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 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.     

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.     

Female Leadership During Migration and the Potential for Sex-Specific Benefits of Mass Spawning in the Brown Surgeonfish (Acanthurus nigrofuscus)

Spawning aggregations in coral-reef fishes have been hypothesized to confer any one of several mutually non-exclusive benefits, largely expected to serve the interests of both sexes simultaneously. Here we provide indication that in the brown surgeonfish, Acanthurus nigrofuscus, spawning aggregations may confer a sex-specific benefit. Following tagged individuals en-route to their daily spawning-aggregation site we found that while migrating groups (≤20 fish) consist of both males and females, females tend to occupy the lead position more often than expected by chance. In addition, we found evidence that female A. nigrofuscus divide their daily egg-stock among several spawning bouts within the aggregations. We propose that female leadership en-route to spawning aggregations, together with the potential benefits of multiple female mating, are consistent with a sex-specific benefit to spawning aggregations.     

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.     

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.     

Ripening of the oocyte of the river lamprey (Lampetra fluviatilis L.) after river entry

River lampreys (Lampetra fluviatilis L.) enter rivers for spawning with their gonads in the final stages of maturation, however the oocytes continue to develop until the spawn. This study was undertaken to depict detailed changes in the oocytes in the period approaching the spawn, using metric analysis and to look for atresia to determine if the time period influences the final fecundity. The study was performed on 37 females caught between October and May in the Rega River, north‐western Poland. Mid‐part sections of fixed ovaries were stained with Heidenhain haematoxylin and PAS. Ten cell structures of the oocytes were measured under light microscope with the aid of a computer image analysis programme. In the autumn, when the first lampreys entered the river, the nuclei of the oocytes were in the polar position. With the approaching mating season many oocyte structures changed significantly. Statistically significant (P < 0.001, Mann–Whitney U‐test) was the increase in oocyte size (up to 1.06 × 0.78 mm in morphometric measurement), enlargement of the yolk platelets (f.a. 8.0 × 5.1 μm), elevation of theca over the chorion followed by the accumulation of glycoconjugates (f. a. 29.2 μm), growth of the zona granulosa at the vegetative pole (f. a. 24.2 μm), and increase in the thickness of the chorion at the animal pole (f. a. 11.1 μm). Other statistically significant changes (P < 0.05 and P < 0.01, Mann–Whitney U‐test) included a decrease in the width of the cortical cytoplasm band (f. a. 20.5 μm), decrease in the ratio of the distance between the nucleus and cell membrane and the long axis of the oocyte (f. a. 7.6 μm), and increase in the thickness of the chorion at the vegetative pole (f. a. 6.0 μm). No statistically significant changes in the area of the nucleus and diameter of the cortical alveoles were noted. Beginning with the lamprey entering the river until the spawn, the oocytes undergo significant growth and maturation. In this period the fecundity of the lamprey decreased only slightly, as atresia was observed sporadically in the ovaries.     

Parasitic Attachments by Overwintering Silver Lampreys, Ichthyomyzon unicuspis, and Chestnut Lampreys, Ichthyomyzon castaneus

We present evidence that at least some parasitic-phase silver lampreys, Ichthyomyzon unicuspis, and chestnut lampreys, I. castaneus, remain attached to host fish during the winter. Lake sturgeon, Acipenser fulvescens, harvested through the ice by spearfishers in the Lake Winnebago system in Wisconsin may bear silver lampreys or fresh lamprey wounds, and sturgeon with lamprey marks were significantly larger than sturgeon without them. Silver lampreys collected on paddlefish, Polyodon spathula, in the Wisconsin River in March were not significantly longer than silver lampreys collected previously in late October, but they were significantly heavier, an indication that they were feeding to at least some extent during the intervening period. Other large fish species, including northern pike, Esox lucius, and flathead catfish, Pylodictus olivaris, have been collected or observed during the winter with silver or chestnut lampreys attached. Although energy and nutrient intake by parasitic lampreys may be reduced during the winter, lampreys attached to hosts may also benefit from the hosts' mobility and ability to avoid potentially harmful situations.     

Variations in the presence of chloride cells in the gills of lampreys (Petromyzontiformes) and their evolutionary implications

Although confined to fresh water, non‐parasitic species of lampreys and the landlocked parasitic sea lamprey, all of which were derived relatively recently from anadromous ancestors, still develop chloride cells, whose function in their ancestors was for osmoregulation in marine waters during the adult parasitic phase. In contrast, such cells are not developed by the non‐parasitic least brook lamprey Lampetra aepyptera, which has been separated from its ancestor for >2 million years, nor by the freshwater parasitic species of the genus Ichthyomyzon. The length of time that a non‐parasitic species or landlocked parasitic form or species has spent in fresh water is thus considered the overriding factor determining whether chloride cells are developed by those lampreys.     

Ontogenetic Behavior and Migration of Dabry's Sturgeon, Acipenser Dabryanus, from the Yangtze River, With Notes on Body Color and Development Rate

We conducted laboratory experiments with Dabry's sturgeon, Acipenser dabryanus, from the upper Yangtze River to develop a conceptual model of early behavior. We daily observed fish from day-0 (embryo, first life interval after hatching) to day-30 feeding larva for preference of bright habitat and cover, swimming distance above the bottom, up- and down-stream movement, and diel activity. Hatchling to day-12 embryos and days 13–24 larvae were similar for ontogenetic behavior, i.e., neither initiated a dispersal migration, both swam within 15cm of the bottom, both preferred bright habitat, and neither strongly preferred cover or open habitat. Embryos and larvae were weakly active day and night. Days 72–76 juveniles had a weak nocturnal downstream migration, indicating wild juveniles disperse from a spawning site. In other sturgeon species yet studied representing three genera on three continents, Dabry's sturgeon is the first that does not disperse as an embryo or larva. Development of Dabry's sturgeon is slow, requiring more cumulative temperature degree days per millimeter of larvae TL than is required for other sturgeons to develop into larvae. Thus, a dispersal migration that diverts energy from development may not be adaptive. The available information suggests the initial dispersal of early life intervals is likely done by females, which spawn in a dispersed spawning style, not the usual aggregated spawning style. Juvenile migrants had a black body and tail with a light line along the lateral scutes. The color of juvenile migrants shows that a dark body and tail is characteristic of Acipenser that migrate downstream as larvae or juveniles.     

Identification of a female spawn‐associated Kazal‐type inhibitor from the tropical abalone Haliotis asinina

Abalone (Haliotis) undergoes a period of reproductive maturation, followed by the synchronous release of gametes, called broadcast spawning. Field and laboratory studies have shown that the tropical species Haliotis asinina undergoes a two‐week spawning cycle, thus providing an excellent opportunity to investigate the presence of endogenous spawning‐associated peptides. In female H. asinina, we have isolated a peptide (5145 Da) whose relative abundance in hemolymph increases substantially just prior to spawning and is still detected using reverse‐phase high‐performance liquid chromatography chromatograms up to 1‐day post‐spawn. We have isolated this peptide from female hemolymph as well as samples prepared from the gravid female gonad, and demonstrated through comparative sequence analysis that it contains features characteristic of Kazal‐type proteinase inhibitors (KPIs). Has‐KPI is expressed specifically within the gonad of adult females. A recombinant Has‐KPI was generated using a yeast expression system. The recombinant Has‐KPI does not induce premature spawning of female H. asinina when administered intramuscularly. However it displays homomeric aggregations and interaction with at least one mollusc‐type neuropeptide (LRDFVamide), suggesting a role for it in regulating neuropeptide endocrine communication. This research provides new understanding of a peptide that can regulate reproductive processes in female abalone, which has the potential to lead to the development of greater control over abalone spawning. The findings also highlight the need to further explore abalone reproduction to clearly define a role for novel spawning‐associated peptide in sexual maturation and spawning. Copyright © 2016 European Peptide Society and John Wiley & Sons, Ltd.     

Disturbance of freshwater habitats by anadromous salmon in Alaska

High densities of habitat modifiers can dramatically alter the structure of ecosystems. Whereas spawning sockeye salmon (Oncorhynchus nerka) dig nests that cover over 2 m² and are at least 20 cm deep, and can spawn at high densities, relatively little attention has been devoted to investigating the impacts of this disturbance. We hypothesized that this temporally and spatially predictable bioturbation has large impacts on the coastal aquatic habitats used by sockeye. We experimentally investigated the impacts of disturbance caused by spawning sockeye in two streams and two lakes in Alaska by excluding salmon from 2.25 m² plots where they traditionally spawn. We sampled exclusions and control plots before, during, and after spawning. During sockeye spawning, fine sediment accumulated in areas where sockeye were excluded from spawning. In addition, sockeye spawning significantly decreased algal biomass by 80% compared to exclusion plots. We found mixed effects of spawning on the invertebrate assemblage. Tricladida and Chironomidae densities increased by 3x in exclusion plots relative to control plots in one creek site. However, for most taxa and sites, invertebrate densities declined substantially as spawning progressed, regardless of experimental treatment. Habitat modification by spawning salmon alters both community organization and ecosystem processes.     

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.     

Natural spawning of a Hawaiian sea urchin,Tripneustes gratilla

Observations of natural spawning events are rare but critically important for the field of fertilization ecology. For broadcast spawners, knowing the natural behavior of spawning, including proximity of animals and the timing of spawning, is essential for informing spawning experiments and directing future study. Here we describe a natural spawning event that took place on the island of Maui in the winter of 2010. Groups of Tripnuestes gratilla were observed to spawn in the late afternoon shortly after a local high tide that was a lower high of a mixed, semidiurnal tide cycle. Urchins appeared to increase the probability of fertilization by ascending to the local high points of the reefs and spawning within groups of 2–5. Although the majority of the urchins that spawned were T. gratilla, two individuals of Echinothrix diadema were observed to spawn at the same time, raising questions about potential hybridization in the wild.     

Mass spawning aggregation of the giant bumphead parrotfish Bolbometopon muricatum

The present study reports a previously undocumented mass spawning aggregation and group spawning phenomena of c. 1200 individual bumphead parrotfish Bolbometopon muricatum in Palau, Micronesia. Although B. muricatum are protected in Palau, it is further recommended that management strategies should consider establishment of no‐take zones at B. muricatum spawning aggregations and concomitant sleeping grounds elsewhere.     

Brood parasitism of an open-water spawning cichlid by the cuckoo catfish

The cuckoo catfish Synodontis multipunctatus and S. grandiops are endemic to Lake Tanganyika and the only known nonavian vertebrates that exhibit obligate interspecific brood parasitism. Seven maternal mouth-brooding cichlid fish species are reported to be natural hosts of the parasitic catfish and share a common reproductive behaviour that the catfish exploits: cichlid females spawn eggs on the bottom, allowing the catfish female to place her eggs near the cichlid eggs, and the cichlid females collect the catfish eggs by mouth together with their own eggs. However, so far it has not been reported that the cuckoo catfish exploit different spawning behaviours. The genus Cyprichromis consists of five maternal mouth-brooding species endemic to Lake Tanganyika, most of which spawn and collect eggs in open water. This study reports that the cuckoo catfish also parasitizes the open-water spawning Cyprichromis coloratus, although it may not be a regular host.     

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.     

Use of behavioral and physiological indicators to evaluate Scaphirhynchus sturgeon spawning success

Thirty gravid, female shovelnose sturgeon (Scaphirhynchus platorynchus) were captured in the Lower Missouri River in March 2004 to evaluate the effectiveness of physiology, telemetry and remote sensor technology coupled with change point analysis in identifying when and where Scaphirhynchus sturgeon spawn. Captured sturgeons were instrumented with ultrasonic transmitters and with archival data storage tags (DST) that recorded temperature and pressure. Female sturgeon were tracked through the suspected spawning period. Thereafter, attempts were made to recapture fish to evaluate spawning success. At the time of transmitter implantation, blood and an ovarian biopsy were taken. Reproductive hormones and cortisol were measured in blood. Polarization indices and germinal vesicle breakdown were assessed on the biopsied oocytes to determine readiness to spawn. Behavioral data collected using telemetry and DST sensors were used to determine the direction and magnitude of possible spawning‐related movements and to identify the timing of potential spawning events. Upon recapture observations of the ovaries and blood chemistry provided measures of spawning success and comparative indicators to explain differences in observed behavior. Behavioral and physiological indicators of spawning interpreted along with environmental measures may assist in the determination of variables that may cue sturgeon reproduction and the conditions under which sturgeon successfully spawn.     

Communal Spawning in the Common Frog Rana temporaria– Egg Temperature and Predation Consequences

Females of the common frog, Rana temporaria, spawn synchronously at communal breeding sites, forming a communal egg mass of individual spawn clumps. A total of four spawning sites were studied. In the centre of the communal egg mass, daily maximum temperature was higher and minimum temperature lower than further out. Spawn clumps in the centre of the communal egg mass developed faster than single spawn clumps. In the centre of the egg masses, some spawn clumps suffered almost total egg mortality. This was probably the consequence of a cold spell with sub‐zero temperatures in combination with a tendency for low minimum temperatures in the centre. In a laboratory study, worm leeches, Erpobdella spp., fed readily on eggs of common frog, while horse leeches, Haemopis sanguisuga, hardly ate any eggs. At the spawning sites, most worm leeches were found in single spawn clumps, far from the communal egg mass. Females which spawned late, when most of the spawning was over, preferred to spawn in the centre of the communal egg mass. This suggests that there is a net advantage for eggs in spawn clumps laid in the centre.