Developmental morphology of the Indian cyprinid fish Barilius canarensis

Embryonic and larval development of an Indian cyprinid fish, Barilius canarensis, is described from laboratory-reared specimens. The eggs, measuring 2.1–2.4 mm in diameter, were demersal, almost spherical in shape, transparent and unpigmented, with a pale yellow yolk without oil globules. Hatching occurred 39–45 h after fertilization at 26.8°–27.4°C. The newly hatched larvae, measuring 4.8–5.1 mm in body length (BL) with 22 + 17 = 39 myomeres, were characterized by melanophores already deposited on the eyes. The eggs of B. canarensis resembled those of the related danionin species Candidia barbatus, Opsariichthys uncirostris uncirostris, Zacco platypus, Z. sieboldii, and Z. temminckii. Although the larvae of B. canarensis were also similar to those of the foregoing species in general morphology, they differed in having a straight notochord tip and pigmentation on the eyes at hatching and the almost entire absence of melanophores on the ventral body surface from the yolk sac to postflexion larval stages. Conversely, melanophores occurred on the anterior abdominal and pericardial cavities from the preflexion to postflexion larval stages.     

Effects of maternal stress coping style on offspring characteristics in rainbow trout (Oncorhynchus mykiss)

Maternal size, age, and allostatic load influence offspring size, development, and survival. Some of these effects have been attributed to the release of glucocorticoids, and individual variation in these stress hormones is related to a number of traits. Correlated traits are often clustered and used to define the proactive and reactive stress coping styles. Although stress coping styles have been identified in a number of animal groups, little is known about the coupling between stress coping style and offspring characteristics. In the present study, plasma cortisol levels in ovulated mothers and cortisol levels in non-fertilized eggs from two rainbow trout (Oncorhynchus mykiss) strains selected for high (HR) and low (LR) post-stress plasma cortisol levels were compared. Offspring characteristics such as egg size, larval growth, and energy reserves also were compared between the two strains. Maternal plasma and egg cortisol levels were correlated, but no difference between the HR and LR strains was detected in either parameter. LR females produced larger eggs, and larvae with larger yolk sacs compared to HR females, however no differences in larval body size (excluding the yolk) was detected between strains. Considering that the HR and LR strains have a number of correlated behavioral and physiological traits that resemble the reactive and proactive stress coping styles, respectively, the results suggest that proactive mothers invest more energy into their offspring, producing larvae with larger energy reserves. It is possible that larger energy reserves in proactive larvae support the energy requirement for establishing and defending territory in salmonid fish. Furthermore, in the present study we found a positive relationship between mother plasma cortisol and egg cortisol; however neither mother plasma cortisol nor egg cortisol differed between strains. These results indicate that cortisol endowment from the mother to the offspring plays a minor role in the transfer of the behavioral and physiological traits which separates these strains.     

Reproductive biology and morphology of eggs and larvae of Stiphodon percnopterygionus (Gobiidae: Sicydiinae) collected from Okinawa Island

Reproductive biology and morphology of eggs and early larvae of the sicydiine goby Stiphodon percnopterygionus were investigated on Okinawa Island, southern Japan. Spawning season was estimated as being from May to December. Standard length at maturity was approximately 20 mm in both sexes, and batch fecundity was approximately 1000–10 000 per female. The egg masses, guarded by the male, were laid on the undersurface of stones in freshwater. The pyriform eggs had long- and short-axis diameters of 0.54–0.58 mm and 0.49–0.50 mm, respectively. Newly hatched larvae (1.20–1.32 mm notochord length: NL) were poorly developed, with large yolk sacs and unopened mouths. Three days after hatching (1.87–2.05 mm NL), eyes were fully pigmented and mouths were opened. An erratum to this article is available at .     

Developmental morphology of the cyprinid fish Inlecypris auropurpureus

Embryonic, larval, and juvenile development of a Myanmarese cyprinid fish, Inlecypris auropurpureus, is described from laboratory-reared specimens. The eggs, measuring 0.9–1.0 mm in diameter, were demersal, almost spherical in shape, transparent and unpigmented, with a pale yellow yolk without oil globules. Hatching occurred 49–56 h after fertilization at 26.2°–27.3°C. The newly hatched larvae, measuring 2.9–3.1 mm in body length (BL) with 17 + 19–20 = 36–37 myomeres, had melanophores on the head and body. A cement organ on the forehead for adhering to objects during the yolk sac and early preflexion larval stages was distinctive. The yolk was completely absorbed at 3.6–4.0 mm BL. Notochord flexion was initiated at 5.1–5.6 mm BL and finished at 7.1 mm BL. Aggregate numbers of all fin rays were completed at 14 mm BL. Squamation was initiated midlaterally on the anterior trunk at 14 mm BL and completed at 27 mm BL. Although the eggs of I. auropurpureus resembled those of the closely related species Chela dadiburjori, Danio rerio, and Devario malabaricus, they differed from those of Danio rerio and Devario malabaricus in having a narrower perivitelline space. The larvae and juveniles of I. auropurpureus were also similar to those of C. dadiburjori, Danio rerio, and Devario malabaricus in general morphology, but they differed from the latter three species in having a series of dark blotches laterally on the body in the juvenile stage. Moreover, I. auropurpureus differed from C. dadiburjori in having more myomeres and a near-single row of melanophores on the body along the dorsal midline from the yolk-sac to early postflexion larval stages, from Danio rerio in having a cement organ on the forehead during the yolk-sac and early preflexion larvae, and a single melanophore on the lower eye margin in the early yolk-sac larvae, and from Devario malabaricus in having a single melanophore on the lower eye margin in the early yolk-sac larvae. The presence of a cement organ on the forehead indicates a close relationship among the genera Inlecypris, Chela, and Devario.     

Cross‐generational costs of compensatory growth in nine‐spined sticklebacks

Compensatory growth (CG) is a form of phenotypic plasticity allowing individuals’ growth trajectories to rebound after a period of resource limitation, but little is known about the reproductive and cross‐generational costs of CG. We studied the potential costs of CG by exposing female nine‐spined sticklebacks Pungitius pungitius to 1) high (favourable), 2) low (stressful), and 3) recovery (initially stressful, subsequently favourable) feeding treatments, and quantified the effects of these treatments on female reproductive traits (clutch, egg and yolk size), and on the size of their offspring. The low feeding treatment reduced female size relative to the high and recovery feeding treatments, which produced equally large females. Hence, females from the recovery treatment demonstrated CG and full growth compensation. Feeding treatments had significant effects on clutch, yolk, egg and larval size, also when the effect of female size was controlled for. However, these effects came about mostly because females from the low feeding treatment produced small clutches with large eggs (containing little yolk) and larvae, whereas females from the recovery feeding treatment produced as large clutches, eggs (with similar amounts of yolk) and larvae as females from the high feeding treatment. Yet, structural equation modelling revealed that while a direct effect of female size on offspring size was positive in the low and high feeding treatments, it was negative in the recovery feeding treatment, independently of egg and clutch size. This indicates a cross‐generational tradeoff between female and offspring sizes in the recovery feeding treatment. Also the tradeoff between clutch and larval size was more pronounced in recovery than in low or high feeding treatments. Hence, apart from demonstrating that environmental influences experienced by females during their development have the potential to influence their size, fecundity and reproductive traits, the results also provide evidence for complex cross‐generational costs of recovery growth.     

Differentiated egg size of the cannibalistic salamander <Emphasis Type="Italic">Hynobius retardatus</Emphasis>

Larvae of the salamander, Hynobius retardatus, are carnivorous, and even though there are two morphs, a typical morph and a broad-headed or “cannibal” morph, both are cannibalistic. They also sometimes eat other large prey, for example larvae of the frog, Rana pirica. In natural habitats, use of both conspecific and R. pirica larvae as food may contribute more strongly to high survival and substantially to fitness when larval densities are higher, because early-stage H. retardatus larvae sometimes experience scarcity of their typical prey. In cannibalistic oviparous amphibians, larger individuals that developed from larger eggs can more efficiently catch and consume larger prey and thus their survival may be better than that of smaller individuals developed from smaller eggs. Populations might therefore diverge in respect of egg size in response to variation in the density of conspecific and R. pirica larvae in natural ponds, with eggs being larger when larval density is higher. I examined how variance in hatchling size correlated with the incidence of cannibalism, and whether increasing larval density in natural ponds correlated with increasing egg size. Variance in initial larval body size facilitated cannibalism, and egg size increased as larval density in the ponds increased. In ponds with high larval density, where cannibalism and large prey consumption is a critical factor in offspring fitness, the production of fewer clutches with larger eggs, and thus of fewer and larger offspring, results in greater maternal fitness. Variation among the mean egg size in populations is likely to represent a shift in optimum egg size across larval density gradients.     

Interfemale variation in egg yolk androgen allocation in the European starling: do high-quality females invest more?

Androgenic hormones occur naturally in bird egg yolk and are known to enhance growth in canary and gull chicks. Gil et al. (1999, Science, 286, 126-128) have recently proposed that female allocation of androgens to eggs represents a form of costly maternal investment. This hypothesis predicts that females of high quality or high reproductive potential should invest more yolk androgens in their eggs than females of low quality or low reproductive potential. We tested these predictions by examining interfemale variation in allocation of the androgenic steroids testosterone and androstenedione to eggs in a wild population of European starlings, Sturnus vulgaris. We collected 30 full clutches of eggs and captured 22 of the females that laid these clutches for phenotypic measurements. In agreement with the hypothesis, we found that there was significant interfemale variation in yolk androgen concentrations. Furthermore, older females deposited more androstenedione and testosterone in eggs than 1-year-old females, and females laying early or large clutches deposited more testosterone in eggs than females laying late or small clutches. However, females in good body condition did not deposit more androgens in eggs than females in poor body condition. Large-yolked eggs had a higher total yolk androgen content than small-yolked eggs, but larger eggs did not have higher concentrations of yolk androgens. The results in general support the investment hypothesis of yolk androgen allocation. However, further data are needed regarding the costs of yolk androgen allocation and the effects of yolk androgens on offspring to substantiate the assumptions of the hypothesis. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Eggs and larvae of <Emphasis Type="Italic">Awaous melanocephalus</Emphasis> (Teleostei: Gobiidae)

The morphology of eggs and larvae of Awaous melanocephalus is described. The eggs measured 0.33–0.35 mm in long-axis diameter and 0.32–0.34 mm in short-axis diameter. Newly hatched larvae (0.90–0.99 mm in notochord length, NL; 0.93–1.04 mm in total length, TL) were poorly developed, lacking a mouth and having a large yolk sac and unpigmented eyes. The mouth opened and the eyes became fully pigmented 3 days after hatching (1.78–2.00 mm NL, 1.88–2.10 mm TL). The yolk sac was completely absorbed 5 days after hatching at a water temperature of 27°–28°C.     

Simultaneous maturation of two discrete oocyte batches in ayu

Ayu Plecoglossus altivelis including amphidromous and landlocked populations, are known for batch spawning. Unusual clutches at ovulation were observed from rearing stocks that were naturally induced to sexual maturation. Two peaks occurred simultaneously in the frequency distribution of egg size, compared to normal fish, which have clutches with unimodal size distribution. The eggs in the small size class could be fertilized, and small-sized larvae hatched from these eggs. To explain the alteration in egg size through divergence from amphidromous to landlocked populations, the role of batch allocation among oocytes is discussed.     

Maternal resource variation across the laying sequence in Canada geese Branta canadensis maxima

Variation in maternal investments to offspring presumably reflects an optimization of resource allocation such that a female's fitness is maximized. In birds, both egg size and yolk constituents are examples of resources that can vary among offspring within a clutch. Egg size and maternally-derived steroid hormone concentrations present in yolk have been characterized for many species that lay small clutches or have altricial young, but little information is available for species that lay moderate to large clutches of precocial young. In this study, we recorded laying position, measured fresh egg mass and determined maternally-derived testosterone and estradiol concentrations present in yolks for whole clutches of free-living Canada geese Branta canadensis maxima to assess variation in maternal resources within clutches. We found that egg size varied non-linearly across the laying sequence such that first laid eggs were small, the largest eggs in the clutch occurred in the second and third positions, and size declined in eggs laid in subsequent positions. Concentration of testosterone in the yolk followed a pattern in which the first and second laid eggs have the highest concentrations within a clutch and declining concentrations in subsequently laid eggs. In contrast, maternally-derived yolk estradiol concentrations (measured in a subset of clutches) did not change across the laying sequence.     

Developmental morphology of the cyprinid fish Tanichthys albonubes

Embryonic, larval, and juvenile development of a small cyprinid species, Tanichthys albonubes, is described from laboratory-reared specimens. The eggs, measuring 1.0–1.2 mm in diameter, were demersal, almost spherical in shape, transparent and unpigmented, with a pale yolk without oil globules. Hatching occurred 45–53 h after fertilization at 25.5°–26.9°C. The newly hatched larvae, measuring 2.2–2.6 mm in body length (BL), had melanophores on the head and body. In particular, a dark vertical streak occurring posterior to the otic capsule and melanophores above the eyes were distinctive. The yolk was completely absorbed at 3.4 mm BL. Notochord flexion was initiated at 5.0 mm BL and finished at 6.0 mm BL. Aggregate numbers of all fin rays were completed at 11 mm BL. Squamation was initiated at 8.4 mm BL and completed at 13 mm BL. Although the eggs of T. albonubes resembled those of other small danionin species, including Aphyocypris chinensis, Chela dadiburjori, Danio rerio, Devario malabaricus, Gobiocypris rarus, Hemigrammocypris rasborella, and Horadandia atukorali, they differed from those of A. chinensis, C. dadiburjori, G. rarus, and Horadandia atukorali in having a wider perivitelline space. The larvae and juveniles of T. albonubes were similar to those of the aforementioned seven species plus Danio albolineatus, Danio kerri, and Devario sp. (cf. D. aequipinnatus) in general morphology. However, the early life stage morphology of T. albonubes differed from them in having a dark vertical streak posterior to the otic capsule and melanophores above the eyes in the yolk sac larval stage, and a dark lateral streak with an unpigmented area just above the former on the body, a dark blotch on the caudal fin, and reddish dorsal, anal, and caudal fins during the postflexion larval and juvenile stages.     

Variations in the sizes of gonads,eggs and larvae of the dace,Leuciscus leuciscus

Synopsis Prespawning female dace were examined in 7 successive years; in 6 years mean egg size (mm³) and egg number were inversely related and hence the ovary weights of equivalent-sized females were constant. Fecundity increased logarithmically with fish length, and an index of reproductive effort (ovary weight ÷ length cubed) also increased. The number of eggs per gram of ovarian tissue decreased with fish length; this was because mean egg size (mm³) increased and not because of a change in the proportion of connective tissue in the ovary. But in 1977, both egg number and mean egg size were low, although very high somatic growth had occurred in the previous, very warm, summer of 1976. Eggs from different-sized female dace were artificially fertilized, and incubated at a constant temperature. Dry weights of larvae, egg dry weights, mean egg size and larval starvation times showed linear correlations with each other and with parental (female) lengths. The progeny from the very smallest parent died several days earlier than those from the other parents. Size-related predation rates may be of more consequence than starvation death in natural populations. The optimum position of dace along the continuum between many small eggs and fewer larger eggs may vary at different levels of reproductive effort.     

Yolk testosterone and estradiol variation relative to clutch size, laying order and hatching asynchrony in Common Grackles

Maternal effects are typically thought to enhance rather than reduce offspring performance, but asynchronous hatching (ASH) in birds typically produces a size hierarchy within a clutch that frequently reduces the growth and survival of nestlings from eggs that hatch later. Given that yolk steroids can significantly affect offspring phenotype and that in many species the levels of yolk steroids have been found to increase with laying order, the maternal transfer of steroids to egg yolk has been proposed as a mechanism for females to offset the deleterious effects of ASH. To test this hypothesis, we determined whether yolk steroids varied with laying order or clutch size in Common Grackles (Quiscalus quiscula). Because ASH varies with clutch size (hatching span averages 48 h in five-egg clutches, 24 h in four-egg clutches) and regularly results in the starvation of later hatched nestlings, we predicted: (1) testosterone and 17?-estradiol levels should increase with laying order in both clutch sizes to mitigate the negative effects of ASH on last-hatched nestlings, and (2) the increase should be greater in five-egg clutches due to more pronounced hatching asynchrony. Using a competitive-binding steroid radioimmunoassay, we found no systematic variation in either testosterone or estradiol levels relative to laying order or clutch size. In the absence of evidence that yolk steroids interact adaptively with ASH, research must look elsewhere for potential benefits that might compensate for the costs these steroids impose on nestlings.     

Patterns and fitness consequences of intraclutch variation in egg provisioning in tropical Australian frogs

Intraclutch variation in offspring size should evolve when offspring encounter unpredictable environmental conditions. This form of bet-hedging should maximise the lifetime reproductive success of individuals that engage it. We documented the numbers of eggs and means and variances of yolk volume in 15 frog species that occur in tropical savanna woodland. We experimentally determined the effects of initial yolk volume on larval growth patterns in four species. Intraclutch variation in yolk volume occurred to some degree in all species surveyed. Some species had very low, others had very high, intraclutch variation in yolk volume, but all species in which some clutches were highly variable also produced clutches with low variability. Species that occur in areas where the offspring environment is likely to be unpredictable had elevated levels of intraclutch variation in egg provisioning. There was no trade-off between egg size and number in any species surveyed. Under benign laboratory conditions, tadpoles from eggs with larger yolk volumes hatched at larger sizes, and these size differences persisted through a substantial proportion of the larval stage. This indicates that intraclutch variation in egg size has major offspring and thus parental fitness consequences, and is therefore a functional selection variable. This study provides evidence in support of models which predict that intraclutch variation in offspring provisioning can evolve in organisms that reproduce in unpredictable habitats.     

Negative effects of elevated testosterone on female fecundity in zebra finches

Although factors influencing androgen deposition in the avian egg and its effects on nestling fitness are recently receiving considerable attention, little is known about the potential costs of high testosterone levels in the females. Our study aimed at determining the effect of injections of testosterone (T) in female zebra finches (Taeniopygia guttata), on clutch size, egg mass, yolk mass, and yolk androgen content. Females were given a single bolus injection of T in a range of doses after laying the first egg. Results show that administration of T negatively affected clutch size; the strength of this effect increased with increasing doses of T. Females injected with the highest testosterone dose showed suppressed oviposition of the third and the fourth eggs. Interestingly, testosterone administration made females produce eggs with relatively large yolks, suggesting that T may mediate the trade-off between number and size of eggs. Testosterone injection resulted in elevated levels of androgen in the eggs, in contrast to control clutches, which showed a decreasing pattern of androgen concentration along the laying sequence. We conclude that high androgen investment in eggs may be limited by physiological requirements of the ovulatory process.     

Maternal androgens in the pied flycatcher: timing of breeding and within-female consistency

Maternal hormones can have substantial phenotypic effects in the progeny of many vertebrates. It has been proposed that mothers adaptively adjust hormone levels experienced by particular young to optimize their reproductive output. In birds, systematic variation in egg hormone levels has been related to different female reproductive strategies. Because in many bird species prospects of the offspring change seasonally and with brood number, strategic adjustment of yolk androgen levels would be expected. To test this idea, we induced pied flycatcher (Ficedula hypoleuca) females to nest twice during the same season by removing their first clutches shortly after clutch completion. We collected eggs of first and replacement clutches to measure yolk concentrations of androstenedione (A4) and testosterone (T) and captured the females that laid these clutches for phenotypic measurements. Although average egg androgen levels were remarkably consistent within females, hormone patterns differed considerably between first and replacement clutches. Eggs of replacement clutches were heavier with larger yolks compared to first clutches, but they contained on average lower levels of androgens. Within clutches, androgen concentration increased over the laying sequence in the first clutch, but decreased or remained more constant over the laying sequence in the replacement clutch. Mean yolk T, but not A4 levels, were negatively associated with laying date for both breeding attempts. Moreover, females in good body condition produced eggs containing lower levels of androgens than females in poor condition. Our results are consistent with the idea that differences in yolk androgen levels may be one mechanism underlying seasonal variation in reproductive success and it is possible that changes in egg androgen patterns may reflect a change in female reproductive strategy. High within-female consistency also highlights the possibility that there may be some underlying genetic variation in yolk androgen levels.     

长江口水域夏季鱼卵和仔稚鱼年间变化

基于2005年、2008年、2009年和2011年8月(夏季)在长江口水域(30°30'—31°45'N,121°15'—123°10'E)4个航次的浮游生物拖网资料,分析了长江口水域鱼卵和仔稚鱼的种类组成、数量分布特征及其年间变化。结果表明:4个航次采集的鱼卵和仔稚鱼鉴定到种的种类有17种,隶属于8目13科,以鲈形目种类最多,11种,其次是鲱形目,5种,其他各目种类均小于5种;种类数存在明显年间差异,2005年种类数最多(鱼卵3种,仔稚鱼8种),其次是2009年和2011年,2008年种类数最少(鱼卵1种,仔稚鱼5种)。优势种年间更替明显,长蛇鲻(Saurida elongata)、虾虎鱼(Gobiidae spp.)和中华小公鱼(Stolephorus chinensis)在2005年是优势种,2008年优势种为鳀鱼(Engraulis japonicus),2009年优势种为鳀鱼、寡鳞飘鱼(Pseudolaubuca engraulis)、虾虎鱼等,2011年虾虎鱼和小公鱼(包括小公鱼属未定种Stolephorus spp.和中华小公鱼Stolephorus chinensis)成为优势种。2005年鱼卵和仔稚鱼数量分布的密集区在嵊泗列岛附近水域,2008年鱼卵和仔稚鱼出现较少,未出现明显的数量密集区;2009年鱼卵数量较少,仔稚鱼数量较多,密集区主要分布在在长江口以外123°E附近水域;2011年鱼卵主要分布在在长江北支口门外附近水域,仔稚鱼在调查区内分布相对均匀。     

The effect of temperature and substrate on the development of the cortisol stress response in the lake sturgeon, <Emphasis Type="Italic">Acipenser fulvescens,</Emphasis> Rafinesque (1817)

Lake sturgeon, Acipenser fulvescens, are considered threatened or endangered throughout most of their North American Range. Current hatchery rearing for re-stocking programs utilise conventional methods with little to no understanding of the relationship between rearing conditions and the development of the hypothalamic-pituitary-interrenal (HPI) stress axis. In the present study we examined the effects of substrate type and temperature on the development of the HPI stress axis in prolarval and larval lake sturgeon. Lake sturgeon raised over either gravel or no substrate did not consistently show an increase in whole body cortisol at the prolarval stage. However, after the onset of exogenous feeding a consistent increase in whole body cortisol following a stress was evident. Lake sturgeon larvae raised in gravel substrate demonstrated a sustained increase in whole body cortisol for at least 240 min post stress whereas whole body cortisol in larvae raised in no substrate returned to baseline within 240 min post stress. Lake sturgeon larvae raised at 9, 12 and 15°C exhibited markedly different cortisol responses with baseline whole body cortisol being, 38.6 ± 3, 5.67 ± 0.41 and 25.38 ± 2.84 ng.g⁻¹ respectively. Furthermore, the chase induced increases in whole body cortisol at the larval stage were significantly different for each temperature treatment. These experiments demonstrate that physical environment has a significant impact on the development of the HPI stress axis in lake sturgeon.     

Early maternal, genetic and environmental components of antioxidant protection, morphology and immunity of yellow-legged gull (Larus michahellis) chicks

Maternal effects mediated by egg quality are important sources of offspring phenotypic variation and can influence the course of evolutionary processes. Mothers allocate to the eggs diverse antioxidants that protect the embryo from oxidative stress. In the yellow-legged gull (Larus michahellis), yolk antioxidant capacity varied markedly among clutches and declined considerably with egg laying date. Analysis of bioptic yolk samples from clutches that were subsequently partially cross-fostered revealed a positive effect of yolk antioxidant capacity on embryonic development and chick growth, but not on immunity and begging behaviour, while controlling for parentage and common environment effects. Chick plasma antioxidant capacity varied according to rearing environment, after statistically partitioning out maternal influences mediated by egg quality. Thus, the results of this study indicate that egg antioxidants are important mediators of maternal effects also in wild bird populations, especially during the critical early post-hatching phase.     

Position of egg within a clutch is linked to size at hatching in a demersal tropical fish

Size variation among propagules is ubiquitous and small initial differences in size can be critical to survival, particularly in taxa where initial survival is variable and strongly size-dependent. Despite this, the sources of size variation among fish at hatching are rarely investigated. This study examined spatial position within egg clutches as a source of size variation at hatching of the benthic spawning fish Amphiprion melanopus. We quantified within-clutch size variation at hatching and found that newly hatched larvae from the periphery (5 mm from edge) of 2-dimensional clutches were smaller in standard length, cranial depth, eye diameter and body area (7%, 8%, 4% and 11%, respectively) than larvae from the interior positions within clutches. To investigate the source of this variation, sizes of embryos at different locations with clutches were measured within 2 h of fertilisation (8 d before hatching). Newly laid embryos from the clutch periphery were smaller in length and volume than embryos from the clutch interior (> 2% and 4-6%, respectively). These eggs from the periphery also had a 33% lower rate of oxygen consumption than did embryos from the clutch interior, throughout development. The relationships between position within a clutch and egg size, oxygen consumption and larval size imply that size variation in larval fish at hatching is partly generated during early embryogenesis, either from maternal endowment or maternal nest design, and was amplified throughout development.