Evolutionary and climatic factors affecting tooth size in the red foxVulpes vulpes in the Holarctic

Research into the geographical pattern of tooth size in the red fox,Vulpes vulpes (Linnaeus, 1758) in the Holarctic was conducted on a sample of 3806 skulls belonging to 41 fox populations. The Nearctic was represented by 948 specimens (249 females, 359 males, 340 specimens of unknown sex) belonging to 13 populations, whereas the Palearctic was represented by 2858 red foxes (1034 females, 1256 males, 568 specimens of unknown sex) from 32 populations. In the Nearctic, the largest foxes live on Kodiak Island (V. v. harrimani) and the Kenai Peninsula (V. v. kenaiensis), while the smallest ones live in California (V. v. necator) and Georgia (V. v. fulvus). In the Palearctic, the largest foxes come from the Far East (V. v. jakutensis, V. v. beringiana, V. v. tobolica), while the smallest are from the southern borders of the Eurasian range (V. v. pusilla, V. v. barbara, V. v. arabica). In both the Palearctic and Nearctic, tooth size in the fox varies depending on the geo-climatic factors. The fox’s tooth size confirms the general basis of Bergmann’s rule. In the Palearctic, specimens with larger teeth occur in cooler habitats with greater seasonality. These are first and foremost Northern and Far Eastern populations. In the Nearctic, tooth size in red foxes depends on the temperature and humidity of their habitat. Competition within the species and between species has important impact on the variation and dimorphism of tooth size in the red fox. Both in the Nearctic and Palearctic, red foxes from regions of sympatric co-occurrence with other closely relatedVulpes species, are more sexually dimorphic in terms of tooth size than red foxes from allopatric regions. Analysis of morphological distance on the basis of the size of dental characteristics shows, that in the Palearctic, the foxes from India (V. v. pusilla), while in the Nearctic, the population from Kodiak Island (V. v. harrimani) are most distant from the remaining populations. Geographic barriers such as the Bering Strait, Parry Channel, Mackenzie River, Kolyma and Omolon River systems have had a critical impact on red fox evolution. The most likely place for the evolution and diversification of the phyletic lineVulpes vulpes seems to be the Middle East region.     

GROWTH AND SEXUAL DIMORPHISM OF ATLANTIC WALRUSES (ODOBENUS ROSMARUS ROSMARUS) IN FOXE BASIN, NORTHWEST TERRITORIES, CANADA

Growth of Atlantic walruses ( Odobenus rosmarus rosmarus ) was investigated using morphological data collected in association with Inuit subsistence walrus hunts. Four growth models were examined. The growth parameters of a constrained Richards model were used to quantify growth and to test for sexual dimorphism. The asymptotic length of male walruses (315.2 cm ± 3.8 (SE), n = 103) was significantly larger ( t = 7.21, df = 191, P < 0.05) than the asymptotic length of females (276.6 cm ± 3.4, n = 90). Sexual size dimorphism in adults was due to a longer growth period and a faster growth rate in males. The predictive equation relating mass ( M , kg) to standard length ( SL , cm) was: Log₁₀ M = -3.74 + 2.68(Log₁₀ SL ), n = 25, r ²= 0.98. There were no significant differences in the size of male walruses from Foxe Basin collected in the 1950s and this study. There were too few data to compare females. There were no significant differences in size between walruses sampled in Greenland and Foxe Basin in the 1980s and 1990s. Foxe Basin walruses were significantly larger than walruses sampled in northern Hudson Bay in the 1950s. Female Atlantic walruses sampled in Foxe Basin were larger than female Pacific walruses ( Odobenus rosmarus divergens ) sampled in Alaska.     

Reproduction in the dusky grouper from the southern Mediterranean

Demographic data and gonad histology confirmed that the dusky grouper Epinephelus marginatus is a protogynous hermaphrodite that follows a monandric pathway to sexual development. Females reached first sexual maturity at 36·7 cm L_s and estimated mean length at first maturity (L₅₀) was 43·8 cm L_s for females and 81·3 cm L_s for males. Adult sex ratios during the reproductive period were c. 3·5: 1 females to males. Females exhibited group-synchronous ovarian development and multiple ovulation occurred over the spawning period. Gonads were ripe from early May and spawning occurred from June until early September. The size of ripe testes (0·6% W )indicated strong oligospermy and suggested a mating system with no sperm competition. Sexual transition was protogynous involving regression of ovarian tissue and proliferation of testicular tissue in the gonads. Transitional individuals occurred from May through November and accounted for 9% of sampled adult population. Sex change occurred in fish 69–93 cm (L_s) long and the size distributions of males and females overlapped over 27% of the L_s range. Special zones were recognized as gathering areas for sexually mature dusky groupers during the reproductive period.     

Evolutionary implications of morphological variation in the lower carnassial of red fox<Emphasis Type="Italic">Vulpes vulpes</Emphasis>

Research on the morphological variability of the occlusal surface of M₁ talonid in the red foxVulpes vulpes (Linnaeus, 1758) in the Holarctic has been carried out on 2271 specimens originating from 42 populations. The Nearctic was represented by 666 specimens belonging to 13 populations, whereas Palearctic was represented by 1605 specimens from 29 populations. Analyses of the developmental level and formation of cristids between the hypoconid and entoconid allowed the differentiation of 34 shape variants of the occlusal surface of the talonid in the red fox. Because of the complicated variation of cristids, 34 variants were assigned to 5 morphotypes of group P. In the Palearctic and Nearctic a significant geographic variation occurred of P morphotypes and their variants. Primitive variants of the talonid structure on M₁ are predominant in populations from the south of the Asian range of the red fox, while more progressive characters of the occlusal surface of the lower carnassial are typical of the northern and centrally located red fox populations in the Palearctic and Nearctic. The geographic differentiation is probably connected with different Pleistocene histories of particular populations.     

Otolith description and age‐and‐growth of Kurtus gulliveri from northern Australia

The sagitta of Kurtus gulliveri was ovate, moderately thick with the following attributes: lateral surface convex, mesial surface flat; dorsal margin sinuate, posterior margin rounded ventrally, ventral margin rounded and irregular; sulcus divided into ostium and cauda by constriction of dorsal and ventral margins, heterosulcoid, colliculum heteromorph; dorsal depression large and distinct, ventral groove close to margin in larger otoliths; rostrum broad and antirostrum small, separated by wide, shallow excisural notch. Otolith size was moderate, average 4·6% standard length ( L _S), typical for a perciform. Annuli on 78 whole sagittae were read, and 15% of these were transversely sectioned for verification of the annuli. Males ranged from 94 to 235 mm L _S and females from 95 to 284 mm L _S. There was little difference in size distribution of the sample between the sexes, perhaps due to a 6 month spawning season over which young were continually added to the population. Some sexual dimorphism was noted, however, as age 2 year females were significantly larger than males of the same age. The largest fish aged was a 284 mm L _S, 3 year‐old female, and the oldest age reached was 4 years by two males. It appears likely that most spawning females are ≥2 years old, but some larger 1 year old fish may attain sexual maturity.     

Reproductive biology and implications for management of the orange-spotted grouper Epinephelus coioides in the southern Arabian Gulf

The reproductive biology of Epinephelus coioides was determined from the examination of 1455 individuals collected between July 2005 and June 2007 in the southern Arabian Gulf. Histological preparations of gonads indicated that males were either derived from a juvenile phase or the transition of postspawning females, confirming a diandric protogynous sexual pattern. The spawning season was well defined, occurring once a year during April and early May. Peaks in spawning occurred after the full and new moons and was completed within a single lunar cycle. The presence of mature males over the entire size and age range and the absence of inactive mature females during the spawning season suggested that the population was not constrained by sperm limitation. While specimens undergoing sexual transition were only observed in size and age ranges of 335–685 mm total length ( L _T) and 5–6 years, patterns in the proportion of males in size and age classes suggested that sex change occurred at a relatively constant rate after female maturation up to the maximum size (1002 mm L _T) and age (11 years). Relationships between reproductive output and capacity with size and age indicated that conventional regulations that equate the mean size at first capture to sexual maturation are unsuitable for the management of E. coioides . The maximum age, small size and young age at sexual maturation ( L _min= 320 mm L _T, 2 years, for females and 242 mm L _T, 1 year, for males) conflict with the general pattern for large epinepheline groupers and may be a direct result of the intensive demersal fishery in the southern Arabian Gulf.     

Differences in size and growth rates of male and female migrating Japanese eels in Pearl River, China

The Sr/Ca ratios in otoliths of silver Japanese eels Anguilla japonica , in Pearl River, China, indicated that both sexes did not stay in brackish water and grew in fresh water from the glass eel stage until spawning migration. This did not support the hypothesis that females tended to distribute upstream and males might be restricted to estuaries. The back-calculated total length of males at glass eel stage was not significantly different from that of females, indicating that the hypothesis that small glass eels became males and larger ones became females may not be true. The mean (±S.D.) age and total length of males at migration were 6·4±1·6 years and 48·3±4·5 cm, which were significantly smaller than for females, 8·3±1·6 years and 61·4±4·1 cm. The age of migration was related inversely to growth rate for both sexes. Growth parameters of the von Bertalanffy growth equation were K =0·21 cm year°1, L _∞=55·7 cm and t _o=-0·55 year for males and K =0·14 cm year⁻¹, L _∞=77·5 cm and t _o=-0·60 year for females. The difference in asymptotic length ( L _∞) between males and females may be because females postpone migration to achieve larger size for maximizing reproductive success.     

Sex ratio and sexual dimorphism of the anchovy Anchoa januaria (Actinopterygii, Engraulidae) in a tropical bay in south-eastern Brazil

Sex ratio and morphological traits of a very abundant anchovy Anchoa januaria were described in a tropical bay in south-eastern Brazil. The aim was to test the hypothesis that sexual dimorphism occurs due to the different reproductive roles of the sexes. A fish sampling programme was carried out between September 1998 and August 1999 at six sites: four sandy beaches and two lower-river sites. Population structure at river sites comprised adults only, ranging from 60 to 80 mm total length ( L _T), while at sandy beaches both juveniles and adults were found, ranging from 32 to 80 mm L _T. Well-balanced 'spawning school' at river sites during reproduction were detected, while female-dominated schools occurred in the bay feeding areas. Males had relatively longer pectoral fins, slightly larger hearts and more somatic mass than females. Females outnumbered males at sizes >67 mm L _T and had significantly longer intestines and heavier livers than males. The largest size reached by females was probably related to a higher growth rate as they have a larger intestinal absorbing area for nutrients. The prediction of higher energetic investment in reproduction by females that should have larger organs associated with food acquisition and processing to produce energy-rich eggs was confirmed for A. januaria in Sepetiba Bay.     

Pelvic dimorphism in relation to body size and body size dimorphism in humans

Many mammalian species display sexual dimorphism in the pelvis, where females possess larger dimensions of the obstetric (pelvic) canal than males. This is contrary to the general pattern of body size dimorphism, where males are larger than females. Pelvic dimorphism is often attributed to selection relating to parturition, or as a developmental consequence of secondary sexual differentiation (different allometric growth trajectories of each sex). Among anthropoid primates, species with higher body size dimorphism have higher pelvic dimorphism (in converse directions), which is consistent with an explanation of differential growth trajectories for pelvic dimorphism. This study investigates whether the pattern holds intraspecifically in humans by asking: Do human populations with high body size dimorphism also display high pelvic dimorphism? Previous research demonstrated that in some small-bodied populations, relative pelvic canal size can be larger than in large-bodied populations, while others have suggested that larger-bodied human populations display greater body size dimorphism. Eleven human skeletal samples (total N: male = 229, female = 208) were utilized, representing a range of body sizes and geographical regions. Skeletal measurements of the pelvis and femur were collected and indices of sexual dimorphism for the pelvis and femur were calculated for each sample [ln(M/F)]. Linear regression was used to examine the relationships between indices of pelvic and femoral size dimorphism, and between pelvic dimorphism and female femoral size. Contrary to expectations, the results suggest that pelvic dimorphism in humans is generally not correlated with body size dimorphism or female body size. These results indicate that divergent patterns of dimorphism exist for the pelvis and body size in humans. Implications for the evaluation of the evolution of pelvic dimorphism and rotational childbirth in Homo are considered.     

Reversed sexual size dimorphism in microtines: Are females larger than males or are males smaller than females?

Summary We analysed sexual size dimorphism for 21 populations of microtine rodents. Female to male size ratio varied considerably among populations from females significantly larger than males (ratio=1.18) to males larger than females (ratio=0.78). In a multiple regression analysis female to male home range size ratio explained 94% of the total variation in body size dimorphism and was the only one of eight independent variables that was selected in a stepwise regression procedure. When females are the larger sex, males have home range sizes much larger than females. We suggest that the relationship between home range size ratio and body weight size dimorphism reflects different selection pressures on males and females in competition for resources and mates.     

Sexual and geographical variation in life history parameters of the shorthorn sculpin

A total of 293 shorthorn sculpins Myoxocephalus scorpius from Tromsø, northern Norway, were sampled between November 1998 and April 1999 to determine sex, total length, age, growth, maturity and mortality. Females grew to larger sizes ( L _∞=26·9 v. 18·5 cm), matured later (2 v. 1 year of age) at larger size (maturation length=16 v. 14 cm L _T), and had lower instantaneous mortality rates (0·93 v. 1·20 year⁻¹) than males. The life history parameters of shorthorn sculpins in northern Norway were more similar to the parameters of short-lived central European populations than to the parameters of the long-lived population of Newfoundland. This study confirms that northern Norwegian shorthorn sculpins exhibit sexual dimorphism as in other shorthorn sculpin populations. The relationships between growth pattern, age at maturity and mortality rates observed in the Tromsø population and in other shorthorn sculpin populations, correspond well with the predictions from a published life history model.     

Taxonomy of the deep-sea eel genus, Histiobranchus (Synaphobranchidae, Anguilliformes), with notes on the ecology of H. bathybius in the eastern North Atlantic

The eel genus Histiobranchus Gill occurs benthopelagically over the continental rise and abyss of the World Ocean, primarily beneath temperate and subpolar surface waters. Its generic status within the subfamily Synaphobranchinae is confirmed by comparison of the structure and topography of its cephalic sensory system and skeletal features with Synaphobranchus. At least three species of Histiobranchus are recognized: H. bathybius (panoceanic), H. bruuni (Tasman Sea and waters south-east of New Zealand) and H. australis (two geographical forms; South Atlantic and south-western Indian Ocean, and South Indo-West Pacific Oceans). Collections totalling 319 specimens of H. bathybius from the eastern North Atlantic (1790–5440 m depth) yielded a size range of 99–1370 mm total length ( L _T), with no apparent sexual dimorphism. Length–frequency distributions indicate a mode of juvenile fish at around 100–200 mm L _T and a further two around 600–700 and 1300–1400 mm L _T among adults. Generally smaller fish occur in shallower regions, although the size range is broad over the whole depth range. No apparent trend occurs in the size distribution with latitude over the range 17–54) N. Females outnumber males (1 male : 1·7 females) and both sexes are largely distinguishable from 300 mm L _T. Ripening eggs occur in females from both adult length modes, with running ripe and spent females of very different size indicating iteroparity.     

Brief communication: Morphometric data for adult lion-tailed macaques (Macaca silenus)

Basic morphometric data were collected from 22 adult lion-tailed macaques (M. silenus) of both sexes. M. silenus is a rare primate species from which adequate morphometric data have not heretofore been available for comparative purposes. Data collected include measures of gross body size (weight; crown-rump and rump-heel length), and for males, measures of secondary sexual characteristics (canine tooth and testes size). Degree of sexual dimorphism was marked, with males significantly larger and heavier than females. The three body size measures were correlated for males but not for females. There was substantial variation among individual males in secondary sex characteristics measurements. The data indicate than lion-tailed macaque morphometrics are consonant with the general pattern of positive allometry for body size and sexual dimorphism characteristic of the primate order.     

The reproductive biology and ecology of the Port Jackson shark Heterodontus portusjacksoni in the coastal waters of eastern Australia

The reproductive biology and ecology of the Port Jackson shark Heterodontus portusjacksoni was investigated at three locations on the central and southern coast of New South Wales (NSW), Australia from January 2002 to December 2005 using underwater visual census surveys and samples obtained from a commercial fishery. Adults displayed sexual dimorphism in total length ( L _T) at sexual maturity, with males maturing between 762 and 772 mm L _T and females between 902 and 905 mm L _T. The mean ovarian fecundity was estimated at 16 offspring per female but was unrelated to female L _T. Male gonado-somatic ( I _G) and hepato-somatic ( I _H) indices and female I _G declined from July to November as did maximum ovarian follicle diameter and the diameter of the three largest follicles. Adults were absent from inshore reefs between December and July. Hence, H. portusjacksoni has a synchronous annual breeding season in NSW, which occurs between July and November (the austral winter to spring), with a peak in oviposition from August to October. Heterodontus portusjacksoni copulatory and ovipository behaviour are reported for the first time. Copulation was observed and involved oral grasping of the female's pectoral fin by a single male, which wrapped his body around hers to insert one clasper. Ovipositing females appeared to search crevices in the reef prior to delivering a single capsule, which was washed into the crevice by water movement, with the female departing very soon after oviposition. This study represents the first rigorously quantitative analysis of H. portusjacksoni reproductive biology and ecology in NSW waters.     

Population variables and life-history characteristics of the alligator pipefish Syngnathoides biaculeatus, in Papua New Guinea

Population structure and life-history variables of the widely distributed alligator pipefish Syngnathoides biaculeatus were characterized in Bootless Bay, Papua New Guinea over the course of 11 months. There was little evidence of seasonality with four focal populations showing no significant change in abundance. Similarly, the sex ratio remained 1:1 for all but 1 month. Reproductive males carrying eggs (148–278 mm in total length, L _T) were found in all months. Brood size was significantly, positively related to male L _T for newly laid broods only. Maximum observed brood size was 351 and mean ± s . d . brood size was 238 ± 57 for newly laid broods. Juveniles and males showed no change in mean L _T over the year while slightly smaller females were captured in November 2006 and September 2007. Males were significantly longer than females so von Bertalanffy growth coefficients were estimated separately for each sex: males L _∞= 285 mm, K = 0·82 year⁻¹ and females L _∞= 261 mm, K = 1·10 year⁻¹. These estimates suggest that this species grows rapidly and has a short-life span. In the context of growing concern about overexploitation of syngnathids, a rapid growth rate combined with year round reproductive activity suggests that the tropical S. biaculeatus may be relatively resilient with regard to fishing pressure.     

Sexual size dimorphism and sexual selection in turtles (order testudines)

Summary This paper combines published and original data on sexual size dimorphism, reproductive behavior, and habitat types in turtles. Our major finding is that observed patterns of sexual size dimorphism correlate with habitat type and male mating strategy. (1) In most terrestrial species, males engage in combat with each other. Males typically grow larger than females. (2) In semiaquatic and bottom-walking aquatic species, male combat is less common, but males often forcibly inseminate females. As in terrestrial species, males are usually larger than females. (3) In truly aquatic species, male combat and forcible insemination are rare. Instead, males utilize elaborate precoital displays, and female choice is highly important. Males are usually smaller than females.We interpret these correlations between sexual behavior and size dimorphism in terms of sexual selection theory: males are larger than females when large male size evolves as an adaptation to increase success in male combat, or to enable forcible insemination of females. In contrast, males are usually smaller than females where small size in males evolves to increase mobility (and hence, ability to locate females), or because selection for increased fecundity may result in increased female size. In turtle species with male combat or forcible insemination, the degree of male size superiority increases with mean species body size.     

Age, growth and reproduction of the black scorpionfish, Scorpaena porcus (Pisces, Scorpaenidae), on the Black Sea coast of Turkey

Age, growth and reproduction of the black scorpionfish, Scorpaena porcus were studied in specimens from the coast of the Sinop Peninsula (Black Sea) between March 2002 and April 2003 in order to characterize these population parameters in comparison to specimens from populations of nearby regions. A total of 1086 specimens was captured by beam trawl at the depths between 0 and 30 m. The total number of females (510) was significantly higher than that of males (373). Total length of males and females ranged between 5.7 and 23.6 cm, and 4.9 and 31.7 cm, respectively. The length–weight relationship showed a positive allometric growth. Females grew faster and reached a larger size at age than males ( L _∞ = 111.9 cm, K  = 0.035 year⁻¹, φ' = 2.64 for females, and L _∞ = 74.6 cm, K  = 0.054 year⁻¹, φ' = 2.49 for males). The age range estimated was up to 8 years for females and 5 years for males. Reproduction likely occurs between June and September. Sex ratio varied greatly with season, perhaps indicating different seasonal migratory patterns in adults of different sex. An inverse correlation between gonadosomatic index (GSI) and hepatosomatic index (HSI) was evident during the reproduction seasons. The mean size at first sexual maturity was 17.5 cm TL for females, and 16.7 cm TL for males.     

Reproductive cycle and sexual maturity of the anglerfish Lophiomus setigerus in the East China Sea with a note on specialized spermatogenesis

The reproductive cycle and sexual maturity of the anglerfish Lophiomus setigerus were examined. Spermatids were released from the germinal cysts into the lumina of the seminal lobules, and both spermatids and spermatozoa were present in the lumina of the seminal lobules and sperm ducts. Spermatogenesis and vitellogenesis occurred throughout most of the year. The testes of males were full of spermatozoa throughout the year, with spawning from May to November. Males and females reached sexual maturity at a mean total length and age of 178 mm, 3.3 years and 303 mm, 6.1 years, respectively. There were clear seasonal cycles in the gonadosomatic index (I_G) and hepatosomatic index ( I _H) in females. The mean I_G of females increased rapidly with ovarian development while the mean I _H decreased from the middle of vitellogenesis to final maturation. Mean values of I _G and I _H in males increased with testicular development.     

Sexual dimorphism in the postcranial skeleton of New World primates

This study examines sexual dimorphism in 24 dimensions of the postcranial skeleton of four platyrrhine species: Callithrix jacchus, Saguinus nigricollis, Saimiri sciureus, and Cebus albifrons. The two callitrichid species show a relatively small amount of variation in the degree of sexual dimorphism among the different dimensions. Variation is considerably higher in the two cebid species as reflected by a mosaic pattern of sexual dimorphisms with males being significantly larger than females in some dimensions, and females significantly larger than males in others. In dimensions of the pectoral girdle and limb bones, males and females in each of the two cebid species are essentially scaled versions of each other, with males being peramorphic compared to females. This pattern is primarily the result of time hypermorphosis, i.e. an extension of the growth period in time in males. Rate hypermorphosis, i.e. an increase in the rate of growth in time in males, appears to play an additional role, however, in S. sciureus. By contrast, in dimensions of the true pelvis, sex differences in shape are dissociated from those in size. They are interpreted as the result of acceleration, i.e. increase in rate of shape change in females, as an adaptation to obstetrical functions. Interspecific analyses indicate positive allometry of mean degree of postcranial dimorphism with respect to body size. This coincides with previous findings by Leutenegger and Cheverud [1982, 1985] on the scaling of sexual dimorphism in body weight and canine size, and thus supports their model which posits selection on body size as the prime mover for the evolution of sexual dimorphism.