Melanin‐Based Coloration Covaries with Hiding and Exploratory Behavior in Male Spanish Terrapins

Melanin‐based coloration reflects mostly social status and dominance in males of several species. However, the relation of melanism with other suites of behavioral traits has received less attention. Here, we examined whether the melanic coloration of the shell of male Spanish terrapins Mauremys leprosa was related to several behavioral patterns. To test this, we simulated predatory attacks of different risk levels and measured the time that the turtles spent hiding entirely in their own shells (i.e. appearance times). We also measured the activity level of the turtles in a novel‐environment test. The results showed that melanin‐based coloration was related to antipredatory behavior. Male terrapins with a greater extent of black coloration in their shells demonstrated shorter appearance times after a predator attack. However, this pattern was significant only when turtles were subjected to a high risk. In addition, darker turtles had longer latency times in a novel environment. The results of the present study support the hypothesis that melanin‐based coloration may be associated with some behavioral functions. Our study provides a good framework for future studies concerning the effect of melanin‐coloration on multiple behavioral traits that could be applied to further research using other animal models.     

浅斑近微蛛体形大小对交配行为的影响

理论和实验表明体形大小是许多动物择偶的重要信号。体形大小与产卵大小、最终产卵数量、后代适合度等都有着密切的关系,因此动物利用体形大小为信号来择偶可以获得高的生殖利益。目前研究交配行为和体形大小的关系的文献较少,特别在微蛛亚科。本文以浅斑近微蛛为实验材料,在室内研究了体形大小对交配行为的影响。测量了体重,体长,头胸甲与腹部的长和宽,以头胸甲的面积(头胸甲的长与宽的乘积)为变量来表示体形大小。以雄蛛的行为为依据,将其交配行为划分为4个时期:潜伏期、求偶期、插入期和纳精期,并记录了各个时期的持续时间。通过分析,发现了一种微蛛新的交配模式,即可将浅斑近微蛛的插入期分为三个阶段:首先是大约25min长插入期,接着是一个大约15min的中度插入期,最后是一个大约5min的短插入期。结果表明,在本试验条件下,浅斑近微蛛的体形大小对其交配行为的潜伏期、求偶期、插入期和纳精期的持续时间都没有影响,对插入期的3个阶段的长短也没有影响。     

Testing the role of background matching and self‐shadow concealment in explaining countershading coloration in wild‐caught rainbowfish

Countershading, or dorsal pigmentary darkening (DPD), describes a form of vertically varying coloration, where an animal typically has a dark dorsal surface and a paler ventral side, and is widespread among mammals, birds, reptiles, fishes and insects. DPD is thought to confer concealment from predators and, in terrestrial systems, there is good evidence that the dark–light transition in body coloration acts to conceal the body's shadow. Surprisingly few studies of DPD have been conducted in aquatic environments, and thus it is not known whether the mechanisms of concealment are similar to those that operate in terrestrial habitats. In this study, we determined the role of the light environment and predation risk in determining DPD in wild‐caught populations of a freshwater fish, the western rainbowfish (Melanotaenia australis). We also examined the underlying mechanisms of DPD for concealment by testing the assumptions of background matching and self‐shadow concealment. In a subsequent experiment, we determined whether any observed variation in DPD was maintained when the visual background was manipulated in the laboratory (to induce a change in body coloration). We found that both the amount of downwelling irradiance and the level of predation risk at the collection site affected skin darkness (dorsal, ventral and overall), whereas the ratio of dorsoventral coloration (DPD) was not affected by the parameters considered. The laboratory experiment revealed that fish changed their body coloration to match their visual background, and did so by altering the relative ratio of dorsoventral skin darkness. In contrast with research on terrestrial animals, our findings suggest that the most likely method of achieving crypsis is through background matching, rather than self‐shadow concealment. It is thus possible that differences in the optical characteristics of terrestrial and aquatic environments, and/or variation in the angles at which prey are typically viewed and attacked, have resulted in divergent mechanisms of using DPD to attain crypsis. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 915–928.     

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.     

Foliose Ulva Species Show Considerable Inter‐Specific Genetic Diversity,Low Intra‐Specific Genetic Variation,and the Rare Occurrence of Inter‐Specific Hybrids in the Wild

Foliose Ulva spp. have become increasingly important worldwide for their environmental and financial impacts. A large number of such Ulva species have rapid reproduction and proliferation habits, which explains why they are responsible for Ulva blooms, known as “green tides”, having dramatic negative effects on coastal ecosystems, but also making them attractive for aquaculture applications. Despite the increasing interest in the genus Ulva, particularly on the larger foliose species for aquaculture, their inter‐ and intra‐specific genetic diversity is still poorly described. We compared the cytoplasmic genome (chloroplast and mitochondrion) of 110 strains of large distromatic foliose Ulva from Ireland, Brittany (France), the Netherlands and Portugal. We found six different species, with high levels of inter‐specific genetic diversity, despite highly similar or overlapping morphologies. Genetic variation was as high as 82 SNPs/kb between Ulva pseudorotundata and U. laetevirens, indicating considerable genetic diversity. On the other hand, intra‐specific genetic diversity was relatively low, with only 36 variant sites (0.03 SNPs/kb) in the mitochondrial genome of the 29 Ulva rigida individuals found in this study, despite different geographical origins. The use of next‐generation sequencing allowed for the detection of a single inter‐species hybrid between two genetically closely related species, U. laetevirens, and U. rigida, among the 110 strains analyzed in this study. Altogether, this study represents an important advance in our understanding of Ulva biology and provides genetic information for genomic selection of large foliose strains in aquaculture.     

Sex differences in sexual partner acquisition, retention, and harassment during female homosexual consortships in Japanese macaques

Female Japanese macaques (Macaca fuscata) in certain populations are unusual in that they exhibit male-typical patterns of mounting behavior and sexual-partner preference. The goal of this study was to determine whether female Japanese macaques, from one such population, employ male-typical behavioral tactics to disrupt existing homosexual consortships, as well as to acquire and retain same-sex sexual partners. "Harassment" of homosexual consortships occurred when a sexually motivated, third-party male or female interrupted a consorting female couple by displacing or aggressing them. Sexual harassment was a male-typical strategy for disrupting existing homosexual consortships, but was rarely performed by females. "Intrusions" occurred when a male or female competitor attempted to acquire exclusive access to a female engaged in a homosexual consortship by targeting that female as the focus of competition and her partner as his/her competitor. "Sexual coercion" occurred when one individual alternately sexually solicited and aggressed another individual as part of the same behavioral sequence during an intrusion. Males employed consortship intrusions and sexual coercion when they attempted to acquire female sexual partners that were already engaged in homosexual consortships, but females rarely did so. However, females did employ male-typical patterns of aggressive competition and sexual coercion to retain same-sex sexual partners when confronted with male competitors' attempts to usurp those partners. These results indicate that female sexual activity during homosexual consortships is not uniformly "masculine" in expression, but rather is a mixture of male- and female-typical behaviors.     

Risk‐taking and survival in the House Sparrow Passer domesticus: are plumage ornaments costly?

An important cost of sexual and social colour signals may be increased conspicuousness of the animals to visual predators. Although such predation costs have repeatedly been proposed for various ornaments of birds, especially for melanised and depigmented signals with low presumed production costs, tests of this hypothesis are rare and inconclusive. In this study we investigated whether individual variation in plumage ornamentation was associated with predator-related risk-taking behaviour and short-term survival in free-living House Sparrows Passer domesticus . In a large sample of birds we measured three aspects of coloration used in sexual and social signalling: the size of the melanised black throat patch in males, and the area and conspicuousness of the depigmented wing-bar in both sexes. We measured risk-taking by manipulating the distance of feeders from shelters, and recorded individually ringed birds feeding close to and far from shelter. Sparrows seemed to perceive the farther feeders as more risky, as indicated by the shorter time spent and smaller groups feeding far from rather than close to shelter. However, the use of the more risky (farther) feeders was not related to any of the colour traits we measured, suggesting that Sparrows do not adjust their risk-taking behaviour to their ornamentation. Males (the more ornamented sex) did not take less risk than females. Furthermore, we found no evidence that larger throat patches or more ornamented wing-bars reduced the probability of survival. Our findings were robust and consistent across multiple approaches, even when we controlled for several potential confounding effects. These results do not support the suggestion that melanised and depigmented plumage ornaments have significant predation costs in House Sparrows.     

Chronology of reproductive investment determines predation risk aversion in a felid‐ungulate system

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Leaf‐footed bugs possess multiple hidden contrasting color signals,but only one is associated with increased body size

Antipredatory displays that incorporate hidden contrasting coloration are found in a variety of different animals. These displays are seen in organisms that have drab coloration at rest, but when disturbed reveal conspicuous coloration. Examples include the bright abdomens of mountain katydids and the colorful underwings of hawk moths. Such hidden displays can function as secondary defenses, enabling evasion of a pursuant predator. To begin to understand why some species have these displays while others do not, we conducted phylogenetic comparative analyses to investigate factors associated with the evolution of hidden contrasting coloration in leaf‐footed bugs. First, we investigated whether hidden contrasting coloration was associated with body size because these displays are considered to be more effective in larger organisms. We then investigated whether hidden contrasting coloration was associated with an alternative antipredatory defense, in this case rapid autotomy. We found that leaf‐footed bugs with hidden contrasting coloration tended to autotomize more slowly, but this result was not statistically significant. We also found that the presence of a body size association was dependent upon the form of the hidden color display. Leaf‐footed bugs that reveal red/orange coloration were the same size, on average, as species without a hidden color display. However, species that reveal white patches on a black background were significantly larger than species without a hidden color display. These results highlight the diversity of forms that hidden contrasting color signal can take, upon which selection may act differently.     

Function‐related Drivers of Skull Morphometric Variation and Sexual Size Dimorphism in a Subterranean Rodent,Plateau Zokor (Eospalax baileyi)

Sexual dimorphism is prevalent in most living organisms. The difference in size between sexes of a given species is generally known as sexual size dimorphism (SSD). The magnitude of the SSD is determined by Rensch's rule where size dimorphism increases with increasing body size when the male is the larger sex and decreases with increasing average body size when the female is the larger sex. The unique underground environment that zokors (Eospalax baileyi) live under in the severe habitat of the Qinghai‐Tibetan Plateau (QTP) could create SSD selection pressures that may or may not be supported by Rensch's rule, making this scientific question worthy of investigation. In this study, we investigated the individual variation between sexes in body size and SSD of plateau zokors using measurements of 19 morphological traits. We also investigated the evolutionary mechanisms underlying SSD in plateau zokors. Moreover, we applied Rensch's rule to all extant zokor species. Our results showed male‐biased SSD in plateau zokors: The body‐ and head‐related measurements were greater in males than in females. Linear regression analysis between body length, body weight, and carcass weight showed significant relationships with some traits such as skull length, lower incisor length, and tympanic bulla width, which might support our prediction that males have faster growth rates than females. Further, the SSD pattern corroborated the assumption of Rensch's rule in plateau zokors but not in the other zokor species. Our findings suggest that the natural underground habitat and behavioral differences between sexes can generate selection pressures on male traits and contribute to the evolution of SSD in plateau zokors.     

Scaling of Size and Dimorphism in Primates II: Macroevolution

Previous researchers found positive scaling of body size and sexual size dimorphism (SSD) in primates, known as Rensch's rule. The pattern is present in Haplorhini, but absent in Strepsirhini. I found that positive evolutionary correlations between size and SSD drive positive scaling relationships within Haplorhini as a whole and Platyrrhini, Cercopithecinae, Colobinae, and Hominoidea individually at the generic level and higher, but that evolutionary correlations within genera in these clades are often nonsignificant or negative. I suggest that positive evolutionary correlations result from greater change in male than in female size, usually because of sexual selection acting on polygynous populations. I suggest that negative evolutionary correlations result from greater change in female size, owing to either natural selection or, in Callitrichidae, sexual selection acting on polyandrous populations. The high incidence of negative evolutionary correlations within Haplorhini suggests a relatively large influence of natural selection on SSD, at least with regard to differences in SSD between congeners. I propose two possible explanations for the difference in intrageneric and supergeneric evolutionary patterns: 1) natural selection is a relatively weak force for modifying SSD and has a noticeable effect only when one compares related species experiencing similar levels of sexual selection, and 2) natural selection is a relatively strong force for modifying SSD but is less likely than sexual selection to affect higher level taxonomic comparisons noticeably because of the cumulative effect over time of marginal differences in mortality rates of these two types of selection. I discuss types of data required to test these explanations and implications for reconstructing fossil behavior.     

Sniffing Behavior in Lemur catta: Seasonality, Sex, and Rank

We analyzed the role of direct olfactory investigation in relation to seasonality, sex, and female rank via of a longitudinal study on 16 adult ring-tailed lemurs living in two groups at the Pistoia Zoo (Tuscany, Italy). The observations took place from May 1997 to March 1999 and lasted >1,500 h. Males were olfactorily more active than females except for skin licking, which also appeared to be associated with affiliative behaviors. Both sexes had peak frequency of direct olfactory monitoring during the reproductive season; contrarily, skin licking showed apparently random fluctuations. There is a significant positive correlation between rank and sniffing genitals performed by females on other females and a significant negative correlation between rank and sniffing genitals received: high-ranking females are mostly actors, while low-ranking females are mostly receivers. The continuous follow-up of the reproductive conditions of potential competitors suggests that sniffing genitals might play a role in female reproductive strategies.     

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捕食风险与动物行为及其决策的关系边疆晖樊乃昌（中国科学院西北高原生物研究所,西宁８１０００１）（浙江师范大学,金华３１００００）ＴｈｅＥｆｅｃｔｏｆＰｒｅｄａｔｉｏｎＲｉｓｋｏｎＡｎｉｍａｌｓ’ＢｅｈａｖｉｏｒａｎｄＴｈｅｉｒＤｅｃｉｓｉｏｎ．Ｂｉａ...     

Inter‐ and intrasex habitat partitioning in the highly dimorphic southern elephant seal

Partitioning resources is a key mechanism for avoiding intraspecific competition and maximizing individual energy gain. However, in sexually dimorphic species it is difficult to discern if partitioning is due to competition or the different resource needs of morphologically distinct individuals. In the highly dimorphic southern elephant seal, there are intersexual differences in habitat use; at Iles Kerguelen, males predominantly use shelf waters, while females use deeper oceanic waters. There are equally marked intrasexual differences, with some males using the nearby Kerguelen Plateau, and others using the much more distant Antarctic continental shelf (~2,000 km away). We used this combination of inter and intrasexual behavior to test two hypotheses regarding habitat partitioning in highly dimorphic species. (a) that intersexual differences in habitat use will not appear until the seals diverge in body size and (b) that some habitats have higher rates of energy return than others. In particular, that the Antarctic shelf would provide higher energy returns than the Kerguelen Shelf, to offset the greater cost of travel. We quantified the habitat use of 187 southern elephant seals (102 adult females and 85 subadult males). The seals in the two groups were the same size (~2.4 m) removing the confounding effect of body size. We found that the intersexual differences in habitat use existed before the divergence in body size. Also, we found that the amount of energy gained was the same in all of the major habitats. This suggests that the use of shelf habitats by males is innate, and a trade‐off between the need to access the large benthic prey available on shelf waters, against the higher risk of predation there. Intrasexual differences in habitat use are another trade‐off; although there are fewer predators on the Antarctic shelf, it is subject to considerable interannual fluctuations in sea‐ice extent. In contrast, the Kerguelen Plateau presents more consistent foraging opportunities, but contains higher levels of predation. Habitat partitioning in this highly dimorphic species is therefore the result of complex interplay of life history strategies, environmental conditions and predation pressure.     

Scaling of Size and Dimorphism in Primates I: Microevolution

I used a new quantitative genetics model to predict relationships between sex-specific body size and sex-specific relative variability when populations experience differences in relative intensity of sex-specific selection pressures—stronger selection on males or females—and direction of selection: increase or decrease in size. I combined Lande's (Evolution 34: 292–305) model for the response of sex-specific means to selection with a newly derived generalization of Bulmer's (Am. Nat. 105: 201–211) model for the response of relative variability to selection. I used this combined response model to predict correlations of sex-specific size and relative variability under various starting conditions, which one can compare to correlations between closely related primate populations. One can then compare predicted patterns of sex-specific selection pressures to social and ecological variables pertaining to those populations to identify likely forces producing microevolutionary change in sexual size dimorphism (SSD). I provide examples of this approach for populations representing three taxa: Papio anubis, Saguinus mystax, and Cercopithecus aethiops pygerythrus. Model results suggest that microevolutionary changes in SSD can result from greater selection acting on males or females, and that natural selection or natural and sexual selection combined, rather than sexual selection alone, may sometimes explain sex-specific selection differentials.     

The detectability of the colour pattern in the aposematic firebug,Pyrrhocoris apterus: an image‐based experiment with human ‘predators’

Crypsis and aposematism are often regarded as two opposite protective strategies. However, there is large variation in prey appearance within both strategies. In this article, we investigated the conspicuousness of the aposematic red‐and‐black firebug, Pyrrhocoris apterus, by presenting images of natural and digitally manipulated phenotypes in their natural habitat on a computer screen to human ‘predators’, and comparing the detection times. We asked whether the natural colour pattern can be made more or less conspicuous by rearranging the spatial distribution of colour elements. Hence, we created a phenotype in which the black colour elements were moved to the body outline to test for a possible disruptive effect. In the ‘black’ and ‘red’ manipulations, we removed one of the two colours, creating two uniform colour variants. We found that some of our manipulations increased, but none reduced, the detection time significantly; this indicates that the naturally coloured firebug is highly conspicuous. The detection time varied among backgrounds and there was a significant relationship between detection time and chromatic similarity between the bug and the background for the natural and black phenotypes. Although background colour composition has an important effect on the signal, we argue that the coloration of P. apterus has evolved for high conspicuousness. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 806–816.     

Sex and social costs of escaping in the striped plateau lizard Sceloporus virgatus

As a predator approaches, prey base decisions about when toflee on a balance between degree of predation risk and costsof escaping. Lost opportunities to perform activities that mayincrease fitness are major escape costs. Paramount among theseare chances to increase fitness by courting and mating and bydriving away sexual rivals. Because sexual selection imposesdifferent social demands on the sexes, social opportunitiescan have different consequences for males and females, but effectsof sex differences in social opportunity costs on escape behaviorare unknown. We conducted a field experiment showing that malestriped plateau lizards (Sceloporus virgatus) given the opportunityto court or perform aggressive behavior permit closer approachbefore fleeing, but females do not. Males allowed a simulatedpredator to approach closer before initiating escape if a tetheredmale or female rather than a control stimulus was introducedto them, but females initiated escape at similar distances inall conditions. For males, a trade-off between the greater predationrisk accepted before fleeing due to the likelihood of enhancingfitness by sexual or aggressive behavior accounts for closerapproach allowed in the presence of conspecifics. Mating opportunitiesare not limiting for females in most species and females oftenhave little to gain by interacting aggressively with other females.Therefore, presence of a conspecific male or female may notjustify taking greater risk. Results confirm the predictionof optimal escape theory that flight initiation decreases ascost of escaping increases. The sex difference in effect ofpresence of conspecifics on flight initiation distance is aconsequence of the sex difference in costs of escaping.