Male and female effects on sperm precedence in the giant sperm species <Emphasis Type="Italic">Drosophila bifurca</Emphasis>

Sperm competition is expected to be a driving force in sexual selection. In internally fertilized organisms, it occurs when ejaculates from more than one male are present simultaneously within the female’s reproductive tract. It has been suggested that greater sperm size may improve the competitive ability of sperm, but studies provide contradictory results depending on the species. More recently, the role of females in the evolution of sperm morphology has been pointed out. We investigate here the male and female effects that influence sperm precedence in the giant sperm species, Drosophila bifurca Patterson & Wheeler. Females were mated with two successive males, and the paternity outcomes for both males were analyzed after determining sperm transfer and storage. We found very high values of last male sperm precedence, suggesting a strong interaction between rival sperm. However, the data also indicate high frequencies of removal of the sperm of the first male from the female reproductive tract prior to any interaction with the second male. This implies that successful paternity depends mainly on successful sperm storage. Knowing what happens to the sperm within females appears to be a prerequisite for disentangling post-copulatory sexual interactions between males and females.     

AGE,GROWTH, AND REPRODUCTIVE PATTERNS OF DALL'S PORPOISE (PHOCOENOIDES DALLI) IN THE CENTRAL NORTH PACIFIC OCEAN

Life history parameters were estimated for Dall's porpoise, Phocoenoides dalli, from biological specimens collected in the western Aleutian Islands, during 1981–1987. Of 2,033 males and 3,566 females examined, reproductive data were available for 1,941 males and 1,906 females; ages were determined for 813 males and 1,297 females. Female sexual maturity was based on the presence of one or more corpus on either ovary; 845 were sexually immature and 1,061 were sexually mature. Two estimates of female average age at sexual maturity (ASM) were 3.8 and 4.4 yr; average length at sexual maturity (LSM) was 172 cm. Males were considered sexually mature when evidence of spermatogenesis was detected; 1,136 were sexually immature and 805 were sexually mature. Two estimates of male ASM were 4.5 and 5.0 yr; LSM was 179.7 cm. Physical maturity was assessed for 246 males and 446 females by examining the degree of fusion in thoracic vertebral epiphyses. For both sexes, the average age at physical maturity was 7.2 yr. Average length at physical maturity was 202.6 cm for males and 192.7 cm for females. Average lengths of physically mature males (x?= 198.1 cm, SE = 0.8566) and females (x?= 189.7 cm, SE = 0.4002) were significantly different(P < 0.0001). Early postnatal growth was rapid in both sexes. A secondary growth spurt in both mass and length was characteristic for both sexes; the increase in length preceded the mass increase by 1–2 yr. Average length at birth (LOB) was approximately 100 cm; birth mass averaged 11.3 kg (SE = 0.0772). By the time the umbilicus had healed (<2 mo), the average length and mass had increased to 114.1 cm and 23.8 kg, respectively. Gestation period based on projections using LOB was 12 mo, but this was considered an overestimate. Calving was modal, centered in early July; an annual reproductive interval was indicated. Among the sexually mature females, 120 were pregnant, 55 were pregnant and lactating, 321 were pregnant with colostrum, and 33 were “resting.” By 3 July (95% CI =x? 1 d), 50% of births had occurred, during each of the seven years sampled. The ovulation rate was estimated at 0.914 ovulations per average reproductive year. Enlarged follicles and recent ovulations were observed in postpartum females in late July.     

Sperm transfer and paternity in the scorpionfly <Emphasis Type="Italic">Panorpa cognata</Emphasis>: large variance in traits favoured by post-copulatory episodes of sexual selection

Post-copulatory episodes of sexual selection can be a powerful selective force influencing the reproductive success of males. In order to understand variation in male fertilisation success, we first need to consider the pattern of sperm utilisation by females following matings with more than one male. Second, we need to study those traits responsible for male success in sperm competition. Here we study both male sperm transfer characteristics as well as offspring paternity of females mated to two males in the scorpionfly Panorpa cognata. By repeatedly mating males to virgin females and interrupting copulation at defined time points, we found for all males that sperm transfer set off after approximately 40 min. During the remaining copulation, sperm transfer of individual males was continuous and with constant rate. Yet the rate of sperm transfer differed between individual males from about one sperm per minute to more than eight sperm per minute for the most successful males. In addition, we measured the fertilisation success in sperm competition of males with known sperm transfer capability. The relative number of sperm transferred by males during copulation, estimated from copulation duration and the males’ individual sperm transfer rate, explained a large proportion of variation in offspring paternity. The mode of sperm competition in this species, thus, conforms largely to a fair raffle following complete mixing of sperm prior to fertilisation. Hence, male differences in both the ability to copulate for long and of rapid sperm transfer will translate directly into differences in reproductive success.     

POLYGYNANDRY IN THE DUSKY PIPEFISH SYNGNATHUS FLORIDAE REVEALED BY MICROSATELLITE DNA MARKERS

In the dusky pipefish Syngnathus floridae, like other species in the family Syngnathidae, ‘pregnant’ males provide all post-zygotic care. Male pregnancy has interesting implications for sexual selection theory and the evolution of mating systems. Here, we employ microsatellite markers to describe the genetic mating system of S. floridae, compare the outcome with a previous report of genetic polyandry for the Gulf pipefish S. scovelli, and consider possible associations between the mating system and degree of sexual dimorphism in these species. Twenty-two pregnant male dusky pipefish from one locale in the northern Gulf of Mexico were analyzed genetically, together with subsamples of 42 embryos from each male's brood pouch. Adult females also were assayed. The genotypes observed in these samples document that cuckoldry by males did not occur; males often receive eggs from multiple females during the course of a pregnancy (six males had one mate each, 13 had two mates, and three had three mates); embryos from different females are segregated spatially within a male's brood pouch; and a female's clutch of eggs often is divided among more than one male. Thus, the genetic mating system of the dusky pipefish is best described as polygynandrous. The genetic results for S. floridae and S. scovelli are consistent with a simple model of sexual selection which predicts that for sex role-reversed organisms, species with greater degrees of sexual dimorphism are more highly polyandrous.     

Does sympatry predict life history and morphological diversification in the Mexican livebearing fish Poeciliopsis baenschi?

Understanding why some species coexist and others do not remains one of the fundamental challenges of ecology. Although there is evidence to suggest that closely‐related species are unlikely to occupy the same habitat because of competitive exclusion, there are many cases where closely‐related species do co‐occur. Research comparing sympatric and allopatric populations of co‐occurring species provides a framework for understanding the role of phenotypic diversification in species coexistence. In the present study, we compare phenotypic divergence between sympatric and allopatric populations of the livebearing fish, Poeciliopsis baenschi. We focus on life‐history traits and body shape, comprising two sets of integrated traits likely to diverge in response to varying selective pressures. Given that males and females can express different phenotypic traits, we also test for patterns of divergence among sexes by comparing size at maturity and sexual dimorphism in body shape between males and females in each population type. We take advantage of a natural experiment in western Mexico where, in some locations, P. baenschi co‐occur with a closely‐related species, Poeciliopsis turneri (sympatric populations) and, in other locations, they occur in isolation (allopatric populations). The results obtained in the present study show that sympatric populations of P. baenschi differed significantly in life‐history traits and in body shape compared to their allopatric counterparts. Additionally, males and females showed different responses for size at maturity in sympatric conditions versus allopatric conditions. However, the amount of sexual dimorphism did not differ between sympatric and allopatric populations of P. baenschi. Hence, we conclude that not all traits show similar levels of phenotypic divergence in response to sympatric conditions. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 608–618.     

Reproductive biology of brown trout, Salmo trutta macrostigma Dumeril 1858, in a tributary of the Ceyhan River which flows into the eastern Mediterranean Sea

The study describes some key elements of the reproductive biology, including spawning season, age at sexual maturity, fecundity and egg diameter of the native brown trout, Salmo trutta macrostigma, in a tributary of the Ceyhan River. A total of 197 brown trout (118 females and 79 males) were captured in 2000–2001 by electric fishing. In observations on monthly changes, the gonadosomatic index (GSI) and the monthly frequency distribution of egg diameter confirmed that spawning lasted from November to January. Some 27.7% of the females and 62.5% of the males attained sexual maturity in their second year. The smallest fork length (FL) of brown trout attaining sexual maturity was 17.4 cm for males and 17.8 cm for females. Mean fecundity in age groups II, III, IV and V were 360, 452, 693 and 1283 eggs per female, respectively. One 9‐year‐old female had a unique 3232 egg count. The mean fecundity of the sampled population was 554 eggs per fish, positively correlated with the FL (mm) (R = 0.8227 ) and body weight (R = 0.8130). The diameter of mature eggs in the spawning season ranged from 3.250 to 5.930 mm, with a 4.146 mm average. Mean egg diameter in age groups II, III, IV and V in the spawning season were 0.813, 3.799, 4.663 and 5.243 mm, respectively. Fecundity, egg weight and diameter were statistically different in all age groups.     

Population structure,spatial distribution and life‐history traits of blacktip reef sharks Carcharhinus melanopterus

During a survey of the population of blacktip reef shark Carcharhinus melanopterus in Moorea (French Polynesia) between 2007 and 2011, population structural characteristics were estimated from 268 individuals. Total length (L_T) ranged from 48 to 139 cm and 48 to 157 cm for males and females, respectively, demonstrating that the average L_T of females was larger than that of males. The C. melanopterus population at Moorea showed an apparent spatial sexual segregation with females preferentially frequenting lagoons and males the fore‐reefs. Mean growth rate was c. 6 cm year^?1. Males reached sexual maturity at 111 cm L_T. This study reports on the population characteristics of this widespread carcharhinid shark species and makes comparisons with other locations, confirming high geographic variability in the population structure of the species.     

Reproductive biology of gag in the southern Gulf of Mexico

Aspects of the reproductive biology of gag Mycteroperca microlepis in the southern Gulf of Mexico were studied by following seasonal variations in the gonado‐somatic index and through histological examination of gonads. Gag were collected from inshore and offshore waters of the Campeche Bank, Yucatan, Mexico, between April 1996 and December 2001. This species is a protogynous hermaphrodite, and appeared to be depth‐size distributed. The smallest gag (9–49 cm L _F) collected were all juvenile females, and were caught in inshore waters (1–10 m depth), while the largest (49–116 cm L _F), mainly adult females, males and transitionals, were captured in offshore waters (33–167 m depth). Overall the offshore male to female ratio was female‐biased (1 : 3·3) and differed significantly from unity. The species spawns at depths of c . 50–53 m, from early winter to mid‐spring, with peak spawning activity occurring between January and March. Fifty per cent of females reached first maturity at 72·1 cm L _F. At 103 cm L _F, 50% of sampled females had changed into males. Gag can be considered a monandric species, and sexual transition for this grouper seemed to occur in fish distributed within a narrow size range (85–111 cm L _F). The results are compared with those of other authors for gag stocks from the south‐east Atlantic coast of the U.S.A. and the north‐east Gulf of Mexico.     

Length–weight and length–length relationships,gonadosomatic indices and size at first maturity of Eugerres mexicanus (Steindachner, 1863) (Percoidei: Gerreidae) from the Usumacinta River,Mexico

This study examines the length‐weight (L‐W) and length‐length (L‐L) relationships, gonadosomatic indices, and size at sexual maturity for individuals of the endemic Mexican mojarra, Eugerres mexicanus (Steindachner, 1863) from the Usumacinta River Basin, Tabasco, Mexico. From April 2008 to January 2010, 360 specimens were examined, including 151 (41.9%) male, 179 (49.7%) female and 30 unsexed (8.3%) individuals. Overall female:male ratio was 1.2 : 1. Gonadosomatic index (GSI) values indicate increased reproductive activity during February, with significant differences between males (GSI = 2.07) and females (GSI = 5.48). The mean size at first maturity (L₅₀) was 17.3 cm TL for males and 20.5 cm TL for females.     

Variable mate-guarding time and sperm allocation by male snow crabs (Chionoecetes opilio) in response to sexual competition, and their impact on the mating success of females.

Two laboratory experiments investigated mate guarding and sperm allocation patterns of adult males with virgin females of the snow crab, Chionoecetes opilio, in relation to sex ratio. Although females outnumbered males in treatments, operational sex ratios were male-biased because females mature asynchronously and have a limited period of sexual attractiveness after their maturity molt. Males guarded females significantly longer as the sex ratio increased: the mean time per female was 2.9 d in a 2 males:20 females treatment compared to 5.6 d in a 6 males:20 females treatment. Female injury and mortality scaled positively to sex ratio. Males that guarded for the greatest number of days were significantly larger, and at experiment's end had significantly smaller vasa deferentia, suggesting greater sperm expense, than males that guarded for fewer days. In both experiments, the spermathecal load (SL)--that is, the quantity of ejaculate stored in a female's spermatheca--was independent of molt date, except in the most female-biased treatment, where it was negatively related. The SL increased as the sex ratio increased, mainly because females accumulated more ejaculates. However, similarly sized males had smaller vasa deferentia and passed smaller ejaculates, such that, at a given sex ratio, the mean SL was 55% less in one experiment than in the other. Some females extruded clutches with few or no fertilized eggs, and their median SL (3-4 mg) was one order of magnitude smaller than that of females with well-fertilized clutches (31-50 mg), indicating sperm limitation. Males economized sperm: all females irrespective of sex ratio were inseminated, but to a varying extent submaximally; each ejaculate represented less than 2.5% of male sperm reserves; and no male was fully exhausted of sperm. Sperm economy is predicted by sperm competition theory for species like snow crab in which polyandry exists, mechanisms of last-male sperm precedence are effective, and the probability that one male fertilizes a female's lifetime production of eggs is small.     

Multiple paternity in a natural population of a salamander with long-term sperm storage

Sperm competition appears to be an important aspect of any mating system in which individual female organisms mate with multiple males and store sperm. Post-copulatory sexual selection may be particularly important in species that store sperm throughout long breeding seasons, because the lengthy storage period may permit extensive interactions among rival sperm. Few studies have addressed the potential for sperm competition in species exhibiting prolonged sperm storage. We used microsatellite markers to examine offspring paternity in field-collected clutches of the Ocoee salamander (Desmognathus ocoee), a species in which female organisms store sperm for up to 9 months prior to fertilization. We found that 96% of clutches were sired by multiple males, but that the majority of females used sperm from only two or three males to fertilize their eggs. The high rate of multiple mating by females suggests that sperm competition is an important aspect of this mating system. Comparison of our data with those of other parentage studies in salamanders and newts reveals that multiple mating may be common in urodele amphibians. Nevertheless, the number of males siring offspring per clutch in D. ocoee did not differ appreciably from that in other species of urodeles with shorter storage periods.     

Biometric and reproductive aspects of the pen shell Atrina seminuda (Bivalvia: Pinnidae) in northeastern Venezuela

The pen shell Atrina seminuda is a target for a small artisanal Caribbean fishery; however, little is known about its biology. The length frequency distribution, biometric relations and reproduction of this species were studied in northeastern Venezuela. Using transects selected from among 360 stations a population was sampled monthly by SCUBA diving. An analysis of 1748 individuals showed a shell height (SH) between 4?cm and 23.6?cm, with an average of 16.7?±?2.5?cm. For both sexes, statistical differences in mean SH were observed (analysis of variance F (1, 1113)?=?69.538; P?SL), total weight (TW) and adductor muscle weight (MW) as a function of SH. In all cases there were statistical differences between males and females, with males presenting positive allometry and females showing isometry. Mean SH at maturity for males (8.24?cm) was lower than for females (11.88?cm), and for sexes combined was estimated at 8.71?cm. Gametogenic activity was observed throughout the year, with two main peaks of spawning (April and August/September 2008). All this information is crucial for demographic modelling and stock assessment for managing this species.     

MEASURING SELECTION ON A POPULATION OF DAMSELFLIES WITH A MANIPULATED PHENOTYPE

The estimation of the relationship between phenotype and fitness in natural populations is constrained by the distribution of phenotypes available for selection to act on. Because selection is blind to the underlying genotype, a more variable phenotypic distribution created by using environmental effects can be used to enhance the power of a selection study. I measured selection on a population of adult damselflies (Enallagma boreale) whose phenotype had been modified by raising the larvae under various levels of food availability and density. Selection on body size (combination of skeletal and mass at emergence) and date of emergence was estimated in two consecutive episodes. The first episode was survival from emergence to sexual maturity and the second was reproductive success after attaining sexual maturity. Female survival to sexual maturity was lower, and therefore opportunity for selection greater, than males in both years. Opportunity for selection due to reproductive success was greater for males. The total opportunity for selection was greater for males one year and for females the other. Survival to sexual maturity was related to mass gain between emergence and sexual maturity. Females gained more mass and survived less well than males in both years but there was no linear relationship between size at emergence and survival for females in either year. However, females in the tails of the phenotype distribution were less likely to survive than those near the mean. In contrast, small males consistently gained more mass than large males and survived less well in one year. There was significant selection on timing of emergence in both years, but the direction of selection changed due to differences in weather; early emerging females were more successful one year and late emerging males and females the other. The number of clutches laid by females was independent of body size. Because the resources used to produce eggs are acquired after emergence and this was independent of size at emergence, female fitness did not increase with size. Small males may have had lower survival to sexual maturity but they had higher mating success than large males. Resources acquired prior to sexual maturity are essential for reproductive success and may in some species alter their success in inter- and intrasexual competition. Therefore, ignoring the mortality associated with resource acquisition will give an incomplete and potentially misleading picture of selection on the phenotype.     

Females Mated without Sperm Transfer Maintain High Sexual Receptivity in the Wolf Spider Schizocosa malitiosa

The mating of the Neotropical lycosid Schizocosa malitiosa is long and complex, involving intense genital stimulation and copulatory courtship. This suggests functions other than just insemination. Previous data indicated that mated females of this species are less sexually receptive than virgins. We hypothesise that copulatory characteristics presented by males during prolonged copulations could be responsible for subsequent female sexual reluctance, and may be selected by cryptic female choice. Our objective was to examine the influence of copulatory behaviour on subsequent female sexual receptivity in S. malitiosa, isolating it from the effects of sperm transfer per se. For this purpose, we obtained males without sperm in their copulatory organs (palpal bulbs), and prevented them from charging their palps by sealing their genital pores immediately after their last moult (treated males). Virgin females were separated into three groups: (i) females exposed once to normal males, (ii) females exposed twice to normal males, and (iii) females exposed first to treated and second to normal males. The results showed that, 3 d after their first mating, females first mated with untreated males were frequently refractory to remating, whereas all those first mated with treated males were receptive. Copulations performed by treated males showed some differences from those performed by normal males, but maintained the basic behavioural pattern with abundant sexual stimulation. The presence of sperm fluids in the female receptacles appears to be the most likely factor generating female remating reluctance. Males may manipulate female responses using receptivity inhibiting substances in their sperm, like those described for insects. Females would first ensure sperm supply, becoming more choosy afterwards. All females, whether mated once or twice, generated similar numbers of progeny, indicating no relationship between number of matings and number of spiderlings.     

Reproductive characteristics and reproductive tract anatomy of the California butterfly ray Gymnura marmorata (Myliobatiformes: Gymnuridae)

The California butterfly ray Gymnura marmorata is frequently caught by artisanal fisheries in north-western Mexico (Baja California Sur; BCS). The aim of this study was to estimate reproductive characteristics and describe the micro and macroscopic reproductive structures of female and male G. marmorata. Sampling was carried out at two fisheries landing sites located on the west coast BCS, from 2008 to 2012. The largest disc width (W_D) ever recorded for this species was 131.0 cm and females were larger than males. The male sample was composed exclusively of adult specimens, with a median size at 50% maturity (W_D50) estimated at 44.9 cm W_D. For females, size at onset of maturity was estimated at 63.0 cm W_D; W_D50, at 64.7 cm W_D; and size at pregnancy at 80.1 cm W_D. Gonadal asymmetry was observed for the first time in this species. Indeed, in males only the left testis was functional (testicular asymmetry), while in females, only the left ovary was functional (ovarian asymmetry).     

Reproductive biology of Bigeye Tuna, <Emphasis Type="Italic">Thunnus obesus</Emphasis>, (Scombridae) in the eastern and central tropical Pacific Ocean

Understanding the reproductive potential of any species is of great importance for resource assessment and management. We studied the reproductive biology of Bigeye Tuna, Thunnus obesus, based on 1283 samples taken from the Chinese longline vessels in the eastern and central Tropical Pacific Ocean during February through November 2006. The female-male ratio was 1.0 : 1.5 and males were predominant in all length classes except for the length class of 166–170 cm (fork length). Males dominated in sizes larger than 171 cm, all specimens of 192 cm or larger were males. The main spawning period of Bigeye Tuna was between March and November. Gonadosomatic rates of males were larger than those of females. Statistically, female and male Bigeye Tuna had no significant reproductive seasonality. The observed minimum length at sexual maturity for female Bigeye Tuna was 94 cm. Length at 50% sexual maturity of female Bigeye Tuna was estimated at 107.8 cm, and maturation rate was 0.106 cm⁻¹. The results derived in this study provide the information critical to our understanding of key life history parameters of Bigeye Tuna in tropic Pacific Ocean.     

Gonadosomatic index‐based size at first sexual maturity of the catfish Eutropiichthys vacha (Hamilton, 1822) in the Ganges River (NW Bangladesh)

This study describes the size at first sexual maturity, length–weight relationships (LWR) in relation to size at first sexual maturity, and Fulton’s condition factor (K_F) of Eutropiichthys vacha in the Ganges River, northwestern Bangladesh. Sampling was done using traditional fishing gear including cast nets, square lift nets and conical traps during January and April, and July to December 2010. For each individual, total length (TL) was measured to the nearest 0.01 cm, and total weight (BW) was determined to the nearest 0.01 g. The gonadosomatic index (GSI) was calculated by the equation, GSI (%) = (Gonad weight in g/BW) × 100. The size at first sexual maturity of males and females was estimated by the relationship between gonadosomatic index and total length. A total of 583 specimens (289 males; 294 females) ranging from 8.30 to 27.00 cm TL and 3.16 to 159.50 g BW were analyzed. Sizes at first sexual maturity for male and female E. vacha were 13.15 and 14.00 cm TL, respectively. The analysis of covariance (ancova ) revealed significant differences in slope and intercept between early and late phases for males (F = 4.532, P < 0.001) and females (F = 21.984, P < 0.001). The K_F was not significantly correlated with TL for males (r_s = 0.052; P = 0.378), but was highly correlated for females (r_s = ?0.165; P = 0.005). This study establishes a strong base for monitoring changes in length at first sexual maturity attributable to high fishing pressures or other reasons within the Ganges and associated river ecosystems.