Incompletely Protective Refuges: Selection and Associated Defences by a Lizard, Cordylus cordylus (Squamata: Cordylidae)

Despite recent interest in refuge use, refuge characteristics and their relationships to refuge-associated antipredatory defences have been relatively neglected. These topics were studied experimentally in the Cape girdled lizard, Cordylus cordylus . Lizards used crevices in novel situations under uncertain risk and when confronted by a human simulated predator. They preferred narrow crevices opening on only one side at ground level and orientated horizontally. Narrowness restricts access by predators, reduces detectability to a small visual angle and permits use of crevice-specific defences, making the lizards difficult to dislodge. Limiting openings to one side decreases detectability by reducing light, limits attacks to one direction and permits further withdrawal from any opening. Crevices at ground level confer greater distance from and reduced visibility to an overhead predator. Horizontal orientation may reduce visibility to an overhead predator, but the preference could be an artefact of the greater ease of entering horizontal crevices. Pheromonal labelling of crevices by conspecific males did not influence short-term crevice choices by males. Within crevices, C . cordylus pressed their dorsal surfaces against crevice roofs, pushing upward with their legs. They also positioned their tails to block access by predators to their bodies. We discuss these and related crevice-associated defences in other animals.     

Structures of five antibiotics bound at the peptidyl transferase center of the large ribosomal subunit

Structures of anisomycin, chloramphenicol, sparsomycin, blasticidin S, and virginiamycin M bound to the large ribosomal subunit of Haloarcula marismortui have been determined at 3.0A resolution. Most of these antibiotics bind to sites that overlap those of either peptidyl-tRNA or aminoacyl-tRNA, consistent with their functioning as competitive inhibitors of peptide bond formation. Two hydrophobic crevices, one at the peptidyl transferase center and the other at the entrance to the peptide exit tunnel play roles in binding these antibiotics. Midway between these crevices, nucleotide A2103 of H.marismortui (2062 Escherichia coli) varies in its conformation and thereby contacts antibiotics bound at either crevice. The aromatic ring of anisomycin binds to the active-site hydrophobic crevice, as does the aromatic ring of puromycin, while the aromatic ring of chloramphenicol binds to the exit tunnel hydrophobic crevice. Sparsomycin contacts primarily a P-site bound substrate, but also extends into the active-site hydrophobic crevice. Virginiamycin M occupies portions of both the A and P-site, and induces a conformational change in the ribosome. Blasticidin S base-pairs with the P-loop and thereby mimics C74 and C75 of a P-site bound tRNA.     

An experimental investigation into the effect of crevice availability on abundance and size-structure in a population of Littorina rudis (Maton): Gastropoda: Prosobranchia

Littorina rudis (Maton) is a common species on a wide variety of rocky shores. At Newhaven L. rudis is generally abundant on boulders, juveniles being reliant upon empty barnacle shells for shelter. The experimental addition of crevices resulted in greatly increased population densities, suggesting that crevice availability is normally a limiting factor on this shore. A relationship is demonstrated between the size of artificially provided crevices and the mean shell lengths of the populations of L. rudis they support. The relevance of crevice availability and quality, including empty barnacle shells, to the abundance and size distribution of L. rudis and other littorines is discussed.     

Experimental analysis of retreat‐site selection by thick‐tailed geckos Nephrurus milii

Abstract Although animals use habitats non‐randomly in the wild, complex correlations among environmental features mean that proximate influences on habitat selection can be identified only by experimental manipulation of potential cues. Thick‐tailed geckos Nephrurus milii are large lizards that are widely distributed through southern Australia. These nocturnally active animals typically spend daylight hours under surface rocks. We presented captive geckos with alternative retreat‐sites (rock crevices) differing in attributes potentially relevant to habitat selection. The lizards showed strong preference for shelter‐sites that enhanced thermoregulation (warm rather than cool) and that reduced the animal's vulnerability to predators (narrow crevices with small openings and not containing the scent of a predatory snake). Horizontal rather than sloping crevices were also preferred. Overall crevice size and thickness of the overlying rock did not influence retreat‐site selection in the laboratory, but could be important in the field because of their influence on thermal regimes under rocks. The present study supports the idea that nocturnal reptiles base their selection of diurnal shelters on multiple aspects related to the fitness consequences of occupancy of alternative available retreat‐sites.     

Coexistence in a sea urchin guild and its implications to coral reef diversity and degradation

Summary Coexistence between the coral reef inhabiting sea urchins Echinometra mathaei, Diadema savignyi and D. setosum was studied by comparing differences in body morphology, distribution, diet, susceptibility to predators, intra- and interspecific competition and settlement. The three species share similar diets and broad within-habitat distributions but differ in their microspatial preferences. E. mathaei is the smallest species, has the highest settlement rates and lives territorially within small burrows or crevices. D. savignyi is intermediate in size and lives frequently in intermediate size crevices or occassionally in social groups. D. setosum is the largest species and occassionally lives in large crevices or more frequently in social groups. Both Diadema have similarily low settlement rates. Competition experiments showed that E. mathaei was consistently the top competitor for crevice space. Diadema species shared larger crevices but competition occured within smaller crevices and was frequently won by the largest individual, regardless of species. D. savignyi may be the top competitor for crevice space between the Diadema species due to a reduced spine length/test size ratio which gives it a larger test for the same crevice size requirement. Predation rates were high for E. mathaei and low for both Diadema species. Coexistence is mediated by predation on the competitive-dominant while predation coupled with different body morphologies and behavior allows spatial resource partitioning of the reef's variable topography. Consequently, the three variables of predation, topographic complexity and differing body shapes create the observed species diversity. A reduction in predators due to stochastic fluctuations or from fishing pressure can lead to E. mathaei population increases and competitive exclusion of Diadema.     

Belly up: Reduced crevice accessibility as a cost of reproduction caused by increased girth in a rock‐using lizard

Costs of reproduction are any aspect of current reproduction that has the potential to reduce survivorship or reproductive output, and may include physiological costs or increased risks. Females of many species experience increased body mass, and increased girth, when gravid. Increased body mass reduces running speed and increases the cost of locomotion during pregnancy, but few studies have examined the cost of increased girth. If increased girth of gravid females reduces access to shelter from predators or the elements, increased girth could constitute a cost of reproduction. In the laboratory, we experimentally tested whether access to crevices was limited in viviparous, saxicolous female lizards (Eulamprus brachysoma), which use crevices for shelter, by measuring access to artificial crevices of known widths, and body height during and after pregnancy. Gravid E. brachysoma had significantly greater body height (11.2% on average), and as a result were forced to use significantly wider crevices (18.4% wider on average) than post‐parturition. Females with larger clutch sizes had wider mid‐bodies and required larger crevices. Control females, which were not gravid at either time of testing, showed no significant change in the size of crevice they could enter over time. If access to narrow crevices provides advantages such as protection from predators, or is important for thermoregulation, then gravid females may suffer a cost of reproduction because their access to narrower crevices is limited.     

Population variation in spawning current speed selection in the blacktail shiner,Cyprinella venusta (Pisces: Cyprinidae)

Synopsis Many stream fishes lay demersal eggs. Some species simply scatter the eggs across the substrate, but many place the eggs in protected sites such as crevices (e.g., Cyprinidae:Cyprinella). Eggs laid in crevices may experience increased development times or lower embryo survival due to reduced water exchange rates compared to more open egg deposition sites. If so, the presence of an appreciable current near the crevice may be important for insuring water exchange. However, currents that are too strong may prevent complete insemination or prevent the eggs from adhering to the substratum before they are washed from the crevice. Thus, current speed preferences for spawning may be under strong stabilizing selection. Representatives of four populations of a crevice spawning stream fish (blacktail shiner,Cyprinella venusta), representing environments ranging from a swift stream to a lake, were studied in stream tanks to evaluate female spawning current use. The observed frequency of use of spawning currents was significantly different from random in all populations. Low current speeds were generally avoided, and a speed of ca. 30 cm sec^–1 was used most often. However, significant differences were observed among populations, and the variation correlated well with the apparent range of current speeds found in the four environments. Although other possible explanations cannot be dismissed, the results are consistent with the hypothesis that the populations have become adapted to their local current regimes.Authorship in order of seniority     

Movement of the Southern Australian abalone Haliotis laevigata in relation to crevice abundance

The long-term movement of the abalone Haliotis laevigata was measured at three sites in Waterloo Bay, South Australia, characterized by differing amounts of available crevice space. Movement was negligible at a site where crevices were abundant, but extensive at a site without crevice space. At a site of limited available crevice space, the amount of movement increased as available crevice space decreased. In addition, the extent of movement was size dependent and movement was oriented in the direction of the approaching swell. Current experimental designs for measuring natural mortality of abalone depend heavily on assumptions about their movement. An understanding of this behaviour is thus a prerequisite for designing experiments to measure the mortality rates of abalone or other sedentary animals. The natural mortality rate at one site was estimated to be 0.59 (s.e.m. = 0.02, i.e. survival = 56% per year) and the disappearance rate (natural mortality and emigration) was independent of sex.     

Refuge utilization and preferences between competing intertidal crab species

Many invertebrates avoid predation risk by seeking and defending refuges that can be in limited supply, producing strong intra- and inter-specific interference competition. Previous experimental studies in central Chile demonstrated that interference competition for refuges is the primary factor driving habitat segregation between the predatory crabs Acanthocyclus gayi and A. hassleri, with the latter species monopolizing galleries inside mussel beds in the mid intertidal zone and limiting A. gayi to rock crevices. Yet, habitat partitioning between rival species can result from differences in habitat preferences and not solely from interference interactions. Moreover, since A. gayi is also known to shelter in turf-forming algae (predominantly Gelidium), which dominates extensive areas in the low intertidal zone, among-sites variation in the turf morphology and abundance could modify habitat preferences and the pattern of inter-specific interactions. We experimentally evaluated refuge habitat preferences of individual, similarly-sized adult A. gayi and A. hassleri in the laboratory, comparing choice patterns across multiple trials with paired combinations of the main refuge types commonly used by crabs in the field: a) mussel galleries, b) rock crevices, c) short algal turf, and d) tall algal turf. Our results showed that both species display a strong ranking of preferences for some refuge habitats over others. In general, mussel galleries were the preferred refuge type for both crab species, but their preference rankings changed depending on turf morphology. When turf was short, A. hassleri and A. gayi made identical refuge choices, strongly preferring mussel galleries over crevices and these over the short turf. In contrast, when the turf was tall A. gayi selected equally the tall turf or mussel galleries, and these were strongly preferred over crevices. A. hassleri, on the other hand, largely ignored tall turf and kept the highest preference for mussel galleries. A field experiment in which crabs were offered to foraging birds demonstrated that A. hassleri is more susceptible than A. gayi to predation by kelp gulls when outside refuges. Differences in patterns of coloration between crabs may underlie between-species differences in predation susceptibility and their habitat choices. These results suggest that the among-site differences in turf morphology (height and shape of fronds), which is largely driven by varying intensity of upwelling, could affect crab preferences for refuge habitats and the relative importance of inter-specific interference competition.     

Enhancement of juvenile Caribbean spiny lobsters: an evaluation of changes in multiple response variables with the addition of large artificial shelters

Shortage of natural crevice shelters may produce population bottlenecks in juvenile Caribbean spiny lobsters (Panulirus argus), a socially gregarious species. We conducted a field experiment to test enhancement of a local population of juvenile P. argus with the addition of artificial shelters (“casitas”) that mimic large crevices (1.1 m² in surface area and 3.8 cm in height). Changes in density and biomass of juvenile lobsters 15–50 mm carapace length (CL) were assessed with a multiple before-after control-impact (MBACI) analysis. Separate analyses were also conducted on small (15–35 mm CL) and large (35.1–50 mm CL) juveniles to assess size-related effects. First, we carried out 13 lobster surveys on nine fixed 1-ha sites over a shallow reef lagoon (“before” period). Then, we deployed ten casitas in each of five sites and left four sites as controls, and conducted 22 further surveys (“after” period). Deployment of casitas resulted in a sixfold increase in juvenile density (76% contributed by small and 24% by large juveniles) and a sevenfold increase in biomass (40 and 60%, respectively). Capture–recapture results revealed that enhancement was achieved not by promoting individual growth but by increasing survival, persistence, and foraging ranges of small and large juveniles. Casitas both mitigated shortage of natural shelter and increased sociality, allowing for cohabitation of smaller, more vulnerable juveniles with larger conspecifics that have greater defensive abilities. Casitas may help enhance local populations of juvenile P. argus in Caribbean seagrass habitats, typically poor in natural crevice shelters. The use of MBACI and the simultaneous assessment of multiple interrelated response variables may be a powerful analytical approach to test shelter limitation in other species and to examine the function of structural habitat in other systems.     

Edaphic and allelochemic aspects of intertidal crevice sediments in relation to habitat recognition by Thalassotrechus barbarae (Horn) (Coleoptera: Carabidae)

Extracts of crevice sediment volatiles were attractive to the intertidal carabid beetle Thalassotrechus barbarae (Horn) suggesting that these compounds serve as chemical cues to adults selecting crevices to enter after their nocturnal excursions on nearby rock surfaces. In this respect, T. barbarae resembles saline lakeshore carabid beetles that recognize their habitats by perceiving volatiles emitted by microorganisms. Analyses of the crevice sediments showed that they were predominantly sand with enough clay (mean 2.7%) and organic matter (mean 1.7%) to have a mean cation exchange capacity of 9.13 cmol(+)·kg⁻¹. Sodium adsorption ratios (mean 40.3), exchangeable sodium percentages (mean 35.9), and electrical conductivities (mean 18.8 dS·m⁻¹) of the sediment solution, as well as cation exchange capacity, were similar to those of sediments collected from nine carabid beetle habitats on the shores of saline lakes and ponds, indicating close ecological affinities between marine and non-marine littoral carabids.     

Initial Effects of Wildfire on Freshwater Turtle Nesting Habitat

Natural wildfire regimes are important for ecosystem succession but can have negative ecological effects depending on fire characteristics. A portion of a granite rock barrens landscape that extends along the eastern shoreline of Georgian Bay, Lake Huron to eastern Ontario, Canada, burned in 2018 during a wildfire that affected >11,000 ha. This landscape is a biodiversity hotspot providing habitat for many species at risk where freshwater turtles nest in soil deposits in cracks and crevices in the bedrock dominated by moss (Polytrichum spp.) and lichen (Cladonia spp.) cover. To assess the initial effect of wildfire on freshwater turtle nesting habitat, we measured soil depths and estimated moss, lichen, and vascular plant cover at 2 morphology types (crevice, flat) in burned and unburned areas of the landscape. The probability that burned flat plots supported soil was near zero; the burned flat plots had 98% less soil volume compared to unburned flat plots. Although crevices were more resistant to soil loss, burned crevices still had a 15% lower probability of having soil and 35% less soil volume compared to unburned crevice plots. We estimated nest site availability by calculating the number of locations with shallow (5–10 cm), intermediate (10–20 cm), and deep (>20 cm) soils required for a small (5 cm × 5 cm) or medium (10 cm × 10 cm) nest chamber. Overall, the burned open rock barrens had 71–73% fewer sites with suitable soil depth and volume for a nest chamber of either size. Furthermore, burned plots had almost no lichen and moss cover but were dominated by bare soil, forbs, and jack pine (Pinus banksiana) seedlings. Although the loss of tree cover in previously forested areas may increase nest site availability for freshwater turtles in newly open areas, we suggest that organic soil combustion and soil erosion may require restoration activities in the post-fire landscape to support successful nesting of at-risk turtles. © 2020 The Wildlife Society.     

To run or hide? Age-dependent escape behaviour in the common flat lizard (Platysaurus intermedius wilhelmi)

Flat lizards Platysaurus intermedius wilhelmi occur on small discrete rock outcrops in Mpumalanga Province, South Africa. These rock outcrops are structurally simple and this, combined with the lizard's behaviour (ambush foraging in the open), make them ideal for field studies of anti-predatory behaviour. Lizards were approached in the field and how escape behaviour was influenced by habitat and age-sex class was recorded. Juveniles ( c. 4 months of age) responded quite differently to an approaching human 'predator' compared with adult males and females (which responded similarly). Compared to adults, juveniles allowed a closer approach by the investigator; took longer to find a refuge and therefore fled further; were more likely to remain visible in the open and maintain visual contact with the investigator; and more likely to flee into vegetation when given the opportunity to take refuge in a crevice. We suggest that because a greater suite of predators (including arthropods living in rock crevices) feed on small juvenile lizards, this may affect their choice of refuge and result in the avoidance of crevices when chased. Finally, because juveniles were frequently found on small rock outcrops, the influence of rock outcrop area on anti-predatory behaviour was tested. Escape behaviour (time to refuge) was independent of rock outcrop area.     

The effect of natural variation in substrate architecture on the survival of juvenile bicolor damselfish

Coral reefs offer settling fish larvae a spatial mosaic of microhabitats that differ not only in structural complexity but also in the abundance and diversity of predators. This paper provides evidence that interactions between predators and prey are causally linked to variation in relevant architectural characteristics of natural substrates. Juvenile bicolor damselfish, Stegastes partitus, experienced greater mortality on Montastrea annularis boulder coral than on piles of Porites porites coral rubble. This pattern was consistent on both back and fore reef habitats. Architectural differences, variable encounter rates with predators, and access to different food sources all contributed to higher mortality. Spatial and structural differences among refuges provided by the two substrates were most important in affecting survival. Porites rubble contained almost three times the number of crevices (39.9 vs. 14.3 crevices m^–2AM²), had a much smaller mean crevice size (15.9 vs. 48.5 cm²), and had a more complex internal structure than Montastrea coral. Survival of juvenile fish living on Montastrea coral was positively correlated with crevice density (r² = 0.41) whereas survival of fish living on Porites rubble was negatively correlated with crevice size (r² = 0.43). These patterns were evident on fore reef habitats only, whereas on back reef habitats no clear patterns emerged. The effect of these natural differences in architecture on mortality rates of juvenile S. partitus was experimentally tested in the field using a combination of natural and artificial substrates. As the number of large shelters was increased I found that the density of potential predators increased but the survival rates of juvenile bicolor damselfish declined. These results highlight the importance of structural architecture among common reef substrates in affecting predator-prey relationships and in determining the survivorship and small-scale distribution patterns of juvenile reef fish.     

The settlement behaviour of planulae larvae of the hermatypic coral Favia fragum (Esper)

Experiments on the settlement behaviour of planulae larvae of the reef coral Favia fragum (Esper) are described. The larvae are positively phototaxic upon release but reverse this and become attracted to dark surfaces, corners, crevices, and the undersides of objects on the bottom. Clean glass surfaces were preferred to surfaces covered with biological slime but there was no preference for rough against smooth surfaces. There was clear evidence of gregarious settlement behaviour, the planulae being able to recognize both adult colonies and previously settled juveniles. A distinction was made between crawling and swimming larvae, and the consequences of their differences in behaviour on the spatial distribution of Favia on reef are discussed. Settlement behaviour of Favia is similar in many respects to that of the Pacific reef coral Pocillopora damicornis (Dana) but is distinguished by gregarious settlement and by a preference for clean surfaces over surfaces covered with biological slime.     

Germination as a determinant of seedling distributions among natural substrates in Picea engelmannii (Pinaceae) and Abies lasiocarpa (Pinaceae)

In Rocky Mountain (USA) subalpine forests, seedlings of Picea engelmannii (Engelmann spruce) colonize logs more frequently than seedlings of its codominant associate Abies lasiocarpa (subalpine fir). We hypothesized that spruce germinates more readily on logs than fir, perhaps because small spruce seeds are more likely to lodge in log crevices than larger fir seeds. Our objectives were to test this hypothesis and compare both species' germination among several substrates to assess germination influences on natural seedling distributions. Spruce and fir seeds were sown on field-collected logs, litter, and soil in the greenhouse and monitored for 36 d. To test the crevice hypothesis, seeds were either scattered on logs or wedged into crevices, assuming that if both species were placed in crevices, interspecific germination differences on logs would decrease. Spruce mean germination percentages were significantly greater than fir's in all substrate treatments except when seeds were wedged in log crevices. The difference in means between the two log treatments was greater for fir (68%) than spruce (21%). Spruce germinated more rapidly than fir on all substrates. We suggest that large seed size reduces fir's success in colonizing logs, and that germination and establishment factors interact to determine natural seedling distributions for these subalpine conifers.     

Detection of bacterial cytotoxic activities from water-damaged ceiling tile material following incubation on blood agar

Samples of ceiling tiles with high levels of bacteria exhibited cytotoxic activities on a HEP-2 tissue culture assay. Ceiling tiles containing low levels of bacterial colonization did not show cytotoxic activities on the HEP-2 tissue culture assay. Using a spread plate procedure on blood agar plates, the levels of bacteria colonizing portions of cellulosic indoor ceiling tiles were easily identified. Levels of bacteria measured by this simple procedure may be a good indicator of microbial colonization of indoor building materials especially in the case of water damage. We suggest that bacterial levels above 150 CFU g⁻¹ of ceiling tile material indicate colonization has occurred. Received 24 December 1998/ Accepted in revised form 26 March 1999     

UV-Green Iridescence Predicts Male Quality during Jumping Spider Contests

Animal colour signals used in intraspecies communications can generally be attributed to a composite effect of structural and pigmentary colours. Notably, the functional role of iridescent coloration that is ‘purely’ structural (i.e., absence of pigments) is poorly understood. Recent studies reveal that iridescent colorations can reliably indicate individual quality, but evidence of iridescence as a pure structural coloration indicative of male quality during contests and relating to an individual’s resource-holding potential (RHP) is lacking. In age- and size-controlled pairwise male-male contests that escalate from visual displays of aggression to more costly physical fights, we demonstrate that the ultraviolet-green iridescence of Cosmophasis umbratica predicts individual persistence and relates to RHP. Contest initiating males exhibited significantly narrower carapace band separation (i.e., relative spectral positions of UV and green hues) than non-initiators. Asymmetries in carapace and abdomen brightness influenced overall contest duration and escalation. As losers retreated upon having reached their own persistence limits in contests that escalated to physical fights, losers with narrower carapace band separation were significantly more persistence. We propose that the carapace UV-green iridescence of C. umbratica predicts individual persistence and is indicative of a male’s RHP. As the observed UV-green hues of C. umbratica are ‘pure’ optical products of a multilayer reflector system, we suggest that intrasexual variations in the optical properties of the scales’ chitin-air-chitin microstructures are responsible for the observed differences in carapace band separations.     

Bubble-Induced Cave Collapse

Conventional wisdom among cave divers is that submerged caves in aquifers, such as in Florida or the Yucatan, are unstable due to their ever-growing size from limestone dissolution in water. Cave divers occasionally noted partial cave collapses occurring while they were in the cave, attributing this to their unintentional (and frowned upon) physical contact with the cave walls or the aforementioned “natural” instability of the cave. Here, we suggest that these cave collapses do not necessarily result from cave instability or contacts with walls, but rather from divers bubbles rising to the ceiling and reducing the buoyancy acting on isolated ceiling rocks. Using familiar theories for the strength of flat and arched (un-cracked) beams, we first show that the flat ceiling of a submerged limestone cave can have a horizontal expanse of 63 meters. This is much broader than that of most submerged Florida caves (~ 10 m). Similarly, we show that an arched cave roof can have a still larger expanse of 240 meters, again implying that Florida caves are structurally stable. Using familiar bubble dynamics, fluid dynamics of bubble-induced flows, and accustomed diving practices, we show that a group of 1-3 divers submerged below a loosely connected ceiling rock will quickly trigger it to fall causing a “collapse”. We then present a set of qualitative laboratory experiments illustrating such a collapse in a circular laboratory cave (i.e., a cave with a circular cross section), with concave and convex ceilings. In these experiments, a metal ball represented the rock (attached to the cave ceiling with a magnet), and the bubbles were produced using a syringe located at the cave floor.