Nocturnal foraging habitats of French and bluestriped grunts,Haemulon flavolineatum and H. sciurus,at Tobacco Caye,Belize

Synopsis Nocturnal foraging habitats of Haemulon flavolineatum and H. sciurus were investigated in the backreef habitat around Tobacco Caye, Belize. Grunts leave the reef at dusk to forage in the grass beds and sand flats surrounding the reef. The hypothesis that French and bluestriped grunts use separate foraging habitats was examined by following tagged fishes from their diurnal territories or schooling sites to nocturnal foraging grounds. The tag consisted of a small, glowing Cyalume light stick sutured to the dorsal musculature of the fish, next to the first dorsal fin. Surveys of foraging habitats were done to support the tracking study. Large quadrats (225 m²) were set out over the sand flats and grass beds during the day. The numbers of French and bluestriped grunts feeding in each habitat were counted one hour after dark. Foraging French grunts used sand flats, whereas bluestriped grunts usually fed in grass beds. Repeated sightings of two French grunts and one bluestriped grunt in the same individual night-time locations support the hypothesis that nocturnal foraging sites may be used repeatedly by the same individuals.     

Microsatellite multiplexes for high‐throughput genotyping of French grunts (Haemulon flavolineatum,Pisces: Haemulidae) and their utility in other grunt species

Grunts of the genus Haemulon are abundant in the western Atlantic and are an important component of regional fisheries. Management efforts for these and other Caribbean reef fish require an understanding of population structure and the extent of dispersal among populations. We characterized 11 polymorphic microsatellite loci for French grunts (Haemulon flavolineatum) and grouped them into three PCR multiplexes to facilitate high‐throughput genotyping. These loci will be useful for population genetic studies of French grunts. Cross‐species amplifications suggest these loci are also suitable for population genetic studies of H. aurolineatum, H. plumieri, and H. sciurus.     

Sound production during competitive feeding in the grey gurnard

The acoustic repertoire of captive grey gurnard Eutrigla gurnardus during competitive feeding consisted of three types of sound: knocks, grunts and growls. Knocks were audible as a single sound, whereas grunts and growls were perceived as longer, pulsed sounds to the human ear. Typically, knocks were composed of 1–2 pulses, grunts of 4–8 pulses and growls >10 pulses. Growls were longer and had shorter pulse periods than grunts. All sound types had peak frequencies of c . 500 Hz. The sequences of behaviours observed during feeding interactions suggest that grey gurnard obtain food both by scramble and contest tactics. Competing fish emitted knocks mainly while grasping a food item and also during other non‐agonistic behaviour, suggesting that knock production may reflect a state of feeding arousal but could also serve as a warning of the forager's presence to nearby competitors. Grunts were mainly emitted during frontal displays, which were the most frequent behavioural act preceding grasps, suggesting that they may play a role in deterring other fish from gaining access to disputed food items.     

Depth-Variable Settlement Patterns and Predation Influence on Newly Settled Reef Fishes (Haemulon spp., Haemulidae)

During early demersal ontogeny, many marine fishes display complex habitat-use patterns. Grunts of the speciose genus Haemulon are among the most abundant fishes on western North Atlantic coral reefs, with most species settling to shallow habitats (≤12 m). To gain understanding into cross-shelf distributional patterns exhibited by newly settled stages of grunts (<2 cm total length), we examined: 1) depth-specific distributions of congeners at settlement among sites at 8 m, 12 m, and 21 m, and 2) depth-variable predation pressure on newly settled individuals (species pooled). Of the six species identified from collections of newly settled specimens (n = 2125), Haemulon aurolineatum (tomtate), H. flavolineatum (French grunt), and H. striatum (striped grunt) comprised 98% of the total abundance; with the first two species present at all sites. Prevalence of H. aurolineatum and H. flavolineatum decreased substantially from the 8-m site to the two deeper sites. In contrast, H. striatum was absent from the 8-m site and exhibited its highest frequency at the 21-m site. Comparison of newly settled grunt delta density for all species on caged (predator exclusion) and control artificial reefs at the shallowest site (8-m) revealed no difference, while the 12-m and 21-m sites exhibited significantly greater delta densities on the caged treatment. This result, along with significantly higher abundances of co-occurring piscivorous fishes at the deeper sites, indicated lower predation pressure at the 8-m site. This study suggests habitat-use patterns of newly settled stages of some coral reef fishes that undergo ontogenetic shifts are a function of depth-variable predation pressure while, for at least one deeper-water species, proximity to adult habitat appears to be an important factor affecting settlement distribution.     

Acoustic communication in the Bocon toadfish (<Emphasis Type="Italic">Amphichthys cryptocentrus</Emphasis>)

The soundscapes of many coastal habitats include vocalizations produced by species of the family Batrachoididae (toadfish and midshipman). We describe the calling and grunting behavior of male Amphichthys cryptocentrus, a tropical toadfish, and predict how these vocalizations are influenced by conspecifics. We recorded individual males, which produced broadband grunts and multi-note, harmonic “boatwhistle” calls. Grunts were either in combination with calls or stand-alone. We used a null model to test if these latter grunts were produced at random or in response to calls from conspecifics. The model supports the hypothesis that grunts were in response to calls from neighboring males, suggesting acoustic competition. Using the most conservative estimate of hearing abilities we predict that males responded to the second harmonic of neighbor’s calls (230 Hz) at amplitudes of approximately 100–125 dB re 1μPa²/Hz. We also observed that call and grunt rates increased when males were exposed to higher rates of acoustic activity from neighboring fish. Fish used grunts to respond to background calls that occurred at different amplitudes, suggesting they responded to the calls of multiple neighboring fish and not just the highest amplitude neighbor. This communication with multiple fish within hearing range suggests a communication network in which the spatial distribution of individual toadfish relative to one another will impact their vocal behavior. Thus, the density and distribution, and not just abundance, of these toadfish at a given site will influence the characteristics of the chorus and the role of this species in the local soundscape.     

Movement patterns,home range size and habitat utilization of the bluespine unicornfish, <Emphasis Type="BoldItalic">Naso unicornis</Emphasis> (Acanthuridae) in a Hawaiian marine reserve

Synopsis We quantified bluespine unicornfish, Naso unicornis, movement patterns, home range size and habitat preferences in a small Hawaiian marine reserve. Bluespine unicornfish were site-attached to home ranges situated within the reserve boundaries and their movements were aligned with topographic features. Two different diel movement patterns (‘commuting’ and ‘foraying’) were observed. Commuters made crepuscular migrations of several hundred meters between daytime foraging areas and nighttime refuge holes. Foraying fish did not partake in crepuscular migrations and utilized refuge holes both day and night. Two bluespine unicornfish were also nocturnally active. There was little direct evidence of dispersal from the reserve but differences in bluespine unicornfish abundance and size among reef habitat zones were consistent with ontogenetic habitat shifts. The influence of habitat topography on bluespine unicornfish movements suggests that gross habitat characteristics could be used to predict reef fish movements. This could provide a simple method for setting marine reserve boundaries at sites for which empirical fish movement data are unavailable.     

Playbacks of food-associated calls attract chimpanzees towards known food patches in a captive setting

Food-associated calls have received much research attention due to their potential to refer to discovered food in a word-like manner. Studies have found that in many species, food-associated calls attract receivers to the food patch, suggesting these calls play roles in food sharing, cooperation and competition. Additionally, in various species, these calls play a role that has received much less attention: mediating social interactions among foragers that are already nearby or within the food patch, independently of whether they attract outside foragers. In order to increase understanding of the function of the chimpanzee (Pan troglodytes) food-associated rough grunt, we conducted captive playback studies testing whether rough grunt playbacks attract, repel or have no effect on the proximity of foragers already familiarized with the presence of food. We tested how acoustic playbacks of rough grunts (or control calls) from one of two known, identical feeding sites affected receivers’ approach and feeding behaviors. More often than expected, participants first approached the feeding site from which rough grunts, but not control calls, were broadcast. However, neither condition increased the likelihood that participants fed first from a given site. Our results support the hypothesis that rough grunts elicit an approach response in receivers, while providing no evidence that they repel. In addition, our study provides evidence that receivers may approach rough grunts even if they do not intend to feed. We discuss the information rough grunts may convey to receivers beyond information about discovered food and the potential benefits signalers may gain from this calling behavior.

     

Patterns of migration between feeding and spawning sites in a coral reef surgeonfish

Many coral reef fishes exhibit regular localised migrations between feeding and spawning areas, but the factors affecting these migration patterns, such as the distance, frequency and spawning site fidelity are poorly understood. The aim of this study was to investigate the patterns of migration to spawning sites of the surgeonfish, Ctenochaetus striatus (Acanthuridae). We explored relationships amongst an individual’s size and sex, the distance and frequency it migrated from its feeding area to spawning sites, fidelity to particular spawning sites and the number of individuals that aggregated to spawn. In order to achieve this, 406 C. striatus were captured and tagged on inshore reefs in Kimbe Bay (5°30′S 150°6′E), New Britain, Papua New Guinea. Tagged individuals were consistently observed within spatially discrete but overlapping feeding areas (maximum diameter averaging <13 m). The mean distance migrated was 58 m (ranging from 2 to 291 m). No tagged individuals were witnessed spawning at more than one site. Whilst most individuals (n = 88) migrated to the spawning site that was closest to their feeding areas, those that migrated to sites further away (n = 9) always spawned at sites where the number of conspecifics aggregating was larger. Neither the size nor the sex of individuals limited migration distance. However, males migrated significantly more frequently than females (on average once every 2 days vs. once every 3 days), and migration frequency was positively correlated with size in females. Migration distance did not affect the frequency with which individuals spawned. Whether patterns of migration are determined by cost-benefit optimisation, tradition, or an alternative mechanism is unknown.     

The meaning and function of grunt variants in baboons

Wild baboons Papio cynocephalus ursinus, give tonal, harmonically rich vocalizations, termed grunts, in at least two distinct, behavioural contexts: when about to embark on a move across an open area ('move' grunts); and when approaching mothers and attempting to inspect or handle their young infants ('infant' grunts). Grunts in these two contexts elicit different responses from receivers and appear to be acoustically distinct (Owren et al. 1997 Journal of the Acoustical Society of America101 2951-2963). Differences in responses to grunts in the two contexts may, then, be due to acoustic differences, reflecting at least a rudimentary capacity for referential signalling. Alternatively, responses may differ simply due to differences in the contexts in which the grunts are being produced. We conducted playback experiments to test between these hypotheses. Experiments were designed to control systematically the effects of both context and acoustic features so as to evaluate the role of each in determining responses to grunts. In playback trials, subjects differentiated between putative move and infant grunts. Their responses based only on the acoustic features of grunts were functionally distinct and mirrored their behaviour to naturally occurring move and infant grunts. However, subjects' responses were in some cases also affected by the context in which grunts were presented, and by an interaction between the context and the acoustic features of the grunts. Furthermore, responses to grunts were affected by the relative rank difference between the caller and the subject. These results indicate that baboon grunts can function in rudimentary referential fashion, but that the context in which grunts are produced and the social identity of callers can also affect recipients' responses. Copyright 1999 The Association for the Study of Animal Behaviour.     

Discrimination of French grunts (Haemulon flavolineatum, Desmarest, 1823) from mangrove and coral reef habitats using otolith microchemistry

Using French Grunts (Haemulon flavolineatum) held captive within mangrove and reef sites, we determined (a) whether otolith microchemical differences existed between mangroves and reefs separated at a biologically relevant spatial scale (0.25-7.1 km), (b) whether patterns in elemental concentrations were consistent across years, and (c) whether it was possible to identify whether a given fish occupied a mangrove as a juvenile.Three sites were established at Great Exuma, Bahamas (two reefs and one mangrove, May 2001) and at Turneffe Atoll, Belize (one reef and two mangroves, August 2001 and 2002). Using concentrations of Sr and Ba, discriminant function analysis (DFA) indicated unique spatial microchemical signatures of fish from each of the three Bahamas sites allowing an average correct classification of 77%. Using concentrations of Sr, Ba, Sn and Pb (2001), and in addition Li, Mg, Cu and Rb (2002), DFA of the three Belize sites indicated an average correct classification of 68% and 85% in 2001 and 2002, respectively. To assess temporal variability in otolith microchemistry, we compared microchemical signatures of Belize fish from 2001 to those from 2002. On average, 42% of fish from 2002 were correctly classified to their captive sites using chemical information from 2001, thus suggesting considerable temporal variability in otolith microchemistry. Finally, to identify whether a given fish occupied a mangrove during its juvenile stage, we ablated the juvenile portion of the otolith taken from reefs in Belize 2002. Results of this analysis indicated that 36% of 39 individuals taken from the reef had a signature more representative of one of the mangrove sites. Although otolith microchemistry varied temporally and our analysis was restricted to the grouping of individuals to only one of three sites, mangroves appeared to contribute to reef populations.     

Effect of naturally changing zooplankton concentrations on feeding rates of two coral species in the Eastern Pacific

Zooplankton concentrations are known to vary by as much as an order of magnitude over a lunar cycle. Here, we conducted an experiment to determine the effect of ambient zooplankton concentrations over a lunar cycle on feeding rates of the corals Pavona gigantea (Verrill) (mounding coral, 3.0 mm diameter polyps) and Pocillopora damicornis (Linnaeus) (branching coral, 1.0 mm diameter polyps) in situ on a shallow reef at Isla Contadora, Gulf of Panamá (Pacific), Panamá. Coral fragments exposed to either enhanced or ambient zooplankton concentrations were allowed to feed for 1 h, collected, and their gut contents dissected. The number of zooplankton captured was counted, feeding rates calculated per cm², and the species composition of captured zooplankton assemblages determined. Although both species captured the same zooplankton assemblage, feeding rates were always significantly higher for P. gigantea than for P. damicornis. Under ambient flow and zooplankton concentrations, feeding rates were highly correlated with zooplankton concentration in the 200-400 μm size class. Under constantly enhanced zooplankton concentrations in the control fragments, feeding rates did not vary significantly over the lunar cycle. As such, coral feeding rates vary not as a result of lunar phase per se, but with changes in zooplankton abundance over the lunar cycle. Coral feeding rates are directly proportional to ambient zooplankton concentrations and may vary by as much as 50% over a lunar cycle, suggesting that corals must cope with major swings in sources of fixed carbon and nutrients over relatively short timescales.     

Foraging behavior and diet composition of Trimma caudomaculata and Trimma caesiura (Gobiidae) on coral reefs in Okinawa, Japan

The feeding habits of two small gobies, Trimma caudomaculata and T. caesiura, were surveyed on coral reefs in Okinawa, Japan. Both species actively exhibited foraging attempts beneath overhanging reef structures during the daytime, and fed on small animals, mainly copepods. Trimma caudomaculata formed feeding schools in water columns to swallow mainly Calanoida and Harpacticoida. Trimma caesiura stayed by themselves on the reef substratum and captured mainly Harpacticoida by taking short jumps into the water column. Diet compositions of these congeneric gobies may reflect differences in their foraging behaviors and use of microhabitats.     

Prevalence of Borrelia burgdorferi Sensu Lato in Ticks Collected from Migratory Birds in Switzerland

The prevalence of ticks infected by Borrelia burgdorferi sensu lato on birds during their migrations was studied in Switzerland. A total of 1,270 birds captured at two sites were examined for tick infestation. Ixodes ricinus was the dominant tick species. Prevalences of tick infestation were 6% and 18.2% for birds migrating northward and southward, respectively. Borrelia valaisiana was the species detected most frequently in ticks, followed by Borrelia garinii and Borrelia lusitaniae. Among birds infested by infected ticks, 23% (6/26) were infested by B. lusitaniae-infected larvae. Migratory birds appear to be reservoir hosts for B. lusitaniae.     

Application of Acoustic Telemetry to Assess Residency and Movements of Rockfish and Lingcod at Created and Natural Habitats in Prince William Sound

Loss and/or degradation of nearshore habitats have led to increased efforts to restore or enhance many of these habitats, particularly those that are deemed essential for marine fishes. Copper rockfish (Sebastes caurinus) and lingcod (Ophiodon enlongatus) are dominant members of the typical reef fish community that inhabit rocky and high-relief substrates along the Pacific Northwest. We used acoustic telemetry to document their residency and movements in the nearshore waters of Prince William Sound, Alaska in order to assess use of created reef habitat in an individual-based manner. A total of 57 fish were surgically implanted with acoustic transmitters. Forty-five fish were captured and monitored in three habitats: artificial reef, low-relief natural reef, and patchy high-relief natural reef. Within each habitat, both rockfish and lingcod exhibited long periods of residency with limited movements. Twelve rockfish were captured at the natural reefs and displaced a distance of 4.0 km to the artificial reef. Five of the 12 rockfish returned within 10 d of their release to their initial capture site. Another five of the 12 displaced fish established residency at the artificial reef through the duration of our study. Our results suggest the potential for artificial reefs to provide rockfish habitat in the event of disturbances to natural habitat.     

Predator Crown-of-Thorns Starfish (Acanthaster planci) Outbreak,Mass Mortality of Corals,and Cascading Effects on Reef Fish and Benthic Communities

Outbreaks of the coral-killing seastar Acanthaster planci are intense disturbances that can decimate coral reefs. These events consist of the emergence of large swarms of the predatory seastar that feed on reef-building corals, often leading to widespread devastation of coral populations. While cyclic occurrences of such outbreaks are reported from many tropical reefs throughout the Indo-Pacific, their causes are hotly debated, and the spatio-temporal dynamics of the outbreaks and impacts to reef communities remain unclear. Based on observations of a recent event around the island of Moorea, French Polynesia, we show that Acanthaster outbreaks are methodic, slow-paced, and diffusive biological disturbances. Acanthaster outbreaks on insular reef systems like Moorea''s appear to originate from restricted areas confined to the ocean-exposed base of reefs. Elevated Acanthaster densities then progressively spread to adjacent and shallower locations by migrations of seastars in aggregative waves that eventually affect the entire reef system. The directional migration across reefs appears to be a search for prey as reef portions affected by dense seastar aggregations are rapidly depleted of living corals and subsequently left behind. Coral decline on impacted reefs occurs by the sequential consumption of species in the order of Acanthaster feeding preferences. Acanthaster outbreaks thus result in predictable alteration of the coral community structure. The outbreak we report here is among the most intense and devastating ever reported. Using a hierarchical, multi-scale approach, we also show how sessile benthic communities and resident coral-feeding fish assemblages were subsequently affected by the decline of corals. By elucidating the processes involved in an Acanthaster outbreak, our study contributes to comprehending this widespread disturbance and should thus benefit targeted management actions for coral reef ecosystems.     

The influence of light on the twilight migrations of grunts

Synopsis Behaviors that precede the daily migrations of mixed-species schools of juvenile grunts (Pomadasyidae), from patch reefs to grass beds at dusk and vice versa at dawn, are defined and utilized to ascertain the precision of the migrations. Although premigratory behaviors differ at dusk and dawn, the migrations are precise twilight events which occur at the same light intensities during dawn and dusk. Histological sections of the retina reveal that both cones and rods are fully exposed to ambient light during the migrations. Under the difficult photic conditions that prevail during migration, the retina is structured photomechanically to maximize the absorption of ambient light. Body colorations of the grunts, which consist mostly of intense colored stripes during the day, are replaced at night by cryptic melanic patterns. The precision of migration, the photomechanical movements in the retina, and the changes in body coloration are considered adaptive because they reduce predation on grunts when they migrate and are most vulnerable to attack. In support of this conclusion, the migrations take place just before the evening and just after the morning quiet period - thus they avoid that period during twilight when predation is highest in tropical fish communities.     

Acoustic communication in an electric fish,Pollimyrus isidori (Mormyridae)

It has been known since von Frisch's work in the 1930's that mormyrid electric fishes are quite sensitive to sound. We now describe a repertoire of natural sounds produced by the mormyrid, Pollimyrus isidori, during breeding and aggression; reception of communication sounds is probably a major function for mormyrid audition. In aquaria, Pollimyrus isidori produce 'grunts', 'moans', 'growls', 'pops' and 'hoots' at various phases during nesting, courtship, and territory defense. All five sounds are produced primarily at night. Territorial males produce grunts, moans and growls during courtship. Vocalizing is stimulated by the presence of a gravid female on the male's territory and decreases with the onset of spawning. Hoots and pops are given during agonistic behavior. Grunts are bursts of acoustic pulses, stereotyped for an individual, with the potential as individual signatures. The electric organ is silent during grunts and moans and is discharged at a reduced rate during growls. The courtship and spawning of Pollimyrus isidori is described.     

Distinctly different behavioral responses of a copepod,Temora longicornis,to different strains of toxic dinoflagellates,Alexandrium spp.

Zooplankton responses to toxic algae are highly variable, even towards taxonomically closely related species or different strains of the same species. Here, the individual level feeding behavior of a copepod, Temora longicornis, was examined which offered 4 similarly sized strains of toxic dinoflagellate Alexandrium spp. and a non-toxic control strain of the dinoflagellate Protoceratium reticulatum. The strains varied in their cellular toxin concentration and composition and in lytic activity. High-speed video observations revealed four distinctly different strain-specific feeding responses of the copepod during 4 h incubations: (i) the ‘normal’ feeding behavior, in which the feeding appendages were beating almost constantly to produce a feeding current and most (90%) of the captured algae were ingested; (ii) the beating activity of the feeding appendages was reduced by ca. 80% during the initial 60 min of exposure, after which very few algae were captured and ingested; (iii) capture and ingestion rates remained high, but ingested cells were regurgitated; and (iv) the copepod continued beating its appendages and captured cells at a high rate, but after 60 min, most captured cells were rejected. The various prey aversion responses observed may have very different implications to the prey and their ability to form blooms: consumed but regurgitated cells are dead, captured but rejected cells survive and may give the prey a competitive advantage, while reduced feeding activity of the grazer may be equally beneficial to the prey and its competitors. These behaviors were not related to lytic activity or overall paralytic shellfish toxins (PSTs) content and composition and suggest that other cues are responsible for the responses.     

Temporal variation and characterization of grunt sounds produced by Atlantic cod Gadus morhua and pollack Pollachius pollachius during the spawning season

Fine‐scale temporal patterning in grunt production and variation in grunt attributes in Atlantic cod Gadus morhua and pollack Pollachius pollachius was examined. Pollachius pollachius produced only a single sound type, the grunt, similar to that previously described for G. morhua. Sound production and egg production were correlated in P. pollachius but not in G. morhua. Only G. morhua displayed a strongly cyclical pattern, producing more grunts at night. Finer‐scale temporal patterning in grunt production was observed in both species which produced significantly fewer grunts following a period of high grunt production. These quieter periods lasted up to 45 min for P. pollachius and up to 1 h in G. morhua. Grunts were not always produced in isolation but organized into bouts in both species. Longer bouts were more frequent during periods of increased sound activity and were linked with changes in grunt characteristics including increased grunt duration, pulse duration and repetition period of each pulse combined with decreased dominant frequency. This study provides the first evidence of acoustic signalling being used by spawning P. pollachius and presents the most detailed analysis of the complexity of gadoid sound production.     

Predation Risk,Resource Quality,and Reef Structural Complexity Shape Territoriality in a Coral Reef Herbivore

For many species securing territories is important for feeding and reproduction. Factors such as competition, habitat availability, and male characteristics can influence an individual’s ability to establish and maintain a territory. The risk of predation can have an important influence on feeding and reproduction; however, few have studied its effect on territoriality. We investigated territoriality in a haremic, polygynous species of coral reef herbivore, Sparisoma aurofrenatum (redband parrotfish), across eight reefs in the Florida Keys National Marine Sanctuary that were either protected or unprotected from fishing of piscivorous fishes. We examined how territory size and quality varied with reef protection status, competition, predation risk, and male size. We then determined how territory size and quality influenced harem size and female size to understand the effect of territoriality on reproductive potential. We found that protected reefs trended towards having more large predatory fishes and that territories there were smaller but had greater algal nutritional quality relative to unprotected reefs. Our data suggest that even though males in protected sites have smaller territories, which support fewer females, they may improve their reproductive potential by choosing nutritionally rich areas, which support larger females. Thus, reef protection appears to shape the trade-off that herbivorous fishes make between territory size and quality. Furthermore, we provide evidence that males in unprotected sites, which are generally less complex than protected sites, choose territories with higher structural complexity, suggesting the importance of this type of habitat for feeding and reproduction in S. aurofrenatum. Our work argues that the loss of corals and the resulting decline in structural complexity, as well as management efforts to protect reefs, could alter the territory dynamics and reproductive potential of important herbivorous fish species.