Seasonal Variations and Diurnal Rhythms of Gonadotropin (GtH2) Levels in Immature and Mature Female Carp (Cyprinus carpio L)

Daily changes of gonadotropin (GtH2) blood levels of immature and mature carp females were investigated in four seasons during the year: spring, summer (spawning period) and in autumn and winter (“resting” period). GtH2 levels were measured using ELISA method. The maturity stage of oocytes was determined histologically and on the base of germinal vesicle position in the oocyte. It appeared shown that GtH2 levels changed daily during the year in the range of 3.56 to 6.61 ng/ml, regardless of the state of ovarian maturity. Cosinor analysis and ellipse of errors revealed the presence of a circadian rhythm in GtH2 levels only in mature females during the reproduction period (L:D=16:8, water temp. 20 to 22°C).     

Size composition and reproductive cycle of Pseudupeneus grandisquamis (Pisces: Mullidae) in the Central Mexican Pacific

Size composition and reproductive cycle of Pseudupeneus grandisquamis (Pisces: Mullidae) in the Central Mexican Pacific. From June 1995 to December 1998 we used shrimp trawl nets to capture 492 Pseudupeneus grandisquamis in soft-bottom grounds off the central Mexican Pacific (Jalisco and Colima). The ength-weight ratio indicates allometric growth (p = 0.0035L(3.46) y r2 = 0.97). The total sex ratio was 1:1.15 (females: males). Mean total length was 153.8 mm (range 77-236 mm). A progression in length was found: the highest means were observed at the end of 1996 and during 1997. The mean length of females (162 mm) was longer than in males (150 mm). Four gonadic maturity stages were observed in both sexes; in females the most frequently stage was stage IV (mature gonads, 48.6%), while in males it was stage III (gonads in maturation, 45.0%). Seven microscopic oocyte development stages were identified. Oocyte development seems to be asynchronous. The mature testicle showed sperm inside the lobular lumen and in the duct. The highest values of the gonadosomatic index, as well as the largest percentages of individuals with mature gonads (at macroscopic and microscopic levels), were observed during winter and summer, suggesting that this species has two reproduction peaks by year. The length at which 50% of the individuals show gonads in maturation (L50) was 183 mm for the females and 181 mm for the males.     

Maturity Ogives for South Pacific Albacore Tuna (Thunnus alalunga) That Account for Spatial and Seasonal Variation in the Distributions of Mature and Immature Fish

Length and age at maturity are important life history parameters for estimating spawning stock biomass and reproductive potential of fish stocks. Bias in estimates of size and age at maturity can arise when disparate distributions of mature and immature fish within a population are not accounted for in the analysis. Here we investigate the spatial and temporal variability in observed size and age at maturity of female albacore tuna, Thunnus alalunga, using samples collected across the South Pacific. Maturity status was identified using consistent histological criteria that were precise enough to allow for mature but regenerating females to be distinguished from immature females during the non-spawning season, permitting year-round sampling for maturity estimation in albacore. Using generalised linear mixed models, we found that the proportion of mature females at length varied significantly with latitude and time of year. Specifically, females at northern latitudes (∼10–20°S, where spawning occurs) were mature at significantly smaller lengths and ages than females at southern latitudes (∼20–40°S), particularly during the spawning season (October–March). This variation was due to different geographic distributions of mature and immature fish during the year. We present a method for estimating an unbiased maturity ogive that takes into account the latitudinal variation in proportion mature at length during a given season (spawning or non-spawning). Applying this method to albacore samples from the western region of the South Pacific gave a predicted length at 50% mature of ∼87 cm fork length (4.5 years).     

Plasma and pituitary concentrations of gonadotropins (FSH and LH) in minke whales (Balaenoptera acutorostrata) during the feeding season

This study investigated plasma and pituitary concentrations of follicle-stimulating hormone (FSH) and luteinizing hormone (LH) and steroid hormones (progesterone: P4, testosterone:T, estradiol-17beta: E2) by enzyme-immunoassay (EIA) in minke whales (Balaenoptera acutorostrata) captured during the feeding season (December to March) in the Antarctic Ocean. Plasma FSH and LH levels in female minke whales were higher (P <0.05) than in male whales. Although the pituitary weight was not significantly different between male and female whales, pituitary FSH and LH levels were higher in females than in males (P<0.01) and mature whales than immature whales (P<0.05). Plasma levels of FSH, T and E2 were not significantly different between immature and mature male whales, but plasma LH and pituitary FSH and LH levels were higher (P<0.05) in mature than in immature whales. In both immature and mature whales regardless of gender, pituitary FSH and LH levels were correlated significantly (r=0.69: P<0.01). In mature male whales, plasma T and E2 levels (r=0.60: P<0.01), and testis weight and plasma T levels (r=0.46: P <0.05) were correlated. In immature female whales, plasma FSH and LH levels were highly correlated (r=0.68: P<0.001), but were not for mature female whales. The results show that gender and maturity influence gonadal and pituitary function of minke whales during the feeding season.     

Seasonal effects of melatonin on ovary and plasma gonadotropin and vitellogenin levels in intact and pinealectomized catfish, Clarias batrachus (Linn)

Effects of daily administration of melatonin for 15 days were evaluated with respect to ovarian activities and plasma gonadotropin (GtH II) and vitellogenin (Vg) levels in intact (INT) and pinealectomized (Px) female catfish, C. batrachus, during preparatory (April), prespawning (May and June), spawning (July) and post-spawning (September) periods. Px (saline control groups) caused a stimulatory effect during preparatory (with respect to Vg synthesis and incorporation) and prespawning (with respect to Vg synthesis) periods whereas no effect was observed during spawning and post-spawning periods with respect to the reproductive parameters studied. During April, melatonin-treatment significantly decreased plasma GtH II levels and percentage of vitellogenic oocytes without any significant changes in plasma Vg levels and gonadosomatic index (GSI). During early prespawning period, in May, 50microg melatonin brought about a significant reduction in plasma GtH II levels in INT group, whereas 100microg caused a decrease in all parameters; on the other hand, in Px groups both dose levels proved to be inhibitory. In June (late prespawning period) melatonin-treatment could not bring about any change in GSI and plasma Vg levels compared to the control groups regardless of Px but plasma GtH II and mean number of yolky oocytes were significantly reduced in melatonin-treated INT group. During spawning period (July) melatonin inhibited the GSI, mean number of yolky oocytes and plasma GtH II levels without affecting plasma Vg levels. In September (post-spawning period), melatonin did inhibit both GSI and plasma GtH II levels. The results, thus, indicate that melatonin showed variable effects (inhibitory and/or no effect) to GSI, mean number of yolky oocytes and plasma Vg levels but a consistent inhibiton of plasma GtH II levels indicating that melatonin may control the reproduction by blocking the GtH II release from the pituitary via affecting the hypothalamo-hypophysial axis.     

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.     

多巴胺能药物对鲤鱼促性腺激素分泌活动的影响

研究多巴胺（ＤＡ）及其激动剂阿扑吗啡（ＡＰＯ）和拮抗剂Ｄｏｍｐｅｒｉｄｏｎｅ（ＤＯＭ）对不同性腺发育时期和不同年龄雌鲤促性腺激素（ＧｔＨ）分泌活动的影响。结果表明：鲤鱼基础ＧｔＨ分泌具有明显的季节性变化,表现为血清ＧｔＨ含量在性腺发育早期较低,性腺成熟期升高,到性腺退化期又降低。ＤＡ和ＡＰＯ对ＧｔＨ分泌的抑制作用及ＤＯＭ对ＧｔＨ分泌的刺激作用在不同性腺发育时期为：性腺发育早期〉性腺成熟期〉性腺退化     

Spawning seasonality and body sizes at sexual maturity in the bluespine unicornfish,Naso unicornis (Acanthuridae)

We herein evaluate several reproductive metrics of Hawaiian Archipelagic populations of the bluespine unicornfish (Naso unicornis), an economically and ecologically important, broadly distributed tropical Pacific reef fish, based on multi-year, fishery-dependent and fishery-independent collections. Sex-specific spawning seasonality was characterized for fish collected mostly from Oahu (Main Hawaiian Islands, MHI) using a gonadosomatic index. Histological slides preparations were used to score gonad developmental phase and to classify individuals of either sex as immature or mature. Sex-specific median body lengths at maturity (L₅₀) were estimated by logistic fits of proportion mature versus length class. Spawning was highly seasonal in Hawaii, with a single brief (May–June) peak spawning period. Proportionate gonad-to-body weight values were relatively low, averaging only about 0.1 % and 0.6 % across all months of year and 0.16 % and 1.03 % during May–June for males and females, respectively. Median lengths at sexual maturity differed between the sexes. L₅₀ values for fish collected throughout all months of year were 30.1 ± 0.5 (standard error) cm Fork Length (FL) for males and 35.5 ± 0.7 cm FL for females. Spawning seasonality and L₅₀ estimates for bluespine unicornfish in Hawaii suggest that the species spawns several months earlier in the calendar year and matures at larger body lengths in Hawaii versus Guam, the Northern Mariana Islands, and Pohnpei in the Federated States of Micronesia. Estimated lengths at sexual maturity are compared to the minimum length (14 inches or 35.6 cm FL) mandated for this species in Hawaii: median size at maturity occurs at a length appreciably less than (males) or approximately equal to (females) minimum legal size. A likely disproportionately large contribution of old females to population replenishment is discussed relative to the minimum size limit.     

Reproductive biology of the prawn Melicertus kerathurus (Decapoda: Penaeidae) in Thermaikos Gulf (N. Aegean Sea)

The reproductive biology including insemination frequency, ovarian maturation, gonadosomatic index (GSI), size at first maturity, and fecundity of Melicertus kerathurus were investigated using monthly samples from Thermaikos Gulf. Insemination was recorded by the presence of a spermatophore in the thelycum, and ovarian development was based on macroscopic ovarian staging (ST1–ST5). Inseminated females were found throughout the year with high percentages recorded from April to July. Although all ovarian stages were represented in mated females, insemination increased with size and ovarian maturation. High percentages of vitellogenic or mature ovarian stages were observed from May to July, while immature and developing ovaries were predominant mainly in winter. Spawned ovaries occurred from May to October. Carapace length at first maturity based on the presence of a spermatophore (CL_50sp) was estimated at 39.20 mm, while that based on the presence of vitellogenic and mature ovaries (CL_50ov) at 40.70 mm. The seasonal peak in the proportion of mature females (ST4) varied with size. Inseminated females at ST4 and GSI peaked in June–July. GSI varied in relation to insemination status and ovarian stage. In large females (>50 mm CL), the decline in mature ovaries and GSI increment with size indicates a relative reduction in the reproductive output. The number of oocytes ranged from 62,742 to 602,947 (mean ± SD: 268,000 ± 113,000). As the prawns are targeted during the spawning season, mainly by the artisanal fishery, and female size at first maturity is selected by artisanal net size, managerial measures toward artisanal fishery should be implemented.     

Serum haptoglobin concentrations in ringed seals (Pusa hispida) from Svalbard, Norway

Haptoglobin (Hp) levels were measured in blood serum from 185 apparently healthy ringed seals (Pusa hispida) from Svalbard (age range, 1-30 yr) collected during the spring seasons of 2002 through 2004. The Hp concentration was 0.26 +/- 0.02 g/l (mean +/- SE; range, 0.0-1.5 g/l). Maturity status, body condition index (CI), interactions between CI and maturity status, and sex and maturity status all had significant influences on Hp levels. Mature males had significantly higher Hp concentrations compared with mature females (0.30 +/- 0.03 vs. 0.17 +/- 0.01 g/l, F(1,16) = 14.9, P<0.01). No differences were found between Hp levels in immature male and female seals (0.33 +/- 0.06 g/l). The Hp levels increased significantly in immature seals when the CI decreased. Despite the fact that mature males had a significantly lower CI than mature females, no direct correlation was found between Hp concentration and CI among adults. The elevated Hp values found in mature males might be related to injuries inflicted among males while fighting during the mating season or to high stress levels related to mating competition. The Hp concentrations of ringed seals measured in this study are useful baseline data for development of a diagnostic tool for future monitoring of the general health of this and other ringed seal populations.     

Gonadal development and plasma steroid levels during pubertal development in captive-reared striped bass, Morone saxatilis

Puberty is the period during which full sexual development occurs and the capacity to reproduce is acquired. Despite its importance, our understanding of the endocrine regulation of puberty in lower vertebrates is still limited. The objective of the present study was to describe the changes in gonadal development and plasma steroid levels in a relatively late maturing species, the striped bass, during the first four years of life. In about 65% of the females, puberty was initiated during the third year. Although gonadosomatic index (GSI) and oocyte diameter increased during this year, this first cycle was characterized by a heterogeneous population of developing oocytes, a relatively low mean maximum oocyte diameter, and an absence of yolk granules in the oocytes. Plasma 17beta-estradiol (E(2)) levels were low in all three-year-old fish, suggesting that an insufficient stimulation of vitellogenin production by E(2) may underlie the lack of vitellogenin incorporation into developing oocytes. All monitored parameters increased during the fourth year, but were still below the values attained by older females. In about 60% of the males, puberty was initiated during the first year and all males were mature by the third year. During the first two years, several immature males initiated spermatogenesis without reaching full maturity. In mature males, mean GSI, plasma testosterone, and 11-ketotestosterone levels increased simultaneously, reaching higher values each subsequent year. Our results indicate that, similar to the situation in mammals, more than one reproductive cycle is required in striped bass before complete adulthood is reached.     

The population ecology of gammarus tigrinus (sexton) in the reed beds of the Tjeukemeer

The population ecology of Gammarus tigrinus (Sexton) was studied in the Tjeukemeer during 1969 and 1970. G. tigrinus reaches very high densities — up to 24,000/m² in parts of the study area. In 1970, the summer densities were 2–21/2 times greater than in 1969. Individuals do not grow to such large sizes in the summer as at other times of the year. Females begin to carry eggs in March or April and reproduction ceases in November. Large females have larger broods than smaller animals and the average size of the brood varies with the time of year. The egg incubation period and growth rate are dependent upon temperature. At summer temperatures females became sexually mature after about four weeks and the egg incubation period is about io days. The entire population is turned over about three times during the year. A combination of rapid growth rate, early onset of sexual maturity and high fecundity are probably responsible for the rapid spread of G. tigrinus throughout much of the Netherlands.     

Differing behavioral and endocrinological effects of two female sex pheromones on male goldfish

Ovulatory female goldfish sequentially release at least two sex pheromones: 17 alpha, 20 beta-dihydroxy-4-pregnen-3-one (17,20 beta P) and a mixture of F prostaglandins (PGFs). This study sought to determine whether these pheromones have different endocrinological and behavioral actions and whether the PGF pheromone, which is released by spawning females, is responsible for increasing the gonadotropin (GtH) and milt (sperm and seminal fluid) levels of spawning males. Grouped and isolated males were exposed to combinations of these pheromones, food odor, and spawning and nonspawning females. 17,20 beta P stimulated GtH increases in both grouped and isolated males but had only minor effects on behavior; because its principal function appears to be physiological it may be considered a "primer" pheromone. In contrast, exposue to the PGFs elicited large increases in sexual behavior but increased GtH only when fish were exposed as groups; this pheromone's principal action appears to be behavioral and it should be considered a "releaser" pheromone. Although males had increased GtH and milt levels after 1 hr of spawning, males allowed to interact with nonspawning females also had elevated GtH; thus, behavioral interactions appear capable of elevating GtH in the absence of either pheromone. The existence of an independent behavioral mechanism which stimulates GtH was supported by the fact that males exposed to 17,20 beta P while spawning had GtH levels much greater than males exposed to only one of these stimuli.     

Reversible reproductive control in harbour seals (Phoca vitulina) with a gonadotropin-releasing hormone agonist

Reproductive control in captive pinnipeds is an important management subject for many facilities. To date reproductive control in harbour seals (Phoca vitulina) has been achieved using anti-androgens, progestagen preparations, castration, and physical separation of the sexes. The harbour seal group at the seal station in Friedrichskoog, Germany consists of three mature females (all >10 years), one older mature male (13 years of age in 2000) and one male who reached maturity during the study (3 years of age in 2000). In 2000 the older mature male received for the first time a 3-month depot injection of a gonadotropin-releasing hormone agonist (buserelin acetetate, 9.9 mg) by subcutaneous injection. This male was subsequently given the gonadotropin-releasing hormone agonist in 2001, 2004 and 2005. The younger male reached maturity during the investigation and received burserelin for the first time in 2004 and again in 2005. No pups were born in 2001, 2002, 2005 or 2006. No reproductive control was performed in 2002 and 2003, resulting in three newborns in 2003 and 2004. Serum levels of testosterone were measured by a routine liquid chromatography coupled with mass spectometry. Pre-burserelin testosterone levels varied between 0.02 and 2.18 ng/ml. Post-burserelin levels were under the detection limit except for the first year of the investigation. No behavioural changes such as changes in social ranking and no clinical side effects were observed. This study shows that the gonadotropin-releasing hormone agonist, burserelin acetate, can be used for reversible reproductive control in harbour seals without observed side effects or detrimental behavioural changes.     

Reproductive biology of the guitarfish <Emphasis Type="Italic">Rhinobatos hynnicephalus</Emphasis> (Batoidea: Rhinobatidae) in Ariake Bay,Japan

Size at sexual maturity, reproductive cycle, and fecundity of the guitarfish, Rhinobatos hynnicephalus, were examined in Ariake Bay, Japan. Females reached sexual maturity at a larger size than males [total length (TL) at 50% sexual maturity: males, 431 mm; females, 476 mm]. Monthly gonadosomatic indices of males decreased abruptly from July to August. Histological examinations confirmed the presence of mature spermatozoa in the testes in July, with semen in seminal vesicles in July and August. Preovulatory ova were observed in females with near-term embryos in August. Parturition occurred in August, immediately followed by mating, ovulation, and fertilization. Gestation period is approximately 1 year. Fertilized uterine eggs without embryonic development were present throughout the annual reproductive cycle. Embryonic development began in June and ended in August, indicating that R. hynnicephalus undergoes embryonic diapause (9 months). Fecundity increased with TL and ranged from 1 to 9 (mean, 4.4) embryos per litter.     

Selectivity Parameters and Size at First Maturity in Deep-Water Shrimps, Aristaeomorpha foliacea (Risso, 1827) and Aristeus antennatus (Risso, 1816), from the North-Western Ionian Sea (Mediterranean Sea)

Selectivity experiments were carried out during trawling targeting deep-water shrimps Aristaeomorpha foliacea (Risso, 1827) and Aristeus antennatus (Risso, 1816) (Crustacea, Decapoda, Aristeidae) in the North-Western Ionian Sea (Eastern-Central Mediterranean). Different criteria were employed to analyse maturity; however, the proportion at 50% of retained, mated and mature specimens was always used to indicate the size, expressed as Carapace Length (CL, mm), at first capture (CL_c), mating (CL_sp) and at first maturity (CL_m), respectively. In order to estimate the size at 50% maturity (CL_m) for females of both species, three criteria were adopted. In particular, CL_m was computed for the mature females not considering the presence of spermatophores, for the mature females with spermatophores and for the mature females intersected by the decreasing proportion with size of females without spermatophores. Three diamond stretched mesh codends of 40, 50 and 60 mm were tested using a cover of 20 mm. The 40-mm stretched mesh size (European Union legal size in the Mediterranean) was not selective for the sampled population of each species. The size at first capture (CL_c), calculated in both species for the two sexes combined, increased significantly with mesh size. Even for the mesh size of 60 mm, the size at first capture was still smaller than the sizes at 50% maturity, whatever the criterion adopted. Since the differences between the size at first maturity and the sizes at first capture are greater in A. foliacea than A. antennatus, the former species appears in this respect to be more vulnerable to trawling than the latter.     

Effects of size and early sexual maturity on downstream migration during smolting in Baltic samon (Salmo salar)

The effects of size and previous sexual maturity on downstream migration in two-summer-old Baltic salmon were studied during their normal seaward migratory period in spring. We used an automatic PIT (Passive Integrated Transponder) tag monitoring system for individual recognition of tagged fish. Size affected the probability of migration among both previously immature and previously mature males while there was no such relationship among females. Nearly all females and 63.6% of the previously immature males migrated during the smolting period, while only 24.5% of the previously mature males did. In migrating fish the individual activity level increased from almost zero in May to about 450 antenna passages per day in mid June, coinciding with a pronounced increase in ambient water temperature. During the period of increasing activity in early June the fish changed from night activity to day activity and finally activity all day and night. The fish moved downstream in schools but no specific school size dominated. This paper demonstrates the strong effect of body size and previous sexual maturity on the probability of migration.     

A presumptive role for GABA in the stimulatory effects of Des-Gly10, [D-Ala6]-LHRH-ethylamide and pimozide on the gonadotropin release in carp

To investigate the effect of endogenous gamma-aminobutyric acid (GABA) on the blood maturating gonadotropin (GtH) levels, or to study its interaction with pimozide (dopamine antagonist) and a luteinizing hormone-releasing hormone analog (LHRH-a), sexually mature male and female carps were treated with drugs that may either inhibit GABA biosynthesis or GABA degradation. In females the irreversible inhibitor of GABA-transaminase, gamma-vinyl GABA (GVG), which was to increase the endogenous GABA-ergic tone, had no influence on GtH release. On the other hand, the increased GtH response to the combination of pimozide (PIM) and LHRH-a was clearly enhanced by the administration of 3-mercaptopropionic acid (MPA), an inhibitor of the rate limiting enzyme of GABA-biosynthesis. In males the GABA-ergic compound, valproic acid (DPA) decreased LHRH-a stimulated GtH levels. In male carps that received PIM to diminish the dopaminergic inhibition of GtH release, the spermiating response to LHRH-a was increased by administration of MPA. These data suggest that GABA interacts with the action of dopamine and the gonadotropin releasing hormone (GnRH) on the release of GtH.     

Reproduction in Bagre marinus (Ariidae) off Pernambuco, northeastern Brazil

Throughout 1997, the catches of artisanal gillnetters working off the coast of northeastern Brazil were sampled monthly for Bagre marinus (Mitchill 1815). Significantly more females (n = 207) than males (n = 82) were caught, although there was no significant difference in their size compositions (21–47 cm fork length, FL). All males sampled (21–40 cm FL) had developed gonads and were classified as sexually mature. According to macroscopic and microscopic examination of their reproductive tract, females were separated into four reproductive stages (immature, maturing, mature, and resting). Size at 50% sexual maturity for females was estimated to be 33 cm FL. A positive linear relationship was detected between the size of mature females and their fecundity (between 11 and 32 oocytes). Clear reproductive progress indicated a spawning period between March and May. We conclude that further fishery‐independent data are required to determine patterns of male abundances and distributions.