Predicting structural complexity of reefs and fish abundance using acoustic remote sensing (RoxAnn)

This study determined whether the acoustic roughness of Caribbean reef habitats is an accurate proxy for their topographic complexity and a significant predictor of their fish abundance. Fish abundance was measured in 25 sites along the forereef of Glovers Atoll (Belize). At each site, in situ rugosity (ISR) was estimated using the “chain and tape” method, and acoustic roughness (E1) was acquired using RoxAnn. The relationships between E1 and ISR, and between both E1 and ISR and the abundance of 17 common species and the presence of 10 uncommon species were tested. E1 was a significant predictor of the topographic complexity (r ² = 0.66), the abundance of 10 common species of surgeonfishes, pomacentrids, scarids, grunts and serranids and the presence of 4 uncommon species of pomacentrids and snappers. Small differences in E1 (i.e. ∆0.05–0.07) reflected in subtle but significant differences in fish abundance (~1 individual 200 m⁻² and 116 g 200 m⁻²) among sites. Although we required the use of IKONOS data to obtain a large number of echoes per site, future studies will be able to utilise RoxAnn data alone to detect spatial patterns in substrate complexity and fish abundance, provided that a minimum of 50 RoxAnn echoes are collected per site.     

Feeding macroecology of territorial damselfishes (Perciformes: Pomacentridae)

The present study provides the first analysis of the feeding macroecology of territorial damselfishes (Perciformes: Pomacentridae), a circumtropical family whose feeding and behavioral activities are important in structuring tropical and subtropical reef benthic communities. The analyses were conducted from data collected by the authors and from the literature. A strong positive correlation was observed between bite rates and sea surface temperature (SST) for the genus Stegastes. A negative correlation was found between bite rates and mean body size for the genera Stegastes and Pomacentrus, but this relationship was not significant when all territorial pomacentrids were analyzed together. A negative correlation between body size and SST was observed for the whole group and for the genera Stegastes, and Pomacentrus. No relationship was found between territory size and feeding rates. Principal Components Analysis showed that differences in feeding rates accounted for most of the variability in the data. It also suggested that body size may be important in characterizing the different genera. In general, tropical species are smaller and have higher bite rates than subtropical ones. This study extended the validity of Bergmann’s rule, which states that larger species or larger individuals within species occur towards higher latitudes and/or lower temperatures, for an important group of reef fishes. The identification of large-scale, robust ecological patterns in the feeding ecology of pomacentrid fishes may establish a foundation for predicting large-scale changes in reef fish assemblages with expected future changes in global SST.     

Biochemical systematics of fishes of the genus Stegastes (Pomacentridae) from the Southern Marianas

The biochemical systematics of the subset of species of Stegastes (Pisces: Pomacentridae) available from coral reefs surrounding Guam, USA, was studied using three species of Pomacentrus as the outgroup. Analyses of allozymic variation at 14 presumptive loci (10 of which were polymorphic) using various evolutionary tree-construction techniques identified S. fasciolatus as the sister species of an assemblage comprised of S. albifasciatus, S. nigricans, and S. lividus: within this assemblage, S. lividus is the sister species of S. albifasciatus and S. nigricans, the pair of species which shared the most synapomorphic alleles and consistently exhibited the lowest degree of genetic distance relative to estimates for the other pairs of species studied. The consistency of ingroup topologies produced by parsimony analyses using Manhattan rectilinear distances computed from allele frequencies with those produced using the distance Wagner procedure and UPGMA (unweighted pair-group method of arithmetic averages) clustering, substantiates strongly the proposed phylogeny. The systematic scheme inferred from molecular data is concordant with predictions based on morphologic differences, i.e., differences in numbers of dorsal spines, that have been used as evidence in support of the proposal to partition Stegastes into at least two subgenera.     

Underwater observations on the reproductive activity of the demoiselleChromis dispilus (Pisces: Pomacentridae)

Reproductive behaviour of the demoiselleChromis dispilus Griffin was studied along the coast and offshore islands of north-eastern New Zealand. Spawning occurred at intervals over the summer months, December to March. Males displayed nursery behaviour and nest-guarding during the incubation period of the eggs. Juvenile growth was initially very rapid. A comparison of reproductive activities with other species ofChromis show a basic similarity in behaviour, although the demoiselle appears to differ in certain behavioural patterns associated with spawning.     

Variability in larval development of the tropical reef fish Pomacentrus amboinensis (Pomacentridae): the parental legacy

The effects of food availability, female size, and social interactions on the quality of Pomacentrus amboinensis larvae at hatching were examined using two field-based experiments. In Experiment 1, food availability and female size significantly influenced size, eye diameter and levels of yolk reserves of larvae at hatching. Small females (47 to 52 mm standard length, SL) whose diets were not supplemented, produced the longest larvae (3.0 ± 0.01 mm total length, TL) with the least yolk reserves (50.1 ± 1.04 μm²). Irrespective of female size, those that received additional food produced larvae with the largest yolk-sacs (large females: 87.60 ± 1.53 μm²; small females: 80.14 ± 1.24 μm²). In Experiment 2, interactions with conspecifics had a greater affect on the somatic development of larvae at hatching than food availability. Increased social interactions resulted in larvae that were ⋍3% longer, with 2% greater head depth, than larvae from females that spawned in isolation on the experimental reefs. Fed females produced larvae with ⋍20% more yolk than larvae from females whose diets were not supplemented. All three factors (food availability, female size, and intensity of social interactions) tested within these experiments vary spatially and temporally among reefs. There is the potential, therefore, for larvae at the onset of the planktonic stage to vary in quality, level of development, and probability of survival. Received: 12 August 1996 / Accepted: 26 August 1996     

Planktonic larval duration of one hundred species of Pacific and Atlantic damselfishes (Pomacentridae)

The plankton larval duration for 100 species of Pacific and Atlantic damselfishes was estimated from daily growth increments on the otolith of juvenile fish collected at various localities between July 1987 and September 1988. For newly-settled fishes, larval duration was determined by counting the entire number of increments present on the otolith, while for older juveniles estimates were made by counting the number of increments between the center of the otolith and a mark corresponding to settlement. We document the development of otolith formation during the period when eggs are incubated on the reef and show that daily increments are only accreted after larvae hatch and enter the planktonic phase. The planktonic larval duration for damselfish is shorter and less variable, both between and within species, compared to other groups of reef fishes such as wrasses and surgeonfishes. Larval duration ranged from 12 to 39 d. Average duration between species ranged from 13.1 to 35.2 d. The time spent in the plankton was not significantly correlated with geographic distribution when evaluated among species, however, genera with confined regional distribution have a shorter mean larval life than do widely distributed genera. Size at settlement was positively correlated with time spent in the plankton among species, but a significant correlation between these variables was only evident within one of ten species. The low variance in planktonic larval duration within species indicates that most damselfish are unable to delay metamorphosis following competency. This inability to postpone settlement limits the potential for dispersal, especially when dispersal time between suitable habitats is greater than about 30 d.     

Genetic relationships of the colour morphs of Acanthochromis polyacanthus (Pomacentridae) on the northern Great Barrier Reef

Acanthochromis polyacanthus is an unusual tropical damselfish because it lacks pelagic larvae and has lost the capacity for broad-scale dispersal among coral reefs. On the modern Great Barrier Reef, this fish has multiple colour morphs and we have previously described steep genetic clines in the central region between three colour morphs. A similar transition of colour morphs has been found in the far northern region (Latitude 11°S), where at least five colour morphs and intergrading hybrids were found during a survey of 14 locations on nine adjacent reefs. On two of these reefs, monochromatic white and bicoloured morphs coexist, partitioning the reef slope vertically when sympatric, but occupying the full depth range elsewhere. In areas of overlap, they interbreed, producing hybrids of intermediate colour. Although the discriminatory power of our genetic analysis was weakened by the low level of variability in these populations (only four polymorphic loci), allozyme electrophoresis revealed significant genetic differentiation within and among reefs. There was neither simple linkage between colour and protein variation, nor correlation between genetic similarity and proximity of sampled populations. We suggest that the observed chaotic structure is the result of differences among the founding populations, although the two colour-based stocks have been homogenized through a long history of introgression during multiple episodes of isolation and secondary contact. Despite genetic mixing, phenotypic diversity (i.e. colour morphs) seems to have been preserved in these populations by sexual selection, perhaps reinforced by natural selection. The long-term stability of this outcome, however, is suggested by the presence of unique morphs on two reefs that are detached from the continental shelf and surrounded by very deep water. The strong identity of these populations can be attributed to long occupation of these isolated sites compared with those on the continental shelf which have only been established in the last 15 000 yr and still retain differences arising from the initial founder events. Received: 15 January 1997 / Accepted: 15 August 1997     

Pleistocene diversification of the Pomacentrus coelestis species complex (Pisces: Pomacentridae): historical biogeography and species boundaries

Pleistocene eustatic changes in sea level are often invoked to explain genetic divergence among marine organisms. However, molecular phylogenies have revealed relatively few examples of speciation events dating to the Pleistocene. We present a species-level hypothesis of the timing of evolution for the Pomacentrus coelestis species complex (Pomacentridae), based on the nuclear S7 intron and the mitochondrial Cytb gene, and reconstruct ancestral range distributions across the timetree. Ancestral range reconstruction suggests the complex originated in the Coral Triangle and East Indian Ocean, with subsequent range expansion outward from this region. We suggest that land barriers to dispersal (e.g., Indo-Pacific barrier) may be responsible for the divergence between Indian (P. alleni, P. similis, P. caeruleopunctatus, and P. caeruleus) and Pacific (P. micronesicus, P. auriventris, and P. coelestis) species groups, and subsequent isolation by Pleistocene sea-level fluctuations in certain areas of the Coral Triangle (glacial refugia) may play an important role in the diversification of this species complex. Additionally, our analyses show cryptic lineages within P. micronesicus and highlight the need for comprehensive sampling within and among species to reveal recent speciation events.     

Mating and its effect on acoustic signalling behaviour in a primitive orthopteran,Cyphoderris strepitans (Haglidae): the cost of feeding females

Summary Males of the primitive orthopteran, Cyphoderris strepitans, provide their mates with two types of nuptial food gift during mating: 1) females feed while coupled on the fleshy metathoracic wings of the male and the resultant flow of haemolymph and 2) the spermatophore transferred by the male includes a gelatinous spermatophylax which the female eats after mating. During the peak breeding interval, virgin males secure significantly more matings than their numbers relative to non-virgin males would predict. We tested the hypothesis that non-virgin males, having lost a substantial portion of available energy through previous investment in females, call significantly less than virgin males. Reduced calling should result in the attraction of fewer females and/or a greater risk of intrusion from competing males and consequently, a lowered mating success. Calling activity of male C. strepitans of varying mating status was monitored with a sound activated relay apparatus for two consecutive nights following their capture. Males were of three experimental groups, virgin, freshly wounded males (mated on the same night of capture), and old-wound males (mated at least one night prior to capture). Our cata showed a significant short-term reduction in signalling activity as a consequence of mating. Whereas there was no difference in the time spent calling by virgins on the first and second nights following capture, freshly wounded and oldwound males called significantly less on the first night than on the second. Furthermore, the proportion of males calling on both nights was significantly greater for the virgin and old-wound groups than for freshly wounded males. When the duration over which males called on the first night was compared with that of the second, virgin males showed no difference whereas old-wound males called for significantly shorter intervals on the first night. These results indicate that the cost of a large nutrient investment in females lowers the energy level in males below the threshold required for a prolonged signalling period. After a refractory interval during which males feed and replenish their energy reserves, calling levels equivalent to those which occur prior to mating are regained. Active female choice may also contribute to the reduced mating success of non-virgins, but this possibility remains untested.     

Auditory and olfactory abilities of pre-settlement larvae and post-settlement juveniles of a coral reef damselfish (Pisces: Pomacentridae)

The propagules of most species of reef fish are advected from the reef, necessitating a return to reef habitats at the end of the pelagic stage. There is increasing evidence of active attraction to the reef but the sensory abilities of reef fish larvae have not been characterized well enough to fully identify cues. The electrophysiological methods of auditory brainstem response (ABR) and electroolfactogram (EOG) were used to investigate auditory and olfactory abilities of pre- and post-settlement stages of a damselfish, Pomacentrus nagasakiensis (Pisces, Pomacentridae). Audiograms of the two ontogenetic stages were similar. Pre-settlement larvae heard as well as their post-settlement counterparts at all but two of the tested frequencies between 100 Hz and 2,000 Hz. At 100 and 600 Hz, pre-settlement larvae had ABR thresholds 8 dB higher than those of post-settlement juveniles. Both stages were able to detect locally recorded reef sounds. Similarly, no difference in olfactory ability was found between the two ontogenetic stages. Both stages showed olfactory responses to conspecifics as well as L-alanine. Therefore, the auditory and olfactory senses have similar capabilities in both ontogenetic stages. Settlement stage larvae of P. nagasakiensis can hear and smell reef cues but it is unclear as to what extent larvae use these sounds or smells, or both, as cues for locating settlement sites. An erratum to this article can be found at     

Photographic and acoustic tracking observations of the behaviour of the grenadier Coryphaenoides (Nematonurus) armatus the eel Synaphobranchus bathybius,and other abyssal demersal fish in the North Atlantic Ocean

Using an autonomous free-fall vehicle (AU-DOS), observations were made of demersal fish attracted to baits and baited acoustic transmitters at two stations in the North Atlantic Ocean. A comparison was made between Station PAP (48°50N; 16°30W), 4800 m deep on the Porcupine Abyssal Plain which is relatively eutrophic, and Station MAP (31°N; 20°W), 4900 m deep on the Madeira Abyssal Plain, which is oligotrophic. Experiments were conducted during summer, in 1989 and 1990. Four species of fish were observed at Station MAP, the grenadier, Coryphaenoides (Nematonurus) armatus, the eel, Synaphobranchus bathybius, and the ophidiids Spectrunculus grandis, and Barathrites sp. At Station PAP, C. (N.) armatus and H. (S.) bathybius were attracted to bait on all deployments and only two other individuals of different species, probably ophidiids, were seen. The mean first grenadier arrival time was 30 and 138 min at Stations PAP and MAP, respectively. Mean first eel arrival time was 29 and 151 min at Stations PAP and MAP, respectively. Estimated population densities of fish were 167 grenadiers km^-2 and 180 synaphobranchid eels km^-2 at Station PAP and 8 grenadiers km^-2 and 7 eels km^-2 at Station MAP. Only the grenadier C. (N.) armatus definitely ingested transmitters, and this species dominated fish activity around the baits. Mean time of departure of grenadiers with transmitters in their stomachs across an acoustic horizon at 1000 m range was 371 and 488 min at Stations PAP and MAP, respectively. Grenadiers had a longer mean staying time at the food source at the more oligotrophic Station MAP (364 min) than at Station PAP (141 min). This corresponds with predictions of optimal foraging theory.     

Competition for the home and aggressive behaviour in the chiton Acanthopleura gemmata (Blainville) (Mollusca: Polyplacophora)

Summary Computerized screening of all the positions recorded during a synodic month on 120 individually marked chitons (Acanthopleura gemmata) pinpointed their preferential resting points. Unlike the majority of intertidal chitons so far studied, A. gemmata rests in well-defined homes actively dug in the rock. Homes proved to be not strictly individual and periodically interchangeable. A complex aggressive behaviour was recorded in the field when two animals came in contact at home. When competing for ownership of a resting site rival chitons may suppress their nocturnal feeding activity. Despite its strong home-related territoriality A. gemmata showed no mutual exclusion on the feeding area. The highly specialized resting habits clearly protects A. gemmata from its most important predator, the toad fish Arothron immaculatus. The behaviour of A. gemmata is compared to that of other chitons and gastropods, and the current hypotheses concerning the adaptive value of the homing behaviour in littoral molluscs are discussed.     

Genetic divergence of Maldivian and Micronesian demes of the damselfishes Stegastes nigricans,Chrysiptera biocellata,C. glauca and C. leucopoma (Pomacentridae)

Samples representing the damselfishes (Perciformes: Pomacentridae) Stegastes nigricans, Chrysiptera biocellata, C. glauca, and C. leucopoma from North Male Atoll, Republic of Maldives, and Guam, Southern Marianas, were surveyed electrophoretically. A sample representing C. leucopoma from Ngemelis Island, Republic of Palau, was included in the analysis to provide a within-Micronesia comparison. On average, products of 23 presumptive protein-coding loci were examined in each sample. In comparisons of Maldivian and Guamanian samples, absolute or virtually fixed allelic differences were detected at: (1) ADA ^* and PEPS ^* for Stegastes nigricans; (2) ADA ^*, AAT-1 ^*, AAT-2 ^*, and mMDH ^* for C. biocellata, and; (3) ADA ^*, AAT-1 ^*, and AAT-2 ^* for C. glauca. At three polymorphic loci, ADA ^*, sMDH-1 ^*, and sMDH-2 ^*, similar allele frequencies were found in Palauan and Guamanian samples representing C. leucopoma, suggesting that these samples share a gene pool. Significantly different allele frequencies at ADA ^* and sMDH-1 ^* and a virtually fixed allelic difference at sMDH-2 ^* were detected between the Maldivian samples and each of the Micronesian samples representing C. leucopoma. The observed patterns of allele frequency differentiation suggest that Maldivian and Micronesian samples of each of the four study species represent separate demes.     

Development of the nasal olfactory organs in the larvae, settlement-stages and some adults of 14 species of Caribbean reef fishes (Labridae, Scaridae, Pomacentridae)

Larval fishes likely use a variety of settlement cues to locate and navigate toward the habitats they will inhabit as juveniles. Information about the morphology and state of development of the sensory organs of larval stages of fishes provides insight into their capabilities at the time of settlement. The peripheral olfactory organ of wild-caught late-stage larvae and early juveniles and some adults of 14 species of the Caribbean reef fishes wrasses (Labridae), parrot fishes (Scaridae) and damselfish (Pomacentridae) were examined using scanning electron microscopy and compared in terms of settlement specificity. Ages in days after hatching and days post-settlement were determined from the otoliths. Morphology of the nares and the olfactory epithelium are described for these species by stage. The separation of the anterior and posterior nares occurred before settlement in the labrids but in some specimens of scarids this separation was not complete by the time of settlement. Densities of ciliated and microvillous receptor cells and non-sensory ciliated epithelial cells were calculated. Densities of ciliated receptor cells ranged from 0.389 μm⁻² in a specimen of Thallasoma bifasciatum to 0.0057 μm⁻² in Bodianus rufus and of microvillous receptor cells from 0.038 μm⁻² in a Clepticus parrae juvenile to 0.266 μm⁻² in a juvenile Doratonotus megalepis. Densities of non-sensory cilia, also associated with high olfactory ability, were also high. The olfactory organ in wrasses is well developed prior to settlement and is comparable to that of adult fishes. The possible role of olfaction in larval schooling, reef cue detection and orientation toward habitat at settlement is discussed.

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Locomotory behaviour and swimming performance of the Norway lobster,Nephrops norvegicus,in the presence of an acoustic tag

Observations have been made on the locomotory behaviour and swimming performances of the Norway lobster, Nephrops norvegicus (L.), fitted with an acoustic transmitter ventrally under the cephalothorax. The walking behaviour of adult males (44 mm carapace length) appeared to be unaffected, but the tag caused significant reductions in certain measures of tail-flip swimming performance such as swimming speed and endurance. Flume-tank experiments in low water currents suggested that the transmitter would increase hydrodynamic drag during swimming by 9 to 32%, depending on lobster size. Given the weight and dimensions of the acoustic transmitters currently available, it is considered advisable to confine acoustic tracking studies to relatively large N. norvegicus.Correspondence to: C. J. Chapman     

Stranding and matting behaviour in Nyctiphanes australis (Euphausiidae: Crustacea)

Numerous stranding events of the euphausiid Nyctiphanes australis Sars on the coast of south-east Tasmania during the winter months of 1984/1985 are documented. The krill were at a reproductive stage of their life cycle. Strandings almost always occurred on calm, sunny days on beaches with a northerly aspect. Typically, barometric pressure was greater than and wind speed was less than the monthly average. No significant differences were found between stranded euphausiids and those from surface swarms, and the relationship between the two forms of behaviour in this area is discussed. A new form of behaviour termed matting was observed, in which the euphausiids aggregate dorsal-side down on the substrate in shallow water. Matting usually occurs synchronously with stranding. Laboratory experiments showed that N. australis responds to changes in both the plane of polarization and intensity of light with behaviour typical of that observed in stranding and matting events. Light appears to be a key factor mediating these types of behaviour. Hypotheses for the relationship between stranding and matting are offered.