Effects of embryonic exposure to conspecific chemical cues on hatching and larval traits in the common frog <Emphasis Type="Italic">(Rana temporaria)</Emphasis>

Summary. Recent studies indicate that amphibian eggs are capable of hatching plasticity in response to chemical cues released by predators feeding on conspecific eggs or larvae. However, information is scarce on the relative importance of predator and conspecific cues in such a process. In particular, no attempt has been made to compare the effects of embryonic exposures to chemical cues indicative of a predation risk for eggs and larvae, although both life stages can co-occur in natural habitats. In this context, common frog embryos (Rana temporaria) were raised until hatching in the presence of crushed conspecific extracts from eggs and tadpoles to assess their respective influences on some hatching and larval traits. While a significant delay in hatching time was observed in embryos exposed to chemical cues from tadpole extract, this life-history shift appeared unaffected by embryonic exposure to egg extract. Hatchlings derived from eggs incubated in the presence of both conspecific extracts showed a significantly greater weight than unexposed controls. However, such an effect was no longer apparent 15, 30 and 50 days after hatching, suggesting that embryonic exposure to chemical cues from damaged conspecific eggs and tadpoles has no influence on larval growth. Lastly, morphological measurements performed on hatchlings and older tadpoles (15, 30 and 50 days old) revealed no significant effect of embryonic treatments on the shape of body and tail.     

Polyandry, multiple mating, and female fitness in a water strider Aquarius paludum

Contrary to Bateman’s principle, polyandry appears to be a common female mating strategy. Several hypotheses have been proposed to explain the evolution of polyandry. It is assumed that females gain either material or genetic benefits from multiple matings, or that they are coerced into mating by males. In water striders, mating is generally assumed to be costly to females, and they are thought to mate for reasons of convenience, adjusting their resistance to mating according to male harassment. Here, we tested the effect of number of matings (with the same male) and number of partners on female fitness in a water strider Aquarius paludum. In the first experiment, we regulated the time females spent with a male and found that females’ egg production increased with multiple matings up to a point. The result supports the existence of an optimal female mating frequency. In the second experiment, we tested how polyandry affects the number of eggs laid and egg hatching success. We conducted three different trials: females mated four times with either a single male, two different males, or with four different males. Females that mated with four different males laid the lowest number of eggs and had the lowest egg hatching success, suggesting that polyandry reduces females’ egg production and egg hatching success in A. paludum. We conclude that A. paludum females probably gain material benefits from mating but no genetic benefits were found in this study.     

Brooding behaviour and reproductive success in two species of free-living simultaneous hermaphrodites

Although polyclads are amongst the most structurally simple of the triploblastic metazoans, they adopt a wide range of reproductive strategies. Parental care behaviour in this group is yet to be quantified for any species. We assessed the significance of brooding behaviour to the reproductive success of two free-living marine flatworms. Echinoplana celerrima and Stylochus pygmaeus were collected from the field and placed in pairs in containers of filtered seawater where they laid batches of eggs. Both parents were then removed from half of the containers and the brooding behaviour and hatching success of eggs were quantified. There were interspecific differences in brooding behaviour. Egg masses were covered by one E. celerrima parent for 12 ± 2% of time, whereas egg masses of S. pygmaeus were covered by one or both parents simultaneously for 85 ± 8% of time. Egg batches were abandoned by both species immediately prior to the onset of hatching (10–12 days). Hatching success was generally high (~90%) and brooding did not enhance the hatching success of eggs. We assessed the significance of parental care to hatching success of E. celerrima egg masses in the presence of three potential egg predators; in the presence of other organisms. E. celerrima devoted less time to brooding; however, hatching success was not affected. The amount of time spent brooding eggs differed greatly between the two polyclad species but was not essential to their reproductive success under benign conditions. Parental care may be of adaptive value under more stressful environmental conditions commonly experienced in estuarine environments such as lowered salinity, increased hypoxia or turbidity. Covering egg batches may play an additional role of advertising sexual status and a willingness to care for eggs.     

New effects of Roundup on amphibians: predators reduce herbicide mortality; herbicides induce antipredator morphology

The use of pesticides is important for growing crops and protecting human health by reducing the prevalence of targeted pest species. However, less attention is given to the potential unintended effects on nontarget species, including taxonomic groups that are of current conservation concern. One issue raised in recent years is the potential for pesticides to become more lethal in the presence of predatory cues, a phenomenon observed thus far only in the laboratory. A second issue is whether pesticides can induce unintended trait changes in nontarget species, particularly trait changes that might mimic adaptive responses to natural environmental stressors. Using outdoor mesocosms, I created simple wetland communities containing leaf litter, algae, zooplankton, and three species of tadpoles (wood frogs [Rana sylvatica or Lithobates sylvaticus], leopard frogs [R. pipiens or L. pipiens], and American toads [Bufo americanus or Anaxyrus americanus]). I exposed the communities to a factorial combination of environmentally relevant herbicide concentrations (0, 1, 2, or 3 mg acid equivalents [a.e.]/L of Roundup Original MAX) crossed with three predator-cue treatments (no predators, adult newts [Notophthalmus viridescens], or larval dragonflies [Anax junius]). Without predator cues, mortality rates from Roundup were consistent with past studies. Combined with cues from the most risky predator (i.e., dragonflies), Roundup became less lethal (in direct contrast to past laboratory studies). This reduction in mortality was likely caused by the herbicide stratifying in the water column and predator cues scaring the tadpoles down to the benthos where herbicide concentrations were lower. Even more striking was the discovery that Roundup induced morphological changes in the tadpoles. In wood frog and leopard frog tadpoles, Roundup induced relatively deeper tails in the same direction and of the same magnitude as the adaptive changes induced by dragonfly cues. To my knowledge, this is the first study to show that a pesticide can induce morphological changes in a vertebrate. Moreover, the data suggest that the herbicide might be activating the tadpoles' developmental pathways used for antipredator responses. Collectively, these discoveries suggest that the world's most widely applied herbicide may have much further-reaching effects on nontarget species than previous considered.     

Food and predators affect egg production in song sparrows

Although the possibility that food and predators may interact in limiting avian populations has long been recognized, there have been few attempts to test this experimentally in the field. We conducted a manipulative food addition experiment on the demography of Song Sparrows (Melospiza melodia) across sites that varied in predator abundance, near Victoria, British Columbia, Canada, over three consecutive breeding seasons. We previously showed that food and predators had interactive effects on annual reproductive success (young fledged per female). Here, we report the effects on egg production. Our results show that food limits the total number of eggs laid over the breeding season ("total egg production") and that interactive food and predator effects, including food effects on nest predation, determine how those eggs are "parceled out" into different nests. Food addition alone significantly affected total egg production, and there was no significant interannual variability in this result. At the same time, both food and predators affected the two determinants of total egg production: "clutch number" (total number of clutches laid) and average clutch size. Both clutch number and size were affected by a food x predator x year interaction. Clutch number was lower at low-predator locations because there was less nest predation and thus less renesting. Food addition also significantly reduced nest predation, but there was significant interannual variation in this effect. This interannual variation was responsible for the food x predator x year interactions because the larger the effect of food on nest predation in a given year, the smaller was the effect of food on clutch number; and the smaller the effect of food on clutch number, the larger was the effect of food on clutch size. Potential predator and year effects on total egg production were thus cancelled out by an inverse relationship between clutch number and clutch size. We suggest that combined food and predator effects on demography could be the norm in both birds and mammals.     

Observations on Octopus joubini: Some aspects of reproductive biology and growth

Observations were made on the brooding behavior of 10 laboratory reared female Octopus joubini, including egg laying, care of the eggs, feeding during brooding, and survival of the female from the onset of egg laying. Data concerning the numbers of eggs laid by each female and hatching (duration, frequency, correlation of duration to water temperature, etc.) are given. The newly hatched octopuses (whose ages were determined within 24 h) were fed Uca spp. to determine their growth rate from hatching to 4 months of age. In addition, some specimens were presented with a variety of foods that were easily accessible in the laboratory or field in an effort to find a food suitable for mass culture.Contribution No. 1830 from the University of Miami, Rosenstiel School of Marine and Atmospheric Science.     

Supplemental food affects egg size but not hatching asynchrony in rollers

Maternal effects can function as a mechanism of transgenerational plasticity by which the environment experienced by parents is translated into the offspring phenotype and fitness. In birds, parents may affect the competitive ability of their offspring, and hence their fitness, by modifying their hatching pattern and/or egg size. However, little is known about how mothers can modify offspring phenotypes and their fitness in response to a sudden change in environmental conditions during egg-laying. Here, we studied the effect of supplemental food during egg-laying on hatching asynchrony and egg size in the Eurasian roller (Coracias garrulus), a species with marked hatching asynchrony. We also explored the effects of maternal investment on offspring fitness. Food supplementation did not affect hatching asynchrony. However, females in food-supplemented nests laid eggs that increased in size with laying order except for an ultimate small egg. Meanwhile, size of eggs laid by females in control nests did not change with laying order. Supplemental food positively affected hatchability of the egg laid just before the last one and negatively affected hatchability of the last laid egg, which seemed to be a side effect of egg size. Consequently, food-supplemented nests produced fewer fledglings and had higher probabilities of suffering brood reduction than control nests. We conclude that egg size in rollers is a plastic trait, sensitive to short-term changes in food conditions. Furthermore, our results show that maternal investment in egg size may potentially affect offspring fitness.     

Incubation start and egg size in relation to body reserves in the common eider

Avian incubation is often initiated before all eggs are laid. In altricial birds this has been proposed to facilitate brood reduction through asynchronous hatching. However, in precocial birds eggs normally hatch synchronously even if incubation has started before all eggs are laid. Patterns of incubation start may be the adaptive trait selected for both in altricial and precocial species. Several hypotheses have been proposed to explain the timing of incubation start in birds. Decreasing egg-size after incubation start may be adaptively related to an early incubation start, either to ensure synchronous hatching or to decrease fitness cost of late hatched eggs. We have measured individual body condition, egg size and start of incubation in common eider Somateria mollissima, a precocial sea-duck which does not feed during the incubation period. Females in poor body condition start to incubate earlier in the laying sequence than those in good body condition. Furthermore females in poor body condition lay smaller final eggs than females in good body condition. The laying of smaller eggs late in the sequence is therefore probably related to energetic or nutritional state. We propose that females in poor body condition start to incubate early to shorten the nest period in order to reduce their mass loss, but at the cost of reduced size and growth of the ducklings from the eggs laid after incubation start. Females in good body condition on the other hand postpone incubation start at the cost of a longer incubation period and a higher mass loss to the benefit of synchronized hatching and a higher survival of ducklings.     

Conservation implications of density-dependent predation by ghost crabs on hatchling sea turtles running the gauntlet to the sea

Protecting eggs from predators is common practice in sea turtle conservation, but routine protection of hatchlings is not. Of 42 loggerhead hatchlings observed emerging from 10 nests on undeveloped Onslow Beach, North Carolina, 24 % were preyed on by ghost crabs. In experimental trials, ghost crabs similarly threatened and captured neonate freshwater sliders, supporting their substitution as proxy for threatened and endangered sea turtle hatchlings in field experiments testing density dependence. Exploiting natural long-shore variation in ghost crab density, we show that a 2.6-fold higher ghost crab density resulted in 5 times more nocturnal threat encounters with sliders and 3.4 times more slider captures. Sliders released in simulated group emergences experienced lower per capita capture risk by ghost crabs than solitary sliders, implying predator dilution. Non-independence of egg and hatchling depredation motivates consideration of merging sea turtle egg and hatchling stages when modeling and managing food web interactions.     

Egg coloration is correlated with female condition in eastern bluebirds (Sialia sialis)

Egg coloration has been hypothesized to reflect female condition. Because of the proposed physiological costs associated with deposition of biliverdin pigments and because of their conspicuousness, eggs with blue-green coloration may reliably convey information about female or brood quality. We tested the hypothesis that expression of blue-green coloration of eastern bluebird (Sialia sialis) eggs positively correlates to female condition. First, we documented the incidence of egg color polymorphism within the population. We observed that 98% of females laid blue-green eggs while less than 2% laid white eggs and less than 1% laid pink eggs. In a subset of clutches, we used full spectrum reflectance spectrometry (300–700 nm) to compare eggshell coloration to measures of female condition. We found that the color of eggs within clutches was more similar than the color of eggs from different clutches, and that the blue-green eggs have spectral peaks that are consistent with the characteristic absorbance spectra of biliverdin pigmentation. Females in better body condition and older females laid more colorful eggs. Moreover, individual females laid more colorful eggs later in the laying sequence. Overall, these data indicate that egg coloration covaries with female condition, suggesting that egg coloration could function as a reliable signal of female quality or that egg coloration may allow females to recognize eggs laid by conspecific brood parasites.     

Physiologische Anpassungen eines marinen insekts. II. Die Eigenschaften von schwimmenden und absinkenden Eigelegen

The gelatinous egg-masses of Clunio marinus (Diptera; Chironomidae) consist of a tube of jelly containing up to 175 eggs. The swimming egg-mass of the Atlantic populations (Race A) and the sinking egg-mass of the Baltic populations (Race B) have a higher density than the sea water, on which the female spawns; salinity is not important to the swimming or sinking of the egg-masses. Density and sinking velocity of gelatinous egg-masses from Atlantic midges are significantly higher than those from Baltic midges if the eggs are laid on water of the same salinity. The density of the eggs and the quantitative relation of eggs to jelly are the same in both races. The jelly of the egg-masses from the Baltic Sea race swells more than that from the Atlantic Sea race (increase of volume by absorption of water = 3-fold and 2.4-fold, respectively). During swelling the jelly alters from a trough-like shape into a tube-like from which closes around the eggs. If the surface tension of the spawning medium is lowered by use of detergents, the A-spawn will sink. A- and B-spawns occupy different positions at an oil-water interface. Interference contrast microscopy revealed that the A-jelly forms a well-defined outer line in relationship to the surrounding aqueous medium; this line is missing in the B-jelly. The morphological fine surface structures, as seen by the scanning electron microscope, do not explain the different properties of the A- and B-spawn. Extracts and hydrolysates of jelly material were separated by thin-layer chromatography. The results indicate that the spawn jelly of C. marinus consists of a polysaccharide-protein-complex, similar to the jellies of other invertebrates. It is rather probable that the chemical structure or the components of the jellies from A- and B-populations are not exactly the same; this would explain the different properties described. The ecological significance of swimming and sinking gelatinous egg-masses and the taxonomical position of both midge populations are discussed.     

Microbial and environmental effects on avian egg viability: do tropical mechanisms act in a temperate environment?

The viability of freshly laid avian eggs declines after several days of exposure to ambient temperatures above physiological zero, and declines occur faster in tropical than temperate ecosystems. Microbial infection during preincubation exposure has recently been shown as a second cause of egg viability decline in the tropics, but whether microbial processes influence the viability of wild bird eggs in temperate ecosystems is unknown. We determined the microbial load on eggshells, the incidence of microbial penetration of egg contents, and changes in the viability of wild bird eggs (Sialia mexicana, Tachycineta bicolor, Tachycineta thalassina) experimentally exposed to temperate-zone ambient conditions in situ in a mediterranean climate in northern California. Initial microbial loads on eggshells were generally low, although they were significantly higher on eggs laid in old boxes than in new boxes. Eggshell microbial loads did not increase with exposure to ambient conditions, were not reduced by twice-daily disinfection with alcohol, and were unaffected by parental incubation. The rate of microbial penetration into egg contents was low and unaffected by the duration of exposure. Nevertheless, egg viability declined very gradually and significantly with exposure duration, and the rate of decline differed among species. In contrast to studies performed in the tropics, we found little evidence that temperature or microbial mechanisms of egg viability decline were important at our temperate-zone site; neither temperatures above physiological zero nor alcohol disinfection was significantly related to hatching success. Delaying the onset of incubation until the penultimate or last egg of a clutch at our study site may maintain hatching synchrony without a large trade-off in egg viability. These results provide insight into the environmental mechanisms that may be responsible for large-scale latitudinal patterns in avian clutch size and hatching asynchrony.     

Food supply during prelaying period modifies the sex-dependent investment in eggs of Eurasian kestrels

The theory of sex allocation suggests that if the reproductive value and the cost of producing/rearing offspring differ between male and female offspring, parents should invest differently in sexes depending on environmental conditions. Female parents could allocate more resources to eggs of one sex to compensate potential sex-dependent constraints later during the nestling period. In this study, we tested the influence of environmental conditions on sexual dimorphism in eggs of Eurasian kestrels (Falco tinnunculus) by experimentally manipulating food availability before laying. We found that an increase in food abundance before laying did not increase egg mass but changed sex-dependent resource distribution in eggs. In food-supplemented pairs, but not in control pairs, egg mass and hatchling mass were similar between males and females. In addition, we found, in the food-supplemented group, that the latest hatched females showed shorter hatching times than in the control group. In control pairs, female eggs, hatchlings and nestlings were heavier than males. In addition, male fledglings in the food-supplemented group gained less mass than those in the control group. As that food abundance was only increased until the onset of laying, female kestrels were expected to invest in eggs taking food abundance before egg formation as a predictor of future conditions during brood rearing. Our study shows that environmental conditions before laying promote a subtle adjustment of the resources invested in both sexes of offspring rather than in other breeding parameters. This adjustment resulted in a shortening of hatching time of the last hatched females that possibly gives them advantages in their competitive capacity with respect to male nest-mates.     

Effects of elevated yolk testosterone levels on survival, growth and immunity of male and female yellow-legged gull chicks

Androgen hormones of maternal origin contained in the eggs of avian species are considered to have positive effects on offspring characteristics and performance. However, negative consequences have also been reported, suggesting that mothers may experience a trade-off between beneficial and detrimental effects of egg androgens to offspring fitness. We studied the effects of elevated yolk testosterone (T) concentration on survival, development and phenotype of male and female yellow-legged gull (Larus michahellis) chicks by injecting egg yolks with physiological doses of the hormone. Elevated yolk T resulted in a male-biased post-hatching sex ratio, T-treated clutches producing a greater proportion of males compared to control ones at day 4 post-hatching, likely resulting from a reduction of female embryonic survival, whereas no effect of hormone treatment on hatching success or short-term chick survival was observed. In addition, T depressed post-hatching body mass in both sexes but had no effects on the intensity of the cell-mediated immune response or skeletal growth. No sex differences in egg characteristics or chick phenotype were detected. Time to hatching was not affected by T, but females originating from first laid eggs hatched earlier than males of the same laying order, independently of hormone treatment. However, the implications of sex differences in hatching times are unclear in the study species. Taken together, our results suggest that female yellow-legged gulls may be constrained in transferring androgens to their eggs by negative consequences on the viability of female offspring and growth of chicks of the two sexes.     

Fertilization and early developmental stages of eggs of Clupea harengus pallasi n. maris-albi under fresh-water conditions

Eggs of the White Sea herring Clupea harengus pallasi Val. are laid in the intertidal zone, where they are liable to come under the influence of fresh water. The present paper deals with the effect of fresh-water influence on the egg of the White Sea herring, under experimental conditions. In fresh water, egg and sperm are in an unactivated state and, after transfer into sea water, are capable of re-activation, even after 212/ h exposure. The treated eggs develop until the hatching of normal and viable larvae. The number of eggs which are able to complete development depends upon the period of freshwater exposure, although the correlation between the length of that period and survival rates is not very close. Apparently, this is due both to the ability of C. harengus pallasi eggs to develop parthenogenetically and different qualities of eggs (which were obtained from different females). The morphological features of White Sea herring eggs, induced by experimental fresh-water influence, are very similar to those typical for herring eggs of the genus Ilisha, obtained from natural brackish waters. However, such changes are not characteristic of the eggs of forms from natural fresh and brackish-waters.     

Factors affecting the allocation of paternal care in waterbugs (Belostoma flumineum Say)

Summary Male giant waterbugs (Belostoma flumineum Say) brood eggs oviposited on their dorsa by conspecific females. Laboratory observations indicate that viable egg pads are occasionally discarded before hatching. Theory predicts that such behavior should occur only if the costs incurred by brooding exceed the benefits of hatching the egg pad. We studied the effects of egg pad size, time invested in brooding, and egg viability upon the continuation of paternal care in the giant waterbug. We found that smaller egg pads are less likely to hatch than larger ones, and males appear to be less likely to discard egg pads as temporal investment increases. However, the inviability of eggs did not appear to affect the probability of an egg pad being discarded. Males of this species appear to have evolved a decision-making process involving the continuation of paternal care. Offprint requests to: K.C. Kruse     

Mating,egg-laying and hatching of Octopus tetricus (Mollusca: Cephalopoda) in the laboratory

Observations on mating, egg-laying and hatching of aquarium-held individuals of Octopus tetricus Gould, 1852 are described. Females were able to store viable spermatozoa for periods up to 114 days, and laid many small eggs in strings 10 to 12 cm long. The duration of embryonic development varied inversely with temperature. The newly hatched planktonic larvae were 2.5 mm long by 1.1 mm wide, but attempts to rear them were unsuccessful, the longest survival being 21 days. The females died following egg hatching.     

Effects of territory quality, food availability and sibling competition on the fledging success of oystercatchers (Haematopus ostralegus)

We investigated the fledging probability of oystercatcher, Haematopus ostralegus, chicks as a function of hatching order, brood size, territory quality and food availability. Sibling dominance was related to the hatching order in both low- (’leapfrogs’) and high-quality (’residents’) territories. Differences in hatchling mass might have aided the establishment of a dominance hierarchy, since breeders produced small late eggs and hatchlings. These mass differences were most pronounced in leapfrogs, and in large broods in years with lower food availability (’poor’ years). Late hatchlings fledged less often and with lower body masses compared to early hatchlings in all situations. Leapfrogs produced smaller broods and hatched their broods more asynchronously in poor years than leapfrogs breeding in years with more available food (’good’ years) and residents breeding in both poor and good years. Large brood sizes resulted in lower survival of hatchlings in poor years. These results favour the ’brood reduction’ hypothesis. However, contrary to the expectations of this hypothesis, hatching order also affected fledging success in residents. Moreover, large brood size resulted in higher survival of hatchlings in good years, particularly in residents. Thus, although large broods experienced losses due to sibling competition in some years, they nevertheless consistently produced more fledglings per brood in all years, both as leapfrogs and residents. We believe this effect is due to parental quality correlating with initial brood size. Most leapfrogs, at best, fledged one chick successfully each year, losing chicks due to starvation. Nevertheless, leapfrog broods were reduced in size after hatching significantly less quickly than resident broods. These results suggest that breeders lay and hatch insurance eggs to compensate for unpredictable losses due to the high predation rates on both nests (ca 50%) and chicks (ca 90%), in accordance with the ’nest failure’ hypothesis. Received: 14 February 2000 / Revised: 27 September 2000 / Accepted: 10 June 2000     

Ecophysiological studies on the developing eggs and embryos of the European lobster Homarus gammarus

During the course of its embryonic development, the European lobster Homarus gammarus Linnaeus exhibits progressive increases in content of water (54.0 to 83.1%), ash (2.7 to 16.7%), protein (47.4 to 50.9%) and non-protein nitrogen (1.0 to 2.4%), and steady decreases in content of fat (43.8 to 25.4%) and energy (6343 to 5431 cal/g dry organic substance). Cumulative yolk utilization efficiency during the total development is 81.8% for dry weight; the corresponding value for energy is 60.1, for protein 75.6 and for fat 47.4%. Energy content of a single egg is 10.49 cal. Of 4.20 cal expended for metabolic processes of the embryo, only 13.3% energy is drawn from protein oxidation; fat oxidation supplies as much as 87.7% energy, that of carbohydrate only 2.3%. Embryonic development results in a remarkable decrease in net yolk utilization efficiency, which falls from 85.5% in the early developmental stages, to less than 70% in later developmental stages. The mean dry weight of a single egg membrane increases from 38 g (2.2% of egg weight) in a freshly laid and attached egg, to 81 g in an egg with an almost completely developed embryo. This result supports the earlier observation of Cheung (1966) that the formation of the inner chitinous egg membrane occurs after the egg is laid and attached to the setum. Protein seems to be the major constituent of the egg membrane (4049 cal/g dry weight), which has the following composition: protein 70.4%, non-protein nitrogen 0.13%, ash 2.83%. Initial permeability of the egg membrane to water (about 6% of the total water requirement is let in) is followed by a period during which the egg membrane is almost impermeable to water (stages I to III); the egg membrane becomes permeable to water again and lets in 85% of the total water requirement (the rest, i.e. 9%, is metabolic water) at a relatively advanced stage of development. These assumed changes in egg membrane permeability appear to be indicative of variations in the egg's osmoconcentration leading to shiftings in net transport of water. Rates of water and salt uptake during embryonic development are essentially parallel (Fig. 1). The egg membrane remains permeable to salts throughout development; salt intake almost doubles after the egg passes through stage III. A single egg, weighing 3.7 mg requires 4.9 mg water for successful completion of embryonic development. The imbibition of water by the developing marine demersal egg seems to (1) serve in osmotic hatching; (2) float the hatched larva by means of specific gravity reduction; (3) aid the larva to quickly adjust its body temperature. The simple osmotic hatching mechanism, proposed by previous workers, seems to be inadequate to account for the events and timing of the hatching process in the lobster. It is suggested that hatching time is determined not solely by increased internal pressure caused by inflow of water and salts, but also by some unknown internal factor. In the lobster egg, as well as in many other marine demersal eggs, protein metabolism is suppressed to a considerable extent, and fat metabolism is geared up Thus, the non-cleidoic lobster eggs exhibit metablic properties which are typical of cleidoic eggs. This finding is discussed in the light of Needham's (1950) concept of cleidoicterrestrial and non cleidoic-aquatic eggs.Dedicated to Professor P. Kochukutta Menon, Montreal, on his 62nd birthday, June 3, 1970.     

Incubation behavior,brood patch formation and obligate brood reduction in Fiordland crested penguins

Summary The first-laid eggs of Eudyptes penguins are much smaller than second eggs, generally hatch later, and experience greater mortality, features that characterize a brood reduction system unlike that of any other avian group. Hypotheses that (1) first-egg mortality is incidental to aggressive behavior by nesting birds and (2) hatching asynchrony is created by preferential egg treatment during incubation were tested in a colony of Fiordland crested penguins (E. pachyrhynchus). Neither hypothesis was supported. Although first eggs experienced greater mortality than second eggs, losses were not associated with aggressive behavior or the degree of nest crowding. First eggs hatched later than second eggs and were found more often in the anterior (previously assumed inferior) nest position. However, that position was not thermally disadvantaged, nor were lower temperatures consistently recorded from the smaller eggs. Parental brood patches did not form until after the first egg was laid; this delay may be a causal factor in the evolution of the unique reversal in eudyptid egg dimorphism.