Habitat utilization and alteration by the invasive burrowing isopod, Sphaeroma quoyanum, in California salt marshes

In recent years the pace of exotic species introduction and invasion has accelerated, particularly in estuaries and wetlands. Species invasions may affect coastal ecosystems in many ways. Alteration of sedimentary environments, through structure formation and burrowing, has particularly dramatic effects on coastal habitats. This study examines modification of channel bank and marsh edge habitat by the burrowing Australasian isopod Sphaeroma quoyanum Milne Edwards, in created and natural salt marshes of San Diego Bay and San Francisco Bay. Abundance and distribution patterns of this isopod species, its relationships with habitat characteristics, and its effects on sediment properties and bank erosion were examined seasonally, and in several marsh microhabitats. Mean isopod densities were 1541 and 2936 individuals per 0.25 m² in San Francisco Bay, and 361 and 1153 individuals per 0.25 m² in San Diego Bay study sites during December and July 1998, respectively. This isopod forms dense, anastomosing burrow networks. S. quoyanum densities did not differ as a function of location within creeks or location in natural versus created marshes. Burrows, which are on average 6 mm wide and 2 cm long, were associated with firm sediments containing high detrital biomass. Although erosion is a natural process along salt marsh banks, enclosure experiments demonstrated that isopod activities can enhance sediment loss from banks. In areas infested with S. quoyanum, losses may exceed 100 cm of marsh edge per year. The effects of habitat alteration by this invading species are likely to increase in severity in the coastal zone as these ecosystems become degraded. Received: 30 March 2000 / Accepted: 21 September 2000     

Recruitment and ontogenetic habitat shifts of the yellow snapper (<Emphasis Type="Italic">Lutjanus argentiventris</Emphasis>) in the Gulf of California

We examined recruitment and ontogenetic habitat shifts of the yellow snapper Lutjanus argentiventris in the Gulf of California, by conducting surveys and collections in multiple mangrove sites and major marine coastal habitats from 1998 to 2007. Over 1,167 juvenile individuals were collected and 516 otoliths were aged to describe the temporal pattern of the settlement. L. argentiventris recruits in mangroves, where juveniles remain until they are approximately 100 mm in length or 300-days-old. Back-calculated settlement dates and underwater surveys indicated a major recruitment peak during September and October, around 8 days before and after the full moon. The majority of mangrove sites in the Gulf of California had a similar L. argentiventris average size at the beginning of the settlement season for the cohort of 2003; although there were significant differences in individual sizes at the end of the nursery stage. When sub-adults leave mangroves, they live in shallow rocky reefs and later become abundant in deeper rocky reefs. The density of migratory individuals (10–20 cm SL) decreased exponentially as the distance between a reef and a nearby mangrove site increased. This finding has important implications for local fishery regulations and coastal management plans.     

Differences in context and function of two distinct waving displays in the fiddler crab, <Emphasis Type="Italic">Uca perplexa</Emphasis> (Decapoda: Ocypodidae)

To respond appropriately to communication signals, animals must have the ability to decipher signal meaning. At a basic level, interpreting the difference between territorial and courtship signals can be vital for the survival and reproduction of social animals. Male and female fiddler crabs communicate with claw-waving displays, but the function of these waves remains uncertain. Species differ in the context in which they wave: Some wave during courtship, some during territorial defence and some during both. In this paper, we provide evidence that males of an Australian species of fiddler crab, Uca perplexa, use two different types of claw waving display, lateral and vertical. Lateral waves are employed solely in a courtship context, whilst vertical waves are employed during courtship as well as territorial interactions. Using video recordings, we show that lateral waves were produced spontaneously (broadcast), and their frequency increased only in the presence of female wanderers. Vertical waves were not broadcast but were elicited by male wanderers during agonistic interactions and female wanderers during close range courtship. Male resident U. perplexa were able to discriminate the sex of wandering crabs on the mudflat at distances of 32 cm. During all resident–wanderer interactions, residents attempted to maintain a position directly between the wanderer and the home burrow and orient themselves to face females and to present the major claw towards males. To our knowledge, this is the first demonstration of the multiple use of waving displays in a fiddler crab species.     

Feeding and burrowing ecology of two East African mangrove crabs

The behavior and ecology of two mangrove crabs,Sesarma meinerti De Man, 1887 andCardisoma carnifex (Herbst, 1794) were investigated at the beginning of the rainy season (October–November 1988) at Mida Creek, Kenya. Both species occupy upper intertidal levels, above mean high-water neap, and completely overlap in their zonation. Each burrow lasts ca. 3 wk, with no significant difference between the species. Both are more active around dusk and dawn and also follow a similar trend in their foraging activity, but differ in that burrowing is mainly diurnal inC. carnifex and nocturnal inS. meinerti. A hierarchy of food preference, established by offeringC. carnifex andS. meinerti leaves of five mangrove species, proved similar for both, withBruguiera gymnorhiza ranking first andAvicennia marina last, but significant only forC. carnifex. A rough estimate of the amount of litter consumed by these two species and of the soil mixed up by their burrowing activity indicates that they play a role of primary importance in the ecology of East African mangroves.     

Interactive effects of predation pressure and insecticide (temefos) toxicity on populations of the marsh fiddler crab Uca pugnax

The interaction of predation pressure with lethal and sublethal effects of temefos (Abate^®) an organophosphorous insecticide, were studied in field populations of Uca pugnax (S.I. Smith). Changes in fiddler crab population densities were followed in open-marsh temefos-treated and untreated test plots and in treated and untreated plots which were caged over to reduce predation by marsh birds. Temefos significantly reduced the population density of U. pugnax in the open test plots but not in the caged plots. These results indicate that temefos has a primarily sublethal effect on the crabs, the effect becoming lethal only after interaction with avian predation. Evidence from the presence of a time-lag effect in the population decrease, from a calculated predation index, and from laboratory studies reported elsewhere of behavioral alteration by temefos also supports the conclusion that temefos primarily impairs the escape response of U. pugnax: this leads to increased predation and subsequently to a decreased fiddler crab population. Such studies of sublethal effects of toxicants and field studies of interactions of lethal and sublethal effects of such compounds with natural population dynamics of affected species are necessary to evaluate possible effects of toxicants on populations.     

Mound building in Ilyoplax pingi (Crustacea: Brachyura: Ocypodidae)

The ocypodid crab Ilyoplax pingi, observed in Kanghwa I., Korea in 1992, builds a mound at the burrow entrance, by piling mud dug out from the burrow. The mounds were made by both waving and non-waving males, and by both ovigerous and non-ovigerous females. The burrow diameter at the widest part tended to be larger in crabs with mounds than in crabs without mounds, whereas the burrow depth was not different between them. The mound density increased in the late exposure period, when the crabs' surface activities declined. Mound removal and rebuilding experiments revealed that the presence of the mounds has the effect of keeping neighboring crabs away.     

Competition and interaction between Neosarmatium smithi (Crustacea: Grapsidae) and Terebralia palustris (Mollusca: Gastropoda) in a Kenyan mangrove

 In a Kenyan mangrove, we studied the interactions between the gastropod Terebralia palustris and the crab Neosarmatium smithi when foraging on decaying mangrove leaves. Interactions are considerable on account of their wide overlap in zonation (Rhizophora mucronata belt), food items (mangrove leaves) and activity window (diurnal low water). The snails find a leaf by a systematic transecting of the platform and eat the leaf after crawling on to it, usually en masse. The crabs rarely venture beyond 80 cm from their burrows, and once they find a leaf they quickly drag it back into their burrow, probably to reduce the strong intra-specific competition. A crab is able to drag a leaf away even if a number of snails are already feeding on it, by strongly pulling the leaf away or by pushing the snails off the leaf. The success of the “thief ” depends on both the crab's size and the number of snails on the leaf. Received: 25 July 1999 / Accepted: 27 April 2000     

Use of time, space and food resources in the mangrove climbing crab Selatium elongatum (Grapsidae: Sesarminae)

The activity patterns and use of space of the mangrove dweller Selatium elongatum were studied in the field, in combination with gut content analyses for evaluation of the crab's natural diet. This sesarmine crab proves to be one of the most adapted to climbing and living on mangroves. Moreover, S. elongatum showed a strong preference for being active at high tide, both during the day and night, always along the trunks a few centimetres above the water level, feeding on floating algae and mangrove leaves. Larger males were always recorded within activity areas into which they allow females and smaller males but exclude males of the same size, which probably compete for females. These results confirm that, in spite of the homogeneous herbivorous diet, little niche overlap exists among the several sesarmine crabs inhabiting the East African mangroves, all being leaf consumers segregated on the basis of their horizontal and vertical zonation, and/or of their activity rhythm. Received: 28 September 1998 / Accepted: 14 June 1999     

Some beavioral characteristics related to food supply and soil texture of burrowing habitats observed on Uca vocans vocans and U. lactea perplexa

From the observations on two species of fiddler crabs, droving is conspicuous in Uca vocans vocans but does not occur in U. lactea perplexa. U. vocans come down to feed at the water's edge with a richer food supply and fine substrate particles. Droving is advantageous to increase feeding efficiency in this species which holds burrows on a relatively sandy habitat where the food supply is less rich. In U. lactea, which burrows at higher levels, on the other hand, the food supply and soil texture of their burrow area are not significantly different from those at the water's edge. Home range relationships of both species are consistent with the occurrence or not of droving behavior and depend on food productivity of burrowing habitats.     

Role of Crab Herbivory in Die-Off of New England Salt Marshes

Abstract:  Die-offs of cordgrass are pervasive throughout western Atlantic salt marshes, yet understanding of the mechanisms precipitating these events is limited. We tested whether herbivory by the native crab , Sesarma reticulatum , is generating die-offs of cordgrass that are currently occurring on Cape Cod, Massachusetts (U.S.A.), by manipulating crab access to cordgrass transplanted into die-off areas and healthy vegetation. We surveyed 12 Cape Cod marshes to investigate whether the extent of cordgrass die-off on creek banks, where die-offs are concentrated, was related to local Sesarma grazing intensity and crab density. We then used archived aerial images to examine whether creek bank die-off areas have expanded over the past 2 decades and tested the hypothesis that release from predation, leading to elevated Sesarma densities, is triggering cordgrass die-offs by tethering crabs where die-offs are pervasive and where die-offs have not yet been reported. Intensity of crab grazing on transplanted cordgrass was an order of magnitude higher in die-off areas than in adjacent vegetation. Surveys revealed that Sesarma herbivory has denuded nearly half the creek banks in Cape Cod marshes, and differences in crab-grazing intensity among marshes explained >80% of variation in the extent of the die-offs. Moreover, the rate of die-off expansion and area of marsh affected have more than doubled since 2000. Crab-tethering experiments suggest that release from predation has triggered elevated crab densities that are driving these die-offs, indicating that disruption of predator–prey interactions may be generating the collapse of marsh ecosystems previously thought to be exclusively under bottom-up control .     

福建洛阳江口红树林湿地恢复对蟹类密度的影响

底栖动物尤其是蟹类对红树林湿地生态系统的生态功能恢复有重要意义,研究检验不同红树林种类及林龄和郁闭度是否对蟹类分布情况存在显著影响。从福建洛阳江口红树林湿地动态变化对蟹类密度的影响可以看出：不同植被类型对蟹类密度的影响达到极显著水平。光滩与成熟的秋茄（Kandelia candel）,桐花树（Aegiceras corniculatum）和白骨壤（Avicennia marina）之间有显著性差异（P〈0.05）,而桐花树、秋茄、白骨壤之间无显著性差异（P〉0.05）。在不同红树林树种和林龄2因素对蟹类密度的影响的双因素分析中：树种、林龄、树种和林龄间互作均对蟹类密度的影响达到极显著水平;在不同红树林树种和郁闭度2因素对蟹类密度的影响的双因素方差分析中：树种对蟹类密度的影响达到极显著水平,郁闭度对蟹类密度的影响达到显著水平;树种和郁闭度之间互作对蟹类密度的影响不显著。     

Adaptations to barnacle fouling in the mangroves <Emphasis Type="Italic">Kandelia obovata</Emphasis> and <Emphasis Type="Italic">Aegiceras corniculatum</Emphasis>

Mangrove trees planted for restoration or rehabilitation often fail to show satisfactory survivals. Although it is generally assumed that barnacle fouling is a factor in mangrove mortality, the sub-lethal response of mangroves to this fouling is not known. In the present study, we compare the leaf morphology (stomata density, leaf thickness, percentage thickness of palisade layer and number of hypodermal layers), lenticel density and the plant fitness (number of flowers or droppers per tree) between mangroves fouled by the acorn barnacle Fistulobalanus albicostatus (the fouled group, number of barnacles ~ 120 individuals per tree) and non-fouled mangroves (the control group). We observed 55 Kandelia obovata from three locations, two in Hong Kong (n = 35, fouled 25 and control 10) and one location in Taiwan (n = 20, fouled 10 and control 10). Aegiceras corniculatum (n = 45) was observed from two sites in Hong Kong, one on the east coast (n = 20, 10/10) and one on the west coast (n = 25, 10/15). In seedlings (K. obovata n = 10, 5/5), fouling density did not affect leaf morphology. In mature trees, however, leaves from fouled trees had significantly higher density of stomata and stem lenticel, a lower percentage thickness of leaf palisade layer and more layers of lower hypodermis in the leaves, probably for compensating the impeded gaseous exchange functions of the lenticels. The number of droppers and flowers on fouled mangroves was significantly lower. It is possible that after compensating for respiratory stress, the plant had less energy available for reproduction. Although there was no obvious barnacle-induced mortality in this study, barnacles appear to be related to a reduction in plant fitness. This may induce sufficient stress that over time fouled mangroves cannot sustain viable populations.     

Potential effects of a non-indigenous predator in its expanded range: assessing green crab, <Emphasis Type="Italic">Carcinus maenas</Emphasis>, prey preference in a productive coastal area of Atlantic Canada

The non-indigenous green crab (Carcinus maenas) is an important predator on bivalve wild beds in coastal areas worldwide. This study explored size-dependent green crab prey preference on American oysters (Crassostrea virginica), blue mussels (Mytilus edulis), and soft-shell clams (Mya arenaria) in a productive coastal system of Atlantic Canada. Using two sizes of prey and three different experimental manipulations, small, medium, and large green crabs were given a choice among these three bivalves, and their daily feeding rates were monitored over the course of 3 days. For both prey sizes, green crabs showed an early feeding preference for soft-shell clams and, only as they declined in numbers, a switch toward mussels and subsequently toward oysters. We found that such changes in the timing (order) of prey preference are related to prey differences in shell thickness, a fairly reliable indicator of prey shell strength.     

Influence of extracts of adult crabs on molting of fiddler crab megalopae (<Emphasis Type="Italic">Uca pugnax</Emphasis>)

Larvae of many benthic invertebrates metamorphose in response to habitat cues, which include the presence of adult conspecifics. Prior research showed that fiddler crab [Uca pugnax (Smith)] megalopae advance molting to the first crab stage in seawater in which conspecific adult crabs were maintained. In the present study, extracts of adult crabs were prepared and the specificity and protein content were characterized. U. pugnax megalopae were reared in the laboratory to minimize their prior exposure to potential molting cues. Then they were presented with extract solutions (in 1995, 1996, 1997, and 2000) to determine the specificity of the molting response to extracts of several crab species and the effects of the protein concentration of the extract, age of the megalopae at exposure to extract, and the duration of exposure on the molting response. Megalopae of U. pugnax molted sooner in seawater containing extract from adult conspecifics than in filtered seawater without extract or in seawater containing extract from the congener U. minax. Extract from the mud crab Dyspanopeus sayi retarded molting of U. pugnax megalopae. The stimulatory effect of U. pugnax extract on molting of megalopae was dependent on extract concentration. U. pugnax megalopae did not respond to extract until 6 days after molting, and only 1 day of exposure was sufficient to stimulate molting. The boiled extract was effective after being frozen (–15°C) for 2 years. The preparation of stable extract provides a uniform stimulus for multiple experiments examining the specificity of molting cues for brachyuran crustacean larvae and the onset of receptivity to cues.Communicated by J.P. Grassle, New Brunswick     

Population ecology of the mud crab <Emphasis Type="Italic">Scylla paramamosain</Emphasis> (Estampador) in an estuarine mangrove system; a mark-recapture study

Mud crabs of the genus Scylla are commercially important mangrove residents that are extensively fished throughout their range in the Indo-West Pacific. Despite this high level of exploitation very little is known about the population dynamics of any of the Scylla species. The present study concentrated on an exploited population of Scylla paramamosain in a natural estuarine mangrove on Can Coc Island in the mouth of the Hau River, Mekong Delta, Vietnam. A total of 6,114 juvenile crabs captured on the seaward mangrove fringe were internally tagged by injection of coded microwire tags and released over a period of 29 days. Recaptures were monitored over the following 144 days. There was little migratory movement within the estuary; of 285 recaptured crabs, 93% were recovered within the island mangrove study area and only two individuals were recovered from mangroves lining the opposite mainland riverbanks. A von Bertalanffy growth function was fitted to growth increment data from recaptured crabs, with L _max = 150 mm to derive a growth constant, k = 2.39 year⁻¹. The theoretical age at which carapace width is zero (t ₀) was derived from known size at recruitment at instar 1, giving a value of −0.0095 years. Previous studies of the same population have shown that female S. paramamosain reach maturity at a mean size of 102 mm carapace width. The present study indicates that they attain this size at around 160 days from first settlement in the mangrove fringe. Abundance of juvenile crabs in the study area was estimated by the Petersen method as 1,101,500 (95% CI 4,17,300–1,785,800, representing 1,102 crabs ha⁻¹ of mangrove. Estimation of mortality from tag returns and from the age-catch curves during a period of constant recruitment were comparable (Z = 1.11 and 1.04 month⁻¹, respectively). Fishing only accounted for 14% of total mortality, suggesting that at the time exploitation was not at a critical level despite the apparent high level of fishing activity.     

Response to perceived predation threat in fiddler crabs: trust thy neighbor as thyself?

Prey require information if they are to respond to predation threat in a risk-sensitive manner. One way that individuals can obtain this information is through the predator-mediated, threat-induced behavior of conspecifics. We examined such a possibility in a refuge-seeking species, the sand fiddler crab (Uca pugilator). Crabs were either exposed directly to a simulated predation threat (a moving cylinder) or the threat-induced response of a near neighbor. We found that fiddler crabs responded to the flight of their neighbors even when they, themselves, were not privy to the stimulus that induced their neighbors response. However, the wider range of behaviors exhibited by these crabs—which included no reaction, freezing, running back to the burrow entrance, and burrow retreat—suggest that non-threatened crabs either (1) perceived the gravity of the predation threat differently from their directly threatened neighbors and/or (2) engaged in behaviors that allowed them to acquire further information in the face of uncertainty. Conspecific behaviors also had an effect on the hiding duration of crabs, with individuals hiding longer if they saw both the predation threat and the flight of their neighbor. Our results suggest that cues provided by conspecifics can play an important role in guiding the antipredator response of refuge-seeking prey.     

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     

Effect of the presence of the shore crab, <Emphasis Type="Italic">Carcinus maenas</Emphasis>, on burrowing behaviour and clearance rate of the common cockle, <Emphasis Type="Italic">Cerastoderma edule</Emphasis>

Bivalves demonstrate various morphological and behavioural adaptations to reduce the risk of being attacked by predators. This paper examines how the presence of the crab Carcinus maenas (L.), a natural predator of the cockle Cerastoderma edule (L.), affects its burrowing depth and clearance or feeding rate. Cockles were placed in experimental tanks and treated with three levels of predatory disturbance: (1) unfed crab loose inside the tank, (2) unfed crab inside a cage suspended in the water column and (3) no crab present. Cockles’ burrowing depth was measured in two sediment types: mud and sand. Cockles burrowed more deeply in treatments with no crabs. Burrowing depth in sand was significantly greater than in mud. Two factors may contribute to the reduction in burial depth of C. edule in the presence of C. maenas: the change in the vertical orientation of the cockle and the ‘cough response’. No significant difference was found in the cockles’ clearance rate among the different levels of predator threat.     

Consumer Control of Salt Marshes Driven by Human Disturbance

Abstract:  Salt marsh ecosystems are widely considered to be controlled exclusively by bottom–up forces, but there is mounting evidence that human disturbances are triggering consumer control in western Atlantic salt marshes, often with catastrophic consequences. In other marine ecosystems, human disturbances routinely dampen (e.g., coral reefs, sea grass beds) and strengthen (e.g., kelps) consumer control, but current marsh theory predicts little potential interaction between humans and marsh consumers. Thus, human modification of top–down control in salt marshes was not anticipated and was even discounted in current marsh theory, despite loud warnings about the potential for cascading human impacts from work in other marine ecosystems. In spite of recent experiments that have challenged established marsh dogma and demonstrated consumer-driven die-off of salt marsh ecosystems, government agencies and nongovernmental organizations continue to manage marsh die-offs under the old theoretical framework and only consider bottom–up forces as causal agents. This intellectual dependency of many coastal ecologists and managers on system-specific theory (i.e., marsh bottom–up theory) has the potential to have grave repercussions for coastal ecosystem management and conservation in the face of increasing human threats. We stress that marine vascular plant communities (salt marshes, sea grass beds, mangroves) are likely more vulnerable to runaway grazing and consumer-driven collapse than is currently recognized by theory, particularly in low-diversity ecosystems like Atlantic salt marshes.