Sexual Selection and Dynamics of Jaw Muscle in Tupinambis Lizards

Sexual dimorphism patterns provide an opportunity to increase our understanding of trait evolution. Because selective forces may vary throughout the reproductive period, measuring dimorphism seasonally may be an interesting approach. An increased male head size may be important in intersexual and intrasexual interactions. In Tupinambis lizards, a big head is attributed in part to a large adductor muscle mass. Competition for mating can differ in species with different sex ratio and different degrees of sexual size dimorphism. We examined sexual differences in mass of the pterygoideus muscle, its temporal variation throughout the reproductive period and the relationship between muscle and reproductive condition in Tupinambis merianae and T. rufescens. We characterized sexual size dimorphism and sex ratio in both species. Mature males had larger jaw muscles than mature females in both species, mainly during the reproductive season. The dimorphism in jaw muscle was due to an increase in muscle mass in sexually active males. Seasonal increases in muscle mass and variation between immature and mature individuals suggest that the jaw muscle might be a secondary sexual character. We propose that the pterygoideus muscle may act as a signal of reproductive condition of males because it is associated with testis size and sperm presence. The patterns of sexual dimorphism in jaw muscle in both species were similar; however, the comparison shows how sexual characters remain dimorphic in different competition contexts and in species with different degrees of body size dimorphism. Our results suggest that jaw muscle as sexual character could be influenced by inter- and intrasexual selective pressures.     

Sexual differences in head form and diet in a population of Mexican lance‐headed rattlesnakes,Crotalus polystictus

Sexual dimorphism of phenotypic traits associated with resource use is common in animals, and may result from niche divergence between sexes. Snakes have become widely used in studies of the ecological basis of sexual dimorphism because they are gape‐limited predators and their head morphology is likely to be a direct indicator of the size and shape of prey consumed. We examined sexual dimorphism of body size and head morphology, as well as sexual differences in diet, in a population of Mexican lance‐headed rattlesnakes, Crotalus polystictus, from the State of México, Mexico. The maximum snout–vent length of males was greater than that of females by 21%. Males had relatively larger heads, and differed from females in head shape after removing the effects of head size. In addition, male rattlesnakes showed positive allometry in head shape: head width was amplified, whereas snout length was truncated with increased head size. By contrast, our data did not provide clear evidence of allometry in head shape of females. Adults of both males and females ate predominately mice and voles; however, males also consumed a greater proportion of larger mammalian species, and fewer small prey species. The differences in diet correspond with dimorphism in head morphology, and provide evidence of intersexual niche divergence in the study population. However, because the sexes overlapped greatly in diet, we hypothesize that diet and head dimorphisms in C. polystictus are likely related to different selection pressures in each sex arising from pre‐existing body size differences rather than from character displacement for reducing intersexual competition. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 633–640.     

Size dimorphism in Rankinia [Tympanocryptis] diemensis (Family Agamidae): sex-specific patterns and geographic variation

Sexual dimorphism has implications for a range of biological and ecological factors, and intersexual morphological differences within a species provide an ideal opportunity for investigating evolutionary influences on phenotypic variation. We investigated sexual size dimorphism (SSD) in an agamid species, Rankinia [Tympanocryptis] diemensis , to determine whether overall size and/or relative morphological trait size differences exist and whether geographic variation in size dimorphism occurs in this species. Relative morphological trait proportions included a range of head, limb, and inter-limb measurements. We found significant overall intersexual adult size differences; females were the larger sex across all sites but the degree of dimorphism between the sexes did not differ between sites. This female-biased size difference is atypical for agamid lizards, which are usually characterized by large male body size. In this species, large female-biased SSD appears to have evolved as a result of fecundity advantages. The size of relative morphological trait also differed significantly between the sexes, but in the opposite direction: relative head, tail, and limb sizes were significantly larger in males than females. This corresponds to patterns in trait size usually found in this taxonomic group, where male head and limb size is important in contest success such as male–male rivalry. There were site-specific morphological differences in hatchlings, including overall body size, tail, inter-limb, thigh, and hindlimb lengths; however, there were no sex-specific differences indicating the body size differences present in the adult form occur during ontogeny.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 699–709.     

Sexual niche partitioning in two species of New Guinean Pachycephala whistlers

Sex differences in foraging behavior have been widely reported in the ornithological literature, but few examples are available from tropical avifaunas. Differences between males and females in foraging behavior have been hypothesized to be a byproduct of sexual size dimorphism or a result of niche partitioning to reduce intersexual competition for food or different reproductive roles. From 2010 to 2013, I used foraging data and mist‐net capture rates from multiple study sites to examine possible sex differences in the foraging behavior of two New Guinean Pachycephala whistlers. I found that male Regent (Pachycephala schlegelii) and Sclater's (Pachycephala soror) whistlers consistently foraged in higher strata than females. It is unlikely that these differences are due to sexual dimorphism because these species exhibit little sexual dimorphism. Sex differences in foraging behavior were consistent across years and study sites and did not appear linked to breeding behavior, supporting the food‐competition hypothesis, but not the reproductive‐roles hypothesis. Male territorial defense often occurs in relatively high strata in Pachycephala whistlers, possibly influencing male foraging strata. However, male territorial behavior cannot explain why females predominately forage in lower strata. Instead, intersexual competition for food resources is likely the primary driver of differences in the foraging behavior of male and female Regent and Sclater's whistlers.     

Sexually dimorphic muscles in the forelimb of the Japanese toad,Bufo japonicus

During the breeding season, male anurans display clasping behavior by holding females with their forelimbs. This behavior is peculiar to males, and may require specializations in forelimb musculature. The present study revealed that five kinds of forelimb muscles were heavier in the male Japanese toad than in the female: the flexor carpi radialis (FCR), the flexor antibrachii medialis caput superius (FAMsup), the abductor indicis longus (AIL), the extensor carpi radialis caput superius (ECRsup), and the flexor antibrachii lateralis superficialis caput superius (FALSsup). In addition, one breast muscle, the coracoradialis (CR), was also heavier in males than in females. A quantitative analysis of muscle fibers processed for myosin ATPase activity showed that, in such “sexually dimorphic muscles” of the female, both fast (twitch) and slow (tonic) muscle fibers were of smaller diameter than in other forelimb muscles of both sexes (all male muscles plus “nondimorphic muscles” of the female). Moreover, both types of fibers were less numerous than in the corresponding muscles of the male. These results suggest that the “sexually dimorphic muscles” are used especially for clasping by the male and are degenerative or subnormal in the female. Slow muscle fibers were neither peculiar to, nor abundant in, these clasping muscles, although they may well be necessary for tonic and prolonged contractions of the forelimb muscles during clasping. The mechanism of sexual dimorphism may be a direct action of androgens on clasping muscles or an indirect action on clasping muscles via the innervating motoneurons.     

Identifying tasks to elicit maximum voluntary contraction in the muscles of the forearm

Maximum voluntary contractions (MVCs) are often used for the normalisation of electromyography data to enable comparison of signal patterns within and between study participants. Recommendations regarding the types of tasks that are needed to collect MVCs for the muscles of the forearm have been made, specifically advocating the use of resisted moment tasks to get better estimates of forearm MVCs. However, a protocol detailing which specific tasks to employ has yet to be published. Furthermore, the effects of limb dominance on the collection of MVCs have not been considered previously. Muscle activity was monitored while 23 participants performed nine isometric, resisted tasks. The tasks that are likely to elicit MVC in the flexor carpi ulnaris, flexor carpi radialis, flexor digitorum superficialis, extensor carpi ulnaris, extensor carpi radialis, extensor digitorum communis, and pronator teres were identified. Thus, targeted protocols can be designed to mitigate against fatigue. Hand dominance had limited effect, with differences being found only in the finger flexors and extensors (p< 0.03). Thus, use of the contralateral flexor digitorum superficialis and extensor digitorum communis muscles to obtain baselines for activation levels and patterns may not be appropriate.     

蓝尾石龙子的头部两性异形和食性

通过测量头、体大小和胃检研究浙江泰顺产蓝尾石龙子 (Eumeceselegans)个体发育过程中两性异形和食性的变化。蓝尾石龙子成体个体大小和头部大小的两性差异显著 ,雄性大于雌性。不同发育阶段雌性头长与SVL的线性回归斜率无显著差异 ,头宽与SVL线性回归斜率的差异显著 ,成体和SVL <5 0mm幼体头宽随SVL的增长速率显著小于SVL为 5 0 - 6 9mm的幼体。雄性头部相对于SVL呈加速式异速生长。两性比较发现 :雌雄幼体头长和头宽随SVL的增长速率无显著差异 ,SVL <5 0mm幼体特定SVL的头长和头宽无显著的两性差异 ,但SVL为 5 0 - 6 9mm的雄性幼体头长和头宽大于SVL相同的雌性幼体 ;雄性成体头长和头宽随SVL的增长速率显著大于雌性。SVL <5 0mm的雌性幼体头部相对小于SVL为 5 0 - 6 9mm的同性幼体 ,性成熟雌体头部相对小于SVL为 5 0 - 6 9mm的同性幼体。雌性幼体、雄性幼体、雌性成体和雄性成体食物生态位宽度分别为 12 3、 12 5、 4 8和 14 4。雌雄幼体食物生态位重叠度最高 ,雌雄成体食物生态位重叠度次之 ,成体与幼体食物生态位重叠度较小。成体摄入食饵的大小 (用胃内完整食物长度的平均值表示 )和变化范围大于幼体。两性成、幼体摄入的食饵大小差异显著。两性个体摄入的食饵大小均与其SVL呈正相关 ,表明较大     

It takes two: Seasonal variation in sexually dimorphic weaponry results from divergent changes in males and females

Sexually dimorphic weaponry often results from intrasexual selection, and weapon size can vary seasonally when costs of bearing the weapon exceed the benefits outside of the reproductive season. Weapons can also be favored in competition over nonreproductive resources such as food or shelter, and if such nonreproductive competition occurs year‐round, weapons may be less likely to vary seasonally. In snapping shrimp (Alpheus angulosus), both sexes have an enlarged snapping claw (a potentially deadly weapon), and males of many species have larger claws than females, although females are more aggressive. This contrasting sexual dimorphism (larger weaponry in males, higher aggression in females) raises the question of whether weaponry and aggression are favored by the same mechanisms in males and females. We used field data to determine whether either sex shows seasonal variation in claw size such as described above. We found sexual dimorphism increased during the reproductive season due to opposing changes in both male and female claw size. Males had larger claws during the reproductive season than during the nonreproductive season, a pattern consistent with sexual selection. Females, however, had larger claws during the nonreproductive season than during the reproductive season—a previously unknown pattern of variation in weapon size. The observed changes in female weapon size suggest a trade‐off between claw growth and reproduction in the reproductive season, with investment in claw growth primarily in the nonreproductive season. Sexually dimorphic weaponry in snapping shrimp, then, varies seasonally due to sex differences in seasonal patterns of investment in claw growth, suggesting claws may be advantageous for both sexes but in different contexts. Thus, understanding sexual dimorphisms through the lens of one sex yields an incomplete understanding of the factors favoring their evolution.     

Sexual differences in morphology and niche utilization in an aquatic snake,Acrochordus arafurae

Filesnakes (Acrochordus arafurae) are large (to 2 m), heavy-bodied snakes of tropical Australia. Sexual dimorphism is evident in adult body sizes, weight/length ratios, and body proportions (relative head and tail lengths). Dimorphism is present even in neonates. Two hypotheses for the evolution of such dimorphism are (1) sexual selection or (2) adaptation of the sexes to different ecological niches. The hypothesis of sexual selection is consistent with general trends of sexually dimorphic body sizes in snakes, and accurately predicts, for A. arafurae, that the larger sex (female) is the one in which reproductive success increases most strongly with increasing body size. However, the sexual dimorphism in relative head sizes is not explicable by sexual selection.The hypothesis of adaptation to sex-specific niches predicts differences in habitats and/or prey. I observed major differences between male and female A. arafurae in prey types, prey sizes and habitat utilization (shallow versus deep water). Hence, the sexual dimorphism in relative head sizes is attributed to ecological causes rather than sexual selection. Nonetheless, competition between the sexes need not be invoked as the selective advantage of this character divergence. It is more parsimonious to interpret these differences as independent adaptations of each sex to increase foraging success, given pre-existing sexually-selected differences in size, habitat or behavior. Data for three other aquatic snake species, from phylogenetically distant taxa, suggest that sexual dimorphism in food habits, foraging sites and feeding morphology, is widespread in snakes.     

Seasonal dimorphism in the horny bills of sparrows

Bill size is often viewed as a species‐specific adaptation for feeding, but it sometimes varies between sexes, suggesting that sexual selection or intersexual competition may also be important. Hypotheses to explain sexual dimorphism in avian bill size include divergence in feeding niche or thermoregulatory demands, intrasexual selection based on increased competition among males, or female preference. Birds also show seasonal changes in bill size due to shifts in the balance between growth rate and wear, which may be due to diet or endogenous rhythms in growth. Insight into the function of dimorphism can be gained using the novel approach of digital x‐ray imaging of museum skins to examine the degree to which the skeletal core or the rhamphotheca contribute to overall dimorphism. The rhamphotheca is ever‐growing and ever‐wearing, varying in size throughout life; whereas the skeletal core shows determinant growth. Because tidal marsh sparrows are more dimorphic in bill size than related taxa, we selected two marsh taxa to investigate dimorphism and seasonality in the size of the overall bill, the skeletal core, and the rhamphotheca. Bill size varied by sex and season, with males having larger bills than females, and bill size increasing from nonbreeding to breeding season more in males. Skeletal bill size varied with season, but not sex. The rhamphotheca varied primarily with sex; males had a larger rhamphotheca (corrected for skeletal bill size), which showed a greater seasonal increase than females. The rhamphotheca, rather than the skeletal bill, was responsible for sexual dimorphism in overall bill size, which was particularly well developed in the breeding season. The size of the rhamphotheca may be a condition‐based character that is shaped by sexual selection. These results are consistent with the evidence that bill size is influenced by sexual selection as well as trophic ecology.     

Intersexual differences in the foraging behavior of Formicivora littoralis (Thamnophilidae), an endangered Neotropical bird

Sex differences in foraging can be explained by avoidance of competition, reproductive strategies or breeding roles. We investigated foraging behavior and potential intersexual differences in the Restinga Antwren over the year, recording foraging tactics, their timing, substrate and their height above the ground. We found significant differences between sexes in foraging tactics, substrates and height of foraging. Females foraged mostly on the ground while males in the middle and top of the vegetation. Smaller differences in foraging height between sexes in the breeding season compared with non-breeding season support the reproductive strategies model, although the avoidance of competition cannot be excluded.     

Quantitative assessment of upper limb muscle fatigue depending on the conditions of repetitive task load

The aim of this study was to discriminate fatigue of upper limb muscles depending on the external load, through the development and analysis of a muscle fatigue index. Muscle fatigue is expressed by a fatigue index based on an amplitude parameter (calculated in the time domain) and a fatigue index based on a frequency parameter (a parameter calculated in the frequency domain). The fatigue index involves a regression function that describes changes in the EMG signal parameter, time elapsing before muscle fatigue and the probability of specific trends in changes in EMG parameters for the population under study.

The experimental study covered a group of 10 young men. During the study, they exerted force at a specific level and for a specific time in 12 load variants. During the study, EMG signals from four muscles of the upper limb were recorded (trapezius pars descendents, biceps brachii caput breve, extensor carpi radialis brevis, flexor carpi ulnaris). For each variant and for each examined muscles, the value of the fatigue index was calculated. Values of that index quantitatively expressed fatigue of a specific muscle in a specific load variant.

A statistical analysis indicated variation in the fatigue of the biceps brachii caput breve, extensor carpi radialis brevis, and flexor carpi ulnaris muscles depending on the external load (load variant) according to the task performed with the upper limb.

The study demonstrated usefulness of the fatigue index in expressing quantitatively muscle fatigue and in discriminating muscle fatigue depending on the external load.     

Sexual size dimorphism in a landlocked Pacific salmon in relation to breeding habitat features

Many animal species exhibit size dimorphism between sexes. Sexual selection, whereby male–male competition favors larger body sizes, has been considered a likely cause of sexual size dimorphism. Habitat features in breeding areas could affect the outcome of male–male competition, yet few attempts have been made to relate breeding habitat features with interpopulation variation in sexual size dimorphism. In this study, we examined interpopulation variation in sexual size dimorphism by studying the landlocked amago salmon (Oncorhynchus masou ishikawae) at a microgeographic scale. We found that female body size was independent of stream size but that male body size decreased with smaller stream sizes. A likely explanation is that the relationship between reproductive success and the size of males is influenced by the availability of refuges that are only available to small-bodied males. Sexual differences in body size increased with decreasing stream sizes, supporting the hypothesis that the reproductive success of larger males is reduced in smaller streams. In contrast, the maturation-length threshold increased with stream size for both sexes. The stream-size-based interpopulation variation in sexual size dimorphism and size at maturity in landlocked amago salmon may therefore have arisen through a combination of sexual and natural selection.     

Diet and reproductive biology of the Starred Agama,Laudakia stellio (Linnaeus, 1758) (Squamata: Agamidae), in the northern Sinai,Egypt

We quantified sexual size dimorphism, diet and reproduction in the Starred Agama, Laudakia stellio, in northern Sinai. Males were larger than females in snout-vent length and head index. The species is a sit-and-wait predator and feeds on insects, mainly coleopterans. About 30% of stomachs included plant material. No difference between sexes existed in terms of prey size preference. The reproductive season is seen to be year round with no distinctive seasonality. The smallest sexually mature female measured 92 mm SVL, whereas the smallest sexually reproductive male was 89 mm SVL. Clutch size ranged from 6 to 18 eggs.     

Intersexual niche segregation among three bark-foraging birds of eucalypt forests

All three species of specialized bark-climbing birds in eucalypt forests of north-eastern New South Wales exhibited sexual differences in their foraging behaviour. The degree of these differences was weakest in the least social species and strongest in the most social species, suggesting that intersexual niche segregation is an adaptation to reduce intraspecific competition for food. In the least social White-throated Treecreeper, the slightly longer-billed males foraged on dead branches more than females only during the breeding season when pair members were forced to share the same area. In the group-living Red-browed Treecreeper and Varied Sittella, males foraged lower and used rough-barked trunks and limbs more than females throughout the year. Sexual differences in the foraging substrates of Sittellas may have been related to sexual dimorphism of the bill, but in the monomorphic Red-browed Tree-creeper, male dominance was probably important. Some evidence suggests that further subdivision of the niche may occur in the highly social Sittella along the vertical dimension according to age and/or reproductive status. This report argues that intersexual niche segregation is just as likely to develop under conditions of increased interspecific competition as under reduced competition.     

Four forearm flexor muscles of the horse, Equus caballus: anatomy and histochemistry.

Two of the forearm flexors of the horse, the superficial and deep digital flexor muscles, are critical to support the digital and fetlock joints, exhibit differing insertions, and are passively supported by the proximal and distal check ligaments, respectively. These two muscles differ in histochemical composition and architecture. The differences are correlated with the different stress levels transmitted through their tendons, and the different frequencies of clinical breakdown that have been reported. Both muscles contain type I and type IIa fibers. A few type IIb fibers occurred in the deep digital flexor. The superficial digital flexor contained approximately 56% type I fibers, extremely short muscle fibers, and extensive connective tissue investment. In contrast, the deep digital flexor had three muscle heads: ulnar, radial, and "long" and "short" regions of the humeral head. The "long" and "short" regions of the humeral head contained 33% and 44% type I fibers, respectively, fiber lengths three to four times as long as those in the superficial digital flexor, and relatively less connective tissue investment. Flexor carpi radialis and flexor carpi ulnaris compared most closely with the humeral head of the deep digital flexor. These data suggest a correlation of the unique architecture of superficial digital flexor with its proposed elastic storage properties during locomotion in horses, and an explanation for the frequent breakdown of the superficial digital flexor in athletic horses.     

Environmental influences on the sexual dimorphism in body size of western bobcats

Sexual size dimorphism might be influenced by environmental constraints on sexual selection or by intraspecific competition between males and females. We studied bobcats (Lynx rufus) in collections of museum specimens from western North America to examine these hypotheses. Structural body size was estimated from several measurements of the skull, ln-transformed and indexed through principal components analysis. Sexual dimorphism in body size was estimated from the difference in size index of males and females, and compared to geographic and climatic variables associated with biotic provinces (ecoregions). Of several climatic variables that were associated with bobcat body size, only seasonality of climate was associated with sexual dimorphism. Sexual size dimorphism, longitude, elevation, and seasonality were intercorrelated. As longitude decreased (moving inland from west-coastal ecoregions), sexual dimorphism decreased with the increased elevation and seasonality of continental climates of the Rocky Mountains. We suggest that increased seasonality and the need for fasting endurance by females may place constraints on the degree of sexual dimorphism in bobcats. Sexual dimorphism of body size and sexual size dimorphism of trophic structures (teeth) exhibited a strong positive association over geography, thus indirectly supporting the hypothesis that intrasexual competition for prey could account for the geographic variation in sexual size dimorphism. Thus, both environmental constraints on sexual selection of body size and intersexual competition were supported as possible explanations of the degree of sexual size dimorphism that occurs in populations of bobcats.     

Sex‐specific phenotypic plasticity in response to the trade‐off between developmental time and body size supports the dimorphic niche hypothesis

Female‐biased sexual size dimorphism (SSD) is widespread in many invertebrate taxa. One hypothesis for the evolution of SSD is the dimorphic niche hypothesis, which states that SSD evolved in response to the different sexual reproductive roles. While females benefit from a larger body size by producing more or larger eggs, males benefit from a faster development, which allows them to fertilize virgin females (protandry). To test this hypothesis, we studied the influence of temperature and intraspecific density on the development of Chorthippus montanus. We reared them in monosexual groups under different conditions and measured adult body size, wing length, nymphal mortality, and development time. The present study revealed an inverse temperature–size relationship: body size increased with increasing temperature in both sexes. Furthermore, we found intersexual differences in the phenotypic response to population density, supporting the dimorphic niches hypothesis. At a lower temperature, female development time increased and male body size decreased with increasing density. Because there was no food limitation, we conclude that interference competition hampered development. By contrast to expectations, mortality decreased with increasing density, suggesting that interference did not negatively affect survival. The present study shows that sex‐specific niche optima may be a major trigger of sexual dimorphisms. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 48–57.     

黑斑侧褶蛙的两性异形和雌性繁殖特征

测定了黑斑侧褶蛙成体的体长、体重、头长、头宽、眼径、鼓膜径、前肢长、后肢长等形态指标以及雌体的怀卵数量。黑斑侧褶蛙雌体的体长和体重显著大于雄体。其它局部形态特征指标与体长呈正相关,协方差分析表明,雄体的鼓膜径大于雌体,其余形态指标不存在明显的两性差异。黑斑侧褶蛙雌体怀卵数量与体长和体重皆成正相关关系,表明黑斑侧褶蛙通过增加个体大小增加繁殖输出。     

Sexual dimorphism of head size in Gallotia galloti: testing the niche divergence hypothesis by functional analyses

1. Two often cited hypotheses explaining sexual head size dimorphism in lizards are: sexual selection acting on structures important in intrasexual competition, and reduction of intersexual competition through food niche separation.
2. In this study some implicit assumptions of the latter hypothesis were tested, namely that an increase in gape distance and bite force should accompany the observed increase in head size. These assumptions are tested by recording bite forces, in vivo , for lizards of the species Gallotia galloti . In this species, male lizards have significantly larger heads than female conspecifics of similar snout–vent length.
3. Additionally, the average force needed to crush several potential prey species was determined experimentally and compared with the bite force data. This comparison clearly illustrates that animals of both sexes can bite much harder than required for most insect food items, which does not support the niche divergence hypothesis. The apparent 'excess' bite force in both sexes might be related to the partially herbivorous diet of the animals.
4. To unravel the origin of differences between sexes in bite capacity, the crushing phase of biting was modelled. The results of this model show different strategies in allocation of muscle tissue between both sexes. The origin of this difference is discussed and a possible evolutionary pathway of the development of the sexual dimorphism in the species is provided.