A phylogenetic analysis of egg size,clutch size,spawning mode,adult body size,and latitude in reef fishes

Theoretical treatments of egg size in fishes suggest that constraints on reproductive output should create trade-offs between the size and number of eggs produced per spawn. For marine reef fishes, the observation of distinct reproductive care strategies (demersal guarding, egg scattering, and pelagic spawning) has additionally prompted speculation that these strategies reflect alternative fitness optima with selection on egg size differing by reproductive mode and perhaps latitude. Here, we aggregate data from 278 reef fish species and test whether clutch size, reproductive care, adult body size, and latitudinal bands (i.e., tropical, subtropical, and temperate) predict egg size, using a statistically unified framework that accounts for phylogenetic correlations among traits. We find no inverse relationship between species egg size and clutch size, but rather that egg size differs by reproductive mode (mean volume for demersal eggs = 1.22 mm³, scattered eggs = 0.18 mm³, pelagic eggs = 0.52 mm³) and that clutch size is strongly correlated with adult body size. Larger eggs were found in temperate species compared with tropical species in both demersal guarders and pelagic spawners, but this difference was not strong when accounting for phylogenetic correlations, suggesting that differences in species composition underlies regional differences in egg size. In summary, demersal guarders are generally small fishes with small clutch sizes that produce large eggs. Pelagic spawners and egg scatterers are variable in adult and clutch size. Although pelagic spawned eggs are variable in size, those of scatterers are consistently small.     

Racial differences in eggs and juveniles of Windermere charr, Salvelinus alpinus

At least four races of charr occur in Windermere, the largest natural lake in England: north basin and south basin autumn spawners, north basin and south basin spring spawners. This study examines racial differences between eggs and juveniles, and relates juvenile size and survival to egg size. There were no major differences between races for egg incubation times and the percentage of eggs hatching successfully, the latter being high (mean values 76–96%) with a negligible proportion of abnormal alevins (<0.8%). Although there were no significant differences in the lengths of the female parents, both eggs and alevins were significantly larger for the autumn spawners than the spring spawners. Size differences in alevins, especially live weight, were positively related to egg size but not female parent size. Mean percentage survival for juveniles attaining the independent feeding stage was higher for the progeny of autumn spawners (32%) than spring spawners (3%). Racial differences in the egg and alevin stages therefore appear to have a significant effect on subsequent survival, and could be ultimately responsible for the relatively small proportion of spring spawners (only 4–6%) in the Windermere population of charr.     

Parent–egg–progeny Relationships in Teleost Fishes: An Energetics Perspective

Age-related variations in chemical composition of egg matter were found in females in some studies, but they do not seem to be a universal phenomenon. In contrast, egg size can be well predicted from female age. The relationship has a parabolic shape, but the predicted size decrease of eggs from old females has not always been documented. Female size is an important contributor to egg size, both at intra- and inter-specific levels. Dependence of fecundity on body size has usually been described by a power function. A trade-off between egg number and size is considered in light of life history strategies. During a spawning season egg size may differ between successive batches, but lack of effects of egg batch sequence was reported in some studies. In yolk-feeding fish three discrete periods of elevated mortality are typically observed: shortly after egg activation, during hatching, and at final yolk resorption. The positive relationships between female size, egg size and offspring size/resistance to starvation and predation are a key pathway in parent–egg–progeny relationships. Both maternal and paternal effects contribute to the total survival of offspring, but they operate in different ways and at different times. In contrast to the importance of female size, no paternal size effects were revealed, but density and motility of spermatozoans were decisive. Typically, paternal effects diminish earlier in ontogeny. Major factors governing embryonic survival (fertilisation success and hatching success) differ from factors to which starvation mortality of yolk-feeding larvae is related. Embryonic survival is affected by female age via egg matter composition, by egg ripeness and paternal factors such as sperm density and motility. In contrast, starvation mortality of yolk-feeding larvae depends largely on female attributes (age, size and fecundity) via egg size, and, in some batch spawners, on egg batch sequence. Among teleost species egg size varies across a wide range (from 0.3 to 85–90 mm in diameter). Species that spawn large eggs are relatively rare. Caloric value of egg dry matter varies within a narrow range of 20–30 J mg⁻¹. Ecosystem and evolutionary components, and reproductive style are factors that contribute to egg endowment and yolk quality. During the last decade considerable progress was made in the methodology and understanding of paternal effect on progeny performance in fishes. This paper reviews these of parent–egg–progeny relationships.     

Relationship between pelagic larval duration and abundance of tropical fishes on temperate coasts of Japan

The influence of pelagic larval duration (PLD) and egg type dispersal capabilities of 35 demersal and pelagic-spawning tropical fish species is examined in relation to their abundance on the temperate coasts of Japan. The PLDs of pelagic spawners were significantly longer than those of demersal spawners, and a high occurrence of pelagic spawners on the temperate coasts suggests that these fishes are more easily transported to temperate coasts than demersal spawners. For demersal spawners, the common species on the temperate coasts had significantly longer PLDs than the rare species; this suggests that PLD is a major factor influencing the distribution patterns of tropical demersal spawners on temperate coasts. Moreover, a negative correlation between PLD and the abundance of some species of pelagic and demersal spawners suggests the presence of reproductively active fishes in northern subtropical and even in temperate waters.     

Broadcasting fables: Is external fertilization really primitive? Sex,size, and larvae in sabellid polychaetes

Traditionally, broadcast spawning and planktonic larvae have been considered the plesiomorphic ‘ground plan’ for the Polychaeta and other metazoan groups. To assess whether this reproductive mode is in fact ‘primitive’, the study of monophyletic groups with various reproductive modes should be informative. A large range of body sizes would allow testing the ideas that aspects of reproductive mode may be functionally constrained. The family Sabellidac is one such group, with sexual reproductive modes ranging from broadcast spawning to intratubular brooding to ovovivi-parity, and a body size range over more than five orders of magnitude. Sabellids have previously been the subject of detailed cladistic analyses (Fitzhugh 1989, 1991); here we introduce several new characters based on morphology of reproductive structures. Larval development in four brooding sabellid species is also described with the aim of introducing new characters for future systematic analyses. Our cladistic analysis of sabellid genera suggests that gonochorism and brooding of direct-developing larvae are plesiomorphic in the Sabellidae, with external fertilization and swimming larvae limited to apomorphie clades in the subfamily Sabellinae. The presence of sperm with elongate heads may be correlated with the presence of intratubular brooding, though an adequate causal explanation for this relationship can not yet be presented. The concept that ‘modified’ sperm must be derived from ‘primitive’ sperm is shown to be false, with ‘modified’ sperm being plesiomorphic for the Sabellidae, from which ‘primitive’ sperm is derived in apomorphic Sabellinae. All sabellids have lecithotrophic development and appear to be phylogenetically constrained in this regard. Data gathered on body size and reproductive variables in the Sabellidac suggests the following (when phylogenetic effects are not controlled): (1) egg number and total egg volume are significantly correlated with body size, with small animals having fewer, larger eggs than large animals; (2) individual egg volume is not correlated with body size; (3) reproductive mode is significantly correlated with body size; intratubular brooders tend to be small-bodied, whereas broadcast spawners are large. However when the effect of body size is controlled for, then (4) egg number, egg volume and total egg volume all vary significantly with reproductive mode. Broadcast spawners expel a large number of small eggs for a high total egg volurne. Intratubular brooders have a few relatively large eggs for a small total egg volume. When statistics arc performed using phylogenetically independent contrasts there is a significant correlation between total egg volume and body size but not for egg number and body size. The effect of non-independence (due to phylogeny) of our data needs to be more fully controlled in future analyses but methods of incorporating continuous data into cladistic analyses should also be investigated. We show that some predictions can be made about reproductive mode based on body size but ad hoc patterns of reproductive character-state transformation should not be made independent of empirical hypotheses of phylogenetic relationship. Further studies of this kind throughout the Annelida are needed to determine the plesiomorphic reproductive mode for the phylum.     

Does within-population variation in fish egg size reflect maternal influences on optimal values?

Various models that assume correlations between maternal phenotype and offspring environment predict adaptive variation in egg size within populations. Here we conduct a comparative test of these models using published data on fish egg size. Intrapopulation variation in egg size was most pronounced in fish with demersal eggs and larvae (median coefficient of variation [CV] at family level = 6.5%), where offspring environment is likely influenced by maternal phenotype, and least so in fish with pelagic eggs (CV = 3.6%), which experience a relatively stochastic spatial distribution during incubation. This difference was significant at various taxonomic levels, was robust to variation in mean egg size and habitat (i.e., freshwater or marine), and was mirrored in independent paired contrasts. Fish with demersal eggs and pelagic larvae were not significantly different from those with pelagic eggs (CV = 3.8%), indicating that selection favoring within-population variation in egg size occurs mainly posthatching and that any such selection occurring prehatching may be less intense. These results suggest that patterns of within-population variation in egg size among fish taxa reflect adaptive processes and that maternal effects on the egg size-fitness function may explain apparent discrepancies from the single-optima Smith-Fretwell model.     

Osmotic changes during the development of eggs and larvae of the lumpsucker, Cydopterus lumpus L.

The demersal eggs of Cyclopterus appear to osmoregulate like the pelagic eggs of cod and plaice. Unfertilized eggs in ovarian fluid exhibited ovoplasm osmolarities similar to those of adult blood and ovarian fluid (356–359 mosmol kg⁻¹). Yolk osmolarities remained virtually constant from fertilization and during development (356–366 mosmol), with a slight decrease near hatching (to 332 mosmol). Yolk and body fluids of larvae (338 mosmol) had osmoconcentrations similar to egg yolk values near hatching. Yolk osmoconcentration of unfertilized eggs remained unchanged during the first 12 h in sea water, with a slow increase thereafter. Fertilized eggs of bad quality cultures exhibited higher yolk osmoconcentrations than eggs of good quality. Cyclopterus eggs were found to develop normally and survive in 20–34%o salinity, larvae seemed to have the same salinity range.     

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

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Population structure of flounder (Platichthys flesus) in the Baltic Sea: differences among demersal and pelagic spawners

We found significant population structure and isolation by distance among samples of flounder (Platichthys flesus) in the Baltic, Kattegat and Skagerrak seas using microsatellite genetic markers. This pattern was almost entirely due to a difference between flounder that have demersal spawning in the northern Baltic, as compared to pelagic spawners in the southern Baltic and on the west coast of Sweden. Among demersal spawners we found neither genetic differentiation nor any isolation by distance among sampling sites. We speculate that demersal flounder are descendants of a population that colonized the Baltic previous to pelagic spawners. The demersal flounder may thus have had longer time to adapt to the low salinity in the Baltic, and accordingly display egg characteristics that make it possible to reproduce at the low salinity levels in the northern Baltic. Among pelagic spawners significant isolation by distance was detected. Pelagic spawners have previously been shown to display clinal variation in egg size, which allows them to float also at the moderate salinity levels up to the region north of the island Bornholm. Management units for harvesting should ideally be based on true biological populations, and for the commercially important flounder up to 15 different management stocks in the Baltic have been suggested. We could not find a population genetic foundation for such a high number of management units, and our data suggest three management units: the northern Baltic (demersal populations), southern Baltic with the Oresund straits and the most northwestern sampling sites (Skagerrak, Kattegat and North Sea).     

The role of progeny quality and male size in the nesting success of smallmouth bass: integrating field and laboratory studies

Smallmouth bass display size-specific variation in reproductive success with larger brood-guarding males in a population more likely to rear offspring to independence than smaller individuals. The exact mechanisms responsible for this size-specific increase in reproductive output have yet to be identified. To assist in this process, we investigated the relationship between the size of brood-guarding male smallmouth bass and offspring quality (in this case, egg physiology, egg morphology, egg size, hatching success and lab survival). Further, we examined how factors such as egg physiology, egg morphology and egg size influenced reproductive success in the wild and hatching success in a controlled laboratory environment. Nesting male smallmouth bass that successfully reared their offspring to independence spawned earliest in the nesting period were the largest individuals, and guarded eggs with greater concentrations of cortisol compared to males that abandoned their offspring prematurely. Offspring survival in the laboratory was not correlated with offspring survival in the wild, indicating that caution should be used interpreting studies that attempt to relate laboratory-derived survival metrics to the wild. Together, results demonstrate size-specific differences in offspring quality for nesting smallmouth bass, which are correlated with higher concentrations of cortisol in eggs. However, hatching success under laboratory conditions was dissimilar to nesting success in the field relative to cortisol concentrations.     

The interrelation of the size of fish eggs, the date of spawning and the production cycle

The variability in size of pelagic and demersal marine and freshwater fish eggs is examined. The difference between the smallest and largest volumes, based on published figures for the diameters, is large in many species. In marine species with planktonic eggs, the median percentage difference is just over 100%, and this is similar in species with demersal eggs and in freshwater fish.
The available evidence suggests that geographical differences in egg size are small, but in marine fish there is a well-known seasonal decline in egg size. In herring it has previously been shown that egg size in different spawning groups can be correlated with the timing of the production cycle. A similar correlation can be seen in the seasonal shift in time and locality of spawning, and egg size, of the plaice. Sufficient data on seasonal freshwater fish egg variations are not available, but the time of spawning does appear to be linked with the availability of food for the larvae in both lake and stream species.     

Size‐dependent reproductive success of wild zebrafish Danio rerio in the laboratory

Size‐dependent reproductive success of wild zebrafish Danio rerio was studied under controlled conditions in the laboratory to further understand the influence of spawner body size on reproductive output and egg and larval traits. Three different spawner size categories attained by size‐selective harvesting of the F₁‐offspring of wild D. rerio were established and their reproductive performance compared during a 5 day period. As to be expected, large females spawned more frequently and had significantly greater clutch sizes than small females. Contrary to expectations, small females produced larger eggs when measured as egg diameter with similar amounts of yolk compared to eggs spawned by large spawners. Eggs from small fish, however, suffered from higher egg mortality than the eggs of large individuals. Embryos from small‐sized spawners also hatched later than offspring from eggs laid by large females. Larval standard length (L_S)‐at‐hatch did not differ between the size categories, but the offspring of the large fish had significantly larger area‐at‐hatch and greater yolk‐sac volume indicating better condition. Offspring growth rates were generally similar between offspring from all size categories, but they were significantly higher for offspring spawned by small females in terms of L_S between days 60 and 90 post‐fertilization. Despite temporarily higher growth rates among the small fish offspring, the smaller energy reserves at hatching translated into lower condition later in ontogeny. It appeared that the influence of spawner body size on egg and larval traits was relatively pronounced early in development and seemed to remain in terms of condition, but not in growth, after the onset of exogenous feeding. Further studies are needed to explore the mechanisms behind the differences in offspring quality between large‐ and small‐sized spawners by disentangling size‐dependent maternal and paternal effects on reproductive variables in D. rerio.     

Predicting fluctuations in the size of newly emerged sea-trout fry in a Lake District stream

The objective was to predict interannual fluctuations in the size of sea-trout fry when they emerged from the redd, using models developed from field data for 70 excavated redds (≥three per year), and from experimental data on egg and alevin development at 30 constant temperatures in the laboratory (range 1·5—10·5) C with 100 naturally fertilized eggs at each temperature). Egg weight increased with female length and also with the number of eggs laid in a redd, both relationships being well described by a power function. Early spawners were the largest females laying the largest and most numerous eggs, whilst late spawners were the smallest females laying the smallest and least numerous eggs, with middle spawners being intermediate between these two extremes. Mean values for egg weight and number of eggs per redd were obtained for these three groups. The numbers of early, middle and late spawners for each year of a 30-year study and the mean values from the excavated redds were used to estimate weighted means for the number of eggs per unit area and egg weight. Mean values varied considerably between years (30-year ranges: 518–7964 eggs per 60 m2; 112–138 mg wet weight). In the laboratory, mean weights of newly hatched alevins and newly emerged fry were both related positively to mean egg weights. Alevin and fry mean weights were independent of the number of days required for 50% of the eggs to hatch or fry to emerge. Models described in a previous paper formed the basis of those used to predict fry weights over the emergence period. Model predictions were validated by field data for the whole emergence period in 8 years (1967–1971, 1974, 1975, 1980), and by pre-fry weights on single dates in 21 years (1967–1987). As pre-fry densities on these single dates were very similar to egg densities for the same year class, mortality in the egg and alevin stages was very low. The chief objective was therefore fulfilled, and the extent of interannual fluctuations for the 30-year study showed some variation in mean fry weight (30-year ranges: 153–193 mg for both the whole emergence period and the date on which 50% of fry emerged) but a progressive decrease in fry weight through the emergence period. Possible reasons for this variation are discussed, and it is concluded that the size of the female spawners is the dominant factor.     

An individual-based model for predicting the emergence period of sea trout fry in a Lake District stream

The objective of this study was to predict interannual fluctuations in the emergence period of sea trout fry, using models developed from field data for 70 excavated redds, and laboratory data on egg and alevin development at 30 constant temperatures (range 1·5–10·5° C with 100 naturally fertilized eggs at each temperature). Egg weight and numbers per redd both increased with female length; a power function described the relationship. Early spawners were the largest females laying the largest and most numerous eggs, whilst late spawners were the smallest females laying the smallest and least numerous eggs; middle spawners being intermediate between these two extremes. Mean values for egg weight and numbers of eggs per redd were obtained for these three groups. Hatching and emergence times in the laboratory decreased with increasing temperature. Of five models tested for hatching time, the best fit was provided by a three-parameter hyperbolic model which formed the asis of the individual-based model used to predict egg hatching and fry emergence. Model development was described in detail and the final equations predicted the times taken for 5, 50 and 95% of the fry to emerge, and hence the period over which 90% of the fry emerged. Analogous models were obtained for egg hatching. All models were excellent fits to the laboratory data. Hatching times for eggs kept in perforated boxes in the stream were almost identical to those kept at similar mean temperatures in the laboratory. Model predictions of fry emergence times were validated by field data for 8 years (1967–1971, 1974, 1975, 1980). The chief objective was therefore fulfilled, and predictions for the 30-year study (1967–1996) revealed a large variation in the timing of emergence (extremes: 11 March–4 April 1989, 15–20 May 1979). Most of the variation in median emergence date was due to variations in water temperature, with spawning dates as a secondary factor; the latter, however, had a greater effect on the length of the emergence period.     

Life histories of closely related amphidromous and non‐migratory fish species: a trade‐off between egg size and fecundity

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Fitness consequences of egg-size variation in the lesser snow goose

We investigated the relationship between eggsize variation and (a) egg hatching success, (b) chick survival to fledging and recruitment, and (c) adult female survival, over 12 years in the lesser snow goose (Anser caerulescens caerulescens). By comparing the means and variances of egg size for successful and unsuccessful eggs, our aim was to assess the relative fitness of eggs of different sizes and to determine the type of selection operating on egg size in this species. As both egg size and reproductive success vary with age in the lesser snow goose we controlled for the effects of female age. Egg-size variation is very marked in this population, varying by up to 52% for eggs hatching successfully. However, there was no relationship between egg size and post-hatching survival of goslings to fledging or recruitment, either within or between broods, pooling across years. Egg size varied significantly between successful and unsuccessful clutches in only 2 of 33 individual year comparisons. First-laid eggs surviving to onset of incubation, and eggs hatching successfully, were on average larger than unsuccessful eggs, but this was probably due to the confounding effects of female age-specific and sequence-specific egg survival. Variance of egg size differed significantly between successful and unsuccessful eggs in only 3 of 24, and 0 of 21, individual year comparisons for pre- and post-hatching survival respectively. We therefore found little evidence for a relationship between egg-size variation and offspring fitness, or for strong directional, normalising or diversifying selection operating on egg size, in the lesser snow goose. In addition, there was only weak support for the hypothesis that egg-size variation is maintained by temporal variation in selection pressure (sensu Ankney and Bisset 1973). It is likely that egg-size variation represents the pleiotropic expression of alleles affecting more general physiological or metabolic processes. While this does not rule out the existence of alleles with more direct effects on egg size we suggest that their contribution to heritable egg size is small.     

Relationship between clutch size,egg volume and hatching success in a Yellow-legged Gull Larus michahellis colony in south-eastern Tunisia

This study aimed to test the hypothesis that clutch size covaries with egg volume and hatching success in the Yellow-legged Gull Larus michahellis. We determined clutch size and egg volume in a sample of 131 nests, and we used the data to check whether egg volume varied among nests according to clutch size, while taking into account the effects of egg laying order. We also estimated hatching success rate and investigated the relationship between hatching success and clutch size. Egg volume varied among clutches according to clutch size, with eggs being larger in three-egg clutches than in two-egg clutches. Moreover, three-egg clutches showed higher daily survival rates, and hence hatching success, than two-egg clutches. Overall, our results suggest that in the Yellow-legged Gull clutch size covaries with egg volume and hatching success, which could possibly reflect an age effect through different mechanisms. Indeed, older females could be hypothesised to exhibit greater breeding performance than younger females because of their higher experience in tapping energy resources for egg formation and defending nests from dangers. Moreover, due to their age, older females are likely to have lower residual reproductive potential and should invest more heavily in current breeding attempts.     

Effects of electromagnetic fields on the reproductive success of American kestrels

Reduced reproductive success of birds nesting near power lines has been documented but never directly attributed to electromagnetic fields (EMFs). Laboratory studies have identified EMF effects on embryonic development, but reproductive success of wild birds is dependent on additional factors, including fertility, egg size, hatching, and fledging success. We tested whether EMFs affect reproductive success of birds. Captive American kestrels (Falco sparverius) were bred for one season per year for 2 yr under either controlled or EMF conditions. EMF exposure was equivalent to that experienced by wild reproducing kestrels and was weakly associated with reduced egg laying in 1 yr only. In both years fertility was higher, but hatching success was lower in EMF pairs than control pairs. Fledging success was higher in EMF pairs than control pairs in 1995 only. Egg composition and embryonic development were examined in 1 yr only, but hatchlings were measured in both years. EMF eggs were larger, with more yolk, albumen, and water, but had thinner egg shells than control eggs. Late-term EMF embryos were larger and longer than control embryos, although hatchlings were similar in body mass and size. EMF exposure affected reproductive success of kestrels, increasing fertility, egg size, embryonic development, and fledging success but reducing hatching success.     

Lifetime egg production and egg mortality in the damselflyPyrrhosoma nymphula (Sulzer) (Zygoptera: Coenagrionidae)

Artificial oviposition sites were used to estimate egg deposition rates in the field. Females laid an average of 10.76 eggs/minute with a mean duration of 22.81 minutes, giving an average clutch size of 245 eggs. Since one mating corresponded to one clutch of eggs, lifetime mating success was used as a measure of the number of clutches produced. Mean lifetime clutch production was 5.91 clutches per female, equating to 1447 eggs per female per lifetime. Eggs were hatched in the laboratory at temperatures comparable with those in the field. Hatching was highly synchronised and the overall hatching success was 75.1%. Causes of egg mortality in the laboratory were limited to infertility and unhatchability. Since no other sources of egg mortality could be found at the study site, this value was a good reflection of hatching success in the field. Lifetime egg production and hatching success were used to estimate the number of viable offspring produced per female, giving a higher order estimate of reproductive success than has previously been published for a zygopteran.     

The life-history strategy of deepwater sculpin, Myoxocephalus thompsoni (Girard), in Lake Michigan: dispersal and settlement patterns during the first year of life

Deepwater sculpin, Myoxocephalus thompsoni (Girard), were sampled from six stations from the 15–100 m depth contours in Lake Michigan between April 1983 and July 1984. In south-eastern Lake Michigan M. thompsoni lay benthic eggs in offshore waters, which hatch between November and August, with peak hatching in March. Abundance of larvae in pelagic samples was higher offshore than inshore, but larval size was greater and development more advanced at inshore stations, indicating an inshore movement after hatching. Larvae reached metamorphosis at 20 mm and settled to the bottom beginning in July. Pelagic larvae 20–40 mm were found in the lower water column at all stations, but newly settled individuals were only captured with bottom trawls at inshore locations (≤60 m depth). Data from ichthyoplankton and bottom trawl samples in 1983 and 1984 indicated that locations for successful settlement of larvae to the bottom extended only as deep as the shallowest fringe of the adult population (> 50 m in 1983). In 1983, maximum density of larvae reached 0.4 individuals m⁻³ by June. Survival from the pelagic larval stage to the demersal young-of-year stage in 1983/1984 was c . 0.1–0.4%. The specific mechanism of mortality at the time of transition to a demersal habit has not been determined.