Characterisation of burrow architecture under natural conditions in the sand-dwelling wolf spider Allocosa brasiliensis

Allocosa brasiliensis (Petrunkevitch, 1910) is a wolf spider that constructs silk-lined burrows along the coastal sand dunes of Argentina, Brazil and Uruguay. This species shows a reversal in typical sex roles and sexual size dimorphism expected for spiders. Females are the smaller and mobile sex, which initiates courtship at the male burrow entrance. Mating occurs in the male burrow, and when it ends, the male leaves and the female stays inside. Females prefer to mate with those males showing longest burrows, so burrow dimensions would be under strong sexual selection pressures. Previous studies in the laboratory indicated that male burrows are longer than those of virgin females, which were described as simple silk capsules. Preliminary observations suggested that juvenile burrows presented differences from those of adults; however, detailed observations of A. brasilienisis burrow characteristics at the field were lacking. The aim of this study was to characterise the burrows of adults and juveniles of A. brasiliensis under natural conditions. We recorded the dimensions of burrows inhabited by males, females and juveniles (n = 30 for each category) and created beeswax moulds that reflected burrow shape. Juveniles inhabited tubular burrows with two branches and two openings; on the contrary, adults were found in tubular burrows with a single entry. Males and females inhabited burrows of similar length and width, but those of juveniles were shorter and narrower. We discuss the results and their possible functional explanations according to the selective pressures expected for each developmental stage and sex.     

Reproductive biology of Limnoria chilensis: another boring peracarid species with extended parental care

Several boring peracarid species engage in extended parental care with parents tolerating small juveniles in their burrows, but only anecdotal observations have been reported from boring isopods of the genus Limnoria. The isopod Limnoria chilensis Menzies is frequently found in holdfasts of the macroalgae Macrocystis pyrifera (L.) C. Agardh and Durvillea antarctica (Chamisso) Hariot. In the present study, holdfasts of D. antarctica that harboured L. chilensis were carefully dissected in order to examine the association pattern of reproductive females and small juveniles. In most infested holdfasts, L. chilensis reached very high densities of 43.7 ± 3.9 (mean ± SE) individuals cm^?2 (range: 0–90 individuals cm^?2). The overall sex ratio (females:males) was close to unity (1.2). Male size varied between 1.5 and 2.6 mm body length (BL) and female size between 1.5 and 3.6mm BL. The percentage of reproductive females was low (19.7% of all females), and only the largest females (>2.6 mm BL) were reproductive. Embryo numbers ranged between six and 19 per female and a significant correlation between female BL and the number of embryos was found. Several females were found with small juveniles (0.8–1.2 mm BL) in the terminal end of their burrows. On several occasions, aggregations of similar-sized juveniles (1.0–1.5 mm BL) in their own burrows were found near a female burrow, indicating that these juveniles initiated their first individual burrows from within the maternal burrows. Some females with small juveniles in the terminal end of their burrows were either accompanied by males or they were ovigerous, indicating that they may have been in the process of producing a second brood. The fact that juveniles build their first burrows in the protection of female burrows suggests that such small juveniles have not yet developed full boring capacity. Extended parental care in this (and other boring) peracarid species represents a mechanism facilitating high juvenile survival rates. Given the highly local recruitment, it is suggested that the reproductive biology of this isopod has strong implications for Its population biology.     

Extended parental care in two endobenthic amphipods

Extended parental care is described for two endobenthic amphipods which inhabit the estuarine soft-bottoms along the Gulf of Maine. The juveniles of both amphipod species, Leptocheirus pinguis and Casco bigelowi, remain in the burrow of their mother after they hatch from the brood pouch. Several consecutive clutches of L. pinguis can inhabit the female's burrow simultaneously, but in C. bigelowi there is always only one clutch of juveniles in the burrow of the female. Larger juveniles of L. pinguis start building their own small tubes at the bottom of the female's burrow. Casco bigelowi females increase the size of their burrows during the time when the juveniles grow. The juveniles of both species leave the female's burrow at about half adult size. The major tasks of female L. pinguis and C. bigelowi during the time of extended parental care is to irrigate and maintain the deep burrow. It is hypothesized that extended parental care in L. pinguis, C. bigelowi and other tube/burrow-living peracarids is a mechanism to protect small offspring from epibenthic predation.     

Behaviour and habitat of Neohela monstrosa (Boeck, 1861) (Amphipoda: Corophiida) in Norwegian Sea deep water

There are few in situ observations of deep-sea macrofauna, due to the remoteness of this ecosystem. Visual surveys conducted for marine management by MAREANO, (marine area database for Norwegian waters) and the petroleum industry (by SERPENTS, scientific and environmental remotely operated vehicle partnership using existing industrial technology) have provided unique material of visual information from large areas in the Norwegian Sea. The distribution, density and behaviour of the deep-sea amphipod Neohela monstrosa (Boeck, 1861) is described based on videos and samples from the Norwegian Sea. This amphipod is common on mud bottoms at 200–2181 m depth in the area. Dense communities were found in stands of the arctic sea pen Umbellula encrinus at more than 1000 m depth where temperatures were below 0° C. The mean density of N. monstrosa observed for larger areas was 4/100 m² but densities of 15–36 individuals per m² were found in local patches. It is domicolous which is characteristic of the superfamily Corophiida and digs burrows in soft muddy bottoms primarily by using large shovel-like gnathopods to scoop the sediment out. The amphipod was observed pushing and rolling sediment balls out of its burrow, which were probably held together with amphipod silk. It digs out an upper 3 to 4 cm wide burrow with a horizontal side burrow a couple of centimetres down. Neohela monstrosa appears to feeds on newly settled detritus that it collects from the surface sediment through the use of its long antennae while the burrow is mainly used for protection against predators such as demersal fish. Newly released juveniles are probably kept in the burrow for protection. Based on the local high density of N. monstrosa together with its habit of making long burrows, we suggest that there is significant bioturbation associated with the presence of N. monstrosa in deep sedimentary habitats of the Norwegian Sea, which likely provides an important ecosystem function.     

Burrow occupancy patterns of the Western Burrowing Owl in southern Nevada

Understanding long-term patterns of burrow occupancy for the Western Burrowing Owl ( Athene cunicularia hypugaea ) is necessary for the conservation of this species, especially in arid, desert ecosystems where burrow occupancy data are lacking. Monthly burrow monitoring was conducted over a 4-year period (1997–2001) in southern Nevada to determine burrow occupancy patterns of Burrowing Owls and to evaluate the effects of burrow type and desert region on burrow occupancy. Burrow occupancy occurred year-round and was most consistent in the Transition region and tended to be lowest in the Mojave Desert region. Peak burrow occupancy occurred during March through May, followed by a gradual decline in occupancy through the summer and fall until January and February, when occupancy was lowest. Occupancy was significantly higher at sites with both culvert and pipe burrows than at sites with earthen burrows in disturbed habitat or earthen burrows in natural habitat. Breeding-season occupancy was not significantly higher in wetter, cooler portions (e.g., Great Basin desert region) of the study area. Results suggest that occupancy is influenced by habitat features—such as suitable burrows in open areas with low vegetation—and climatic regime.     

Juvenile Sphaeroma quadridentatum invading female-offspring groups of Sphaeroma terebrans

Female isopods Sphaeroma terebrans Bate 1866 are known to host their offspring in family burrows in aerial roots of the red mangrove Rhizophora mangle. During a study on the reproductive biology of S. terebrans in the Indian River Lagoon, Florida, USA, juvenile S. quadridentatum were found in family burrows of S. terebrans. Between September 1997 and August 1998, each month at least one female S. terebrans was found with juvenile S. quadridentatum in its burrow. The percentage of S. terebrans family burrows that contained juvenile S. quadridentatum was high during fall 1997, decreased during the winter, and reached high values again in late spring/early summer 1998, corresponding with the percentage of parental female S. terebrans (i.e. hosting their own juveniles). Most juvenile S. quadridentatum were found with parental female S. terebrans, but a few were also found with reproductive females that were not hosting their own offspring. Non-reproductive S. terebrans (single males, subadults, non-reproductive females) were never found with S. quadridentatum in their burrows. The numbers of S. quadridentatum found in burrows of S. terebrans ranged between one and eight individuals per burrow. No significant correlation between the number of juvenile S. quadridentatum and the numbers of juvenile S. terebrans in a family burrow existed. However, burrows with high numbers of juvenile S. quadridentatum often contained relatively few juvenile S. terebrans. The majority of juvenile S. quadridentatum found in family burrows of S. terebrans were smaller than the juvenile S. terebrans that were cared for by their mothers. The results indicate that the presence of S. quadridentatum in S. terebrans family burrows may negatively affect the duration of extended parental care in S. terebrans. It is not known why parental female S. terebrans are not able to discriminate against juvenile S. quadridentatum. Possibly, the fact that the two species are closely related facilitates S. quadridentatum sneaking into S. terebrans family burrows.     

Burrowing Owl nest success and burrow longevity in north central Oregon

We studied nest success, burrow longevity, and rates of burrow reuse for a migratory population of Burrowing Owl ( Athene cunicularia ) in north central Oregon from 1995 to 1997. Nest success varied annually from 50% to 67%. Principal causes of nest failure were desertion (26%) and depredation by badgers ( Taxidea taxus ; 13%). Reuse of available nest and satellite burrows in subsequent years was 87% in 1996 and 57% in 1997. Reuse was highest at burrows in sandy soils, which may indicate that nest-site availability is a limiting factor in sandier soil types. Trampling by livestock resulted in the loss of 24% of all burrows between one season and the next, and natural erosion resulted in closure of 17%. Both causes of burrow failure occurred more frequently in soils with a sand component due to their friable nature. We recommend that habitat used by livestock be evaluated for use by Burrowing Owls, that occupied areas be managed to minimize destruction of burrows by livestock, and that predator-control efforts be revised to exclude mortality of badgers.     

Sexual dimorphism in chelicerae,forelegs and palpal traits in two burrowing wolf spiders (Araneae: Lycosidae) with sex-role reversal

Allocosa alticeps and Allocosa brasiliensis are two sex-role-reversed spiders. Females locate males and initiate courtship. Copulation occurs inside male burrows and after copulation the males deliver their burrows to the females. We explored the occurrence of differences in morphological traits related to sex roles in both Allocosa species and compared the results with two non-burrowing wolf spiders with typical sex roles. We measured the length of the foreleg's tibia-patella and the chelicerae. Scanning electron micrographs of palpal tarsus distal sections were taken. Males showed higher values than females in all the traits considered. Adult and penultimate males lacked true claws compared with juveniles and females, but showed modified spines. The palpal organ was more proximal to the tibia compared with the non-burrowing lycosids considered in this study. Spines and palpal organ location could be associated with more effective digging. We discuss how natural and sexual selection could interact on morphology in this scenario.     

Nest-site characteristics of Burrowing Owls ( Athene cunicularia ) in the Snake River Birds of Prey National Conservation Area, Idaho, and application to artificial burrow installation

Burrowing Owl ( Athene cunicularia ) populations are declining in many portions of their range, and research and management efforts into stemming declines are underway. One tool with promise is the artificial burrow, which can supplement nesting opportunities and play a role in research, mitigation, translocation, and reintroduction studies. However, few studies directly assess important burrow and surrounding topographic features upon which owls choose sites and then construct and install artificial burrows accordingly. In this study we (1) measure physical, vegetative, and topographic characteristics of Burrowing Owl nest sites in the Snake River Birds of Prey National Conservation Area (SRBPNCA); (2) compare used and unused burrows to determine features important in nest-site selection; and (3) use this information to help guide current and future construction and placement of artificial burrows in the SRBPNCA. Owls nested in abandoned American badger ( Taxidea taxus ) burrows in areas with more than one burrow, close to roads and irrigated agricultural fields, and characterized by sparse and low vegetation dominated by nonnative plant species. Only one feature studied, tunnel entrance angle, corresponded with choice by owls; odds of burrow use decreased 17% with each 1° increase in slope of the tunnel entrance. Owls nesting near irrigated agricultural fields also had higher productivity. We discuss applications of our results to construction and placement of artificial burrows in the SRBPNCA and similar shrub-steppe environs in western North America.     

Burrow morphology of the echiuran worm Maxmuelleria lankesteri (Echiura: Bonelliidae), and a brief review of burrow structure and related ecology of the Echiura

The burrow morphology of the echiuran worm Maxmuelleria lankesteri (Herdman) was investigated in situ using a resin casting technique. Work was carried out in Lochs Sween and Creran on the west coast of Scotland in predominantly fine mud sediments. Burrow casts typically had only 1 opening, although there is evidence to suggest that a second opening may exist. In 58% of burrows, the opening, which was small and funnel-shaped, was associated with a surface mound reaching up to 20 cm in height. The tunnel below the burrow opening, the ‘neck’, was narrow and circular in transverse section with a smooth wall, possibly due to the action of the mucus-laden proboscis which emerges during feeding. The maximum burrow depth recorded was 80 cm and tunnel orientation became more horizontal with increasing depth. In some larger burrows, the tunnel began to orientate upwards towards its end. Below the neck, the tunnel was much wider but had a sub-circular transverse section. The burrows had distinct striations on the walls of lower tunnels, possibly caused by movements of the animal within. In some cases, a community of symbiotic organisms had developed, including polychaetes and 2 species of bivalve, Mysella bidentata (Montagu) and Saxicavella jeffresii Winckworth. Evidence of burrow modification by the crustacean Jaxea nocturna Nardo was noted and gobies, including Gobius niger Linnaeus, were also responsible for some alterations to the upper burrow around the opening. Evidence suggested that burrows were permanent structures which changed little in position.     

Temporal changes in burrow structure of the thalassinidean ghost shrimps Trypaea australiensis and Biffarius arenosus

The thalassinidean ghost shrimps Trypaea australiensis and Biffarius arenosus are dominant burrowing macroinvertebrates of soft‐sediment habitats in Western Port, Victoria, Australia. Burrow structure has been described for both species but little is known about how the burrows change over time. This study used resin casting to investigate temporal variation by making burrow casts each month over a 12‐month period. It was found that the overall burrow morphology of both T. australiensis and B. arenosus was consistent over time, but in the warmer months (summer and spring), both species constructed burrows that were deeper and of greater volume than burrows constructed in winter and autumn. Possible reasons for this variation are discussed.     

Species richness and California voles in an annual and a perennial grassland

Populations of a common burrowing rodent, Microtus californicus (the California vole), thrive in ungrazed or lightly grazed grasslands in coastal California. Two sites ungrazed by livestock, one dominated by native perennial grasses and another dominated by invasive annuals, were evaluated over 2 consecutive years for the relationship between plant species richness and location of M. californicus burrow entrances (burrows). Plant species and burrows were sampled as present or absent in contiguous 1-m 2 quadrats on a 100-m 2 grid. Quadrats with burrows averaged significantly more plant species than quadrats without them (11.3 vs. 9.9 species, P < 0.001). Burrows found in 1996 were not correlated with species richness in 1995, suggesting that voles affect richness rather than seek it out. Vole burrow locations showed significant clumping on the annual site and trended toward clumping on the perennial site in both 1995 and 1996. Because voles seem to create a clumped pattern with their burrow entrances, the associated increase in plant species richness may have a strong effect on the overall structure of the plant community. A quantitative comparison of the 2 sites showed that the plant matrix of the perennial site contained flora of the annual site. This similarity in plant species composition may allow for similar treatment of our 2 types of sites and potentially other California grasslands. Undetected increases in vole populations with livestock grazing reduction may account for the erratic results from grasslands management research and the inconsistent success of derived management practices.     

Another migid in the wall: natural history of the endemic and rare spider Calathotarsus simoni (Mygalomorphae: Migidae) from a hill slope in central Argentina

A population of Calathotarsus simoni Schiapelli and Gerschman (1975) was discovered on a hillside in the Ventania system, Argentina. Our objectives were to quantify burrow density, record burrow morphology and door characteristics and describe the micro-habitat. We counted 57 burrows and report a density of 0.01 burrows/m². Aggregation indices suggest that burrows are aggregated under some area plots but more evenly distributed on others. The trapdoor is thick and rigid with bevelled edges connected to the entrance rim by a narrow articulated hinge. Two egg sacs from females were obtained and data on eggs and spiderlings are presented. We registered six burrows of an undetermined species of Actinopus. While a few specimens of Actinopus sp. were found inhabiting this hillside, the highest proportion of burrows belonged to C. simoni. Spider diversity on the hillside shows the predominance of Linyphiidae, Nemesiidae and Gnaphosidae. One juvenile of C. simoni was captured using pitfall traps.     

Across-shore distribution of Ocypode quadrata burrows in relation to beach features and human disturbance

Spatial distribution patterns of the ghost crab Ocypode quadrata were analysed using different approaches at one disturbed and two preserved reflective sandy beaches of the Mexican Caribbean. We used spatial correlation analysis to describe the across-shore distribution of O. quadrata and their patch dynamics by beach and sampling time. In addition, we analysed the across-shore variability of habitat extent of O. quadrata (habitat envelope) and its relation to human disturbance and beach features. The spatial structure of burrow density was consistent through time, showing a bimodal distribution with very low abundance in the mid distribution range and a discrete increment of burrows to landward. Nonetheless, the size of patches varied temporally for the three beaches. Burrow size increased from seaward to landward. The location of the first burrow and the habitat envelope varied among sampling times and beaches. The location of the first burrow is mostly mediated by the swash climate, while the habitat envelope is mainly controlled by the level of human disturbance. Despite the low number of disturbed and control beaches, our results suggest that the habitat envelope could be included in future studies when using ghost crabs as indicators of human disturbance. Polynomial models and variographic analysis proved to be useful tools to describe the across-shore distribution and patch dynamics of the ghost crab.     

Foraging by the flower-dwelling spider,Misumenops argenteus (Thomisidae), at high prey density sites

In this study, the foraging sites of the crab spider Misumenops argenteus on Trichogoniopsis adenantha (Asteraceae) were investigated over a 2-year period. The choice of sites was examined in relation to prey density in rainy and dry seasons, in leaves versus stems, reproductive branches versus vegetative branches, and in flowerheads in different phenophases. Misumenops argenteus generally occupied sites where the prey frequency was high, indicating that this species was able to evaluate the quality of the foraging sites, as predicted by optimal foraging theory. One exception to the rule was that adult males occurred at all sites, independently of prey density, as also observed in other spider species in which males search for females and not for prey. In addition, young (third and fourth instar) spiders occupied flowerheads with few prey, but which provided shelter.     

Spatial distribution of the ghost crab Ocypode quadrata in low‐energy tide‐dominated sandy beaches

The spatial distribution of the ghost crab Ocypode quadrata (Crustacea, Brachyura) was described in five low‐energy tide‐dominated sandy beaches in São Sebastião Channel (Grande, Zimbro, Pitangueiras, Cabelo Gordo, and Segredo). On each beach, the zonation study was performed on five random 1?m wide transects sampled from the water line to the vegetation. In general, individuals occurred above 1?m in height in relation to mean low water (MLW) and peaked between 1.5 and 2.0?m. The sandy beaches were then divided into three 1?m wide strips (tidal levels) parallel to the water line and equivalent to medium intertidal, upper intertidal, and subterrestrial fringe to test the effect of beach and tidal level on the abundance and size of O. quadrata. These strips were sampled as a whole using adjacent 1 m² squares. A previous evaluation showed a positive significant relationship between carapace length and burrow diameter, thus supporting crab size estimates from burrow openings. The smallest individuals occurred mainly in the medium intertidal, but were also recorded in the subterrestrial fringe, which was occupied mainly by large‐sized individuals. In general, the individuals were randomly dispersed within the strips and concentrated in the upper intertidal zone on all the sandy beaches. The density of O. quadrata varied among beaches, with a tendency to lower values on areas with very fine and poorly or moderately sorted sediments and on that most used for recreational activities (Grande). Burrow size also varied among beaches, with a tendency of smaller burrows in areas with coarser sand grains and higher tourism. These results indicate that the effect of human impact on density and size of O. quadrata may be confounded by the high environmental heterogeneity of the studied system, which may make it difficult to utilize this species as an indicator of the conservation status of low‐energy tide‐dominated sandy beaches.     

Habitat use patterns within the home range of pygmy rabbits ( Brachylagus idahoensis ) in southeastern Idaho

Pygmy rabbits ( Brachylagus idahoensis ) are a small sagebrush ( Artemisia tridentata ) obligate lagomorph found within the Great Basin of northwestern United States. Because of its reliance on sagebrush, this species is thought to be experiencing a major range reduction as a result of loss of sagebrush habitat. To aid in conservation of this species, we need to better understand its use of the sagebrush environment. We estimated summer home range use patterns by relocating 5 radio-collared pygmy rabbits (3 females and 2 males) over a 24-hour cycle. We then compared soil texture, shrub density, height, and canopy cover between areas close to burrow entrances and areas of high use and low use. Mean home range sizes of female and male rabbits were 37.2 and 67.9 ha, respectively. Rabbits had disproportionate amounts of time (68.4% ± 9.1, s￣x ) and travel (63.0% ± 5.7, s￣x ) in areas within a 60-m radius of their burrows. Soil texture did not differ among the 3 areas, but shrub density, specifically big sagebrush, and forb density were significantly higher close to the burrow than in the high- and low-use areas. We conclude that pygmy rabbits are possibly burrow obligates and that their abundance and distribution are likely limited by available burrow sites.     

The parasitoid complex and parasitoid‐induced mortality of spiders (Araneae) in a Central European woodland

The parasitoid complex of spiders was studied in three different sites of a forest and, for comparison, in open habitats in north‐western Lower Saxony (Germany). Qualitative data and, for four spider species, detailed quantitative rearing data were obtained. At least 25 parasitoids of spiders were recorded (23 species of Hymenoptera and two species of Diptera). External parasitoids of spiders were rare in the woodlands; only up to 1% of the spiders collected by hand searching and sweep netting had ectoparasitoids. During the investigation 23 parasitoid species were reared from spider egg masses, 10 of them occurring in woodlands. Two species were secondary parasitoids. Several parasitoid–host relationships were recorded for the first time. Egg masses of Floronia bucculenta were parasitized up to 5% by two species of the genus Gelis. Up to 10% of the egg masses of Linyphia hortensis were parasitized by Aclastus species. Egg masses of the spider genus Ero were parasitized by three species, causing an overall mortality of 40%. For egg masses of Agroeca spp. parasitization at some locations was up to 60%. An overview of the Central European parasitoid complex of spiders is provided.     

Correlates of burrow location in Beechey ground squirrels

Partial correlation analysis indicates that Beechey ground squirrels show a strong preference for digging burrows under and around large objects, may show a weaker tendency to locate their burrows under the cover of tree canopies, and avoid digging burrows in areas with both treecanopy and ground cover (stones, logs). These relationships hold for large but not small burrow systems. The need for unobstructed visual surveillance and an autumn food supply are proposed to be determinants of these preferences.