Deterring predators, daunting opponents or drawing partners? Signaling rates across diverse contexts in the lizard Anolis sagrei

Although the visual display behavior in Anolis lizards has received ample attention, the function of dewlap extensions (DE), push-ups (PU), and head-nods (HN) in general, and in Anolis sagrei in particular, remains highly equivocal. Therefore, our primary goal was to quantify the display rates of these visual signal types (DE, PU, and HN) in a variety of contexts, using A. sagrei as study species. To our knowledge, this is the first study to test individuals of both sexes in a repeated-measures design across multiple contexts, including predator, non-predator and social interactions (mirror, male–male, male–female, female–male). We found that males have an overall higher signaling rate than females across all contexts. In addition, we found that lizards of both sexes exhibited higher display rates in the presence of conspecifics than when confronted with a predator or non-predator, suggesting that DE, PU, and HN function in intraspecific communication, not in predator deterrence. Whereas females did not significantly raise display rates in a consexual and heterosexual context with respect to subject-alone context, males did. The PU signal type only appears to play a major role for A. sagrei males during aggressive encounters. During heterosexual interactions, increased frequencies of all signal types suggest that DE, PU, and HN are essential for male courtship. Finally, we suggest that intersexual selection is probably a driving force for frequency-related dewlap use in both sexes. In contrast, pronounced intersexual differences were detected for PU and HN rates within a social context.     

Consensus among females in their choice of males in the guppy Poecilia reticulata

We studied female guppies from two populations (Trinidad and Jemez Springs, New Mexico) to determine their mating preferences, the degree of consensus among females for particular male phenotypes, and the variation among females for the types of traits they prefer. We recorded the visual responses of 68 Trinidad and 10 Jemez females in paired male trials. The three sexually-selected male traits quantified were the area of orange color and iridescence on the body, and display behavior. Females from the Trinidad population agreed in their choice of males in three of the eight replicates, and this agreement was based on display rate rather than male color patterns. Females from the Jemez population showed no agreement in their preferences of males. Mating preferences of females varied both between populations and among females of a population. In both populations, female preferences were based primarily on courtship intensity and only secondarily on color pattern. However, females from both populations differed in the relative importance of orange color and iridescence. The fact that females differ in criteria for evaluating males has important implications for selection and maintenance of color polymorphisms and for the interactions among multiple secondary sexual traits of males in the guppy. Received: 5 December 1995/Accepted after revision: 7 June 1996     

Frequency-dependent male mate harassment and intra-specific variation in its avoidance by females of the damselfly Ischnura elegans

We focused on male harassment on different female color morphs of the damselfly Ischnura elegans and on variation in morph-specific mating avoidance tactics by females. In I. elegans, one of the female morphs is colored like the conspecific male (andromorphs) while the other morphs are not (gynomorphs). Our first goal was to quantify morph-specific male mating attempts, hence male harassment, in populations with manipulated population parameters (densities, sex ratios, and proportion of andromorphs). Second, we examined the female's perspective by looking for potential differences in morph-specific mating avoidance tactics and success of those tactics in a natural population. Differences in population conditions did influence the number of male mating attempts per morph. The less frequent female morph was always subject to fewer mating attempts, which contradicts earlier hypotheses on mimicry, but supports those that assume that males learn to recognize female morphs. Gynomorphs occupy less open habitat and often fly away when a male approaches, while andromorphs use more open habitat, do not fly large distances and directly face approaching males. Female morphs did not differ in the proportion of successful mating-avoidance attempts. Our results suggest that the maintenance of the color polymorphism is most probably the result of interactive selective forces depending on variation in all population conditions, instead of solely density- or frequency-dependent selection within populations.     

Resource distributions among habitats determine solitary bee offspring production in a mosaic landscape.

Within mosaic landscapes, many organisms depend on attributes of the environment that operate over scales ranging from a single habitat patch to the entire landscape. One such attribute is resource distribution. Organisms' reliance on resources from within a local patch vs. those found among habitats throughout the landscape will depend on local habitat quality, patch quality, and landscape composition. The ability of individuals to move among complementary habitat types to obtain various resources may be a critical mechanism underlying the dynamics of animal populations and ultimately the level of biodiversity at different spatial scales. We examined the effects that local habitat type and landscape composition had on offspring production and survival of the solitary bee Osmia lignaria in an agri-natural landscape in California (U.S.A.). Female bees were placed on farms that did not use pesticides (organic farms), on farms that did use pesticides (conventional farms), or in seminatural riparian habitats. We identified pollens collected by bees nesting in different habitat types and matched these to pollens of flowering plants from throughout the landscape. These data enabled us to determine the importance of different plant species and habitat types in providing food for offspring, and how this importance changed with landscape and local nesting-site characteristics. We found that increasing isolation from natural habitat significantly decreased offspring production and survival for bees nesting at conventional farms, had weaker effects on bees in patches of seminatural habitat, and had little impact on those at organic farm sites. Pollen sampled from nests showed that females nesting in both farm and seminatural habitats relied on pollen from principally native plant species growing in seminatural habitat. Thus connectivity among habitats was critical for offspring production. Females nesting on organic farms were buffered to isolation effects by switching to floral resources growing at the farm site when seminatural areas were too distant. Overall local habitat conditions (farm management practices) can help bolster pollinators, but maintaining functional connectivity among habitats will likely be critical for persistence of pollinator populations as natural habitats are increasingly fragmented by human activities.     

Negative correlation between male ornament size and female preference intensity in a wild guppy population

Although traditional sexual selection theories for the evolution of ornamental male traits often assume consistency in female preferences for the traits over time, recent theories predict plasticity in female mate preferences and the contribution of plastic female preferences to the maintenance of polymorphism in male sexual ornaments. However, the plasticity of female preferences and its influences on male ornaments are almost unknown in natural populations. Here we examined both the intensity of female preferences and the exaggeration of a male ornament (relative area of orange spots) in a wild population of the guppy (Poecilia reticulata) every few months over a period of 3 years. Results indicated that female preference intensity was negatively correlated with the relative area of orange spots of males. In addition, we found a positive correlation between female preference intensity and the relative area of orange spots of males of their offspring generation. Because the relative area of orange spots of male guppies is a heritable trait, female preferences might have a strong influence on male orange spot size in the next generation. This study provides the first evidence of plastic female preferences depending on the scarcity of males with large ornaments in a natural population. Results of this study imply a possible contribution of female preferences for the maintenance of variation in male ornamentation.     

Adjustment of female reproductive investment according to male carotenoid-based ornamentation in a gallinaceous bird

Carotenoid-based ornaments (many yellow–orange–red colourations) may signal the genetic or parental quality of the bearer. Thus, their expression could influence the amount of resources/energy that the mate will invest in the production of offspring, thereby optimising its reproductive fitness. The differential allocation hypothesis (DAH) predicts that females mated with more attractive males should lay more and better eggs. This has been explored only in few bird species with carotenoid-based traits. We tested this hypothesis in the red-legged partridge (Alectoris rufa), a gallinacean with very variable laying capacity. Both sexes display carotenoid-based ornamentation that gradually fades throughout the laying period. Here, the redness of beak and eye rings of captive males was intensified after mating by means of paint. The proportion of females that laid eggs did not differ between treatments. Amongst laying females, those mated with colour-enhanced males (experimental females) tended to lay earlier and produced significantly more eggs than controls, but of similar quality (egg mass and composition). We additionally investigated whether male attractiveness influenced egg components depending on the clutch size and laying sequence. The testosterone level in eggs from experimental females was positively related to the laying order, whereas control eggs did not show any trend. Our results provided mixed support for the DAH, but nevertheless revealed that female red-legged partridges may adjust their breeding investment according to male carotenoid-based ornamentation.     

Mate choice games, context-dependent good genes, and genetic cycles in the side-blotched lizard, Uta stansburiana

According to mate choice models, a female should prefer males with traits that are reliable indicators of genetic quality which the sire can pass on to their progeny. However, good genes may depend on the social environment, and female choice for good genes should be context dependent. The side-blotched lizard, Uta stansburiana, exhibits genetically based throat colors (orange, blue, or yellow) that could be used as a sexually selected signal since they reliably predict the genetic quality of mates. The frequencies of male and female morphs cycle between years, and both male and female morphs have an advantage when rare; thus genetic quality will depend on morph frequency. A female should choose a sire that maximizes the reproductive success of both male and female progeny. We examine a game theoretical model that predicts female mate choice as a function of morph frequency and population density. The model predicts the following flexible mate choice rule: both female morphs should prefer rare males in ’boom years’ of the female cycle (e.g., ’rarest-of-N rule’), but prefer orange males in ’crash years’ of the female cycle (’orange-male rule’). Cues from the current social environment should be used by females to choose a mate that maximizes the future reproductive success of progeny, given the social environment of the next generation. We predict that the cue is the density of aggressive orange females. In the side-blotched lizard, cycling mate choice games and context-dependent mate choice are predicted to maintain genetic variation in the presence of choice for good genes. Received: 8 March 2000 / Revised: 26 August 2000 / Accepted: 4 September 2000     

Is female promiscuity constrained by the presence of her social mate? An experiment with bluethroats <Emphasis Type="Italic">Luscinia svecica</Emphasis>

Extra-pair paternity is common in birds and much research has focussed on the selective advantage of extra-pair matings for both sexes. In contrast, little attention has been given to the fact that in most species the majority of offspring are sired by the social male. We investigated whether extra-pair matings of female bluethroats (Luscinia svecica) are constrained by the presence of the pair male, by detaining males in cages on their territories for one morning during the peak of female fertility. The proportion of offspring sired by extra-pair males was higher in broods where males had been detained (35%) than in control broods (16%), while the proportion of broods that had at least one extra-pair offspring did not differ significantly between experimental (65%) and control broods (44%). Within the experimental group, levels of extra-pair paternity were not related to the day of experiment in relation to start of egg laying, but males caught early in the morning lost more paternity than males caught later on. Our results show that pair males exert constraints on the frequency of extra-pair paternity by being present during the period of peak fertility, which could be a direct effect of their mate guarding effort and/or due to an advantage in sperm competition for pair males.     

Armaments and ornaments in the rock sparrow: a possible dual utility of a carotenoid-based feather signal

Females often base their mating preferences on male sexual secondary traits that are used to settle contests among males. Such traits are likely to be honest indicators of male quality if they are constantly used during costly male–male agonistic interactions. Carotenoid signals have been shown to work as a handicap because they are costly to produce. However, the role of carotenoids as “honest” signals during male contests is less clear, and it is not known whether a carotenoid-based trait can serve in both male–male competition and female choice. In this study, we studied the dual function of a carotenoid feather ornament in the rock sparrow (Petronia petronia), a bird species in which both sexes have a yellow throat patch whose size positively correlates with phenotypic measures. First, we investigated, in a field study, whether the size of a male’s yellow patch correlates with his ability to acquire a territory. Second, we tested the signal function of the yellow patch in two male–male interaction in captivity experiments. Finally, we measured female preference for males differing in throat patch size in a mate choice experiment. Our experiments revealed that the size of a male’s throat patch positively correlated with the number of nest boxes he was able to defend. Moreover, in controlled conditions, males with relatively large yellow patches had earlier access to food than those with small patches. Also, in an experiment in which a dummy rock sparrow with an experimentally manipulated yellow patch was positioned near a feeder, latency to feed by focal birds positively correlated with dummy patch size. Lastly, in a dichotomous mate choice experiment, females showed a proximity preference for males whose patch was experimentally enlarged. Taken together, these results suggest that the same carotenoid feather signal may be used in both male–male competition and female choice in this passerine bird.     

Spatial scale of local breeding habitat quality and adjustment of breeding decisions

Experimental studies provide evidence that, in spatially and temporally heterogeneous environments, individuals track variation in breeding habitat quality to adjust breeding decisions to local conditions. However, most experiments consider environmental variation at one spatial scale only, while the ability to detect the influence of a factor depends on the scale of analysis. We show that different breeding decisions by adults are based on information about habitat quality at different spatial scales. We manipulated (increased or decreased) local breeding habitat quality through food availability and parasite prevalence at a small (territory) and a large (patch) scale simultaneously in a wild population of Great Tits (Parus major). Females laid earlier in high-quality large-scale patches, but laying date did not depend on small-scale territory quality. Conversely, offspring sex ratio was higher (i.e., biased toward males) in high-quality, small-scale territories but did not depend on large-scale patch quality. Clutch size and territory occupancy probability did not depend on our experimental manipulation of habitat quality, but territories located at the edge of patches were more likely to be occupied than central territories. These results suggest that integrating different decisions taken by breeders according to environmental variation at different spatial scales is required to understand patterns of breeding strategy adjustment.     

Experimental increase in food supply influences the outcome of within-family conflicts in Tengmalm’s owl

Within a family there are conflicts of interest between parents and offspring, and between male and female parents, over the supply of parental care. The observed pattern of parental care is the outcome of negotiations within the family, and may be influenced by environmental factors such as food abundance. We experimentally increased food supply to ten Tengmalm’s owl (Aegolius funereus) nests from hatching to fledging, mimicking natural cached prey. Ten un-supplemented nests served as controls. Parents and offspring were fitted with radio-tags. Food provisioning by parents was measured both in the (1) mid- and (2) late nestling stage and in the (3) early and (4) late post-fledging stage. In response to food supplementation, both males and females reduced food provisioning, but the effect was more pronounced in females. Females generally contributed much less to food provisioning than males, and food supplementation increased the difference between the sexes. Mass loss during the brooding stage was substantially lower for supplemented than for control females. Food supplementation did not improve offspring survival, and had no effect on body measurements of nestlings. In conclusion, parents of both sexes used the increased food supply to reduce the costs of caring for their current offspring, but females responded more strongly than males.     

Cave molly females (Poecilia mexicana, Poeciliidae, Teleostei) like well-fed males

We examined the preference of Atlantic molly females (Poecilia mexicana) to associate with a well-fed or a starved male in simultaneous choice tests. Females from three different populations were tested in three treatments: (1) the females could choose on the basis of multiple cues from the males (visual plus non-visual); (2) only non-visual cues could be perceived in darkness, (3) only visual cues were presented. The three tested populations differ clearly in their ecology: one population occurs in a typical river habitat, the second one in a milky sulfur creek outside a cave, and the third population occurs in a cave habitat (cave molly). In the river-dwelling population, females never showed a preference. In the population from the sulfur creek, females preferred to associate with the well-nourished male when visual cues from the males were available. Only cave molly females exhibited a strong preference for well-nourished males in all treatments. A morphological comparison demonstrated that wild-caught males from river habitats are typically in a good nutritional state. In the sulfur creek, males showed signs of starvation. Cave molly males were in an even worse nutritional state. In the cave population, saturated males probably indicate high fitness, thereby driving the evolution of the preference for good male nutritional state.Communicated by K. Lindström     

Male bill colour and age are associated with parental abilities and breeding performance in blackbirds

In monogamous bird species, male parental investment may influence offspring fitness and females may gain advantages through mating with males providing extensive paternal care. However, paternal care is a benefit that can only be assessed indirectly because mate choice precedes paternal activities. Individual quality and age, both signalled by morphological characteristics, may reflect parental abilities. Because they may reflect individual foraging abilities, carotenoid-based colorations have been proposed to honestly signal parental quality. The blackbird (Turdus merula), a socially monogamous species, exhibits biparental care and males show bills that vary from pale yellow to orange due to carotenoid pigments. In this study, we investigated whether male bill colour and age are associated with parental ability. Our results suggest that males with more orange bills and older males are better fathers. Indeed, male visit rate increased with their bill colour index independently of age, and brood condition was higher for adult males, compared to yearlings, independently of bill colour. Overall, the number of fledglings produced was positively influenced by both the age of males and the colour intensity of their bills. Males with more orange bills and adults had a greater number of fledglings and these males also had higher levels of prolactin, a hormone known to promote parental care. This latter finding suggests that prolactin may be the link between carotenoid based colorations and the intensity of paternal effort. Thus, male bill colour seems to honestly reveal male physiological adjustment to paternal activities.     

Female reproductive qualities affect male painted turtle (Chrysemys picta marginata) reproductive success

Male reproductive success (RS) in polygamous species with minimal social systems is often determined by the number of mates. However, because male RS is translated through females, the number of offspring sired can also be influenced by female qualities. Empirically sufficient data to document how tradeoffs between mate number and quality influence male RS are seldom available for long-lived, iteroparous species. We combined long-term life history data (1983–2006) on the E. S. George Reserve (ESGR, MI, USA) with parentage data from 155 clutches of 59 female painted turtles (Chrysemys picta marginata) of varying reproductive frequencies (2003–2006) to determine the relative contribution of female numbers and qualities on male RS. One previously documented trait of female painted turtles that can have substantial influences on male RS is repeat paternity through the use of stored sperm to fertilize over 95 % of within-year clutches. In addition, our study found that second-clutch producing female painted turtles on the ESGR have higher among-year reproductive frequencies than do first-clutch only females. Multiple paternity was detected in 14.1 % of clutches (min-max?=?6.1–30.0 % annually), and the number of mates of both sexes was low annually (males 1.0; females 1.2) and over 4 years (males 1.1; females 1.7). Among successful males, RS varied substantially (1–32 offspring) and was strongly influenced by the combination of female reproductive frequency and repeat paternity (>38 % among years), but not mate number. Low mate number for both sexes was unexpected in a species without complex mating behaviors or parental care.     

Sex-specific costs of hatching last: an experimental study on herring gulls (<Emphasis Type="Italic">Larus argentatus</Emphasis>)

An organism’s pattern of development can have important long-term fitness effects. In species where the sexes differ in size or other phenotypic traits, they may also have different optimal developmental rates. This influences both parental sex allocation strategies and susceptibility of the sexes to early developmental conditions. However, sex differences in developmental rate and vulnerability to environment during the embryonic period are not well understood. In birds, sibling competition and hatching asynchrony may select for accelerated embryonic development of the last offspring in order to reduce their competitive disadvantage after hatching. They may advance their hatching in response to vocal stimuli by the older siblings. It is, however, unclear whether this flexibility in developmental rates is sex specific. In this study, we experimentally manipulated between-embryo contact and tested whether this affected the pre-natal developmental rate and post-hatching performance of male and female offspring from last-laid eggs in the herring gull. Post-hatching performance was measured both in competitive and non-competitive situations. Among young incubated in isolation, males hatched faster than females, but both sexes fledged in similar, relatively good condition. Among young incubated with normal between-embryo contact, hatching time did not differ between sexes, but males fledged in poorer condition than females, regardless of whether they were reared singly or in a brood. These results suggest that male and female offspring differ in their ability to mitigate the costs of hatching asynchrony.     

Effectiveness of Corridors Relative to Enlargement of Habitat Patches

Abstract:  The establishment of biological corridors between two otherwise isolated habitat patches is a common yet contentious strategy for conserving populations in fragmented landscapes. We compared the effectiveness of corridors with the effectiveness of an alternate conservation strategy, the enlargement of existing habitat patches. We used a spatially explicit population model that simulated population size in two kinds of patches. One patch had a corridor that connected it to a larger "source" patch and the other patch was unconnected and enlarged at the periphery by an area the same size as the corridor. Patch isolation, corridor width, patch size, and the probability that individuals would cross the border from habitat to matrix were varied independently. In general, population size was greater in enlarged patches than in connected patches when patches were relatively large and isolated. Corridor width and the probability of crossing the border from habitat to matrix did not affect the relative benefit of corridors versus patch enlargement. Although biological corridors may mitigate potential effects of inbreeding depression at long time scales, our results suggest that they are not always the best method of conserving fragmented populations.     

Male lagoon gobies, <Emphasis Type="Italic">Knipowitschia panizzae</Emphasis>, prefer more ornamented to larger females

Female ornamentation may be directly sexually selected, by male choice or female competition, or occurs as the result of a genetic correlation, arising from sexual selection on males. However, increasing evidence supports the former hypothesis, suggesting that males actively choose their partner preferring traits indicative of female quality. In the lagoon goby, Knipowitschia panizzae, a polygynous species whose males perform parental care to eggs, body length and the size of a sex-specific yellow patch on the belly are known to be reliable indicators of female fecundity. In this paper, we tested, using dummies, the male’s mating preferences for female body and yellow belly patch sizes. The two experimental trials in which a single female trait was variable showed that males prefer a larger belly patch and a larger body size, indicating that both these characters are selected by male mate choice. However, when faced with dummies exhibiting an inverse combination of body and belly patch sizes (experiment 3), males significantly preferred the smaller ones with larger yellow belly patches. A calculation of dummy theoretical fecundity reveals that in the first two experiments, males would have received an immediate benefit from their choice in terms of egg number, whereas in the third one, males chose partners that would have provided them with fewer eggs. The male lagoon goby preference for females with larger belly patches, regardless of their size, suggests that this trait, in addition to indicating fecundity, conveys information about other aspects of female and/or egg quality.     

The relationship between dewlap size and performance changes with age and sex in a Green Anole (Anolis carolinensis) lizard population

Ornaments are believed to signal an individual's ability to reproduce successfully and/or survive. Since an individual's fitness is often influenced by multiple traits (e.g. number of copulations, ability to acquire nest sites or to escape predators), which are difficult to quantify simultaneously, we examine performance traits (bite force, jumping performance) believed to be relevant to an individual's fitness. Specifically, we ask whether variation in dewlap size is related to variation in body size, bite force and jumping ability in the lizard Anolis carolinensis. Our results show that dewlap size is correlated with jumping capacity across all individuals, whereas the relationships between dewlap size, body size and bite force differ depending on sex/age class. We argue that selection against relatively large dewlaps at the transition between small mature and large mature males might be responsible for the lack of a relationship within large males. The absence or presence of a correlation between dewlap size and bite force, on the other hand, might be explained by differences in behaviour, such as territory establishment, anti-predator tactics, and/or mate choice. Our work thus suggests that selective forces influencing the evolution of ornaments may operate differently on different sexes and life-history stages.     

Role of Patch Size, Disease, and Movement in Rapid Extinction of Bighorn Sheep

Abstract: The controversy (  Berger 1990, 1999 ; Wehausen 1999 ) over rapid extinction in bighorn sheep ( Ovis canadensis ) has focused on population size alone as a correlate to persistence time. We report on the persistence and population performance of 24 translocated populations of bighorn sheep. Persistence in these sheep was strongly correlated with larger patch sizes, greater distance to domestic sheep, higher population growth rates, and migratory movements, as well as to larger population sizes. Persistence was also positively correlated with larger average home-range size ( p = 0.058, n = 10 translocated populations) and home-range size of rams ( p = 0.087, n = 8 translocated populations). Greater home-range size and dispersal rates of bighorn sheep were positively correlated to larger patches. We conclude that patch size and thus habitat carrying capacity, not population size per se, is the primary correlate to both population performance and persistence. Because habitat carrying capacity defines the upper limit to population size, clearly the amount of suitable habitat in a patch is ultimately linked to population size. Larger populations (250+ animals) were more likely to recover rapidly to their pre-epizootic survey number following an epizootic ( p = 0.019), although the proportion of the population dying in the epizootic also influenced the probability of recovery ( p = 0.001). Expensive management efforts to restore or increase bighorn sheep populations should focus on large habitat patches located ≥23 km from domestic sheep, and less effort should be expended on populations in isolated, small patches of habitat.     

Males in the shade: habitat use and sexual segregation in the Galápagos sea lion (Zalophus californianus wollebaeki)

In many sexually dimorphic species adult sexes tend to segregate socially, spatially, or in habitat use. Several hypotheses have been formulated regarding underlying mechanisms. We investigated terrestrial habitat use and sexual segregation in a tropical otariid, the Galápagos sea lion (Zalophus californianus wollebaeki), where most of the hypotheses can be ruled out a priori. Factors relating to thermoregulation and costs of locomotion were of prime importance for habitat use. Habitats directly adjacent to the sea, with simple structured flat surfaces, shade, and tide pools were most frequented, but sexes and age classes differed in their usage patterns. Sexual segregation, both spatial and by habitat was pronounced in the reproductive period (RP), but remained high during the nonreproductive period (NRP). A GLM model of habitat use showed that in both seasons adult males frequented habitat types that adult females and other age classes used much less. Males were most abundant in suboptimal inland habitats, which offered only shade for cooling. Females with newborns differed in habitat use from females with older offspring and lone females. Spatial and habitat segregation are explained most parsimoniously as by-products of social processes, primarily intrasexual competition and female avoidance of male harassment, linked to the polygynous mating system.Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.