Field-based video observations of wild barnacle cyprid behaviour in response to textural and chemical settlement cues

Many marine invertebrate larvae respond behaviourally to environmental settlement cues, yet behaviours are often only inferred from settlement patterns or are limited to laboratory studies. The behaviour of wild cypris larvae of Semibalanus balanoides L. was filmed on settlement tiles in the field. Tiles were of five different textures with a nested treatment of crude conspecific adult extract (AE). The effects of texture and AE on eleven defined behaviours were analysed. Texture affected the gross and net exploratory distances, velocity, acceleration and time spent exploring. AE attracted more cyprids during the first minute of immersion and increased the time spent on surfaces. Relatively few arrivals that either travel far and fast, or exit the surface rapidly, may indicate a lower chance of settlement. An increase in time spent on a surface may increase the probability of being in contact with the surface when the sign stimulus to settle occurs.     

Drift and settlement of stream insects in a complex hydraulic environment

1. The hydraulic and geomorphic characteristics of stream patches are often associated with distinctive assemblages or densities of stream invertebrates, and it is routinely presumed that these patterns reflect primarily species‐specific habitat requirements. An alternative hypothesis is that such patterns may be influenced by constraints on movement, such as the results of departure and settlement processes. We describe a manipulative experiment that examined how the hydraulic environments created by topographic bedforms influenced the drift behaviour and potential settlement sites for two species of mayfly (Baetis rhodani and Ecdyonurus torrentis). These species are common in the drift and often co‐occur in streams, but differ in their small‐scale distribution patterns, body shape and movement behaviour. 2. Flume experiments were carried out to determine how the hydraulic environments conditioned by a step bedform influence the behaviour of mayflies in the drift (swimming, posturing, tumbling), and the consequences of those behaviours (drift distance and time), compared to drift over a plane bed. The ramped step in the flume mimicked step bedforms that are common in coarse‐grained, high‐gradient streams. In contrast to the plane bed, a zone of recirculating flow was created downstream of the step, above which flow was faster and more turbulent. Uniform flows are used in most flume studies of drift; our approach is novel in recreating a complex hydraulic environment characteristic of stream channels. 3. Both species had some behavioural control over drift, and drift distances and times were shorter for live larvae than for dead larvae over the plane bed. The step had no impact on drift time or distance for live Baetis, but dead larvae were trapped in the flow separation eddy and drift time increased accordingly. Some Ecdyonurus also became trapped in the eddy, but live larvae drifted farther than dead larvae, and farther over the step than the plane bed. 4. Whilst in the drift, larvae altered their behaviour according to the ambient hydraulic environment, but in a species‐specific manner. Over the plane bed, Baetis had occasional swimming bursts, but primarily postured (maintained a stable body orientation), whereas Ecdyonurus spent roughly equal time posturing and swimming. In the more turbulent flows generated by the step, Baetis spent proportionately more time swimming, whereas Ecdyonurus spent more time posturing and often tumbling as body orientation became unstable. 5. In a high‐gradient stream, Baetis was more abundant close to steps than in plane bed patches with less complex flow, whereas the opposite pattern held for Ecdyonurus. Thus, the small‐scale distribution patterns of these species within streams correspond to their drift behaviours and ability to access various hydraulic patch types in our flume. These results are consistent with the hypothesis that constraints on movement and settlement may be important driver of distribution patterns within streams.     

Conflict between antipredator and antiparasite behaviour in larval damselflies

 Larval damselflies resist infestation by parasitic larval mites by exhibiting behaviours such as grooming, crawling, swimming, and striking at host-seeking mites. Larval damselflies are known to increase time spent in these behaviours in the presence of mites but reduce time spent in these behaviours in the presence of fish predators. The presence of both fish and larval mites presents an obvious conflict: a larval damselfly may actively avoid parasitism by mites, thus increasing its risk of predation, or it may reduce its activity when fish are present, thus increasing its risk of parasitism. We analysed the behaviour of larval Ischnura verticalis in an experiment where we crossed presence and absence of fish with presence and absence of larval mites. Presence of mites induced a large increase in activity of larval I. verticalis but fish had no effect and there were no interpretable interactions between effects of mites and fish. Subsequent experiments indicated that larval I. verticalis in the presence of both mites and fish were more likely to be attacked and killed by fish than those exposed only to fish. The high activity level of I. verticalis larvae in the presence of both fish and mites may suggest that costs of parasitism are high, or that under field conditions it is rare for larvae to be in the immediate presence of both fish predators and potentially parasitic mites. Received: 28 March 1996 / Accepted: 6 September 1996     

Strong behavioural defensive responses of endemic Rana latastei tadpoles induced by a native predator's odour

Prey species must constantly acquire information on predator identity, abundance and dangerousness from the environment. In aquatic habitats, this information is mainly propagated by water-borne chemical signals, either predator-specific odours or prey alarm cues. Anuran larvae innately respond to conspecific alarm cues and are able to associate them to predator cues during their lifetime. In this study, we investigated the anti-predatory responses of endemic Italian agile frog (Rana latastei) tadpoles exposed to either conspecific or heterospecific alarm cues and a native predator's (Anax imperator larvae) odour. Pre-and post-stimulus behaviours of each tadpole were recorded by a digital camera and analysed by a source executable software for image-based tracking. We found that Italian agile frog tadpoles responded to fasted dragonfly odour by strongly reducing their activity, both in terms of the amount of time they spent active and path length covered in comparison to control groups. Contrary to previous studies, predators' diet had a negligible effect on tadpole response and our experiment did not bring any evidence of the phylogenetic-relatedness hypothesis. The innate or early-in-development recognition of dragonfly larvae is clearly adaptive and may increase tadpole survival with relatively low costs, but, at the same time, may increase the risk of ignoring novel potential threats.     

The importance of honeydew as food for larvae of Chrysoperla carnea in the presence of aphids

Larvae of the common green lacewing Chrysoperla carnea are predacious and feed on a wide range of small, soft‐bodied arthropods. In addition to their feeding on prey arthropods to cover their nutritional requirements for growth and development, the consumption of non‐prey foods such as honeydew has been reported. It is commonly believed that these food supplements are primarily exploited by the larvae when prey is scarce or of low nutritional quality. Here, we assess whether C. carnea larvae also use honeydew when high‐quality aphid prey are readily available. In a choice experiment, the feeding behaviour of C. carnea larvae was observed in the presence of both aphids and honeydew. The larvae were starved, aphid‐fed, or honeydew‐fed prior to the experiment. The time spent feeding on honeydew compared with feeding on aphids was highest for starved larvae and lowest for honeydew‐fed larvae. Among the three treatments, the aphid‐fed larvae spent the most time resting and the least time searching. In an additional experiment food intake was assessed in terms of weight change when larvae were provided with an ad libitum supply of either aphids or honeydew. Larvae yielded a significant lower relative weight increase on honeydew compared with aphids. The reduced weight increase on honeydew was compensated when larvae were subsequently provided with aphids, but not when honeydew was provided again. This study showed that (i) prior honeydew feeding reduces overall aphid consumption, and (ii) larvae do consume honeydew even after they have been given ad libitum access to aphids. The fact that larvae of C. carnea still use honeydew as a food source in the presence of suitable prey underlines the importance of carbohydrates as foods.     

Larval settlement of the common Australian sea urchin Heliocidaris erythrogramma in response to bacteria from the surface of coralline algae

Bacterial biofilms are increasingly seen as important for the successful settlement of marine invertebrate larvae. Here we tested the effects of biofilms on settlement of the sea urchin Heliocidaris erythrogramma. Larvae settled on many surfaces including various algal species, rocks, sand and shells. Settlement was reduced by autoclaving rocks and algae, and by treatment of algae with antibiotics. These results, and molecular and culture-based analyses, suggested that the bacterial community on plants was important for settlement. To test this, approximately 250 strains of bacteria were isolated from coralline algae, and larvae were exposed to single-strain biofilms. Many induced rates of settlement comparable to coralline algae. The genus Pseudoalteromonas dominated these highly inductive strains, with representatives from Vibrio, Shewanella, Photobacterium and Pseudomonas also responsible for a high settlement response. The settlement response to different bacteria was species specific, as low inducers were also dominated by species in the genera Pseudoalteromonas and Vibrio. We also, for the first time, assessed settlement of larvae in response to characterised, monospecific biofilms in the field. Larvae metamorphosed in higher numbers on an inducing biofilm, Pseudoalteromonas luteoviolacea, than on either a low-inducing biofilm, Pseudoalteromonas rubra, or an unfilmed control. We conclude that the bacterial community on the surface of coralline algae is important as a settlement cue for H. erythrogramma larvae. This study is also an example of the emerging integration of molecular microbiology and more traditional marine eukaryote ecology.     

Rapid settlement in broadcast spawning corals: implications for larval dispersal

Broadcast spawning of gametes with planktonic development of larvae is the most common reproductive mode in tropical corals, and is generally thought to optimize the dispersal potential of larvae. To this end, many previous studies of coral larval dispersal have focused on the maximum time larvae can remain competent to settle and consequently how far they might disperse. However, dispersal ability of broadcast-spawned coral larvae will be linked, at least in part, to the minimum time to settlement competency as well as the length of the planktonic period—although estimates of minimum time to competency remain largely anecdotal, with few rigorous studies of the pre-competent period. To determine the minimum time to larval settlement in two species of broadcast-spawning coral ( Platygyra daedalea and Goniastrea favulus), we monitored larval settlement rates in aquaria every 6 h from the time larvae commenced swimming (i.e. were ciliated, fully developed larvae) for a period of approximately 10 days. For P. daedalea, peak settlement occurred between 60 and 66 h following fertilization (2.5 and 2.75 days), which is markedly earlier than the 4- to 6-day time period commonly cited as the minimum time before broadcast-spawned coral larvae are competent to settle. Surprisingly, it was also clear from our experimental results that settlement in P. daedalea occurred as a distinct pulse during the 60- to 66-h period, rather than continuously throughout the study period. G. favulus larvae also appear to be able to settle quickly (from 54 h following fertilization). We argue, on the basis of these short competency times and apparently rapid settlement, that dispersal in broadcast-spawning coral larvae may not be as great as has previously been assumed.     

Ontogenetic changes in responses to settlement cues by Anemonefish

Population connectivity for most marine species is dictated by dispersal during the pelagic larval stage. Although reef fish larvae are known to display behavioral adaptations that influence settlement site selection, little is known about the development of behavioral preferences throughout the larval phase. Whether larvae are attracted to the same sensory cues throughout their larval phase, or exhibit distinct ontogenetic shifts in sensory preference is unknown. Here, we demonstrate an ontogenetic shift in olfactory cue preferences for two species of anemonefish, a process that could aid in understanding both patterns of dispersal and settlement. Aquarium-bred na?ve Amphiprion percula and A. melanopus larvae were tested for olfactory preference of relevant reef-associated chemical cues throughout the 11-day pelagic larval stage. Age posthatching had a significant effect on the preference for olfactory cues from host anemones and live corals for both species. Preferences of olfactory cues from tropical plants of A. percula, increased by approximately ninefold between hatching and settlement, with A. percula larvae showing a fivefold increase in preference for the olfactory cue produced by the grass species. Larval age had no effect on the olfactory preference for untreated seawater over the swamp-based tree Melaleuca nervosa, which was always avoided compared with blank seawater. These results indicate that reef fish larvae are capable of utilizing olfactory cues early in the larval stage and may be predisposed to disperse away from reefs, with innate olfactory preferences drawing newly hatched larvae into the pelagic environment. Toward the end of the larval phase, larvae become attracted to the olfactory cues of appropriate habitats, which may assist them in identification of and navigation toward suitable settlement sites.     

Are behavioural responses to predation cues linked across life cycle stages?

1. Prey organisms can perceive cues to predation hazard and adopt low‐risk behaviours to increase survival. Animals with complex life cycles, such as insects, can exhibit such anti‐predatory behaviours in multiple life stages. 2. Cues to predation risk may induce ovipositing females to choose habitats with low predation risk. Cues to predation risk may also induce larvae to adopt facultative behaviours that reduce risk of predation. 3. One hypothesis postulates that anti‐predation behaviours across adult and larval stages may be negatively associated because selection for effective anti‐predator behaviour in one stage leads to reduced selection for avoidance of predators in other stages. An alternative hypothesis suggests that selection by predation favours multi‐component defences, with both avoidance of oviposition and facultative adoption of low‐risk behaviours by larvae. 4. Laboratory and field experiments were used to determine whether defensive responses of adult and larval mosquitoes are positively or negatively associated. The study tested effects of waterborne cues from predatory Toxorhynchites theobaldi on oviposition choices and larval behaviours of three of its common prey: Culex mollis, Limatus durhamii and Aedes albopictus. 5. Culex mollis shows strong anti‐predator responses in both life stages, consistent with the hypothesis of a multi‐component behavioural defence. The other two species showed no detectable responses to waterborne predator cues in either adult or larval stages. Larvae of these unresponsive species were significantly more vulnerable to this predator than was C. mollis. 6. For these mosquitoes, species appear either to have been selected for multi‐component defences against predation or to act in ways that could be called predator‐naïve.     

Behavior and development of larvae in the sponge Haliclona amboinensis

Sponges play important roles in marine ecosystems by contributing to habitat complexity and benthopelagic coupling of nutrients. Yet, the reproduction and settlement behaviors of diverse sponge species are not well understood. Here, we examined the brooding demosponge Haliclona amboinensis, which is common on shallow reefs in Bolinao, northwestern Philippines. Gravid sponges were found between the months of May and August, coinciding with warmer sea surface temperature. Sponges released parenchymella larvae from brood chambers in the mid‐morning, suggesting that light and temperature may serve as cues to initiate hatching. Larvae immediately swam toward the surface upon emergence and migrated to the bottom of the tanks 1–2 hr after release. The presence of light and crustose coralline algae induced high larval settlement. Metamorphosis proceeded rapidly in vitro, with larval cells spreading laterally on the substrate. The osculum was first visible at 3 days after settlement. The short pelagic duration of larvae in H. amboinensis promotes local recruitment and may be important for the maintenance of sponge populations in the face of disturbances.     

Temporal settlement patterns of larvae of the broadcast spawning reef coral Favites chinensis and the broadcast spawning and brooding reef coral Goniastrea aspera from Okinawa, Japan

The faviid corals, Favites chinensis and Goniastrea aspera are widely distributed in the Indo-Pacific region. Both corals are hermaphroditic broadcast spawners, but G. aspera is also known to brood planula larvae in Okinawa. This study investigated the temporal settlement patterns of planula larvae of the scleractinian corals F. chinensis and G. aspera that developed from spawned gametes, and planula release and settlement of brooded larvae of G. aspera from Okinawa, Japan. Some of the broadcast-spawned larvae of F. chinensis and G. aspera had very short pre-competency periods of 1–2 and 2–3 days after spawning, and relatively long maximum settlement-competency periods of 56–63 and 63–70 days after spawning, respectively. These pre-competency periods are among the shortest reported for larvae of broadcast spawning coral species, and appear to be negatively correlated with seawater temperature. F. chinensis larvae tended to settle rapidly with 34–39% of larvae settling in the first week after spawning, while broadcast-spawned G. aspera larvae had a slower settlement pattern with 11–15% of larvae settling in the first week after spawning. Brooded larvae of G. aspera settled more rapidly, with settlement rates of 27–31% within the first 24 h and 45–65% within the first week after the start of the experiment. The production of planula larvae with rapid settlement capabilities may enable F. chinensis and G. aspera to establish and maintain populations in shallow reef sites at Okinawa. The release of the brooded planulae for up to 2 months may explain why G. aspera is locally more dominant on shallow reefs in Okinawa than F. chinensis. On a broader scale, the longer settlement competency periods of some of the broadcast-spawned larvae of these species increase their potential for longer-distance dispersal and may partly explain the wide biogeographic distribution of these species in the Indo-Pacific region.     

Surface films as mosquito larvicides: partitioning the mode of action

Oils and other surface films used against mosquito larvae may act by flooding the tracheal system with oil, by disrupting the surface forces that allow larvae to rest at the surface, by toxicity, or by eliciting chemosensory responses. In an attempt to identify diagnostic symptoms of these modes of action we treated fourth-stage larvae of Culex pipiens L. form molestusForskål (Diptera: Culicidae: Culicinae) with agents operating in a single mode (Ondina oil for flooding with oil, nicotine for toxicity, and the detergent Triton X-100 for disruption of surface forces), in two modes (silicone oil) or in three (eucalyptus oil, citronellal, or caproic acid ethyl ester). We monitored the time course of flooding and immobilisation, used experiments to separate volatile toxicity from toxicity of the aqueous solution, and used video and The Observer software to analyse larval behaviour.Larvae that experienced tracheal flooding applied their mouthparts to the siphon (tail nibbling), often losing contact with the surface while doing so and falling to the bottom. Nicotine immobilised larvae without interfering with surface forces, and the larvae remained immobile hanging from the air-water interface for long periods. In detergent, larvae made repeated unsuccessful attempts to thrust the siphon through the surface. The behaviour-modifying chemicals caproic acid ethyl ester, eucalyptus oil and citronellal all operated in more than one mode. They all decreased the proportion of time larvae spent at the surface, whereas Ondina oil and silicone fluid increased it.Using this approach it should be possible to identify the modes of action of novel larvicidal agents. This could form a basis for rational design of larvicides giving an optimal compromise between efficacy against mosquito larvae and minimal damage to non-target organisms.     

Ocean acidification boosts larval fish development but reduces the window of opportunity for successful settlement

Locating appropriate settlement habitat is a crucial step in the life cycle of most benthic marine animals. In marine fish, this step involves the use of multiple senses, including audition, olfaction and vision. To date, most investigations of larval fish audition focus on the hearing thresholds to various frequencies of sounds without testing an ecological response to such sounds. Identifying responses to biologically relevant sounds at the development stage in which orientation is most relevant is fundamental. We tested for the existence of ontogenetic windows of reception to sounds that could act as orientation cues with a focus on vulnerability to alteration by human impacts. Here we show that larvae of a catadromous fish species (barramundi, Lates calcarifer) were attracted towards sounds from settlement habitat during a surprisingly short ontogenetic window of approximately 3 days. Yet, this auditory preference was reversed in larvae reared under end-of-century levels of elevated CO₂, such that larvae are repelled from cues of settlement habitat. These future conditions also reduced the swimming speeds and heightened the anxiety levels of barramundi. Unexpectedly, an acceleration of development and onset of metamorphosis caused by elevated CO₂ were not accompanied by the earlier onset of attraction towards habitat sounds. This mismatch between ontogenetic development and the timing of orientation behaviour may reduce the ability of larvae to locate habitat or lead to settlement in unsuitable habitats. The misinterpretation of key orientation cues can have implications for population replenishment, which are only exacerbated when ontogenetic development decouples from the specific behaviours required for location of settlement habitats.     

Effect of hunger on the allocation of time among pea plants by the larvae of an aphidophagous hover fly,Eupeodes corollae [Dipt: Syrphidae]

We observed the movement of predatory larvae of the syrphid flyEupeodes corollae (F.) (formerlyMetasyrphus corollae) among small pea plants with and without aphids. Starved larvae spent longer time than well-fed larvae on similar plants and both groups of larvae stayed longer on plants with aphids than on plants without aphids. On plants with aphids, larvae which failed to capture prey left the plant sooner than those which captured aphids. The capture of at least one aphid on a plant increased the persistence of syrphid larvae. The average rate of energy gain was higher for well-fed larvae than for starved larvae because starved larvac stayed on plants even when their rate of return was lower. When larvae that had captured aphids left plants, their rate of energy gain, tended to be lower than at any time following capture of the 2^nd, aphid. The 1^st aphid was captured in less time than similar larvae spent on plants without aphids. Time between captures of aphids by well-fed larvae was less than the time such larvae spent on plants without aphids. Among starved larvae, the intercatch intervals were similar to the time on plants without aphids. We discuss the significance of these results relative to current predator foraging theory and the efficiency ofE. corollae as a biological control agent.      

Complex behaviour by coral-reef fish larvae in open-water and near-reef pelagic environments

We present the first in situ observations of the pelagic larvae of coral-reef fishes feeding, schooling and being preyed upon. In addition, we report on their behavioural interactions with adult and juvenile fishes. Observations on over 500 larvae of over 50 species (mostly from four families) near the end of their pelagic interval were made in both open water (> 1 km offshore) and near-reef environments. Nearly 10% of larvae were seen to feed in open water, but < 1% fed near the reef. Presettlement schooling was observed in five species of four families. We observed no predation upon larvae in open water except near the bottom. Near the reef, 8.5% of larvae were eaten. The main predators near and on the reef were a species of wrasse and lizardfishes. Rates of predation seem to differ among genera of pomacentrids, perhaps related to differences in behaviour when settling. When confronted with adult fishes, which happened largely near the reef, larvae reacted with a limited range of behaviours, including sheltering near the observer, swimming to the surface, slowing or stopping, or swimming offshore. The frequency of these behaviours differed among larvae of three pomacentrid genera. Interactions with reef residents, particularly pomacentrids, were common, and usually involved aggression by the resident toward settling larvae. This may act to discourage settlement during the day when such residents are active. These data show that behaviour of late larvae of coral-reef fishes is complex and can greatly influence survival and recruitment. Further, behaviour differs among taxa, showing that not only are larvae not passive, but also that a generalised behaviour of larvae does not exist.     

Settlement behavior of cyphonautes larvae of the bryozoan Membranipora membranacea in response to two algal substrata

Abstract. Settlement is an important process in the biphasic life histories of many marine invertebrates. Little is known regarding the fine-scale behavioral mechanisms for finding and attaching to a suitable substratum, particularly under conditions that may impose a potential challenge, such as flow. In this study, we examined the settlement behavior of cyphonautes larvae of the bryozoan, Membranipora membranacea, in response to two different algal substrata. Larvae showed a strong preference for settling on the kelp Nereocystis luetkeana over the red alga Mazzaella splendens. We then tested whether the behavioral mechanisms used by larvae to attach to these algae differed when presented with the challenge of flowing water during settlement. We found that larvae exhibited different behaviors on the two species of algae in flowing water. Larvae were more often observed in direct contact with the preferred alga (N. luetkeana) exhibiting fine-scale active search behaviors. On the less preferred alga (M. splendens), larvae were less frequently observed in direct contact with the alga, and appeared to be exhibiting broad-scale passive search behaviors along the surface of the blade. Our results suggest that cyphonautes larvae alter their behavior in response to their preferred settlement habitat.     

Settlement behaviour of larvae of the Stripey Snapper, <Emphasis Type="Italic">Lutjanus carponotatus</Emphasis> (Teleostei: Lutjanidae)

Larval behaviour is important to dispersal and settlement, but is seldom quantified. Behavioural capabilities of larval Lutjanus carponotatus in both offshore pelagic and reef environments at Lizard Island, Great Barrier Reef were observed in situ to determine if they were sufficient to influence dispersal. Offshore, larvae swam with higher directional precision and faster on the windward side of the island (28 cm.s⁻¹) than on the leeward side (16 cm s⁻¹). Most larvae swam directionally. Mean swimming directions were southerly in the windward area and northerly in the leeward area. Larvae avoided the surface and remained mostly between 3–15 m. Larvae released near reefs were 2–3 times faster swimming away from reefs (19 cm s⁻¹) than swimming toward or over them (6–8 cm s⁻¹). Speed swimming away was similar to that offshore. Of 41 larvae released near reefs, 73% reached the reef, 59% settled, and 13% of those reaching the reef were eaten. Larvae settled onto hard and soft coral (58%), topographic reef features (29%) and sand and rubble (13%). Settlement depth averaged 5.5 m (2–8 m). Before settling larvae spent up to 800 s over the reef (mean 231 s) and swam up to 53 m (mean 14 m). About half of the larvae interacted with reef residents including predatory attacks and aggressive approaches by residents and aggressive approaches by settling larvae. Settlement behaviour of L. carponotatus was more similar to a serranid than to pomacentrids. Settlement-stage larvae of L. carponotatus are behaviourally capable, and have a complex settlement behaviour.     

Chemical‐Mediated Predator Avoidance in the European House Cricket (Acheta domesticus) is Modulated by Predator Diet

Chemical information often mediates interactions between predators and prey, and threat‐sensitivity theory includes predictions that prey species should respond to chemical signatures of predators in a manner that is commensurate with the level of the assessed threat. Using the European house cricket (Acheta domesticus), we explored the influence of diet‐derived cues from the centipede Scolopocryptops sexspinosus on anti‐predator behavior in three laboratory experiments. In experiment 1, we compared the amount of time that adult female crickets spent on untreated filter paper and filter paper exposed to centipedes fed either the larvae of Hermetia illucens (black soldier fly), crickets, or a mixture of fly larvae and crickets. We discovered that crickets spent significantly less time on filter paper exposed to centipedes fed crickets only or a mixture of crickets and fly larvae compared with blank filter paper or filter paper exposed to centipedes fed fly larvae only. In our second experiment, we compared the amount of time that crickets spent on blank filter paper and filter paper exposed to adult female conspecifics to rule out the possibility that crickets simply avoid all filter paper exposed to metabolic by‐products, and crickets exhibited no discrimination. In our third experiment, we tested the potential effects of diet order on anti‐predatory behaviors. Specifically, we compared the amount of time that adult female crickets spent on filter paper exposed to centipedes fed fly larvae followed by crickets and filter paper exposed to centipedes fed crickets followed by fly larvae. We discovered no diet sequence effect. Our study demonstrates that European house crickets are sensitive to the chemical cues of their centipede predators, but only when centipedes have fed upon crickets.     

Growth versus molting time of caterpillars as a function of temperature,nutrient concentration and the phenolic rutin

Summary A factorial experiment tested the effects of varying nutrient concentration (normal versus diluted), presence or absence of the phenolic allelochemical rutin and daytime temperature (20, 25 and 30° C) on growth, molting and food utilization efficiencies of tobacco hornworms (Manduca sexta). Two of the utilization efficiencies (approximate digestibility and efficiency of conversion of ingested food) were unaffected by temperature; the third one, efficiency of conversion of digested food, was affected by temperature but there was no consistent effect. Lower temperatures significantly increased the proportion of the stadium spent molting, with larvae at a daytime temperature of 20° C spending 9% more of the stadium in molting than larvae at 30° C. Growth time was not influenced by nutrient concentration. When temperature was low and rutin absent, molt time and the proportion of the stadium spent molting were affected by nutrient concentration. Addition of the phenolic rutin did not have an appreciable effect on growth time or digestive processes. However, it increased molting time by 7 to 14% and thus increased the duration of the stadium and reduced relative growth rate. These results indicate that the effect of food quality on growth rate is a function of the thermal conditions of insect herbivores. At cooler temperatures, a disproportionate increase in time spent molting, rather than altered food utilization efficiencies, contributed to lower growth rates. The consequences for larval growth of fluctuating temperatures due to diurnal cycles and the presence of predators forcing larvae into thermally suboptimal microhabitats are discussed.     

Diel timing of planulation and larval settlement in the coral Isopora cuneata (Scleractinia: Acroporidae)

The prevailing environmental conditions when reproductive products are released in the water column can have a profound influence on the biology and ecology of marine organisms. In reef-building corals, brooding species that release azooxanthellate larvae are expected to release their larvae before sunrise, similar to species releasing zooxanthellate larvae as established in previous studies. This study investigated the diel timing of release of asymbiotic larvae by Isopora cuneata in northwestern Philippines during 2 years through ex situ observations on gravid colonies collected from the reef. Planulation mainly occurred after sunset until midnight, with grand mean hour of release at 19:35 h (95% confidence limit: 18:44–21:02 h) in April 2010 and 19:41 h (18:10–20:59 h) in March 2011. This pattern of release timing is distinct from what has been previously reported for brooding scleractinian coral species with zooxanthellate larvae (i.e., release close to sunrise). The release coincides with the low tidal stand, which may provide opportunity for the prompt settlement of newly released, buoyant larvae to shallow reef areas where adult colonies are generally distributed. The larvae were able to settle with 20 ± 25% success within a day of release, though a distinct settlement peak at 3 days post-release (70 ± 25%). This study provides new information on the early life strategies of the reef-building coral Isopora cuneata. The unique pattern of larval release time and the settlement behavior may influence the population dynamics and success of the species through space and time.