Does Life History Shape Sexual Size Dimorphism in Anurans

DOES LIFE HISTORY SHAPE SEXUAL SIZE DIMORPHISM IN ANURANS:

A COMPARATIVE ANALYSIS

A Thesis

Presented to

The Faculty of Graduate Studies

The University of Guelph

XU HAN

In partial fulfilment of requirements

for the degree of

Master of Science

March, 2008

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While these forms may be included Bien que ces formulaires in the document page count, aient inclus dans la pagination, their removal does not represent il n'y aura aucun contenu manquant. any loss of content from the thesis. Canada ABSTRACT

DOES LIFE HISTORY SHAPE SEXUAL SIZE DIMORPHISM IN ANURANS:

A COMPARATIVE ANALYSIS

Xu Han Advisor: University of Guelph, 2008 Professor Jinzhong Fu

The evolution of sexual size dimorphism (SSD) is constrained by life history traits, and therefore, mating combat, length of breeding season, female fecundity and parental care are expected to be correlated with male and/or female body sizes. Using phylogenetic comparative methods, correlations between SSD and life history traits were examined in 545 anuran species. Data were analyzed with phylogenetic independent contrasts and a maximum likelihood method. A positive correlation was found between mating combat and body size. Egg size, clutch size and clutch volume were also positively correlated with female body size and SSD. Furthermore, body size and SSD were positively correlated in all anurans and in the family Bufonidae, but were negatively correlated in the family Dicroglossidae. Female fecundity may mainly influence the evolution of SSD, and the general pattern of SSD in anurans does not follow Rensch's rule, although SSD does so follow in the family Dicroglossidae. ACKNOWLEDGEMENTS

I would like to express my gratitude to all those who have supported me during the completion of this thesis. First, I would like to thank my supervisor, Dr. Jinzhong Fu, for his guidance, suggestions and encouragement during my research. His perpetual energy and enthusiasm in science has motivated me. In addition, he was always accessible and willing to help throughout my Master's study.

Dr. James Bogart and Dr. Beren Robinson deserve special thanks as my thesis committee members. Dr. Bogart's encyclopedic knowledge of anurans was an invaluable resource. His patience and kindness are greatly appreciated. Dr. Robinson gave up hours of his time to advise me, and I cannot thank him enough for his attention and passion in training new scientists. Dr. Paul Herbert and Dr. Andrew MacDougall sat on my examination committee. Their excellent suggestions resulted in substantial improvement in the final version of this thesis.

I was delighted to interact with Dr. Thomas Nudds by attending his "Scientific

Communication" classes and with Dr. Robert Hanner in his "Molecular Evolution" class.

Dr. Nudds' instructions on critical thinking and proposal design were extremely helpful for this project and will be good for my future scientific career. Dr. Hanner taught me phylogenetic methodologies, which were very useful in the data analyses of this project.

I was also grateful to the Department of Integrative Biology at the University of

Guelph for giving me permission to commence this thesis in the first case. Moreover, this project would not have been possible without the McLaughlin Library at the University

1 of Guelph. It allowed me to access all of the data relevant to this research. Thanks as well

to the Natural Sciences and Engineering Research Council of Canada, who funded this project through my advisor Dr. Jinzhong Fu.

Many other people assisted me with this project. My labmates in the Fu labs,

especially Kate Crosby, Daniel Noble, Ke Bi, John Urquhart, Marina Amato, Owen

Lonsdale, and Zhonge Hou, gave me valuable hints and assistance throughout the project.

Members in the Boulding lab, including Dr. Elizabeth Boulding, Matthew Lemay, Hyuk

Je Lee, Mark Culling, and Heather Freamo provided constructive criticism and

stimulating suggestions on my research through lab meetings. Thanks also go to Daniel

Noble, Mark Sherrard, Heather Freamo, and my comrades in Zoology House, Alison

Fischer and Anibal Castillo, who looked closely at early versions of the thesis for English

style and grammar, and offered suggestions for improvement. My experience of being a teaching assistant with Graham Nancekivell in the "Vertebrate Structure and Function" course has been proven to be invaluable to this research. I learned many important

evolutionary concepts and terminologies in this course and enjoyed the work due to

Graham's cheerful personality. In addition, I would like to thank all my friends in Canada

for the support and good company they have provided me with.

My deepest gratitude goes to my family for their unflagging love and support throughout my life. Without them, I would never have made it this far. I am indebted to my father, Zongqi Han, for his care and love. As a typical father in a Chinese family, he worked industriously to support the family and spared no effort to provide the best possible environment for me to grow up and attend school. My mother, Donghua Lin,

11 was extremely influential in encouraging me to study abroad. Her constant support encouraged me to keep moving forward when I encountered difficulties. I would like to give my special thanks to my husband, Renji Lu, whose patient love and thoughtfulness enabled me to complete this work.

iii TABLE OF CONTENTS

ACKNOWLEDGEMENTS i

TABLE OF CONTENTS iv

LIST OF TABLES vii

LIST OF FIGURES .....viii

INTRODUCTION 1

Mating Related Life History Traits 4

Mating Combat. 4

Length of Breeding Season 6

Breeding Related Life History Traits 7

Egg Size, Clutch Size and Clutch Volume 7

Parental Care 8

Rensch's Rule 9

Objectives 10

METHODS 13

Taxa Included for Comparative Analyses 13

Body Size and Life History Traits 13

Body Size 13

Sexual Size Dimorphism .14

Mating Combat 17

Length of Breeding Season 18

Egg Size, Clutch Size, and Clutch Volume 19

Parental Care ..20

iv Phytogeny Reconstructions 20

Criteria for Choosing Phytogenies.... 21

Supertree Constructions 22

Phylogenetic Comparative Analyses 24

Phylogenetic Independent Contrasts 24

BayesDiscrete 29

Comparative Analyses 32

RESULTS 35

Sexual Size Dimorphism 35

SSD and Mating Related Life History Traits 38

Correlation between SSD and Mating Combat 38

Correlation Between SSD and Length of Breeding Season. 39

SSD and Breeding Related Life History Traits 39

Correlation between SSD and Clutch Volume 39

Correlation between SSD and Egg Size 40

Correlation between SSD and Clutch Size 41

Correlation between SSD and Parental Care 41

Rensch'sRule 42

Summary of Correlations 43

DISCUSSION: 47

Sexual Size Dimorphism 47

SSD and Mating Related Life History Traits 48

Correlation between SSD and Mating Combat 48

v Correlation between SSD and Length of Breeding Season -. 49

SSD and Breeding Related Life History Traits 50

Correlation between SSD, Clutch Volume, Egg Size, and Clutch Size 50

Correlation between SSD and Parental Care 51

Rensch's Rule 53

Suggestions for Future Research 56

CONCLUSION 61

REFERENCES..... 62

APPENDIX 1 Data of body size, egg size, clutch size, reversed sexual size dimorphism, mating combat, breeding season and parental care in anurans 93

APPENDIX 2 References for body size, egg size, clutch size, mating combat, length of breeding season, and parental care in anurans species 108

APPENDIX 3 A manually constructed phylogenetic supertree for 545 anuran species across 37 anuran families 133

APPENDIX 4 Result of the correlation tests among body size, body size dimorphism and

life history traits in the all-anuran analysis and in the within-family analyses using the program of Comparative Analysis via Independent Constrasts (CAIC) and the

BayesDiscrete program 143

vi LIST OF TABLES

Table 1. Summary of character states for body size and body size dimorphism 15

Table 2. Phylogenetic comparative analyses, indicating correlations tested, corresponding programs applied and predictor variables used 33

Table 3. Correlations ranked by the order of statistical significance (high to low) 46

APPENDIX 4 Table 1. Correlations between body size and mating related life history traits 144

APPENDIX 4 Table 2. Correlations between body size dimorphism and mating related life history traits 145

APPENDIX 4 Table 3. Correlations between body size and breeding related life history traits.... 147

APPENDIX 4 Table 4. Correlations between body size dimorphism and breeding related life history traits 148

APPENDIX 4 Table 5. Correlations among breeding related life history traits 150

APPENDIX 4 Table 6. Correlations between female body size and male body size as well as correlations between body size dimorphism and body size 151

vn LIST OF FIGURES

Figure 1. Hypothetical relations among body size ratio (F/M), body size, female combat, male scramble competition, male territory defense, breeding season, parental care (female parental care and male parental care), clutch volume, egg size, and clutch size in anurans.

See text for detailed discussion of hypotheses used to generate this predicted relations 12

Figure 2. A hypothetical phylogeny among three taxa. X; represents the value for the node or species i. The branch length between the node or species i and its ancestral node is labelled as Vj 26

Figure 3. A, A hypothetical polytomy among three taxa. The value for the node or species i is represented by Xj. The branch length between the node or species i and its ancestor node is labeled as v;; B, Two sub-nodes assigned by CAIC 28

Figure 4. Parameters estimated in the independent model (upper) and the dependent

(lower) model in BayesTraitsDiscrete 31

Figure 5. Frequency distribution of body size in anurans with mean=45.89mm,

SD=21.26, and n=259 species 36

Figure 6. Frequency distribution of body size ratio (F/M) in anurans with mean=1.15,

SD=0.15 and n=259 species 37

Figure 7. Correlations between sexual size dimorphism, body size and life history traits in anurans. 44

Figure 8. A hypothetical example showing incorrect branch length changes when species with missing values are removed from comparative analyses under the assumption that all branch lengths equal to one unit .60

vin APPENDIX 3 Figure 1. Partial supertree of anurans, showing the relationships among

57 species and four major clades: the superfamily Dendrobatoidea (family Dendrobatidae and family Aromobatidae), and the families Megophryidae, Hylidae and Bufonidae. ..133

APPENDIX 3 Figure 2. Partial supertree of anurans, showing the relationships among

37 species and two major clades: the families Dicroglossidae and Ranidae 134

APPENDIX 3 Figure 3. Supertree of the family Megophryidae, showing the relationships among 25 species 135

APPENDIX 3 Figure 4. Basal portion of the phylogeny of the family Hylidae, showing relationships among 28 species (Redrawn after Wiens etal, 2006) 135

APPENDIX 3 Figure 5. Partial phylogeny of the family Hylidae, showing relationships among 65 species within the subfamily Hylidae (Redrawn after Wiens etal, 2006).... 13 6

APPENDIX 3 Figure 6. Partial phylogeny of the family Hylidae, showing relationships among 66 species within the tribe Hylini (Redrawn after Wiens et al, 2006) 137

APPENDIX 3 Figure 7. Partial phylogeny of the family Hylidae, showing relationships among 39 species within the subfamily Hylidae (Redrawn after Wiens et al, 2006).. ..138

APPENDIX 3 Figure 8. Phylogeny of the superfamily Dendrobatoidea, showing the relationships among 62 species 139

APPENDIX 3 Figure 9. Supertree of the family Bufonidae, showing the relationships among 58 species 140

APPENDIX 3 Figure 10. Basal portion of the supertree of the family Ranidae, showing the relationships among 48 species 141

APPENDIX 3 Figure 11. Partial supertree of the family Ranidae, showing the relationships among 33 species 142

IX APPENDIX 3 Figure 12. Supertree of the family Dicroglossidae, showing the relationships among 27 species 142

x INTRODUCTION

Sexual size dimorphism (SSD), where males and females differ in body size, is the most conspicuous difference between two sexes in many animals. Females are larger than males in most animals, particularly in species that grow throughout their lives

(Andersson, 1994). However, males are predominantly larger than females in some animal groups with determinate growth, such as mammals and birds (Fairbairn et ah,

2007). SSD is generally considered to reflect the adaptations of males and females either to their disparate reproductive roles or to ecological differences (Shine, 1989; Fairbairn et ah, 2007). Many factors may influence the direction and degree of SSD biased towards either sex (Andersson, 1994). SSD is thought to be the consequence of the net balance of sexual selection and natural selection on both sexes.

The common explanation of a female-biased SSD is that fecundity selection favors large females, because they can produce more eggs (Andersson, 1994). In contrast, a male-biased SSD is expected when natural selection for female fecundity is less intense than sexual selection, because large size gives an advantage for males in contests for females or because female choose larger males (Fairbairn et ah, 2007).

However, more contemporary research suggests that it is unlikely that SSD can be simplistically interpreted by female fecundity or male-male competition (Andersson,

1994; Fairbairn et ah, 2007). The relative effects of non-reproductive and reproductive fitness in males and females will ultimately determine the degree of sexual dimorphism.

To understand the relative importance of natural selection and sexual selection on SSD, the balance between life history traits should be taken into account (Andersson and

Iwasa, 1996).

1 Amphibians display indeterminate growth (Halliday and Verrell, 1988). Most of the species exhibit a female-biased SSD (Shine, 1979) and are renowned for variable life histories (Duellman and Trueb, 1994). Unlike mammalian and avian species, where sexual dimorphism may reflect adaptation of the two sexes to different ecological niches

(Andersson, 1994), ecological differences between male and female amphibians are not obvious and are rarely reported in the literature. SSD in amphibians more likely reflects adaptation of the two sexes to their different reproductive roles and so is expected to be influenced explained by the differences between the two sexes in their life history.

Anurans are the most diverse clade of living amphibians, representing over 5400 of the more than 6000 species (Wells, 2007). Their great life history diversity and great variation in the ratio of female to male body size make them a good candidate to study

SSD evolution. The only comprehensive review on overall patterns of SSD in anurans is by Shine (1979). Assessing 589 anuran species, Shine concluded that a female-biased

SSD is the common pattern in anurans with only 10% species showing weak SSD or male-biased SSD. Although there are a number of studies either explaining or predicting the presence of SSD in anurans, the effect of life history on the evolution of SSD is still poorly understood.

Mating, breeding, and development related life history traits are all thought to affect the body size ratio between male and female anurans (Shine, 1979; Woolbright,

1983; Arak, 1988; Emerson; 1996; Monnet and Cherry, 2002). For mating related traits, male combat and the duration of breeding season are both considered important for the evolution of SSD. Larger male may have more ability to defeat other males during competition for females (Shine, 1979). Prolonged breeding periods may affect male body

2 size through energetic constraints on males' reproductive activities (Woolbright, 1983).

Female preference for male call frequency and the population's operational sex ratio are also expected to affect SSD (Ryan, 1980; Arak, 1988). However, male call frequency and operational sex ratio are highly influenced by environmental changes, and the existing data are not suitable for testing these hypotheses in a cross-species analysis. Although female-biased SSD owing to fecundity advantages is known (Shine, 1979), we do not know how interactions between breeding related life history traits, such as egg size, clutch size, and parental care, influence SSD. Furthermore, differences in developmental rate or developmental time between males and females may also result in relative size differences, due to the indeterminate growth in anurans (Roff, 2001). However, based on

Duallman and Trueb's (1994) summary on 45 anuran species, the ages at first reproduction did not differ between sexes in most of species in their data set. Despite this summary, Monnet and Cherry (2002) found mean female-male body size ratio was correlated with mean female-male age difference among 17 anuran species using a phylogenetic independent contrasts method. They proposed that differences between the sexes in age at maturity and longevity in the two sexes could explain SSD in anurans, because the fecundity advantage might favor females that delay reproduction. However, with only 17 species available for analyses, it remains unclear whether their findings can be extrapolated to all anurans.

This study made use of the currently available data on mating and breeding related life history traits to assess whether life history variation among anuran species influences SSD evolution. The life history traits included in the study are mating combat, length of breeding season, clutch size, egg size, clutch volume and parental care. After

3 understanding the importance of these life history traits in shaping SSD, I further tested whether SSD in anurans follows a widespread SSD evolution pattern in animal kingdom named Rensch's rule, where SSD is predicted to decrease with size when females are the larger sex, but increases with size when males are the larger sex (Rensch, 1950).

Mating Related Life History Traits

I. Mating Combat

The milestone research that tested adaptive hypotheses of sexual size dimorphism

in relation to mating combat in anurans was performed by Shine (1979). He concluded that reversed sexual size dimorphism (RSSD), where male body size is equal to or larger than female body size, was strongly correlated with the presence of male combat and

appearance of male weaponry, such as spines and tusks. However, Shine's research had

five weaknesses. First, no criteria were provided for defining a female-biased SSD or a

male-biased SSD. Second, Shine overestimated the percentage of female-biased SSD by

including species without SSD descriptions into this category. Third, Shine assumed that

secondary characters of males, such as spines or tusks, were good indicators of male

combatT because they could be used as weapons during combats. However, the spines

may not evolve as signal to indicate combat potential. For example, males of the tree frog

Hyla albomarginata, H. granosa and H. miliaria groups have projecting prepollical

spines on thumbs, but no evidence shows that they partake in combat (Duellman and

Trueb, 1994). Also, males of the spiny toad Vibrissaphora boringiae present labial spines

during the breeding season (Zheng et ai, 2007), but no record of male combat has been

4 reported for this species. Moreover, spines on males' thumbs are typically modified nuptial excrescences whose primarily function is maintaining amplexus in torrential

streams or avoiding other males' dislodgement (Duellman and Trueb, 1994). The assumption that these nuptial spines are necessarily used as weapons in male combat might not be supported by any evidence. Fourth, Shine assumed that males of a species did not partake in combat if there was no record of male combat. However, no record of

combat may simply mean that the observation on mating behavior has not been made.

Shine's assumption may therefore under estimation the frequency of male combat

behaviors. Fifth, Shine ignored the evolutionary relationships between species, and

viewed each species as an independent data point in his analyses. However, closely related species share phenotypic similarities inherited from a common ancestor (Harvey

and Pagel, 1991). Therefore, his cross-species analysis likely overestimated the true

number of evolutionary of traits. As a consequence, Shine's hypothesis of male-biased

SSD in anurans needs to be retested in a phylogenetic context that controls for the effects

of common ancestry and with more cautious classification of male combat behaviors.

In addition to these weaknesses, Shine (1979) neglected to consider two other

factors that could influence size dimorphism. The first is that different mating systems

might favor different male body size depending on the form of male-male competition.

For example, two forms of male combat are commonly observed in anurans: scramble

competition and territory defense (Wells, 1977). In scramble competition, where males

fight amongst themselves for possession of females in dense breeding aggregations, the

average male mating success is low. Large body size may not be crucial for a male's

fitness in this mating situation, because there is a trade-off of being large and being small.

•5 Large males who delay their maturity may have more chances to dislodge other males in

one breeding season; however, small males who can mature earlier are likely to have

more mating chances during their lifespan. Therefore, selection on large male body size

may not be strong in this mating system. Territory defense occurs when residents

aggressively defend calling sites or oviposition sites against intruders and females choose male territories. In this mating system, chances for a successful mating and multiple mating are higher. Larger males likely have greater mating success, because they can better defend the territories for breeding (Wells, 2007). Hence, RSSD in anurans is predicted to be correlated with male territory defense only and not related to reproductive success in species with male scramble competition breeding situations.

Second, Shine (1979) only focused on the influence of male and not female combat on SSD, probably because combat behaviors are most common among male anurans. However, in some species, such as the poison frog Dendrobates azureus and the midwife toad Alytes obstetricans, females are also known to partake in combat (Wells,

1977; Een and Pinxten, 2000). Since mating combat can also favor large body size in

females, sexual selection acting on female might increase the magnitude of SSD in

species with female combat roles.

II. Length of Breeding Season

Sexual selection alone may be inadequate to explain SSD in anurans, because it ignores factors that many constrain evolutionary responses, such as growth can be restricted by energetic limitations on growth. Woolbright (1983) proposed that prolonged breeding periods would create energetic constraints that prevented males from

6 evolutionarily responding to sexual selection favoring large body size. However, the data in his model were insufficient to affirm or reject these assumptions (Sullivan, 1984).

Duration of stay at the breeding site is essential to male mating success in some anuran species (Duellman and Trueb, 1994). For example, males of the gray treefrog Hyla chrysoscelis that spent more time calling at night mated more often (Godwin and Roble,

1983). Larger males that reserve more energy may be favored in species with a prolonged breeding. In addition, RSSD is observed in many prolonged breeding species, such as the poison frog Dendrobates pumilio (Silverstone, 1975; Savage, 2002) and the fanged frog

Limnonectes kuhlii (Chen, 1991; Yang, 1991). Therefore, if breeding season has an impact on male body size in anurans, then there should be an either positive association between increased breeding season length and increased male body size.

Breeding Related Life History Traits

I. Egg Size, Clutch Size and Clutch Volume

Larger females are often thought to have more internal space for either more eggs or larger eggs (Andersson, 1994). However, variation in egg size is low in most anurans, possibly owing to the developmental constraints of lower survival rate for smaller eggs and longer development time for larger eggs (Duellman and Trueb, 1994). Variation in clutch size on the other hand is very high within and among species. For example, clutch size varies from 580 to 11,600 in the Chinese bullfrog Hoplobatrachus chinensis, depending on female age (Chen, 1991). Therefore, large body size in females is typically associated with more eggs rather than larger eggs. Since the every a female can invest in

7 reproduction is limited, egg size and clutch size are typically negatively correlated

(Messina and Fox, 2001). So clutch volume should be a better estimator of the total reproductive fitness of a female.

II. Parental Care

Parental care, defined as any form of post-ovipositional parental investment that increases the survival of offspring (Wells, 2007), is thought to be associated with the body size of the care-providing parent and can influence SSD in many animal groups

(Messina and Fox, 2001; Shreeves and Field, in press). Selection is thought to favor larger parents because they provide more resources to and better defense of offspring, which improves offspring survival. For some mammals in which females are larger than males, large females are better at carrying or defending the young (Ralls, 1976). In wasps and bees, a high degree of female-biased SSD was found in taxa where females gather and transport heavy loads of provisions to nests (Shreeves and Field, in press). In contrast, parental care activities may result in the small body size of the care-providing parents because the energy expenditure on the breeding activities constrains the growth.

For example, studies on salamanders suggested that brooding adults in the red-backed salamander Plethodon cinereus suffered reductions in growth rates and body mass (Ng and Wilbur, 1995). However, few studies in anurans have attempted to estimate the cost of parental care relative to alternative uses of energy for growth (Wells, 2007).

In addition, selection for parental care can relax fecundity selection on female and so indirectly results in smaller female body size relative to male body size. Parental care increases survival of offspring, thereby reducing selection for high offspring production

8 from large females. Parental care may also have an evolutionary association with egg size. Larger eggs take longer to develop, leading to selection in favor of parental care behaviors that reduce the increased offspring mortality (Nussbaum, 1987). Summers, et al. (2006) investigated 640 anuran species and found a positive correlation between egg size and parental care using phylogenetic comparative methods, and concluded that large egg size typically preceded the evolution of parental care.

In many anuran species where males are larger than females, males are observed to provide care. For example, in the fanged frog genus Limnonectes, males in some species carry the tadpoles on their backs during tadpole development (Emerson, 1996).

Males also provide care by guarding egg clutches in the moustache toad Vibrissaphora boringiae (Zheng et al, in press). Male African bullfrog Pyxicephalus adspersus protect tadpoles by defending them against potential predators (Balinsky and Balinsky, 1954). In light to these observations, I predict that the presence of parental care in a taxon is associated with a decreased clutch size or an increased egg size compared to a sister taxa.

I also anticipated that the presence of parental care is correlated with an increased body size in the care providing parent. Hence, male parental care will be positively correlated with RSSD in.anurans.

Rensch's Rule

Rensch (1950) described a relationship between SSD and body size, observing that SSD decreases with size when females are the larger sex, but increases with size when males are the larger sex. It has been suggested that Rensch's rule may be driven primarily by sexual selection for large male size in combination with a typically high

9 genetic correlation in body size between the sexes (Fairbairn et al. 2007). Abouheif and

Fairbairn (1997) reviewed the evidence for Rensch's rule in many animals and found support for it in the majority of taxa. However, anurans were greatly underrepresented in their survey. Different from Rensch's rule assumption on sexual selection for large male size predominant in the evolution of body size of both sexes, fecundity selection is expected to be a dominant selective force on body size evolution in animals where females are the large sex (Andersson, 1994). For example, insects and spiders often display size dimorphism inconsistent with Rensch's rule, as females appear to have diverged more in size from males over evolutionary time (Blanckenhorn et al, 2007;

Foellmer and Moya-Larano, 2007). If the fecundity hypothesis holds in anurans, then an opposite trend of Rensch' rule may be found. Whether anurans are consistent with this prevalent rule is still unknown and can be tested by including a large number of taxa in a phylogenetic framework (Fairbairn et al, 2007).

Objectives

In the last two decades, data on body size and life history have accumulated for many anuran species. Also, the advent of high-throughput sequence analyses and the development of phylogenetic methods that lead to better molecular phylogenies provides an excellent opportunity to re-examine the role of life history in shaping SSD utilizing phylogenetically corrected analysis. This study has three objectives. First, I will assess the distribution and pattern of SSD among anurans to test Shine's assertion that 90% of anuran species have female-biased size dimorphism. Second, I will test Shine's prediction that RSSD is correlated with male combat, and further examine whether

10 correlated evolution between SSD and mating and breeding related life history traits

occur in anurans, including male scramble competition, male territory defense, length of

breeding season, egg size, clutch size, clutch volume and parental care. Finally, I will test

for Rensch's rule in anurans. All my tests will control for evolutionary relations among

species.

On the basis of traditional assumptions about SSD evolution and observations of

SSD in anurans, the hypothesized correlations among SSD, body size and life history

traits are represented in Figure 1. Female combat, egg size, clutch size, clutch volume,

and female parental care are expected to be positively correlated with female body size

and so result in the evolution of a female-biased-SSD. On the other hand, male combat, prolonged breeding season and male parental care are expected to be positively correlated with male body size and so contribute to the evolution of a male-biased SSD. Alternate

forms of male-male competition are expected to have different effects on SSD. Male

scramble competition is not expected to be associated with SSD, while male territory defense is expected to.be positively related to SSD. Rensch's rule predicts a greater

interspecific variation in male body size compared to female body size, and a negative correlation between the ratio of female to male body size and the mean body size of a species.

11 Body Size

Female/Male Body Size Ratio

Female Body Size Male Body Size

Femal Combat Clutch Volume Parental Care Breeding Season Scramble Competition Territory Defense

Egg Size Clutch Size

Figure 1 Hypothetical relations among body size ratio (F/M), body size, female combat, male scramble competition, male territory defense, breeding season, parental care (female parental care and male parental care), clutch volume, egg size, and clutch size in anurans. See text for detailed discussion of hypotheses used to generate this predicted relations.

12 METHODS

Taxa Included for Comparative Analyses

Body size and life history data were collected from 545 anuran species for comparative analyses (Appendix 1; Appendix 2). Most species were from a superfamily and five families: Superfamily Dendrobatoidea (family Aromobatidae and family

Dendrobatidae; 62 species), Family Bufonidae (58 species), Family Dicroglossidae (27 species), Family Hylidae (198 species), Family Megophryidae (25 species) and Family

Ranidae (81 species). The remaining 94 species were scattered across 30 other families.

Species names and family names follow the "Amphibian species of the world 4.0" website as of December 2006 (Frost, 2006).

Body Size and Life History Traits

I. Body Size

Snout-vent length (SVL) in millimeters was used to represent body size. Female and male body sizes in a species were collected separately. Different publications often provide different levels of details for the body size data, and consequently, the data were separated into three categories of decreasing precision: mean with standard deviation

(mean with SD), mean without standard deviation (mean without SD), and range data

(minimum and maximum values). "Mean with SD" referred to body size data given as a mean SVL with a standard deviation. Data with a mean SVL that lacked a standard deviation fell into the category "mean without SD". Samples sizes for data in these two

13 categories had to consist of measurements of SVL for no less than three individuals in each sex per species. "Range" included body size of a species provided as maximum and minimum value, and also included body size measured on less than three individuals in each sex. Mean data can represent body size more precisely than range data as it indicates the most common body size, so only data in the first two categories were used to estimate body size for each sex. If more than one study described body size for two sexes and provided mean values (with or without SD), then a mean body size across studies was calculated for each sex. Body size of a species was represented by the mean body size of both sexes.

II. Sexual Size Dimorphism

Sexual size dimorphism (SSD) was treated as either a continuous trait or a discrete trait according to the categories of body size data (Table 1) and as such used in two different comparative analyses. SSD treated as a continuous trait was estimated as the ratio of female to male body size ratio, where female body size (SVL mm) was divided by male body size (SVL mm). This required the mean body size to be known for each sex. Estimating SSD using a body size ratio is preferred irt many studies, because it indicates the direction and degree of dimorphism (Fairbairn et ah, 2007). However, it excludes many species where body size data are provided only as a range.

14 Table 1 Summary of character states for body size and body size dimorphism.

Traits Data fype Data Category Mean Mean with SD Range without SD Maximum and minimum value Mean value & Mean value (n>=3); SD (n>=3) Measurement of (n>=3) individuals (n<3) Body Size of Mean of two Mean of two Continuous N/A Species sexes sexes Body size Body size Continuous N/A ratio (F/M) ratio (F/M) Following RSSD One way t-test Body size Body Size authors' Present onF>M ratio (F/M) Dimorphism 64 1 5) descriptions of p>=0.05 <1.02 of Species Discrete RSSD RSSD One way t-test Body size No evidence of Absent onF>M ratio (F/M) RSSD "0" p<0.05 >=1.02

Note. SD = standard deviation; RSSD = Reversed sexual size dimorphism (male-biased sexual size dimorphism).

15 In order to make use of range data, SSD was also estimated as a discrete trait, by coding the presence or absence of larger males into "1" or "0". The common state of SSD in anurans is female biased, species with only range data were assigned "0" unless authors' descriptions suggested otherwise. As mentioned above, body size ratio (F/M) was used to estimate SSD in species with mean data. In order to treat SSD as a discrete trait, a transformation of body size ratio (F/M) into binary states was required and was conducted in two different ways. In 168 species where "mean with SD" body size data was available, the presence or absence of large males was determined using a one tailed t- test between two sexes at a 0.05 significance level. T-tests could not be conducted for the species with "mean without SD" body size data and so a threshold was needed to assign the body size ratio (F/M) into either of two character states. The threshold body size ratio

(F/M) was derived by the t-test results of "mean with SD" data above. Among 119 species not having a reversed sexual size dimorphism (RSSD), the lowest body size ratio

(F/M) was 1.02. Using this value as the threshold, an overestimation of the incidence of

SSD could be avoided. Therefore, body size ratios (F/M) greater than or equal to 1.02 were coded as "0" (female-biased SSD), and body size ratio (F/M) less than 1.02 were coded as "1" (male-biased SSD). Species without sexual dimorphism information were excluded from analyses.

16 III. Mating Combat

The presence (1) or absence (0) of mating combat was classified based on reviewing the literature on mating behavior in each species. The presence of mating combat was defined as observing aggressive physical contests between individual of the same sex to gain access to mates or attractive territories (Futuyma, 1993). Field observations recorded under natural conditions were prefered, but combat behaviors from laboratory experiments were also included when necessary. For example, males of poison frog Colostethus awa aggressively jumped to a recorder playing conspecific courtship calls (Coloma, 1995) and this was considered as male combat for territory. Aggressive behaviors without physical contact, such as aggressive calls, defensive postures and territorial behaviors or defensive behaviors against predators were not considered as mating combat. The absence of a record of mating combat may occur for two reasons: either the species does not partake in mating combat or no observations have been made to date. To avoid incorrectly placing species into the absence category because of no proper observations, I only included species for which other mating information had been recorded, but which lacked evidence of combat. This mating information included descriptions on nest building, mate searching, mating call, and breeding season. Species with no mating information (10.46%) were removed from the correlation analyses between mating combat and other traits.

Male combat and female combat were treated as different traits and were separately analyzed when testing the correlations between mating combat and body size as well as the correlations between mating combat and SSD. Species, in which both

17 males and females exhibited combat behavior, were included in both male and female

combat analyses.

Male combat was artificially divided into two types: scramble competition and

territory defense, according to descriptions in the literature on male mating strategies and

the purpose of combat (Wells, 1977). These two types of combat were analyzed

separately to test if different male combat strategies were correlated with male body size

and SSD. Species where males were observed to have both combat forms were included

in both analyses. A separate analysis of the two types of mating combat was not applied

to females where combat was rare.

IV. Length of Breeding Season

Following Wells (1977), one month was used as a cut-off value to categorize

length of breeding season into a binary trait. Breeding seasons shorter than one month

were considered as "explosive" (0), and longer than one month were considered as

"prolonged" (1). The length of breeding season in this study was the breeding period of a

single population rather than the breeding season of a species as a whole, as some species

may breed over several months, if individual populations have different explosive breeding periods. If breeding periods varied among populations, the species were

included in both the explosive and prolonged categories. When the length of breeding

season was not clearly described in the literature, a species was assumed to be an

explosive breeder or a prolonged breeder based on the following supporting evidence.

Large congregations in breeding ponds, sharp emergence after heavy rainfall, or quick

larval development were considered to be traits associated with explosive breeding. Low

18 density of mating congregation, no mating congregation, or long-term larval development was assumed to reflect prolonged breeders.

V. Egg Size, Clutch Size, and Clutch Volume

Egg size was considered to be the diameter of the ovum (millimetres). Egg size data were divided into two types: 1) mean ovum size of a clutch, and 2) range for ovum size for multiple clutches. The mean data were preferentially used to generate egg size values, and the median of a range was also accepted if mean data were not available.

When more than one publication had data for a single species, a mean egg size was estimated across studies to represent egg size of the species.

Clutch size was defined as the total number of eggs a female laid in one year, and was used to estimate annual fecundity of a species (Rose and Mueller, 2006). Similar to egg size, a single clutch size value was generated using the mean value of a species, or when unavailable, the median value of the range for a species. Mean value across sources was also calculated for species with multiple records. When a species produced more than one clutch in a year, annual clutch size was generated as mean clutch size times the number of clutches laid per year.

Egg size and clutch size are both related to fecundity, but are also negatively correlated with each other in many anuran species (Wells, 2007). To estimate the total ovum-producing efforts of females, the covariance between egg size and clutch size was controlled by calculating annual clutch volume, given by mean or median egg size multiplied by mean or median annual clutch size.

19 VI. Parental Care

Parental care in anurans was classified as either present (1) or absent (0) by

reviewing literature descriptions on parental attendance of clutches, building of foam

nests when laying eggs, carrying eggs, transporting tadpoles, larval developing in parents' bodies, and female deposition of unfertilized eggs to feed tadpoles (Wells,

2007). Any one or combination of these behaviours was considered as evidence of parental care. The absence of parental care was defined as no evidence of any of these behaviors only when other breeding information was also available. This breeding

information included descriptions of breeding habitat, egg development habitat, breeding behaviors during egg laying, egg development time, egg size or clutch size. Species with no breeding information (23.30%) were excluded from analyses.

When testing correlations between parental care and fecundity traits, both female parental care and male parental care were considered to be parental care in a

species. Female parental care and male parental care were analyzed separately in tests of the correlation between parental care and body size or SSD. Species with both parents exhibiting care were included in both analyses.

Phylogeny Reconstructions

Since species cannot be treated as statistically independent because of their shared evolutionary history, I employed phylogenetic comparative methods (PCMs) to test correlations between traits (Harvey and Pagel, 1991). This differs from traditional correlation analysis by controlling for phylogenetic associations among species. Since

PCMs require mapping phenotypic traits of species to a phylogenetic tree, I built a

20 supertree representing the phylogenetie relationships of as many species from my data set as possible.

I. Criteria for Choosing Phylogenies

The following preferences were used to choose phylogenies from published resources. First, if the phylogenies were constructed using molecular characters, I preferentially chose the ones based on more than 1000 base pairs of DNA sequences.

Second, phylogenies that covered large number of taxa were preferred to phylogenies of small groups. Third, when several different phylogenies were published on the same group of organisms, the most recent phylogeny was used. Fourth, I preferred phylogenies that were analyzed using maximum likelihood, maximum parsimony or Bayesian method. I started by constructing a family supertree of seven families relatively well studied in terms of evolutionary relationships and life history, and then added family supertrees into a framework phylogeny of Anura. The final supertree construction utilized the following references for the following groups: Anura: Frost et al, 2006; Bufonidae:

Pramuk, 2006, Cunningham and Cherry, 2004, and Pauly et al, 2004; Dendrobatiodea

(including family Dendrobatidae and family Aro'mobatidae): Grant et al, 2006;

Dicroglossidae: Roelants et al, 2004, Bossuyt et al, 2006, Emerson et al, 2000, and

Evans etal, 2003; Hylidae: Wiens et al, 2006; Megophryidae: Fu et al, 2007 and Zheng et al, 2007; and Ranidae: Bossuyt et al, 2006, Che et al, 2007abc, Roelants et al, 2004,

Veith et al, 2003, Hillis and Wilcox, 2005, and Matsui et al, 2006.

21 II. Supertree Constructions

Supertrees were constructed manually. The congruent phylogenies for the same anuran group were combined to increase the number of species in the analyses. Hard polytomies, where conflicting information on species relationships occurred among different phylogenies, were solved by preferentially choosing phylogenies with higher branch support values, or constructed based on longer DNA sequences. This solution of hard polytomies differs from current supertree construction programs, such as RadCon

(Thorley and Page, 2000). RadCon cannot solve conflicting phylogenies; rather, it generates a strict reduced cladistic consensus supertree by excluding taxa that create ambiguous relationship of the remaining taxa. Since there were many conflicts between the phylogenies included in this study, a supertree produced by this program would contain a large number of unsolved nodes, and so this approach was precluded in this study. The final anuran supertree was close to full resolution, although two soft polytomies remained in the family Ranidae, where not enough information was available on the relationships among species.

Branch lengths were not available for the supertrees, since they were constructed from the results of multiple phylogenetic analyses based on different characters.

Therefore, I set all branch lengths as equal. The phylogenies of Dendrobatoidae and

Hylidae were obtained from original phylogenetic studies, where the branch lengths could be obtained by re-analyzing the DNA data provided by the authors (Grant et al.,

2006; Wiens et al, 2006). However, branch lengths in these two phylogenies cannot accurately represent the number of character changes along branches due to the incomplete sequence for most taxa (the amount of sequence analyzed per taxon varied

22 from 426 bp to 6245 bp in Dendrobatiodea and from 316 bp to 8276 bp in Hylidae).

Many taxa did not share the same DNA characters or only shared a small portion of DNA sequences. To test if the results of comparative analyses were different using these inaccurate branch lengths and using equal branch lengths, I analyzed data in Hylidae by applying two different branch length treatments: 1) using branch lengths generated by maximum likelihood method based on DNA sequence data provided by Dr. John Wiens, and 2) setting all branch lengths to be equal. The two sets of results were highly congruent. Therefore, the branch lengths in the Dendrobatoidae and Hylidae phylogenies were not used and all branch lengths were set as equal in the analyses.

Anuran classification is undergoing major rearrangements (Frost et al, 2006).

According to the current systematic literature, some formerly recognized species were split into several species. For example, the poison frog Dendrobates ventrimaculata includes five undescribed species, one at Rio Ituxi, Brazil, a second at Manaus, Brazil, a third at Leticia, Colombia and French Guiana, a fourth at Pompeya, Ecuador, and a fifth at Porto Walter, Brazil (Grant, et al. 2006). Since body size (Rodriguez and Duellman,

1994) and life history data (Poelman and Dicke, 2006) in the current study regarding D. ventrimaculata matched the French Guiana clade, this clade was chosen to represent the species Dendrobates ventrimaculata. The phylogenetic supertree for all anurans is represented in Appendix 3.

23 Phylogenetic Comparative Analyses

I. Phylogenetic Independent Contrasts

Felsenstein (1985) first proposed the use of phylogenetic independent contrasts

(PIC). PIC removes statistical dependence among species by comparing standardized

differences (contrasts) between characters of two daughter taxa of each node in a

phylogeny (Garland et ah, 1992). The process of computing contrasts begins at the tips of

the tree and progresses to the root. It allows n-1 independent contrasts to be computed

from a data set of n species. Regression of standardized contrasts through the origin

assesses a correlation between two traits by testing if the slope differs significantly from

zero (Garland et al., 1992).

. To calculate the value at each node, PIC assumes the evolution of characters

along branches of a phylogeny can be modeled using a Brownian motion process

(random walk) with the character change variance accumulating linearly with time. The

value of each daughter taxa is weighted by its branch length, which estimates the total variance of character change. The value at each node is calculated as a weighted sum of the values for the two daughter taxa as:

Xk=(vjXi + ViXj)/(vi + Vj), (1)

Where the X is the value of a variable at the node or species, and v \ or j is the branch

length between the daughter taxon Xi or) and its ancestral node XR. The branch length

leading to the ancestral node Vk is lengthened, reflecting the error in the estimation of Xk.

Thus the lengthened branch length Vk' is given by:

vk' = vk + (vi VJ) / (vi + Vj). (2)

24 For example, in Figure 2, the trait value for node X4 is calculated as (V2 Xi + V| X2) / (vi + v2) = 2.8, the branch length leading to this node is v4' = v4 + (vi v2) / (vi + v2) = 1.8. The variance of all the sets of contrasts should be standardized so as to apply to regression analyses (Garland et ah, 1992). The standardized contrasts are calculated as contrast values divided by the square root of the sum of the branch lengths linking the two daughter taxa to their ancestral node as:

(Xi-XjV.VCvi + Vj) (3)

A PIC is implemented in the computer program, Comparative Analysis by

Independent Contrasts (CAIC v 2.6.9) (Purvis and Rambaut, 1995). This program outputs all independent standardized contrasts and parameters in regression analyses. CAIC is more powerful than likelihood based phylogenetic comparative methods, such as

BayesContinuous (Pagel, 1998; Pagel, 1999), since it is easier for operators to detect outlier contrasts in regressions. Procedures CRUNCH and BRUNCH are provided in this program for analyzing two different types of data.

25 V2=1 Xi=2

X2=3

X3=10

Figure 2 A hypothetical phylogeny among three taxa. X; represents the value for the node or species i. The branch length between the node or species i and its ancestral node is labeled as Vj.

26 The CRUNCH procedure can test for a correlation between two continuous characters using the PIC method. It requires operators to set a predictor (independent) variable in the regression statistics. In addition, CRUNCH handles soft polytomies using the mean value among all taxa in the unsolved clade to assign the taxa into two groups and calculating a contrast between the groups (Pagel, 1992). To do this, the mean of independent variable X for all the daughter taxa is calculated, then those taxa with X above this mean value are put into a group or a sub-node and the rest are assigned into another group (Figure 3). Within each sub-node, the mean value for each character is calculated as a weighted mean by:

Xk=2(l/vi)Xi/S(l/.Vi). (4)

The new branch length leading to the sub-node vk is arbitrarily set to 1.0 initially and then the branch length is transformed according to:

.vk'=.vk+l/2(l/vi). (5)

As the example in Figure 3 shows, taxon 1 and taxon 3 are assigned into one sub-node because their values are lower than the mean value 5. The value of this new sub-node is

Xa= {[l/(v,-l)]Xi + [l/(v3-l)]X3} /[l/(v1-l) + (17v3-l)] = 2.8.The branch length of va' = 1 •+ 1 / [ 1 / (vrl) + (1 / v3 -1) ] =1.8.

27 A B Vi=5 V1 - 1 = 4 X1 = 2 Xi=2

Va =J^X< v3-1=1 X4 / v2=6

X2=10 x^C X3 = 3

S \VV2 = 6

X3 = 3 X2=10

Figure 3 A, A hypothetical polytomy among three taxa. The value for the node or species i is represented by X;. The branch length between the node or species i and its ancestor node is labeled as v,; B, Two sub-nodes assigned by CAIC.

28 In the BRUNCH procedure, CAIC can test the correlation between one binary trait and one continuous trait when the binary trait is the predictor variable. It estimates a mean value of the continuous traits on the clade, which share the same state with the binary traits. The formula for calculating this value and the related branch lengths are identical to formulae 4 and 5. Pairs of species that differ in binary traits are compared independently along the phylogeny. By doing this, BRUNCH tests if the higher value of the continuous trait is correlated with a certain state of the binary trait or vice versa. In this procedure, the categorical trait is automatically set to be independent, because the contrasts only take place on the nodes where the state of categorical traits differs.

II. BayesDiscrete

I employed BayesDiscrete method, which was implemented in the program

BayesTraits, for testing correlated evolution between pairs of binary traits using maximum likelihood methods (Pagel, 1994; Pagel and Meade, 2006). Correlated evolution of two traits is tested by comparing the fits (log-likelihood) of the independent model and the dependent model. These two models calculate the likelihood of different states in two traits at each node on a phylogeny by estimating transition rate parameters according to branch lengths.

In the independent model, two transition rates are estimated per trait. Alpha represents the rate of state changing from "0" to "1" and beta is for the rate' of state changing from "1" to "0". The dependent model allows the traits to evolve in a correlated fashion such that the rate of change in one trait depends upon the background state of the other. The dependent model can "adopt four states (00, 01, 10, 11). The transition rates are

29 represented by eight parameters (ql2, q21, ql3, q31, q24, q42, q34, q43), which describe all possible changes in one state holding the other constant (Figure 4). If the likelihood of observing the trait data under the model of dependent trait evolution exceeds that for the model of independent trait evolution, then the state change of one trait depends on the state of the other.

The null hypothesis of no correlation is rejected in favor of the view that the traits have coevolved. The likelihood ratio (LR) that tests the goodness of fit of two models to the data is defined as

LR = -2 loge [ Independent Model / Dependent Model ]

The LR statistic is distributed as a chi-squared variant with four degrees of freedom and evaluated at a 0.05 significance level. In order to reduce the variance between runs, I increased the "multries" parameter to 25 following the suggestions in the program manual (Pagel and Meade, 2006), to increase the precision of the optimization algorithm.

I re-analyzed the data with all alternative solutions for the polytomies, on the requirement of bifurcating branching structure of the phylogeny in BayesTraits. I found no significant difference between results when testing all these possible topologies of polytomies, and so only representative results are reported for each correlation in Appendix 4.

30 Independent transitions in two binary traits a1 Y 0 1 |31

a2 X 0 <- 1 |32

Dependent transitions between two binary traits

X.Y q12 0,0 ^ 0,1 q21

q31 q13 q42 q24

q34 1,0. 1,1 q43

Figure 4 Parameters estimated in the independent model (upper) and the dependent (lower) model in BayesDiscrete.

31 III. Comparative Analyses

All correlation analyses performed are listed in Table 2, along with the information on analytical programs used and variable settings. Anuran families differ in mating systems and reproductive strategies (Wells, 1977), and so the correlated evolution of certain life history traits may be specific to certain families. In order to detect family- level correlations, analyses on each pair of traits were conducted at two levels: across all anuran species (all-anuran analysis) and within each family (superfamily), where supertrees had been built (within-family analyses). Within-family analyses were conducted in the families Bufonidae, Dicroglossidae, Hylidae, Megophryidae, Ranidae and superfamily Dendrobatoidea. As in Grant et al. (2006), the superfamily

Dendrobatoidea consists of two families Dendrobatidae and Aromobatidae. Frogs in this superfamily shared many life history similarities, so they were analyzed together

(Duellman and Trueb, 1994). Missing data was pruned from the comparative analyses.

Consequently, for each independent analysis between two traits, only species with a complete data set for these traits were included. Since the data of each species was not consistently used in every comparison, a Bonferroni correction, which adjusts significance levels for each test depending on the number of tests, performed on the same data set, was not applied (Zar, 1996).

32 Table 2 Phylogenetic comparative analyses, indicating correlations tested, corresponding programs applied and predictor variables used.

Correlations Traits tested for correlations Analysis Programs Predictor variables Log male body size & male combat CAIC (BRUNCH) Male combat RSSD & male combat BayesDiscrete N/A SSD and mating Log male body size & male scramble competition CAIC (BRUNCH) Male scramble competition related life history Log male body size & male territory defense CAIC (BRUNCH) Male territory defense traits RSSD & male territory defense BayesDiscrete N/A ' Male body size & prolonged breeding season CAIC (BRUNCH) Breeding season RSSD & prolonged breeding season BayesDiscrete N/A Log female body size & egg size CAIC (CRUNCH) Egg size Female body size & log clutch size CAIC (CRUNCH) Clutch size Female body size & log clutch volume CAIC (CRUNCH) Clutch volume SSD and breeding Body size ratio (F/M) & log egg size CAIC (CRUNCH) Egg size related life history Body size ratio (F/M) & log clutch size CAIC (CRUNCH) Clutch size traits Body size ratio (F/M) & log clutch volume CAIC (CRUNCH) Clutch volume Log male body size & male parental care CAIC (BRUNCH) Male parental care Log female body size & female parental care CAIC (BRUNCH) Female parental care RSSD & male parental care BayesDiscrete N/A

33 Correlation Egg size & log clutch size CAIC (CRUNCH) Egg size between egg size, Parental care & log egg size CMC (BRUNCH) Parental care clutch size and parental care Parental care & log clutch size CAIC (BRUNCH) Parental care SSD and body Log male body size & log female body size CAIC (CRUNCH) Female body size* size Body size ratio (F/M) & log body size CAIC (CRUNCH) Body size

Note. SSD = Sexual Size Dimorphism (female-biased SSD); CAIC = the Program of Comparative Analysis by Independent Contrasts;

RSSD = Reversed Sexual Size Dimorphism (male-biased SSD); *Female body size was set as the predictor variable in the female and male body size correlation analysis, because female body size was assumed to diverge faster than male body size in anurans (refer to

Discussion).

34 RESULTS

Sexual Size Dimorphism

Body size distribution was positively skewed in anurans (mean=45.89mm,

SD=21.26, n=259 species), and ranged from 13.35mm in the blue-bellied poison frog

Dendrobates minutus (Silverstone, 1975) to 134.05mm in the Surinam toad Pipapipa

(Trueb and Cannatella, 1986) (Figure 5). The distribution of the body size ratio (F/M) in

anurans was nearly normal (mean=1.15, SD=0.15, n=259 species) (Figure 6). The mean

of body size ratio (F/M) was larger than 1.0, indicating that sexual size dimorphism

(SSD) was biased towards females. Male size exceeds the size of conspecific females

only in eight species. By applying the body size threshold ratio of 1.02, 62 species show reversed sexual size dimorphism (RSSD). It included 23.9% of anurans with size

dimorphism data. The degree of reversed sexual size dimorphism (RSSD) was much

lower than the degree of normal sexual size dimorphism. Females could be two times

(188.31%) bigger than males in the odorous frog Rana schmacker (Ye et al, 1993).

However, the skipping frog Euphlyctis cyanophlyctis demonstrated the most extreme

RSSD in anurans, where male size was 38% larger than conspecific female's size

(Gramapurohit et al, 2005).

Species with RSSD were present in 18 different anuran families (Aromobatidae,

Bufonidae, Bombinatoridae, Centrolenidae, Cryptobatrachidae, Dendrobatidae,

Dicroglossidae, Hylidae, Leiuperidae, Leptodactylidae, Limnodynastidae, Megophryidae,

Microhylinae, Myobatrachidae, Pelobatidae, Pipidae, Ranidae, and Scaphiopodidae) and

35 2.00E-02 1.80E-02 A 1.60E-02 : Jf \ 1.40E-02 - / > 1.20E-02 c • = l.OOE-02 -

2.00E-03 - \ O.OOE+00 - *••• • () 50 100 150 Body Size (mm)

Figure 5 Frequency distribution of body size in anurans with mean=45.89mm, SD=21.26, and n=259 species.

36 3.00E+00 -|

2.50E+00 - f \ • 2.00E+00 - !

• nc y $ 3 1.50E+00 -

• \ 5.00E-01 - • •

• • • ^ 4 O.OOE+00 - • i 0 0.5 1 1.5 2 Body Size Ratio (Fe male/Male)

Figure 6 Frequency distribution of body size ratio (F/M) in anurans with mean=1.15, SD=0.15 and n=259 species.

37 distributed across six continents (Asia, Africa, Australia, Europe, North America, and

South America) in both temperate and tropical zones (data not shown).

SSD and Mating Related Life History Traits

I. Correlation between SSD and Mating Combat

Mating combat behaviors have been recorded in 101 (20.53%) of the 492 species where mating descriptions are available. Male combat was more prevalent than female combat. For example, 98 species exhibit male combat, whereas only 12 species show female combat. Among the species having male combat, 42 species (41.58%) have scramble competition, 54 (53.47%) of them defend territories and three species (2.97%) use both of these combat forms.

A positive correlation between male body size and male combat was found in the superfamily Dendrobatiodea (p=0.0061, R2=0.63, slope =0.09, n=8) (Appendix 4, Table

1). This correlation was not detected in any other family or in the all-anuran analysis. In addition, female body size was positively correlated with female combat in

Dendrobatoidea (pO.OOOl, R2=0.50, slope =0.07, n=8). A positive correlation between female body size and female combat was also detected in the all-anurans analysis

(p=0.044, R2=0.46, slope =0.07, n=8). However, this correlation was driven by the species with female combats in Dendrobatoidea, as most female combats occurred in this superfamily. No significant correlation was found between male body size and male scramble competition (p=0.43, n=21) or male territory defense (p=0.36, n=25).

38 Body size ratio (F/M) was not correlated with male or female mating combat in either the all-anuran analysis or any of the within-family analyses (Appendix 4, Table 2).

The likelihood ratios in BayesDiscrete analyses did not suggest a dependent evolution between RSSD and male combat. Also, no correlation between body size ratio (F/M) and male scramble competition was found. Likewise, no dependent evolution between RSSD and territory defense was detected.

II. Correlation Between SSD and Length of Breeding Season

No correlation was found between male body size and length of breeding season in the all-anuran analysis and any of the within-family analyses (Appendix 4, Table 1). If breeding season was not associated with male body size, it should not contribute to the evolution of size dimorphism. Congruent with this prediction, no correlation between

SSD and length of breeding season was found in either all-anuran analysis or within- family analyses in both CAIC and BayesDiscrete analyses (Appendix 4, Table 2).

SSD and Breeding Related Life History Traits

I. Correlation between SSD and Clutch Volume

Egg size was negatively correlated with clutch size (p=0.0085, R2=0.03, slope=-

0.33, n=223) (Appendix 4, Table 5). To estimate the total reproductive benefit for females from increased body size, the correlation between female body size and clutch volume was tested. Female body size was positively correlated with clutch volume in the all-anuran analysis (pO.OOOl, R2=0.38, slope=8.23, n=125) and in four of six anuran families (superfamily) (Dendrobatoidea: p=0.004, R2=0.51, slope=8.20, n=13; Hylidae:

39 pO.OOOl, R2=0.53, slope=7.62, n=44; Megophryidae: p<0.0024, R2=0.75, slope=23.47, n=8; Ranidae: p=0.0001, R2=0.46, slope=10.18, n=27) (Appendix 4, Table 3).

II. Correlation between SSD and Egg Size

Positive correlations between female body size and egg size were found in the all- anuran analysis (pO.OOOl, R2=0.18, slope=0.01, n=157) and in the families

Dicroglossidae (p=0.049, R20.33, slope= 0.21, n=l 1), Hylidae (pO.OOOl, R2=0.53, slope=7.62, n=54), Megophryidae (p=0.025, R2=0.49, slope= 0.40, n=9), and Ranidae

(p=0.044, R =0.12, slope= 0.17, n=32), but not in the family Bufonidae and superfamily

Dendrobatiodea (Appendix 4, Table 3). The significant correlation between female body size and egg size in the all-anuran analysis and in the family Hylidae was possibly due to outliers. After removing these outlier contrasts, the p-values in the regressions increased to 0.08 in the all-anuran analysis and 0.76 in the Hylidae, and the slopes were no longer significant. The occurrence of outliers might reflect the lack of body size and egg size data for many anuran species, which lead to an underestimation of branch lengths close to the root. After estimating missing values using mean body size and egg size value in each clade, the correlations were all still significant (All-anuran analysis: pO.OOOl, R2=0.085, slope=0.20, n=534; Hylidae: pO.OOOl, R2=0.16, slope=0.35, n=179). It was likely that with more body size data, these two outliers would not be dissimilar with other contrasts, and so the outliers were kept in for regression analyses.

The correlation between female body size and egg size might have contributed to body size dimorphism in the family Hylidae, because the body size ratio (F/M) was also positively correlated with egg size (p=0.015, R2=0.11, slope=0.22, n=54) (Appendix 4,

40 Table 4). However, the correlation between body size ratio (F/M) and egg size was not found in other within-family analyses or in the all-anuran analysis.

III. Correlation between SSD and Clutch Size

Increased of female body size was correlated with increased clutch size in the all anuran analysis (pO.OOOl, R2=0.33, slope=7.02, n=141) (Appendix 4, Table 3). This positive correlation was prevalent in many anuran families (superfamily), such as the superfamily Dendrobatoidea (p=0.003, R2=0.35, slope=7-5, n=22), and the families

Hylidae (pO.OOOl, R2=0.48, slope=6.65, n=46), Megophryidae (p<0.0094, R2=0.64, slope=25.87, n=8), and Ranidae (pO.OOOl, R2=0.048, slope=9.55, n=29).

The positive correlation between female body size and clutch size might have contributed to the correlation between the body size ratio (F/M) and clutch size in the all- anuran analysis (pO.OOOl, R =0.15, slope=0.053, n=T41), as well as in the superfamily

Dendrobatoidea (p=0.007, R2=0.30, slope=0.035, n=22) and the family Hylidae

(pO.OOOl, R2=0.37, slope=0.079, n=46) (Appendix 4, Table 4). Two outliers might drive the correlation between female body size and clutch size in Hylidae. Since the correlation was still significant after filling in missing values of female body size and clutch size with the clade mean value, these outliers were kept in the analysis.

IV. Correlation between SSD and Parental Care

Among 492 species with breeding information, 98 species (19.92%) demonstrated parental care. Among them, 22 species (22.45%) exhibited female parental care, 61 species (62.24%) showed male parental care, and the other 15 species (15.31%) showed

41 care from both parents. No correlation was found between female body size and female

parental care or male body size and male parental care (Appendix 4, Table 3). A

correlation between egg size and parental care was not found (p=0.38, n=32) (Appendix

4, Table 5). In contrast, the presence of parental care was negatively correlated with

clutch size in anurans (p=0.01, R2=0.19, slope=-0.33, n=35) (Appendix 4, Table 5).

However, no correlation was found between the body size ratio (F/M) and female or male

parental care (Appendix 4, Table 4). In addition, the likelihood ratios in BayesDiscrete

did not provide strong evidence on dependent evolution between RSSD and male parental

care (Appendix 4, Table 4).

Rensch's Rule

The independent contrasts of female body size and male body size were

significantly correlated with each Other in all of the within-family analyses and the all- anuran analysis when female body size was used as an independent variable (Anura: pO.0001, R2=0.96, slope=1.01, n=247; Bufonidae: p<0.0001, R2=0.98, slope= 0.916, n=16; Dendrobatiodea: pO.0001, R2=0.95, slope=1.04, n=42; Dicroglossidae: p<0.0001,

R2=0.91, slope=0.85, n=15; Hylidae: pO.0001, R2=0.98, slope=1.01, n=89;

Megophryidae: pO.0001, R2=0.65, slope=1.14, n=14; Ranidae: p<0.0001, R2=0.72, slope=0.98, n=39) (Appendix 4, Table 6). The interspecific variances of female body size

(SD=22.60) were significantly greater than the variances of male body size (SD=21.31)

(variance ratio test, p<0.05), indicating that female body size was more variable. Most anuran families followed this trend, however in Dicroglossidae, male size was more variable than female size (female SD=23.03, male SD=25.27; variance ratio test, p<0.05).

42 A positive correlation was detected between SSD and body size in the all-anuran

analysis (pO.OOOl, R2=0.33, slope=0.20, n=247) (Appendix 4, Table 6). This positive

correlation was also found in the family Bufonidae (p=0.0231, R2=0.30, slope=0.16,

n=16). On the contrary, there was a negative correlation between these two traits in the

family Dicroglossidae (p=0.0319, R2=0.29, slope=-0.18, n=15). A positive correlation

between SSD and body size was also found in Hylidae, but this correlation was driven by two outliers (pO.OOOl, R2=0.48, slope=0.21, n=89). After removing them from the regression analysis, the slope was not significant different from zero (p=0.69, R2=0.002, n=87). Both of these outliers were in the Hypsiboas siblezi clade, and there is currently no biological explanation for their occurrence, but missing body size data in many species may be a potential reason. After filling the missing values using the mean body size and body size ratio (F/M) values in each clade, the correlation became non significant (p=0.84, n=544), and so I removed these two outliers from the regression analysis.

Summary of Correlations

All statistically significant correlations found in this study are summarized in

Figure 7. Male body size was positively associated with male combat, but this did not result in SSD. Also, male body size was not correlated with either male scramble competition or territory defense. No correlation was found between male body size and length of breeding season. Female body size was positively correlated with female combat, egg size, clutch size and clutch volume in anurans. Among these traits, female combat did not contribute to the evolution of SSD, but egg size, clutch size and clutch

43 Body Size

An + Di Bu

Female/Male Body Size Ratio + ^^~ ^~-»-^^—

/ 1 + AnDeHy Female Body Size Male Body Size •+/ • + /' Hy/ An/ + \ + D / 7 De An/ ** / 7 )/An / Male Combat \De / (Scramble Competition Femal Combat \Hy / & Territory Defense) Me / 1 ~ \Me 7 1 \Ra / Ra / An — An Egg Size Clutch Size Parental Care

•^ + / "t> •

Clutch Volume

Figure 7 Correlations between sexual size dimorphism, body size and life history traits in anurans. An: all anurans; Bu: Bufonidae;

De: Dendrobatoidea; Di: Dicroglossidae; Hy: Hylidae; Me: Megophryidae; Ra: Ranidae.

44 volume were positively correlated with SSD in the all-anuran analysis and in some families. Parental care was not correlated with body size and egg size, but was negatively correlated with clutch size, so it might have indirectly contributed to SSD. SSD was positively correlated with body size in the all-anuran analysis and in the family

Bufonidae, but negatively correlated with body size in the family Dicroglossidae.

' Fourteen pair-wise comparisons of the 15 traits of interest were statistically significant and are listed in order of significance in Table 3. Among the 14 significant correlations, four were still statistically significant after applying the Bonferroni correction, including the correlations between female and male body size, female body size and clutch size, body size ratio (F/M) and clutch size, and female body size and female combat. However, other p-values were still suggestive even though they were not significant after Bonferroni corrections (refer to Methods).

45 Table 3 Correlations ranked by the order of statistical significance (high to low).

Trend of Correlation Tested Groups P Correlation Female Body Size- All anurans 1. Positive <0.0001 Male Body size All families All anurans Female body size- Hylidae 2. Positive <0.0001 Clutch size Megophryidae Ranidae Body size ratio (F/M)- All anurans 3. Positive <0.0001 Clutch size Hylidae Female body size- 4. Positive Dendrobatoidea <0.0001 Female combat Egg size- 5. Negative Bufonidae 0.0014 Clutch size Female body size- 6. Positive Dendrobatoidea 0.0030 Clutch size Body size ratio (F/M)- 7. Positive Dendrobatoidea 0.0070 Clutch size Egg size- 8. • Negative All anurans 0.0085 Clutch size Body size ratio (F/M)- 9. Positive Bufonidae 0.023 Bodysize Female body size- 10. Positive Megophryidae 0.025 Egg size Body size ratio (F/M)- 11. Negative Dicroglossidae 0.032 Body size Male body size- Positive Dendrobatoidea 0.036 12. Male combat Female body size- 13. Positive Ranidae 0.044 Egg size Female body size- 14. Positive All anurans 0.044 Female combat

Note. P-values of correlations after applying the Bonferroni correction are in bold.

46 DISCUSSION

Sexual Size Dimorphism

The positively skewed body size distribution suggested that large body sizes are uncommon in anurans. Either anurans are under a physical or environmental constraint that limits body size or small body size has some adaptive advantage (Wells, 2007).

Regardless, the giant females observed in spiders or giant males observed in some mammals do not occur in anurans (Fairbairn et ai, 2007).

This study confirmed Shine's assertion that most anurans exhibit female-biased sexual size dimorphism (SSD), but RSSD in anurans might be more common than originally thought. Based on the data collected and the 1.02 threshold body size ratio

(F/M) applied here, males were larger than or equal to females in 23.9% of anuran species. My use of 1.02 as a SSD threshold likely underestimated the RSSD (reversed sexual size dimorphism), so that the actual percentage of RSSD is probably even greater than the 23.9% found here. Nevertheless, the frequency of SSD here is higher than the estimation of 10% proposed by Shine (1979). I removed species without body size information, while Shine assumed that these species exhibited a normal SSD (F>M), perhaps leading him to underestimate the percentage of RSSD. A female-biased SSD and an asymmetric SSD in anurans indicated that the selection for large females was more dominant than the selection for large males. RSSD probably has independently evolved multiple times and the occurrence of it was not likely to be restricted in certain distribution area.

47 SSD and Mating Related Life History Traits

I. Correlation between SSD and Mating Combat

The result that only 20.53% of species displayed mating combat behavior confirms Shine's (1979) observation that mating combat is uncommon in anurans.

However, this percentage was higher than the 5% reported by Shine. The different percentages estimated in these two studies might arise from the different estimations on the absence of mating combat. Shine (1979) may have underestimated mating combat by categorizing species without mating and combat records as absence of combat. This study may be more accurate because I excluded species without any mating record.

Large body size was favored in species where mating combat was observed, because female and male body sizes were positively correlated with female and male combat, respectively. Although the correlation between male body size and male combat was only found in Dendrobatiodea, the association between these two traits was supported in many species in other families, such as Bufonidae, Ranidae, Myobatrachidae and Leiuperidae (Cherry, 1992). For example, in the wood frog Rana sylvatica, male reproductive success strongly increased with body size (Howard and Kluge, 1985). The results were congruent with my predictions that male body size was not correlated with scramble competition. However, the prediction that male body size was positively correlated with territory defense was not supported. This may be due to the relatively small number of contrasts which reduce the statistical power by separately analyzing two different forms of mating combat.

Even though male body size and male mating combat were correlated, the presence of male combat was not always associated with the presence of RSSD. This

48 result is not consistent with Shine's (1979) prediction of a correlation between RSSD and

male combat in anurans. Natural selection favoring large females with high fecundity

may be stronger than sexual selection for the advantage of large male body size during

contests for mates in anurans. For example, in the wood frog {Rana sylvatica), reproductive success increased with body size more strongly in females than males.

Although males displayed severe scramble competition in this species, SSD was still

female-biased presumably because of the intense fecundity selection (Howard, 1988).

II. Correlation between SSD and Length of Breeding Season

Unlike Woolbright's (1983) suggestion, the prediction that SSD in anurans is related to breeding period was not supported in this study. This casts doubt on the assumption that energy increasingly constrains male growth as breeding season lengthens. For example, large males have enhanced mating success in many anurans with an explosive breeding season (Howard, 1988). In addition, the operational sex ratios as well as temporal constraints might influence energetic expenditure and growth of males

(Sullivan, 1984). No evidence supports the prediction that SSD evolves in response to the length of breeding season in anurans, although this may also reflect a reduction of statistical power by treating the length of breeding season as a binary variable. Further tests of the correlation between SSD and breeding season should use quantitative measures of the length of breeding season.

49 SSD and Breeding Related Life History Traits

I. Correlation between SSD, Clutch Volume, Egg Size, and Clutch Size

The positive correlation between body size ratio (F/M) and clutch volume indicated that selection on the total investment in gametes likely favored large females and resulted in the evolution of a female-biased SSD in anurans. The fitness value of both egg size and clutch size likely favor an increase in female body size. Either or both of these traits can contribute to the increase of body size ratio (F/M) in different families

(superfamily): egg size correlated with SSD in Hylidae, clutch size correlated with SSD in the superfamily Dendrobatoidae and the family Hylidae. In the all-anuran analysis, clutch size was positively correlated with SSD but no correlation between egg size and

SSD was found. Therefore, an increase of female body size might be mainly associated with increased clutch size in anurans. My data suggests that releasing females from fecundity selection may cause a reduced degree of female-biased dimorphism. For example, 84% of species demonstrating RSSD have less than 2000 eggs per clutch. In addition, RSSD is always found in lineages where females do not produce large number of eggs, such as the families Bombinatoridae, and Megophrydae, the superfamily

Dendrobatoidae and some clades in the family Hylidae. A problematic aspect of this study was that with the precision of my estimate of clutch size. It was highly variable within species depending on the size of females and environmental conditions, and so my estimated mean or median value likely has low precision. However, it is the best estimation of clutch size currently available.

The hypothesis that fecundity selection explains the evolution of SSD was supported in anurans here and has been supported in various other animal groups. For

50 example, female body size in spiders is positively correlated with clutch size, both at interspecific and intraspecific levels (Foellmer and Moya-Larano, 2007). Furthermore,

SSD in spiders correlates positively with female body size and clutch size. However, the fecundity selection hypothesis is not supported in mammals, birds and reptiles. In mammals, selection favoring small female size, because smaller body size require less energy to maintain their metabolism when brooding the young (Lindenfors et al, 2007).

In birds, no correlation has been found between SSD and fecundity. Differences between species in foraging ecology, parental roles and demands imposed by egg production may outweigh the effects of fecundity on shaping SSD in birds (Szekely et al, 2007). In reptiles, cross-species comparative analyses provide only weak and inconsistent supports for fecundity advantage as an explanation for SSD. There, fecundity selection may favor increased reproductive frequency rather than the per-clutch fecundity advantage of large female size (Cox et al, 2007). Further tests of when and how fecundity selection affects the evolution of SSD are required in a variety of animal groups in order to better understand how often fecundity is the primary factor determining SSD or how often it is offset by other factors.

II. Correlation between SSD and Parental Care

My results did not support the hypothesis that parental care selects either for larger or smaller parents. Parental care was not an essential factor in the increase of either male or female body size in anurans, so it does not appear to be a major cause of SSD.

Most modes of parental care in anurans might not select for the strength or size of parents, and large size might not be particularly favored. Some large frog species, such

51 as the ocellated frog Leptodactylus ocellatus and the giant bullfrog Pyxicephalus adspersus, defend tadpoles against potential vertebrate predators; however, the evidence

for parental protection against large predators is very rare and mostly circumstantial

(Wells, 2007). Moreover, the growth of the care-providing parents may not be limited by the metabolic costs of brooding activities. Parents may experience little reduction in energetic intake when looking after the young or perhaps the additional energy consumed during parental care was trivial (Wells, 2007). Wickramasinghe et al. (2004) found that brooding males in the streamlined frog Nannophrys ceylonensis would come out of their breeding sites and foraged in the vicinity. Parental care in anurans was mostly short-term, such as egg attendance and transporting tadpoles (Pough et al., 2001), and the metabolic costs of the short-termed care were more likely to be trivial (Wells, 2007). For example, there was no difference in fat body weight between calling males and the males transporting larvae in the poison frog Mannophryne trinitatis (Downie et al, 2005).

Two explanations could count for the correlation between the presence of parental care and decreased clutch size: 1) parental care evolves followed by a subsequent decrease in clutch size because smaller clutch sizes were less energetically expensive to defended or tended (Messina and Fox, 2001); or 2) a reduction in clutch size was followed by the evolution of parental care in order to ensure higher offspring survival

(Zug et al., 2001). As it stands, more research is needed to determine which trait evolved first. Given that clutch size was negatively correlated with egg size, a positive correlation between parental care and egg size was expected, but I found no evidence that the presence of parental care was related to increased egg size. This might be because

52 parental care was mainly associated with a decrease in clutch size, which trades off against egg size.

One possible reason for the difference in results between this study and that of

Summers etal. (2006)'on the relationship between egg size and parental care might be that egg size and parental care were coded differently. In Summers et al., all species without descriptions of breeding activities were assumed to have no parental care behavior which likely underestimated the frequency of parental care. I avoided this problem by removing species with no a description of breeding activity from my analysis.

Summers et al. treated egg size as a binary trait by assigning egg size in a species larger than the mean value across all species as "large", and egg size smaller than the mean value as "small". I treated egg size as a continuous trait. This treatment would reflect gradual changes of egg size, which might be a more plausible evolutionary scenario compared to binary changes. In summary, the data in my study was of higher quality than in Summers et al. (2006). The inconsistent findings between these two studies suggest that more tests of the correlation between egg size and parental care are needed in anurans before a strong conclusion can be reached.

Rensch's Rule

Anurans exhibited an allometry for SSD that was the reverse of Rensch's rule where mean body size was positively correlated with SSD. The correlation between female and male body size indicated genetic linkage between two sexes (Fairbairn et al,

2007). Greater variation in female body size than male body size was opposite to

Rensch's rule's prediction. This suggests that fecundity selection favoring increased

53 female body size may be the dominant factor for increasing SSD, because the magnitude

of the dimorphism was proportional to the intensity of selection on females. However,

male size was more divergent than female size in the family Dicroglossidae, which

suggests that in some anuran clades, the evolutionary pattern of SSD may be congruent

with Rensch's rule.

Relaxation of fecundity selection in females might explain why females and

males have similar sizes, but could not explain why males become larger than females in

some species. Many factors could promote an increase in male body size, such as male- male combat and male parental care. However, by adapting to different mating or breeding strategies, different species might respond to selection on male size differently

and there might be no general trend for the increase of male body size in anurans. More

studies on male size are needed to understand the general pattern of SSD in anurans.

Interestingly, the trend of correlation between SSD and body size in the family

Dicroglossidae was the reverse of the family Bufonidae. The differences between these two families might arise from the different sexes changing faster in size. Male size may

diverge faster than female body size in Dicroglossidae because of two factors: 1) intense

sexual selection for large males, and 2) relaxation of fecundity selection for large

females. Although male body size was not significantly correlated with male combat in this family, perhaps because of a small number of contrasts (n=6), other evidence

supported an association between these traits. For example, studies show that male

mating success is positively correlated with male body size in some dicroglossid species.

In the large-headed frog Limnonectes kuhlii, a large male has mating advantage through a

size-dependent spatial movement and through male competition in a polygyny mating

54 system (Tsuji, 2004). Female body size was not significantly correlated with clutch size

in Dicroglossidae perhaps because of the small number of contrasts (n=7). The mean

clutch size in this family is relatively small, varying from 11.5 to 2000 depending on the

species (n=13). Small clutches reduce fecundity selection for large females, which

probably can make an increased male body size more obvious. However, future studies

are needed to shed more light on SSD and life history evolution in this family, as the life

histories of very few species had been studied in-depth.

Greater variation in female body size compared to male body size in Bufonidae

might be explained by the fecundity advantages of large females. In contrast to

Dicroglossidae, species in Bufonidae always had large clutches. The mean clutch sizes in

81.5% species were larger than 2000. Mean clutch size in 44.4% of the species exceeded

10,000. Although there was no significant correlation between female body size and

clutch size detected in Bufonidae (p=0.276, n=7), the trend between these two traits was positive (slope=7.11). Taking into account the positive correlation between female body

size and female fecundity in many Bufo species, such as the common toad Bufo calamita

(Tejedo, 1992) and Bufo bufo (Reading and Clarke, 1995), fecundity selection on females

is expected to result in the greater interspecific variance of female body size in this

family.

Rensch's rule has not been supported in some other animal groups, such as

ectotherms or organisms where there is no strong sexual selection on male body size. For

example, among snakes, Rensch's rule occurred only in those lineages in which male

combat and male-biased SSD were common, whereas reversed Rensch's rule tended to

occur when female-biased SSD was prevalent (Cox et al, 2007). As well, Pacific salmon

55 and trout (Oncorhynchus) do not exhibit combat between males and do not follow

Rensch's rule (Young, 2005). In insects, support for Rensch's rule is mixed; and it probably does not deserve the attribute "rule", a pattern also true for spiders

(Blanckenhorn et al, 2007; Foellmer and Moya-Larano, 2007). Thus, researchers should be cautious when applying Rensch's rule to ectotherms and organisms that are not under strong selection for male competition, as fecundity selection for increased female body size might contribute most to the evolution of SSD.

Suggestions for Future Research

I found considerable heterogeneity between the results in within-family analyses and the all-anuran analysis. Correlations found within families did not necessarily apply to all anurans and vise versa. There might be four reasons for this significant heterogeneity between the two different levels of analyses: 1) different clades facing different selection pressures or responding to the same selection pressures differently; 2) difficulties in generalizing an overall pattern of evolution for all anurans based on the existing species due to the lack of information for extinct ancestral amphibian groups; 3) small numbers of contrasts in within-family analysis reducing the statistical power; 4) artificial effects in data collections, data treatment and analyses. Among these four reasons, the artificial analytical effects may be of greatest concern. The comparative net might have been cast too widely in all-anuran analyses, where supertree construction, assumption of unit branch length, non-random sampling of taxa, and assigning traits as binary variables could have introduced errors into my analyses. Such errors may have

56 been relatively small at the intra-family level but could be greatly inflated in the all-

anuran analyses.

Comparative analyses such as this one can be improved in various ways. First,

larger data sets and better-resolved phylogenies are needed for many groups. The data set

in the current study only included about 10% of all anuran species due to restricted data

availability and time constraints in data compilation. Life history data were also typically

biased towards diurnal species and species living in the northern hemisphere. Researchers need to pay more attention to nocturnal species and species living in the southern hemisphere. Furthermore, more detailed phylogenies including branch length information

on most anuran clades are urgently needed. Frost et al. (2006)'s most comprehensive phylogeny on amphibians only included 7% of the described anuran species in the world!

Phylogenetic studies lack many families, especially those found in Asia, Africa and

Australia. Branch lengths, which are essential to phylogenetic comparative methods, need to be reported in all published phylogenies for future use in comparative analyzes.

Second, more detailed descriptions on body size and life history data are needed

in order to increase the statistical power of comparative analyses. Better resolution of

body size dimorphism and length of breeding season will likely boost the statistical power of such comparative analyses. When authors report their measurements on

continuous variables, such as body size, egg size and clutch size, they should provide the

mean with variance (standard deviation or standard error) and sample'size of these data.

Range data (maximum and minimum) is of much lower quality. If traits have to be

reported as a categorical variable, such as the presence versus absence of RSSD and

prolonged versus explosive breeding season, a detailed definition of the criteria used to

57 assign taxa to each category should be provided. Moreover, these judgments should reasonably estimate trait states, especially the absence of a behavior.

Third, alternative functional explanations of SSD need further attention. A variety of factors can select for SSD in addition to the life history traits that were examined in this study. For example, female choice of large males can favor the evolution of increased male size. Ryan (1980) has shown that female tungara frogs Physalaemus pustulosus preferentially chose mating calls with lower fundamental frequencies, which were associated with large male size. Reproductive modes may impose selection on female fecundity and male mating behaviors (Wells, 2007). Explanations for the evolution of body size may also benefit from taking into account the effects of age, because anurans typically continue to grow throughout life and so selection may vary over ontogeny

(Monnet and Cherry, 2002). Selection from ecological conditions may also favor dimorphism. For example, if species live in resource-poor habitats, then resource limitation could prevent either sex from responding to selection for becoming large

(Blanckenhorn et al, 1995).

Fourth, I suggest that large scale cross-species analyses should focus at lower taxonomic levels (family or genus) for three reasons. First, imprecise estimations of trait values at the root of a phylogeriy reduce the predictive power (R ) of regression models.

Taking the correlation between female body size and clutch size as an example, the predictive power was much lower in the all-anuran analysis than within-family analyses.

The R2 values vary from 0.48 to 0.64 in the four families having significant p-values in regressions, while it was reduced to 0.33 in the all-anuran analysis. Second, the data measurements of different researchers become more inconsistent as the numbers of

58 species increases. However, analyses at lower taxonomic levels will avoid this problem, as researchers can relatively easily collect data on their own and classify data more consistently. Third, well-resolved phylogenies with relatively accurate branch length information are usually only available for small clades. In contrast, creating phylogenetic hypotheses for large animal groups generally require supertree approaches and these are always associated with hard polytomies and no branch length information. So, large-scale comparative analyses will generally suffer from analytical problems that reduce the accuracy of the phylogenies.

Finally, programs for phylogenetical comparative methods (PCMs) should consider changing branch lengths when removing species with missing values when all branch lengths are assumed to be equal to one. Setting all branch lengths equal to one is a common solution for no branch lengths in supertrees and inaccurate branch lengths in some phylogenies. Many PCMs programs, such as CAIC and BayesTraits, assume the total branch length from the root to tips remain the same after the species with missing values are removed. This results in the branch lengths leading to the remaining species not being equal to one any more. For example, in the hypothetical example in Figure 8, after removing species B, the branch length leading to species A will be two times longer that the one leading to species C. As a consequence, when calculating the trait value at node "b", the trait value of species C will be weighted two times larger than it is expected to be.

59 Species Species a b A b A B C C

Figure 8 A hypothetical example showing incorrect branch length changes when species with missing values are removed from comparative analyses under the assumption that all branch lengths equal to one unit.

60 CONCLUSION

Sexual size dimorphism (SSD) is female-biased in most anuran species. The results in the current study are inconsistent with the common hypothesis generated from mammals and birds that mating competition between males causes the evolution of male- biased SSD. The degree of sexual size dimorphism in anurans may not directly reflect the magnitude of sexual selection on male body size. Instead, it likely reflects the fecundity advantages of larger female body sizes. Breeding activities and parental care do not appear to constrain the growth of male anurans. Moreover, the presence of parental care may have an indirect impact on SSD because of its evolutionary association with female fecundity. The general pattern of SSD in anurans does not follow Rensch's rule as derived from other organisms, although this rule only apply to some anuran lineages.

Phylogenetic comparative analyses of other life history traits and ecological features are needed to better understand the evolutionary patterns of SSD in anurans. The correlations discovered here will hopefully stimulate further tests of adaptive or functional hypotheses of SSD in anurans using field or laboratory experiments.

Patterns of SSD in anurans shed new light on the processes of SSD evolution in animals. Most prior research on SSD has focused on mammals and birds, where there is strong sexual selection on male body size. This and other recent studies indicate that the allometry for SSD in the animal groups where fecundity selection is stronger may be different from the trend predicted by Rensch's rule. This suggests that further research on

SSD in fish, amphibians and invertebrates may be helpful in determining how physiological or developmental factors affect the evolution of SSD in animals.

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92 APPENDIX 1 Mean body size, mean egg size, mean clutch size, reversed sexual size dimorphism, mating combat, length of breeding season and parental care in anurans. * RSSD = Reversed Sexual Size Dimorphism (F<=M). ** Female and male body size that were recorded as range are not reported. ***The presence of a traits is coded as "1", the absence of a trait is coded as "0"; missing data is represented by "-". Seaso n RSSD * Famil y Specie s Defens e Eg g Siz e Clutc h Siz e Competitio n Mal e Comba t Paterna l Car e Mal e Scrambl Mal e Territor y Mal e Bod y Siz Femal e Comba t Femal e Bod y Siz Mal e parenta l Car Prolonge d Breedin g Femal e parenta l Car

Alytidae Alytes obstetricans - r- 2.55 53 0** 0 1 0 0 0 1 1 0 Alytidae Discoglossus pictus - - 1.25 750 0 0 0 - - 0 0 0 0 Aromobatidae Allobates femoralis 26.90 24.94 2.00 43 0 0 0 0 - 1 1 0 Aromobatidae Allobates zaparo 28.00 27.80 - - 1 - - - - 1 - - Aromobatidae Colostethus awa - - - 21 0 0 0 1 - 1 1 0 Aromobatidae Colostethus beebei - - 2.10 4 - 0 0 0 0 - 1 1 1 Aromobatidae Colostethus brunneus 19.60 17.90 - - 0 ------Aromobatidae Colostethus caeruleodactylus 16.30 15.50 - - 0 0 0 1 - 1 1 0 Aromobatidae Colostethus elachyhistus 21.20 18.30 - 19 0 0 0 0 0 - 1 1 0 Aromobatidae Colostethus flotator - - - - 0 0 0 1 - 1 1 0 Aromobatidae Colostethus idiomelus 25.10 22.00 - - 0 ------Aromobatidae Colostethus infraguttatus - - , - - 0 - "' ------Aromobatidae Colostethus inguinalis - - - - 0 1 0 1 - 1 0 1

Aromobatidae Colostethus insulatus 18.30 21.25 - - 1 - - - - •- - - - Aromobatidae Colostethus nexipus 21.40 19.30 - - 0 ------Aromobatidae Colostethus nidicola 20.20 19.60 2.40 4 1 0 0 0 0 1 0 0 0 Aromobatidae Colostethus nubicola - - - - 0 0 0 0 0 - 1 1 0 Aromobatidae Colostethus palmatus - - 2.38 22 - 0 0 0 0 - 1 1 0 Aromobatidae Colostethus panamensis - - - - - 0 0 0 0 - 1 - - Aromobatidae Colostethus pratti - - - - - 0 0 0 0 - 1 0 1 Aromobatidae Colostethus stepheni 17.10 16.60 - 4 0 1 0 0 1 1 1 1 0

93 Aromobatidae Colostethus subpunctatus 24.30 21.40 1.50 21 0 0 0 0 0 1 1 1 0 Aromobatidae Colostethus sylvaticus 28.55 j 23.70 - - 0 - - -' - - - - - Aromobatidae Colostethus talamancae - - - 18 0 1 0 0 1 1 1 Aromobatidae Colostethus vertebralis 17.87 16.06 - - 0 0 0 0 0 - 0

Aromobatidae Mannophryne collaris -• - - - 0 1 1 0 1 - 0 Aromobatidae Mannophryne trinitatis 25.60 23.10 10 0 1 1 0 1 - 0 Aromobatidae Phyllobates aurotaenia 30.40 26.90 - - 0 - - - - - , - Aromobatidae Phyllobates bicolor 39.23 36.16 - 17 0 0 0 0 0 1 0 Aromobatidae Phyllobates lugubris 22.20 19.20 - 18 0 0 0 0 0 - 0 Aromobatidae Phyllobates terribilis 43.23 41.05 2.50 15 0 0 1 0 0 - 0 Aromobatidae Phyllobates vittatus 27.70 24.40 2.50 19 0 1 1 1 1 - 0 Arthroleptidae Trichobatrachus robuslus - - - - 0 0 0 - - - 0 Bombinatoridae Bombina bombina - - 1.80 190 1 1 0 1 1 0 0 0 0 Bombinatoridae Bombina microdeladigitora 79.70 72.10 - - 0 0 0 - - 0 - - - Bombinatoridae Bombina orientalis 47.00 42.10 1.50 165 1 1 0 1 0 0 0 0 0 Bombinatoridae Bombina variegata - - - 72 1 1 0 1 1 0 0 0 0 Brachycephalidae Eleutherodactylus augusti - - - 59 0 0 0 - - 1 1 1 0

Brachycephalidae Eleutherodactylus binotatus - - - - • 0 - - - - - 0 0 0 Brachycephalidae Eleutherodactylus bufoniformis - - - - 0 0 0 ------Brachycephalidae Eleutherodactylus planirostris - - - 14 0 0 0 - - 1 0 0 0 Brachycephalidae Ischnocnema quixensis 57.60 43.80 4 35 0 - - - - - 0 0 0 Brevicipitidae Breviceps mossambicus - - 4 25 0 0 0 - - - 0 0 0 Bufonidae Bufo alvarius - - 1.40 7750 0 1 0 1 0 0 0 0 0 Bufonidae Bufo americanus 53.10 58.60 1.20 6000 0 1 0 1 0 0 0 0 0 Bufonidae Bufo andrewsi 71.3 81.20 2 - 0 0 0 0 0 1 0 0 0 Bufonidae Bufo arenarum - - - 4500 0 0 0 0 0 1 0 0 0 Bufonidae Bufo arunco - - - - - 0 0 0 0 - 0 0 0 Bufonidae Bufo asper 83.02 92.85 1.26 12792 0 0 0 0 0 1 0 0 0 Bufonidae Bufo biporcatus 48.30 53.50 - - 0 0 0 0 0 - 0 0 0 Bufonidae Bufo boreas ".- - 1.70 5213 0 1 0 1 0 1 0 0 0 Bufonidae Bufo bufo - - 1.75 5332 0 1 0 1 0 0 0 0 0 Bufonidae Bufo californicus - - 1.42 4714 0 0 0 0 0 0 0 0 0 Bufonidae Bufo canorus - - 2.10 - 0 1 0 1 0 0 0 0 0 Bufonidae Bufo castaneoticus - - 1.50 178 0 0 0 0 0 0 0 0 0 Bufonidae Bufo chavin 50.17 54.82 2.44 276 0 0 0 0 0 1 0 0 0 Bufonidae Bufo coccifer 51.40 59.55 1.29 3000 0 0 0 0 0 0 0 0 0

Bufonidae Bufo cognatus - - • 1.19 22148 0 1 0 1 0 0 0 0 0 Bufonidae Bufo coniferus - - 1.80 - 0 0 0 0 0 1 0 0 0 Bufonidae Bufo crucifer - - - - - 0 0 0 0 - 0 0 0

94 • Bufonidae Bufo dapsilis - - - - 0 0 0 0 0 - - - - Bufonidae Bufo debilis 35.10 35.80 1 - 0 0 0 0 0 0 0 0 0 Bufonidae Bufo exsul 51.61 51.25 - - 1 1 0 1 0 1 0 0 0 Bufonidae Bufojastidiosus - - 4.30 85 0 1 0 1 0 0 0 0 0 Bufonidae Bufo fowleri 55.70 63.60 • - 8000 0 0 0 0 0 1 0 0 0 Bufonidae Bufo garmani - - 1.20 - 0 0 0 0 0 - 0 0 0 Bufonidae Bufo glaberrimus - - - - 0 0 0 0 0 - - - - Bufonidae Bufo granulosus 41.60 44 1 9000 0 0 0 0 0 0 0 0 0 Bufonidae Bufo gutlatus - - - - - 0 0 0 0 - - - -

Bufonidae Bufo gutturalis -• - 1.45 20000 0 1 0 1 0 - 0 0 0 Bufonidae Bufo haemaliticus - - - - 0 0 0 0 0 0 0 0 0 Bufonidae Bufo hemiophrys - - - - 0 0 0 0 0 1 0 0 0 Bufonidae Bufo houstonensis - - - - - 1 0 1 0 0 0 0 0 Bufonidae Bufo juxtasper - - - - 0 0 0 0 0 - - - - Bufonidae Bufo kisoloensis - - - - 0 0 0 0 0 - 0 0 0 Bufonidae Bufo luetkenii - - 1.50 3300 0 0 0 0 0 0 0 0 0 Bufonidae Bufo maculatus - • - 1.30 J - 0 1 0 1 0 - 0 0 0 Bufonidae Bufo margaritifer 51.30 62 2 1537 0 1 0 1 0 0 0 0 0 Bufonidae Bufo marinus 87.90 96.05 1.87 35000 0 1 0 1 0 1 0 0 0 Bufonidae Bufo mauritanicus - - 1.50 - 0 0 0 0 0 0 0 0 0 Bufonidae Bufo melanochlorus - - - - 0 0 0 0 0 0 0 0 0 Bufonidae Bufo melanostictus 68.52 71.86 - - 0 0 0 0 0 1 0 0 0 Bufonidae Bufo microscaphus - - 1.44 3650 0 0 0 0 0 0 0 0 0

Bufonidae Bufo nelsoni - - • - - 0 0 0 0 0 0 0 0 0 Bufonidae Bufo pardalis 93.40 100.80 - - 0 1 0 1 0 - 0 0 0 Bufonidae Bufo punctatus - - 1.15 - 0 0 0 0 0 0 0 0 0 Bufonidae Bufo quercicus 25.70 25.45 1 - 0 0 0 0 0 0 0 0 0 Bufonidae Bufo rangeri - - 1.30 - - 0 0 0 0 0 0 0 0 Bufonidae Bufo regular is - - 1.50 15000 0 1 0 1 1 0 0 0 0 Bufonidae Bufo retiformis 44.4 48.45 - - 0 0 0 0 0 0 0 0 0 Bufonidae Bufo schneideri - - 1.80 - - 0 0 0 0 1 0 0 0 Bufonidae Bufo speciosus - - - - - 1 0 1 0 0 0 0 0 Bufonidae Bufo spinulosus - - - - - 0 0 0 0 1 0 0 0 Bufonidae Bufo terrestris - - 1.20 4000 0 0 0 0 0 0 0 0 0 Bufonidae Bufo valliceps - . - 1.20 - 0 0 0 0 0 0 0 0 0 Bufonidae Bufo variegatus - '• - - - 138 - 0 0 0 0 0 0 0 0 Bufonidae Bufo viridis - - 1.65 11000 0 0 0 0 0 0 0 0 0 Bufonidae Bufo woodhousii - - 1.25 10500 0 1 0 1 0 0 0 0 0 Bufonidae Bufo xeros - - - - 0 0 0 0 0 - 0 0 0

95 Bufonidae Ollotis ibarrai 62.20 70.30 - - 0 ------Bufonidae Schismaderma carens - - 2 6264 0 0 0 0 0 0 0 0 0 Centrolenidae Allophryne ruthveni - - - 300 0 0 0 - - - 0 0 0 Centrolenidae Centrolenella prosoblepon 26.40 24.10 - 20 0 1 0 0 1 0 0 1 Centrolenidae Hyalinobatrachium fleischmanni - - 2 30 0 0 0 - - 0 0

Ceratobatrachidae Ingerana baluensis - •, 2.35 - 0 ------Cryptobatrachidae Stefania evansi 63.90 45.80 1 23 0 0 0 - - 1 0 1 Cycloramphidae Rhinoderma darwinii - - 3.60 40 0 0 0 0 0 1 0 Dendrobatidae Dendrobates arboreus - . - - - - 0 0 0 0 - 0 1 Dendrobatidae Dendrobates auratus 33.72 30.29 2 8 0 1 1 1 1 - 0 Dendrobatidae Dendrobates azureus 42.70 39 3.50 6 1 0 1 - - - 1 Dendrobatidae Dendrobates fantasticus - - - - 0 0 0 0 0 - 1 Dendrobatidae Dendrobates fulguritus 15.20 14.50 - - 1 ------Dendrobatidae Dendrobates galactonotus • 35.70 33.20 - - 0 ------Dendrobatidae . Dendrobates granuliferus 20.80 20.70 - 3 1 1 0 0 1 1 1 Dendrobatidae Dendrobates histrionicus 32.69 33.02 1.50 - 1 1 1 1 0 - 0 1

Dendrobatidae Dendrobates imitator - - - - • 1 0 0 1 - 0 Dendrobatidae Dendrobates lehmanni 32.85 33.40 1.50 - 1 1 0 - - - 0 1 Dendrobatidae Dendrobates leucomelas 35.11 32.73 3.93 5 0 1 0 1 0 - 0 Dendrobatidae Dendrobates minutus 13.50 13.20 - - 0 0 0 0 0 - 0 Dendrobatidae Dendrobates pumilio 20.74 20.94 1.10 3 1 1 1 0 1 1 1 Dendrobatidae Dendrobates quinqueviltatus 17.30 16.60 - - 1 0 0 0 0 - 0 Dendrobatidae Dendrobates reticulatus 15.11 14.37 2 2 0 0 0 0 ' 0 - 0 Dendrobatidae Dendrobates speciosus 29.10 28.40 - - 0 1 1 1 0 - 0 1 Dendrobatidae Dendrobates tinctorius 46.50 41 - 5 0 0 0 0 0 - 0 Dendrobatidae Dendrobates truncatus 28.70 25.60 - 5 0 0 0 0 0 - 0 Dendrobatidae Dendrobates vanzolinii - - - - 0 0 0 0 0 - 1 Dendrobatidae Dendrobates variabilis - - - 6 - 0 0 0 0 - 0 Dendrobatidae Dendrobates ventrimaculatus - - 2 2 1 1 0 0 1 1 1 Dendrobatidae Epipedobates anthonyi 20 18.10 - - 0 0 0 0 0 - 0 Dendrobatidae Epipedobates boulengeri 18.50 16.90 - - 0 0 0 0 0 - 0 Dendrobatidae Epipedobates espinosai - - - - 0 0 0 0 0 - 0 Dendrobatidae Epipedobates hahneli - - 2.30 22 0 0 0 0 0 - 0 Dendrobatidae Epipedobates parvulus 21.20 19 3 8 0 0 0 0 0 1 0 Dendrobatidae Epipedobates petersi 28 24.80 - - 0 0 0 0 0 - 0 Dendrobatidae Epipedobates pictus 24.40 23.20 2 18 0 0 0 0 0 - 1 Dendrobatidae Epipedobates pulchripectus 26 22.60 - - 0 ------Dendrobatidae Epipedobates silverstonei 41.83 35.85 2 30 0 0 0 0 0 - 0 Dendrobatidae Epipedobates tricolor 23.32 21.04 - 22 0 1 0 0 1 - 0

96 Dendrobatidae Epipedobates trivittatus 43.60 37.88 3.20 50 0 1 1 0 1 - 1 1 0 Dicroglossidae Chaparana unculuanus 77.90 74.55 4 97 0 0 0 0 0 - - - - Dicroglossidae Euphlyctis cyanophlyctis 48 67 - - 0 0 0 0 0 1 0 0 0 Dicroglossidae Fe/ervarya cancrivora 68.59 58.75 1.63 - 0 0 0 0 0 1 0 0 0 Dicroglossidae Fejervarya limnocharis 48.75 38.95 1 1560 0 1 0 1 0 1 - - - - Dicroglossidae Hoplobatrachus chinensis 71.73 82.44 1.75 2000 1 0 0 0 0 1 - - - Dicroglossidae Hoplobatrachus occipitalis - - 5 - 0 - - - - - 0 0 0 Dicroglossidae Limnonectes bfythii - - 2.10 1852 1 1 0 0 1 - 1 1 0 Dicroglossidae Limnonectes finchi - - 1.96 - 0 - - - - - 1 1 0 Dicroglossidae Limnonectes gyldenstolpei 52.92 58.50 - - 1 0 0 0 0 - - - Dicroglossidae Limnonectes ibanorum - - - 1122 1 ------Dicroglossidae Limnonectes kuhlii 54.85 55.55 2.25 80 1 1 0 0 1 1 0 0 0 Dicroglossidae Limnonectes laticeps 40.80 43.80 - - 1 0 0 0 0 1 - - - Dicroglossidae Limnonectes macrodon - - 1.4 1660 0 0 0 0 0 1 - - - Dicroglossidae Limnonectes microdiscus 37.34 32.37 4.5 53 0 - - - - - 1 1 0 Dicroglossidae Limnonectes palavanensis - - 2.29 - 0 - - - - - 1 1 0 Dicroglossidae Limnonectes paramacrodon - - - - 1 0 0 0 0 - - - - Dicroglossidae Limnonectes parvus - - 2 11 0 ------Dicroglossidae Nannophrys ceylonensis 48.7 46.6 - - 0 1 0 0 1 - 1 1 0 Dicroglossidae Nanorana parkeri 49.40 45.65 2 - 0 0 0 0 0 1 - - - Dicroglossidae Nanorana pleskei 37.3 36.6 - - . 0 0 0 0 0 0 - - - Dicroglossidae Occidozyga laevis 32.62 24.40 - - 0 ------Dicroglossidae Occidozyga lima 27.63 21.17 1 707 0 ------Dicroglossidae Occidozyga martensii 24.97 20.40 1 - 0 ------Dicroglossidae Paa boulengeri 103.30 100.55 3.90 - 0 0 0 0 0 1 0 0 0 Dicroglossidae Paa spinosa 93.17 89.15 4.25 228 0 0 0 0 0 1 0 0 0 Dicroglossidae Paa yunnanensis 95 95.50 3.50 633 1 0 0 0 0 1 0 0 0 Hemiphractidae Hemiphraclus helioi - - - - 0 0 0 - - - 1 0 1 Hemisotidae Hemisus marmoratum - - 2 j 200 0 0 0 - - - 1 0 1' Hylidae Acris crepitans 24.15 22.80 1.53 500 0 1 0 0 1 1 0 0 0 Hylidae Acris gryllus - - 1.13 232 0 0 0 0 0 1 0 0 0 Hylidae Agalychnis callidryas 62.50 48.20 3.20 265 0 1 0 0 1 1 0 0 0 Hylidae Agalychnis litodryas - - - - 0 0 0 0 0 - 0 0 0 Hylidae Agalychnis saltator - - 5 46 0 1 0 1 0 0 0 0 0 Hylidae Agalychnis spurrelli 74.95 62 - - 0 0 0 0 0 - 0 0 0 Hylidae Anotheca spinosa 63.40 61.10 1.63 316 0 0 0 0 0 1 1 0 1 Hylidae Aparasphenodon brunoi - - - - - 0 0 0 0 - 1 0 1 Hylidae , Aplastodiscus perviridis - - 1.80 227 - 0 0 0 0 - 1 1 0 Hylidae Cruzlohyla calcarifer 65 52 3.75 80 0 0 0 0 0 1 0 0 0

97 Hylidae Cyclorana alboguttata - . 1.30 - 0 0 0 0 0 - 0 0 0 Hylidae Cyclorana australis - - 1.60 550 0 0 0 0 0 1 0 0 0 Hylidae Cyclorana brevipes - - - 1.30 - 0 0 0 0 0 - 0 0 0 Hylidae Dendropsophus aperomeus 25.0 19.83 - - 0 ------Hylidae Dendropsophus parviceps 23.35 16.80 1 234 0 0 0 0 0 - 0 0 0 Hylidae Duellmanohyla rufioculis 36.65 28.20 2.36 112 0 0 0 0 0 1 0 0 0 Hylidae Ecnomiohyla miliarie - - - - 0 1 0 0 1 - - - - Hylidae Hyla albofrenata - - - - 0 0 0 0 0 - - - - Hylidae Hyla albopunctata - - - - 0 0 0 0 0 - - - - Hylidae Hyla allenorum - - - - 0 - - - - - 0 0 0 Hylidae Hyla anceps 40.33 37 1 - 0 0 0 0 0 - 0 0 0 Hylidae Hyla andersonii - - 1.30 500 0 0 0 0 0 0 0 0 0 Hylidae Hyla andina 49.8 46.5 - - 0 0 0 0 0 - 0 0 0 Hylidae Hyla annectans 38.80 34.15 - - 0 0 0 0 0 0 - - - Hylidae Hyla arborea 33 24.80 1.20 1282 1 0 0 0 0 1 0 0 0 Hylidae Hyla arborescandens 45.57 34.78 2.15 - 0 0 0 0 0 - 0 0 0 Hylidae Hyla arenicolor 47.20 35.30 2.07 - 0 0 0 0 0 0 0 0 0 Hylidae Hyla avivoca - - 1.17 632 0 1 0 0 1 1 0 0 0 Hylidae Hyla balzani 48.5 40.4 - - 0 0 0 0 0 - - - - Hylidae Hyla bifurca 31.80 25.10 1.50 186 0 0 0 0 0 - 0 0 0 Hylidae Hyla bipunctata - - - - 0 0 0 0 0 - - - - Hylidae Hyla bischoffi - - - - 0 0 0 0 0 - - - Hylidae Hyla boans 102.10 101 2 3154 1 0 0 0 0 0 1 1 0 Hylidae Hyla brevifrons 21.36 18.49 I 79 0 0 0 0 0 - 0 0 0 Hylidae Hyla bromeliacia - - - 14 0 0 0 0 0 - 1 0 1 Hylidae Hyla calcarata 53.90 35.90 1.50 1143 0 - - - - - 0 0 0 Hylidae Hyla chinensis 34.17 30.60 1.28 459 1 0 0 0 0 0 0 0 0 Hylidae Hyla chrysoscelis - 1.20 2000 0 1 0 0 0 1 0 0 0 Hylidae Hyla cinerea 48.90 53.10 1.20 875 1 1 0 0 1 1 0 0 0 Hylidae Hyla circumdata - - - - 0 0 0 0 0 - . 0 0 0 Hylidae Hyla colymba 36.23 34.65 - - 0 0 0 0 0 1 - - - Hylidae Hyla crepitans 65.50 53.90 1.45 1064 0 0 0 0 0 1 0 0 0 Hylidae Hyla ebraccata - 1.30 - 0 1 0 0 1 1 0 0 0 Hylidae Hyla ehrhardti 35.3 33.0 - - 0 0 0 0 0 1 - - - Hylidae Hyla elegans - - - - 0 1 0 0 1 - 0 0 0 Hylidae Hyla euphorbiacea 36.4 34.7 - - 0 0 0 0 0 0 0 0 0 Hylidae Hyla eximia 28.5 27.8 - - 0 0 0 0 0 0 0 0 0 Hylidae Hyla faber - - 1.50 - 1 1 0 0 1 1 1 1 0 Hylidae Hyla fasciata 46.20 35.60 1.50 569 0 - - - - - 0 0 0

98 Hylidae Hyla femoralis - - 0.90 - 0 0 0 0 0 1 0 0 0 Hylidae Hyla geographica 66.60 44.20 1 2797 0 0 0 0 0 - 0 0 0 Hylidae Hyla gratiosa - .- 1.85 - 0 0 0 0 0 1 0 0 0 Hylidae Hyla guentheri - - - - 0 0 0 0 0 - - - ' - Hylidae Hyla heilprini 49.3 49.4 - - 1 0 0 0 0 - 0 0 0 Hylidae Hyla japonica 28 32.70 1.30 920 1 0 0 0 0 1 0 0 0 Hylidae Hyla koechlini - - - - 0 0 0 0 .0 - 0 0 0 Hylidae Hyla labialis - - - - - 0 0 0 0 - - - - Hylidae Hyla lanciformis 87 74.90 2 1617 0 - - - - - 0 0 0 Hylidae Hyla leali - - - - 0 ------Hylidae Hyla lemai - - 2.50 54 0 0 0 0 0 - 0 0 0 Hylidae Hyla leucophyllata 42 33.90 1.12 587 0 1 0 0 1 1 0 0 0 Hylidae Hyla loquax 40.50 39.30 - 250 0 0 0 0 0 0 0 0 0 Hylidae Hyla marginata - - - - 0 - - - - - 0 0 0 Hylidae Hyla marianitae 51.4 47.6 - - 1 0 0 0 0 - - - - Hylidae Hyla marmorata 49.30 39.10 1.50 979 0 - - - - - 0 0 0 Hylidae Hyla martinsi - - - - 0 0 0 0 0 - - - - Hylidae Hyla melanomma 30.10 27.10 - - .0 0 0 ' 0 0 1 - - - Hylidae Hyla meridionalis - - 1.30 - 0 - - - - - 0 0 0 Hylidae Hyla microcephala 26.76 22.83 1 180 0 0 0 0 0 1 0 0 0 Hylidae Hyla minuscula 23.3 20.0 - - 0 0 0 0 0 - 0 0 0 Hylidae Hyla minuta 23.80 21.30 1 211 0 0 0 0 0 1 0 0 0 Hylidae Hyla miotympanum 39.7 28.5 2.41 120 0 0 0 0 0 - 0 0 0 Hylidae Hyla mlyatai - - - - 0 0 0 0 0 - 0 0 0 Hylidae Hyla nana 20.50 18.50 1.20 100 0 - - - - - 0 0 0 Hylidae Hyla partialis 70 60 - - 0 1 0 0 1 - 1 1 0 Hylidae Hyla pelidna - - - - 0 0 0 0 0 - - - - Hylidae Hyla pentheter - - - - 0 0 0 0 0 - - - - Hylidae Hyla phyllognatha 36.87 32.96 3.08 - 0 0 0 0 0 - 0 0 0 Hylidae Hyla plicata 43.8 39.70 - - 0 0 0 0 0 - - - - Hylidae Hyla polytaenia - - - 150 0 0 0 0 0 - 0 0 0 Hylidae Hyla prasina - - - - 0 0 0 0 0 - - - - Hylidae Hyla pseudopseudis 42.75 46.75 - - 1 - - - - - 0 0 0 Hylidae Hyla pseudopuma 44.30 39.70 1.70 2150 0 1 0 1 0 0 - - - Hylidae Hyla pulchella - - 1.50 - 0 0 0 0 0 0 0 0 0 Hylidae Hyla punctata 36.80 36.40 1.50 324 1 0 0 0 0 - 0 0 0 Hylidae Hyla raniceps - - 1.35 2100 1 0 0 0 0 - 0 0 0 Hylidae Hyla rhodopepla 26.67 20.73 1 285 0 - - - - - 0 0 0 Hylidae Hyla riveroi 22.80 18.50 - - 0 0 0 0 0 - - - -

99 Hylidae Hyla robertmertensi 26.6 24.70 - - 0 0 0 0 0 - - - - Hylidae Hyla rosenbergi - - 1.95 2400 0 1 0 0 1 1 1 1 0 Hylidae Hyla rubicundula - - - - 0 0 0. 0 0 - - - - Hylidae Hyla rufitela - - 1.80 - 0 0 0 0 0 1 0 0 0 Hylidae Hyla sanborni - - 0.90 60 0 0 0 0 0 0 0 0 0 Hylidae Hyla sarayacuensis 33.50 25 2 113 0 0 0 0 0 1 0 0 0 Hylidae Hyla sartori - - - - 0 0 0 0 0 0 - - - Hylidae Hyla savignyi - - 1.25 600 1 0 0 0 0 0 0 0 0 Hylidae Hyla semiguttata - - - - 0 0 0 0 0 - - - - Hylidae Hyla senicula - - - - 0 0 0 0 0 - - - - Hylidae Hyla sibleszi 35.0 32.3 2.30 42 0 0 0 0 0 - 0 0 0 Hylidae Hyla simmons i -• - - - 0 - - - - - 0 0 0 Hylidae Hyla smithii 27.7 24.3 - - 0 0 0 0 0 - - - - Hylidae Hyla squirella - - 0.90 950 0 1 0 0 0 0 0 0 0 Hylidae Hyla sumichrasti 30.20 26.20 1.61 - 0 0 0 0 0 - 0 0 0 Hylidae Hyla taeniopus 64.2 58.0 2.19 - 0 0 0 0 0 1 0 0 0 Hylidae Hyla thorecles - - 5.10 10 0 0 0 0 0 - 0 - 0 0 Hylidae Hyla tica 38.90 31.60 2.32 192 0 0 0 0 0 . 1 0 0 0 Hylidae Hyla triangulum 35.40 25.35 1.50 501 0 0 0 0 0 - 0 0 0 Hylidae Hyla versicolor - - 1.15 1800 0 1 0 0 1 1 0 0 0 Hylidae Hyla walked 31.6 32.0 - - 1 , 0 0 0 0 - - - - Hylidae Hyla wrightorum - -• 1.20 - 0 0 0 0 0 0 0 0 0 Hylidae Hyla zeteki 25.4 22.5 - 24 0 0 0 0 0 - 1 0 1 Hylidae Hyloscirtus palmeri - - - - 0 0 0 0 0 - 0 0 0 Hylidae Hypsiboas albomarginatus 57.30 48.60 - ' 0 0 0 0 0 - - - Hylidae Hypsiboas cinerascens 41.20 39 1.50 426 0 0 0 0 0 - 0 0 0 Hylidae hthmohyla rivularis 35.70 32.30 2.36 90 0 0 0 0 0 1 - - - Hylidae Litoria arfakiana - - - - 0 0 0 0 0 - - - - Hylidae Litoria armatus 64.30 59.60 - 0 0 0 0 0 - - - Hylidae Litoria aurea - - 1.40 - 0 0 0 0 0 1 0 0 0 Hylidae Litoria caerulea - - 1.30 1800 0 0 0 0 0 1 0 0 0 Hylidae Litoria freycineti - - 1.50 - 0 0 0 0 0 1 0 0 0 Hylidae Litoria infrafrenata - - 3 300 - 0 0 0 0 1 0 0 0 Hylidae Litoria meiriana - - - 37 - 0 0 0 0 0 0 0 0 Hylidae Litoria peronii - - 1.50 - 0 1 0 0 1 -" 0 0 0 Hylidae Litoria rubella - - 1.05 170 0 0 0 0 0 0 0 0 0 Hylidae Nyclimantis rugiceps 61.3 61.87 - - 1 0 0 0 0 - 1 0 1 Hylidae Nyctimystes cheesmanae - - - - 0 0 0 0 0 - - - - Hylidae Nyctimystes foricula - - - - 0 0 0 0 0 - - - -

100 Hylidae Nyctimysles kubori • - - - - 0 0 0 0 0 - - - - Hylidae Nyctimystes papua - - - - 0 0 0 0 0 - 0 0 0 Hylidae Osteocephalus buckleyi 61.70 43.30 1.35 1600 0 0 0 0 0 - 0 0 0 Hylidae Osteocephalus langsdorffii - - 1 - 0 0 0 0 0 - - - - Hylidae Osteocephalus leprieurii 57.10 44.70 1 848 0 0 0 0 0 - 0 0 0 Hylidae Osteocephalus oophagus - - 1.74 516 - 0 0 0 0 1 1 0 1 Hylidae Osteocephalus taurinus 81.01 69.90 1 550 0 0 0 0 0 - 0 0 0 Hylidae Osteocephalus verruciger 64.5 53.0 - - 0 0 0 0 0 - - - - Hylidae Osteopilus brunneus 60.5 45.5 0.71 552 0 0 0 0 0 - 1 0 1 Hylidae Osteopilus crucialis - - - - 0 0 0 0 0 - 1 0 1 Hylidae Osteopilus dominicensis 76.8 57.8 - - 0 0 0 0 0 - - ' -

Hylidae Osteopilus marianae - - - - 0 0 0 0 0 - - •- - Hylidae Osteopilus pulchrilineatus 40.10 31.6 - - 0 0 0 0 0 - - - - Hylidae Osteopilus septentrionalis 71.23 53.90 3 130 0 1 0 1 0 0 0 0 0 Hylidae Osteopilus vastus - - 2.50 - 0 0 0 0 0 - - - - Hylidae Osteopilus wilderi 27.3 25.8 - - 0 - ' ------. Hylidae Pachymedusa dacnicolor 79.2 67.60 2.55 467 0 1 0 1 1 1 0 0 0 Hylidae Phrynohyas coriacea - - 2 1430 0 - - - - - 0 0 0 Hylidae Phrynohyas mesophaea - - - 700 0 1 0 1 0 - 0 0 0 Hylidae Phrynohyas resinifictrix - - 1.58 436 0 1 0 0 1 1 1 0 1 Hylidae Phrynohyas venulosa 98.10 88.60 2.75 5635 1 0 0 0 0 0 0 0 0 Hylidae Phyllodytes luteolus - - 1.10 13 ------Hylidae Phyllomedusa atelopoides - - 3 20 0 0 0 0 0 1 I 0 1 Hylidae Phyllomedusa bicolor - - - - 0 0 0 0 0 - 0 0 0 Hylidae Phyllomedusa hypochondrialis - - - - 0 1 0 0 1 - 0 0 0 Hylidae Phyllomedusa lemur 42.63 33.80 3.25 45 0 1 0 1 0 1 0 0 0 Hylidae Phyllomedusa palliata 46.30 42.10 2.50 60 0 - - - - - 0 0 0 Hylidae Phyllomedusa tarsia 104.90 86.90 3 548 0 0 0 0 0 - 0 0 0 Hylidae Phyllomedusa tomopterna 57.20 45.30 3.38 71 0 - - - - - 0 0 0 Hylidae Phyllomedusa vaillantii 78.80 52.10 2.23 1114 0 0 0 0 0 1 0 0 0 Hylidae Plectrohyla glandulosa 44.3 44.6 - - 1 0 0 0 0 - - - - Hylidae Plectrohyla guatemalensis 48.60 47.50 2 - 1 0 0 0 0 - 0 0 0 Hylidae Plectrohyla matudai - - - - 0 0 0 0 0 - 0 0 0 Hylidae Pseudacris brachyphona 30.30 24.60 1.55 950 0 0 0 0 0 1 0 0 0 Hylidae Pseudacris brimleyi - - - 300 0 0 0 0 0 • 1 0 0 0 Hylidae Pseudacris cadaverina 40.9 33.0 1.95 - 0 0 0 0 0 1 0 0 0 Hylidae Pseudacris clarkii . - 0.78 1000 0 0 0 0 0 1 0 0 0 Hylidae Pseudacris crucifer 31.80 25.20 1.13 750 0 1 0 0 1 1 0 0 0 Hylidae Pseudacris feriarum - - 1 - 0 0 0 0 0 1 0 0 0

101 Hylidae Pseudacris maculata - - 1.30 - 0 0 0 0 0 0 0 0 0 Hylidae Pseudacris nigrita - - 0.95 59 0 0 0 0 0 1 0 0 0 Hylidae Pseudacris ocularis - - 0.70 150 0 . 0 0 0 0 1 0 0 0 Hylidae Pseudacris ornata - - 0.95 55 0 0 0 0 0 1 0 0 0 Hylidae Pseudacris regilla 37.20 32.50 1.30 807 0 1 0 0 1 1 0 __i 0 0 Hylidae Pseudacris streckeri - - 1.38 450 0 0 0 0 0 1 0 0 0 Hylidae Pseudacris triseriata - - 0.98 449 0 0 0 0 0 0 0 0 0 Hylidae Pseudis paradoxa - - - - - 0 0 0 0 - 0 0 0 Hylidae Pternohyla fodiens 61.1 49.4 - - 0 0 0 0 0 1 - - - Hylidae Ptychohyla euthysanota 38.2 35.0 - - 0 0 0 0 0 1 0 0 0

Hylidae Ptychohyla hypomykter •- - 2.38 ------0 0 0 Hylidae Ptychohyla leonhardschultzei 39.90 31.60 - - 0 0 0 0 0 1 - - - Hylidae Ptychohyla spinipollex 42.8 37.1 - - 0 0 0 0 0 - - - - Hylidae Scarthyla goinorum - - - - 0 - - - - - 0 0 0 Hylidae Scinax berthae 25 19 0.90 - 0 0 0 0 0 0 0 0 0 Hylidae Scinax boulengeri 49.40 41.6 1.55 650 0 0 0 0 0 1 0 0 0 Hylidae Scinax catharinae 42.67 33 - - 0 0 0 0 0 - 0 0 • 0 Hylidae Scinax elaeochraoa 35 30.60 - - 0 1 0 0 1 0 0 0 0 Hylidae Scinax fuscovarius 47.55 44.58 1 2892 0 0 0 0 0 1 0 0 0 Hylidae Scinax garbei 42 35.40 1.50 550 0 0 0 0 0 - 0 0 0 Hylidae Scinax nasicus - - - - 1 0 0 0 0 - - - - Hylidae Scinax ruber 39.13 32.60 1.50 591 0 0 0 0 0 1 0 0 0 Hylidae Scinax squalirostris - - 1 - 0 0 0 0 0 - 0 0 0 . Hylidae Scinax staufferi 26.60 24.10 - - 0 0 0 0 0 1 . - - - Hylidae Smilisca baudinii - - 1.30 2225 0 0 0 0 0 0 0 0 0 Hylidae Smilisca cyanosticta - - 1.24 1147 0 0 0 0 0 1 0 0 0 Hylidae Smilisca phaeota - - - 1829 0 0 0 0 0 1 0 0 0 Hylidae Smilisca puma - - - - 0 0 0 0 0 1 0 0 0 Hylidae Sphaenorhynchus lacteus - - 1 - 0 0 0 0 0 - 0 0 0 Hylidae Tlalocohyla picta - - - - 0 0 0 0 0 - - - - Hylidae Triprion petasatus 70.70 54.60 - - 0 0 o. 0 0 0 - - - Hylidae Xenohyla truncata - - - - 0 0 0 0 0 - - - - Hyperoliidae Afrixalus fornasini - - 1.60 80 Q 0 0 - . - 0 0 0 Hyperoliidae Alexteroon obstelricans - - - - 0 0 0 - - - 1 0 1 Hyperoliidae Hyperolius puncticulatus - - - - 0 0 0 - - - - ' - - Hyperoliidae Hyperolius tuberilinguis - - 1.50 350 0 0 0 - - - 0 0 0 Hyperoliidae Kassina senegalensis - - 1.60 400 1 - - - - - 0 0 0 Leiopelmatidae Ascaphus truei - - 4.50 28 0 0 0 - - 1 0 0 0 Leiopelmatidae Leiopelma archeyi - - 4.50 - 0 - - - - - 1 1 0

102 Leiopelmatidae Leiopelma hochstetteri - - 5.50 - 0 0 0 - - - 1 1 0 Leiuperidae Edalorhina perezi 32.20 26.90 2 93 0 - - - - - 1 - - Leiuperidae Pleurodema brachyops - - - 0 0 0 - - - 1 - - Leiuperidae Pseudopaludicola falcipes - - 1 - 0 0 0 - - 1 0 0 0 Leptodactylidae Leptodactylus fuscus 42.60 41.10 2 - 1 0 0 - - - 1 1 0 Leptodactylidae Leptodactylus gracilis 43 43 - - 1 0 ' 0 - - - 1 - - Leptodactylidae Leptodactylus hylaedactylus 23.20 22.70 - - 1 0 0 ------Leptodactylidae Leptodactylus ocellalus - - - 1 0 0 - - 1 1 1 1 Leptodactylidae Lithodytes Meatus - - 2 195 0 - - - - - 1 - - Leptodactylidae Vanzolinius discodactylus 33.60 27.40 1 234 0 - - - - - 0 0 0 Limnodynastidae Adelotus brevis 32.75 • 35.53 1.98 - 1 i 0 0 1 1 1 1 0 Limnodynastidae Lechriodus fletcheri - - 1.70 300 0 0 0 - - 1 1 - - Limnodynastidae Limnodynastes dumerilii - - 1.70 3950 0 1 0 0 1 1 1 0 1 Limnodynastidae Limnodynastes ornatus - - 1.20 645 0 0 0 - - 1 1 - - Limnodynastidae Limnodynastes peronii 46.60 51.70 1.50 850 1 1 0 0 1 1 1 - - Limnodynastidae Limnodynastes salmini - - - 2000 0 0 0 - - 1 0 0 0 Limnodynastidae Neobatrachus pictus - - - 1000 0 0 0 - - 1 0 0 0 Limnodynastidae Notaden melanoscaphus - - 1.40 833 0 0 0 - - 0 0 0 0 Limnodynastidae Philoria sphagnicolus - - - 48 0 0 0 - - - 1 1 0 Aglyptodactylus Mantellidae 1.35 1600 0 0 0 0 0 0 madagascariensis - - - - - Mantel lidae Boophis lephraeomyslax - - 1.35 - 0 0 0 - - - 0 0 0 Mantellidae Mantella aurantiaca - - 1.75 40 0 0 0 - - - 1 0 1

Mantellidae Mantidactylusjemoralis - • - 2.50 - 0 0 0 - - 1 0 0 0 Megophryidae Leptobrachium banae - - - - 0 0 0 0 0 - - - - Megophryidae Leptobrachium hasseltii 74.28 55.73 - - 0 0 0 0 0 - - - - Megophryidae Leptobrachium montanum 60.20 52.10 - - 0 - - . ------Megophryidae Leptobrachium mouholi - - - - 0 0 0 0 0 1 - - - Megophryidae Leptolalax liui - - - 1 0 0 0 0 - - - - Megophryidae Leptolalax pelodyloides 29.23 25.95 2 144 0 0 0 0 0 - - - - Megophryidae Oreolalax jingdongensis - - 3.60 170 1 0 0 0 0 0 - - - Megophryidae Oreolalax liangbeiensis 60 51.6 3.50 350 0 0 0 0 0 0 - - - Megophryidae Oreolalax multipunctatus - - 3.5.0 - - 0 o . 0 0 . 0 1 1 0 Megophryidae Oreolalax omeimontis - - - - - 0 0 0 0 0 - - - Megophryidae Oreolalax pingii 45.60 45.20 3.40 175 1 0 0 0 0 0 - - - Megophryidae Oreolalax popei 61.90 65.2 3.25 350 1 . 0 0 0 0 0 - - -

Megophryidae Oreolalax rugosus 49.70 47.10 - • - 0 0 0 0 0 0 - - - Megophryidae Oreolalax schmidli - - 4 120 0 0 0 0 0 0 - - - Megophryidae Scutiger boulengeri 53.80 47.43 3 380 0 0 0 0 0 0 - - -

103 Megophryidae Scutiger glandulatus 66.50 67 3.50 - 0 0 0 0 0 1 0 Megophryidae Scutiger mammatus 69.16 70.46 2.85 606 0 0 0 0 0 1 0 Megophryidae Scutiger muliensis - - - 200 0 0 0 0 0 - - Megophryidae Scutiger tuberculatus - - 3 - 0 0 0 0 0 0 - - Megophryidae Vibrissaphora ailaonica 72.90 75.55 3.50 245 0 0 0 0 0 1 0 Megophryidae Vibrissaphora boringiae 66.8 76.7 3 298 0 0 0 0 0 1 0 Megophryidae Vibrissaphora chapaense 71.60 54.35 - - 0 0 0 0 0 0 - - Megophryidae Vibrissaphora leishanensis - - - 3.90 295 0 0 0 0 0 0 0 Megophryidae Vibrissaphora liui 70.30 86.70 3.40 334 0 0 0 0 0 1 0 Megophryidae Vibrissaphora promustache 61.10 56.70 - - 0 0 0 0 0 - - - - Microhylidae Chaperina fusca 22.40 19.17 - - 0 0 0 - - 1 - - - Microhylidae Gastrophryne olivacea - - 0.85 622 0 0 0 - - 0 0 0 0 Microhylidae Kalophrynus pleurosligma 44.83 40.38 1 4000 0 - - - - - 0 0 0 Microhylidae Kaloula pulchra 63.55 60.88 - - 0 ------Microhylidae Microhyla heymonsi 22.19 19.83 0.70 157 0 0 0 - - 0 0 0 0 Microhylidae Phrynomantis bifasciatus - - 1.30 600 0 0 0 - - - 1 1 0 Myobatrachidae Crinia nimbus - - 3.50 10 0 0 0 - - - 1 1 0 Myobatrachidae Criniasignifera 21.73 18.80 1.50 150 0 0 0 - - 0 0 0 0 Myobatrachidae Geocrinia vicloriana - - 2.20 126 0 0 0 - - - 0 0 0 Myobatrachidae Metacrinia nichollsi - - - 27 0 0 0 - - - 0 0 0 Myobatrachidae Myobatrachus gouldii - 5.06 40 0 0 0 - - 0 0 0 0 Myobatrachidae Paracrinia haswelli - - - - 0 0 0 - - 0 0 0 Myobatrachidae Pseudophryne bibroni - - 2.05 82 0 1 0 0 1 0 1 .1 0 Myobatrachidae Rheobatrachus silus - - 4.70 25 0 0 0 - - 1 1 0 1 Myobatrachidae Spicospina flammocaerulea - - - 200 0 0 0 - - 1 0 . 0 0 Myobatrachidae Uperoleia laevigata - - - - 0 0 0 - - 1 0 0 0 Pelobatidae Pelobates fuscus - - 1.65 1740 1 1 0 1 0 0 0 0 0 Phrynobatrachidae Phrynobatrachus mababiensis - - 0.85 - 0 0 0 - - 1 0 0 0 Phrynobatrachidae Phrynobatrachus natalensis - - 1 650 0 0 0 - - 0 0 0 0 Pipidae Hymenochirusboettgeri - - 0.90 649 0 1 0 0 1 0 0 0 0 Pipidae Pipa carvalhoi 51.40 45.70 2.05 140 0 1 0 0 1 1 1 0 1 Pipidae Pipa pipa 139.40 128.70 6 80 1 1 0 0 1 - 1 0 1 Pipidae Xenopus laevis - - 1.40 17000 0 1 0 0 1 1 0 0 0 Ptychadenidae Ptychadena anchielae - - - - 0 0 0 ------Ptychadenidae Ptychadena mascareniensis - - 1.10 - 0 0 0 - - - 0 0 0 Pyxicephalidae Afrana angolensis - - 1.50 - 0 0 0 - - 0 0 0 0 Pyxicephalidae Afrana fuscigula - - 1.50 15000 0 0 0 - - 0 0 0 0 Pyxicephalidae Anhydrophryne rattrayi - - - 15 0 0 0 - - - 0 0 0 Pyxicephalidae Aphanlophryne pansa - - - 17 0 0 0 - - - 0 0 0

104 Pyxicephalidae Natalobatrachus bonebergi - - 2.60 87 0 0 0 - - - 0 0 0

Pyxicephalidae Strongylopus grayii - • - - - 0 0 0 - - - 0 0 0 Pyxicephalidae Tomopterna delalandii - - 1.50 .-' 0 0 0 - - - 0 0 0 Ranidae Amolops chapaensis 92.20 81.10 '- - 0 ------Ranidae Amolops chunganensis 48.05 39.59 3 436 0 1 0 1 0 0 0 0 0 Ranidae Amolops cremnobatus - - 3.60 - 0 0 0 0 0 - 0 0 0 Ranidae Amolops manlzorwn 65.80 52.20 2.50 - 0 0 0 0 0 0 0 0 0 Ranidae Amolops ricketti 60.67 55.87 3 - 0 0 0 0 0 0 0 0 0 Ranidae Amolops viridimaculatus 98 74.80 - - 0 0 0 0 0 - 0 0 0 -Ranidae Amolops wuyiensis 48.19 41.82 2 600 0 0 0 0 0 0 0 0 0 Ranidae Fejervarya nicobariensis 50.11 42.77 - - 0 0 0 0 0 1 0 0 0 Ranidae Hydrophylax galamensis - - 1.70 5135 0 0 0 0 0 1 0 0 0 Ranidae Lankanectes corrugatus - - - - 0 0 0 0 0 - 0 0 0 Ranidae Meristogenys jerboa 67.09 36.40 1.30 2462 0 0 0 0 0 1 0 0 0 Ranidae Meristogenys kinabaluensis 88 63.57 2.50 - 0 ------Ranidae Rana adenopleura 52.15 50.20 2.20 352 0 0 0 0 0 1 1 1 0 Ranidae Rana amurensis 52.75 41 2.30 2045 0 0 0 0 0 0 0 0 0 Ranidae Rana andersonii 77.25 63.50 - 0 0 0 0 0 0 - - - Ranidae Rana areolata - - 2.50 7000 0 1 0 1 0 1 0 0 0 Ranidae Rana arvalis - - 2 1599 1 1 0 1 0 1 0 0 0 Ranidae Rana asiatica 53.80 53.40 2.30 1150 1 0 0 0 0 0 0 0 0 Ranidae Rana aurora - - 3.56 800 0 1 0 1 0 0 0 0 0 Ranidae Rana berlandieri - - - - 0 0 0 0 0 1 0 0 0 Ranidae Rana blairi - - - 5250 - 0 0 0 0 1 0 0 0 Ranidae Rana boylii 73 56 '2.48 980 0 0 0 0 0 1 0 0 0 Ranidae Rana capita - - 2 7000 0 0 0 0 0 1 0 0 0 Ranidae Rana cascadae - - 2.80 400 0 0 0 0 0 0 0 0 0 Ranidae Rana catesbeiana 119.30 119.30 1.70 18171 1 1 0 0 1 1 0 0 0 Ranidae Rana chalconota - - - - 0 0 0 0 0 - - - - Ranidae Rana chaochiaoensis 58.90 - - - 0 0 0 0 0 - 0 0 0 Ranidae Rana chensinensis 41.90 48 - 1400 1 0 0 0 0 0 0 0 0

Ranidae Rana chiricahuensis - - • - 892 - - 0 0 0 0 1 0 0 0 Ranidae Rana clamitans - - - 1.50 5500 0 1 0 0 1 1 0 0 0 Ranidae Rana daemeli - - - - - 1 0 1 0 - 0 0 0 Ranidae Rana dalmatina 56.70 45.20 3 1068 1 1 0 0 1 0 0 0 0 Ranidae Rana dybowskii - - 3 1550 1 0 0 0 0 1 0 0 0 Ranidae Rana emeljanovi 55.40 46 1.80 1130 0 0 0 0 0 0 0 0 0 Ranidae Rana erythraea 65.73 39.66 2 - 0 0 0 0 0 1 0 0 0 Ranidae Rana forreri - - - - 0 0 0 0 0 1 - - ~N -

105 Ranidae Rana graeca - - 2.75 1100 - 0 0 0 0 1 1 1 0 Ranidae Rana grahami 93.30 74.30 2.50 2446 0 0 0 0 0 0 0 0 0 Ranidae Rana grylio .- - - 7504 0 0 0 0 0 1 0 0 0 Ranidae Rana guentheri 74.93 69.80 1.50 3000 0 0 0 0 0 0 0 0 0 Ranidae Rana heckscheri - - 1.75 7000 0 0 0 0 0 1 0 0 0 Ranidae Rana japonica 53.65 48.05 2.10 1500 0 1 0 1 0 1 0 0 0 Ranidae Rana latastei - - - - - 1 0 1 0 0 - - - Ranidae Rana lessonae - - 1.80 1829 1 1 0 1 0 0 0 0 0 Ranidae Rana livida 100.38 56.50 3.60 1433 0 0 0 0 0 0 0 0 0 Ranidae Rana luteiventris - - 2.12 2400 - 0 0 0 0 1 0 0 0 Ranidae Rana macrocnemis - - - 2040 0 0 0 0 0 0 0 0 0 Ranidae Rana margaretae - - - - 0 0 0 0 0 - - - - Ranidae Rana minima 28.60 27 - 274 0 0 0 0 0 1 0 .0 0 Ranidae Rana muscosa - - 2.30 223 0 0 0 0 0 1 0 0 0 Ranidae Rana nigromaculata 70.39 64.36 2.20 3250 1 0 0 0 0 0 0 0 0 Ranidae Rana nigrovittata 47.20 42.70 1.40 330 0 0 0 0 0 - 0 0 0 Ranidae Rana okinavana - - - 500 0 0 0 0 0 1 0 0 0 Ranidae Rana omeimontis - - 2 - - 0 0 0 0 0 0 0 0 Ranidae Rana onca - - - 200 0 0 0 0 0 1 0 0 0 Ranidae Rana ornativentris 58.46 49.25 1.70 12000 0 0 0 0 0 - 0 0 0 Ranidae Rana palmipes 116.80 92.30 2 2860 0 0 0 0 0 1 0 0 0 Ranidae Rana palustris 66.40 53 1.60 2500 0 1 0 1 0 1 0 0 0 Ranidae Rana pipiens 78.60 69.80 2 3045 0 1 0 1 0 1 0 0 0 Ranidae Rana pleuraden 49.80 51 2 606 1 0 0 0 0 1 0 0 0 Ranidae Rana schmackeri 76.83 40.80 2 1549 0 0 0 0 0 0 0 0 0 Ranidae Rana septentrionalis - - - 1.68 500 0 0 0 0 0 0 0 0 0 Ranidae Rana sevosa 85.35 80.55 1.84 6600 1 0 0 0 0 - •0 0 0 Ranidae Rana shuchinae 44.50 39.90 - 180 0 0 0 0 0 0 0 0 0 Ranidae Rana signata 56.17 39.43 1.70 - 0 0 ' 0 0 0 - 0 0 0 Ranidae Rana sphenocephala 63.70 55.80 1.60 1500 0 0 0 0 0 1 0 0 0 Ranidae Rana sylvatica 61.10 53.60 2.40 781 0 1 0 1 0 0 0 0 0 Ranidae Rana taipehensis 39.20 - - - 0 0 0 0 0 1 0 0 0 Ranidae Rana iarahumarae - - 2.20 2200 - 0 0 0 0 0 0 0 0 Ranidae Rana temporalis - - 2.50 - 0 0 0 0 0 - 0 0 0 Ranidae Rana temporaria 61.20 64.20 2.35 2247 1 1 0 1 0 0 0 0 0 Ranidae Rana iienlaiensis 31.27 30.68 1.50 1350 1 0 0 0 0 0 0 0 0 Ranidae Rana tsushimensis 49.75 41.10 2 450 1 0 0 0 0 1 0 0 0 Ranidae Rana vaillanti - - - - 0 0 0 0 0 1 - - - Ranidae Rana versabilis 62.30 46.40 3 101 0 0 0 0 0 - 0 0 0

106 Ranidae Rana vibicaria - - 2.50 - 0 0 0 0 0 1 0 0 0 Ranidae Rana virgatipes - - 1.80 400 0 1 0 0 1 1 0 0 0 Ranidae Rana warszewitschii - - - - 0 0 0 0 0 - - - - Ranidae Rana yavapaiensis - - - - - 0 0 0 0 1 0 0 0 Ranidae Staurois latopalmatus 61.38 45.26 - - 0 ------Ranidae Staurois natator 45.83 36.01 1.90 - 0 0 0 0 0 1 0 0 0 Rhacophoridae Buergeria japonica 34.91 29.92 1.30 - 0 0 0 - - 1 0 0 0 Rhacophoridae Chirixalus doriae 27.41 21.61 1.35 - 0 0 0 - - - 0 0 0 Rhacophoridae Chirixalus vittatus 25.67 23.60 1.50 217 0 0 0 - - 1 0 0 0 Rhacophoridae Chiromantis xerampelina - - , 1.75 192 0 1 0 1 0 - 1 0 1 Rhacophoridae Philautus gracilipes - - - - 0 - - - - - 0 0 0 Rhacophoridae Polypedates leucomyslax 62.73 43.54 1.50 337 0 0 0 - - 1 0 0 0 Rhacophoridae Rhacophorus bipunctatus - - - - 0 0 0 - - - 0 0 0 Rhinophrynidae Rhinophrynus dorsalis - - 4.50 5000 0 0 0 - - 0 0 0 0 Scaphiopodidae Scaphiopus couchii - - 1.50 3310 0 0 0 - - 0 0 0 0 Scaphiopodidae Scaphiopus holbrookii - - 1.70 - 0 0 0 - - 0 0 0 0 Scaphiopodidae Spea hammpndii 49.25 48.25 1.55 400 1 0 0 - - 0 0 0 0 Sooglossidae Sooglossus sechellensis - '- - - 0 - - - - - 1 1 0

107 APPENDIX 2 References for body size, egg size, clutch size, mating combat, length of breeding season, and parental care in anurans species.

Mating Breeding Descriptions Length of Descriptions (For Defining Species Body Size Egg Size Clutch Size Mating Combat Breeding Parental Care (For Defining Absence of Season Absence of Mating Parental Care) Combat) Bush, 1996; Alytes obstetricans Summers, 2006 Eens, 2000 Bush, 1996 N/A AmphibiaWeb Marquez, 1993 N/A Duellman, 1994 Discoglossus pictus N/A AmphibiaWeb AmphibiaWeb •N/A AmphibiaWeb Wells, 1977 N/A AmphibiaWeb Crump, 1974; A llobales femoralis Silverstone, Crump, 1974 Crump, 1974 Narin, 2004 N/A N/A Rodriguez, 1994 N/A 1976 Silverstone, Allobates zaparo , N/A N/A N/A N/A N/A AmphibiaWeb N/A 1976 Colostethus awa Coloma, 1995 N/A Coloma, 1995 Coloma, 1995 N/A N/A Coloma, 1995 N/A Colostethus beebei N/A Bourne, 2001 Bourne, 2001 • N/A Prohl, 2005 N/A Bourne, 2001 N/A Colostethus Caldwells, 2003 N/A N/A N/A N/A N/A N/A N/A brunneus Colostethus Caldwells, 2003 N/A N/A Lima, 2002 ' N/A N/A Lima, 2002 N/A caeruleodactylus Colostethus Duellman, 1993 N/A Edwards, 1971 N/A N/A N/A Coloma, 1995 N/A elachyhistus Colostethus flotator Savage, 2002 N/A N/A Savage, 2002 N/A N/A Savage, 2002 N/A Colostethus Duellman, 2004 N/A N/A N/A N/A N/A N/A N/A idiomelus Colostethus Coloma, 1995 N/A N/A N/A N/A N/A N/A N/A infraguttatus Colostethus N/A N/A N/A Duellman, 1966 N/A Duellman, 1966 N/A inguinalis N/A

108 Colostethus Duellman, 2004 N/A N/A N/A N/A N/A N/A N/A insulatus Coloma, 1995; Colostethus nexipus N/A N/A N/A N/A N/A N/A N/A Duellman, 2004 Colostethus nidicola Caldwells, 2003 Caldwells, 2003 Caldwells, 2003 N/A Caldwells, 2003 Caldwells, 2003 N/A Caldwells, 2003 Colostethus Summers, 2000; Savage, 2002 N/A N/A N/A N/A Duellman, 1994 N/A nubicola Prohl, 2005 Colostethus N/A Stebbins, 1959 Stebbins, 1959 N/A Bernal, 2005 N/A Duellman, 1994 N/A palmatus Colostethus N/A N/A N/A N/A Duellman, 1966 N/A AmphibiaWeb N/A panamensis Colostethus pratti N/A N/A N/A N/A N/A N/A Duellman, 2004 N/A Colostethus stepheni Caldwells, 2003 N/A Junca, 2006 Junca, 2006 N/A Junca, 2006 Junca, 2006 N/A Colostethus Stebbins, 1959 Stebbins, 1959 Stebbins, 1959 N/A Navas, 2001 Stebbins, 1959 Stebbins, 1959 N/A subpunctatus Colostethus Duellman, 1993; N/A N/A N/A N/A N/A N/A N/A sylvaticus Duellman, 2004 Colostethus N/A Savage, 2002 Prohl, 2005 N/A Savage, 2002 Duellman, 2004 talamancae Vences, 1998 N/A Colostethus Coloma, 1995; Global N/A N/A N/A Edwards, 1971 N/A Amphibian vertebralis Edwards, 1971 N/A Assessment Mannophryne Global N/A N/A N/A Wells, 1977 collaris N/A N/A Amphibian N/A Assessment Mannophryne Sexton, 1960 N/A Jowers, 2005 Sexton, 1960 N/A N/A Duellman, 1994 trinitatis N/A Phyllobates Silverstone, N/A N/A N/A N/A N/A N/A N/A aurotaenia 1976 Silverstone, Global 1976; Animal Animal Animal Phyllobates bicolor N/A Amphibian N/A Myers, 1996; Diversity Web N/A Diversity Web Diversity Web Assessment Myers, 1978 Silverstone, www.poison- Phyllobates lugubris N/A N/A Savage, 2002 N/A Savage, 2002 N/A 1976 frogs.com

109

N N V/ N N Phyllobates V/ Zimmermannn,

Myers, 1978 Myers, 1978 Myers, 1978 Zimmermann, ' N/A

terribilis 1985 1985

N N Silverstone, V/ Silverstone, Phyllobates vittalus Summers, 2006 Savage, 2002 1976;

1976 N/A Duellman, 1994 N/A

Summers, 2000

N N Trichobatrachus V/ robustus AmphibiaWeb . N/A N/A N/A AmphibiaWeb Beck, 1998 N/A Bombina bombina N/A Kuzmin, 1999 Kuzmin, 1999 Kuzmin, 1999 N/A Kuzmin, 1999 N/A N/A Bombina Yang, 1991 N/A N/A N/A Fei, 1999 Yang, 1991 N/A microdeladigitora N/A Bombina orientalis Okada, 1966 Kuzmin, 1999 Kuzmin, 1999 Kuzmin, 1999 N/A Kuzmin, 1999 N/A N/A j . N/A N/A Kuzmin, 1999 Seidel, 1999 N/A Wells, 1977 N/A N/A Degenhardt, Eleutherodactylus N/A N/A 1996; N/A

augusti N/A Stebbins, 2003 Duellman, 1994 N/A

Wright, 1995

N N Eleutherodactylus V/ N/A N/A N/A N/A N/A binotatus N/A AmphibiaWeb Eleutherodactylus Savage, 2002 N/A N/A N/A N/A N/A bufoniformis Savage, 2002 Savage, 2002 Eleutherodactylus

N/A N/A

planirostris Schwartz, 1991 N/A Schwartz, 1991 Schwartz, 1991 Schwartz, 1991 N/A

N N Ischnocnema V/

Duellman, 1996 Crump, 1974 Crump, 1974 N/A N/A N/A N/A

quixensis

N N Breviceps V/

Stewart, 1967 Stewart, 1967 Stewart, 1967

mossambicus N/A AmphibiaWeb N/A N/A

N N Degenhardt, Degenhardt, V/ Bufo alvarius Wright, 1995 Wright, 1995 Wright, 1995 N/A N/A 1996 1996 Licht, 1976;

Bufo americanus Wilbur, 1978 Wright, 1995 Wright, 1995 Cherry, 1992 N/A N/A Wright, 1995

Wright, 1995

I N N | Bufo andrewsi Yang, 1991 Fei, 1999 N/A N/A AmphibiaWeb Fei, 1999 N/A N/A V/ 1

N N 1 Bufo arenarum Cei, 1980 N/A Cei, 1980 N/A V/ Penna, 1990 Cei, 1980 N/A

| BH/O arunco N/A N/A N/A Penna, 1990 N/A N/A AmphibiaWeb

N N | Bufo asper Inger, 1966 Inger, 1968 Inger, 1968 N/A N/A Inger, 196V/ 8 N/A N/A | Bufo biporcalus Inger, 1966 N/A N/A N/A Marquez, 2006 N/A N/A Degenhardt, Bufo boreas Wright, 1995 Wright, 1995 Cherry, 1992 N/A Stebbins, 1951 N/A 1996 Wright, 1995 Beebee, 2000; Beebee, 2000;

Bufo bufo Beebee, 2000 Kuzmin, 1999 Davies, 1979 N/A N/A

Hettyey, 2005 Kuzmin, 1999 Hettyey, 2005

N N Bufo californicus Wright, 1995 Wright, 1995 AmphibiaWeb N/A V/ AmphibiaWeb Wright, 1995 N/A N/A

Bufo canorus Wright, 1995 Summers, 2006 N/A Wright, 1995 Wells, 1977 N/A N/A

N N Caramaschi, V/ Bufo castaneoticus Caldwells, 2004 Caldwells, 2004 N/A N/A Caldwells, 2004 2003 Caldwells, 2004 Bufo chavin Lehr, 2001 Lehr, 2001 Lehr, 2001 N/A N/A Lehr, 2001 N/A N/A fi«/b coccifer N/A Savage, 2002 Savage, 2002 N/A Savage, 2002 Savage, 2002 N/A Savage, 2002

Stebbins, 1951; Degenhardt, Degenhardt,

Bufo cognatus Wright, 1995 Cherry, 1992 N/A N/A N/A N N V/ N Wright, 1995 1996 1996 V/

N N V/ N Bufo coniferus Savage, 2002 Savage, 2002 N/A Savage, 2002 Savage, 2002 Savage, 200V/ 2

N N V/ N N 5«/b crucifer N/A N/A N/A Schwartz, V/ 1991 Bertoluci, 2002 N/A

Sw/b dapsilis Rodriguez, 1994 N/A N/A N/A N/A N/A

N N

V/ Degenhardt, Degenhardt,

Bufo debilis Bogert, 1962 Summers, 2006 N/A N/A N/A N N 1996 1996 V/

fi«/b exsul Murphy, 2003 AmphibiaWeb N/A Schuierer, 1962 N/A AmphibiaWeb N/A

N N Bufo fastidiosus Savage, 1972 Savage, 2002 , Savage, 2002 Savage, 2002 N/A Savage, 2002 N/AV/ Savage, 2002

Bufo fowleri Hulse, 2001 N/A Hulse, 2001 N/A Given, 2002 Wright, 1995 Wright, 1995

N N Bufo garmani Channing, 2001 Summers, 2006 N/A N/A Keith, 1968 N/A N/A V/ N/A

N N fit//b glaberrimus Bartlett, 2003 N/A N/A N/A AmphibiaWeb N/A V/ N/A Bufo granulosus Stebbins, 1959 Summers, 2006 Cei, 1980 N/A Cei, 1980 Cei, 1980 Cei, 1980

Bufo guttatus Duellman, 1997 N/A N/A N/A AmphibiaWeb N/A N/A N/A

N N 5u/b gutturalis N/A Summers, 2006 Cherry, 1992 Passmore, 1981 N/A N/A V/ N/A

S«/b haematiticus Savage, 2002 N/A N/A N/A Savage, 2002 Savage, 2002 Savage, 2002

N N Bufo hemiophrys Wright, 1995 N/A N/A N/A Cocroft, 1995 Stebbins, 2003 N/A N/A V/

N N V/ N V/ N N 5«/b houstonensis N/A • N/AV/ N/A Wells, 1977 N/A Wells, 1977 N/A

N N V/ N V/ N N £H/O juxtasper Inger, 1966 V/ N/A N/A AmphibiaWeb N/A

N N Bufo kisoloensis Loveridge, 1936 N/A N/A Keith, 1968 N/A V/

N N Bufo luetkenii Savage, 2002 Savage, 2002 Savage, 200V/ 2 N/A Savage, 2002 Savage, 2002 Savage, 2002 Bufo maculatus Stewart, 1967 Summers, 2006 Cherry, 1992 N/A N/A N/A N/A f

5 Duellman, 1996 Crump, 1974 Crump, 1974 Cherry, 1992 N/A Wells, 1977 N/A N/A

N N Crump, 1974; V/ Savage, 2002; Bufo marinus Stebbins, 1959 Savage, 2002; Hoser, 1989 Savage, 2002 N/A Savage, 2002 Stebbins, 1959 Wright, 1995 Batraciens et Batraciens et Batraciens et Bufo mauritanicus N/A Summers, 2006 reptiles du N/A reptiles du reptiles du N/A N/A monde monde monde Bufo melanochlorus Savage, 2002 N/A N/A N/A Savage, 2002 Savage, 2002 N/A N/A Bufo melanostictus Inger, 1966 N/A N/A N/A Marquez, 2006 Ye, 1993 N/A N/A Degenhardt, Degenhardt, Degenhardt, Bufo microscaphus N/A Stebbins, 1951 N/A N/A N/A 1996 1996 1996 Bufo nelsoni Wright, 1995 N/A N/A N/A Wright, 1995 Wright, 1995 N/A • Wright, 1995 Bufo pardalis Cherry, 1992 N/A N/A Cherry, 1992 N/A N/A N/A N/A Degenhardt, Degenhardt, Bufo punctalus Wright, 1995 Wright, 1995 N/A N/A N/A Wright, 1995 1996 1996 Bufo quercicus Wilbur, 1978 Wright, 1995 N/A N/A Wright, 1995 Wright, 1995 N/A N/A Bufo ranged N/A Wager, 1965 N/A N/A AmphibiaWeb Wells, 1977 N/A N/A Bufo regularis Loveridge, 1936 Wager, 1965 Wager, 1965 Tandy, 1972 N/A Stewart, 1967 N/A N/A Bufo retiformis Bogert, 1962 N/A N/A N/A Bogert, 1962 Stebbins, 2003 N/A N/A Bufo schneideri N/A Cei, 1980 N/A N/A N/A Cei, 1980 N/A Cei, 1980 Degenhardt, N/A N/A N/A Wells, 1977 N/A Bufo speciosus N/A 1996 N/A Bufo spinulosis N/A N/A N/A N/A Penna 1990 Cei, 1980 N/A N/A Bufo terrestris Wright, 1995 Wright, 1995 Wright, 1995 N/A Wright, 1995 Smith, 1950 . N/A Wright, 1995 Savage, 2002; Bufo valliceps Wright, 1995 N/A N/A Wells, 1977 N/A N/A Wright, 1995 Wells, 1977 Bufo variegatus N/A N/A • Cei, 1980 N/A Cei, 1980 Cei, 1980 N/A Cei, 1980 Bufo viridis N/A Kuzmin, 1999 Kuzmin, 1999 N/A AmphibiaWeb Ye, 1993 N/A N/A Degenhardt, Degenhardt, Degenhardt, Bufo woodhousii Wright, 1995 Stebbins, 1951 N/A N/A N/A 1996 1996 1996 Bufo xeros N/A N/A N/A N/A Tandy, 1976 N/A N/A N/A Ollotis ibarrai N/A N/A N/A N/A N/A N/A N/A N/A Schismaderma Stewart, 1967; Stewart, 1967; Stewart, 1967 N/A AmphibiaWeb Stewart, 1967 N/A N/A carens Wager, 1965 Wager, 1965 Atlophryne ruthveni Duellman, 1997 N/A Duellman, 1997 N/A Duellman, 1997 N/A N/A AmphibiaWeb Centrolenella Lynch, 1973 Savage, 2002 Savage, 2002 N/A Savage, 2002 Zug,2001 N/A prosoblepon N/A

112 Hyalinobatrachium Savage, 2002 Savage, 2002 Savage, 2002 N/A Savage, 2002 Savage, 2002 Savage, 2002 N/A fleischmanni Ingerana baluensis Inger, 1966 Inger, 1966 N/A N/A N/A N/A N/A N/A Stefania evansi Duellman, 1984 Summers, 2006 Duellman, 1984 N/A N/A Duellman, 1984 Duellman, 1984 N/A Rhinoderma The Web Site of Animal Life AmphibiaWeb AmphibiaWeb N/A AmphibiaWeb AmphibiaWeb N/A darwinii Huina-pukios Resource Dendrobates N/A N/A N/A N/A Myers, 1984 N/A Summers, 1999b N/A arboreus Silverstone, Dendrobates Dunn, 1941; 1975; Summers, 2006 Eens, 2000 N/A N/A Duellman, 1994 N/A auratus Savage, 2002 Summers, 1989 Summers, Dendrobates Silverstone, 1999b; Animal Duellman, 1994; Pino, 2007 Wells, 1977 N/A N/A azureus 1975 Animal Diversity Web Wells, 1977 Diversity Web Dendrobates www.poison- N/A N/A N/A N/A N/A Summers, 1999a N/A fantasticus frogs.com Dendrobates Silverstone, N/A N/A N/A N/A N/A N/A N/A fulguritus 1975 Dendrobates Silverstone, N/A N/A N/A N/A N/A N/A N/A galactonotus 1975 Dendrobates Silverstone, Savage, 2002; N/A Savage, 2002 Savage, 2002 N/A Savage, 2002 N/A granuliferus 1975 Summers, 1999a Silverstone, Dendrobates 1975; Duellman, 1994; Summers, 2006 N/A Summers, 1992b N/A N/A N/A histrionicus Summers, 1989; Summers, 1992b Myers, 1996 Dendrobates www.poison- www.poison- N/A N/A N/A N/A N/A N/A imitator frogs.com frogs.com Silverstone, Dendrobates 1975; Summers, 2006 Summers, 1992a Summers, 1992b N/A N/A Summers, 1992a N/A leucomelas Summers, 1989 Dendrobates Silverstone, N/A N/A N/A Myers, 1976 N/A Beck, 1998 N/A minulus 1975

113 Silverstone, Summers, Dendrobates 1975; Summers, 2006 Prohl, 2005 Wells, 1977 N/A Savage, 2002 1999a; N/A pumilio Summers, 1989 Duellman, 1994 Silverstone, Dendrobates Zimmermann, 1975; N/A N/A N/A N/A Beck, 1998 N/A quinquevittatus 1988 Myers, 1982 Dendrobates Myers, 1982 Rodriguez, 1994 Rodriguez, 1994 N/A Myers, 1982 N/A Rodriguez, 1994 N/A reticulatus Dendrobates Silverstone, N/A N/A Jungfer, 1985 N/A N/A . Summers, 1999b N/A speciosus 1975 Dendrobates Silverstone, N/A . Summers, 1999b N/A N/A N/A tinctorius 1975 Summers, 1999b N/A Dendrobates Silverstone, N/A Summers, 1999b N/A N/A N/A Summers, 1999b N/A truncatus 1975 Dendrobates www.poison- Caldwells, 1999; Myers, 1982 N/A N/A N/A . Caldwells, 1999 N/A . vanzolinii frogs.com Prohl, 2005 Dendrobates Zimmermann, www.poison- N/A N/A N/A N/A N/A N/A variabilis 1988 frogs.com Dendrobates Rodriguez, 1994 ventrimaculatus Summers, 2006 Poelman, 2007 Poelman, 2007 N/A Poelman, 2007 Poelman, 2007 N/A Epipedobates Silverstone, N/A N/A anthonyi 1976 N/A N/A N/A Beck, 1998 N/A Epipedobates Silverstone, N/A N/A N/A boulengeri 1976 N/A N/A Beck, 1998 N/A Epipedobates N/A N/A N/A espinosai N/A N/A N/A Beck, 1998 N/A Epipedobates Gottsberger, www.poison- Rodriguez, 1994 Rodriguez, 1994 Rodriguez, 1994 N/A N/A hahneli 2004 frogs.com N/A Silverstone, Epipedobates 1976; Crump, 1974 Crump, 1974 N/A N/A Zug,2001 Rodriguez, 1994 parvulus N/A Crump, 1974 Silverstone, Epipedobates petersi N/A N/A N/A N/A N/A Beck, 1998 N/A 1976

114 Silverstone, Epipedobates pictus 1976; Crump, 1974 Crump, 1974 N/A N/A N/A Beck, 1998 N/A Crump, 1974 Epipedobates Silverstone, N/A N/A N/A N/A N/A N/A .N/A pulchripectus 1976 Epipedobates Zimmermann, Myers, 1979 Summers, 2006 Myers, 1979 N/A N/A Myers, 1979 N/A silverslonei 1988 Duellman, 1993; Epipedobates www.poison- www.poison- Silverstone, N/A Hermans, 2002 N/A N/A N/A tricolor frogs.com frogs.com 1976 Silverstone, Roithmair, 1994; Rodriguez, Epipedobates Myers, 1979; 1976; Roithmair, 1994 Silverstone, . N/A N/A 1994; N/A trivittatus Rodriguez, 1994 Duellman, 1996 1976 Summers, 1999a Chaparana Yang, 1991 Ye, 1993 Yang, 1991 N/A AmphibiaWeb N/A N/A N/A unculuanus Euphlyctis Gramapurohit, N/A N/A N/A AmphibiaWeb Daniel, 1963 N/A AmphibiaWeb cyanophlyctis 2005 Fejervarya Inger, 1966 Ye, 1993 N/A Inger, 1966 N/A Alcala, 1986 cancrivora N/A N/A Fejervarya Chen, 1991; Inger, 1966 Ye, 1993 Ye, 1993 N/A N/A Ye, 1993 limnocharis Ye, 1993 Ye, 1993 Hoplobatrach us Huang, 1990; Chen, 1991 Huang, 1990 N/A N/A Huang, 1990 N/A N/A chinensis Ye, 1993 Hoplobatrachus N/A AmphibiaWeb occipitalis N/A N/A AmphibiaWeb N/A . N/A AmphibiaWeb Summers, 2006; Limnonectes blythii N/A Summers, 2006 Inger, 1968 Emerson, 1992 N/A N/A • AmphibiaWeb N/A Limnonectes finchi N/A Summers, 2006 N/A N/A N/A N/A Beck, 1998 N/A Limnonectes Ohler, 2002 N/A N/A N/A AmphibiaWeb N/A N/A gyldenstolpei N/A Limnonectes N/A N/A Inger, 1968 N/A N/A N/A N/A ibanorum N/A Chen, 1991; Limnonectes kuhlii Yang, 1991 Huang, 1990; Chen, 1991 Tsuji, 2004 N/A Chen, 1991 N/A AmphibiaWeb Pope, 1931 Limnonectes laticeps Drinfc 1979 N/A N/A N/A Dring, 1979 Inger, 1966 N/A N/A Limnonectes Inger, 1966 Summers, 2006 Inger, 1966 N/A N/A Inger, 1966 N/A N/A macrodon Limnonectes Inger, 1966 Summers, 2006 Inger, 1966 N/A N/A N/A Inger, 1966 N/A microdiscus Limnonectes Beck, 1998; N/A Summers, 2006 N/A N/A N/A N/A N/A palavanensis Summers, 2006 Limnonectes Inger, 1966 N/A- N/A N/A AmphibiaWeb N/A N/A N/A paramacrodon Limnonectes parvus N/A Inger, 1954 Inger, 1954 N/A N/A N/A N/A N/A Nannophrys Wickramasinghe Wickramasinghe Wickramasinghe N/A N/A N/A N/A N/A ceylonensis ,2004 ,2004 ,2004 Nanorana parkeri Hu, 1987 Ye, 1993 N/A N/A N/A Hu, 1987 N/A N/A Nanorana pleskei Ye, 1993 N/A N/A N/A AmphibiaWeb AmphibiaWeb N/A N/A Occidozyga laevis Inger, 1966 N/A N/A N/A N/A N/A N/A N/A Occidozyga lima Yang, 1991 Yang, 1991 Yang, 1991 N/A N/A N/A N/A N/A Occidozyga N/A N/A N/A N/A . N/A N/A martensii Pope, 1931 Yang, 1991 Paa boulengeri Yang, 1991 Liu, 1950 N/A N/A AmphibiaWeb Ye, 1993 N/A N/A Chen, 1991; Chen, 1991; Paa spinosa Chen, 1991 N/A N/A AmphibiaWeb Huang, 1990; Pope, 1931 AmphibiaWeb Huang, 1990 Pope, 1931 Yang, 1991; Paa yunnanensis Pope, 1931 Ye, 1993 N/A N/A AmphibiaWeb N/A AmphibiaWeb Ye, 1993 Hemiphractus helioi • N/A N/A N/A N/A AmphibiaWeb N/A AmphibiaWeb N/A Hemisus AmphibiaWeb Wager, 1965 Wager, 1965 marmoratum N/A Wager, 1965 N/A Wager, 1965 N/A Degenhardt, Wright, 1995; Acris crepitans Duellman, 2001 Summers, 2006 1996; Wagner, 1989 N/A Smith, 1950 N/A N/A Smith, 1950 Acris gryllus Wright, 1995 Stebbins, 1951 Stebbins, 1951 N/A Wright, 1995 Wright, 1995 N/A Wright, 1995 Agalychnis Duellman, 2001; Duellman, 2001; Duellman, 2001 Wells, 1977 N/A N/A N/A callidryas Savage, 2002 Savage, 2002 Zug, 2001 Agalychnis lilodryas Duellman, 2001 N/A N/A N/A Duellman, 2001 N/A N/A AmphibiaWeb Agalychnis saltalor Duellman, 2001 Savage, 2002 Savage, 2002 Savage, 2002 N/A Pough, 2001 N/A Savage, 2002 Agalychnis spurrelli Duellman, 2001 N/A N/A N/A Duellman, 2001 N/A N/A Savage, 2002

116 Jungfer, 1999; Savage, 2002; Anotheca spinosa • Duellman, 2001 N/A Savage, 2002 Duellman, 2001 Duellman, 1994 N/A Savage, 2002 Schiesari, 2003 Aparasphenodon N/A N/A N/A N/A AmphibiaWeb N/A Duellman, 1994 N/A brunoi Aplastodiscus N/A . Hadded, 2005 Hadded, 2005 N/A Haddad, 2005 N/A Hadded, 2005 N/A perviridis Cruzlohyla Duellman, 2001; Duellman, 2001 Savage, 2002 N/A Savage, 2002 Savage, 2002 N/A Savage, 2002 calcarifer Savage, 2002 Cyclorana Parker, 1940 Summers, 2006 N/A N/A Hoser, 1989 N/A N/A N/A alboguttata Cyclorana australis Parker, 1940 Tyler, 1983 Tyler, 1983 N/A Tyler, 1983 Tyler, 1983 N/A Tyler, 1983 Cyclorana brevipes Parker, 1940 Summers, 2006 N/A N/A AmphibiaWeb N/A N/A AmphibiaWeb Dendropsophus Duellman, 1982 N/A N/A N/A N/A N/A N/A N/A aperomeus Crump, 1974; Dendropsophus Duellman, Duellman, Crump, 1974 Crump, 1974 N/A N/A • N/A N/A parviceps 1974b 1974b Duellmanohyla Duellman, 2001 Savage, 2002 Savage, 2002 N/A Duellman, 2001 Savage, 2002 N/A N/A rufioculis Ecnomiohyla Duellman, 2001 N/A N/A Savage, 2002 N/A N/A N/A N/A miliarie Hyla albofrenata Lutz, 1973 N/A N/A N/A J Lutz, 1973 N/A N/A N/A Hyla albopunctata Lutz, 1973 N/A N/A N/A Lutz, 1973 N/A N/A N/A Hyla allenorum Rodriguez, 1994 N/A N/A N/A N/A N/A N/A AmphibiaWeb Hyla anceps Lutz, 1973 Lutz, 1973 N/A N/A Lutz, 1973 N/A N/A N/A Hulse, 2001; Hyla-andersonii Wright, 1995 Wright, 1995 Hulse, 2001 N/A Wright, 1995 N/A Hulse, 2001 Wright, 1995 Hyla andina Duellman, 1997 N/A N/A N/A Duellman, 1997 N/A N/A AmphibiaWeb Yang, 1991; Hyla annectans N/A N/A N/A Xu, 2005 AmphibiaWeb N/A N/A Ye, 1993 • Kuzmin, 1999; Hyla arborea Chen, 1991 Kuzmin, 1999 N/A Brzoska, 1982 Oseen, 2002 N/A AmphibiaWeb Okada, 1966 Hyla Caldwells, 1974; Summers, 2006 N/A N/A Duellman, 2001 N/A N/A • N/A arborescandens Duellman, 2001 Degenhardt, Degenhardt, Hyla arenicolor Wright, 1995 Stebbins, 1951 N/A N/A N/A N/A 1996 1996

117 Hyla avivoca Wright, 1995 Summers, 2006 AmphibiaWeb Wright, 1995 N/A Wright, 1995 N/A N/A //y/a balzani Duellman, 1997 N/A N/A N/A Duellman, 1997 N/A N/A N/A Hyla bifurca Crump, 1974 Crump, 1974 Crump, 1974 N/A N/A N/A N/A N/A //y/a bipunctata Lutz, 1973 N/A N/A N/A Lutz, 1973 N/A N/A N/A Hyla bischoffl Lutz, 1973 N/A N/A N/A Lutz, 1973 N/A N/A N/A Crump, 1974; Hyla boans Duellman, 1996; Crump, 1974 Crump, 1974 N/A Duellman, 2001 Duellman, 2001 Beck, 1998 N/A Duellman, 2001 Crump, 1974; Duellman,

Hyla brevifrons Duellman, Crump, 1974 Crump, 1974 N/A N/A N/A N/A

1974b

N N 1974b V/

. M M Hyla bromeliacia Lee, 2000 N/A Duellman, 2001 - N/A Duellman, 2001 Duellman, 1994 N/AV/ Hyla calcarata Crump, 1974 Cramp, 1974 Crump, 1974 N/A N/A N/A N/A Pope, 1931; Hyla chinensis Ye, 1993; Huang, 1990 Chen, 1991 N/A N/A Chen, 1991 N/A N/A Huang, 1990

Hyla chrysoscelis Wright, 1995 Hulse, 2001 Hulse,2001 Fellers, 1979 N/A Wright, 1995 N/A Hulse,2001

N N Hyla cinerea Garton, 1975 Wright, 1995 Garton, 1975 Garton, 1975 N/A Garton, 197V/ 5 N/A Wright, 1995

N N //y/a circumdata Lutz, 1973 N/A N/A N/A V/ Lutz, 1973 N/A N/A Hylacolymba Duellman, 1972 N/A N/A Savage, 2002 Savage, 2002 N/A N/A Duellman, 2001;

Hyla crepitans

Stebbins, 1959 Stebbins, 1959 Stebbins, 1959 N/A Stebbins, 1959 Wright, 1995 N/A N/A

N N Hyla ebraccata Duellman, 2001 Duellman, 2001 N/A Savage, 200V/ 2 N/A Zug, 2001 N/A Duellman, 2001

N N Hyla ehrhardti Hartmann, 2004 N/A N/A Hartmann, 2004 Hartmann,V/ 2004 N/A AmphibiaWeb 1 1 N/A N/A N/A Bastos, 1996 N/A N/A N/A //y/a euphorbiacea Duellman, 2001 ' N/A N/A N/A Duellman, 2001 Duellman, 2001 N/A N/A Degenhardt, Degenhardt, Hyla eximia Duellman, 2001 N/A N/A N/A N/A 1996 1996 AmphibiaWeb Hyla faber N/A Lutz, 1973 N/A Martins, 1998 N/A Martins, 1998 N/A Martins, 1998

Crump, 1974;

Hylafasciata Crump, 1974 Crump, 1974 N/A N/A N/A N/A N/A N N Duellman, 1996 V/ Hyla femoralis Wright, 1995 Wright, 1995 'N/A N/A Wright, 1995 Wright, 1995 N/A J. Bogart's

Hyla geographica Crump, 1974 Cramp, 1974 Crump, 1974 N/A Lutz, 1973 personal N/A. N/A

N N V/ communication t Hyla gratiosa Wright, 1995 Wright, 1995 N/A Murphy, 2003 Wright, 1995 N/A N N

Hyla guentheri Lutz, 1973 N/A N/A N/A Lutz, 1973 V/ N/A N/A Hyla heilprini Trueb, 1974 N/A N/A N/A Schwartz, 1991 N/A N/A AmphibiaWeb

Hyla japonica Okada, 1966 Kuzmin, 1999 Kuzmin, 1999 N/A Park, 1998 Kuzmin, 1999 N/A N/A

N N V/ N V/ N N //y/

Hyla labialis N/A N/A. N/A Stebbins, 1959 N/A

N N Hyla lanciformis Crump, 1974 Crump, 1974 Crump, 1974 N/A N/A N/A V/ N/A AmphibiaWeb

N N //y/fl kali Rodriguez, 1994 N/A N/A N/A N/A N/A V/ N/A Hyla lemai Duellman, 1997 Duellman, 1997 Duellman, 1997 N/A Duellman, 1997 N/A N/A Hyla leucophyllala Crump, 1974 Crump, 1974 Crump, 1974 Savage, 2002 N/A Savage, 2002 N/A Savage, 2002

Hyla loquax Duellman, 2001 N/A Duellman, 2001 N/A Savage, 2002 Savage, 2002 N/A N/A 1 I

N N N/A . N/A N/A N/A N/A N/A V/ N/A AmphibiaWeb

N N //y/a marianitae Duellman, 1997 N/A N/A N/A Duellman, 1997 V/ N/A N/A

N N V/ N Hyla marmorata Crump, 1974 Crump, 1974 Crump, 1974 N/A N/A V/ N/A N/A

//y/a martinsi Lutz, 1973 N/A N/A Lutz, 1973 N/A N/A

N N V/ N N Z/y/a melanomma Duellman,V/ 2001 N/A N/A N/A Duellman, 2001 Duellman, 2001 N/A N/A

Hyla meridionalis Summers, 2006 N/A N/A N/A N/A N/A

N N

Stebbins, 1959; V/

Hyla microcephala Stebbins, 1959 Stebbins, 1959 N/A Stebbins, 1959 Savage, 2002 N/A N N Duellman, 2001 V/

_ //y/a minuscula Duellman, 1997 N/A N/A N/A Duellman, 1997 N/A N/A J

N N Crump, 1974 Crump, 1974 Crump, 1974 N/A N/A Cei, 198V/ 0 N/A N/A

N N | Hyla miotympanum Duellman, 2001 Duellman, 2001 Duellman, 2001 N/A Duellman, 2001 V/ N/A N/A

N N | Hyla miyatai Bartlett, 2003 Summers, 2006 N/A N/A AmphibiaWeb V/ N/A AmphibiaWeb

N N V/ N N | Hyla nana Lutz, 1973 Cei, 1980 Cei, 198V/ 0 N/A N/A N/A N/A

N N | Hyla partialis Lutz, 1973 N/A Lutz, 1973 N/A V/ Lutz, 1973 N/A

N N | Hyla pelidna Duellman, 1989 N/A N/A - N/A AmphibiaWeb V/ N/A N/A

[ Hyla pentheter Duellman, 2001 N/A N/A N/A AmphibiaWeb N/A N/A -5

N N V/ N V/ N N V/ N Duellman, 1972 Summers, 2006 N/A N/A V/ Duellman, 1972 N/A N/A N/A

N N V/ N N | Hyla plicala Duellman,V/ 2001 N/A Duellman, 2001 N/A

N N V/ N N | //y/a polytaenia V/ N/A Lutz, 1973 N/A Lutz, 1973 N/A N/A

N N V/ N N | Hyla prasina N/AV/ N/A N/A Lutz, 1973 N/A N/A

1 Hyla pseudopseudis Lutz, 1973 N/A N/A N/A N/A Savage, 2002

N N [ Hyla pseudopuma Duellman, 2001 Savage, 2002 Savage, 200V/ 2 Crump, 1990 N/A . Savage, 2002 N/A Savage, 2002

N N | Hyla pulchella Lutz, 1973 Cei, 1980 N/A Lutz, 1973 Cei, 198V/ 0 N/A N/A | Hyla punctata Crump, 1974 Crump, 1974 Crump, 1974 N/A Cei, 1980 N/A N/A Cei, 1980; Cei, 1980; Hyla raniceps Lutz, 1973 Cei, 1980 N/A N/A Summers, 2006 Summers, 2006 Lutz, 1973 N/A OS Crump, 1974; Hyla rhodopepla Duellman, 1996; Crump, 1974 Crump, 1974 N/A N/A N/A N/A N/A Rodriguez, 1994 Hyla riveroi Duellman, 1996 N/A N/A N/A AmphibiaWeb N/A N/A AmphibiaWeb Hyla robertmertensi Duellman, 2001 N/A N/A N/A Duellman, 2001 N/A N/A N/A Hyla rosenbergi Duellman, 2001 Savage, 2002 Savage, 2002 Savage, 2002 N/A Savage, 2002 Savage, 2002 N/A Hyla rubicundula Lutz, 1973 N/A N/A N/A Lutz, 1973 N/A N/A N/A . Hyla rufltela Duellman, 2001 Savage, 2002 N/A N/A Savage, 2002 Savage, 2002 N/A Savage, 2002 Hyla sanborni Lutz, 1973 Cei, 1980 Cei, 1980 N/A Lutz, 1973 Cei, 1980 N/A N/A Hyla sarayacuensis Crump, 1974 Crump, 1974 Crump, 1974 N/A N/A Zug, 2001 N/A N/A Hyla sartori Duellman, 2001 N/A N/A N/A Duellman, 2001 Duellman, 2001 N/A N/A Hyla savignyi N/A Kuzmin, 1999 Kuzmin, 1999 N/A Brzoska, 1982 Kuzmin, 1999 N/A N/A Hyla semiguttata Lutz, 1973 N/A N/A N/A Lutz, 1973 N/A N/A N/A Hyla senicula Lutz, 1973 N/A N/A N/A Lutz, 1973 N/A N/A • N/A Hyla sibleszi Duellman, 1997 Duellman, 1997 Duellman, 1997 N/A Duellman, 1997 N/A N/A N/A Hyla simmonsi Duellman, 1989 N/A N/A N/A N/A N/A N/A AmphibiaWeb Hyla smithii Duellman, 2001 N/A N/A N/A Duellman, 2001 N/A N/A ' N/A Hyla squirella Wright, 1995 Wright, 1995 Wright, 1995 Fellers, 1979 N/A Wright, 1995 N/A N/A Hyla sumichrasti Duellman, 2001 Summers, 2006 N/A N/A Duellman, 2001 N/A N/A N/A Hyla laeniopus Duellman, 2001 Summers, 2006 N/A N/A N/A Duellman, 2001 N/A N/A Hyla thorectes Duellman, 2001 N/A Duellman, 2001 N/A Duellman, 2001 N/A N/A N/A Hyla tica Duellman, 2001 Savage, 2002 Savage, 2002 N/A Duellman, 2001 Savage, 2002 N/A N/A Crump, 1974; Hyla triangulum Duellman, Crump, 1974 Crump, 1974 N/A AmphibiaWeb N/A N/A AmphibiaWeb 1974b Hyla versicolor Wright, 1995 Summers, 2006 Smith, 1950 Fellers, 1979 N/A ' Wright, 1995 N/A Wright, 1995 Hyla walkeri Duellman, 2001 N/A N/A N/A Duellman, 2001 . N/A N/A N/A Hyla wrightorum Wright, 1995 Stebbins, 1951 N/A N/A Wright, 1995 Wright, 1995 N/A N/A Hyla zeteki Duellman, 2001 N/A Savage, 2002 N/A Savage, 2002 N/A N/A Savage, 2002 HyloscirXus palmeri Savage, 2002 N/A N/A N/A Savage, 2002 N/A N/A AmphibiaWeb Hypsiboas Hartmann, 2004 N/A N/A N/A Lutz, 1973 N/A N/A N/A albomarginatus Hypsiboas Crump, 1974 Crump, 1974 Crump, 1974 N/A AmphibiaWeb N/A N/A AmphibiaWeb cinerascens Isthmohyla rivularis Duellman, 2001 Summers, 2006 Savage, 2002 N/A Savage, 2002 Savage, 2002 N/A N/A | Litoria arfakiana Menzies, 1976 N/A N/A N/A Menzies, 1976 N/A N/A N/A .

120 Litoria armatus Duellman, 1997 N/A N/A N/A Duellman, 1997 N/A N/A N/A Litoria aurea Moore, 1961 Moore, 1961 N/A N/A Moore, 1961 Moore, 1961 N/A N/A Moore, 1961; Tyler, 1983 N/A Moore, 1961 Tyler, 1983 N/A Tyler, 1983 Litoria caerulea Moore, 1961 Tyler, 1983 Litoria freycineti Moore, 1961 Summers, 2006 N/A N/A Moore, 1961 AmphibiaWeb N/A N/A Animal Animal Litoria infrafrenata N/A N/A Menzies, 1976 AmphibiaWeb N/A AmphibiaWeb Diversity Web Diversity Web Tyler, 1983; N/A N/A N/A Tyler, 1983 Tyler, 1983 N/A Tyler, 1983 Litoria meiriana AmphibiaWeb Litoria peronii Tyler, 1978 Summers, 2006 j N/A Wells, 1977 N/A N/A N/A N/A Litoria rubella Tyler, 1978 Tyler, 1983 N/A N/A Hoser, 1989 Hoser, 1989 N/A Tyler, 1978 Nyctimantis Duellman, 2001 N/A N/A N/A Duellman, 1976 N/A AmphibiaWeb N/A rugiceps Nyctimystes Menzies, 1976 N/A N/A N/A Zweifel, 1980 N/A N/A. N/A cheesmanae Nyctimystes foricula Menzies, 1976 N/A _, N/A N/A Menzies, 1976 N/A N/A N/A Nyctimystes kubori Menzies, 1976 N/A N/A N/A Menzies, 1976 N/A N/A N/A Nyctimystes papua Menzies, 1976 N/A N/A N/A Menzies, 1976 N/A N/A • AmphibiaWeb Osteocephalus Crump, 1974; Trueb, 1971 Jungfer, 1999 N/A AmphibiaWeb N/A N/A N/A buckleyi Jungfer, 1999 Osteocephalus N/A N/A N/A Lutz, 1973 N/A N/A N/A langsdorffii Lutz, 1973 Osteocephalus Trueb, 1971 Crump, 1974 Crump, 1974 N/A AmphibiaWeb N/A N/A leprieurii N/A Osteocephalus Jungfer, 1999; N/A Jungfer, 1999 Jungfer, 1999 N/A Jungfer, 1999 oophagus Schiesari, 2003 Jungfer, 1999 AmphibiaWeb Osteocephalus Duellman, 1997; Crump, 1974; Crump, 1974 taurinus Trueb, 1971 Summers, 2006 N/A Duellman, 1997 N/A N/A N/A Osteocephalus Trueb, 1971 N/A N/A N/A AmphibiaWeb N/A N/A verruciger N/A Osteopilus brunneus Trueb, 1974 Jungfer, 1999 Schiesari, 2003 N/A N/A Schwartz, 1991 N/A AmphibiaWeb Osteopilus crucialis Trueb, 1974 N/A N/A N/A Schwartz, 1991 N/A Duellman, 1994 N/A Osteopilus Trueb, 1974 N/A N/A N/A Schwartz, 1991 N/A N/A N/A dominicensis Osteopilus marianae Trueb, 1974 N/A N/A N/A Schwartz, 1991 N/A N/A N/A

121 Osteopilus Trueb, 1974 N/A N/A N/A N/A Schwartz, 1991 N/A pulchrilineatus AmphibiaWeb Osteopilus Trueb, 1974; Savage, 2002 Savage, 2002 Salinas, 2006 N/A Savage, 2002 Salinas, 2006 N/A septentrionalis Wright, 1995 Osteopilus vastus Trueb, 1974 Schwartz, 1991 N/A N/A Schwartz, 1991 N/A N/A AmphibiaWeb Osteopilus wilderi Trueb, 1974 N/A N/A N/A N/A N/A N/A AmphibiaWeb Pachymedusa Bagnara, 1986; Bagnara, 1986; Bagnara, 1986; Bagnara, 1986; Duellman,2001 N/A Wiewandt, 1971 N/A dacnicolor Wiewandt, 1971 Duellman, 2001 Duellman, 2001 Duellman, 2001 Phrynohyas J. Bogart's Rodriguez, 1994 Crump, 1974 Crump, 1974 N/A N/A personal N/A coriacea N/A communication J. Bogart's Phrynohyas Lutz, 1973 Lutz, 1973 Lutz, 1973 N/A personal N/A mesophaea N/A Lutz, 1973 communication Phrynohyas Lutz, 1973 Schiesari, 2003 Schiesari, 2003 Schiesari, 2003 N/A Schiesari, 2003 Schiesari, 2003 N/A resinificlrix Phrynohyas Savage, 2002 venulosa Duellman, 2001 Rodrigues, 2005 Rodrigues, 2005 N/A Savage, 2002 N/A N/A Phyllodytes luteolus N/A Summers, 2006 Schiesari, 2003 N/A N/A N/A N/A AmphibiaWeb Phyllomedusa Rodriguez, 1994 AmphibiaWeb N/A atelopoides AmphibiaWeb AmphibiaWeb AmphibiaWeb AmphibiaWeb N/A Phyllomedusa bicolor Rodriguez, 1994 N/A N/A N/A AmphibiaWeb N/A N/A AmphibiaWeb Phyllomedusa Duellman, 1997 N/A N/A Abrunhosa, 2004 N/A N/A hypochondrialis N/A AmphibiaWeb Abrunhosa, Phyllomedusa lemur Duellman, 1956 Savage, 2002 Savage, 2002 2004; N/A Savage, 2002 N/A Savage, 2002 Savage, 2002 Phyllomedusa Crump, 1974 Crump, 1974 Crump, 1974 N/A palliata N/A N/A N/A AmphibiaWeb Phyllomedusa tarsia Crump, 1974 Crump, 1974 Crump, 1974 N/A AmphibiaWeb N/A N/A ' AmphibiaWeb Phyllomedusa Crump, 1974 Crump, 1974 Crump, 1974 N/A N/A N/A N/A AmphibiaWeb tomoplerna Phyllomedusa vaillantii Crump, 1974 Crump, 1974 Crump, 1974 N/A N/A Zug, 2001 N/A N/A

122 Plectrohyla Duellman, 2001 N/A N/A N/A AmphibiaWeb N/A N/A N/A glandulosa Plectrohyla Duellman, 1992 Summers, 2006 N/A N/A AmphibiaWeb N/A N/A N/A guatemalensis Plectrohyla matudai Duellman, 2001 N/A N/A N/A AmphibiaWeb N/A N/A AmphibiaWeb Hulse, 2001; Hulse, 2001; Pseudacris Hulse, 2001 Hulse, 2001 N/A Wright, 1995 N/A Wright, 1995 brachyphona Wright, 1995 Wright, 1995 Pseudacris brimleyi Wright, 1995 N/A AmphibiaWeb N/A • Wright, 1995 AmphibiaWeb N/A AmphibiaWeb Pseudacris Duellman, 2001 AmphibiaWeb N/A N/A Duellman, 2001 Stebbins, 2003 N/A AmphibiaWeb cadaverind Pseudacris clarkii Wright, 1995 Wright, 1995 Smith, 1950 N/A Duellman, 2001 Smith, 1950 N/A Wright, 1995 Hulse, 2001; Hulse, 2001; Wright, 1995 Hulse, 2001 N/A Wright, 1995 N/A N/A Pseudacris crucifer Wright, 1995 Wright, 1995 Pseudacris feriarum Wright, 1995 Wright, 1995 N/A N/A Wright, 1995 Wright, 1995 N/A N/A Pseudacris maculata Wright, 1995 Wright, 1995 N/A N/A Wright, 1995 Wright, 1995 N/A N/A Pseudacris nigrita Wright, 1995 Wright, 1995 Stebbins, 1951 N/A Dullman, 1994 Wells, 1977 N/A Stebbins, 1951 Pseudacris ocularis Wright, 1995 Wright, 1995 AmphibiaWeb N/A Wright, 1995 Wright, 1995 N/A N/A Pseudacris ornata Wright, 1995 Wright, 1995 AmphibiaWeb N/A Wright, 1995 AmphibiaWeb N/A N/A Stebbins, 2003; Pseudacris regilla Duellman, 2001 Stebbins, 1951 Wright, 1995 Wells, 1977 N/A N/A Wright, 1995 Wright, 1995 Pseudacris streckeri Wright, 1995 Wright, 1995 Wright, 1995 N/A Wright, 1995 Wright, 1995 N/A N/A Pseudacris Nussbaum, Wright, 1995 Morrison, 2003 triseriata Morrison, 2003 N/A Nussbaum, 1983 1983; N/A Wright, 1995 Wright, 1995 Pseudis paradoxa N/A N/A N/A N/A Cei, 1980 N/A N/A Cei, 1980 Pternohyla fodiens Duellman, 2001 N/A N/A N/A Duellman, 2001 Duellman, 2001 N/A N/A Ptychohyla Duellman, 2001 N/A N/A N/A N/A Duellman, 2001 euthysanota N/A N/A Ptychohyla N/A Summers, 2006 hypomykter N/A N/A N/A N/A N/A N/A Ptychohyla Duellman, 2001 N/A N/A N/A Duellman, 2001 N/A N/A leonhardschultzei Duellman, 2001 Ptychohyla Duellman, 2001 N/A spinipollex N/A . N/A Duellman, 2001 N/A N/A N/A Scarthyla goinorum Rodriguez, 1994 N/A N/A N/A N/A N/A N/A AmphibiaWeb Scinax berthae Lutz, 1973 Cei, 1980 N/A N/A Cei, 1980 Cei, 1980 N/A N/A

123 Duellman, 2001; Scinax boulengeri Duellman, 2001 Summers, 2006 Savage, 2002 N/A Savage, 2002 N/A Savage, 2002 Savage, 2002 Scinax catharinae Lutz, 1973 N/A N/A N/A Lutz, 1973 N/A N/A AmphibiaWeb Scinax elaeochraoa Duellman, 2001 N/A N/A Savage, 2002 N/A Zug, 2001 N/A Savage, 2002 Lutz, 1973; Delariva, 1993; Scinax fuscovarius Rodrigues, 2005 Rodrigues, 2005 N/A 1 N/A Rodrigues, 2005 Lutz, 1973 Rodrigues, 2005 N/A Scinax garbei Crump, 1974 Crump, 1974 Crump, 1974 N/A AmphibiaWeb N/A N/A N/A Scinax nasicus Lutz, 1973 N/A N/A N/A Delariva, 1993 N/A N/A N/A Crump, 1974; Scinax ruber Duellman, 1996; Crump, 1974 Crump, 1974 N/A Duellman, 2001 Zug, 2001 N/A N/A Duellman, 2001 Scinax squalirostris N/A Cei, 1980 N/A N/A Lutz, 1973 N/A N/A Cei, 1980 Duellman, 2001; Duellman, 2001; Scinax staufferi Duellman, 2001 N/A N/A N/A N/A Savage, 2002 Savage, 2002 N/A Duellman, 2001; Smilisca baudinii Duellman, 2001 N/A Duellman, 2001 N/A Savage, 2002 Duellman, 2001 Savage, 2002 Zug, 2001 Smilisca cyanosticta Duellman, 2001 Duellman, 2001 Duellman, 2001 N/A Duellman, 2001 Duellman, 2001 N/A Duellman, 2001 Smilisca phaeota Savage, 2002 . N/A Savage, 2002 N/A Savage, 2002 Savage, 2002 N/A N/A Smilisca puma Savage, 2002 N/A N/A N/A Savage, 2002 Savage, 2002 N/A N/A Sphaenorhynchus Rodriguez, 1994 Summers, 2006 N/A N/A AmphibiaWeb N/A N/A N/A lacleus Tlalocohyla picta Duellman, 2001 N/A N/A N/A Duellman, 2001 N/A N/A N/A Triprion petasatus Duellman, 2001 N/A N/A N/A Duellman, 2001 Duellman, 2001 N/A N/A Xenohyla Iruncala Lutz, 1973 N/A N/A N/A Lutz, 1973 N/A N/A N/A Afrixalus fornasini Schiotz, 1975 Wager, 1965 Wager, 1965 N/A ' Wager, 1965 N/A N/A N/A Alexteroon N/A N/A N/A N/A AmphibiaWeb N/A Duellman, 1994 N/A obstetricans Hyperolius Clanning, 2001 N/A N/A N/A Wager, 1965 N/A N/A N/A puncticulatus Hyperolius N/A Wager, 1965 Wager, 1965 N/A Wager, 1965 N/A N/A N/A tuberilinguis Kassina AmphibiaWeb Wager, 1965 Wager, 1965 N/A N/A N/A N/A N/A senegalensis Ascaphus truei Wright, 1995 Nussbaum, 1983 Stebbins, 1951 N/A Stebbins, 2003 AmphibiaWeb N/A N/A Leiopelma archeyi AmphibiaWeb Summers, 2006 N/A N/A N/A N/A Duellman, 1994 N/A

124 Leiopelma hochstetteri AmphibiaWeb . Summers, 2006 N/A N/A AmphibiaWeb N/A. Duellman, 1994 N/A Edalorhina perezi Duellman, 1996 Crump, 1974 Crump, 1974 N/A N/A . N/A AmphibiaWeb N/A Pleurodema Rivero, 1961 N/A N/A N/A Duellman, 1977 N/A brachyops N/A AmphibiaWeb Pseudopaludicola N/A falcipes Cei, 1980 N/A N/A Cei, 1980 Cei, 1980 N/A Cei, 1980 Leplodactylus fuscus Heyer, 1978 Summers, 2006 N/A N/A Martins, 1988 N/A Martins, 1998 N/A Leptodactylus Heyer, 1978 N/A N/A N/A gracilis Cei, 1980 N/A AmphibiaWeb N/A Leptodactylus Heyer, 1973 N/A N/A N/A Heyer, 1973 N/A N/A hylaedactylus N/A Leptodactylus ocellatus Rivero, 1961 Stebbins, 1951 Stebbins, 1951 N/A Stebbins, 1951 Stebbins, 1951 AmphibiaWeb N/A Lithodytes lineatus Rodriguez, 1994 Crump, 1974 Crump, 1974 N/A N/A N/A AmphibiaWeb N/A Vanzolinius Crump, 1974 Crump, 1974 Crump, 1974 discodactylus N/A N/A N/A N/A AmphibiaWeb Adelotus brevis Katsikaros, 1997 Summers, 2006 N/A Katsikaros, 1997 - N/A AmphibiaWeb Duellman, 1994 N/A Lechriodus fletcheri Parker, 1940 Moore, 1961 AmphibiaWeb N/A Moore, 1961 Hoser, 1989 AmphibiaWeb N/A Limnodynastes Hoser, 1989; Parker, 1940 Summers, 2006 AmphibiaWeb Duellman, 1994 N/A Hoser, 1989 N/A dumerilii Tyler, 1978 Limnodynastes Parker, 1940 Tyler, 1983 Tyler, 1983 N/A Hoser, 1989 Hoser, 1989 Tyler, 1983 N/A ornatus Limnodynastes Schauble, 2004 Summers, 2006 AmphibiaWeb - Wells, 1977 N/A Hoser, 1989 AmphibiaWeb N/A peronii Frogs of the Limnodynastes Parker, 1940 N/A AmphibiaWeb N/A Greater Brisbane AmphibiaWeb N/A N/A salmini Region Neobatrachus pictus N/A N/A AmphibiaWeb N/A Moore, 1961 AmphibiaWeb N/A N/A Notaden N/A Tyler, 1983 Tyler, 1983 N/A Tyler, 1983 Tyler, 1983 N/A Tyler, 1983 melanoscaphus Philoria Knowles, 2004 N/A Moore, 1961 N/A Moore, 1961 N/A AmphibiaWeb N/A sphagnicolus Aglyptodactylus N/A AmphibiaWeb AmphibiaWeb N/A AmphibiaWeb N/A N/A N/A madagascariensis

125 Boophis Blommers- Blommers- Summers, 2006 N/A N/A N/A N/A N/A tephraeomystax Schlosser, 1979 Schlosser, 1979 Mantella aurantiaca AmphibiaWeb AmphibiaWeb AmphibiaWeb N/A AmphibiaWeb N/A Glaw, 2000 N/A Mantidactylus N/A AmphibiaWeb N/A N/A AmphibiaWeb AmphibiaWeb N/A AmphibiaWeb femoralis Leptobrachium N/A N/A N/A Lathrop, 1998 N/A N/A N/A banae Lathrop, 1998 Leptobrachium Inger, 1966 N/A N/A N/A Dring, 1979 N/A N/A N/A hassettii Leptobrachium Ohler, 2004 N/A N/A N/A N/A . N/A N/A N/A montanum Leptobrachium N/A N/A N/A Fei, 1999 Stuart, 2006 N/A N/A mouhoti Stuart, 2006 Leptolalax liui Fei, 1999 N/A N/A N/A Fei, 1999 N/A N/A N/A Leptolalax Huang, 1990 Pope, 1931 Pope, 1931 N/A N/A Huang, 1990 N/A N/A pelodytoides Oreolalax N/A Fei, 1999 Fei, 1999 N/A Fei, 1999 Fei, 1999 N/A N/A jingdongensis Oreolalax N/A liangbeiensis Ye, 1993 Ye, 1993 Fei, 1999 N/A Fei, 1999 N/A Ye, 1993 Oreolalax N/A Fei, 1999 N/A N/A Fei, 1999 Fei, 1999 N/A multipunctatus Fei, 1999 Oreolalax N/A N/A N/A N/A . N/A N/A omeimontis Fei, 1999 Fei, 1999 Fei, 1999; Oreolalax pingii Ye, 1993 Ye, 1993 N/A N/A Ye, 1993 N/A N/A Ye; 1993 Oreolalax popei Ye, 1993 Ye, 1993 Ye, 1993 N/A Fei, 1999 Ye, 1993 . N/A N/A Oreolalax rugosus Ye, 1993 Ye, 1993 N/A N/A N/A Ye, 1993 N/A N/A Oreolalax schmidli Fei, 1999 Fei, 1999 Liu, 1950 N/A Fei, 1999 Fei, 1999 N/A N/A Hu, 1987; Scutiger boulengeri Ye, 1993 Ye, 1993 N/A Fei, 1999 Ye, 1993 N/A N/A Ye, 1993 Scutiger glandulalus Yang, 1991 Fei, 1999 N/A N/A Fei, 1999 Liu, 1950 N/A Liu, 1950 Scutiger mammalus Ye, 1993 Ye, 1993 Ye, 1993 N/A Fei, 1999 Hu, 1987 N/A N/A Scutiger muliensis Fei, 1999 N/A Fei, 1999 N/A N/A Fei, 1999 N/A N/A

126 Scutiger Ye, 1993 Ye, 1993 N/A N/A N/A Ye, 1993 N/A Ye, 1993 tuberculatus Vibrissaphora Ye, 1993 Fei, 1999 N/A N/A ailaonica Ye, 1993 Fei, 1999 Ho, 1999 Ye, 1993 Vibrissaphora Ye, 1993 Ye, 1993 Ye, 1993 N/A Fei, 1999 Ye, 1993 Ye, 1993 boringiae N/A Vibrissaphora Lathrop, 1998 N/A N/A N/A Fei, 1999 Fei, 1999 N/A N/A chapaense Vibrissaphora Ye, 1993 Fei, 1999 Ye, 1993 N/A Ye, 1993 Ye, 1993 N/A Ye, 1993 leishanensis Vibrissaphora liui Ye, 1993 Ye, 1993 Ye, 1993 N/A Fei, 1999 Huang, 1990 Huang, 1990 N/A Vibrissaphora Rao, 2006 N/A N/A N/A Rao, 2006 N/A N/A N/A promustache Chaperina fusca Inger, 1966 N/A N/A N/A N/A Inger, 1966 N/A N/A Degenhardt, Gastrophryne Degenhardt, N/A Wright, 1995 1996; N/A Wells, 1977 N/A AmphibiaWeb olivacea 1996 Smith, 1950 Kalophrynus Inger, 1966 Pope, 1931 Pope, 1931 N/A N/A N/A N/A Inger, 1966 pleurostigma Kaloula pulchra Inger, 1966 N/A N/A N/A N/A N/A N/A N/A Chen, 1991; Chen, 1991; Yang, 1991 Huang, 1990 N/A Dring, 1979 N/A N/A Microhyla heymonsi Huang, 1990 Ye, 1993 Phrynomantis N/A Wager, 1965 Wager, 1965 N/A Wager, 1965 N/A Beck, 1998 N/A bifasciatus Crinia nimbus N/A Mitchell, 2002 AmphibiaWeb N/A AmphibiaWeb N/A Mitchell, 2002 N/A Crinia signifera Parker, 1940 Moore, 1961 Moore, 1961 N/A Hoser, 1989 Hoser, 1989 N/A AmphibiaWeb Geocrinia victoriana N/A Morrison, 2003 AmphibiaWeb N/A Parker, 1940 N/A N/A AmphibiaWeb Metacrinia nichollsi Parker, 1940 AmphibiaWeb N/A N/A N/A N/A N/A N/A Myobatrachus Main, 1965; Parker, 1940 Summers, 2006 AmphibiaWeb N/A N/A N/A AmphibiaWeb gouldii AmphibiaWeb Paracrinia haswellsi Parker, 1940 N/A N/A N/A Moore, 1961 N/A • N/A AmphibiaWeb Moore, 1961; Pseudophryne Parker, 1940 Moore, 1961 Parker, 1940; Duellman, 1994 N/A Hoser, 1989 Duellman, 1994 N/A bibroni Tyler, 1978

127 Rheobatrachus silus N/A Summers, 2006 AmphibiaWeb N/A AmphibiaWeb AmphibiaWeb Deullman, 1994 N/A Spicospina N/A N/A AmphibiaWeb N/A Smith, 2003 AmphibiaWeb N/A AmphibiaWeb flammocaerulea Uperoleia laevigata N/A N/A N/A N/A Hoser, 1989 AmphibiaWeb N/A AmphibiaWeb Pelobates fuscus N/A Kuzmin, 1999 Kuzmin, 1999 Kuzmin, 1999 N/A Wells, 1977 N/A AmphibiaWeb Phrynobatrachus Stewart, 1967; Stewart, 1967 Wager, 1965 N/A N/A N/A N/A N/A mababiensis Wager, 1965 Phrynobatrachus Stewart, 1967 Wager, 1965 Wager, 1965 N/A Wager, 1965 Stewart, 1967 N/A AmphibiaWeb natalensis Hymenochirus boettgeri Rabb, 1963 Rabb, 1962 Rabb, 1962 Rabb, 1962 N/A Rabb, 1962 N/A Rabb, 1962 Pipa carvalhoi Trueb, 1986 Weygoldt, 1976 Weygoldt, 1976 Duellman, 1994 N/A Weygoldt, 1976 Duellman, 1994 N/A Pipa pipa Trueb, 1986 Summers, 2006 N/A Rabb, 1963 N/A Honolulu Zoo Duellman, 1994 N/A Xenopus laevis . N/A Wager, 1965 AmphibiaWeb Rabb, 1963 N/A Stebbins, 2003 N/A Wager, 1965 Ptychadena N/A N/A N/A N/A Wager, 1965 N/A N/A N/A anchietae Ptychadena Stewart, 1967 mascareniensis Channing, 2001 Summers, 2006 N/A N/A Wager, 1965 N/A N/A Afrana angolensis N/A Wager, 1965 N/A N/A N/A Wager, 1965 N/A N/A Afrana fuscigula Stewart, 1967 Wager, 1965 Wager, 1965 N/A Wager, 1965 Stewart, 1967 N/A N/A Anhydrophryne N/A AmphibiaWeb rattrayi N/A N/A Wager, 1965 N/A N/A AmphibiaWeb Aphantophryne N/A Zweifel, 1956 N/A N/A N/A pansa N/A N/A N/A Natalobatrachus N/A bonebergi Wager, 1965 Wager, 1965 N/A Wager, 1965 N/A N/A AmphibiaWeb Strongylopus grayii Channing, 2001 - - N/A Wager, 1965 N/A N/A N/A Tomopterna N/A delalandii Summers, 2006 - N/A N/A AmphibiaWeb N/A N/A N/A Amolops chapaensis Yang, 1991 N/A N/A N/A AmphibiaWeb N/A N/A N/A Amolops Ye, 1993 Ye, 1993 Inger, 1968 Liu, 1950 N/A Ye, 1993 N/A Liu, 1950 chunganensis Amolops Inger, 1998 Inger, 1998 N/A N/A N/A N/A N/A N/A cremnobatus Liu, 1950; Yang, 1991 N/A N/A N/A N/A Amolops mantzorum Ye, 1993 AmphibiaWeb Ye, 1993

128 Amolops ricketti Pope, 1931 Ye, 1993 N/A N/A N/A Ye, 1993 N/A N/A Amolops Yang, 1991 N/A N/A N/A AmphibiaWeb N/A N/A N/A viridimaculatus Amolops wuyiensis Chen, 1991 Huang, 1990 Huang, 1990 . N/A AmphibiaWeb Huang, 1990 N/A N/A Fejervarya Inger, 1966 N/A N/A N/A AmphibiaWeb Inger, 1966 N/A N/A nicobariensis Hydrophylax Stewart, 1967 AmphibiaWeb AmphibiaWeb N/A AmphibiaWeb AmphibiaWeb N/A AmphibiaWeb galamensis Lankanectes N/A N/A N/A N/A Alcala, 1986 N/A N/A Alcala, 1986 corrugatus Meristogenys jerboa Inger, 1966 Inger, 1968 Inger, 1968 N/A AmphibiaWeb Inger, 1966 N/A N/A Meristogenys Inger, 1966 Inger, 1966 N/A AmphibiaWeb N/A N/A kinabaluensis N/A N/A Chen, 1991; Liu, 1950; Huang, 1990; Pope, 1931; Rana adenopleura Yang, 1991 Liu, 1950; N/A Xu, 2005 Liu, 1950 Liu, 1950 N/A Yang, 1991; Yang, 1991; Ye, 1993 Ye, 1993 Rana amurensis Pope, 1931 Kuzmin, 1999 Kuzmin, 1999 N/A AmphibiaWeb Kuzmin, 1999 N/A N/A Rana andersonii Pope, 1931 N/A N/A N/A . N/A Liu, 1950 N/A N/A Smith, 1950; Rana areolata Wright, 1995 Wright, 1995 Wright, 1995 Wells, 1977 N/A Wright, 1995 N/A N/A Inger, 1968; Rana arvalis N/A Kuzmin, 1999 Wells, 1977 N/A Kuzmin, 1999 Kuzmin, 1999 N/A N/A Rana asiatica Pope, 1931 Kuzmin, 1999 Kuzmin, 1999 N/A N/A Kuzmin, 1999 N/A N/A Rana aurora Wright, 1995 Wright, 1995 Nussbaum, 1983 Wells, 1977 N/A Stebbins, 1951 N/A N/A Degenhardt, Rana berlandieri N/A- N/A AmphibiaWeb N/A N/A 1996 N/A AmphibiaWeb Rana blairi N/A N/A AmphibiaWeb N/A Frost, 1977 Stebbins, 2003 N/A AmphibiaWeb Nussbaum, 1983; Rana boylii Nussbaum, 1983 Nussbaum, 1983 N/A Wright, 1995 Stebbins, 1951; N/A N/A N/A . Wright, 1995 Rana capito Wright, 1995 Wright, 1995 Goin, 1940 N/A N/A Wright, 1995 N/A N/A

129 Nussbaum, Nussbaum, Rana cascadae Wright, 1995 1983; Nussbaum, 1983 N/A N/A 1983; N/A N/A Stebbins, 1951 Wright, 1995 Degenhardt, 1996; Wright, 1995; Degenhardt, Rana catesbeiana Hulse,2001 Hulse, 2001; Wells, 1977 N/A N/A Ye, 1993 Ye, 1993 1996 Smith, 1950; Ye, 1993 Rana chalconola Bain, 2003 N/A . N/A N/A Marquez, 2006 N/A N/A N/A Rana Yang, 1991 N/A N/A' N/A N/A chaochiaoensis Liu, 1950 N/A AmphibiaWeb Inger, 1968; Rana chensinensis Okada, 1966 N/A N/A N/A Ye, 1993 N/A Ye, 1993 Ye, 1993 Degenhardt, N/A N/A AmphibiaWeb N/A N/A N/A AmphibiaWeb Rana chiricahuensis 1996 Nussbaum, Oseen, 2002; Rana clamitans Wright, 1995 Wright, 1995 1983; Wells, 1977 N/A N/A Stebbins, 1951 Wright, 1995 Wright, 1995 Rana daemeli Wells, 1977 N/A AmphibiaWeb N/A N/A N/A N/A AmphibiaWeb Rana dalmatina Lode, 2005 Kuzmin, 1999 Kuzmin, 1999 Lode, 2005 N/A Lode, 2005 N/A N/A Rana dybowskii N/A Kuzmin, 1999 Kuzmin, 1999 N/A N/A AmphibiaWeb N/A AmphibiaWeb Rana emeljanovi Ye, 1993 Ye, 1993 Ye, 1993 N/A . N/A Ye, 1993 N/A N/A Rana erythraea Inger, 1966 N/A N/A N/A N/A Inger, 1966 N/A N/A Rana forreri Savage, 2002 N/A N/A N/A Savage, 2002 Savage, 2002 N/A N/A Rana graeca N/A AmphibiaWeb AmphibiaWeb N/A AmphibiaWeb AmphibiaWeb AmphibiaWeb N/A Rana grahami Yang, 1991 Ye, 1993 Yang, 1991 N/A N/A Ye, 1993 N/A Ye, 1993 Rana grylio Wright, 1995 N/A AmphibiaWeb N/A Schwartz, 1991 AmphibiaWeb N/A AmphibiaWeb Huang, 1990; Huang, 1990; Huang, 1990; Rana guentheri Pope, 1931 N/A Xu, 2005 N/A N/A Pope, 1931 Pope, 1931 Ye, 1993 Rana heckscheri Wright, 1995 AmphibiaWeb AmphibiaWeb N/A AmphibiaWeb Wright, 1995 N/A AmphibiaWeb Huang, 1990; Ranajaponica Pope, 1931 Chen, 1991 Wells, 1977 N/A Huang, 1990 N/A N/A Okada, 1966 Rana latastei N/A N/A N/A Hettyey, 2003 AmphibiaWeb AmphibiaWeb N/A N/A Beebee, 2000; Rana lessonae N/A Kuzmin, 1999 Beebee, 2000 N/A Kuzmin, 1999 N/A N/A Kuzmin, 1999

130 Chen, 1991; Chen, 1991; Rana livida Chen, 1991 Huang, 1990; N/A N/A Yang, 1991 N/A N/A Huang, 1990 Yang, 1991 Rana luteiventris N/A Wright, 1995 Wright, 1995 N/A N/A Stebbins, 2003 N/A N/A Rana macrocnemis N/A N/A Kuzmin, 1999 N/A N/A Kuzmin, 1999 N/A N/A Rana margaretae Bain, 2003 N/A N/A N/A AmphibiaWeb N/A N/A N/A Rana minima Ye, 1993 N/A . Ye, 1993 N/A N/A Ye, 1993 N/A Ye, 1993 Stebbins, 1951; Rana muscosa Wright, 1995 Wright, 1995 N/A N/A Wright, 1995 N/A N/A Wright, 1995 Chen, 1991; Chen, 1991; Kuzmin, 1999; Rana nigromaculata Chen, 1991 Kuzmin, 1999; Huang, 1990; N/A Kuramoto, 1977 N/A N/A Ye, 1993 Liu, 1950 Ye, 1993 Rana nigrovillata Yang, 1991 Yang, 1991 Yang, 1991 N/A N/A N/A N/A N/A Rana okinavana Inger, 1947 N/A AmphibiaWeb N/A AmphibiaWeb AmphibiaWeb N/A ' N/A Rana omeimontis N/A Ye, 1993 N/A N/A N/A Ye, 1993 N/A N/A Rana onca Wright, 1995 N/A AmphibiaWeb N/A AmphibiaWeb Wright, 1995 N/A AmphibiaWeb Rana ornativenlris Okada, 1966 Okada, 1966 Okada, 1966 N/A AmphibiaWeb N/A N/A N/A Rana palmipes Crump, 1974 Crump, 1974 Crump, 1974 N/A N/A Zug, 2001 N/A N/A Rana palustris Wright, 1995 Wright, 1995 Wright, 1995 Wells, 1977 N/A Hulse, 2001 N/A Smith, 1950 Degenhardt, Rana pipiens Hu!se,2001 Stebbins, 1951 Wells, 1977 N/A Wright, 1995 N/A N/A 1996 Rana pleuraden Yang, 1991 Ye, 1993 Yang, 1991 N/A N/A Yang, 1991 N/A N/A Chen, 1991; Rana schmackeri Ye, 1993 Chen, 1991 N/A N/A Chen, 1991 N/A N/A Huang, 1990 Rana septentrionalis Wright, 1995 Hulse, 2001 Hulse, 2001 N/A N/A Wright, 1995 N/A N/A Rana sevosa Wright, 1995 AmphibiaWeb AmphibiaWeb N/A AmphibiaWeb N/A N/A N/A Rana shuchinae Yang, 1991 N/A Yang, 1991 N/A N/A Yang, 1991 N/A Rana signata Inger, 1966 Alcala, 1986 N/A N/A Dring, 1979 N/A N/A , AmphibiaWeb Rana sphenocephala Hulse,2001 Wright, 1995 Hulse, 2001 N/A Schwartz, 1991 Wright, 1995 N/A N/A Morrison, 2003; Rana sylvatica Hulse, 2001 Morrison, 2003 Wells, 1977 N/A Hulse, 2001 N/A Hulse, 2001 Stebbins, 1951 Yang, 1991; Rana taipehensis Yang, 1991 N/A N/A N/A Ye, 1993 Ye, 1993 N/A N/A Rana tarahumarae N/A Stebbins, 1951 Stebbins, 1951 N/A N/A Stebbins, 2003 . N/A N/A

131 Rana temporalis N/A Summers, 2006 N/A N/A AmphibiaWeb N/A N/A N/A Beebee, 2000; Rana temporaria Yang, 1991 Morrison, 2003 Morrison, 2003 Wells, 1977 N/A N/A N/A Pough, 2001 Rana tientaiensis Chen, 1991 Huang, 1990 Chen, 1991 N/A N/A Huang, 1990 N/A Huang, 1990 Rana tsushimensis Okada, 1966 AmphibiaWeb AmphibiaWeb N/A AmphibiaWeb AmphibiaWeb N/A AmphibiaWeb Rana vaillanti N/A N/A N/A N/A Savage, 2002 Savage, 2002 N/A Savage, 2002 Rana versabilis Chen, 1991 Chen, 1991 Chen,1991 N/A AmphibiaWeb N/A N/A N/A Rana vibicaria Savage, 2002 Savage, 2002 N/A N/A Savage, 2002 Savage, 2002 N/A N/A Given, 1988; Rana virgatipes Wright, 1995 Wright, 1995 Wright, 1995 Given, 1988 N/A N/A N/A Wright, 1995 Rana warszewitschii Savage, 2002 N/A N/A N/A Savage, 2002 N/A N/A N/A Degenhardt, Rana yavapaiensis N/A N/A N/A N/A AmphibiaWeb N/A AmphibiaWeb 1996 Staurois Inger, 1966 N/A N/A N/A N/A latopalmatus N/A N/A N/A Staurois natator Inger, 1966 Inger, 1966 N/A N/A AmphibiaWeb Inger, 1966 N/A AmphibiaWeb Buergeria japonica Okada, 1966 Fei, 1999 N/A N/A N/A Fei, 1999 N/A Fei, 1999 Chirixdlus doriae Pope, 1931 Fei, 1999 N/A N/A N/A Fei, 1999 N/A Fei, 1999 Hu, 1987; Chirixalus vittatus Yang, 1991 Hu, 1987 N/A • N/A Hu, 1987 N/A Fei, 1999 Ye, 1993 Chiromantis Wager, 1965; Wager, 1965; Beck, 1998; Schiotz, 1999 Pough, 2001 N/A N/A N/A xerampelina Yang, 1991 Yang, 1991 Duellman, 1994 Philautus gracilipes Yang, 1991 N/A N/A N/A N/A N/A N/A AmphibiaWeb Polypedates Inger, 1966; Inger, 1966 Yang, 1991 Yang, 1991 N/A Marquez, 2006 N/A Yang, 1991 leucomystax Yang, 1991 Rhacophorus N/A N/A N/A N/A Dring, 1979 N/A N/A AmphibiaWeb bipunctatus Rhinophiynus N/A Savage, 2002 Savage, 2002 N/A N/A Savage, 2002 N/A Savage, 2002 dorsalis Degenhardt, Degenhardt, Scaphiopus couchii Wright, 1995 Wright, 1995 N/A Wright, 1995 N/A Wright, 1995 1996 1996 Scaphiopus Wright, 1995 Hulse, 2001 N/A N/A Wright, 1995 Hulse, 2001 N/A Wright, 1995 holbrookii Spea hammondii Wright, 1995 Stebbin, 1951 Stebbin, 1951 j N/A Stebbin, 1951 Wells, 1977 N/A Wright, 1995 Sooglossus N/A AmphibiaWeb N/A sechellensis AmphibiaWeb N/A N/A N/A N/A

132 APPENDIX 3 A manually constructed phylogenetic supertree for 545 anuran species across 37 families. Species names applied as in "Amphibian Species of the World'' website (December 2006). All branch lengths were set to one unit.

Ascaphus truei Leiopelma archeyi Leiopelma hochstetteri Rhinophrynus dorsalis Hymenochirus boettgeri Pipa carvalhoi Pipa pipa Xenopus laevis Atytes obstetricans Discoglossus pictus Bombina microdeladigitora Bombina orientals Bombina variegata Bombina bombina Spea hammondii Scaphiopus couchii Scaphiopus holbrooKii Pelobates fuscus Megrophryidae Soogiossus sechellensis Philoria sphagnicolus Ade lotus brevis Neobatrachus pictus Notaden melanoscaphus Limnodynastes ornatus Lechriodus fletcheri Limnodynastes dumehlii Limnodynastes peronii Limnodynastes salmini Rheobatrachus silus Cnnia signifera Crinia nimbus Paracrinia haswelli Geocrinia victoriana Uperoleia laevigata Spicospina flammocaerulea Pseudophryne bibroni Myobatrachus gouidii Metacrinia nichollsi Hemiphractus heliot Ischnocnema quixensis Eleutherodactylus binotalus Eleutherodactylus ptanirosths Eleutherodactylus bufoniformis Eleutherodactylus augusti Stefania evansi Hylidae Allophryne ruthveni Centrolenella prosoblepon Hyalinobatrachium fleischmanni Pleurodema brachyops Leptodactylus gracilis Edalorhina perezi Pseudopaludicota faidpes Leptodactylus hylaedactylus Lithodytes lineatus Leptodactylus fuscus Leptodactylus oceilatus Vanzolinius discodactylus Rhinoderma darwinii Dendrotatoidea Bufonidae Other Anura

Figure 1 Partial supertree of anurans, showing the relationships among 57 species and four major clades: the superfamily Dendrobatoidea (family Dendrobatidae and family

Aromobatidae), and the families Megophryidae, Hylidae and Bufonidae.

133 Kalophrynus pleurostigma Phrynomantis bifasciatus Gastrophryne olivacea Kaloula pulchra Microhyla heymonsi Chaperina fusca Aphantophryne pansa Hemisus marmoratum Breviceps mossambicus r~ ^Kassin a senegalensis Afrixalus fornasini Alexteroon obstetricans Hyperolius tuberilinguis -^ Hyperolius puncticulatus Trichobatrachus robustus i— Ptychadena mascareniensis i Ptychadena anchietae Ingerana baluensis Phrynobatrachus mababiensis "- 1 cz Phrynobatrachus natalensis Tomopterna delalandii Natalobatrachus bonebergi Afrana angolensis Afrana fuscigula Strongylopus grayii Anhydrophryne rattrayi Dicroglossidae Boophis tephraeomystax Aglyptodactylus madagascariensis Mantella aurantiaca —^ Mantidactylus femoralis Buergeria japonica _j CZ Philautus gracilipes Rhacophorus bipunctatus Polypedates leucomystax Chirixalus vittatus L Chirixalus doriae ^ Chiromantis xerampelina Ranidae

Figure 2 Partial supertree of anurans, showing the relationships among 37 species and two major clades: the families Dicroglossidae and Ranidae

134 Leptolalax liui HZ Leptolalax pelodytoides Vibrissaphora liui Vibrissaphora leishanensis Vibrissaphora boringiae r^ Vibrissaphora ailaonica Vibrissaphora chapaense Vibrissaphora promustache Leptobrachium banae Leptobrachium mouhoti Leptobrachium montanum Leptobrachium hasseltii Oreoialax pingli Oreolalax schmidti -c Oreoialax rugosus Oreolalax liangbeiensis r-cEOreoiala x jingdongensis Oreolalax omeimontis Oreolaiax popei Oreolalax multipunctatus Scutiger boulengeri -tz Scutiger glandulatus Scutiger muiiensis Scutiger tubercuiatus LcEScutige r mammatus

Figure 3 Supertree of the family Megophryidae, showing the relationships among 25 species.

Litoria peronii Litoria freycineti Litoria arfakiana Litoria rubella Litoria meiriana Litoria caerulea Litoria aurea Cyclorana brevipes Cyclorana australis Litoria infrafrenata Nyctimystes cheesmanae Nyctimystes foricula Nyctimystes papua Nyctimystes kubori Phyllomedusa tarsia Phyllomedusa vaillantii Phyllomedusa bicolor Phyllomedusa tomopterna Phyllomedusa atelopoides Phyllomedusa hypochondrialis Phyllomedusa palliata Cruzlohyla calcarifer Phyllomedusa lemur Pachymedusa dacnicolor Agalychnis callidryas Agalychnis saltator Agalychnis spurrelli Agalychnis litodryas Hylinae

Figure 4 Basal portion of the phylogeny of the family Hylidae, showing relationships among 28 species (Redrawn after Wiens et al, 2006).

135 Scinax catharinae Scinax berthae Scinax garbei Scinax boulengeri Scinax squalirostris Scinax elaeochraoa Scinax staufferi Scinax fuscovarius j Scinax ruber Scinax nasicus Phyllodytes lutaolus Osteocephalus langsdorffii Nyctimantis rugiceps Aparasphenodon brunoi Phrynehyas coriacea Phrynehyas mesophaea Phrynohyas venulosa Phrynehyas resinHictrix Osteocephalus buckleyi Osteocephalus verruciger Cyclorana alboguttata Osteocephalus taunnus Osteocephalus oophagus Osteocephalus leprieurii Osteopilus dominicensis Osteopilus crucialis Osteopilus marianae Osteopilus septentrionalis Osteopilus pulchrilineatus Osteopilus brunneus Osteopilus vastus Osteopilus wilderi Sphaenorhynchus lacteus Scarthyla goinorum Pseudis paradoxa Xenohyla truncata Hyla marmorata Hyla senicula

Dendropsophus aperomeus Hyia ebraccaia Hyla bifurca Hyla sarayacuensis Hyla leucophyllata Hyla triangulum Hyla pelidna Hyla labialis Dendropsophus parviceps Hyla koechlini Hyla brevifrons Hyla minuta Hyla riveroi Hyla allenorum Hyla aiegans Hyla miyatai Hyla rhodopepla Hyla microeephala Hyla leali Hyla rcbertmertensi Hyla sartori Hyla anceps Hyla bipunctata Hyla nana Hyla minuscula Hyla rubicundula Hyla sanborni Hylini

Figure 5 Partial phylogeny of the family Hylidae, showing relationships among 65 species within the subfamily Hylidae (Redrawn after Wiens et al, 2006).

136 •Acris crepitans - Acris gryllus • Pseudacris cadaverina - Pseudacris regilla • Pseudacris crucifer " Pseudacris ocularis - Pseudacris ornata • Pseudacris streckeri • Pseudacris brachyphona • Pseudacris brimleyi - Pseudacris nigrita " Pseudacris feriarum - Pseudacris triseriata . "Pseudacris maculata • Pseudacris clarkii -Hyla melanomma - Hyla sumichrasti - Plectrohyla glandulosa - Plectrohyla guatemalensis -Plectrohyla matudai - Hyla arborescandens - Hyla thorectes -Hyla pentheter - Hyla taeniopus - Hyla miotympanum -Ecnomiohyla miliarie - Ptychohyla spinipollex - Hyla bromeliacia - Duellmanohyla rufioculis - Ptychohyla hypomykter - Ptychohyla euthysanota - Ptychohyla leonhardschultzei -Tlalocohylapicta - Hyla smithii -Hyla loquax -Hyla pseudopuma - Hyla zeteki - Isthmohyla rivularis - Hyla tica -Anotheca spinosa -Triprion petasatus -Smilisca baudinii - Pternohyla fodiens - Smilisca cyanosticta - Smilisca phaeota -Smilisca puma -Hyla annectans • Hyla chinensis - Hyla meridionalis - Hyla savignyi - Hyla arborea - Hyla cinerea - Hyla squirella - Hyla andersonii - Hyla femoralis - Hyla gratiosa - Hyla chrysoscelis - Hyla versicolor -Hylaavivoca - Hyla japonica - Hyla walkeri - Hyla arenicolor - Hyla wrightorum - Hyla eximia - Hyla plicata - Hyla euphorbiacea

Figure 6 Partial phylogeny of the family Hylidae, showing relationships among 66 species within the tribe Hylini (Redrawn after Wiens et al, 2006).

137 other Hylinae Hyloscirtus palmeri Hyla phyllognatha Hyla colymba Hyla simmonsi Litoria armatus Hyla martinsi Hyla circumdata Hyla pseudopseudis Hyla ehrhardti Hyla albofrenata Aplastodiscus perviridis Hyla lemal Hyla boans Hyla geographies Hyla sibleszi Hypsiboas cinerascens Hyla punctata Hyla heilprini Hyla ranjceps. Hyla fasciata Hyla calcarata Hyla lanciformis Hyla albopunctata Hypsiboas albomarginatus Hyla rufitela Hyla rosenbergi Hyla crepitans Hyla faber Hyla pardalis Hyla semiguttata Hyla polytaenia Hyla andina Hyla balzani Hyla marianitae Hyla bischoffi Hyla marginata Hyla guentheri Hyla pulchella ^ Hyla prasina

Figure 7 Partial phylogeny of the family Hylidae, showing relationships among 39 species within the subfamily Hylidae (Redrawn after Wiens et al, 2006).

138 "Colostethus palmatus • Colostethus stepheni -Colostethus beebei -Mannophryne trinitatis - Mannophryne collaris - Colostethus talamancae • Colostethus nubicola " Colostethus brunneus •Allobateszaparo -Allobatesfemoralis •Colostethus caeruleodactylus "Colostethus nidicola • Colostethus flotator " Epipedobates boulengeri * Epipedobates espinosai • Epipedobates tricolor • Epipedobates anthonyi - Colostethus pratti • Colostethus inguinalis - Colostethus panamensis • Epipedobates trivittatus • Epipedobates silverstonei • Epipedobates parvulus • Epipedobates pictus - Epipedobates petersi - Epipedobates pulchripectus " Epipedobates hahneli -Colostethus subpunctatus -Colostethus sylvaticus -Colostethus nexipus - Colostethus vertebralis -Colostethus idiomelus -Colostethus insulatus - Colostethus awa -Colostethus infraguttatus -Colostethus elachyhistus - Phyllobates bicolor - Phyllobates aurotaenia - Phyllobates terribilis - Phyllobates vittatus - Phyllobates lugubris - Dendrobates fulguritus - Dendrobates minutus - Dendrobates imitator - Dendrobates vanzolinii -Dendrobates ventrimaculatus "Dendrobates variabilis "Dendrobates reticulatus "Dendrobates fantasticus "Dendrobates galactonotus -Dendrobates quinquevittatus - Dendrobates granuliferus -Dendrobates arboreus , -Dendrobates speciosus •Dendrobates pumilio -Dendrobates histrionicus "Dendrobates lehmanni • Dendrobates tinctorius •Dendrobates azureus -Dendrobates leucomelas • Dendrobates truncatus •Dendrobates auratus

Figure 8 Phylogeny of the superfamily Dendrobatoidea, showing the relationships among 62 species.

139 Bufo viridis Bufo americanus Bufo houstonensis Bufo fowleri Bufo hemiophrys Bufo woodhousii Bufo terrestris Bufo californicus Bufo microscaphus Bufo cognatus Bufo speciosus Bufo retiformis Bufo debilis Bufo quercicus Bufo punctatus Bufo canorus Bufo exsul Bufo boreas Bufo nelsoni •£ Bufo luetkenii Bufo melanostictus Bufo valliceps Ollotis ibarrai Bufo coccifer Bufo fastidiosus Bufo coniferus Bufo alvanus Bufo margaritifer Bufo castaneoticus Bufo dapsilis Bufo chavin Bufo spinulosus Bufo arunco Bufo marinus Bufo schneideri Bufo arenarum Bufo crucifer Bufo granulosus Bufo juxtasper Bufo asper Bufo andrewsi - Bufo bufo Bufo regularis Bufo maculatus •c Bufo kisoloensis Bufo gutturalis Bufo xeros u±Buf o mauritanicus Bufo garmani Bufo rangeri Bufo pardalis cESchismaderm a carens Bufo melanochlorus Bufo biporcatus Bufo glaberrimus Bufo guttatus Bufo haematiticus CEBuf o variegatus

Figure 9 Supertree of the family Bufonidae, showing the relationships among 58 species.

140 Lankanectes corrugatus Staurois latopalmatus Staurois natator Meristogenys jerboa Meristogenys kinabaluensis Amolops ricketti Amolops wuyiensis Amolops cremnobatus Amolops viridimaculatus Amolops chunganensis Amolops mantzorum Rana lessonae Rana nigromaculata Hydrophylax galamensis Rana signala Rana chalconota Fejervarya nicobariensis Rana temporalis Rana daemeli Rana nigrovittata Rana guentheri Rana erythraea Rana taipehensis Rana minima Rana emeljanovi Rana tientaiensis Rana adenopleura Rana pleuraden Amolops chapaensis Rana margaretae Rana andersonii Rana grahami Rana schmackeri Rana liyida Rana versabilis Rana shuchinae Rana temporaria Rana arvalis . Rana macrocnemis Rana graeca Rana latastei Rana dalmatina Rana asiatica Rana okinavana Rana tsushimensis Rana japonica Rana chaochiaoensis Rana omeimontis Other Ranidae

Figure 10 Basal portion of the supertree of the family Ranidae, showing the relationships among 48 species.

141 amurensis omativenifis chensinensis d/bowskii boylii iuteiventris muscosa aurora cascadae sylvatica virgatipes septentrionalis gfylio clamitans catesbeiana heckscheri vibicaria warszewitschii vaillanti palmipes tarahumarae pipiens chiricahuensis palustris areolata sevosa capita berlandieri blairi sphenocephala forreri yavapaiensis onca

Figure 11 Partial supertree of the family Ranidae, showing the relationships among

33 species.

Occidozyga martensii Occidozyga lima Occidozyga laevis Limnonectes kuhlii Limnonectes laticeps Limnonectes gyldenstolpei Limnonectes microdiscus Limnonectes ibanorum Limnonectes finchi Limnonectes palavanensis Limnonectes parvus Limnonectes paramacrodon Limnonectes macrodon Limnonectes blythii Nannophrys ceylonensis Euphlyctis cyanophlyctis Hoplobatrachus occipitalis Hoplobatrachus chinensis Sphaerotheca breviceps Fejervarya limnocharis Fejervarya cancrivora Paa yunnanensis Chaparana unculuanus Nanorana parkeri Nanorana pleskei Paa spinosa Paa boulengeri

Figure 12 The supertree of the family Dicroglossidae, showing the relationships

among 27 species.

145 APPENDIX 4 Result of the correlation tests among body size, body size dimorphism and life history traits in the all-anuran analysis and in the within-family analyses using the program of Comparative Analysis via Independent Constrasts (CAIC) and the

BayesDiscrete program. The within-family analyses were conducted in the superfamily Dendrobatoidea, the families Bufonidae,

Dicroglossidae, Hylidae, Megophryidae, and Ranidae. In GAIC, the correlations between two continuous traits were tests in the

CHRUNCH procedure; the correlations between one continuous trait and one binary trait were tests in the BRUNCH procedure. In

BayesDiscrete, the correlated evolution of two binary traits was tested by comparing the log-maximum likelihoods of the independent and dependent models. Results with numbers of comparisons smaller than three are not presented. Results of significant correlations are in bold. SSD = sexual size dimorphism (F>M); RSSD = reversed sexual size dimorphism (F<=M).

143 Table 1 Correlations between body size and mating related life history traits in anurans.

. Hypotheses Correlations Programs All Anurans Bufonidae Dendrobatoidea Dicroglossidae Hylidae Megophryidae Ranidae P 0.28 0.085 0.036 0.33 0.23 ... 0.26 Log Male Body Male Body Size 2 Size CMC R 0.03 0.56 0.49 0.45 0.14 ... 0.24 (BRUNCH) Male Combat Number of Contrasts 37 5 8 6 11 ... 6 Male Combat Slope 0.02 0.1 0.07 -1.87 1.05 — -0.05 P 0.044 — <0.0001 — — — — Female Body Log Female 2 Size Body Size CMC R 0.46 — 0.50 — ...... — (BRUNCH) Number of Contrasts 8 — 8 — ... — — Female Combat Female Combat Slope 0.07 — 0.01 — ... — —

Log Male Body P 0.43 0.08 0.75 — 0.40 — 0.30 Male Body Size Size 2 CMC R 0.03 0.57 0.04 — 0.24 — 0.21 Male Scramble (BRUNCH) Male Scramble Number of Contrasts 21 5 4 — 4 ... 6 Competition Competition Slope 0.03 0.1 -0.01 — 0.10 ... -0.06

Log Male Body P 0.36 — 0.52 — 0.23 — Male Body Size Size 2 CMC R 0.04 — 0.06 — 0.17 — — Male Territory (BRUNCH) Male Territory Number of Contrasts 25 — 8 — 9 — — Defense Defense Slope 0.03 — 0.03 — 0.05 ... —

Male Body Size Male Body Size P 0.77 — — — 0.51 — 0.70 2 CMC R O.01 — — — 0.06 — 0.02 Length of Prolonged (BRUNCH) ' Breeding Breeding Number of Contrasts 29 — — — 9 — 9 Season Season Slope 0.01 — — — -0.03 — -0.02

144 Table 2 Correlations between body size dimorphism and mating related life history traits.

Hypotheses Correlations Programs All Anurans Bufonidae Dendrobatoidea Dicroglossidae Hylidae Megophryidae Ranidae P 0.85 0.56 0.79 0.43 0.15 — 0.51 Body Size Ratio 2 CMC R 0.001 0.090 0.011 0.71 0.19 — 0.090 (BRUNCH) Male Combat Number of Contrasts 37 5 8 6 11 — 6 SSD Slope -0.001 0.013 -0.001 0.064 -0.016 — 0.090 Male Combat Independent j -438.38 -38.48 -61.77 -24.23 -125.73 ... -71.85 RSSD BayesTraits Dependent 1 -434.07 -37.14 -60.50 -23.83 -124.78 ... -68.99 Discrete Male Combat Likelihood Ratio 4.31 1.34 1.27 0.40 0.96 — 2.86 Correlation n.s. - n.s. n.s. n.s. n.s. ... n.s.

SSD P 0.84 0.56 0.053 — 0.50 — 0.73 Body Size Ratio 2 CMC R 0.002 0.090 0.65 — 0,16 — 0.025 Male Male Scramble (BRUNCH) Scramble Number of Contrasts 21 5 4 — 4 — 6 Competition Competition Slope -0.002 0.013 -0.010 — -0.010 — -0.009 P 0.624 — 0.569 — 0.098 — 0.716 Body Size Ratio 2 CMC R 0.010 ... 0.042 — 0.306 ... 0.186 Male Territory (BRUNCH) Number of Contrasts 25 — 8 — 9 ... 6 SSD Defense Slope -0.005 — -0.003 — -0.023 — 0.034 Male Territory Independent -354.00 ... -58.97 ... -115.89 ... -47.86 Defense RSSD BayesTraits Dependent -351.64 — -58.01 — -115.33 ... -46.19 Male Territory Discrete Likelihood Ratio 2.36 ... 0.96 0.56 — 1.67 Defense Correlation n.s. ... n.s. — . n.s. — n.s. P 0.92 — 0.92 ... — — — SSD Body Size Ratio 2 CMC R 0.002 — 0.002 — — — ... Female (BRUNCH) Female Combat Number of Contrasts 8 — 8 — — — ... Combat ...... Slope | -0.001 — -0.001 -

145 Body Size P 0.89 — — 0.40 — 0.50 Ratio 2 CAIC R 0.001 — — .— 0.09 ... 0.059 Prolonged (BRUNCH) Number of Contrasts 29 — — — 9 ... 9 SSD . Breeding Season Slope 0.002 ... — — -0.02 — 0.031 Length of ... — Breeding RSSD Independent -370.07 — -109.59 — -71.71 Season BayesTraits Dependent -367.31 ... — — -106.46 ... -69.70 Prolonged Discrete Breeding Likelihood Ratio 2.76 ... — ... 3.13 2.01 Season Correlation n.s. — — ... n.s. ... n.s.

146 Table 3 Correlations between body size and breeding related life history traits in anurans.

All Anurans Hylidae Bufonidae Dendrobatoidea Dicroglossidae Megophryidae Ranidae Hypotheses Correlations Programs Outliers Outliers All Data All Data excluded excluded P <0.0001 0.080 0.25 0.43 0.049 <0.0001 0.76 0.025 0.044 Female Log Female R2 0.18 0.02 0.16 0.05 0.33 0.53 0.002 0.49 0.12 Body Size Body Size CMC Number of (CRUNCH) 157 155 8 15 11 54 52 9 32 Egg Size Egg Size Contrasts Slope 0.01 0.20 0.33 0.009 0.21 7.62 0.009 0.4 0.17 P <0.0001 0.28 0.003 0.78 <0.0001 0.0094 <0.0001 Female Body Female Size R2 0.33 0.19 0.35 0.01 0.48 0.64 0.48 Body Size CMC (CRUNCH) Number of Log Clutch 141 7. . 22 7 46 8 29 Clutch Size Contrasts Size Slope 7.02 7.11 7.5 -1.87 6.65 25.87 9.55 P <0.0001 — 0.17 0.004 0.06 <0.0001 ^^ 0.0024 0.0001 Female Female Body Body Size Size 2 CMC R 0.38 — 0.34 0.51 0.48 0.53 ^^ 0.75 0.46 Number of (CRUNCH) 125 6 13 7 44 8 27 Clutch Log Clutch Contrasts — ^^ Volume Volume Slope . 8.23 — -0.13 8.2 12.26 7.62 ^^ 23.47 10,18 Male Body P 0.58 — 0.11 — — — — Log Male Size Body Size R2 0.004 — ... CMC ^^ — 0.31 — — I Male (BRUNCH) Number of Male 12 — 3 — — — — 1 Parental Contrasts ^^ Parental Care | Care Slope -0.01 — -0.08 .— — ^^* — — Female P 0.84 — 0.11 — — — — Log Female Body Size Body Size R2 0.04 — 0.80 — — — — CMC Female (BRUNCH) Number of Female 10 /^• — 3 — — — — J Parental Contrasts Parental Care Care Slope -0.02 ^^ — -0.08 — — — —

147 Table 4 Correlations between body size dimorphism and breeding related life history traits in anurans.

All Hylidae Hypotheses Correlations Programs Bufonidae Dendrobatoidea Dicroglossidae Outliers Megophryidae Ranidae Anurans All Data excluded P 0.70 0.61 0.70 0.064 0.015 ^^ 0.48 0.33 Body Size SSD 2 Ratio CMC R 0.001 0.039 0.011 0.30 0.11 0.065 0.03 (CRUNCH) Number of Egg Size 156 8 15 11 54 9 32 Log Egg Size Contrasts Slope -0.015 0.069 0.015 -0.13 0.218 -0.16 -0.17 P <0.0001 0.49 0.007 0.49 <0.0001 0.81 0.33 Body Size o.n 2 SSD Ratio CMC R 0.152 0.082 0.30 0.083 0.37 0.001 0.13 0.80 Number of (CRUNCH) 141 7 22 7 46 139 30 Clutch Size Log Clutch Contrasts 8 Size Slope 0.053 0.020 0.035 0.033 0.079 0.01 -0.094 ' 0.05 P <0.0001 0.498 0.012 0.849 <0.0001 0.61 0.390 0.14 Body Size SSD 2 Ratio CMC R 0.183 0.096 0.425 0.007 0.549 0.01 0.107 0.08 Number of Clutch (CRUNCH) 125 6 13 7 44 123 8.000 28 Log Clutch Contrasts Volume Volume Slope 0.066 0.023 0.035 0,012 0.109 0.02 -0.070 0.06 P 0.94 0.81 — — — ... Body Size — 2 — Ratio CMC R 0.001 — 0.04 — — — Number of (BRUNCH) 12 ' 3 ...... SSD Male Parental Contrasts — — — Care Slope 0.001 — 0.003 — — ^^ — ... Male Parental Independent -333.33 — -47.47 — ...... Care RSSD -327.19 -44.64 ..." ... — ... BayesTraits Dependent — Likelihood Male Parental Discrete 6.14 — 2.83 — ...... Care Ratio Correlation n.s. — n.s...... — ...

148 P 0.64 ... 0.019 — -72.26 ^^ ... — SSD Body Size Ratio 2 ... — — — CMC R 0.026 0.96 -69.69 Female Number of (CRUNCH) 10 ... 3 2.57 Parental Female Contrasts — — — Care Parental Care Slope -0.005 ... -0.028 — n.s. — —

149 Table 5 Correlations among breeding related life history traits m anurans.

Hypotheses Correlations Programs All Anurans Bufonidae Dendrobatoidea Dicroglossidae Hylidae Megophryidae Ranidae P 0.0085 0.0014 0.78 0.13 0.69 0.52 0.34 Egg Size 2 Egg Size CAIC R 0.03 0.41 0.005 0.26 0.05 0.04 0.02 Log Clutch (CRUNCH) Number of Contrasts 223 21 17 9 72 11 42 Clutch Size Size Slope -0.33 -0.36 0.08 -0.69 0.51 -0.18 -0.31 P 0.38 — — ... 0.48 — — Parental Care 2 CAIC R 0.02 ... — ... 0.06 ... — (BRUNCH) Parental Log Egg Size Number of Contrasts 32 — ... — 9 ... — Care Slope 0.04 ...... -0.06 ... — Fecundity P 0.01 — ... — 0.52 ... — Related Parental Care Traits R2 0.19 — — ... 0.05 ...... CAIC • Log Clutch (BRUNCH) Number of Contrasts 35 — — — 9 — ... Size Slope -0.33 ...... -0.21 ... — Table 6 Correlations between female body size and male body size as well as correlations between body size dimorphism and body size in anurans.

Hylidae Hypotheses Correlations AH Anurans Bufonidae Dendrobatoidea Dicroglossidae Megophryid Ranidae Programs Outlier ae AH Data excluded P <0.0001 O.0001 O.0001 O.0001 O.0001 Female Log Female O.0001 O.0001 Body Size 2 Body Size CAIC R 0.96 0.98 0.95 0.91 0.98 0.65 0.72 Number of (CRUNCH) 247 16 42 15 89 14 39 Male Body Log Male Contrasts Size Body Size Slope J 1.01 0.916 1.04 0.85 1.01 1.14 0.98 P j <0.0001 0.023 0.15 0.032 O.0001 0.69 0.46 0.52 Body Size SSD Ratio 2 CAIC R 0.33 0.30 0.05 0.29 0.48 0.002 0.04 0.01 (CRUNCH) Number of J 247 16 42 15 89 87 14 39 Body Size Log Body Contrasts 1 Size Slope J 0.20 0.16 0.04 -0.18 0.21 -0,02 -0.12 -0.12 .

151