Within-clutch egg size asymmetry covaries with embryo sex in the yellow-legged gull Larus michahellis

Non-random sex allocation may occur whenever the expected reproductive value of sons and daughters differs, as is the case when the sexes differ in susceptibility to environmental conditions or maternal effects (e.g. egg size and hatch order). Under such circumstances, covariation between egg and clutch characteristics and egg sex may be expected, and this covariation should vary with maternal state or ecological conditions. In this 2-year study (2007–2008), we examined sex allocation in relation to egg and clutch traits in the yellow-legged gull Larus michahellis, a species where male chicks are larger and more susceptible to harsh rearing conditions than female ones. In 2008, eggs were more likely male early in the season in two- but not three-egg clutches, and large eggs were more likely males late in the season. No egg/clutch traits predicted egg sex in 2007. Within-clutch egg mass asymmetry (the difference in egg mass between the first- and last-laid eggs) predicted sex in both years. In 2007, clutches with smaller egg mass variation were more likely to contain males, while in 2008 this relationship held for the last-laid egg and was reversed for the preceding egg(s). Laying order and sex of the previous egg did not predict egg sex, providing no evidence of sex-specific oocyte clustering. Thus, the relationships between egg sex and egg/clutch traits differed among years, suggesting a phenotypically plastic response of females to extrinsic conditions, and involved within-clutch egg mass asymmetry, a trait likely reflecting variation in maternal quality and/or reproductive tactics, which has been largely neglected in previous studies of sex allocation.     

Fecundity of the Barents Sea capelin (Mallotus villosus)

The fecundity and oocyte diameter of 70 female capelin (Mallotus villosus) from the Barents Sea were investigated. The average fecundity (±SE) was 11 542 (±261). Oocyte diameter and fecundity increased with increasing weight and length of the capelin. Interannual variation in capelin fecundity is large, and corresponds to differences in body weight as revealed by a comparison with previous studies. Received: 10 May 1997 / Accepted: 1 August 1997     

Size at maturity,fecundity and reproductive potential of a protected population of the spiny lobster<Emphasis Type="Italic"> Palinurus elephas</Emphasis> (Fabricius, 1787) from the western Mediterranean

Protected lobster populations are expected to contribute to the replenishment of fished populations through increased egg production. We studied the reproductive biology and egg production potential of a population of the spiny lobster Palinurus elephas protected from fishing since 1990 in the Columbretes Islands Marine Reserve (western Mediterranean). An index of spawning potential was derived to compare egg production potential in the Reserve and in western Mediterranean exploited populations. Females' physiological maturity (ability to reproduce) and functional maturity (ability to mate and bear eggs) occurred at a carapace length (CL) of 76–77 mm. Males' physiological maturity occurred at a slightly larger size, 82.5 mm CL. In the Reserve, P. elephas' individual fecundity increases linearly with body size up to the females' maximum size, although maximum reproductive yield (eggs per body gram) was reached at intermediate sizes. Size-specific fecundity in the protected population was similar to that of lightly fished populations off Ireland and greater than that of western Mediterranean exploited populations. The female size class of 105–110 mm CL contributed most to egg production in the protected population and is well above the minimum landing size (MLS) for western Mediterranean fisheries. Newly mature females (below MLS) generate a very small fraction (1%) of the egg production from the Reserve. Given the pattern of exploitation in western Mediterranean fisheries, egg production potential depends more on the quantity than on the mean size and fecundity of the available females. The role of the greater availability of large males for mating in unfished populations is discussed in terms of the females' individual fecundity and mating success.     

Safe betting: males help dull females only when they raise high-quality offspring

In evolutionary biology, whether parents should enhance or reduce parental care according to mate ornamentation is a subject of great debate. However, the evolution of female ornaments can shed light on this question. In theory, female ornamentation should be traded off against fecundity and thus cannot be wholly informative to males without a direct indication of fecundity. Hence, direct cues of offspring quality should affect the relationship between male investment and female ornamentation. Under this hypothesis, we manipulated two direct cues of offspring quality (egg size and color) after first egg laying in the blue-footed booby and registered male incubation patterns. In this species, foot color is a dynamic signal of current condition and in females is traded off with egg size. We found that males spent more time incubating when paired with dull females but only in nests with large eggs. Males also spent less time incubating small dull eggs. Results indicate that egg size, a direct cue of reproductive value, affected the relationship between male effort and female ornamentation. Males may be willing to help females that have invested in offspring at the expense of ornamentation, which suggests compensation when females are in low condition. Another possibility is that males relax their effort when paired with highly ornamented and fecund females because they have high parenting abilities. Our findings suggest that the information conveyed by female ornaments may depend on direct cues of fecundity. Results also highlight that parental decisions are complex, modulated by a combination of information sources.     

Distribution of marine predator hotspots explained by persistent areas of prey

Investigations of distributional and density patterns of marine predators often reveal areas where high abundances of one or many species overlap in space and time (‘biological hotspots’); however, mechanisms underlying hotspot formation are often unclear, leading to difficulties determining spatial and temporal boundaries of protected areas. On the northeast Newfoundland coast, I previously described annually persistent aggregations of a key forage fish species, capelin (Mallotus villosus): (1) two pre- and post-spawning staging areas in deep (>150 m) bathymetric channels, (2) a cluster of four persistently used demersal spawning sites (17–40 m), and (3) a coastal migratory route (<50 m). Through at-sea surveys repeated over 8 years (2000–2003, 2007, and 2009–2011), I show that the majority of predator hotspots identified were spatially associated with (i.e., within 10 km) these persistent capelin areas for breeding seabirds (common murre: 85.2 ± 4.6 %; northern gannet: 66.9 ± 6.6 %), overwintering seabirds (great and sooty shearwaters: 88.0 ± 6.9 %), and baleen whales (humpback, minke, and fin whales: 86.8 ± 8.6 %). Most predator hotspots were closer to the spawning (3.8–14.0 km) relative to the staging areas (13.1–27.6 km), especially for murres and shearwaters. Interspecific differences were attributed to variation in maximum dive depths and dietary preferences. Predators only aggregated within the spawning area, while capelin were spawning, suggesting that interannual variation in association with predator and capelin hotspots was attributed to variation in survey timing relative to capelin spawning. As these areas of persistent capelin are bound by static bathymetric and large-scale oceanographic features and can be delimited in time based on the capelin spawning period, they may be important candidate areas for protection.     

Effect of diet on reproductive characteristics of Ophryotrocha labronica (Polychaeta: Dorvilleidae)

Effects of food quality on fecundity (defined as the number of eggs deposited in the entire life of a female), egg size and egg energy content (measured by differential scanning calorimeter were studied in two populations of Ophryotrocha labronica (La Greca and Bacci): one taken from the natural environment immediately prior to the experiment, and the other from the same original environment, but bred for more than 5␣years in the laboratory. Results indicated that fecundity is mediated both by food quality and population origin. Specimens fed Tetramin generally spawned more frequently than those receiving the other two diets, while individuals bred in the laboratory since 1990 spawned more frequently than the wild population. Also the number of eggs per spawning and the total fecundity vary with the quality of the diet; Tetramin gives the highest fecundity values. The differences in egg size and energy content seem essentially related to the population origin, in fact, with all diets laboratory specimens produced larger eggs with higher energy contents than wild specimens. Received: 1 August 1997 / Accepted: 29 April 1998     

Male lagoon gobies, <Emphasis Type="Italic">Knipowitschia panizzae</Emphasis>, prefer more ornamented to larger females

Female ornamentation may be directly sexually selected, by male choice or female competition, or occurs as the result of a genetic correlation, arising from sexual selection on males. However, increasing evidence supports the former hypothesis, suggesting that males actively choose their partner preferring traits indicative of female quality. In the lagoon goby, Knipowitschia panizzae, a polygynous species whose males perform parental care to eggs, body length and the size of a sex-specific yellow patch on the belly are known to be reliable indicators of female fecundity. In this paper, we tested, using dummies, the male’s mating preferences for female body and yellow belly patch sizes. The two experimental trials in which a single female trait was variable showed that males prefer a larger belly patch and a larger body size, indicating that both these characters are selected by male mate choice. However, when faced with dummies exhibiting an inverse combination of body and belly patch sizes (experiment 3), males significantly preferred the smaller ones with larger yellow belly patches. A calculation of dummy theoretical fecundity reveals that in the first two experiments, males would have received an immediate benefit from their choice in terms of egg number, whereas in the third one, males chose partners that would have provided them with fewer eggs. The male lagoon goby preference for females with larger belly patches, regardless of their size, suggests that this trait, in addition to indicating fecundity, conveys information about other aspects of female and/or egg quality.     

Effects of size and nuptial gifts on butterfly reproduction: can females compensate for a smaller size through male-derived nutrients?

In butterflies and other insects, fecundity generally increases with female adult weight. Hence, most butterflies are essentially "capital breeders", because nutrients acquired during the larval stage are stored and subsequently used for egg production during the adult stage. However, in some species, males transfer a large nutritious ejaculate to the female at mating. These females can partly be characterized as "income breeders", and female mass can potentially be decoupled from fecundity to some extent. In the gift-giving green-veined white butterfly Pieris napi, it has been shown that female fecundity and longevity increase with number of matings and also that females mature at smaller size under poor food conditions compared to males. So it has been suggested that females can compensate for their smaller size through nuptial feeding. Here we test this hypothesis in P. napi by assessing female fecundity and longevity in relation to female mass and polyandry. The results showed no support for the hypothesis. Smaller females were not capable of increasing their mating rate to compensate for a low weight at eclosion. Instead, larger females remated sooner. Also, smaller females suffered from both a reduced daily and total fecundity compared to larger females and this decrease in fecundity was independent of female mating status, i.e. females allowed to mate only once and multiply mated females suffered to the same extent from their smaller size.     

Brood size and chemical composition of Pareuchaeta norvegica (Crustacea: Copepoda) in Loch Etive,Scotland

Brood size and chemical composition (carbon, nitrogen, and ash) of Pareuchaeta norvegica in Loch Etive, Scotland have been determined. From 23 to 37 eggs were found in each egg mass (mean=30 eggs). The eggs are oval; the mean size of the longer axis is 438 m. Variation in egg mass (as dry weight) is related to increase in dry weight of the maternal body. an increase of 1 mg in maternal body dry weight produces an increase of 0.03 mg dry weight in the egg mass. Carbon content increases with increasing body dry weight both in pre-spawning and spent females, but nitrogen content decreases with increasing body weight. Thus, the carbon:nitrogen ratio increases with increasing body dry weight. The carbon content of an egg mass ranges from 61.4 to 65.3%, and nitrogen content from 9.6 to 13.2% of the total dry weight. Some examples of partitioning of chemical components of the maternal body into the egg mass are also described.     

Geographic variation in maternal investment: acidity affects egg size and fecundity in Rana arvalis

Environmental-stress-mediated geographic variation in reproductive parameters has been little studied in natural vertebrate populations outside the context of climatic variation. Based on life-history theory, an increase in the degree of environmental stress experienced by a population should lead to (1) a shift in reproductive allocation from fecundity to offspring quality, (2) stronger trade-offs between reproductive parameters, and (3) changes in the relationship between female phenotype and maternal investment. To test these predictions, we investigated geographic variation in maternal investment of moor frogs (Rana arvalis) in relation to breeding site acidity (pH 4-8). We found that mean egg size increased and clutch size and total reproductive output (TRO) decreased with increasing acidity among 19 Swedish moor frog populations. Tests for variation and co-variation in maternal investment and female size and age in 233 females from a subset of four acid origin (AO) and four neutral origin (NO) populations revealed that clutch size and TRO increased with female size in both acid and neutral environments. However, in AO populations, egg size also increased with female size, and clutch size and TRO with female age, whereas in NO populations, egg size increased with female age. The strength of the egg-size-clutch-size tradeoff tended to be stronger in AO than in NO females as expected if the former experience stronger environmental constraints. All in all, these results suggest that environmental acidification selects for investment in larger eggs at a cost to fecundity, imposes negative effects on reproductive output, and alters the relationship between female phenotype and maternal investment.     

Sperm allocation in relation to male traits, female size, and copulation behaviour in freshwater crayfish species

Sperm competition is a well-recognised agent in the evolution of sperm and ejaculate structure, as well as variation in female quality. Models of the evolution of ejaculate expenditure predict that male body condition, female fecundity and the risk and intensity of sperm competition may be the ultimate factors shaping optimal ejaculate size. We investigated sperm allocation in Austropotamobius italicus, a freshwater crayfish exhibiting a coercive mating system and external fertilisation, in relation to male and female traits and copulation behaviour under laboratory conditions. We found that mating males were sensitive to female size and produced larger ejaculates when mating with larger females, which were more fecund in terms of number of eggs produced. We found no evidence for female egg production being sperm-limited, as the number of eggs was not dependent on male sperm expenditure. Copulation duration and number of ejaculations reliably predicted the amount of sperm transferred, and both these behavioural measures positively covaried with female body size. These results indicate that male freshwater crayfish can modulate their sperm expenditure in accordance with cues that indicate female fecundity. In addition, a novel finding that emerged from this study is the decrease in sperm expenditure with male body size, which may either suggest that large, old male crayfish are better able than small males to economise sperm at a given mating to perform multiple matings during a reproductive season, or that they experience senescence of their reproductive performance.     

Energy content and fecundity of capelin (<Emphasis Type="Italic">Mallotus villosus</Emphasis>) along a 1,500-km latitudinal gradient

Capelin is a planktivorous key fish species in many subarctic ecosystems representing the link between lower trophic levels and apex predators. Little is known, however, of capelin in Greenland waters, but it has been suggested that size and growth increases due to difference in food availability and temperature conditions along a 1,500 km south–north gradient on the west coast. It is presently unknown how the qualitative state of capelin energy content is affected along this gradient. Based on 2007 and 2008 samples, we show that energy content increases with capelin length in both spawning and non-spawning fish and that it varies with latitude in spawning fish along West Greenland (60–71°N). Combining our results on energy content with information on capelin growth along the same latitudinal climate gradient demonstrates that less and lower-quality food is available to capelin predators in the south than in the north.     

Geographic covariation of chemical quality of the host alga<Emphasis Type="Italic"> Fucus vesiculosus</Emphasis> with fitness of the herbivorous isopod<Emphasis Type="Italic"> Idotea baltica</Emphasis>

Environmental and/or genetic among-site variation in plant quality may influence growth and fecundity of specialized herbivores inhabiting a particular site. Such variation is important as it generates spatial variation in selection for traits related to plant–herbivore interaction. Littoral macroalgae are known to respond plastically to environmental variation by modifying their chemistry or morphology. We studied geographic variation in phlorotannin, nitrogen, protein, and sugar (fucose, mannitol, and melibiose) concentrations of the brown alga Fucus vesiculosus at 12 sites separated by 0.5 to 40 km in the naturally fragmented Archipelago Sea in the northern Baltic Sea. By this regional variation in algal chemistry we attempted to explain among-population variation in size and fecundity of the crustacean herbivore Idotea baltica. We observed high spatial variation in all the measured chemical characteristics of F. vesiculosus, as well as in female size and the number of eggs produced by the herbivores. Spatial variation in nitrogen or protein contents of the alga did not explain the variation of herbivore traits. However, egg size positively covaried with spatial variation in the concentration of mannitol, the major storage carbohydrate of the alga. Such a positive relationship may arise if I. baltica can utilize the nutritive value of a mannitol-rich diet thereby being better able to provision the developing eggs with energy-rich metabolites. Unexpectedly, the concentration of phlorotannins, secondary metabolites having a putative role in defense against herbivory, positively covaried with the size of the herbivore. Among-population variation in host plant chemistry and covariation of that with herbivore growth and reproduction imply that herbivores respond to the local quality of their host plants, and that geographical structuring of populations has to be taken into account in studies of plant–herbivore interactions.Communicated by M. Kühl, Helsingør     

Multivariate trade-offs, succession, and phenological differentiation in a guild of colonial invertebrates

For competing species limited by one or few resources, diversity is thought to be maintained by trade-offs that allow niche differentiation without resource partitioning. However, few studies have quantified multiple key traits for each species in a guild and shown that trade-offs among these traits apply across the guild. Here we document strong bivariate and multivariate relationships among growth rate, fecundity, longevity, and overgrowth ability for six co-occurring colonial invertebrates. We find that all four of these traits are constrained to a single "fast-slow" niche axis that mechanistically relates life history variation to a colonization-competition trade-off. The location of species on this axis strongly predicts the timing of their peak abundance during succession. We also find that species closer to each other on the fast-slow axis are more likely to differ in reproductive phenology, suggesting a secondary dimension of niche differentiation for otherwise similar species.     

Maternal allocation of lipid classes and fatty acids with seasonal egg production in Atlantic cod (Gadus morhua) of wild origin

A suite of characteristics is often used to assess egg quality as these properties potentially play important roles in progeny survival and growth. Our objective was to assess egg characteristics including lipid biocomposition of an iteroparous, batch-spawning teleost of wild origin. Maternal allocation to egg number was generally dome-shaped (5 of 8 females) and egg size declined over the breeding season for eight breeding pairs of wild Atlantic cod (Gadus morhua) (n = 43 batches). Egg lipid composition ranged considerably among females and between egg batches within females (e.g., phospholipids 40–86 %; polar 47–87 % and neutral lipids 15–52 % of total lipids; polyunsaturated fatty acids 16–50 % of total fatty acids). Principal component analyses revealed significant inter-relationships among maternal traits, batch sequence and fecundity, and egg size and composition. Seasonal trends with regard to lipid deposition were variable; three females showed consistent declines in lipid parameters (μg egg⁻¹) with both batch number and egg diameter, one female showed consistent increase and the four remaining females showed no trend. The three females that exhibited seasonal declines in egg lipid content were characterized as having high fertilization success (>75 %). Our findings highlight the variability in lipid allocation to eggs of batch spawners of wild origin and characterize the composition of endogenous reserves available during embryogenesis and yolk sac larval stages.     

Strong maternal effects and extreme heterogeneity of progeny development in the caridean shrimp Sclerocrangon boreas (Crangonidae)

Sclerocrangon boreas is uncommon among marine coastal carideans in having a non-dispersing, abbreviated (2-stage) larval phase. We investigated the implications of this strategy in terms of fecundity, offspring provisioning and brood care in S. boreas from the St. Lawrence Gulf and Estuary in 2009–2010. Fecundity scaled positively to female body size but was low due to the production of large, lipid-rich eggs. Offspring size at all stages of development was positively related to female size. Larval traits and lipid dynamics indicate obligatory lecithotrophic development from hatching to juvenile. The larva becomes a juvenile on the mother and remains associated with her for sometime after. The co-occurrence of early egg stages among many juveniles in some clutches raises the possibility that maternal care of juveniles includes the provisioning of trophic eggs or eggs reclaimed from other females.     

Relationship between egg features and maternal body size in the simultaneous hermaphrodite <Emphasis Type="Italic">Oxynoe olivacea</Emphasis> (Mollusca,Opisthobranchia, Sacoglossa)

This paper provides information on spawn morphology and egg features of the stenophagous planktotrophic Mediterranean sacoglossan Oxynoe olivacea. Smith and Fretwell’s hypothesis, predicting that individuals of the same population growing in the same environmental conditions and varying in size should spawn eggs of a constant size, was tested in a population of O. olivacea living in the Straits of Messina. To determine whether (a) spawn mass size, (b) total egg number per spawn, and (c) egg size were related to parent size of O. olivacea, 21 egg masses (seven egg masses deposited by seven different 20 mm animals, seven egg masses deposited by seven different 25 mm animals and seven egg masses by seven different 30 mm animals) were randomly chosen and examined. Results showed that both spawn mass width and number of eggs per spawn mass increased across O. olivacea body size and apart from the significant variation of the short capsule diameter, there was no consistent variation of egg features in O. olivacea. In conclusion the species allocates constant amounts of energy to individual embryos and thus supports the prediction designed by Smith and Fretwell.     

Reproductive Investment of a Lacertid Lizard in Fragmented Habitat

Abstract:  We studied the effect of habitat fragmentation on female reproductive investment in a widespread lacertid lizard (  Psammodromus algirus ) in a mixed-forest archipelago of deciduous and evergreen oak woods in northern Spain. We captured gravid females in fragments (≤10 ha) and forests (≥ 200 ha) and brought them to the laboratory, where they laid their eggs. We incubated the eggs and released the first cohort of juveniles into the wild to monitor their survival. Females from fragments produced a smaller clutch mass and laid fewer eggs (relative to mean egg mass) than females of similar body size from forests. Lizards did not trade larger clutches for larger offspring, however, because females from fragments did not lay larger eggs (relative to their number) than females from forests. Among the first cohort of juveniles, larger egg mass and body size increased the probability of recapture the next year. Thus, fragmentation decreased the relative fecundity of lizards without increasing the quality of their offspring. Reduced energy availability, increased predation risk, and demographic stochasticity could decrease the fitness of lizards in fragmented habitats, which could contribute to the regional scarcity of this species in agricultural areas sprinkled with small patches of otherwise suitable forest. Our results show that predictable reduction of reproductive output with decreasing size of habitat patches can be added to the already known processes that cause inverse density dependence at low population numbers.     

Yolk androgens in great tit eggs are related to male attractiveness,breeding density and territory quality

Females can adaptively adjust phenotype of their offspring via deposition of various compounds into eggs, including androgens and other hormones. Here, I investigated how egg yolk androgens (testosterone and androstenedione) related to environmental conditions and parental traits in the great tit (Parus major) across three breeding seasons. Male and female traits studied included age, condition and multiple feather ornaments, both carotenoid- and melanin-based (carotenoid and UV chroma of yellow breast feathers, area of black breast band and white cheek immaculateness). Yolk mass increased with laying temperature, laying date and area of male black breast band. Concentration of androgens increased with breeding density, territory quality and carotenoid chroma of male yellow breast feathers and was higher in mates of 1 year old as compared to older males. Yolk androgens were not related to any of the female traits analysed. These patterns were thus consistent with (1) social and environmental effects on yolk mass and composition and (2) both positive and negative differential allocation strategies of resource allocation in females. Overall, male traits were the most important predictors of egg yolk characteristics in this socially monogamous songbird.     

Concordance between short-term experiments and long-term censuses in tide pool fishes

Sixteen years of annual censuses (1990-2005) of an assemblage of tide pool sculpins at Tatoosh Island, Washington, USA, revealed relative constancy in the numbers of recruits and adults. However, the dominant species as a recruit (Clinocottus globiceps) was always replaced numerically as an adult by another species (Oligocottus maculosus). When mechanisms of coexistence were evaluated, little support was found for either a lottery model, because competitive interactions are hierarchical, or a storage-effect model of coexistence, because the relative ranking of recruitment varied little among years and did not covary among species with environmental variables such as ocean temperature, upwelling indices, or an estimator of ocean productivity (oyster condition index). There was also little evidence of niche partitioning based on habitat affinities. Additionally, predation-mediated coexistence had little support, given that the competitive dominant did not have the greatest rates of mortality. Instead, a competition-colonization trade-off may contribute to the coexistence of these species, where C. globiceps always recruits in the greatest numbers, while O. maculosus dominates the adult assemblage via competitive processes. The importance of post-recruitment processes in this assemblage is further suggested by some negative associations among species, the presence of density dependence, some habitat affinities, and previously published experimental work that demonstrated the competitive dominance of O. maculosus over C globiceps.