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.     

Multiple spawning in the tropical squid Photololigo sp.: what is the cost in somatic growth?

This study assesses the potential of the tropical loliginid squid Photololigo sp. to lay multiple batches of eggs and examines changes in somatic growth during reproduction. Histological analysis of the ovary and the relative size of the oviduct to mantle weight and ovary weight were used to determine the potential for multiple spawning. Ovaries of mature females always had immature and mature oocytes present, suggesting that not all the oocytes were maturing simultaneously and that multiple batches of eggs were being produced. Furthermore, poor correlations of oviduct weight with body size and ovary weight indicated that mature oocytes were not accumulating in the oviduct for a single spawning event. Both these observations supported the hypothesis that Photololigo sp. has the potential to lay multiple batches of eggs throughout its life. Specific growth rates, length-weight relationships, relative growth of somatic and reproductive tissue and microscopic assessment of muscle tissue were compared between immature and mature females. Growth rates of immature females were almost twice as fast as those of mature females. Mature females also had no large muscle fibres present, suggesting that energy for reproduction was mobilised from the muscle tissue.     

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.     

Reproduction in the deepwater squid<Emphasis Type="Italic"> Moroteuthis ingens</Emphasis>, what does it cost?

The cost of reproduction for the terminal spawning onychoteuthid squid, Moroteuthis ingens, was analysed using measures of condition and tissue biochemistry. Both males and females showed a dramatic drop in the weight of the gonad in stage 6 (spent) individuals. The mantle weight and nidamental gland weight of females also decreased during the maturation process. Males, however, had a marked increase in both the penis and spermatophoric complex weight in spent individuals, while female oviducal gland weight and nidamental gland length also increased in stage 6 individuals. Residual analysis indicated that testis growth was not developing at the expense of mantle growth, although there was a suggestion of cost to the fins. Females showed that the development of the ovary occurred at a cost to both the mantle and fins. Overall body condition also declined with maturity stage for both males and females, with stage 6 individuals of both sexes in poor condition. Very few females had eggs in the oviducts, suggesting that the oviducts are used as ducts instead of storage organs. Proximal analysis revealed a loss of constituents within the mantle during maturation, with an associated increase in water, indicating the remobilization of energy from the mantle to fuel reproduction. This study suggests that the digestive gland is not used as an energy store in this species.Communicated by M.S. Johnson, Crawley     

Fecundity and spawning in a deep-water cirromorph octopus

 Reproductive characteristics of cirromorph octopuses, assigned to the species Opisthoteuthis grimaldii, were sampled as a commercial fishing by-catch on the Hebrides Slope, west of Scotland. A total of 254 specimens (99 female, 155 males), retrieved from bottom trawls fished at 750 to 1500 m depth, were examined. A maximum of 2097 eggs was counted in a single female ovary (mean female body weight 1242.8 g), most of them <1 mm in length. At egg lengths over 1 mm, diminishing numbers of eggs were present in 1 mm size categories up to a maximum of ≈10 mm. At body sizes >500 g (wet wt), and in every female >750 g (max. female weight recorded in the sample was 2959 g), a succession of unattached eggs was present in the proximal oviduct and a single, unattached mature egg occupied the tip of the distal oviduct. These females were assumed to be in spawning condition and the characteristics of egg distribution in the reproductive tract to be consistent with sequential release of individual eggs and continuous spawning throughout the growth period and lifespan of the mature octopus. In pre-spawning females there was a positive relationship between estimated egg numbers and maximum egg size. After the onset of spawning there was no significant further increase in estimated potential fecundity over the body-size range 500 to 3000 g. Follicular sheaths remaining in the ovary after release of eggs into the proximal oviduct were counted and used to estimate the total number of eggs released up to the time of capture. Follicular sheaths first appeared at 500 to 650 g body weight and increased steeply in number to >1000 in females >1500 g. Two individuals were found with ovarian follicular sheaths but with no terminal egg in the distal oviduct; these were assumed to have released their egg just before capture. Summation of the number of follicular sheaths counted plus the number of eggs estimated as remaining attached in the ovisac, provided a revised estimate of total potential fecundity and raised the estimate for any individual to a maximum of 3202 eggs (mean = 1396 eggs). Received: 3 February 2000 / Accepted: 17 May 2000     

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.     

Fecundity of the Argentinean mackerel Scomber japonicus marplatensis

Research on the spawning and fecundity of the Argentinean mackerel Scomber japonicus marplatensis was carried out during the spawning season 1966/67. The purpose of the present study was to obtain data for the estimation of the number of eggs which can be produced and spawned by one female of a known size, during one season of reproduction: 152 females were studied. All yolked ova of the biggest size group were taken into consideration. On the basis of the relatively short spawning season, the great homogeneity in meristic characteristics, and histological examination, it is assumed that the Argentinean mackerel spawns once a year. The fecundity of the smallest female of 315 mm in total length was calculated to be 101,859 eggs, and the total number of yolked ova 181,000; for the biggest female of 470 mm in length, 522,370 and 853,500, respectively. The relation fecundity/length and fecundity/weight is almost rectilineal. The equations: F=8.17 ^0.008L for fecundity/length relationship, and F=0.42 W ^1.043 for fecundity/weight relationship, were calculated. Prediction curves were established in each case. It would appear that the fecundity of smaller fish is associated more with length, and the fecundity of bigger fish more with weight. Large differences in fecundity of specimens of the same size were observed.Communicated by M. Vannucoi, So Paulo     

Feeding dynamics of Octopus mimus (Mollusca: Cephalopoda) in northern Chile waters

The natural feeding of 485 Octopus mimus (164 to 3088 g) was studied in relation to the species' life cycle and environmental seasonal variations off the north of Chile from autumn 1991 to summer 1992. Analysis of digestive-tract contents revealed that O. mimus preyed upon 25 different prey items belonging to five zoological groups (Teleostei, Mollusca, Crustacea, Echinodermata and Polychaeta). Cannibalism was only occasional. The results indicate that the diet and food intake of this species are significantly affected by sex and maturation. Senescent individuals ingest a small amount of food, and their diet is mainly based on small, not very motile prey. The food intake, expressed as body weight, of non-senescent individuals is higher in females than in males. Seasonal changes in sea-water temperature seem to be followed by adjustments in food intake. Like other Octopus species, O. mimus appears to be an opportunistic predator.     

Maturation,fecundity, and spawning strategy of the thornback ray, <Emphasis Type="Italic">Raja clavata</Emphasis>: do reproductive characteristics vary regionally?

In Portuguese waters, thornback ray Raja clavata spawns mainly between May and January, although females and males in spawning condition are found throughout the year. The maturation process can be divided into three main phases by using information on gonad weight, oviducal gland and uterus width in females and on gonad weight, clasper length and sperm duct width in males. Females attain length-at-first-maturity at 784 mm and males at 676 mm, at ages of 7.5 and 5.8 years, respectively. In females larger than length-at-first-maturity, a resting stage was identified characterized by low gonadosomatic index and well-developed oviducal glands and uteri. These results along with the low proportion of adult females that are effectively reproducing per month demonstrated that the thornback ray cannot be considered a continuous spawner as described in other studies. Fecundity was determinate with about 35 eggs released per batch. During the spawning season, a total of four batch episodes occur indicating that the total fecundity was approximately 136 eggs per female. Regional differences may exist in the reproductive strategy of the species, namely on the duration of the spawning season, length-at-first-maturity, and fecundity, which can be related to a more intense fishing pressure in northern European waters.     

Temporal shifts in the allocation of energy in the arrow squid,<Emphasis Type="Italic"> Nototodarus gouldi</Emphasis>: sex-specific responses

Squid typically display considerable intra-specific plasticity in size and age-at-maturity in response to ambient environmental conditions, yet little is known of the mechanisms driving these variations. We examined the intra-specific variability in Nototodarus gouldi reproductive traits to determine patterns of energy allocation between somatic and reproductive processes over short temporal scales. Females caught during the cool months of May and July were larger, had slower lifetime growth, lower gonad investment, and better somatic condition than females caught during the warmer months, suggesting a trade-off between gonad investment and somatic condition in females. On the other hand, males showed a tight coupling between somatic condition and gonad investment for most months, with increases in somatic and gonad tissue occurring concurrently. In male squid, an increase in lifetime growth rate was coupled with an increase in the relative weights of somatic and reproductive structures, whereas in females, percent increase in body weight per day was correlated only with gonad development. Patterns of repro-somatic investment in mature females had implications for spawning strategies, since female squid with higher levels of gonad investment apparently released batches of eggs together as a group, regardless of body size, whereas females with low gonad investment possibly spawned their eggs independently of one another. In terms of life-history theory, male squid were able to respond rapidly to environmental fluctuations without compromising either the gonad or the soma. However, although mature females did not appear to respond as quickly to ambient conditions, female squid possibly produced two different reproductive strategies, possibly to maximise offspring survival in either a stable or a variable environment. It seems from our study that monthly variations in ambient conditions may have large effects on life-history strategies.Communicated by M.S. Johnson, Crawley     

Does the spawning of Calanus finmarchicus in high latitudes follow a reproducible pattern?

In the spring of 1989, an experimental study of the spawning behaviour of Calanus finmarchicus was carried out in Malangen, northern Norway. Here, a single cohort of females reproduce from mid-March to May, approximately coinciding with the wax and wane of the spring phytoplankton bloom. An evaluation of population characteristics such as the proportion of adults, sex ratio, as well as gonad maturation and daily productivity of the females clearly reveals three phases within the population's reproductive period. In between incline and decline, the highest spawning rates (on average >20 eggs female^-1 d^-1, equivalent to 5.7% body C d^-1) occur after the males have disappeared from the population and almost all females have mature gonads. During this period, the ratio of adults to copepodid Stage Vs changes from dominance of adults to that of CVs. Although first egg production was observed prior to the phytoplankton increase, it is suggested that the onset of the phytoplankton spring bloom in the first few days of April enhances the final maturation of ovaries in the females and therefore triggers the onset of the main spawning period. The clutch sizes (max. 95 eggs clutch^-1) vary with the age of the females, while the spawning frequencies depend on the available food quantities. The overlap of an estimated minimal 4 wk spawning period for the individuals leads to a main reproductive phase for the population of ca. 3 wk, during which time mean clutch sizes and spawning frequencies are maximal (highest average clutch size: 70 eggs female^-1 clutch^-1, 100 to 60% of the females spawning). This period ends before the end of the phytoplankton bloom. Calculated by stepwise interpolation and summation of the mean daily egg production in the population, an average female produced ca. 600 eggs during the spring bloom in Malangen 1989. We suggest that reproduction and population development of C. finmarchicus in spring follows a reproducible pattern for a given temperature regime and non-limiting food conditions. In the case of clearly identifiable cohorts, it seems possible to trace the state of reproduction by evaluating population parameters.     

Reproductive status and body condition of Atlantic bluefin tuna in the Gulf of Maine, 2000–2002

The reproductive status and body condition of 195 (≥185 cm curved fork length, CFL; assigned age 7 and above) Atlantic bluefin tuna were assessed in the Gulf of Maine during the commercial fishing season of June–October, 2000–2002. Given the distance between known spawning and feeding grounds, the prevailing paradigm for Atlantic bluefin tuna (Thunnus thynnus thynnus, L.) suggests that the most likely histological state for females arriving in the Gulf of Maine after spawning would be a resting or quiescent state with little or no perigonadal fat. Alternatively, the presence of mature or mature-inactive histological states in some females supports a more varied or individualistic model for bluefin reproduction. No relationship was found between body condition and reproductive status. Males were found in all reproductive stages, but were more likely to be in spawning condition (stages 4 and 5) or a mature-inactive state (stage 6) in June and July. Female bluefin tuna were found in stage 1 (immature or non-spawning) and stage 6 (mature-inactive). Stage 6 females were only present in June and July and smaller females (<235 cm CFL) were more likely to be in stage 6 than large females (>235 cm CFL) sampled during those same months. The presence of smaller females in stage 6 arriving at the same time as larger females in stage 1 indicates that Western Atlantic bluefin tuna may have an asynchronous reproductive schedule and may mature at a smaller size than the currently accepted paradigm suggests.     

Paternity and condition affect cannibalistic behavior in nest-tending bluegill sunfish

Theory of parental care evolution predicts that a parent should invest more in a brood when its fitness value is greater than alternative investments such as the parent's own survivorship or future broods. In fish, filial cannibalism (eating one's own offspring) is widespread and represents a challenge to parental care evolution. In this study, I investigated filial cannibalism in bluegill sunfish (Lepomis macrochirus). Bluegill are characterized by alternative mating tactics referred to as "parentals" and "cuckolders". Parentals delay maturation, construct nests, court females and provide sole parental care for the developing offspring. Cuckolders mature precociously and parasitize parentals using two tactics called "sneakers" and "satellites". I found that parentals that obtained fewer eggs during spawning appeared more likely to completely cannibalize their brood (total filial cannibalism: P=0.07), regardless of their condition. Among parentals that provided care, partial cannibalism was greater during the egg phase as compared to the fry phase of care, but it was unrelated to brood size. Throughout the care period, parentals in better condition were less likely to partially cannibalize their brood, indicating that parentals use cannibalism to replenish energy reserves. Independent of condition, parentals that were cuckolded more were more likely to eat part of their brood. This relationship was evident only after the eggs had hatched, which is consistent with data showing that parentals can use olfactory cues produced by fry but not eggs to assess their paternity. This latter result proposes that parentals may be selectively culling cuckolder offspring from their nest. These data provide empirical support for parental care theory, and the first evidence for the importance of paternity on cannibalistic behavior.Communicated by M. Abrahams     

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

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

Continuous spawning in the cirrate octopods Opisthoteuthis agassizii and O. vossi: features of sexual maturation defining a reproductive strategy in cephalopods

The reproductive strategy of the cirrate octopods Opisthoteuthis agassizii and O. vossi (collected off Namibia from 1988 to 1990) was analyzed. Ovarian oocyte size frequency analysis for both species revealed continuous egg production over the entire adult life span. Mature eggs were stored in the single oviducal gland and distal oviduct, but oviducal gland fullness was not related to body size (p>0.2). All O. agassizii male specimens from 95 to 5400 g total weight were sexually mature, as were all females from 190 to 1650 g, indicating that considerable growth takes place after the onset of sexual maturity. Continuous spawning is defined as a single, extended and continuous period of egg maturation and spawning. This model of reproductive strategy is previously unreported in cephalopods. All O. vossi male specimens from 750 to 3050 g total weight, and females from 800 to 1300 g, were sexually mature. Mature males and females of both species were collected in all seasons of the year. The adaptation of cirrate octopods to non-scasonal deep-sea environments is considered. The sexual maturity characteristics of males were analyzed, and examination of the spermatophore revealed opercular structures previously unreported in cephalopods. For females, the micropyle of the eggs are described and the mineral analysis of the egg shell disclosed that sulphur was the major element present.     

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.     

Spawning pattern and type of fecundity in relation to ovarian allometry in the round herring <Emphasis Type="Italic">Etrumeus teres</Emphasis>

Spawning pattern (assessed by seasonal changes in ovarian developmental stages) and type of fecundity (assessed by analysis of oocyte-size frequency distributions) of the round herring Etrumeus teres were studied in relation to ovarian growth and seasonal changes in the gonadosomatic (GSI), hepatosomatic (HSI) and liposomatic (LSI) index as well as the somatic condition of spawners (CS) in a spawning ground of southern Japan. Except for summer, mature and recently spawned ovaries occurred all year round. Oogonia and primary oocytes were present in all ovaries, and cortical alveoli stage (CA) oocytes occurred in all mature, hydrated and partially spent (PS) females (PS: females containing post-ovulatory follicles). Before hydration, a clutch of larger yolked oocytes, undergoing synchronous growth (range 0.7–1.1 mm), was present in mature ovaries which was completely separated from a more heterogeneous clutch of oogonia, primary and secondary oocytes (<0.150 mm) and oocytes in the CA stage (range 0.15–0.60 mm). As vitellogenesis progressed, the yolked clutch increased in size but the CA oocytes remained arrested. The latter entered into the secondary growth phase when hydration started in the advanced batch. Ovarian growth was isometric in all developmental stages, validating the use of GSI, which showed a consistent monthly evolution among years. Spawning stopped in summer (July and August) and peaked in winter and spring. HSI correlated positively with GSI on both a monthly mean basis (r = 0.76) and individual fish basis (liver weight explained 67–83% of the variability in ovary weight when females were grouped into 1-unit GSI intervals) suggesting a significant role of liver in vitellogenesis. LSI and CS also showed marked seasonal changes peaking from summer to middle autumn. Overall results suggest that E. teres is a multiple spawner with a group-synchronous ovarian development and indeterminate annual fecundity, with the three processes linked to an isometric growth of the ovary. We propose that such a reproductive pattern is an adaptation to produce batches of large pelagic eggs through a protracted spawning season.     

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     

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.     

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.