First record of a spawning aggregation for the tropical eastern Pacific endemic grouper Mycteroperca olfax in the Galapagos Marine Reserve

This study provides direct and indirect evidence of temporally and spatially consistent spawning aggregations for the grouper Mycteroperca olfax. Recently reported declines in population numbers, probably related to the direct targeting of aggregations by artisanal fishermen, highlight the urgent need for species‐specific management actions in the Galapagos Marine Reserve, such as minimum and maximum landing sizes, and the importance of protecting key aggregation sites with the declaration of no‐take areas and the establishment of total fishing bans during the reproductive season.     

Observations of a Nassau grouper, Epinephelus striatus, Spawning Aggregation Site in Little Cayman, Cayman Islands, Including Multi-Species Spawning Information

Mass spawning aggregations of Caribbean grouper species are a conservation priority because of declines due to over-fishing. Previous studies have documented five historical aggregation sites in the Cayman Islands. Today, three of these sites are inactive or commercially extinct. In January 2002, the Reef Environmental Education Foundation led an expedition to Little Cayman Island to document a recently re-discovered spawning aggregation of Nassau grouper, Epinephelus striatus. A team of divers estimated the abundance, color phase composition, and courtship and spawning behavior of the aggregating grouper. The color phase composition of the aggregation shifted both during the course of each evening and throughout the 10-day project. Divers documented atypical coloration and courtship behavior in 10 additional fish species, of which five were seen spawning. Artisanal fishing occurred daily on the aggregation. The Cayman Islands Department of the Environment collected landings data and sampled catches to obtain length and sex ratios. The Cayman fishing fleet, while small, had a significant impact on the aggregation with a harvest of almost 2000 Nassau grouper during the 10-day project. The study site supports the largest known Nassau grouper aggregation in the Cayman Islands. The relatively large size of fish and the high proportion of males indicate that this site supports a relatively healthy aggregation compared to other Nassau grouper aggregation sites throughout the Caribbean.     

Spawning aggregations of three protogynous groupers in the southern Gulf of Mexico

Spawning aggregations of red hind Epinephelus guttatus, tiger grouper Mycteroperca tigris and yellowfin grouper Mycteroperca venenosa were identified at two coral‐reef systems: Arrecife Alacranes (emergent bank reef) and Bajos del Norte (submerged bank reef) on the continental shelf of the Yucatan Peninsula (Campeche Bank), Mexico. At both reefs, E. guttatus forms large spawning aggregations between February and March. At Bajos del Norte, M. tigris reproduces in a small, low‐density aggregation in May, while M. venenosa aggregates at high densities for spawning between March and May. Multi‐species use of an aggregation site by E. guttatus and M. venenosa was observed at Bajos del Norte. The identified spawning aggregations are apparently stable in location over time, and all three species were commonly observed to spawn within 1 week following the full moon. Development and survival of the larvae spawned in these aggregations are probably aided by a seasonal (spring–summer) upwelling in the north‐east Campeche Bank. A permanent area closure at Bajos del Norte, currently outside any specific fisheries management area or regulations, would provide protection needed for the spawning aggregations of these three species.     

Aggregations of <Emphasis Type="Italic">Plectropomus areolatus</Emphasis> and <Emphasis Type="Italic">Epinephelus fuscoguttatus</Emphasis> (groupers,Serranidae) in the Komodo National Park,Indonesia: Monitoring and Implications for Management

Synopsis We identify fishery management implications from a long-term monitoring program focusing on spawning aggregations of high valued reef fish in Komodo National Park (KNP), Eastern Indonesia. Management objectives of KNP are not only to protect biodiversity, but also to conserve spawning stocks of high-valued commercial species for the replenishment of surrounding fishing grounds. We monitored two sites twice monthly over five years for two species of grouper, Epinephelus fuscoguttatus and Plectropomus areolatus. One site had an aggregation of both E. fuscoguttatus and P. areolatus, whereas the other site contained an aggregation of P. areolatus only. Over the five years monitoring period, aggregations typically formed during each full moon between September and February. Additionally, P. areolatus occasionally aggregated during new moons between April and July. We observed spawning only once, but because formation of aggregations were correlated to a higher incidence of behavior and signs indicative of reproduction and because most fish present were adults, it is likely that the formation of aggregations was associated with spawning. Over the five years monitoring period there was a reduction in mean fish size of up to 8 cm for P. areolatus, and a reduction in numbers of aggregating E. fuscoguttatus. Despite limited protection initiated in 2001, both sites are still heavily fished by local artisanal fishers. Because the observed reductions in size and in numbers could be caused by fishing pressure, managers should follow the precautionary principle by putting additional protective management in place. Since both species are relatively long-lived, at least five years of continued monitoring may be necessary to determine the outcome of management intervention. The variability in timing of aggregation in respect to season and moon phase in P. areolatus indicates that long-term monitoring must cover the entire year and both moon phases.     

The utility of a long‐term acoustic recording system for detecting white seabass Atractoscion nobilis spawning sounds

This study reports the use of a long‐term acoustic recording system (LARS) to remotely monitor white seabass Atractoscion nobilis spawning sounds at three sites along the southern California coastline, adjacent to Camp Pendleton. On the basis of previous studies of A. nobilis sound production relative to periods of known spawning activity, LARS were set to continuously record ambient sounds for a 2 h period around sunset from April to June 2009. Acoustic analyses identified A. nobilis courtship sounds on 89, 28 and 45% of the days at the three locations, respectively. From 474 h of acoustic data, spawning‐related sounds (chants) were detected on 19 occasions in 2009 with an additional 11 spawning chants recorded during a 2007 validation period. Most spawning chants occurred within 30 min of sunset during the months of May and June at a mean ±s.d . surface temperature of 18·2 ± 1·2° C. Consecutive daily spawning activity was not apparent at any sites in 2009. Atractoscion nobilis spawning chants were recorded at all three sites, suggesting that shallow rocky reefs which support kelp forests provide suitable A. nobilis spawning habitat. Results confirm the utility of passive acoustic recorders for identifying A. nobilis spawning periods and locations.     

Patagonian grouper Acanthistius patachonicus (Jenyns, 1840) forms massive spawning aggregations in the Southwest Atlantic

Here we describe massive spawning aggregations and seasonal changes in the large-scale distribution of the Patagonian grouper Acanthistius patachonicus in the Southwest Atlantic based on three sources of information: (a) data from bottom trawl research surveys covering the distributional range of the species within the Argentine continental shelf; (b) folk ecological knowledge gathered from experienced captains of the Argentine industrial trawl fisheries; and (c) sampling of an artisanal trap fishery targeting the Patagonian grouper in a specific location off the coast of Buenos Aires Province. The trawl surveys showed a general pattern of aggregation of Patagonian grouper towards the coast during the reproductive period (September–December). Captains described massive aggregations of the species at specific near-shore locations, where trawl catches of up to 15 t in a single haul were registered during the reproductive season. At a local scale, the artisanal trap fishery described operates exclusively during the reproductive period, targeting near-shore aggregations with a high proportion of individuals releasing gametes onboard. These three sources of information provide evidence of the existence of massive transient spawning aggregations of Patagonian grouper in the Argentine shelf. This is the first report of a reef fish spawning aggregation in the southern region of the Southwest Atlantic. Anecdotal information gathered in this study points to the depletion of many of the aggregations targeted during the 1980s and 1990s by the industrial fleet. At the same time, the spawning aggregation site off Buenos Aires Province has been productive for the last 34 years, being exploited exclusively by traps. The Patagonian grouper is classified as Data Deficient by the International Union for Conservation of Nature, which highlights the need for further research to determine its stock status. Mapping its current spawning aggregations should be a priority to inform the design of a targeted monitoring program and management plan for this species.     

Seasonal patterns and individual differences in the calls of male haddock Melanogrammus aeglefinus

Haddock Melanogrammus aeglefinus males observed in captivity proved to be vocally active throughout the whole year, although the highest levels of vocal activity were reached in the spawning season between February and April. During this season, the most common type of sound consisted of sequences of regularly repeated, almost identical, double‐pulsed knocks (KN). These sequences were mostly associated with a particular behaviour pattern, the patrolling display, which has previously been shown to play a key role in territoriality and spawning. Differences in the double‐pulsed KNs produced by three individual males were demonstrated by two different statistical methods and it is suggested that these differences may allow individual males to be discriminated by other fish, perhaps conveying fitness‐related information on the sender. Some aspects of the calls changed seasonally, perhaps reflecting observed variations in the mass of the drumming muscles. When using passive listening to locate M. aeglefinus spawning grounds, the physical characteristics of the sounds may indicate how close the individual males are to spawning.     

Yellow-eyed piranhas produce louder sounds than red-eyed piranhas in an invasive population of Serrasalmus marginatus

Serrasalmus marginatus is a piranha species native from the lower Paraná River basin and has been invasive in the upper Paraná River basin since the 1980s. In piranhas, sounds of different species have different features. The aim of this study was to investigate if the sounds produced by this species could be used to distinguish two morphotypes: red- and yellow-eyed S. marginatus from the Araguari River (upper Paraná River basin). All the temporal and frequency features of the sounds were equivalent in both groups of eye colour; it corresponds to the species-specific signature described for S. marginatus. Nonetheless, the amplitude features were all statistically different between red- and yellow-eyed piranhas. Yellow-eyed specimens produced louder sounds. In different fish species, colour change in eyes can be due to the absence or the presence of a dominant allele. It can also be involved in social rank or during reproduction. Different hormones and neuropeptides can modulate vocal features. It is hypothesized that a mutation or different hormonal concentrations could explain both sound amplitude and eye colour playing a role in animal communication in S. marginatus.     

Reproduction in an aggregating grouper,the red hind,epinephelus guttatus

Synopsis We examined the reproductive pattern of an aggregating grouper, the red hind,Epinephelus guttatus, in Puerto Rico. Macroscopic and histological examination of gonads confirmed that, although mature, ripe ovaries are found over a three-month period, spawning activity is limited to about 2 weeks each year. Females are determinate spawners and individuals are able to spawn more than once during the course of the annual spawning season. The red hind is protogynous and 50% maturity is attained at 215 mm fork length. In western Puerto Rico, spawning occurs in aggregations at several sites within loosely defined areas located towards the edge of the insular platform. Sex ratios of individuals taken by hook and line at one of the aggregation areas over a consecutive six-year period, suggest considerable intra- and inter-annual variation most likely attributable to a combination of differential ingress and egress by males and females during the course of an aggregation and to fluctuations in recruitment, differential mortality by sex and sex change among years. A comparative assessment of the reproductive patterns of seven western AtlanticEpinephelus spp. suggests that aggregation-spawning is associated with medium- to large-sized groupers, while smaller groupers do not aggregate. Mating systems vary among congeners in association with trends in male-female size dimorphism, sexual pattern and sperm competition. The short-term, localized, nature of spawning aggregations renders species with this reproductive mode particularly vulnerable to fishing pressure at spawning sites. Aggregating species, therefore, may require special management consideration.     

Temporal and spatial mapping of red grouper Epinephelus morio sound production

The goals of this project were to determine the daily, seasonal and spatial patterns of red grouper Epinephelus morio sound production on the West Florida Shelf (WFS) using passive acoustics. An 11 month time series of acoustic data from fixed recorders deployed at a known E. morio aggregation site showed that E. morio produce sounds throughout the day and during all months of the year. Increased calling (number of files containing E. morio sound) was correlated to sunrise and sunset, and peaked in late summer (July and August) and early winter (November and December). Due to the ubiquitous production of sound, large‐scale spatial mapping across the WFS of E. morio sound production was feasible using recordings from shorter duration‐fixed location recorders and autonomous underwater vehicles (AUVs). Epinephelus morio were primarily recorded in waters 15–93 m deep, with increased sound production detected in hard bottom areas and within the Steamboat Lumps Marine Protected Area (Steamboat Lumps). AUV tracks through Steamboat Lumps, an offshore marine reserve where E. morio hole excavations have been previously mapped, showed that hydrophone‐integrated AUVs could accurately map the location of soniferous fish over spatial scales of <1 km. The results show that passive acoustics is an effective, non‐invasive tool to map the distribution of this species over large spatial scales.     

Sound Production in Codoma ornata (Girard) (Cyprinidae)

Male Codoma ornata produce sounds associated with aggression and spawning activities during the breeding season. Females do not produce sounds. Males most often produced sounds associated with escalated displays of aggression, courtship and the spawning act. C. ornata spawn in crevices, but previously were reported to spawn as egg-clusterers in cavities. Structurally, sounds are low frequency, vary in duration according to context and are not harmonic. The mechanism of sound production is unknown.     

Reproductive movement,residency and fisheries vulnerability of brown-marbled grouper, <Emphasis Type="Italic">Epinephelus fuscoguttatus</Emphasis> (Forsskål, 1775)

The brown-marbled grouper, Epinephelus fuscoguttatus, is a long-lived, late-maturing protogynous species listed as Near Threatened by the IUCN. In Pohnpei, Micronesia, reproductively active brown-marbled grouper were tagged with acoustic and spaghetti-type tags at a multi-species fish spawning aggregation (FSA) site to establish patterns of movement, residency and seasonality. Telemetry confirmed the use of common reproductive migratory corridors and significant sex-specific variations in residency at the FSA. Combined underwater visual census and telemetry data verified a 3-month peak aggregation period, with aggregations forming and persisting over ca. 12 days prior to full moon between January and May. FSA formation coincided with seasonally low and relatively stable seawater temperatures. Some males frequented the FSA site during each aggregation month over two consecutive years. Conversely, most females were present at the FSA during only a single aggregation period, with the month of visitation consistent among years. Nearly two-thirds of tagged fish were relocated or recaptured within 11 km of the aggregation site, with a maximum detected distance of 26 km and a minimum estimated catchment area of 100–175 km². Findings highlight the need for a combined approach to management that prohibits the capture and sale of reproductive adults and protects both spawning sites and common reproductive migratory corridors during aggregation periods.     

A spawning aggregation of Nassau grouperEpinephelus striatus (Pisces: Serranidae) in the Mexican Caribbean

Synopsis An annual spawning aggregation of Nassau grouper,Epinephelus striatus, observed off the southern coast of Quintana Roo, Mexico, on full moon days in December and January, was surveyed during the winters from 1991 to 1993. A fish aggregation had been appearing at one traditional site off Mahahual for more than 80 years, but currently it forms at undetermined sites far from the original site. Fish migration was northwards in the fore reef border along the 14 m isobath and isolated groups were observed. Through visual assessment fish aggregations were found to include 200 to 500 individuals. The fishing on Nassau grouper aggregations has been active since 1910 by using hook and line; however, recently fishermen used gill nets. It is necessary to continue the research on this aggregation and establish management guidelines to protect the species.     

Reproductive biology of squaretail coralgrouper Plectropomus areolatus using age‐based techniques

The squaretail coralgrouper Plectropomus areolatus was identified as a fast‐growing, early maturing and relatively short‐lived aggregation‐spawning epinephelid. Examinations of sectioned otoliths found females and males first maturing at 2 and 3 years, respectively, suggesting protogynous hermaphroditism; however, no transitionals were observed in samples. Age distribution for the two sexes was similar and both were represented in the oldest age class; however, significant sex‐specific differences in size‐at‐age were identified. Both sexes fully recruit into the fishery at age 4 years and reach 90% of asymptotic length by age 3 years. Underwater visual assessments, combined with the gonado‐somatic indices, revealed a 5 month reproductive season, with interannual variability observed in the month of highest density within the spawning aggregation. Catch restrictions on adults during spawning times and at reproductive sites, combined with gear‐based management and enhanced enforcement, are recommended to maintain spawning stocks. Based on the available evidence, the sexual pattern for this species is unresolved.     

Spatial,temporal, and environmental dynamics of a multi-species epinephelid spawning aggregation in Pohnpei,Micronesia

Long-term and short-term underwater visual censuses using SCUBA, technical Nitrox, and closed circuit rebreathers (CCR) were carried out in Pohnpei, Micronesia, to define spatial and temporal dynamics within a semi-protected multi-species epinephelid (fish) spawning aggregation (FSA) of brown-marbled grouper, Epinephelus fuscoguttatus, camouflage grouper, Epinephelus polyphekadion, and squaretail coralgrouper, Plectropomus areolatus. Results identified species-specific patterns of habitat use, abundance, residency, and dispersal of FSAs. Fish spawning aggregations formed and dispersed monthly within a 21–160-d period after winter solstice within adjacent yet distinct outer reef habitats. The reproductive season coincided with periods of seasonally low sub-surface seawater temperature. Peaks in density varied among species both within the calendar year and relative to the winter solstice. Significant long-term declines in FSA density were observed for all three species, suggesting population-level fishery-induced impacts, similar to those previously reported for E. polyphekadion. Differences in density estimates were also observed between dive gear, with a threefold difference in densities measured by CCR for E. polyphekadion versus SCUBA that suggest a disturbance effect from exhaled SCUBA bubbles for this species. CCR also allowed surveys to be conducted over a larger area in a single dive, thereby improving the potential to gauge actual abundance and density within FSAs. Based on these findings, a combination of long-term and intensive short-term monitoring strategies is recommended to fully characterize trends in seasonal abundance and habitat use for aggregating species at single or multi-species FSA sites. Inherent variations in the timing and distribution of species within FSA make fine-scale temporal management protocols less effective than blanket protective coverage of these species at (e.g., marine protected areas covering FSAs and adjacent migratory corridors) and away from (i.e., temporal sales and catch restrictions) FSA sites.     

Aspects of the spawning of western Atlantic butterflyfishes (Pisces: Chaetodontidae)

Synopsis The status of knowledge of spawning among the five shallow waterChaetodon species in the western Atlantic is reviewed. Spawning has been observed for three species in Puerto Rico, St. Croix and the Bahamas, with possible courtship in a fourth.Chaetodon aculeatus spawned near the time of sunset over objects on the reef as single female/male pairs or as two females and one male, with pair spawning in rapid succession. Spawning occurred during much of the lunar month from February to April and it is uncertain whether any lunar periodicity to spawning exists. Male-male aggression was noted. Spawning sites (coral heads) were alternated daily and it is likely that females spawn only once every two days. A single female produced as many as 2090 eggs in a single spawning.Chaetodon capistratus spawned during much of the lunar month from February to April. It spawned about 5 min afterC. aculeatus, occasionally using the same sites, and alternated sites daily. A female produced as many as 3710 eggs in one spawning.Chaetodon striatus spawned from February to April but it is unknown if it has any lunar spawning cycle. No predation attempts by piscivores on spawning adults were seen. Predation byMelichthys niger on eggs ofC. striatus occurred. No egg predation was observed forC. aculeatus andC. capistratus. With an assumed four month reproductive season, alternate day spawning and observed egg production values,C. aculeatus andC. capistratus produce respectively about 100 000 and 200 000 eggs per large female per year. The reproductive strategy of smaller species may be to produce moderate numbers of eggs per day over a spawning season of at least a few months while larger species may produce more eggs per day for a shorter period.     

Comparative study of reproductive biology in single and multiple-spawner cyprinid fish. II. Sex steroid and plasma protein phosphorus concentrations

The dynamics and regulation of oogenesis in single and multiple-spawner cyprinid fish showing group-synchronous oocyte development, was investigated in three species from the River Meuse (Belgium): the roach Rutilus rutilus as a single spawner, and the bleak Alburnus alburnus and the white bream Blicca bjoerkna as multiple spawners. This paper compares the seasonal profiles of sex steroids (oestradiol-17β, testosterone and 17,20β-dihydroxy-4-pregnen-3-one) and plasma alkali-labile protein phosphorus. Different patterns of plasma oestradiol-17β (E2) and plasma protein phosphorus (PPP) have been observed not only between the single and the multiple spawner fish, but also among the two multiple spawners. In roach, two increases of E2 levels were observed. The first occurred in September after a short gonadal quiescent period, and coincided with the increase of the PPP at the onset of exogenous vitellogenesis. The second took place in spring before the spawning season. The low PPP recorded during that period probably reflected its rapid incorporation by the oocytes. In both multiple spawners, highest values of PPP were recorded just before the spawning season. In white bream, the PPP declined progressively once the differentiation of exogenous vitellogenic oocytes was completed before the onset of the spawning season. In bleak, PPP levels remained high throughout the spawning season and corresponded to a sustained oocyte recruitment during the whole of this period. Regardless of the pattern of oocyte growth recruitment, the E2 concentrations were high in both multiple spawner species during the breeding season. In the three species, testosterone levels remained low regardless of the maturation stage (ranging from 0.6 to 13.4ng ml^?1). Except for relatively high concentrations of 17,20βP in roach during final maturation and postspawning stages (20 and 28 ng ml^?1, respectively), low levels of this steroid were measured in these cyprinids, and especially in the multiple spawner fish. The role of this progestogen as the maturation inducing Steroid is discussed.     

The fitness of the Brazilian damsel <Emphasis Type="Italic">Stegastes fuscus</Emphasis> is increased by sharing the territory with the dusky grouper <Emphasis Type="Italic">Epinephelus marginatus</Emphasis>

Stegastes fuscus and Epinephelus marginatus are known for co-habiting shelters. The damselfish S. fuscus uses the territory for nesting and must protect its eggs from grazers; the grouper E. marginatus is an omnivorous sit-and-wait predator. This study aims to evaluate the effect of juvenile groupers on the reproductive success of the Brazilian damsel. Twenty-five hours of underwater observations were done in São Sebastião and Ilhabela, Northern shore of São Paulo, Brazil. Fitness increase was measured by the egg-clutch area and number of contributing females in 130 nests shared by groupers and another 130 where damselfishes stood alone. An egg predator crab was placed into the damselfish territory, and behavioural responses during 2 min were recorded for nests with or without E. marginatus, 80 replicates each. Nests shared by the dusky groupers had more eggs and received eggs from more females too. While fathers who were alone in the territory had to deal with the egg predator crab, in shared nests, the grouper would take care of the intruder, sometimes feeding on it. Therefore, the Brazilian damsel may benefit from the presence of the dusky grouper by increasing the fitness and diminishing the costs of parental care.