Footdrumming and other anti-predator responses in the bannertail kangaroo rat (Dipodomys spectabilis)

Summary Anti-predator behaviors of the bannertail kangaroo rat (Dipodomys spectabilis) toward snake predators were investigated. We induced responses by presenting a live (tethered) snake and an inflatable snake decoy to rats of known ages and sex in their territories. Comparisons of behaviors during spontaneous activity (baseline), naturally occurring snake-rat interactions, and experimental procedures further clarified anti-predator behaviors. The tethered snake immediately induced high rates of footdrumming (the repeated striking of the hind feet on the ground) in all rats (n=24) (Table 2), and individuals of both sexes and all ages footdrummed significantly more at the tethered snake than at the decoy. Other antipredator behaviors included alert postures, jumpback, kicking sand and avoidance. Juvenile rats exhibited more avoidance behavior and alert postures than adults and footdrummed at the decoy only if they experienced the snake first. We conclude that footdrumming functions as an individual alarm signal against predation by snakes in D. spectabilis. The drumming probably informs a snake that it has been detected and may cause it to leave. Because rats did not footdrum in response to aerial predators (owls), we suggest that kangaroo rats use different anti-predator strategies that depend on the kind of threat and the type of predator.     

Territorial defense in the bannertail kangaroo rat (Dipodomys spectabilis): footdrumming and visual threats

Summary Footdrumming and visual displays associated with territorial defense in the bannertail kangaroo rat (Dipodomys spectabilis) were studied in the field in Arizona. We predicted that foot-drumming functioned like bird song in territorial advertisement. Rats were tested in their territories for their responses to the presentation of four different stimuli: a playback of airborne footdrumming patterns (audio); a stuffed and mounted kangaroo rat in an aggressive posture (visual); a combination of both (audio-visual); and a playback of cricket stridulation as a control. Kangaroo rats investigated all experimental stimuli equally by becoming alert, approaching, and retreating (Fig. 2). Rats, therefore, seemed to assess the amount of threat and footdrummed at higher rates and sooner in response to the more threatening visual stumulus of a stuffed rat than to the apparently less threatening ones of a airborne footdrumming playback or cricket stridulation (Fig. 1). We conclude that rats footdrum at long ranges to repel a potential intruder and also at close ranges when territories are threatened by a persistent intruder. Chase and attack, as in territorial birds, are the final lines of defense.     

Use of different signaling modalities to communicate status by dominant and subordinate Heermann’s kangaroo rats (<Emphasis Type="Italic">Dipodomys heermanni</Emphasis>)

Dominance relationships in solitary species may be an important factor in the maintenance of long-term, stable relationships among territorial neighbors. We examined the mediation of intraspecific interactions in a solitary, territorial kangaroo rat, Dipodomys heermanni, and tested whether unfamiliar kangaroo rats establish a dominance hierarchy and then decrease aggression and increase communication (via footdrumming and sandbathing) after initial interactions and the establishment of a social structure. Results revealed that both dominance hierarchies and familiarization with particular individuals are likely to mediate social interactions. After only one pairing per dyad, an almost linear dominance hierarchy emerged, which became perfectly linear after a 90-min familiarization period. During the course of subsequent interactions between dyad partners, fighting decreased and non-agonistic communication increased. Dominant kangaroo rats sandbathed at higher rates than subordinates, possibly to deposit scent to advertise competitive ability, whereas subordinate kangaroo rats footdrummed from inside the burrow, which seemed to indicate an unwillingness to interact. We suggest the kangaroo rats use a conditional strategy when deciding to fight (be dominant) or withdraw (be subordinate) by employing different modes to communicate status and minimize the potential risk of injury during unnecessarily prolonged fights.     

Fitness costs of neighborhood disruption in translocations of a solitary mammal

Translocation is used to reestablish wild populations of animals, but translocation projects often do not meet their objectives because postrelease mortality of animals is high. One reason for translocation failure is that the behavioral or ecological requirements of released animals are unmet. Maintaining founder-group social relationships during release can affect reestablishment of social species. Solitary territorial species with stable neighbors (restricted dispersal and lifetime occupation of a home range) of the same species may also benefit from the maintenance of these social relationships during translocation. We translocated Stephens' kangaroo rats (Dipodomys stephensi), a solitary species listed as endangered under the U.S. Endangered Species Act, with and without neighboring kangaroo rats. We compared the settlement (establishment of a stable home range) decisions and fitness of kangaroo rats between the 2 treatments. Kangaroo rats translocated with neighbors traveled shorter distances before establishing territories, had higher survival rates, and had significantly higher reproductive success than kangaroo rats translocated without neighbors. Number of offspring was 24-fold higher for kangaroo rats translocated with neighbors than those translocated without neighbors. Differences in behavior following release may partially explain differences in survival between the 2 groups. Immediately following release, animals translocated with neighbors fought less and spent significantly more time foraging and digging burrows than animals translocated without neighbors. Our results indicate that even for solitary species, maintaining relationships among members of a translocated group of animals can influence translocation success. This study is the first empirical demonstration of the fitness consequences of disrupting social relationships among territorial neighbors.     

Inter-specific variation in anti-predator behavior in sympatric species of kangaroo rat

Kangaroo rats, Dipodomys, occupy desert habitats with little cover and thus are under high predation risk from diverse predators. The behavior used to assess predation risk or to escape capture is unknown. We therefore compared anti-predator behavior of two sympatric species of kangaroo rat of different sizes, D. merriami and D. spectabilis. We first examined whether kangaroo rats use olfaction as a first line of defense against snake predation and tested the rats for their responses to scent extracted from two species of snake that live sympatrically with the kangaroo rats, the Mojave rattlesnake (Crotalus scutulatus) and the gopher snake (Pitophis melanoleucus). We also tested for species differences in anti-predator behavior through 15-min interactions between the kangaroo rats and free-moving gopher snakes. We found that D. spectabilis actively approached the scent of both rattlesnakes and gopher snakes more than controls of vegetable oil and evaporated solvent (Fig. 1). In contrast, D. merriami did not differentiate snake odors from controls in the experimental arena, but they sniffed the sand where a free-moving snake had passed more than D. spectabilis. Both species successfully avoided predation in encounters with live snakes. Although total numbers of approaches and withdrawals were similar (Fig. 2), D. spectabilis spent significantly more time within striking distance of the snake than D. merriami. D. spectabilis approached the head of the snake in 93% of its approaches and often engaged in nose to snout contact with the snake. If the snake struck, D. spectabilis jumped directly backward to avoid a strike and footdrummed at a safe distance. In contrast, D. merriami oriented to the snake more than D. spectabilis, but approached the head in only 41% of the approaches and rarely engaged in nose-to-snout contact. The snakes struck, hissed and decreased predatory approaches with D. spectabilis but not with D. merriami (Fig. 3). These results show that kangaroo rats can behaviorally influence the risk of being preyed on by snakes. The two species differ, however, in how they react to snakes. The larger D. spectabilis confronts snakes while the smaller D. merriami monitors snakes from a safe distance and avoids them.     

Antipredator behavior of a social desert rodent: footdrumming and alarm calling in the great gerbil, Rhombomys opiums

We sought to understand why a social, desert rodent, the great gerbil, Rhombomys opimus, expends energy and possible risk of predation by footdrumming and vocalizing in the presence of a diversity of terrestrial predators: snakes, monitor lizards, polecats, foxes, and humans. Behavioral observations, human approaches, and experiments with tethered predators revealed that both male and female gerbils called and footdrummed in the presence of offspring, close relatives, and potential mates. Because adults called more often when pups were present, and solitary gerbils seldom gave an alarm, the alarm behavior probably warns conspecifics, especially vulnerable offspring, of potential danger. We also found that gerbils altered alarm behavior with the type of predator. They drummed more in the burrow when a dog that could not enter the burrow was present, and they drummed more out of the burrow in response to a snake that could enter the burrow. Gerbils vocalized and stood in an alert posture in response to all stimuli. The different footdrumming responses of gerbils to terrestrial predators seems related to the hunting style and type of risk posed by the predator, especially its ability to enter the burrow system. Received: 23 August 1999 / Received in revised form: 6 December 1999 / Accepted: 25 February 2000     

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.     

An Age-Structured Demographic Model for the Endangered Stephens' Kangaroo Rat

Effective conservation of endangered species often is hampered by inadequate knowledge of demography. We extracted information on survival and fecundity from an 18-month, live-trapping study of Dipodomys stephensi , and from this we developed an age-structured demographic model to assess population viability. Adult Stephens' kangaroo rats persisted longer than juveniles, and adult females persisted longer than adult males. Disappearance rates were high in the first months after initial capture. Thereafter, the fraction of animals persisting decreased slowly and in an approximately linear fashion on a semilogarithmic scale, suggesting age-independent mortality factors such as predation. Juvenile persistence did not differ substantially between two years of strikingly different rainfall. Onset of breeding followed the start of winter rains. Length of the breeding season, average number of litters per female, and the fraction of first-year females breeding were much greater in the year of higher rainfall. We propose a birth-pulse demographic model for D. stephensi that distinguishes juvenile and adult age classes. Temporal environmental variation can be modeled adequately with a constant survivorship schedule and variable fecundity determined by yearly precipitation. Several issues should be resolved, however, before conservation decisions are based on the model. Better estimates of juvenile survivorship are critical, the quantitative relationship between precipitation and fecundity must be determined, and the potential for density dependence and source-sink population dynamics must be evaluated.     

Evenly dispersed social groups and intergroup competition for juveniles in a coral-reef fish

Summary A model based on current knowledge of recruitment of settling juveniles of coral-reef fishes suggests that adult social groups far removed from their nearest neighbors can expect to attract larger numbers of juveniles than groups that are close to nearest neighbors. For protogynous fishes in which females change to males when intragroup, female-to-male sex ratios rise above threshold values, selection should favor individuals occupying groups that attract many juveniles. The model predicts that social groups should, in consequence, be regularly dispersed and that, in effect, groups compete with each other to attract juveniles. Dispersion of social groups was measured in two populations of the protogynous fish Anthias squamipinnis. The location of all groups were plotted on maps of the reef. A set of nearest-neighbor analyses applied to the data all showed significant departures of observed nearest-neighbor distances from randomness, in the direction of regularity. At both sites, neighboring groups were separated by much unoccupied space. In one population, the substrate consisted of discrete coral aggregates scattered over open sand bottom, while in the second population groups occupied a relatively continuous vertical reef face. Although regular dispersion could, in theory, result from regular distribution of food patches, current evidence does not favor this explanation. It is more likely that regular dispersion results either from the reproductive advantage of occupying groups that are highly attractive to juveniles, as seen in the model, or from limited perceptual and locomotor abilities of juveniles at the time of settlement.     

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     

Chihuahuan Desert kangaroo rats: nonlinear effects of population dynamics, competition, and rainfall

Using long-term data on two kangaroo rats in the Chihuahuan Desert of North America, we fitted logistic models including the exogenous effects of seasonal rainfall patterns. Our aim was to test the effects of intraspecific interactions and seasonal rainfall in explaining and predicting the numerical fluctuations of these two kangaroo rats. We found that logistic models fit both data sets quite well; Dipodomys merriami showed lower maximum per capita growth rates than Dipodomys ordii, and in both cases logistic models were nonlinear. Summer rainfall appears to be the most important exogenous effect for both rodent populations; models including this variable were able to predict independent data better than models including winter rainfall. D. merriami was also negatively affected by another kangaroo rat (Dipodomys spectabilis), consistent with previous experimental evidence. We hypothesized that summer rainfall influences the carrying capacity of the environment by affecting seed availability and the intensity of intraspecific competition.     

Ontogeny of space use and diet of two temperate damselfish species, <Emphasis Type="Italic">Parma microlepis</Emphasis> and <Emphasis Type="Italic">Parma unifasciata</Emphasis>

Studies of reef fish herbivory have mainly focused on the impacts and behaviour of adults of tropical species. In this study, the ontogenetic shifts in home range, aggression, feeding rate, diet and gut morphology in juveniles and adults of two temperate territorial damselfishes, Parma microlepis and Parma unifasciata, were determined. Both P. microlepis and P. unifasciata juveniles under 80 mm TL exhibited no aggressive chases towards conspecifics or other species, while above 80 mm TL aggressive chase frequency increased in conjunction with an increase in home range, defended as a territory. Ontogenetic diet shifts, characterised by an increase in herbivory (P. unifasciata: juveniles: 64% plant material, adults: 95% plant material; P. microlepis: juveniles: 43% plant material, adults: 67% plant material) were observed for both species. The ratio of digestive tract length to body length, which often accompanies a switch to herbivory, increased significantly with ontogeny for both species. Compared to tropical confamilial grazers, these temperate damselfish species feeding rates were lower, and they had larger territories which were not as strongly defended (fewer aggressive chases).     

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.     

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.     

Operational sex ratio and alternative reproductive behaviours in the European bitterling,<Emphasis Type="Italic"> Rhodeus sericeus</Emphasis>

We investigated the effects of male population density and male-biased operational sex ratio (OSR) with constant and limited resource density on male mating tactics shown by a freshwater fish, the European bitterling, Rhodeus sericeus. This species spawns inside living unionid mussels. Large males defended territories and were aggressive towards conspecifics under equal sex ratios. They also monopolised pair spawnings with females, releasing 98% of all sperm clouds during mating. However, the mating tactic changed at high male density where large males ceased to be territorial and instead competed with groups of smaller males to release sperm when females spawned. Large, medium and small males now obtained 61%, 33%, and 6% of sperm releases respectively, thereby reducing the opportunity for sexual selection by half. Females spawned at equal rates in the two densities of males, despite lower courtship at high density. These results run counter to the usual expectation that an increasingly male-biased OSR should lead to higher variance in male mating success. Instead, the use of alternative reproductive behaviours by males can lead to lower resource competition and mating variance at high male densities.     

Growth and population structure in the infaunal bivalve Yoldia eightsi in relation to iceberg activity at Signy Island,Antarctica

Growth rates in the infaunal nuculanid bivalve mollusc Yoldia eightsi (Couthouy) were assessed for field populations in Factory Cove, Signy Island, Antarctica, between. February and April 1992. Daily increments in length (maximum shell dimension) ranged from 2.3 m d^-1 for a 30 mm individual to 5.1 m d^-1 for a 10 mm specimen. These growth rates were converted to annual increments, based on a growing season for the Signy population of around 5 mo, and ages for the largest individuals in the population (35 mm in length), were calculated to be 65 yr. Specimens of 43 mm in length have been found near this site and, if their growth rates are similar to this population, their ages would be in the order of 120 yr. Size distributions from two sites in the same Y. eightsi bed 300 m apart showed significant differences. At the more exposed site the distribution was dominated by small juveniles, with 86% of the population 10 mm in length, while 13% were >20 mm in length. At the less exposed site the values were 27% <10 mm and 56% >20 mm. Icebergs have often been seen grounded on the Y. eightsi bed in this study, especially on the exposed outer portion. This factor, in association with inhibition of larval settlement by high densities of large individuals, is proposed as an explanation of the observed population distributions and the absence of very large specimens in the Factory Cove population.     

Twenty years of changes in spatial association and community structure among desert perennials

I present results from analyses of 20 years of spatiotemporal dynamics in a desert perennial community. Plants were identified and mapped in a 1-ha permanent plot in Joshua Tree National Park (California, USA) in 1984. Plant size, mortality, and new seedlings were censused every five years through 2004. Two species, Ambrosia dumosa and Tetracoccus hallii, were dominant based on their relative abundance and ubiquitous distributions. Spatial analysis for distance indices (SADIE) identified regions of significantly high (patches) or low (gaps) densities. I used SADIE to test for (1) transience in the distribution of patches and gaps within species over time and (2) changes in juvenile-adult associations with conspecific adults and adults of the two dominant species over time. Plant performance was quantified in patches and gaps to determine plant responsiveness to local spatial associations. Species identity was found to influence associations between juveniles and adults. Juveniles of all species showed significant positive spatial associations with the dominant A. dumosa but not with T. hallii. The broad distribution of A. dumosa may increase the spatial extent of non-dominant species that are facilitated by this dominant. The spatial location of patches and gaps was generally consistent over time for adults but not juveniles. Observed variability in the locations of juvenile patches and gaps suggested that suitable locations for establishment were broad relative to occupied regions of the habitat, and that conditions for seed germination were independent of conditions for seedling survival. A dramatic change in spatial distributions and associations within and between species occurred after a major drought that influenced data from the final census. Positive associations between juveniles and adults of all species were found independent of previous associations and most species distributions contracted to areas that were previously characterized by low density. By linking performance to spatial distribution, results from this study offer a spatial context for plant-plant interactions within and among species. Community composition could be influenced both by individual species tolerances of abiotic conditions and by the competitive or facilitative interactions individuals exert over neighbors.     

Density-dependent settlement and mortality structure the earliest life phases of a coral population

The local densities of heterospecifics and conspecifics are known to have profound effects on the dynamics of many benthic species, including rates of settlement and early post-settlement survivorship. We described the early life history of the Caribbean coral, Siderastrea radians by tracking the population dynamics from recently settled planulae to juveniles. Through three years of observation, settlement correlated with the abundance of other benthic organisms, principally turf algae (negatively) and crustose coralline algae (positively). In addition, adult density showed independent effects on coral settlement and early post-settlement survivorship. Settlement rates increased across low levels of adult cover and saturated at a maximum around 10% cover. Early post-settlement survivorship decreased with adult cover, revealing structuring density dependence in coral settlers. The earliest life stages of corals are defined by low survivorship, with survivorship increasing appreciably with colony size. However, recent settlers (one-polyp individuals, < 1-year-old) are more likely to grow into two-polyp juveniles than older single polyps (> 1-year-old) that were delayed in their development. The early benthic phase of corals is defined by a severe demographic bottleneck for S. radians, with appreciable density-dependent and density-independent effects on survivorship. For effective management and restoration of globally imperiled coral reefs, we must focus more attention on this little studied, but dynamic, early life history period of corals.     

Behavioral responses of territorial red squirrels to natural and experimental variation in population density

The relative scarcity of studies at the intersection of behavioral and population ecology is surprising given the presumed importance of behavior in density-dependent population regulation. Here we tested whether North American red squirrels (Tamiasciurus hudsonicus) adjust their behavior in response to local population density and whether they use rates of territorial vocalizations in their local neighborhood to assess density. We examined these relationships using 18?years of live trapping and 20?years of behavioral data that were collected across natural variation in local population density. To disentangle the effects of population density on behavior from those due to changes in per capita food abundance or changes in the frequency of antagonistic interactions with neighbors, we also experimentally manipulated population density with long-term food supplementation as well as perceived population density with long-term playbacks of territorial vocalizations. The frequency with which squirrels emitted territorial vocalizations was positively associated with local population density. In contrast, antagonistic physical interactions observed between squirrels and territorial intrusions were rare and the frequency of intrusions was weakly and negatively, not positively, associated with population density. Squirrels experiencing naturally and experimentally high density conditions spent less time in the nest and feeding but more time being vigilant. Similar density-dependent changes in behavior were observed in response to our manipulations of perceived population density, indicating that vocalization rates and not physical interactions or food abundance were the mechanism by which squirrels assessed and responded behaviorally to changes in local density.