Transfer of a Pathogen from Fish to Amphibians

Abstract: Ecological studies of exotic species focus primarily on how invaders directly affect particular resident species. In contrast, little is known about the indirect effects of introduced species on native communities, including how pathogens may be spread by introduced species. We provide evidence suggesting that introduced fish may serve as a vector for a pathogenic oomycete, Saprolegnia ferax , that has been associated with embryonic mortality of amphibians in the Cascade Mountains of Oregon, U.S.A. In laboratory experiments, mortality induced by S. ferax was greater in western toad (   Bufo boreas ) embryos exposed directly to hatchery-reared rainbow trout ( Oncorhynchus mykiss ) experimentally infected with S. ferax and hatchery-reared trout not experimentally infected than in control embryos. Embryos also developed significant S. ferax infections when raised on soil that was exposed to trout experimentally infected with S. ferax . Furthermore, toad embryos exposed to S. ferax isolated from sites where Saprolegnia outbreaks are common experienced higher mortality than embryos exposed to S. ferax isolated from sites where Saprolegnia outbreaks have not occurred. Given the widespread practice of introducing hatchery-reared fishes, we suggest that fish used in stocking programs could be an important vector for diseases responsible for amphibian losses.     

Interspecific Variation in Susceptibility of Frog Tadpoles to the Pathogenic Fungus Batrachochytrium dendrobatidis

Abstract:  As part of an overall biodiversity crisis many amphibian populations are in decline throughout the world. Numerous causes have been invoked to explain these declines. These include habitat destruction, climate change, increasing levels of ultraviolet radiation, environmental contamination, and the introduction of non-native species and diseases. Several types of pathogens have been implicated in contributing to amphibian population declines: viruses, bacteria, oomycetes, and fungi. One particular fungus, the chytridiomycete Batrachochytrium dendrobatidis may have caused amphibian population declines in several regions. This pathogen causes chytridiomycosis, which is fatal to newly metamorphic and adult amphibians of certain species. We present experimental evidence that larval stages may also be susceptible to exposure to Batrachochytrium . There was, however, differential sensitivity to B. dendrobatidis in larvae we examined. In laboratory experiments, larval western toads (  Bufo boreas ) exposed to B. dendrobatidis experienced increased mortality and behaviors that suggested they were affected by exposure compared with unexposed control tadpoles. Larvae of Cascades frogs (  Rana cascadae ), bullfrogs ( R. catesbeiana ), and Pacific treefrogs ( Hyla regilla ) did not die after exposure to Batrachochytrium and appeared to behave normally. R. cascadae larvae exposed to B. dendrobatidis , however, showed an increase incidence in mouthpart abnormalities, a characteristic effect of chytridiomycosis, compared with unexposed controls. These results show that Batrachochytrium can negatively affect some species of amphibians at the larval stage and not others. The implications of interspecific variation in susceptibility to fungal infection are broad.     

Amphibian embryo and parental defenses and a larval predator reduce egg mortality from water mold

Water molds attack aquatic eggs worldwide and have been associated with major mortality events in some cases, but typically only in association with additional stressors. We combined field observations and laboratory experiments to study egg stage defenses against pathogenic water mold in three temperate amphibians. Spotted salamanders (Ambystoma maculatum) wrap their eggs in a protective jelly layer that prevents mold from reaching the embryos. Wood frog (Rana sylvatica) egg masses have less jelly but are laid while ponds are still cold and mold growth is slow. American toad (Bufo americanus) eggs experience the highest infection levels. They are surrounded by thin jelly and are laid when ponds have warmed and mold grows rapidly. Eggs of all three species hatched early when infected, yielding smaller and less developed hatchlings. This response was strongest in B. americanus. Precocious hatching increased vulnerability of wood frog hatchlings to invertebrate predators. Finally, despite being potential toad hatchling predators, R. sylvatica tadpoles can have a positive effect on B. americanus eggs. They eat water mold off infected toad clutches, increasing their hatching success.     

An alternative mechanism to reduce intracapsular hypoxia in ovicapsules of Fusitriton oregonensis (Gastropoda)

Oxygen supply inside egg masses has been reported as a constraint of embryo development. Many species that enclose their eggs in jelly masses or ovicapsules have strategies to avoid or reduce intracapsular hypoxia. In some amphibian species, a decrease in the wall thickness of the egg capsule over time produces an increase in oxygen conductance of the wall, reducing the problem of intracapsular oxygen limitation. Previous studies of gastropods have reported a decrease in the thickness of the capsules during development. However, there are no studies relating capsule thinning to the oxygen limitation problem in this group. This study links the thinning of egg capsules with oxygen diffusion as a possible mechanism to reduce or avoid hypoxia inside the capsules of the gastropod Fusitriton oregonensis. Capsule thickness, capsule area, oxygen partial pressure inside and outside the capsule, and oxygen consumption of the embryos at early and late developmental stages were measured. The conductance and the diffusion coefficients of the capsule were estimated using these measurements. Results showed that (1) capsule thickness decreased throughout development by about 50%, (2) oxygen consumption of embryos increased from early to late stages, (3) oxygen partial pressure inside the capsule did not change during development, (4) conductance coefficient increased with time, and (5) estimation of diffusion coefficient was lower than amphibian egg jelly, shark capsules, egg fishes, and eggs of giant cuttlefish. The reduction in the thickness of the capsule wall and the associated increase in its conductance during embryonic development may reduce oxygen constraints, especially at late developmental stages.     

Factors Mediating Co‐Occurrence of an Economically Valuable Introduced Fish and Its Native Frog Prey

Habitat characteristics mediate predator–prey coexistence in many ecological systems but are seldom considered in species introductions. When economically important introduced predators are stocked despite known negative impacts on native species, understanding the role of refuges, landscape configurations, and community interactions can inform habitat management plans. We measured these factors in basins with introduced trout (Salmonidae) and the Cascades frog (Rana cascadae) to determine, which are responsible for observed patterns of co‐occurrence of this economically important predator and its native prey. Large, vegetated shallows were strongly correlated to co‐occurrence, and R. cascadae larvae occur in shallower water when fish are present, presumably to escape predation. The number of nearby breeding sites of R. cascadae was also correlated to co‐occurrence, but only when the western toad (Anaxyrus boreas) was present. Because A. boreas larvae are unpalatable to fish and resemble R. cascadae, they may provide protection from trout via Batesian mimicry. Although rescue‐effect dispersal from nearby populations may maintain co‐occurrence, within‐lake factors proved more important for predicting co‐occurrence. Learning which factors allow co‐occurrence between economically important introduced species and their native prey enables managers to make better‐informed stocking decisions. Factores que Median la Co‐Ocurrencia de un Pez Introducido con Valor Económico y su Presa, una Rana Nativa     

The dynamics of brood desertion among communally breeding females in the treehopper, Publilia concava

In species that exhibit extended parental care, females sometimes lay eggs communally in order to redistribute the costs of offspring care. Communal egg-laying often involves redundancy in female effort, such that the number of females contributing to reproduction is more than is needed to provide adequate parental care. As a result, a subset of females will often abandon the communal brood, with the time of departure ranging from immediate abandonment after egg-laying (brood parasitism) to delayed abandonment with prolonged care (cooperative breeding). In this paper I approach the parental care dynamics of female-female broods as a desertion game similar to that of mate desertion in species with bi-parental care. I describe a field study of the insect Publilia concava (Hemiptera: Membracidae), a species that exhibits communal oviposition and a full range of egg guarding. This species exhibits full redundancy in female care, with no difference in survival between singly and doubly guarded broods. I find that double guarding is extremely rare in the population, with most communal broods having only one female guard. While this guard was usually the female that initiated the brood, these same females were more likely to abandon when secondary females arrived and when secondary females exhibited longer guarding durations. Paradoxically, the secondary females usually abandoned the broods they visited, resulting in up to 50% of broods with double abandonment. These unguarded broods suffered a 50% reduction in hatching success, reflecting an important risk for primary females that abandon egg masses to secondary females. Overall, P. concava exhibits desertion dynamics similar to mate desertion in vertebrates and it is likely that the theoretical work in this area will be useful for future work that addresses the allocation of parental care among communal breeders.     

Hypothesis Concerning the Causes of the Disappearance of Boreal Toads from the Mountains of Colorado

Recent reports of world-wide declines and extinctions of amphibian populations have raised questions about the relation of environmental change to the demise of certain amphibian species. Between 1974 and 1982, 11 populations of boreal toads ( Bufo boreas boreas ) in the West Elk Mountains of Colorado totally disappeared. The apparent cause of extinction of these toads was infection with the bacteria Aeromonas hydrophila . In this paper, the presence of disease in declining populations of these toads is used in conjunction with a variety of data from the literature to formulate a working hypothesis for explaining the cause of the decline of this species, and perhaps others:

     

Synergistic effects of ultraviolet radiation and conditions at low tide on egg masses of limpets (<Emphasis Type="Italic">Benhamina obliquata</Emphasis> and <Emphasis Type="Italic">Siphonaria australis</Emphasis>) in New Zealand

The effects of ultraviolet radiation (UVR), desiccation and conditions in tidal pools on embryonic survival were examined for two common pulmonate limpets that lay intertidal benthic egg masses on rocky shores in New Zealand: Benhamina obliquata and Siphonaria australis. Field surveys and manipulative experiments were conducted between December 2006 and September 2007 in the Wellington region of New Zealand (41°17′S, 174°47′E). Egg mass deposition sites in the field were species-specific: B. obliquata deposited eggs primarily in shaded crevices, whereas S. australis predominantly deposited egg masses in the sun and in tidal pools. For both species, however, embryonic mortality was greater in egg masses that had been in full sun compared to shade. For S. australis, there was also high mortality in egg masses in tidal pools or desiccated compared to those that remained submerged in flowing seawater at low tide. In outdoor experiments, embryonic mortality was also always greatest for egg masses exposed to full sun, and lowest for those in shaded treatments. Mortality was also higher if egg masses were in simulated tidal pools, and for S. australis, if desiccated, compared to those submerged in flowing seawater. Periods of particularly sunny conditions with high temperatures also resulted in higher overall mortality. Finally, egg masses of both species that were initially deposited in the shade had greater mortality in response to subsequent UV exposure compared to egg masses initially deposited in full sun. Results from this study suggest that the egg masses of these two species are highly vulnerable to UVR, as well as other intertidal stressors. Embryos of both of these species may be at risk of high mortality particularly during summer when extreme conditions of UV intensity and high temperature coincide with low tide cycles. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of the Interaction between Genetic Diversity and UV-B Radiation on Wood Frog Fitness

Abstract:  Genetic diversity may buffer amphibian populations against environmental vicissitudes. We hypothesized that wood frogs (  Rana sylvatica ) from populations with lower genetic diversity are more susceptible to ultraviolet-B (UV-B) radiation than those from populations with higher diversity. We used RAPD markers to obtain genetic diversity estimates for 12 wood frog populations. We reared larval wood frogs from these populations and exposed experimental groups of eggs and larvae to one of three treatments: unfiltered sunlight, sunlight filtered through a UV-B-blocking filter (Mylar), and sunlight filtered through a UV-B-transmitting filter (acetate). In groups exposed to UV-B, larval mortality and deformity rates increased significantly, but egg mortality did not. We found a significant negative relationship between genetic diversity and egg mortality, larval mortality, and deformity rates. Furthermore, the interaction between UV-B treatment and genetic diversity significantly affected larval mortality. Populations with low genetic diversity experienced higher larval mortality rates when exposed to UV-B than did populations with high genetic diversity. This is the first time an interaction between genetic diversity and an environmental stressor has been documented in amphibians. Differences in genetic diversity among populations, coupled with environmental stressors, may help explain patterns of amphibian decline.     

Possible mutualism between females of the subsocial membracid Polyglypta dispar (Homoptera)

Summary Female Polyglypta dispar membracids facultatively guard egg masses or oviposit into masses guarded by other females. Defending females repel at least some enemies of both eggs and nymphs. Eggs of guarding females may be partially protected from parasitism by the presence of additional eggs laid by other females. Females sometimes desert egg masses or groups of nymphs, and undefended masses are sometimes adopted by other females. Some individuals remain on or near the plant where they grew up, and littermates sometimes oviposit into the same egg mass. Some copulating pairs are littermates, while others are not. Females with larger numbers of mature and nearly mature eggs in their ovaries are less likely to defend egg masses, and females desert smaller groups of nymphs more often.     

Does spawning behavior minimize exposure to environmental stressors for encapsulated gastropod embryos on rocky shores?

Adults of motile intertidal invertebrates are able to seek shelter to avoid environmental stress associated with low tides, but embryos within egg masses are effectively sessile for the duration of their encapsulation. Gastropod egg masses from 34 taxa on two rocky shores in SE Australia (34°37′08″S, 150°92′03″E and 34°35′45″S, 150°53′20″E) were surveyed over 2 years (June 2002–May 2004) to test the hypothesis that eggs are deposited in patterns that minimize exposure to environmental stress. Egg masses were expected to be predominantly deposited in shaded habitats not prone to environmental extremes. It was also anticipated that the deposition of egg masses in habitats exposed to UVR, desiccation, and/or extremes in temperature would occur when exposure to these abiotic factors was minimized. Among the taxa investigated, only four species spawned in full sun (Bembicium nanum, Nerita morio, Siphonaria zelandica and S. denticulata). Summer had the highest UVR index, water temperature, and air temperature as well as the lowest daytime tides. Univariate and multivariate analyses confirmed that egg mass abundance was highest during summer, with no change in egg mass size. This study shows that those species depositing egg masses on the surfaces of rock platforms do not adjust the seasonal timing or macrohabitat location of their spawning to avoid physiologically stressful conditions, particularly UVR. Alternate reasons for the evolution of egg mass deposition behavior in apparently sub-optimal habitats are discussed, and it is almost certainly the complex interplay of a variety of highly species-specific factors that is responsible for the patterns observed.     

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.     

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

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

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.     

Inter-sexual differences in T-cell-mediated immunity of black-headed gull chicks (<Emphasis Type="Italic">Larus ridibundus</Emphasis>) depend on the hatching order

Hatching asynchrony in avian species leads to age and size differences between nestlings within a brood, handicapping last-hatched chicks in the sibling rivalry. Starvation due to this competitive disadvantage has been regarded as the primary cause of an increase in mortality with hatching order. However, for gulls it has also been suggested that disease is the cause of mortality for last-hatched chicks, possibly through reduced immunocompetence and thereby an enhanced susceptibility to infection. In addition, the male-biased mortality reported for several gull species may be related to a higher vulnerability to diseases in males compared to females. To determine the potential influence of the immune system on these mortality patterns, we investigated the T-cell-mediated immunity (CMI) of black-headed gull chicks in relation to hatching order and sex. We found a significant decrease in the CMI with hatching order. This result may be causally related to systematic changes in maternal yolk steroids and carotenoids within the laying sequence. For second-laid eggs, male CMI was significantly lower than female CMI. This is possibly linked to higher plasma levels of testosterone in male embryos which might have an immunosuppressive effect. If so, this effect is masked in eggs of either high (first egg) or low (last egg) quality. Chicks with low CMI showed enhanced mortality rates. Thus the differences in immune response are likely to contribute to the observed mortality patterns. However, hatching order significantly affected mortality independently of CMI, suggesting that competitive disadvantage due to hatching asynchrony is also important.Communicated by M. Webster     

Reproductive Investment of a Lacertid Lizard in Fragmented Habitat

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

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.     

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.     

The ecological significance of lipid/fatty acid synthesis in developing eggs and newly hatched larvae of Pacific cod (Gadus macrocephalus)

The lipid/fatty acid composition of marine fish eggs and larvae is linked with buoyancy regulation, but our understanding of such processes is largely restricted to species with pelagic eggs. In this study, we examined developmental changes in the lipid/fatty acids of eggs and embryos of Pacific cod (Gadus macrocephalus), a species that spawns demersal eggs along coastal shelf edges, but as larvae must make a rapid transition to the upper reaches of the water column. Adult Pacific cod were collected in the Gulf of Alaska during the spawning season and eggs of two females were artificially fertilized with sperm from three males for each female. The eggs were subsequently reared in the laboratory to determine (1) how lipids/fatty acids were catabolized during egg and larval development, and (2) whether lipid/fatty acid catabolism had measurable effects on egg/embryo density. Eggs incubated at 4°C began hatching after 3-weeks and continued to hatch over a 10-day period, during which there was a distinct shift in lipid classes (phospholipids (PL), triacyglycerols (TAG), and sterols (ST)) and essential fatty acids (EFAs: 22:6n-3 (DHA), 20:5n-3 (EPA), and 20:4n-6 (AA)). In the egg stage, total lipid content steadily decreased during the first 60% of development, but just prior to hatch we observed an unexpected 2–3-fold lipid increase (~6–9 μg individual⁻¹) and a significant drop in egg density. The increase in lipids was largely driven by PL, with evidence of long-chained fatty acid synthesis. Late-hatching larvae had progressively decreasing lipid and fatty acid reserves, suggesting a shift from lipogenesis to lipid catabolism with continued larval development. Egg density measures suggest that lipid/fatty acid composition is linked to buoyancy regulation as larvae shift from a demersal to a pelagic existence following hatch. The biochemical pathway by which Pacific cod are apparently able to synthesize EFAs is unknown, therefore representing a remarkable finding meriting further investigation.     

A field test of the intraspecific brood parasitism hypothesis in the golden egg bug (<Emphasis Type="Italic"> Phyllomorpha laciniata</Emphasis>)

In natural populations of golden egg bugs ( Phyllomorpha laciniata), females lay eggs on plants where they develop unattended, or on conspecifics, where they remain firmly glued until the nymphs hatch and start an independent life. Mortality rates among eggs laid on plants are higher than among eggs carried by adults. Because females cannot lay eggs on themselves, in order to improve offspring survival, they have to lay eggs on other individuals. Two hypotheses have been proposed to explain egg carrying: (1) the mating pair intraspecific brood parasitism hypothesis suggests that females dump eggs on copulating pairs, and (2) the paternal care hypothesis suggests that the system is driven mainly by males accepting eggs to improve the survival rates of their own offspring. Our data from the field show that 77% of the eggs are carried by males, because more males than females carry eggs, and because males carry a greater number of eggs. In addition, we show that mating males carry more recently laid eggs than single males. These results support the view that egg carrying is performed predominantly by males and that eggs are laid on males by their current mating partner, probably between repeated copulations. Males are likely to accept eggs, despite intermediate levels of paternity, because they cannot discriminate in favour of their own eggs, because rejected eggs will face 97% mortality rates on plants, and because they do not suffer mating costs when they carry eggs. However, females carry 23% of the eggs, but no differences in egg carrying have been found between mating and single females, suggesting that this is not the result of egg dumping while females are copulating. Egg carrying by females could reflect low levels of intraspecific parasitism, which is likely to reflect the low rate of successful attempts by egg-laying females who try to oviposit on other conspecifics rather indiscriminately, in an effort to improve the survival of their offspring.