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1.
Anolis carolinensis has two aggressive displays involving movements of the hyoid apparatus: erection of the throat and extension of the dewlap. Erection of the throat is an enlargement of the gular region and dewlap extension consists of a vertical erection of the gular flap. Cinefluoroscopy and high speed cinematography show that the dewlap is extended in three phases: 1) protraction of the entire hyoid apparatus; 2) forward pivoting movement of the ceratobranchials II; and 3) retraction of the ceratobranchials II and the entire hyoid apparatus. The cartilaginous elements of the hyoid apparatus are variably mineralized. The entoglossal process and the hypohyals are the most calcified elements. The mineralized portion of the hyoid body, to which the other elements articulate, presents a complex pattern. The calcification of entoglossal process and the hypohyals stop just where they are fused with the hyoid body. The hyoid body presents four mineralized masses, two central corresponding to the base of the ceratobranchials II and two lateral being the head of the ossified ceratobranchials I. The lateral masses articulate on the central masses by a synovial joint. Morphologically, the ceratobranchials II form the hyoid body and become separated at the mid length of the synovial articulation of the ceratobranchials I and the hyoid body. The calcified matrix of the ceratobranchials II gradually changes from a large calcified mass (within the hyoid body) to a semicircle, opened ventrally, which permits their bending during dewlap extension. The highly mineralized posterior tip of the entoglossal process and the hyoid body serve as a pivot to pivoting forward movement of the ceratobranchials II producing at the change of the pattern of mineralization. Forward movement of the ceratobranchials II is produced by electrical stimulation of the M. branchio hyoideus. The opposition of the throat skin to the movement of the ceratobranchials II produces the bending of those longest elements. Electrical stimulation of the hyoid muscles confirms the key role of M. branchiohyoideus during dewlap extension. Simultaneous contractions of all the hyoid and extrinsic tongue (retractor and protractor) muscles with the M. branchiohyoideus during dewlap extension may be a possible motor pattern for dewlap extension in Anolis lizards.  相似文献   

2.
During the breeding season, male green anoles (Anolis carolinensis) court females by extending a red throat fan called a dewlap. Motoneurons controlling this sexually dimorphic behavior are located in two portions of the brain stem: (a) the vagal portion of nucleus ambiguus (AmbX), and (b) the region containing the glossopharyngeal portion of nucleus ambiguus and the ventral motor nucleus of the facial nerve (AmbIX/VIImv). These motoneurons project to the ceratohyoideus muscle via the ramus pharyngo-laryngeus IX+X. To investigate the effects of season on and androgen regulation of neural and peripheral structures controlling dewlap extension, two experiments were conducted: (a) During the breeding and nonbreeding seasons, motoneuron number, soma size, and nucleus size were investigated in intact males and females and in castrated males treated with a testosterone propionate (TP) or a blank Silastic capsule. (b) Cross-sectional area of the nerve and muscle fiber size, number, and density were investigated in the four treatment groups during the breeding season only. No significant differences were found in motoneuron number. In the breeding season, subtle male-biased sex differences existed in both AmbX and AmbIX/VIImv soma size. Nerve cross-sectional area and muscle fiber size and number were substantially larger in males than females. Muscle fiber density was higher in females. No consistent effects due to season or androgen treatment were detected, although characteristics of motoneurons were in some cases slightly larger in the nonbreeding season. These results suggest that, while parallels to behavior exist between the sexes, morphological changes in adulthood in the dewlap motoneurons and muscle do not normally regulate courtship behavior in the male green anole.  相似文献   

3.
During the breeding season, male green anoles (Anolis carolinensis) court females by extending a red throat fan called a dewlap. Motoneurons controlling this sexually dimorphic behavior are located in two portions of the brain stem: (a) the vagal portion of nucleus ambiguus (AmbX), and (b) the region containing the glossopharyngeal portion of nucleus ambiguus and the ventral motor nucleus of the facial nerve (AmbIX/VIImv). These motoneurons project to the ceratohyoideus muscle via the ramus pharyngo‐laryngeus IX+X. To investigate the effects of season on and androgen regulation of neural and peripheral structures controlling dewlap extension, two experiments were conducted: (a) During the breeding and nonbreeding seasons, motoneuron number, soma size, and nucleus size were investigated in intact males and females and in castrated males treated with a testosterone propionate (TP) or a blank Silastic capsule. (b) Cross‐sectional area of the nerve and muscle fiber size, number, and density were investigated in the four treatment groups during the breeding season only. No significant differences were found in motoneuron number. In the breeding season, subtle male‐biased sex differences existed in both AmbX and AmbIX/VIImv soma size. Nerve cross‐sectional area and muscle fiber size and number were substantially larger in males than females. Muscle fiber density was higher in females. No consistent effects due to season or androgen treatment were detected, although characteristics of motoneurons were in some cases slightly larger in the nonbreeding season. These results suggest that, while parallels to behavior exist between the sexes, morphological changes in adulthood in the dewlap motoneurons and muscle do not normally regulate courtship behavior in the male green anole. © 1999 John Wiley & Sons, Inc. J Neurobiol 40: 202–213, 1999  相似文献   

4.
Data from a diversity of sources are consistent with the hypothesis that the Grand Cayman anole, Anolis conspersus, is descended directly from Anolis grahami of Jamaica. Although the two species have remained morphologically similar, coloration in A. conspersus has changed considerably from that of its ancestor. The most dramatic difference is seen in dewlap colour, where A. conspersus has evolved a blue and highly UV-reflective dewlap from the ancestral orange-and-yellow colour state. In addition, variation in normal (non-metachrosis) dorsum coloration in A. grahami populations is limited to shades of green (olive, emerald, teal), whereas in A. conspersus dorsum coloration varies from green to blue and to brown. This increased colour variation occurs despite Grand Cayman being a small, relatively featureless island only 35 km in length. Results of this study suggest that ambient light differences associated with precipitation-related vegetation structure may have played an important role in the evolution of A. conspersus body colour variation. Evidence is presented to show how geological, ecological, and physiological factors could have interacted to select for a short wavelength-reflective dewlap from a long wavelength-reflective precursor following the colonization of Grand Cayman from Jamaica by A. grahami between 2 and 3 Mya.  相似文献   

5.
The dewlap of male anoline lizards is an important component of the mate recognition system. Much attention has focussed on the role dewlap colour plays in the interactions between species of Anolis but little has been paid to the effects of intraspecific variability in this feature. Patterns of intrasubspecific variability in this feature are described for several subspecies of Anolis distichus and A. brevirostris . In most cases variability in dewlap colour appears unrelated to the amount of electrophoretic variability and the level of population differentiation, although there is some evidence for large scale geographic subdivision within three of the taxa examined. Female choice and the implications for various species concepts are discussed.  相似文献   

6.
Caribbean Anolis lizards are often cited as a textbook example of adaptive radiation. Similar morphologies (ecomorphs) have originated in similar ecological settings on different large islands in the West Indies. However, relatively little is known about one of the morphologically most specialized and divergent ecomorphs: the twig anoles. Here, we investigate aspects of morphology, dewlap size, locomotor and bite performance, structural habitat and diet of the poorly known twig anole, Anolis sheplani from Hispaniola. Few observations have previously been made of this species in its natural habitat, and few quantitative data on its natural history are available. A. sheplani is an extreme twig anole with respect to its morphology, performance capacities, and ecological niche. Males and females of this species do not differ from each other in body dimensions, performance or habitat use, but males do have a bigger dewlap than females. We present data for 25 individuals and compare them with data for other Greater Antillean anoles. It becomes apparent that twig anoles constitute a large component of the morphological, functional, and ecological diversity of Anolis lizards. Small twig anoles such as A. sheplani appear to be pushing the boundaries of morphospace and are thus crucial in our understanding of the evolution of phenotypic diversity.  相似文献   

7.
Lizards of the genus Anolis extend and retract a large and often brightly colored throat fan called a dewlap. The dewlap in most anoles is a sexually dimorphic structure. It is larger in males than females and males use the dewlap more frequently and in more contexts than do females. In the present study we investigated whether plasma testosterone (T) levels and season affect the frequency of dewlap use in male--male interactions in the brown anole, Anolis sagrei. We manipulated plasma T levels by implanting adult, castrated males with pellets that delivered no T, a moderate dose of T, or a high dose of T. In tests with stimulus males, castrated males that were treated with a moderate or with a high dose of T had a significantly higher frequency of dewlap extensions than did castrated males that received no T. However, the frequency of dewlap extensions in castrated males that received the high dose of T did not differ significantly from castrated males that received the moderate dose of T or from non-castrated control males. Males captured during the breeding season and tested in the laboratory had a significantly higher frequency of dewlap extensions than did males captured in the postbreeding season and tested in the same manner. These results suggest that plasma T levels affect how frequently males of A. sagrei extend their dewlaps in male--male interactions and that seasonal changes in male dewlap use may be due to seasonal differences in plasma T levels.  相似文献   

8.
We describe a new species of Anolis from the southeastern slopes of the Andes of Ecuador, province of Zamora-Chinchipe, Parque Nacional Podocarpus. It belongs to (1) the aequatorialis species-group by being of moderate to large size with narrow toe lamellae, and (2) the eulaemus sub-group by having a typical Anolis digit, in which the distal lamellae of phalanx II distinctly overlap the proximal scales of phalanx I. The new species is most similar morphologically to Anolis fitchi but differs from it mainly by having a dewlap with longitudinal rows of 2-5 granular, minute scales separated by naked skin (longitudinal rows of one or two keeled, large scales separated by naked skin in Anolis fitchi) and a vertically shorter dewlap (longer dewlap in Anolis fitchi).  相似文献   

9.
The highly specialized feeding apparatus of modern birds is characterized in part by paraglossalia, triangular bones or cartilages in the tongue that constitute part of the rarely fossilized hyobranchial apparatus. Here, we report on a new, juvenile specimen of the ankylosaurid dinosaur Pinacosaurus grangeri Gilmore, 1933 that provides the first evidence of paraglossalia outside of crown group Aves. The specimen is remarkable in preserving a well‐ossified hyobranchial apparatus, including paired paraglossalia, first and second ceratobranchials, epibranchials, and evidence of a median cartilaginous basihyal. Reassessment of Edmontonia, another ankylosaur, also reveals evidence of bony paraglossalia. Ankylosaur paraglossalia closely resemble those of birds, but are relatively larger and bear prominent muscle scars, supporting the hypothesis that ankylosaurs had fleshy, muscular tongues. The other hyobranchial elements, surprisingly, resemble those of terrestrially feeding salamanders. Ankylosaurs had reduced, slowly replacing teeth, as evidenced from dental histology, suggesting that they relied greatly on their tongues and hyobranchia for feeding. Some curved, rod‐like elements of other dinosaur hyobranchia are reinterpreted as second ceratobranchials, rather than first ceratobranchials as commonly construed. Ankylosaurs provide rare fossil evidence of deep homology in vertebrate branchial arches and expose severe biases against the preservation and collection of the hyobranchial apparatus. In light of these biases, we hypothesize that paraglossalia were present in the common ancestor of Dinosauria, indicating that some structures of the highly derived avian feeding apparatus were in place by the Triassic Period. © 2015 The Linnean Society of London  相似文献   

10.
The dewlap in the lizard Anolis carolinensis is made of scales separated by large interscale regions capable of broad stretching during fan extension. This indicates that the skin contains proteins that allow extension of interscale regions. The immunocytochemical analysis of the epidermis indicates that HgG5, a glycine‐rich hydrophobic beta‐protein poor in cysteine is localized only in the stiff beta‐layer of the outer scale surface, but is completely absent in mesos and alpha‐layers and in hinge regions. HgGC10, a cysteine‐medium‐rich beta‐protein is present in beta‐layers but especially in alpha‐layers of interscale epidermis that presents folds and lacks a beta‐layer. HgGC3 is weakly localized in the alpha‐layer, but is mainly found in hinge regions. HgGC8 and HgG13 are low to absent in the alpha‐ and beta‐layer. The immunolocalization of cysteine‐rich beta‐proteins such as HgGC10/3 in alpha‐layers and interscale epidermis suggests that these small proteins are involved in the formation of a corneous material compatible with dewlap extension. The basement membrane underneath scales is joined to bundles of collagen fibrils in the dermis through anchoring fibrils that likely determine flattening of the epidermis during the extension of the throat fan.  相似文献   

11.
The use of conspicuous communication signals often increases a signaler's risk of predation. Many species communicate with a repertoire of signals that may differ in their conspicuousness to predators. Few studies have examined the ability of prey to selectively decrease the use of individual signals in their displays under heightened predation risk. Here, I examined the behavior of male brown anole lizards (Anolis sagrei) in response to a simulated predatory attack from a model kestrel. This species communicates with three major visual signal types, the head‐bob, pushup, and dewlap extension, which vary in their motion and spectral characteristics. I predicted that lizards would decrease frequencies of the dewlap extension and pushup following the attack, but not the head‐bob. Males modulated their use of individual signals by decreasing pushup rates, but not head‐bob rates. Decreases in dewlap frequency were marginally significant. One explanation for these results is that lizards decrease frequencies of signal types based partly on their conspicuousness. The energetic cost of each signal type may be an equally important factor that determines the signaler's response to predators, particularly if a predatory attack is perceived as imminent.  相似文献   

12.
Animal signalling systems are extremely diverse as they are under different, often conflicting, selective pressures. A classic textbook example of a diverse signal is the anoline dewlap. Both at the inter‐ and intraspecific levels, dewlap size, colour, shape and pattern vary extensively. Here, we attempt to elucidate the various factors explaining the diversity in dewlap size and pattern among seven Anolis sagrei populations from different islands in the Bahamas. The seven islands differ in the surface area, number and kind of predators, sexual size dimorphism and Anolis species composition. In addition, we investigate whether selective pressures acting on dewlap design differ between males and females. Whereas dewlap pattern appears to serve a role in species recognition in both sexes, our data suggest that relative dewlap size is under natural and/or sexual selection. We find evidence for the role of the dewlap as a pursuit‐deterrence signal in both males and females as relative dewlap size is larger on islands where A. sagrei occurs sympatrically with predatory Leiocephalus lizards. Additionally, in males relatively large dewlaps seem to be selected for in a sexual context, whereas in females natural selection, for instance by other predators than Leiocephalus lizards, appears to constrain relative dewlap size.  相似文献   

13.
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.  相似文献   

14.
The expression of male secondary sexual traits can be dynamic, changing size, shape, color, or structure over the course of different seasons. However, the factors underlying such changes are poorly understood. In male Anolis carolinensis lizards, a morphological secondary sexual signal called the dewlap changes size seasonally within individuals. Here, we test the hypothesis that seasonal changes in male dewlap size are driven by increased use and extension of the dewlap in spring and summer, when males are breeding, relative to the winter and fall. We captured male green anole lizards prior to the onset of breeding and constrained the dewlap in half of them such that it could not be extended. We then measured dewlap area in the spring, summer, and winter, and dewlap skin and belly skin elasticity in summer and winter. Dewlaps in unconstrained males increase in area from spring to summer and then shrink in the winter, whereas the dewlaps of constrained males consistently shrink from spring to winter. Dewlap skin is significantly more elastic than belly skin, and skin overall is more elastic in the summer relative to winter. These results show that seasonal changes in dewlap size are a function of skin elasticity and display frequency, and suggest that the mechanical properties of signaling structures can have important implications for signal evolution and design.  相似文献   

15.
A key hypothesis explaining the existence of dawn and dusk choruses in acoustically communicating animals centers on the need to advertise continued territorial occupancy after and before a period of nocturnal inactivity. If this hypothesis is correct, it follows that similar dawn and dusk choruses should occur in territorial animals exploiting other signal modalities. Adult male Anolis lizards defend territories by using elaborate head-bobbing displays and extensions of a throat fan or dewlap. Males are inactive at night and return to their territories at dawn, remaining there until dusk. I quantified the production of visual displays as a function of time of day for four species on the island of Jamaica: Anolis lineatopus, Anolis sagrei, Anolis grahami, and Anolis opalinus. All exhibited dawn and/or dusk peaks in display behavior. These patterns have remarkable parallels with the dawn and dusk choruses reported for many acoustically communicating animals.  相似文献   

16.
We describe morphological variation (scalation and coloration) observed among eight individuals of the Panamanian lizard species Anolis casildae. This variation was not observed in the holotype and aids in identification of this recently described species (originally described on the basis of a single, male specimen). This species occurs only in the Reserva Forestal Fortuna (Chiriquí Province) and the adjacent Bosque Protector Palo Seco (Bocas del Toro Province) in western Panama. Anolis casildae can be distinguished from all other Panamanian anole species via six features: (1) two enlarged superciliary scales (the first larger than the second); (2) an anterior nasal scale in contact with the rostral scale or separated from the rostral by one scale; (3) 6-8 sublabial scales to the center of the eye; (4) 3-4 scales between the supraobital semicircles; (5) unique coloration (4-6 oblique brown bands interspersed by blue-outlined yellow patches; dewlap is a dirty cream color with broad yellow scale rows irregularly interspersed with smaller emerald green scales) and (6) A. casildae occurs from 1,050 to 1,400 m in the Cordillera Central. We also compare our natural history observations of A. casildae to a similar large anole, A. frenatus, a species which we believe A. casildae to be closely related.  相似文献   

17.
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. © 2002 Wiley Periodicals, Inc. J Neurobiol 50: 24–30, 2002  相似文献   

18.
The pattern of reproductive character displacement (RCD)—in which traits associated with reproductive isolation are more different where two species occur together than where they occur in isolation—is frequently attributed to reinforcement, a process during which natural selection acting against maladaptive mating events leads to enhanced prezygotic isolation between species or incipient species. One of the first studies of RCD to include molecular genetic data was described 40 years ago in a complex of Haitian trunk anole lizards using a small number of allozyme loci. In this example, Anolis caudalis appears to experience divergence in the color and pattern of an extensible throat fan, or dewlap, in areas of contact with closely related species at the northern and southern limits of its range. However, this case study has been largely overlooked for decades; meanwhile, explanations for geographic variation in dewlap color and pattern have focused primarily on adaptation to local signalling environments. We reinvestigate this example using amplified fragment length polymorphism (AFLP) genome scans, mtDNA sequence data, information on dewlap phenotypes and GIS data on environmental variation to test the hypothesis of RCD generated by reinforcement in Haitian trunk anoles. Together, our phenotypic and genetic results are consistent with RCD at the southern and northern limits of the range of A. caudalis. We evaluate the evidence for reinforcement as the explanation for RCD in Haitian trunk anoles, consider alternative explanations and provide suggestions for future work on the relationship between dewlap variation and speciation in Haitian trunk anoles.  相似文献   

19.
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.  相似文献   

20.
Subspecies of Anolis lizards are often defined on the basis of geographic variation in the color and pattern of the dewlap, an extensible throat fan considered central to species recognition and sexual selection. Among the most impressive examples of this phenomenon are two species of trunk anoles found across Hispaniola and the Bahamas: Anolis distichus is divided into 16 subspecies with dewlap colors ranging from deep wine red to pale yellow while Anolis brevirostris is divided into three subspecies with dewlaps ranging from pale yellow to orange. Limited sampling of allozyme data indicates some genetic divergence among subspecies and suggests that they may deserve recognition at the species-level. Our goal here is to use more comprehensive geographic sampling of mtDNA haplotypes to test whether the five subspecies of A. distichus and three subspecies of A. brevirostris that occur in the Dominican Republic correspond with genetically distinct populations that may warrant recognition under the general lineage concept. We obtain an aligned dataset of 1462bp comprised of the genes encoding ND2 and adjacent tRNAs from 76 individuals of A. distichus from 28 localities and 12 individuals of A. brevirostris from five localities. We find that haplotypes sampled from each Dominican subspecies of A. distichus form well-supported and deeply divergent clades (>10% uncorrected sequence divergence). Strong concordance between mtDNA haplotype structure and previously diagnosed phenotypic variation in traits central to interspecific communication (i.e., the dewlap) leads us to hypothesize that each of the presently recognized Dominican subspecies of A. distichus and A. brevirostris deserves elevation to full species status under the general lineage concept.  相似文献   

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