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On the Structure and Function of Tails in Snakes: Relative Length and Arboreality

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PAGE 1

1 ON THE STRUCTURE AND FUNCTION OF TAILS IN SNAKES: RELATIVE LENGTH AND ARBOREALITY By COLEMAN MATTHEW SHEEHY III A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2006

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2 Copyright 2006 By Coleman Matthew Sheehy III

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3 To the increased understanding, respect, and conservation of snakes worldwide

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4 ACKNOWLEDGMENTS I thank my committee members Harvey B. Lillywhite (chair), James Albert, and Max A. Nickerson for their guidance and support. Harvey B. Lillywhite first inspired this work by suggesting a link between snake tails and the gravita tional hypothesis. I thank Harvey B. Lillywhite and Michael B. Harvey for providing unpublished snake length data, Florida Museum of Natural History (FLMNH) curators F. Wayne King and Max A. Nickerson for access to the Herpetology collection, Roy McDiarmid and George Zug at the United States National Museum (USNM) for permission to use the Herpetology collection, and Steve Gotte at the USNM and Kenney L. Krysko at the FLMNH for supplying data from the Herpetology collection databases. I thank Michael B. Harvey, Laurie Vitt, James McCranie, Rom Whitaker, Bob Henderson, Bob Powell, Blair Hedges, Ming Tu, Max A. Nickerso n, Richard Sajdak, Bill Love, and John Rossi for providing difficult to find snak e natural history information. I thank Michael B. Harvey and Ron Gutberlet for information on snake phylogenies, Michael McCoy for assistance with various statistic al programs and analys es, and Kent Vliet for the use of a Macintosh computer. I thank A ndres Lopez and Griffin Sheehy for help with illustrating programs, Andres Lopez for help wi th phylogenetic programs, and Jason Neville for computer assistance. I thank Glades Herp In c. for permission to measure live snakes and, perhaps more importantly, I am grateful to Russel L. Anderson, Sam D. Floyd and Ryan J. R. McCleary for assistance in handling and measur ing live snakes, many of which were large, highly venomous, and uncooperative. In addition to my committee members, I thank Ryan J. R. McCleary, David A. Wooten, Leslie Babonis, Chris Samu elson, Bruce Jayne, and Roy McDiarmid for stimulating discussions regarding snake natural history and evolution. I thank Griffin E. Sheehy and Andrea Martinez for all their patience, love and support. Finally, I want to

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5 thank my parents Coleman M. Sheehy, Jr. and Elle n R. Sheehy for never failing to support and encourage a young boy’s endless passion for snak es. I could not have been more lucky.

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6 TABLE OF CONTENTS page ACKNOWLEDGMENTS...............................................................................................................4 LIST OF TABLES................................................................................................................. ..........8 LIST OF FIGURES................................................................................................................ .........9 ABSTRACT....................................................................................................................... ............10 CHAPTER 1 INTRODUCTION................................................................................................................. .12 Gravitational Influence on Tail Morphology..........................................................................13 Length Limitations............................................................................................................. .....14 Relative Tail Length and Macrohabitat Use...........................................................................15 Multiple Functions of Tail Use...............................................................................................17 Locomotion..................................................................................................................... .17 Caudal Luring.................................................................................................................. 20 Defense........................................................................................................................ ....21 Morphology and Reproduction........................................................................................24 Sexual Dimorphism and Ontogenetic Shifts...................................................................25 2 MATERIALS AND METHODS............................................................................................27 Categorizing Climbing in Snak es: Gravitational Habitat.......................................................27 Analyses....................................................................................................................... ...........31 Frequency Distribution....................................................................................................31 Gravitational Habitat and Total Body Length.................................................................31 Gravitational Habitat and Tail Length.............................................................................32 Constructing the Phylogeny............................................................................................32 Relative Tail-Length, Gravitati onal Habitat, and Phylogeny..........................................34 3 RESULTS...................................................................................................................... .........48 Frequency Distribution......................................................................................................... ..48 Gravitational Habitat and Total Body Length........................................................................48 Gravitational Habitat and Tail Length....................................................................................49 Relative Tail-Length, Gravitati onal Habitat, and Phylogeny.................................................49 4 DISCUSSION................................................................................................................... ......63 Relative Tail Length, Gravity, and Climbing in Snakes.........................................................63 Snake Tails and Defense: Speed and Pseudautotomy............................................................67 APPENDIX STENOTOPICA LLY ARBOREAL SPECIES........................................................71

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7 LIST OF REFERENCES............................................................................................................. ..73 BIOGRAPHICAL SKETCH.........................................................................................................85

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8 LIST OF TABLES Table page 2-1 Taxa and associated data sources for 227 snake species included in this study................36 2-2 Total variation in relative tail -lengths of 26 species of snakes..........................................43 3-1 The 227 species included in this study..............................................................................50 A-1 Stenotopically arboreal species..........................................................................................71

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9 LIST OF FIGURES Figure page 2-1 Composite phylogeny of 227 snake taxa...........................................................................44 3-1 Snake total length frequency distribution..........................................................................56 3-2 Coefficients of variation of log10 transformed total length................................................58 3-3 Regression of log10 tail length on log10 total length...........................................................59 3-4 Analysis of covariance (ANCOVA) on corrected tail-length for 227 species of stenotopically arboreal.......................................................................................................6 0 3-5 Analysis of covariance (ANCOVA) on corrected tail-length for 227 species of scansorial..................................................................................................................... .......61 3-6 Independent contrasts...................................................................................................... ...62 4-1 Regression of absolute tail length on relativ e tail-length...................................................70

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10 Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science ON THE STRUCTURE AND FUNCTION OF TAILS IN SNAKES: RELATIVE LENGTH AND ARBOREALITY By Coleman Matthew Sheehy III December 2006 Chair: Harvey B. Lillywhite Major Department: Zoology The pervasive effect of gravity on blood circ ulation is an important consequence for elongate animals such as snakes that utilize arbor eal habitats. Upright pos tures create vertical gradients of hydrostatic pressures within circulatory vessels, and the magnitude of the pressure change is proportional to the tota l length of the blood column. In air, this potentially induces blood pooling and edema in dependent tissues and a decrease in the volume of blood reaching the head and vital organs of animals that are ta ll or elongate. Arboreal sn akes exhibit a suite of behavioral, morphological, and phys iological adaptations for counter ing the effects of gravity on blood circulation including rela tively non-compliant tissue compartments in the tail. Comparative studies involving arboreal, te rrestrial, and aquatic species have demonstrated that blood pooling in dependent tissues of arboreal snakes can be tenfold lower than in terrestrial and aquatic species. In addition to non-comp liant tail compartments, arboreal species appear to have longe r tails relative to their non-cl imbing terrestrial and aquatic counterparts. However, the generality of tail le ngth patterns related to arboreal habitats and gravity has not been previously studied in a br oad range of taxa. Here I examine the hypothesis that there are constraints limiting the total length of arboreal snakes and that arboreal snakes

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11 have relatively longer tails to help counter the effects of gravity wh ile also helping to adjust total length to within the ra nge of limitations. I used macrohabitat and behavior to create f our gravitational habitats to categorize the amount of climbing and, therefore, the amount of gravitational st ress experienced: stenotopically arboreal, eurytopically arboreal/terre strial, stenotopically terrestrial and stenotopically aquatic. The effects of sexual dimorphism, and ontogenetic shifts in allometry and habitat use, were avoided by using only adult females. Data were acquired from the literature, museum specimens, and live snakes. Frequency distribu tions and analysis of variance (ANOVA) were used to evaluate the assumption that total lengt h limitations exist in stenotopically arboreal species. I tested for differences in relative tail-length (RTL) among the four gravitational habitats (227 species representing almost all snake families and subfamilies) by using analyses of covariance (ANCOVA). I tested for correlations between macrohabitat use and RTL within a phylogenetic framework by independent contrasts anal yses after constructing a composite tree. The RTL for the 227 species included in this study ranged from 1.1% ( Rhinotyphlops episcopus ) to 48.1% ( Uromacer frenatus ). The total length of st enotopically arboreal species appears to be constrained to between 50 a nd 200 cm, with few exceptions. The RTL between stenotopically arboreal and eury topically arboreal/terrestrial sp ecies did not differ and were therefore combined into a single scansorial category. Scansorial species had RTLs on average two times longer than non-scansori al species. Snakes with relativ ely longer tails have a larger percent of dependent vessels contai ned within the tight integument of the tail and are thus more resistant to blood pooling experienced during clim bing. Therefore, the relatively long tails of scansorial snakes can be regarded as one of a su ite of characters likely to be adaptive responses to the cardiovascular constraints on blood circulation imposed by gravity.

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12 CHAPTER 1 INTRODUCTION Macrohabitat use is thought to strongly influence the evol ution of vertebrate morphology (Miles and Ricklefs 1984; Wikramanayake 1990; Lillywhite 1996; Martins, Araujo, Sawaya, and Nunes 2001). Even in limbless and elongate vert ebrates such as snak es, the evolution of morphology is strongly influenced by the type of macrohabitat utilized (Vitt and Vangilder 1983; Guyer and Donnelly 1990; Cadle and Greene 1993; Lillywhite a nd Henderson 2001; Martins et al. 2001). The roughly 3000 extant snake species ha ve demonstrated a remarkable propensity for radiating into a wide variety of fossorial, arboreal, terrestrial, fr eshwater, and marine habitats in every part of the biosphere excluding only th e deep sea and polar regions. This diverse ecological radiation, along with the loss of limbs, makes sn akes excellent animals for investigating the effects of macrohabitat use on the evolution of morphology. Arboreality has evolved numer ous times independently among snakes, and several studies have identified various behavioral, morphological and physiological charac teristics associated with snakes in arboreal habitats (e.g., Johnson 1955; Marx and Rabb 1972; Henderson and Binder 1980; Lillywhite 1993; Lillywhite and Henderson 2001; Ma rtins et al. 2001). While it is often difficult to demonstrate adaptation (e.g., Gould and Lewontin 1979; Bock 1980), the fact that a phylogenetically di verse array of arboreal sp ecies share strikingly similar, if not identical, morphological, behavioral, and phys iological specializations suppor ts the inference that these traits are adaptive and, in many cases, the re sult of convergent evolution (Lillywhite 1996; Lillywhite and Henderson 2001). Studies investigating (directly or indirectly) snakes’ tails have incl uded a wide range of topics including cardiovascular adaptations (e.g., Lillywhite 1985, 1993, 2005; Lillywhite and Henderson 2001), sexual dimorphism (e.g., Kl auber 1943; King 1989; Shine 1993, 2000), mating

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13 success (e.g., Shine et al. 2000), use in loco motion (e.g., Jayne 1988; Jayne and Bennett 1989), the evolution of vertebral elements (e.g., Johnson 1955; Lindell 1994; Shine 2000), defensive adaptations (e.g., Greene 1973, 1988; Arnold 1988; Mendelson 1991; Savage and Slowinski 1996), caudal luring (e.g., Greene 1997), deve lopment (Polly, Head, and Cohn 2001), and ecological correlations (e.g., Guyer and Donnell y 1990; Martins et al. 2001; Lobo, Pandav, and Vasudevan 2004; Wster, Duarte, and da Graa Sa lomo 2005; Wiens, Brandley, and Reeder 2006). However, the adaptive role of the tail in arboreal habita ts is poorly understood. For example, the importance of the tail in snake locomotion (particularly in arboreal habitats) remains unclear (Jayne 1988; see below). Gravitational Influence on Tail Morphology An important consequence of utilizing complex, three-dimensional vertical habitats is the pervasive effect of gravity on blood circulation, which can be pa rticularly pronounced in tall or long organisms such as giraffes and snakes (Lillywhite 1993,1996). Up right postures create vertical gradients of hydrostatic pressures within circulatory vesse ls. In air, this increases transmural pressures related to the absolute lengt h of the fluid column, leading to a tendency for blood pooling and for fluid to filter from capillaries into surrounding tissue compartments resulting in edema in dependent tissues (Li llywhite 1985, 1993). As blood pooling and edema increase, the volume of blood reaching the heart decreases, thereby redu cing the central blood pressure and consequently the amount of blood fl ow to the head and vital organs (Lillywhite 1993b, 1996, 2005; Lillywhite and Henderson 2001). Arboreal snakes exhibit a suite of behavioral, mor phological, and physiological adaptations for countering the e ffects of gravity on blood circulat ion including stereotypical body movements, small mass/length ratio s, relatively anterior hearts, tightly applied integument, and relatively non-compliant tissue compartments in the tails (Lillywhite 1985, 1993; Lillywhite and

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14 Henderson 2001). Comparative st udies involving arboreal, terr estrial, and aquatic species demonstrate that blood pooling in dependent tissue s of arboreal snakes during vertical posture is significantly less than in terrestrial and aquati c species, and these differences can approach tenfold (Lillywhite 1985, 1993; Lillywhite and He nderson 2001). The ability for arboreal snakes to defend against edema and blood pooling is almost certainly at tributed to the aforementioned characters, one of which being relatively non-compliant tissue comp artments in the tail (Lillywhite and Henderson 2001; Lillywhite 2005). However, the magnitude of the compliance does not appear to be related to the length of th e snake or its tail (Lillywhite 1993). There does, however, appear to be a relati onship between tail length and macr ohabitat use in that arboreal species generally have longer tails relative to their nonclimbing terrestrial and aquatic counterparts (see below). What th en are the selective pressures re sponsible for the interspecific variation in tail length? Length Limitations Body size may be the most fundamental charac ter of an organism, because nearly all aspects of an organism’s biology are correlated with this variable (Naganuma and Roughgarden 1990; Boback and Guyer 2003). The body sizes (i.e., to tal length) of snakes in general could be evolving toward an optimal length of 1.0 m (Bob ack and Guyer 2003), suggesting that selection and constraint might be influencing the total le ngth of many snake species. However, the body size distribution is not obviously skewed in either direction, and idiosyncratic features of the natural history of snakes may be creating this distribution pattern (B oback and Guyer 2003). Macrohabitat use likely imposes locomotor constraints upon snakes (Boback and Guyer 2003), and there is considerable evidence that arboreal snakes are limite d in their use of habitat as a consequence of interactions between their mo rphology and the physical size of branches (see Lillywhite and Henderson 2001, for a review on arboreal snake functional ecology). Arboreal

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15 locomotion involves various combinations of undul atory, rectilinear, and concertina movements along a complex and often unstable vertical th ree-dimensional substrate (Gans 1974; Edwards 1985; Lillywhite and Henderson 2001). Short snakes might not be able to adequately span gaps, whereas large or heavy bodied snakes might not be supported by smaller branches (Henderson and Nickerson 1976). Long snakes might be fu rther limited physiologically by cardiovascular constraints. Because foraging snakes must be ab le to span gaps between stems and branches and approach prey without revealing their presen ce (Lillywhite and Henderson 2001), it is reasonable to expect there are upper and lower limitations on total length in arbor eal snake species. Assuming there are length limitations imposed on snakes living in arboreal habitats, one way snakes could have evolved total lengths with in the acceptable range while still counteracting blood pooling and assuring adequate blood flow to th e head is by lengthening the tail relative to the body since perivascular tissues are tighter in the tail than in the body cavity (Lillywhite 2005). Lengthening the tail relati ve to the body could potentially be achieved evolutionarily by adding vertebrae to the tail or by elongating the caudal vertebrae themselves (Johnson 1955; Shine 2000). Relative tail-length (RTL), which is the proportional ratio of tail length/total length, is commonly used when performing inters pecific comparisons in tail length to account for total length differences (Klauber 1943). Rela tive tail-length in snakes appears to have low intraspecific variation (i.e., it is a stable charact er), and this variation is further reduced when investigating the sexes separate ly (Klauber 1943). Therefore, interspecific similarities and differences in RTL are potentially useful s ources of ecological and taxonomic information (Klauber 1943). Relative Tail-Length and Macrohabitat Use Correlations between RTL and m acrohabitat use have been su ggested previously, but the supporting data are weak or inconclusive. Klau ber (1943) suggested that thick-bodied snakes

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16 have relatively shorter tails than do thin-bodied snakes. Clark (1967) stated that fossorial habits seem to be accompanied by a shortening of the tail. Marx and Rabb (1972) investigated 962 species of 195 colubrid genera and found that the maximum number of subcaudal scales occurred in six species of snakes that were all arboreal (i.e., vi ne snakes). Because these six species are not all closely related, they conclude d that these characteristics represent derived ecological specializations for ar boreal habitats. King (1989) proposed that RTL in snakes is highly variable interspecifically a nd appears to be correlated with ecological factors. He also stated that other considerations such as mode of locomotion, habita t, and risk of predation, which while apparently correlated with tail length in li zards, have not been investigated in snakes. Greene (1997) stated qualita tively that the tails of phyl ogenetically basal snakes (scolecophidians) and most vipers are especially short, while the tails of many colubrids and elapids are longer. Martins et al (2001) investigat ed 20 species of Bothrops and concluded that an increase in RTL occurred along with an increase in arboreality in some clades. A strong method for assessing the influence of habitat use on the evol ution of body form in snakes is to analyze monophyletic clades so th at characters can be interpreted within an explicitly phylogenetic framework (Harvey and Pagel 1991; Martins et al. 2001). Although earlier studies have found a corr elation between RTL and arboreality in snakes, most have not separated species by lineages and thus might be confounded by phylogenetic effects. Martins et al. (2001) addressed this issu e using the monophyletic genus Bothrops and their results corroborate previous studies. However, the st udy investigated only 20 species within a single clade, making the results difficult to apply to snak es in general. Similar studies using additional monophyletic groups are needed to determine whethe r this trend is widesp read in snakes.

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17 Herein, I investigate the relationship between tail length and macrohab itat use in arboreal species of snakes. The purposes of the present in vestigation are threefold. First, I evaluate the assumption that there are limitations imposed on th e total length of arbor eal species. Second, I test the hypothesis that arboreal species have relatively longer tails than nonclimbing species. Third, I discuss the relationship between RTL a nd climbing within the c ontext of gravitational adaptation, and hypothesize that addi tional selective pressures might be directing the evolution of relatively long tails in snakes. Multiple Functions of Tail Use Most vertebrate structures have multiple functions (Moon 2000), and snake tails are an excellent example. As a likely consequence of limblessness, several selective pressures are simultaneously acting on the tails of many snake species. For ex ample, many juvenile pitvipers (e.g., Agkistrodon piscivorus ) use caudal luring to attract prey, but also vibrate th eir tails rapidly in defense when threatened. In order to unders tand the functional versatil ity, as well as possible constraints of complex vertebrate structures such as snake tails, information about how they are used in diverse environments and behaviors is required (Moon 2000). Locomotion Snakes likely use their tails to assist in locomotion (e.g., balance, propulsion, holding or climbing) (Klauber 1943); however, the extent to which this occurs is poorly understood (Jayne 1988; Lillywhite and Henderson 2001). Many fosso rial species have short blunt tails (e.g., Eryx Sympholis lippiens scolecophidians, and uropeltids), and so me also have strongly keeled caudal scales (e.g., Sonora aemula and uropeltids). The apparent commonality of these characteristics among fossorial species suggests th ey are adaptations to fossorial habitats (e.g., locomotion and defense). However, in many cases the functi onal morphology is not well studied. Clark (1967) posits that the short tails of many fossorial snakes (i.e., scolecophidians) f acilitate subterranean

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18 locomotion by acting as an anchor against which th e snake can push, since a long tail in this case would likely bend and be unable to serve this purpose. However, Klauber (in Clark 1967) suggests instead that burrowing woul d render the tail practically us eless, and that the tails of fossorial snakes are shortened secondarily due to non-use, rather than from direct selective pressures. In a combination of experimental and co rrelative analyses, Jayne and Bennett (1989) demonstrated the difficulties in determini ng the effect of tail morphology on locomotor performance in terrestrial snakes. Jayne and Bennett demonstrated that losses ranging from 0.03% to 80.4% tail length had no significant effect on terrestrial locomotory performance in 52 garter snakes ( Thamnophis sirtalis ) with naturally incomplete tails. Locomotor performance was not affected by the experimental removal of the di stal one third of the tail. The experimental removal of the distal two thirds of the tail only caused a small (4.5 %) but significant average decrease in speed. However, the same study also found that burst speed in T. sirtalis performing terrestrial lateral undulation was fastest in individu als with intermediate re lative tail-lengths. Jayne and Bennett concluded that minor deviations from intermediate relative tail length do not affect locomotor performance among snakes. Th ese results demonstrat e that long tails can perhaps be slightly disadvantageous in terms of locomotor performance in T. sirtalis and suggest that the long tails of some snake species are possi bly due to other additional selective pressures. For example, high incidence of tail loss is found in the genera Nerodia and Thamnophis suggesting tail loss from predation attempts (King 1987; Jayne and Bennett 1989; Mendelson 1991; see section below on tail lo ss). Furthermore, juvenile Nerodia harteri have been observed using their long tails to anchor th emselves to rocks at the water’s edge while fishing (Rossi, pers.

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19 comm.), whereas Nerodia sipedon has been observed using their ta ils to anchor themselves to submerged sticks and rocks while fishing in curre nts (M. A. Nickerson pers. comm.; pers. obs). Snake caudal vertebrae are complex, highly variab le in size within individuals, and clearly distinguishable from the trunk vertebrae (Johnson 1955). Because the number of caudal vertebrae corresponds to the number of subcaudal scales by a ratio of 1:1 in most snake species, snakes with longer tails typically have more s ubcaudals and thus more caudal vertebrae than snakes with shorter tails (R uthven and Thompson 1908; Gans and Taub 1965; Alexander and Gans 1966; Voris 1975; Shine 2000). Differences in the relative size and number of caudal vertebrae suggest that th e effectiveness of the ta il in locomotory propulsi on varies considerably among snake taxa (Jayne 1988). However, the consequences of caudal morphology likely vary with mode of locomotion (Jayne 198 8). More research is needed to determine the extent of tail use in the various forms of snake locomotion. The tails of most arboreal boids, viperids, and many colubrids are prehen sile and assist in climbing and securing to branches (Lilly white and Henderson 2001). Tree boas ( Corallus ) often wrap their prehensile tails around branches dur ing prey capture and consumption (Henderson 2002; pers. obs.). Henderson (2002) observed tree boas ( Corallus grenadensis ) on Grenada hanging by their tails from vegetation, with the forepart of their bodie s in typical ambush posture, presumably hunting for bats. I ha ve observed similar hunting behavior in Corallus ruschenbergerii on Tobago and in Corallus caninus in captivity. However, there appears to be no correlation between long RTL and tail prehen sion for snakes in general (Lillywhite and Henderson 2001). Specializations in snake locomotion, such as highly efficient sw imming and sidewinding, have likely evolved multiple times and employ tail use to varying degrees. Sea snakes and sea

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20 kraits have evolved several adaptations to a ma rine existence, one of which being flattened, oarlike tails used for effective and rapid propul sion (Heatwole 1999). Yellow-bellied sea snakes ( Pelamis platurus ) are well adapted to a pelagic existence and have subsequently lost the ability to crawl on land, spending their entire lives at sea (Greene 1997). Acroch ordids have a muscle that shapes the skin of the body and tail into a keel while sw imming (Heatwole 1999). However, not all marine snakes have oarlike tails. Some homalopsines have tails that are only slightly compressed, whereas natricines and other homalopsin es have tails similar in size and shape to many terrestrial species (Heatwole 1999). Sidewinding is often considered to be a sp ecialized mode of locomotion (Jayne 1988). However, it occurs in a surprisingly wide divers ity of taxa including booid and colubroid snakes (Gans and Mendelssohn 1972). Jayne (1988), in a study of snake locomotion, concluded that the tails of snakes are unlikely ab le to produce the movements and forces necessary for sidewinding and thus are likely contributing little during this form of locomotion. Caudal Luring Several snake species within the Boidae, Vipe ridae, Elapidae, and Colubridae use caudal luring to attract insectivorous pr ey by wiggling the often contra stingly colored tail tip (Sazima and Puorto 1993; Greene 1997). These taxa include Boa constrictor (Radcliffe, Chiszar, and Smith 1980) Morelia viridis (Murphy, Carpenter, and Gillingham 1978), Calloselasma rhodostoma (Schuett 1984; Daltry, Wuster, and Thorpe 1998) Cerastes vipera (Heatwole and Davison 1976) Daboia russelii (Henderson 1970) Agkistrodon (Neill 1948; Allen 1949; Wharton 1960; Carpenter and Gillingham 1990) Crotalus (Kauffeld 1943), Hypnale (Whitaker and Captain 2004), Sistrurus (Jackson and Martin 1980; Rabatsky and Farrell 1996) Bothriopsis (Greene and Campbell 1972), Bothrops (Sazima 1991) Acanthophis (Carpenter, Murphy and Carpenter 1978; Chiszar, Boyer, Lee, Murphy, and Radcliffe 1990) Alsophis portoricensis (Leal

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21 and Thomas 1994) Pantherophis obsoleta (Tiebout 1997) Tropidodryas striateceps (Sazima and Puorto 1993) and Madagascarophis (W. Love pers. comm.). Caudal luring has been reported primarily in juveniles (80% of the sp ecies known to caudal lure) to attract small, visually oriented prey such as anurans and lizards into striking range (Neill 1960; Parellada and Santos 2002). Cessation of this behavior often occurs within the first year or two with an ontogenetic shift in diet towards larger endothermic prey items such as mammals and birds (Neill 1960; Jackson and Martin 1980; Daltry et al. 199 8). However, in some species such as Acanthophis antarcticus (Carpenter et al. 1978), Bothriopsis bilineata (Greene and Campbell 1972), Cerastes vipera (Heatwole and Davison 1976), and Sistrurus miliarius (Jackson and Martin 1980), the behavior persists into adulthood. Pe rsistence of caudal lu ring in these species has been attributed to the importance of insectiv orous prey items in their adult diets (Jackson and Martin 1980). Defense Snakes exhibit the most elaborate antipreda tor behaviors among rept iles (Greene 1988). These behaviors range from generalized to specia lized, and many involve the tail. Postcloacal scent glands are found in all snake species and exude an offensive odor (Whiting 1969). These noxious secretions likely enhance the effectiven ess of cloacal discharge (fecal material) in deterring predators (Greene 1988). Defensive tail displays are widespread in snakes; Greene (1973) identified 73 snake species in at least six families known to pe rform unusually conspicuous tail displays. Characteristics of these disp lays include elevating (e.g., Eryx and Charina ), tightly coiling (e.g., Diadophis and Farancia ), or waving (e.g., Micrurus and Micruroides ) the tail, which may be long or blunt, and brightly colo red or drab. When threatened, Rhadinaea decorata often hides its head beneath coils while raising and wri ggling its tail (Campbell 1998), whereas Oligodon

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22 albocinctus and Sinomicrurus macclellandi flatten their bodies and curl up the end of their tails (Whitaker and Captain 2004). Calliophis melanurus and C. nigrescens raise and coil their tails when disturbed (Whitaker and Captain 2004). Chilorhinopus butleri and C. gerardi hide their heads within coils and wave their tails in the ai r in defense (Spawls, Howell, Drewes, and Ashe 2002). Tail length and thickness correlate with tail in jury rate and defensive behaviors in the genera Eryx and Gongylophis (Greene 1973; R. Sajdak pers. co mm.). Species exhibiting tail displays (e.g., E. jaculus E. johnii E. miliaris and E. tataricus) have longer and thicker tails, small heads, and a relatively high frequency of tail injury. However, species that rely on a biting defense instead of tail displays (e.g., G. colubrinus G. conicus, and possibly E. somalicus ) have shorter, thinner tails, larger and more distinct heads, and a lower frequency of tail injury. These displays likely disorient potenti al predators and, in many species serve as a decoy to divert attack from the head to the tail (Greene 1973). Many species have tails with specialized external morphological characters used for defense. Several terrestria l and fossorial species (e.g., Carphophis, Contia tenuis Farancia Oligodon affinis and several typhlopids) posse ss a tail spine, which is pressed against an attacker (Leonard and Stebbins 1999; Whitaker and Captain 2004). Juvenile Farancia abacura have also been observed using this tail spine to impale or “pop” tadpol es normally too large to be swallowed (Rossi 1992: 97). Uropeltids have enlarg ed, reinforced tail tips with specialized scale morphology that collect dirt to form protective plugs while burrowing (Gans 1976; Gans and Baic 1977). Cutaneous photorecepti on, or dermal light sense, in the tail of the sea snake Aipysurus laevis aides in concealment from visually oriented predators (Zimmerman and Heatwole 1990). These sea snakes often hi de in clumps of coral and use cutaneous

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23 photoreception to avoid having thei r tails exposed to light, and thus vulnerable, during the daytime. The genera Crotalus and Sistrurus form the monophyletic group of approximately 31 species of New World pitvipers characterized by the presence of a ratt le (Greene 1988). The rattle is associated with a suite of anatomical, physiological, and be havioral specializations and is generally accepted as being used primarily in defensive signaling (see Moon 2001, for summary of rattle evolution). However, the evolution of this unique stru cture remains unresolved (Greene 1988; Moon 2001). A potential beha vioral precursor to the rattle is tail vibrating, a defensive behavior widespread in snakes where the tail is rapidly vibrated against loose surface debris to produce a buzzing sound (Greene 1988). Caudal pseudautotomy occurs in several snake species as a defensive strategy (Savage and Slowinski 1996). However, this behavior has on ly been documented in snakes with relatively long tails and is thought to be rare (Arnold 1988; Marco 2002). Enulius Scaphiodontophis and Urotheca possess morphological specializa tions that likely facilita te pseudautotomy such as long, fragile tails with thick bases (Savage and Sl owinski 1996). However, high incidence of tail loss has also been observed in several othe r typically long-taile d genera such as Alsophis (Seidel and Franz 1994), Amphiesma and Boiga (Whitaker and Captain 2004), Coluber (Marco 2002) Coniophanes, Rhadinaea Sibynophis and Thamnophis (Jayne and Bennett 1989; see Mendelson 1991, for review), Dendrophidion, (Duellman 1979), Drymobius (Mendelson 1991), Gastropyxis (as Hapsidophrys) Grayia and Mehelya (Spawls et al. 2002), Nerodia (King 1987), Natriceteres and Psammophis (Broadley 1987), Pliocercus (Smith and Chizar 1996) and Xenochrophis (Ananeva and Orlov 1994), although these genera do not appear to possess the morphological

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24 specializations for tail loss present in Enulius Scaphiodontophis and Urotheca (Savage and Slowinski 1996). While both intervertebral pseudautotomy and intr avertebral autotomy occur in lizards, only intervertebral pseudautotomy is known to o ccur in snakes (Arnold 1988; see Savage and Slowinski 1996, for a review of terminology). Unlike many lizard species, pseudautotomy in snakes does not appear to be unde r neural control (Arnold 1988) a nd requires physical resistance, which is often facilitated by twis ting or rotating the body in one direction until the tail snaps off (Savage and Slowinski 1996; Marco 2002). However, reflex action often allows the segment of lost tail to continue thrashi ng, thus likely distracting the pr edator and allowing the snake to escape (Marco 2002; Savage 2002). Although caudal autotomy is a lepidosaurian synapomorphy, the ability to regenerate any portion of the tail has been lost in all snake species (Pough et al. 2001). Morphology and Reproduction Peters (1964) defines the snake tail as the sec tion of the body posterior to the cloaca. In males, the hemipenes and associated retractor mu scles are located in the tail, while the testes (and ovaries in females) are located within the body cavity. In both sexes of all species, the tail also contains a pair of scent glands (Whiting 196 9). Male snakes typically use their tails during courtship to gain access to the female’s cloa ca by performing a tail-sear ch copulatory attempt (TSCA) until the cloacas meet (G illingham, Carpenter, and Murphy 1983). This TSCA is often preceded by various tail moveme nts by the female, such as tail-whipping and tail-waving (Carpenter and Ferguson 1977) and is followe d by intromission and coitus (Gillingham et al. 1983). In most boids, courtship is facilitated by the use of two cl aw-like spurs, wh ich are located on either side of the male’s cloaca. Male Madagascan tree boas ( Sanzinia madagascariensis ) use their spurs in combat bouts with other male conspecifics (Carpenter Murphy, and Mitchell

PAGE 25

25 1978). During these bouts, the males entwine thei r tails and vigorously flex the erected spurs against the scales of the opponent while hanging from branches. Larger male garter snakes ( Thamnophis sirtalis ) and European grass snakes ( Natrix natrix ) usually achieve more matings apparently because they can physic ally displace the tails of smalle r rival males by tail wrestling (Luiselli 1996; Shine et al. 2000). Tail wrestling in this context may be widespread in snake species that display “mating balls,” or mating aggregations where multiple males simultaneously attempt copulation (Shine et al. 2000: F9). Sexual Dimorphism an d Ontogenetic Shifts Although RTL is a stable character, sexual di morphism in snake tail length is common (Klauber 1943). Males typically have longer, more attenuate tails than females (King 1989; Shine 1993). However, the degree of these differen ces varies interspecifi cally (King 1989; Shine 1993). King (1989) proposed three hypotheses that could help e xplain the existence of sexual dimorphism in tail length among snakes: (1) mo rphological constraint (the hemipenes and retractor muscles of males are located in the tail); (2) female reproductive output (a more posterior cloaca might increase body cavity volume and allow increased fecundity); and (3) male mating ability (longer tails in males may be adva ntageous during courtship). These predictions were tested using 56 colubrid genera, and the re sults supported both the morphological constraint and female reproductive output hypotheses (Ki ng 1989). Additional studies involving tail wrestling in several natricine species provide support for the male mating ability hypothesis (Luiselli 1996; Shine et al. 2000). However, se lection could act on more than one of these hypotheses simultaneously. Whereas RTL is stable among individuals of the same species, sex, and age, RTL is nonetheless ontogenetically variable in most snake taxa (Klauber 1943). Therefore, the tail proportions (allometry) of many sp ecies change as they grow. The majority of snake species

PAGE 26

26 have relative tail-lengths that increase as they age (Klauber 1943); howev er, some species (e.g., Pituophis and Lampropeltis ) have relative tail-lengths that become shorter (Klauber 1943). Furthermore, some snake species demonstrate ont ogenetic shifts in macrohabitat use, with one age class (e.g., juveniles or a dults) found more frequently utili zing arboreal macrohabitats than another (Martins and Oliveira 1999). This behavioral shift has been well documented in several large boid species such as Python sebae (Spawls et al. 2002) and Boa constrictor (Campbell 1998), some colubrids such as Pseustes poecilonotus (Boos 2001), some viperids such as Bothrops jararaca (Sazima 1992; Martins et al. 2001) and some elapids such as Ophiophagus hannah (R. Whitaker pers. comm.). Usually, juvenile s are more arboreal than adults in those snake species known to exhibit behavioral shifts in habitat use. However, the opposite trend has been observed in Leptophis depressirostris (Nickerson, Sajdak, Henderson, and Ketcham 1978) and in some populations of Boa constrictor (Campbell 1998). Ontoge netic shifts involving arboreal macrohabitat use likely occur in many snak e clades, but more detailed natural history information is needed to know the ex tent to which this is the case.

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27 CHAPTER 2 MATERIALS AND METHODS Categorizing Climbing in Sna kes: Gravitational Habitat Snakes occupy a wide variety of habitat types, and it is often us eful to divide habitat usage into generalized categories when discussing snak e community assemblages. Perhaps the most widely utilized categories for snake habitat use incl ude fossorial, terrestrial, aquatic, semiaquatic, arboreal, and semiarboreal, or some subset of these (e.g., Johnson 1955; Shine 1983; Guyer and Donnelly 1990; Dalrymple, Bernardino, Steine r, and Nodell 1991; Lindell 1994; Conant and Collins 1998; Vidal, Kindl, Wong, and Hedges 2000). However, as correctly noted by Johnson (1955) these categories can become inadequate when performing interspecific ecomorphological st udies because patterns in habitat usage are often obscured. This is because th ese categories describe the habitat per se and not the behaviors snakes exhibit while ut ilizing these habitats. For exam ple, watersnakes in the genus Nerodia typically live and obtain food near wate r (Conant and Collins 1998). Consequently, they are usually categorized as aqua tic or semiaquatic (e.g., Vidal et al 2000). However, many Nerodia species spend a significant amount of time out of the water and off the ground climbing among branches to bask (Conant and Collins 1998) In fact, during parts of the year, some Nerodia species can spend as much or more time above the ground in shrubs and trees than Coluber constrictor a snake typically categorized as semiarboreal (Mushinsky, Hebrard, and Walley 1980; Plummer and Congdon 1994). Separating Nerodia and Coluber into distinct ecological categories (aquatic or semiaquatic versus semiarboreal) obscures these similarities in habitat usage. Similarly, categorizing both Nerodia and sea snakes such as the entirely pelagic Pelamis platurus as aquatic is equally misl eading. But in this case it is because the grouping suggests strong similarity in habitat usage when in actuality there is very little. Therefore, a

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28 different system was needed to categorize climbi ng in snakes that combines habitat use with behavior, which I call gravitational habitat. In order to categorize the am ount of climbing, and therefor e the amount of gravitational stress imposed, I divided gravitational habitat in to four categories: st enotopically arboreal, eurytopically arboreal/terrestrial, stenotopically terrestrial, and st enotopically aquatic. These categories were based on adult habitat use info rmation compiled from the literature and from personal observations. I define stenotopically arboreal species as those primarily living in arboreal habitats. These species are rarely found on the ground and should experience the greatest gravitational stress. Eu rytopically arboreal/t errestrial species are often found on the ground, but regularly climb for reasons in cluding hunting, escaping predators, and thermoregulation (e.g., Masticophis and Coluber ). The stenotopically terrestrial category includes both terrestrial and fossori al species that rarely if ever climb and, thus, these species should experience relatively little gravitational stress. Some characte ristically terres trial species, or populations of species, do climb occasionally (e.g., Bitis arietans, B. armata Bothrops asper Crotalus horridus, and Thamnophis sirtalis ). However, I consider th is behavior at ypical and do not consider them eurytopically arboreal/terrestri al. I define stenotopically aquatic species as those primarily found in water. They typi cally exhibit one or more morphological specializations accepted as adaptations to aquatic habitats such as a flattened or oarlike tail, more dorsally positioned eyes and nostrils, salt excretin g glands, and valvular nostrils (Heatwole, 1999). Laticaudines (genus Laticauda ), the homalopsine snake Enhydris enhydris and Helicops angulatus are included within this category because they are clearly adapted to an aquatic lifestyle and are usually found in water even though they occasionally sojourn onto land. Stenotopically aquatic species should experience the least gr avitational stress because the

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29 surrounding water column acts as an “antigravity suit” (Lillywh ite 1993: 561). Importantly, all categorical placements were based entirely on th e ecology and behavior of each species and not on morphology. Habitat use and other natural history informa tion for each species were collated from the literature. Whereas many of these sources were various journal publica tions, a large amount of this information came from the following: Af rica (Broadley and Cock 1975; Broadley 1983; Branch 1998; Schmidt and Noble 1998; Spawls et al. 2002); Australia (Shine 1995); Central America (Campbell 1998; Savage 2002); India (Wh itaker and Captain 2004); Madagascar (Glaw and Vences 1994; Henkel and Schmidt 2000); Nort h America (Smith and Brodie 1982; Conant and Collins 1998); South America (Murphy 1997; Boos 2001; Duellman 2005); and the West Indies (Schwartz and Henderson 1991). However, the natural history of some of the species included in this study is incompletely known, thereby making categorizati on difficult. For example, Alsophis antillensis and closely related Antillophis parvifrons are West Indian racers with relatively long tails (27.8% and 31.1% respectively). However, whether these species climb or not is unknown. Several other species of Alsophis are known to climb well, but because th is information is not available for A antillensis and A parvifrons I chose not to assume they clim bed and thus categorized them as stenotopically terrestrial ev en though they may actually climb to some extent. In order to avoid the confounding effects of se xual dimorphism and ontogenetic shifts in allometry and habitat use, only adult females were used for all aspects of this study except for the frequency distribution (see below). A snake was determined to be an adult if its total length was within, or very near, the publishe d adult range for that species. Tail length data were collected from the literature, museum specimens, and live sn akes (see Table 2-1 for species used in this

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30 study and sources). Museum specimens were ac quired from the Florida Museum of Natural History (FLMNH) and National Museum of Na tural History, Smithsonian (USNM). For museum specimens, sex was determined by subca udal incision. Species in which females are known to possess well-developed he mipenis-like structures (e.g., Pseudoficimia and Gyalopium ; Hardy 1972; Smith and Brodie 1982) were not used in this study. Some Bothrops insularis females have hemipenis-like structures (Hoge, Be lluomini, Schreiber, and Penha 1961), and this species was included in the study under the a ssumption that length da ta reported by Klauber (1943) were from functional females. Klauber (1 943) states that differe ntial shrinkage between the tail and body can occur as a resu lt of preservation, but that the am ount is insignif icant as long as the preservation methods are consistent. Live snakes were measured at Glades Herp Inc., Florida, and were probed to determine sex. Snout-vent length (SVL) was measured from the tip of the snout to th e posterior edge of the anal scale. Tail length was measured from the posterior edge of the an al scale to the tip of the tail, and only snakes with comp lete tails were used in this study. Snout-vent length and tail length were measured using either a meter stic k ( 1.0 mm) or a caliper ( 0.1 mm). A string was used to follow the body contours of live snak es and rigid specimens and was subsequently measured with a meter stick. Length measurements were repeated on individuals up to ten times when possible and then averaged. I used mean SVL and tail lengths when those data were available from the literature or from multiple sp ecimens, but otherwise I used data representing single specimens. To verify the stability of relative tail-length (RTL), I gathered large samples of data from the literature on the total variation in adult female RTL for 26 species (5 families, 9 subfamilies, 26 genera) and found the variation to be small (m ean variation SE, 1% 0.3%,

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31 95% C.I.) (Table 2-2). Furthermore, the RTL of museum specimens was checked to ensure the data were consistent with published adult lengths for that species. Analyses Frequency Distribution Frequency distributions for total lengths were produced for each of the four gravitational habitat categories: stenotopically arboreal (n=75), eurytopically arboreal/terrestrial (n=74), stenotopically terrestrial (n=94) and stenotopically aquatic (n= 35)(Table A-1). All categories comprise both Old and New World species. Data used for the frequency distributions were based on raw total lengths. Stenotopically ar boreal species should sp end the most time in arboreal habitats and, thus, were analyzed sepa rately from eurytopically arboreal/terrestrial species. Furthermore, eurytopi cally arboreal/terrestrial species might not climb enough to have their lengths affected by such limitations. Because there is much interspecific variati on among snakes regarding which sex is longer (Shine 1993), I used length measurements repres enting the longest adult total lengths regardless of sex. Maximum total lengths for each specie s were used when possible unless stated by the authors that the maximum size is extremely unusual. In that case, the ne xt largest measurement was used. Maximum adult total length was used because whereas juvenile length is likely also under related selective pressures, they differ from adults in two important ways: (1) juveniles may utilize a different macrohabitat until adult size is reached or until a certain length is attained; and (2) juveniles are shorter and are thus less affected by gravita tional stresses on blood circulation. Gravitational Habitat and Total Body Length To test for differences in mean total length amon g the four categories, I used an analysis of variance (ANOVA). To meet the assumpti on of normality, the length data were log10

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32 transformed. Following the ANOVA, I compared the means among the four categories, using Tukey’s Honestly Significant Difference test. Coe fficients of variation (C V) were calculated and used to compare variance among the four categories. Gravitational Habitat and Tail Length I tested for differences in tail length among 227 species within 138 genera, 12 families, and 18 subfamilies using an analysis of covarian ce (ANCOVA) with total length treated as a covariate. Differences in tail length corrected for body length (correct ed tail-length) were inferred from comparisons of the Y-intercepts in the ANCOVA. Data in this and all subsequent analyses were collected independently from thos e used in the frequenc y distribution described above. Data were log10 transformed to meet the assumptions of linearity and normality before performing the ANCOVA analyses. All statistical analyses were performed using the statistical program package R (R Development Core Team 2006). Constructing the Phylogeny Phylogenies constructed by Lawson, Slowinsk i, Crother, and Burbrink (2005), Lawson, Slowinski, and Burbrink (2004), a nd Vidal et al. (2000) were co mbined and used as a base phylogeny, to which additional phylogenies from deta iled studies were added (Fig. 2-1). When possible, the trees chosen were those recommended by the authors. However, these were often consensus trees and, as such, occasionally contained polytomies. Polytomy resolution was often achieved by either using another tree in the same study with high bootstrap support, or by using a tree from a different study. Fo r different trees containing sim ilar species but with conflicting relationships, I used the tree with the strongest bootst rap support and, when available, the more extensive taxon sampling. However, phylogeneti c relationships for some clades remain unresolved (e.g., Dipsas Langaha Leptodeira Prosymna and Psammophis ), and uncertain relationships were in the end reflected as polytomies. Many of these polytomies contain

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33 congeners categorized within the same gravitational habitat and, thus are not likely to affect the results (Fig 2-1). In some cases, I used species thought closely related to those in the composite tree to replace species for which no specime ns or length data were available (e.g., Hydrops triangularis was replaced by H. caesurus ; Xenoxybelis argenteus was replaced by X. boulengeri ). The following studies were used to add taxa to the base phylogeny (Fig. 2-1). The scolecophidian clade was replaced using the phylogeny of Vidal and Hedges (2002). The phylogeny of Lenk, Kalyabina, Wink, and Joger (2001) was used for placement of true vipers ( Atheris Bitis Causus Macrovipera and Vipera ). The phylogeny of Malhotra and Thorpe (2004) was used for Asian pitviper ( Protobothrops Trimeresurus and Tropidolaemus ) placement. The phylogeny of Parkinson, Campbell, and Chippindale (2002, in Gutberlet and Harvey 2004) was used for New World pitviper relationships, and the phylogeny of Murphy, Fu, Lathrop, Feltham, and Kovac (2002) wa s used for relationships within Crotalus The phylogeny of Nagy, Joger, Wink, Glaw, and Vences (2003) was used for pseudoxyrhophine relationships. The phylogeny of Kelly, Barker, and Villet (2003) was used for Dasypeltis and Dendrelaphis placement and psammophine relationships. Th e phylogeny of Alfaro and Arnold (2001) was used for thamnophiine relationships, and the phylogeny of De Quieroz, Lawson, and LemosEspinal (2002) was used for relationships within Thamnophis The phylogeny of Lawson et al. (2005) was used for placement of the Homalopsinae, as we ll as the following genera: Ahaetulla, Carphophis, Crotaphopeltis, Dinodon, Dips adoboa, Drymarchon, Gonyosoma, Grayia, Lycophidion, Masticophis, Mehelya, Natricet eres, Opheodrys, Prosymna, Philothamnus, Phyllorynchus, Pseudaspis, Sonora, Spilote s, Telescopus, Thelotornis, Thrasops, and Xenochrophis The phylogeny of Utiger et al. (20 02) was used for the placement of Arizona

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34 Elaphe Lampropeltis Pantherophis and Pituophis The phylogeny of Vidal et al (2000) was used for the placement of the following genera: Helicops Hydrodynastes Hydrops Ialtris Oxyrhopus and Pseudoeryx The phylogeny of Kraus and Brown (1998) was used for the placement of the genera Chilomeniscus and Dispholidus Subfamily designation followed that of the EMBL Reptile Database (Uetz 2006). Howe ver, recent phylogenetic analyses by Lawson et al (2005) provide further evidence that considerab le changes in these cl assifications need to occur in order to recognize monophyletic groups. Relative Tail-Length, Gravitati onal Habitat, and Phylogeny Interspecific comparative analyses can be confounded by pseudoreplication caused by phylogenetic relatedness among species samples (Price 1997). However, these effects can be partially resolved using indepe ndent contrasts (Felsenstein 1 985). I assessed the relationship between RTL and macrohabitat use using the pr ogram Mesquite version 1.06 (Maddison and Maddison 2005). Because there is no comprehensive phylogeny available for snakes, I constructed an estimate of relatedness by co mbining data from 14 recent molecular snake phylogenies into a composite tree containing 227 species within 138 genera, 12 families, and 18 subfamilies (Fig. 2-1). These data represent almost all snake families and subfamilies (see below). Branch lengths could not be estim ated because the final phylogenetic tree was composed from a variety of sources. Independent contrasts were therefore generated with branch lengths assigned as either equal or with the assumption that the ag e of a clade is proportional to the number of species it contains using Graf en’s branch lengths (Grafen 1989; Maddison and Maddison 2005). However, plots of standardized contrasts against the variance of untransformed contrasts showed strong significan t correlations for Grafen’s, but not equal, branch lengths. As significant correlations violate a key assumption of independent c ontrast analysis (Diaz-Uriarte and Garland 1996), I used equal branch lengths on ly in the final analysis. Contrasts were

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35 calculated between nodes for macrohabitat use and log10 transformed RTL, and relationships were examined between the variables by calculati ng regressions on these standardized contrasts using least-squared change op timization parsimony (Garland, Harvey, and Ives 1992; Grafen 1992).

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36 Table 2-1. Taxa and associated data sources for 227 snake species included in this study. Family Subfamily Species Source Acrochordidae Acrochordus granulatus Lillywhite unpubl. data Aniliidae Anilius scytale UF62496 Atractaspididae Aparallactinae Amblyodipsas polylepis Broadley and Cock 1975 Amblyodipsas ventrimaculatus Broadley and Cock 1975 Aparallactus capensis Broadley and Cock 1975 Aparallactus guentheri Broadley and Cock 1975 Aparallactus lunulatus Broadley and Cock 1975 Xenocalamus mechowii Broadley and Cock 1975 Xenocalamus sabiensis Broadley and Cock 1975 Atractaspis bibronii Broadley and Cock 1975 Boidae Boinae Acrantophis dumerili Sheehy unpubl. data Boa constrictor Lillywhite unpubl. data Corallus caninus Lillywhite unpubl. data Corallus ruschenbergerii Sheehy unpubl. data Epicrates striatus Sheehy unpubl. data Erycinae Charina trivirgata Klauber 1943 Gongylophis muelleri Sheehy unpubl. data Pythoninae Morelia viridis Sheehy unpubl. data Python molurus Lillywhite unpubl. data Python regius Lillywhite unpubl. data Python sebae Broadley and Cock 1975 Colubridae Boodontinae Duberria lutrix Broadley and Cock 1975 Grayia smythii UF57047 Lamprophis fuliginosus Broadley and Cock 1975 Lamprophis lineatus Klauber 1943 Lycodonomorphus leleupi Broadley and Cock 1975 Lycophidion capense Broadley and Cock 1975 Lycophidion variegatum Broadley and Cock 1975 Mehelya capensis Schmidt and Noble 1998 Pseudaspis cana Broadley and Cock 1975

PAGE 37

37 Table 2-1. Continued Family Subfamily Species Source Colubridae Colubrinae Ahaetulla nasuta Lillywhite unpubl. data Arizona elegans UF48734 Boiga blandingii UF69248 Boiga cynodon UF55217 Chilomeniscus stramineus Klauber 1943 Chironius carinatus UF34779 Coluber constrictor UF47855 Crotaphopeltis hotamboeia Broadley and Cock 1975 Dasypeltis fasciata UF61301 Dasypeltis medici Broadley and Cock 1975 Dasypeltis scabra Broadley and Cock 1975 Dendrelaphis pictus Lillywhite unpubl. data Dinodon rufozonatum Lillywhite unpubl. data Dinodon semicarinatum UF24089 Dipsadoboa aulica Broadley and Cock 1975 Dipsadoboa unicolor Schmidt and Noble 1998 Dispholidus typus Broadley and Cock 1975 Drymarchon corais UF83818 Elaphe radiata Lillywhite unpubl. data Gastropyxis smaragdina UF52721 Gonyosoma oxycephalum Lillywhite unpubl. data Lampropeltis getula Klauber 1943 Lampropeltis triangulum UF68828 Lycodon laoensis UF69087 Masticophis flagellum Klauber 1943 Masticophis lateralis Klauber 1943 Mastigodryas boddaerti UF91619 Opheodrys aestivus UF43623 Oxybelis fulgidus UF56396 Oxybelis aeneus Sheehy unpubl. data Pantherophis obsoleta Lillywhite unpubl. data Philothamnus dorsalis Schmidt and Noble 1998 Philothamnus irregularis Broadley and Cock 1975 Philothamnus semivariegatus Broadley and Cock 1975 Philothamnus angolensis UF53342 Philothamnus hoplogaster Broadley and Cock 1975 Phyllorynchus browni Klauber 1943 Phyllorynchus decurtatus Klauber 1943

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38 Table 2-1. Continued Family Subfamily Species Source Colubridae Colubrinae Pituophis catenifer Klauber 1943 Pituophis melanoleucus UF132981 Prosymna ambigua Schmidt and Noble 1998 Prosymna bivittata Broadley and Cock 1975 Prosymna stuhlmannii Broadley and Cock 1975 Prosymna sundevallii Broadley and Cock 1975 Sonora michoacanensis Klauber 1943 Spilotes pullatus UF56320 Stilosoma extenuatum Lillywhite unpubl. data Tantilla planiceps Klauber 1943 Tantilla relicta Lillywhite unpubl. data Tantilla hendersoni Stafford 2004 Tantilla melanocephala UF40711 Telescopus semiannulatus Broadley and Cock 1975 Thelotornis capensis Broadley and Cock 1975 Thrasops jacksonii UF52696 Dipsadinae Adelphicos quadrivirgatus Klauber 1943 Atractus depressiocellus Myers 2003 Atractus hostilitractus Myers 2003 Atractus clarki Myers 2003 Atractus darienensis Myers 2003 Carphophis amoenus UF43842 Diadophis punctatus UF56868 Dipsas catesbyi Harvey unpubl. data Dipsas indica Harvey unpubl. data Dipsas peruana Harvey unpubl. data Dipsas boettgeri Harvey unpubl. data Dipsas chaparensis Harvey unpubl. data Hypsiglena torquata Klauber 1943 Imantodes cenchoa UF11284 Leptodeira annulata Duellman 1958 Leptodeira septentrionalis Duellman 1958 Leptodeira frenata Duellman 1958 Leptodeira bakeri Duellman 1958 Leptodeira nigrofasciata Duellman 1958 Leptodeira maculata Duellman 1958 Leptodeira punctata Duellman 1958

PAGE 39

39 Table 2-1. Continued Family Subfamily Species Source Colubridae Colubrinae Taeniophallus brevirostris Schargel et al. 2005 Thamnodynastes pallidus UF88631 Tretanorhinus variabilis UF119241 Homalopsinae Cerberus rynchops Lillywhite unpubl. data Enhydris enhydris Lillywhite unpubl. data Natricinae Natriciteres olivacea Broadley and Cock 1975 Natrix natrix UF116535 Nerodia fasciata UF41022 Nerodia floridana UF109763 Nerodia sipedon UF63519 Nerodia taxispilota Lillywhite unpubl. data Nerodia rhombifer Lillywhite unpubl. data Regina alleni UF99998 Regina septemvittata UF68506 Seminatrix pygaea UF42444 Storeria dekayi Lillywhite unpubl. data Thamnophis hammondii Klauber 1943 Thamnophis ordinoides UF57050 Thamnophis sirtalis UF42468 Virginia valeriae UF54122 X enochrophis flavipunctatus Lillywhite unpubl. data Psammophiinae Psammophis biseriatus UF52841 Psammophis punctulatus UF52832 Psammophis sibilans Broadley and Cock 1975 Psammophis angolensis Broadley and Cock 1975 Psammophis brevirostris Broadley and Cock 1975 Psammophis crucifer Broadley and Cock 1975 Psammophis jallae Broadley and Cock 1975 Psammophylax tritaeniatus Broadley and Cock 1975 Rhamphiophis oxyrhynchus UF59339 Rhamphiophis rostratus Broadley and Cock 1975 Pseudoxyrhophiinae Dromicodryas bernieri USNM 499459 Ithycyphus miniatus USNM 149366

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40 Table 2-1. Continued Family Subfamily Species Source Colubridae Pseudoxyrhophiinae Langaha pseudoalluaudi Domergue 1988 Langaha madagascariensis Guib 1949 Langaha alluaudi Guib 1949 Leioheterodon madagascariensis Sheehy unpubl. data Leioheterodon modestus Sheehy unpubl. data Micropisthodon ochraceus Domergue 1993 Xenodontinae Alsophis antillensis UF53983 Alsophis cantherigerus UF44287 Alsophis portoricensis UF57211 Alsophis vudii UF99946 Antillophis parvifrons UF23131 Arrhyton funereum UF113638 Clelia clelia UF34331 Darlingtonia haetiana UF55681 Drepanoides anomalus Dixon and Soini 1986 Erythrolamprus aesculapii UF115664 Farancia abacura Lillywhite unpubl. data Helicops angulatus UF88643 Heterodon nasicus UF56015 Heterodon platirhinos UF44172 Hydrodynastes gigas UF20576 Hydrops caesurus Scrocchi et al. 2005 Hypsirhynchus ferox UF60786 Ialtris dorsalis UF85307 Liophis typhlus UF32985 Oxyrhopus petola UF62069 Philodryas baroni Lillywhite unpubl. data Pseudoboa coronata UF117617 Pseudoeryx plicatilis UF86909 Siphlophis cervinus UF95759 Uromacer catesbyi UF23158 Uromacer frenatus UF23170 Uromacer oxyrhynchus UF23172 X enodon severus UF87926 X enoxybelis boulengeri Duellman 2005 Cylindrophiidae Cylindrophis ruffus UF51669

PAGE 41

41 Table 2-1. Continued Family Subfamily Species Source Elapidae Elapinae Aspidelaps scutatus Broadley and Cock 1975 Bungarus fasciatus Sheehy unpubl. data Dendroaspis angusticeps Broadley and Cock 1975 Dendroaspis polylepis Broadley and Cock 1975 Dendroaspis viridis Sheehy unpubl. data Elapsoidea guentheri Broadley and Cock 1975 Elapsoidea semiannulata Broadley and Cock 1975 Elapsoidea sundevallii Broadley and Cock 1975 Elapsoidea nigra Klauber 1943 Hemachatus haemachatus Broadley and Cock 1975 Micrurus fulvius Lillywhite unpubl. data Naja melanoleuca Broadley and Cock 1975 Naja haje Broadley and Cock 1975 Naja mossambica Broadley and Cock 1975 Naja atra Lillywhite unpubl. data Naja kaouthia Lillywhite unpubl. data Hydrophiinae Aipysurus laevis Lillywhite unpubl. data Emydocephalus annulatus Lillywhite unpubl. data Hydrophis melanocephalus Guinea 1981 Laticauda colubrina Guinea 1981 Notechis scutatus Lillywhite unpubl. data Pelamis platurus Lillywhite unpubl. data Leptotyphlophidae Leptotyphlops alfredschmidti Lehr et al. 2002 Leptotyphlops macrops Broadley and Wallach 1996 Tropidophiidae Tropidophiinae Tropidophis morenoi Hedges et al 2001 Tropidophis haetianus Sheehy unpubl. data Typhlopidae Rhinotyphlops episcopus Franzen and Wallach 2002 X enotyphlops grandidieri Wallach and Ineich 1996 Viperidae Crotalinae Agkistrodon contortrix Lillywhite unpubl. data Agkistrodon mokeson Klauber 1943 Agkistrodon piscivorus Lillywhite unpubl. data Atropoides nummifer Sheehy unpubl. data Bothriechis schlegelii Lillywhite unpubl. data

PAGE 42

42 Table 2-1. Continued Family Subfamily Species Source Viperidae Crotalinae Bothrops insularis Klauber 1943 Crotalus atrox Lillywhite unpubl. data Crotalus horridus Lillywhite unpubl. data Crotalus ruber Lillywhite unpubl. data Crotalus cerastes Boundy and Balgooyen 1988 Protobothrops cornutus Herrmann et al. 2004 Trimeresurus gramineus Klauber 1943 Trimeresurus purpureomaculatus Lillywhite unpubl. data Trimeresurus albolabris Lillywhite unpubl. data Tropidolaemus wagleri Sheehy unpubl. data Viperinae Atheris squamigera Klauber 1943 Atheris chlorechis Sheehy unpubl. data Bitis arietans Broadley and Cock 1975 Bitis atrops Broadley and Cock 1975 Bitis caudalis Broadley and Cock 1975 Bitis gabonica Lillywhite unpubl. data Bitis nasicornis Lillywhite unpubl. data Causus defilippii Broadley and Cock 1975 Causus rhombeatus Broadley and Cock 1975 Macrovipera mauritanica Sheehy unpubl. data Vipera xanthina Lillywhite unpubl. data Xenopeltidae X enopeltis unicolo r UF50268

PAGE 43

43 Table 2-2. Total variation in rela tive tail-lengths (RTL) of 26 sp ecies of snakes. The data are from adult females only and include speci es typically categorized as fossorial, terrestrial, semiarboreal and arboreal. In species where the RTL becomes shorter with age, the absolute value of the di fference is shown. For sources (Sc), 1 = Broadley 1959; 2 = Klauber 1943. Family Subfamily Species n min RTL max RTL max min Sc Atractaspididae Aparallactinae A mblyodipsas unicolor 130.060.071%1 Boidae Erycinae Charina trivirgata 570.110.121%2 Colubridae Boodontinae D uberria lutrix 100.100.122%1 L amprophis fuliginosus 340.110.132%1 P seudaspis cana 50.130.163%1 Colubrinae A rizona elegans 250.100.111%2 Conopsis nasus 700.110.101%2 L ampropeltis getula 2490.110.101%2 Masticophis flagellum 340.190.201%2 P hyllorhynchus decurtatus 1480.090.081%2 R hinocheilus lecontei 230.120.120%2 Salvadora grahamiae 230.180.180%2 Sonora michoacanensis 570.150.150%2 Tantilla planiceps 150.150.183%2 Dipsadinae A delphicos quadrivirgatus 620.110.110%2 D iadophis punctatus 1830.130.141%2 Geophis nasalis 870.110.132%2 H ypsiglena torquata 300.120.120%2 N atricinae Thamnophis hammondii 1530.190.181%2 Elapidae Elapinae E lapsoidea nigra 290.050.061%2 Micrurus nigrocinctus 320.090.090%2 Viperidae Crotalinae B othrops insularis 940.110.110%2 Trimeresurus gramineus 160.140.140%2 Viperinae A theris squamigera 250.120.131%2 B itis arietans 270.060.093%1 Causus defilippii 220.050.072%1

PAGE 44

44 Figure 2-1. Composite phylogeny of 227 snake taxa. Species in gray boxes are stenotopically arboreal. Bolded branches denote eurytopical ly arboreal/terrestrial species. Asterisks denote stenotopically aquatic species. Bo lded species exhibit pseudautotomy. Remaining unmarked species ar e stenotopically terrestrial.

PAGE 45

45 Figure 2-1. Continued

PAGE 46

46 Figure 2-1. Continued

PAGE 47

47 Figure 2-1. Continued

PAGE 48

48 CHAPTER 3 RESULTS Frequency Distribution The frequency distribution of raw total lengths for all four gravitational habitat categories is shown in Fig. 3-1. The fr equency distribution mean (mm) median, mode and standard deviation within each of th e four categories were: ste notopically arboreal (1212, 1050, 1050, 506.5), eurytopically arboreal /terrestrial (1644, 1215, 1680, 1103.6), stenotopically terrestrial (855.5, 652, 480, 557.7), and stenotopically aqua tic (1105, 1000, 1000, 530). Stenotopically arboreal total lengths had the lowest sta ndard deviation, whereas the eurytopically arboreal/terrestrial total lengths ha d the greatest by more than twof old. Slightly more than half (52%) of the stenotopically arboreal species were between 60-120 cm total length, and 93.5% were between 50-200 cm total length There were no stenotopically arboreal specie s shorter than 50 cm, four species between 201-240 cm (in order of increasing length: Dendroaspis viridis Boiga cynodon Boiga forsteini and Gonyosoma oxycephalum ), and only one species ( Leptophis ahaetulla ) longer than 240 cm. Seve ral non-stenotopically arboreal snake species have total lengths < 50 cm and > 200 cm (Fig. 3-1). Gravitational Habitat and Total Body Length There were differences in total length between the four gravitational habitat categories (P < 0.0001, df = 3, F = 19.433, Fig. 3-1). Mean total le ngth was longer in st enotopically arboreal species than in stenotopically terrestrial species (P < 0.00001), longer in eurytopically arboreal/terrestrial species than in stenotopically terrestrial species (P < 0.00001), and longer in stenotopically aquatic species th an in stenotopically terrestrial species (P = 0.0015). However, I did not find a difference in mean total length be tween stenotopically arbo real and eurytopically arboreal/terrestrial species (P = 0.32), between stenotopically arbor eal and stenotopically aquatic

PAGE 49

49 species (P = 0.96), or between eu rytopically arboreal/t errestrial and stenotopically aquatic species (P = 0.24). There were differences in the variance am ong the four categories (Fig. 3-2). The coefficients of variation in order of increasing value are: stenotopically arboreal (CV = 0.011), stenotopically aquatic (CV = 0.014), eurytopi cally arboreal/terrestrial (CV = 0.023), and stenotopically terrestrial (CV = 0.027). Gravitational Habitat and Tail Length Perhaps not surprisingly, tail length is correlated with total length (r2 = 0.65, P < 0.0001, n = 277, Fig. 3-3). Because I did not find a di fference in corrected tail-length between stenotopically arboreal and eurytopically arbor eal/terrestrial species (P = 0.30, Fig. 3-4), I combined these categories into a single scansori al category (Fig. 3-5). The scansorial species had corrected tail-lengths on average 1.9 times l onger than the two non-scan sorial categories (P < 0.0001) (Fig. 3-5). However, I did not find a difference in corrected tail-length between stenotopically terrestrial and sten otopically aquatic speci es (P = 0.94, Fig. 3-4). The relative taillengths for the 227 species included in this study ranged from 1.1% ( Rhinotyphlops episcopus ) to 48.1% ( Uromacer frenatus ) (Table 3-1). Relative Tail-Length, Gravitati onal Habitat, and Phylogeny The analysis of covariance (ANCOVA) result s supported using the s cansorial gravitational habitat category for the independent contrast anal yses instead of the separate analyses of the stenotopically and eurytopically arboreal/terrestri al categories. After accounting for the effect of phylogeny on the data, I found a significant rela tionship between independent contrasts of scansorial macrohabitat use and log10 transformed RTL (r2 = 0.31, P < 0.0001, Fig. 3-6).

PAGE 50

50 Table 3-1. The 227 species include d in this study in order of decreasing relative tail-length (RTL), and with associated categorizations used in analyses. Range (Rg), OW = Old World, NW = New World. Gravitational habi tat (Gh), 3 = stenotopically arboreal; 2 = eurytopically arboreal/terrestrial; 1 = st enotopically terrestrial; 0 = stenotopically aquatic. Family Subamily Species RTL Rg Gh Colubridae Xenodontinae Uromacer frenatus 0.481NW3 Colubridae Xenodontinae Uromacer catesbyi 0.455NW3 Colubridae Xenodontinae Uromacer oxyrhynchus 0.440NW3 Colubridae Pseudoxyrhophiinae Langaha alluaudi 0.427OW3 Colubridae Pseudoxyrhophiinae Micropisthodon ochraceus 0.414OW3 Colubridae Pseudoxyrhophiinae Langaha pseudoalluaudi 0.403OW3 Colubridae Xenodontinae X enoxybelis boulengeri 0.386NW3 Colubridae Colubrinae Gastropyxis smaragdina 0.383OW3 Colubridae Colubrinae Oxybelis aeneus 0.380NW3 Colubridae Psammophiinae Psammophis punctulatus 0.377OW2 Colubridae Xenodontinae Antillophis parvifrons 0.371NW2 Colubridae Pseudoxyrhophiinae Langaha madagascariensis 0.368OW3 Colubridae Colubrinae Thelotornis capensis 0.365OW3 Colubridae Colubrinae Opheodrys aestivus 0.355NW3 Colubridae Psammophiinae Psammophis biseriatus 0.352OW2 Colubridae Xenodontinae Helicops angulatus 0.340NW0 Colubridae Colubrinae Ahaetulla nasuta 0.333OW3 Colubridae Psammophiinae Psammophis brevirostris 0.333OW2 Colubridae Colubrinae Philothamnus semivariegatus 0.325OW3 Colubridae Colubrinae Chironius carinatus 0.320NW2 Colubridae Colubrinae Oxybelis fulgidus 0.319NW3 Colubridae Pseudoxyrhophiinae Ithycyphus miniatus 0.315OW3 Colubridae Xenodontinae Alsophis vudii 0.312NW2 Colubridae Xenodontinae Arrhyton funereum 0.311NW1 Colubridae Psammophiinae Rhamphiophis oxyrhynchus 0.308OW2 Colubridae Dipsadinae Imantodes cenchoa 0.306NW3 Colubridae Colubrinae Philothamnus angolensis 0.302OW3 Colubridae Xenodontinae Alsophis portoricensis 0.302NW2 Colubridae Colubrinae Dendrelaphis pictus 0.300OW3 Colubridae Colubrinae Masticophis lateralis 0.298NW2 Colubridae Psammophiinae Psammophis angolensis 0.296OW1 Colubridae Colubrinae Philothamnus irregularis 0.295OW3 Colubridae Xenodontinae Alsophis cantherigerus 0.293NW2 Colubridae Psammophiinae Psammophis jallae 0.293OW2 Colubridae Xenodontinae Ialtris dorsalis 0.293NW1 Colubridae Colubrinae Mastigodryas boddaerti 0.288NW2 Colubridae Psammophiinae Rhamphiophis rostratus 0.285OW2 Colubridae Dipsadinae Leptodeira punctata 0.283NW2 Colubridae Psammophiinae Psammophis sibilans 0.280OW2 Colubridae Colubrinae Spilotes pullatus 0.279NW2

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51 Table 3-1. Continued Family Subamily Species RTLRgGh Colubridae Natricinae X enochrophis flavipunctatus 0.278OW1 Colubridae Xenodontinae Alsophis antillensis 0.278NW1 Colubridae Boodontinae Grayia smythii 0.277OW1 Colubridae Colubrinae Dispholidus typus 0.276OW3 Colubridae Colubrinae Thrasops jacksonii 0.275OW3 Colubridae Colubrinae Philothamnus hoplogaster 0.275OW2 Colubridae Colubrinae Masticophis flagellum 0.269NW2 Elapidae Elapinae Dendroaspis viridis 0.260OW3 Elapidae Elapinae Dendroaspis angusticeps 0.258OW3 Colubridae Dipsadinae Dipsas catesbyi 0.257NW3 Colubridae Xenodontinae Philodryas baroni 0.256NW3 Colubridae Dipsadinae Dipsas boettgeri 0.256NW3 Colubridae Colubrinae Philothamnus dorsalis 0.255OW3 Colubridae Colubrinae Coluber constrictor 0.250NW2 Colubridae Dipsadinae Dipsas chaparensis 0.248NW3 Colubridae Natricinae Nerodia taxispilota 0.247NW2 Colubridae Dipsadinae Dipsas bucephala 0.245NW3 Colubridae Natricinae Nerodia fasciata 0.240NW2 Colubridae Pseudoxyrhophiinae Dromicodryas bernieri 0.240OW1 Colubridae Natricinae Regina septemvittata 0.239NW2 Colubridae Colubrinae Tantilla hendersoni 0.239NW1 Colubridae Colubrinae Gonyosoma oxycephalum 0.236OW3 Colubridae Natricinae Thamnophis ordinoides 0.234NW1 Colubridae Xenodontinae Drepanoides anomalus 0.232NW1 Colubridae Xenodontinae Hypsirhynchus ferox 0.230NW1 Colubridae Colubrinae Boiga cynodon 0.229OW3 Colubridae Dipsadinae Dipsas peruana 0.229NW3 Colubridae Natricinae Thamnophis hammondii 0.229NW1 Colubridae Dipsadinae Thamnodynastes pallidus 0.228NW2 Colubridae Natricinae Natriciteres olivacea 0.228OW1 Colubridae Natricinae Nerodia rhombifer 0.227NW2 Colubridae Colubrinae Dipsadoboa aulica 0.226OW3 Colubridae Dipsadinae Leptodeira bakeri 0.226NW2 Colubridae Colubrinae Boiga blandingii 0.226OW3 Colubridae Natricinae Nerodia floridana 0.224NW2 Colubridae Dipsadinae Leptodeira annulata 0.224NW3 Colubridae Natricinae Nerodia sipedon 0.224NW2 Colubridae Xenodontinae Darlingtonia haetiana 0.223NW1 Colubridae Xenodontinae Siphlophis cervinus 0.222NW3 Colubridae Psammophiinae Psammophis crucifer 0.220OW1 Colubridae Dipsadinae Leptodeira septentrionalis 0.219NW2 Colubridae Natricinae Thamnophis sirtalis 0.217NW1 Colubridae Natricinae Regina alleni 0.215NW1 Colubridae Colubrinae Dinodon semicarinatum 0.214OW2 Colubridae Colubrinae Tantilla planiceps 0.211NW1

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52 Table 3-1. Continued Family Subamily Species RTL Rg Gh Colubridae Dipsadinae Leptodeira frenata 0.209 NW 2 Colubridae Natricinae Storeria dekayi 0.208 NW 1 Boidae Boinae Corallus ruschenbergerii 0.207 NW 3 Colubridae Psammophiinae Psammophylax tritaeniatus 0.207 OW 1 Colubridae Colubrinae Tantilla relicta 0.203 NW 1 Colubridae Dipsadinae Diadophis punctatus 0.203 NW 1 Colubridae Dipsadinae Leptodeira maculata 0.202 NW 2 Elapidae Elapinae Dendroaspis polylepis 0.199 OW 2 Colubridae Xenodontinae Pseudoboa coronata 0.199 NW 1 Colubridae Xenodontinae Clelia clelia 0.198 NW 1 Atractaspididae Aparallactinae Aparallactus lunulatus 0.198 OW 1 Colubridae Dipsadinae Tretanorhinus variabilis 0.197 NW 1 Colubridae Pseudoxyrhophiinae Leioheterodon modestus 0.196 OW 1 Colubridae Colubrinae Tantilla melanocephala 0.195 NW 1 Colubridae Dipsadinae Taeniophallus brevirostris 0.194 NW 1 Colubridae Xenodontinae Oxyrhopus petola 0.193 NW 2 Colubridae Homalopsinae Enhydris enhydris 0.192 OW 0 Atractaspididae Aparallactinae Aparallactus capensis 0.190 OW 1 Colubridae Boodontinae Lycodonomorphus leleupi 0.189 OW 1 Atractaspididae Aparallactinae Aparallactus guentheri 0.188 OW 1 Colubridae Natricinae Natrix natrix 0.188 OW 2 Colubridae Colubrinae Dipsadoboa unicolor 0.187 OW 2 Colubridae Dipsadinae Leptodeira nigrofasciata 0.186 NW 2 Colubridae Colubrinae Elaphe radiata 0.184 OW 2 Viperidae Crotalinae Protobothrops cornutus 0.184 OW 3 Colubridae Colubrinae Sonora michoacanensis 0.181 NW 1 Colubridae Colubrinae Dinodon rufozonatum 0.179 OW 2 Colubridae Xenodontinae Liophis typhlus 0.177 NW 2 Colubridae Colubrinae Dasypeltis medici 0.174 OW 2 Elapidae Hydrophiinae Notechis scutatus 0.173 OW 2 Elapidae Elapinae Naja melanoleuca 0.168 OW 2 Colubridae Homalopsinae Cerberus rynchops 0.165 OW 1 Colubridae Colubrinae Lycodon laoensis 0.164 OW 2 Viperidae Crotalinae Trimeresurus gramineus 0.162 OW 3 Colubridae Pseudoxyrhophiinae Leioheterodon madagascariensis 0.160 OW 1 Colubridae Natricinae Seminatrix pygaea 0.160 NW 1 Colubridae Xenodontinae Heterodon platirhinos 0.160 NW 1 Elapidae Elapinae Naja mossambica 0.159 OW 2 Elapidae Elapinae Naja haje 0.158 OW 2 Viperidae Crotalinae Agkistrodon piscivorus 0.157 NW 1 Colubridae Xenodontinae Hydrops caesurus 0.157 NW 1 Viperidae Crotalinae Bothriechis schlegelii 0.157 NW 3 Boidae Boinae Corallus caninus 0.156 NW 3 Viperidae Crotalinae Trimeresurus purpureomaculatus 0.156 OW 3 Colubridae Colubrinae Lampropeltis triangulum 0.151 NW 1

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53 Table 3-1. Continued Family Subamily Species RTLRgGh Viperidae Viperinae Atheris chlorechis 0.151OW3 Viperidae Crotalinae Trimeresurus albolabris 0.151OW3 Colubridae Colubrinae Pantherophis obsoleta 0.150NW2 Colubridae Boodontinae Pseudaspis cana 0.150OW1 Viperidae Crotalinae Tropidolaemus wagleri 0.149OW3 Colubridae Colubrinae Telescopus semiannulatus 0.148OW2 Elapidae Elapinae Hemachatus haemachatus 0.148OW1 Colubridae Dipsadinae Hypsiglena torquata 0.147NW1 Elapidae Elapinae Naja kaouthia 0.146OW1 Viperidae Viperinae Atheris squamigera 0.144OW3 Elapidae Hydrophiinae Emydocephalus annulatus 0.144OW0 Colubridae Colubrinae Drymarchon corais 0.143NW2 Colubridae Colubrinae Arizona elegans 0.143NW1 Colubridae Colubrinae Prosymna ambigua 0.143OW1 Elapidae Elapinae Naja atra 0.141OW1 Elapidae Elapinae Aspidelaps scutatus 0.136OW1 Colubridae Colubrinae Pituophis catenifer 0.135NW1 Elapidae Hydrophiinae Aipysurus laevis 0.134OW0 Colubridae Natricinae Virginia valeriae 0.134NW1 Colubridae Dipsadinae Carphophis amoenus 0.133NW1 Boidae Erycinae Charina trivirgata 0.133NW1 Colubridae Colubrinae Lampropeltis getula 0.131NW1 Colubridae Boodontinae Mehelya capensis 0.130OW2 Colubridae Boodontinae Lamprophis fuliginosus 0.130OW1 Tropidophiidae Tropidophiinae Tropidophis morenoi 0.129NW1 Colubridae Xenodontinae Pseudoeryx plicatilis 0.129NW1 Viperidae Crotalinae Bothrops insularis 0.128NW2 Colubridae Xenodontinae X enodon severus 0.127NW1 Tropidophiidae Tropidophiinae Tropidophis haetianus 0.127NW2 Colubridae Xenodontinae Hydrodynastes gigas 0.126NW1 Boidae Boinae Epicrates striatus 0.125NW2 Colubridae Xenodontinae Heterodon nasicus 0.125NW1 Colubridae Boodontinae Lamprophis lineatus 0.123OW1 Colubridae Colubrinae Chilomeniscus stramineus 0.123NW1 Boidae Pythoninae Python molurus 0.122OW2 Colubridae Colubrinae Dasypeltis fasciata 0.122OW2 Colubridae Colubrinae Crotaphopeltis hotamboeia 0.121OW1 Colubridae Colubrinae Dasypeltis scabra 0.120OW2 Colubridae Dipsadinae Atractus clarki 0.116NW1 Viperidae Viperinae Vipera xanthina 0.115OW1 Viperidae Viperinae Macrovipera mauritanica 0.114OW1 Colubridae Boodontinae Duberria lutrix 0.113OW1 Colubridae Xenodontinae Erythrolamprus aesculapii 0.111NW1 Viperidae Crotalinae Agkistrodon contortrix 0.110NW1 Colubridae Dipsadinae Atractus hostilitractus 0.107NW1

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54 Table 3-1. Continued Family Subamily Species RTLRg Gh Colubridae Boodontinae Lycophidion variegatum 0.105OW 1 Boidae Pythoninae Python sebae 0.104OW 2 Elapidae Hydrophiinae Pelamis platurus 0.104OW 0 Colubridae Colubrinae Prosymna stuhlmannii 0.103OW 1 Colubridae Dipsadinae Atractus depressiocellus 0.103NW 1 Colubridae Xenodontinae Farancia abacura 0.102NW 1 Boidae Pythoninae Morelia viridis 0.102OW 3 Xenopeltidae X enopeltis unicolor 0.100OW 1 Acrochordidae Acrochordus granulatus 0.099OW 0 Elapidae Elapinae Elapsoidea sundevallii 0.099OW 1 Colubridae Colubrinae Phyllorynchus decurtatus 0.097NW 1 Colubridae Boodontinae Lycophidion capense 0.096OW 1 Elapidae Hydrophiinae Laticauda colubrina 0.096OW 0 Viperidae Viperinae Causus rhombeatus 0.096OW 1 Colubridae Dipsadinae Atractus darienensis 0.095NW 1 Elapidae Hydrophiinae Hydrophis melanocephalus 0.093OW 0 Boidae Boinae Boa constrictor 0.090NW 2 Colubridae Colubrinae Pituophis melanoleucus 0.090NW 1 Viperidae Crotalinae Atropoides nummifer 0.088NW 1 Elapidae Elapinae Bungarus fasciatus 0.085OW 1 Colubridae Colubrinae Prosymna sundevallii 0.085OW 1 Leptotyphlophidae Leptotyphlops macrops 0.082OW 1 Colubridae Colubrinae Phyllorynchus browni 0.081NW 1 Viperidae Crotalinae Crotalus ruber 0.080NW 1 Viperidae Viperinae Bitis nasicornis 0.080OW 1 Colubridae Colubrinae Prosymna bivittata 0.080OW 1 Viperidae Crotalinae Crotalus horridus 0.076NW 1 Elapidae Elapinae Micrurus fulvius 0.075NW 1 Viperidae Viperinae Bitis arietans 0.075OW 1 Elapidae Elapinae Elapsoidea guentheri 0.075OW 1 Viperidae Crotalinae Crotalus atrox 0.073NW 1 Colubridae Colubrinae Stilosoma extenuatum 0.073NW 1 Boidae Boinae Acrantophis dumerili 0.071OW 1 Atractaspididae Aparallactinae X enocalamus sabiensis 0.069OW 1 Atractaspididae Aparallactinae Amblyodipsas ventrimaculatus 0.068OW 1 Boidae Pythoninae Python regius 0.066OW 1 Boidae Erycinae Gongylophis muelleri 0.064OW 1 Atractaspididae Aparallactinae Amblyodipsas polylepis 0.064OW 1 Elapidae Elapinae Elapsoidea nigra 0.064OW 1 Viperidae Viperinae Bitis gabonica 0.063OW 1 Viperidae Crotalinae Crotalus cerastes 0.063NW 1 Viperidae Viperinae Bitis caudalis 0.063OW 1 Viperidae Viperinae Bitis atrops 0.062OW 1 Atractaspididae Aparallactinae X enocalamus mechowii 0.060OW 1 Viperidae Viperinae Causus defilippii 0.060OW 1

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55 Table 3-1. Continued Family Subamily Species RTLRg Gh Elapidae Elapinae Elapsoidea semiannulata 0.055OW 1 Atractaspididae Atractaspidinae Atractaspis bibronii 0.052OW 1 Aniliidae Anilius scytale 0.044NW 1 Leptotyphlophidae Leptotyphlops alfredschmidti 0.042NW 1 Typhlopidae X enotyphlops grandidieri 0.035OW 1 Cylindrophiidae Cylindrophis ruffus 0.017OW 1 Typhlopidae Rhinotyphlops episcopus 0.011OW 1

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56 Figure 3-1. Snake total length fr equency distribution. A) Stenot opically arboreal species. B) Eurytopically arboreal/terrestrial species. C) Stenotopically terre strial species. D) Stenotopically aquatic species. 0 2 4 6 8 10 12 14 16 40 80 120 160200240280 Total Length (cm) N umber of Species 0 5 10 15 20 25 30 40 100 200 300 400500600700800900 Total Length (cm) N umber of Species A B

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57 Figure 3-1. Continued 0 5 10 15 20 25 30 20 60 100 140180220260 Total Length (cm) N umber of Species 0 1 2 3 4 5 6 7 8 9 10 40 80 120 160200240280 Total Length (cm) N umber of Species D C

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58 Figure 3-2. Coefficients of variation of log10 transformed total le ngth among the four gravitational habitats. 0.000 0.005 0.010 0.015 0.020 0.025 0.030 Stenotopically Arboreal Eurytopically Arb/Terr Stenotopically Terrestrial Stenotopically Aquatic Gravitational HabitatCoefficient of Variatio n

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59 2.22.42.62.83.03.23.43.6 0.51.01.52.02.5 Log(Total Length)Log(Tail Length) Figure 3-3. Regression of log10 tail length on log10 total length for 227 snake species (r2 = 0.65; P < 0.0001).

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60 2.22.42.62.83.03.23.43.6 0.51.01.52.02.5 Log(Total Length)Log(Tail Length) Figure 3-4. Analysis of cova riance (ANCOVA) on corrected ta il-length for 227 species of stenotopically arboreal (upper dotted line, open triangles), eurytopically arboreal/terrestrial (l ower dotted line, solid circles), stenotopically terrestrial, and stenotopically aquatic (overlapping solid lines, solid circles and open circles respectively) snakes.

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61 2.22.42.62.83.03.23.43.6 0.51.01.52.02.5 Log(Total Length)Log(Tail Length) Figure 3-5. Analysis of covari ance (ANCOVA) on corrected ta il-length for 227 species of scansorial (dotted line, open triangles), and stenotopically terrestrial and aquatic (overlapping solid lines, solid circles and open squares respectively) snakes.

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62 0 0.5 1 1.5 2 2.5 -1.75-1.5-1.25-1-0.75-0.5-0.25Log RTLMacrohabitat use Figure 3-6. Independent contrast s of macrohabitat use and log10 transformed relative tail-length (RTL; r2 = 0.31, P < 0.0001). For m acrohabitat use, 0 = sten otopically aquatic; 1 = stenotopically terrestr ial; 2 = scansorial.

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63 CHAPTER 4 DISCUSSION Relative Tail-Length, Gravit y, and Climbing in Snakes Gravity is a pervasive force that clearly prov ides acute challenges to blood circulation in terrestrial vertebrates. These challenges are pa rticularly pronounced in elongate vertebrates such as snakes utilizing complex three-dimensional vertical habitats (Lil lywhite 1996). Thus, the ability to regulate blood circulat ion in vertical habitats or up right postures potentially imposes important constraints on the behavior and morpho logy of scansorial snake species. Overcoming these cardiovascular constraint s has likely resulted in the evolution of novel morphological adaptations as species radiated from terrestrial into arboreal habitats. A likely consequence of limblessness in snakes is an increase in the selective pressures imposed on various aspects of the body and tail (Polly et al. 2001). This is particularly true in species adapted to the extreme challenges associ ated with arboreal environments (Lillywhite 1996; Lillywhite and Henderson 2001). Therefore, differences in body and tail characteristics should exist among snake species inhabiting diffe rent macrohabitats and, thus, experiencing different selective pressures. One such morphological change likely involves th e total length of ste notopically arboreal species. The distribution freque ncy of total length for adult stenotopically arboreal species suggests there are constraints limiting total length to between 50 and 200 cm, with few exceptions. There were no stenotop ically arboreal species with tota l lengths < 50 cm as adults. Because gradients of intravascular and transmur al pressures increase with absolute vertical length of a blood column, relatively short species are not strongly affected by the cardiovascular constraints imposed by gravity. Carotid blood fl ow in the stenotopica lly terrestrial viper Agkistrodon piscivorus was less impaired by tilting in sma ller than in larger individuals

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64 (Lillywhite and Smits, 1992). Furthermore, Lill ywhite and Smits (1992) suggest that gravity induced cardiovascular constraints on blood circulation limit the tota l length of arboreal vipers to about 1 m. For snakes in general, my results in dicate that constraints lim it total length in adult stenotopically arboreal species to > 50 cm and that these constraints are not likely cardiovascular. Insofar as gravity would not seem to be a f actor at smaller body size, species < 50 cm may have lesser ability to adequately span gaps, which might result in ecological constraints such as increased vulnerability to predation or decreas ed hunting success. Because several of the arboreal species included in the distribution freq uency have closely rela ted terrestrial species with total lengths < 50 cm (e.g., Trimeresurus ), the comparative data support the inference that there are lower limitations on the total length of stenotopically arboreal snakes to > 50 cm. Similarly, there were few stenot opically arboreal snake species (n = 5) with total lengths > 200 cm suggesting upper level constr aints on the total leng th of these species. The four species between 201-240 cm (in orde r of increasing length: Dendroaspis viridis Boiga cynodon Boiga forsteini and Gonyosoma oxycephalum ) all have relatively long tails, with three having relative tail-lengths between 23-26% ( no length data available for Boiga forsteini although its relative tail-length (RTL) should be within this range as well). Only one species ( Leptophis ahaetulla ) slightly exceeded 240 cm in total length (247.5 cm), and it also has a relati vely long tail (39% of total length). Snakes > 200 cm are perhaps also negatively influenced by ecological constraints in complex vertical three-dimensional habitats. Their relatively large mass (particularly in boids) might restrict movements to thicker branches. The variance in total length among eurytopically arboreal/terrest rial species was more than twofold greater than that of st enotopically arboreal species. In fact, the stenotopically arboreal

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65 species had the lowest variance in total length of all four categories. These results further suggest there are constraints limiting total length of stenotopically arboreal species, particularly because I did not find differences in either the mean total length or the mean corrected tail-length between stenotopically and eurytopica lly arboreal/terrestrial species. Corrected tail-length of scanso rial species was on average 1.9 times longer than that of non-scansorial species (Fig. 3-5). Lengthening the tail relative to the body in creases the relative length of blood vessels that ar e surrounded by tight (low compliance) tissues. This helps to mitigate posterior blood pooling and thereby facilita tes blood flow to the h eart, brain, and vital organs during the upright positions experienced during climbing (Lillywhite and Gallagher 1985; Lillywhite 2005). Therefore, the long tails of scan sorial snakes can be regarded as one of a suite of characters likely to be adaptive responses to the cardiovascular constraints on blood circulation imposed by gravity (Lillywhite 1987; Lillywhite 2005). Relative tail-length is positively correlated with absolute tail-length (Fig. 4-1) and likely more important in countering the effects of grav ity than absolute tail-length. Snakes with relatively longer tails have a larger percent of dependent vessels contained within the tight integument of the tail and are thus more resistan t to dependent transmural pressures experienced during climbing. I did not find a difference in RTL be tween stenotopically and eurytopically arboreal/terrestrial species (Fig. 3-4). There are two possible expl anations for this. (1) Even though eurytopically arboreal/terre strial species climb less fre quently than stenotopically arboreal species, they still need adaptations to counter the effects of gravity on blood circulation. This is the most likely explana tion, and it could help explain w hy many eurytopic species that periodically climb (e.g., Coluber constrictor and Masticophis flagellum ) have relatively long

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66 tails, but lack prehensile tail s. (2) Stenotopically arboreal sp ecies might have evolved from eurytopically arboreal/terrestrial species. He nderson and Binder (1980) proposed this hypothesis in a review of the ecology and be havior of vine snakes. Howeve r, the composite tree in this study does not support this hypothesis due to lack of resolution (F ig. 2-1). The tree shows two instances where stenotopically arboreal sp ecies likely evolved from eurytopically arboreal/terrestrial ancestors, and at least four instances where eurytopically arboreal/terrestrial species likely evolved from stenotop ically arboreal ance stors (Fig. 2-1). Ho wever, there are at least three instances in which th e ancestral state is unclear. Analyses using better resolved phylogenies at the generic and species level for mu ltiple clades are necessary to thoroughly test this hypothesis. The relative tail-lengths for the stenotopically aquatic speci es (9.3% to 19.2%) fall well within the range for the stenotopi cally terrestrial species (1.1% to 38.3%). This is likely because the tails of stenotopically a quatic species are affected by the unique selective pressures associated with an essentially weightless environm ent. In most aquatic environments, the tails do not function in countering the effects of grav ity on blood circulation, bu t rather in propulsion. Consequently, these species should be expected to have relative tail-lengths long enough to effectively propel the snake through water, ye t short enough to minimize drag. Therefore, similarity in RTL between stenotopically aquatic and stenotopically terrestrial species is likely the result of entirely different selective pressures. Several patterns emerged from the data regard ing RTL at the family level. First, the Viperidae in general have rela tively short body lengths, absolute tail lengths, and relative taillengths, and these characteristics appear to be re tained in arboreal species (Lillywhite and Smits 1992). The results from this study support these obs ervations, with the highest RTL value for an

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67 arboreal viperid in this study being 18.4% for Protobothrops cornutus (Table 3-1). However, even though vipers tend to be relatively shor t, Martins et al. (2001) demonstrate that RTL increases with increasing arboreality in the genus Bothrops This trend for increased RTL in scansorial species is further supported by the 25 viperid species included in this study, and it likely characterizes viperids in general. Simila rly, the 11 boid species used in this study tended to have relatively short tails, although many speci es are relatively long bodied. The highest RTL value for a stenotopically arboreal boid was 16.2% for Corallus ruschenbergerii (Table 3-1). The species with the longest relativ e tail-lengths were clearly memb ers of the family Colubridae. The 87 species with the longest RTL values were all colubrids (ranging from 20.8% 48.1%), with the only exception of two arboreal elapids ( Dendroaspis viridis and Dendroaspis angusticeps ) with RTL values of 26.0% and 25.8% re spectively (Table 3-1). Although Dendroaspis polylepis is fast moving and an agile climber, it is primarily terrestrial and has a shorter RTL (19.9%) than its stri ctly arboreal congeners even t hough they attain similar total lengths (Branch 1998). Snake Tails and Defense: Speed and Pseudautotomy In addition to scansorial species, relatively long tails were also noted in several nonscansorial species characterized by defensive tail loss (pseudautotomy) and/or using speed as a primary means of capturing prey and escaping predators (e.g., Darlingtonia Drepanoides and Thamnophis ). In fact, many of these species had RTL values similar to many scansorial species (RTLs of 29 species included in this study were between 20% and 34%). Included within this group of species that use th eir tails to escape predators are species that often swim, but evidence suggests that relatively long tails are not particularly advantageous for either rapid terrestrial loco motion or swimming (Jayne 1988; Jayne and Bennett 1989). Jayne (1988) found that RTL in Nerodia and Elaphe were about equally effective for swimming,

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68 suggesting the relatively long tails of natricines are not an adaptation for swimming. However, of the 29 species of stenotopically terrestrial species with relatively long tails included in this study, at least 19 are known from the literature to lose their tails in defense. While the relative frequency of tail loss in wild populations cannot always be accura tely determined from museum specimens alone due to potential collection bi as (Arnold 1988; Jayne and Bennett 1989), these results certainly warrant further investigation. Many long-tailed terrestrial species are also active, diurnal pred ators that rely on speed to procure prey (Greene 1997). However, if fast moving terrestrial snakes possess adaptations for speed, and long RTL is not required for speed, th en alternative selectiv e pressures are likely responsible for maintaining long tails in these species. Furthermore, because many of these species are known to possess stub ta ils (Greene 1997), it is reasonabl e to infer that the long tails of these terrestrial speci es might be a mechanism for defense. Many of these species rely on speed for escaping predators, and quickly fleeing increases the lik elihood of a predatory attempt being directed to the tail instead of the body (G reene 1973). A long tail can subsequently break allowing the snake to escape. Guyer and Donnelly (1990) used length-mass rela tionships to divide a Costa Rican snake assemblage of 27 species into four distin ct morphological groups and found that, among 603 individuals, the highest frequenc ies of tail breaks occurred in th e leaf litter and terrestrial categories (mean = 0.19 and 0.08 respectively) w ith the arboreal and semiarboreal categories having the lowest (mean = 0.01 and 0.0 respectivel y). In fact, the leaf-litter habitat was characterized by having long RTL a nd high frequency of tail loss (e.g., Coniophanes Dendrophidion Pliocercus Rhadinaea and Scaphiodontophis ) and, thus, highlighting the apparent prevalence of pseudautotomy in many terrestrial snake species within a single

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69 assemblage. Additional studies of snake assemblages in other areas would be of considerable use in determining the frequency of pseuda utotomy among snake assemblages in diverse ecosystems. The current dogma regarding ps eudautotomy suggests it is a rare phenomenon in snakes (Arnold 1988; Marco 2002). However, the paucit y of information on this topic renders such a conclusion premature. As more natural history data become available, the number of snake species observed exhibiting relatively high freque ncies of tail loss in wi ld populations continues to increase. The sum of these findings suggests that pseudautotomy in snakes is possibly more common than presently thought and potentially important in understand ing the evolution of relatively long tails in many terrestrial snake species. However, different functions of tail length need not be mutually exclusive. It is likely that the long RTL of many eurytopically arboreal/terrestrial species (e.g., Coluber constrictor ) serve the dual purposes of aiding in defense by pseuda utotomy, while also countering the effects of gravity during climbing. Further studies comp aring tail characteristic s of scansorial and terrestrial species (e.g., prehen sion, tail length, presence of frac ture planes, size of caudal vertebra, and muscle mass) would be of considerable interest a nd I intend to pursue these ideas in a later study.

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70 -2.500 -2.000 -1.500 -1.000 -0.500 0.000 0.000.501.001.502.002.503.00Log10 (absolute tail-length)Log10 (RTL) Figure 4-1. Regression of absolu te tail length on rela tive tail-length (RTL) for 227 snake species (r2 = 0.64, P < 0.0001).

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71 APPENDIX A STENOTOPICALLY ARBOREAL SPECIES Table A-1. Stenotopically arboreal species (4 families, 10 subfamilies, 34 genera, 77 species) used for total length frequency distri butions; NW = New World, OW = Old World. Family Subfamily Species Rang e Boidae Boinae Corallus annulatus NW Corallus caninus NW Corallus ruschenbergerii NW Pythoninae Morelia viridis OW Colubridae Colubrinae Ahaetulla dispar OW Ahaetulla nasuta OW Ahaetulla prasina OW Ahaetulla pulverulenta OW Boiga cynodon OW Boiga blandingii OW Boiga cyanea OW Boiga forsteini OW Boiga multifasciata OW Boiga ocellata OW Chrysopelea ornata OW Chrysopelea paradisi OW Dendrelaphis cyanochloris OW Dendrelaphis pictus OW Dipsadoboa aulica OW Dispholidus typus OW Drymobius margaritiferus NW Elaphe frenata OW Elaphe prasina OW Gastropyxis smaragdina OW Gonyosoma oxycephalum OW Leptophis ahaetulla NW Leptophis depressirostris NW Opheodrys aestivus NW Oxybelis aeneus NW Oxybelis fulgidus NW Philothamnus angolensis OW Philothamnus dorsalis OW Philothamnus hoplogaster OW Philothamnus irregularis OW Philothamnus semivariegatus OW Thelotornis capensis OW Thelotornis kirtlandii OW Thrasops jacksonii OW Dipsadinae Dipsas boettgeri NW Dipsas catesbyi NW Dipsas chaparensis NW Dipsas indica NW

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72 Table A-1 Continued Family Subfamily Species Rg Colubridae Dipsadinae Dipsas peruana NW Imantodes cenchoa NW Imantodes inornatus NW Leptodeira annulata NW Leptodeira septentrionalis NW Sibon nebulata NW Sibon sartorii NW Psammophiinae Psammophis biseriatus OW Pseudoxyrhophiinae Langaha alluaudi OW Langaha madagascariensis OW Langaha pseudoalluaudi OW Micropisthodon ochraceus OW Stenophis gaimardi OW Stenophis granuliceps OW Xenodontinae Philodryas baroni NW Siphlophis cervinus NW Uromacer catesbyi NW Uromacer frenatus NW Uromacer oxyrhynchus NW Elapidae Elapinae Dendroaspis angusticeps OW Dendroaspis viridis OW Viperidae Crotalinae Bothriechis schlegelii NW Trimeresurus cornutus OW Trimeresurus erythrurus OW Trimeresurus gramineus OW Trimeresurus macrolepis OW Trimeresurus malabaricus OW Trimeresurus medoensis OW Trimeresurus purpureomaculatus OW Tropidolaemus wagleri OW Viperinae Atheris chlorechis OW Atheris hispida OW Atheris nitschei OW Atheris rungweensis OW Atheris squamigera OW

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83 Stafford, P. J. 2004. A new species of Tantilla (Serpentes; Col ubridae) of the Taeniata group from southern Belize. J. Herpetol. 38:43–52. Tiebout, H. M., III. 1997. Caudal luri ng by a temperate colubrid snake, Elaphe obsoleta and its implications for the evolution of the rat tle among rattlesnakes. J. Herpetol. 31:290–290. Uetz, P. The EMBL Reptile Database. [upda ted 14 November 2006; cited 16 November 2006]. Available from http://www.embl-heidelberg.de /~uetz/LivingReptiles.html Utiger, U., N. Helfenberger, B. Schtti, C. Sc hmidt, M. Ruf, and V. Ziswiler. 2002. Molecular systematics and phylogeny of old and new world ratsnakes, Elaphe Auct., and related genera (Reptilia, Squamata, Colubrid ae). Russian J. Herpetol. 9:105–124. Vidal, N., and S. B. Hedges. 2002. Higher-level relationships of snakes inferred from four nuclear and mitochondrial ge nes. C. R. Biol. 325:977–985. Vidal, N., S. G. Kindl, A. Wong and S. B. Hedges. 2000. Phylogenetic relationships of Xenodontine snakes inferred from 12S and 16S ribosomal RNA sequences. Mol. Phylogenet. Evol. 14:389–402. Vitt, L. J., and L. D. Vangilder. 1983. Ecology of a snake community in northeastern Brazil. Amphibia-Reptilia 4:273–296. Voris, H. K. 1975. Dermal scale-vertebra relatio nships in sea snakes (Hydrophiidae). Copeia 1975:746–755. Wallach, V., and I. Ineich. 1996. Redescrip tion of a rare Malagasy blind snake, Typhlops grandidieri Mocquard, with placement in a ne w genus (Serpentes: Typhlopidae). J. Herpetol. 30:367–376. Wharton, C. H. 1960. Birth and behavi or of a brood of cottonmouths, Agkistrodon piscivorus piscivorus with notes on tail-luri ng. Herpetologica 16:125–129. Whitaker, R., and A. Captain. 2004. Snakes of I ndia: the field guide. Draco Books, Chennai, India. Whiting, A. M. 1969. Squamate cloacal glands : morphology, histology, and histochemistry. Doctoral dissertation, Pe nnsylvania State Univ. Wiens, J. J., M. C. Brandley, and T. W. R eeder. 2006. Why does a trait evolve multiple times within a clade? Repeated evolution of snake like body form in squamate reptiles. Evolution 60:123–141. Wikramanayake, E. D. 1990. Ecomorphology and biogeography of a tropical stream fish assemblage: evolution of assemb lage structure. Ecology 71:1756–1764.

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84 Wster, W., M. R. Duarte, and M. da Graa Salomo. 2005. Morphological correlates of incipient arboreality and ornithophagy in isla nd pitvipers, and the phyl ogenetic position of Bothrops insularis J. Zool. Lond. 266:1–10. Zimmerman, K., and H. Heatwole. 1990. Cutane ous photoreception: a new sensory mechanism for reptiles. Copeia 1990:860–862.

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85 BIOGRAPHICAL SKETCH Coleman Sheehy has always had a passionate cu riosity about the natural world. Growing up in Richmond, Virginia, his earliest interests centered around dinosaurs but these interests quickly expanded to include living reptiles and am phibians and, in particular, snakes. Thanks to an encouraging and supportive family, Coleman’ s basement, while growing up, was constantly filled with native wildlife he collected on local field excursions. Most of these animals were reptiles and amphibians and it was during thes e years that Coleman honed many of the observational and field skills that would become such an important aspect of his academic career. After acquiring his General Education Diploma (GED), Coleman was invited to work at James River Park in Richmond, where his responsi bilities included leadin g interpretative hikes and float trips through miles of undeveloped, island-dotted habita t along the James River. He also gave hundreds of educational programs focuse d on introducing the public of all ages to the fascinating reptiles and amphibians native to Virginia and to the importance of their conservation. Coleman later began working at a local pet store specializing in exotic reptiles and amphibians and, after several years, became genera l manager of the primary store in Richmond. It was during this time that Coleman realized ma ny of the problems associated with the pet trade and he sought to resolve many of these by in itiating captive breeding programs and by only purchasing or trading captive bred animals. Also during this time, Coleman saved enough money to fund a month-long trip exploring Madagas car. It was during this trip that he met two prominent American herpetologists (Drs. Rona ld Nussbaum and Christopher Raxworthy) who strongly influenced his career pa th. The experience left Coleman with the realization that he could incorporate his love of herpetology and traveling into a successful career through

PAGE 86

86 academia. Coleman subsequently enrolled at J. Sargeant Reynolds Community College in Virginia and graduated summa cum laude with an Associate in Science in the year 2000. He then transferred to the University of Florida, where he graduated magna cum laude with a bachelor’s degree in zoology and a minor in wildlife ecology and conservation in 2002. Coleman has traveled to Madagascar, H onduras, South Africa, Tobago, the Bahamas, Colombia, Brazil, and Taiwan as well as many states within the US. While working on his master’s, Coleman taught the laboratories for many classes, including Herpetology, the Natural History of Amphibians and Reptiles, Verteb rate Zoology, Functional Vertebrate Anatomy, Ecology, Advanced Island Biogeography, and Introductory Biology. He has worked in the herpetology collections of the Florida Museum of Natural History and the Transvaal Museum in South Africa, and he has presente d several talks at scientific m eetings. He has also published regularly in peer-reviewed jour nals since 2001 and, to date, has ten publications including the description of a new species of frog from Bolivia. Coleman has been accepted into a Ph.D. program at the Univer sity of Texas at Arlington, where he will pursue his interest s in snake systematics, evolu tional biology, ecomorphology, and functional morphology by studying several groups of Central and South American snakes. His long-term goal is to secure a tenured position in a Museum or University, where he can conduct research and teach in the areas of herpetology and vertebrate evolution, while also actively promoting the conservation of wildlife worldwide.


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ON THE STRUCTURE AND FUNCTION OF TAILS IN SNAKES:
RELATIVE LENGTH AND ARBOREALITY




















By

COLEMAN MATTHEW SHEEHY III


A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
MASTER OF SCIENCE
UNIVERSITY OF FLORIDA
2006


























Copyright 2006

By

Coleman Matthew Sheehy III




































To the increased understanding, respect, and conservation of snakes worldwide









ACKNOWLEDGMENTS

I thank my committee members Harvey B. Lillywhite (chair), James Albert, and Max A.

Nickerson for their guidance and support. Harvey B. Lillywhite first inspired this work by

suggesting a link between snake tails and the gravitational hypothesis. I thank Harvey B.

Lillywhite and Michael B. Harvey for providing unpublished snake length data, Florida Museum

of Natural History (FLMNH) curators F. Wayne King and Max A. Nickerson for access to the

Herpetology collection, Roy McDiarmid and George Zug at the United States National Museum

(USNM) for permission to use the Herpetology collection, and Steve Gotte at the USNM and

Kenney L. Krysko at the FLMNH for supplying data from the Herpetology collection databases.

I thank Michael B. Harvey, Laurie Vitt, James McCranie, Rom Whitaker, Bob Henderson, Bob

Powell, Blair Hedges, Ming Tu, Max A. Nickerson, Richard Saj dak, Bill Love, and John Rossi

for providing difficult to aind snake natural history information.

I thank Michael B. Harvey and Ron Gutberlet for information on snake phylogenies,

Michael McCoy for assistance with various statistical programs and analyses, and Kent Vliet for

the use of a Macintosh computer. I thank Andres Lopez and Griffin Sheehy for help with

illustrating programs, Andres Lopez for help with phylogenetic programs, and Jason Neville for

computer assistance. I thank Glades Herp Inc. for permission to measure live snakes and,

perhaps more importantly, I am grateful to Russel L. Anderson, Sam D. Floyd and Ryan J. R.

McCleary for assistance in handling and measuring live snakes, many of which were large,

highly venomous, and uncooperative. In addition to my committee members, I thank Ryan J. R.

McCleary, David A. Wooten, Leslie Babonis, Chris Samuelson, Bruce Jayne, and Roy

McDiarmid for stimulating discussions regarding snake natural history and evolution. I thank

Griffin E. Sheehy and Andrea Martinez for all their patience, love and support. Finally, I want to










thank my parents Coleman M. Sheehy, Jr. and Ellen R. Sheehy for never failing to support and

encourage a young boy's endless passion for snakes. I could not have been more lucky.












TABLE OF CONTENTS
page

ACKNOWLEDGMENTS .............. ...............4.....


LIST OF TABLES .........__.. ..... .__. ...............8....


LIST OF FIGURES .............. ...............9.....


AB S TRAC T ............._. .......... ..............._ 10...


CHAPTER


1 INTRODUCTION ................. ...............12.......... ......


Gravitational Influence on Tail Morphology ................. ...............13........... ...
Length Limitations ................. ................. ...............14......
Relative Tail Length and Macrohabitat Use ................. ...............15...............
Multiple Functions of Tail Use ................. ...............17........... ...
Locomoti on ................. ...............17........... ....
Caudal Luring ................. ...............20.................
Defense .................. .............. ...............21.......

Morphology and Reproduction................... ............2
Sexual Dimorphism and Ontogenetic Shifts .............. ...............25....

2 MATERIALS AND METHODS ................. ...............27................


Categorizing Climbing in Snakes: Gravitational Habitat ................. ......... ................27
A nalyses......................... ...........3
Frequency Distribution ................... ... ... ........ ...............3.. 1....
Gravitational Habitat and Total Body Length ................ ...............31...............
Gravitational Habitat and Tail Length ................. ...............32........... ...
Constructing the Phylogeny ........................ ............ ........3
Relative Tail-Length, Gravitational Habitat, and Phylogeny ................. ................ ...34

3 RE SULT S .............. ...............48....


Frequency Distribution ................... ... .. ....... ...............48.......
Gravitational Habitat and Total Body Length .............. ...............48....
Gravitational Habitat and Tail Length ................. .............. ...............49.....
Relative Tail-Length, Gravitational Habitat, and Phylogeny .............. ....................4

4 DI SCUS SSION ................. ...............63................


Relative Tail Length, Gravity, and Climbing in Snakes............... ...............63.
Snake Tails and Defense: Speed and Pseudautotomy .............. ...............67....

APPENDIX STENOTOPICALLY ARBOREAL SPECIES............. ..............71..













LIST OF REFERENCES ................. ...............73................


BIOGRAPHICAL SKETCH .............. ...............85....










LIST OF TABLES


Table page
2-1 Taxa and associated data sources for 227 snake species included in this study. ...............36

2-2 Total variation in relative tail-lengths of 26 species of snakes ................. ............... ....43

3-1 The 227 species included in this study .............. ...............50....

A-1 Stenotopically arboreal species............... ...............71










LIST OF FIGURES


Figure page

2-1 Composite phylogeny of 227 snake taxa. ............. ...............44.....

3-1 Snake total length frequency distribution. ............. ...............56.....

3-2 Coefficients of variation of loglo transformed total length ................. ............ .........58

3-3 Regression of loglo tail length on loglo total length ................. ................ ......... .59

3-4 Analysis of covariance (ANCOVA) on corrected tail-length for 227 species of
stenotopically arboreal. ............. ...............60.....

3-5 Analysis of covariance (ANCOVA) on corrected tail-length for 227 species of
sc ansori al ................. ...............61................

3-6 Independent contrasts............... ...............6

4-1 Regression of absolute tail length on relative tail-length............... ..............7









Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Science

ON THE STRUCTURE AND FUNCTION OF TAILS IN SNAKES:
RELATIVE LENGTH AND ARBOREALITY

By

Coleman Matthew Sheehy III

December 2006

Chair: Harvey B. Lillywhite
Maj or Department: Zoology

The pervasive effect of gravity on blood circulation is an important consequence for

elongate animals such as snakes that utilize arboreal habitats. Upright postures create vertical

gradients of hydrostatic pressures within circulatory vessels, and the magnitude of the pressure

change is proportional to the total length of the blood column. In air, this potentially induces

blood pooling and edema in dependent tissues and a decrease in the volume of blood reaching

the head and vital organs of animals that are tall or elongate. Arboreal snakes exhibit a suite of

behavioral, morphological, and physiological adaptations for countering the effects of gravity on

blood circulation including relatively non-compliant tissue compartments in the tail.

Comparative studies involving arboreal, terrestrial, and aquatic species have

demonstrated that blood pooling in dependent tissues of arboreal snakes can be tenfold lower

than in terrestrial and aquatic species. In addition to non-compliant tail compartments, arboreal

species appear to have longer tails relative to their non-climbing terrestrial and aquatic

counterparts. However, the generality of tail length patterns related to arboreal habitats and

gravity has not been previously studied in a broad range of taxa. Here I examine the hypothesis

that there are constraints limiting the total length of arboreal snakes and that arboreal snakes









have relatively longer tails to help counter the effects of gravity while also helping to adjust total

length to within the range of limitations.

I used macrohabitat and behavior to create four gravitational habitats to categorize the

amount of climbing and, therefore, the amount of gravitational stress experienced: stenotopically

arboreal, eurytopically arboreal/terrestrial, stenotopically terrestrial, and stenotopically aquatic.

The effects of sexual dimorphism, and ontogenetic shifts in allometry and habitat use, were

avoided by using only adult females. Data were acquired from the literature, museum

specimens, and live snakes. Frequency distributions and analysis of variance (ANOVA) were

used to evaluate the assumption that total length limitations exist in stenotopically arboreal

species. I tested for differences in relative tail-length (RTL) among the four gravitational

habitats (227 species representing almost all snake families and subfamilies) by using analyses of

covariance (ANCOVA). I tested for correlations between macrohabitat use and RTL within a

phylogenetic framework by independent contrasts analyses after constructing a composite tree.

The RTL for the 227 species included in this study ranged from 1.1% (Rhinotyiphlops

episcopus) to 48.1% (Uromacer frenatus). The total length of stenotopically arboreal species

appears to be constrained to between 50 and 200 cm, with few exceptions. The RTL between

stenotopically arboreal and eurytopically arboreal/terrestrial species did not differ and were

therefore combined into a single scansorial category. Scansorial species had RTLs on average

two times longer than non-scansorial species. Snakes with relatively longer tails have a larger

percent of dependent vessels contained within the tight integument of the tail and are thus more

resistant to blood pooling experienced during climbing. Therefore, the relatively long tails of

scansorial snakes can be regarded as one of a suite of characters likely to be adaptive responses

to the cardiovascular constraints on blood circulation imposed by gravity.









CHAPTER 1
INTTRODUCTION

Macrohabitat use is thought to strongly influence the evolution of vertebrate morphology

(Miles and Ricklefs 1984; Wikramanayake 1990; Lillywhite 1996; Martins, Araujo, Sawaya, and

Nunes 2001). Even in limbless and elongate vertebrates such as snakes, the evolution of

morphology is strongly influenced by the type of macrohabitat utilized (Vitt and Vangilder 1983;

Guyer and Donnelly 1990; Cadle and Greene 1993; Lillywhite and Henderson 2001; Martins et

al. 2001). The roughly 3000 extant snake species have demonstrated a remarkable propensity for

radiating into a wide variety of fossorial, arboreal, terrestrial, freshwater, and marine habitats in

every part of the biosphere excluding only the deep sea and polar regions. This diverse

ecological radiation, along with the loss of limbs, makes snakes excellent animals for

investigating the effects of macrohabitat use on the evolution of morphology.

Arboreality has evolved numerous times independently among snakes, and several studies

have identified various behavioral, morphological, and physiological characteristics associated

with snakes in arboreal habitats (e.g., Johnson 1955; Marx and Rabb 1972; Henderson and

Binder 1980; Lillywhite 1993; Lillywhite and Henderson 2001; Martins et al. 2001). While it is

often difficult to demonstrate adaptation (e.g., Gould and Lewontin 1979; Bock 1980), the fact

that a phylogenetically diverse array of arboreal species share strikingly similar, if not identical,

morphological, behavioral, and physiological specializations supports the inference that these

traits are adaptive and, in many cases, the result of convergent evolution (Lillywhite 1996;

Lillywhite and Henderson 2001).

Studies investigating (directly or indirectly) snakes' tails have included a wide range of

topics including cardiovascular adaptations (e.g., Lillywhite 1985, 1993, 2005; Lillywhite and

Henderson 2001), sexual dimorphism (e.g., Klauber 1943; King 1989; Shine 1993, 2000), mating









success (e.g., Shine et al. 2000), use in locomotion (e.g., Jayne 1988; Jayne and Bennett 1989),

the evolution of vertebral elements (e.g., Johnson 1955; Lindell 1994; Shine 2000), defensive

adaptations (e.g., Greene 1973, 1988; Arnold 1988; Mendelson 1991; Savage and Slowinski

1996), caudal luring (e.g., Greene 1997), development (Polly, Head, and Cohn 2001), and

ecological correlations (e.g., Guyer and Donnelly 1990; Martins et al. 2001; Lobo, Panday, and

Vasudevan 2004; Wilster, Duarte, and da Graga Salomio 2005; Wiens, Brandley, and Reeder

2006). However, the adaptive role of the tail in arboreal habitats is poorly understood. For

example, the importance of the tail in snake locomotion (particularly in arboreal habitats)

remains unclear (Jayne 1988; see below).

Gravitational Influence on Tail Morphology

An important consequence of utilizing complex, three-dimensional vertical habitats is the

pervasive effect of gravity on blood circulation, which can be particularly pronounced in tall or

long organisms such as giraffes and snakes (Lillywhite 1993,1996). Upright postures create

vertical gradients of hydrostatic pressures within circulatory vessels. In air, this increases

transmural pressures related to the absolute length of the fluid column, leading to a tendency for

blood pooling and for fluid to filter from capillaries into surrounding tissue compartments

resulting in edema in dependent tissues (Lillywhite 1985, 1993). As blood pooling and edema

increase, the volume of blood reaching the heart decreases, thereby reducing the central blood

pressure and consequently the amount of blood flow to the head and vital organs (Lillywhite

1993b, 1996, 2005; Lillywhite and Henderson 2001).

Arboreal snakes exhibit a suite of behavioral, morphological, and physiological

adaptations for countering the effects of gravity on blood circulation including stereotypical body

movements, small mass/length ratios, relatively anterior hearts, tightly applied integument, and

relatively non-compliant tissue compartments in the tails (Lillywhite 1985, 1993; Lillywhite and









Henderson 2001). Comparative studies involving arboreal, terrestrial, and aquatic species

demonstrate that blood pooling in dependent tissues of arboreal snakes during vertical posture is

significantly less than in terrestrial and aquatic species, and these differences can approach

tenfold (Lillywhite 1985, 1993; Lillywhite and Henderson 2001). The ability for arboreal snakes

to defend against edema and blood pooling is almost certainly attributed to the aforementioned

characters, one of which being relatively non-compliant tissue compartments in the tail

(Lillywhite and Henderson 2001; Lillywhite 2005). However, the magnitude of the compliance

does not appear to be related to the length of the snake or its tail (Lillywhite 1993). There does,

however, appear to be a relationship between tail length and macrohabitat use in that arboreal

species generally have longer tails relative to their nonclimbing terrestrial and aquatic

counterparts (see below). What then are the selective pressures responsible for the interspecific

variation in tail length?

Length Limitations

Body size may be the most fundamental character of an organism, because nearly all

aspects of an organism's biology are correlated with this variable (Naganuma and Roughgarden

1990; Boback and Guyer 2003). The body sizes (i.e., total length) of snakes in general could be

evolving toward an optimal length of 1.0 m (Boback and Guyer 2003), suggesting that selection

and constraint might be influencing the total length of many snake species. However, the body

size distribution is not obviously skewed in either direction, and idiosyncratic features of the

natural history of snakes may be creating this distribution pattern (Boback and Guyer 2003).

Macrohabitat use likely imposes locomotor constraints upon snakes (Boback and Guyer 2003),

and there is considerable evidence that arboreal snakes are limited in their use of habitat as a

consequence of interactions between their morphology and the physical size of branches (see

Lillywhite and Henderson 2001, for a review on arboreal snake functional ecology). Arboreal









locomotion involves various combinations of undulatory, rectilinear, and concertina movements

along a complex and often unstable vertical three-dimensional substrate (Gans 1974; Edwards

1985; Lillywhite and Henderson 2001). Short snakes might not be able to adequately span gaps,

whereas large or heavy bodied snakes might not be supported by smaller branches (Henderson

and Nickerson 1976). Long snakes might be further limited physiologically by cardiovascular

constraints. Because foraging snakes must be able to span gaps between stems and branches and

approach prey without revealing their presence (Lillywhite and Henderson 2001), it is reasonable

to expect there are upper and lower limitations on total length in arboreal snake species.

Assuming there are length limitations imposed on snakes living in arboreal habitats, one

way snakes could have evolved total lengths within the acceptable range while still counteracting

blood pooling and assuring adequate blood flow to the head is by lengthening the tail relative to

the body since perivascular tissues are tighter in the tail than in the body cavity (Lillywhite

2005). Lengthening the tail relative to the body could potentially be achieved evolutionarily by

adding vertebrae to the tail or by elongating the caudal vertebrae themselves (Johnson 1955;

Shine 2000). Relative tail-length (RTL), which is the proportional ratio of tail length/total

length, is commonly used when performing interspecific comparisons in tail length to account

for total length differences (Klauber 1943). Relative tail-length in snakes appears to have low

intraspecific variation (i.e., it is a stable character), and this variation is further reduced when

investigating the sexes separately (Klauber 1943). Therefore, interspecific similarities and

differences in RTL are potentially useful sources of ecological and taxonomic information

(Klauber 1943).

Relative Tail-Length and Macrohabitat Use

Correlations between RTL and macrohabitat use have been suggested previously, but the

supporting data are weak or inconclusive. Klauber (1943) suggested that thick-bodied snakes









have relatively shorter tails than do thin-bodied snakes. Clark (1967) stated that fossorial habits

seem to be accompanied by a shortening of the tail. Marx and Rabb (1972) investigated 962

species of 195 colubrid genera and found that the maximum number of subcaudal scales

occurred in six species of snakes that were all arboreal (i.e., vine snakes). Because these six

species are not all closely related, they concluded that these characteristics represent derived

ecological specializations for arboreal habitats. King (1989) proposed that RTL in snakes is

highly variable interspecifically and appears to be correlated with ecological factors. He also

stated that other considerations such as mode of locomotion, habitat, and risk of predation, which

while apparently correlated with tail length in lizards, have not been investigated in snakes.

Greene (1997) stated qualitatively that the tails of phylogenetically basal snakes

(scolecophidians) and most vipers are especially short, while the tails of many colubrids and

elapids are longer. Martins et al. (2001) investigated 20 species of Bothrops and concluded that

an increase in RTL occurred along with an increase in arboreality in some clades.

A strong method for assessing the influence of habitat use on the evolution of body form in

snakes is to analyze monophyletic clades so that characters can be interpreted within an

explicitly phylogenetic framework (Harvey and Pagel 1991; Martins et al. 2001). Although

earlier studies have found a correlation between RTL and arboreality in snakes, most have not

separated species by lineages and thus might be confounded by phylogenetic effects. Martins et

al. (2001) addressed this issue using the monophyletic genus Bothrops, and their results

corroborate previous studies. However, the study investigated only 20 species within a single

clade, making the results difficult to apply to snakes in general. Similar studies using additional

monophyletic groups are needed to determine whether this trend is widespread in snakes.









Herein, I investigate the relationship between tail length and macrohabitat use in arboreal

species of snakes. The purposes of the present investigation are threefold. First, I evaluate the

assumption that there are limitations imposed on the total length of arboreal species. Second, I

test the hypothesis that arboreal species have relatively longer tails than nonclimbing species.

Third, I discuss the relationship between RTL and climbing within the context of gravitational

adaptation, and hypothesize that additional selective pressures might be directing the evolution

of relatively long tails in snakes.

Multiple Functions of Tail Use

Most vertebrate structures have multiple functions (Moon 2000), and snake tails are an

excellent example. As a likely consequence of limblessness, several selective pressures are

simultaneously acting on the tails of many snake species. For example, many juvenile pitvipers

(e.g., Agkistrodon piscivorus) use caudal luring to attract prey, but also vibrate their tails rapidly

in defense when threatened. In order to understand the functional versatility, as well as possible

constraints of complex vertebrate structures such as snake tails, information about how they are

used in diverse environments and behaviors is required (Moon 2000).

Locomotion

Snakes likely use their tails to assist in locomotion (e.g., balance, propulsion, holding or

climbing) (Klauber 1943); however, the extent to which this occurs is poorly understood (Jayne

1988; Lillywhite and Henderson 2001). Many fossorial species have short blunt tails (e.g., Eryx,

Sympholis lippiens, scolecophidians, and uropeltids), and some also have strongly keeled caudal

scales (e.g., Sonora aemula and uropeltids). The apparent commonality of these characteristics

among fossorial species suggests they are adaptations to fossorial habitats (e.g., locomotion and

defense). However, in many cases the functional morphology is not well studied. Clark (1967)

posits that the short tails of many fossorial snakes (i.e., scolecophidians) facilitate subterranean










locomotion by acting as an anchor against which the snake can push, since a long tail in this case

would likely bend and be unable to serve this purpose. However, Klauber (in Clark 1967)

suggests instead that burrowing would render the tail practically useless, and that the tails of

fossorial snakes are shortened secondarily due to non-use, rather than from direct selective

pressures.

In a combination of experimental and correlative analyses, Jayne and Bennett (1989)

demonstrated the difficulties in determining the effect of tail morphology on locomotor

performance in terrestrial snakes. Jayne and Bennett demonstrated that losses ranging from

0.03% to 80.4% tail length had no significant effect on terrestrial locomotory performance in 52

garter snakes (Thamnnophis sirtalis) with naturally incomplete tails. Locomotor performance was

not affected by the experimental removal of the distal one third of the tail. The experimental

removal of the distal two thirds of the tail only caused a small (4.5%) but significant average

decrease in speed. However, the same study also found that burst speed in T. sirtalis performing

terrestrial lateral undulation was fastest in individuals with intermediate relative tail-lengths.

Jayne and Bennett concluded that minor deviations from intermediate relative tail length do not

affect locomotor performance among snakes. These results demonstrate that long tails can

perhaps be slightly disadvantageous in terms of locomotor performance in T. sirtalis, and suggest

that the long tails of some snake species are possibly due to other additional selective pressures.

For example, high incidence of tail loss is found in the genera Nerodia and Thamnnophis

suggesting tail loss from predation attempts (King 1987; Jayne and Bennett 1989; Mendelson

1991; see section below on tail loss). Furthermore, juvenile Nerodia harteri have been observed

using their long tails to anchor themselves to rocks at the water' s edge while fishing (Rossi, pers.









comm.), whereas Nerodia sipedon has been observed using their tails to anchor themselves to

submerged sticks and rocks while fishing in currents (M. A. Nickerson pers. comm.; pers. obs).

Snake caudal vertebrae are complex, highly variable in size within individuals, and clearly

distinguishable from the trunk vertebrae (Johnson 1955). Because the number of caudal

vertebrae corresponds to the number of subcaudal scales by a ratio of 1:1 in most snake species,

snakes with longer tails typically have more subcaudals and thus more caudal vertebrae than

snakes with shorter tails (Ruthven and Thompson 1908; Gans and Taub 1965; Alexander and

Gans 1966; Voris 1975; Shine 2000). Differences in the relative size and number of caudal

vertebrae suggest that the effectiveness of the tail in locomotory propulsion varies considerably

among snake taxa (Jayne 1988). However, the consequences of caudal morphology likely vary

with mode of locomotion (Jayne 1988). More research is needed to determine the extent of tail

use in the various forms of snake locomotion.

The tails of most arboreal boids, viperids, and many colubrids are prehensile and assist in

climbing and securing to branches (Lillywhite and Henderson 2001). Tree boas (Corallus) often

wrap their prehensile tails around branches during prey capture and consumption (Henderson

2002; pers. obs.). Henderson (2002) observed tree boas (Corallus grenadensis) on Grenada

hanging by their tails from vegetation, with the forepart of their bodies in typical ambush

posture, presumably hunting for bats. I have observed similar hunting behavior in Corallus

ruschenbergerii on Tobago and in Corallus caninus in captivity. However, there appears to be

no correlation between long RTL and tail prehension for snakes in general (Lillywhite and

Henderson 2001).

Specializations in snake locomotion, such as highly efficient swimming and sidewinding,

have likely evolved multiple times and employ tail use to varying degrees. Sea snakes and sea









kraits have evolved several adaptations to a marine existence, one of which being flattened,

oarlike tails used for effective and rapid propulsion (Heatwole 1999). Yellow-bellied sea snakes

(Pelamnis platurus) are well adapted to a pelagic existence and have subsequently lost the ability

to crawl on land, spending their entire lives at sea (Greene 1997). Acrochordids have a muscle

that shapes the skin of the body and tail into a keel while swimming (Heatwole 1999). However,

not all marine snakes have oarlike tails. Some homalopsines have tails that are only slightly

compressed, whereas natricines and other homalopsines have tails similar in size and shape to

many terrestrial species (Heatwole 1999).

Sidewinding is often considered to be a specialized mode of locomotion (Jayne 1988).

However, it occurs in a surprisingly wide diversity of taxa including booid and colubroid snakes

(Gans and Mendelssohn 1972). Jayne (1988), in a study of snake locomotion, concluded that the

tails of snakes are unlikely able to produce the movements and forces necessary for sidewinding

and thus are likely contributing little during this form of locomotion.

Caudal Luring

Several snake species within the Boidae, Viperidae, Elapidae, and Colubridae use caudal

luring to attract insectivorous prey by wiggling the often contrastingly colored tail tip (Sazima

and Puorto 1993; Greene 1997). These taxa include Boa constrictor (Radcliffe, Chiszar, and

Smith 1980), M~orelia viridis (Murphy, Carpenter, and Gillingham 1978), Callosela~sma

rhodostoma (Schuett 1984; Daltry, Wuster, and Thorpe 1998), Cera~stes vipera (Heatwole and

Davison 1976), Daboia russelii (Henderson 1970), Agkistrodon (Neill 1948; Allen 1949;

Wharton 1960; Carpenter and Gillingham 1990), Crotalus (Kauffeld 1943), Hypnale (Whitaker

and Captain 2004), Sistrurus (Jackson and Martin 1980; Rabatsky and Farrell 1996), Blesibliespr\i\

(Greene and Campbell 1972), Bothrops (Sazima 1991), Acanthophis (Carpenter, Murphy and

Carpenter 1978; Chiszar, Boyer, Lee, Murphy, and Radcliffe 1990), Alsophis portoricensis (Leal









and Thomas 1994), Pantherophis obsolete (Tiebout 1997), Tropidodryas striateceps (Sazima

and Puorto 1993), and Ma'~daga;scarophis (W. Love pers. comm.). Caudal luring has been

reported primarily in juveniles (80% of the species known to caudal lure) to attract small,

visually oriented prey such as anurans and lizards into striking range (Neill 1960; Parellada and

Santos 2002). Cessation of this behavior often occurs within the first year or two with an

ontogenetic shift in diet towards larger endothermic prey items such as mammals and birds (Neill

1960; Jackson and Martin 1980; Daltry et al. 1998). However, in some species such as

Acanthophis antarcticus (Carpenter et al. 1978), Bothriopsis bilineata (Greene and Campbell

1972), Cera~stes viper (Heatwole and Davison 1976), and Sistrurus miliarius (Jackson and

Martin 1980), the behavior persists into adulthood. Persistence of caudal luring in these species

has been attributed to the importance of insectivorous prey items in their adult diets (Jackson and

Martin 1980).

Defense

Snakes exhibit the most elaborate antipredator behaviors among reptiles (Greene 1988).

These behaviors range from generalized to specialized, and many involve the tail. Postcloacal

scent glands are found in all snake species and exude an offensive odor (Whiting 1969). These

noxious secretions likely enhance the effectiveness of cloacal discharge (fecal material) in

deterring predators (Greene 1988).

Defensive tail displays are widespread in snakes; Greene (1973) identified 73 snake

species in at least six families known to perform unusually conspicuous tail displays.

Characteristics of these displays include elevating (e.g., Eryx and Charina), tightly coiling (e.g.,

Diadophis and Farancia),~~FFF~~~FF~~~FF or waving (e.g., M~icrurus and M~icruroides) the tail, which may be

long or blunt, and brightly colored or drab. When threatened, Rhadinaea decorate often hides its

head beneath coils while raising and wriggling its tail (Campbell 1998), whereas Oligodon









albocinctus and Sinomicrurus macclellandil~~~~~1111~~~~ flatten their bodies and curl up the end of their tails

(Whitaker and Captain 2004). Calliophis melan2urus and C. nigrescens raise and coil their tails

when disturbed (Whitaker and Captain 2004). Chilorhinopus butler and C. gerardi hide their

heads within coils and wave their tails in the air in defense (Spawls, Howell, Drewes, and Ashe

2002).

Tail length and thickness correlate with tail injury rate and defensive behaviors in the

genera Eryx and Gongylophis (Greene 1973; R. Saj dak pers. comm.). Species exhibiting tail

displays (e.g., E. jaculus, E. johnii, E. miliaris, and E. Qtatricus) have longer and thicker tails,

small heads, and a relatively high frequency of tail injury. However, species that rely on a biting

defense instead of tail displays (e.g., G. colubrinus, G. conicus, and possibly E. somalicus) have

shorter, thinner tails, larger and more distinct heads, and a lower frequency of tail injury. These

displays likely disorient potential predators and, in many species, serve as a decoy to divert

attack from the head to the tail (Greene 1973).

Many species have tails with specialized external morphological characters used for

defense. Several terrestrial and fossorial species (e.g., Calrphophis, Contia tenuis, FaranciaF~~FFF~~~FF~~~FF

Oligodon afJinis, and several typhlopids) possess a tail spine, which is pressed against an attacker

(Leonard and Stebbins 1999; Whitaker and Captain 2004). Juvenile Farncia abacura have also

been observed using this tail spine to impale or "pop" tadpoles normally too large to be

swallowed (Rossi 1992: 97). Uropeltids have enlarged, reinforced tail tips with specialized scale

morphology that collect dirt to form protective plugs while burrowing (Gans 1976; Gans and

Baic 1977). Cutaneous photoreception, or dermal light sense, in the tail of the sea snake

Aipysurus laevis aides in concealment from visually oriented predators (Zimmerman and

Heatwole 1990). These sea snakes often hide in clumps of coral and use cutaneous










photoreception to avoid having their tails exposed to light, and thus vulnerable, during the

daytime.

The genera Crotalus and Sistrurus form the monophyletic group of approximately 31

species of New World pitvipers characterized by the presence of a rattle (Greene 1988). The

rattle is associated with a suite of anatomical, physiological, and behavioral specializations and is

generally accepted as being used primarily in defensive signaling (see Moon 2001, for summary

of rattle evolution). However, the evolution of this unique structure remains unresolved (Greene

1988; Moon 2001). A potential behavioral precursor to the rattle is tail vibrating, a defensive

behavior widespread in snakes where the tail is rapidly vibrated against loose surface debris to

produce a buzzing sound (Greene 1988).

Caudal pseudautotomy- occurs in several snake species as a defensive strategy (Savage and

Slowinski 1996). However, this behavior has only been documented in snakes with relatively

long tails and is thought to be rare (Arnold 1988; Marco 2002). Enulius, Scaphiodontophis and

Urotheca possess morphological specializations that likely facilitate pseudautotomy such as

long, fragile tails with thick bases (Savage and Slowinski 1996). However, high incidence of tail

loss has also been observed in several other typically long-tailed genera such as Alsophis (Seidel

and Franz 1994), Amphiesma and Boiga (Whitaker and Captain 2004), Coluber (Marco 2002),

Coniophanes, Rhadinaea, Sibynophis and Thamnnophis (Jayne and Bennett 1989; see Mendelson

1991, for review), Dendrophidion, (Duellman 1979), Drymobius (Mendelson 1991), Gastropyxis

(as Hapsidophrys), Grayia and M~ehelya (Spawls et al. 2002), Nerodia (King 1987), Natriceteres

and Psamnmophis (Broadley 1987), Pliocercus (Smith and Chizar 1996), and Xenochrophis

(Ananeva and Orlov 1994), although these genera do not appear to possess the morphological









specializations for tail loss present in Enulius, Scaphiodontophis and Grotheca (Savage and

Slowinski 1996).

While both intervertebral pseudautotomy- and intravertebral autotomy occur in lizards, only

intervertebral pseudautotomy- is known to occur in snakes (Arnold 1988; see Savage and

Slowinski 1996, for a review of terminology). Unlike many lizard species, pseudautotomy in

snakes does not appear to be under neural control (Arnold 1988) and requires physical resistance,

which is often facilitated by twisting or rotating the body in one direction until the tail snaps off

(Savage and Slowinski 1996; Marco 2002). However, reflex action often allows the segment of

lost tail to continue thrashing, thus likely distracting the predator and allowing the snake to

escape (Marco 2002; Savage 2002). Although caudal autotomy is a lepidosaurian

synapomorphy, the ability to regenerate any portion of the tail has been lost in all snake species

(Pough et al. 2001).

Morphology and Reproduction

Peters (1964) defines the snake tail as the section of the body posterior to the cloaca. In

males, the hemipenes and associated retractor muscles are located in the tail, while the testes

(and ovaries in females) are located within the body cavity. In both sexes of all species, the tail

also contains a pair of scent glands (Whiting 1969). Male snakes typically use their tails during

courtship to gain access to the female's cloaca by performing a tail-search copulatory attempt

(TSCA) until the cloacas meet (Gillingham, Carpenter, and Murphy 1983). This TSCA is often

preceded by various tail movements by the female, such as tail-whipping and tail-waving

(Carpenter and Ferguson 1977) and is followed by intromission and coitus (Gillingham et al.

1983). In most boids, courtship is facilitated by the use of two claw-like spurs, which are located

on either side of the male' s cloaca. Male Madagascan tree boas (Sanzinia madaga;scariensis) use

their spurs in combat bouts with other male conspecifics (Carpenter, Murphy, and Mitchell









1978). During these bouts, the males entwine their tails and vigorously flex the erected spurs

against the scales of the opponent while hanging from branches. Larger male garter snakes

(Thamnnophis sirtalis) and European grass snakes (Natrix natrix) usually achieve more matings

apparently because they can physically displace the tails of smaller rival males by tail wrestling

(Luiselli 1996; Shine et al. 2000). Tail wrestling in this context may be widespread in snake

species that display "mating balls," or mating aggregations where multiple males simultaneously

attempt copulation (Shine et al. 2000: F9).

Sexual Dimorphism and Ontogenetic Shifts

Although RTL is a stable character, sexual dimorphism in snake tail length is common

(Klauber 1943). Males typically have longer, more attenuate tails than females (King 1989;

Shine 1993). However, the degree of these differences varies interspecifically (King 1989; Shine

1993). King (1989) proposed three hypotheses that could help explain the existence of sexual

dimorphism in tail length among snakes: (1) morphological constraint (the hemipenes and

retractor muscles of males are located in the tail); (2) female reproductive output (a more

posterior cloaca might increase body cavity volume and allow increased fecundity); and (3) male

mating ability (longer tails in males may be advantageous during courtship). These predictions

were tested using 56 colubrid genera, and the results supported both the morphological constraint

and female reproductive output hypotheses (King 1989). Additional studies involving tail

wrestling in several natricine species provide support for the male mating ability hypothesis

(Luiselli 1996; Shine et al. 2000). However, selection could act on more than one of these

hypotheses simultaneously.

Whereas RTL is stable among individuals of the same species, sex, and age, RTL is

nonetheless ontogenetically variable in most snake taxa (Klauber 1943). Therefore, the tail

proportions allometryy) of many species change as they grow. The majority of snake species









have relative tail-lengths that increase as they age (Klauber 1943); however, some species (e.g.,

Pituophis and Lampropeltis) have relative tail-lengths that become shorter (Klauber 1943).

Furthermore, some snake species demonstrate ontogenetic shifts in macrohabitat use, with one

age class (e.g., juveniles or adults) found more frequently utilizing arboreal macrohabitats than

another (Martins and Oliveira 1999). This behavioral shift has been well documented in several

large boid species such as Python sebae (Spawls et al. 2002) and Boa constrictor (Campbell

1998), some colubrids such as Pseustes poecilonotus (Boos 2001), some viperids such as

Bothrops jararaca (Sazima 1992; Martins et al. 2001), and some elapids such as Ophiophagus

hannah (R. Whitaker pers. comm.). Usually, juveniles are more arboreal than adults in those

snake species known to exhibit behavioral shifts in habitat use. However, the opposite trend has

been observed in Leptophis depressirostris (Nickerson, Saj dak, Henderson, and Ketcham 1978)

and in some populations of Boa constrictor (Campbell 1998). Ontogenetic shifts involving

arboreal macrohabitat use likely occur in many snake clades, but more detailed natural history

information is needed to know the extent to which this is the case.









CHAPTER 2
MATERIALS AND METHODS

Categorizing Climbing in Snakes: Gravitational Habitat

Snakes occupy a wide variety of habitat types, and it is often useful to divide habitat usage

into generalized categories when discussing snake community assemblages. Perhaps the most

widely utilized categories for snake habitat use include fossorial, terrestrial, aquatic, semiaquatic,

arboreal, and semiarboreal, or some subset of these (e.g., Johnson 1955; Shine 1983; Guyer and

Donnelly 1990; Dalrymple, Bernardino, Steiner, and Nodell 1991; Lindell 1994; Conant and

Collins 1998; Vidal, Kindl, Wong, and Hedges 2000).

However, as correctly noted by Johnson (1955), these categories can become inadequate

when performing interspecific ecomorphological studies because patterns in habitat usage are

often obscured. This is because these categories describe the habitat per se, and not the

behaviors snakes exhibit while utilizing these habitats. For example, watersnakes in the genus

Nerodia typically live and obtain food near water (Conant and Collins 1998). Consequently,

they are usually categorized as aquatic or semiaquatic (e.g., Vidal et al. 2000). However, many

Nerodia species spend a significant amount of time out of the water and off the ground climbing

among branches to bask (Conant and Collins 1998). In fact, during parts of the year, some

Nerodia species can spend as much or more time above the ground in shrubs and trees than

Cohiber constrictor, a snake typically categorized as semiarboreal (Mushinsky, Hebrard, and

Walley 1980; Plummer and Congdon 1994). Separating Nerodia and Cohiber into distinct

ecological categories (aquatic or semiaquatic versus semiarboreal) obscures these similarities in

habitat usage. Similarly, categorizing both Nerodia and sea snakes such as the entirely pelagic

Pelamnis platurus as aquatic is equally misleading. But in this case it is because the grouping

suggests strong similarity in habitat usage when in actuality there is very little. Therefore, a









different system was needed to categorize climbing in snakes that combines habitat use with

behavior, which I call gravitational habitat.

In order to categorize the amount of climbing, and therefore the amount of gravitational

stress imposed, I divided gravitational habitat into four categories: stenotopically arboreal,

eurytopically arboreal/terrestrial, stenotopically terrestrial, and stenotopically aquatic. These

categories were based on adult habitat use information compiled from the literature and from

personal observations. I define stenotopically arboreal species as those primarily living in

arboreal habitats. These species are rarely found on the ground and should experience the

greatest gravitational stress. Eurytopically arboreal/terrestrial species are often found on the

ground, but regularly climb for reasons including hunting, escaping predators, and

thermoregulation (e.g., M~asticophis and Coluber). The stenotopically terrestrial category

includes both terrestrial and fossorial species that rarely if ever climb and, thus, these species

should experience relatively little gravitational stress. Some characteristically terrestrial species,

or populations of species, do climb occasionally (e.g., Bitis arietans, B. armata, Bothrops asper,

Crotalus horridus, and Jhamnnophis sirtalis). However, I consider this behavior atypical and do

not consider them eurytopically arboreal/terrestrial. I define stenotopically aquatic species as

those primarily found in water. They typically exhibit one or more morphological

specializations accepted as adaptations to aquatic habitats such as a flattened or oarlike tail, more

dorsally positioned eyes and nostrils, salt excreting glands, and valvular nostrils (Heatwole,

1999). Laticaudines (genus Laticauda), the homalopsine snake Enhydris enhydris, and Helicops

angulatus are included within this category because they are clearly adapted to an aquatic

lifestyle and are usually found in water even though they occasionally soj ourn onto land.

Stenotopically aquatic species should experience the least gravitational stress because the









surrounding water column acts as an antigravityy suit" (Lillywhite 1993: 561). Importantly, all

categorical placements were based entirely on the ecology and behavior of each species and not

on morphology.

Habitat use and other natural history information for each species were collated from the

literature. Whereas many of these sources were various journal publications, a large amount of

this information came from the following: Africa (Broadley and Cock 1975; Broadley 1983;

Branch 1998; Schmidt and Noble 1998; Spawls et al. 2002); Australia (Shine 1995); Central

America (Campbell 1998; Savage 2002); India (Whitaker and Captain 2004); Madagascar (Glaw

and Vences 1994; Henkel and Schmidt 2000); North America (Smith and Brodie 1982; Conant

and Collins 1998); South America (Murphy 1997; Boos 2001; Duellman 2005); and the West

Indies (Schwartz and Henderson 1991).

However, the natural history of some of the species included in this study is incompletely

known, thereby making categorization difficult. For example, Alsophis antillensis and closely

related Antillophis parvifr~ons are West Indian racers with relatively long tails (27.8% and 31.1%

respectively). However, whether these species climb or not is unknown. Several other species

ofAlsophis are known to climb well, but because this information is not available for A.

antillensis and A. parvifr~ons, I chose not to assume they climbed and thus categorized them as

stenotopically terrestrial even though they may actually climb to some extent.

In order to avoid the confounding effects of sexual dimorphism and ontogenetic shifts in

allometry and habitat use, only adult females were used for all aspects of this study except for the

frequency distribution (see below). A snake was determined to be an adult if its total length was

within, or very near, the published adult range for that species. Tail length data were collected

from the literature, museum specimens, and live snakes (see Table 2-1 for species used in this










study and sources). Museum specimens were acquired from the Florida Museum of Natural

History (FLMNH) and National Museum of Natural History, Smithsonian (USNM). For

museum specimens, sex was determined by subcaudal incision. Species in which females are

known to possess well-developed hemipenis-like structures (e.g., Pseudoficimia and Gyalopium;

Hardy 1972; Smith and Brodie 1982) were not used in this study. Some Bothrops insularis

females have hemipenis-like structures (Hoge, Belluomini, Schreiber, and Penha 1961), and this

species was included in the study under the assumption that length data reported by Klauber

(1943) were from functional females. Klauber (1943) states that differential shrinkage between

the tail and body can occur as a result of preservation, but that the amount is insignificant as long

as the preservation methods are consistent. Live snakes were measured at Glades Herp Inc.,

Florida, and were probed to determine sex.

Snout-vent length (SVL) was measured from the tip of the snout to the posterior edge of

the anal scale. Tail length was measured from the posterior edge of the anal scale to the tip of

the tail, and only snakes with complete tails were used in this study. Snout-vent length and tail

length were measured using either a meter stick (a 1.0 mm) or a caliper (A 0.1 mm). A string

was used to follow the body contours of live snakes and rigid specimens and was subsequently

measured with a meter stick. Length measurements were repeated on individuals up to ten times

when possible and then averaged. I used mean SVL and tail lengths when those data were

available from the literature or from multiple specimens, but otherwise I used data representing

single specimens. To verify the stability of relative tail-length (RTL), I gathered large samples of

data from the literature on the total variation in adult female RTL for 26 species (5 families, 9

subfamilies, 26 genera) and found the variation to be small (mean variation & SE, 1% + 0.3%,









95% C.I.) (Table 2-2). Furthermore, the RTL of museum specimens was checked to ensure the

data were consistent with published adult lengths for that species.

Analyses

Frequency Distribution

Frequency distributions for total lengths were produced for each of the four gravitational

habitat categories: stenotopically arboreal (n=75), eurytopically arboreal/terrestrial (n=74),

stenotopically terrestrial (n=94), and stenotopically aquatic (n=35)(Table A-1). All categories

comprise both Old and New World species. Data used for the frequency distributions were

based on raw total lengths. Stenotopically arboreal species should spend the most time in

arboreal habitats and, thus, were analyzed separately from eurytopically arboreal/terrestrial

species. Furthermore, eurytopically arboreal/terrestrial species might not climb enough to have

their lengths affected by such limitations.

Because there is much interspecific variation among snakes regarding which sex is longer

(Shine 1993), I used length measurements representing the longest adult total lengths regardless

of sex. Maximum total lengths for each species were used when possible unless stated by the

authors that the maximum size is extremely unusual. In that case, the next largest measurement

was used. Maximum adult total length was used because whereas juvenile length is likely also

under related selective pressures, they differ from adults in two important ways: (1) juveniles

may utilize a different macrohabitat until adult size is reached or until a certain length is attained;

and (2) juveniles are shorter and are thus less affected by gravitational stresses on blood

circulation.

Gravitational Habitat and Total Body Length

To test for differences in mean total length among the four categories, I used an analysis of

variance (ANOVA). To meet the assumption of normality, the length data were loglo









transformed. Following the ANOVA, I compared the means among the four categories, using

Tukey's Honestly Significant Difference test. Coefficients of variation (CV) were calculated and

used to compare variance among the four categories.

Gravitational Habitat and Tail Length

I tested for differences in tail length among 227 species within 138 genera, 12 families, and

18 subfamilies using an analysis of covariance (ANCOVA) with total length treated as a

covariate. Differences in tail length corrected for body length (corrected tail-length) were

inferred from comparisons of the Y-intercepts in the ANCOVA. Data in this and all subsequent

analyses were collected independently from those used in the frequency distribution described

above. Data were loglo transformed to meet the assumptions of linearity and normality before

performing the ANCOVA analyses. All statistical analyses were performed using the statistical

program package R (R Development Core Team 2006).

Constructing the Phylogeny

Phylogenies constructed by Lawson, Slowinski, Crother, and Burbrink (2005), Lawson,

Slowinski, and Burbrink (2004), and Vidal et al. (2000) were combined and used as a base

phylogeny, to which additional phylogenies from detailed studies were added (Fig. 2-1). When

possible, the trees chosen were those recommended by the authors. However, these were often

consensus trees and, as such, occasionally contained polytomies. Polytomy resolution was often

achieved by either using another tree in the same study with high bootstrap support, or by using a

tree from a different study. For different trees containing similar species but with conflicting

relationships, I used the tree with the strongest bootstrap support and, when available, the more

extensive taxon sampling. However, phylogenetic relationships for some clades remain

unresolved (e.g., Dipsa~s, Langaha, Leptodeira, Prosymna, and Psamnmophis), and uncertain

relationships were in the end reflected as polytomies. Many of these polytomies contain










congeners categorized within the same gravitational habitat and, thus, are not likely to affect the

results (Fig 2-1). In some cases, I used species thought closely related to those in the composite

tree to replace species for which no specimens or length data were available (e.g., Hydrops

triangularis was replaced by H. caesurus; Xenoxybelis argenteus was replaced by X.

boulengeri) .

The following studies were used to add taxa to the base phylogeny (Fig. 2-1). The

scolecophidian clade was replaced using the phylogeny of Vidal and Hedges (2002). The

phylogeny of Lenk, Kalyabina, Wink, and Joger (2001) was used for placement of true vipers

(Atheris, Bitis, Causus, Macrovipera and Vipera). The phylogeny of Malhotra and Thorpe

(2004) was used for Asian pitviper (Protobothrops, Trimeresurus, and Tropid'olaemus)

placement. The phylogeny of Parkinson, Campbell, and Chippindale (2002, in Gutberlet and

Harvey 2004) was used for New World pitviper relationships, and the phylogeny of Murphy, Fu,

Lathrop, Feltham, and Kovac (2002) was used for relationships within Crotalus. The phylogeny

of Nagy, Joger, Wink, Glaw, and Vences (2003) was used for pseudoxyrhophine relationships.

The phylogeny of Kelly, Barker, and Villet (2003) was used for Dasypeltis and Dend'relaphis

placement and psammophine relationships. The phylogeny of Alfaro and Arnold (2001) was

used for thamnophiine relationships, and the phylogeny of De Quieroz, Lawson, and Lemos-

Espinal (2002) was used for relationships within Hzamnophis. The phylogeny of Lawson et al.

(2005) was used for placement of the Homalopsinae, as well as the following genera: Ahaetulla,

Calrphophis, Crotaphopeltis, Dinodon, Dipsad'oboa, Drymarchon, Gonyosoma, Grayia,

Lycophidion, M~asticophis, M~ehelya Natriceteres, Opheodrys, Prosymna, Philothamnus,

Phyllorynchus, Pseudaspis, Sonora, Spilotes, Telescopus, Hzelotornis, Thrasops, and

Xenochrophis. The phylogeny of Utiger et al. (2002) was used for the placement of Arizona,










Elaphe, Lampropeltis, Panthero~~PP~~Pp his,~PP~~PP and Pituophis. The phylogeny of Vidal et al. (2000) was

used for the placement of the following genera: Helicops, Hydrodyna~stes, Hydrops, Ialtris,

Oxyrhopus, and Pseudoeryx. The phylogeny of Kraus and Brown (1998) was used for the

placement of the genera Chilomeniscus and Dispholidus. Subfamily designation followed that of

the EMBL Reptile Database (Uetz 2006). However, recent phylogenetic analyses by Lawson et

al. (2005) provide further evidence that considerable changes in these classifications need to

occur in order to recognize monophyletic groups.

Relative Tail-Length, Gravitational Habitat, and Phylogeny

Interspecific comparative analyses can be confounded by pseudoreplication caused by

phylogenetic relatedness among species samples (Price 1997). However, these effects can be

partially resolved using independent contrasts (Felsenstein 1985). I assessed the relationship

between RTL and macrohabitat use using the program Mesquite version 1.06 (Maddison and

Maddison 2005). Because there is no comprehensive phylogeny available for snakes, I

constructed an estimate of relatedness by combining data from 14 recent molecular snake

phylogenies into a composite tree containing 227 species within 138 genera, 12 families, and 18

subfamilies (Fig. 2-1). These data represent almost all snake families and subfamilies (see

below). Branch lengths could not be estimated because the final phylogenetic tree was

composed from a variety of sources. Independent contrasts were therefore generated with branch

lengths assigned as either equal or with the assumption that the age of a clade is proportional to

the number of species it contains using Grafen's branch lengths (Grafen 1989; Maddison and

Maddison 2005). However, plots of standardized contrasts against the variance of untransformed

contrasts showed strong significant correlations for Grafen's, but not equal, branch lengths. As

significant correlations violate a key assumption of independent contrast analysis (Diaz-Uriarte

and Garland 1996), I used equal branch lengths only in the final analysis. Contrasts were









calculated between nodes for macrohabitat use and loglo transformed RTL, and relationships

were examined between the variables by calculating regressions on these standardized contrasts

using least-squared change optimization parsimony (Garland, Harvey, and Ives 1992; Grafen

1992).










































































i


Table 2-1. Taxa and associated data sources for 227 snake species included in this study.
Family Subfamily Species Source


Acrochordidae


Acrochordus granulatus


Lillywhite unpubl. data


Aniliidae


Anilius scytale


UF62496


Atractaspididae


Aparallactinae


Amblyodipsas polylepis
Amblyodipsas ventrimaculatus
Aparallactus capensis
Aparallactus guentheri
Aparallactus lunulatus
Xenocalamus mechowii
Xenocalamus sabiensis
Atractaspis bibronii


Acrantophis dumerili
Boa constrictor
Corallus caninus
Corallus ruschenbergerii
Epicrates striatus


Charina trivirgata
Gongylophis muelleri


Morelia viridis
Python molurus
Python regius
Python sebae


Duberria lutrix

Grayia smythii
Lamprophis fuliginosus
Lamprophis lineatus
Lycodonomorphus leleupi
Lycophidion capense
Lycophidion variegatum
Mehelya capensis
Pseudaspis cana


Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975


Sheehy unpubl. data
Lillywhite unpubl. data
Lillywhite unpubl. data
Sheehy unpubl. data
Sheehy unpubl. data


Klauber 1943
Sheehy unpubl. data


Sheehy unpubl. data
Lillywhite unpubl. data
Lillywhite unpubl. data
Broadley and Cock 1975


Broadley and Cock 1975
UF57047
Broadley and Cock 1975
Klauber 1943
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
Schmidt and Noble 1998
Broadley and Cock 1975


Boidae


Boinae


Erycinae




Pythoninae


Colubridae


Boodontinae












Species
Ahaetulla nasutta
Arizona elegans
Boiga blandingli
Boiga cynod'on
Chilonveniscus stranzineus
Chironius carinatus
Coluber constrictor
Crotaphopeltis hotanyboeia
Dasypeltis fasciata
Dasypeltis medici
Dasypeltis scabra
Dendrelaphis pictus
Dinodon rufozonatum
Dinodon senricarinatun?
Dipsadoboa aulica
Dipsadoboa unicolor
Dispholidus typus
Drymarchon corals
Elaphe radiata
Gastropyxis snzaragdina
Gonyosonza oxycephalum
Lanmpropeltis getula
Lampropeltis triangulum
Lycodon laoensis
Mlasticophis flagellum
M~asticophis lateralis
\ ,u,r rn nu\ boddaerti
Opheodrys aestivus
Oxybelis f lgidus
Oxybelis aeneus
Pantherophis obsoleta
Philothamnus dorsalis
Philothamnus irregularis
Philothamnus semivariegatus
Philothamnus angolensis
Philothamnus hoplogaster
Phyllorynchus brown
Phyllorvnchus decurtatus


Source
Lillywhite unpubl. data
UF48734
UF69248
UF55217
Klauber 1943
UF34779
UF47855
Broadley and Cock 1975
UF61301
Broadley and Cock 1975
Broadley and Cock 1975
Lillywhite unpubl. data
Lillywhite unpubl. data
UF24089
Broadley and Cock 1975
Schmidt and Noble 1998
Broadley and Cock 1975
UF83818
Lillywhite unpubl. data
UF52721
Lillywhite unpubl. data
Klauber 1943
UF68828
UF69087
Klauber 1943
Klauber 1943
UF91619
UF43623
UF56396
Sheehy unpubl. data
Lillywhite unpubl. data
Schmidt and Noble 1998
Broadley and Cock 1975
Broadley and Cock 1975
UF53342
Broadley and Cock 1975
Klauber 1943
Klauber 1943


I I


Table 2-1. Continued
Family
Colubridae Col


Subfamily
ubrinae





Source
Klauber 1943
UFl32981
Schmidt and Noble 1998
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
Klauber 1943
UF56320
Lillywhite unpubl. data
Klauber 1943
Lillywhite unpubl. data
Stafford 2004
UF40711
Broadley and Cock 1975
Broadley and Cock 1975
UF52696


Klauber 1943
Myers 2003
Myers 2003
Myers 2003
Myers 2003
UF43842
UF56868
Harvey unpubl. data
Harvey unpubl. data
Harvey unpubl. data
Harvey unpubl. data
Harvey unpubl. data
Klauber 1943
UF11284
Duellman 1958
Duellman 1958
Duellman 1958
Duellman 1958
Duellman 1958
Duellman 1958
Duellman 1958





Table 2-1. Continued
Family
Colubridae Col


Subfamily
ubrinae


Species
Pitatophis catenifer
Pitatophis melanoletecis
Prosymna ambigta
Prosymna bivittata
Prosymna stithlmannii
Prosymna sundevallii
Sonora michoacanensis
Spilotes pullatus
Stilosoma extenuatum
Tantilla planiceps
Tantilla relicta
Tantilla hendersoni
Tantilla melanocephala
Telescopus semiannulants
Thelotornis capensis
Thrasops jacksonii


Adelphicos quadrivirgatus
Atractics depressiocellus
Atractics hostilitractus
Atractics clarki
Atractics darienensis
Carphophis amoenus
Diadophis princtatus
Dipsas catesbyi
Dipsas indica
Dipsas pentana
Dipsas boettgeri
Dipsas chaparensis
Hypsiglena torquata
Imantodes cenchoa
Leptodeira annulata
Leptodeira septentrionalis
Leptodeira frenata
Leptodeira bakeri
Leptodeira ; i;, IIf,;,a Aite
Leptodeira maculata
Leptodeira punctata


Dipsadinae





_ _


Table 2-1. Continued
Family
Colubridae Col


Subfamily
ubrinae


Species
Taeniophalhts brevirostris
Thamnodynastes pallidus
Tretanorhimes variabilis


Source
Schargel et al. 2005
UF88631
UF119241


Homalopsinae


Cerberus rimchops
Enhydris enhydris


Natricinae


Natriciteres olivacea
Natrix natrix
Nerodia fasciata
Nerodia floridana
Nerodia sipedon
Nerodia taxispilota
Nerodia rhombifer
Regina alleni
Regina septemvittata
Seminatrix pygaea
Storeria dekavi
Thamnophis hammondii
Thamnophis ordinoides
Thamnophis sirtalis
Bjlrginia valeriae
Xenochrophis flavipunctatus


Psammophis biseriatcs
Psammophis princtulatus
Psammophis sibilans
Psammophis angolensis
Psammophis brevirostris
Psammophis crucifer
Psammophis jallae
Psammophylax tritaeniants
Rhamphiophis oxvrhvnchus
Rhamphiophis rostrants

Dromicodrvas bernieri
Ithvcvphus miniatcs


Lillywhite unpubl. data
Lillywhite unpubl. data

Broadley and Cock 1975
UF116535
UF41022
UF109763
UF63519
Lillywhite unpubl. data
Lillywhite unpubl. data
UF99998
UF68506
UF42444
Lillywhite unpubl. data
Klauber 1943
UF57050
UF42468
UF54122
Lillywhite unpubl. data

UF52841
UF52832
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
UF59339
Broadley and Cock 1975

USNM 499459
USNM 149366


Psammophiinae














Pseudoxyrhophiinae


































































Cvlindrophis rutlis


UF51669


_ I __


Table 2-1. Continued
Family
Colubridae Psel


Subfamily
udoxyrhophiinae


Species
Langaha pseudoalluaudi
Langaha nmadagascariensis
Langaha alluaudi
Leioheterodon nmadagascariensis
Leioheterodon nmodestus
Mlicropisthodon ochraceus


Alsophis antillensis


Alsophis portoricensis
Alsophis vudit
Antillophis parvifrons
Arrhyton Jitnereun2
Clelia clelia
Darlingtonia haetiana
Drepanoides anonzalus
Erythrolanmprus aesculapti
Farancia abacura
Helicops angulatus
Heterodon nasicus
Heterodon platirhinos
Hydrodynastes gi gas
Hydrops caesurus
Hypsirhynchus ferox
Ialtris dorsalis
Liophis typhlus
Oxvrhopus petola
Philodryas baroni
Pseudoboa coronata
Pseudoervx plicatilis
Siphlophis cervinus
Uronzacer catesby
Uronzacer frenatus
Uronzacer oxyrhynchus
Xenodon severus
Xenoxybelis boulengeri


Source
Domergue 1988
Guibe 1949
Guibe 1949
Sheehy unpubl. data
Sheehy unpubl. data
Domergue 1993


Xenodontinae


UF53983
UF44287
UF57211
UF99946
UF23131
UF113638
UF34331
UF5568 1
Dixon and Soini 1986
UF115664
Lillywhite unpubl. data
UF88643
UF56015
UF44172
UF20576
Scrocchi et al. 2005
UF60786
UF85307
UF32985
UF62069
Lillywhite unpubl. data
UF117617
UF86909
UF95759
UF23158
UF23170
UF23172
UF87926
Duellman 2005


Cvlindrophiidae















































Leptotyphlophidae


I


Table 2-1. Continued
Family
Elapidae ElaI


Subfamily
pinae


Species
Aspidelaps scutatus
Bungants fasciatus
Dendroaspis o, irg,l0,llps
Dendroaspis polylepis
Dendroaspis viridis
Elapsoidea gatentheri
Elapsoidea semiannulata
Elapsoidea sundevallii
Elapsoidea nigra
Hemachatics haemachants
Mlicntrats f lvists
Naja melanolerica
Naja haje
Naja mossambica
Naja atra
Naja kaouthia


Source
Broadley and Cock 1975
Sheehy unpubl. data
Broadley and Cock 1975
Broadley and Cock 1975
Sheehy unpubl. data
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
Klauber 1943
Broadley and Cock 1975
Lillywhite unpubl. data
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
Lillywhite unpubl. data
Lillywhite unpubl. data


Tropidophiidae



Typhlopidae



Viperidae


Tropidophiinae


Aipysunts laevis
Emydocephahts annulants
Hydrophis melanocephahts
Laticateda cohibrina
Notechis scretants
Pelamis plattets


Leptotyphlops alfredschmidti
Leptotyphlops macros


Tropidophis morenoi
Tropidophis haetiames

Rhinotyphlops episcopus
Xenotyphlops grandidieri


Hedges et al. 2001
Sheehy unpubl. data


Franzen and Wallach 2002
Wallach and Ineich 1996


Lillywhite unpubl. data
Klauber 1943
Lillywhite unpubl. data
Sheehy unpubl. data
Lillywhite unpubl. data


Crotalinae


Agkistrodon contortrix
Agkistrodon mokeson
Agkistrodon piscivorus
Atropoides nummifer
Bothriechis schlegelii


Hydrophiinae


Lillywhite unpubl. data
Lillywhite unpubl. data
Guinea 1981
Guinea 1981
Lillywhite unpubl. data
Lillywhite unpubl. data

Lehr et al. 2002
Broadley and Wallach 1996





Source
Klauber 1943
Lillywhite unpubl. data
Lillywhite unpubl. data
Lillywhite unpubl. data
Boundy and Balgooyen 1988
Herrmann et al. 2004
Klauber 1943
Lillywhite unpubl. data
Lillywhite unpubl. data
Sheehy unpubl. data





Table 2-1. Continued
Family
Viperidae Cro


Subfamily
talinae


Species
Bothrops insularis
Crotalus atrox
Crotalus horridus
Crotalus ruber
Crotalus cerastes
Protobothrops cornutus
Trimeresurus gramineus
Trimeresurus purpureomaculatus
Trimeresurus albolabris
Tropidolaemus wagleri


Viperinae


Atheris squamigera
Atheris chlorechis
Bitis arietans
Bitis atrops
Bitis caudalis
Bitis gabonica
Bitis nasicornis
Causus defil~ipi
Causus rhombeatus
Mlacrovipera mauritanica
Vipera xanthina


Klauber 1943
Sheehy unpubl. data
Broadley and Cock 1975
Broadley and Cock 1975
Broadley and Cock 1975
Lillywhite unpubl. data
Lillywhite unpubl. data
Broadley and Cock 1975
Broadley and Cock 1975
Sheehy unpubl. data
Lillywhite unpubl. data


Xenoveltidae


Xenoveltis unicolor


UF50268












Table 2-2. Total variation in relative tail-lengths (RTL) of 26 species of snakes. The data are
from adult females only and include species typically categorized as fossorial,
terrestrial, semiarboreal and arboreal. In species where the RTL becomes shorter
with age, the absolute value of the difference is shown. For sources (Sc), 1 =
Broadley 1959; 2 = Klauber 1943.


_


mm
nRTL
13 0.06
57 0.11
10 0.10
34 0.11
5 0.13
25 0.10
70 0.11
249 0.11
34 0.19
148 0.09
23 0.12
23 0.18
57 0.15
15 0.15
62 0.11
183 0.13
87 0.11
30 0.12
153 0.19
29 0.05
32 0.09
94 0.11
16 0.14
25 0.12
27 0.06
22 0.05


max
RTL
0.07
0.12
0.12
0.13
0.16
0.11
0.10
0.10
0.20
0.08
0.12
0.18
0.15
0.18
0.11
0.14
0.13
0.12
0.18
0.06
0.09
0.11
0.14
0.13
0.09
0.07


max -
min
1%
1%
2%
2%
3%
1%
1%
1%
1%
1%
0%
0%
0%
3%
0%
1%
2%
0%
1%
1%
0%
0%
0%
1%
3%
2%


Family
Atractaspididae
Boidae
Colubridae



















Elapidae

Viperidae


Subfamily Species
Aparallactinae Amblyodipsas unicolor
Erveinae Charina trivirgata
Boodontinae Duberria lutrix
Lamprophis fliginosts
Pseridaspis cana
Colubrinae Arizona elegans
Conopsis nasus
Lampropeltis getula
Mlasticophisflagellum
Phydlorhymchus decurtants
Rhinocheilus lecontei
Salvadora grahamiae
Sonora michoacanensis
Tantilla planiceps
Dipsadinae Adelphicos quadrivirgatus
Diadophis princtatus
Geophis nasalis
Hypsiglena torquata
Natricinae Thamnophis hammondii
Elapinae Elapsoidea nigra
Mlicrients nigrocinctus
Crotalinae Bothrops insularis
Trimeresunts gramineus
Viperinae Atheris squamigera
Bitis arietans
Cartsus defilippii











Leptotyphlops macrops
Leptotyphlops alfredschmidti
Rhinotyphlops episcopus
Xenotyphlops grandidied'
Anilius scytale
Tropidophis haetianus
Tropidophis morenoi
Xenopeltis unicolor
Cylindmphis auffus
M~orelia viridia
Python sebae
Python molurus
Python regius
Charina trivingata
Gangylophis muelleni
Acrantophis dumedili
Boa constrictor
Epicrates striatus
Corallus caninus
Corallus ruschenbergeril
Acrochordus granulatus *
Causus defilippli
Causus rhombeatus
M~acrovipera mauritanica
Vipera xanthina
Atheria chlorechis
Atheria squamigera
Bitis arietans
Bitis atrops
Bitis caudalis
Bitis gabonica
Bitis nasicornis
Tropidolaemnus wagled'
Trimneresurus gramineus
Trimeresurus albolabria
Trimeresurus purpureomaculatus
~Protobothrops cornutus
Crotalus cerastes
Crotalus honidus
Crotalus ruber
Crotalus atrox
Agkistrodon contortlix
Agkistrodon piscivorus
Bothriechis schlegelii
Bothrops insularis
Atmopaides nummifer
Enhydris enhydris *
Cerberus rynchops
(Continued on next page)


Figure 2-1. Composite phylogeny of 227 snake taxa. Species in gray boxes are stenotopically
arboreal. Bolded branches denote eurytopically arboreal/terrestrial species. Asterisks
denote stenotopically aquatic species. Bolded species exhibit pseudautotomy.
Remaining unmarked species are stenotopically terrestrial.











Prosymna bivittata
Prosymna sundevallii
Pmosymna ambigua
Prosymna stuhimannii
Dromicodryas hernieri
Dubenia luttix
Leicheterodon madagascadiensis
Leicheterodon modestus
Micropisthodon achraceus
Ithycyphus miniatus
Langaha pseudoalluaudi
Langaha alluaudi
Langaha madagascariensis
Pseudaspis cana
Lamprophis lineatus
Lamprophis fuliginosus
Lycodonomorphus teleupi
Mehetya capensis
Lycophidion capense
Lycophidian variegatum
Rhamphiophis oxyrhynchus
Rhamphiophis rostratus
-Psammaphis jallae
-Psammaphispunctulatus
-Psammophis crucifer
-Psammaphis angolensis
-Psammophis sibilans
-Psammophis tritaeniatus
-Psammaphis biseriatus
-Psammaphis brevirostris
Atractaspis bibronii
Amblyodipsas polylepis
Amblyodipsas ventrimaculatus
Xenocalamus mechowii
Xenocalamus sabiensis
Aparallactus capensis
Aparallactus guentheti
Aparallactus lunulatus
Notechis scutatus
Laticauda colublina *
Alpysurus laevis *
Hydmophis melanocephalus *
Pelamis platuus *
Emydocephalus a~nnulatus *
~Micmmrs fulvius
Hemachatus haemachatus
Naja mossambica
Naja kaouthia
Naja afra
Naja melanoleuca
Naja haje
Aspidelaps scutatus
Elapsoidea guentheti
Elapsoidea semiannulata
Elapsoidea sundevalliii
Elapsoidea nigra
Bungaris fasciatus
Dendreaspis viridis
Dendreaspis angusticeps
Dendreaspis polylepis
(Continued on next page)



Figure 2-1. Continued











Heterodon nasicus
Heterodon platirhinos
Carphophis amoenus
Farancia abacura
Xenachrophis flavipunctatus
Natriciteres alivacea
Natrix nlatrix
Thamnophis sirtalis
Thamnophis hammondii
Thamnophis ordinoides
Storeria dekayi
Virginia valeriae
Regina alleni
Seminatrix pygaea
Regina septemvitrata
Nerodia floridana
Nerodia taxispilota
Nerodia rhambiter
Nerodia sipedonl
Nerodia fasciata
Imantodes canchoa
Hypsiglena torquata
Tretanlorhinus variabilis
Diadophis punctatus
Leptodeira nigrafasciata
Leptodeira maculate
Leptodeira septantrionalis
Leptodeira bakeri
Leptodeira frenata
Leptodeira annulata
Leptodeira punctata
Atractus depressiocellus
Atractus hostilitractus
Atractus darienensis
Atractus clarki
Dipsas peruana
-Dipsas boattgeri
Dipsas catesbyi
-Dipsas chaparensis
-Dipsas indica
Helicops angulatus It
Pseudoeryx plicatilis
Hydrops triangularis
Uromnacer catesbyi
Uromacer frenatus
Uromacer oxyrhynchus
Alsophis anltillenlsis
Arrhyron funereum
Hypsirhynchus ferox
Anltillophis panrvitrals
Darlinlgtonia haerianla
Laltris darsalis
Alsophis partoricansis
Alsophis canltherigerus
Alsophis vudii
Thamnodynastes pallidus
Taeniophallus brevirostris
Xenodon severus
Liophis typhlus
Erythralamprus aesculapil
Philodryas baroni
Xenoxybelis boulengeri
Hydrodynastes gigas
Siphlophis cervinus
Oxyrhopus petola
Drepanoides anomalus
Pseudoboa coronata
Glelia clelia
(Continued on next page)
Figure 2-1. Continued












Graylia smythii
AAhatulla nasuta
Dendrelaphis pictus
M~astigodryas boddaerfi
Chironius carinaturs
Coluber constrictor
Sonora michoacanensis
Tantilla melanocephala
Tantilla pianiceps
Tantilla handersoni
Tantilla relicta
Drymarchon corals
~Masticophis lateralis
~Masticophis flagellum
Spilotes pullatus
Phyllorhynchus brown
Phyllorhynchus decurtatus
Opheodrys aestivus
Oxybelis fulgidus
Oxybelis aeneus
T~helotomis caponsis
Thrasops jacksonii
Gastropyxis smaragdina
-Philothamnus hopiogaster
-Philothamnus irregularis
-Philothamnus angolensis
-Philothamnus dorsalis
-Philothamnus semivariegatus
Elaphe radiata
Stilosoma extenuatum
Lampropeltrs getula
Lampropeltrs triangulum
Arizona elegans
Panthorophis obsolete
Pituophis catenifer
Pituophis melanoleucus
Gonyosoma oxycephalum
Lycodon lacensis
Chilomeniscus straminous
Dinodon rufozonatum
Dinodon semicarinatum
Telescopus semiannufatus
Crotaphopeltis hotamboeia
Dipsadoboa unicolor
Dipsadoboa aulica
Dispholidus typus
Boiga cynodon
Boiga blandingli
Dasypeltrs fasciata
Dasypeltrs scabra
Das~pelfts medici

Figure 2-1. Continued









CHAPTER 3
RESULTS

Frequency Distribution

The frequency distribution of raw total lengths for all four gravitational habitat categories

is shown in Fig. 3-1. The frequency distribution mean (mm), median, mode and standard

deviation within each of the four categories were: stenotopically arboreal (1212, 1050, 1050,

506.5), eurytopically arboreal/terrestrial (1644, 1215, 1680, 1103.6), stenotopically terrestrial

(855.5, 652, 480, 557.7), and stenotopically aquatic (1105, 1000, 1000, 530). Stenotopically

arboreal total lengths had the lowest standard deviation, whereas the eurytopically

arboreal/terrestrial total lengths had the greatest by more than twofold. Slightly more than half

(52%) of the stenotopically arboreal species were between 60-120 cm total length, and 93.5%

were between 50-200 cm total length. There were no stenotopically arboreal species shorter than

50 cm, four species between 201-240 cm (in order of increasing length: Dendroaspis viridis,

Boiga cynodon, Boiga forsteini, and Gonyosoma oxycephahtm), and only one species (Leptophis

ahaetulla) longer than 240 cm. Several non-stenotopically arboreal snake species have total

lengths < 50 cm and > 200 cm (Fig. 3-1).

Gravitational Habitat and Total Body Length

There were differences in total length between the four gravitational habitat categories (P <

0.0001, df = 3, F = 19.433, Fig. 3-1). Mean total length was longer in stenotopically arboreal

species than in stenotopically terrestrial species (P < 0.00001), longer in eurytopically

arboreal/terrestrial species than in stenotopically terrestrial species (P < 0.00001), and longer in

stenotopically aquatic species than in stenotopically terrestrial species (P = 0.0015). However, I

did not find a difference in mean total length between stenotopically arboreal and eurytopically

arboreal/terrestrial species (P = 0.32), between stenotopically arboreal and stenotopically aquatic









species (P = 0.96), or between eurytopically arboreal/terrestrial and stenotopically aquatic

species (P = 0.24).

There were differences in the variance among the four categories (Fig. 3-2). The

coefficients of variation in order of increasing value are: stenotopically arboreal (CV = 0.01 1),

stenotopically aquatic (CV = 0.014), eurytopically arboreal/terrestrial (CV = 0.023), and

stenotopically terrestrial (CV = 0.027).

Gravitational Habitat and Tail Length

Perhaps not surprisingly, tail length is correlated with total length (r2 = 0.65, P < 0.0001, n

= 277, Fig. 3-3). Because I did not find a difference in corrected tail-length between

stenotopically arboreal and eurytopically arboreal/terrestrial species (P = 0.30, Fig. 3-4), I

combined these categories into a single scansorial category (Fig. 3-5). The scansorial species

had corrected tail-lengths on average 1.9 times longer than the two non-scansorial categories (P

< 0.0001) (Fig. 3-5). However, I did not find a difference in corrected tail-length between

stenotopically terrestrial and stenotopically aquatic species (P = 0.94, Fig. 3-4). The relative tail-

lengths for the 227 species included in this study ranged from 1.1% (Rhinotyphlops episcopus) to

48.1% (Uromacer frenatus) (Table 3-1).

Relative Tail-Length, Gravitational Habitat, and Phylogeny

The analysis of covariance (ANCOVA) results supported using the scansorial gravitational

habitat category for the independent contrast analyses instead of the separate analyses of the

stenotopically and eurytopically arboreal/terrestrial categories. After accounting for the effect of

phylogeny on the data, I found a significant relationship between independent contrasts of

scansorial macrohabitat use and loglo transformed RTL (r2 = 0.31, P < 0.0001, Fig. 3-6).










Table 3-1. The 227 species included in this study in order of decreasing relative tail-length
(RTL), and with associated categorizations used in analyses. Range (Rg), OW = Old
World, NW = New World. Gravitational habitat (Gh), 3 = stenotopically arboreal; 2
= eurytopically arboreal/terrestrial; 1 = stenotopically terrestrial; 0 = stenotopically
aquatic.
Family Subamily Species RTL Rg Gh
Colubridae Xenodontinae Uromacer frenatus 0.481 NW 3
Colubridae Xenodontinae Uromacer catesbyi 0.455 NW 3
Colubridae Xenodontinae Uromacer oxyrhynchus 0.440 NW 3
Colubridae Pseudoxyrhophiinae Langaha alluaudi 0.427 OW 3
Colubridae Pseudoxyrhophiinae Mlicropisthodon ochraceus 0.414 OW 3
Colubridae Pseudoxyrhophiinae Langaha pseudoalluaudi 0.403 OW 3
Colubridae Xenodontinae Xenoxybelis boulengeri 0.386 NW 3
Colubridae Colubrinae Gastropyxis smaragdina 0.383 OW 3
Colubridae Colubrinae Oxybelis aeneus 0.380 NW 3
Colubridae Psammophiinae Psammophis punctulatus 0.377 OW 2
Colubridae Xenodontinae Antillophis parwfrons 0.371 NW 2
Colubridae Pseudoxyrhophiinae Langaha madagascariensis 0.368 OW 3
Colubridae Colubrinae Thelotornis capensis 0.365 OW 3
Colubridae Colubrinae Opheodrys aestivus 0.355 NW 3
Colubridae Psammophiinae Psammophis biseriatus 0.352 OW 2
Colubridae Xenodontinae Helicops angulatus 0.340 NW 0
Colubridae Colubrinae Ahaetulla nasuta 0.333 OW 3
Colubridae Psammophiinae Psammophis brevirostris 0.333 OW 2
Colubridae Colubrinae Philothamnus semivariegatus 0.325 OW 3
Colubridae Colubrinae Chironius carinatus 0.320 NW 2
Colubridae Colubrinae Oxybelis f dgidus 0.319 NW 3
Colubridae Pseudoxyrhophiinae Ithycyphus miniatus 0.315 OW 3
Colubridae Xenodontinae Alsophis vudit 0.312 NW 2
Colubridae Xenodontinae Arrhyton f nereum 0.311 NW 1
Colubridae Psammophiinae Rhamphiophis oxyrhynchus 0.308 OW 2
Colubridae Dipsadinae Imantodes cenchoa 0.306 NW 3
Colubridae Colubrinae Philothamnus angolensis 0.302 OW 3
Colubridae Xenodontinae Alsophis portoricensis 0.302 NW 2
Colubridae Colubrinae Dendrelaphis pictus 0.300 OW 3
Colubridae Colubrinae M~asticophis lateralis 0.298 NW 2
Colubridae Psammophiinae Psammophis angolensis 0.296 OW 1
Colubridae Colubrinae Philothamnus irregularis 0.295 OW 3
Colubridae Xenodontinae Alohsu~d ~e a0.293 NW 2
Colubridae Psammophiinae Psammophis jallae 0.293 OW 2
Colubridae Xenodontinae Ialtris dorsalis 0.293 NW1
Colubridae Colubrinae Me~\lrrie, n ew, boddaerti 0.288 NW 2
Colubridae Psammophiinae Rhamphiophis rostratus 0.285 OW 2
Colubridae Dipsadinae Leptodeira punctata 0.283 NW 2
Colubridae Psammophiinae Psammophis sibilans 0.280 OW 2
Colubridae Colubrinae Spilotes pullatus 0.279 NW 2





1 1


Table 3-1. Continued
Family Subamily
Colubridae Natricinae
Colubridae Xenodontinae
Colubridae Boodontinae
Colubridae Colubrinae
Colubridae Colubrinae
Colubridae Colubrinae
Colubridae Colubrinae
Elapidae Elapinae
Elapidae Elapinae
Colubridae Dipsadinae
Colubridae Xenodontinae
Colubridae Dipsadinae
Colubridae Colubrinae
Colubridae Colubrinae
Colubridae Dipsadinae
Colubridae Natricinae
Colubridae Dipsadinae
Colubridae Natricinae


Species
Xenochrophis flavipunctatus
Alsophis antillensis
Grayia smythii
Dispholidus typus
Thrasops jacksonii
Philothammes hoplogaster
Mlasticophis flagellum
Dendroaspis viridis
Dendroaspis w W s, is ps lll
Dipsas catesbyi
Philodryas baroni
Dipsas boettgeri
Philothammes dorsalis
Coluber constrictor
Dipsas chaparensis
Nerodia taxispilota
Dipsas bucephala
Nerodia fasciata


RTL Rg Gh
0.278 OW1
0.278 NW1
0.277 OW1
0.276 OW 3
0.275 OW 3
0.275 OW 2
0.269 NW 2
0.260 OW 3
0.258 OW 3
0.257 NW 3
0.256 NW 3
0.256 NW 3
0.255 OW 3
0.250 NW 2
0.248 NW 3
0.247 NW 2
0.245 NW 3
0.240 NW 2
0.240 OW 1
0.239 NW 2
0.239 NW 1
0.236 OW 3
0.234 NW1
0.232 NW1
0.230 NW1
0.229 OW 3
0.229 NW 3
0.229 NW1
0.228 NW 2
0.228 OW 1
0.227 NW 2
0.226 OW 3
0.226 NW 2
0.226 OW 3
0.224 NW 2
0.224 NW 3
0.224 NW 2
0.223 NW1
0.222 NW 3
0.220 OW1
0.219 NW 2
0.217 NW 1
0.215 NW 1
0.214 OW 2
0.211 NW 1


Colubridae Pseudoxyrhophiinae Dromicodryas bernieri


Colubridae Natricinae
Colubridae Colubrinae
Colubridae Colubrinae
Colubridae Natricinae
Colubridae Xenodontinae
Colubridae Xenodontinae
Colubridae Colubrinae
Colubridae Dipsadinae
Colubridae Natricinae
Colubridae Dipsadinae
Colubridae Natricinae
Colubridae Natricinae
Colubridae Colubrinae
Colubridae Dipsadinae
Colubridae Colubrinae
Colubridae Natricinae
Colubridae Dipsadinae
Colubridae Natricinae
Colubridae Xenodontinae
Colubridae Xenodontinae
Colubridae Psammophiinae
Colubridae Dipsadinae
Colubridae Natricinae
Colubridae Natricinae
Colubridae Colubrinae
Colubridae Colubrinae


Regina septemvittata
Tantilla hendersoni
Gonyosoma oxycephahtm
Thamnophis ordinoides
Drepanoides anomahts
Hypsirhynchus ferox
Boiga cynodon
Dipsas pentana
Thamnophis hammondii
Thamnodynastes pallidus
Natriciteres olivacea
Nerodia rhombifer
Dipsadoboa aulica
Leptodeira bakeri
Boiga blandingii
Nerodia floridana
Leptodeira annulata
Nerodia sipedon
Darlingtonia haetiana
Siphlophis cervimes
Psammophis crucifer
Leptodeira septentrionalis
Thamnophis sirtalis
Regina alleni
Dinodon semicarinatum
Tantilla planiceps





1 1 II


Table 3-1. Continued
Family Subamily
Colubridae Dipsadinae
Colubridae Natricinae
Boidae Boinae
Colubridae Psammophiinae
Colubridae Colubrinae
Colubridae Dipsadinae
Colubridae Dipsadinae
Elapidae Elapinae
Colubridae Xenodontinae
Colubridae Xenodontinae
Atractaspididae Aparallactinae
Colubridae Dipsadinae
Colubridae Pseudoxyrhophiinae
Colubridae Colubrinae
Colubridae Dipsadinae
Colubridae Xenodontinae
Colubridae Homalopsinae
Atractaspididae Aparallactinae
Colubridae Boodontinae
Atractaspididae Aparallactinae


Species
Leptodeira frenata
Storeria dekayi
Corallus ruschenbergerii
Psammophylax tritaeniants
Tantilla relicta
Diadophis princtatus
Leptodeira maculata
Dendroaspis polylepis
Pseudoboa coronata
Clelia clelia
Aparallactus lunulatus
Tretanorhimes variabilis
Leioheterodon modestus
Tantilla melanocephala
Taeniophallus brevirostris
Oxyrhopus petola
Enhydris enhydris
Aparallactus capensis
Lycodonomorphus leleupi
Aparallactus gaentheri
Natrix natrix
Dipsadoboa unicolor
Leptodeira ; i;, II f;,\l;, i,
Elaphe radiata
Protobothrops cormitus
Sonora michoacanensis
Dinodon rufozonatum
Liophis typhhts
Dasypeltis medici
Notechis scretants
Naja melanoletica
Cerberus rynchops
Lycodon laoensis


RTL Rg
0.209 NW
0.208 NW
0.207 NW
0.207 OW
0.203 NW
0.203 NW
0.202 NW
0.199 OW
0.199 NW
0.198 NW
0.198 OW
0.197 NW
0.196 OW
0.195 NW
0.194 NW
0.193 NW
0.192 OW
0.190 OW
0.189 OW
0.188 OW
0.188 OW
0.187 OW
0.186 NW
0.184 OW
0.184 OW
0.181 NW
0.179 OW
0.177 NW
0.174 OW
0.173 OW
0.168 OW
0.165 OW
0.164 OW
0.162 OW
0.160 OW
0.160 NW
0.160 NW
0.159 OW
0.158 OW
0.157 NW
0.157 NW
0.157 NW
0.156 NW
0.156 OW
0.151 NW


Colubridae
Colubridae
Colubridae
Colubridae
Viperidae
Colubridae
Colubridae
Colubridae
Colubridae
Elapidae
Elapidae
Colubridae
Colubridae
Viperidae
Colubridae
Colubridae
Colubridae
Elapidae
Elapidae
Viperidae
Colubridae
Viperidae
Boidae
Viperidae
Colubridae


Natricinae
Colubrinae
Dipsadinae
Colubrinae
Crotalinae
Colubrinae
Colubrinae
Xenodontinae
Colubrinae
Hydrophiinae
Elapinae
Homalopsinae
Colubrinae
Crotalinae
Pseudoxyrhopl
Natricinae
Xenodontinae
Elapinae
Elapinae
Crotalinae
Xenodontinae
Crotalinae
Boinae
Crotalinae
Colubrinae


Trimeresunts gramineus
hiinae Leioheterodon madagascariensis
Seminatrix pygaea
Heterodon platirhinos
Naja mossambica
Naja haje
Agkistrodon piscivorus
Hydrops caesunts
Bothriechis schlegelii
Coralhts canimes
Trimeresurus patrpureomaculants
Lampropeltis triangntem










Table 3-1. Continued


Family
Vipeidae
Viperidae
Colubridae
Colubridae

Viperidae
Colubridae
Elapidae
Colubridae
Elapidae
Viperidae
Elapidae
Colubridae
Colubridae
Colubridae
Elapidae
Elapidae
Colubridae
Elapidae
Colubridae
Colubridae
Boidae
Colubridae
Colubridae
Colubridae


Subamily
Viperinae
Crotalinae
Colubrinae
Boodontinae
Crotalinae
Colubrinae
Elapinae
Dipsadinae
Elapinae
Viperinae
Hydrophiinae
Colubrinae
Colubrinae
Colubrinae
Elapinae
Elapinae
Colubrinae
Hydrophiinae
Natricinae
Dipsadinae
Erveinae
Colubrinae
Boodontinae
Boodontinae


Species
Atheris chlorechis
Trimeresurus albolabris
Pantherophis obsoleta
Pseridaspis cana
Tropidolaemus wagleri
Telescopus semiannulants
Hemachatics haemachants
Hypsiglena torquata
Naja kaouthia
Atheris squamigera
Emydocephahts annulants
Drymarchon corais
Arizona elegans
Prosymna ambigta
Naja atra
Aspidelaps scutatus
Pitatophis catenifer
Aipystents laevis
Virginia valeriae
Carphophis amoenus
Charina trivirgata
Lampropeltis getula
Mlehelya capensis
Lamprophis Jitliginossts


RTL Rg Gh
0.151 OW 3
0.151 OW 3
0.150 NW 2
0.150 OW1
0.149 OW 3
0.148 OW 2
0.148 OW 1
0.147 NW 1
0.146 OW 1
0.144 OW 3
0.144 OW 0
0.143 NW 2
0.143 NW1
0.143 OW1
0.141 OW1
0.136 OW1
0.135 NW 1
0.134 OW 0
0.134 NW 1
0.133 NW 1
0.133 NW 1
0.131 NW 1
0.130 OW 2
0.130 OW1
0.129 NW1
0.129 NW1
0.128 NW 2
0.127 NW1
0.127 NW 2
0.126 NW 1
0.125 NW 2
0.125 NW 1
0.123 OW 1
0.123 NW 1
0.122 OW 2
0.122 OW 2
0.121 OW1
0.120 OW 2
0.116 NW1
0.115 OW1
0.114 OW 1
0.113 OW 1
0.111 NW 1
0.110 NW 1
0.107 NW 1


Tropidophiidae Tropidophiinae Tropidophis morenoi
Colubridae Xenodontinae Pseudoeryx plicatilis
Viperidae Crotalinae Bothrops insularis
Colubridae Xenodontinae Xenodon sevents
Tropidophiidae Tropidophiinae Tropidophis haetiames
Colubridae Xenodontinae Hydrodynastes gigas
Boidae Boinae Epicrates striants
Colubridae Xenodontinae Heterodon nasicus
Colubridae Boodontinae Lamprophis lineatus
Colubridae Colubrinae Chilomeniscus stramineus
Boidae Pvthoninae Python moherus
Colubridae Colubrinae Dlasyieltisfasciata
Colubridae Colubrinae Crotaphopeltis hotamboeil
Colubridae Colubrinae Dasypeltis scabra
Colubridae Dipsadinae Atractics clarki
Viperidae Viperinae Vipera xanthina
Viperidae Viperinae Mlacrovipera mauritanica
Colubridae Boodontinae Duberria httrix
Colubridae Xenodontinae Erythrolamprus aesculapii
Viperidae Crotalinae Agkistrodon contortrix
Colubridae Dipsadinae Atractics hostilitractus


a










Table 3-1. Continued


j 11


Family
Colubridae
Boidae
Elapidae
Colubridae
Colubridae
Colubridae
Boidae
Xenopeltidae
Acrochordidae
Elapidae
Colubridae
Colubridae
Elapidae
Viperidae
Colubridae
Elapidae
Boidae
Colubridae
Viperidae
Elapidae
Colubridae
Leptotyphlophidae
Colubridae
Viperidae
Viperidae
Colubridae
Viperidae
Elapidae
Viperidae
Elapidae
Viperidae
Colubridae
Boidae
Atractaspididae
Atractaspididae
Boidae
Boidae
Atractaspididae
Elapidae
Viperidae
Viperidae
Viperidae
Viperidae
Atractaspididae
Viperidae


Subamily
Boodontinae
Pythoninae
Hydrophiinae
Colubrinae
Dipsadinae
Xenodontinae
Pythoninae


Elapinae
Colubrinae
Boodontinae
Hydrophiinae
Viperinae
Dipsadinae
Hydrophiinae
Boinae
Colubrinae
Crotalinae
Elapinae
Colubrinae

Colubrinae
Crotalinae
Viperinae
Colubrinae
Crotalinae
Elapinae
Viperinae
Elapinae
Crotalinae
Colubrinae
Boinae
Aparallactinae
Aparallactinae
Pythoninae
Erveinae
Aparallactinae
Elapinae
Viperinae
Crotalinae
Viperinae
Viperinae
Aparallactinae
Viperinae


Species
Lycophidion variegatum
Python sebae
Pelamis platunts
Prosymna stithlmannii
Atractics depressiocellus
Farancia abacura
Mlorelia viridis
Xenopeltis unicolor
Acrochordus granulatus
Elapsoidea sundevallii
Phyllorymchus decurtants
Lycophidion capense
Laticateda colubrina
Causus rhombeatus
Atractics darienensis
Hydrophis melanocephahts
Boa constrictor
Pitatophis melanoleucus
Atropoides nummf fer
Bungants fasciatus
Prosymna sundevallii
Leptotyphlops macros
Phyllorymchus brown
Crotahts ntber
Bitis nasicornis
Prosymna bivittata
Crotahts horridus
Mlicrunts Jilyius
Bitis arietans
Elapsoidea gatentheri
Crotahts atrox
Stilosoma extenuatum
Acrantophis dumerili
Xenocalamus sabiensis
Amblyodipsas ventrimaculants
Python regius
Gonglphismuelleri
Amblyodipsas polylepis
Elapsoidea nigra
Bitis gabonica
Crotahts cerastes
Bitis catedalis
Bitis atrops
Xenocalamus mechowii
Causus defilippii


RTL
0.105
0.104
0.104
0.103
0.103
0.102
0.102
0.100
0.099
0.099
0.097
0.096
0.096
0.096
0.095
0.093
0.090
0.090
0.088
0.085
0.085
0.082
0.081
0.080
0.080
0.080
0.076
0.075
0.075
0.075
0.073
0.073
0.071
0.069
0.068
0.066
0.064
0.064
0.064
0.063
0.063
0.063
0.062
0.060
0.060


Rg
OW
OW
OW
OW
NW
NW
OW
OW
OW
OW
NW
OW
OW
OW
NW
OW
NW
NW
NW
OW
OW
OW
NW
NW
OW
OW
NW
NW
OW
OW
NW
NW
OW
OW
OW
OW
OW
OW
OW
OW
NW
OW
OW
OW
OW










Table 3-1. Continued
Family Subamily Species RTL Rg Gh
Elapidae Elapinae Elapsoidea semiannulata 0.055 OW1
Atractaspididae Atractaspidinae Atractaspis bibronii 0.052 OW1
Aniliidae Anilius scytale 0.044 NW1
Leptotyphlophidae Leptotyphlops alfredschmidti 0.042 NW1
Typhlopidae Xenotyphlops grandidieri 0.035 OW 1
Cylindrophiidae Cylindrophis ruffus 0.017 OW 1
Typhlopidae Rhinotyphlops episcopus 0.011 OW 1





111


14 +


E 12

S10

O 8


O'
40 80 120


160 200 240


Total Length (cm)


25 +1


Q0 20
a
15

a 10


-
0 +


40 100 200 300 400 500 600 700 800 900

Total Length (cm)


Figure 3-1. Snake total length frequency distribution. A) Stenotopically arboreal species. B)
Eurytopically arboreal/terrestrial species. C) Stenotopically terrestrial species. D)
Stenotopically aquatic species.





25

2 20

S15

a 10


60 100 140 180

Total Length (cm)


220 260


80 120 160 200 240 280

Total Length (cm)


Figure 3-1. Continued










0.030

-a0.025

S0.020

S0.015




0.000
Stenotopically Eurytopically Stenotopically Stenotopically
Arboreal Arb/Terr Terrestrial Aquatic
Gravitational Habitat

Figure 3-2. Coefficients of variation of loglo transformed total length among the four
gravitational habitats.





















coo

a,
5o O
-o oooo o

E- a Ouv G




to o



o

2.2 2.4 2.6 2.8 3.0 3.2 3.4 3.6

Log(Total Length)

Figure 3-3. Regression of loglo tail length on loglo total length for 227 snake species (r2 = 0.65;
P < 0.0001).
























on a *S




O. e


aC
llllllll






2.2 2.4 2.6 2.8 3.0 3.2 3.4 3.6

Log(Total Length)

Figure 3-4. Analysis of covariance (ANCOVA) on corrected tail-length for 227 species of
stenotopically arboreal (upper dotted line, open triangles), eurytopically
arboreal/terrestrial (lower dotted line, solid circles), stenotopically terrestrial, and
stenotopically aquatic (overlapping solid lines, solid circles and open circles
respectively) snakes.
























o e~L *8 e




allllllll
2.2 2. .

Lo(otlLegh
Fiue3-.Aalsso cvr iac (A C V )o orce al-eghfr27seiso
scasoia (dte ie pntinge) n tntpcly ersra n qai







(cnov rlapn sdolid lines, solid crircles) and open squiares erespe trively sn akes.

















O OO O
O O


1.5 -

O

1-I O OO0


O O


-1.75


-1.25


-1 -0.75


-0.25


Log RTL


Figure 3-6. Independent contrasts of macrohabitat use and loglo transformed relative tail-length
(RTL; r2 0.31, P < 0.0001). For macrohabitat use, O stenotopically aquatic; 1
stenotopically terrestrial; 2 = scansorial.









CHAPTER 4
DISCUSSION

Relative Tail-Length, Gravity, and Climbing in Snakes

Gravity is a pervasive force that clearly provides acute challenges to blood circulation in

terrestrial vertebrates. These challenges are particularly pronounced in elongate vertebrates such

as snakes utilizing complex three-dimensional vertical habitats (Lillywhite 1996). Thus, the

ability to regulate blood circulation in vertical habitats or upright postures potentially imposes

important constraints on the behavior and morphology of scansorial snake species. Overcoming

these cardiovascular constraints has likely resulted in the evolution of novel morphological

adaptations as species radiated from terrestrial into arboreal habitats.

A likely consequence of limblessness in snakes is an increase in the selective pressures

imposed on various aspects of the body and tail (Polly et al. 2001). This is particularly true in

species adapted to the extreme challenges associated with arboreal environments (Lillywhite

1996; Lillywhite and Henderson 2001). Therefore, differences in body and tail characteristics

should exist among snake species inhabiting different macrohabitats and, thus, experiencing

different selective pressures.

One such morphological change likely involves the total length of stenotopically arboreal

species. The distribution frequency of total length for adult stenotopically arboreal species

suggests there are constraints limiting total length to between 50 and 200 cm, with few

exceptions. There were no stenotopically arboreal species with total lengths < 50 cm as adults.

Because gradients of intravascular and transmural pressures increase with absolute vertical

length of a blood column, relatively short species are not strongly affected by the cardiovascular

constraints imposed by gravity. Carotid blood flow in the stenotopically terrestrial viper

Agkistrodon piscivorus was less impaired by tilting in smaller than in larger individuals










(Lillywhite and Smits, 1992). Furthermore, Lillywhite and Smits (1992) suggest that gravity

induced cardiovascular constraints on blood circulation limit the total length of arboreal vipers to

about 1 m. For snakes in general, my results indicate that constraints limit total length in adult

stenotopically arboreal species to > 50 cm, and that these constraints are not likely

cardiovascular.

Insofar as gravity would not seem to be a factor at smaller body size, species < 50 cm may

have lesser ability to adequately span gaps, which might result in ecological constraints such as

increased vulnerability to predation or decreased hunting success. Because several of the

arboreal species included in the distribution frequency have closely related terrestrial species

with total lengths < 50 cm (e.g., Trimeresurus), the comparative data support the inference that

there are lower limitations on the total length of stenotopically arboreal snakes to > 50 cm.

Similarly, there were few stenotopically arboreal snake species (n = 5) with total lengths >

200 cm suggesting upper level constraints on the total length of these species. The four species

between 201-240 cm (in order of increasing length: Dendroaspis viridis, Boiga cynodon, Boiga

forsteini, and Gonyosoma oxycephalum) all have relatively long tails, with three having relative

tail-lengths between 23-26% (no length data available for Boiga forsteini, although its relative

tail-length (RTL) should be within this range as well). Only one species (Leptophis ahaetulla)

slightly exceeded 240 cm in total length (247.5 cm), and it also has a relatively long tail (39% of

total length). Snakes > 200 cm are perhaps also negatively influenced by ecological constraints

in complex vertical three-dimensional habitats. Their relatively large mass (particularly in boids)

might restrict movements to thicker branches.

The variance in total length among eurytopically arboreal/terrestrial species was more than

twofold greater than that of stenotopically arboreal species. In fact, the stenotopically arboreal









species had the lowest variance in total length of all four categories. These results further

suggest there are constraints limiting total length of stenotopically arboreal species, particularly

because I did not find differences in either the mean total length or the mean corrected tail-length

between stenotopically and eurytopically arboreal/terrestrial species.

Corrected tail-length of scansorial species was on average 1.9 times longer than that of

non-scansorial species (Fig. 3-5). Lengthening the tail relative to the body increases the relative

length of blood vessels that are surrounded by tight (low compliance) tissues. This helps to

mitigate posterior blood pooling and thereby facilitates blood flow to the heart, brain, and vital

organs during the upright positions experienced during climbing (Lillywhite and Gallagher 1985;

Lillywhite 2005). Therefore, the long tails of scansorial snakes can be regarded as one of a suite

of characters likely to be adaptive responses to the cardiovascular constraints on blood

circulation imposed by gravity (Lillywhite 1987; Lillywhite 2005).

Relative tail-length is positively correlated with absolute tail-length (Fig. 4-1) and likely

more important in countering the effects of gravity than absolute tail-length. Snakes with

relatively longer tails have a larger percent of dependent vessels contained within the tight

integument of the tail and are thus more resistant to dependent transmural pressures experienced

during climbing.

I did not find a difference in RTL between stenotopically and eurytopically

arboreal/terrestrial species (Fig. 3-4). There are two possible explanations for this. (1) Even

though eurytopically arboreal/terrestrial species climb less frequently than stenotopically

arboreal species, they still need adaptations to counter the effects of gravity on blood circulation.

This is the most likely explanation, and it could help explain why many eurytopic species that

periodically climb (e.g., Coluber constrictor and M\~a;sticophis flagellum) have relatively long









tails, but lack prehensile tails. (2) Stenotopically arboreal species might have evolved from

eurytopically arboreal/terrestrial species. Henderson and Binder (1980) proposed this hypothesis

in a review of the ecology and behavior of vine snakes. However, the composite tree in this

study does not support this hypothesis due to lack of resolution (Fig. 2-1). The tree shows two

instances where stenotopically arboreal species likely evolved from eurytopically

arboreal/terrestrial ancestors, and at least four instances where eurytopically arboreal/terrestrial

species likely evolved from stenotopically arboreal ancestors (Fig. 2-1). However, there are at

least three instances in which the ancestral state is unclear. Analyses using better resolved

phylogenies at the generic and species level for multiple clades are necessary to thoroughly test

this hypothesis.

The relative tail-lengths for the stenotopically aquatic species (9.3% to 19.2%) fall well

within the range for the stenotopically terrestrial species (1.1% to 38.3%). This is likely because

the tails of stenotopically aquatic species are affected by the unique selective pressures

associated with an essentially weightless environment. In most aquatic environments, the tails

do not function in countering the effects of gravity on blood circulation, but rather in propulsion.

Consequently, these species should be expected to have relative tail-lengths long enough to

effectively propel the snake through water, yet short enough to minimize drag. Therefore,

similarity in RTL between stenotopically aquatic and stenotopically terrestrial species is likely

the result of entirely different selective pressures.

Several patterns emerged from the data regarding RTL at the family level. First, the

Viperidae in general have relatively short body lengths, absolute tail lengths, and relative tail-

lengths, and these characteristics appear to be retained in arboreal species (Lillywhite and Smits

1992). The results from this study support these observations, with the highest RTL value for an









arboreal viperid in this study being 18.4% for Protobothrops cornutus (Table 3-1). However,

even though vipers tend to be relatively short, Martins et al. (2001) demonstrate that RTL

increases with increasing arboreality in the genus Bothrops. This trend for increased RTL in

scansorial species is further supported by the 25 viperid species included in this study, and it

likely characterizes viperids in general. Similarly, the 11 boid species used in this study tended

to have relatively short tails, although many species are relatively long bodied. The highest RTL

value for a stenotopically arboreal boid was 16.2% for Corallus ruschenbergerii (Table 3-1).

The species with the longest relative tail-lengths were clearly members of the family Colubridae.

The 87 species with the longest RTL values were all colubrids (ranging from 20.8% 48.1%),

with the only exception of two arboreal elapids (Dendroaspis viridis and Dendroaspis

angusticeps) with RTL values of 26.0% and 25.8% respectively (Table 3-1). Although

Dendroa;spis polylepis is fast moving and an agile climber, it is primarily terrestrial and has a

shorter RTL (19.9%) than its strictly arboreal congeners even though they attain similar total

lengths (Branch 1998).

Snake Tails and Defense: Speed and Pseudautotomy

In addition to scansorial species, relatively long tails were also noted in several non-

scansorial species characterized by defensive tail loss (pseudautotomy) and/or using speed as a

primary means of capturing prey and escaping predators (e.g., Darlingtonia, Drepanoides, and

Thamnnophis). In fact, many of these species had RTL values similar to many scansorial species

(RTLs of 29 species included in this study were between 20% and 34%).

Included within this group of species that use their tails to escape predators are species that

often swim, but evidence suggests that relatively long tails are not particularly advantageous for

either rapid terrestrial locomotion or swimming (Jayne 1988; Jayne and Bennett 1989). Jayne

(1988) found that RTL in Nerodia and Elaphe were about equally effective for swimming,









suggesting the relatively long tails of natricines are not an adaptation for swimming. However,

of the 29 species of stenotopically terrestrial species with relatively long tails included in this

study, at least 19 are known from the literature to lose their tails in defense. While the relative

frequency of tail loss in wild populations cannot always be accurately determined from museum

specimens alone due to potential collection bias (Arnold 1988; Jayne and Bennett 1989), these

results certainly warrant further investigation.

Many long-tailed terrestrial species are also active, diurnal predators that rely on speed to

procure prey (Greene 1997). However, if fast moving terrestrial snakes possess adaptations for

speed, and long RTL is not required for speed, then alternative selective pressures are likely

responsible for maintaining long tails in these species. Furthermore, because many of these

species are known to possess stub tails (Greene 1997), it is reasonable to infer that the long tails

of these terrestrial species might be a mechanism for defense. Many of these species rely on

speed for escaping predators, and quickly fleeing increases the likelihood of a predatory attempt

being directed to the tail instead of the body (Greene 1973). A long tail can subsequently break

allowing the snake to escape.

Guyer and Donnelly (1990) used length-mass relationships to divide a Costa Rican snake

assemblage of 27 species into four distinct morphological groups and found that, among 603

individuals, the highest frequencies of tail breaks occurred in the leaf litter and terrestrial

categories (mean = 0.19 and 0.08 respectively) with the arboreal and semiarboreal categories

having the lowest (mean = 0.01 and 0.0 respectively). In fact, the leaf-litter habitat was

characterized by having long RTL and high frequency of tail loss (e.g., Coniophanes,

Dendrophidion, Pliocercus, Rhadinaea, and Scaphiodontophis) and, thus, highlighting the

apparent prevalence of pseudautotomy- in many terrestrial snake species within a single









assemblage. Additional studies of snake assemblages in other areas would be of considerable

use in determining the frequency of pseudautotomy- among snake assemblages in diverse

ecosystems.

The current dogma regarding pseudautotomy suggests it is a rare phenomenon in snakes

(Arnold 1988; Marco 2002). However, the paucity of information on this topic renders such a

conclusion premature. As more natural history data become available, the number of snake

species observed exhibiting relatively high frequencies of tail loss in wild populations continues

to increase. The sum of these findings suggests that pseudautotomy in snakes is possibly more

common than presently thought and potentially important in understanding the evolution of

relatively long tails in many terrestrial snake species.

However, different functions of tail length need not be mutually exclusive. It is likely that

the long RTL of many eurytopically arboreal/terrestrial species (e.g., Cohiber constrictor) serve

the dual purposes of aiding in defense by pseudautotomy, while also countering the effects of

gravity during climbing. Further studies comparing tail characteristics of scansorial and

terrestrial species (e.g., prehension, tail length, presence of fracture planes, size of caudal

vertebra, and muscle mass) would be of considerable interest and I intend to pursue these ideas

in a later study.














0.000


-0.500




C -1.000




C1-1.500




-2.000


O
O


-2.500
0.00 0.50


1.00


1.50 2.00


2.50 3.00


Log,, (absolute tail-length)


Figure 4-1. Regression of absolute tail length on relative tail-length (RTL) for 227 snake species
(r2 0.64, P < 0.0001).










APPENDIX A
STENOTOPICALLY ARBOREAL SPECIES

Table A-1. Stenotopically arboreal species (4 families, 10 subfamilies, 34 genera, 77 species)
used for total length frequency distributions; NW = New World, OW = Old World.
Family Subfamily Species Rang

Boid ae Boinae Corallus annulatus NW
Corallus caninus NW
Corallus ruschenbergerii NW
Pythoninae Morelia viridis OW
Colubridae Colubrinae Ahaetulla dispar OW
Ahaetulla nasuta OW
Ahaetulla prasina OW
Ahaetulla pulverulenta OW
Boiga cynodon OW
Boiga blandingii OW
Boiga cyanea OW
Boiga forsteini OW
Boiga multifasciata OW
Boiga ocellata OW
Chrysopelea ornata OW
Chrysopelea paradisi OW
Dendrelaphis cyanochloris OW
Dendrelaphis pictus OW
Dipsadoboa aulica OW
Dispholidus typus OW
Drymobius margaritiferus NW
Elaphe frenata OW
Elaphe prasina OW
Gastropyxis smaragdina OW
Gonyosoma oxycephalum OW
Leptophis ahaetulla NW
Leptophis depressirostris NW
Opheodrys aestivus NW
Oxybelis aeneus NW
Oxybelis fulgidus NW
Philothamnus angolensis OW
Philothamnus dorsalis OW
Philothamnus hoplogaster OW
Philothamnus irregularis OW
Philothamnus semivariegatus OW
Thelotornis capensis OW
Thelotornis kirtlandii OW
Thrasops jacksonii OW
Dipsad inae Dipsas boettgeri NW
Dipsas catesbyi NW
Dipsas chaparensis NW
Dipsas indica NW










Table A-1 Continued
Family Subfamily
Colubridae Dipsadinae


Species
Dipsas peruana
Imantodes cenchoa
Imantodes inornatus
Leptodeira annulata
Leptodeira septentrionalis
Sibon nebulata
Sibon sartorii
Psammophis biseriatus
Langaha alluaudi
Langaha madagascariensis
Langaha pseudoalluaudi
Micropisthodon ochraceus
Stenophis gaimardi
Stenophis granuliceps
Philodryas baroni
Siphlophis cervinus
Uromacer catesbyi
Uromacer frenatus
Uromacer oxyrhynchus
Dendroaspis angusticeps
Dendroaspis viridis
Bothriechis schlegelii
Trimeresurus cornutus
Trimeresurus erythrurus
Trimeresurus gramineus
Trimeresurus macrolepis
Trimeresurus malabaricus
Trimeresurus medoensis
Trimeresurus purpureomaculatus
Tropidolaemus wagleri
Atheris chlorechis
Atheris hispida
Atheris nitschei
Atheris rungweensis
Atheris squamigera


Rg
NW
NW
NW
NW
NW
NW
NW
OW
OW
OW
OW
OW
OW
OW
NW
NW
NW
NW
NW
OW
OW
NW
OW
OW
OW
OW
OW
OW
OW
OW
OW
OW
OW
OW
OW


Psammophiinae
Pseudoxyrhophiinae






Xenodontinae





Elapidae Elapinae

Viperidae Crotalinae










Viperinae










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BIOGRAPHICAL SKETCH

Coleman Sheehy has always had a passionate curiosity about the natural world. Growing

up in Richmond, Virginia, his earliest interests centered around dinosaurs, but these interests

quickly expanded to include living reptiles and amphibians and, in particular, snakes. Thanks to

an encouraging and supportive family, Coleman's basement, while growing up, was constantly

filled with native wildlife he collected on local field excursions. Most of these animals were

reptiles and amphibians and it was during these years that Coleman honed many of the

observational and field skills that would become such an important aspect of his academic

career.

After acquiring his General Education Diploma (GED), Coleman was invited to work at

James River Park in Richmond, where his responsibilities included leading interpretative hikes

and float trips through miles of undeveloped, island-dotted habitat along the James River. He

also gave hundreds of educational programs focused on introducing the public of all ages to the

fascinating reptiles and amphibians native to Virginia and to the importance of their

conservation.

Coleman later began working at a local pet store specializing in exotic reptiles and

amphibians and, after several years, became general manager of the primary store in Richmond.

It was during this time that Coleman realized many of the problems associated with the pet trade

and he sought to resolve many of these by initiating captive breeding programs and by only

purchasing or trading captive bred animals. Also during this time, Coleman saved enough

money to fund a month-long trip exploring Madagascar. It was during this trip that he met two

prominent American herpetologists (Drs. Ronald Nussbaum and Christopher Raxworthy) who

strongly influenced his career path. The experience left Coleman with the realization that he

could incorporate his love of herpetology and traveling into a successful career through









academia. Coleman subsequently enrolled at J. Sargeant Reynolds Community College in

Virginia and graduated sunana cunt laude with an Associate in Science in the year 2000. He

then transferred to the University of Florida, where he graduated magna cunt laude with a

bachelor' s degree in zoology and a minor in wildlife ecology and conservation in 2002.

Coleman has traveled to Madagascar, Honduras, South Africa, Tobago, the Bahamas,

Colombia, Brazil, and Taiwan as well as many states within the US. While working on his

master' s, Coleman taught the laboratories for many classes, including Herpetology, the Natural

History of Amphibians and Reptiles, Vertebrate Zoology, Functional Vertebrate Anatomy,

Ecology, Advanced Island Biogeography, and Introductory Biology. He has worked in the

herpetology collections of the Florida Museum of Natural History and the Transvaal Museum in

South Africa, and he has presented several talks at scientific meetings. He has also published

regularly in peer-reviewed j ournals since 2001 and, to date, has ten publications including the

description of a new species of frog from Bolivia.

Coleman has been accepted into a Ph.D. program at the University of Texas at Arlington,

where he will pursue his interests in snake systematics, evolutional biology, ecomorphology, and

functional morphology by studying several groups of Central and South American snakes. His

long-term goal is to secure a tenured position in a Museum or University, where he can conduct

research and teach in the areas of herpetology and vertebrate evolution, while also actively

promoting the conservation of wildlife worldwide.