The adaptive significance of larval dispersal in coral reef fishes

Synopsis Coral reef fishes almost universally disperse over relatively great distances during a pelagic larval phase. Barlow (1981) suggested that this dispersal is adaptive because adult fishes inhabit a patchy, uncertain environment. This reiterated an older idea that the random extinction of local populations necessarily favours dispersal, since ultimately all populations of non-dispersers will disappear. Whereas this view is based on adult survival, we emphasize a less frequent view that substantial larval dispersal may be adaptive when offspring experience patchy and unpredictable survival in the pelagic habitat. We do not address the question of why these animals ‘broadcast’ rather than ‘brood’, but suggest that species committed to pelagic offspring will be under selection to disperse siblings to spread the risk of failure among members of a cohort. Our arguments are supported by a heuristic computer simulation.     

Influence of salinity on the larval development of the fiddler crab <Emphasis Type="Italic">Uca vocator</Emphasis> (Ocypodidae) as an indicator of ontogenetic migration towards offshore waters

Larvae of many marine decapod crustaceans are released in unpredictable habitats with strong salinity fluctuations during the breeding season. In an experimental laboratory study, we investigated the influence of seven different salinities (0, 5, 10, 15, 20, 25 and 30) on the survival and development time of fiddler crab zoea larvae, Uca vocator, from northern Brazilian mangroves. The species reproduces during the rainy season when estuarine salinity strongly fluctuates and often reaches values below 10 and even 5. Salinity significantly affected the survival rate and development period from hatching to megalopa, while the number of zoeal stages remained constant. In salinities 0 and 5, no larvae reached the second zoeal stage, but they managed to survive for up to 3 (average of 2.3 days) and 7 days (average of 5.1 days), respectively. From salinity 10 onwards, the larvae developed to the megalopal stage. However, the survival rate was significantly lower (5–15%) and development took more time (average of 13.5 days) in salinity 10 than in the remaining salinities (15–30). In the latter, survival ranged from 80–95% and development took 10–11 days. Given the 100% larval mortality in extremely low salinities and their increased survival in intermediate and higher salinities, we conclude that U. vocator has a larval ‘export’ strategy with its larvae developing in offshore waters where salinity conditions are more stable and higher than in mangrove estuaries. Thus, by means of ontogenetic migration, osmotic stress and resulting mortality in estuarine waters can be avoided.     

Functional response of suspension feeding anuran larvae to different particle sizes at low concentrations (Amphibia)

The influence of particle size, initial particle concentration and larval stage on the ingestion rate, ‘retention efficiency’, and filtering rate of anuran larvae with varying filter apparatus anatomy and different life histories was investigated for four species. Larvae of premetamorphic Stages 28 and 32 and prometamorphic Stage 40 were selected for filtering experiments on the basis of their different growth rates. Three different sizes of silica gel particles were offered as mock food. Particle concentration was measured photometrically. The Michaelis-Menten model was used to describe the dependency of ingestion rate, filtering rate, and ‘retention efficiency’ upon initial particle concentration, and to calculate maximum ingestion rate, threshold concentration, and the half-saturation constant. (1) The highest ingestion rates, filtering rates and ‘retention efficiencies’ were achieved by Xenopus laevis larvae, followed by Bufo calamita larvae. Bufo bufo larvae lay at the opposite end of the scale. Rana temporaria larvae were placed between B. calamita and B. bufo larvae. This order is attributed to differences in life histories, especially the different breeding environments in which these larvae occur. (2) The larger the particle size and the older the stage, the greater the tendency toward saturation of the ingestion rate, filtering rate and ‘retention efficiency’. These filtration parameters are graded according to particle size. The ingestion rate (number of particles), filtration rate and ‘retention efficiency’ are greatest for PS3. Ingestion volume is greatest for PS 1. The difference between PS3 and PS2 on the one hand, and PS1 on the other, is often great; for Stage 28 X. laevis it is very great. This shows that larvae ingest large particles more effectively, and that the most effective ingestion takes place at Stages 28 and 32, owing to the growth function of these stages. The ability of larvae to ingest large particles effectively is possibly a very basic phylogenetic characteristic. (3) The threshold concentration is lowest when the particles are at their largest. In accordance with conclusions drawn by other authors, threshold feeding is attributed to regulation by buccal pumping and mucus production. Considerable importance is attributed to threshold feeding with respect to larval adaptation to oligotrophic environments.     

Ontogeny of behaviour in larvae of marine demersal fishes

The development of behaviours that are relevant to larval dispersal of marine, demersal fishes is poorly understood. This review focuses on recent work that attempts to quantify the development of swimming, orientation, vertical distribution and sensory abilities. These behaviours are developed enough to influence dispersal outcomes during most of the pelagic larval stage. Larvae swim in the ocean at speeds similar to the currents found in many locations and at 3–15 body lengths per second (BL s⁻¹), although, based on laboratory measurements, species from cold environments swim slower than those from warm environments. At least in warm-water species, larvae swim in an inertial hydrodynamic environment for most of their pelagic period. Unfed swimming endurance is >10 km from about 8–10 mm, and reaches more than 50 km before settlement in several species. Larval fishes are efficient swimmers. In most species, a large majority of larvae have orientated swimming in the ocean, but the precision of orientation does not improve with growth. Swimming direction of the larvae frequently changes ontogenetically. Vertical distribution changes ontogenetically in most species, and both ontogenetic ascents and descents are found. Development of schooling is poorly understood, but it may influence speed, orientation and vertical distribution. Sensory abilities (hearing, olfaction, vision) form early, are well developed and are able to detect cues relevant to orientation for most of the pelagic larval stage. All this indicates that the passive portion of the pelagic larval duration will be short, at least in most warm-water species, and that behaviour must be taken into account when considering dispersal, and in particular in dispersal models. Although quantitative information on the ontogeny of some behaviours is available for a relatively small number of species, more research in this field is required, especially on species from colder waters.     

A Second Look at Mitochondrial DNA Variability in European Anchovy (Engraulis encrasicolus): Assessing Models of Population Structure and the Black Sea Isolation Hypothesis

Genetic architectures of marine fishes are generally shallow because of the large potential for gene flow in the sea. European anchovy, however, are unusual among small pelagic fishes in showing large differences among sub-basins and in harbouring two mtDNA phylogroups (‘A’ & ‘B’), representing 1.1–1.85 million years of separation. Here the mtDNA RFLP dataset of Magoulas et al. [1996, Mol. Biol. Evol. 13: 178–190] is re-examined to assess population models accounting for this subdivided population structure and to evaluate the zoogeographical origins of the two major phylogroups. Haplotype and nucleotide diversities are highest in the Ionian Sea and lowest in the Aegean and Black seas. However, this gradient is absent when ‘A’ and ‘B’ haplotypes are examined separately. Neither the self-sustaining nor the basin population models adequately describe anchovy population behaviour. Tests for neutrality, mismatch and nested clade analyses are concordant in depicting recent expansions of both phylogroups. Unimodel mismatch distributions and haplotype coalescences dating to the last (Eemian) interglacial (‘B’) and the Weichselian pleniglacial period (‘A’) indicate separate colonizations of the Mediterranean Basin. Phylogroup ‘A’ is unlikely to have arisen through continuous long-term isolation in the Black Sea because of climate extremes from displaced subpolar weather systems during the ice ages. Ancestors of both groups appear to have colonized the Mediterranean from the Atlantic in the late Pleistocene. Hence, zoogeographic models of anchovy in the Mediterranean must also include the eastern (and possibly southern) Atlantic.     

Beta diversity of a Gulf of California rocky-shore fish community

Synopsis Reef fish community composition in three segments of a peninsular rocky shoreline in the Gulf of California was estimated over four periods by visual observation. ‘Point’ and ‘bay’ segments had regular and distinct species compositions over most periods while a ‘middle’ segment was least distinct but consistently had the greatest number of species. Compositional change along the peninsula was least regular during the coldest sea temperature period (April). Mean species turnover between segments was highest between point and bay. Within segments, the point had greater compositional predictable composition (lowest species turnover). When species with regular frequency of encounter were classified into ‘point’, ‘middle’, ‘bay’, and ‘no trend’ groups it was found that comparatively more ‘point’ species had pelagic eggs and comparatively more ‘bay’ species had demersal eggs. Beta diversity of rocky-shore fishes along the physical gradient of the Punta Doble peninsula reflects a transition between exposed and protected rocky shoreline communities. The correlated physical environmental characteristics associated with exposed and protected habitats are discussed in relation to diversity regulation and dispersal strategies in reef fishes.     

Recruitment and post-recruit immigration affect the local population size of coral reef fishes

 This study quantifies the contributions of larval recruitment and post-recruit (juvenile and adult) immigration to net increases in population size for 150 species of fishes found on ten isolated coral patches or ‘bommies’ (108–267 m²) within a typical reef of the Great Barrier Reef system. At least one third of the total number of recruits and immigrants to all bommies were post-recruit fishes, and movement between bommies in 136 species was detected at some time during the 22 month sampling period. The relative numbers of recruits and post-recruit immigrants per species varied widely within the assemblage, and between the replicate bommies. Populations of 95 species received both types of immigrants, 41 species had only post-recruit immigrants, and 14 species received only larval recruitment. In most species, recruitment occurred over the austral summer between October and February, while post-recruit movements occurred in both summer and winter. Rates of post-recruit immigration varied temporally within bommies, and pulses of post-recruits were less temporally concordant between bommies than pulses of recruits. This study is further evidence that post-settlement processes can have a significant effect on the local population size of reef fishes. Accepted: 30 January 1997     

Seasonality in fish larval assemblage structure within marine reef National Parks in coastal Kenya

Variability of larval assemblages within marine habitats has an effect on population structure of adults, and on patchiness of populations in space and time. However, despite this effect, few studies have examined this variability in the Western Indian Ocean (WIO) especially in relation to marine park boundaries. In this study, we tested the hypothesis that seasonality and biophysical variables influence fish larval assemblage structure within two of the oldest marine parks in Kenya of differing reef structure. Monthly samples were obtained from the parks using oblique plankton tows for 14–24 months. A total of 38 families and 52 genera of fish larvae were obtained from Malindi Marine Park, while 20 genera in 20 families were sampled from the more enclosed Watamu Marine Park but with a connection to an estuarine system. The preflexion larvae comprised over 80% of the total larvae in the parks with the dominant groups being; Gobiidae, Blenniidae, Engraulidae and Apogonidae. The availability of a wide size range of larvae for some groups like Blenniidae, Gobiidae, Engraulidae, Labridae and Scaridae indicated the possibility of some larvae completing their pelagic phase within the parks. Larval densities differed significantly between seasons and stations in both parks, with higher densities ranging between 95–311 larvae/100 m³ during the calm northeast monsoon (NEM) season, compared to 5–58 larvae/100 m³ during the stormy southeast monsoon (SEM) season. Stepwise multiple regression analysis indicated a significant positive influence of zooplankton density on fish larval abundance in Malindi Park, while a positive and significant influence of temperature was found in Watamu Park. Distinct assemblages of larvae occurred within the parks indicating within park patchiness in larval distribution and, perhaps existence of retention features. These results provide useful baseline data on larval distribution in the WIO, and will be useful in understanding population dynamics of adult populations when complimented with additional oceanographic data.     

Preliminary evidence of delayed spawning and suppressed larval growth and condition of the major carps in the Yangtze River below the Three Gorges Dam

Commercial catches of the ‘major carps’ (grass carp Ctenopharyngodon idella, silver carp Hypophthalmichthys molitrix, black carp Mylopharyngodon piceus and bighead carp Aristichthys nobilis) in the middle and lower reaches of the Yangtze River have declined precipitously since construction of the Three Gorges Dam (TGD). To assess specific environmental effects, young-of-the-year major carps were collected below TGD in the Yangtze River at Jianli (larvae) and E’zhou (juveniles) and in the Dongting Lake (juveniles) during June–August 2007. Lapillar otoliths were analyzed to determine early growth. There were no discernible growth increments in otoliths for a majority of the larvae (38 of 63 grass carp and 40 of 47 silver carp), while increments were evident in otoliths formed in the larval stage for all the juveniles, indicating that larvae without discernible increments failed to recruit to the juvenile stage. Back-calculated hatch dates of major carps were between 31 May and 24 July. The initiation of the spawning season was delayed about 1 month compared to pre-TGD records. Larval growth rates, as reflected by otolith-increment width, were lower at Dongting Lake (closer to TGD) than at E’zhou (farther from TGD), indicating that TGD-moderated effects on early growth and development of carps are more apparent for fishes hatched nearer to the dam. The delay in spawning and decreased early growth may partly explain the recent decline of the major carp resources in the middle and lower Yangtze River.     

Gender Specific Brood Cells in the Solitary Bee Colletes halophilus (Hymenoptera; Colletidae)

We studied the reproductive behaviour of the solitary bee Colletes halophilus based on the variation in cell size, larval food amount and larval sex in relation to the sexual size dimorphism in this bee. Brood cells with female larvae are larger and contain more larval food than cells with males. Occasionally males are reared in female-sized cells. We conclude that a female C. halophilus in principal anticipates the sex of her offspring at the moment brood cell construction is started. Additionally a female is able to ‘change her mind’ about the sex of her offspring during a single brood cell cycle. We present a model that can predict the sex of the larvae in an early stage of development.     

Laboratory spawning cues in Menidia beryllina,and M. peninsulae (Pisces,Atherinidae) with notes on survival and growth of larvae at different salinities

Synopsis Spawning patterns of inland silversides, Menidia beryllina, and tidewater silversides, Menidia peninsulae, were examined in the laboratory under several combinations of ‘tidal’ and diel light cycle cues. M. beryllina showed a high frequency of spawning throughout the day when held under constant conditions (24L: OD, current velocity 8 cm sec⁻¹) and when ‘tidal’ and diel light cycles were presented singly or in combination. In contrast, M. peninsulae demonstrated a high frequency of spawning only when presented a combination of ‘tidal’ and diel light cycle cues and spawned predominantly at night. Menidia beryllina embryos were euryhaline. Hatching ranged from 73 to 78% at salinities of 5,15 and 30‰ M. peninsulae embryos showed an inverse relationship between the percentage hatch and the incubation salinity, 90% at 5‰ and only 65% at 30‰ Survival and growth of larval M. beryllina from the day of hatching through 16 days old was optimal at 15‰ Although survival of M. peninsulae larvae was optimal at 30%, no trend was apparent in growth of larvae held for 16 days at 5, 15 or 30‰ salinity. Contribution No. 508 from the Gulf Breeze Environmental Research Laboratory     

Ant benefits in a seed dispersal mutualism

Myrmecochorous plant seeds have nutrient rich appendages, elaiosomes, which induce some ant species to carry the seeds back to their nest where the elaiosome is consumed and the seed is discarded unharmed. The benefits to plants of dispersal of their seeds in this way have been well documented, but the benefits to the ants from consuming the elaiosomes have rarely been measured and are less clear. Ant benefits from myrmecochory were investigated in a laboratory experiment using the ant Myrmica ruginodis and seeds of Ulex species. To separate the effects of elaiosome consumption on the development of newly produced larvae versus existing larvae, ten ‘Queenright’ colonies containing a queen were compared to ten ‘Queenless’ colonies. Six measures of colony fitness over a complete annual cycle were taken: sexual production, larval weight and number, pupal weight and number, and worker survival. Queenless colonies fed with elaiosomes produced 100.0±29.3 (mean ± SE) of larvae compared to non-elaiosome fed colonies which produced 49.6±19.0; an increase of 102%. Larval weight increased in both Queenright and Queenless colonies. In colonies fed with elaiosomes, larvae weighed 1.02±0.1 mg, but in non-elaiosome fed colonies larvae weighed 0.69±0.1 mg; an increase of 48%. The food supplement provided by Ulex elaiosomes was trivial in energetic terms, under the conditions of an ample diet, suggesting that these effects might be due to the presence of essential nutrients. Chemical analysis of Ulex elaiosomes showed the presence of four essential fatty acids and four essential sterols for ants.     

Relations of homology between higher cognitive emotions and basic emotions

In the last 10 years, several authors including Griffiths and Matthen have employed classificatory principles from biology to argue for a radical revision in the way that we individuate psychological traits. Arguing that the fundamental basis for classification of traits in biology is that of ‘homology’ (similarity due to common descent) rather than ‘analogy’, or ‘shared function’, and that psychological traits are a special case of biological traits, they maintain that psychological categories should be individuated primarily by relations of homology rather than in terms of shared function. This poses a direct challenge to the dominant philosophical view of how to define psychological categories, viz., ‘functionalism’. Although the implications of this position extend to all psychological traits, the debate has centered around ‘emotion’ as an example of a psychological category ripe for reinterpretation within this new framework of classification. I address arguments by Griffiths that emotions should be divided into at least two distinct classes, basic emotions and higher cognitive emotions, and that these two classes require radically different theories to explain them. Griffiths argues that while basic emotions in humans are homologous to the corresponding states in other animals, higher cognitive emotions are dependent on mental capacities unique to humans, and are therefore not homologous to basic emotions. Using the example of shame, I argue that (a) many emotions that are commonly classified as being higher cognitive emotions actually correspond to certain basic emotions, and that (b) the “higher cognitive forms” of these emotions are best seen as being homologous to their basic forms.     

Is metamorphosis a critical interval in the early life of marine fishes?

Synopsis Small volumes of food in ‘stomach’, large proportions of empty ‘stomachs’ and small maximum sizes of prey during metamorphosis of cod,Gadus morhua, all point to an energy crisis during this developmental interval. The small size of the developing stomach of the 10–14 mm long cod may be partly responsible for the above mentioned difficulties during metamorphosis. A hypothesis based on Hjort's ‘critical stage theory’ is proposed, adding another critical interval at metamorphosis. Based on the presented data on cod and reports on other species in the literature, the new critical interval is discussed for larvae of marine fish species in general.     

‘Placebos’ and the logic of placebo comparison

Robin Nunn has argued that we should stop using the terms ‘placebo’ and ‘placebo effect’. I argue in support of Nunn’s position by considering the logic of why we perform placebo comparisons. Like all comparisons, placebo comparison is just a case of comparing one thing with another, but it is a mistake, I argue, to think of placebo comparison as a case where something is compared to ‘a placebo’. Rather, placebo comparison should be understood as a situation which sets-up the treatment and control groups in a particular way; not as a case involving objects or procedures called ‘placebos’ employed in order to control for ‘placebo effects’.     

Taxonomy and host-specificity of Gyrodactylus aideni n. sp. and G. pleuronecti (Monogenea: Gyrodactylidae) from Pseudopleuronectes americanus (Walbaum) in Passamaquoddy Bay, New Brunswick, Canada

Wild and cultured winter flounder Pseudopleuronectes americanus (Walbaum) from Passamaquoddy Bay were surveyed for species of Gyrodactylus Nordmann, 1832. Two species were found: G. pleuronecti Cone, 1981 and G. aideni n. sp, both members of Malmberg’s ‘groenlandicus group’. Although the hard parts in the haptor are very similar in the two species, hamuli of G. aideni are consistently shorter than those of G. pleuronecti. The two species differed by 35 base pairs in the ITS 1, 5.8 and ITS 2 region. A BLAST search identified a variety of species of Gyrodactylus from marine fishes in the Atlantic Ocean as closest matches, indicating the ‘groenlandicus group’ is part of a major marine lineage within Gyrodactylus (sensu lato) that has successfully radiated among coastal percid, pleuronectid, cottid and anarhichadid fishes. Exposure experiments suggested that winter flounder is the primary host of both species of parasites and that three other pleuronectid species in the bay may potentially serve only as occasional transport hosts.     

Hatching rhythms and dispersion of decapod crustacean larvae in a brackish coastal lagoon in Argentina

Mar Chiquita, a brackish coastal lagoon in central Argentina, is inhabited by dense populations of two intertidal grapsid crab species,Cyrtograpsus angulatus andChasmagnathus granulata. During a preliminary one-year study and a subsequent intensive sampling programme (November–December 1992), the physical properties and the occurrence of decapod crustacean larvae in the surface water of the lagoon were investigated. The lagoon is characterized by highly variable physical conditions, with oligohaline waters frequently predominating over extended periods. The adjacent coastal waters show a complex pattern of semidiurnal tides that often do not influence the lagoon, due to the existence of a sandbar across its entrance. Besides frequently occurring larvae (exclusively freshly hatched zoeae and a few megalopae) of the two dominating crab species, those of three other brachyurans (Plathyxanthus crenulatus, Uca uruguayensis, Pinnixa patagonica) and of one anomuran (the porcellanidPachycheles haigae) were also found occasionally. Caridean shrimp (Palaemonetes argentinus) larvae occurred in a moderate number of samples, with a maximum density of 800·m⁻³. The highest larval abundance was recorded inC. angulatus, with almost 8000°m⁻³. Significantly moreC. angulatus andC. granulata zoeae occurred at night than during daylight conditions, and more larvae (statistically significant only in the former species) during ebb (outflowing) than during flood (inflowing) tides. In consequence, most crab zoeae were observed during nocturnal ebb, the least with diurnal flood tides. Our data suggest that crab larvae do not develop in the lagoon, where the adult populations live, but exhibit an export strategy, probably based upon exogenously coordinated egg hatching rhythms. Zoeal development must take place in coastal marine waters, from where the megalopa eventually returns for settlement and metamorphosis in the lagoon. Significantly higher larval frequency ofC. granulata in low salinities (≤12‰) and at a particular sampling site may be related to local distribution patterns of the reproducing adult population. Unlike crab larvae, those of shrimp (P. argentinus) are retained inside the lagoon, where they develop from hatching through metamorphosis. They significantly prefer low salinity and occur at the lagoon surface more often at night. These patterns cannot be explained by larval release rhythms like those in brachyuran crabs, but may reflect diel vertical migrations to the bottom. It is concluded that osmotic stress as well as predation pressure exerted by visually directed predators (small species or life-cycle stages of estuarine fishes) may be the principal selection factors for the evolution of hatching and migration rhythms in decapod larvae, and that these are characteristics of export or retention mechanisms, respectively.     

Pacific Coral-reef Fishes: The Implications of Behaviour and Ecology of Larvae for Biodiversity and Conservation, and a Reassessment of the Open Population Paradigm

The two-phase life history of most marine fishes and invertebrates has enormous implications for dispersal, population connectivity, and resource management. Pelagic dispersal larvae of marine animals traditionally thought to ensure that populations are widespread, that chances of local extinction are low, and that marine protected areas (MPA) can easily function to replenish both their own populations and those of unprotected areas. Traditionally, dispersal is considered to depend primarily on two variables: pelagic larva duration and far-field currents. These conclusions arise from the open population paradigm and are usually accompanied by a simplifying assumption: larvae are distributed passively by far-field currents. Unfortunately, they ignore the complex reality of circulation and hydrological connectivity of reefs, and do not consider newly-demonstrated behavioural capabilities of coral-reef fish larvae. Far-field circulation varies with depth and often excludes water bodies where propagules are released, and this has important implications for predicting trajectories of even passive larvae. However, larvae are not passive: late-stage larvae of coral-reef fishes can swim faster than currents for long periods, can probably detect reefs at some distance, and can actively find them. This behaviour is flexible, which greatly complicates modelling of larval fish trajectories. Populations at ecological (as opposed to evolutionary) scales are probably less open and more subdivided than previously assumed. All this means that dispersal predictions based solely on far-field water circulation are probably wrong. An emerging view of larval-fish dispersal is articulated that takes these new data and perspectives into account. This emerging view shows that re-evaluation of traditional views in several areas is required, including the contribution of larval-fish biology and dispersal to biodiversity patterns, the way reef fishes are managed, and the way in which MPA are thought to operate. At evolutionary and zoogeographic scales, reef-fish populations are best considered to be open.     

Development of the Acoustically Evoked Behavioral Response in Larval Plainfin Midshipman Fish,Porichthys notatus

The ontogeny of hearing in fishes has become a major interest among bioacoustics researchers studying fish behavior and sensory ecology. Most fish begin to detect acoustic stimuli during the larval stage which can be important for navigation, predator avoidance and settlement, however relatively little is known about the hearing capabilities of larval fishes. We characterized the acoustically evoked behavioral response (AEBR) in the plainfin midshipman fish, Porichthys notatus, and used this innate startle-like response to characterize this species'' auditory capability during larval development. Age and size of larval midshipman were highly correlated (r² = 0.92). The AEBR was first observed in larvae at 1.4 cm TL. At a size ≥1.8 cm TL, all larvae responded to a broadband stimulus of 154 dB re1 µPa or −15.2 dB re 1 g (z-axis). Lowest AEBR thresholds were 140–150 dB re 1 µPa or −33 to −23 dB re 1 g for frequencies below 225 Hz. Larval fish with size ranges of 1.9–2.4 cm TL had significantly lower best evoked frequencies than the other tested size groups. We also investigated the development of the lateral line organ and its function in mediating the AEBR. The lateral line organ is likely involved in mediating the AEBR but not necessary to evoke the startle-like response. The midshipman auditory and lateral line systems are functional during early development when the larvae are in the nest and the auditory system appears to have similar tuning characteristics throughout all life history stages.