Evidence for sexual dichromatisms in spawning aggregations of yellowfin grouper Mycteroperca venenosa and tiger grouper Mycteroperca tigris from the southern Gulf of Mexico

Colour pattern characteristics and gonad histology were used to detect sexual dichromatisms in yellowfin grouper Mycteroperca venenosa and tiger grouper Mycteroperca tigris from the Campeche Bank, Mexico. Specimens were obtained from commercial catches between March and May during 2002 and 2004. All specimens were examined dead. Ninety-seven per cent of males had different sex-associated colour patterns. Male yellowfin grouper displayed a bright yellow blotch on both sides of the lower jaw while females retained a reddish lower jaw. Male tiger grouper had uniform dark pectoral fins while females had bright orange pectoral fins. In situ observations of live fishes at fishing sites showed the lower jaw and pectoral fin colourations to be clearly visible underwater at a depth of 35 m. All males of both species and most females (80% yellowfin grouper and 98% tiger grouper) were sexually active and probably caught during their spawning season. This suggests that distinct colourations observed for male M. venenosa and M. tigris may be seasonal displays associated with spawning. Both the lower jaw and pectoral fin colourations were still visible in dead fishes after several days on ice. Differences observed for ray length of exserted vertical fins in tiger grouper specimens were probably not a sex-associated characteristic.     

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.     

Prediction and verification of possible reef‐fish spawning aggregation sites in Los Roques Archipelago National Park,Venezuela

This study attempts to predict and verify possible spawning aggregation sites and times in the Los Roques Archipelago National Park, Venezuela, based on physical reef characteristics and the knowledge of experienced local fishermen. Three possible aggregation sites were selected for monitoring based on satellite images, low‐cost bathymetric mapping and interviews with experienced local fishermen. Abundances and sizes of 18 species that are known to form reproductive aggregations were monitored at these sites using underwater visual census for 7 days after each full moon from February to August, 2007. While spawning events were not observed, possible indirect evidence of spawning aggregations was found for Lutjanus analis at Cayo Sal and Boca de Sebastopol, Lutjanus apodus at Cayo Sal, Lutjanus cyanopterus at Cayo Sal and Piedra La Guasa and Epinephelus guttatus at Bajo California and Cayo de Agua. Additionally, indirect evidence was identified for the past existence of a spawning aggregation of Epinephelus striatus in the northern part of the archipelago, which may have been eliminated by overfishing c.15 years ago. Bathymetric mapping showed that the shelf edge at sites monitored in this study was shallower than at spawning aggregation sites in other parts of the Caribbean, and that sites were not proximal to deep water. While this study does not prove the existence or locations of spawning aggregations of reef fishes in the archipelago, it does add insight to a growing understanding of generalities in the relationship between seafloor characteristics and the locations of transient reef‐fish spawning aggregations in the Caribbean.     

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.     

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.     

Spawning behavior of the tiger grouper (<Emphasis Type="Italic">Mycteroperca tigris</Emphasis>) in a Caribbean atoll

Many grouper (Epinephelidae) species aggregate to spawn at specific times of year, at consistent locations, and at specific times of day and phases of the moon. The tiger grouper (Mycteroperca tigris) is widely distributed in the Caribbean Sea and aggregates to spawn just after the full moon. In January and February 2003–2005, we conducted visual surveys of a tiger grouper spawning aggregation at Glover’s Reef, Belize. The primary objectives of this study were to describe grouper spawning activities with respect to the timing of the lunar cycle, identify and describe male and female spawning behaviors, and estimate the number of grouper spawning events each day. The peak number of both male and female tiger groupers counted each year occurred from 6 to 10 d after the full moon. The peak count of males was 1.6–2.2 times greater than the peak count of females. Six categories of male–male and six categories of male–female interactions were identified. All male tiger groupers displayed an ability to rapidly change color; this was often triggered by a close encounter with other male or female tiger groupers. The daily spawning activity was remarkably predictable. The first spawning occurred on average 8.1 min before sunset, and the last occurred 5.1 min after sunset. On average, less than half of the females at the spawning site spawned each day, but some females spawned more than once in a given day. Females appeared to choose their mates.     

Spatial and temporal patterns of movement and migration at spawning aggregations of red hind, <Emphasis Type="Italic">Epinephelus guttatus</Emphasis>, in the U.S. Virgin Islands

This study compared the spatial and temporal patterns of red hind, Epinephelus guttatus, movement and migration from annual spawning aggregations on St Thomas (STT) and St Croix (STX), United States Virgin Islands. Around STT E. guttatus migrated 6–33 km from a functional spawning migration area of 500 km² and around STX E. guttatus migrated 5–18 km from an area of 90 km². Similarities between sites were found in regards to timing of movement, temporal and spatial changes in sex ratios, annual and lunar predictability and were synchronized with environmental cues. E. guttatus spawning aggregations in the Virgin Islands occur between the winter solstice (i.e., after December 20) and about February 20 of any year and show a distinctive peak 20–40 days after the winter solstice. Spawning typically occurred during periods of declining seawater temperature and slacking currents within a temperature range of 26–27.5°C and current speed of 2.5–3.5 cm s⁻¹. Males arrived early to spawning sites and stayed longer than females. These gender-based behavioral patterns are important to E. guttatus reproductive dynamics and must be factored into future studies and the design of fisheries regulations to ensure sustainability of spawning aggregation sites. The predictability of E. guttatus spawning aggregations relative to the winter solstice will be extremely beneficial for defining the temporal and spatial aspects of area closures. The consistency and synchrony of movement and migration will improve both the efficiency of planning research and monitoring programs and directing enforcement activities during critical time periods. Applying this knowledge strategically will maximize the limited resources available for research and enforcement and lead to greater protection of spawning aggregations.     

Spatial distribution of red grouper Epinephelus morio (Serranidae) catchability on the Campeche Bank of Mexico

Distribution patterns of catchability were evaluated for the red grouper Epinephelus morio (Serranidae) on the Campeche Bank off the northern Yucatan coast of Mexico. Catchability‐at‐length values were estimated with a method based on a transition matrix. Five years of catch and effort data for the Mexican mid‐fleet were used for the estimation procedure. Two main patterns were found within the bank. The first occurred in the deepest areas and in the eastern part of the bank, where catchability increased with size. The second was close to the coasts, where lower values of catchability were observed, even in large sizes. These results suggest that the main area of aggregation for reproduction with higher values of catchability is in the eastern part of the bank (22–23°N, 87–88°W). Catchability differences suggest that spatial management policies must be implemented to allow depleted stock to recover.     

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.     

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.     

Reproductive styles of shallow-water groupers (Pisces: Serranidae) in the eastern Gulf of Mexico and the consequences of fishing spawning aggregations

Synopsis Seasonal and spatial aspects of spawning for three commercially important grouper species in the northeastern Gulf of Mexico are detailed. These species — all of which are protogynous hermaphrodites - spawn in deep water (> 25 m for red grouper,> 40 m for gag and scamp), making it difficult to observe spawning behaviors without ROV or submersible support. They respond to intense fishing pressure in ways that are directly related to their respective reproductive styles. Species that aggregate appear to be more susceptible to such pressures than those that do not, as evidenced by marked skewing of sex ratios in favor of females. Gag, Mycteroperca microlepis, have suffered a drop in the proportion of males from 17% to 1% in the last 20 years; scamp, Mycteroperca phenax, have dropped from 36% to 18%; and red grouper, Epinephelus morio, which do not aggregate, have shown little change in the sex ratio over the past 25–30 years.     

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.     

Evaluation of a conservation strategy: a spawning aggregation closure for red hind, Epinephelus guttatus, in the U.S. Virgin Islands

Many tropical reef fishes spawn in large aggregations, which are readily targeted by fishers. By the 1980s, at least two grouper spawning aggregations were eliminated by intensive fishing off the island of St. Thomas, U.S. Virgin Islands, and another aggregating species, red hind, was intensively targeted. By 1988, the average length of red hind had greatly decreased to 295 mm, and the sex ratio was extremely skewed to 15 females per male, suggesting a heavily fished stock. Since this species is a protogynous hermaphrodite, the loss of large individuals (primarily males) could potentially result in sperm limitation in spawning aggregations. In 1990, a spawning aggregation closure was implemented. By 1997, average size of red hind had increased to 395 mm and sex ratio had shifted to 4 females per male. Fish were observed aggregating only in structurally complex habitat along the insular shelf edge. This habitat type is apparently not common along most of the shelf edge off St. Thomas and may provide shelter while reducing risk of predation during aggregation periods. These data suggest that protection of spawning aggregations is a sound management strategy with considerable potential for aiding the sustainable use of reef fish resources.     

The ups and downs of coral reef fishes: the genetic characteristics of a formerly severely overfished but currently recovering Nassau grouper fish spawning aggregation

The Nassau grouper (Epinephelus striatus) has sustained large declines across its distribution, including extirpation of many of its fish spawning aggregations (FSAs). Within US Virgin Islands (USVI) waters, Nassau grouper FSAs were overfished until their disappearance in the 1970s and 1980s. In the early 2000s, however, Nassau grouper were found gathering at Grammanik Bank, USVI, a mesophotic coral reef adjacent to one of the extinct aggregation sites, and regulatory protective measures were implemented to protect this fledgling FSA. The population genetic dynamics of this rapid FSA deterioration followed by protection-facilitated, incipient recovery are unknown. We addressed two objectives: (1) we explored which factors (i.e., local vs. external recruitment) might be key in shaping the USVI FSA recovery; and (2) we examined the consequences of severe past overfishing on this FSA’s current genetic status. We genotyped individuals (15 microsatellites) from the USVI FSA comprising three successive spawning years (2008–2010), as well as individuals from a much larger, presumably less impacted, Nassau grouper FSA in the Cayman Islands, to assess their comparative population dynamics. No population structure was detected between the USVI and Cayman FSAs (F _ST = −0.0004); however, a temporally waning, genetic bottleneck signal was detected in the USVI FSA. Parentage analysis failed to identify any parent–offspring matches between USVI FSA adults and nearby juveniles, and relatedness analysis showed low levels of genetic relatedness among USVI FSA individuals. Genetic diversity across USVI FSA temporal collections was relatively high, and no marked differences were found between the USVI and Cayman FSAs. These collective results suggest that external recruitment is an important driver of the USVI FSA recovery. Furthermore, despite an apparent genetic bottleneck, the genetic diversity of USVI Nassau grouper has not been severely compromised. Our findings also provide a baseline for future genetic monitoring of the nascent USVI aggregation.

     

Coexistence of congeneric spiny lobsters on coral reefs: differences in conspecific aggregation patterns and their potential antipredator benefits

Den sharing by conspecific spiny lobsters (aggregation) is modulated by chemical attraction but may confer several, not necessarily mutually exclusive, antipredator byproduct benefits: a “guide effect”, which only benefits the individual attracted to a sheltered conspecific; a “dilution effect”, which reduces per-capita risk of predation simply through aggregation; or active “group defense”. Each potential benefit has a different set of predictors (relationships between aggregation and conspecific or predator densities), but conflicting results could suggest the simultaneous operation of more than one benefit. These predictions were tested for coexisting Panulirus guttatus (a reef-obligate) and Panulirus argus (a temporary reef-dweller) using data collected during 11 surveys on fixed sites over a coral reef in Mexico. P. guttatus greatly outnumbered P. argus, but P. argus showed a greater tendency to aggregate. All three benefits of den sharing operated for the more social P. argus, with “group defense” being of the most benefit for larger individuals, and the “guide” and “dilution” effects for smaller individuals recently immigrating into the reef habitat and sharing dens with larger conspecifics. P. guttatus did not display “group defense” and its aggregations appeared to be modulated by the interplay between attraction and aggressive behaviors. This species relied more on solitary crypticity, especially at larger sizes, but appeared to benefit from a “guide effect” at high conspecific densities. In experimental tanks, each species tended to aggregate when tested separately, but when tested jointly, aggregation among P. guttatus was significantly reduced. The experimental results reflect the differential patterns of aggregation between the fore-reef, where P. guttatus dominated, and the back-reef, where coexistence of both species was greater.     

Reproductive biology and management of two commercially important groupers in the SW Atlantic

The reproductive biology of Epinephelus morio (red grouper) and Mycteroperca bonaci (black grouper) were evaluated based on 533 specimens collected from artisanal fisheries landings in the Abrolhos Bank, Brazil, between May 2005 and September 2012. Sex ratio for the black grouper was 1:14 (n = 155 females and 11 males; 26.1–147 cm TL) and 1:10 for the red grouper (n = 334 females and 33 males; 15.0–96.0 cm TL). For both species, highest values of the gonadosomatic index (GSI) for females were recorded between July and October, indicating spawning during the austral winter. The length at first maturity (L₅₀) for females was estimated at 62.0 and 47.0 cm TL for the black and red grouper, respectively. Batch fecundity based on TL and TW ranged from 2 to 15.4^?10⁶ and 1.5 to 13.7^?10⁶ for the black and red grouper, respectively. Interviews with experienced fishers revealed that spawning seasons of both groupers are largely unrecognized. Results demonstrate a positive relationship between GSI peaks, lower temperatures and stronger winds. The information provided herein may help decision-making regarding fisheries management and conservation for E. morio and M. bonaci at various levels of governance in the Abrolhos Bank, the region with the largest and richest coralline reefs in the South Atlantic.     

Mass spawning of Caesio teres (Pisces: Caesionidae) at Enewetak Atoll,Marshall Islands

Synopsis Mass spawning of Caesio teres occurred between March and August, 1983 on a reef emerging from deep water just inside the East Channel of Enewetak Atoll, Marshall Islands. Aggregations and spawning were observed from one day before until three days after the full moon. Spawning aggregations were not observed during the new or quarter moons. Spawning commenced after high tide, when the current began to flow out the pass from lagoon to ocean. The spawning aggregation of close to 1000 individuals migrated to the spawning site. Spawning occurred when the aggregate ascended to near the water's surface. Subgroups dashed horizontally within the aggregate, releasing a highly visible gamete cloud. Predation on spawning adults was not observed. Predation on spawned eggs was noted. The eggs of C. teres are described. Mid-Pacific Research Laboratory, Enewetak Atoll, Republic of the Marshall Islands Present address: Motupore Island Research Station, P.O. Box 320, University of Papua New Guinea, Papua New Guinea     

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.