Ethotypic Variation of Prey Recognition in Juvenile Anolis lineatopus (Reptilia: Iguanidae)

Among prey-naive Anolis lineatopus from Jamaica, individuals reject ants most commonly, crickets less commonly, and waxmoth larvae almost never. This study investigates the individual differences in the mechanism underlying prey perception in two of many ethotypes found, i.e., cricket-rejectors and cricket-acceptors. In one experiment artificially coloured live waxmoth larvae, crickets and ants were presented, in another experiment crickets and waxmoth larvae were moved passively with different speeds. Ethotypes differ in their evaluation of the interaction of prey key stimuli. Cricket-rejectors and cricket-acceptors differ in their responses to both crickets and waxmoth larvae that were either unicoloured yellow or black, but not in their responses to two types of ants of either colouration. Both ethotypes also differ in their preferences for artificially moved crickets and waxmoth larvae as a function of the interaction of prey type and velocity. In all but two prey items there is interaction in the perceptual pathways concerned. The first presentation in ontogeny of an ant inhibits attack on a cricket presented the next day. Some results suggest that ethotypes differ but gradually in their evaluation of stimuli presented, with cricket-rejectors having a higher threshold for attack. However, other results suggest that cricket-rejectors evaluate certain prey in a way qualitatively different from that of acceptors. At present there is little evidence for an understanding of the ethotypic differences of prey selection at a neural level, probably because neuroethologists have failed to look at individual differences of prey recognition; this failure is discussed in some detail.     

Epiermal morphology and sloughing frequency in normal and prolactin treated Anolis carolinensis (Iguanidae, Lacertilia)

The morphology of the epidermal generations throughout the sloughing cycle of Anolis carolinensis is essentially the same as that described for other squamates, whether studied in control or hormone-injected animals. Readily identifiable granules in the clear layer of this and other lizard species emphasize the eventual hardening of this layer prior to its separation from the subjacent Oberhautchen, which brings about sloughing. Eosinophils are described for the first time in the lacunar tissue of lizard, but their functional role remains problematical. Daily 25 μg injections of prolactin (alone or in combination with gonadotropins) raise the frequency of sloughing by shortening the resting phase as compared with controls and animals receiving 1.0 I.U./day injections of gonadotropins. The duration of the proliferation-renewal phase and the pattern of histological changes occurring therein is unchanged by experimental treatments. The results show that neither endogenous nor exogenous gonadotropins affect the sloughing frequency, but exogenous prolactin produces a similar effect to that described for thyroxine. Although the mechanisms of action are unknown, the results provide another example of the cutaneous action of prolactin in vertebrates.     

Differential Response to Colors by the Lizard Uta palmeri (Reptilia: Iguanidae)

Many lizards have brightly colored body parts thought to be signals, but color perception in lizards has not been well studied. We present data documenting differential response to colors in an iguanid lizard. Free-living Uta palmeri differentially approached and bit at a red-colored object when simultaneously presented objects of four different colors (red, blue, yellow, black). This response to red was statistically significant for all three age-sex classes of lizards tested (adult males, adult females, juveniles). In addition, juveniles and adult females exhibited intermediate levels of responses (approaching and sniffing, and licking) to the yellow object but adult males did not. Seasonally important food items of this species are red (fish regurgitated by nesting seabirds) and yellow (yolk from broken seabird eggs), and response behavior (biting versus licking) was similar to that exhibited during feeding. Thus, as presented in our experimental paradigm, red and yellow elicited feeding responses. The spectral reflectance curves of the red and yellow stimuli were found to exhibit peaks at wavelengths of 630 nm (range 570–670), and at 600 nm (range 550–700), respectively. These values overlap with the spectral range of the gravid coloration of female U. palmeri (peak 615 nm, range 550–690) that is thought to be a signal to conspecifics. Furthermore, the reflectance curves are similar in shape and height (intensity). Thus, the similarities of the reflectance curves of the red paint and of the natural coloring suggest that palmeri can detect the female gravid coloration. Whether the response to red objects is due to discriminating differences in hue or in relative brightness remains to be examined.     

Ethotypic Variation of Prey Recognition in Juvenile Anolis lineatopus (Reptilia: Iguanidae) Revisited

A previous study (von Brockhusen -Holzer & Curio 1990) had led to the major conclusion that prey-naive Anolis lineatopus hatchlings differ in terms of a “hard-wired” perceptual mechanism underlying the recognition of cricket prey. We reconsider here one result leading to that conclusion since it rested on negative evidence in one treatment group as compared to another. A newly applied discriminant analysis utilizing the complete prey rejection/acceptance data of individuals in the two treatment groups, i.e. cricket-rejectors and prefed cricket-acceptors, again permits the repudiation of a motivation-based alternative hypothesis explaining the rejector-acceptor dichotomy. Thereby we reconfirm the original hypothesis of a “hard-wired”, preprogrammed polyethism underlying prey recognition.     

Dewlap and other variation in the lizards Anolis distichus and A. brevirostris (Reptilia: Iguanidae)

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.     

An Analysis of Inter-Population Divergence in Visual Display Behavior of the Green Anole Lizard (Anolis carolinensis)

Variation in habitat structure can profoundly affect the evolution of visual displays in animals. We compared the field display behavior of two populations of the green anole lizard (Anolis carolinensis) in Southern Louisiana. These two populations occupy habitats varying markedly in structure (clumped vs. continuous), ecological setting (urban vs. natural) and the degree of male competition and predation. We filmed 56 large adult male lizards during the active spring reproductive period and detected substantial divergence in display behavior between populations. We found that (1) relative dewlap areas did not differ significantly between populations, (2) Tulane anoles spent about twice as much time displaying compared with Good Hope field anoles, although the average display duration did not differ significantly between populations, and (3) the two populations differed significantly in the relative frequency of display types: lizards from the Tulane population used a higher proportion of A and B display types, whereas Good Hope Field anoles used C displays more often. Finally, we detected differences in other aspects of display behavior (use of display modifiers, volley length distributions). We suggest that these display differences may result from differences in green anole male density (three times higher at Tulane), combined with habitat differences (clumped vs. continuous).     

A functional interpretation of the varanid dentition (Reptilia, Lacertilia, Varanidae)

A cinéradiographic analysis of the feeding movements in Varanus bengalensis produced the following results. The mouth is opened by raising the head (upper jaw) rather than by lowering the lower jaw. Starting from the resting position, the muzzle unit is elevated around 9 degrees relative to the rest of the skull during jaw opening; the quadrate swings anteriorly around 21 degrees. During jaw closure, the snout is depressed around 15 degrees relative to the rest of the skull, hence 6 degrees beyond the resting position. The quadrate swings backwards around 27 degrees. Amphikinesis is interpreted as allowing a stronger posterior recurvature of the maxillary teeth in Varanus. This increases the holding effect of the teeth without increasing their length, an adaptation of Varanus to capture relatively large prey. The formation of plicidentine (dentine infolding) in the teeth of Varanus increases the surface of attachment of the teeth on the supporting bone. Moreover, the dentine lamellae take up tensile and compressive stresses along their long axes upon axial or vertical loading of the teeth. The slope of pleurodonty is modelled so as to minimalize shear stress on the surface of ankylosis upon axial or vertical loading of the teeth.     

The mechanism of dewlap extension in Anolis carolinensis (Reptilia: Iguanidae) with histological analysis of the hyoid apparatus

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.     

Global diversity of lizards in freshwater (Reptilia: Lacertilia)

No lizards are strictly aquatic, but at least 73 species in 11 different families can be considered to regularly utilize freshwater habitats. There are no aquatic lizards in the Nearctic or Palearctic regions, whereas the Neotropics, Southeast Asia, and the Indo-Australian Archipelago support the greatest diversity of freshwater forms, particularly in the families Gymnophthalmidae, Scincidae and Varanidae. A number of larger aquatic lizards are harvested for food and for the reptile skin trade and several are CITES listed. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Functional morphology of dewlap extension in the lizard Anolis equestris (Iguanidae)

The dewlap is an extendible flap of skin ordinarily folded under the throat. Lizards, particularly those in the genus Anolis, extend their dewlaps during interactions with conspecifics, other lizards, and potential predators. Dewlap extension is effected by movements of elements of the hyoid apparatus. This paper describes the anatomy of the hyoid and associated musculature in Anolis equestris, a large arboreal lizard with a prominent dewlap. A mechanism for dewlap extension is proposed based on results of morphological and experimental techniques. Specializations of the hyoid skeleton for dewlap extension include elongated second ceratobranchials and highly movable joints between the ceratohyals and the hypohyals and between the first ceratobranchials and the body of the hyoid. A well developed M. ceratohyoideus extends between the ceratohyals and the first ceratobranchials of the hyoid apparatus. During dewlap extension, the hyoid apparatus acts as a first order lever. Contraction of M. ceratohyoideus pulls the ceratohyals posteriorly causing the hypohyals and the body of the hyoid to rotate dorsally around the first ceratobranchial/body joints. This movement results in the second ceratobranchials swinging forward and down, unfolding the dewlap. The relative immobility of the first ceratobranchials provides stability to the hyoid apparatus during dewlap extension. A comparison is made of dewlap extension and other hyoid displays.     

Evolution of Anoline Lizard Display Behavior

Based on my conceptual framework of anoline display behavior,I am suggesting the following evolutionary trends. Lateral presentationduring display was probably promoted by monocular vision. Alongwith lateral presentation, postures evolved to increase lateraloutline. These postures which magnified body size were probablyof selective advantage within aggressive social contexts sincelarger animals tend to dominate smaller ones through bluff.Body movement evolved along with lateral orientation and size-enhancingpostures. These movements would be most effective if they complementedlateral orientation. Effectors available for such movementswere primarily pre-adapted for vertical motion. The patternsof movement generated were probably simple oscillatory bobbingmovements by the head which were weakly stereotyped, interspecificallysimilar, appearing in many contexts, and having a weakly definedinformation content. Events having selective advantage for speciesrecognition promoted stereotypy of bobbing behavior into species-uniquedisplays; each species had its unique signature display whichserved in a manifold communicatory capacity. The signature displayappeared in assertion, courtship, and challenge contexts. Itsinformation content varied depending upon context and recipientof the display (e.g., male or female). Besides the stereotypedaspects of the display, certain features remained variable withpotential information significance. Core variability (see text)promotes individual recognition and may be the origin of newunique display patterns as sibling species emerge. Display modifiers(see text) are variable display features shared by members ofa population (many being shared interspecifically) that providea graded appearance to display performance; modifiers can indicatelevel of arousal and facilitate interspecific communication.For some species display repertoire size seems to have evolvedfrom a single display (signature display) to repertoires ofmultiple displays; these subsequent displays are generally restrictedto aggressive interactions.     

Foraging ecology in the lizard Anolis oculatus (Iguanidae) from Dominica, West Indies

In general, populations of Anolis lizards on West Indian islands face few predators, are at high density and are thought to be limited by food. This paper describes how the foraging ecology of Anolis oculatus , a solitary species confined to the island of Dominica, Lesser Antilles, varies with habitat and season in relation to the quantity and quality of available food.
Availability of invertebrate food (determined using pitfall traps and sticky traps) was greater in a dry scrub woodland site, Cabrits National Park (CNP), compared with a montane rainforest site, Palmiste Ridge. In the former, there were general increases in abundance, volumes of softbodied prey and sizes of invertebrates from dry season to wet season. Concomitant dietary changes, as determined principally by stomach flushing, included an increase (by volume) in the proportion of soft-bodied prey. Dietary analyses confirmed the importance of ants (Formicidae) in the diet of A. oculatus , although for large individuals (mainly adult males) at the rainforest site, soft-bodied prey such as Oligochaeta and Orthoptera were, in volumetric terms, more important. Prey capture observations showed that large A. oculatus fed mainly above ground. Anolis oculatus at the montane rainforest site used higher perch heights than those in dry scrub woodland, although in both habitats, small individuals (mainly juveniles) fed mainly at ground level on ants. In the dry season in CNP, the diet (in volumetric terms) of smaller Anolis was dominated by hardbodied prey such as ants, springtails (Collembola), barklice (Psocoptera) and beetles (Coleoptera). large Anolis used springtails and barklice to a lesser extent, resulting in relatively low food niche overlap values between size classes and a reduced potential for intraspecific competition compared with the wet season.     

Coordinated Ontogeny of Food Preference and Responses to Chemical Food Stimuli by a Lizard Ctenosaura pectinata (Reptilia: Iguanidae)

When important ecological factors change predictably during the life of an organism, the ontogeny of related behaviors must be timed to maintain appropriate behavioral responsiveness to current ecological conditions. In the brown iguana, Ctenosaura pectinata , hatchlings in natural populations eat primarily insects, consuming little plant matter, whereas adults eat primarily plants, consuming some insects as well. We conducted laboratory experiments on diet preferences and responses to chemical cues that the lizards sampled by tongue-flicking and used to identify food. All hatchlings ate crickets, but only one of six ate romaine lettuce. They responded strongly to chemical cues from prey, as indicated by elevated tongue-flick rates, but not from romaine lettuce. All older individuals ate both crickets and romaine lettuce. They responded much more strongly to chemical cues from both crickets and romaine lettuce than to control chemicals, as indicated by higher proportions of individuals that bit and higher tongue-flick attack scores.
Thus, an ontogenetic change to increased responsiveness to plant chemical stimuli was coordinated with an ontogenetic change to an herbivorous diet. The mechanisms underlying these ontogenetic changes are unknown, but folivory may be unprofitable before juveniles acquire intestinal flora that degrade cellulose by ingestion of feces of adult conspecifics. Possible mechanisms are discussed, including the detection of chemical cues from appropriate food plants during consumption of feces from older individuals. Studies of other squamate reptiles suggest that exposure to these chemicals might affect both future responsiveness to the chemical cues and a tendency to eat the corresponding plants.     

The Effects of Social Experience on Aggressive Behavior in the Green Anole Lizard (Anolis carolinensis)

To understand how context-specific aggression emerges from past experience, we examined how consecutive aggressive encounters influence aggressive behavior and stress responses of male green anole lizards ( Anolis carolinensis ). Animals were shown a video clip featuring an aggressively displaying conspecific male, which provoked aggressive responding, while control animals viewed a neutral video. After 5 d of interaction with the videos, both the subject and control groups were presented with a live conspecific. As a non-invasive assay of stress responses, we measured changes in body color and eyespot darkness, two features known to be strongly correlated with titers of stress hormones. Our results demonstrate that experience increased aggression in male anoles, but that increases in aggression to a repeated stimulus were transient. Tests with a novel conspecific indicate that the experienced animals remained aggressive when presented with novel stimuli. Although there were differences in the morphological indicators of the stress response between experimental and control groups during video presentations, there were no differences when presented with novel conspecifics. These data indicate that experience-dependent differences were not mediated by differences in the 'stressfulness' of aggressive interaction, as thought to be the case for animals in chronic subordinate/dominant dyads. We suggest that habituation and reinforcement interact to promote aggressive responding and to restrict it to novel individuals. Such context specificity is a hallmark of natural patterns of aggression in territorial species.     

Reproductive Biology of the Rock Lizard (Darevskia saxicola,Reptilia, Lacertidae)

Biology Bulletin - Data on the reproduction of the rock lizard (Darevskia saxicola (Eversmann 1834)) in the gorges of Bol’shaya Laba (Karachay-Cherkessia) and Malaya Laba (Krasnodar...     

Structure of shells of eggs of Callisaurus draconoides (Reptilia, Squamata, Iguanidae)

Female zebra-tailed lizards (Iguanidae: Callisaurus draconoides ) lay roughly ovoid eggs with thin, highly extensible shells. The outer surface of the eggshell is a thin, calcareous crust of calcium carbonate in the calcite morph. Immediately beneath the crystalline matrix is a shell membrane composed of multiple layers of fibres organized into an undulating series of troughs and crests, apparent in both cross-section and surface view. The outer surface of the shell membrane is differentiated into a tightly woven fibrous mat that may serve to anchor the calcareous layer to the membrane. Organization of the eggshell into a series of troughs and crests serves to increase the surface area available for contact with the substrate and, presumably, to increase the capacity of the eggshell to stretch as the egg absorbs water.     

A New Species of Japalura(Agamidae: Lacertilia: Reptilia) from Central Highland,Vietnam

A new species of the agamid genus Japalura is described based on three specimens from southern part of Central Vietnam.It is distinguished from remaining congeners by the following combination of characters: adult size(SVL females 68–69 mm),tail length/SVL ratio 226%–239%,HW/SVL ratio 17%–18%; FLL/SVL ratio 41%–43%; HLL/SVL ratio 72%–73%; 7–9 supralabials,7–9 infralabials,54–56 middorsal scales,20–22 lamellae under finger IV,24–26 lamellae under toe IV,1 scale between nasal and supralabials; tympanum concealed; absence of transverse gular fold.The geographical distribution of Japalura genus in general and of a new species in particular is discussed.