Differences in feeding ecology among three co-occurring species of wrasse (Teleostei: Labridae) on rocky reefs of temperate Australia

The foraging behaviours and dietary compositions of three co-occurring labrids (Ophthalmolepis lineolatus, Notolabrus gymnogenis and Pictilabrus laticlavius), which are conspicuous on rocky reefs in temperate south-eastern Australia, were investigated between 2003 and 2005. SCUBA observations at two locations showed that the feeding intensity, and hence the associated effects of these fishes on rocky reef invertebrate prey, was temporally consistent. Relative differences in the contributions of ingested prey and use of different feeding microhabitats demonstrated that the feeding ecology differed significantly among the three species. Thus, O. lineolatus fed on proportionately higher volumes of polychaetes, polyplacophorans, marginellid gastropods (especially Austroginella sp.), bivalves and echinoids, which were sighted opportunistically in a wide selection of microhabitats, but particularly in sand/rubble. Ambush hunting was used regularly by smaller N. gymnogenis and all sizes of P. laticlavius to forage on amphipods, small decapods and small gastropods at algal bases or fronds and Diopatra dentata tubes. Amphipods were similarly important in the diet of smaller O. lineolatus. Larger N. gymnogenis foraged opportunistically over an increased reef area and made greater use of microhabitats that offered minimal prey refuge (e.g. sand/rubble, bare rock/steel) from which common prey, in particular decapods, were obtained. The significant intra- and inter-specific differences in dietary compositions, allied with differences in the use of feeding microhabitats, would facilitate co-occurrence of these three conspicuous species and contribute to maintaining high richness of labrid species in reef systems. Echinoids were regularly consumed by each species but they made a moderate contribution to the diet of only O. lineolatus, which suggests that only one of the three labrids is likely to play a significant role in regulation of echinoid densities in these rocky reef habitats. However, the broad diets and diverse forging strategies employed by these labrid species imply that they have a system-wide influence on invertebrate prey on rocky reefs.     

Mechanisms of benthic prey capture in wrasses (Labridae)

Teleost fishes capture prey using ram, suction, and biting behaviors. The relative use of these behaviors in feeding on midwater prey is well studied, but few attempts have been made to determine how benthic prey are captured. This issue was addressed in the wrasses (Labridae), a trophically diverse lineage of marine reef fishes that feed extensively on prey that take refuge in the benthos. Most species possess strong jaws with stout conical teeth that appear well-suited to gripping prey. Mechanisms of prey capture were evaluated in five species encompassing a diversity of feeding ecologies: Choerodon anchorago (Bloch, 1791), Coris gaimard (Quoy and Gaimard, 1824), Hologymnosus doliatus (Lacepède, 1801), Novaculichthys taeniourus (Lacepède, 1801) and Oxycheilinus digrammus (Lacepède, 1801). Prey capture sequences were filmed with high-speed video at the Lizard Island Field Station (14°40′S, 145°28′E) during April and May 1998. Recordings were made of feeding on pieces of prawn suspended in the midwater and similar pieces of prawn held in a clip that was fixed to the substratum. Variation was quantified among species and between prey types for kinematic variables describing the magnitude and timing of jaw, hyoid, and head motion. Species differed in prey capture kinematics with mean values of most variables ranging between two and four-fold among species and angular velocity of the opening jaw differing seven-fold. The kinematics of attached prey feeding could be differentiated from that of midwater captures on the basis of faster angular velocities of the jaws and smaller movements of cranial structures which were of shorter duration. All five species used ram and suction in combination during the capture of midwater prey. Surprisingly, ram and suction also dominated feedings on attached prey, with only one species making greater use of biting than suction to remove attached prey. These data suggest an important role for suction in the capture of benthic prey by wrasses. Trade-offs in skull design associated with suction and biting may be particularly relevant to understanding the evolution of feeding mechanisms in this group. Published online: 11 July 2002     

Water flow and prey capture by three scleractinian corals, Madracis mirabilis, Montastrea cavernosa and Porites porites, in a field enclosure

Scleractinian corals experience a wide range of flow regimes which, coupled with colony morphology, can affect the ability of corals to capture zooplankton and other particulate materials. We used a field enclosure oriented parallel to prevailing oscillatory flow on the forereef at Discovery Bay, Jamaica, to investigate rates of zooplankton capture by corals of varying morphology and polyp size under realistic flow speeds. Experiments were carried out from 1989 to 1992. Particles (Artemia salina cysts) and naturally occurring zooplankton attracted into the enclosures were used as prey for the corals Madracis mirabilis (Duchassaing and Michelotti) (narrow branches, small polyps), Montastrea cavernosa (Linnaeus) (mounding, large polyps), and Porites porites (Pallas) (wide branches, small polyps). This design allowed corals to be used without removing them or their prey from the reef environment, and avoided contact of zooplankton with net surfaces. Flow speed had significant effects on capture rate for cysts (M. mirabilis), total zooplankton (M. mirabilis, M. cavernosa), and non-copepod zooplankton (M. mirabilis). Zooplankton prey capture increased with prey concentration for M. mirabilis and M. cavernosa, over a broad range of concentrations, indicating that saturation of the feeding response had not occurred until prey density was over 10⁴ items m⁻³, a concentration at least an order of magnitude greater than the normal range of reef zooplankton concentrations. Location of cyst capture on coral surfaces was not uniform; for M. cavernosa, sides and tops of mounds captured most particles, and for P. porites, capture was greatest near branch tops, but was close to uniform for M. mirabilis branches in all flow conditions. The present study confirms laboratory flume results, and field results for other species, suggesting that many coral species experience particle flux and encounter rate limitations at low flow speeds, decreasing potential zooplankton capture rates. Received: 17 September 1996 / Accepted: 22 November 1997     

Alternative feeding mechanisms in megalopae of the blue crab Callinectes sapidus

The apparent mismatch between the energy requirements for planktotrophic growth and prey availability has long been paradoxical. One hypothesis to explain this paradox is that planktotrophic larvae display plasticity in feeding mechanisms in response to variable prey types and concentrations. This hypothesis was tested by videotaping megalopae of the brachyuran crab Callinectes sapidus Rathbun feeding on various-sized prey. Frame-by-frame analysis of the videotapes indicated that C. sapidus megalopae used both raptorial and suspension feeding to capture prey while in the water column. Raptorial feeding was used to capture macro-zooplankton, including copepods. The swimming form of suspension feeding was based on a modified fling-and-clap mechanism using the chelipeds. Suspension feeding while at rest utilized a weak current generated by the mouthparts to direct prey to the mouth. Both suspension-feeding mechanisms resulted in the efficient capture of rotifer-sized particles. The energy/handling time ratios for all three feeding mechanisms are very similar (E/H range 0.016-0.019 µg C s^-1) for the natural prey tested. These results support the hypothesis that feeding in brachyuran larvae is plastic and includes mechanisms of both raptorial and suspension feeding. The ability to suspension feed at rest is adaptive, since megalopae use selective tidal transport to re-invade an estuary and may spend up to 18 h day^-1 clinging to a benthic substrate. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00227-002-0781-1.     

Feeding ecology and niche segregation in oceanic top predators off eastern Australia

We examined the feeding ecology and niche segregation of the ten most abundant fish species caught by longline operations off eastern Australia between 1992 and 2006. Diets of 3,562 individuals were examined. Hook timer data were collected from a further 328 fish to examine feeding behaviour in relation to depth and time of day. Prey biomass was significantly related to predator species, predator length and year and latitude of capture. Although the fish examined fed on a mix of fish, squid and crustacea, fish dominated the diet of all species except small albacore (Thunnus alalunga) which fed mainly on crustacea and large swordfish (Xiphias gladius) and albacore which fed mainly on squid. Cannibalism was observed in lancetfish (Alepisaurus spp.). Multidimensional scaling identified three species groups based on their diet composition. One group consisted of yellowfin tuna (T. albacares), striped marlin (Tetrapturus audax) and dolphinfish (Coryphaena hippurus); a second group consisted of bigeye tuna (T. obesus), swordfish and albacore; and a third consisted of southern bluefin tuna (T. maccoyii) and blue shark (Prionace glauca). Of note was the separation of mako shark (Isurus oxyrhynchus) and lancetfish from all other predators. Prey length generally increased with increasing predator length although even large predators fed on a wide range of prey lengths including very small prey. Overall, differences in prey type and size, feeding times and depths were noted across the range of species examined to the extent that predators with overlapping prey, either in type or size, fed at different times of the diel period or at different depths. Taken together these data provide evidence for feeding niche segregation across the range of oceanic top predators examined.     

Effects of water motion and prey behavior on zooplankton capture by two coral reef fishes

Water motion is an important factor affecting planktivory on coral reefs. The feeding behavior of two species of tube-dwelling coral reef fish (Chaenopsidae) was studied in still and turbulent water. One species of blenny, Acanthemblemaria spinosa , lives in holes higher above the reef surface and feeds mainly on calanoid copepods, while a second, A. aspera , lives closer to the reef surface, feeds mainly on harpacticoid copepods, and is exposed to less water motion than the first. In the laboratory, these two blenny species were video recorded attacking a calanoid copepod ( Acartia tonsa, evasive prey) and an anostracan branchiopod (nauplii of Artemia sp., passive prey). Whereas A. spinosa attacked with the same vigor in still and turbulent water, A. aspera modulated its attack with a more deliberate strike under still conditions than turbulent conditions. For both fish species combined, mean capture success when feeding on Artemia sp. was 100% in still water and dropped to 78% in turbulent water. In contrast, when feeding on Acartia tonsa, mean capture success was 21% in still water and rose to 56% in turbulent water. We hypothesize that, although turbulence reduces capture success by adding erratic movement to Artemia sp. (passive prey), it increases capture success of Acartia tonsa (evasive prey) by interfering with the hydrodynamic sensing of the approaching predator. These opposite effects of water motion increase the complexity of the predator-prey relationship as water motion varies spatially and temporally on structurally complex coral reefs. Some observations were consistent with A. aspera living in a lower energy benthic boundary layer as compared with A. spinosa: slower initial approach to prey, attack speeds modulated according to water velocity, and lower proportion of approaches that result in strikes in turbulent water.Communicated by P.W. Sammarco, Chauvin     

Potential effects of a non-indigenous predator in its expanded range: assessing green crab, <Emphasis Type="Italic">Carcinus maenas</Emphasis>, prey preference in a productive coastal area of Atlantic Canada

The non-indigenous green crab (Carcinus maenas) is an important predator on bivalve wild beds in coastal areas worldwide. This study explored size-dependent green crab prey preference on American oysters (Crassostrea virginica), blue mussels (Mytilus edulis), and soft-shell clams (Mya arenaria) in a productive coastal system of Atlantic Canada. Using two sizes of prey and three different experimental manipulations, small, medium, and large green crabs were given a choice among these three bivalves, and their daily feeding rates were monitored over the course of 3 days. For both prey sizes, green crabs showed an early feeding preference for soft-shell clams and, only as they declined in numbers, a switch toward mussels and subsequently toward oysters. We found that such changes in the timing (order) of prey preference are related to prey differences in shell thickness, a fairly reliable indicator of prey shell strength.     

Thermal constraints on prey-capture behavior of a burrowing spider in a hot environment

Summary Seothyra henscheli (Eresidae) is a burrowing spider that lives in the dune sea of the southern Namib Desert, Namibia. Prey capture by these spiders involves a foray from a cool subterranean retreat to the undersurface of a capture web that can be lethally hot. Striking, disentangling and retrieving prey from the capture web typically involves several short trips to the capture web, alternating with retreats to the cool burrow. It has been suggested that this behavior limits the increase of body temperature a spider must experience while working at the hot capture web. We used biophysical models in conjunction with direct observations of prey-capture behavior and distributions of sand temperature to estimate body temperatures experienced by S. henscheli during prey capture. In the circumstances we observed, only the relatively long post-strike retreat from the capture web is important in keeping spiders' body temperatures from exceeding their lethal limits. After the post-strike retreat, shuttling appreciably limits the increase in body temperature of small individuals, but may have little effect on body temperature increase in larger spiders. Correspondence to: J.S. Turner at the present address     

Zooplankton capture by two scleractinian corals,Madracis mirabilis andMontastrea cavernosa,in a field enclosure

Capture of zooplankton by scleractinian corals has been noted for several species, yet quantitative information on rates of capture and differential capture by prey taxon has been lacking. We used field enclosures to examine prey capture for two coral species,Madracis mirabilis (Duchassaing and Michelotti) andMontastrea cavernosa (Linnaeus), on the north coast of Jamaica (Discovery Bay) in November 1989, February and March 1990, and January 1992.M. mirabilis has small polyps and a branching colony morphology (high surface/volume ratio), whereasM. cavernosa has large polyps and mounding colonies (low surface/volume ratio). Corals were isolated front potential prey, then were introduced into enclosures with enhanced zooplankton concentrations for 15- to 20-min feeding periods. Corals were fixed immediately after the experiment to prevent digestion, and coelenteron contents were examined for captured zooplankton. Plankton pumps were used to sample ambient zooplankton in the enclosures near the end of each run. Selectivity and capture rates were calculated for each prey taxon in each experiment; both indices were high for relatively uncommon large prey, and low for copepods, which were often the most common items in the plankton. Sizes of zooplankton captured by both species were generally larger than those available considering all prey taxa combined, but were almost the same for both coral species, even though the corals' polyp sizes are very different. This occurred primarily because small copepods, with low capture rates, dominated most plankton samples. For specific prey species, or group of species, there were few significant differences in size between the prey available and the prey captured.M. mirabilis, with small polyps, also captured far more prey per unit coral biomass than didM. cavernosa, with much larger polyps. We hypothesize that the large differences in capture rate of prey taxa are related to escape or avoidance behavior by those potential prey, and to the mechanics of capture, rather than to any selectivity by the corals.     

Effects of symbiotic status,flow speed,and prey type on prey capture by the facultatively symbiotic temperate coral <Emphasis Type="Italic">Oculina</Emphasis><Emphasis Type="Italic">arbuscula</Emphasis>

Symbiotic temperate corals can supplement prey capture by the coelenterate host with autotrophic carbon production by endosymbiotic zooxanthellae. To test the relationship between heterotrophic consumption and photosynthetic energy, prey capture by symbiotic and aposymbiotic specimens of the temperate scleractinian coral Oculina arbuscula (Verrill) was measured in January-April 2001. Corals were tested in a laboratory flume at five flow speeds, using Artemia franciscana cysts and nauplii as prey. Per-polyp capture rate and feeding efficiency were independent of symbiotic condition. Capture rate increased with flow speed, while capture efficiency declined. The location of capture shifted from the upstream to downstream side of the coral as flow speed increased. Differences in capture rate, location, and feeding efficiency for cysts and live brine shrimp nauplii were likely due to prey size rather than swimming ability.     

The effects of zooplankton swimming behavior on prey-capture kinematics of red drum larvae, <Emphasis Type="Italic">Sciaenops ocellatus</Emphasis>

Most marine fishes undergo a pelagic larval phase, the early life history stage that is often associated with a high rate of mortality due to starvation and predation. We present the first study that examines the effects of prey swimming behavior on prey-capture kinematics in marine fish larvae. Using a digital high-speed video camera, we recorded the swimming velocity of zooplankton prey (Artemia franciscana, Brachionus rotundiformis, a ciliate species, and two species of copepods) and the feeding behavior of red drum (Sciaenops ocellatus) larvae. From the video recordings we measured: (1) zooplankton swimming velocity in the absence of a red drum larva; (2) zooplankton swimming velocity in the presence of a red drum larva; and (3) the excursion and timing of key kinematic events during prey capture in red drum larvae. Two-way ANOVA revealed that: (1) swimming velocity varied among zooplankton prey; and (2) all zooplankton prey, except rotifers and ciliates, increased their swimming velocity in the presence of a red drum larva. The kinematics of prey capture differed between two developmental stages in S. ocellatus larvae. Hyoid-stage larvae (3–14 days old) fed on slow swimming B. rotundiformis (rotifers) while hyoid-opercular stage larvae (15 days and older) ate fast moving A. franciscana. Hyoid-opercular stage red drum larvae had a larger gape, hyoid depression and lower jaw angle, and a longer gape cycle duration relative to their hyoid-stage conspecifics. Interestingly, the feeding repertoire within either stage of red drum development was not affected by prey type. Knowledge of the direct relationship between fish larvae and their prey aids in our understanding of optimal foraging strategies and of the sources of mortality in marine fish larvae.     

A comparison of strike and prey capture kinematics of three species of piscivorous fishes: Florida gar ( Lepisosteus platyrhincus), redfin needlefish ( Strongylura notata), and great barracuda ( Sphyraena barracuda)

Ram feeding is the process by which a predatory fish uses a high-velocity lunge or chase to overtake its prey. This study compares the strike and prey capture behaviors and kinematics of three species of ram-feeding fishes: Florida gar Lepisosteus platyrhincus, redfin needlefish Strongylura notata, and great barracuda Sphyraena barracuda. These ram-feeding piscivores are morphologically similar with fusiform bodies, posteriorly placed dorsal and anal fins, and large, conical teeth. Strike and prey capture kinematics for five individuals of each species were recorded with high-speed video. Pre-strike behavior in L. platyrhincus consists of a slow stalk, resulting in the close, lateral positioning of the predators head relative to the prey. Lepisosteus employ a sideways lunge of the head during the strike, which lasts only 25–40 ms and is the most rapid strike among these three species. Strongylura notata and Sphyraena barracuda exhibit longitudinal orientation to the prey before the strike, followed by a high velocity, head-on lunge, initiated by an s-start in Sphyraena barracuda. Prey capture in adult L. platyrhincus and Strongylura notata is characterized by the jaws closing on the prey, with the prey held orthogonal to the jaws. This is followed by manipulation using the inertia of the prey to reposition the prey head first, and then suction transport into the buccal cavity. Prey capture in juvenile Sphyraena is accomplished by closing the jaws after the prey has entered the buccal cavity, resulting in possible ram transport of the prey with no oral manipulation under these experimental conditions. Although these three species all employ ram feeding for prey capture of elusive prey, each species has a unique repertoire that appears to minimize hydrodynamic constraints and prey response, utilize locomotory capabilities, and may be suited to each species specific habitat.Communicated by P.W. Sammarco, Chauvin     

Habitat use pattern of tidal mud and sand flats by shorebirds (charadriiformes) wintering in southern India

Habitat use pattern and status of shorebirds in southern India were investigated in six tidal flats comprising three mudflats and three sand flats on the east coast of southern India. Totally, 7757 shorebirds belonging to 21 species were recorded during September 2000–March 2002. The birds preferred mudflats over sand flats. Density, diversity and richness of shorebirds were relatively higher in tidal flats, especially in mudflats. The shorebirds preferred mudflats over sand flats as stopover sites and sites for refueling the required energy during migration. Rhythmic changes in tidal pattern also influenced the allochthonous nutrients and prey species, which also attracted the migratory shorebirds. However, the abundance of shorebirds with reference to tidal flats *season*tide and habitats*seasons * tide had strong variations (P < 0.001). During migratory season, bird density, diversity and species richness were observed higher than in other seasons.     

Morphology and kinematics of prey capture in the syngnathid fishes Hippocampus erectus and Syngnathus floridae

The mechanism of prey capture in two syngnathid fishes, the lined seahorse Hippocampus erectus (Perry) and the dusky pipefish Syngnathus floridae (Jordan and Gilbert), is described based on anatomical observations and high-speed video recordings (200 and 400 images s⁻¹) of feeding events by four seahorses and three pipefish. The fish were collected near Turkey Point, Florida, U.S.A., in January 1994 to March 1995. The dominant features of the morphology of these and many other syngnathiform fishes include extreme elongation of the suspensorium and neurocranium with a small mouth located at the anterior tip of the head. In the seahorse, a preparatory phase of prey capture consisted of slow ventral head flexion. This was followed by rapid elevation of the head and snout as the prey was drawn into the mouth by suction. Both H. erectus and S. floridae capture prey rapidly, with peak head excursions and mouth opening occurring within 5 to 7 ms of the onset of the strike. There was no upper jaw protrusion. In both species the time to recovery of the cranium and hyoid apparatus to resting positions was highly variable but took at least 500 ms. Manipulations of freshly dead specimens indicated a biomechanical linkage between head elevation and hyoid depression. However, the predictions of a previously proposed four-bar linkage model that couples hyoid depression to head elevation were not fully supported by kinematic data from one seahorse, suggesting that additional linkages act during the expansive phase of prey capture. These species exhibit the generalized kinematic pattern of prey capture in bony fishes, with head elevation, hyoid depression and mouth opening occurring almost simultaneously. The derived morphology results in a unique feeding behav‐ior, in which prey are captured during a sudden up-swing of the head, which brings the mouth to the prey. Suction is used to draw the prey into the buccal cavity. Received: 4 August 1996 / Accepted: 27 August 1996     

Feeding mechanism of the brisingid starfish Novodinia antillensis

This paper describes the food and feeding mechanisms of the bathyal brisingid sea-star Novodinia antillensis (A. H. Clark, 1934). N. antillensis is shown to be an opportunistic suspension feeder capable of capturing living planktonic crustaceans of a wide range of sizes. The sea-star extends its 10 to 14 arms into water currents to form a feeding fan with a large surface area, and uses highly retentive pedicellaria on the lateral arm spines to grasp and retain prey. The flexible arms form arm loops to complete the capture. The pedicellaria have a structure very similar to that used by other species known to capture macroscopic prey and, although they are an order of magnitude smaller, they probably function in a similar manner.     

Cercopagis pengoi and Mysis spp. alter their feeding rate and prey selection under predation risk of herring (Clupea harengus membras)

The anti-predator behaviour of Baltic crustacean planktivores was studied in feeding experiments under predation pressure of herring. The experiments were conducted with pelagic mysids: Mysis mixta and Mysis relicta, and with Cercopagis pengoi, a non-indigenous cladoceran, which invaded the Baltic Sea in 1992. Zooplankton was offered as prey. Two kinds of experiments were performed in the absence and presence of chemical predator cues: (1) two-prey experiments with prey, which have poor or good escape responses and all three planktivores and (2) natural prey experiments with mysids in natural zooplankton assemblages. The results showed that all three species reacted to the chemical cue of herring by decreasing their feeding rate and altering prey selection. C. pengoi selected easily captured prey (rotifers) in two-prey experiments under predation risk while selection for any prey was evident in mysids in natural prey experiments only in the absence of predator cues. This indicates that planktivores have different anti-predator strategies, which are modified by their own prey capture abilities. C. pengoi was a very efficient predator on small prey with size-specific prey consumption rate 5 to 18 times the rate of mysids. Results show that the studied planktivores are capable of adjusting their feeding behaviour to decrease their conspicuousness in order to increase survival under predation risk. Further, results support the view that C. pengoi has adapted well to the Baltic ecosystem, sharing food niche with pelagic mysids and most probably having a strong influence on the whole pelagic food web.     

Predator shoaling moderates the confusion effect in blue-green chromis,Chromis viridis

Summary In experiments, blue-green chromis [Chromis viridis (Cuvier 1830)] were fed on either scattered or aggregated swarms of brine shrimp (Artemia sp.). Ten runs with each prey dispersion treatment were performed with shoals of one, two, five and ten chromis. The mean lag in reaching peak feeding rate for fish fed on aggregated prey was significantly shorter in the larger chromis shoals. In contrast, with the scattered treatment all such lags were similar and very short. As foraging proceeded, higher feeding rates were observed in the larger feeding shoals, regardless of prey dispersion. Prey capture success (i.e. the rate of retention of intercepted prey) declined with time, but was significantly higher in groups of ten fish. Two main conclusions emerge. Firstly, grouping facilitated initiation of feeding by individuals preying on concentrated swarms and reduced the delay in reaching a maximum feeding level. This may have been due to a suppression of the confusion effect through reduced reliance upon vigilance. Secondly, reduced vigilance allowed larger shoals of chromis to feed effectively over more extended periods. Trends of increasing shoal cohesion and decreasing prey retention rate with time were consistent with a postulated increase in antipredator vigilance with declining feeding motivation.     

Effects of limiting access to prey on development of first zoeal stage of the brachyuran crabs Cancer magister and Hemigrapsus oregonensis

We tested the influence of limiting access to prey on larval development of the crabs Cancer magister and Hemigrapsus oregonensis by raising their Stage 1 larvae in the laboratory on different prey densities and with various periods of access to prey. Experiments were conducted in 1995 and 1996 at the Shannon Point Marine Center in Anacortes, Washington, USA. Our results show that crab larvae do not require continuous access to prey for optimal development nor do they appear to require light for prey capture. Survival and duration of Stage 1 C. magister fed continuously on only one-fourth the amount of the control density of prey and those fed at the control density for only 6 h per day were the same as for larvae fed continuously at the control density (20 ml⁻¹). Larvae with cyclic access to prey at the control density for 24 h and then starved for 72 h showed significantly lower survival and longer instar duration to Stage 2. Experiments on Stage 1 H. oregonensis which investigated a combination of prey density, period of access to prey and light/dark conditions during feeding revealed that survival decreased with decreasing prey density or with decreasing feeding period, but no differences were observed during periods of limited prey availability as a function of light or dark conditions. Stage duration was not affected by reduced prey density nor by the light/dark condition at the time of feeding, but it was prolonged when the period of access to prey was limited. The period of access to prey did not affect the weight of Day 1 Stage 2 larvae. Larvae fed high densities of prey for 4 h followed by 20 h of reduced-density diet exhibited the same survival and stage duration as controls that were continuously fed high-density prey. Our results define sub-optimal diets that can be used experimentally to determine the nutritional contributions made by naturally-occurring prey organisms during larval development in the two species. In nature, larvae may satisfy nutritional requirements through periodic encounters with dense prey patches during vertical migrations by day or night. Received: 12 August 1997 / Accepted: 5 February 1998     

Parent-offspring aggression in moorhens

Summary Colonial orb-weaving spiders from Mexico were studied to test predictions of risksensitive foraging theory: 1. group foraging increases prey capture/individual, and reduces prey variance; 2. spiders should be expected to exhibit risk-averse behavior (forage in groups) when the average level of prey exceeds individual needs, and exhibit risk-prone behavior (forage solitarily) when prey are searce. Laboratory and field studies show that group foraging increases capture efficiency and reduces variability in prey captured per spider. In desert/mesquite grassland habitat, where prey availability is low, M. atascadero forage solitarily in most cases. In tropical rainforest/agriculture sites, M. increassata forage in large colonies of thousands of webs. In intermediate habitats, M. spinipes forages solitarily or in groups, depending on prey availability. Over a range of sites with varying levels of prey, M. spinipes shifts from a risk-prone to a risk-averse group foraging strategy as prey increases.Group foraging behavior observed in colonial Metepeira fits the predictions of risk-sensitive foraging models. These findings explain why spiders tend to group webs together only in areas of superabundant prey. The role of risk-sensitivity in the evolution of coloniality in spiders is discussed.