Reproductive biology of deep-water calanoid copepods from the Arctic Ocean

Reproductive modes and egg production were studied in 15 species of meso- and bathypelagic copepods from nine Calanoida families in the Arctic Ocean. During shipboard incubation, females of seven species released eggs freely into the water and females of three species produced membrane-bound egg sacs. One species, Aetideopsis rostrata, produced a mass of eggs looking like an egg sac, but the “sac” lacked a membrane and disintegrated within 2 h. Females of four additional species were encountered with membrane-bound egg sacs in the preserved samples. In most families, only one reproductive mode, either egg-carrying (the Euchaetidae, Augaptilidae), or broadcast spawning (the Heterorhabdidae, Spinocalanidae, Scolecitrichidae, Tharybidae, Bathypontiidae) was observed. In contrast, different genera of the Aetideidae family demonstrated different reproductive modes, with broadcast spawning predominant in the benthopelagic species, and both broadcast spawning and egg-brooding in the planktonic species. Clutch size and egg diameter varied widely between species in both broadcast spawners and egg-brooders. In broadcast spawners, the clutch size varied from 1 to 95 eggs female⁻¹, while the average egg diameter ranged from 152 to 440 μm. The clutch size for egg brooders varied between 3 and 82, while average egg diameter varied from 258 to 732 μm. Deep-water broadcast spawners produced much larger eggs compared to surface-dwelling broadcast spawning species. This larger egg size may result in a reduction, or elimination, of feeding during naupliar stages, thereby improving the survival potential of deep-water species.     

Observations on the embryonic development of some deep-living decapod crustaceans,with particular reference to species of Acanthephyra

Eggs of meso- and bathypelagic decapod crustaceans were removed from gravid females and allowed to develop in vitro. The time course of embryonic development, as indicated by the appearance of certain morphological and physiological features, was determined for most species at a constant temperature of 12°C. The embryonic period, and the time between the appearance of certain key features such as naupliar and compound eyes and heartbeats, showed little variation in any one species. Significant differences were observed in the development times of different species. In general, species with large eggs have a longer embryonic period than those with small eggs, but there also variations between species with eggs of similar size. During the development of Acanthephyra eggs there is an approximate doubling of the egg volume, correlated with a decrease in egg density and an increase in the water content. The increase in the water content and egg volume is similar to that of other decapod eggs. It is concluded that those species with small eggs could have several broods in 1 year, whereas those species with large eggs are likely to be restricted to one, or in some cases possibly two, broods.     

Mechanisms of avian egg recognition: Which egg parameters elicit responses by rejecter species?

Summary Some species of North American passerines nearly always reject nonmimetic eggs placed in their nests and have apparently evolved this behavior in response to brood parasitism. Experiments presented here examined the specific egg parameters to which rejecter species respond, the relative tolerances rejecters show towards nonmimetic eggs and the degree to which rejection is limited to eggs of the brown-headed cowbird (Molothrus ater), the only parasitic bird widespread in North America.Relative to cowbird eggs, American robin (Turdus migratorius) eggs are larger, blue rather than white and immaculate rather than spotted. Experiments using 10 egg models at 137 nests showed that robins respond to each of these differences (Figs. 2 and 3) but do not usually reject an egg that deviates from their own by only one difference. Eggs that differ in ant two of the three parameters are usually rejected. This built-in tolerance reduces the likelihood that robins will reject their own eggs if these are atypical in size or coloration.Small egg size was the most important parameter eliciting rapid rejections (i.e. within 1 day), probably because differnces in size can be detected by both visual and tactile perception. By contrast, small egg size was the least important parameter determining whether eggs were eventually rejected (i.e. within 5 days, Tables 2 and 3). In terms of their eventual response, robins may be more sensitive to egg coloration than to size because the latter parameter is less reliable in distinguishing between robin and cowbird eggs.Experiments were also carried out at 37 nests of the gray catbird (Dumetella carolinensis), whose immaculate blue-green egg is only slightly larger than a cowbird egg. Catbirds are much more responsive to white ground color than to maculation (Table 4), perhaps because color is more reliable in distinguishing between catbird and cowbird eggs.Rejecter species exhibit degrees of tolerance towards foreign eggs that are proportional to the divergence between their eggs and those of the cowbird. Birds with eggs strongly divergent from cowbird eggs benefit from being relatively tolerant because they avoid rejecting their own eggs but still act against cowbird eggs. Species with cowbird-like eggs must be relatively intolerant to maximize the chances that cowbird eggs are rejected.Experiments show that rejection is not specific to cowbird eggs. Thus, birds have apparently responded evolutionarily to brood parasitism by developing recognition of their own eggs, rather than by developing recognition and rejection specific to parasitic eggs.     

Differences in reproductive strategies of morphs of the genus Echinaster (Echinodermata: Asteroidea) from the Eastern Gulf of Mexico

Two morphs of the asteroid genus Echinaster, E. Type 1 and E. Type 2, have reciprocal annual cycles of reproduction and nutrient storage. The pyloric caeca (storage organs) reach maximal size over winter then decrease with increasing gametogenic activity in the spring. E. Type 1 broadcasts buoyant eggs in late May to early June. E. Type 2 deposits benthic eggs in late April. Lecithotrophic larval development is similar in both morphs except that larvae of E. Type 1 undergo a brief planktonic phase. E. Type 1 reaches sexual maturity at a larger size, expends a lower reproductive effort but has a greater absolute reproductive output than E. Type 2. E. Type 2 produces fewer, larger eggs and has a greater parental investment per egg. In both morphs, females have much higher gonadal lipid levels than males and expend a higher reproductive effort in terms organic matter and energy. Reproductive effort, reproductive output, nutrient storage in the pyloric caeca and body size varied between populations and between years in each morph. Egg size and parental investment per egg were constant. Within-morph variability is attributed to differences in nutritional state. Differences in reproductive strategy support the hypothesis that the morphs are separate species.     

Evolution of egg size and fertilisation efficiency in sea stars: large eggs are not fertilised more readily than small eggs in the genus<Emphasis Type="Italic"> Patiriella</Emphasis> (Echinodermata: Asteroidea)

Fertilisation kinetics theory suggests that, when sperm are limiting, the larger eggs of broadcast-spawning marine organisms ought to be fertilised more frequently than smaller eggs, because they provide a bigger target for searching sperm. Whilst this effect has been demonstrated within species, it is not known if this pattern holds among species. We tested whether a large difference in egg size between congeneric seastars with contrasting planktotrophic and lecithotrophic modes of development results in differences in the likelihood of eggs being fertilised in sperm-limiting situations. Measurement of egg sizes and sperm swimming speeds led to the prediction that the sperm–egg collision rate constant for Patiriella calcar (420-µm-diameter egg) should be nine times greater than for P. regularis (140-µm-diameter egg). Although the eggs of P. calcar should be fertilised at greater rates in low sperm concentrations, they were not. When gametes were allowed to mix for 10 s, the hypothesis that P. calcar eggs required less sperm than P. regularis to ensure 50% of eggs were fertilised was rejected. When gametes were mixed for 5 min, P. regularis eggs were more frequently fertilised, but the difference was not statistically significant. We conclude there must be a difference between these species in the likelihood that when a sperm finds a conspecific egg it can successfully fertilise. This apparent uncoupling of egg size and likelihood of fertilisation suggests that fertilisation is not a major constraint on the evolution of egg size in these seastars.Communicated by G.F. Humphrey, Sydney     

Experimentally manipulated brood sex ratios: growth and survival in the black-headed gull (Larus ridibundus), a sexually dimorphic species

In sexually size dimorphic species, individuals of the larger sex often suffer from enhanced mortality during the nestling period. This has been attributed to higher nutritional requirements of the larger sex, which may render this sex more vulnerable to adverse food conditions. However, sex-biased mortality might not exclusively depend on the differences in food demand but also on other phenotypic differences, e.g., in competitiveness. Interference competition between the sexes and position in the laying sequence in particular may be essential components contributing to biased mortality.By creating synchronously-hatched unisex broods in the sexually size dimorphic black-headed gull, we specifically tested the effect of sex-specific food demand by excluding interference competition between the sexes as well as hatching asynchrony. To test the effect of egg quality, which varies with the position in the laying sequence, we composed each nest of chicks from eggs of all different positions in the laying sequence.All-male nests showed significantly enhanced mortality compared to all-female nests from the beginning of the development of the sexual size dimorphism onwards. This underlines the role of a higher food demand in biased mortality of the larger sex.In males but not females, asymptotic body mass and skeletal size were negatively associated with position in the laying sequence, while survival was not affected by position. As a consequence, sexual size dimorphism at the end of the nestling period was less pronounced compared to the natural situation. These data show that, although male growth is more sensitive to a decrease in egg quality, the higher mortality of last hatched chicks in natural nests is mainly due to hatching asynchrony and egg size but not egg content.     

Time and recruitment costs as currencies in manipulation studies on the costs of reproduction

Life history theory predicts that parents will have lower Darwinian fitness if they tend clutches that are above or below the size they naturally produce. We experimentally tested for relationships between fitness and clutch size in Tree Swallow (Tachycineta bicolor) offspring and parents. Over 130 trios of nests initiated on the same day were randomly divided among reduce (-3 eggs), control (3 eggs picked up and replaced), or add (+3 eggs) manipulations. Pre-manipulation modal clutch size was six eggs (range before manipulations was 1-10; afterwards, it was 1-11). Hatching took longer in larger clutches, but the proportion of eggs hatching and fledging was similar for clutches from 4 to 10, so that clutches of 10 produced the maximum number of fledgling. Parental feeding rates were higher for larger broods, but per capita feeds to nestlings were fewer, and nestlings were smaller. Nonetheless, survival of both young and adults, based on recaptures in subsequent years, was not significantly affected by manipulations. Manipulations also had no significant effect on subsequent reproduction, including the number of fledglings produced by either local recruits or returning breeders. Collectively, our results failed to detect fitness costs associated with tending larger clutches for either parents or the offspring reared and suggested directional selection for larger clutch size. However, because clutches that hatch later produce fewer recruits, the extra days required to lay more eggs and to fledge extra young may eliminate a large part of the advantage that would accrue to parents producing enlarged clutches. For example, our data suggest that there may be less than a 16% benefit to producing nine instead of six eggs, rather than 50%, as is suggested by experimentally manipulated egg numbers alone. Thus, time, rather than costs of reproduction, may be the crucial constraint selecting against Tree Swallows laying larger clutches.     

Resources and offspring provisioning: a test of the Trexler-DeAngelis model for matrotrophy evolution

Theoretical models of the evolution of matrotrophy from a lecithotrophic ancestor suggest that resource availability plays a major role in selective scenarios favoring a change in offspring provisioning. We examined effects of feeding level on embryo provisioning in the livebearing fish Gambusia geiseri, a species with dual provisioning of embryos via both yolk sequestered in large eggs and post-fertilization mother-to-embryo nutrient transfer. Females were fed either once per day or once every three days for three months. Females fed daily had marginally larger brood size, significantly larger embryos, and a higher rate of nutrient transfer (assayed directly by injection of radiolabeled nutrients) than females fed every third day. There was no difference in the size of unfertilized eggs between the feeding treatments. Resource effects on matrotrophic provisioning in G. geiseri suggest that matrotrophy plays an important role in provisioning and allows females to adjust offspring size in response to resource availability.     

Egg size and laying order in relation to offspring sex in the extreme sexually size dimorphic brown songlark,<Emphasis Type="Italic"> Cinclorhamphus cruralis</Emphasis>

In some bird species, mothers can advantage the offspring of one sex either by elevating them in the laying order to promote earlier hatching or by allocating greater resources to eggs of the preferred sex. In size dimorphic species, the predictions as to which sex should benefit most from such pre-laying adjustments are ambiguous. The smaller sex would benefit from an initial size advantage to help compensate for the faster growth rate of the larger sex. However, an early advantage to offspring of the larger sex might have a greater effect on their lifetime reproductive success than an equivalent advantage to offspring of the smaller sex. We investigated these hypotheses in the polygynous brown songlark, Cinclorhamphus cruralis, which is one of the most sexually size dimorphic birds known. We conducted within-clutch comparisons and found that females hatched from larger eggs and were initially heavier (but not structurally larger) than their brothers. This may afford females an early competitive advantage, as egg volume remained correlated with chick mass until at least 5 days of age. Similarly, we found that hatch order was still positively associated with nestling mass and size when the brood was 10 days of age, but there was no clear relationship between offspring sex and hatching order. During this study, food was plentiful and there were few obvious cases of nestling starvation. When food is limited, we suggest that the greater nutrient reserves of female hatchlings could not only help compensate for their slower growth, but could also give them a survival advantage over their brothers early in the nestling period. Consequently, egg size dimorphism may be an adaptation that facilitates an early shift in brood sex-ratio towards cheaper daughters in conditions of low food availability.     

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.     

Factors influencing egg size in the gammarid amphipod Gammarus insensibilis

Developmental and seasonal changes in egg volume were examined in a population of the amphipod Gammarus insensibilis Stock occurring on the south coast of England, towards its northern limit of distribution. Results showed a marked increase in egg volume during development (2.9 times by Egg Stage V), resulting from water uptake and from the conversion of yolk reserves into structural elements. The maximum rate of increase coincides with the period of organ and limb development. At hatching, after initial rupture of the egg membrane by urosome spines, egg volume increases rapidly over a short period (15 to 20 min) by a further 30% (uptake rate 3.6×10^–5 mm³s^–1), followed by a post-hatching decrease in juvenile volume. Increase in size at hatching is the result of drinking by embryos, although changes in body-wall permeability may contribute. Females carrying eggs in an advanced stage of development exhibit egg-collecting behaviour. This is seen as an adaptation to an increased likelihood of egg loss with increase in volume of the brood as hatching approaches. Seasonal changes in Stage I (early) egg size are marked in this species, with winter eggs as much as 60% greater in volume than summer eggs. Egg size is inversely related to the temperature during oocyte development. A simple model has been derived to account for the observed seasonal pattern in egg size. The consequences of seasonal variation in egg and juvenile size are considered.     

Life histories of large and small murexes (Prosobranchia: Muricidae)

One can predict the major features of a muricid's life history from its adult size. Most adults do not grow, and juveniles of all species grow 1 to 2 mm/month, so that larger adults have had fonger juvenile periods. Larger females deposit larger egg capsules and, since each of these contains more eggs, their clutches are larger. Small females deposit several clutches each year, and thus have relatively large annual fecundities. However, large females live longer, so each spawns many more eggs in her life-time than would a smaller female. From 90 to 99% of the juveniles die within their first year. However hatchings of small species are much more likely to complete their first year; a newly hatched Urosalpinx cinerea is 25 times more likely to survive long enough to breed once than a newly hatched Ceratostoma foliatum (Gmelin, 1791).     

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     

Two modes of development in the morphospecies Crepidula dilatata (Gastropoda: Calyptraeidae) from Southern Chile

In Chinquihue Bay, a sheltered locality in Southern Chile, female Crepidula dilatata Lamarck brood egg masses which differ in intracapsular development. In some, all eggs develop and are hatched as free veliger larvae (indirect development). In others, only some of the eggs develop and hatch as young adults (direct development), the rest being consumed as nurse eggs. Two possible interpretations are considered: intrapopulation variation of developmental pattern in a species, and the coexistence of two sibling species. Stages of the intracapsular development are described and illustrated. Fecundity varies according to size of the female. For females brooding eggs with indirect development it is estimated at between 3840 and 85575 embryos per spawn; for females with eggs undergoing direct development, embryo production per egg mass is estimated at between 70 and 812. A comparison is made between females brooding the two kinds of egg masses. Minor differences exist in adult coloration, shape of egg capsules, egg diameters, spawning season and their abundance and distribution in the intertidal. Brooding females with eggs displaying indirect development are significantly larger than those with eggs of direct development. The latter females form chains subtidally but not intertidally as do females bearing eggs with indirect development. On these females with pelagic larvae, chains frequently include sedentary males which reach large sizes. On basal females with direct development, chains contain mostly other females, suggesting that matings are temporary and by errant males. These results are compared with existing information in the literature for other species both in this genus and in other marine invertebrates. It is concluded that two sympatric sibling species with different modes of development are present in the morphospecies C. dilatata Lamarck in Southern Chile.Devoted to Dr. E.F. Kilian, Zoologisches Institut der Universität Gießen, FRG.     

Ecological costs on local adaptation of an insect herbivore imposed by host plants and enemies

Herbivore populations may become adapted to the defenses of their local hosts, but the traits that maximize host exploitation may also carry ecological costs. We investigated the patterns and costs of local adaptation in the pine processionary moth, Thaumetopoea pityocampa, to its host plants, Pinus nigra and P. sylvestris. The two hosts differ in needle toughness, a major feeding impediment for leaf-eating insects. We observed a west-to-east gradient of increasing progeny size in the Italian Alps, matching the pattern in toughness of their respective local host plant. Eastern populations that feed on the native P. nigra with tough needles had larger eggs, and neonate larvae with larger head capsules, than western populations that feed on the native P. sylvestris and the introduced P. nigra with softer foliage. In a reciprocal transfer experiment that involved the eastern-most and the western-most populations of T. pityocampa from this region, and excluded natural enemies, we found evidence for local adaptation to the host plant. Specifically, larvae from the western population only performed well when raised on their local hosts with soft needles, and they suffered near-complete mortality on the tough foliage at the eastern site. In contrast, larvae from the eastern population survived equally well at both sites. Local adaptation involved a trade-off between progeny size and the number of offspring. We hypothesized that an additional cost, imposed by natural enemies, may be associated with increased egg size: we also observed a west-to-east gradient of increased egg parasitism. We tested this hypothesis in a common garden by exposing eggs of both populations to parasitism by two native egg parasitoids, Ooencyrtus pityocampae and Baryscapus servadeii. The eastern population suffered a higher level of parasitoid attack by O. pityocampae than the western population, and performance of hatched adults of both parasitoids was enhanced in large eggs. Thus, increased neonate quality (larger eggs yielding larger larvae) confers an advantage on tough foliage but incurs the ecological cost of increased parasitism, which may constrain further adaptation by this herbivore.     

Fuels for development: evolution of maternal provisioning in asterinid sea stars

For marine invertebrates, larval developmental mode is inseparably linked to the nutritional content of the egg. Within the asterinid family of sea stars there have been multiple, independent, evolutionary transitions to lecithotrophic development from the ancestral, planktotrophic state. To investigate the evolution of maternal investment and development within the Asterinidae, we quantified individual lipid classes and total protein for eggs and larval stages of closely related species representing three developmental modes (planktotrophy, planktonic lecithotrophy and benthic lecithotrophy). Within species, maternal provisioning differed between females indicating that egg quality varied with parentage. Maternal investment was related to egg size but, after correcting for egg volume, we identified two major oogenic modifications associated with the evolution of lecithotrophic development: (1) a reduction in protein deposition that probably reflects the reduced structural requirements of nonfeeding larvae, (2) an increase in deposition of a single class of energetic lipid, triglyceride (TG). The exception was Parvulastra exigua, which has benthic, lecithotrophic development and lays eggs with a lipid to protein ratio close to that of planktotrophs. This oogenic strategy may provide P. exigua larvae with a protein “weight-belt” that assists in maintaining a benthic existence. Asterinids with planktotrophic development used a significant portion of egg TG to build a feeding bipinnaria larva. For Meridiastra mortenseni, female-specific differences in egg TG were still evident at the bipinnaria stage indicating that egg quality has flow-on effects for larval fitness. In lecithotrophic asterinids, TG reserves were not depleted in development to the larval stage whereas protein stores may help fuel early larval development. Available data indicate that there may be two evolutionarily stable egg lipid profiles for free-spawning, temperate echinoderms.     

Eggshell strength of an obligate brood parasite: a test of the puncture resistance hypothesis

Eggs of several brood parasites have thicker and stronger shells than expected for their size. The present study evaluated the puncture resistance hypothesis for the occurrence of thick-shelled eggs in common cuckoos Cuculus canorus by investigating costs of cuckoo egg ejection in four Acrocephalus warblers—the great reed warbler A. arundinaceus, reed warbler A. scirpaceus, marsh warbler A. palustris and sedge warbler A. schoenobaenus. The three latter species all suffered ejection costs, while ejection was not costly in the larger great reed warbler. The occurrence of ejection costs was negatively related to host bill size. In the marsh warbler, we compared ejection costs in naturally parasitized nests and two experimental treatments, in which broods were parasitized artificially with great reed warbler and conspecific eggs. Hosts damaged their own eggs significantly more often when ejecting the thick-shelled cuckoo eggs than when ejecting the similarly sized but thinner-shelled great reed warbler eggs, providing some support for the puncture resistance hypothesis. Ejection of conspecific eggs did not involve any costs. Furthermore, contrary to predictions derived from the laying damage hypothesis, there was no evidence that egg damage was associated with cuckoo egg laying. Hosts damaging their own eggs during ejection were more likely to subsequently desert their clutches than those that did not. The frequency of clutches smeared with the contents of the ejected egg were positively related to the hypothesized difficulty of foreign egg puncturing. Potential advantages of thicker shells in common cuckoo eggs are discussed.     

Reproductive biology of the common shrimp Crangon crangon (Decapoda: Crangonidae) in the central Irish Sea

Aspects of the reproductive biology of the common shrimp Crangon crangon (L.) were studied in Port Erin Bay, Isle of Man, Irish Sea. Size at sexual maturity was determined from the proportions of ovigerous females and of females with maturing ovaries. The size at which 50% of females are mature is estimated (±95% confidence intervals) as 12.5±0.48 mm carapace length. Based on the proportions of ovigerous females and of mature females, the main breeding season was from January to June. Mean ovarian dry weights indicated two broods (winter and summer), with females bearing winter broods (WB) having higher gonad indices than those with summer broods (SB). WB females with non-eyed eggs and with eyed eggs differed in the regression of ovarian dry weight on carapace length, indicating preparation for laying a second brood. In both broods the moult stages of berried females were related to egg stage. Moulting will occur following the release of the brood. During embryonic development, mean egg length and egg volume were larger in all stages, and the mean dry weight of individual eggs of all stages heavier, in WB than in SB; there was no difference in egg number, however. Consequently, reproductive investment, the proportion of female weight devoted to egg production, was 67% higher in WB (0.20±0.04) than in SB (0.12±0.03). There is a significant effect of egg volume on brood weight, but not on egg number. In both broods, egg number was a negatively allometric function of female body size in non-eyed eggs and an isometric function of female body size in eyed eggs. Brood mortality during incubation was higher in SB (17%) than WB (10%). Differences in the reproductive variables and investment between the two broods of C. crangon are discussed in the light of reproductive strategies and life history.Communicated by J.P. Thorpe, Port Erin     

Embryo development and maternal–embryo nutritional relationships of piked spurdog (<Emphasis Type="Italic">Squalus</Emphasis><Emphasis Type="Italic"> megalops</Emphasis>)

The maternal–embryo relationship was determined for the piked spurdog (Squalus megalops). In addition, the increase in offspring size with maternal size was studied and the embryonic development was described. Wet weight of in utero eggs and offspring size was correlated with maternal size; larger females produced larger embryos which would have higher survival rate and reproductive value. All embryos present in a female were at a similar stage of development. The external yolk sac is reabsorbed late in gestation, suggesting that embryos are mostly nourished by yolk sac reserves. Embryo size-at-birth varied considerably (180–244 mm total length) as a result of the significant variability in ova size at ovulation. The amounts of water, organic and inorganic matter of embryos at different stages of development were measured to determine possible maternal contributions during embryonic development. Total wet weight from smallest and largest in utero eggs to smallest and largest term embryos changed by +46 and +58%, respectively. This pattern was due to a change in water content by +137 and +154%, and inorganic matter by +100 and +156%. Organic matter of smallest and largest in utero eggs changed by −23 and −17%, respectively. The uterus of pregnant females became specialised for water and mineral transport, not nutrient provision. These results indicate that S. megalops is a strict yolk-sac viviparous species with no maternal contribution of organic matter during development.     

Tactics of parasitic American coots: host choice and the pattern of egg dispersion among host nests

Summary I examined the tactics adopted by a conspecific brood parasite, the American coot (Fulica americana), and the degree to which these tactics reflect sources of mortality for parasitic eggs. Only 8% of parasitic eggs produced independent offspring, compared to a 35% success rate for non-parasitic eggs, and most mortality was due to egg-rejection by hosts or the consequences of laying eggs too late in the host's nesting cycle. Parasites usually laid parasitically before initiating their own nests and usually parasitized immediate neighbours. Parasites did not remove host eggs before laying their own egg, and egg disappearance in general was not more common at parasitized nests. I found no evidence for non-random host choice, either on the basis of stage of the host's nesting cycle or the host's brood size. The absence of adaptive host choice is likely a consequence of the fact that, due to host limitation, only a small proportion of parasites had meaningful variation among potential hosts to choose from. The pattern of egg dispersion among host nests by individual parasites appears to be a compromise between constraints imposed by host limitation and the increased success obtained from spreading eggs among nests. Most females laying fewer than five parasitic eggs laid them in a single host nest while females laying five or more eggs normally parasitized two or more hosts. An examination of egg rejection and survival rates showed that parasites would maximize success by laying a single egg per host nest, and the pattern of laying several eggs per host nest is likely a consequence of host limitation. However, no egg that was the fifth laid, or later, parasitic egg in a host nest was ever successful and this probably explains why most females laying five or more eggs parasitized more than one host.