Effects of temperature on activity,food consumption rates,and gut passage times of seaweed-eating<Emphasis Type="Italic"> Tegula</Emphasis> species (Trochidae) from California

Previous feeding studies on herbivorous marine snails rarely have focused on temperature effects on food intake. If temperature affects food intake, ectothermic snails may experience difficulty obtaining sufficient nutritional resources, limiting their ability to sustain populations at suboptimal temperatures. We hypothesized that the feeding responses of Tegula species would correspond with temperatures characteristic of their geographic distributions. We determined activity, consumption rates, and gut passage times at 11°C, 15°C, 19°C, and 23°C for three Tegula species with distinct thermal distributions: T. brunnea (cold water), T. aureotincta (warm water), and individuals from warm- and cold-water populations of T. funebralis, a broadly distributed species. Activity and consumption rates of T. aureotincta increased with increasing temperature, but were highest for T. brunnea at 19°C, a temperature rarely achieved in habitats occupied by this species, and lowest at 11°C. Warm-water T. funebralis showed significantly lower activity and consumption rates at 11°C, whereas cold-water T. funebralis consumed food fastest at 15°C and were most active at 23°C. Temperature affected gut passage time only in T. aureotincta. These data suggest that temperature might influence the northern limit of T. aureotincta by affecting activity and food consumption rates. T. brunneas activity and ability to consume food were not hindered by warmer temperatures despite the present day restriction of this species to colder waters. Also, widely separated (>300 km) T. funebralis populations may be adapted to regional conditions based on the different temperature responses of northern and southern snails.Communicated by P.W. Sammarco, Chauvin     

Spawning and larval development of the black turban snail Tegula funebralis (Prosobranchia: Trochidae)

An understanding of spawning and larval development can be fundamental to interpreting the abundance, distribution, and population structure of marine invertebrate taxa. Tegula funebralis (A. Adams, 1855), the black turban snail, has been the focus of numerous ecological studies on the Pacific coast of North America. To date, there are only conflicting and anecdotal reports of spawning, and there is no information on larval or juvenile development for this conspicuous and abundant species. On 19 September 1995, two individuals of T. funebralis were observed free-spawning gametes into seawater in tanks at the Oregon Institute of Marine Biology. Embryos and larvae were subsequently reared to metamorphosis and beyond. Development was pelagic and similar to development described for other trochids, and larvae were observed not to feed at any stage. Larvae began to metamorphose at 5.7 to 6.7 d and settled at 260 μm shell length. Juveniles grew ≃ 10 μm in shell length per day and appeared to feed on detritus. Juveniles lacked some adult diagnostic shell characters, including two columellar nodes and a closed umbilicus. In the field, small (<3 mm) juveniles occurred in the adult habitat on all sampling dates between October and March. Small juveniles were found only under rocks and were most abundant under rocks partially buried in coarse sand, suggesting that juveniles may utilize a specific microhabitat within the adult T. funebralis habitat. Received: 7 October 1996 / Accepted: 17 October 1996     

Influence of habitat on abundance and size structure of a large temperate-reef fish, Achoerodus viridis (Pisces: Labridae)

The relationship between densities of Achoerodus viridis (Pisces: Labridae) and reef habitats at various localities within New South Wales (NSW), Australia was examined. Types of habitats were quantified at inner, mid and outer estuarine locations in each of two estuaries (Botany Bay and Port Jackson) to determine whether purported patterns of movement from estuaries could be related to differences in habitat. Although the same types of habitat were generally found at all locations, differences in the proportion of habitat types were found between shallow and deep reefs and among inner, mid and outer estuarine locations for both estuaries. Shallow habitats were usually dominated by Ecklonia radiata, turf and/or fringe habitat in Botany Bay, whereas deep sites were generally dominated by urchin-grazed barrens habitat and, sometimes, sponge- and ascidian-dominated deep reef. Shallow sites in Port Jackson were dominated by a mixture of habitats, as were deep reefs at mid-estuarine locations. Other deep reefs in Port Jackson were dominated by E. radiata (inner estuarine) or barrens (outer estuarine) habitat. Thus, patterns of habitat cover were not consistent between estuaries and numbers of fish could not be related to proportional representation of habitat on reefs along estuarine gradients. Univariate and multivariate analyses showed that there was little evidence that any size class of fish was correlated with the proportional representation of a particular habitat or group of habitats. Counts of fish that focused on barrens and E. radiata forest habitats over a period of 10 yr showed that similar numbers and all sizes of fish were found in the two types of habitat. Greater numbers of small fish were, however, found in the E. radiata forest habitat than in the barrens habitat. Estimates of abundance along the coast of NSW (100s to 1000 km) in a range of habitats (e.g. ascidian-dominated reef, kelp forest, urchin-grazed barrens) showed that there was no indication that a particular habitat consistently had greater numbers of A. viridis than other habitats. Therefore, A. viridis of a range of sizes appears to be flexible in its use of habitats on reefs. Received: 24 December 1997 / Accepted: 23 June 1998     

A neutral DNA marker suggests that parallel physiological adaptations to open shore and salt marsh habitats have evolved more than once within two different species of gastropods

Local adaptation is an important mechanism generating physiological diversity and can be especially pronounced in species with restricted dispersal and gene flow such as direct developing snails of the genus Littorina. We compared physiological responses to salinity and desiccation stress in two co-occurring species of northeastern Pacific Littorina (L. subrotundata and L. sitkana) with salt marsh and open shore ecotypes. The animals from salt marsh populations were significantly more tolerant to low salinities and significantly less resistant to desiccation stress than their open shore counterparts. The lower resistance to desiccation in salt marsh animals was not associated with a higher rate of water loss during air exposure or with lower body water reserves, but instead reflected a lower tolerance to high salinities. These habitat-related physiological differences occurred in parallel in the two studied species of Littorina and persisted after prolonged laboratory acclimation, suggesting that they may reflect selection for markedly different local optima in the salt marsh habitats than in the open shore habitats. We used a neutral polymorphic nuclear DNA marker (intron of aminopeptidase N) to estimate the level of gene flow between the populations from different habitats and found isolation by distance regardless of the habitat from which the snails were collected. Our molecular data suggest that physiological cohesiveness of ecotypes can arise despite different genetic backgrounds, and could potentially be due to parallel evolution of convergent phenotypes in similar habitats.Communicated by O. Kinne, Oldendorf/Luhe     

Some beavioral characteristics related to food supply and soil texture of burrowing habitats observed on Uca vocans vocans and U. lactea perplexa

From the observations on two species of fiddler crabs, droving is conspicuous in Uca vocans vocans but does not occur in U. lactea perplexa. U. vocans come down to feed at the water's edge with a richer food supply and fine substrate particles. Droving is advantageous to increase feeding efficiency in this species which holds burrows on a relatively sandy habitat where the food supply is less rich. In U. lactea, which burrows at higher levels, on the other hand, the food supply and soil texture of their burrow area are not significantly different from those at the water's edge. Home range relationships of both species are consistent with the occurrence or not of droving behavior and depend on food productivity of burrowing habitats.     

Behaviour moderates climate warming vulnerability in high-rocky-shore snails: interactions of habitat use, energy consumption and environmental temperature

High-rocky-shore intertidal animals are predicted to be generally more vulnerable to climate warming than lower-shore species, because their thermal tolerances lie closer to maximum environmental temperatures (T _e). However, this prediction is based on taxonomically and ecologically limited information. The present study investigated the effect of habitat use on climate warming vulnerability of the tropical high-shore snail, Echinolittorina malaccana (from Brunei Darussalam, 5°N), which aestivates in sun-exposed or shaded habitats. The thermal regimes of these habitats differed vastly, but snails showed similar daily energy consumption in either habitat, due to temperature-insensitive metabolism (TIM) between 35 and 46 °C in the sun-resting snails. However, maximum T _e values in the shade and the sun were 35 and 46 °C, respectively, suggesting that sun-resting snails, which presently experience temperatures near the incipient lethal temperature range (46–56 °C), should be more threatened by further warming than shade-resting snails, which have an 11 °C ‘safety margin’. Thus, vulnerability of high-shore species to climate warming could be moderated by availability of shaded habitat, making predictions for these species more complex than previously realized.     

No evidence for homeoviscous adaptation in intertidal snails: analysis of membrane fluidity during thermal acclimation, thermal acclimatization, and across thermal microhabitats

Many eurythermal organisms alter composition of their membranes to counter perturbing effects of environmental temperature variation on membrane fluidity, a process known as homeoviscous adaptation. Marine intertidal gastropods experience uniquely large thermal excursions that challenge the functional integrity of their membranes on tidal and seasonal timescales. This study measured and compared membrane fluidity in marine intertidal snail species under three scenarios: (1) laboratory thermal acclimation, (2) thermal acclimatization during a hot midday low tide, and (3) thermal acclimatization across the vertical intertidal zone gradient in temperature. For each scenario, we used fluorescence polarization of the membrane probe DPH to measure membrane fluidity in individual samples of gill and mantle tissue. A four-week thermal acclimation of Tegula funebralis to 5, 15, and 25°C did not induce differences in membrane fluidity. Littorina keenae sampled from two thermal microhabitats at the beginning and end of a hot midday low tide exhibited no significant differences in membrane fluidity, either as a function of time of day or as a function of thermal microhabitat, despite changes in body temperature up to 24°C within 8 h. Membrane fluidities of a diverse group of snails collected from high, middle, and low vertical regions of the intertidal zone varied among species but did not correlate with thermal microhabitat. Our data suggest intertidal gastropod snails do not exhibit homeoviscous adaptation of gill and mantle membranes. We discuss possible alternatives for how these organisms counter thermal excursions characteristic of the marine intertidal zone.     

Echolocation behavior of the bat <Emphasis Type="Italic">Vespertilio murinus</Emphasis> reveals the border between the habitat types “edge” and “open space”

We test the hypothesis that echolocation behavior can be used to find the border between bat habitats. Assuming that bats react to background targets in “edge space” but not in “open space”, we determined the border between these two habitat types for commuting individuals of the parti-colored bat Vespertilio murinus. We recorded sequences of bats’ echolocation signals while they flew parallel to the walls of large buildings and to the ground and determined the signals’ average bandwidth, duration, and pulse interval. These parameters varied systematically with the estimated horizontal and vertical distances between the bats and the background. A distinct effect of horizontal distance to the background on echolocation behavior was found for horizontal distances of less than 6 m, thus indicating the border between edge and open space. Only a few bats flew at vertical distances below 5 m. However, enough passages at vertical distances of 5 m and above indicated that the vertical border is somewhere below a distance of 5 m. Within edge space, V. murinus reacted to the background by reducing signal duration, increasing bandwidth at closer distances, and often emitting one signal per wing beat. In open space, signal parameters did not vary as a function of distance to the background. There, V. murinus emitted the longest signals with the narrowest bandwidth and often made one or two wing beats without emitting a pulse. With our data we support with statistical methods the hypothesis that echolocation behavior reveals the border between the habitat types “edge” and “open space”.     

Ecology and characteristics of bdellovibrios from three tropical marine habitats

Marine bdellovibrios have not previously been reported from the southern hemisphere, and knowledge of their occurrence in marine ecosystems is rudimentary. This study examined quantitative and qualitative aspects of bdellovibrios parasitic to the bacterium Vibrio alginolyticus at each of three representative tropical marine habitats of the Great Barrier Reef. Bdellovibrios were found in the water column throughout a 12 mo period from May 1992 at a sandy beach, a mangrove and a fringing coral reef. Their abundance was correlated with water temperature (P<0.001) and was highest in summer, lowest in winter and intermediate in spring and autumn. Over the sampling period, bdellovibrios were most abundant at the mangrove habitat (36.6 ml^-1) and least abundant at the reef (9.5 ml^-1), but there was substantial variability in numbers at all habitats among seasons and months of the year. On some occasions no bdellovibrios were found in replicate samples from the beach and reef habitats, while on others the maximum detectable by the method used (180 ml^-1) was sometimes found at the beach and mangrove habitats. Bdellovibrios within each habitat were uniformly distributed among sampling sites (P>0.05). They were more abundant in sub-surface than bottom waters in summer, but the reverse occurred in winter. Midwater samples usually had least bdellovibrios. Bdellovibrio numbers were significantly correlated with those of potential host bacteria—colony-forming bacteria at all habitats and total bacteria at the beach and reef habitats. Strain characteristics, primarily based on host range, indicated qualitative differences in bdellovibrio populations among habitats. Pseudomonas atlantica, P. aeruginosa, P. marina, Cytophaga marinoflava, Vibrio gazogenes, V. mimicus and a Spirillum-like bacterium were not parasitised by bdellovibrios from any habitat. Of the other 25 Vibrio spp. tested, most were parasitised by the majority of bdellovibrio strains from each habitat. Strain differences were principally with respect to parasitism of non-Vibrio bacteria. All strains required Na⁺ and grew at 35°C, but some failed to grow at 15°C.     

The indirect effects of eutrophication on habitat choice and survival of fish larvae in the Baltic Sea

The structure of the habitat is usually crucial for growth and survival of young life stages. Presently, some nursery areas of fish larvae are changing due to eutrophication, e.g. due to enhanced growth of ephemeral filamentous algae at the expense of perennial species. We studied the influence of two habitats, one with filamentous algae (Cladophora glomerata) and the other with bladder wrack (Fucus vesiculosus), on habitat choice of pike larvae (Esox lucius) in the absence/presence of a predator or a competitor. We further tested whether the habitat choice is adaptive in increasing survival under predation threat. In contrast to expectations, pike larvae preferred the habitat with ephemeral filamentous algae to the bladder wrack, thriving in clean waters, independent of the presence/absence of both predator/competitor. In addition, the survival of the larvae was higher in the filamentous algae in the presence of predators, which suggested that the habitat preference of the larvae was adaptive. The structure of the bladderwrack habitat was probably too open for newly hatched larvae, which implies that F. vesiculosus and other large brown algae are not as important refuges for young larvae as previously thought.     

Habitat use and group size of pied cormorants (<Emphasis Type="Italic">Phalacrocorax varius</Emphasis>) in a seagrass ecosystem: possible effects of food abundance and predation risk

Both food abundance and predation risk may influence habitat use decisions. However, studies of habitat use by birds in marine environments have focused only on food abundance. I investigated the possible influences of food abundance and predation risk from tiger sharks (Galeocerdo cuvier) on habitat use by pied cormorants (Phalacrocorax varius) over two spatial scales and on cormorant group size. Cormorants were usually solitary, but group size was highest in shallow habitats during months when shark density was low. Regardless of season, cormorant density within shallow habitats was higher over seagrass than sand, and cormorants were distributed between these two microhabitats proportional to prey density. Therefore, cormorants appear to respond to prey abundance at a relatively narrow spatial scale (i.e., tens of meters). At the habitat-patch scale (~1 km), the density of cormorants and their prey (teleosts) was higher in shallow habitats than in deep ones, but the density of cormorants was influenced by an interaction between water temperature (i.e., season) and habitat. There was decreased use of shallow habitats as water temperature, and the density of tiger sharks, increased. When shark density was low, cormorants were distributed across habitats roughly in proportion to the abundance of fish, suggesting that cormorants respond to food abundance at the scale of habitat patches. However, as shark abundance increased, the relative density of cormorants dropped in the dangerous shallow habitats such that there was a greater density of cormorants relative to their food in deep habitats when sharks were abundant. This suggests that pied cormorants trade-off food and risk by accepting lower energetic returns to forage in safer habitats. This study provides the first evidence that marine habitat selection by birds may be influenced by such a trade-off, and provides further evidence that tiger sharks are important in determining habitat use of their prey and mediating indirect interactions within Shark Bay.Communicated by P. W. Sammarco, Chauvin     

The chemosensory ability of the predatory leech Whitmania laevis (Arhynchobdellida: Haemopidae) for prey searching

Although prey-detecting and searching abilities of predatory leeches of rhynchobdellid or the Erpobdelliformes of arhynchobdellid species have been studied in the past, hirudiniformes leeches are rarely mentioned. In this study, we investigated the chemosensory ability for prey-detecting and searching in Whitmania laevis, a hirudiniformes species that mainly preys on freshwater snails, and examined if such ability aided in their prey selection. Five sympatric snail species, i.e., apple snail Pomacea canaliculata, thiarid snail Thiara tuberculata, viviparid snail Sinotaia quadrata, ear pond snail Radix auricularia swinhoei and tadpole snail Physa acuta were used as prey. Our results showed that W. laevis has the chemosensory ability to detect the waterborne odors of snails. However, they follow the snails by their mucus trails, and not by the odor that the snails leave in the water. Of these five snail species, W. laevis only followed the trails of the thiarid snails, ear pond snails and tadpole snails, and did not show a different response to the trails produced by snails of different sizes. Our results suggest that W. laevis can use waterborne odors to detect the existence of prey. They rely on mucus trails to follow their preferred prey, but do not distinguish between snails of a preferred size by their mucus trails. In addition, when following the trail of a preferred snail, W. laevis exhibits a newly described searching behavior, i.e., head tapping, and may use it to locate a snail trail and increase its probability of finding the trail-laying snail nearby.     

Spatial repartition and ontogenetic shifts in habitat use by coral reef fishes (Moorea,French Polynesia)

This study explores the extent to which ontogenetic habitat shifts modify spatial patterns of fish established at settlement in the Moorea Island lagoon (French Polynesia). The lagoon of Moorea Island was divided into 12 habitat zones (i.e. coral seascapes), which were distinct in terms of depth, wave exposure, and substratum composition. Eighty-two species of recently settled juveniles were recorded from March to June 2001. Visual censuses documented changes in the distribution of juveniles of each species over time among the 12 habitats. Two patterns of juvenile habitat use were found among species. Firstly, some species settled and remained in the same habitat until the adoption of the adult habitats (i.e. recruitment; e.g. Chaetodon citrinellus, Halichoeres hortulanus, Rhinecanthus aculeatus). Secondly, others settled to several habitats and then disappeared from some habitats through differential mortality and/or post-settlement movement (e.g. 65–70 mm size class for Ctenochaetus striatus, 40–45 mm size class for Epinephelus merra, 50–55 mm size class for Scarus sordidus). A comparison of the spatial distribution of juveniles to that of adults (61 species recorded at both stages) illustrated four patterns of subsequent recruitment in habitat use: (1) an increase in the number of habitats used during the adult stage (e.g. H. hortulanus, Mulloidichthys flavolineatus); (2) a decrease in the number of habitats adults used compared to recently settled juveniles (e.g. Chrysiptera leucopoma, Stethojulis bandanensis); (3) the use of different habitat types (e.g. Acanthurus triostegus, Caranx melampygus); and (4) no change in habitat use (e.g. Naso litturatus, Stegastes nigricans). Of the 20 most abundant species recorded in Moorea lagoon, 12 species modified the spatial patterns established at settlement by an ontogenetic habitat shift.Communicated by T. Ikeda, Hakodate     

Patch use and vigilance by sympatric lemmings in predator and competitor-driven landscapes of fear

Prey living in risky environments can adopt a variety of behavioral tactics to reduce predation risk. In systems where predators regulate prey abundance, it is reasonable to assume that differential patterns of habitat use by prey species represent adaptive responses to spatial variation in predation. However, patterns of habitat use also reflect interspecific competition over habitat. Collared (Dicrostonyx groenlandicus) and brown (Lemmus trimucronatus) lemmings represent such a system and possess distinct upland tundra versus mesic meadow habitat preferences consistent with interspecific competition. Yet, we do not know whether this habitat preference might also reflect differences in predation risk or whether the two species differ in their behavioral tactics used to avoid predation. We performed experiments where we manipulated putative predation risk perceived by lemmings by increasing protective cover in upland and meadow habitats while we recorded lemming activity and behavior. Both lemming species preferentially used cover more than open patches, but Dicrostonyx was more vigilant than Lemmus. Both species also constrained their activity to protective patches in upland and meadow habitats, but during different periods of the day. Use of cover and vigilance were independent of habitat, suggesting that both species live in a fearsome but flattened landscape of fear at Walker Bay (Nunavut, Canada), and that their habitat preference is a consequence of competition rather than predation risk. Future studies aiming to map the contours of fear in multi-prey–predator systems should consider how predation and competition interact to modify prey species’ habitat preference, patch use, and vigilance.     

Genetic and environmental effects on the growth rate of Littorina saxatilis

Transfer experiments with two morphs of Littorina saxatilis Olivi (=L. rudis) typically inhabiting exposed and sheltered localities, showed a between-morph difference in shell growth in the same type of habitat, and a withinmorph difference between exposed and sheltered environments. The former indicates a genetic difference between the two morphs, although growth rate has an environmental component as shown by the latter. Juvenile snails of the exposed morph were on average slightly larger than sheltered morph juveniles on hatching, but at 20 wk, when raised in identical environments, the sheltered morph juveniles had grown significantly larger than the exposed ones. A rise in temperature from 5° to 10°C enhanced growth rate for snails raised in the laboratory. Temperature alone could not however explain increased growth during the spring and summer in natural populations.     

Chemical mediation of invertebrate defensive behaviors and the ability to distinguish between foraging and inactive predators

When the predatory sea star Pycnopodia helianthoides was placed upstream, the sea urchin Strongylocentrotus purpuratus responded defensively by extending and opening its globiferous pedicellariae. No pedicellaria response was given in control seawater or when the sea star was downstream. The snail Tegula funebralis responded by moving up vertical surfaces when Pycnopodia helianthoides or when Pisaster ochraceus were placed upstream. When these sea stars were introduced downstream, the snail's response was not significantly different from that in control seawater. Water collected from an aquarium containing a single sea star was sufficient to trigger the response of S. purpuratus and T. funebralis; the physical presence of the sea star was not essential. This indicated that a chemical stimulus was involved, and the lack of responses when sea stars were downstream argued strongly against the possible additional involvement of visual or vibrational stimuli. S. purpuratus gave stronger pedicellaria responses to water flowing over an active Pycnopodia helianthoides than to water flowing over the same sea star when it was inactive. The significance of the ability to distinguish between actively foraging and inactive predators is discussed, and a mechanism is proposed to explain differences in the amount of stimulatory chemicals released by active and inactive sea stars.     

Ecological implications of the presence of the tube-building polychaete <Emphasis Type="Italic">Lanice conchilega</Emphasis> on soft-bottom benthic ecosystems

The common tube-building polychaete Lanice conchilega is known as a habitat structuring species and can form dense aggregations. The effects of L. conchilega on the surrounding benthic community have received little attention, especially in subtidal areas. Therefore, the presence of L. conchilega in different habitats in the North Sea and its effect on the abundance, species richness, diversity and community structure in these habitats are evaluated in the present paper, based on data from the ICES North Sea Benthos Survey of 2000. Lanice conchilega has a wide geographical distribution and a low habitat specialization, but optimally occurs in shallow fine sands. In the present study, the presence of L. conchilega resulted in a density increase and a significant (positive) correlation of the benthos density with the density of L. conchilega. Furthermore, the species richness (number of species) increased with increasing density of L. conchilega. This trend was, however, not consistent: the number of species reached more or less an asymptotic value or even decreased after reaching a critical density of L. conchilega (>500–1,000 ind/m2), as observed in shallow fine sands. The same overall pattern was detected concerning the expected number of species. The N ₁-diversity index showed similar or slightly higher values in L. conchilega patches compared to patches without L. conchilega. From the results of the community analysis, it can be concluded that the species, which were responsible for the increase of the diversity, belonged to the overall species-pool of that habitat. The effects on density and diversity differed between the four discerned habitats (shallow muddy sand, shallow fine sand, shallow medium sand and deep fine sand), and were most pronounced in shallow fine sands. These patterns can be attributed to the habitat structuring capacity of L. conchilega. The mechanisms responsible for the increase of the habitat quality in patches of L. conchilega can be summarized as (1) changes in the hydrodynamics, (2) increases of the habitat stability and oxygen supply, and (3) a creation of habitat heterogeneity in a uniform environment. In this way, L. conchilega alters the habitat characteristics and affects other organisms, and can therefore even be considered as an ecosystem engineer. In other words, L. conchilega patches are responsible for an increased habitat quality in an otherwise uniform habitat, which results in a higher survival of the surrounding benthic species.     

The effects of castration, sex ratio and population density on social segregation and habitat use in Soay sheep

We analysed 16 years of census data gathered on the island of Hirta (archipelago of St. Kilda) to investigate the effects of castration, population density, sex ratio, season and group type on habitat use and social segregation of Soay sheep. From 1978 to 1980, 72 male lambs were castrated. We used this experiment to study how a change in reproductive status could affect sociality and habitat choice of these males. Males, females and castrates were all segregated outside the rutting season in autumn. Castrates were the least segregated from females in spring and summer but were most segregated from them during the pre-rut. The more equal the sex ratios, the higher was the degree of social segregation. The three sex classes used similar habitat types, namely, Holcus agrostis, Agrostis festuca and Calluna habitats. Holcus agrostis and Agrostis festuca were top- and second-ranked in female and castrate habitat use, while Holcus agrostis and Calluna were the two top habitat types used by rams. It is unclear why males included Calluna heath habitats, but it cannot be excluded that they might have shifted their use depending on forage availability. A lack in differences in habitat use between castrates and females suggests that body size differences alone cannot be the driving factor for habitat segregation in male and female Soay sheep and that there are reasons other than body size that could motivate reproductive males to use additional habitat types, such as Calluna heath. Although habitat use shifted from one habitat type to the next between low- and high-population-density years and between seasons, there was no clear link between population density and how different groups (male, female or castrate) used these areas. We discuss effects of reproductive status, population density and sex ratio on social segregation and habitat use and suggest that these factors need to be taken into account when investigating causes of sexual segregation in ungulates.     

The effect of conspecifics on habitat selection in territorial species

Summary Despite widespread interest in habitat selection, many of the assumptions about how territorial animals choose habitats have not been tested. This study of juvenile Anolis aeneus lizards focuses on the relationship between the number of previous settlers in a habitat and the subsequent behavior of new arrivals at that habitat. Clearings containing the types of microhabitat preferred by juveniles were established in the field, several juvenile residents were allowed to establish territories in enclosures in the center of each clearing, and then naturally occurring immigrants were allowed access to the empty microhabitat surrounding the enclosures. Arrival rates and the probability of settlement were monitored on a daily basis from the day the first juveniles arrived until several days after the last juvenile had settled (=saturation). In each of seven trials, arrival rates were comparable early and late in the settlement process, and were unrelated to the degree of habitat saturation. Arrival rates did vary on a temporal basis, probably as a result of environmental factors affecting egg laying and hatching schedules, and habitats with high arrival rates saturated more quickly than those with lower arrival rates. All of the individuals arriving at the clearings did not settle, but settlers and non-settlers did not differ with respect to competitive ability, as measured by body size. The probability of settlement increased as settlement proceeded in each of seven trials, up to the day of saturation. These results refute the commonly held assumptions that prospective territory owners avoid entering relatively full habitats, and that they prefer to settle in relatively unsaturated habitats. The discussion considers why assumptions about the behavioral processes of habitat selection are so widely accepted, given the dearth of empirical information on the subject.     

Species richness patterns along environmental gradients in island land molluscan fauna

The relationship between species richness and environmental variables may change depending on habitat structure, dispersal ability, species mixing, and community adaptation to the environment. It is crucial to know how these factors regulate the environment-diversity relationship. The land molluscan fauna of the Ogasawara Islands in the West Pacific is an excellent model system to address this question because of the high species endemicity (> 90%), small area, and simple habitat structure of the islands. I examined relationships among indigenous species composition, richness, and habitat condition, and especially productivity and forest moisture on the island of Anijima. Two major communities of snails could be distinguished by detrended correspondence analysis (DCA): one group dominated in a moist habitat with high productivity, and the other group dominated in a dry habitat with low productivity. However, species richness became highest at the intermediate condition between the habitats in which the two snail communities were dominant, so that species richness showed a hump-shaped relationship with moisture and productivity. In contrast, the species richness of the snail community in the moist habitat showed a monotonically positive correlation, and that in the dry habitat showed a monotonically negative correlation with moisture and productivity. Thus, the greater species richness in intermediate moisture and productivity resulted from the ecotone effect or community overlap at the transitional areas, where faunas with different ecologies can meet in a single site. These findings suggest that hump-shaped productivity-diversity relationships in land Mollusca would reflect the ecotone effect as a result of the mixing of species adapted to either fertile habitats or sterile habitats.