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     

The effect of maternal body size on embryo survivorship in the broods of pregnant male pipefish

The occurrence of male pregnancy in the family Syngnathidae (seahorses, pipefishes, and sea dragons) provides an exceptionally fertile system in which to investigate issues related to the evolution of parental care. Here, we take advantage of this unique reproductive system to study the influence of maternal body size on embryo survivorship in the brood pouches of pregnant males of the broad-nosed pipefish, Syngnathus typhle. Males were mated with either two large females, two small females, a large then a small female, or a small then a large female. Our results show that offspring survivorship depends on an interaction between female body size and the number of eggs transferred by the female. Eggs of larger females deposited in large numbers are more likely to result in viable offspring than eggs of smaller females laid in large numbers. However, when females deposited smaller numbers of eggs, the eggs from smaller females were more likely to produce viable offspring compared to those from larger females. We found no evidence that this result was based on mating order, the relative sizes of competing females, or egg characteristics such as dry weight of eggs. Additionally, male body size did not significantly influence the survivorship of offspring during brooding. Our results suggest that the factors underlying offspring survivorship in pipefish may be more complex than previously believed, with multiple factors interacting to determine the fitness of individual offspring within the broods of pregnant males.     

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.     

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.     

Costs and benefits of surplus offspring in the lesser kestrel (Falco naumanni )

Lesser kestrels (Falco naumanni) lay clutches which appear excessive as only 3% of them yield as many young as eggs laid. Four hypotheses may explain the adaptive value of producing surplus eggs: (1) the bet-hedging hypothesis assumes that the environment varies unpredictably and surplus eggs serve to track uncertain resources; (2) the ice-box hypothesis suggests that surplus offspring serve as a reserve food during a period of shortage; (3) the progeny choice hypothesis says that parents produce surplus offspring in order to choose these with higher fitness; and (4) the insurance-egg hypothesis proposes that extra eggs are an insurance against the failure of any egg. To test the significance of this strategy in the lesser kestrel, an experiment manipu-lating brood size at hatching was carried out over 2 years, with good and bad feeding conditions. The experiment consisted of adding a chick to experimental broods where one egg failed to hatch or removing a randomly selected chick from experimental broods where all eggs had hatched. Independently of annual food availability, pairs with brood sizes reduced by one chick fledged more nestlings than pairs with brood size equalling their clutch sizes. Body condition of young was also better in the former group, but only in 1993 (a high-food year). Independently of year, mean local survival of parents with complete broods at hatching was lower than for parents raising reduced broods. These results supported only the insurance-egg hypothesis which says that surplus eggs may be an insurance against the failure of any egg, but parents may suffer reproductive costs when all eggs hatch. Received: 17 January 1997 / Accepted after revision: 27 April 1997     

Studies on the growth,reproduction, and life cycle of the supralittoral isopod Ligia pallasii

A two-year study on a field population of Ligia pallasii Brandt has shown that the isopods live for 1.5 to 2 years. Breeding occurs in the spring and early summer, with some females carrying winter broods of eggs. The mean length of breeding females is 22.5±2.2 mm (standard deviation) and the mean brood size is 48±11 eggs. Mature males are larger than mature females (900 and 300 mg live weight, respectively), and are disproportionately broader (31×17 mm and 22.5×9 mm, respectively). The larger size and breadth of the males is an adaptation for copulation, and may be atributed in some measure to slower growth of the female due to the extra energy demands of reproduction. The overall 1:1 ratio of males to females in the population represents the balance between an equal sex ratio in the immature stages, more females in the 18 to 24 mm length category, and more males in the larger length categories. This condition is attributed mainly to the faster growth of the males.     

Reproduction and growth of the freshwater prawn, Palaemon paucidens (Decapoda: Palaemonidae) in a lake of Korea

This study analyzed the reproduction and growth of freshwater prawn, Palaemon paucidens De Haan 1844 from Suk-dang lake(Korea). The analysis of the sex ratio showed a higher proportion of males than that of females. The average size of the eggs was 6.12 (+/-0.55) mm3 in the non-eyed stage (stageA) and 7.20 (+/-0.86)mm3 in the eyed stage (stage B). The reproductive output (RO) calculated with the dried-body weight of an incubating female and the weight of the eggs in dried condition was 26.97% (n = 17) of the average females weight. Incubating prawn appeared in April, and the gonadosomatic indices(GSI) showed the highest value during three months from January to March. Based on the month when there was a high proportion of incubating females with a high GSI, the estimated spawning season of P. paucidens was April. The maturity of the female was evaluated by the development of the ovaries and the existence of eggs. The average body length when 50% of the females in the group reached maturity was 8.55 (+/-2.74) mm. The analysis of the length-frequency distribution showed that the life span of P. paucidens ranged from 12 to 13 months. Females mature faster than males.     

Latitudinal variation in the reproductive biology of the commensal crab Pinnaxodes chilensis (Decapoda: Pinnotheridae) along the Chilean coast

The pinnotherid crab Pinnaxodes chilensis is a common commensal of the edible sea urchin Loxechinus albus along the Chilean coast. Several aspects of the reproductive biology of P. chilensis were examined between April and June 1999, along temperature and salinity gradients, at three sampling sites along the Chilean coast (23°45'S-39°24'S). Results demonstrated significant differences in egg number, egg volume, dry egg weight, and reproductive output of ovigerous females, between the studied populations of northern and central-southern Chile. Egg volume, egg dry weight, and reproductive output of females decreased from high to low latitudes, while egg number increased from high to low latitudes, exhibiting a clear trade-off with egg volume. It has been shown that changes in environmental conditions (e.g. temperature, salinity) along a latitudinal gradient, can generate clines in reproductive characteristics in both free-living and parasitic species.     

Life-history traits of Plesionika martia (Decapoda: Caridea) from the eastern-central Mediterranean Sea

Life-history traits of Plesionika martia (Milne Edwards, 1883) were studied through data collected during six seasonal trawl surveys carried out in the Ionian Sea (eastern-central Mediterranean) between July 1997 and September 1998. P. martia was found at between 304 and 676 m depth, with the highest density in the 400-600 m range. Intraspecific, size-related depth segregation was shown. Recruitment occurred in summer at the shallowest depths. Juveniles moved to the deepest grounds as they grew. The largest female and male were 26 and 25 mm carapace length, respectively. The sex ratio was slightly in favour of females at depths >400 m. Although a seasonal spawning peak was shown, the reproduction appears to be rather prolonged throughout the year. Females with ripe gonads were found from spring to autumn. Ovigerous females with eggs in late maturity stage were found year round. Large females could spawn more than one time within their annual reproductive cycle. The size at first maturity (50% of the ovigerous females) was 15.5 mm CL. Average brood size of eggs with a well-developed embryo was 2,966ǃ,521. Iteroparity, low fecundity and large egg size patterns were observed. Brood size increased according to the carapace length. Two main annual groups were found in the field population of the Ionian Sea. Estimates of the Von Bertalanffy growth parameters are: L_X=30.5 mm, k=0.44 year^-1 in females; L_X=28.0 mm, k=0.50 year^-1 in males. A negative allometry was detected mostly in the ovigerous females. The life cycle of P. martia is discussed in the light of life-history adaptations shown in other deep-water shrimp species.     

Reproductive biology of the spiny lobster, <Emphasis Type="Italic">Panulirus Penicillatus</Emphasis>, in the southeastern coastal waters off Taiwan

The reproductive biology of spiny lobster, Panulirus penicillatus, was studied based on 2,068 lobsters, ranging from 34.28 to 131.60 mm carapace length (CL), sampled in Taitung coastal waters from September 2003 to December 2004. The overall sex ratio approximated 1:1 (χ² = 0.02, P > 0.05), but the monthly sex ratios in 2004 showed significant differences and males were predominant in sizes larger than 80 mm CL. Reproductive activity, assessed using histology, a gonadosomatic index and percentage of ovigerous females, indicated that the mature females could be found in every month and that the major spawning occurred from May to September. The presence of re-developing/re-ripe ovaries by month and size-specific spawning time suggest that larger mature females (>60 mm CL) spawn at least three times a year while smaller new mature females spawn at least once a year. For females, the estimated sizes at 50% physiological and functional maturity were (mean ± SE) 56.46 ± 0.56 mm CL and 66.63 ± 1.07 mm CL. The estimated sizes at functional maturity were between 72 and 74 mm CL for males. The number of eggs per spawning event (brood size, BS) was related to CL by the equation Y _BS = 2.4 × 10^-3CL^4.18 (r ² = 0.902, n = 12). Female lobsters with CL ranging from 60 to 80 mm made the greatest contributions to egg production because of their high brood size and active reproductive activity. A minimum legal size should be established for the fishery to protect egg production potential of lobster population in the southeastern coastal waters off Taiwan.     

Demography of <Emphasis Type="Italic">Nebalia</Emphasis> sp. (Crustacea: Leptostraca) determined by carrion bait trapping in Lobster Bay,Cape d‘Aguilar Marine Reserve,Hong Kong

The feeding ecology of leptostracans is not well-known. In the present study, species of Nebalia (Crustacea: Leptostraca) were captured every month for 1 year from Lobster Bay, Hong Kong, using baited traps. Captured individuals fell into a size range of 0.36–3.64 mm and 0.95–5.52 mm in carapace and total lengths, respectively. They were divided into different reproductive stages based on the morphology of the second pair of antennae and thoracopods. Sexual dimorphism becomes apparent between 0.7–1.5 mm (CL) and 1.4–3.2 mm (TL). Most individuals collected were sexually differentiated, but immature (79.7% of the captured individuals), followed by post-ovigerous (11.4%) and ovigerous (8.9%) females. Sexually undifferentiated juveniles were rare (3.9%). The sex ratio of the immature individuals approached 1:1 with a propensity towards male dominance. Not one mature male was captured. Although females with setose thoracopod endopods, indicating that they were ovigerous, were captured frequently, only two were carrying eggs. The two brooding females had clutch sizes of 11 and 17 eggs. Alternate peaks in relative abundance of ovigerous and post-ovigerous females were identified, the latter following the former. Reproductive activities occurred mainly in April–May and October–December, indicating that two cohorts are recruited. Catches using baited traps could be explained by the reproductive season of Nebalia sp. High catches were usually associated with the months of highest reproductive activity, low catches with peaks in post-ovigerous females. It is therefore concluded that not all individuals at various reproductive stages are attracted to carrion and hence adopt a scavenging feeding mode. Catches of Nebalia sp. in Hong Kong possibly reflect their annual cycle of reproductive activity     

Life history strategy of a southern European population of brown shrimp (Crangon crangon L.): evidence for latitudinal changes in growth phenology and population dynamics

This study provides a comprehensive examination of the life cycle dynamics of the brown shrimp (Crangon crangon L.) in the Mondego estuary, Portugal, a habitat located near the southern edge of the range of this species in European waters. The phenology of all life stages was documented (from the occurrence of ovigerous females, developing embryos, planktonic larvae and benthic post-larvae), and migration patterns of larvae into and out of the estuary were examined. Temperature-dependent functions of egg and larval development were combined with data on field abundance to predict the timing and magnitude of occurrence of larvae and recruits. Compared to brown shrimp at higher latitudes, southern conspecifics grow slower, mature earlier and have smaller brood sizes, and larvae have a more protracted settlement period. The Mondego estuary, besides acting as nursery area, is a dynamic platform for C. crangon to use in different stages, sizes and seasons.     

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.     

Allometric constraints and variables of reproductive effort in brachyuran crabs

Allometric relationships of reproductive output were compared in 20 species from 7 families of brachyuran crabs from the east and west coasts of North America, using regression analysis of log reproductive parameters versus log body weight. Comparisons of crabs spanning 4 orders of magnitude in body weight indicated that female body size was the principal determinant of reproductive output: 95% of the variance in brood weight, 79% of the variance in the number of eggs per brood, 63% of the variance in annual brood weight, and 74% of the variance in annual fecundity were explained by body weight. Brood weight exhibited an isometric constraint to about 10% of body weight. Allometric limitations on space available for yolk accumulation in the body cavity appeared to be the main constraint on brood size. Ovum size increased significantly, but only slightly, with increasing body size. There was a significant trade-off between ovum size and the number of eggs per brood. There was no significant relationship between the number of broods per year and body size. The number of eggs per brood was significantly better than brood weight as a predictor of the number of broods produced per year by a species, indicating that demographic pressure on fecundity rather than energetic considerations is the primary selective mechanism influencing annual reproductive effort. Each of the 7 families of crabs exhibited trends toward distinct patterns for the suite of co-adapted reproductive traits. However, no interspecific reproductive patterns were apparent with respect to the variables of feeding type, salinity tolerance, habitat, and geographic range represented by the 20 species.     

Causes of couspecific nest parasitism in the northern masked weaver

Summary Conspecific nest parasitism is a common reproductive strategy in the northern masked weaver (Ploceus taeniopterus). Parasites appear to be females with nests of their own who lay an additional egg in another female's nest as a way to enhance their reproductive success. Brood size in the northern masked weaver is practically constrained to three: starvation in four-chick broods is very common. The constraint on brood size is probably imposed by the extreme hatching asynchrony characteristic of this species: the last egg in a four-egg clutch hatches 72 h after the first egg, and when a chick starves, it is almost always the youngest chick. Late-hatching chicks also grow more slowly than do early-hatching chicks. If a female were to lay a fourth egg in her own nest, there is little chance that it would succeed. However; if she places it in another female's nest before that host lays her third egg, then it may have a greater chance of success.     

Brood defense and filial cannibalism in the spottail darter (<Emphasis Type="Italic">Etheostoma squamiceps)</Emphasis>: the effects of parental status and prior experience

Male spottail darters (Etheostoma squamiceps) defend nest sites in which females deposit eggs over the course of several weeks. In laboratory experiments, I tested three predictions of the hypothesis that the presence of eggs increases the value of a nest site to male spottail darters: (1) males should preferentially defend nests with eggs over empty nests, and males that sired the eggs should (2) spend more time defending them and (3) consume less of their broods than should alloparent males. Parent males were tested using eggs they were guarding in the field; alloparents included males that were not defending eggs immediately prior to the experiment (inexperienced), and males that were defending eggs in the field when captured but were given eggs sired by a different male (experienced). In choice experiments in which males were offered nest sites differing only in the presence of eggs, males preferred to defend nest sites with eggs regardless of parental status. Parent males spent significantly more time defending eggs and consumed less of their broods than did inexperienced alloparent males. Experienced alloparent males were similar to parent males in brood defense but had levels of filial cannibalism similar to inexperienced males, suggesting that prior experience may influence brood defense but not egg consumption.     

Comparative fecundity and reproductive strategies in seventeen species of the Cumacea (Crustacea: Peracarida)

The fecundity of 17 species of cumaceans belonging to 5 families from the eastern and western temperate regions of the Atlantic and the Canadian Arctic and Subarctic Oceans are discussed separately and then compared. The species range from 0.78 to 10.11 mm (mean carapace length) with fecundities ranging from 15.1 to 168.9 (mean number of eggs brood^-1 female^-1). Interspecific comparisons show that carapace volume versus volume of brood yields a very strong relationship (r=0.99). Therefore, it is a combination of carapace volume plus egg volume which determines the fecundity in the Cumacea. A discussion of the various life-history and reproductive strategies employed by the Cumacea is included.     

Reproductive dominance and differential ovicide in the communally breeding burying beetle Nicrophorus tomentosus

Communal breeding can be characterized by the division of reproduction among cooperating individuals and, if the distribution of reproduction is inequitable, by the mechanisms for achieving skewed reproductive success. The burying beetle (Nicrophorus tomentosus) is a facultative communal breeder. Unrelated adults, especially females, provide extensive parental care to broods of mixed parentage. The frequency and degree of reproductive skew between two females were examined experimentally. On medium-size carcasses, the proportion of eggs attributed to each female was not significantly different from random in 42% of the broods, skewed in 42% and not shared in 16%. Although reproduction was usually skewed in favor of the larger female, the relative sizes of the two females did not predict the degree of skew. On large carcasses, the proportion of eggs attributed to each female was not different from random in 87% of the broods and weakly skewed in 13%. Several mechanisms for biasing reproductive success were investigated. Females increase the proportion of their offspring in the brood by committing differential ovicide. Secondly, burying and preparing a carcass cooperatively stimulates ovarian development of the larger female and slows it for the smaller female, reducing or delaying oviposition by the subordinate. Thirdly, larger females are more likely to be dominant and are more fecund than smaller females. Received: 20 July 1996 / Accepted after revision: 30 November 1996     

Brooding of the sub-Antarctic heart urchin, <Emphasis Type="Italic">Abatus cavernosus</Emphasis> (Spatangoida: Schizasteridae), in southern Patagonia

The reproductive cycle of the sub-Antarctic spatangoid sea urchin, Abatus cavernosus, was examined during a 2-year period in southern Patagonia, Argentina. The population studied is the northernmost known coastal population in the austral oceans, and is influenced by a greater annual range of seawater temperature than other Abatus species. The sex ratio of the overall A. cavernosus population did not differ significantly from 1:1, but was not homogeneous across size classes. A clearly defined annual reproductive cycle was found. Spawning occurred from May to July and was synchronous between sexes. Females were observed to be brooding within a 9-month period, from May to February. Juveniles were released during the austral summer, from January to February. The length at which 50% of the females were brooding occurred at a test length of 25.9 mm. A. cavernosus had a large egg size (mean diameter = 1.4 mm) and low fecundity (maximum = 57 eggs per female) compared to closely related species. The number of eggs within each brood pouch was highest in larger anterior brood pouch, which is close to two gonopores, as opposed to the other anterior and two smaller posterior pouches. Significant interannual variation was observed in gonad cycles, fecundity, and embryo development such as: brood size decreased during 2001; adjusted gonad dry weight and fecundity were higher during 2003. Hypotheses concerning the gonadal and brooding cycles and fecundity of sub-Antarctic and Antarctic Schizasteridae are discussed. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Changes in physiology,structure and histochemistry occurring during the life-history of the sexually dimorphic isopod Dynamene bidentata (Crustacea: Peracarida)

The main food reserve of the intertidal hebivorous isopod Dynamene bidentata is lipid. Only algal-dwelling juveniles exhibit rhythms of oxygen consumption and defecation related to the tidal cycle. Males undergo the moult to maturity in the algal habitat and are, therefore, able to replenish their food reserves before entering upon a nonfeeding reproductive phase in empty barncle tests and erevices. Storage sites are increased at this moult so that males are able to survive on their food reserves for 2 breeding seasons. Females enter the reproductive habitat in the preovigerous stage and cease to feed. Lipid food reserves are largely used up in the formation of eggs and the final stage cuticle. Consequently, after the ovigerous moult, females are required to utilise lipoprotein-containing tissue as an energy source, and are able to survive only for a single breeding season. At the final moult, the posterior end of the male becomes highly thickened, rugose, and supplied with tegumental glands. These modifications are correlated with the need for self protection, and protection of the vulnerable females, against various physical and biological factors in the intertidal zone. There is a large increase in the dry weight of the brood over the period of incubation, and this is due mainly to the development of the exoskeleton at the last marsupial moult. The source of materials for the weight increase is attributable to a combination of factors, of which salt uptake probably plays the major part.