexity to the shorebird–human interaction that should be taken into account for management and conservation. This is because SW Atlantic mudflats are dominated by high densities (up to 60 crabs m⁻²) of a strong bioturbator, the burrowing crab Neohelice granulata (=Chasmagnathus granulatus). These crabs affect the habitat use and foraging performance of shorebird species. The two-banded plover Charadrius falklandicus feed more in burrow areas whereas the yellowlegs Tringa spp. and the white-rumped sandpiper Calidris fuscicollis tend to avoid burrow areas. We evaluated the effect of human recreational activities on shorebirds, taking into account shorebird–crab interaction, during shorebird migratory periods (November to April) of 2001, 2002, and 2003. This study was performed at the Mar Chiquita coastal lagoon (37° 40′ S, Argentina). Results showed that people only rarely used the extensive soft bottom intertidals dominated by burrowing assemblages of N. granulata because the soft sediments make walking difficult. Therefore, human impact is reduced for the two-banded plover. However, for several migratory shorebirds that also avoid burrow areas, these areas act as suboptimal alternative areas when human impact is important in their preferred habitats. When human activity is low, shorebirds remain foraging in the area but they decrease their feeding intakes. The extensions of burrow areas differ among estuaries and are likely to generate between-estuaries differences as stopover sites for shorebirds. Also, other species that form large burrowing assemblages can have a similar impact on shorebird–human interactions.     

Swimming behavior of Dungeness crab,Cancer magister Dana,megalopae in still and moving water

In the Grays Harbor estuary, juvenile Dungeness crab (Cancer magister Dana) are found at higher densities in epibenthic shell deposits compared to open mud flat. Differences in predation rate between habitats have been suggested to be due to habitat preference and differential survival. Megalopae preferred shell over open space in still-water conditions. However, it is not known whether megalopae are able to select shell in flowing water since larval preference is known to differ between still and flowing water. Here we report the first experimental study of swimming behavior of Dungeness crab megalopae in a range of current velocities (0–40 cm s^?1) equivalent to natural flow in Grays Harbor estuary. Experiments were conducted in daylight using a recirculating flume. Megalopae swimming speeds ranged from 8.5 cm s^?1 (8 body lengths s^?1) in still water to 44.8 cm s^?1 (44 body lengths s^?1) at flow speeds of 40 cm s^?1, Neither swimming behavior nor sheltering behavior in shells showed any flow-related pattern. Megalopae spent a large proportion of time swimming against the current and made headway upstream against all current velocities tested. The results suggest that Dungeness crab megalopae are able to maneuver and actively search for settlement sites under current velocities found in natural habitats, including intertidal shell deposits, and support the hypothesis of active selection of shell by megalopae.     

Relationship between waterfowl and the seagrassRuppia maritima in a Southwestern Atlantic coastal lagoon

We evaluated the distribution of waterfowl in relation to a seagrass (Ruppia maritima) patch, to infauna, and on its relationship with substrate characteristics. An experiment performed in the Mar Chiquita coastal lagoon (37°46′S, 57°27′W; Argentina) was used to evaluate the effect of herbivory on widgeon grass,R. maritima. Control plots of equal size, located between bird exclosures, were exposed to herbivory. The experiment ran for 31 d, starting on December 1, 1994. Censuses showed that black-necked swan ( $\bar x = 50$ birds ha^?1, SD = 37, n = 10) and coots ( $\bar x = 42$ birds ha^?1, SD = 28, n = 10) were the most important (always present) of the waterfowl seen feeding onR. maritima. The results of the experiment showed greater leaf lengths, lower belowground (rhizomes and roots) biomass, greater aboveground (leaves and shoots) biomass, and greater abundance of the polychaeteHeteromastus similis in exclosure plots. There were no exclosure effects on total biomass (belowground plus aboveground), reproductive parts (fruits and pre- and postpollination flowers), or abundance of most benthic infauna. Topographic surveys showed that sediment surface was higher within theR. maritima patches, but there were no significant differences in granulometric composition. Surveys also showed that bird distribution was strongly associated with the presence ofR. maritima.     

Diversity and Habitat Distribution of Birds in Coastal Marshes and Comparisons with Surrounding Upland Habitats in Southeastern South America

We studied variation in bird assemblages with plant associations for three different coastal marshes from Southeastern South America (SESA) and assessed how marsh bird assemblages related to nearby upland bird assemblages. We surveyed bird species and plant structure along the tidal gradient of each locality from the low tide level to the upper habitats bordering coastal marshes. Twenty species frequently used coastal marshes, including relatively few migratory species. We found that birds occurring in SESA coastal marshes do not have distributions constrained to coastal marshes. Nonetheless, four bird assemblages were recognized in association with vegetation types and/or sites. Among the recorded coastal marsh species, the bay-capped wren-spinetail (Spartonoica maluroides) is both the most frequent and the most habitat constrained. Bird richness increases steadily along the tidal gradient associated with the increase in vegetation structure, suggesting that bird richness is directly explained by vegetation and indirectly by the physical conditions influencing vegetation structure. Results highlight the importance of SESA middle marshes as habitat for conservation of some threatened SESA grassland birds.     

Population Structure of the SW Atlantic Estuarine Crab Neohelice granulata Throughout Its Range: a Genetic and Morphometric Study

Phylogeographic patterns of the SW Atlantic estuarine crab Neohelice granulata were examined using mitochondrial DNA cytochrome oxidase I sequences and analyzed together with morphometric data. Specimens were sampled during a 4-year period (2001?C2004) from 11 localities encompassing the full distributional range of this species along the SW Atlantic (22°57?? S to 42°25?? S). DNA sequences were obtained from 69 individuals belonging to seven localities, and morphometric variation in 12 continuous characters was analyzed for 646 crabs from ten localities. Strong genetic differentiation, consistent with a pattern of isolation-by-distance, was detected among all localities indicating that gene flow occurs mainly between neighboring populations. Analyses of molecular variance showed genetic subdivision between the southern (Argentina) and the northern (Brazil) sites, suggesting restricted gene flow at a regional scale. The genetic structure of this species could be divided into two distinct groups due to a limited gene flow between southern and northern regions as a consequence of larval dispersal patterns. Coastal currents in the vicinity of the Rio de la Plata likely act as a barrier to dispersal within the species range. Moreover, genetic data indicate that populations of N. granulata might have undergone a northward demographic expansion since the late Pleistocene. The morphometric analysis showed no geographical pattern of morphological differentiation, although there were differences among sampling sites.     

Origin of resources and trophic pathways in a large SW Atlantic estuary: An evaluation using stable isotopes

The Río de la Plata (34° 36′ S, 55° 58′ W; Argentina and Uruguay) estuary, one of the most important South American estuarine environments, is characterized by weak seasonal freshwater discharge, low tidal amplitude (<1 m), a wide and permanent connection to the sea, and a salt-wedge regime. Using stable isotope analysis, we explored the relative importance of the different sources of primary production in the food web. Our results show that phytoplankton and macrodetritus from terrestrial salt and freshwater marshes both contribute to the food web of the Río de la Plata estuary. On the basis of the sampled species, we identified four trophic levels. The clam Mactra isabelleana, the calanoid copepod Acartia tonsa, and the opossum shrimp Neomysis americana are the primary consumers. The rays Atlantoraja castelnaui and Squatina guggenheim and the shark Galeorhinus galeus are the top predators. The Río de la Plata food web shows an important input of nutrients derived from phytoplankton. Rays, sharks, and predatory gastropods reveal an important contribution of C4 plants (likely Spartina spp.). However, production derived from C3 plants is also important for some species. The fishes Brazilian menhaden, Brevoortia aurea; the stripped weakfish Cynoscion guatucupa; and the whitemouth croaker, Micropogonias furnieri, showed differences in their isotopic signatures as juveniles and adults, indicating different food sources, and they were therefore treated as different components of the food web. Our data suggest that detritus from salt and freshwater marshes is reaching the Río de la Plata estuary and can be an important allocthonous source of energy to this environment.     

The Intertidal Burrowing Crab <Emphasis Type="Italic">Neohelice</Emphasis> (=<Emphasis Type="Italic">Chasmagnathus</Emphasis>) <Emphasis Type="Italic">granulata</Emphasis> Positively Affects Foraging of Rodents in South Western Atlantic Salt Marshes

The role of positive and indirect interactions is often crucial in communities with intense abiotic stress such as salt marshes. The burrowing crab, Neohelice (=Chasmagnathus) granulata, is the dominant benthic macroinvertebrate of southwest Atlantic marshes (southern Brazil to Northern Argentinean Patagonia), having strong direct and indirect effects on marsh soil and, in consequence, on marsh vegetation and primary consumers. In this work, we investigate if this crab indirectly modifies habitat use by the granivorous rodents, Akodon azarae and Oligoryzomys flavescens, by increasing nutrient availability and thus enhancing seed production by the marsh plant Spartina densiflora. The study was conducted at the Mar Chiquita Coastal Lagoon, Argentina (37°32′ S). Rodent frequencies in S. densiflora were positively correlated with crab densities throughout the low and middle marsh. Additionally, the highest quality of S. densiflora and inflorescence density was recorded at the highest crab densities. Experimental manipulation of crab densities shows that N. granulata indirectly enhances the performance of S. densiflora (e.g., decreased fiber content and C/N ratios) and increases density of seeds. Moreover, N. granulata also facilitates S. densiflora seed availability to rodents by concentrating them in sediment mound at their burrows entrances. Experimental rodent exclusions showed that rodent species used S. densiflora seeds, a variable positively related to crab burrow density. Thus, our results show that N. granulata drives the granivorous rodent distribution and the intensity of seeds–rodent interaction trough facilitative and indirect interactions in marsh community.     

Field accumulative behavior of organochlorine pesticides. The role of crabs and sediment characteristics in coastal environments

The influence of intertidal crab beds on the concentrations of organochlorine (OC) pesticides in sediment was studied in two different coastal environments in Argentina. Samples of male burrowing crabs (Chasmagnathus granulatus) were collected for this study. Our field data showed lower bioaccumulation of OC pesticides in crabs from sediments with a higher total organic carbon (TOC) and higher clay content. Thus, concentrations in crabs depend on the physico-chemical characteristics of the sediment where they live more than on the OC pesticide concentrations in the environment. The distribution patterns in sediment from inside and outside crab burrows were similar for both coastal areas being HCHs  γ-chlordane > p,p′-DDE for San Antonio Bay (SAO), and HCHs > p,p′-DDE  γ-chlordane for Mar Chiquita (MCh) coastal lagoon. OC pesticide concentrations in sediment were significantly lower inside than outside crab burrows, irrespective of the sediment physico-chemical characteristics due to the bioturbation activity of C. granulatus.     

The Southwest Atlantic intertidal burrowing crab Neohelice granulata modifies nutrient loads of phreatic waters entering coastal area

Along the coastal areas of the Southwest Atlantic estuaries and embayments, phreatic water often circulates through very extended areas (up to several hundred meters perpendicular to the coast), dominated by dense assemblages of deep burrows of the crab Neohelice granulata (formerly Chasmagnathus granulatus). This crab inhabits the intertidal area, from mudflats to marshes vegetated by species of Spartina, Sarcocornia and Juncus, generating extensive burrowing beds where burrow density may reach up to 60 burrows m⁻². Since the lower limit of the crab burrows is usually the water table, we investigated through field experiments the effect of N. granulata and their burrows on the chemical characteristics of this phreatic water. Water analysis from experimental (1) occupied burrows (with crabs), (2) unoccupied burrows (where crabs were excluded), and (3) sediment pore water show remarkable differences. Water oxygenation, and nitrate, ammonium and sulphate concentrations inside occupied burrows were higher than in the water inside unoccupied burrows or pore waters. Moreover, directed sampling of phreatic water entering and leaving the crab bed, shows that dissolved inorganic nitrogen concentration is enhanced as the water crosses the crab bed. These results may be ascribed to the fact that in the salt marsh the crabs spend most of their time within burrows, where presumably they store food (plants) and defecate. These activities generate an area of accumulation of excrements and nutrients in different decomposition states. The present work shows a novel way by which bioturbating organisms can affect nutrients exportation from salt marshes to the open waters.     

Indirect effects of the intertidal burrowing crab<Emphasis Type="Italic">Chasmagnathus granulatus</Emphasis> 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.