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     

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.     

Neighbor burrow-plugging in Ilyoplax pusillus (Crustacea: Brachyura: Ocypodidae)

The burrow living ocypodid crab Ilyoplax pusillus (De Haan, 1835) sometimes plugs the burrow of neighbors situated from 1.0 to 8.3 cm away with surface mud, while the neighbor, termed the pluggee, is in the burrow. Most pluggers were large males, whereas pluggees were smaller than pluggers and had a sex ratio close to 1:1. After being plugged, most pluggees usually reemerged on the surface within 5 min, but occasionally took up to 77 min to do so. The plugger always foraged or performed waving displays around the burrow of the pluggee while the pluggee stayed inside the burrow. Although the activity site of the pluggee was originally directed toward the burrow of the plugger or the adjacent area, after reemergence its activity site was oriented toward another direction. From these observations, it is suggested that neighbor burrow-plugging is a behavior adopted mainly by large males as a means of maintaining the area of their surface activities against smaller neighbors. My observations were made from June 1984 to June 1985 at Fukuro River Estuary, central Japan.     

Morphology and orientation of tube extensions on aggregations of the polychaete annelid Phragmatopoma californica

The morphology of organisms can influence fluid flow near their surfaces which, in turn, can influence the rates of mass exchange as well as the ability of the animals to capture and retain particles. In this paper, the morphologies and orientations of extensions on the tubes of aggregated individuals of the sabellariid polychaete Phragmatopoma californica from different habitats are described. The effects of these structures on fluid exchange and feeding-tentacle deflection are also assessed. The tube shape of aggregated P. californica varies among habitats. In exposed habitats, extensions on tubes (flares) encircle the entire circumference of the worms' apertures. In more protected habitats, extensions (hoods) partially encircle the circumference of apertures and are oriented so that the opening of the hood faces into the predominant direction of water flow over the aggregation. After the surface structures are damaged, hoods are repaired more quickly than flares. In this study, all damaged hodds were repaired within 24 h while flares were fully repaired only after 240 h. The repaired hoods are oriented in the same direction as the original structures Both hoods and flares decrease the rate of fluid exchange between the aggregation surface and the mainstream water flow. These structures also decrease the deflection of the feeding tentacles by the water flow. The orientation of the hoods relative to the direction of water flow affects both the rates of fluid exchange and the deflection of the tentacles. Present address: Hawaii Institute of Marine Biology, P.O. Box 1346, Kaneohe, Hawaii 96744, USA     

The burrows of alpheid shrimp associated with gobiid fish in the northern Red Sea

The burrows of 4 sympatric associations of alpheid shrimp (Alpheus djiboutensis 1 and 2, and Alpheus sp. A and B) and non-burrowing gobiid fish were studied in the lagoon of a narrow fringing reef, in the northern Red Sea. Epoxy resin casts were used to determine the subterranean structure of burrows. The burrows were irregular, and in close contact with hard objects within the substrate. The number and structure of openings and their daily displacement were type-specific. On the other hand, no type specificity was observed in the deeper parts of the burrow. The mechanism of the formation of new openings by the combined activities of the fish Cryptocentrus sungami and the shrimp A. djiboutensis 2 is described. A positive correlation in the size of the partners seems to be due to competition of fish for larger burrows.     

Interspecific attractiveness of structures built by courting male fiddler crabs: experimental evidence of a sensory trap

Male fiddler crabs Uca musica sometimes build sand hoods and male Uca beebei sometimes build mud pillars next to their burrows to which they attract females for mating. Mate-searching females preferentially approach these structures and subsequently mate with structure builders. Here we show that the preference for structures is not species-specific and argue that it may not have evolved for mate choice. When not near burrows, many species of fiddler crabs approach and temporarily hide near objects, suggesting that hoods and pillars may attract females because they elicit this general predator-avoidance behavior. To test this sensory trap hypothesis we individually released female U. musica, U. beebei and Uca stenodactylus, a non-builder, in the center of a circular array of empty burrows to which we added hoods and pillars and then moved a model predator toward the females. All species ran to structures to escape the predator and the two builders preferred hoods. Next, we put hood replicas on male U. beebei burrows and pillar replicas on male U. musica burrows. When courted, females of both species preferentially approached hoods as they did when chased with a predator. However, males of both species with hoods did not have higher mating rates than males with pillars perhaps because hoods block more of a male's visual field so he sees and courts fewer females. Sexual selection may often favor male signals that attract females because they facilitate general orientation or navigation mechanisms that reduce predation risk in many contexts, including during mate search.     

Behavioural ecology of the mud-burrowing crab Goneplax rhomboides

Diving studies at Fishguard harbour in South Wales (UK) combined with laboratory studies, provided information on the burrowing behaviour of Goneplax rhomboides (L.). Polyester resin casts indicated that burrows had from 1 to 6 openings and tunnels were horizontal at about 10 cm depth, with branches usually at right angles to each other. Burrowing was studied in the laboratory and related to variations in burrow casts taken in the field. Preliminary laboratory experiments suggest that social interactions occur between crabs, but analysis of the spatial distribution of burrows in the field suggested randomness. Burrows in the field showed some similarity of orientation, although the reasons for this were uncertain.     

Functional burrow morphology of Biffarius arenosus (Decapoda: Callianassidae) from southern Australia

Burrows of the thalassinidean shrimp Biffarius arenosus (Poore, 1975) were investigated by in situ resin-casting on an intertidal sandflat in Western Port, southern Australia. Even though burrow casts exhibited interspecific variation, all had at least two openings, a U-shaped top and a series of tunnels and chambers. The space occupied by the burrows was deeper than it was wide; a maximal depth of 58 cm was recorded. Inhalant and exhalant shafts were restricted relative to tunnel diameter, and were often arranged in a multi-layered U-shape. Additional shafts branching from deeper sections of the burrows were occasionally observed. The main section of a burrow, extending from the base of the U, usually consisted of a downward spiral or an irregular spiral combined with some straight tunnel sections, all with a circular cross-section. The central spiral branched into further tunnels and chambers. Nearly all casts possessed peripheral chambers positioned either at the edges of horizontal lattices or in the deepest sections of the burrow. No evidence of plant material was found in peripheral chambers. Major burrow features were consistent with B. arenosus adopting a deposit-feeding trophic mode and collecting its food below the sediment surface. Received: 17 July 1998 / Accepted: 26 November 1998     

Ecological and social determinants of spacing behaviour in desert baboon groups

This paper investigates the determinants of individual spacing behaviour in a desert baboon population (Papio cynocephalus ursinus). Patterns of neighbour proximity and neighbour density were examined among adults in four groups under different ecological and social conditions (through instantaneous sampling during focal follows). Initial analysis of these data shows that (1) the use of vertical substrates (refuges such as tall trees and cliff faces) can confound patterns of spacing, and (2) individual differences in spacing can depend on the spatial scale over which it is measured. To minimise these substrate and scale effects, this analysis focuses on animals which are off refuges and examines spacing behaviour through its underlying statistical `dimensions' (identified through factor analysis). Analysis of these dimensions indicates that sex, group size, activity-habitat and female reproductive state can all have independent effects on spacing: (1) males are more dispersed than females in smaller groups, (2) male and female dispersion increases with time spent in foraging habitats, and (3) female dispersion is reduced during lactation. According to the hypotheses tested, these results indicate that feeding competition only affects spacing behaviour during foraging while predation risk plays little or no role in spacing. Most aspects of spacing behaviour are best explained by male reproductive strategies and their social repercussions. Received: 25 May 1998 / Accepted after revision: 18 July 1998     

Observations on the burrows and burrowing behaviour of two mud-dwelling decapod crustaceans,Nephrops norvegicus and Goneplax rhomboides

The burrows of the Norway lobster Nephrops norvegicus (L.) and of the crab Goneplax rhomboides (L.) were studied in Loch Torridon, Scotland. Polyester resin casts of burrows in the sea were made by divers to reveal their subsurface form. Tunnels made by N. norvegicus were usually simple, with two or more openings on the mud surface, and penetrated to a depth of about 30 cm. G. rhomboides burrows did not descend more than about 15 cm beneath the surface, but were usually more complex than the lobster burrows and had several openings. The methods of burrow construction used by the two crustaceans are described from aquarium observations. Neither N. norvegicus nor G. rhomboides show obvious morphological adaptations for burrowing, and it is suggested that the fossorial habit was adopted very early by decapods. The burrows of N. norvegicus do not seem to have assumed any functions in addition to the original one of providing refuge from predators. There is not sufficient known of the biology of the crab to indicate whether the same is true in its case.     

How a subsocial intertidal beetle,Bledius spectabilis,prevents flooding and anoxia in its burrow

Summary All stages of Bledius spectabilis Kratz (Staphylinidae) dig wine-bottle shaped burrows in the intertidal Salicornia zone. The adult female lays her eggs around her burrow and, by remaining with them, prevents both flooding by the tide and anoxia. Flooding is prevented by an ever-ready burrow which exploits a surface tension effect and by blocking the burrow once the tide comes in. The burrow has a narrow neck (about 2 mm in diameter) leading to a living chamber 5 mm in diameter. Tides over artificial tubes in agar showed that a critical minimum neck diameter of 2–3 mm prevented sudden flooding, giving time to block the neck with mud. Blocking took about four minutes. Ensuring the burrow is reopened at each low tide is a vital role of the brooding female in anaerobic or impermeable soils. Field models of burrows became anoxic in 4 days, much less than the 4 week long egg stage. A lab model system with anaerobic agar and calculations of oxygen uptake and diffusion supported this conclusion. Mortality of orphaned larvae may be much lower, however, in burrows within large Bledius aggregations because of mitigating good drainage and soil aeration: larval mortality from physical causes in these burrows did not increase over 3 weeks but without the mother, 14% of the eggs were attacked by mould, and two burrows were taken over by a predatory carabid, Dichierotrichus gustavi, and all larvae eaten. The surface tension effects of small air-filled openings may be used by small air-breathing animals in many intertidal habitats. Like Bledius, other intertidal animals, including large ones, may block their burrows at high tide to keep them full of air. Maternal care, in particular the combination of behaviours which protect the brood from the tide and anoxia, enables this airbreathing insect to colonise the inhospitable habitat of the intertidal saltmarsh.     

Meiofaunal thiobios and “the Arenicola negation”: Case not proven

It has been alleged recently that the distribution of meiofauna around Arenicola burrows disproves the existence of a specifically anaerobic marine meiofauna. Critical appraisal of the evidence on which this denial was based shows that the conclusion was untenable. It may have arisen mistakenly from inadequate sampling of the burrow micro habitats, from lack of complementary data on burrow wall oxygen regime and from prejudgement (by denying the possibility of primary anaerobiosis) of the reasons why species held to be anaerobic by other authors occur in proximity to oxic conditions. These points and some new data on Arenicola burrow oxygenation are discussed. It is shown that at present the evidence from meiofaunal distribution around Arenicola burrows may be taken as much in favour of thiobiotic theories as against them.     

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.     

Reproduction, larval development and functional relationships of the burrowing, spionid polychaete Dipolydora armata with the calcareous hydrozoan Millepora complanata

Dipolydora armata (Langerhans, 1880) is a small (4 to 5 mm) spionid polychaete found burrowing in the calcareous hydrozoan Millepora complanata Lamarck, 1816, on coral reefs at Barbados, West Indies. It excavates complex networks of interconnecting burrows and forms aggregations of worms in cavities within branches of the coral. Adult worms have a mixed feeding mode (suspension feeding and deposit feeding). Size–frequency distributions of worms in branch samples suggest that they mature in a single year and that reproduction occurs throughout the year. Burrow openings on the surface of the coral develop distinctive, erect spines caused by combined growth of worm tubes and host tissue. Millepore zooids were absent in the vicinity of tube openings and on spines, and thus the potential feeding surface of the coral will be reduced in heavily colonized branches. Burrows and openings were densest at the bases of millepore branches where weakening of the skeleton would be expected to occur. The absence of openings near the branch tips suggests difficulty in larval settlement there, amongst stinging zooids. Reproduction␣and larval development of the worms were examined, and a sequence of larval stages from one to 20 segments and a juvenile stage of 22 segments are described. Eggs are deposited in brood sacs attached to the burrow wall, and the larvae feed upon nurse eggs (adelphophagy). The presence of larvae and juveniles occurring free in the burrows suggests that larval development may be completed within the host coral as an alternative or in addition to a planktonic larval phase. Lack of provisional larval setae, early development of adult capillary setae, production of special spermatophores and a protracted breeding cycle in D. armata are all traits which would favour complete development within the host skeleton. Received: 6 March 1997 / Accepted: 25 October 1997     

Microbial activity and accumulation of organic matter in the burrow of the mud shrimp, <Emphasis Type="Italic">Upogebia major</Emphasis> (Crustacea: Thalassinidea)

Microbial activity and accumulation of organic matter in the burrow of the thalassinidean mud shrimp, Upogebia major, were studied on a tidal flat along the northern coast of Tokyo Bay, Japan. The burrow of U. major is Y-shaped with an upper U-shaped part plus a lower I-shaped part. Its lower part can extend to a depth of 2 m. In the present study, we compare electron transport system activity (ETSA), bacterial abundance and organic matter content [total organic carbon (TOC), total nitrogen (TN) and chlorophyll a (chl. a)] of the burrow wall sediment with the tidal flat surface and non-burrow sediments. We also compared the U- and I-shaped part in terms of these parameters. ETSA in the burrow wall was higher than at the tidal flat surface in the warmer season, and was always higher than at surrounding non-burrow sediments. Bacterial abundance in the burrow wall was higher than at the tidal flat surface and surrounding sediment. TOC and TN contents in the burrow wall were two to three times higher than those at the tidal flat surface and non-burrow sediments, regardless of season. However, there was no significant difference in chl. a content between burrow wall and tidal flat surface. These results suggest that organic enrichment of the burrow wall is a result of organic matter particles such as phytodebris accumulation along the burrow wall. For all parameters of the burrow walls, no statistical differences were found between the two parts. Present results indicate that U. major actively transports the water containing suspended organic particles not only through the U-part but also into the deeper I-part. These indicate that burrow of the mud shrimp provides a dynamic environment for microbial community in tidal flat sediment.     

The influence of group size on natal dispersal in the communally rearing and semifossorial rodent, Octodon degus

In social or group living species, members of groups are expected to be affected differentially by competition through the effect of group size (i.e., the “social competition hypothesis”). This hypothesis predicts an increase in the probability of dispersal with increasing size of social groups. At a more mechanistic level and based on the known effects of competition on stress hormone levels, a positive relationship between group size and glucocorticoids of juveniles should be observed. We used a demographic approach to test these predictions on a natural population of the communally rearing and semifossorial rodent—Octodon degus. Burrow systems provide degus with places to rear offspring and to evade stressful thermal conditions and predators. Thus, we predicted dispersal to increase with increasing number of degus per main burrow system used, a measure of habitat saturation in degus. The probability of dispersal increased with increasing number of degus per main burrow system used. Mean fecal metabolites of cortisol in offspring increased, yet not statistically significantly, with the number of juveniles in groups. These results were consistent with a scenario in which competition drives natal dispersal in juveniles in social degus. In particular, competition would be the consequence of high degu abundance in relation to the abundance of burrow systems available at the time of offspring emergence.     

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     

Dispersion of Tellina tenuis from Kames Bay,Millport, Scotland

The dispersion of Tellina tenuis da Costa in the laboratory was analysed by the Clark and Evans nearest-neighbour test and by the Kolmogorov-Smirnov one-sample test. The dispersion was random, with a slight tendency toward aggregation that was independent of density. In the field, the dispersion was analysed by the Clark and Evans nearest-neighbour test and by the X² approximation to Fisher's coefficient of dispersion. The dispersion was again random, this time with a slight tendency toward spacing out that was independent of density. The tendency toward aggregation displayed in the laboratory was independent of the dispersion pattern shown at the start of the experiment, and also unaffected by the edge of the container. The apparent randomness suggests that T. tenuis is primarily a suspension feeder, but may be a deposit feeder under certain environmental conditions.     

Spacing and group coordination in a nocturnal primate, the golden brown mouse lemur (Microcebus ravelobensis): the role of olfactory and acoustic signals

In order to remain stable, dispersed social groups have to solve two fundamental problems: the coordination of movement and cohesiveness within a group and the spacing between the groups. Here, we investigate mechanisms involved in intra-group coordination and inter-group spacing using the golden brown mouse lemur, Microcebus ravelobensis, as a model for a nocturnal, solitary foraging mammal with a dispersed social system. By means of radiotelemetry and bioacoustics we studied the olfactory and vocal behaviour during nocturnal dispersal and reunion of five sleeping groups.All groups used 3–17 sleeping sites exclusively, suggesting a sleeping site-related territoriality and competition for them. The occurrence of olfactory and vocal behaviour showed an asymmetrical temporal distribution. Whereas marking behaviour was observed exclusively during dispersal, a particular call type, the trill, was used by all groups during reunions. Interestingly, these trills carried group-specific signatures.Our findings provide the first empirical evidence for nocturnal primates in a natural environment that olfactory signals represent an important mechanism to regulate the distribution of different groups in space, whereas acoustic signals control intra-group cohesion and coordination.     

Burrow morphology,tube formation,and microarchitecture of shell dissolution by the spionid polychaete Polydora websteri

Burrow morphology, detrital tube formation within the burrow, and ultrastructure of the surface of the burrow of Polydora websteri Hartman, 1943, are described in the shells of Crassostrea virginica (Gmelin) and of Mytilus edulis Linné. Observations were made on live worms living normally in artificial preparations of polished shell covered with transparent plastic film. P. websteri is capable of settling wherever there are crevices in shell surfaces, and slowly penetrates the shell forming a U- or flask-shaped cavity. A detrital tube is formed within the burrow from detritus collected and pressed in place by the worm. During this process, the modified setae of the 5th setiger are thrust against the wall of the inner tube and help to keep the diameter of the inner tube constant throught its length. Individuals of P. websteri burrow into the shell when placed on preparations of polished shell of Mytilus edulis under plastic film. As they do so, they secrete a viscous fluid which dissolves interprismatic and interlamellar organic matrices and then dissolves the exposed crystals. Etched prisms and lamellae reveal a complex pattern of internal dissolution. Enlargement of the burrow by the worm thus takes place by chemical dissolution of shell and also probably by flushing of loosened partially dissolved prisms and lamellae from the tube. Some shell fragments become incorporated in the walls of the detrital tube. There is no evidence of setal abrasion on the shell walls of the burrow even at the ultrastructural levelUniversity of Delaware College of Marine Studies Contribution No. 99.