Territorial defense and advertisement by footdrumming in bannertail kangaroo rats (Dipodomys spectabilis) at high and low population densities

Summary Behaviors associated with territorial defense of large dirt mounds by bannertail kangaroo rats (Dipodomys spectabilis) at high and low population densities are reported. Rats were observed for 640 h during three summers and one spring from 1980–1982 in SE Arizona.Bannertail kangaroo rats defended their territories by footdrumming, as a long-distance warning signal, and chasing as a closer-distance threat. They footdrummed on or near their mounds spontaneosly during the night, in response to neighbors' footdrums, and durint mound challenges. There were no sexual differences in footdrumming, and juveniles frequently performed the behavior. The low frequency sounds were within the auditory sensitivities of the rats and footdrumming patterns differed somewhat between individuals. A playback experiment provided preliminary evidence that rats can differentiate between the footdrums of neighbors and strangers. When a bannertail visited the mound of another bannertail the mound owner actively defended its mound and immediately chased the visitor away. Fighting and mound challenges occurred infrequently, and the uniformly dispersed distribution of mounds suggested an effort by rats to avoid mutual interference.When population densities were high, dispersing juveniles built new mounds, consequently, distances between mounds decreased. Rats responded by tolerating closer neighbors, spending more time active on their mounds, and visiting neighboring mounds less. Adults increased footdrumming rates at high population densities, but juveniles footdrummed at high rates regardless of the population size.     

Males are attracted by their own courtship signals

Courting male fiddler crabs Uca terpsichores (1 cm carapace width) sometimes build mounds of sand called hoods at the entrances to their burrows. Males wave their single enlarged claws to attract females to their burrows for mating. It was shown previously that burrows with hoods are more attractive to females and that females visually orient to these structures. In this study, we test whether males also use their hoods to find their burrows. We first determined the maximum distance that males can see and find a burrow opening without a hood. Males were removed from their burrows and placed on the sand at a range of distances from a burrow opening. If they were more than about 8 cm (seven units of eye-height) away, they were unable find the burrow. In contrast, males that were burrow residents used a non-visual path map to return to their burrows from much greater distances. To determine if hoods help males find their burrows when there are errors in their path maps, we moved residents 1–49 cm on sliding platforms producing errors equal to the distances they were moved. Males with self-made hoods or hood models at their burrows relocated their burrows at significantly greater distances than did males with unadorned burrows. Hood builders also relocated their burrows faster. Hence, hoods have two functions: they attract females and they provide a visual cue that males use to find their burrows quickly and reliably when their path maps fail. An erratum to this article can be found at     

Feeding and mouthpart morphology in the semaphore crabHeloecius cordiformis (Decapoda: Brachyura: Ocypodidae)

The mouthparts of the Australian semaphore crabHeloecius cordiformis (H. Milne-Edwards, 1837) are described. Male and female semaphore crabs were observed and collected from mangrove swamps around the south side of Botany Bay, Australia between 1983 and 1987. Semaphore crabs inhabit mangrove mud/sand flats between the midlittoral intertidal zone and the supralittoral fringe of bordering saltflats. Scanning electron micrographs provide detailed pictures of mouthpart morphology and the setae present on the third, second, and first maxillipeds; the second and first maxillae; and the mandibles. Amongst other setal types, spoon-tipped, corn, and bottle-brush setae are described. While semaphore crabs are primarily deposit feeders (microphagous), they have well developed mandibles and are, therefore, also able to eat larger plant and animal material (macrophagous). A brief comparison is made between the mouthparts of semaphore crabs and fiddler crabs (Uca spp.), since the two genera are closely related, occupy similar niches, and resemble each other in appearance. This comparison reveals that while the second maxilliped setal morphology differs between the two genera (Heloecius possesses corn setae,Uca possesses spoontipped setae) these setae may have similar functions. The feeding behaviour ofH. cordiformis is related to substrate composition, burrow construction and various mouthpart specialisations. It is conlcuded that, within the context of the present study, certain structural e.g. mouthpart modifications allowing both scavenging and deposit feeding, and behavioural modifications e.g. mound building in wet muddy habitats, enable semaphore crabs to occupy a wide range of habitat types within the mangrove.Please address all correspondence and requests for reprints to Dr Maitland at his present address at the University of the Witwatersrand     

Sand disposal of the painted ghost crab Ocypode gaudichaudii (Decapoda: Ocypodidae): A possible role in courtship

In 1991/1992, we studied the sand disposal behavior of the painted ghost crabs Ocypode gaudichaudii on the Pacific coast of Panamá. O. gaudichaudii either kick, dump or tamp sand they excavate from their burrows. Here we relate these three kinds of sand disposal to burrow structure and distribution, as well as to crab size and sex. Our objective was to determine whether tamping may be a male courtship signal. Burrows whose owners tamped sand were on average longer, deeper, and higher on the beach than were burrows whose owners kicked or dumped sand. Five burrow shapes were distinguished, with half-spiral and spiral shapes being most common among tamped burrows. All crabs excavated from tamped burrows were males. Tamped burrows peaked in abundance around full and new moons. These observations, together with what is known about mating and breeding behavior of other ghost crabs, suggest that tamping may be involved in O. gaudichaudii courtship.     

Role of food subsidies and habitat structure in influencing benthic communities of shell mounds at sites of existing and former offshore oil platforms

Mussels (Mytilus californianus, M. galloprovincialis) and other organisms sloughed from offshore oil platforms provide a food subsidy to benthic consumers and alter underlying soft bottom habitat by creating hard substrate. The removal of overlying platforms eliminates this food subsidy, but large shell mounds remain. The distribution, abundance, and population characteristics of mobile macroinvertebrates differed among shell mounds beneath existing offshore oil platforms, shell mounds at the former sites of offshore oil platforms, and soft bottom. Predatory and omnivorous echinoderm and mollusk species were more abundant and generally larger on shell mounds under platforms than on shell mounds without platforms. Omnivorous and deposit feeding echinoderms were the most abundant macroinvertebrate taxa sampled on mound-only sites. The brown rock crab (Cancer antennarius), known to have a strong preference for hard substrate, was significantly more abundant on shell mounds, with or without platforms, than adjacent soft bottom sites. Results suggest that the effects of platform removal differed among benthic species according to trophic level, degree of mobility, and substrate preference. Although the shell mound habitat persists after removal of platform structures, species abundance and the composition of the associated benthic community is altered by removal of the platform structure.Communicated by P.W. Sammarco, Chauvin     

Burrow morphology of the ghost shrimp<Emphasis Type="Italic"> Nihonotrypaea petalura</Emphasis> (Decapoda: Thalassinidea: Callianassidae) from western Kyushu,Japan

The callianassid shrimp Nihonotrypaea petalura (Stimpson, 1860) is a common member on boulder beaches in Japanese waters. Its burrow morphology was investigated, based on 28 resin casts collected from a steeply sloping beach with dense boulders and 30 from a more gently sloping beach with less dense boulders in Ariake Sound, southern Japan. The structure and dimensions of the burrows from the two beaches were basically the same, and thus a combined mean value could be given to most of their constituent elements. In its entire dimensions, the burrow is greater in lateral extent than depth, with a mean maximum horizontal extension of 145 mm and a mean maximum depth of 119 mm for the shrimp with a mean total length of 31.8 mm. The burrow winds along boulders or cobbles and consists of, from top to bottom, a single surface opening with an ejected mound, a top shaft leading to the uppermost chamber at a mean depth of 48–56 mm, a passage with a regular cross section that is wider than that of the top shaft, and bulbous chambers (mean no.=4.7) with an irregular cross section associated with branches (mean no. per burrow=1.2). Bulbous chambers are much larger than the uppermost chamber and are usually connected by passages, with some directly attached to each other. The combined architecture of these features is unique and relatively simple among the burrows of all callianassid species, the majority of which inhabit bare soft sediments. The structure and function of the N. petalura burrow are discussed in relation to lifestyle.Communicated by T. Ikeda, Hakodate     

Hood construction as a spacing mechanism inCleistostoma kuwaitense (Crustacea: Ocypodidae)

The ocypodid crabCleistostoma kuwaitense inhabits the upper shores of the mudflats of Kuwait, where it constructs a semi-permanent mud hood over the entrance to its burrow. Between September and November 1986, data were collected on the distribution and orientation of the hoods and their openings to investigate the crab's social spacing system. Irrespective of density, the distribution of burrow hoods is non-random, tending towards a regular dispersion. A contributory factor for this dispersion pattern is that the surface hoods tend to increase the distance between adjacent burrows. The hood openings are randomly distributed with respect to compass direction but avoid directly facing those of nearest neighbours. In the absence of burrow plugging and eviction as used by other ocypodid crabs, construction of elongated hoods appears to be a good alternative for ensuring social spacing.     

Restoring coastal wetlands that were ditched for mosquito control: A preliminary assessment of hydro-leveling as a restoration technique

The wetlands surrounding Tampa Bay, Florida were extensively ditched for mosquito control in the 1950s. Spoil from ditch construction was placed adjacent to the wetlands ditches creating mound-like features (spoil-mounds). These mounds represent a loss of 14% of the wetland area in Tampa Bay. Spoil mounds interfere with tidal flow and are locations for non-native plants to colonize (e.g., Schinus terebinthifolius). Removal of the spoil mounds to eliminate exotic plants, restore native vegetation, and re-establish natural hydrology is a restoration priority for environmental managers. Hydro-leveling, a new technique, was tested in a mangrove forest restoration project in 2004. Hydro-leveling uses a high pressure stream of water to wash sediment from the spoil mound into the adjacent wetland and ditch. To assess the effectiveness of this technique, we conducted vegetation surveys in areas that were hydro-leveled and in non-hydro-leveled areas 3 years post-project. Adult Schinus were reduced but not eliminated from hydro-leveled mounds. Schinus seedlings however were absent from hydro-leveled sites. Colonization by native species was sparse. Mangrove seedlings were essentially absent (≈2 m⁻²) from the centers of hydro-leveled mounds and were in low density on their edges (17 m⁻²) in comparison to surrounding mangrove forests (105 m⁻²). Hydro-leveling resulted in mortality of mangroves adjacent to the mounds being leveled. This was probably caused by burial of pneumatophores during the hydro-leveling process. For hydro-leveling to be a useful and successful restoration technique several requirements must be met. Spoil mounds must be lowered to the level of the surrounding wetlands. Spoil must be distributed further into the adjacent wetland to prevent burial of nearby native vegetation. Finally, native species may need to be planted on hydro-leveled areas to speed up the re-vegetation process.     

Adaptive droving behavior observed in the fiddler crab Uca vocans vocans

Droving is conspicuous in Uca vocans vocans in summer. The crabs burrowing on the upper habitat have a higher tendency to wander compared to the ones burrowing on the lower habitat. Most of the wandering crabs captured on the low tide levels are relatively large and male. Larger crabs and males prefer to burrow on the upper zones of U. vocans habitat, but the smaller ones and females prefer the lower habitat. The upper level of the U. vocans habitat has relatively low N-content compared to the water's edge. Therefore droving is advantageous in crabs that have burrows on the upper level. Female and smaller resident crabs have faster feeding motions than male and larger ones, and can satisfy their feeding demands more rapidly. Therefore, for males and larger crabs it is advantageous to move away from the burrow area and forage in areas of higher food content near the low tide level where the number of feeding motions increases.     

Dispersal,population structure and burrow shape of Ocypode cursor

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

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.     

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.     

Chemical Stability of Capped Dredged Material Disposal Mounds in Long Island Sound, Usa

Geochemical analysis and visual inspection of cores collected from capped dredged material mounds revealed that in many cores, cap material was clearly distinguishable, both visually and chemically, from mound material. Contaminated dredged sediments were disposed in Long Island Sound eleven and seven years prior to sampling, and capped with uncontaminated dredged sediments. Core data provided no conclusive evidence of physical disturbance of, or chemical migration from, the contaminated mound sediments. Obvious chemical gradients of contaminant concentrations, which we propose are indicators of chemical migration, were not detected in the cores. Heterogeneity of the cap and mound sediments in some cores made the interface less distinct. We postulate that the preservation of spatial variability of sediment texture and associated contaminants observed within dredged material mounds is a result of the dredging process. Preservation of the textural and contaminant history of dredged sediments within finegrained cohesive cap materials provides evidence of the absence of physical or chemical disturbance.     

Chemical Stability of Capped Dredged Material Disposal Mounds in Long Island Sound,Usa

Geochemical analysis and visual inspection of cores collected from capped dredged material mounds revealed that in many cores, cap material was clearly distinguishable, both visually and chemically, from mound material. Contaminated dredged sediments were disposed in Long Island Sound eleven and seven years prior to sampling, and capped with uncontaminated dredged sediments. Core data provided no conclusive evidence of physical disturbance of, or chemical migration from, the contaminated mound sediments. Obvious chemical gradients of contaminant concentrations, which we propose are indicators of chemical migration, were not detected in the cores. Heterogeneity of the cap and mound sediments in some cores made the interface less distinct. We postulate that the preservation of spatial variability of sediment texture and associated contaminants observed within dredged material mounds is a result of the dredging process. Preservation of the textural and contaminant history of dredged sediments within finegrained cohesive cap materials provides evidence of the absence of physical or chemical disturbance.     

Fiddler crab burrowing affects growth and production of the white mangrove (<Emphasis Type="Italic">Laguncularia racemosa</Emphasis>) in a restored Florida coastal marsh

Positive plant–animal interactions are important in community ecology, but relatively little attention has been paid to their effect on the production of mangroves, dominant halophytic trees in tropical coastal marshes. Here, the role of fiddler crab (Uca spp.) burrowing on the growth and production of the white mangrove, Laguncularia racemosa (<2 years old), was examined in a restored marsh in Tampa Bay, Florida (27°41.65 N, 82°30.34 W) with manipulative experiments from June 2006 to May 2007. Fiddler crab burrowing significantly increased mangrove height by 27%, trunk diameter by 25%, and leaf production by 15%, compared to mangroves in crab exclusion enclosures. Additionally, the exclusion of fiddler crabs significantly increased interstitial water salinity from 32.4 to 44.2, and decreased the oxidation–reduction potential of the low organic sediments, but did not affect soil pH or sulfide concentration. Mangrove height, trunk diameter, and leaf production along a transect that varied in crab burrow density were positively associated with the number of crab burrows. Further, the density of sympatric Spartina alterniflora shoots was positively correlated with crab burrow density along the transect. As in temperate marshes, fiddler crabs can have significant ecological effects on mangrove communities, serving as ecological engineers by modulating the amount of resources available to marsh plants, and by altering the physical, chemical, and biological state of these soft sediment communities. In restored coastal systems that typically have very poor sediment quality, techniques such as soil amendment could be used to facilitate a more natural interaction between crabs and mangroves in ecosystem development.     

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     

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.     

How do weaponless male fiddler crabs avoid aggression?

Mimicry of females enables weaker males in many species to avoid intrasexual aggression. In fiddler crabs (Uca annulipes), males use their major claw in aggressive interactions to acquire and defend a territory. Males that have autotomised their major claw will be disadvantaged in fighting, but might use their temporary resemblance to females to avoid costly aggressive encounters with other males. We investigated whether: (1) courting males mistake clawless male fiddler crabs for females; (2) clawless males are able to acquire, defend and retain territories as successfully as intact males; and (3) clawless males are more cautious than intact males. Clawless and intact males differed in burrow acquisition methods and fighting behaviour, but were equally successful at acquiring and retaining burrows. While courting males treated clawless males as female, we found no evidence that clawless males mimic the behaviour of females, or that they exploit the advantage of their mistaken identity. Clawless males further appear to avoid male aggression by altering their territorial strategies to minimise the potential for conflict.     

Colonisation process of vegetative fragments of <Emphasis Type="Italic">Posidonia oceanica</Emphasis> (L.) Delile on rubble mounds

Seagrass colonise new areas via the dispersion of seeds or vegetative fragments. Independent of the manner of colonization, habitat requirements need to be met for the successful establishment of seagrasses. Here we report on the colonization process of Posidonia oceanica in a highly disturbed area: a gas pipeline trench at Capo Feto (SW Sicily, Italy). A trench dredged through a P. oceanica bed was back-filled with rubble added from dump barges leading to the formation of a series of rubble mounds on the seabed. Over time, these mounds became colonised with P. oceanica. I>. In order to understand the pattern of P. oceanica colonization, shoot density was quantified over 3 years (2001–2003) on different mound locations (crests, sides, valleys). Seagrass coalescence was observed only in valleys between mounds where shoot density averaged 133±50 shoots m⁻², while values for sides and crests were significantly lower (30.5±14 and 5.8±2.6 shoots m⁻², respectively). Although sediment accumulated on both crests and valleys, a significantly thicker sediment layer was recorded in the valleys (9.8±0.4 cm) than on crests (1.1±0.2). Plaster dissolution rate (an indicator of the hydrodynamic regime) tended to decrease from crests to valleys but even in the valleys, the currents were still higher than in the adjacent vegetated control location. This pattern was constant over time and depths. This is the first study to report on P. oceanica vegetative recruitment on artificial rubble after a disturbance event. It appears that the valleys between the rubble mounds are suitable for seagrass recruitment as sediment deposited between the rubble provides the necessary resources for plant settlement and growth. Once the seagrass patches are established, they may start a positive feedback of attenuation of currents, sediment accumulation and seagrass patch expansion.     

Perceived predation risk and mate defense jointly alter the outcome of territorial fights

Virtually all animal conflicts occur over access to mates or resources that affect survival, the two key components of fitness. In this paper, we report that predation risk and mate defense jointly affect the outcomes of contests between male sand crabs (Scopimera globosa) for burrows in which crabs mate and take shelter from predators. We observed the contests under three different conditions: (1) the natural condition of low predation risk and without the presence of a female; (2) the first experiment in which we imposed upon only intruding males the perception of predation risk—by digging them from their burrows, capturing and handling them, and placing them into other males’ burrows—to increase the value of the burrows for the intruders as shelter, and (3) the second experiment in which we repeated this treatment but increased the resource value of the burrow to the resident by placing a female in his burrow. The difference in body size between contestants was the main determinant of victory in all analyses. However, perceived predation risk also partly affected the outcomes of the fights: The motivated intruders were likely to win even when they were a little smaller than the residents. In addition, defense of a female had a significant effect on the outcomes of fights: The motivated residents won more fights than the motivated intruders, indicating that these two treatments caused asymmetric increases of the resource value. This is the first report of two external factors simultaneously raising resource value, affecting motivation of contestants, and altering the outcome of fights.