They change color depending on their mood. They possess uniquely adapted hands and feet distinct from other tetrapods. They feature independently movable eyes. This comprehensive volume delves into these fascinating details and thorough research about one of the most charismatic families of reptilesChameleonidae.
Written for professional herpetologists, scholars, researchers, and students, this book takes readers on a voyage across time to discover everything that is known about chameleon biology: anatomy, physiology, adaptations, ecology, behavior, biogeography, phylogeny, classification, and conservation.
A description of the natural history of chameleons is given, along with the fossil record and typical characteristics of each genus. The state of chameleons in the modern world is also depicted, complete with new information on the most serious threats to these remarkable reptiles.
|Publisher:||University of California Press|
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About the Author
Krystal A. Tolley is Principal Scientist, Molecular Ecology Program in Applied Biodiversity Research of the South African National Biodiversity
Institute. Her research focuses on understanding the historical processes that generate patterns of diversity and lead to species radiations of reptiles and amphibians in southern Africa. Anthony Herrel is a permanent researcher at the Centre Nationale de la Recherche Scientifique (CNRS) and the head of the Function and Evolution team of the Unite Mixed de Recherche (UMR) 7179, based at the Museum national d’Histoire naturelle in Paris.
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The Biology of Chameleons
By Krystal A. Tolley, Anthony Herrel
UNIVERSITY OF CALIFORNIA PRESSCopyright © 2014 The Regents of the University of California
All rights reserved.
Biology of the Chameleons
KRYSTAL A. TOLLEY and ANTHONY HERREL
Chameleons so easily capture the imagination, and have done so for centuries. In scientific writings, they first appear with Aristotle (350 BC), and although they were recognized as being similar to lizards in some ways, he also likened them to fish, baboons, and crocodiles. Since then, they have since been the subject of strange myths, amusing tales, and nature documentaries and have even taken the form of popular cartoon characters. But what makes them so fascinating that they have infiltrated the common psyche more than other reptiles? Indeed, they are set apart to such a degree that many people are unaware they are even lizards at all. They are simply, "chameleons." In this book, we draw together and review the body of literature that covers chameleons over several centuries by exploring and summarizing our knowledge on this intriguing group of lizards.
Chameleons are highly specialized animals characterized by a suite of morphological, physiological and functional adaptations. Much of what makes chameleons immediately identifiable as such are these morphological specializations. Among these, the laterally compressed body, the prehensile feet and tail, the independently moveable eyes (Figs. 1.1 and 1.2 in color insert), and a long tongue capable of being projected from the mouth are the most striking. As detailed in Chapters 2 (Chameleon Anatomy) and 3 (Chameleon Physiology), many of these features are intimately related to the unique behavior and lifestyle of chameleons as highly specialized arboreal predators. The sensory system of chameleons, for example, is highly tuned toward visual stimuli, as chameleons rely on visual signals in both a social and a feeding-related context. As part of the specialization of the visual system, chameleons have a negatively powered lens and use accommodation cues to judge distance, features unique among lizards (Fig. 1.3 in color insert). Because of their visual specialization, the auditory system is less developed and shows lower sensitivity as compared with that of other lizards. Little is known about the olfactory (smell), vomerolfactory, and gustatory (taste) systems in chameleons, but all appear reduced as compared with other lizards.
Associated with the predominantly arboreal habitat of chameleons comes a number of functional specializations such as their prehensile feet and tails. Chapter 4, Function and Adaptation of Chameleons, examines how these specializations allow chameleons to effectively explore the three-dimensional (mostly) arboreal habitat. However, as a consequence of their specializations, chameleons are the slowest of all lizards, with sprint speeds about 10 times slower than those of other lizards. The slow locomotion of chameleons is a result of the contractile capacities of the locomotor muscles, changes in limb posture that allows them to move effectively on narrow substrates, and a lower overall muscle mass. In contrast to the limb muscles, the tongue muscles of chameleons are anything but slow; they produce high forces for their cross-sectional area. These unusually fast, ballistic tongues permit them to capture a wide variety of prey. The supercontractile tongue-retractor muscles are unique among vertebrates and allow them to reel in even large vertebrate prey.
Chameleons are found among widely varying thermal regimes and climatic conditions, including hot and dry desert habitats, tropical rainforests, Mediterranean climates, and high-altitude environments. Similar to other lizards, however, chameleons carefully regulate body temperatures using behavioral thermoregulation and color change to maintain temperatures close to their preferred temperature of around 30 to 32°C. The preferred temperatures of chameleons are low as compared with those of most other diurnal lizards. The temperature invariant function of the ballistic tongue protraction and their low preferred temperatures may consequently have allowed chameleons to invade high mountain habitats rarely accessible to other lizards.
Ecologically, chameleons have taken control of the arboreal niche across Africa and Madagascar. No other reptile (aside from their avian cousins) dominates to such a degree. Walk through any montane forest in Africa or Madagascar with a spotlight at night, and you are sure to see a sleeping chameleon perched on a branch or twig. Although they are best known for their arboreal lifestyle, a number of species, indeed entire genera, are mainly terrestrial (Fig. 1.4 in color insert). While chameleons are primarily associated with forests, multiple species have capitalized on emerging habitats (grasslands and heathlands). In fact, in some places, they occupy the entire strata from ground to high canopy. Chapter 5, Ecology and Life History of Chameleons, summarizes the considerable body of studies on the interactions of chameleons with their environment. Here, we learn not only the more recognizable aspects of chameleon ecology, but are introduced to the lesser-known details. We are provided with an overview of their reproductive traits, which range from seasonal to year-round reproduction plus entire clades that are viviparous rather than oviparous. We find that although they are primarily insectivorous, their diet can be wide and may on occasion include their own kind. Even their foraging mode is thought to be exceptional among reptiles, being a combination of active foraging with sit-and-wait, termed "cruise foraging," which allows them to remain cryptic, while actively in pursuit of prey.
Despite their intensely cryptic lifestyle (Fig. 1.5 in color insert), chameleons have evolved some interesting behavioral traits. Indeed, chameleons are famous for their sometimes flamboyant ornamentation and their ability to change color. Chapter 6, Chameleon Behavior and Color Change, dispels the popular myths about chameleons and examines the how and why of their behavior. We are introduced to the physiological aspects of color change, and find that it is related to communication, camouflage, and thermoregulation (Fig. 1.6 in color insert). We also discover how ornaments are involved in sexual selection through display and aggression (Fig. 1.7 in color insert). Chameleon behavior is not limited to conspecific interactions, and includes some tricks for predator avoidance that go beyond crypsis, including free-falling from perches to avoid predation.
Moving from the individual level to the landscape level, chameleons have an interesting history and are an unusually diverse group of lizards. They are, in fact, a young clade of lizards, dating back only to the beginning of the Cenozoic, whereas most other lizard families are much older (Chapter 7, Evolution and Biogeography of Chameleons). Despite their young age, there are over 190 described species, and most experts agree that there are multiple species awaiting discovery and description (Chapter 8, Overview of the Systematics of the Chamaeleonidae). Actually, if the age of the clade is considered, the number of chameleon species is relatively high as compared with other many other squamate families (Fig. 1.8), which suggests a history of rapid lineage diversification. Chameleons are exceeded in this respect by their relatives, the Agamidae, but also Gekkonidae, Lacertidae, Scincidae, Colubrid snakes (Colubridae), and vipers (Viperidae). It appears that chameleon diversification is linked to their invasion of the arboreal habitat some 45 million years ago. Today, there are distinct centers of diversity, particularly in East Africa and Madagascar, that have elevated species richness and endemism. Although the fossil record is scarce, the clues provided by fossils that lead up to chameleons is rich, giving us insight to the early history of the group and ancestors. Chapter 9, Fossil History of Chameleons, takes us on a journey through time, beginning in the Cretaceous with the ancestors of chameleons, the priscagamids, and ends in the Holocene with fossils of extant species from Africa, Madagascar, Europe, and the Middle East.
Chapter 10, Chameleon Conservation, provides a comprehensive and up-to-date examination on the state of these lizards in our modern world. The conservation status and major threat categories are quantified and species are examined for their vulnerability. An alarming statistic that emerges, is that two thirds of chameleons already assessed by the International Union for Conservation of Nature (IUCN) are considered Threatened or Near Threatened, a figure that is much higher than that for other groups of reptiles. Of the tangible threats, habitat alteration appears to be the most prevalent, with a disproportionate impact on narrow-range endemics. Chameleons are also popular in the pet trade, and the more than 30 years of trade statistics have been summarized in this chapter. The majority of exported chameleons are harvested from wild populations, rather than through captive breeding programs, and the largest importer of chameleons is the United States, which is responsible for two thirds of the market.
The scientific literature available is surprisingly large and has filled this volume. Despite this, there is still much to learn about these intriguing animals. Even simple facts about their anatomy, physiology, and function remain unknown. Because of their cryptic nature, we know little of their daytime activities, and our impressions of their interactions with conspecifics and other animals are based on precious few studies. Aspects of their life history appear to be the most well known, yet for most species we have no understanding of their reproductive cycles, diets, home range sizes, and dispersal ability. Poorly explored forests in Africa and Madagascar reveal new species with every survey. The number of species described every year continues to rise (Fig. 1.9), and this is certainly not the end of a trend, given molecular phylogenies that show numerous undescribed lineages. There are huge gaps in the fossil record, particularly early in the history of this family, and new finds would certainly shed light on the morphology of early arboreal chameleon lineages. In exploring what we do know about chameleons, it seems we have succeeded in bringing to light the vast gap of what we do not know. But these are exciting times in the world of chameleons, and with the tools we have at our disposal today, it is clear that it will not take several more centuries to uncover many of these mysteries.
We would like to thank the authors of the these book chapters for contributing their time, expertise and energy toward making this book possible.CHAPTER 2
CHRISTOPHER V. ANDERSON and TIMOTHY E. HIGHAM
The family Chamaeleonidae is a distinctive clade of squamate reptiles with a plethora of unusual structural adaptations. Chameleons exhibit numerous distinctive features, including a laterally compressed body, forcep-like feet with toes grouped in opposing bundles, prehensile tail, enlarged casque, independently rotating eyes, and long tongue capable of being projected from the mouth. While chameleons are unique animals, they are also extremely diverse, with species spanning an approximate 20-fold range in adult total length and a 2000-fold range in body mass. Moreover, chameleons exhibit an extensive range of ornamentation. In addition, chameleons live over incredible ranges of habitats and demonstrate an abundance of variation in their behavior and ecology (Chapters 5 and 6), features of which are often predicated on anatomical specializations.
As a result of their unique nature, people have been interested in the biology of chameleons for centuries. In order to understand many aspects of chameleon biology, however, understanding the associated morphological underpinnings can be of vital importance. Here we summarize what is known about the anatomy of chameleons, emphasizing the differences between chameleons and other reptiles and the differences among chameleons.
2.1 MUSCULOSKELETAL MORPHOLOGY
The lateral compression of the chameleon in conjunction with their enlarged eyes and the formation of an enlarged casque have strong influences on the structure of the chameleon skull and the distribution and orientation of various cranial muscles. The extent to which these characteristics are developed, however, is also variable within the family, and thus examination of the anatomy of the skull and its musculature is not only of interest relative to other lizard groups, but also within the chameleons.
Skull and Teeth The extensive studies of the chameleon skull have included discussion and examination of the structure of the skull in Archaius (Hillenius, 1988; Rieppel and Crumly, 1997), Bradypodion (Fig. 2.1c,d,f) (Parker, 1881; Methuen and Hewitt, 1914; Brock, 1941; Engelbrecht, 1951; Rieppel, 1981; Hillenius, 1988), Brookesia (Fig. 2.1a,b) (Siebenrock, 1893; Methuen and Hewitt, 1914; Rieppel 1987; Rieppel and Crumly, 1997), Calumma (Methuen and Hewitt, 1914; Hillenius, 1988; Rieppel and Crumly, 1997), Chamaeleo (Parker, 1881; Siebenrock, 1893; Methuen and Hewitt, 1914; Prasad, 1954; Rieppel, 1987; Hillenius, 1988), Furcifer (Methuen and Hewitt, 1914; Hillenius, 1988; Rieppel and Crumly, 1997), Kinyongia (Hillenius, 1988), Rhampholeon (Werner, 1902b; Methuen and Hewitt, 1914; Frank, 1951; Rieppel, 1987; Hillenius, 1988), Rieppeleon (Rieppel, 1987; Hillenius, 1988), and Trioceros (Fig. 2.1e) (Rieppel, 1981, 1987, 1993; Hillenius, 1988). These studies have amassed a list of variations between the skulls of different genera and developmental stages. However, they have also resulted in varying interpretations of the skull bones, particularly of the temporal region. Here we summarize the morphology of the adult skull in chameleons following the terminology and interpretations of Rieppel (1981).
The premaxilla of chameleons is unpaired (fused) and lies medially between the maxillae (Fig. 2.1b,d) (Parker, 1881; Siebenrock, 1893; Werner, 1902b; Brock, 1941; Engelbrecht, 1951; Frank, 1951; Rieppel, 1981, 1987). The premaxilla in Brookesia (Siebenrock, 1893), Chamaeleo (Siebenrock, 1893), Rieppeleon (Rieppel; 1987), and Rhampholeon (Werner, 1902b; Frank, 1951) bear two vestigial teeth, whereas the premaxillae in Trioceros do not bear teeth (Rieppel, 1981). Engelbrecht (1951) reports that Bradypodion pumilum also lack premaxillary teeth. Rieppel (1981), on the other hand, observed indications of two vestigial teeth fused with the premaxilla, noting, however, that histological investigation is required to verify whether these are true teeth or paired bony projections on the transverse process of the premaxilla.
Ventrally, the vomerine (palatal) process of the premaxilla is reduced (Fig. 2.1f) (Romer, 1956), extending only a short distance posteriorly, and in Bradypodion (Fig. 2.1f) (Engelbrecht, 1951; Frank, 1951; Rieppel, 1981), it does not contact the vomer, as the palatal process of the maxillae meets behind the premaxilla. The maxillae do not meet behind the premaxilla in Brookesia (Siebenrock, 1893; Rieppel and Crumly, 1997), however. Typically, the vomers are fused (unpaired) in chameleons (Frank, 1951; Rieppel, 1981); however, in Bradypodion pumilum, the vomers are paired for most of their length and fused only anteriorly where they join with the maxillae (Engelbrecht, 1951; Rieppel, 1981). In Archaius, the vomer is paired (Rieppel and Crumly, 1997). At the posterior end, the vomer joins the palatines (Fig. 2.1f) (Engelbrecht, 1951; Rieppel, 1981). The palatines extend posteriorly and flare laterally to join the maxillaries (Fig. 2.1f) (Werner, 1902b; Engelbrecht, 1951; Rieppel, 1981).
Dorsally, the nasal process of the premaxilla extends posteriorly and fully separates the maxillae in most species (Fig. 2.1b,d) (Werner, 1902b; Engelbrecht, 1951; Frank, 1951;Rieppel, 1981, 1987; Rieppel and Crumly, 1997). In Bradypodion, Brookesia, and Chamaeleo, the posterior edge of the nasal process of the premaxilla meets the fused nasals (Fig. 2.1b,d) (Seibenrock, 1893; Camp, 1923; Parker, 1942; Engelbrecht, 1951; Romer, 1956; Rieppel, 1981, 1987). However, the nasal process of the premaxilla separates the nasals and meets an anterior process of the frontal in Rhampholeon and Rieppeleon (Werner, 1902b; Parker, 1942; Frank, 1951, Rieppel, 1981, 1987). The nasals are paired and variably separated from the frontals by the premaxilla in Calumma and Furcifer (Rieppel and Crumly, 1997).
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Table of Contents
Contributors Foreword 1 Biology of the Chameleons: An
Introduction Krystal A. Tolley and Anthony Herrel 2 Chameleon Anatomy Christopher V. Anderson and Timothy E. Higham 2.1 Musculoskeletal Morphology 2.2 External Morphology and
Integument 2.3 Sensory Structures 2.4 Visceral Systems 3 Chameleon Physiology Anthony Herrel 3.1 Neurophysiology 3.2 Muscle Physiology 3.3 Metabolism, Salt, and Water Balance 3.4 Temperature 3.5 Skin Pigmentation, Color Change, and the Role of Ultraviolet Light 3.6 Developmental Physiology 4 Function and Adaptation of Chameleons Timothy E. Higham and Christopher V. Anderson 4.1 Locomotion 4.2 Feeding 5 Ecology and Life History of Chameleons G. John Measey, Achille Raselimanana, and Anthony Herrel 5.1 Habitat 5.2 Life-History Traits 5.3 Foraging and Diet 5.4 Predators 6 Chameleon Behavior and Color Change Devi Stuart-Fox 6.1 Sensory Systems and Modes of Communication 6.2 Color Changes 6.3 Social and Reproductive Behavior 6.4 Sexual Dimorphism: Body Size and Ornamentation 6.5 Antipredator Behavior 7 Evolution and Biogeography of Chameleons Krystal A. Tolley and Michele Menegon 7.1 Evolutionary Relationships 7.2 Diversity and Distribution 7.3 Regional Diversity 7.4 Patterns of Alpha Diversity 7.5 Patterns of Beta Diversity 8 Overview of the Systematics of the Chamaeleonidae Colin R. Tilbury 8.1 Evolution of Methodology in Chameleon Taxonomy 8.2 Current Status of Taxonomy of the Chamaeleonidae 8.3 Subfamilial Groupings within Chamaeleonidae 8.4 Overview of Extant Genera 9 Fossil History of Chameleons Arnau Bolet and Susan E. Evans 9.1 Phylogenetic Relationships of Iguania and Acrodonta 9.2 Fossil Record of Acrodonta 9.3 Origins of Acrodonta 9.4 Origins of Chamaeleonidae 10 Chameleon Conservation Richard Jenkins, G. John Measey, Christopher V. Anderson, and Krystal A. Tolley 10.1 Conservation Status of Chameleons 10.2 Trade in Chameleons 10.3 Chameleons and Global Change 10.4 The Way Forward Appendix Abbreviations References Photo Credits