Temporal settlement patterns of larvae of the broadcast spawning reef coral Favites chinensis and the broadcast spawning and brooding reef coral Goniastrea aspera from Okinawa, Japan

The faviid corals, Favites chinensis and Goniastrea aspera are widely distributed in the Indo-Pacific region. Both corals are hermaphroditic broadcast spawners, but G. aspera is also known to brood planula larvae in Okinawa. This study investigated the temporal settlement patterns of planula larvae of the scleractinian corals F. chinensis and G. aspera that developed from spawned gametes, and planula release and settlement of brooded larvae of G. aspera from Okinawa, Japan. Some of the broadcast-spawned larvae of F. chinensis and G. aspera had very short pre-competency periods of 1–2 and 2–3 days after spawning, and relatively long maximum settlement-competency periods of 56–63 and 63–70 days after spawning, respectively. These pre-competency periods are among the shortest reported for larvae of broadcast spawning coral species, and appear to be negatively correlated with seawater temperature. F. chinensis larvae tended to settle rapidly with 34–39% of larvae settling in the first week after spawning, while broadcast-spawned G. aspera larvae had a slower settlement pattern with 11–15% of larvae settling in the first week after spawning. Brooded larvae of G. aspera settled more rapidly, with settlement rates of 27–31% within the first 24 h and 45–65% within the first week after the start of the experiment. The production of planula larvae with rapid settlement capabilities may enable F. chinensis and G. aspera to establish and maintain populations in shallow reef sites at Okinawa. The release of the brooded planulae for up to 2 months may explain why G. aspera is locally more dominant on shallow reefs in Okinawa than F. chinensis. On a broader scale, the longer settlement competency periods of some of the broadcast-spawned larvae of these species increase their potential for longer-distance dispersal and may partly explain the wide biogeographic distribution of these species in the Indo-Pacific region.     

The parachute function of the hull in eggs of Mopalia kennerleyi (Chitonida: Mopaliidae), and swimming of its larvae through ontogeny

Free‐spawning species of chitons produce eggs enclosed in a coating known as the hull. In Chitonida, several studies have shown that the hull helps to direct sperm to specific areas of the egg surface, facilitating fertilization. One study has found evidence that this structure also serves to reduce the sinking rates of the eggs. To clarify how the presence of the hull modifies sinking rates in chiton eggs, here we compare sinking speeds and densities of eggs of Mopalia kennerleyi with and without the hull. Sinking rates of eggs with the hull were approximately one‐third of those without it. This structure acts as a flotation device because it has a density very close to that of seawater, and it increases the effective diameter and therefore the drag on the negatively buoyant egg. Since there is limited knowledge about morphology and behavior of chiton larvae, we also analyzed changes during ontogeny in behavior, swimming speeds, and body shape of larvae of M. kennerleyi. Over time, the larvae decreased their upward swimming tendency and preferred to stay near the bottom, and their bodies became elongated and dorso‐ventrally compressed. These changes may be related to preparation for settlement and metamorphosis. Further studies of these subjects are required in chitons, since movement of early stages, as eggs/embryos sinking or larvae swimming in the water column, may affect their survival.     

Rapid settlement in broadcast spawning corals: implications for larval dispersal

Broadcast spawning of gametes with planktonic development of larvae is the most common reproductive mode in tropical corals, and is generally thought to optimize the dispersal potential of larvae. To this end, many previous studies of coral larval dispersal have focused on the maximum time larvae can remain competent to settle and consequently how far they might disperse. However, dispersal ability of broadcast-spawned coral larvae will be linked, at least in part, to the minimum time to settlement competency as well as the length of the planktonic period—although estimates of minimum time to competency remain largely anecdotal, with few rigorous studies of the pre-competent period. To determine the minimum time to larval settlement in two species of broadcast-spawning coral ( Platygyra daedalea and Goniastrea favulus), we monitored larval settlement rates in aquaria every 6 h from the time larvae commenced swimming (i.e. were ciliated, fully developed larvae) for a period of approximately 10 days. For P. daedalea, peak settlement occurred between 60 and 66 h following fertilization (2.5 and 2.75 days), which is markedly earlier than the 4- to 6-day time period commonly cited as the minimum time before broadcast-spawned coral larvae are competent to settle. Surprisingly, it was also clear from our experimental results that settlement in P. daedalea occurred as a distinct pulse during the 60- to 66-h period, rather than continuously throughout the study period. G. favulus larvae also appear to be able to settle quickly (from 54 h following fertilization). We argue, on the basis of these short competency times and apparently rapid settlement, that dispersal in broadcast-spawning coral larvae may not be as great as has previously been assumed.     

Isolation of microsatellites from two species of scleractinian coral

Microsatellites were isolated from two broadcast‐spawning species of scleractinian coral (Platygyra daedalea and Goniastrea favulus) from Australia's Great Barrier Reef. We found 27 microsatellites across both species, although only five loci were polymorphic in each species. Microsatellite loci displayed a wide range of diversity levels with four to 11 alleles per locus (H_O = 0.26–0.91) in P. daedalea and two to seven alleles per locus (H_O = 0.16–0.96) in G. favulus. Most loci showed departures from Hardy–Weinberg equilibrium which may reflect nonrandom mating but may also be related to difficulties associated with coral DNA.     

The choice of oviposition site and the effects of density and oviposition timing on survivorship in<Emphasis Type="Italic"> Rana japonica</Emphasis>

In species that lack parental care and whose larvae are restricted to a given space throughout development, choice of egg-laying or reproductive sites often greatly influences fitness. Furthermore, reproductive success of individuals may be affected not only by conditions of the breeding area but also by the time when they start to breed, because intraspecific variation in the timing of breeding may result in asymmetric competition between larvae. We addressed these issues in an experimental and field study of the Japanese brown frog, Rana japonica. We tested whether the survival success of larvae decreased with an increase in the number of deposited egg masses and with a greater delay in oviposition. We found that the number of egg masses and timing of oviposition together significantly predicted larval survival per clutch. Moreover, we observed the natural oviposition of R. japonica to examine the prediction that if the density of larvae and the timing of oviposition affect survivorship of larvae, R. japonica will avoid depositing eggs in pools in which egg masses have already been deposited. We found that for small pools only, R. japonica tended to favor oviposition at unoccupied pools (those lacking egg masses and larvae) in the spawning season. For large pools, however, adults favored those already occupied by egg masses in the spawning period. The density of egg masses and tadpoles or the timing of oviposition may have less effect on spawning or the survival of tadpoles in large pools than in small pools.     

The effect of selected trace metals on the fertilization success of several scleractinian coral species

This study provides new information on the effects of various concentrations of the trace metals copper, lead, zinc, cadmium, and nickel on fertilization success of gametes from the scleractinian reef corals Goniastrea aspera,Goniastrea retiformis, Acropora tenuis, and Acropora longicyathus. The EC50 values (the concentration that reduces the fertilization rate by 50% relative to the control fertilization) for copper effects on fertilization success of these coral species range from 15 to 40 μg/L, which is similar to responses of other marine invertebrates. The EC50 values for lead were 1450–1800 μg/L for the Acropora species, and >2400 μg/L for G. aspera gametes, which indicates that lead was much less toxic than copper. Fertilization responses to zinc and nickel were variable and a significant reduction in fertilization success for A. tenuis gametes was found only at very high cadmium concentrations. The data from this study and other recent research clearly demonstrate that some trace metals impair the fertilization success of gametes from faviid and acroporiid reef corals. Trace metal inputs into reef waters should be limited and controlled to avoid potential interference with sexual reproductive processes of reef corals.     

The evolution of offspring size and number: a test of the Smith-Fretwell model in three species of crickets

Most models of parental investment in offspring assume a trade-off between propagule size and number, and an increasing concave down function relating offspring fitness to propagule size. In this study, we test these two fundamental assumptions, using three closely related species of crickets, Gryllus firmus, G. veletis, and G. pennsylvanicus. Egg weight, 35-day fecundity and 35-day egg biomass were estimated in a population of each species, and the relationships between these reproductive traits and date of egg laying and body size were estimated. The relationships between egg weight and offspring survival were also sought for eggs buried at different depths, soil moistures, and soil types (G. firmus and G. veletis), as well as in the field (G. pennsylvanicus). A trade-off between egg weight and 35-day fecundity was revealed in a multivariate analysis taking into account among-species variation in egg weight and body size. Independent of the environmental conditions affecting the eggs, a positive correlation existed between the number of larvae that emerged from the soil and propagule weight in each species. Therefore, these findings provide partial support for the assumptions considered in the models mentioned above. A single optimal egg size was favored in two out of the three sets of conditions in which the functions relating egg weight to larval survival could be derived. The conditions encountered by the eggs, however, influenced the average survival of the larvae, as well as the shape of the relationship between egg weight and offspring survival. This suggests that cricket eggs frequently face heterogeneous environments with respect to egg and hatchling survival; the implication of habitat heterogeneity on the evolution of an optimal egg size is considered. The relationships between the reproductive components and female age and size, as well as between egg size and variation in cricket life-history, are discussed in an ecological and evolutionary context.     

Spawning strategy in Atlantic bobtail squid <Emphasis Type="Italic">Sepiola atlantica</Emphasis> (Cephalopoda: Sepiolidae)

This study aimed to determine the spawning strategy in the Atlantic bobtail squid Sepiola atlantica, in order to add new information to the knowledge of its reproductive strategy. A total of 12 females that spawned in aquaria were examined. Characteristics of the reproductive traits and egg clutches were similar to those of other known Sepiolidae. Clutch size varied from 31 up to 115 eggs. Females of this species had incorporated up to 1.58 times of their body weight into laid eggs. The size of laid eggs showed a positive correlation with maternal body size, supporting the idea that female size is a determinant of egg size. Our data suggest that S. atlantica is an intermittent terminal spawner, and that its spawning strategy comprises group-synchronous ovary maturation, multiple egg laying, and deposition of egg clutches in different locations. The obtained data provide insights for future comparative studies on reproductive allocation.     

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.     

Dispersion of vendace eggs and larvae around potential nursery areas reveals their reproductive strategy

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Reproductive biology of four large riverine fishes (Cyprinidae) in a man-made lake,Orange River,South Africa

The reproductive biology ofBarbus holubi, B. kimberleyensis, Labeo capensis andL. umbratus was examined in a large reservoir on the Orange River, South Africa. The findings are integrated into the existing knowledge on largeBarbus andLabeo species, which coexist in most river systems in Africa and Asia.LargeBarbus spawn on gravel beds within the river channel during floods in spring or summer. In continuously flowing regulated rivers, time of spawning is governed by water temperatures. They have moderate fecundity; large eggs, incubation time of several days and the larvae are initially immobile with large yolk sacs. In the impoundment, they spawn in the inflowing regulated river withB. kimberleyensis spawning four to six weeks later than the more cold-tolerantB. holubi, the dominant largeBarbus. Survival is generally good and the juveniles disperse throughout the lake, but unseasonal release of cold water from an upstream impoundment may cause poor reproductive success.In contrast,Labeo species generally spawn on newly flooded ground, usually leaving the main river channel. Spawning may or may not be preceded by a longitudinal migration. Labeos are relatively fecund with small (30%Barbus size) eggs which hatch quickly and the larvae swim in bursts up into the water column before sinking down again.L. capensis does not require a longitudinal spawning migration and breeds throughout the lake, depending on local conditions. Large temporal variation in gonadal development within the population can result in more than one spawning. Dispersal within the lake is poor.L. umbratus uses larger inflowing tributaries for spawning thanL. capensis and its juveniles have a much greater power of dispersal. Early dependence on external feeding and undependable occurrence of conditions for spawning and juvenile feeding makes for variable reproductive success.     

Reproductive allocation and reproductive ecology of seven species of Diptera

1. Variation in resource allocation to egg size and number was investigated in seven sympatric species of Piophilidae that oviposit on carcasses or discarded cervid antlers: Liopiophila varipes (Meigen), Prochyliza xanthostoma Walker, Protopiophila latipes (Meigen), Protopiophila litigata Bonduriansky, Stearibia nigriceps (Meigen), and two unidentified species of Parapiophila McAlpine. 2. Following optimal reproductive allocation theory, relatively larger, fewer eggs were expected in (1) species that oviposit on antlers, where larvae probably experience lower risk of predation and greater competition than larvae in carcasses, and (2) species with aggressive males and male-biased sex ratios on the oviposition substrate, where risk of injury during oviposition may have favoured females laying fewer eggs. 3. Variation in reproductive allocation strategies could not be explained by known differences in larval or adult environment, but congeneric species clustered by reproductive allocation patterns. The Parapiophila species produced larger, fewer eggs than the other species, and egg number increased slowly with body size. The Protopiophila species did not deviate from expected egg sizes and numbers, and egg number increased steeply with body size. 4. An interspecific egg size–egg number trade-off resulted in a tight linear scaling of ovary volume to body size, suggesting common physiological constraints on relative ovary mass. 5. Within each species, egg size was nearly constant whereas egg number increased with female body size, suggesting species-specific stabilising selection on egg size.     

Population structure is not a simple function of reproductive mode and larval type: insights from tropical corals

1. For a wide range of organisms, heritable variation in life-history characteristics has been shown to be strongly subject to selection, reflecting the impact that variation in characters such as genotypic diversity, duration of larval development and adaptations for dispersal can have on the fitness of offspring and the make-up of populations. Indeed, variation in life-history characteristics, especially reproduction and larval type, have often been used to predict patterns of dispersal and resultant population structures in marine invertebrates. 2. Scleractinian corals are excellent models with which to test this relationship, as they exhibit almost every possible combination of reproductive mode and larval type. Some general patterns are emerging but, contrary to expectations, genetic data suggest that while populations of broadcast spawning species may be genotypically diverse they may be heavily reliant on localized recruitment rather than widespread dispersal of larvae. 3. Here we use microsatellites to test the importance of localized recruitment by comparing the genetic structure of populations of two broadcast spawning corals with contrasting modes of reproduction and larval development; Goniastrea favulus is self-compatible, has sticky, negatively buoyant eggs and larvae and is expected to have restricted dispersal of gametes and larvae. In contrast, Platygyra daedalea is self-incompatibile, spawns positively buoyant egg-sperm bundles and has planktonic development. 4. Surprisingly, spatial-autocorrelation revealed no fine-scale clustering of similar genotypes within sites for G. favulus, but showed a non-random distribution of genotypes in P. daedalea. Both species showed similar levels of genetic subdivision among sites separated by 50-100 m (F(ST) = 0.03), suggesting that larval dispersal may be equivalent in both species. 5. Interestingly, as fragmentation has been considered rare in massive corals, our sample of 284 P. daedalea colonies included 28 replicated genotypes that were each unlikely (P < 0.05) to have been derived independently from sexual reproduction. 6. We conclude that the extreme life history of G. favulus does not produce unusually fine-scale genetic structure and subsequently, that reproductive mode and larval type may not be not good predictors of population structure or dispersal ability.     

Ontogenetic change in the lipid and fatty acid composition of scleractinian coral larvae

Some scleractinian coral larvae have an extraordinary capacity to delay metamorphosis, and this is reflected in the large geographic range of many species. Coral eggs typically contain a high proportion of wax esters, which have been hypothesized to provide a source of energy for long-distance dispersal. To better understand the role of lipids in the dispersal of broadcast spawning coral larvae, ontogenetic changes in the lipid and fatty acid composition of Goniastrea retiformis were measured from the eggs until larvae were 30 days old. Egg biomass was 78.8 ± 0.5% lipids, 86.3 ± 0.2% of which were wax esters, 9.3 ± 0.0% polar lipids, 4.1 ± 0.2% sterols, and 0.3 ± 0.1% triacylglycerols. The biomass of wax esters declined significantly through time, while polar lipids, sterols and triacylglycerols remained relatively constant, suggesting that wax esters are the prime source of energy for development. The most prevalent fatty acid in the eggs was palmitic acid, a marker of the dinoflagellate Symbiodinium, highlighting the importance of symbiosis in coral reproductive ecology. The proportion of polyunsaturated fatty acids declined through time, suggesting that they are essential for larval development. Interestingly, triacylglycerols are only abundant in the propagules that contain Symbiodinium, suggesting important differences in the energetic of dispersal among species with vertical and horizontal transmission of symbionts.     

Delayed maturation in the colonial coralGoniastrea aspera (Scleractinia): whole-colony mortality, colony growth and polyp egg production

The present study examines (1) the cost of reproduction on colony growth, and (2) relationships among sexual maturity, whole-colony mortality rate and colony growth rate inGoniastrea aspera free from external influences by macrobenthos. Survival of colonies in permanent plots was followed for two years. Egg production by polyps in colonies collected just before the first spawning of a year was estimated by dissecting the polyps. Growth of the colonies (increase in number of polyps) was followed over one annual reproductive cycle. The cost of egg production on colony growth was apparent through colony ontogeny: (1) immature colonies had a greater annual growth rate than mature colonies, but produced almost no eggs; (2) in mature colonies, growth rate was negatively correlated with NE/PV (number of eggs per polyp volume mm^-3). Annual whole-colony mortality was high in colonies with fewer than11 polyps in initial colony size, while mortality was extremely low once a colony grew beyond this size. This critical size for low whole-colony mortality was much smaller than the colony size (40 polyps) which would attain maturity one year later. Age at maturity was estimated as six years. While survival to maturity may be a selective force for the evolution of delayed maturation, the present data suggest that high colony fecundity, achieved after a long growth period as an immature colony, and an abrupt decrease of colony growth rate after maturation are the crucial forces.     

The behaviour of Abudefduf zonatus (Pisces,pomacentridae) at Heron Island,Great Barrier Reef

Social organization and reproductive behaviour of Abudefduf zonatus were studied at Heron Island, Great Barrier Reef. This species is closely associated with the sand and coral substrate of the inner reef flat zone. Territorial, clustering, nest-digging, breeding and parental care behaviours are described. Mature males maintain territories for several weeks or longer around empty clam shells in which periodic spawning occurs. Females and sub-adult males hold territories interspersed among those of adult males. No pair bonds are formed, and eggs are guarded only by males. Breeding is roughly synchronized among fish of one area. Clustering behaviour may contribute to breeding syndrony in local populations.     

The reef coral Goniastrea aspera: a ‘winner’ becomes a ‘loser’ during a severe bleaching event in Thailand

The reef coral Goniastrea aspera is regarded as one of the most environmentally tolerant species on Indo-Pacific reefs. Its demise, following a severe bleaching event in the Andaman Sea in the north-eastern Indian Ocean in 2010, was surprising in view of the rapid recovery of co-existing species such as Porites lutea. Demographic studies of G. aspera at this site showed the population was mainly composed of large individuals, which recruited in the early 1990s. These results, and size-specific mortality observed in G. aspera, post-bleaching, suggest that factors, related to size and age, may have contributed to the coral’s marked decline.     

DOES BATEMAN'S PRINCIPLE APPLY TO BROADCAST-SPAWNING ORGANISMS? EGG TRAITS INFLUENCE IN SITU FERTILIZATION RATES AMONG CONGENERIC SEA URCHINS

Evolutionary biologists generally invoke male competition and female choice as mechanisms driving sexual selection. However, in broadcast-spawning organisms sperm may be limiting and females may compete, in the Darwinian sense, for increased mating success. In this study, I investigate how species differences in egg and sperm traits result in different patterns of fertilization among three closely related sea urchins (Strongylocentrotus purpuratus, S. franciscanus, and S. droebachiensis). Field studies demonstrate that all three species achieve similar percentages of eggs fertilized when eggs and sperm are released simultaneously. However, when sperm must disperse before encountering eggs, differences arise among species such that those with the smaller eggs and faster but shorter-lived sperm achieve relatively fewer fertilizations than do species with larger eggs and slower but longer-lived sperm. A field hybridization experiment, field estimates of sperm dispersal, correlations of egg size to field rates of fertilization, laboratory studies of fertilization kinetics, and a simulation model all suggest that it is attributes of the egg (probably egg size) that are responsible for the differences. These patterns of fertilization match the species' patterns of dispersion; species that do well only when sperm and eggs are released in close proximity are more aggregated, species that do relatively well when sperm and eggs are released farther apart are more dispersed. These results are consistent with the notion that eggs of different species are adapted to maximize reproductive success under different degrees of sperm limitation and suggest that male competition and female choice may not be an appropriate dichotomy in broadcast-spawning organisms.     

Seasonality and life histories of two endemic New Zealand carabid beetles (Coleoptera: Carabidae): Mecodema oconnori Broun and Megadromus capito (White)

Abstract

Seasonal activity and life histories of two large endemic carabid beetles, Mecodema oconnori Broun and Megadromus capito (White), were investigated in two lowland forest remnants in the Horowhenua region, North Island, New Zealand. Seasonal activity was estimated from manual searches of surface refuges, plus pitfall trapping from October to March. Adults of both species were present throughout the year, although M. oconnori was much more abundant in spring. Megadromus capito larvae were caught from October to March, and the three instars overlapped temporally, implying a long period of larval emergence and development. Adult females were dissected to investigate their reproductive phenology. Both species carried eggs for at least 6 months of the year. M. oconnori females contained very few eggs, with a maximum of three eggs each and a mean of 1.6 eggs per female. Mg. capito females had up to 28 eggs each and a mean of 10.5. Both species appear to have low egg loads, a relatively long period of reproductive activity and long life spans, but longer‐term studies on the pre‐adult stages and reproductive activity of adults are required to complete understanding of their life cycles.     

Mating behavior in the black hamlet — gamete trading or egg trading?

Synopsis The black hamlet (Hypoplectrus nigricans, Serranidae) is a simultaneous hermaphrodite, like many other serranines. It has external fertilization and planktonic eggs and engages in a kind of reciprocal spawning consisting of three components: (1) The clutch is divided into sequentially spawned parcels. (2) Partners regularly alternate release of parcels in a spawning bout. (3) Courtship is associated with the female spawning role. Two hypotheses have been proposed to account for this pattern. The egg trading hypothesis states that the pattern results from competition for fertilizations and assumes that reproductive success (RS) as a male is limited by access to eggs and that female RS is not limited by access to sperm. The gamete trading hypothesis states that the pattern results from eggs being at substantial risk of not being fertilized — i.e. female RS is limited by access to sperm. An analysis was performed of data from the black hamlet and three other serranines to determine whether significant sperm limitation occurs. The evidence fails to support the hypothesis that access to sperm limits female RS. Unspawned eggs were not found in fish collected outside the spawning period, but fish without eggs always had milt (sperm). The percentage of eggs fertilized did not decrease over the course of a spawning bout. There was also a slight positive correlation between the number of eggs released in a spawning and the percentage that were fertilized, and serranines that divide the clutch into parcels do not tend to have higher fecundities than those that do not. The gamete trading hypothesis can therefore be rejected for H. nigricans.