Maturation of female common snook Centropomus undecimalis: implications for managing protandrous fishes

The assumption for hermaphroditic fish species that mature individuals of the terminal sex arise directly from mature individuals of the primary sex has led to the use of sex ratios as a proxy for age at maturity (A₅₀). The timing of transition and deficient energy reserves, however, can result in a delay between transition and spawning. To test the assumption of female maturity and investigate the relationship between maturation and energy reserves, common snook, Centropomus undecimalis, a protandrous hermaphrodite, were collected from rivers, estuaries, inlets and offshore habitats on the east coast of Florida during 2010–2015. Immature females were observed every month, with lowest proportions during the peak spawning months of July and August. When calculated based on sex ratio, A₅₀ (8.1 years) overestimated the age at which 50% of the female population was, in fact, mature (4.1–4.9 years). Best-fit models indicate that mesenteric fat index (I_F) and hepato-somatic index (I_H) were significantly affected by gonad phase, month and size and weakly by habitat. In post hoc analysis, immature female I_F did not differ significantly from developing and regenerating females, but immature female I_H was significantly lower than that for all mature phases except animals in the regressing phase. Although immature females may have sufficient energy in terms of fat, it appears that energy is not allocated to reproductive processes, as evidenced by lower I_H. Nonetheless, approximately 95% of females were spawning-capable during peak spawning months, suggesting that the energy threshold at which immature females reach maturity is met by most females each spawning cycle.     

Strong sexual selection in males against a mutation load that reduces offspring production in seed beetles

Theory predicts that sexual reproduction can increase population viability relative to asexual reproduction by allowing sexual selection in males to remove deleterious mutations from the population without large demographic costs. This requires that selection acts more strongly in males than females and that mutations affecting male reproductive success have pleiotropic effects on population productivity, but empirical support for these assumptions is mixed. We used the seed beetle Callosobruchus maculatus to implement a three‐generation breeding design where we induced mutations via ionizing radiation (IR) in the F₀ generation and measured mutational effects (relative to nonirradiated controls) on an estimate of population productivity in the F₁ and effects on sex‐specific competitive lifetime reproductive success (LRS) in the F₂. Regardless of whether mutations were induced via F₀ males or females, they had strong negative effects on male LRS, but a nonsignificant influence on female LRS, suggesting that selection is more efficient in removing deleterious alleles in males. Moreover, mutations had seemingly shared effects on population productivity and competitive LRS in both sexes. Thus, our results lend support to the hypothesis that strong sexual selection on males can act to remove the mutation load on population viability, thereby offering a benefit to sexual reproduction.     

Fat cycling in the mosquitofish (Gambusia affinis): fat storage as a reproductive adaptation

Summary We argue, based on reviewed literature covering reptiles, amphibians, birds, and fish, that fat storage may represent a life history adaptation because it enables an organism to shift in time when resources are allocated to reproduction. We applied these arguments to fat and population cycles in three populations of the mosquito fish, Gambusia affinis. For males, there appeared to be a constant size at maturation during the reproductive season. Mature males became scarce late in the summer. At the same time, immature males delayed maturity and attained much larger sizes; they matured in large numbers in the fall. The amount of stored fat tended to be equal for immature and mature males at all times except in the late summer. In the August samples, when mature males were relatively rare, they also had the lowest level of fat reserves. It appears that the older generation of mature males did not store fat and did not overwinter. At the same time, immature males registered a two to three fold increase in fat reserves. These differences in fat content between mature and immature males disappeared by September, probably because of the recruitment of a new generation of mature males. The reserves were gradually utilized during the winter. Females reproduced from the late spring through mid- to late-summer. They stopped reproducing in the late summer, when there was ample time to produce an additional litter of young. There was an inverse relationship between resources devoted to reproduction and fat reserves. As reproductive allotment decreased in the late summer, fat reserves increased. The magnitude of the change in fat reserves was similar to that displayed by males. The reserves were depleted over the winter. Significant reserves remained at the beginning of the reproductive season the following spring. Reproducing females utilized the remaining reserves significantly more rapidly than non-reproducing females. An analysis of resource availability revealed an overall decrease in food availability in the late summer, coincident with the increase in fat reserves. These cycles are therefore not attributable to changes in resource availability. They instead indicate a change in how resources are allocated by the fish. The trends in the data indicate that fat reserves are used to shift investment in reproduction from the late summer to the following spring. In males, deferring maturity, rather than maturing in August, allows them to store the necessary reserves to survive the winter so that they can mate the following spring. In females, a subset of the fat reserves is intended for producing the first clutch of eggs the following spring. The female pattern corresponds to those reported for a diversity of organisms. The possible advantages of shifting reproductive effort from the fall to the following spring include higher fecundity and higher offspring fitness. The limitations of the methodology and potential directions for future research are discussed.     

Older triploid fish retain impaired reproductive endocrinology in the European sea bass Dicentrarchus labrax

This paper reports on an evaluation of growth, gonadal development and reproductive endocrinology of older triploid (3n) European sea bass Dicentrarchus labrax in comparison with their diploid (2n) counterparts throughout their fifth and seventh annual cycle of life. While older triploids retained impaired reproductive endocrinology, a sexually related dimorphic growth was observed with 3n females attaining the largest sizes. Comparisons of some body indexes showed that 3n females had a significantly lower hepato‐somatic index (I_H) than 2n females but a significantly higher viscero‐somatic index (I_F). In contrast, both male and female triploids showed significantly lower gonado‐somatic index (I_G) than diploids. Accordingly, diploids produced mature gametes but triploids did not, demonstrating that despite the longer time given to triploids for gonadal development, they could not reproduce. Furthermore, older triploids had lower levels of plasma sex steroids (testosterone, T; 11‐ketotestosterone, 11‐KT and oestradiol‐17β, E₂) and luteinizing hormone (LH) than their 2n counterparts with 3n females showing drastic effects of triploidization on their reproductive endocrinology. Vitellogenin (VTG) was undetectable in 3n females. Gonadal content of steroid hormones and Sparus aurata‐type gonadotropin‐releasing hormone (sbGnRH) in the brain and pituitary were also lower in triploids compared with diploids. These results suggest that older 3n D. labrax retain functional sterility in both sexes, and 3n females might reach larger sizes than 3n males and their 2n counterparts in this species.     

Lipid dynamics in the viperine snake, Natrix maura, from the Ebro Delta (NE Spain)

Body reserves play a major role in several aspects of vertebrate biology. Accurate identification and quantification of body reserves constitute a useful contribution to the better understanding of the energetic costs of reproduction and the implication of food availability in life history traits of organisms. In this study, lipid content in fat bodies, liver and muscle of the viperine snake (Natrix maura) were measured along an active season. Samples were collected monthly from a natural population of the Ebro Delta Natural Park (NE Spain). This methodology pointed out that lipids stored in fat bodies were the main energetic source during reproduction. In addition, lipids stored in liver appeared to be critical for vitellogenesis, while lipids stored in muscle turned out to be a supplementary energetic resource to fuel reproductive effort. Relationships between changes in body reserves and prey availability in canals of the Ebro Delta were also considered. In males, lipid reserves presented a positive correlation with food availability. On the contrary, lipid reserves in female viperine snakes decreased during vitellogenesis even though food availability increased in this period, which suggests a quick transfer of body lipids to clutch. In April, when rice fields of the Ebro Delta were dry and aquatic prey was scarce for viperine snakes, males and females presented a lower lipid content in fat bodies, liver and muscle than they did in other months, showing a clear link between prey availability and body reserves during food shortage. Thus, patterns of variation in fat levels indicated that Natrix maura is a capital breeder since it acquires resources in advance and stores them until they are invested during the reproductive period. Nevertheless, the shortage in April forces Natrix maura to turn into a facultative income breeder to fuel vitellogenesis. Finally, fat reserves in body components were compared with an estimate of body condition calculated by the residuals from the regression of body mass on body length. In male viperine snakes, the estimate of body condition was correlated with fat levels, revealing that this index is useful to measure condition in living individuals. On the contrary, body condition in females was not correlated with fat levels, which suggests that it is not appropriate to infer condition in female viperine snakes since it depends on the reproductive status of the individuals.     

Annual reproductive activity of Caiman crocodilus fuscus in captivity

We describe the gonadal and abdominal fat storage cycles for a population of Caiman crocodilus fuscus kept in captivity at a breeding farm. The reproductive cycles of males and females are clearly seasonal. Vitellogenic follicles were observed in females year‐round; however, the follicular growth that produces the preovulatory follicles for the breeding season occurs in November–May (most of the dry season to the beginning of the rainy season). These months correspond with the presence of physiologically reproductive males. Oviposition takes place during the wet season (April–August) and the births occur at the end of this season (September–November). Fat storage was not related to reproductive activity, and did not vary by month or between sexes. Although environmental and hormonal conditions have been improved for these captive animals, they maintain a strongly seasonal reproductive cycle, historically fixed to oviposit in the most favorable period for nest construction and hatching during the rainy season. Zoo Biol 22:121–133, 2003. © 2003 Wiley‐Liss, Inc.     

Age-based life history of humpback red snapper,Lutjanus gibbus,in New Caledonia

This paper examines the life history of humpback red snapper Lutjanus gibbus, an important fishery species for coastal communities across the Indo-Pacific, in southern New Caledonia, where the species is lightly exploited. A total of 243 L. gibbus were sampled between January 2013 and December 2016 from occasional harvests of commercial fishers. Examination of sectioned otoliths revealed that opaque increment formation occurred annually between November and March, coinciding with the species' spawning season. Estimates of maximum age were similar between sexes, with observed ages of 38 and 36 years for females and males, respectively, extending the reported longevity of this species by at least 11 years. Growth differed significantly between sexes, with males reaching greater length at age and greater asymptotic length than females (38.88 v. 31.46 cm fork length (L_F). Total mortality for all samples was estimated as 0.13 and was slightly higher for males (0.16) than females (0.11). Estimates of natural and fishing mortality were low and slightly higher for males than females. Male L. gibbus were found to mature at slightly greater lengths and younger ages than females, with the length and age at which 50% of individuals attained maturity estimated to be 25.8 cm L_F and 3.9 years of age for females and 26.8 cm L_F and 3.4 years of age for males. The results provide key baseline information from which to assess the effect of fishing on the species for populations in New Caledonia and adjacent locations and, when viewed with those of other studies, highlight the importance of understanding spatial patterns in demography of harvested fish species across gradients of exploitation and environmental influences.     

Age and growth of the sand smelt, Atherina boyeri (Risso 1810), in the Mar Menor coastal lagoon (SE Iberian Peninsula)

The age and growth of sand smelt, Atherina boyeri (Risso 1810), in the Mar Menor (SE Iberian Peninsula) were studied in samples taken from catches of local fishermen obtained between November 1997 and September 1998. The maximum lengths were 94 mm (FL, fork length) in females and 87 mm (FL) in males. Age determination based on scale readings and validated by length frequency analysis shows that the population has a 3‐year life cycle. Females were significantly longer than males in each of the age classes. Both sexes of the sand smelt grow allometrically (b = 3.113 males; b = 3.043 females) and attain approximately 56.2% of their maximum fork length in their immature first year, after which the annual growth rate drops quickly. The highest growth rate was observed from winter to spring (G_L = 16.01 males 1+; G_L = 11.25 males 2+; G_L = 17.18 females 1+; G_L = 9.62 females 2+). The condition cycles were similar for both sexes, with a minimum in June–July and two maximums in April and November.     

Reproductive biology of deep-sea catsharks (Chondrichthyes: Scyliorhinidae) in the eastern North Pacific

Apristurus brunneus, A. kampae, and Parmaturus xaniurus were caught by trawl and longline between Washington and southern California (48°N to 32°N latitude) from June 2001 through October 2004. In females, oviducal gland width increased with TL in all three species but remained within 4 to 6% of TL. Oocyte diameter increased with TL during reproductive development in all species, and continued to increase with size after reproductive maturity was reached. The weight-length relationships differed significantly among species but not between sexes of any species from the eastern North Pacific combined. Using logistic regression, it was determined that total length at 50% maturity (TL₅₀) of A. brunneus males was 514 mm TL and all A. brunneus males were mature at 547 mm TL. Female A. brunneus reached TL₅₀ at 501 mm TL and all female A. brunneus were mature at 581 mm TL. Apristurus kampae males and females reached TL₅₀ at approximately 485 mm TL and 490 mm TL, respectively. Parmaturus xaniurus males reached TL₅₀ at 444 mm TL and females at 501 mm TL. Gravid female A. brunneus were found in all months of the year, but gravid P. xaniurus were found primarily June through September. Adult A. kampae females were found only in July through November and there were gravid females in all these months. Based upon gonadosomatic indices, I _G, these species may mate within a defined period of time (April through June); however, A. brunneus appear to reproduce throughout the year and P. xaniurus reproduce within a defined season (approximately July through September).     

Sexual competition in an inbreeding social spider,Stegodyphus dumicola (Araneae: Eresidae)

Summary Sexual competition is shown to occur in the social spiderStegodyphus dumicola (Eresidae). While the secondary sex ratio inS. dumicola was female-biased, the overall operational sex ratio (numbers of breeding males to breeding females over the season) showed no strong female bias. Males matured before females and had a shorter lifespan than the females. Mating took place in the natal colony. Males fought over access to the few mature females available early in the reproductive season, but females appeared to control the duration of mating. Later in the season, some adults of both sexes dispersed alone to breed elsewhere. We conclude that different rates of maturation between the sexes within a colony provide the opportunity for females that mature early in the season to be choosy in selecting a mate and this forces males to compete. Early reproduction may be beneficial for both females and males, because the offspring of females that reproduce early may have a competitive advantage over later (and smaller) offspring in the colony.     

Variation in body condition of breeding Savi’s Warblers Locustella luscinioides: the reproductive stress and flight adaptation hypothesis revisited

Current theory suggests that mass change in adult birds while breeding may be adaptive (to reduce wing-loading during nestling feeding) or result from physiological stress. To test which might be more important in determining mass loss in breeding Savi’s Warblers (Locustella luscinioides), we used a new approach in which the variation in four indices of body condition was described: weight, fat score, muscle score and lean weight (i.e. excluding fat and muscle). We expected weight variations to be adaptive if they involved changes in fat and lean weight, whereas physiological stress should influence the muscle score to a greater extent. As in other species, females showed a greater variation in weight, and carried more fat, than males during the breeding cycle. During incubation, females had greater weight and fat score than males. The weight remained constant and lean weight declined in both sexes, whereas females increased in muscle, which probably reflects the regression of the reproductive organs. During the nestling stage, both sexes declined significantly in all four indices of condition, showing evidence of physiological stress. However, the greater decline in weight in females than in males is consistent with the flight-adaptation hypothesis, as are the cyclic changes in lean weight associated with the various nesting attempts. The fact that both sexes declined significantly in weight, muscle and lean weight with an increasing number of nesting attempts, but not in fat, which was recovered after each nestling period, also indicates that both reproductive stress and adaptive changes occur during breeding. When the whole breeding season was considered, females showed a greater decline in muscle than males, which we interpret to be evidence for a greater reproductive stress in females. We suggest that the small breast muscle size and depleted protein reserves at the end of the breeding period might influence future survival through impaired flight ability and a compromised post-breeding moult.     

Why Callosobruchus rhodesianus causes limited damage during storage of cowpea seeds in a tropical humid zone in Togo

The sympatric bruchids Callosobruchus rhodesianus (Pic) and Callosobruchus maculatus (F.) (Coleoptera: Bruchidae) colonize cultures of cowpea, Vigna unguiculata (Walp.) (Papilionaceae), in Togo at the end of the rainy season. Seeds containing larvae of the two bruchid species were introduced into the storage systems. Callosobruchus rhodesianus adults emerged from the seeds at the beginning of storage, but this species disappeared quickly and C. maculatus became dominant. Analysis of the reproduction of females collected at the beginning of storage made it possible to distinguish between four groups of females differing in their reproductive activity. Three groups were sexually active but they differed in their fecundity and the duration of the latency period before the first oviposition phase. The fourth group of females was in reproductive diapause. In interspecific competition, the presence of C. maculatus larvae reduced the survival chances of C. rhodesianus and therefore is disadvantageous for this species. The main factors explaining the rapid decrease of the C. rhodesianus populations under storage conditions were the variability of female reproductive activity with the emergence of insects in reproductive diapause and the significant larval mortality in interspecific competition with C. maculatus.     

Juvenile hormone favors sexually-selected traits but impairs fat reserves and abdomen mass in males and females

The physiological mechanism underlying resource allocation in sexual selection studies has been little studied. One candidate is hormones as these favor resource allocation to reproductive traits but impair survival due to a resource over-expenditure directed to the former traits. We have investigated whether a juvenile hormone analog (JHa, methoprene) administrated topically is involved in the resource allocation to wing pigmentation (an ornamental trait), fat reserves and flight muscle mass in both sexes of Calopteryx haemorrhoidalis and C. virgo. We also investigated the possible negative effect of such implementation on abdomen mass (an indirect measure of egg production) and field-based survival in adult males of C. haemorrhoidalis and C. splendens. We found that males and females treated with JHa, against a control group, developed higher wing pigmentation and showed reduced fat reserves but had no change in muscle mass. In females, JHa decreased abdominal weight (an indicator of fecundity) and in males, survival was impaired only in C. splendens. These results support the idea that JH induces resource allocation to wing pigmentation, a sexually selected trait in both sexes. Thus, this study suggests that the action of JH could be a mechanistic link between ornaments and physiological condition in both males and females.     

The reproductive biology of the two-banded sea bream (Diplodus vulgaris) from the southwest coast of Portugal

The study of the reproduction of Diplodus vulgaris (Geoff.) as part of a base‐line study of the fishery resources of the south‐west coast of Portugal, was based on the analysis of the spawning season, gonad maturation, size at maturity, fecundity and hermaphroditism. The spawning season is relatively long, from December to March, with peaks in January and February. No significant differences were found either in the sex ratio (M : F = 1.01) over the year or by size. The size at first maturity (L₅₀) for all sexes and undetermined individuals combined was 18.33 cm total length (TL), with no significant differences between males and females. The estimated L₅₀ is considerably greater than the minimum legal size in Portugal of 15.0 cm. Mean absolute fecundity (F_a) was 131 127 oocytes, ranging from 31 523 to 250 608. The relationship between absolute fecundity and total length (TL) (F_a = 25 398 TL – 484 426) and somatic weight (SW) (F_a = 878.8SW – 71 416) was of the linear type. The mean number of oocytes per gram of somatic weight was 526, ranging from 194 to 887. The reproductive strategy of this species is characterized by a rudimentary hermaphroditism with possible protandry, as evidenced by the existence of individuals in sexual transition.     

Reproductive biology of hardhead catfish Ariopsis felis: evidence for overwintering oocytes

Hardhead catfish Ariopsis felis are a common marine catfish in the coastal waters of the Gulf of Mexico (GOM). The low economic value of this species has depressed interest and research, and although the species is known for its extremely low fecundity and large oocytes, little else is known about this catfish species. A total of 1230 samples across all months of the year from 2016 to 2018 resulted in 681 females, and analysis of gonado-somatic index (I_G) revealed 1% to be a clear cut-off indicating maturity. Females are considered capable of spawning from April to June when I_G averaged 4–8%. Both atresia and post-ovulatory follicles were present in July, suggesting that spawning ends in July in the northern GOM. The 1% I_G cut-off was used to designate maturity, and from that an L₅₀ of 253 mm was estimated. Batch fecundity from 41 females estimated a mean batch size of 36 oocytes. Perhaps the most interesting finding was the presence of secondary growth stage oocytes (e.g., cortical alveoli) from July through November, well outside the spawning capable period. Furthermore, 78% of females had some early vitellogenic oocytes present during the non-spawning season, and the distribution of these relatively large (2–5 mm) oocytes did not change over time. The results here are not only important as reproductive biology information for a common and abundant species, but also present interesting and unusual patterns of non-spawning season oocyte development that is not commonly seen in Western Hemisphere subtropical fish species.     

Sexual differences in biomass and nutrient allocation of first-year Silene dioica plants

Reproductive and somatic biomass, nitrogen (N), and phosphorus (P) pools were compared between females and males in 1st-year plants of Silene dioica. We estimated irretrievable resources allocated to seeds, pollen, flowers, and unrecovered summer leaf investment by collecting plant parts at abscission throughout the season. At the end of the season, we determined resources lost through senescent stems and autumn leaf turnover and resources stored in perennial roots and overwintering buds. Sexual differences in allocation patterns depended on the resource used for comparison, and whether absolute or proportional resource pools were assessed. Total resource pools in terms of biomass and N were similar for females and males. However, male plants acquired relatively more P. The proportional reproductive investment, i.e., reproductive effort, was similar for males and females in terms of biomass and N. In terms of P, male reproductive effort was higher. There was no difference between sexes in the proportional and relative biomass allocated to perennial roots and overwintering buds. However, in terms of absolute and relative N allocation to below-ground parts, females had larger reserves than males. Females, moreover, had a larger proportion of their P in below-ground parts. However, as male total P pools were larger, absolute P reserves did not differ between sexes. The high reproductive effort and N depletion of below-ground parts in males resulted largely from higher flower production compared to females. In females, seeds were the major component of reproductive effort. These results show that if biomass and nutrient allocation are assessed in parallel for dioecious plants, we obtain a more complete view of their sexual differences. Received: 07 May 1998 / Accepted: 30 October 1998     

Age‐based demographic and reproductive assessment of orangespine Naso lituratus and bluespine Naso unicornis unicornfishes

Bluespine unicornfish Naso unicornis and orangespine unicornfish Naso lituratus were sampled in Pohnpei and Guam, Micronesia, over 13 months to identify reproductive and age‐based demographic features necessary for informed management. Age and reproductive information were derived from analysis of sagittal otoliths and gonads. Both species had moderate life spans [maximum ages of 23 (N. unicornis) and 14 years (N. lituratus)] compared with published estimates of conspecifics from other locations (>30 years) and of other Naso species. Length at maturation for N. unicornis was similar between Pohnpei and Guam while females consistently matured at a larger size [c. 30 cm fork length (L_F)] than males (c. 27 cm L_F). This sex‐specific pattern was reversed in N. lituratus for which estimates of maturation length (females: 15 cm L_F; males: 18 cm L_F) were only obtained from Guam. Developmental patterns in female gonads of both species suggested that initiation of maturation occurs very early. Growth patterns of N. lituratus displayed rapid asymptotic growth compared with N. unicornis and other congeners as well as slight sex‐specific patterns of length‐at‐age. Results highlight the considerable spatial variation that may occur in the population biology of these species across various scales. Additionally, proper management remains complicated without improved knowledge of fishery trends and reproductive behaviour in unicornfishes, species that are prime fishery targets in Micronesia and elsewhere.     

The costs of reproduction to males and females of a paternal mouthbrooding cardinalfish Apogon notatus

In the cardinalfish Apogon notatus , starving males mouthbrood the eggs, and females are more active than males in courtship and in attacks against conspecific intruders upon their breeding territory, but sexual ornaments are developed in males. Condition factor, hepatosomatic index and fat body-somatic index declined in both sexes during the breeding season. The deterioration of somatic condition was more severe in females, suggesting that the overall energetic costs were larger for females which produced more than twice as many clutches as males brooded in a season. Selective mortality of females could be due primarily to the depletion of their energy reserves. The increased mortality in females is a possible mechanism for the male-biased operational sex ratio which may provide the potential for sexual selection on males.     

Reproductive periodicity and steroid hormone profiles in the sex-changing coral-reef fish, <Emphasis Type="Italic">Plectropomus leopardus</Emphasis>

The reproductive biology of coral trout, Plectropomus leopardus, from the Great Barrier Reef (Australia) was investigated by correlating gonadal condition with plasma levels of gonadal steroids. Female fish were found to be regressed from mid-summer to early spring, after which rapid and cyclical increases in gonado-somatic index (I _G), maximum oocyte diameter (MOD) and plasma concentrations of estradiol-17β and testosterone were detected. Male fish, in contrast, commenced recrudescence slightly earlier in winter and responded with less dramatic increases in both I _G and plasma concentrations of testosterone and 11-ketotestosterone. The mode of oocyte development was multiple group-synchronous, and cyclical fluctuations in reproductive parameters (I _G, MOD and gonadal steroid concentrations) were synchronized with new-moon lunar phases. It is likely, therefore, that individual P. leopardus have the capacity to spawn on multiple occasions, with lunar periodicity. However, evidence suggests that early bouts of reproduction may be more important in terms of reproductive investment than subsequent bouts later in the same season. It is concluded that patterns of gametogenesis and steroidogenesis in P. leopardus are similar to the patterns displayed by other tropical groupers, suggesting that management regimes and propagation protocols developed for these fishes may also be appropriate for use with P. leopardus.