Dawn chorus and female behaviour in the Willow Tit Parus montanus

We studied the effect of female behaviour on the dawn chorus of Willow Tits Parus montanus. Ten females were temporarily kept in their nest holes in order to delay their dawn emergence. Simultaneously, their mates' singing behaviour was measured. We compared the control dawn behaviour (during which the females emerged when they wanted) with that during the experimental delay. The dawn song by the males lasted longer when the females' emergences were delayed. Our results show that the duration of the dawn chorus is directly related to the female's behaviour. We suggest that at least one function of the dawn song is intrapair signalling, which is often linked to sperm competition.     

New insights into sex chromosome evolution in anole lizards (Reptilia,Dactyloidae)

Anoles are a clade of iguanian lizards that underwent an extensive radiation between 125 and 65 million years ago. Their karyotypes show wide variation in diploid number spanning from 26 (Anolis evermanni) to 44 (A. insolitus). This chromosomal variation involves their sex chromosomes, ranging from simple systems (XX/XY), with heterochromosomes represented by either micro- or macrochromosomes, to multiple systems (X₁X₁X₂X₂/X₁X₂Y). Here, for the first time, the homology relationships of sex chromosomes have been investigated in nine anole lizards at the whole chromosome level. Cross-species chromosome painting using sex chromosome paints from A. carolinensis, Ctenonotus pogus and Norops sagrei and gene mapping of X-linked genes demonstrated that the anole ancestral sex chromosome system constituted by microchromosomes is retained in all the species with the ancestral karyotype (2n?=?36, 12 macro- and 24 microchromosomes). On the contrary, species with a derived karyotype, namely those belonging to genera Ctenonotus and Norops, show a series of rearrangements (fusions/fissions) involving autosomes/microchromosomes that led to the formation of their current sex chromosome systems. These results demonstrate that different autosomes were involved in translocations with sex chromosomes in closely related lineages of anole lizards and that several sequential microautosome/sex chromosome fusions lead to a remarkable increase in size of Norops sagrei sex chromosomes.     

Hormones, sexual signals, and performance of green anole lizards (Anolis carolinensis)

The evolutionary processes that result in reliable links between male signals and fighting capacity have received a great deal of attention, but the proximate mechanisms underlying such connections remain understudied. We studied a large sample of male green anole lizards (Anolis carolinensis) to determine whether testosterone or corticosterone predicted dewlap size and/or bite-force capacity, as dewlap size is known to be a reliable predictor of bite-force capacity in territorial males. We also examined whether these relationships were consistent between previously described body size classes ("lightweights" and "heavyweights"). Heavyweights had 50% higher testosterone concentrations than lightweights during the breeding season, suggesting a mechanism for the disproportionately larger heads and dewlaps and higher bite-forces of heavyweights. Plasma testosterone concentrations were positively correlated with dewlap size and bite-force performance in lightweights (but not heavyweights) but only because of mutual intercorrelation of all three variables with body size. We suggest two possibilities for the relationship between testosterone levels and body size: (1) testosterone promotes growth in this species or (2) smaller sexually mature males are unable to compete with larger males such that the benefits of elevated testosterone do not outweigh the costs. Corticosterone levels did not differ between the male morphs, and lightweights, but not heavyweights, showed an inverse relationship between testosterone levels and corticosterone levels. Our results suggest that testosterone is important for traits related to dominance in adult male green anoles and may influence the ability to compete with rivals via fighting ability or through the use of signals.     

A performance-based cost to honest signalling in male green anole lizards (Anolis carolinensis)

Sexual signals are considered costly to produce and maintain under the handicap paradigm, and the reliability of signals is in turn thought to be maintained by these costs. Although previous studies have investigated the costly nature of signal production, few have considered whether honesty might be maintained not by the costliness of the signal itself, but by the costs involved in producing the signalled trait. If such a trait is itself costly to produce, then the burden of energetic investment may fall disproportionately on that trait, in addition to any costs of signal maintenance that may also be operating. Under limited resource conditions, these costs may therefore be great enough to disrupt an otherwise reliable signal-to-trait relationship. We present experimental evidence showing that dietary restriction decouples the otherwise honest relationship between a signal (dewlap size) and a whole-organism performance trait (bite force) in young adult male Anolis carolinensis lizards. Specifically, while investment in dewlap size is sustained under low-resource condition relative to the high-resource treatment, investment in bite force is substantially lower. Disruption of the otherwise honest dewlap size to bite force relationship is therefore driven by costs associated with the expression of performance rather than the costs of signal production in A. carolinensis.     

Social influences on female choice in green anole lizards (Anolis carolinensis)

We conducted an experiment on female Anolis carolinensis lizards to investigate whether social factors influenced their selection of an end-chamber in a test arena. We tested (1) whether characteristics of males previously seen in the end-chambers would influence female choice and (2) whether the presence of other females simultaneously choosing would influence choice. In experiment one, females observed a large and a small male in the end-chambers prior to choosing. Females were tested individually and in pairs. When tested individually, females preferred the end-chamber previously inhabited by the larger male. When females were tested in pairs, however, in each case one female chose the large male's end-chamber and the other female failed to make a choice. In experiment two, we conducted the same paired-choice test, but prior to the test we evaluated the dominance relationships between the pair of females. In the majority of cases, the more dominant female was the one to enter the large male's end-chamber. Results indicate that females are influenced by the presence and characteristics of males, but that female competition also plays a role in choice.     

Relationships between hormones and aggressive behavior in green anole lizards: an analysis using structural equation modeling

We investigated the relationship between aggressive behavior and circulating androgens in the context of agonistic social interaction and examined the effect of this interaction on the androgen-aggression relationship in response to a subsequent social challenge in male Anolis carolinensis lizards. Individuals comprising an aggressive encounter group were exposed to an aggressive conspecific male for 10 min per day during a 5-day encounter period, while controls were exposed to a neutral stimulus for the same period. On the sixth day, their responses to an intruder test were observed. At intervals, individuals were sacrificed to monitor plasma androgen levels. Structural equation modeling (SEM) was used to test three a priori interaction models of the relationship between social stimulus, aggressive behavior, and androgen. Model 1 posits that exposure to a social stimulus influences androgen and aggressive behavior independently. In Model 2, a social stimulus triggers aggressive behavior, which in turn increases circulating levels of androgen. In Model 3, exposure to a social stimulus influences circulating androgen levels, which in turn triggers aggressive behavior. During the 5 days of the encounter period, circulating testosterone (T) levels of the aggressive encounter group followed the same pattern as their aggressive behavioral responses, while the control group did not show significant changes in their aggressive behavior or T level. Our SEM results supported Model 2. A means analysis showed that during the intruder test, animals with 5 days of aggressive encounters showed more aggressive responses than did control animals, while their circulating androgen levels did not differ. This further supports Model 2, suggesting that an animal's own aggressive behavior may trigger increases in levels of plasma androgen.     

Sexual dimorphism in neuromuscular junction size on a muscle used in courtship by green anole lizards.

The green anole lizard exhibits seasonal courtship behavior that is sexually dimorphic. This courtship consists of the extension of a bright red throat fan (dewlap) associated with head-bobbing display behavior. While males extend their dewlaps in aggressive encounters as well as in courtship, females use their considerably smaller dewlaps much less frequently and mainly in agonistic encounters. In parallel, a number of components of the neuromuscular system controlling dewlap extension are greater in males than in females during the breeding season, including dewlap motoneuron soma size and muscle fiber size and number. These features do not seem to change substantially in adulthood, despite a dramatic decline in dewlap use during the nonbreeding season. We explored the morphology of this neuromuscular system in more detail in the present experiment in males and females during both the breeding and nonbreeding seasons. Fiber and whole muscle length (approximately perpendicular to the fibers) were measured. Acetylcholinesterase histochemistry was used to visualize neuromuscular junctions (NMJs), and the surface area and density of NMJs were assessed for each animal. During the breeding season, NMJ size was larger in males than in females, but NMJ density along each fiber was equivalent between the sexes. In addition, whole muscle length and that of individual muscle fibers, was larger in males than in females. However, when corrected for body size, the sex difference in muscle fiber length disappeared. In the nonbreeding season, the sexual dimorphisms were maintained, suggesting that these features do not change substantially due to differences in circulating testosterone or a difference in use across seasons. Overall, these results are consistent with the idea that enhanced NMJ size is a relatively stable feature of the dewlap muscle in adulthood that either facilitates or is a consequence of using a larger muscle to extend a bigger dewlap in males compared to females.     

The effects of progesterone on sexual behavior in male green anole lizards (Anolis carolinensis).

It is well known that androgen-dependent sexual behaviors in male mammals and birds are inhibited by exogenous progesterone (P). However, recent research on male whiptail lizards (Cnemidophorus inornatus) indicates that P can stimulate sexual and copulatory behavior. We report here both antiandrogenic and synandrogenic actions of P on sexual behavior in males of another reptile, the green anole lizard (Anolis carolinensis). Earlier reports on birds and mammals are reviewed and discussed in relation to a possible physiological role of P in influencing sexual behavior in male vertebrates.     

The effects of progesterone on sexual behavior in male green anole lizards (Anolis carolinensis)

It is well known that androgen-dependent sexual behaviors in male mammals and birds are inhibited by exogenous progesterone (P). However, recent research on male whiptail lizards (Cnemidophorus inornatus) indicates that P can stimulate sexual and copulatory behavior. We report here both antiandrogenic and synandrogenic actions of P on sexual behavior in males of another reptile, the green anole lizard (Anolis carolinensis). Earlier reports on birds and mammals are reviewed and discussed in relation to a possible physiological role of P in influencing sexual behavior in male vertebrates.     

Seasonal plasticity in the copulatory neuromuscular system of green anole lizards: a role for testosterone in muscle but not motoneuron morphology

The copulatory system of green anoles is highly sexually dimorphic. Males possess bilateral copulatory organs called hemipenes, each independently controlled by two muscles: the transversus penis (TPN) and retractor penis magnus (RPM). The TPN everts the hemipene through the cloaca and the RPM retracts it. Adult females do not possess hemipenes or either of these two muscles. The spinal nucleus projecting to the TPN and RPM contains more and larger motoneurons in males than females. Because anoles breed seasonally, two experiments were designed to test whether adult copulatory morphology varies with environmental condition, and if so, whether the effect is mediated by testicular androgens. Three groups of adult males were used in each experiment: males from breeding environmental conditions with reproductive testes (BS); males in breeding conditions with regressed testes (BS-X); and males in nonbreeding conditions with regressed testes (NBS). Experiment 1 compared gonadally intact males and Experiment 2 compared castrated males treated with either testosterone (T) or an empty implant. In both experiments, copulatory and control motoneurons appeared smaller in NBS males, but T did not affect their size. In contrast, while hemipene and RPM muscle fiber size were not plastic across season in gonadally intact males, T in castrated males significantly increased both measures under BS and BS-X, but not NBS, conditions. These results demonstrate that neuron soma size might change on a general level and environmental cues can mediate T-induced changes in peripheral structures, suggesting that plasticity across copulatory system components is regulated by different mechanisms.     

Seasonal and sexual dimorphisms in the green anole forebrain

During the breeding season, male green anole lizards extend a throat fan (dewlap) in courtship. This behavior is facilitated by testosterone (T). Females extend a much smaller dewlap less often, even with the same dose of T. During the nonbreeding season when T is low, dewlap extension is reduced. To determine if parallels exist between structure and function, we investigated neuron soma size and density in the preoptic area (POA) and ventromedial nucleus of the amygdala (AMY), which are involved in the display behavior, in breeding and nonbreeding males and females. Cells from breeding animals were larger than cells from nonbreeding animals, but they were not sexually dimorphic. No significant effects existed in neuron density. This experiment indicates that portions of the anole forebrain important for sexual behavior are plastic and might be influenced by seasonal changes in steroid hormones. To investigate whether T can reverse the seasonal difference in soma size in both sexes, gonadectomized nonbreeding anoles were implanted with an empty or T propionate-filled capsule; animals were also tested for male-typical courtship behavior. Males and females treated with T had higher rates of dewlap extension, but across treatment groups these rates were greater in males. Neuron soma size in the POA and AMY was larger in males than females, but no effects of treatment were detected. Taken together, the results indicate that T can stimulate behavior in the nonbreeding season and suggest that a dissociation exists between the regulation of the courtship display and soma size of relevant brain regions.     

An incipient invasion of brown anole lizards (<Emphasis Type="Italic">Anolis sagrei</Emphasis>) into their own native range in the Cayman Islands: a case of cryptic back-introduction

Human-mediated dispersal has reshaped distribution patterns and biogeographic relationships for many taxa. Long-distance and over-water dispersal were historically rare events for most species, but now human activities can move organisms quickly over long distances to new places. A potential consequence of human-mediated dispersal is the eventual reintroduction of individuals from an invasive population back into their native range; a dimension of biological invasion termed “cryptic back-introduction.” We investigated whether this phenomenon was occurring in the Cayman Islands where brown anole lizards (Anolis sagrei) with red dewlaps (i.e., throat fans), either native to Little Cayman or invasive on Grand Cayman, have been found on Cayman Brac where the native A. sagrei have yellow dewlaps. Our analysis of microsatellite data shows strong population-genetic structure among the three Cayman Islands, but also evidence for non-equilibrium. We found some instances of intermediate multilocus genotypes (possibly 3–9% of individuals), particularly between Grand Cayman and Cayman Brac. Furthermore, analysis of dewlap reflectance data classified six males sampled on Cayman Brac as having red dewlaps similar to lizards from Grand Cayman and Little Cayman. Lastly, one individual from Cayman Brac had an intermediate microsatellite genotype, a red dewlap, and a mtDNA haplotype from Grand Cayman. This mismatch among genetic and phenotypic data strongly suggests that invasive A. sagrei from Grand Cayman are interbreeding with native A. sagrei on Cayman Brac. To our knowledge, this is the first evidence of cryptic back-introduction. Although we demonstrate this phenomenon is occurring in the Cayman Islands, assessing its frequency there and prevalence in other systems may prove difficult due to the need for genetic data in most instances. Cryptic back-introductions may eventually provide some insight into how lineages are changed by the invasion process and may be an underappreciated way in which invasive species impact native biodiversity.     

Caudal autotomy and regeneration in lizards

Caudal autotomy, or the voluntary self-amputation of the tail, is an anti-predation strategy in lizards that depends on a complex array of environmental, individual, and species-specific characteristics. These factors affect both when and how often caudal autotomy is employed, as well as its overall rate of success. The potential costs of autotomy must be weighed against the benefits of this strategy. Many species have evolved specialized behavioral and physiological adaptations to minimize or compensate for any negative consequences. One of the most important steps following a successful autotomous escape involves regeneration of the lost limb. In some species, regeneration occurs rapidly; such swift regeneration illustrates the importance of an intact, functional tail in everyday experience. In lizards and other vertebrates, regeneration is a highly ordered process utilizing initial developmental programs as well as regeneration-specific mechanisms to produce the correct types and pattern of cells required to sufficiently restore the structure and function of the sacrificed tail. In this review, we discuss the behavioral and physiological features of self-amputation, with particular reference to the costs and benefits of autotomy and the basic mechanisms of regeneration. In the process, we identify how these behaviors could be used to explore the neural regulation of complex behavioral responses within a functional context.