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.     

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.     

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.

     

Reproduction of the Nassau grouper,Epinephelus striatus (Pisces: Serranidae) and its relationship to environmental conditions

Synopsis Spawning aggregations of the Nassau grouper, Epinephelus striatus, occurring at the southern end of Long Island were investigated during the winters of 1987–1988 and 1988–1989. Fish aggregate at two specific sites around the time of the full moon in December and January, possibly not during November and almost certainly not during February. At one site the fish aggregate well inside the shelf edge while at the other they are on the dropoff into deep water. Fish migrate, probably as distinct groups of up to about 500 individuals, to the sites on or before the full moon and remain for several days. There was no significant difference in the size of males and females caught from the aggregations between years or months. Female/male sex ratio was between 5 : 1 and 3 : 1. Courtship occurred in late afternoon with spawning commencing shortly before sunset. Most spawning occurred within 10 min of sunset. Water temperatures were 25.0–25.5°C during a period of gradual decrease towards the annual minimum in February and March. Two color patterns were important in courtship and spawning. The bicolor pattern is a submissive coloration indicating a non-aggressive state acquired by both males and females near the time of spawning. The dark phase is acquired by females who are followed by numerous bicolor fish during courtship and they lead spawning events in this pattern. Spawning occurred among subgroups of the aggregation numbering 3–25 fish; gamete release was well above the bottom. Drogues deployed with the gametes either moved inshore or did not move far away from the shelf edge over the course of several days. Nassau groupers may not strictly be protogynous hermaphrodites and other groupers which form spawning aggregations may not be so either. Latitudinal shifts in spawning time may be related to water temperatures. Currents at aggregation sites do not appear to favor offshore transport of eggs. Questions of whether spawning aggregations should be protected need to be answered.     

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.     

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.     

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.     

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.     

Isopod infestation of postspawning Nassau grouper around Little Cayman Island

Attacks by the isopod Excorallana tricornis tricornis on Nassau grouper Epinephelus striatus caught in Antillean fish traps around Little Cayman Island, Cayman Islands, are recorded. Divers reported heavily parasitized fish during the study period, suggesting that trapping did not exclusively precipitate parasitism. Parasitic infestations were reported to be common in Nassau grouper immediately following their spawning season in mid-winter.     

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.     

Temporal and Spatial Trends in Spawning Aggregations of Camouflage Grouper, Epinephelus Polyphekadion, in Pohnpei, Micronesia

The aims of this investigation were to document the temporal and spatial dynamics of aggregated camouflage grouper, Epinephelus polyphekadion in Pohnpei, Micronesia, and to assess seasonal and daily patterns of spawning and reproductive behavior. Camouflage grouper aggregated and spawned adjacent to the Kehpara Marine Sanctuary (KMS), Pohnpei, Micronesia, during two consecutive months in 1998 and 1999. A combination of gonadosomatic indices, oocyte diameter analysis, and histological evidence confirmed that camouflage grouper spawned 1–2 days prior to full moon over an 8–9h period after dusk in each of the four study months. Males entered the site approximately 7 days prior to females and color variations were frequently observed. Significant differences were detected in mean size between the sexes. A series of management initiatives was initiated by the government in the 1980s to reduce fishing during aggregation periods, such as a March–April grouper sales ban, but these were largely ineffective. Following the 1998–1999 survey, several recommendations were made to halt aggregation fishing, including a commercial and subsistence catch and sales ban during spawning months, and the extension of the KMS to incorporate two additional grouper aggregations directly adjacent to it.     

Morphometric analysis of orange roughy (Hoplostethus atlanticus) off the continental slope of southern Australia

Morphometric variation was used to examine the stock structure, in southern Australian waters, of the deepwater marine teleost Hoplostethus atlanticus , orange roughy. Seven samples were collected from non-spawning aggregations in 1989–1990. Three samples were also collected in the winter of 1992, two from the main spawning site off the eastern coast of Tasmania (St Helens), and the third from the other main fishing ground south of Tasmania. The 38 morphometric measurements taken from each of over 1300 fish were size-standardized by an allometric formula and analysed by univariate and multivariate statistics. The results indicate significant variation in the morphology of orange roughy caught from geographically distinct aggregations. They further suggest that the main spawning aggregation may consist of fish from different groups at different times of the spawning period. There appear to be at least seven morphologically distinguishable stocks of orange roughy in southern Australian waters, despite genetic data indicating appreciable levels of gene flow between them.     

Reproductive behavior of the halfmoon grouper, <Emphasis Type="Italic">Epinephelus rivulatus</Emphasis>, at Ningaloo Reef,Western Australia

Behaviors associated with spawning by the halfmoon grouper, Epinephelus rivulatus, at Ningaloo Reef, Western Australia, are described from in situ observations made each evening throughout most of a lunar cycle. Spawning occurred after sunset on six consecutive evenings during the full moon period. During this time males were particularly aggressive toward one another and maintained high levels of activity among the gravid females that rested within each of their territories. Spawning occurred when a male swam alongside a responsive female and the pair rose in a tight spiral 1–1.5 m into the water column before releasing gametes and returning to the seabed. Spawning activity was followed by a longer nonspawning period (ca. 20 days), when fewer intraspecific interactions were observed and gamete reserves were replenished. Histological and behavioral evidence suggests that this cycle of spawning and replenishment may occur on a monthly basis. Although individual fish, particularly females, moved into certain areas to spawn, E. rivulatus did not form spawning aggregations as do larger species of grouper.     

Characterization of large basking shark Cetorhinus maximus aggregations in the western North Atlantic Ocean

Cetorhinus maximus aggregations recorded during extensive aerial survey efforts off the north‐eastern United States between 1980 and 2013 included aggregations centring on sightings with group sizes of at least 30 individuals. These aggregations occurred in summer and autumn months and included aggregation sizes of up to 1398 individuals, the largest aggregation ever reported for this species. The aggregations were associated with sea surface temperatures of 13–24° C and chlorophyll‐a concentrations of 0·4–2·6 mg m^?3 and during one aggregation, a high abundance of zooplankton prey was present. Photogrammetric tools allowed for the estimation of total body lengths ranging between 4 and 8 m. Characterization of these events provides new insight into the potential biological function of large aggregations in this species.     

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.     

The spatial distribution of zooplankton populations in the presence and absence of fish

The spatial distribution patterns of the main zooplankton groups in two small experimental ponds, one which contained fish and one which did not, were measured over a twelve month period from March 1983. Populations of all zooplankton groups were generally distributed non-randomly and with varying degrees of aggregation. The aggregation ofDaphnia longispina populations was inversely related to ambient wind speed, indicating a breakdown of behavioural clumping by wind-induced water currents. Evidence that populations ofDaphnia aggregate in response to fish predation was inconclusive. In contrast cyclopoid nauplii showed no evidence of behavioural clumping and the degree of aggregation was not related to wind speed. Nauplii aggregations were however greater in the absence than in the presence of fish.     

The seabird tick, <Emphasis Type="Italic">Ixodes</Emphasis><Emphasis Type="Italic">uriae</Emphasis>, uses uric acid in penguin guano as a kairomone and guanine in tick feces as an assembly pheromone on the Antarctic Peninsula

In the vicinity of Palmer Station, Antarctica, the seabird tick, Ixodes uriae, forms large aggregations under rocks at the periphery of Adelie penguin rookeries. When the adult penguins return to the rookeries at the beginning of the nesting season the ticks leave their off-host aggregation site, attach to the penguins for a period of feeding, and then subsequently return to the aggregation site. In this study, we searched for chemical cues that may be used by the ticks to locate their aggregation sites as well as cues involved in finding penguins. Tick excreta and soil extracts from beneath tick aggregations contained a pheromone that elicited assembly behavior in unfed larvae, non-fed nymphs and non-fed adults. Guanine, the major excretory product of ticks, elicited assembly behavior, thus, guanine is likely an active component involved in assembly. Non-fed stages also responded positively to penguin guano and uric acid, the primary excretory product of penguins, suggesting that uric acid and other components of penguin guano function as a kairomone used by the non-fed ticks to locate their host. After feeding, the immature ticks’ response to both the assembly and kairomones is switched off for several days, and the ticks regain responsiveness only after they have molted. Fed adult females lay eggs and die without ever regaining responsiveness. Thus, I. uriae relies on two closely related chemicals to regulate two critical aspects of its life: assembly and host-finding. Guanine and other components of tick excreta function as an assembly pheromone in promoting the formation of off-host aggregations, while uric acid and other components of penguin guano function as a kairomone used by the tick to locate its host.     

Layering with depth in a heterospecific fish aggregation

Synopsis The schooling behavior and interspecific interactions within a nearshore planktivorous fish aggregation in Bermuda was observed during the summer (July–Sept) and winter (Dec–Jan) of 1986–87. Four species of morphologically and ecologically similar fish were, studied: the silverside Allanetta harringtonensis, the round herring Jenkensia lamprotaenia, the anchovy Anchoa choerostoma, and the pilchard Harengula humeralis. Heterospecific aggregations of these fish layered with depth into discrete horizontal bands 30 to 60 cm in width. Layers were categorized by unique sets of species as well as by age offish (measured as fish length). The median depth of the layer was also a function of whether the fish were feeding or resting. The median depth and the width of each layer were consistent across three sites and midmorning versus midafternoon observation times. Reduction in predation per individual is suggested as a possible force acting to maintain the structure of this fish aggregation.     

Aggregation and spawning of the humphead wrasse Cheilinus undulatus (Pisces: Labridae): general aspects of spawning behaviour

The humphead wrasse Cheilinus undulatus formed resident spawning aggregations daily after high tide at specific locations along the seaward edge of the Palau barrier reef. The location and extent of one aggregation site remained consistent for 6 years with no physical features distinguishing it from adjacent areas. Spawning was documented most months and probably occurred year round with possible seasonal and lunar variation. Spawning males arrived first at the site, followed by females and potentially small primary males. The aggregation female to male sex ratio was estimated to be between 6:1 and 10:1. A maximum of 15 males and 100–150 females were observed at the site. A male courtship posture with the anal fin pointed, the caudal fin folded down and the dorsal fin folded against the body was maintained while swimming a few metres off the bottom in view of females. When ready to spawn females rose up as the posturing male passed and the pair released gametes in a relatively sedate fashion near the surface along the shelf break. No attempted predation on spawning adults was seen. Egg predation after spawning was uncommon. On days with early to mid‐day high tides the spawning period started 2·0–2·5 h after high tide when the speed of lagoon–ocean tidal currents peaked and lasted c. 1 h. On days with later afternoon high tides, spawning occurred sooner after high tide and before current speeds peaked. Other fishes with planktonic eggs spawned at the site as pairs or small groups in a rough succession after high tide with C. undulatus, the last species to spawn.