Microhabitat segregation in two desert rodent species: the relation of prey availability to diet

Summary I studied diet in relation to microhabitat use in two desert rodents:Microdipodops megacephalus, the dark kangaroo mouse, andPeromyscus maniculatus, the deer mouse. Contrary to expectation, both species ate primarily arthropods, which were most abundant near shrubs.Peromyscus used the area near shrubs, in contrast toMicrodipodops, which used open microhabitat. As a consequence, the diet ofPeromyscus was narrower and more concentrated on abundant prey types than that ofMicrodipodops. Thus microhabitat segregation, which is frequently reported for desert rodents, is related to a diet-breadth difference between these rodents. The use of open microhabitat and low density resources byMicrodipodops, when compared with the large bipedalDipodomys and small quadrupedalPerognatus, suggests that bipedal locomotion in desert rodents is related to use of open microhabitat, and that body size is related to density of food resources.     

When perception reflects reality: Non‐native grass invasion alters small mammal risk landscapes and survival

Modification of habitat structure due to invasive plants can alter the risk landscape for wildlife by, for example, changing the quality or availability of refuge habitat. Whether perceived risk corresponds with actual fitness outcomes, however, remains an important open question. We simultaneously measured how habitat changes due to a common invasive grass (cheatgrass, Bromus tectorum) affected the perceived risk, habitat selection, and apparent survival of a small mammal, enabling us to assess how well perceived risk influenced important behaviors and reflected actual risk. We measured perceived risk by nocturnal rodents using a giving‐up density foraging experiment with paired shrub (safe) and open (risky) foraging trays in cheatgrass and native habitats. We also evaluated microhabitat selection across a cheatgrass gradient as an additional assay of perceived risk and behavioral responses for deer mice (Peromyscus maniculatus) at two spatial scales of habitat availability. Finally, we used mark‐recapture analysis to quantify deer mouse apparent survival across a cheatgrass gradient while accounting for detection probability and other habitat features. In the foraging experiment, shrubs were more important as protective cover in cheatgrass‐dominated habitats, suggesting that cheatgrass increased perceived predation risk. Additionally, deer mice avoided cheatgrass and selected shrubs, and marginally avoided native grass, at two spatial scales. Deer mouse apparent survival varied with a cheatgrass–shrub interaction, corresponding with our foraging experiment results, and providing a rare example of a native plant mediating the effects of an invasive plant on wildlife. By synthesizing the results of three individual lines of evidence (foraging behavior, habitat selection, and apparent survival), we provide a rare example of linkage between behavioral responses of animals indicative of perceived predation risk and actual fitness outcomes. Moreover, our results suggest that exotic grass invasions can influence wildlife populations by altering risk landscapes and survival.     

Competition and habitat selection in Namib desert tenebrionid beetles

Summary We tested whether intra- and interspecific competition could affect habitat selection in the two most abundant tenebrionid beetles,Physadesmia globosa andOnymacris rugatipennis, in a dry riverbed in the Namib desert. The spatial distributions of these beetles at the microhabitat scale were negatively correlated. We performed a removal experiment, progressively removing first 25% and then a further 25% of the population of the most abundant species,P. globosa, under the trees where most of the preferred food of both species is concentrated. There was no response ofO. rugatipennis to this removal in the tree habitat. In the open, barely-vegetated habitat where mostO. rugatipennis are found, the number of this species caught in pitfall traps increased following both removals and decreased followingP. globosa replacement under the trees. It appears that intraspecific competition forces someP. globosa to occupy the open habitat. Interspecific competition betweenP. globosa andO. rugatipennis in the open habitat reduces the number ofO. rugatipennis that can co-exist withP. globosa there. Removal ofP. globosa under the trees allows conspecifics in the open habitat to move under the trees, releasingO. rugatipennis in the open habitat from competition. This then results in an increase in the numbers ofO. rugatipennis in the open habitat as a result of immigration from neighbouring areas. We found that differences in foraging efficiency, measured as giving-up times in artificial food patches, create a likely mechanism of co-existence that explains the distinct preferences of these two species for tree and open habitats.     

The effects of owl predation on the foraging behavior of heteromyid rodents

Summary Researchers have documented microhabitat partitioning among the heteromyid rodents of the deserts of North America that may result from microhabitat specific predation rates; large/bipedal species predominate in the open/risky microhabitat and small/quadrupedal species predominate in the bush/safer microhabitat. Here, we provide direct experimental evidence on the role of predatory risk in affecting the foraging behavior of three species of heteromyid rodents: Arizona pocket mouse (Perognathus amplus; small/quadrupedal), Bailey's pocket mouse (P. baileyi; large/quadrupedal), and Merriam's kangaroo rat (Dipodomys merriami; large/bipedal). Both kangaroo rats and pocket mice are behaviorally flexible and able to adjust their foraging behavior to nightly changes in predatory risk. Under low levels of perceived predatory risk the kangaroo rat foraged relatively more in the open microhabitat than the two pocket mouse species. In response to the presence of barn owls, however, all three species shifted their habitat use towards the bush microhabitat. In response to direct measures of predatory risk, i.e. the actual presence of owls, all three species reduced foraging and left resource patches at higher giving up densities of seeds. In response to indirect indicators of predatory risk, i.e. illumination, there was a tendency for all three species to reduce foraging. The differences in morphology between pocket mice and kangaroo rats do appear to influence their behavioral responses to predatory risk.     

Macro- and microhabitat use of Telfair’s skink (Leiolopisma telfairii) on Round Island, Mauritius: implications for their translocation

The successful eradication of introduced rodents from islets off the coast of Mauritius has led to local conservation bodies investigating the possibility of translocation as a measure of safeguarding endemic reptile populations. The present study was the first to determine the habitat and microhabitat requirements of Telfair's skinks (Leiolopisma telfairii) on Round Island, Mauritius, with a view to aiding future translocation projects to islands within their historic range. Contrasting preferences found for Telfair's skink at macro- and micro- habitat levels underline the importance of sampling at multiple ecological scales in such investigations. Significantly fewer sightings of L. telfairii were recorded in bare rock habitats compared to more vegetated habitats. Conversely, at a microhabitat scale principal component analysis indicated structural characteristics were the primary determinant of microhabitat choice. The first dietary analysis of Telfair's skinks confirmed their status as omnivores. Cockroaches (Blattodea spp.) appeared to be a primary food source. Four exotic plant species were also present in faecal samples and the potential for L. telfairii to aid their dispersal is discussed. Implications for the long-term management and proposed translocation of Telfair's skinks are discussed.     

The role of microhabitat preference and social organization in determining the spatial distribution of a coral reef fish

Both habitat preferences and social organization can influence the spatial distributions of individuals. We explored effects of individual behavior and social organization on distributions of arc-eye hawkfish (Paracirrhites arcatus) in lagoons of French Polynesia. Analysis of habitat selectivity data obtained during surveys revealed that the most highly preferred microhabitat of arc-eye hawkfish was large Pocillopora coral with an open branching morphology. However, such corals were rare and most hawkfish occupied smaller, less preferred Pocillopora. Indeed, total abundance of Pocillopora explained nearly two thirds of the lagoon-wide variation in abundance of hawkfish and the derived relationship between the numbers of hawkfish and Pocillopora predicted 86% of the spatial variation in hawkfish abundance during subsequent surveys. In contrast, large, open-branched Pocillopora explained little of the spatial variation in abundance of hawkfish. Individual behavior and social organization significantly impacted use of the most highly preferred Pocillopora. During a colonization experiment set up outside hawkfish home ranges, all colonizers resided on the most highly preferred corals. Following addition of the most highly preferred Pocillopora corals to areas occupied by hawkfish, only the largest, socially dominant individuals obtained access to added corals, spending significantly more time and increasing both aggressive acts and prey attacks from these substrates. These results illustrate the importance of understanding the modulating effects of social behavior on habitat use and explain why most hawkfish individuals do not occupy their most preferred microhabitat type.     

Microhabitat Selection of three Forest Understory Birds in the Brazilian Atlantic Rainforest

When assessing fragmentation effects on species, not only habitat preferences on the landscape scale, but also microhabitat selection is an important factor to consider, as microhabitat is also affected by habitat disturbance, but nevertheless essential for species for foraging, nesting and sheltering. In the Atlantic Rainforest of Brazil we examined microhabitat selection of six Pyriglena leucoptera (white-shouldered fire-eye), 10 Sclerurus scansor (rufous-breasted leaftosser), and 30 Chiroxiphia caudata (blue manakin). We radio-tracked the individuals between May 2004 and February 2005 to gain home ranges based on individual fixed kernels. Vegetation structures in core plots and fringe plots were compared. In C. caudata, we additionally assessed the influence of behavioural traits on microhabitat selection. Further, we compared microhabitat structures in the fragmented forest with those in the contiguous, and contrasted the results with the birds' preferences. Pyriglena leucoptera preferred liana tangles that were more common in the fragmented forest, whereas S. scansor preferred woody debris, open forest floor (up to 0.5 m), and a thin closed leaf litter cover which all occurred significantly more often in the contiguous forest. Significant differences were detected in C. caudata for vegetation densities in the different strata; the distance of core plots to the nearest lek site was significantly influenced by sex and age. However, core sites of C. caudata in fragmented and contiguous forests showed no significant differences in structure. Exploring microhabitat selection and behavior may greatly support the understanding of habitat selection of species and their susceptibility to fragmentation on the landscape scale.     

Microhabitat preference constrains invasive spread of non-native natal grass (<Emphasis Type="Italic">Melinis repens</Emphasis>)

Preventing the establishment of a non-native species is critical for ensuring the species does not become invasive, yet most non-native species will have little impact on their environment. Despite this, little is known about what influences whether a species will remain relatively benign, or whether it will cause economic or ecological harm. Understanding a plant’s microhabitat provides insight into the necessary conditions for establishment and the current distribution limitations of a population. We investigated microhabitat preference of the non-native natal grass (Melinis repens (Willd.) Zizka) in Florida scrub using microhabitat sampling to measure vegetation composition. We examined the extent to which microhabitats were associated with natal grass presence and biomass in invaded disturbed scrub and roadside plots using backwards stepwise logistic regression and general linear models to identify significant microhabitat variables. We further compared these plots with those in undisturbed, uninvaded scrub to characterize vegetation across habitat types, and used our model to predict the probability of natal grass invasion in undisturbed scrub. Natal grass preferred microhabitats with high litter volume and distance to shrubs and intermediate cactus, graminoid, and vine cover. Roadside natal grass achieved higher biomass and was less microhabitat limited than disturbed scrub natal grass. We determined that undisturbed scrub plots represent distinct microhabitats that natal grass is unlikely to invade. Microhabitat sampling provides land-managers a non-intrusive technique to assess potential habitat suitability based non-native plant preferences before a costly invasion occurs.     

Microhabitat Characteristics of Preble's Meadow Jumping Mouse High-Use Areas

ABSTRACT Riparian wetlands are complex ecosystems containing species diversity that may easily be affected by anthropogenic disturbances. Preble's meadow jumping mouse (Zapus hudsonius preblei) is a federally threatened subspecies dependent upon riparian wetlands along the Front Range of Colorado and southeastern Wyoming, USA. Although habitat improvements for Preble's meadow jumping mouse are designed at multiple spatial scales, most knowledge about its habitat requirements has been described at a landscape scale. Our objective was to improve our understanding of Preble's meadow jumping mouse microhabitat characteristics within high-use areas (hotspots), which are essential for the development of effective management and conservation strategies. We evaluated Preble's meadow jumping mouse habitat by describing areas of high use and no use as determined from monitoring radiocollared individuals. A comparison of microhabitat characteristics from random samples of high-use and no-use areas indicated that mice use areas closer to the center of the creek bed and positively associated with shrub, grass, and woody debris cover. Distance to center of the creek bed, and percent of shrub and grass cover also had the greatest relative importance of habitat variables modeled when describing high-use areas. High-use areas contained 3 times more grass cover than forb cover, and overall had a greater proportion of wetland shrub and grass cover. However, proportion of cover type (shrub or grass) did not vary greatly between high-use and no-use areas. Our results suggest that management and conservation efforts should continue to focus on establishment of native wetland vegetation near streams and creeks. For example, vegetation should include shrubs such as willow (Salix spp.), narrowleaf cottonwood (Populus angustifolia), alder (Alnus incana), grasses such as fescue (Fescue spp.), sedges (Carex spp.), and rush (Juncus spp).     

Habitat selection within species ofDrosophila: a review of experimental findings

Summary Habitat selection whereby individuals within a population have different microhabitat preferences is possibly important for genetic diversity and stability. It has received a good deal of attention, especially forDrosophila populations. The literature is summarized and an attempt is made to reconcile what appear to be contradictory findings. Some generalizations are drawn: (1) individualDrosophila choose habitats in consistent and characteristic ways; (2) some microhabitat differences are more important to flies than others; (3) genetic factors influence habitat preferences; (4) experiential factors influence habitat preferences; (5) differences in processing ability moderate the influence of experience on habitat choice. It is argued that the traditional analysis of variance models are frequently not very helpful for visualizing the process, and a computational model is suggested instead.     

Microhabitat use by rodent species in a central Brazilian cerrado

We analysed microhabitat use by the rodents Calomys tener, Necromys lasiurus and Thalpomys lasiotis and the factors that may influence their abundances in “murundu” grasslands (open fields with termite mounds) at Aguas Emendadas Ecological Station, Planaltina, Federal District, Brazil. Two grids with 100 sampling points were established and traps were placed at each intersection of the grid, where five microhabitat variables were also measured. Rodents were trapped from June through October 2008. Microhabitat explained 21% of the variation in community structure, with grass density and the number of termite mounds explaining most of the variation. Necromys lasiurus was most often captured in areas with dense grasses, whereas T. lasiotis and C. tener were most often in areas less dense with grasses.     

Conservation implications of competition between generalist and specialist rodents in Mediterranean afforested landscape

Density dependent habitat selection at the community level is regarded as a major determinant of biodiversity at the local scale, and data on these processes and how they are affected by human activities is highly applicable to conservation. By studying the competitive relationships between a specialist and a generalist we can acquire valuable insights about how different environmental elements determine species abundance and distribution and consequently biodiversity. Here we describe a study of density dependent processes that determine the community structure of two rodents: a specialist—the broad toothed mouse (Apodemus mystacinus), and a generalist—the common spiny mouse (Acomys cahirinus) in a Mediterranean maqui habitat, and how this structure is impacted by anthropogenic planting of pine stands. We carried out two field experiments: The first, based on open field trapping, looking at how rodent communities change with habitat structure. The second experiment was an enclosure study aimed at validating the habitat preferences and competitive relationship between the specialist and the generalist. We identified asymmetric competition relationships in which the specialist was dominant over the generalist. Competition intensity was lower in maqui with >10% oak cover, although both species abundances were high. Competition was found only during the limiting season (summer). Based on these findings we produced management recommendations to keep indigenous small mammals’ biodiversity high. Density dependent habitat selection processes play a central role in determining biodiversity, and understanding the mechanisms motivating these processes is needed if alterations in biodiversity in response to human disturbance are to be understood.     

Microhabitat use by wild-ranging Cabrera voles <Emphasis Type="Italic">Microtus cabrerae</Emphasis> as revealed by live trapping

The conservation problems affecting the endangered rodent Microtus cabrerae are mainly related to habitat loss and fragmentation. In the present study, we examined the presence/absence of the species in a group of potentially suitable habitat patches, and then evaluated the microhabitat attributes of locations used by wild-ranging individuals within occupied patches. Our results revealed that shallow flooding and abundant grass cover appear to be important factors in determining the presence among localities, but high rush cover strongly determined the presence and habitat use by voles both among and within localities. The preservation of these microhabitat characteristics may be required to effectively ensure the occurrence and stability of M. cabrerae colonies.     

Habitat selection of the long‐nosed bandicoot,Perameles nasuta (Mammalia,Peramelidae), in a patchy urban environment

Habitat selection in an omnivorous marsupial, the long‐nosed bandicoot, Perameles nasuta Geoffroy, was investigated in an urban environment with both natural and highly modified habitats at North Head, New South Wales, Australia. Habitat use at both macro‐ and microhabitat scales was determined using live‐trapping, and P. nasuta was shown to be a habitat specialist at this site. At night, animals used open grass macrohabitats disproportionately more for foraging than other macrohabitats. Trap‐revealed macrohabitat use was supported by radiotracking three males. Lack of understorey and absence of leaf litter were the major microhabitat features affecting habitat choice, although soil type probably also had some effect. Open areas may provide a more abundant and/or accessible food supply for P. nasuta, although better manoeuvrability or increased visibility to detect predators may also be important. Diurnal nest sites, located using radiotracking, were primarily in dense scrub vegetation, often comprising introduced species of plants. The dependence of P. nasuta on: (i) dense undergrowth for diurnal nesting and temporary nocturnal sheltering; and (ii) open areas for foraging indicates the importance of conserving a mosaic of open and dense vegetation to ensure the continued persistence of this endangered population at North Head.     

Aspects of habitat selection in the mosquitofish gambusia affinis

Laboratory experiments were performed to determine behavioral preferences of Gambusia affinis for habitat variables in partitioned aquaria. The fish preferred calm water, dark-colored substrates and subsurface vegetation providing lateral concealment. Floating cover, which obstructed access to the surface, was avoided. Lack of preference for real over plastic plant cover indicates that visual rather than chemical cues are involved. These laboratory preferences correspond to the reported microhabitat distribution of G. affinis in nature and indicate the presence of an active habitat preference, as opposed to passive habitat correlation, in this species. Species-specific habitat preferences, which may be narrower where a species occurs sympatrically with its congeners, probably form part of a reproductive isolating mechanism.     

Fire refugia: The mechanism governing animal survivorship within a highly flammable plant

Identifying factors that influence the survival of individuals during disturbance is critical to understanding patterns of species reassembly within ecological communities. Although most studies of recovery of populations post‐burning acknowledge the potentially important contribution of animals surviving in situ, few have measured the effectiveness of refugia. This paper tests the hypothesis that some plants with tightly packed leaf‐bases provide a refuge for invertebrates during fire (even when the plants themselves burn) by using the highly flammable grass tree (Xanthorrhoeaceae: Xanthorrhoea). Invertebrates were sampled from four unburnt and five experimentally burnt grass trees (Xanthorrhoea preissii Endl.). Also collected were invertebrates fleeing during burning. The dataset comprises 949 specimens, representing 81 species from 18 orders, of which 749 individuals were from unburned plants. Slaters (Isopoda), silverfish (Thysanura), spiders (Araneae) and bugs (Hemiptera) dominated assemblages of the unburnt grass trees. Despite grass trees burning at temperatures of up to 515°C, some invertebrates survived in situ. Species‐specific microhabitat preferences within the plant appeared to influence survivorship. Species collected in the crown of unburned plants were found more often alive on burnt plants than species typically inhabiting the dead skirt of decaying leaves (thatch). We contend that the mechanism causing differential mortality is fire temperature. In the dead skirt, temperatures reached 225.33 ± 66.57°C. In contrast, a region of mild temperature (25.00 ± 3.54°C) persisted throughout burning near the apical meristem (within the crown). We conclude that grass trees are a potential reservoir from which invertebrates might re‐colonize recently burnt areas. However, owing to species‐specific microhabitat preferences and differential mortality across microhabitats, the invertebrate assemblage remaining in situ will be restricted taxonomically compared with the original grass tree fauna. Moreover, different fire regimes might mediate the effectiveness of grass trees as refugia. Finally, we argue that in situ survival of invertebrates within plants with tightly packed leaf‐bases is an unrecognized global phenomenon applicable to a wide array of plant taxa.     

Microhabitat and time-since-fire: effects on demography oF Eryngium cuneifolium (Apiaceae), a Florida scrub endemic plant

Eryngium cuneifolium Small. (Apiaceae) is a narrowly distributed endemic found only in Ceratiola ericoides (Florida rosemary)-dominated Florida scrub, a periodically burned, shrub-dominated habitat. Multivariate analyses using 22 ∗∗∗microhabitat characteristics indicated significant microhabitat and time-since-fire effects on survival, growth, and fecundity of 1287 individuals over a 4-yr period. Survival increased with distance to the nearest shrub, and plants in larger open patches had greater survival rates. Neighboring shrubs of Ceratiola ericoides and Calamintha ashei were associated with a higher mortality of E. cuneifolium than other neighboring shrub species. Survival was reduced by two-thirds over 4 yr (14% vs. 42%) for E. cuneifolium near C. ericoides. Sand accretion increased growth and fecundity. With greater time since fire, woody shrubs increasingly dominate and open patches shrink, significantly reducing survival, growth, and fecundity of E. cuneifolium. Effects were particularly dramatic between 2 and 7 yr postfire, when annual mortality increased from <10% to >30% (r = 0.74). This herbaceous species is dependent on an open habitat maintained by periodic fire. Belowground competition or allelopathy from shrubs probably restricts E. cuneifolium to recently burned, open patches within the most xeric parts of Florida scrub.     

Habitat utilization patterns of sympatric populations of Pseudomys gracilicaudatus and Rattus lutreolus in coastal heathland: A multivariate analysis

Abstract Habitat usage characteristics of two species of native murid rodents, Pseudomys gracilicaudatus and Rattus lutreolus were investigated on an area of coastal heathland at Myall Lakes National Park. A grid of 151 trap stations comprising 17 traplines was positioned across a mosaic of habitats. At each trap station 19 structural vegetation and physical variables known to affect the microdistribution of small mammals were measured. Multivariate statistical procedures identified those microhabitat variables that contribute to individual species' habitat use and habitat partitioning, and reduce potential competition for space. Cluster analysis classified trap stations into one of six habitat types that were mapped on the study area, identifying a heterogeneous assemblage of interlocking habitats. The pattern is a consequence of topographic variation on the site and, to a lesser extent, its fire history. Trapping results show P. gracilicaudatus and R. lutreolus exhibit similar macrohabitat selection, preferring topographically low habitats, with both species predominantly occupying short dense heath with dense sedge cover. The high overlap in macrohabitat use is greatly reduced when considered trap station by trap station, so that discriminant function and multiple regression analyses demonstrate marked microhabitat selection. Elevation was a highly significant variable, accounting for 41% and 27% of the variance in the habitat used by P. gracilicaudatus and R. lutreolus, respectively. This variable represents a soil moisture gradient that determines changes in the floristic and structural components of the biotic environment. Two other structural vegetation variables and vegetation height contributed 30% of the variance in P. gracilicaudatus distribution. Sedge cover was found to be significant and explained 13% of the variance in R. lutreolus distribution. Within-habitat separation was explained best with a linear combination of variables in a discriminant function, rather than by any single variable. Differential microhabitat selection, interference competition and diet separation appear to be the major factors facilitating coexistence of these two species.     

Characterization of benthic microhabitat: An experimental system for aquatic insects

An experimental system used to determine microhabitat current velocity and microhabitat selection by aquatic insects is described. The experimental system includes a microvelocity probe and a hydraulically calibrated artificial substrate. A thermistor velocity probe detects flow velocities to 0.5 m s^–1 near the surface of substrates at locations inhabited by aquatic insects. The artificial substrate was designed to provide two major habitat types, highly turbulent vortex areas and regions with unidirectional, near laminar flow. Substrate calibration and microhabitat characteristics of the substrates are demonstrated. Experimental studies of Simulium sp. location on substrates indicated that while simuliid larvae are characteristic of lotic, erosional habitats, actual microhabitats selected are governed by substantially lower flow velocity.     

Depth distributions of coral reef fishes: the influence of microhabitat structure,settlement, and post-settlement processes

Many coral reef fishes have restricted depth ranges that are established at settlement or soon after, but the factors limiting these distributions are largely unknown. This study examines whether the availability of microhabitats (reef substrata) explains depth limits, and evaluates whether juvenile growth and survival are lower beyond these limits. Depth-stratified surveys of reef fishes at Kimbe Bay (Papua New Guinea) showed that the abundance of new settlers and the cover of coral substrata differed significantly among depths. A field experiment investigated whether settling coral reef fishes preferred particular depths, and whether these depth preferences were dependent on microhabitat. Small patch reefs composed of identical coral substrata were set up at five depths (3, 6, 10, 15 and 20 m), and settlement patterns were compared to those on unmanipulated reef habitat at the same five depths. For all species, settlement on patch reefs differed significantly among depths despite uniform substratum composition. For four of the six species tested, depth-related settlement patterns on unmanipulated habitat and on patch reefs did not differ, while for the other two, depth ranges were greater on the patch reefs than on unmanipulated habitat. A second experiment examined whether depth preferences reflected variation in growth and survival when microhabitat was similar. Newly settled individuals of Chrysiptera parasema and Dascyllus melanurus were placed, separately, on patch reefs at five depths (as above) and their survival and growth monitored. D. melanurus, which is restricted to shallow depths, had highest survival and growth at the shallowest depth. Depth did not affect either survival or growth of C. parasema, which has a broader depth range than D. melanurus (between 6 and 15 m). This suggests that the fitness costs potentially incurred by settling outside a preferred depth range may depend on the strength of the depth preference.