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.     

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.     

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.     

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.     

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.     

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.     

A not so natural history of the tarantula Brachypelma vagans: Interaction with human activity

We describe the structure of a population of Brachypelma vagans (Ausserer, 1875) in relation to the intensity of human activity and report characteristics of the burrows in Campeche, Mexico. During September and October 2003, we established sampling areas in five different classes of vegetation type/land use: mature forest (MF), secondary forest (SF), backyard (BY), and a football field divided into corner area (FC) and goal area (FG). The densities of spiders and the proportion of different age/gender classes of individuals on the sites were compared. Morphological data among adults and juveniles were contrasted, and differences in morphology between juveniles were tested according to land use class. We compared the nearest distances between neighbouring burrows and between burrows and trees. Also, we studied the orientation of the burrows, and compared the diameter of the burrow entrance. Brachypelma vagans was found exclusively in the open areas with densities that ranked from 0.02 to 0.1 individuals per square metre, being among the highest ever reported for Theraphosidae. However, there was a negative relationship between density and intensity of human activity. The population of this tarantula shows segregation in occupation of space. Females occupied exclusively the backyards, whereas juveniles occupied sites according to their stage of development. The youngest juveniles occupied the backyards, while the pre‐adults occupied the football field. The distance between burrows was highly variable at all the sites. However it tended to be shorter in the backyards. The orientation of burrows was in all sites preferentially directed northwards. The diameter of the burrow entrances was a relatively good indicator of the sex and age of its occupier, and related almost directly to the dimension of the body. This study provides better knowledge of the structure of B. vagans populations in a human‐modified environment and gives new information on the natural history of these spiders.     

The bioacoustics of the burrowing frog Heleioporus (Leptodactylidae)

Summary

The frog Heleioporus (Anura: Leptodactylidae) calls from a burrow. This paper examines the acoustic properties of the burrow by experimentally actuating plaster casts taken from five species: H. eyrei, H. psammophilus, H. albopunctatus, H. inornatus and H. barycragus. These findings are correlated with the properties of both natural calls of the frogs from the burrow and calls induced from the frog outside of the burrow. The morphology of the burrow enhances the principal frequency component of the call by resonance. Frogs call from a slight bulge beneath the burrow entrance. This calling site corresponds to the optimum position for generating a sound pressure maximum within the burrow. This was determined experimentally in an artificially constructed burrow with a complex internal sound source. In addition to optimizing the acoustic properties of the burrow the frog, in this position, has an open visual field from the burrow entrance. We conclude that the burrow system may compensate for the lack of a vocal sac in these frogs.     

Ecology and behaviour of the ‘road tarantulas’ Eupalaestrus weijenberghi and Acanthoscurria suina (Araneae,Theraphosidae) from Uruguay

Spatial distribution, burrow morphology, phenology, defensive displays and predation of two theraphosid spiders living in meadows were studied in Uruguay. Exhaustive field studies complemented with laboratory observations were carried out. Eupalaestrus weijenberghi occurred throughout the country while Acanthoscurria suina occurred only in the southern half. Both species seemed to be the most abundant theraphosid spiders in Uruguay as they live in the dominant landscape of the country. Burrows from both species showed a similar morphological pattern but a terminal narrow tube was found only in E. weijenberghi. We observed that A. suina always co‐occurred with E. weijenberghi but this species frequently occurred in the absence of the former. When co‐occurring A. suina showed an aggregated distribution while E. weijenberghi was randomly distributed. A restricted reproductive period was found for both species, mainly between March and April, evidenced by the massive presence of walking adult males and then confirmed by pit‐fall traps. Moulting, oviposition and longevity were also studied. Conspicuous species‐specific defensive behaviours were displayed by the tarantulas in all cases, when collected. These tarantulas showed spatial co‐occurrence and temporal coexistence with the beetle Diloboderus abderus, an important pest of grasslands. Tarantula predation on adult beetles was confirmed, these spiders being one of the few beetle predators. Their maximal efficiency in beetle predation could reach 65?000 prey captured per hectare by month. These polyphagous tarantulas seem to be potential agents for the biological control of beetle pests.     

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.     

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.     

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.     

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.     

Asymmetric larval competition between two species of neotropical foam‐nesting frogs: Leptodactylus fuscus and Engystomops pustulosus

Larval competition was investigated in foam‐nesting frogs with contrasting breeding strategies. Leptodactylus fuscus nests in burrows with moderate numbers of large eggs, mainly on dry nights; Engystomops pustulosus makes floating nests with large numbers of small eggs, mainly on wet nights. Both use the same temporary pools in open habitat over an extended breeding season. Larval growth to metamorphosis was assessed in a semi‐natural experiment and in the laboratory. Tanks contained either single species or equal numbers of both species. Inter‐specific competition was asymmetric; L. fuscus survived as well as or better in competition with E. pustulosus than in intra‐specific competition. Leptodactylus fuscus attained larger sizes with E. pustulosus present. In interspecific competition E. pustulosus showed worse survival and grew slower and to a smaller size. Outdoors, both species tolerated water temperatures that often rose to 42°C and reached metamorphosis in very short times: 14 days after oviposition in L. fuscus and 17 days in E. pustulosus.     

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.     

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.     

The taxonomy and biology of the Polystomatidae (Monogenea) in Australian freshwater turtles (Chelidae,Pleurodira)

Polystomoides scottae n. sp. is described from the urinary and accessory bladders of Chelodina expansa. Four species of Neopolystoma are described from the conjunctival sacs of Australian freshwater turtles (Chelidae): Neopolystoma cribbi n. sp. from Emydura signata, E. macquarii, Elseya latisternum and Chelodina expansa; Neopolystoma spratti n. sp. from Chelodina longicollis; Neopolystoma tinsleyi n. sp. from Chelodina expansa; and Neopolystoma queenslandensis n. sp. from Emydura signata and E. macquarii. Polystomoides australiensis, Neopolystoma macleayi and N. chelodinae are redescribed from new individuals. Neopolystoma australis is made a junior synonym of N. kreffti. P. australiensis, N. macleayi and N. cribbi produced on average 5·6, 4·3 and 0·7 eggs per day, respectively. Eggs of P. australiensis take 33–57 days at 18–24°C and 30–33 days at 24–28°C to develop and hatch. Oncomiracidia of P. australiensis and N. macleayi had 64 ciliated cells; the patterns of the ciliated cells are the same for both species. Different sizes of worms in different sites in the hosts suggest strongly that the oncomiracidia of P. australiensis and N. macleayi enter the host, migrate to the ureters, possibly to the kidneys, then to the urinary bladder and finally to either the accessory bladders or cloaca. Successful cross-infection experiments between host genera (Emydura to Elseya) indicated that P. australiensis and N. macleayi have a low degree of host specificity.     

On the structure and funtion of white-tailed prairie dog burrows

The architecture of burrows of the white-tailed prairie dog ( Cynomys leucurus ) is poorly known. For this reason and for comparative purposes, one recenlty active burrow of this species was excavated in southern Montana; the detailed methodology is described. Data were compiled on the dimensions of 29.3 m of excavated passages, and interpretations of several features are discussed. A ""turning bay,"" sleeping quarters, two hibernacula, and a maternity area are described, the last feature for the first time in print. In addition, we report Cynomys using their teeth to dig, also for the first time. Further, an inadvertent remodeling of the burrows is ascribed to normal animal traffic and appears to confirm a prediction based on late Pleistocene fossil burrows in Alberta.     

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.