Energy expenditure for mouthbrooding in a cichlid fish

Summary Energy costs of mouthbrooding were investigated in the East African maternal mouthbrooder Pseudocrenilabrus multicolor by measuring rates of oxygen consumption. Mothers with their brood in the mouth expended 15.7% more energy than mothers without their brood and 13.8% more than starving nonreproductive controls. After subtracting the energy expenditure of the brood, the excess is reduced to 4.7% and 3.2%, respectively, a difference that is not statistically significant. By contrast, feeding nonreproductive females expended more than twice as much energy as the other groups. We conclude that mouthbrooding is a low-cost strategy profiting from investments made during the preceding nonreproductive phase. The implications for the evolution of mouthbrooding are discussed within the context of ecological constraints.     

Parental care and the cost of reproduction in a Mediterranean fish

Summary Symphodus tinca is a common near-shore Mediterranean labrid fish in which females may sometimes spawn their eggs over hundreds of square meters, or alternatively spawn into well-defined algal nests. Eggs spawned in either manner are fertilized, but widely scattered eggs receive no parental care, whereas eggs spawned into nests are usually guarded by the male until they hatch. Here, I report weight changes of individual marked fish that engaged in a variety of different reproductive behaviors during three breeding seasons. Males gained weight at 0.15% per day outside the spawning season, and added 29–78% to their overall body weight between reproductive seasons, even following substantive weight losses in a spawning season (up to 20% among nesting males). Nesting and nest-guarding males lost an average of 0.32% and 0.41% of their body weight per day in 1986 and 1987. This cost is four times greater than reproduction for nonnesting males, which registered a 0.03% daily weight gain. Actively spawning females lost 0.06% of their body weight daily during the spawning season. While long-term growth rates did not appear to be substantially affected by reproduction in either sex or by parental care in males, present work does not exclude the possibility that such long-term effects may exist.     

Laboratory observations on spawning,brood size and egg hatchability of the Antarctic krill Euphausia superba from Prydz Bay,Antarctica

Forty-three female Euphausia superba were collected from three sites in Prydz Bay, Antarctica in January 1985. Most gravid females took 3 to 14 d to spawn after capture and some less mature females released eggs after 21 d. Most females spawned only once; overall mean brood size was 2 809 eggs and mean hatchability for eggs from the premoult females was 69.7%. For seven females spawning a second time, overall mean brood size was 597 eggs. Mean hatchability for second-spawn eggs from five premoult females was 67.1%. Possible origins of the second-spawned eggs were small numbers of matured eggs left in the ovaries after the first spawning or eggs developed from those of peripheral nucleolus and vitellogenic stages. Throughout the 8 wk of observations, there was no apparent development of eggs in females which had released all their eggs at one spawning.     

Filial cannibalism in a nest-guarding fish: females prefer to spawn in nests with few eggs over many

In fish, fecundity correlates with female body size and egg-tending males often eat small broods. Therefore, small females may prefer to spawn in nests that already contain many eggs, to ensure the brood is as large as possible. In contrast, large females may prefer nests with few eggs, if high egg number or density has a negative effect on egg survival, or if there are drawbacks of spawning last in a nest. To test the hypothesis that female body size affects nest (and male mate) choice, using the sand goby (Pomatoschistus minutus), we allowed small and large females to choose between two males that were matched in size — one guarding a small clutch and the other a large clutch, respectively. We recorded where females spawned (measure of female preference), the combined brood size, male courtship, egg care and nest building. We also quantified the effect of brood size and egg density on egg survival in a separate data set. Although the combined broods did not exceed the small brood sizes that are at risk of being eaten, both small and large females preferred to spawn in nests with smaller clutch sizes. This preference could not be explained by more courtship or male parental effort, nor by reduced survival of larger or denser broods. Instead, our result might be explained by females avoiding the danger of cannibalism of young eggs by males or the risk of reduced egg health associated with being near the nest periphery.     

Male-male interference competition decreases spawning rate in the European bitterling (<Emphasis Type="Italic">Rhodeus sericeus</Emphasis>)

We investigated the consequences of male-male interference competition associated with alternative male mating tactics in a freshwater fish, the European bitterling (Rhodeus sericeus). Male bitterling defend territories around living mussels and attract females to lay their eggs in the gill cavities of mussels. We experimentally manipulated spawning-site abundance and male density at two spatial scales. We showed that the total number of eggs spawned by females was constrained by the number of mussels available for oviposition. The effect was mediated by behavioral interactions among competing males because of variation in the Operational Sex Ratio (OSR) in close proximity to a mussel and not by a direct limitation in mussel capacity to accommodate the eggs. Both total and local male densities affected spawning behavior, and interacted in their effect on female spawning rate. Territorial male aggression caused courtship interruptions that prolonged the time until successful spawning and increased with male density. However, territoriality broke down at the highest male density, with a consequent stabilizing effect on spawning rate.Communicated by J. Krause     

Factors affecting buoyancy in the eggs of Bairdiella icistia (Pisces: Sciaenidae)

Several aspects of buoyancy were studied in the normally pelagic eggs of the sciaenid fish Bairdiella icistia (Jordan and Gilbert). Spawning fish acclimated to a low salinity (15‰) produced larger, more buoyant eggs which had a higher water content than eggs produced by fish living in ordinary sea water (33‰ S). Eggs fertilized in lower salinities were larger and more buoyant than eggs fertilized in higher salinities. The salinity of the medium during the first 5 to 7 min after fertilization had a lasting effect on egg buoyancy, but subsequent transfer to a different salinity also influenced buoyancy. Although egg buoyancy in this species can be influenced by the ambient salinity both before and after spawning, the capacity for adjusting buoyancy is limited, and eggs spawned in salinities lower than 30‰ would probably sink.     

Reproduction of the intraovarian brooding apodid Leptosynapta clarki (Echinodermata: Holothuroidea) in British Columbia

Reproduction in the viviparous apodid sea cucumber Leptosynapta clarki was examined in Grappler Inlet, Bamfield, Vancouver Island, Canada from July 1989 through July 1991. The reproductive cycle of L. clarki is annual with gonad growth during the summer months. Male sea cucumbers spawn during November and December and internally fertilize, through as yet undetermined means, the oocytes within the ovary. Embryonic development is rapid, so that within 2 wk fertilized eggs reach the pentactula stage. Pentactulae have an open mouth and anus and actively move in the ovarian tubules feeding on droplets and fluid derived from the resorption of unfertilized eggs. Histological examination of gametogenesis shows similar development to that found in non-brooding holothurians. However, the protandric sex change observed in some male L. clarki after spawning and changes to the overian wall structure prior to fertilization are previously undescribed in sea cucumbers.     

Sequential polyandry affords post-mating sexual selection in the mouths of cichlid females

Females mating with multiple males may obtain direct benefits such as nuptial gifts or paternal care or indirect (i.e. genetic) benefits resulting in higher-quality offspring. While direct benefits are easily identified, it is difficult to determine indirect benefits, and it is hence largely unclear how they are obtained. This is particularly true in species with external fertilisation, where females seem to have little control over fertilisation. In cichlids, most maternal mouthbrooders show sequential multiple mating, where females visit several males for egg deposition. Genetic data revealed that multiple paternity of eggs and young in the mouth of females is common, but behavioural data of female spawning decisions are missing. Here, we test four hypotheses to explain female multiple mating in the maternally mouthbrooding cichlid, Ophthalmotilapia ventralis: (1) fertilisation insurance, (2) genetic bet-hedging, (3) female choice and (4) ‘sperm shopping’ (i.e. induction of sperm competition resulting in sexually selected sperm). Detailed observations of spawning behaviour in the field combined with histological analyses of the male reproductive organs suggest that fertilisation insurance, genetic bet-hedging and pre-mating female choice are unlikely to explain the sequential female multiple mating in O. ventralis. Instead, cryptic female choice by sperm shopping, i.e. post-mating sexual selection, is most compatible with our data and might be the major ultimate cause of multiple mating in females of this species and of mouthbrooding cichlids with maternal care in general. Our study provides new insight into ultimate causes of sequential polyandry in species with external fertilisation, as hitherto post-mating sexual selection by cryptic female choice has been assumed to be incompatible with external fertilisation mechanisms except by components of the ovarian fluid.     

Spawning and development in <Emphasis Type="Italic">Osedax</Emphasis> boneworms (Siboglinidae,Annelida)

We report observations on spawning and early development in bone-eating worms of the genus Osedax. Individual females of Osedax rubiplumus were observed at 1820 m depth freely spawning hundreds of oocytes, and females of an undescribed species, Osedax “orange collar”, were observed spawning in laboratory aquaria. Cytological and molecular analysis of the spawned oocytes of two Osedax species revealed no evidence for the bacterial endosymbionts that the female worms require for their nutrition, suggesting that the bacteria must be acquired later from the environment, as they are in other siboglinids. Individual O. “orange collar” females released an average of 335 (±130) eggs per day, but the number of oocytes spawned per day varied greatly, suggesting that not all the females spawned daily. Fertilization rates of the spawned oocytes varied from 0 to 100%, though most females showed nearly 100% fertilization rates. Oocytes spawned in the laboratory at 4–6°C were negatively buoyant. If fertilized, these oocytes extruded polar bodies and then after at least four hours cleaved unequally. Subsequent cleavages occurred in a spiral pattern at roughly 2-h intervals, resulting in free-swimming trochophore larvae after 24 h. These lecithotrophic trochophores swam for 9–16 days before settling with several hooked chaetae, similar to those of dwarf Osedax males. The larval life span of the Osedax species studied in the laboratory appears to be shorter than in closely related Vestimentifera. Osedax rubiplumus, on the other hand, has much larger oocytes and so may have greater dispersal potential than these other Osedax species. The high fecundity and apparently continuous reproduction of Osedax boneworms permits the opportunistic exploitation of sunken vertebrate bones. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Lipids in atlantic halibut (Hippoglossus hippoglossus) eggs from planktonic samples in Northern Norway

Fertilized Atlantic halibut (Hippoglossus hippoglossus L.) eggs in different developmental stages (Days 0 to 18) were sampled from plankton in North Norway in February 1986 and analysed for lipid classes and fatty acid content. In unfertilized ovulated eggs taken from ripe fish caught in 1983/1984, polar and neutral lipids comprised ca. 71 and 30% of the total lipids, respectively, decreasing and increasing to 67 and 33%, respectively, in Stage III (11 to 18 d old) fertilized eggs. Of the polar lipids, phosphatidylcholine decreased markedly from 62% in unfertilized ovulated eggs to ca. 40% in Stage I (0 to 3 d old) fertilized eggs, while phosphatidylethanolamine increased from ca. 7 to 33%. Triacylglycerols, the major neutral lipids, increased from ca. 13% in unfertilized ovulated eggs to 16% in Stage III fertilized eggs. The total lipid in Stage I fertilized eggs had relatively low levels of polyunsaturated fatty acids (PUFA), with (n-3) PUFA accounting for only ca. 25% of the total fatty acids. The (n-3) PUFA increased to ca. 40% of the total fatty acids in Stage III, while the (n-3): (n-6) ratio increased from 4.1 to 7.0.     

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     

Apparent resource competition among embryos in the brood pouch of a male pipefish

Embryo success was studied in the paternally brooding pipefish Syngnathus typhle. During brooding, which lasts about a month, males provide embryos in their brood pouch with nutrients and oxygen via a placenta-like structure. Egg size depends on female size. In aquaria, males were mated with differently sized females to give the following treatments: M, mixed-egg-size broods of approximately half large and half small eggs; L, single-egg-size broods of large eggs; S, single-egg-size broods of small eggs; and F, field mated males. All males were kept in aquaria for a full brooding period. For each egg-size category, the number of newborn was compared with the number of eggs the male initially fertilized in his brood pouch. Within mixed-egg-size broods, a higher proportion of large eggs survived and large eggs resulted in heavier newborn than small eggs. Indeed, small eggs from a mixed-egg-size brood had significantly lower relative success (proportion of embryos surviving to birth) than those from a brood entirely composed of small eggs. The implication is that embryos compete for resources within the brood pouch, and that competitive success depends on egg size. Given that females produce eggs corresponding in size to their body size, and that females are known to compete indirectly for access to mates (i.e., the sex-roles are reversed), this intrabrood competition could be seen as an extension of female-female competition, but alternative explanations are discussed. Received: 28 April 1995/Accepted after revision: 28 October 1995     

Calanoid copepod eggs in sea-bottom muds. V. Seasonal changes in hatching of subitaneous and diapause eggs of Tortanus forcipatus

A warm-water neritic calanoid copepod, Tortanus forcipatus, has been found to lay two physiologically different types of eggs (subitaneous and diapause eggs), which are separately spawned in response to different seasonal environmental conditions. Subitaneous eggs are produced when the planktonic populations are at their numerical maxima, and are only shed until late summer. After peak spawning, true resting eggs are spawned which undergo diapause for 1 to 3 months. The eggs recover from their diapause around mid-winter, but hatching does not take place until the water temperature at the sea bottom exceeds 15°C in early summer.     

Intraspecific brood parasitism as a conditional reproductive tactic in the treehopper<Emphasis Type="Italic"> Publilia concava</Emphasis>

In species exhibiting egg guarding as well as communal egg laying, females may adopt the strategy of laying eggs in the nests of conspecifics and leaving without providing care (termed intraspecific brood parasitism). This study is the first to describe such a behavior in the insect Publilia concava (Hemiptera: Membracidae) through field studies that followed 849 marked females across 1,828 brood associations. While brood parasitism increased the total number of eggs in a host brood, it did not reduce the overall hatching success of host broods. Solitary females exhibited a range of guarding durations while parasitic females rarely remained to guard eggs. Females exhibiting the parasitic tactic increased their lifetime number of clutches without decreasing the number of solitary clutches that they were able to initiate. Estimates of egg number for these individual broods revealed that females adopting the parasitic tactic (in addition to solitary breeding) had higher lifetime fecundity relative to females that did not parasitize. In females that exhibited both tactics (solitary and parasitic), the parasitic tactic yielded a higher rate of oviposition. The major component of oviposition rate was the time to find hosts and this time decreased with increasing host availability across 222 replicate groups. Females exhibited a shift toward the parasitic tactic when host broods were more abundant (i.e. in larger groups and later in the season). However, the time to find hosts increased as the frequency of the parasitic tactic increased, suggesting that this tactic may be maintained through negative frequency dependence. The results of this study suggest that brood parasitism may be the preferred tactic, as part of a conditional strategy, when hosts are readily available with solitary breeding being the preferred tactic when hosts are in short supply.     

Spawning behaviour and egg capsules of Concholepas concholepas (Mollusca: Gastropoda: Muricidae)

Observations on the spawning behaviour of the commercially important muricid snail Concholepas concholepas are reported. The sequence of events relating to external transportation, molding and hardening, and attachment of the egg capsules to a substratum were timed and divided into 6 major events. The snails studied in the laboratory spawned between April and Septemer. These results and data from the literature reveal an all-year-round spawning activity for the species. The frequency, size and time of spawning were studied over 1 year. Spawning activity always began at night, suggesting that the species retains its spawning rhythm in the laboratory. These was a direct relationship between length of the snail and size of the capsules spawned. No nutritive eggs were found; the number per capsule varied between 668 and 14,250 according to capsule size. Hatching time varied between 69 and 128 days at 13.5° to 14.5°C.     

Costs and benefits of temporary brood desertion in a Neotropical harvestman (Arachnida: Opiliones)

Brood desertion is a life history strategy that allows parents to minimize costs related to parental care and increase their future fecundity. The harvestman Neosadocus maximus is an interesting model organism to study costs and benefits of temporary brood desertion because females abandon their clutches periodically and keep adding eggs to their clutches for some weeks. In this study, we tested if temporary brood desertion (a) imposes a cost to caring females by increasing the risk of egg predation and (b) offers a benefit to caring females by increasing fecundity as a result of increased foraging opportunities. With intensive field observations followed by a model selection approach, we showed that the proportion of consumed eggs was very low during the day and it was not influenced by the frequency of brood desertion. The proportion of consumed eggs was higher at night and it was negatively related to the frequency of brood desertion. However, frequent brood desertion did not result in higher fecundity, measured both as the number of eggs added to the current clutch and the probability of laying a second clutch over the course of the reproductive season. Considering that harvestmen are sensitive to dehydration, brood desertion during the day may attenuate the physiological stress of remaining exposed on the vegetation. Moreover, since brood desertion is higher during the day, when egg predation pressure is lower, caring females could be adjusting their maternal effort to the temporal variation in predation risk, which is regarded as the main cost of brood desertion in ectotherms.     

Gametogenesis and larval development in Sabella spallanzanii (Polychaeta: Sabellidae) from the Mediterranean Sea

Sabella spallanzanii (Gmelin) is one of the best known and widely distributed Mediterranean polychaetes, but available data on its general biology has been inferred mostly from populations recently introduced to Australia. In the present paper, data on gametogenesis and larval development of a Mediterranean population from the Gulf of Taranto (Ionian Sea, Italy) are reported. Histological and electron microscopical analysis of gametogenesis showed that oogenesis, a long process beginning soon after spawning in February, is of extraovarian type. By contrast spermatogenesis is a very fast process lasting from September to December, when the coelomic cavity is completely packed with mature gametes and almost devoid of coelomocytes. During the period of spawning, some experiments of in vitro fertilisation were performed, always without success, while some of the collected females naturally spawned fertilised eggs (100%). Particularly interesting was the discovery of a small percentage of eggs which appeared fertilised but unspawned within the coelomic cavity of female. Investigation of the S. spallanzanii life cycle, from the naturally spawned fertilised eggs to embryonic and juvenile stages, was performed by daily microscopical observations and by SEM. Despite the presence of large eggs giving rise to a lecithotrophic larva, a long pelagic phase was observed, the longest found among Sabellidae. The larval development pattern is also described. Settlement began after about 2 weeks; metamorphosis occurred 10 d after settlement, when mucus was secreted abundantly and an external tube was formed. The long larval pelagic period and the development pattern, suggesting a high potential for dispersal, support both the introduction and invasive behaviour of the species in Australian waters. Received: 28 May 1999 / Accepted: 10 December 1999     

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.     

Cannibalize or care? The role of perceived paternity in the sand goby, <Emphasis Type="Italic">Pomatoschistus minutus</Emphasis>

Parental care is a costly part of reproduction. Hence, natural selection should favor males which avoid caring for unrelated young. However, the decision to abandon or reduce care requires cues which are evaluated to give information on potential reproductive value of the offspring. The prediction that male sand gobies, Pomatoschistus minutus, care for foreign eggs as long as they were spawned in their own nest and at least some of such cues are fulfilled was tested. Egg-guarding males that had recently taken part in a spawning event were given a clutch of eggs that was sired either by themselves or another male, in either their own or another male’s aquarium. Males that had not taken part in a spawning event were used as controls and were given eggs sired by another male. We measured the amount of filial cannibalism and nest building. Control group males did not care for eggs and ate them all before rebuilding the nest. In the other treatments, there were no significant effects of paternity, though males moved to another male’s aquarium increased their clutch area threshold and completely consumed larger clutches than males that were not moved. There was no intermediate response in any treatment in the form of increased partial filial cannibalism or less well-constructed nests. Our results suggest that egg-guarding males cannot distinguish between eggs sired by themselves and those sired by other males but are able to react to cues indicating paternity state. Males do not adopt eggs to attract females in P. minutus.