Morphology and Systematics of the Solomon Island Ranid Frogs

Morphology and Systematics of the Solomon Island Ranid Frogs

Rachel M. Norris

Department of Environmental Biology

The University of Adelaide

A thesis submitted for the degree of Doctor of Philosophy, at the University of Adelaide December, 2002

i Table of Contents

Abstract iv Declaration vi Acknowledgements vii Publications resulting from this thesis viii Chapter 1. Introduction 1 Chapter 2. Historical Account 6 2.1. Taxonomy of the Solomon Island ranids: Species accounts 6 2.2. Taxonomy of the Solomon Island ranids: Subfamily accounts 22 2.3. Mode of Life History: a uniting feature? 29 2.4. Zoogeography of the Solomon Islands and surrounding islands 31 Chapter 3. Morphometrics 38 3.1. Introduction 38 3.2. Methods 39 3.2.1. How many PCs? 41 3.3. Results 42 3.3.1. Analysis of Separate Sexes 42 3.3.2. Analysis of Sexes Combined 50 3.3.3. Further Analysis: Discriminant Function Analysis 65 3.4. Discussion 76 Chapter 4. Osteology of Solomon Island Ranids 79 4.1. Introduction 79 4.2. Methods 79 4.2.1. Material Examined 80 4.3. Osteological Descriptions 81 4.3.1. Batrachylodes Boulenger, 1887 81 4.3.1.1. Batrachylodes elegans Brown and Parker, 1970 81 4.3.1.2. Batrachylodes mediodiscus Brown and Parker, 1970 83 4.3.1.3. Batrachylodes trossulus Brown and Myers, 1949 86 4.3.1.4. Batrachylodes vertebralis Boulenger, 1887 92 4.3.1.5. Batrachylodes wolfi (Sternfeld, 1918) 97 4.3.2. Ceratobatrachus Boulenger, 1884 102 4.3.2.1. Ceratobatrachus guentheri Boulenger, 1884 102 4.3.3. Discodeles Boulenger, 1918 109 4.3.3.1. Discodeles bufoniformis (Boulenger, 1884) 109 4.3.3.2. Discodeles guppyi (Boulenger, 1884) 115 4.3.4. Palmatorappia Ahl, 1927 120 ii 4.3.4.1. Palmatorappia solomonis (Sternfeld, 1920) 120 4.3.5. Platymantis Günther, 1859 "1858" 126 4.3.5.1. Platymantis guppyi (Boulenger, 1884) 126 4.3.5.2. Platymantis myersi Brown, 1949 132 4.3.5.3. Platymantis neckeri (Brown and Myers, 1949) 138 4.3.5.4. Platymantis parkeri (Brown, 1965) 143 4.3.5.5. Platymantis solomonis (Boulenger, 1884) 149 4.3.5.6. Platymantis weberi Schmidt, 1932 155 4.3.6. Rana Linnaeus, 1758 160 4.3.6.1. Rana kreffti Boulenger, 1882 160 Chapter 5. Karyology of the Solomon Island Ranids 167 5.1. Introduction 167 5.2. Methods 167 5.2.1. Material examined 167 5.2.2. Preparation of Chromosome spreads 168 5.2.3. Giemsa Banding (G-banding) 168 5.2.3.1. Procedure 169 5.2.4. Constitutive Heterochromatin Banding (C-banding) 169 5.2.4.1. Procedure 170 5.2.5. Silver Staining - Staining for active nucleolus organiser regions 170 5.2.6. Photomicroscopy 170 5.3. Results 171 5.3.1. Solomon Island species 171 5.3.2. Comparison Species from literature 178 5.4. Discussion 181 Chapter 6.Phylogenetic Analysis 184 6.1. Introduction 184 6.2. Methods 185 6.2.1. Character Selection 191 6.2.2. Outgroup selection 192 6.2.3. Assumptions for phylogeny reconstruction 193 6.2.3.1. Character weighting 193 6.2.3.2. Missing data 193 6.2.3.3. Character ordering 194 6.2.4. Support and Confidence 194 6.3. Results 195 6.3.1. Analysis 1 195 6.4. Character Reassessment 217 iii 6.4.1. Analysis 2 217 6.4.2. Analysis 3 221 6.4.3. Analysis 4 224 6.5. Discussion 225 References 230 Appendix 1. Material Examined 242 Appendix 2. 273 Discriminant Function Analysis 273 Boxplots genera females 277 Boxplots genera males 280 Boxplots species females 283 Boxplots species males 300 Appendix 3. Character Descriptions 315 iv Abstract

The ranid frogs of the Solomon Islands have been known in the taxonomic literature for over 100 years. The Solomon Islands and surrounding islands contain a unique array of direct-developing ranids, most of which are endemic to the island masses on which they live. However, their phylogenetic relationships to each other and to other ranids in the southwest Pacific region have not been defined. Their assumed relationships are based on a restricted sample of osteological characters and the assumption of direct development, inferred from egg size and number. This study validates the Solomon Island taxa (using morphometrics) and explores the biology of the Solomon Island ranids, with detailed osteological descriptions, external morphology and karyology. Using characters from these data sets a cladistic analysis using parsimony reconstructed a phylogeny of these frogs. Morphometric analyses were undertaken on two levels, firstly using principal components analysis (PCA) to assess sexual dimorphism, and secondly using discriminant function analysis (DFA) to test the identity of the taxa. Sexual dimorphism was not recognised in morphometric characters and so sexes were pooled for the DFA. For Batrachylodes 93.08% of individuals were successfully classified to their actual groups, Discodeles 89.77% and for Platymantis 84.55%. Osteological descriptions of Batrachylodes elegans, B. mediodiscus, B. trossulus, B. vertebralis, B. wolfi, Ceratobatrachus guentheri, Discodeles bufoniformis, D. guppyi, Palmatorappia solomonis, Platymantis guppyi, P. myersi, P. neckeri, P. parkeri, P. solomonis, P. weberi and Rana kreffti are provided. Karyotypes of eight species of ranid frogs from the Solomon Islands were compared and contrasted with the 2n = 26, FN = 52 karyotype of Rana, the typical karyotype of the subfamily Raninae. This karyotype was found in the Solomon Island species of Rana, R. kreffti. Ceratobatrachus guentheri, had an increased 2n of 30, a lower FN of 38, and altered relative lengths and centromere positions of pairs 1-5. These changes could have resulted from centric fissions and pericentric rearrangements which produced an increase in the number of telocentric chromosomes. The remaining seven species (Batrachylodes vertebralis, Discodeles bufoniformis, D. guppyi, Platymantis myersi, P. neckeri, P. solomonis and P. weberi) had reduced diploid numbers and FN. The means by which reduction in 2n and FN has occurred in these species is unknown, but may either involve centric fissions to produce telocentrics, followed by translocation onto other v chromosomes, or a process involving pericentric rearrangements to produce telocentric chromosomes followed by fusions of these products. With the exception of Rana, the level of chromosome rearrangements in these frogs that are endemic to the Solomon Islands is high compared with that observed in the continental lineages of this subfamily. Phylogenetic analysis using maximum parsimony found three equally parsimonious trees. Subsequent character reanalysis (successive weighting) produced one parsimonious tree. The phylogenies indicate multiple invasion events into the Solomon Islands by these ranid frogs and despite the high level of endemism, monophyly is not supported.

vi Declaration

This work contains no material which has been accepted for the award of any other degree or diploma in any university or tertiary institution and, to the best of my knowledge and belief, contains no material previously published or written by another person, except where due reference has been made in the text.

I give consent to this copy of my thesis, when deposited in the University Library, being made available for loan and photocopying.

vii ACKNOWLEDGEMENTS

I would like to thank the following people and institutions who have waited patiently for this thesis.

■ Associate Professor Michael Tyler, for supervising the thesis, for your advice and support and willingness to read it and help. ■ Dr Margaret Davies, for tirelessly pursuing me to keep on track. Without your advice and support and editing skills I doubt I could have finished it. ■ The Department of Environmental Biology (formerly Zoology) for their patience. I'm sorry Bill that you didn’t get to read it. ■ Professor Maciej Henneberg and the Department of Anatomical Sciences, for support and study leave. ■ Professor Arnold Kluge, thank you so much for your advice and help throughout this thesis. ■ Dr Tom Burton, for the visit all those years ago, only to find out that in Solomon Island ranids, big frogs have big muscles and little frogs have little muscles. ■ Dr Michael Mahony, University of Newcastle, for support and advice on the chromosomes. ■ Dr Steve Donnellan, South Australian Museum, for collecting frogs and trying to get some molecular work up and running, alas just not my thing. ■ Dr Mark Hutchinson, South Australian Museum, for discussions on cladistics and reading a draft of chapter 6. ■ Mr Jens Vindum, Dr Alan Leviton, Department of Herpetology, California Academy of Sciences; Mr José Rosado, Museum of Comparative Zoology, Harvard; Prof. Arnold Kluge, Department of Zoology, University of Michigan; Dr Darrel Frost, Department of Herpetology, American Museum of Natural History; Mr Alan Resetar, Dr Harold Voris, Field Museum of Natural History, Chicago; Mr Ross Sadlier, Dr Alan Greer, Division of Herpetology, Australian Museum, Sydney; Ms Adrienne Edwards, Dr Mark Hutchinson, Division of Herpetology, South Australian Museum, Adelaide, for allowing me to visit their departments and the loan of material. ■ The Royal Society of South Australia, The American Museum of Natural History, The Field Museum of Chicago, The DR Stranks Travelling Fellowship and the University of Adelaide Overseas Travelling Scholarship for various funding to allow me to pursue all aspects of this thesis and the support of The University of Adelaide Postgraduate Scholarship. ■ My family. ■ My friends especially Carmel, Anna and Genevieve. viii Publications Resulting from this Thesis

Mahony, M.J., Norris, R.M. and Donnellan, S.C. (1996) Karyotypes of Southwest Pacific Ranid Frogs (Anura: Ranidae). Australian Journal of Zoology, 44, 119- 128.

Norris, R.M. (1999) Testing multiple species hypothesis on frogs. Perspectives in Biology, 4(1):51-64

Chapter 1. Introduction

The Solomon Islands have been regarded as an amphibian outpost (Boulenger, 1886a; Brown, 1952). Boulenger (1886a:35) considered the Islands to be “on the limits of two great zoological districts... where many Papuasian and Polynesian forms intermingle”. Brown (1952) suggested that the amphibian fauna of the Solomon Islands originated from two ancestral elements: an older element of highly specialised endemic frogs, Batrachylodes, Ceratobatrachus, Discodeles, Palmatorappia and Platymantis (in part) and a more recent element including Rana and one or two species of Platymantis. This hypothesis has never been tested. Uncertainties as to the number of ancestral invasions by ranid frogs into the Solomon Islands, raise a number of questions. It is possible that there has been: i one invasion and then radiation, if one accepts Brown’s older highly specialised ranid element, whose centre of abundance is in the Solomon Islands and surrounding islands (Brown, 1952). ii two invasions from one ancestral stock, possibly from the more widely dispersed and hence ancestral genus, Platymantis (Noble, 1931). or iii Two invasions from two different ancestral stocks as evidenced from the two elements identified by Brown (1952), possibly as an older invasion resulting in the evolution of Batrachylodes, Ceratobatrachus, Discodeles, Palmatorappia and some members of Platymantis, followed by a more recent invasion indicated by Rana and some members of Platymantis.

The ranid frogs of the Solomon Islands are well known taxonomically but their phylogenetic relationships have not been investigated. They are endemic to the region and are highly diverse morphologically although they do share the same reproductive strategy, that of direct development, as extrapolated from the presence of large unpigmented eggs. Brown (1952) concluded that a detailed phylogenetic analysis is needed in order to elucidate the relationships of the Solomon Island ranids. Currently 25 species distributed among six genera (Table 1.1) are recognised from the Solomon Islands and surrounding islands (Frost, 2002).

1 Table 1.1. Current taxonomic status of Solomon Island ranid frogs. Species Author Batrachylodes elegans Brown and Parker, 1970 Batrachylodes gigas Brown and Parker, 1970 Batrachylodes mediodiscus Brown and Parker, 1970 Batrachylodes minutus Brown and Parker, 1970 Batrachylodes montanus Brown and Parker, 1970 Batrachylodes trossulus Boulenger, 1884 Batrachylodes vertebralis Boulenger, 1887 Batrachylodes wolfi (Sternfeld, 1918) Ceratobatrachus guentheri Boulenger, 1884 Discodeles bufoniformis (Boulenger, 1884) Discodeles guppyi (Boulenger, 1884) Discodeles malukuna Brown and Webster, 1969 Discodeles opisthodon (Boulenger, 1884) Discodeles vogti (Hediger, 1934) Palmatorappia solomonis (Sternfeld, 1918) Platymantis acrochorda (Brown, 1965) Platymantis aculeodactylus Brown, 1952 Platymantis guppyi (Boulenger, 1884) Platymantis macrops (Brown, 1965) Platymantis myersi Brown, 1949 Platymantis neckeri (Brown and Myers, 1949b) Platymantis parkeri (Brown, 1965) Platymantis solomonis (Boulenger, 1884) Platymantis weberi Schmidt, 1932 Rana kreffti Boulenger, 1882

Although Bougainville Island is politically a part of Papua New Guinea, this thesis recognises this island mass as historically and geologically belonging to the Solomon Islands (Fig. 1.1).

2

Fig. 1.1. The Solomon Islands including Bougainville Island and its surroundings (map downloaded from: http://www.janeresture.com/kiribati_phoenix/map.htm).

The Solomon Islands form two island chains bracketed by Bougainville in the west and San Cristobal in the east. The entire island chain is 650 miles in length (Thompson and Hackman, 1969). The islands that make up the Solomon Island chain did not all form at the same time with the oldest deposits in Malaita determined from the Upper Cretaceous whilst basalts in Choiseul and Guadalcanal dated at the end of the Oligocene (Thompson and Hackman, 1969). Volcanism throughout the Oligocene and Miocene formed the remainder of the island chain with the exception of New Georgia where the geology is younger having been deposited in the Pliocene (Thompson and Hackman, 1969). The emergence of all the islands continued throughout the Pleistocene to the Recent. In addition, Thompson and Hackman (1969) record that sea levels have been much lower in the past than at present and suggested land links between the islands of Bougainville, Choiseul, Santa Isabel, the Shortlands group and possibly Florida and Guadalcanal. However this lowering of the sea level did not influence

3 the distance and hence separation of the islands of Malaita, New Georgia and San Cristobal (Thompson and Hackman, 1969). Such an arrangement may explain the distribution of the ranid frog taxa occurring on these islands, especially the assumed depauperate nature of that on San Cristobal. The Solomon Islands has a diverse array of fauna including many species of insects, birds, land molluscs, reptiles and frogs. Many are endemic with restricted distributions to one or a few islands. Whilst studying birds and insects from the islands, Greenslade (1969), noticed that the overall movement of taxa throughout the islands was in a west to east direction. Greenslade (1969) also found that movement occurred from larger islands (i.e. Bougainville) to smaller islands (i.e. San Cristobal). The dominating effect of larger islands was reflected in the number of species occurring on the islands, with larger islands have more species (Greenslade, 1969). Such observations are mimicked in the ranid frog taxa with the majority of species being found in Bougainville and gradually decreasing in an west to east direction to San Cristobal. It may be that Bougainville is a centre of radiation for these ranid frogs. The aim of this research is to investigate the phylogenetic relationships of the ranid frogs from the Solomon Islands, using characters derived mainly from gross morphology. There are other species of frogs on these islands, two are hylids, Litoria thesaurensis, L. infrafrenata and the other is the introduced marine toad, Bufo marinus, they will not be considered in this thesis. Chapter 2 provides an historical account of the taxonomy of the Solomon Island ranid frogs. Such literature identifies and details many morphological features, both external and internal, of these animals and highlights the taxonomic and phylogenetic problems associated with this fauna. Chapter 3 uses morphometrics to confirm existing taxonomic groupings within the Solomon Islands and also examined areas of sexual dimorphism. Such tasks are necessary before considering species for phylogenetic analysis. Chapter 4 provides detailed osteological descriptions of all species that were available for examination and dissection. This chapter provides a basis for the character analysis in the phylogenetic reconstruction. Previously, no detailed osteology has been described as a species by species account for these animals. Chapter 5 examines the karyology of the Solomon Island ranid frogs. Many studies use molecular biology to provide characters for phylogenetic analysis and hence examining the

4 chromosomes of these frogs might provide some phylogenetic information. Chapter 6 details the phylogenetic analysis based on the characters obtained from osteology, morphometrics and external morphology. Outgroups are discussed, characters and character states are described in detail and phylogenetic trees are presented. The chapter also discusses the pros and cons of a variety of methods dealing with character reassessment. The results are considered in light of the historical predictions of the geographic origins of the Solomon Island ranid frogs.

5 Chapter 2. Historical Account Early literature containing accounts of Solomon Island ranids recognised several genera that have subsequently been synonymised. Cornufer comprised Platymantis guppyi, P. neckeri and others. Hylorana and/or Hylarana comprised Rana kreffti. Hylodes comprised some members of Platymantis. Discodeles bufoniformis, D. guppyi and D. opisthodon were originally placed in Rana. For the purpose of discussion of this literature the later names will be used, corresponding to Frost (1985, 2002), the only exception being that for historical accuracy the genus Cornufer will be used for the large-disced species of Platymantis prior to Zweifel (1967).

2.1 Taxonomy of the Solomon Island ranids: Species accounts The Solomon Island frogs in this study currently belong in the family Ranidae, subfamily Raninae and are distributed amongst the following genera: Batrachylodes, Ceratobatrachus, Discodeles, Palmatorappia, Platymantis and Rana (Frost, 1985, 2002). The species and their published distributions are summarised in Table 2.1.

6 Table 2.1. List of species and published distributions Species Distribution Batrachylodes elegans Bougainville I., Solomon Is. Batrachylodes gigas Bougainville I., Solomon Is. Batrachylodes mediodiscus Bougainville Is., Solomon Is. Batrachylodes minutus Bougainville I., Solomon Is. Batrachylodes montanus Bougainville I., Solomon Is. Batrachylodes trossulus Bougainville I.; Choiseul I., Solomon Is. Batrachylodes vertebralis Fauro I.; Bougainville I.; Buka I; New Georgia I.; Malaita I.; Santa Isabel I.; Ugi I.; Tulagi I.; Kolombangara I., Solomon Is. Batrachylodes wolfi Buka I.; Bougainville Is., Solomon Is. Ceratobatrachus guentheri Bougainville I.; Shortland I.; Fauro I.; Malaita I.; Tulagi I.; Santa Isabel I.; Rendova I.; Guadalcanal I.; Kolombangara I.; Choiseul I.; Russell Ids; Florida I.; Mono I.; Vella Lavella I., Solomon Is. Discodeles bufoniformis Bougainville I.; Fauro I.; Malaita I.; Tulagi I.; Santa Isabel I.; Mono I.; Ronongo I.; Ugi I.; Choiseul I.; Kolombangara I.; Vella Lavella I., San Cristobal I.; Santa Ana I., Solomon Is. Discodeles guppyi Shortland I.; Fauro I.; Malaita I.; Guadalcanal I.; Rendova I.; Gatukai I.; Bougainville I.; Santa Isabel I.; New Georgia I.; Florida I., Solomon Is.; New Britain I. Discodeles malukuna Guadalcanal I., Solomon Is. Discodeles opisthodon Bougainville I.; Fauro I.; Shortland I.; Mono I.; Kolombangara I.; San Cristobal I.; Ugi I.; New Georgia I.; Santa Ana I., Solomon Is. Discodeles vogti Rambutyo I., Admiralty Is. Palmatorappia solomonis Buka I.; Santa Isabel I.; Solomon Is. Platymantis acrochorda Bougainville I., Solomon Is. Platymantis aculeodactylus Bougainville I.; Choiseul I., Solomon Is. Platymantis guppyi Mono I.; Santa Isabel I.; Malaita I.; Bougainville I.; Choiseul I.; Guadalcanal I.; Arnavon I.; Ronongo I., Solomon Is. Platymantis macrops Bougainville I., Solomon Is. Platymantis myersi Bougainville I., Solomon Is. Platymantis neckeri Bougainville I., Solomon Is Platymantis parkeri Buka I.; Bougainville I., Solomon Is. Platymantis solomonis Stirling I.; Santa Isabel I.; Tulagi I.; Malaita I.; New Georgia I.; Bougainville I.; Mono I.; Shortland I.; Fauro I.; Kolombangara I.; Choiseul I.; Arnavon I.; Vangunu I., Solomon Is. Platymantis weberi Tulagi I.; Santa Isabel I.; Malaita I.; Guadalcanal I.; Bougainville I.; Vangunu I.; Narovo I.; Vella Lavella I., Solomon Is. Rana kreffti Buka I. to San Cristobal I.' Solomon Is.; New Ireland; Bismarck Archipelago

The Ranidae was diagnosed by Cope (1865) as follows:

“Sacral diapophyses cylindrical; simple coccyx, attached by two cotyloid cavities. Manubrium with a strong bony style; the xiphisternum similar. Fronto-parietal bones never embracing a fontanelle. Tongue extensively free, more or less deeply notched. Ear perfectly developed, no parotoids.” (Cope, 1865:114-115).

7 The first taxonomic account of a ranid frog from the Solomon Islands was that of Rana kreffti by Boulenger (1882) in his “Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum”. The species description was a redescription from an earlier study by Günther (1859) who had examined the specimens as Hylarana erythraea. In 1884, Boulenger published taxonomic descriptions of frogs from the Solomon Islands, in particular, the Treasury, Shortland and Faro Islands from a collection made by H.B. Guppy. In this paper he defined the family Ceratobatrachidae based on the “presence of teeth in both the upper and lower jaw, and by the diapophyses of the sacral vertebra not being dilated” (Boulenger, 1884:212). He diagnosed the family as follows:

“Pupil horizontal. Tongue deeply notched and cordiform, extensively free behind. Vomerine teeth. Head large, strongly ossified. Tympanum distinct. Fingers and toes free, with non-dilated tips. Outer metatarsals united. Precoracoids present; omosternum and sternum with bony style. Terminal phalanges simple.” (Boulenger, 1884:212).

Within this new family Ceratobatrachidae, Boulenger (1884) defined the genus Ceratobatrachus and the monotypic species, guentheri. Three new species of Discodeles: bufoniformis, guppyi and opisthodon were also described, although there was no generic description of Discodeles as Boulenger (1884) had assigned them to the genus Rana. He aligned D. bufoniformis with the east Indian frog, Rana kuhlii, and D. guppyi and D. opisthodon with the New Guinean frog, Rana grunniens. According to Boulenger (1884), Discodeles bufoniformis and D. guppyi were similar to each other but different from D. opisthodon in the placement of the vomerine teeth, the width of the interorbital space, the presence of an outer metatarsal tubercle and texture of the dorsum, whereas Discodeles guppyi and D. opisthodon were similar to each other but different from D. bufoniformis in the size of the tympanum, the length of the fingers and the shape of the inner metatarsal tubercle. All three species differed from each other in the amount of toe webbing and the length of the hind leg. Finally two new species of Cornufer: C. guppyi and C. solomonis were described. These species differed from each other by the extent of dilation of finger and toe discs, being very large in the former and small if not absent in the

8 latter. As with the new species of Discodeles, Boulenger did not discuss the generic placement of these two species because they were assigned to the already known genus Cornufer sensu Tschudi (1838). In 1886, Boulenger provided another account of the herpetological fauna of the Solomon Islands based on his previous taxonomic work (Boulenger, 1882, 1884). He included more detailed taxonomic descriptions and illustrations of the following ranids: Discodeles bufoniformis, D. guppyi, D. opisthodon, Rana kreffti, Cornufer guppyi, and C. solomonis and Ceratobatrachus guentheri. In the case of Ceratobatrachus guentheri, Boulenger discussed characters other than those pertaining to external morphology such as “the ova are very large” (Boulenger, 1886a:58) and the skeleton being ranoid in appearance, although differing in that the omosternum was bifurcated basally. He detailed the skull as follows:

“… strongly ossified, rough, with bony granules on the fronto-parietal and temporal regions, and the sutures nearly obliterated. The fronto-parietals expand in a large triangular postorbital process. The anterior process of the temporal, the maxillary and the quadrato-jugal have coalesced without leaving traces of sutures; the latter bone sends off a curved spine directed outwards and forwards. The posterior process of the temporal is expanded horizontally, subtriangular, and its posterior border is divided into several knobs. The lower jaw, … remarkable for the presence of acrodont teeth, is also interesting for the extreme shortness of dentary…'' (Boulenger, 1886a:58).

The genus Batrachylodes was first reported by Boulenger (1887) from a collection made by C.M. Woodford from Faro Island, Solomon Islands. Batrachylodes was described as follows:

“Pupil horizontal. Tongue oval, free and feebly nicked behind. No vomerine teeth. Tympanum distinct. Fingers and toes free, the tips dilated into large discs. Distal phalanges T-shaped. Omosternum and sternum with a bony style'' (Boulenger, 1887:337).

Both the genus Batrachylodes and the nominate species vertebralis, were described and illustrated from a single adult female. Interestingly Boulenger (1887) reported Cornufer dorsalis also from Faro Island. This frog was subsequently identified as Cornufer guppyi (Barbour, 1921), a frog already known to Boulenger (1884, 1886a). Also in 1888, Boulenger recorded for the first time Dumeril’s Cornufer corrugatus from the same region. However this was later found to be a Platymantis weberi (Schmidt, 1932).

9 Ogilby (1890) recorded the presence of Discodeles guppyi, D. opisthodon and Ceratobatrachus guentheri from Howla Island, Shortland group, Solomon Islands, thereby extending their known distribution. A new species of Rana was described by Werner in 1894 and subsequently amplified in 1900. Found in New Britain and New Guinea, the species differed from other members of the genus by having a shorter leg and snout. He named it Rana novae-britanniae. This discovery has implications for Solomon Island ranids as this frog has either been synonymised or allied with Rana kreffti. Roux (1905) summarised the family Ranidae to date. Since Boulenger’s 1882 Catalogue the number of genera recognised within the Ranidae had risen from 23 to 36. Roux (1905) added a short description of Batrachylodes (taken from Boulenger (1887)) to the list, and finished the paper with a key to all the genera known in the Ranidae. Rana, Batrachylodes and Cornufer appeared in the same part of the key united by the following characters: the absence of an intercalary bone in the digits and a developed omosternum and sternum with a bony style. However Rana and Batrachylodes were further separated from Cornufer because the latter had united metatarsals. In 1912, Vogt reported on the reptiles and amphibians from the South Sea Islands. In this report he described a new taxon, Discodeles ventricosus, from a single female specimen. According to Vogt (1912) this frog differed from other known species of Discodeles (D. bufoniformis, D. guppyi, D. opisthodon) by having more webbing between the toes and shorter hind limbs (compared with D. guppyi). A smaller tympanum and no glandular folds distinguished it from Rana kreffti. Vogt (1912) also recorded the presence of Rana novae-britanniae in these islands. Three new taxa from Buka Island in the Solomon Islands were described by Sternfeld (1918). He placed one in the genus Chaperina (C. friedericii), one in Sphenophryne (S. wolfi) and the third in Hylella (H. solomonis). The first two species he placed within the family Engystomatidae and the latter in Hylidae and provided brief taxonomic descriptions of each. It was later found (Ahl, 1927; Mertens, 1929) that these species were more appropriately associated as Batrachylodes vertebralis, B. wolfi and Palmatorappia solomonis respectively. Boulenger (1918a) began to recognise distinct divisions amongst species groups of Rana in his revision of the south-eastern-most representatives. He

10 erected “natural subgenera”, namely Rana, Discodeles and Hylorana (Boulenger, 1918a:236) within the genus Rana. A key was provided and based on characters relating to vomerine teeth, canthus rostralis shape, finger length, disc dilation, toe webbing, size of nasals and shape of omosternum. He commented that the subgenus Discodeles led to the genus Cornufer (i.e. Cornufer was derived from Discodeles). He synonymised Werner’s (1894, 1900) Rana novae-britanniae with the Solomon Island Rana kreffti, and considered Vogt's (1912) Discodeles ventricosus to be a synonym of Discodeles opisthodon. Roux (1918) confirmed Boulenger’s (1918a) assertions that the specimens described by Werner (1894, 1900) as Rana novae-britanniae from New Britain were Rana kreffti, thus extending the range of the Solomon Island species. Boulenger later clarified his earlier statement that the subgenus Discodeles led to the genus Cornufer, by reassessing whether the genus Cornufer should also be reclassified as a subgenus of Rana (Boulenger, 1918b). He noted that Cornufer differed from Rana in the possession of free or feebly-webbed toes and a complete union of the outer metatarsals. However, these characters were shared with the Melanesian species of the subgenus Discodeles. From these observations Boulenger concluded that Cornufer was “unquestionably connected to [the subgenus] Discodeles'' but that it [Cornufer] could be differentiated from [the genus] Rana [and therefore Discodeles] using Günther's (1859) notion that “species with the large digital discs (Cornufer Tschudi) should be kept distinct from those with small discs and practically free toes (Platymantis Gthr.)'' (Boulenger, 1918b:372). Boulenger (1918b) observed that the digital discs of the Solomon Island representatives of Cornufer (C. solomonis), were small and did not differ from those seen in Discodeles whereas the discs of species assigned to Cornufer had a

“transverse groove, extending across the lower surface and corresponding in position with the horizontal limb of the T-shaped terminal phalanx, is continuous with the crescentic or horseshoe-shaped groove between the upper and lower surface, thus defining a hemispherical area within the disc'' (Boulenger, 1918b:372).

Since Platymantis was a valid genus as defined by Günther (1859) who removed it from Dumeril’s (1853) Hylodes, with the following generic description:

“Vomerine teeth. Skin smooth, or with narrow folds; no large gland. Disks rather small; fingers and toes free. Tympanum distinct; eustachian tubes moderate; tongue

11 large, free and deeply notched behind. Male without vocal sac. Feejee and Philippine Islands.” (Günther, 1859:93). then Cornufer also remained a valid genus (Boulenger, 1918b). As a result this delineation in finger disc size, Boulenger (1918b) transferred the species solomonis from Cornufer to Platymantis, thus adding the latter genus to the Solomon Island herpetofauna. In 1920, Boulenger again provided a synopsis of the genus Rana. In this monograph Boulenger reiterated his concept of subgenera within Rana, by expanding the number of divisions from three to nine. He identified from Melanesia, Rana (Hylorana) kreffti, Rana (Discodeles) bufoniformis, R. (D.) guppyi and R. (D.) opisthodon. He again regarded the genera Cornufer and Platymantis as being separate but derived from Discodeles (Boulenger, 1920). He also related the species of Discodeles to Rana via the New Guinean Rana grunniens. As well as the Solomon Island species Boulenger aligned another six species to the subgenus Discodeles and then further partitioned them into the groups Ranae guppianae for the Solomon Island species and Ranae beddomianae for the remainder. The only other Solomon Island species detailed in this monograph was Rana kreffti, which was placed within the subgenus Hylorana along with other New Guinean ranids, which in turn were placed within the group Ranae papuae. Standard taxonomic descriptions were provided. Barbour (1921) reviewed all the known species from the Solomon Islands beginning with Ceratobatrachus guentheri. He suppressed the family Ceratobatrachidae on the grounds that the character, 'teeth on the upper and lower jaw', was not enough to warrant familial status. This was the only character that distinguished the Ceratobatrachidae from the Ranidae. Barbour used parallels within the Leptodactylidae to support his claim, however, he admitted to not being able to suggest any close relatives to this frog, and suggested Van Kampen’s (1919) idea of a subfamily may be a solution. He noted the variation in both coloration and pattern of Batrachylodes vertebralis and that males possessed a snout that projected anteriorly at the tip and was slightly thickened at the lip margin. Barbour (1921) suggested that this feature may aid in burrowing or pushing under logs or stones. Barbour (1921) reassessed Boulenger's (1887) record of Cornufer dorsalis from the Solomon Islands finding it to be erroneous

12 and that it was a misidentified C. guppyi. He regarded Rana kreffti as common as it was widespread throughout Papuasia, having been recorded in New Britain and the Solomon Islands whereas Discodeles guppyi had a far more restricted distribution having not been found outside the Solomon Islands. One major taxonomic rearrangement that Barbour (1921) proposed was the synonymy of Discodeles opisthodon with that of D. bufoniformis. Having examined the characters that differ between the two species (e.g. tympanum diameter, length of hind limb, dorsum texture), Barbour could not see any reason to keep them separate as they differed “by degree and not of kind” (Barbour, 1921:99). He therefore recognised only one species: D. bufoniformis. The concept of subgenera within the genus Rana was continued in Van Kampen’s (1923) work on “The Amphibia of the Indo-Australian Archipelago". In this study, Van Kampen recognised the genera Rana, Ceratobatrachus, Cornufer and Batrachylodes as having some or all of their representative species in the Solomon Islands. He also identified four subgenera within Rana: Rana, Discodeles, Hylarana (not Hylorana) and Platymantis. This was the first instance of the genus Platymantis being subsumed into a subgenus of Rana. Keys were provided to substantiate the taxonomic claims. Van Kampen reinstated the species Discodeles opisthodon and D. ventricosa (not ventricosus) although he conceded that these species may be D. bufoniformis (after Barbour, 1921) and D. opisthodon (after Boulenger, 1918a) respectively. In 1927, Ahl reviewed Sternfeld’s (1918) work on Hylella solomonis and Van Kampen’s (1923) diagnosis of Hyla nigropunctata, both reported from the Solomon Islands. After viewing cotypes he concluded that the frogs were not Hylella nor Hyla and did not belong in the family Hylidae. The specimens were representatives of the family Ranidae and were related to Rappia (Ahl, 1927). These species both had webbed feet and hands and Ahl’s assertion was that they were not used for swimming but for increased adhesion similar to that seen in the gecko, Palmatogecko. Hence he renamed them as Palmatorappia solomonis. He provided a more detailed taxonomic description of the new genus and existing species, than had either of the two previous authors. Kinghorn's 1928 publication on the “Herpetology of the Solomon Islands” was written to

13 “assemble and modify previous descriptions, to republish old and new figures, and to compile keys to the species, so that future workers will have a complete reference to the Solomon Island herpetofauna” (Kinghorn, 1928:123).

Material described came from the islands of Ysabel and Guadalcanal. Kinghorn began with keys to the Batrachia and Ranidae followed by detailed taxonomic descriptions of genera and species. Unlike Van Kampen (1923), Kinghorn retained the synonymy of Discodeles opisthodon within D. bufoniformis, and recognised Platymantis as a genus not a subgenus. He also defined a new genus and species of frog with webbed hands and feet: Hypsirana heffernani, unaware of Ahl's 1927 work describing Palmatorappia solomonis. He continued to recognise the separate familial status of Ceratobatrachus guentheri placing it in the Ceratobatrachidae, thus not following the work of Barbour (1921). Kinghorn’s work did not add to the Solomon Island amphibian fauna but it did extend the ranges of the species within the islands. Mertens (1929) re-examined Sternfeld’s (1918) work on Solomon Island ranids, in particular the specimens of Chaperina friedericii and Sphenophryne wolfi. His observations led him to conclude that both were of the same species just at different age stages and that they did not belong in the family Brevicipitidae but the Ranidae. In fact they were congeneric with Batrachylodes (Boulenger, 1887). Due to the poor preservation of the specimens Mertens (1929) found it difficult to decide whether the two species were identical to the, then, only known species of Batrachylodes, B. vertebralis. Rather than describe a new species based on poor material Mertens referred them both to Batrachylodes vertebralis. Mertens (1929) also recognised that Sternfeld’s (1918) Hylella solomonis from Buka Island was actually Ahl’s (1927) Palmatorappia solomonis and that Kinghorn (1928) had made an error in erecting Hypsirana heffernani as this taxon also was Palmatorappia solomonis. To recognise the uniqueness of the south-east Asian ranids, Noble (1931) raised all of the taxa from the Solomon Islands to generic status and placed them into a different subfamily to Rana, the Cornuferinae. Burt and Burt (1932) reversed Noble's generic status for all Solomon Island ranids when they compiled a list of Pacific amphibians, collected during the Whitney South Seas Expedition. They reverted back to Boulenger's (1884, 1920)

14 and Van Kampen's (1923) subgenera, placing species of Discodeles and Platymantis within Rana. Schmidt (1932:178) when examining collections of ranids from the islands of Ugi, Tulagi, Malaita, Ysabel, Kulambangra and New Georgia stated: “the interiors and higher altitudes of the larger islands are still very little known”. He extended the distribution of known ranids and provided information about living animals including call data. He noted that there was considerable variation in the calls of Ceratobatrachus guentheri. He described a new species, Platymantis weberi, and distinguished it from P. solomonis as having a more acutely shaped snout, a rugose upper eyelid, numerous dorsal folds and smaller body size. Schmidt (1932) placed Boulenger's (1888) Cornufer corrugatus and Van Kampen's (1923) Rana rugata also in the synonymy of Platymantis weberi. He also found two specimens of Discodeles bufoniformis which corresponded to the defunct D. opisthodon but was discouraged from recognising D. opisthodon as a distinct taxon because of the

“lack of material, and by the conviction that such close allied forms require discrimination by field study together with a check on sexual differences which can only by supplied by mated pairs” (Schmidt 1932:181).

Hediger (1934) recorded the herpetology and zoogeography from New Britain to the north of the Solomon Islands. He diagnosed a new subspecies of Discodeles bufoniformis: D. b. cognatus, describing it from a single female specimen. He differentiated it from the nominal form by its smooth skin lacking warts and a strongly projecting inner metatarsal tubercle. This subspecies differed from D. ventricosa (= D. opisthodon Barbour, 1921; = D. bufoniformis Van Kampen, 1923) by the arrangement of vomerine teeth, length of fingers 1 and 2, length of hind limb, and amount of toe webbing. Based on the uncertainty of the classification of species within Discodeles, Hediger (1934) noted that D. bufoniformis and D. guppyi could be distinguished from each other, and hence his reason for the erection of a subspecies for his specimen. He also changed the nomenclature of D. ventricosus. Originally named Rana ventricosus, the gender was changed by Van Kampen (1923) to Rana (Discodeles) ventricosa. Hediger (1934) realised that this latter name was preoccupied by an Indian species, and hence was a primary homonym. He renamed ventricosa as Rana (Discodeles) vogtii after its author, Vogt. Hediger (1934) also found five examples of frogs that

15 he could not distinguish between Rana papua and Rana kreffti. These frogs shared characters of both species and so he followed the advice of Brongersma and placed them within Rana papua as a subspecies: kreffti. Loveridge (1948) resurrected Werner's (1894) Rana novae-britanniae from the synonymy of R. kreffti (Boulenger 1918a, 1920; Roux, 1918; Van Kampen 1923) stating:

“…the white-bellied Rana novaebritanniae is perfectly distinct from the mottled- bellied kreffti, and its uniformly white underside appears to separate it also from Rana p. papua'” (Loveridge 1948:413).

However, he did not recognise its specific status placing it as a subspecies of Rana papua, as Rana papua novaebritanniae, following the format of Hediger (1934) thus indicating the intimacy of the New Britain and New Guinean species. Brown and Myers (1949a) described Cornufer neckeri from a collection of frogs from the Whitney South Seas Expedition. Earlier, Burt and Burt (1932) had placed 22 specimens from this voyage into the only known large-disced species on the Solomon Islands, Cornufer guppyi, however Brown and Myers (1949a) demonstrated that these specimens belonged to a new species: Cornufer neckeri. They suggested a possible close relationship with C. guppyi and C. vitiensis from Fiji (formerly, C. dorsalis, Boulenger (1887)). Cornufer neckeri had a more projecting snout, a less angular canthus rostralis and wider subarticular tubercles on the fingers than C. vitiensis and a narrower head than C. guppyi. Brown and Myers gave a diagnostic account of the finger pads of these species and provided a key to the Cornufer of the Solomon Islands and Fiji. A new species of Platymantis, P. myersi also collected from the Whitney South Seas Expedition was described by Brown (1949). He related this new species to the Fijian species, P. vitianus and P. unilineatus (although doubting the validity of the latter) and not to the two species from the Solomon Islands, P. solomonis and P. weberi. Batrachylodes remained monotypic until Brown and Myers (1949b) described B. trossulus. They agreed with Merten’s (1929) synonymy of Sternfeld's (1918) Chaperina friedericii and Sphenophryne wolfi with B. vertebralis, and described a second species of Batrachylodes: B. trossulus, collected by Necker, of which there were six specimens. This was a small species (the largest female being 20 mm),

16 and the males had small white nuptial tubercles on the underside of the chin. The species had a distinctive colour pattern comprising a black band on the lateral surface of the head and body. Brown and Myers (1949b) synonymised Burt and Burts’ (1932) Rana rugata with Batrachylodes trossulus which differed from B. vertebralis in not having broadly-dilated finger discs. Tanner (1951) erroneously placed Palmatorappia solomonis Ahl, 1927, within the Hylidae, and recognised Kinghorn's (1928) Hypsirana heffernani within the Ranidae. Following Noble (1931), Brown (1952) elevated Discodeles to generic status and agreed that it was closely related to Platymantis. Because Noble (1931) had implied that all species of Discodeles developed directly, Brown recommended that all the Asiatic species of Rana placed in the subgenus Discodeles by Boulenger (1920) should be removed as they did not share this character. Brown (1952) showed that Discodeles opisthodon had shorter hind limbs and different ratios of head width:tibia length and tibia length:snout-vent length to D. bufoniformis. He thus reversed Barbour’s (1921) synonymy and erected D. opisthodon to full species. Barbour also reexamined Hediger's (1934) Discodeles bufoniformis cognatus and Vogt's (1912) unique specimen of D. ventricosus concluding that they were also D. opisthodon, thus recognising three species: bufoniformis, opisthodon and guppyi within Discodeles. Brown reinforced Ahl’s (1927) notion that Palmatorappia solomonis was a member of the Ranidae not of the Hylidae as the omosternum was forked at the base. Brown (1952) recognised four species of Platymantis distinguishing them from Cornufer as

“ranids possessing small or moderate discs on the tips of toes and fingers…; a distinct ventral pad … on toes … and fingers, separated from the dorsal part by crescentic groove…, toes free…” (Brown 1952:45-46).

He also described a new species, Platymantis aculeodactylus and reassessed P. weberi as a subspecies of P. papuensis. This latter decision was based on the fact that Schmidt (1932) when describing P. weberi compared his specimens with P. corrugatus (= P. corrugatus papuensis Loveridge (1948)), citing differences in the tarsal fold, folds on the dorsum and rugosity of the solar area. When Brown (1952) compared P. weberi with P. papuensis from New Guinea he found that they "probably represented geographically isolated and subspecifically differentiated populations of the same species" (Brown, 1952:51), differing only by

17 snout-vent length. Brown (1952) also followed Hediger (1934) in recognising the Solomon Island species of Rana as two subspecies of Rana papua: Rana papua kreffti which occupied only the extreme south-western islands of the San Cristobal group and Rana papua novaebritanniae which occupied the northernmost islands plus New Britain. In 1960, Zweifel extended the known distribution of Discodeles guppyi. In his examination of frogs collected during the 1958-1959 Gilliard New Britain Expedition, he found six specimens that correlated with the Solomon Island species D. guppyi. The specimens possessed external vocal sacs (lacking in D. bufoniformis and D. opisthodon), a narrow head and long hind limbs. He also synonymised D. vogti Hediger, 1934 from the Admiralty Islands with D. guppyi. Nevertheless he could not demonstrate that the other species in New Britain (and identified as Discodeles bufoniformis cognata by Hediger (1934)) were also D. guppyi. Hediger (1934) had given no measurements, but the specimen did have long legs eliminating it from being bufoniformis as suggested by Brown (1952). The vomerine teeth did not extend beyond the inner angles of the choanae, a character used by Brown (1952) to distinguish guppyi from opisthodon. Zweifel (1960) thought that the species bufoniformis cognata was inadequately described by Hediger (1934) and because there was nothing to distinguish it from D. guppyi he placed it in synonymy with Discodeles guppyi and not D. opisthodon as suggested by Brown (1952). In 1965, Brown synonymised Platymantis with Cornufer. Brown agreed with Inger’s (1954) work on Philippine ranids that the distinction of two genera based on one character (that of digital pad size) was not warranted, especially when a wider range of species was considered and taken into account. In addition, the species of these genera did not seem to differ ecologically. Brown placed all of the previously assigned species of Platymantis into Cornufer. Thus the known Solomon Island species became: Cornufer aculeodactylus, C. guppyi, C. myersi, C. neckeri, C. papuensis weberi and C. solomonis. He went on to describe three new species from the Solomon Islands: C. acrochordus, C. macrops, C. parkeri bukaensis, and C. parkeri parkeri, thus increasing the known number of taxa from six to nine. Cornufer parkeri bukaensis and C. parkeri parkeri Brown regarded as geographic subspecies of one polytypic species. They were closely related but were based on a small sample size. If the population had been overlapping then

18 Brown (1965) would have been more confident in their specific status. Brown (1965) presented a key to the species of Cornufer of the Solomon Islands. Recognising only one genus resulted in wider distributions which in turn had implications for ancestry, leading Brown to suggest that:

“Cornufer appears to represent a relict, peripheral group of ranid frogs with the greatest number of species at present fringing Philippine and Solomon archipelagos and a smaller remnant in New Guinea and related islands, as well as Palau and Fiji islands in the outer Pacific” (Brown 1965:3).

Zweifel (1966, 1967) examined the type of Cornufer: C. unicolor described by Tschudi (1838:28). On examining the original specimen he found it to be not a ranid but a leptodactylid and more specifically Eleutherodactylus inoptatus. Cornufer Tschudi, 1838, antedated Eleutherodactylus Dumeril and Bibron, 1841, and under the Law of Priority the former should replace the latter, but Zweifel (1960) suggested that this would lead to taxonomic confusion as over 200 species were currently recognised in Eleutherodactylus. He proposed suppressing the name Cornufer as use of this name for the ranid frogs had not yet reached stability, having only been proposed a year earlier by Brown (1965). Hence Zweifel recommend that Platymantis be used instead and therefore moved all species from Cornufer to Platymantis. Brown and Tyler (1968) described a new subspecies: Platymantis papuensis schmidti from material collected from New Britain. This then identified three geographically isolated sibling populations of Platymantis papuensis: New Guinea (P. p. papuensis), New Britain (P. p. schmidti) and the Solomon Islands (P. p. weberi) implying some degree of relationship. Another new species of Discodeles, D. malukuna was described by Brown and Webster (1969). This species was found in the mountains of Guadalcanal and was identified as having extremely reduced webbing between the toes, lack of warty tubercles on the dorsum and urn-shaped patterns formed by a pair of folds on the anterior part of dorsum. It was also relatively small in body size, compared with congeners (with males 43 – 58 mm in snout-vent length). Based on the degree of toe webbing, Brown and Webster (1969) regarded this species as an intermediate form between Discodeles and Platymantis implying a close relationship between the two.

19 In re-examining the known species of Batrachylodes: B. vertebralis and B. trossulus, and by examining their own recent extensive collection, Brown and Parker (1970) noticed that the formerly described Sphenophryne wolfi described by Sternfeld (1918) and synonymised with B. vertebralis by Mertens (1929) was in fact a distinct species, B. wolfi. They then increased the currently known species from three to eight with the descriptions of five new species. These were B. minutus, B. mediodiscus, B. gigas, B. elegans and B. montanus. All were endemic to the Solomon Islands but found widely throughout the island chain. Brown and Parker (1970) arbitrarily divided the genus into two groups: those with broadly dilated finger discs and those with finger discs moderate or not dilated. In 1978, the International Commission of Zoological Nomenclature accepted the work of Zweifel (1967) in suppressing the genus Cornufer and giving Platymantis precedence over it. Platymantis vitiensis became the type species. Using multivariate analyses and supported by call data, Menzies (1982a, 1982b) redefined the Platymantis papuensis subspecies complex. He found that the three island subspecies: papuensis (New Guinea), schmidti (New Britain) and weberi (Solomon Islands) were all valid distinct species and he elevated them to this rank. He also found that P. weberi was ecologically distinct from the other two species. Using information gained from call data, Menzies (1982b) postulated the phylogenetic relationships of species of Platymantis from the New Guinean region. By assuming that the most highly structured call was the most advanced, Menzies (1982b) indicated that P. solomonis from the Solomon Islands was the most primitive platymantid in the region. Menzies could derive the call of P. magnus (New Ireland) from P. solomonis using the reduced first pulsed syllable heard in the former as evidence. Although the two species, P. solomonis (Solomon Islands) and P. magnus (New Ireland) were similar in morphology and call their allopatric distribution maintained their species distinctness. The calls of P. schmidti (New Britain) and P. papuensis (New Guinea) showed further reduction of the pulsed syllable. The call of another Solomon Island species, P. neckeri, was very different to all the others in lacking the first pulsed syllable, leading Menzies (1982b) to suggest the call had been derived from a different evolutionary pathway. This species had an arboreal habit unlike the other four species and Menzies (1982b) suggested that this may be the impetus for differing calls, the arboreal species needing less recognition than the terrestrial forms.

20 Discodeles ventricosus was used to replace Rana ventricosus (Vogt, 1912) when Discodeles was recognised as a full genus by Brown (1952). However, according to Dubois (1986) this was a junior primary homonym as the name was preoccupied by Rana ventricosa Linneaus, 1758. The Code (Art. 52) states that junior primary homonyms are permanently invalid and Dubois (1986) regarded the valid name of this species as Discodeles vogtii (Hediger, 1934), and ignored the synonymy by Brown (1952). Menzies (1987) revised the Papuan species of Rana. He used call data and the multivariate statistical test, discriminant function analysis, to support his findings. His synopsis included all species of Rana distributed between the Indonesian Islands of Timor and Seram to the Solomon Islands; this area included the mainland of New Guinea and its eastern islands of New Ireland and New Britain. Prior to this analysis, Menzies (1987) recognised 22 species of Rana including New Britain’s R. novaebritanniae and the Solomon’s R. kreffti. Results from the cluster analysis failed to identify R. novaebritanniae. Menzies (1987) noted that the holotype of the R. novaebritanniae was in poor condition and that the exact locality and the collector were unknown. He found that the specimens attributed to R. novaebritanniae clustered closely with R. daemeli, the species found on New Guinea and northern Australia. This interpretation was supported by call data from Talasea on northern New Britain which were identical with the New Guinea mainland voice recordings. On the basis of this evidence Menzies synonymised R. novaebritanniae with R. daemeli and concluded that the latter was the only species to be found on New Britain. As for the Solomon Island species, R. kreffti, Menzies (1987) found that the female representatives in his study formed a wide cluster in the analysis, possibly indicating more than one species. However the female syntype fell within the cluster which satisfied Menzies of the species status. Of the four male representatives one fell far outside the cluster; this was a syntype of R. kreffti. Confounding this Menzies had obtained two different call types from the Solomon Islands. One was identical to that heard in New Ireland, whilst the other from Bougainville was different. Menzies (1987) tentatively suggested that there was at least one other species of Rana in the Solomon Islands but, because he did not have specimen data to support his call data, he did not name another species nor did he indicate whether this ‘other’ species was Brown’s (1952) R. novaebritanniae. Menzies' (1987)

21 findings led him to recognise a single species from the Solomon Islands, R. kreffti and a single species from New Britain, the wide spread R. daemeli. He placed both within the Rana papua complex.

2.2 Taxonomy of the Solomon Island ranids: Subfamily accounts According to Frost (2002), the Solomon Island ranids are currently placed within the large subfamily, the Raninae. This higher taxonomic placement has been challenged on a number of occasions. Noble (1931) recognised the distinctness of the Solomon Island ranids by placing them in a separate subfamily, the Cornuferinae. This new subfamily comprised the following ten genera: Hylarana, Micrixalus, Staurois, Platymantis, Cornufer, Discodeles, Ceratobatrachus, Batrachylodes, Palmatorappia, Simomantis and extended from “southern China, the Philippines, the Fijis and Solomons, westward across New Guinea, and the Indo-Australian Archipelago to India” (Noble, 1931:521). It represented a uniform group possessing digital tips more or less dilated, with a groove around the edge of the ventral surface, indicating some form of friction pad. The group had arisen from Rana although the definition of some species was hard to define as they appeared to grade into one another (Noble, 1931). According to Noble (1931) the Solomon Island genera parallelled Staurois in the possession of a partial or complete finger disc, however, they differed from Staurois in that the omosternum forked posteriorly. The least specialised was Platymantis, which had only the lateral groove on the finger disc as seen in Micrixalus. Despite this evidence Noble (1931) suggested that Platymantis was more likely to have arisen directly from Rana not Micrixalus as its distribution ranged from the Philippines, Halmahera, Kei Island, New Guinea, New Britain, the Solomon Islands and Fiji. Noble (1931) reported that Discodeles differed from Platymantis in that the central area of the tongue exhibited obtuse papillae and the toes were extensively webbed in the former. The genus Cornufer had much larger discs and the friction pads were complete as seen in Staurois and Polypedates but Noble did not consider Cornufer to be closely related to either of these genera due to its slightly webbed toes, forked omosternum and large nasals. Cornufer was reported from Burma, the Philippines, Borneo, Solomons and Fiji.

22 Noble (1931:523) agreed with Barbour (1921) by not placing Ceratobatrachus in a separate family, its teeth being only “excess bony growths of the lower jawbones”, stating that:

“it is obviously related to Platymantis solomonis … both [are] large-headed species with small discs and short webs between the toes. Their shoulder girdles are essentially alike” Noble (1931:523).

Noble (1931) also recognised that whilst Batrachylodes and Palmatorappia were similar in that they lacked vomerine teeth, they differed in that species of Batrachylodes lacked webbing, whereas Palmatorappia had extensively webbed fingers and toes and a forked omosternum. Laurent (1951) considered the subfamily Raninae to be an archaic group due to its vast distribution. He recognised Noble's (1931) subfamily Cornuferinae but unlike Noble he placed the now recognised genus, Hylarana, into the Cornuferinae. This gave the Cornuferinae, whose origin was likely from tropical Asia, African representatives. Savage (1973) continued with Noble’s (1931) concept of a separate subfamilial status for Solomon Island frogs (and others) by placing them in the Platymantinae (converted from the Cornuferinae). Ranid frogs within this new subfamily were distributed in warm temperate regions from western China through tropical Malaya, Indo-Malayan islands, New Guinea to northern Australia and the Solomon, Palau, and Fiji Islands. He regarded the Ranidae as “clearly African in origin” (p. 425) that had radiated into tropical Eurasia by the Eocene. The Platymantinae were a highly derived group descended from non-African ranines that had invaded the Indian subcontinent by the Oligocene-Miocene. Dubois (1975) commented on the phylogeny of the Ranidae in his paper describing a new subgenus and species of Rana. He discussed the ideas of Boulenger (1918a, 1918b, 1920) that Discodeles was a subgenus of Rana, that Platymantis was a distinct genus and that both were descended from the Rana grunniens group. Using characters based on egg size and number, mode of development and skeletal features such as forked omosternum and large nasals, Dubois recognised the uniqueness of the Solomon Island ranids. He mentioned Noble’s (1931) idea of placing them in a separate subfamily, the Cornuferinae, which in turn had been followed by Laurent (1951) and Savage (1973). Dubois (1975) whilst agreeing that this separation made zoogeographical sense

23 disagreed that it made systematic sense. Using examples from Hylarana he could not fathom how the Asian species, Rana temporalis, could be conspecific with the African species, Rana albolabris. If this scenario was correct it would make the subgenus Hylarana heterogenous and artificial. He referred to the work of Deckert (1938) which indicated close relationships occurred between groups of species in Asia. Deckert (1938) like Boulenger (1918a), concentrated on skeletal characters such as the pectoral girdle, including omosternum and size of the nasals. Species of Platymantis, Discodeles, Palmatorappia and Batrachylodes were united by having an arciferal pectoral girdle, a forked omosternum and large, wide, joined nasals. Dubois rejected the use of a separate subfamily for these species (and others e.g. Dicroglossus) reverting to Boulenger’s (1918a, 1920) concept of subgenera for the Ranidae. In his paper on nomenclatural changes at the family, subfamily, generic and subgeneric level within the Ranoidea, Dubois (1981) was again reluctant to accept a division of the Raninae to accommodate Noble’s (1931) subfamily Cornuferinae and regarded the Cornuferinae (sensu Noble (1931)) as heterogeneous and artificial (Dubois, 1975) stating the close affinities between Discodeles and Platymantis with Amolops, Hylarana and Staurois as evidence. He again preferred the subgenera of Boulenger (1918a, 1920) for the Raninae. However, following the work of Laurent (1981) in the allocation of tribes for dividing the Rhacophoridae, Dubois suggested recognising three tribes for the Raninae: the Ranini Gray, 1825; the Limnodytini Fitzinger, 1843 and the Ceratobatrachini Boulenger, 1884. Yet Dubois (1981) stressed caution in accepting these new subdivisions as the Asiatic species of Rana were not as easily divided into lineages as the African species of Rana. Dubois (1981) strongly advised the use of Boulenger’s (1920) concept of subgenera for Rana until further phylogenetic work was performed. Zweifel and Tyler (1982) recognised the distinctness of the Solomon Island ranids by following the classification of Savage (1973) and recognising the separate subfamily, the Platymantinae. They did this on the assumption that all platymantines have the same mode of development, that is direct development. Dubois (1983, 1984) again refuted the idea of a separate subfamily for the Solomon Island ranids, subsuming the Platymantinae into the already existing Raninae for all Asiatic ranids.

24 Laurent (1986) also rejected Noble’s (1931) classification of Cornuferinae (= Platymantinae of Savage, 1973), although he did concede that with further investigations this may be in reality a phylogenetic group. He did not consider Noble’s (1931) assertion that Hylarana belonged within the Cornuferinae as this genus was more closely related to the genus Rana than any of the other genera that comprised the Cornuferinae. On this basis he erected two tribes within the subfamily Raninae: the Ranini and the Platymantini and distinguished between these tribes based on the following characters: dilation of the digital extremities, the presence or absence of the circummarginal groove, and the shape of the terminal phalanx. He provided a key to the tribes and placed all the Solomon Island ranid genera within the Platymantini. Dubois (1986) also used tribes within his classification of the Ranidae. He based his classification not on cladistic principles but on genetic, morphological and ecological criteria and did not recognise holophyletic (= monophyly as used by cladists) groupings but homophyletic groupings (= both monophyly and paraphyly as used by synthesists). In reviewing the classification of Anurans, Dubois (1986) gave particular importance to those characters extracted from mode of reproduction and life history, stating that such characters demarcated genera and represented clear discontinuities at the genetic level. Such characters had remained little employed in phylogenetic reconstructions (Dubois, 1986; Duellman, 1985) and constituted real evolutionary jumps according to Dubois (1986). In order to promote his concept of a genus he had to dismantle many existing Amphibian genera (Dubois, 1988). Dubois’ classification of the Superfamily Ranoidae recognised five families, one of which was the Ranidae. Within this family he recognised four subfamilies: Raninae, Phrynobatrachinae, Mantellinae and Rhacophorinae. Within the Raninae, Dubois then recognised tribes. Again he did not recognise the subfamily Cornuferinae of Noble (1931) nor the tribes of Laurent (1986) as these classifications were based on the expansion of digital pads and the presence of adhesive pads and not on supposedly more conservative characters such as the mode of reproduction. Dubois (1986) considered these former groupings to be polyphyletic. In reference to Laurent’s (1986) work, Dubois stated that although Laurent, like Dubois recognised one subfamily, the Raninae, Laurent’s recognition of two tribes, the Ranini and Platymantini repeated the classification error of Noble (1931) but at a lower taxonomic level. Dubois argued

25 that the work done by researchers on African Rana, in which most of Boulenger’s (1918a, 1920) subgenera had been converted to genera, could not be readily applied to forms from southeast Asia. As a result Dubois proposed a new classification. He subdivided the Raninae into six tribes each representing a different phylogenetic lineage. In order to do this however, the genus Rana (considered heterogeneous by Dubois) had to be re-evaluated. In addition, Dubois (1986) claimed his classification was constructed using the group as a whole and not based on narrow geographic regions. The six tribes defined by Dubois (1986) within the Raninae were Ranini, Ptychadenini, Tompternini, Dicroglossini, Pyxicephalini, and Ranixalini. Members of the Solomon Island ranids were placed in the Ranini and Dicroglossini. The genus Batrachylodes was placed in the tribe Ranini unlike the rest of the Solomon Island ranids. Members of the tribe Ranini had a firmisternal pectoral girdle, an unforked or slightly forked omosternum and reduced nasals that were separate from one another. Dubois placed Batrachylodes in this tribe based on Brown’s observations of the omosternum and his suggestion that the genus could be derived from a ‘hylaranid stock’. There is no evidence that Dubois examined specimens. The Asiatic members of the genus Rana were also placed in the Ranini but were distributed amongst five sub-genera. The tribe Dicroglossini included the genera Ceratobatrachus, Discodeles, Palmatorappia and Platymantis. Members of this tribe had a firmisternal pectoral girdle, an omosternum moderately or strongly forked at the base and very large nasals, that were in general in contact with one another and with the frontoparietals. This group partially corresponded to Noble’s (1931) Cornuferinae and Laurent’s (1986) tribe, the Platymantini. Although Dubois considered both these latter groupings to be artificial and heterogenous if certain taxa were removed e.g. Hylarana, Amolops and Staurois they could well be a homogenous lineage. As a result of Dubois placing Rana in another tribe, many species currently maintained in this genus, but now placed within the Dicroglossini, were elevated to generic status, as they could not remain in Rana because this genus would then be polyphyletic. The new genera to accommodate such species were Limnonectes, and Ingerana. In justifying the name Dicroglossini for the tribe, Dubois considered the names Ceratobatrachidae Boulenger, 1884, Cornuferinae Noble, 1931 and Platymantinae Savage, 1973 as synonyms.

26 In 1989, Ohler and Dubois further studied the tribes Ranini and Dicroglossini, using characters pertaining to finger discs. Previous classifications of Anurans had considered the presence of fingers and toes with rounded or blunt discs and lacking differentiated structures as plesiomorphic, whereas those species with finger and toes expanded and differentiated into adhesive pads were considered more derived. However, Ohler and Dubois (1989) noticed that when examined in detail these adhesive pads were alike in histology and cytology regardless of phylogenetic origin and hence were a result of convergence. Ohler and Dubois (1989) noted that within the two tribes, Ranini and Dicroglossini (Dubois, 1986), there were species with and without adhesive pads and they found that both tribes exhibited morphoclines in adhesive pad morphology, ranging from no adhesive pads to incomplete and complete adhesive pads. If the absence of adhesive pads was a plesiomorphic trait and if Dubois’ (1986) recent tribes were valid and corresponded to different lineages, then they hypothesised that the presence of adhesive pads found in both tribes arose independently and were not homologous. A superficial examination of those species with complete adhesive pad morphology showed similar morphologies in both tribes, thus refuting their hypothesis. It wasn’t until Ohler and Dubois (1989) studied the morphology of frogs with incomplete adhesive pads that they recognised the existence of two separate morphoclines, from a blunt to a complete adhesive pad on finger/toe. It was the position of the grooves in incomplete pads that differed, being latero-terminal in the Ranini and dorso- terminal in the Dicroglossini even though both tribes had species with similar morphologies of complete adhesive pad and extremely dilated discs with grooves in the marginal position. According to Ohler and Dubois (1989) such evidence strengthened the phylogenetic hypotheses separating the two tribes and confirmed that species with complete adhesive pads in the different tribes, despite their resemblance were not equivalent but the result of convergence. Only the ventral part of the disc participated in forming the sole in the Ranini but it was the ventral and dorsal parts that contributed to the disc in the Dicroglossini. Due to this demonstration of convergence in both tribes the simple character of presence and absence of adhesive pads could not be used as a character for phylogenetic analysis.

27 In 1992, Dubois revised his classification of the Ranidae. He added characters pertaining to the tadpole, especially information regarding denticulate rows, and also merged in his data on adhesive pads (Ohler and Dubois, 1989). He was still unconvinced by the localised attempts at the classification of the ranids, and maintained his global approach to the superfamily Ranoidae, now classified under a new rank, Epifamily. In this classification Dubois raised his tribe Discroglossini to that of a subfamily, the Dicroglossinae. Within this subfamily he erected four new tribes: Ceratobatrachini, Conrauini, Dicroglossini and Limnonectini. Ceratobatrachini Boulenger, 1884 was diagnosed as having direct development, without a free tadpole stage and without a lateral line in the adult. There were six genera in this tribe: Ceratobatrachus, Discodeles, Ingerana, Palmatorappia, Platymantis and Taylorana. Dubois retained the subfamily Raninae in this present classification in which he recognised two tribes: Ranini and Paini. The tribe Ranini was diagnosed as Raninae in which males lack big black folded thorns on chest, forearms, and digits I to III, no thorny projections around the anus and a generalised tadpole. The genus Batrachylodes was again placed within this tribe along with five other genera, one of which was Rana. Again because of his globalised approach to the systematics of the Ranoidea, African and Asiatic forms were members of the same phylogenetic lineage. The taxonomic and nomenclatural decisions reported here have been undertaken in the absence of robust phylogenetic underpinning. Questions of phylogenetic relationships of the ranid frogs of the Solomon Islands are addressed in this thesis together with consideration of their position within the tribes proposed by Dubois (1992) and their zoogeographical origins. Brown (1996) maintained the subfamily Platymantinae but restricted it to the genera: Ceratobatrachus, Discodeles, Palmatorappia and Platymantis. He placed Batrachylodes and Rana kreffti in the Raninae. In regards to Batrachylodes Brown (1996) stated that its relationships to other ranines is unknown. In the absence of any detailed phylogenetic analysis, Frost (2002) maintains conservatism placing all species of Solomon Island ranids in the one subfamily Raninae.

28 2.3 Mode of Life History: a uniting feature? Although the ranid frogs of the Solomon Islands are very diverse in their external morphology there is one assumed feature that unites these frogs, a direct developing mode of reproduction. This mode of reproduction has only been seen in a few species of ranids frogs from these islands but is extrapolated from the presence of large, unpigmented eggs which are few in number. The first recorded instance of the direct developing habit in ranid species on the Solomon Islands was by Boulenger (1886a) who published the field notes of Guppy. These notes detailed the larval development of Discodeles opisthodon. Boulenger (1886a) gave measurements of the ova (6 to 10 mm) and noted the absence of a tail and gills. Boulenger (1886b) further described the direct development seen in Discodeles opisthodon, in a key published in The Annals and Magazine of Natural History. He placed D. opisthodon in the key under the following conditions:

“II. The yolk-sac is very large, and the young undergoes the whole or part of the metamorphosis within the egg; at any rate the larva does not assume an independent existence until after the loss of the external gills. A. The ova are deposited in damp situations or on leaves, and the embryo leaves the egg in the perfect air-breathing form. Rana opisthodon, Blgr. Hylodes martinicensis D. & B.” (Boulenger, 1886b:464).

Vogt (1912) noted that a gravid specimen of Discodeles ventricosus had eggs that were not as large as those found in D. opisthodon, leading him to surmise that direct development did not occur in this species. However, when he examined the eggs of Platymantis solomonis he found that they were large in size and few in number and following Boulenger (1884) confirmed direct development for this species. The first attempt at grouping the frogs of the Solomon Islands on the basis of their reproductive habit was that by Noble (1925). In this work he described the direct development of Discodeles opisthodon and compared it to that of a known direct-developing frog, Cornufer guentheri, from the Philippines. Noble (1925) described the eggs and development of these frogs and ascertained that they were very similar. From inference, he recorded that the assumed respiratory folds of D. opisthodon were the same structures found in C. guentheri, that is, foldings of the somatopleure for rapid absorption and readjustment of the yolk just before hatching.

29 The importance of reproductive habit reflecting common ancestry was again discussed by Noble (1927) who noted that in the previous ten years prior to his publication, studies on the Amphibia had increased and that within these studies more data on larvae and life history had been documented. He suggested that these data were useful in discovering the broader relationships of forms and gave important clues to the evolution of various groups. He defined ‘mode of life history’ as “the breeding habits of the adults and, the development and larval habits of the offspring” (Noble, 1927:32). In discussing the Amphibia, Noble concentrated on the Ranidae, lamenting that not much was known beyond the genus Rana. He again discussed his previous work (Noble, 1925) on the similarity of development between Discodeles opisthodon and Cornufer guentheri. Similar structures (e.g. abdominal folding) occurring in two species with identical breeding habits, inferred a close affinity between the two. He went on to say that Cornufer was very closely allied to Rana (although it is unsure whether he was referring to Discodeles, as he was using Boulenger’s (1886a, 1920) classification of Rana). One interesting statement was:

“Cornufer very probably gave rise to a number of different groups but the life histories of none of these are known” (Noble, 1927:109).

Noble (1927) suggested that Ceratobatrachus may have a similar mode of development, and had directly arisen from Cornufer. Evidence of a difference in larval development between frogs of the genus Platymantis and Discodeles came from a paper by Atoda (1950). He described the development of an as yet unidentified direct-developing frog from the island of Palau (later identified as Platymantis pelewensis). He compared the development to Discodeles opisthodon, the major difference being that the larvae of his material develop with a tail. In 1992, Schmidt confirmed direct development in Ceratobatrachus guentheri. Although bred in captivity and subjected to poor conditions, i.e. fungal infections, Schmidt was able to witness the development of frogs within the eggs and hatching after 32 days. In all cases of egg laying Schmidt observed large white eggs about 5 mm in diameter, he counted 22 eggs per clutch in one case and then multiple clusters of eight eggs in another clutch, thus confirming that the presence of few large unpigmented eggs was evidence of direct development.

30 2.4 Zoogeography of the Solomon Islands and surrounding islands Boulenger (1888) whilst documenting the Solomon Island herpetofauna from the islands of Guadalcanal and New Georgia commented on the considerable heterogeneity between the fauna of the two extreme most islands, Faro and San Cristoval (Cristobal) in the Solomon Island chain, saying that the fauna of the former is more Papuasian, while the latter more Polynesian in origin. Barbour (1921) reviewed the herpetofauna of the Solomon Islands and reported that the collection verified the proposition of Boulenger (1888, 1890) in that the knowledge of the island’s fauna was fast approaching completion. However, little was known from the interior of the islands. The fauna of the Solomon Islands “is obviously Papuan not Australian in its origin” (Barbour, 1921:92). Barbour (1921) disagreed with Boulenger's (1888) claim of heterogeneity stating that “the fauna of the group is more homogeneous than at first appeared” (Barbour, 1921:92). He also noted that species occurring widely in the Solomon Islands were, in general, absent from the neighbouring islands of New Britain and New Ireland, suggesting a long period of isolation. In addition, the phylogenetic development of Ceratobatrachus and Batrachylodes must have been slow as they were widely distributed throughout the Solomon Islands (Barbour, 1921), also indicating a long period of isolation for the Solomon Island chain, leading Barbour to state:

“It still appears an ancient, somewhat depauperate, continental, and not an oceanic, fauna which probably has spread under rather adverse conditions'” (Barbour, 1921:92).

The type of amphibians inhabiting the islands were those that could

“disperse themselves and reproduce in areas where standing water … is practically non-existent, and where heavy rainfall and steep hills combine to form torrents which would carry off larvae rather than permit their leisurely development” (Barbour, 1921:92).

Burt and Burt (1932) stated that the reptiles and amphibians occurring in the Pacific Ocean were derived from East Indian elements, which in turn showed strong Australian-Asiatic affinities using as an example the species occurring on islands surrounding New Guinea. Such species were identical to those found on the large land mass of New Guinea, with a certain percentage becoming distinct

31 due to isolation. Burt and Burt (1932) found that the fauna of the South Seas Islands were Papuan in origin with an eastward migration and that a

“rather well-marked highway of dispersal … extends through the Solomon Islands, Banks, Santa Cruz, and New Hebrides to New Caledonia and the Loyalty Islands … and then to Fiji, … [with a] progressive and continual reduction in the number of species inhabiting the more distant areas” (Burt and Burt, 1932:478).

As the Solomon Islands were rich in endemic forms, this indicated a possible centre of differentiation. According to Burt and Burt (1932) Rana kreffti and Platymantis solomonis became restricted on their eastward journey, whereas, Batrachylodes vertebralis, Ceratobatrachus guentheri, Cornufer guppyi, Palmatorappia solomonis, Discodeles bufoniformis and D. guppyi became endemic as a result of speciation, after an apparent period of isolation (Burt and Burt 1932). Tanner (1951) discussed the herpetofauna of Guadalcanal in the Solomon Islands, in particular species found in grasslands on the northern side of the island. Despite high endemicity, Tanner (1951) thought that the Solomon Island fauna was essentially New Guinean in origin and he suggested this was due to a past connection with New Guinea. He stated that

“after the Solomon Islands landmass separated from New Guinea evidence supports the belief that the present two chains of islands developed which has contributed to the endemism [sic] of the several islands” (Tanner 1951:57).

Tanner also believed the Solomon Islands to be continental in origin. In his 1952 monograph on the Solomon Island ranids Brown stated:

“There is a great need of generic and family revisions in both the amphibian and reptilian faunas which would include all the forms known to inhabit any part of this island region and also include their Australian and Asiatic relatives” (Brown, 1952:4-5).

He acknowledged that relatively little was known of the herpetofauna from the interior of islands and thus could not agree with either Boulenger (1888) or Barbour (1921) that the Solomon Islands herpetology was well known. He suggested that the highly endemic Solomon Island ranid fauna were closely related to each other, with the exception of Batrachylodes and Rana, the latter two having descended from a single more primitive ranid stock. Yet, Brown acknowledged that more work from anatomical and embryological approaches

32 was needed in order to outline the probable lines of evolution and to reconstruct hypothetical ancestor(s). Close relationships of the ranids was borne out in their osteology and reproductive habit. Batrachylodes was possibly of hylaranid stock because of a differing omosternum yet, its reproductive habit was similar to the other genera. Brown (1952) used similarity as a measure of relatedness, allying the species of Platymantis and Cornufer in the Solomon Islands with species on other landmasses e.g.

 P. solomonis — P. vitianus (Fiji)

 P. aculeodactylus — P. cheesmanae (New Guinea)

 P. myersi — P. beauforti (New Guinea)

 P. papuensis weberi — P. papuensis papuensis (New Guinea)

 C. guppyi — C. vitiensis (Fiji) leaving the remaining species, C. neckeri as distinct from all others, being "Rhacophorus-like in appearance although lacking an intercalary cartilage". He also considered that the two species of Rana were subspecifically related to the New Guinean Rana papua. Based on these observations Brown suggested the amphibian fauna was comprised of two elements:

“…the older element … closely related, possibly diphyletic, endemic group of highly specialized ranid frogs, … Batrachylodes, Ceratobatrachus, Cornufer, Discodeles, Palmatorappia and Platymantis (in part)” and a “more recent element … include members of Hyla, Rana (Hylarana) and Platymantis” (Brown 1952:12).

Brown (1952) postulated that the older element might have its centre of origin in the Solomon Islands/Bismarck Archipelago as all but two genera were confined to this region. He gave two hypothetical explanations to support the widely dispersed ranges of Platymantis and Cornufer:

“(1) that they represent the older, more widely dispersed, possibly ancestral genera as held by Noble (1931) or (2) that they have simply been more successful in spreading through an island range” (Brown 1952:12-13).

Using the three Solomon Island species of Platymantis as examples, Brown (1952) suggested that the recent element, had closer affinities with New Guinea

33 and thus comprised late arrivals, only being able to maintain themselves in the Solomon Islands when suitable breeding habitats became available. He compared the fauna of the different island groups within the Solomon Islands and then between the Solomon Islands and the Bismarck Archipelago/New Guinea and recognised homogeneity modified by:

“(1) the northern group of islands, Bougainville and Choiseul, appear to have a richer amphibian fauna and San Cristobal in the extreme south the poorest; (2) inter-island subspecific populations are known, … and when more extensive collections become available, may be substantiated for others” (Brown 1952:13).

The poor representation of material present in San Cristobal, suggested to Brown (1952) that the southern chain was always completely isolated, lending support to the submarine plateaux having been separated by marine barriers. When examining Batrachylodes Brown (1952) indicated that Noble (1931) never viewed a specimen and had classified it as a “diminutive Cornufer lacking vomerine teeth'' (Brown 1952:24). Yet, Brown noticed that the omosternum of Batrachylodes did not support this derivation, being unforked at the base, and thus suggested a different origin possibly from a hylaranid group. Myers (1954) considered that Ceratobatrachus guentheri must have been a resident of the Solomon Islands for a very long time due to its endemicity and specialisations. He postulated that this 'common frog' of the Solomon Islands must have had ancestors in New Guinea although nothing remotely resembling it had yet been found. He regarded the ranids as once excellent island-hoppers that were now long restricted by sea-barriers. Zweifel (1960) described Platymantis gilliardi, from New Britain and compared it (and others) to the Solomon Island species of Platymantis. Using ratios of body measurements such as that of inter-narial distance to eye-naris distance, Zweifel distinguished P. gilliardi from platymantids in New Guinea and the Solomon Islands. Due to the lack of endemicity on New Britain and its poor representation of ranid species compared with New Guinea and the Solomon Islands, Zweifel (1960) did not think that land connections had ever occurred between either New Britain and New Guinea nor New Britain and the Solomon Islands, as the fauna on New Britain would have been richer. To explain the occurrence of an ecologically underpopulated island (New Britain) between two regions of more diverse and highly endemic faunas (New Guinea and the

34 Solomon Islands), Zweifel accepted that the Solomon Islands were of volcanic origin, recently deposited in geological time (citing David, 1950), and colonised by chance dispersal at an earlier date, as frogs from these islands were particularly adept at island hopping as suggested by Myers (1954). The Bismarck Archipelago including New Britain formed much later between New Guinea and the Solomon Islands, and was populated by over-water dispersal from east and west, resulting in the mixed arrangement of fauna seen in New Britain today (Myers, 1954). Brown and Tyler (1968) suggested that the New Britain ranid fauna had a closer affinity with the fauna found on the Solomon Islands rather than that of New Guinea noting that whilst there has been a relatively long period of isolation between New Britain and other island complexes, its separation had been less from the Solomon Islands. However, subspecies of Platymantis from New Guinea and the Solomon Islands exhibited greater similarity to each other than either did to the New Britain population, further supporting Zweifel's (1960) idea of the geological recency of the latter. Tyler (1972) interpreted the incredible divergence of ranids in the Solomon Islands as evidence that they represent a relictual fauna, which implied a direct relationship between the extent of morphological divergence and time. However, the worldwide distribution of the Ranidae suggested that this family entered New Guinea from the Oriental Region. If ranids have existed in the Solomon Islands for a considerable period of time, Tyler found it difficult to understand why only one species, (Rana papua) had succeeded in entering Australia and was restricted to the Cape York Peninsula. Tyler (1972) thought it more likely that on arriving on islands devoid of frogs, there was virtually an evolutionary explosion followed by rapid morphological and biological divergence associated with the exploitation and occupation of a wide variety of available niches. He was hesitant in dating the entry to the islands any earlier than the Pleistocene. Zweifel (1975) described two new species of Platymantis from New Britain and compared these new species to others in the region. He reiterated Brown and Tyler's (1968) idea that New Britain had a closer affinity to the Solomon Islands than to New Guinea. Brown and Menzies (1978) followed this trend when describing a new species of Platymantis from New Ireland. In summarising the knowledge of amphibian fauna so far from the Bismarck Archipelago, they noted that Zweifel’s (1960) notion of an impoverished region was not supported. Since

35 1960 much field work had added to the amphibian fauna increasing it from eight to twenty species (although not all ranids). Again using the distribution of Platymantis as an example, Tyler (1979) proposed an Oriental origin for ranid species in the Solomon Islands. The Philippines had to be a centre of ranoid radiation, with Tyler suggesting that Platymantis evolved there in the Late Tertiary dispersing south-eastwards into New Britain, the Solomon Islands and Fiji by rafting. Secondary radiations then occurred from within the Solomon Islands and within New Britain thus explaining the anomaly of the apparent closeness of relationships between species on different islands e.g. P. batantae (New Guinea) with P. mimicus (New Britain) and P. punctata (New Guinea) with P. myersi (Solomon Islands). This also accounted for the depauperate number of ranid species on New Guinea. Tyler (1979) consider Rana a more recent arrival to the region as the number of species reduced from west to east. Brown (1996) reassessed the biogeography of amphibians in the southwest Pacific and divided the anuran fauna into three components, (i) I, anurans resulting from colonisation of present day Asian stocks, (ii) II, anurans resulting from colonisation by present day Australian stocks and (iii) III, anurans in distinct subfamilies that are centred in islands. He included the Solomon Island ranid genera, Ceratobatrachus, Discodeles, Palmatorappia and Platymantis in Component III, placing them within the subfamily, Platymantinae. Whilst the genus Batrachylodes and the endemic species of Rana, R. kreffti were placed in Component I, in the subfamily Raninae. The presence of Rana kreffti in the Solomon Islands was as a result of secondary colonisation from New Guinea however the relationship of the eight endemic species of Batrachylodes to other ranines was unknown to Brown (1996). By implication of Brown's (1996) component groups their ancestry must be derived from an Asiatic stock. For the genera within the Platymantinae, Brown (1996) concluded that Asia must be an immediate source area, because despite their endemism the ranid family is well know throughout Asia. The Platymantinae of Brown (1996) is restricted to the Solomon and Bismarck Islands except for one genus, Platymantis, which extends to the Philippines, an arrangement already recognised by Tyler (1979) and Zweifel and Tyler (1982). Brown (1996) agrees with these

36 authors that the south east Asian ranid genera (e.g. Staurois, Ingerana) do not belong in the Platymantinae. Restricting the Platymantinae to only Solomon Island genera (including all species of Platymantis) better reflects an island with ancestry originating from southeast Asia that was colonised very early (Allison, 1996, Brown, 1996; Tyler, 1979, Zweifel and Tyler, 1982). However, Brown (1996) notes that this still does not give a satisfactory explanation for their dispersal as far as Fiji. Brown (1996) hypothesised ultimately a Gondwanan origin for these island frogs where rift blocks separated from eastern Australia prior to the separation of Australia and Gondwana. Ancestors could then have been transported on such blocks.

37 Chapter 3 Morphometrics

3.1 Introduction The statistical tests, principal component analysis and discriminant function analysis were used to explore the large multivariate data set obtained from the external morphology of the ranids of the Solomon Islands. The aim of this exercise was to examine whether groups of frogs could be identified from measurements of external morphology in the data set and whether these groups corresponded with already existing taxonomic groups. There is no definitive key for the Solomon Island ranids and hence identification of individuals was obtained by using a montage of keys provided by the following authors (Boulenger, 1884, 1886a, 1887, 1918a, 1920; Brown, 1949, 1952, 1965; Brown and Myers, 1949a, 1949b; Brown and Parker, 1970; Kinghorn, 1928; Sternfeld, 1918; Van Kampen, 1923). Because this was not an ideal situation, in that identification of species was essential for the phylogenetic analyses, it was decided to examine the data set morphometrically to reinforce the identification obtained using the keys. How do the Solomon Island ranids occupy morphological space with respect to each other? Could this be used to satisfy species identification of individuals before character selection for phylogenetic reconstructions? Principal component analysis (PCA) is a non-model-based statistical technique of involves data reduction. It reduces the dimensionality of data sets which contain a large number of variables that are interrelated. At the same time it retains as much as possible of the variation present in the original data set (Jolliffe, 1986). This is achieved by computing a new set of variables, called principal components (PCs), which are uncorrelated but ordered so that the first few components retain most of the variation present in the original variables. Conversely the last few variables account for very little variation, i.e. they identify near constant linear relationships (Jolliffe, 1986). The main use of PCA is descriptive rather than inferential, hence, it is a descriptive, dimension-reducing technique. Discriminant function analysis (DFA), like principal components analysis, involves solving for eigenvalues and eigenvectors of a particular matrix

38 (Dunteman, 1989). Where DFA differs from PCA is in its determination of one or more linear functions, measured on a number of variables from defined groups, that maximise the between groups variation relative to within group variation. The number of derived linear composites or discriminant functions equals the number of groups minus one or the number of variables, whichever is less (Dunteman, 1989). For example, for 14 variables amongst six groups there would be five discriminant functions. If there is a large number of groups and a large number of highly correlated variables then conducting a DFA on a subset of PCs derived from a PCA is recommended. This is because the PCs are uncorrelated and each one can therefore be tested independently for statistical significance by a F-ratio, using a one-way analysis of variance. Only those PCs that individually discriminate between groups are then used in the DFA analysis. This is not possible with highly correlated variables (Jolliffe, 1986; Duntemen, 1989).

3.2 Methods A total of 445 specimens from three genera representing 17 species were measured for morphometric analyses. Only the genera, Batrachylodes, Discodeles and Platymantis, each of which had three or more species, were analysed (see Appendix 1: specimens are identified by the letter m). Measurements taken are summarised in Table 3.1. Dial calipers (Mitutoyo Mfg Co., Ltd) were used to take measurements (measuring to 0.05 mm).

39 Table 3.1. Variable abbreviations and descriptions used in this study. Abbreviation Description SV snout-vent length; from tip of snout to vent TIB tibia length; length of tibia with leg flexed TAR Tarsus length; length of tarsus HL head length; from tip of snout to posterior margin of tympanum (in line with angle of jaw) HW head width; width between tympana EYD eye diameter; from anterior corner to posterior corner of eye EN eye to naris distance; from anterior corner of eye to posterior edge of external naris IN internarial distance; minimum distance between the external nares (i.e. from inner edge) TYM tympanum diameter; taken in the horizontal plane to the head IOW interorbital width; minimum distance between the eyes, taken from the inner edge of the eyelid FL3 3rd finger length; from the tip of the pad to the base of the middle metacarpal tubercle FD3 3rd finger disc width; taken from widest part of disc TL4 4th toe length; from the tip of the pad to the base of the outer metatarsal tubercle TD4 4th toe disc width; taken from widest part of disc SEX male; female

All measurements were entered into a database (Microsoft Excel™). The statistical packages SPSS (ver. 6.1, Macintosh) and SAS JMP (ver. 3.1, Macintosh) were used for data analysis. Descriptive statistics were calculated for every species. These include box plots to indicate mean, median and standard deviations (Appendix 2). T-tests were performed to identify sexual dimorphism. Multivariate analyses (principal components analysis (PCA) and discriminant function analysis (DFA)) were performed to determine species groups. Each genus containing three or more species was analysed. Females and males were firstly analysed separately and their principal components examined to see whether combining these data sets (and thus increasing sample size) could be warranted. For the PCA, correlation matrix was preferred over a covariance matrix so that all variables could be treated equally (Jolliffe, 1986). A covariance matrix would have given greater weight to larger (and hence more variable) measurements such as snout-vent length (SV) and less weight to smaller less variable measurements such as internarial distance (IN) and eye to naris distance (EN). Measurements were log transformed prior to the analysis, to increase the probability of linear relationships between variables.

40 3.2.1 How many PCs? As the aim of a principal components analysis is data reduction, choices must be made in selecting the number of principal components that best explain the variance of the original data set. According to Jolliffe (1986) and Dunteman (1989), there are various rules which are mostly ad hoc and at the discretion of the researcher. A PCA will give back as many principal components as there are original variables; this is undesirable if the aim is data reduction. Jolliffe (1986) outlined various methods of selecting the number of PCs. The “cumulative percentage of total variation” is where the researcher selects the number of PCs that account for a cumulative percentage of total variation between 80 and 90%. The required PCs are then the smallest number for which this chosen percentage is exceeded. Another method outlined by Jolliffe (1986) is the “size of variance of principal components” which is a modification of ‘Kaisers rule’, where only those PCs whose eigenvalues are less than or equal to 1 are retained. Jolliffe (1986) suggests that choosing those PCs with an eigenvalue less than or equal to 0.7 is more appropriate as much information is lost when the cut-off is at 1. Dunteman’s (1989) methods of selecting the number of PCs require further data manipulation. His “correlations of variables with principal components” involves multiplying the elements of a particular column of eigenvectors by the square root of the corresponding eigenvalue for that column. Then to determine the “proportion of variance explained” columns are added together until the number of PCs needed to obtain a value exceeding 80% are found. These PCs are then retained. This study used all the above four methods to ascertain the number of PCs required to best explain the variance of the original data.

41 3.3 Results

3.3.1 Analysis of Separate Sexes Tables 3.2 - 3.7 show the eigenvalues and eigenvectors for females and males for the genera Batrachylodes, Discodeles and Platymantis respectively. Tables 3.8 - 3.10 are summary tables of the eigenvector elements for the first three principal components taken from Tables 3.2 - 3.7 and were constructed using the format detailed by Jolliffe (1986). A ‘+’ or ‘-‘ indicates an eigenvector whose absolute value is greater than half the maximum eigenvector of the relevant PC. A ‘(+)’ or ‘(-)’ indicates an eigenvector whose absolute value is between a quarter and a half of the maximum eigenvector for the relevant PC (Jolliffe, 1986). When comparing the summary tables (Tables 3.8 - 3.10) for both female and male species of Batrachylodes, Discodeles, and Platymantis, it can be seen that for all genera the male and female data sets, have the same eigenvector loadings for PC1; and similar loadings on PC2 and PC3. This similarity in arrangement allows the data sets to be combined for each genus. Admittedly there are other variables influencing the 2nd and 3rd PC loadings but to a lesser extent. Consequently females and males were then analysed together for each genus and results are presented below.

42 Table 3.2. Eigenvalues and Eigenvectors for females of the genus Batrachylodes, for all 14 principal components. EigenValue: 11.65 0.71 0.39 0.36 0.23 0.17 0.15 0.10 0.07 0.06 0.05 0.03 0.02 0.01 Percent: 83.23 5.10 2.78 2.54 1.67 1.20 1.05 0.72 0.51 0.42 0.33 0.23 0.15 0.08 CumPercent: 83.23 88.33 91.11 93.65 95.32 96.52 97.57 98.28 98.80 99.21 99.54 99.77 99.92 100.00 Eigenvectors: SV 0.28 0.06 -0.04 -0.18 0.18 0.09 -0.03 -0.21 -0.10 -0.59 0.03 0.65 -0.14 0.06 TIB 0.28 -0.12 -0.10 0.20 -0.06 -0.17 -0.24 0.41 0.19 -0.11 0.13 -0.09 -0.19 0.69 TAR 0.28 -0.04 -0.08 0.27 -0.05 -0.24 -0.22 0.47 0.12 -0.20 -0.08 0.10 0.31 -0.59 HL 0.28 0.07 -0.05 -0.15 0.18 -0.31 -0.06 -0.07 -0.26 -0.11 -0.06 -0.46 -0.62 -0.27 HW 0.28 -0.02 -0.06 -0.29 0.20 -0.22 -0.14 -0.15 -0.42 -0.01 -0.01 -0.23 0.65 0.23 EYD 0.27 0.03 -0.15 -0.36 0.19 0.46 -0.02 -0.04 0.62 -0.10 -0.10 -0.33 0.10 -0.06 EN 0.27 0.06 -0.01 0.03 -0.56 -0.39 0.48 -0.34 0.27 -0.15 -0.08 -0.07 0.09 0.05 IN 0.26 -0.30 -0.28 0.27 0.45 0.06 0.65 0.10 -0.08 0.22 0.02 0.09 0.00 0.01 TYM 0.18 0.91 0.06 0.09 0.12 0.02 0.14 0.16 -0.00 0.20 0.14 0.07 0.03 0.06 IOW 0.24 -0.18 0.89 -0.11 0.20 -0.10 0.08 0.08 0.14 0.13 -0.00 0.07 -0.01 -0.00 FL3 0.28 -0.00 -0.12 0.02 -0.06 -0.02 -0.29 -0.18 0.04 0.51 -0.65 0.30 -0.11 0.05 FD3 0.27 -0.15 -0.14 -0.46 -0.35 0.07 -0.03 0.16 -0.09 0.40 0.53 0.21 -0.10 -0.15 TL4 0.27 -0.05 0.06 0.54 0.04 0.18 -0.31 -0.53 0.05 0.10 0.42 -0.09 0.02 -0.10 TD4 0.27 -0.01 0.18 0.15 -0.40 0.59 0.13 0.19 -0.45 -0.16 -0.22 -0.19 -0.01 0.03

43 Table 3.3. Eigenvalues and Eigenvectors for males of the genus Batrachylodes, for all 14 principal components. EigenValue: 11.19 0.72 0.56 0.33 0.30 0.19 0.15 0.15 0.11 0.09 0.07 0.06 0.04 0.03 Percent: 79.91 5.11 4.02 2.36 2.15 1.37 1.10 1.04 0.77 0.67 0.53 0.45 0.30 0.20 CumPercent: 79.91 85.02 89.04 91.41 93.56 94.95 96.05 97.09 97.86 98.52 99.05 99.50 99.80 100.00 Eigenvectors: SV 0.28 -0.12 -0.14 -0.04 0.08 -0.08 -0.16 0.06 -0.36 -0.45 0.60 -0.35 0.13 0.07 TIB 0.28 0.05 -0.19 0.11 -0.13 -0.14 -0.28 -0.31 0.10 0.02 0.24 0.54 -0.42 0.34 TAR 0.28 0.17 -0.24 0.26 -0.14 -0.20 -0.08 -0.31 -0.10 0.05 -0.20 0.17 0.71 -0.18 HL 0.29 0.02 -0.02 0.01 0.06 -0.02 -0.00 0.07 -0.03 -0.26 -0.10 0.17 -0.36 -0.82 HW 0.28 -0.19 0.02 0.01 0.23 0.36 -0.05 -0.02 0.18 -0.56 -0.50 0.04 0.07 0.32 EYD 0.28 -0.05 -0.13 -0.12 0.22 0.48 0.33 -0.53 -0.21 0.36 0.05 -0.18 -0.12 -0.04 EN 0.27 -0.01 0.00 -0.28 0.12 -0.66 0.32 -0.19 0.36 -0.02 -0.12 -0.31 -0.09 0.08 IN 0.26 0.19 -0.09 0.60 -0.31 0.09 0.51 0.31 0.06 -0.03 0.04 -0.14 -0.15 0.14 TYM 0.21 0.70 0.34 -0.06 0.49 0.07 -0.03 0.17 0.08 0.06 0.17 0.13 0.12 0.07 IOW 0.21 -0.47 0.71 0.38 0.10 -0.10 -0.07 -0.10 0.02 0.17 0.11 0.03 0.07 -0.05 FL3 0.28 -0.08 -0.09 -0.06 0.14 -0.23 -0.16 0.39 -0.57 0.33 -0.40 -0.02 -0.16 0.18 FD3 0.26 -0.38 -0.27 -0.24 0.10 0.13 0.13 0.44 0.37 0.26 0.25 0.29 0.27 -0.05 TL4 0.28 0.13 -0.02 -0.01 -0.28 0.21 -0.57 0.05 0.37 0.24 -0.06 -0.50 -0.07 -0.08 TD4 0.25 0.09 0.40 -0.51 -0.62 0.12 0.17 0.02 -0.19 -0.09 -0.01 0.16 0.09 0.06

44 Table 3.4. Eigenvalues and Eigenvectors for females of the genus Discodeles, for all 14 principal components. EigenValue: 13.29 0.29 0.12 0.09 0.05 0.04 0.04 0.03 0.03 0.01 0.01 0.00 0.00 0.00 Percent: 94.92 2.10 0.89 0.61 0.34 0.31 0.27 0.22 0.19 0.06 0.04 0.03 0.02 0.01 CumPercent: 94.92 97.02 97.91 98.52 98.85 99.17 99.43 99.65 99.84 99.91 99.94 99.97 99.99 100.00 Eigenvectors: SV 0.27 -0.12 0.05 -0.02 0.19 -0.32 -0.02 -0.03 0.14 0.22 -0.30 -0.62 0.40 0.25 TIB 0.27 0.14 -0.30 -0.04 -0.06 -0.08 0.25 -0.14 -0.23 0.22 -0.59 0.51 0.16 0.00 TAR 0.27 0.10 -0.26 0.00 -0.10 -0.09 0.35 -0.18 -0.17 0.24 0.68 0.00 0.10 0.34 HL 0.27 -0.12 0.14 -0.11 0.17 -0.22 -0.11 -0.05 0.06 0.13 -0.10 0.08 -0.79 0.36 HW 0.27 -0.20 0.29 -0.02 0.25 -0.39 -0.06 -0.03 0.25 0.18 0.27 0.36 0.17 -0.51 EYD 0.26 -0.05 0.44 -0.62 -0.47 0.29 -0.09 -0.13 -0.05 0.06 -0.00 -0.00 0.12 0.03 END 0.27 0.04 -0.26 -0.17 0.20 -0.03 -0.57 0.35 -0.56 -0.08 0.12 -0.02 0.06 -0.08 IND 0.27 0.01 -0.44 0.07 -0.04 0.29 -0.46 -0.41 0.50 -0.07 0.03 0.04 0.03 -0.00 TYM 0.25 -0.63 0.03 0.16 0.30 0.56 0.25 0.18 -0.05 -0.03 -0.02 0.07 0.07 0.08 IOW 0.27 -0.19 0.01 0.48 -0.68 -0.18 -0.10 0.39 0.07 -0.01 -0.03 0.02 -0.04 0.01 FL3 0.27 0.00 0.02 -0.10 0.03 -0.26 0.21 -0.09 0.02 -0.88 -0.01 0.05 0.07 0.10 FD3 0.26 0.55 0.03 -0.11 0.16 0.21 0.17 0.58 0.41 0.03 0.01 0.04 0.01 0.04 TL4 0.27 0.06 -0.20 -0.04 -0.08 0.08 0.31 -0.11 -0.11 0.01 -0.05 -0.46 -0.36 -0.64 TD4 0.26 0.40 0.50 0.53 0.14 0.21 -0.12 -0.31 -0.27 -0.03 -0.01 -0.03 0.03 0.01

45 Table 3.5. Eigenvalues and Eigenvectors for males of the genus Discodeles, for all 14 principal components. EigenValue: 11.82 1.45 0.32 0.14 0.08 0.07 0.05 0.03 0.02 0.01 0.01 0.00 0.00 0.00 Percent: 84.44 10.35 2.30 0.99 0.58 0.51 0.35 0.18 0.15 0.06 0.04 0.02 0.02 0.01 CumPercent: 84.44 94.79 97.09 98.08 98.66 99.17 99.52 99.70 99.85 99.91 99.95 99.97 99.99 100.00 Eigenvectors: SV 0.26 0.11 0.04 -0.08 -0.10 -0.34 -0.08 0.11 0.66 -0.52 0.03 0.07 0.00 0.21 TIB 0.28 -0.19 -0.04 0.01 -0.38 -0.17 -0.05 -0.08 -0.35 0.04 -0.56 0.16 0.13 0.48 TAR 0.29 -0.09 -0.03 -0.05 -0.17 -0.09 -0.02 -0.07 -0.20 0.01 0.38 -0.56 -0.56 0.25 HL 0.28 0.20 -0.04 -0.16 0.07 -0.16 0.24 0.11 -0.11 -0.14 -0.53 -0.28 -0.18 -0.58 HW 0.26 0.33 0.25 -0.37 0.34 -0.27 0.20 0.04 0.05 0.53 0.10 -0.01 0.21 0.26 EYD 0.27 0.17 -0.48 -0.01 0.50 0.21 0.05 0.06 -0.34 -0.41 0.12 0.13 0.11 0.21 END 0.28 -0.08 -0.33 0.21 0.27 0.09 -0.56 -0.17 0.33 0.39 -0.23 -0.10 -0.10 -0.06 IND 0.28 -0.01 -0.43 -0.26 -0.45 0.31 0.25 0.24 0.18 0.27 0.17 0.30 -0.07 -0.14 TYM 0.19 0.59 0.14 0.71 -0.19 0.15 0.16 0.03 0.01 0.08 0.02 -0.00 -0.01 0.06 IOW 0.27 0.07 0.51 -0.19 -0.04 0.38 -0.51 0.38 -0.19 -0.10 0.03 0.12 -0.03 -0.10 FL3 0.29 -0.04 0.16 0.04 0.03 -0.27 -0.03 -0.54 -0.16 -0.03 0.24 0.55 -0.23 -0.31 FD3 0.22 -0.53 0.07 0.41 0.21 -0.29 0.18 0.54 -0.05 0.11 0.13 0.09 0.04 -0.10 TL4 0.29 -0.10 -0.05 0.02 -0.23 -0.04 -0.10 -0.21 -0.08 -0.06 0.27 -0.36 0.72 -0.26 TD4 0.25 -0.35 0.32 0.02 0.18 0.53 0.43 -0.32 0.25 -0.08 -0.15 -0.09 -0.00 0.11

46 Table 3.6. Eigenvalues and Eigenvectors for females of the genus Platymantis, for all 14 principal components. EigenValue: 12.89 0.70 0.13 0.09 0.06 0.04 0.03 0.02 0.02 0.01 0.01 0.01 0.00 0.00 Percent: 92.06 4.97 0.91 0.64 0.42 0.29 0.22 0.16 0.11 0.08 0.05 0.05 0.03 0.02 CumPercent: 92.06 97.02 97.93 98.57 98.99 99.28 99.50 99.66 99.77 99.85 99.91 99.95 99.98 100.00 Eigenvectors: SV 0.28 -0.05 -0.08 -0.07 -0.04 -0.04 -0.09 0.00 -0.31 -0.15 -0.80 0.27 0.14 0.23 TIB 0.28 -0.08 0.08 0.06 -0.06 -0.26 -0.02 -0.18 0.09 -0.03 0.26 0.65 -0.51 0.21 TAR 0.28 -0.11 -0.13 0.18 -0.26 -0.24 -0.08 -0.04 0.26 0.77 -0.19 -0.19 -0.02 -0.05 HL 0.28 -0.13 0.01 -0.05 0.02 0.06 -0.12 -0.15 -0.18 -0.11 -0.06 0.05 -0.17 -0.89 HW 0.28 -0.00 -0.15 -0.13 0.00 -0.06 -0.19 -0.20 -0.70 0.11 0.33 -0.36 -0.09 0.25 EYD 0.26 -0.27 0.56 -0.01 -0.27 0.62 -0.20 0.00 0.07 0.06 0.09 0.02 0.12 0.14 END 0.27 -0.13 0.31 -0.12 0.35 -0.34 -0.03 0.73 -0.08 0.06 0.11 0.01 0.11 -0.04 IND 0.27 -0.15 -0.47 0.21 -0.33 0.28 0.48 0.42 -0.05 -0.16 0.11 -0.02 -0.11 0.01 TYM 0.26 -0.28 -0.20 0.12 0.76 0.27 0.15 -0.25 0.20 0.07 -0.03 -0.06 0.03 0.11 IOW 0.26 0.23 -0.19 -0.85 -0.06 0.09 0.08 -0.03 0.30 0.04 0.05 0.01 0.05 0.01 FL3 0.28 0.04 0.28 0.08 -0.08 -0.25 0.12 -0.12 0.28 -0.44 -0.18 -0.57 -0.32 0.11 FD3 0.23 0.64 0.33 0.18 0.09 0.08 0.52 -0.13 -0.19 0.21 -0.01 0.11 0.12 -0.08 TL4 0.28 -0.10 -0.09 0.15 -0.17 -0.32 -0.02 -0.25 0.14 -0.27 0.27 0.09 0.72 -0.03 TD4 0.24 0.55 -0.24 0.29 0.09 0.19 -0.60 0.21 0.19 -0.13 0.05 0.01 -0.04 0.02

47 Table 3.7. Eigenvalues and Eigenvectors from 14 external measurements for males of the genus Platymantis, for all 14 principal components. EigenValue: 12.64 0.91 0.14 0.08 0.07 0.06 0.03 0.03 0.02 0.01 0.01 0.01 0.00 0.00 Percent: 90.30 6.47 0.96 0.55 0.47 0.45 0.20 0.20 0.12 0.10 0.08 0.05 0.029 0.02 CumPercent: 90.30 96.77 97.74 98.29 98.75 99.20 99.40 99.60 99.72 99.82 99.90 99.95 99.98 100.00 Eigenvectors: SV 0.28 -0.09 -0.04 -0.10 -0.05 0.06 -0.12 -0.18 -0.59 -0.05 -0.07 0.52 -0.27 0.39 TIB 0.28 -0.05 0.12 -0.34 0.11 -0.14 -0.14 -0.16 0.22 0.19 -0.41 -0.05 -0.56 -0.38 TAR 0.28 -0.07 0.23 -0.34 -0.01 0.04 -0.54 0.17 0.32 -0.13 0.51 0.02 0.05 0.20 HL 0.28 -0.12 -0.05 0.01 -0.03 0.01 0.01 -0.09 -0.27 0.10 -0.00 -0.82 -0.09 0.35 HW 0.28 -0.02 -0.15 -0.02 -0.06 -0.04 -0.12 -0.24 -0.40 0.15 0.39 -0.03 0.27 -0.65 EYD 0.27 -0.17 0.19 0.67 -0.27 -0.32 -0.33 0.26 0.01 -0.03 -0.22 0.05 0.02 -0.07 END 0.27 -0.15 -0.24 -0.16 -0.00 -0.32 0.44 0.57 0.03 0.37 0.18 0.13 -0.06 0.07 IND 0.27 -0.22 0.46 0.19 -0.21 0.51 0.42 -0.20 0.20 0.21 0.15 0.11 -0.05 -0.01 TYM 0.26 -0.30 -0.43 0.32 0.65 0.24 -0.05 -0.08 0.22 -0.11 0.01 0.07 0.00 0.02 IOW 0.27 0.21 -0.58 -0.04 -0.59 0.12 -0.01 -0.19 0.35 -0.09 -0.11 0.04 0.05 0.08 FL3 0.28 0.09 0.16 -0.01 0.07 -0.39 0.42 -0.16 0.04 -0.72 0.13 -0.03 -0.05 -0.05 FD3 0.21 0.68 0.13 0.21 0.28 -0.23 -0.00 -0.26 0.12 0.39 0.08 0.08 0.10 0.22 TL4 0.28 -0.11 0.17 -0.32 0.10 0.02 0.03 0.02 -0.01 -0.00 -0.51 0.04 0.71 0.06 TD4 0.24 0.51 0.04 0.02 0.08 0.49 -0.00 0.53 -0.20 -0.21 -0.12 -0.08 -0.12 -0.20

48 Table 3.8. Summary table for females and males of the genus Batrachylodes for the first three principal components. See text for explanation of ‘+’ and ‘-‘. Batrachylodes Females Batrachylodes Males Variable PC1 PC2 PC3 Variable PC1 PC2 PC3 SV + SV + TIB + TIB + (-) TAR + TAR + (-) HL + HL + HW + HW + (-) EYD + EYD + EN + EN + IN + (-) (-) IN + (+) TYM + + TYM + + (+) IOW + + IOW + - + FL3 + FL3 + FD3 + FD3 + - (-) TL4 + TL4 + TD4 + TD4 + +

Table 3.9. Summary table for females and males of the genus Discodeles for the first three principal components. See text for explanation of ‘+’ and ‘-‘. Discodeles Females Discodeles Males Variable PC1 PC2 PC3 Variable PC1 PC2 PC3 SV + SV + TIB + (-) TIB + - TAR + TAR + - HL + (+) HL + (+) HW + + (+) HW + (-) + EYD + (+) - EYD + + END + - END + - IND + - IND + - TYM + + (+) TYM + - IOW + + IOW + (-) FL3 + (+) FL3 + FD3 + - FD3 + + TL4 + TL4 + (-) TD4 + - + TD4 + + +

Table 3.10. Summary table for females and males of the genus Platymantis for the first three principal components. See text for explanation of ‘+’ and ‘-‘. Platymantis Females Platymantis Males Variable PC1 PC2 PC3 Variable PC1 PC2 PC3 SV + SV + TIB + TIB + TAR + TAR + (+) HL + HL + HW + (-) HW + (-) EYD + (-) + EYD + (-) (+) END + + END + (-) IND + - IND + (-) + TYM + (-) (-) TYM + (-) - IOW + (+) (-) IOW + (+) - FL3 + + FL3 + (+) FD3 + + + FD3 + + TL4 + TL4 + (+) TD4 + + (-) TD4 + +

49 3.3.2 Analysis of Sexes Combined For all genera the correlations in Tables 3.11 - 3.13 are positive ranging from 0.39 to 0.96 for Batrachylodes; 0.68 to 0.99 for Discodeles and 0.62 to 0.99 for Platymantis. For all genera there are 14 variables and therefore 14 principal components were extracted from the correlation matrices. However, the correlations among the variables can be considered to be all high so it would be reasonable to assume that only a few PCs are needed to explain most of the variation (Dunteman, 1989). The eigenvalues and eigenvectors for each of the three genera are presented in Tables 3.14 - 3.16.

50 Table 3.11. Correlations among 14 external body measurements for six species of Batrachylodes, sexes combined (n=159) Variable logSVL LogTIB logTAR logHL LogHW logEYD logEN logIN logTYM logIOW logFL3 logFD3 logTL4 logTD4 SV 1.00 0.91 0.89 0.95 0.93 0.91 0.87 0.82 0.63 0.69 0.92 0.89 0.86 0.81 TIB 1.00 0.96 0.93 0.88 0.87 0.87 0.86 0.61 0.66 0.91 0.85 0.91 0.81 TAR 1.00 0.92 0.86 0.85 0.86 0.87 0.63 0.63 0.90 0.81 0.90 0.80 HL 1.00 0.95 0.90 0.89 0.86 0.70 0.70 0.93 0.88 0.88 0.82 HW 1.00 0.90 0.85 0.81 0.61 0.73 0.90 0.90 0.84 0.79 EYD 1.00 0.82 0.79 0.62 0.65 0.88 0.87 0.82 0.79 EN 1.00 0.76 0.63 0.66 0.88 0.84 0.82 0.82 IN 1.00 0.57 0.59 0.81 0.75 0.83 0.74 TYM 1.00 0.40 0.62 0.46 0.63 0.60 IOW 1.00 0.68 0.65 0.66 0.69 FL3 1.00 0.88 0.88 0.81 FD3 1.00 0.78 0.76 TL4 1.00 0.84 TD4 1.00

Table 3.12. Correlations among 14 external body measurements for three species of Discodeles, sexes combined (n=88) Variable SV TIB TAR HL HW EYD END IND TYM IOW FL3 FD3 TL4 TD4 SV 1.00 0.97 0.96 0.99 0.97 0.95 0.97 0.96 0.92 0.96 0.99 0.87 0.98 0.90 TIB 1.00 0.99 0.96 0.93 0.93 0.97 0.97 0.85 0.94 0.98 0.93 0.99 0.93 TAR 1.00 0.97 0.94 0.94 0.97 0.97 0.87 0.95 0.99 0.92 0.99 0.92 HL 1.00 0.99 0.97 0.97 0.96 0.93 0.96 0.99 0.86 0.98 0.90 HW 1.00 0.95 0.94 0.93 0.93 0.96 0.97 0.82 0.95 0.87 EYD 1.00 0.94 0.92 0.88 0.92 0.95 0.85 0.95 0.87 END 1.00 0.97 0.87 0.94 0.97 0.90 0.96 0.90 IND 1.00 0.88 0.94 0.96 0.88 0.97 0.88 TYM 1.00 0.91 0.89 0.68 0.88 0.75 IOW 1.00 0.96 0.84 0.96 0.89 FL3 1.00 0.90 0.99 0.92 FD3 1.00 0.92 0.94 TL4 1.00 0.92 TD4 1.00

51 Table 3.13. Correlations among 14 external body measurements for eight species of Platymantis, sexes combined (n=198) Variable SV TIB TAR HL HW EYD END IND TYM IOW FL3 FD3 TL4 TD4 SV 1.00 0.99 0.98 0.99 0.99 0.95 0.97 0.96 0.94 0.93 0.97 0.76 0.99 0.84 TIB 1.00 0.99 0.99 0.98 0.95 0.97 0.95 0.93 0.91 0.98 0.76 0.99 0.83 TAR 1.00 0.98 0.98 0.94 0.96 0.96 0.93 0.90 0.97 0.74 0.99 0.82 HL 1.00 0.99 0.96 0.98 0.96 0.96 0.91 0.97 0.73 0.99 0.81 HW 1.00 0.93 0.97 0.95 0.93 0.95 0.97 0.78 0.98 0.86 EYD 1.00 0.95 0.92 0.92 0.84 0.94 0.67 0.94 0.72 END 1.00 0.93 0.94 0.90 0.96 0.72 0.97 0.78 IND 1.00 0.93 0.87 0.93 0.67 0.97 0.77 TYM 1.00 0.83 0.90 0.62 0.94 0.71 IOW 1.00 0.93 0.84 0.91 0.90 FL3 1.00 0.83 0.97 0.87 FD3 1.00 0.74 0.95 TL4 1.00 0.82 TD4 1.00

Table 3.14. Eigenvalues and Eigenvectors for 14 external measurements for six species of Batrachylodes, sexes combined (n=159) EigenValue: 11.42 0.66 0.48 0.34 0.27 0.19 0.16 0.11 0.09 0.08 0.08 0.07 0.04 0.03 Percent: 81.59 4.70 3.42 2.43 1.90 1.35 1.13 0.80 0.63 0.60 0.56 0.46 0.26 0.18 CumPercent: 81.59 86.28 89.70 92.13 94.03 95.38 96.51 97.31 97.94 98.54 99.10 99.56 99.82 100.00 Eigenvectors: SVL 0.28 -0.05 -0.08 -0.18 0.08 0.05 -0.11 0.08 -0.27 -0.05 -0.51 0.70 0.16 0.09 TIB 0.28 -0.00 -0.19 0.20 -0.01 -0.24 -0.18 -0.25 -0.21 0.12 0.31 0.15 -0.54 0.47 TAR 0.28 0.08 -0.20 0.27 0.03 -0.31 -0.11 -0.34 -0.23 0.27 0.14 -0.08 0.58 -0.31 HL 0.29 0.05 -0.02 -0.11 0.14 -0.05 0.02 0.16 -0.32 -0.03 -0.08 -0.15 -0.51 -0.68 HW 0.28 -0.12 0.00 -0.25 0.21 0.12 -0.11 0.21 -0.42 -0.29 0.04 -0.52 0.24 0.37 EYD 0.28 -0.02 -0.11 -0.31 0.11 0.40 -0.11 -0.68 0.35 -0.12 -0.12 -0.12 -0.07 -0.06 EN 0.27 0.02 0.03 -0.13 -0.35 -0.53 0.54 -0.11 0.14 -0.39 -0.17 -0.07 0.01 0.07 IN 0.26 0.05 -0.24 0.49 0.43 0.30 0.55 0.15 0.15 -0.01 -0.04 0.01 0.00 0.09 TYM 0.20 0.82 0.39 -0.21 0.14 0.01 0.03 0.07 0.07 0.09 0.21 0.10 0.06 0.08 IOW 0.22 -0.47 0.75 0.12 0.30 -0.13 0.00 -0.09 0.11 0.07 0.08 0.09 0.01 -0.03 FL3 0.28 -0.03 -0.09 -0.10 -0.05 -0.18 -0.16 0.29 0.39 0.59 -0.39 -0.29 -0.06 0.14 FD3 0.27 -0.28 -0.23 -0.39 -0.13 0.11 0.13 0.28 0.20 0.11 0.61 0.25 0.13 -0.12 TL4 0.27 0.08 -0.02 0.39 -0.16 -0.01 -0.53 0.26 0.36 -0.51 0.04 0.04 0.04 -0.11 TD4 0.26 -0.01 0.27 0.22 -0.67 0.50 0.12 -0.02 -0.24 0.19 -0.04 -0.07 -0.00 0.01

52 Table 3.15. Eigenvalues and Eigenvectors for 14 external measurements for three species of Discodeles, sexes combined (n=88) EigenValue: 13.11 0.45 0.12 0.10 0.05 0.05 0.04 0.03 0.03 0.01 0.01 0.01 0.00 0.00 Percent: 93.66 3.24 0.83 0.69 0.37 0.33 0.28 0.25 0.19 0.06 0.04 0.03 0.02 0.02 CumPercent: 93.66 96.90 97.73 98.41 98.78 99.11 99.38 99.63 99.83 99.89 99.93 99.97 99.98 100.00 Eigenvectors: SV 0.27 -0.11 0.05 -0.01 -0.13 -0.31 0.15 0.04 0.06 0.22 -0.66 -0.44 0.10 0.28 TIB 0.27 0.16 -0.28 0.05 -0.03 -0.15 -0.21 -0.25 -0.19 0.21 -0.24 0.70 0.24 0.12 TAR 0.27 0.10 -0.22 0.07 -0.06 -0.14 -0.25 -0.31 -0.12 0.23 0.54 -0.36 -0.18 0.40 HL 0.27 -0.15 0.11 -0.14 -0.11 -0.18 0.19 -0.01 0.06 0.18 -0.02 0.26 -0.80 -0.22 HW 0.27 -0.25 0.30 -0.06 -0.25 -0.35 0.26 0.08 0.22 0.11 0.44 0.13 0.48 -0.15 EYD 0.27 -0.11 0.23 -0.74 -0.05 0.44 -0.30 -0.06 0.03 0.02 -0.04 -0.01 0.09 0.10 END 0.27 0.03 -0.30 -0.15 0.02 0.10 0.37 0.48 -0.64 -0.05 0.11 -0.05 0.06 -0.01 IND 0.27 -0.01 -0.47 0.09 -0.12 0.45 0.42 -0.18 0.50 -0.13 -0.00 0.01 0.03 0.05 TYM 0.25 -0.58 0.06 0.22 0.73 0.08 -0.04 -0.03 -0.02 -0.01 0.01 0.04 0.02 0.08 IOW 0.27 -0.13 0.13 0.50 -0.40 0.32 -0.43 0.44 0.04 0.03 -0.03 0.03 -0.06 -0.00 FL3 0.27 0.01 0.03 -0.03 -0.08 -0.31 -0.12 -0.07 -0.02 -0.88 -0.04 0.02 -0.08 0.13 FD3 0.25 0.58 0.02 -0.08 0.43 -0.15 -0.11 0.45 0.41 0.08 0.06 0.01 -0.02 0.01 TL4 0.27 0.07 -0.19 0.03 0.06 -0.05 -0.23 -0.23 -0.08 0.01 -0.08 -0.32 0.12 -0.80 TD4 0.26 0.41 0.59 0.27 0.07 0.28 0.31 -0.33 -0.24 -0.02 -0.02 -0.02 0.01 0.02

53 Table 3.16. Eigenvalues and Eigenvectors for 14 external measurements for eight species of Platymantis, sexes combined (n=198) EigenValue: 12.78 0.76 0.12 0.10 0.08 0.06 0.03 0.03 0.02 0.01 0.01 0.01 0.01 0.00 Percent: 91.31 5.43 0.82 0.69 0.57 0.39 0.21 0.18 0.12 0.10 0.07 0.05 0.04 0.02 CumPercent: 91.31 96.74 97.56 98.26 98.83 99.22 99.43 99.61 99.72 99.82 99.89 99.94 99.98 100.00 Eigenvectors: SV 0.28 -0.07 -0.11 -0.04 -0.05 -0.02 -0.06 -0.07 -0.41 -0.07 0.01 -0.83 0.09 -0.15 TIB 0.28 -0.07 0.04 0.09 -0.14 -0.29 0.07 -0.17 0.08 0.02 0.59 0.08 -0.56 -0.31 TAR 0.28 -0.09 -0.14 0.26 -0.21 -0.23 -0.13 -0.25 0.41 0.57 -0.37 -0.12 0.01 0.10 HL 0.28 -0.13 0.01 -0.05 -0.00 0.03 -0.07 -0.11 -0.24 -0.09 0.07 0.06 -0.28 0.86 HW 0.28 -0.01 -0.15 -0.13 -0.03 -0.03 -0.06 -0.21 -0.62 0.17 -0.30 0.49 0.03 -0.29 EYD 0.27 -0.23 0.58 0.04 -0.17 0.60 -0.30 -0.11 0.09 0.04 0.08 0.05 0.11 -0.13 END 0.27 -0.14 0.26 -0.26 0.02 -0.35 -0.14 0.75 -0.03 0.23 0.02 0.04 0.09 0.01 IND 0.27 -0.18 -0.41 0.43 -0.03 0.44 0.39 0.44 -0.00 0.02 0.00 0.06 -0.08 -0.05 TYM 0.26 -0.29 -0.02 -0.20 0.84 0.02 0.11 -0.19 0.22 -0.01 -0.05 -0.02 0.01 -0.08 IOW 0.26 0.22 -0.32 -0.72 -0.30 0.24 0.14 -0.04 0.32 -0.04 0.05 0.00 0.05 -0.00 FL3 0.28 0.06 0.28 0.08 -0.14 -0.20 0.21 0.03 0.16 -0.61 -0.54 0.00 -0.20 -0.09 FD3 0.23 0.66 0.37 0.12 0.16 0.05 0.46 -0.06 -0.10 0.30 0.10 -0.05 0.11 0.10 TL4 0.28 -0.10 -0.10 0.20 -0.11 -0.28 0.05 -0.14 0.07 -0.30 0.32 0.18 0.72 0.09 TD4 0.24 0.53 -0.24 0.21 0.24 0.07 -0.65 0.14 0.10 -0.19 0.03 0.05 -0.08 -0.05

54

In Batrachylodes, the first PC had a large variance of 11.42 which accounted for 81.56% of the variance of the 14 measurements (Table 3.14). The 2nd PC had an eigenvalue of 0.66, just below the cut-off suggested by Jolliffe (1986) and accounted for 4.70% of the total variation. The 3rd and succeeding components accounted for considerably less variance ranging from 3.42% to a negligible 0.05% for the 14th PC. This pattern of eigenvalues is typical of highly correlated variables (Dunteman, 1986). Discodeles had similar component loadings to Batrachylodes (Table 3.15). The 1st PC had a large variance of 13.11 which accounted for 93.66% of the total variance of the 14 measurements. The 2nd and succeeding components accounted for considerably less variance ranging from 3.24% to a negligible 0.02% for the 14th PC. In Platymantis, the 1st PC had an eigenvalue of 12.78 and accounted for 91.31% of total variation of the 14 original variables (Table 3.16). The 2nd PC eigenvalue was 0.76, exceeding Jolliffe’s cut-off score, and accounted for 5.43% of total variation. The remaining 12 PCs had negligible contributions ranging from 0.82% to 0.02% of the total variation. For all the three genera, the results corresponded with having highly correlated original variables. As was found in the preliminary analyses of separate sexes, the 1st PC dominated in all genera. All the eigenvectors on this 1st component were positive and evenly loaded (Tables 3.14 - 3.16). This is symbolised in the summary tables (Tables 3.17 - 3.19) for the three genera. To ascertain whether the 2nd and perhaps 3rd components had any contributions, the ‘correlations of the variables with the principal components’ and the ‘proportion of variation accounted for’ were determined. Using Dunteman’s (1989) methods Tables 3.20 - 3.22 were obtained by multiplying the elements of a particular column of eigenvectors by the square root of the corresponding eigenvalue.

55

Table 3.17. Summary table for six species of the genus Batrachylodes, both sexes combined, for the first three principal components. See text for explanation of ‘+’, ‘-‘, ‘(+)’ and ‘(-)’. Batrachylodes PC1 PC2 PC3 SV + TIB + TAR + (-) HL + HW + EYD + EN + IN + (-) TYM + + + IOW + - + FL3 + FD3 + (-) (-) TL4 + TD4 + +

Table 3.18. Summary table for species of the genus Discodeles, both sexes combined, for the first three principal components. See text for explanation of ‘+’, ‘-‘, ‘(+)’ and ‘(-)’. Discodeles Variable PC1 PC2 PC3 SV + TIB + (+) (-) TAR + (-) HL + (-) HW + (-) + EYD + (+) END + - IND + - TYM + - IOW + FL3 + FD3 + + TL4 + (-) TD4 + + +

Table 3.19. Summary table for eight species of the genus Platymantis, both sexes combined, for the first three principal components. See text for explanation of ‘+’, ‘-‘, ‘(+)’ and ‘(-)’. PC1 PC2 PC3 SV + TIB + TAR + (-) HL + HW + (-) EYD + (-) + END + IND + (-) - TYM + (-) IOW + (+) - FL3 + + FD3 + + + TL4 + TD4 + + (-)

56

Table 3.20. Principal Component loadings for 14 external measurements and percent variance explained for six species of Batrachylodes. PC1 PC2 PC3 SV 0.96 -0.03 -0.06 TIB 0.96 -0.00 -0.13 TAR 0.95 0.07 -0.13 HL 0.98 0.04 -0.01 HW 0.95 -0.10 -0.00 EYD 0.93 -0.02 -0.08 EN 0.92 0.01 0.02 IN 0.88 0.04 -0.16 TYM 0.69 0.66 0.27 IOW 0.74 -0.38 0.52 FL3 0.96 -0.03 -0.06 FD3 0.90 -0.23 -0.16 TL4 0.93 0.06 -0.01 TD4 0.88 -0.01 0.19

% variance explained 81.77 4.65 3.38 individually % variance explained 81.77 86.42 89.80 cumulatively

Table 3.21. Principal Component Loadings for 14 external measurements and percent variance explained for three species of Discodeles. Variable PC1 PC2 PC3 SV 0.99 -0.07 0.02 TIB 0.99 0.11 -0.09 TAR 0.99 0.06 -0.08 HL 0.99 -0.10 0.04 HW 0.97 -0.17 0.10 EYD 0.96 -0.07 0.08 END 0.98 0.02 -0.10 IND 0.97 -0.01 -0.16 TYM 0.90 -0.39 0.02 IOW 0.97 -0.09 0.04 FL3 0.99 0.01 0.01 FD3 0.91 0.39 0.01 TL4 0.99 0.05 -0.07 TD4 0.93 0.28 0.20

% variance explained 93.66 3.24 0.83 individually % variance explained 93.66 96.90 97.73 cumulatively

57

Table 3.22. Principal Component loadings for 14 external measurements and percent variance explained for eight species of Platymantis, sexes combined. PC1 PC2 PC3 SV 0.99 -0.06 -0.04 TIB 0.99 -0.06 0.01 TAR 0.98 -0.08 -0.05 HL 0.99 -0.11 0.00 HW 0.99 -0.01 -0.05 EYD 0.95 -0.20 0.20 END 0.97 -0.12 0.09 IND 0.96 -0.16 -0.14 TYM 0.94 -0.25 -0.01 IOW 0.94 0.19 -0.11 FL3 0.99 0.05 0.10 FD3 0.80 0.57 0.12 TL4 0.99 -0.09 -0.03 TD4 0.87 0.46 -0.08

% variance explained 91.31 5.43 0.82 individually % variance explained 91.31 96.74 97.56 cumulatively

For species of Batrachylodes it can be seen that all of the variables have high correlations with the dominant 1st PC (Table 3.20). With the exception of the variables tympanum diameter (TYM) and interorbital width (IOW) this 1st PC has large correlations with the original variables. It can be interpreted as an overall size dimension. The 1st PC scores (Table 3.20) are an equally weighted average of the 14 standardised measurements. Because the variables are all positively correlated the eigenvectors are also positive (Table 3.14); also if the correlations are all about the same magnitude then the elements of the first eigenvector are all positive and about the same magnitude (Dunteman, 1989). Therefore it is reasonable to call this PC a size factor. Since this PC also accounts for 81.59% of the total variance and its eigenvalue is high (11.42) (Table 3.14), this is the most appropriate to summarise the data. However, due to the lower scores for some variables (TYM, IOW) the scores for the 1st PC, 2nd PC and 3rd PCs were added together, in order for the proportion of variance accounted for to be over 80% in all variables (Table 3.23). On this evidence, even though the 1st component has such domination, the 2nd and 3rd components were also retained, hence giving a better indication of shape. The graphical clustering of the 1st and 2nd components for the six species of Batrachylodes can be seen in Fig. 3.1, for 2nd and 3rd Fig. 3.2 and for 1st and 3rd Fig. 3.3. Using the summary table (Table

58

3.17) and Figs 3.1 - 3.3 it can be seen that species of Batrachylodes are mainly separated by overall size (PC1). It appears that as the diameter of the tympanum (TYM) increases, the width between the orbits (IOW) decreases and to a lesser extent the width of the finger discs (FD3) decreases. On the 3rd component, TYM, IOW and TD4 are contrasted against FD3, IN and TAR, the former increase whilst the latter decrease. However, these latter two components are negligible when compared to the dominant 1st PC. It would be safe to conclude that species of Batrachylodes can be separated by overall size.

Table 3.23. Proportion of variance accounted for in 14 external measurements by the first three principal components of six species of Batrachylodes. Variable Proportion of variance accounted for SV 0.93 TIB 0.93 TAR 0.92 HL 0.96 HW 0.91 EYD 0.87 EN 0.85 IN 0.80 TYM 0.97 IOW 0.97 FL3 0.92 FD3 0.89 TL4 0.86 TD4 0.81

59

Batrachylodes

2.0

me ve 1.5 me veve mi ve ve me me ve me ve tr tr tr mi ve 1.0 ve ve ve mi ve tr ve tr tr el meme veve me ve ve wo ve veve vevewo 0.5 trtr me veveveve el mitr me mo tr tr me wo tr ve el ve ve wowo el tr ve wo ve ve veve el wo el mi mi 0.0 tr me wo wo mi tr tr el Prin2 tr wo wo el wo wo wo ve ve el mi tr ve ve ve wowo elel tr ve wo -0.5 veveve ve ve el ve el wo wo wo woel el ve ve el wo tr ve me el el el woel ve -1.0 veveve wo el tr ve el ve ve el woel el ve ve

ve veve ve ve -1.5 ve

-2.0 ve -10 -8 -6 -4 -2 0 2 4 6 Prin1

Fig. 3.1. Principal Component 1 against Principal Component 2 for six species of Batrachylodes (el = B. elegans, me = B. mediodiscus, mi = B. minutus, tr = B. trossulus, ve = B. vertebralis, wo = B. wolfi). Batrachylodes

2.5

wo 2.0

mi 1.5 wo el wo tr mi tr wo wo me woel 1.0 elwo el mi me tr tr tr wo tr wo wo tr ve elwo me tr me wo tr wo wo tr 0.5 wo ve wo el me el el tr tr woel wo tr el tr wo me ve ve tr wo ve mi mi Prin3 wo me miwo eltr ve veve mi me 0.0 veve el wowo el tr tr ve el ve me wo ve el wo tr me el ve veve ve ve el ve ve el ve ve el ve vetr me vemeve ve ve el ve ve ve ve el ve el ve tr ve ve me -0.5 ve ve el me ve ve ve ve ve ve el ve elve mi ve ve ve ve ve veve -1.0 ve ve ve ve ve ve

-1.5 ve ve veve ve -2.0 -2.5 -2.0 -1.5 -1.0 -0.5 0.0 0.5 1.0 1.5 2.0 Prin2

Fig. 3.2. Principal Component 2 against Principal Component 3 for six species of Batrachylodes (el = B. elegans, me = B. mediodiscus, mi = B. minutus, tr = B. trossulus, ve = B. vertebralis, wo = B. wolfi).

60

Batrachylodes

6 el el el el elel el el ve ve elel 4 el wo el veel wo wo elve elve ve ve ve wo el wowo wo el ve wo elwo wowowo wo ve ve ve wo veve ve ve ve el wo ve ve ve wowo wo wo ve veveel 2 ve wo wo ve wo el el el elve wo wo ve el ve ve me 0 ve me vewo ve ve me meve ve ve ve me me me me ve ve me ve memeve veve veme ve ve veve ve ve ve veve -2 ve ve ve ve ve ve me tr Prin1 ve me ve tr ve tr ve tr tr -4 tr tr ve tr ve tr tr tr tr tr tr tr tr tr tr mi tr tr -6 tr mi mi mi mi mi mi -8

mi -10 -2.0 -1.5 -1.0 -0.5 0.0 0.5 1.0 1.5 2.0 2.5 Prin3

Fig. 3.3. Principal Component 1 against Principal Component 3 for six species of Batrachylodes (el = B. elegans, me = B. mediodiscus, mi = B. minutus, tr = B. trossulus, ve = B. vertebralis, wo = B. wolfi).

For Discodeles again the 1st PC dominates the results (Table 3.15). This component represents 93.66% of the total variation of the original variables and has a large eigenvalue of 13.11. Only this component satisfies Jolliffe’s (1986) criteria for PC selection. It, like in Batrachylodes, can be considered an overall size component. The 2nd PC loaded positively for 4th toe disc size (TD4), 3rd finger disc size (D3) and negatively for tympanum diameter (TYM) and to a lesser extent head length (HL) and head width (HW) (Table 3.21). This PC accounts for 3.24% of total variation (Tables 3.15 and 3.21). The 3rd PC loaded positively for TD4, HW and eye diameter (EYD), negatively for eye to naris distance (EN) and internarial distance (IN), and to a lesser extent, the length of the tibia (TIB) and tarsus (TAR). This component accommodates only 0.83% of the variation in the original variables (Tables 3.15 and 3.21). When correlating the PC loadings with the original variables all the variables have high correlations with the 1st dominant PC (Table 3.24) and when adding the 1st and 2nd components together (Table 3.21) all variables account for over 90% of variance. On this basis the 3rd component was removed from the analysis. However, due

61

to the overwhelming size influence of the 1st component, the 2nd component was retained for an indication of shape.

Table 3.24. Proportion of variance accounted for in 14 external measurement variables by the first two Principal Components in three species of Discodeles. Variable Proportion of variance accounted for SVL 0.99 TIB 0.98 TAR 0.98 HL 0.99 HW 0.98 EYD 0.93 END 0.96 IND 0.95 TYM 0.97 IOW 0.95 FL3 0.99 FD3 0.98 TL4 0.99 TD4 0.94

Table 3.24 indicates that all the variables have a substantial proportion of their variance explained by the largest two PCs. Based on the above it was decided to retain just two PCs. Discodeles

2.0 gu

gu 1.5 gu gu gu gu gu gu gu gu 1.0 gu gu

gu gu gu 0.5 gu gu P bu gu op gu ri bu gu n op op bu 2 bu bu bu gu bu bu opbu op op gu 0.0 bu bu bubu bu bu bu bu bubu bu bu bu bu bu op gu bu bu bu bu bu bubu bu bu bu bu -0.5 bu bu bu bu bu bu bubu bu bu bu bu bu bu bu bu bu bu bu bu bu bu -1.0 bu

-6 -4 -2 0 2 4 6 8 Prin1

Fig. 3.4. Principal Component 1 against Principal Component 2 for three species of Discodeles (bu = D. bufoniformis, gu = D. guppyi, op = D. opisthodon).

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The graph of the 1st and 2nd components for three species of Discodeles is shown in Fig. 3.4. It can be seen that D. bufoniformis and D. guppyi can be separated on the 2nd PC. The variables FD3, TD4 have high positive loadings on the component indicating that D. bufoniformis (small discs) and D. guppyi (large discs) can be separated by these two variables. In addition, tympanum diameter loaded negatively on this component indicating that D. bufoniformis has a larger tympanum than D. guppyi. To a lesser extent the two species can be separated by D. bufoniformis having a slightly larger head (both length and width) and smaller leg length (tibia) than D. guppyi. Interestingly, the species D. opisthodon could not be well separated from D. bufoniformis both morphometrically and in the key identifications, but could from D. guppyi. For Platymantis the 1st PC had a large eigenvalue (12.78: Table 3.16) and represents 91.36% of the total variation of the original variables. The 2nd PC had an eigenvalue of 0.76, accounting for 5.43% of total variation. These two components satisfy Jolliffe’s (1986) criteria having a cumulative percentage of 96.79%. Again the 1st component can be considered an overall size component with all eigenvectors loading positively and evenly. This mimics the situation found in Batrachylodes and Discodeles. The 2nd PC loaded positively for toe and finger disc size, contrasted with eye diameter, eye-naris distance and tympanum diameter which all loaded negatively (Table 3.22). The 3rd PC loaded positively for finger length, finger disc width and eye diameter whilst interorbital width and internarial distance loaded negatively. However, this component accommodates only 0.82% of the variation in the original variables (Tables 3.16 and 3.22). When correlating the PC loadings with the original variables, all the variables have high correlations (>80%) with the 1st dominant PC (Table 3.25). So when adding the 1st and 2nd components together (Table 3.22) all variables account for over 90% of variance. On this basis the 3rd component was removed from the analysis and only the 1st and 2nd components graphed. The graphical display of the 1st and 2nd components for the eight species of Platymantis can be seen in Fig. 3.5. It shows that P. parkeri and P. aculeodactylus can be well separated from all other Platymantis on the basis of overall size. Platymantis neckeri and P. guppyi can be separated by the possession of large finger and toe discs. Unfortunately, the

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remaining four species: solomonis, weberi, myersi and acrochorda cannot be well separated using this analysis.

Table 3.25. Proportion of variance accounted for in 14 external measurements by the first two principal components for eight species of Platymantis Variable Proportion of variance accounted for SV 0.99 TIB 0.99 TAR 0.97 HL 0.99 HW 0.99 EYD 0.93 END 0.96 IND 0.94 TYM 0.94 IOW 0.93 FL3 0.98 FD3 0.97 TL4 0.98 TD4 0.97

Platymantis

gu

2.0 gu gu gu gu gu ne gup gu gu gup gup gu gup gu gup ne ne gupgup gu ne nene ne gu ne ne ne ne ne ne gugup ne ne gup 1.0 nene ne gu par par pa pa pa pa acac Prin2 pa par pa ac acac par ac pa par ac ac my papa ac acacac acr ac we pa ac my 0.0 par ac we par pa ac ac wemy pa ac acacacr so so we wemywe ac ac acrwe we we ac we so acrwe we par so we we ac myso so wesoso we wemy mywe we so we mymy weso wemy soso sosomyso so we so my so we sowe so soso so soso sowe wewe so so my so we so so so somy so so so wesoso so so -1.0 so wesomyso so soso so soso my we so so so so

0 Prin1

Fig. 3.5. Principal Component 1 against Principal Component 2 for eight species of Platymantis (acr = P. acrochorda, ac = P. aculeodactylus, gu = P. guppyi, my = P. myersi, ne = P. neckeri, par = P. parkeri, so = P. solomonis, we = P. weberi).

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3.3.3 Further Analysis: Discriminant Function Analysis Species of the three genera, Batrachylodes, Discodeles and Platymantis were further analysed using the statistical technique, discriminant function analysis (DFA). The principal components (PCs) derived from the preceding PCA analyses were used as variables for the DFA. This ensured uncorrelated statistically independent variables (Dunteman, 1989). For all PCA analyses for all genera, 14 PCs were found. A pooled within-groups correlation matrix was constructed (Tables 3.26 - 3.28) and the variables subjected to a one-way analysis of variance.

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Table 3.26. Pooled within-groups correlation matrix for six species of Batrachylodes

PRIN1 PRIN10 PRIN12 PRIN13 PRIN14 PRIN2 PRIN3 PRIN4 PRIN5 PRIN6 PRIN7 PRIN8 PRIN9

PRIN1 1.00 PRIN10 -.00 1.00 PRIN12 .03 -.03 1.00 PRIN13 .04 -.03 -.04 1.00 PRIN14 -.08 .08 -.03 .01 1.00 PRIN2 .58 .06 .01 -.01 .015 1.00 PRIN3 .17 .20 .11 .11 -.10 -.03 1.00 PRIN4 .40 -.07 -.10 -.08 .04 -.07 -.10 1.00 PRIN5 .04 -.08 .03 -.00 .10 .01 .06 -.06 1.00 PRIN6 .03 .00 .01 -.01 .05 -.10 .14 .06 -.01 1.00 PRIN7 .06 -.09 -.04 -.03 .04 .04 .23 .06 -.03 -.01 1.00 PRIN8 -.08 -.02 .06 .02 .09 -.02 .02 .09 -.06 -.06 .01 1.00 PRIN9 .12 -.02 -.05 -.02 -.06 .03 .11 .10 .05 .04 -.07 .08 1.00

Table 3.27. Pooled within-groups correlation matrix for three species of Discodeles PRIN1 PRIN10 PRIN11 PRIN12 PRIN13 PRIN14 PRIN2 PRIN3 PRIN4 PRIN5 PRIN6 PRIN7 PRIN8 PRIN9

PRIN1 1.00 PRIN10 -.00 1.00 PRIN11 .01 .00 1.00 PRIN12 -.01 -.00 .01 1.00 PRIN13 .01 .00 -.01 .01 1.00 PRIN14 .01 -.00 -.01 .01 .01 1.00 PRIN2 -.10 -.03 .12 -.12 .15 .03 1.00 PRIN3 -.00 .01 -.00 -.01 -.08 .09 .33 1.00 PRIN4 -.00 .00 -.00 -.00 -.02 .03 .10 -.14 1.00 PRIN5 .00 -.00 .00 .00 .01 -.01 -.04 .07 .02 1.00 PRIN6 .00 -.00 -.00 .01 .02 -.03 -.06 .15 .04 -.02 1.00 PRIN7 .02 .00 -.03 .03 -.01 -.04 .27 .10 .03 -.01 -.04 1.00 PRIN8 .01 -.00 -.01 .01 .00 -.03 .09 .08 .02 -.01 -.03 -.05 1.00 PRIN9 .01 .00 -.01 .01 -.01 -.01 .10 .00 -.00 -.00 -.00 -.02 -.01 1.00

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Table 3.28. Pooled within-groups correlation matrix for eight species of Platymantis. PRIN1 PRIN10 PRIN11 PRIN12 PRIN13 PRIN14 PRIN2 PRIN3 PRIN4 PRIN5 PRIN6 PRIN7 PRIN8 PRIN9

PRIN1 1.00 PRIN10 .07 1.00 PRIN11 -.30 -.01 1.00 PRIN12 -.05 .02 .00 1.00 PRIN13 .02 .00 -.00 .02 1.00 PRIN14 -.13 .01 .00 .04 -.00 1.00 PRIN2 -.21 -.04 .17 .07 .13 .03 1.00 PRIN3 -.45 .11 .04 .04 .11 .10 -.08 1.00 PRIN4 -.13 .05 -.07 -.05 -.06 .10 .25 .05 1.00 PRIN5 -.25 -.08 -.05 .06 -.02 .02 -.27 .21 -.06 1.00 PRIN6 .37 .02 .02 .07 -.11 .07 -.25 -.16 -.20 -.01 1.00 PRIN7 -.05 .07 .00 .04 .02 -.07 -.05 -.12 -.16 .08 .00 1.00 PRIN8 -.18 -.02 -.02 .09 -.00 -.04 .15 -.03 .03 -.06 -.00 -.04 1.00 PRIN9 -.17 -.01 -.03 .11 .01 .01 .16 -.20 -.13 -.07 -.08 -.08 -.10 1.00

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Since not all PCs are needed to explain the total variation within the original data set, an analysis of variance determined those variables that were statistically significant and hence used in the DFA analysis (Tables 3.29 - 3.31). In addition, DFA’s were run using the ‘minimising Wilks’ lambda’ method.

Table3.29. Wilks' Lambda (U-statistic) and univariate F-ratio with 5 and 153 degrees of freedom, for six species of Batrachylodes

Variable Wilks' F Sig. Lambda

PRIN1 .26 86.86 .00 PRIN10 .89 3.60 .00 PRIN12 .94 2.06 .07 PRIN13 .97 .97 .44 PRIN14 .89 3.75 .00 PRIN2 .82 6.64 .00 PRIN3 .58 22.21 .00 PRIN4 .54 26.01 .00 PRIN5 .92 2.73 .02 PRIN6 .89 3.63 .00 PRIN7 .90 3.28 .01 PRIN8 .91 3.12 .01 PRIN9 .88 4.02 .00

Table 3.30. Wilks' Lambda (U-statistic) and univariate F-ratio with 2 and 85 degrees of freedom for three species of Discodeles Variable Wilks' F Sig. Lambda PRIN1 .99 .32 .72 PRIN10 1.00 .03 .97 PRIN11 .99 .40 .67 PRIN12 .99 .48 .62 PRIN13 .98 .85 .43 PRIN14 .98 1.03 .36 PRIN2 .37 72.71 .00 PRIN3 .67 21.24 .00 PRIN4 .96 1.78 .18 PRIN5 .99 .36 .70 PRIN6 .95 2.00 .14 PRIN7 .91 4.01 .02 PRIN8 .97 1.18 .31 PRIN9 .99 .29 .75

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Table 3.31. Wilks' Lambda (U-statistic) and univariate F-ratio with 7 and 190 degrees of freedom for eight species of Platymantis Variable Wilks’ F Sig Lambda PRIN1 .14 160.98 .00 PRIN10 .95 1.35 .23 PRIN11 .97 .86 .55 PRIN12 .88 3.67 .00 PRIN13 .92 2.22 .03 PRIN14 .92 2.40 .02 PRIN2 .10 248.74 .00 PRIN3 .44 35.01 .00 PRIN4 .57 20.82 .00 PRIN5 .84 5.29 .00 PRIN6 .75 8.90 .00 PRIN7 .82 6.00 .00 PRIN8 .92 2.24 .03 PRIN9 .84 5.04 .00

Wilks’ lambda tests for the equality of group means and is the ratio of within- groups sum of squares to the total sum of squares (Dunteman, 1989). If Wilks’ lambda approaches or equals 1 then the group means are equal, conversely if it approaches or equals 0 the within variability is small compared to the total sum of squares (SPSS, 1993). Hence it is the smallest value of Wilks’ lambda that indicates a difference in group means. When ‘minimising Wilks’ lambda’ the analysis uses a forward selection, backward elimination technique (SPSS, 1993). The 1st variable that is accepted for analysis has the largest selection criterion, in this case the value of the F-statistic. After accepting the 1st variable the value of the criterion is re-evaluated for all the variables not in the analysis and the variable with the largest acceptable criterion is entered next. The variable entered first is then re-evaluated to determine whether it meets the removal criterion. If it does, it is removed from the analysis. These steps are repeated until no more variables meet the entry or removal criteria. For the three genera of Solomon Island ranids the minimum F statistic to enter the analysis was 3.84 and the maximum F statistic to remove was 2.71. This was set for all analyses. The position of entry and or removal for each genus can be seen in the summary tables (Tables 3.32 - 3.34).

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Table 3.32. Summary Table for Batrachylodes Action Step Entered Removed Vars Wilks’ P in Lambda

1 PRIN1 1 .26 .000 2 PRIN4 2 .12 .000 3 PRIN2 3 .05 .000 4 PRIN3 4 .03 .000 5 PRIN7 5 .02 .000 6 PRIN10 6 .02 .000 7 PRIN9 7 .02 .000 8 PRIN14 8 .01 .000 9 PRIN6 9 .01 .000 10 PRIN8 10 .01 .000 11 PRIN5 11 .01 .000

Table 3.33. Summary Table for Discodeles Action Vars Wilks' Step Entered Removed in Lambd P a

1 PRIN2 1 .37 .000 2 PRIN3 2 .23 .000 3 PRIN7 3 .19 .000

Table 3.34. Summary Table for Platymantis Action Vars Wilk’s Step Entered Removed in Lambda P

1 PRIN2 1 .098 .000 2 PRIN1 2 .014 .000 3 PRIN3 3 .005 .000 4 PRIN4 4 .002 .000 5 PRIN6 5 .001 .000 6 PRIN5 6 .001 .000 7 PRIN9 7 .001 .000 8 PRIN7 8 .001 .000 9 PRIN11 9 .000 .000 10 PRIN8 10 .000 .000 11 PRIN12 11 .000 .000 12 PRIN13 12 .000 .000 13 PRIN14 13 .000 .000

For Batrachylodes 11 variables were entered into the analysis. Variables PC11, PC12 and PC13 were removed from the analysis. Two of these PC12 and PC13 were not statistically significant from the ANOVA (Table 3.29). Even more data reduction was seen in the results for Discodeles, only three variables were used in the DFA analysis after minimising using Wilks’ lambda, namely PC2, PC3 and PC7 (Table 3.33). Interestingly, PC1 was removed from the analysis, yet this variable had the largest eigenvalue and dominated the PCA analysis but was not statistically significant at the 0.05 level in the ANOVA. By itself this largest principal

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component, although accounting for 93.66% of total variation of the 14 original variables did not significantly discriminate among the three species of Discodeles (Table 3.30). Thus this component, which can be regarded as an overall size component, did not separate out the species and the smaller less influential PCs: 3 and 7 which were discarded in the PCA analysis, better represent the data. This finding is cause enough to include all 14 PCs in the DFA analysis and not just those PCs that satisfied the requirements of group delineation in the PCA. For Platymantis all but one PC, PC11, were entered into the analysis (Table 3.31). PC2 was entered into the analysis first followed by the dominant 1st component. Canonical discriminant functions were then derived for each genus (Tables 3.36 – 3.38). The number of discriminant functions found equals the number of groups minus one or the number of variables entered into the analysis, whichever is the smallest number. For Batrachylodes five functions were found (6 species - 1), Discodeles three functions found (number of variables) and for Platymantis there were seven functions (8 species - 1). Like PCA, the DFA calculated eigenvalues and the percentage of variance explained by the data. Jolliffe’s (1986) methods of selecting PCs were again used to select discriminant functions that best represent the original data. Three functions were chosen for Batrachylodes, two for Discodeles and three for Platymantis (Tables 3.36 - 3.38).

Table 3.36. Canonical Discriminant Functions for Batrachylodes Fcn Eigenvalue Pct of Cum Canonical After Wilks’ Chi- df P Variance Pct Corr Fcn Lambda squared

: 0 .01 705.14 55 .000 1* 11.78 78.80 78.80 .96 : 1 .11 324.28 40 .000 2* 1.63 10.91 89.72 .79 : 2 .30 179.65 27 .000 3* .83 5.53 95.24 .67 : 3 .55 89.66 16 .000 4* .48 3.23 98.47 .57 : 4 .81 30.83 7 .001 5* .23 1.53 100.00.43 : * Marks the 5 canonical discriminant functions remaining in the analysis.

Table 3.37. Canonical Discriminant Functions for Discodeles Fcn Eigenvalue Pct of Cum Canonica After Wilks' Chi- df P l Variance Pct Corr Fcn Lambda squared

: 0 .19 141.35 6 .000 1* 2.62 84.38 84.38 .85 : 1 .67 33.24 2 .000 2* .49 15.62 100.00 .57 : * Marks the 2 canonical discriminant functions remaining in the analysis.

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Table 3.38. Canonical Discriminant Functions for Platymantis Fcn Eigenvalue Pct of Cum Canonical After Wilks' Chi- df P Variance Pct Corr Fcn Lambda squared

: 0 .000 1590.49 91 .000 1* 19.53 52.66 52.66 .98 : 1 .004 1026.96 72 .000 2* 13.08 35.26 87.92 .96 : 2 .057 533.76 55 .000 3* 2.13 5.73 93.65 .83 : 3 .179 321.29 40 .000 4* 1.30 3.51 97.16 .75 : 4 .411 165.92 27 .000 5* .51 1.36 98.52 .58 : 5 .618 89.73 16 .000 6* .36 .98 99.49 .52 : 6 .842 32.11 7 .000 7* .19 .51 100.0 .40 : * Marks the 7 canonical discriminant functions remaining in the analysis.

The graphical displays of group identification are shown in Figs 3.6 – 3.11. Please refer to Appendix 2 for the data that generates such graphs. For Batrachylodes good clustering can be seen in Fig. 3.6 where the 1st discriminant function is plotted against the 2nd. Batrachylodes vertebralis clusters more tightly in the DFA than in the PCA and there is further separation of the species with large discs: elegans, wolfi and vertebralis. The three species of Discodeles are best represented by the 1st discriminant function graphed against the 2nd function (Fig. 3.9). Good separation can be seen including a better representation of D. opisthodon and D. bufoniformis. This could not be ascertained from the PCA. The eight species of Platymantis appear to cluster into three groups and within these groups species clusters can be seen (Figs 3.10, 3.11). Again the small SVL frogs, P. parkeri, P. acrochorda and P. aculeodactylus are well separated from the other species. The species with large discs: P. neckeri and P. guppyi also form a cluster and again the species: P. solomonis, P. myersi and P. weberi overlap considerably. These similar body-sized ranids must be separated by other qualitative features.

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Batrachylodes

4 ve 3 ve veve ve mi ve ve ve ve ve ve veve ve veve 2 ve veve ve ve veve ve ve mi me ve elve ve veveveve ve tr veveveveve ve 1 me me ve ve mi me veveveveel tr mi tr ve veve veveve veve tr ve veveel el me ve veelwo ve tr mi me me veve ve el elel 0 mi tr tr elel el me ve el el el tr tr el wo tr tr el tr mi mi me el wo el -1 tr tr me me el tr me me wo wowo el tr tr wo DIS2_1 me tr wo el -2 el tr wowoel wo wo el wo wo tr wowo tr wo wowo -3 wo wo wo el wo wo wo -4

-5 wo

-6 -12 -10 -8 -6 -4 -2 0 2 4 6 DIS1_1

Figure 3.6 Discriminant Function 1 against Discriminant Function 2 for six species of Batrachylodes

Batrachylodes

el 3 el elel el el el el tr el 2 el el el el tr el el ve metr el el tr el veve tr me ve ve ve tr el el ve 1 me me tr elveel ve ve wo me tr el tr ve el trme el ve mive me tr ve wo tr ve me ve ve ve ve ve tr wo tr meve ve wo ve ve ve tr ve ve mi ve 0 tr ve wo veve wo mi tr ve DIS3_1 me ve veve ve ve veve miwo veme ve wo veve ve wo tr tr ve ve wo veve veve -1 wo wo wome ve wowo wo wo mi ve ve ve ve tr veve ve ve wo wo me wo wo mi -2 wo wo tr mive ve wo ve mi ve wo vewo -3 ve -6 -5 -4 -3 -2 -1 0 1 2 3 4 DIS2_1

Figure 3.7 Discriminant Function 2 against Discriminant Function 3 for six species of Batrachylodes

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Batrachylodes

el 3 el elel el el el tr el el 2 el el el el tr el ve el el tr me elel tr veve tr me ve veve tr el ve el 1 me me ve tr veve elel el tr tr me ve wo tr me el mi ve ve el tr me me ve tr me veve ve wo ve ve ve tr tr me ve wove ve ve ve wo mi tr ve ve ve 0 tr ve veve wo mi tr ve wo DIS3_1 me ve ve ve ve ve ve wo mi me veve ve veve wo trtr ve wo ve ve ve ve ve wo -1 wo me vewo wo wo wo mi ve ve wo wo ve ve tr ve veve ve me wowo mi wowo -2 tr wowo mi veve wo ve mi ve wo ve wo -3 ve -12 -10 -8 -6 -4 -2 0 2 4 6 DIS1_1

Figure 3.8 Discriminant Function 1 against Discriminant Function 3 for six species of Batrachylodes

Discodeles

4

op 3 op op op op op op 2 bu

bu gu bu bu gu gu bu bu gu gu 1 bu bu bu gu bu bu bu bu bu gu bu gu bu bu gu DIS2_1 bu bubu bu bu bu gu gu 0 bu bu bu bu bu gu bu bu gu bu bu bu bu bu gu gu gu bu gu gu gu bu bu bu bu bu -1 bu bu bu bu bu gu bu gu bu bu bu bu gu bu bu bu -2 bu

gu bu -3 -3 -2 -1 0 1 2 3 4 5 DIS1_1

Figure 3.9 Discriminant Function 1 against Discriminant Function 2 for three species of Discodeles

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Platymantis

6 so so 5 mywe so so somy my sowe we so sososo 4 sowe so mywe sosososososo we mywe sosososososososo weso soso so somysosoweso somysosomysoso wesososowewe 3 we mysoso we we sowemywe we so soso we we so so we we mywe so 2 my wewe mywe sowe we wewe we we so so my 1 ac we my wemy ac 0 ac ac ac ac ac ac -1 ac ac acr acr DIS2_1 pa ac acacacr -2 ac acr acac pa pa ac acac ne ac ne -3 pa ac ne pa ne pa ac ne gu pa pa ac ne -4 pa pa gu ne pa pa ne nene pa pa pa ne ne ne ne gu gugu pa ne gugu pa pa gu gu -5 nene ne gugu pa gu pa gu gu gugu gu -6 gu gu gu pa gu gu -7

-8 gu -10 0 10 DIS1_1

Figure 3.10 Discriminant Function 1 against Discriminant Function 2 for eight species of Platymantis

Platymantis

5 nene ne ne 4 pa ne

ne ne ne 3 pa pa wene ne pa ne pa my ne my we ne ne 2 pa so papa ne ne pa we we ne papa we gu pa sosoweso pa mymy pa soso my we 1 pa somy my we ne pa so so we my mymy so so weweso papa weso wesowewe we pa so so wewe we so so gu pa pa sosomywemy we we 0 wemysoweso we we soso sososo we gu so so so DIS3_1 acr so my so so ac we sosososo sowe gu gu ac so so sowe soweso soso so acr so soso we gu gu -1 ac acr so gu my we gu ac ac ac so so ac so gu ac gu acac ac -2 ac ac gugu acac ac ac ac so my ac gu gu so gu gugu ac gugu -3 ac gu ac gu gu ac gu -4 ac

-5 -10 0 10 DIS1_1

Figure 3.11 Discriminant Function 2 against Discriminant Function 3 for eight species of Platymantis

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3.4 Discussion Discriminant function analysis, unlike PCA, will classify groups based on actual group membership and predicted group membership. For Batrachylodes 93.08% of individuals were successfully classified to their actual groups. For Discodeles it was 89.77% and for Platymantis 84.55% (Tables 3.39 – 3.41).

Table 3.39. Classification results for six species of Batrachylodes. Percent of "grouped" cases correctly classified for six species of Batrachylodes: 93.08% No. of Predicted Group Membership Actual Group Cases elegans mediodiscus minutus trossulus vertebralis wolfi elegans 25 23 0 0 0 2 0 92.0%0.0% 0.0%0.0% 8.0% 0.0% mediodiscus 14 0 13 0 0 1 0 0.0%92.9% 0.0%0.0% 7.1% 0.0% minutus 8 0 0 7 1 0 0 0.0%0.0% 87.5%12.5% 0.0% 0.0% trossulus 21 0 0 0 21 0 0 0.0%0.0% 0.0%100.0% 0.0% 0.0% vertebralis 65 4 0 0 0 60 0 6.2%0.0% 0.0%0.0% 92.3% 0.0% wolfi 26 1 0 0 0 1 24 3.8%0.0% 0.0%0.0% 3.8% 92.3%

Table 3.40. Classification results for three species of Discodeles. Percent of "grouped" cases correctly classified for three species of Discodeles: 89.77% No. of Predicted Group Membership Actual Group Cases bufoniformis guppyi opisthodon bufoniformis 58 51 1 6 87.9% 1.7% 10.3% guppyi 23 2 21 0 8.7% 91.3% 0.0% opisthodon 7 0 0 7 0.0% 0.0% 100.0%

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Table 3.41. Classification results for eight species of Platymantis. Percent of "grouped" cases correctly classified for eight species of Platymantis: 84.85% Actual Group No of Predicted Group Membership Cases acrochodus aculeodactylus guppyi myersi neckeri parkeri solomonis weberi acrochordus 4 4 0 0 0 0 0 0 0 100.0% 0.0% 0.0% 0.0% 0.0% 0.0% 0.0% 0.0% aculeodactylus 24 1 23 0 0 0 0 0 0 4.2% 95.8% 0.0% 0.0% 0.0% 0.0% 0.0% 0.0% guppyi 24 0 0 23 0 1 0 0 0 0.0% 0.0% 95.8% 0.0% 4.2% 0.0% 0.0% 0.0% myersi 16 0 0 0 12 0 0 1 3 0.0% 0.0% 0.0% 75.0% 0.0% 0.0% 6.3% 18.8% neckeri 18 0 0 0 0 18 0 0 0 0.0% 0.0% 0.0% 0.0% 100% 0.0% 0.0% 0.0% parkeri 22 0 0 0 0 0 22 0 0 0.0% 0.0% 0.0% 0.0% 0.0% 100% 0.0% 0.0% solomonis 56 0 0 0 5 0 0 42 9 0.0% 0.0% 0.0% 8.9% 0.0% 0.0% 75% 16.1% weberi 34 0 0 0 2 0 0 8 24 0.0% 0.0% 0.0% 5.9% 0.0% 0.0% 23.5% 70.6%

These percentages of groups correctly classified are high and so taxa can be confidently identified. Interestingly 10% of individuals originally classified as D. bufoniformis were classified as D. opisthodon (Table 3.40), thus adding to speculation about the status of D. opisthodon. Encouragingly the individuals classified as D. opisthodon were all classified as D. opisthodon, however, there were only seven individuals and a larger sample size is needed before the status of D. opisthodon can be recognised. For the remainder of this thesis individuals identified as D. opisthodon were removed from the study. For Batrachylodes (Table 3.39) only the species trossulus had a 100% match of actual to predicted group membership. However the other five species had well over 85% correctly classified group membership. For Platymantis three species had 100% correctly identified individuals: acrochorda, parkeri and neckeri (Table 3.41). Two species: aculeodactylus and guppyi had over 95% correct identification. These species can be confidently ascertained by morphometrics. The remaining three species: myersi (75%), solomonis (75%) and weberi (70.6%) have less correctly assigned individuals. Yet again these three species overlap considerably and it should be reinforced that other equally important qualitative features are required to separate out these species. In summary, and in the absence of a definitive taxonomic key for the Solomon Island ranids, specimens within the following genera: Batrachylodes, Discodeles and Platymantis, that were used in this thesis, were confidently placed

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in their appropriate taxa from the results of the morphometric analyses. This then enabled further character analysis (i.e. osteological examination and external morphological descriptions) and ultimately their relationships to each other to be tested in a phylogenetic analysis. The remaining Solomon Island species, Ceratobatrachus guentheri and Palmatorappia solomonis, are monotypic with distinct morphologies so their identification was not ambiguous. Identification of the species, Rana kreffti, was made based on the findings of Menzies (1987) that only one species of Rana exists in the Solomon Islands, all species identified from New Britain having been reclassified as Rana daemeli.

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Chapter 4. Osteology of Solomon Island Ranids

4.1 Introduction The first osteological account of any of the ranid frogs from the Solomon Islands was by Laurent (1943) of Ceratobatrachus guentheri. He described the unusual skull with its secondary hyperossification, noting that the bones were not fused as Boulenger (1886a) had suggested due to the clearly distinguished suture lines. He also briefly described the orientation of the diapophyses of the vertebrae. Other than this there have been no detailed osteological descriptions of the Solomon Island ranids, although authors have noted the uniting features of a forked omosternum and large, medially fused nasals (Boulenger, 1886a; Brown, 1952; and Noble, 1931).

4.2 Methods 70 specimens from 16 species were examined. 52 were cleared and double- stained for skeletal examination after the Alcian Blue/Alizarin Red staining technique of Dingerkus and Uhler (1977) which was modified by increasing the amount of alcian blue and heating the trypsin to 37°C. This technique revealed bone (red) and cartilage (blue). 18 specimens were prepared using the Alizarin red technique of Davis and Gore (1947), to stain bones (see Appendix 1, specimens are identified by the letter o). All skeletal specimens were examined using a Wild M5 stereo dissecting microscope. A representative of each species was then illustrated using the camera lucida attachment. Osteological descriptions follow the format of Trueb (1979) and the nomenclature of Trueb (1973) with the exception of the use of vomers rather than prevomers. Descriptions of the pelvic girdle follow Tyler (1976). Descriptions of the hyoid follow Trueb (1966) except that posterior cornua was used instead of thyrohyal.

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4.2.1 Material Examined Material is deposited in the following institutions: CAS = California Academy of Sciences, San Francisco, California, USA. MCZ = Museum of Comparative Zoology, Harvard University, Boston, USA. SAM = South Australian Museum, Adelaide, Australia.

Batrachylodes elegans: CAS 117452, m; SAM R8210A, m, Mutahi, Bougainville, Solomon Islands. Batrachylodes mediodiscus: SAM R8204A, m; SAM R8204B, m, Mutahi, Bougainville, Solomon Islands. Batrachylodes trossulus: SAM R5167C, m, Kunua, Bougainville, Solomon Islands (05°49’S 154°48’E); SAM R8209A, f; SAM R8209D, f, Mutahi, Bougainville, Solomon Islands; SAM R47122, f, Pavora River, Choiseul Island, Solomon Islands (06°46’S, 125°32’E). Batrachylodes vertebralis: CAS 117473, m, Mutahi, 3000-3600ft, Bougainville, Solomon Islands; MCZ 41834, m; MCZ 41836, m; MCZ 41839, m; MCZ 41846, m; MCZ 41848, f, Bougainville, 4,000ft above Popoheirarai, Solomon Islands; SAM R4243, m; SAM R5162B, m; SAM R5162F, m, Kunua, Bougainville, Solomon Islands (05°49’S 154°48’E); SAM R47108, f; SAM R47110, m, Pavora River, Choiseul Island, Solomon Islands (06°46’S, 125°32’E). Batrachylodes wolfi: SAM R4927A, m; SAM R5164B, m; SAM R5164C, m, Kunua, Bougainville, Solomon Islands (05°49’S 154°48’E); SAM R8195C, f, Mutahi, Bougainville, Solomon Islands. Ceratobatrachus guentheri: MCZ 42114, juvenile; MCZ 42118, juvenile, Kunua, Bougainville, Solomon Islands (05°49’S 154°48’E); MCZ 62222, f; MCZ 62225, f; MCZ 62226, f, Mutahi 1800- 2700ft, Bougainville, Solomon Islands. Discodeles bufoniformis: CAS 109895, sex unknown, Matsiogu, 2100ft, Bougainville, Solomon Islands; MCZ 38551, f; MCZ 38556, f; MCZ 38557, f; MCZ 38578, m; MCZ 38587, m, Kunua, Bougainville, Solomon Islands (05°49’S 154°48’E). Discodeles guppyi: SAM R47091, f; SAM R47092, m, Pavora River, Choiseul Island, Solomon Islands (06°46’S, 125°32’E); Palmatorappia solomonis: MCZ 60852, f; MCZ 60858, f; MCZ 60859, m; MCZ 60863, m; MCZ 60864, m, Mutahi, 2,200-2,700ft, Bougainville, Solomon Islands; SAM R4426, m; SAM R4925B, m; SAM R4925C, Kunua, Bougainville, Solomon Islands (05°49’S 154°48’E). Platymantis guppyi: SAM R47128, f, Isabel, Solomon Islands (06°00’S 159°29’E); SAM R4917B, m; SAM R4922C, Kunua, Bougainville, Solomon Islands (05°49’S 154°48’E). Platymantis myersi: SAM R47157, f, Pavora River, Choiseul Island, Solomon Islands (06°46’S, 125°32’E). Platymantis neckeri: SAM R4926D, m; SAM R4928B, m, Kunua, Bougainville, Solomon Islands (05°49’S 154°48’E). Platymantis parkeri: SAM R5169A, m; SAM R5169B, f, Kunua, Bougainville, Solomon Islands (05°49’S 154°48’E); SAM R8199, m, Mutahi, Bougainville, Solomon Islands. Platymantis solomonis: SAM R47134, f; SAM R47135, f, Pavora River, Choiseul Island, Solomon Islands (06°46’S, 125°32’E); MCZ 58558, juvenile; MCZ 58559, f; MCZ 58656, f; MCZ 58568, f; MCZ 58569, f, Turiboiru, Bougainville, Solomon Islands. Platymantis weberi: SAM R4412, m, Kunua, Bougainville, Solomon Islands (05°49’S 154°48’E). Rana kreffti: MCZ 38083, f; MCZ 38084, f; MCZ 38087, m; MCZ 38089, m; MCZ 38207, f; SAM R4240, ;SAM R5157, F; SAM R5161A, f, Kunua, Bougainville, Solomon Islands (05°49’S 154°48’E).

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4.3 Osteological Descriptions

4.3.1 Batrachylodes Boulenger, 1887

4.3.1.1 Batrachylodes elegans Brown and Parker, 1970 Fig. 4.1 Skull robust, with well-ossified neurocranium. Nasal plate found in males covering sphenethmoid dorsally, merging with it ventrally. Nasal plate absent in females. Sphenethmoid ossified anteriorly to nasals. Prootic fused with exoccipital dorsally but not ventrally to form otoccipital. Crista parotica well developed but not articulated with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, posteriorly fused with otoccipital, no flanges posterolaterally. Nasals large, broad, bulbous medially, juxtaposed medially, overlying bony sphenethmoid. Maxillary process of nasal short, acuminate, lying adjacent to preorbital process of pars facialis of maxilla, not reaching palatines. Palatines well ossified, long, slender, rounded medially, terminating on sphenethmoid at a level between anterior extremity of cultriform process of parasphenoid and vomers; lateral expansion extending beneath pars facialis of maxilla. Parasphenoid robust, 1/2 width of frontoparietals; cultriform process rounded terminally, extending anteriorly to level posterior to anterior margin of frontoparietals. Parasphenoid alae broad, horizontal. Pterygoid well developed; anterior ramus extending to level of palatine, ventrally overlapping pars dentalis of maxilla; posterior ramus acuminate; medial ramus separated from crista parotica by cartilage. Quadratojugal robust, fully articulated. Squamosal well developed. Otic ramus distinct, longer than zygomatic ramus, not articulating with crista parotica. Zygomatic ramus slender, tapering to point. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Premaxilla broad; alary processes long, separated medially. Palatal shelf of premaxilla broad, expanded laterally; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla distinctly V-shaped. Vomers reduced medially, articulating with ventral surface of sphenethmoid; anterior process reduced; posterior process long, spine-like, converging medially; lateral alae forming anterior and medial margins of choanae; dentigerous processes absent.

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Lateral margin of hyoid plate indented; plate cartilaginous, as long as wide. Anterior cornua slender, cartilaginous, curving anteromedially, expanded at midpoint; anterior processes absent. Alary processes broad, blade-like, cartilaginous lateral extensions directly arising from above anterior margin of hyoid plate. Posterolateral processes long (approximately 64% length of posterior cornua), straight with distal notch. Posterior cornua bony, except distally. Posterior cornua abuts hyoid plate, narrowly separated proximally, separated at angle of 20°. Both proximal and distal ends of posterior cornua undilated.

Fig. 4.1. Hyoid apparatus of Batrachylodes elegans CAS117452, ventral. Scale bar = 1 mm.

Firmisternal pectoral girdle robust. Epicoracoids ossified. Omosternum present, unforked. Sternum ossified, well developed, short, approximately as long as broad, with large cartilaginous xiphoid process. Clavicles large, slender, straight. Coracoids well developed, anterior margin concave, posterior margin straight, expanded medially and well separated medially; oriented horizontally. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

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Eight presacral vertebrae. Neural spines present on Presacrals II, III, IV decreasing in size anterior to posterior. Transverse processes of Presacrals III and IV moderately expanded distally. Lengths of transverse processes: III > IV > V = VI = VII > II = VIII. Orientation of transverse processes: II angled anteriorly, III, VIII horizontal, IV, V, VI, VII angled posteriorly. Sacral diapophyses undilated, cylindrical, with anterior edge oriented posterolaterally. Urostyle bearing dorsal crest along entire length. Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements slightly T-shaped. Phalangeal formula of foot: 2-2-3-4-3. Distal tips of phalangeal elements T-shaped. Sesamoid bone at junction of tibiotarsus.

4.3.1.2. Batrachylodes mediodiscus Brown and Parker, 1970 Figs 4.2, 4.3 Skull broad, robust, with well-ossified neurocranium. Nasal plate found in males covering sphenethmoid dorsally, merging with it ventrally. Nasal plate absent in females. Sphenethmoid ossified dorsally to level beyond anterior margin of nasals just posteriorly to premaxilla. Prootic fused with exoccipital dorsally but not ventrally to form otoccipital. Crista parotica well developed juxtaposed with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, posteriorly fused with otoccipital, no flanges posterolaterally. Nasals large, broad, slightly separated medially, overlying bony sphenethmoid. Maxillary process of nasal short, acuminate, lying adjacent to preorbital process of pars facialis of maxilla, not reaching palatines. Palatines well ossified, long, slender, rounded medially, terminating on sphenethmoid at a level between anterior extremity of cultriform process of parasphenoid and vomers; lateral expansion overlapped by pars facialis of maxilla. Parasphenoid robust, 2/3 width of frontoparietals; cultriform process truncate terminally, not reaching palatines. Parasphenoid alae broad, horizontal. Pterygoid well developed; anterior ramus extending to level of palatine, ventrally overlapped by pars dentalis of maxilla; posterior ramus acuminate; medial ramus articulating with crista parotica via cartilage. Anterior and posterior rami forming a substantial brace; inner margins of rami forming a crescentic curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal, oriented approximately horizontal to midline. Quadratojugal robust, anteriorly separated from maxilla, posteriorly by a cartilaginous connection with

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squamosal. Squamosal well developed. Otic ramus distinct, longer than zygomatic ramus, not articulating with, but juxtaposed with crista parotica. Zygomatic ramus slender. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Premaxilla broad; alary processes short, narrowly separated. Palatal shelf of premaxilla broad, expanded laterally; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla distinctly V-shaped. Vomers reduced medially, articulating with ventral surface of sphenethmoid; anterior process reduced; posterior process long, spine-like, converging medially; lateral alae forming anterior and medial margins of choanae; dentigerous processes absent.

a

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b

Fig. 4.2. Skull of Batrachylodes mediodiscus SAM8204b, male. (a) dorsal (b) ventral. Nasal Plate has been removed for clarity. Scale bar = 1 mm

Firmisternal pectoral girdle robust. Epicoracoid cartilage present. Omosternum present, unforked. Sternum ossified, well developed, short, approximately as long as broad, with large cartilaginous xiphoid process. Clavicles large, slender, straight. Coracoids well developed, anterior margin concave, posterior margin straight, expanded medially and well separated medially; oriented horizontally. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified. Eight presacral vertebrae. Neural spines present on Presacrals II, III, IV, decreasing in size anterior to posterior. Transverse processes of Presacrals III and IV moderately expanded distally. Lengths of transverse processes: III > IV > V = VI = VII = VIII > II. Orientation of transverse processes: II, III, VIII horizontal, IV, V, VI, VII angled posteriorly. Sacral diapophyses undilated, cylindrical, with anterior edge oriented horizontally. Urostyle bearing dorsal crest along 2/3 length. Ilia cylindrical, shaft curved dorso-ventrally, approximately 70% of total length of pelvic girdle with distinct crest on shaft (approximately 50% width of shaft, 86% length of shaft). Prominent round, dorsal protuberance proximally, arising anteriorly to acetabulum. Dorsal acetabular expansion concave, angled approximately 45° to ilial shaft. Ischium well developed, bearing a crest, forming

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posterior and posterodorsal acetabulum. Ventral portion of acetabulum moderately developed with ossified pubis fused with ilium and ischium. Acetabulum large, width approximately 44% of ilium (minus shaft) and ischium; anterior edge of rim approximately 1/2 above the base of ilial shaft.

Fig. 4.3. Pelvic girdle of Batrachylodes mediodiscus SAM8204b, male, lateral. Sutures not identifiable. Scale bar = 1 mm.

Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements T- shaped. Prepollical elements well developed. Sesamoid bones ventrally at 1st metacarpophalangeal joint. Phalangeal formula of foot: 2-2-3-4-3. Distal tips of phalangeal elements T-shaped. Prehallux well developed, elements cartilaginous. Sesamoid bone at junction of tibiotarsus and ventrally between interphalangeal joints on toes 1,2,5.

4.3.1.3. Batrachylodes trossulus Brown and Myers, 1949 Figs 4.4 - 4.9 Skull robust, with well-ossified neurocranium. Nasal plate found in males covering sphenethmoid dorsally, merging with it ventrally. Nasal plate absent in females. Sphenethmoid ossified to level with the centre of nasals forming a rounded apex anteriorly; ossified portion extending 1/3 length of orbit in ventral view. Prootic fused with exoccipital dorsally but not ventrally to form otoccipital. Crista parotica well developed but not articulated with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, posteriorly fused with otoccipital, no flanges posterolaterally. Nasals large, broad, narrowly separated medially, overlying bony sphenethmoid. Maxillary process of nasal short, acuminate, lying adjacent to preorbital process of pars facialis of

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maxilla, not reaching palatines. Palatines well ossified, long, slender, rounded medially, terminating on sphenethmoid at a level between anterior extremity of cultriform process of parasphenoid and vomers; lateral expansion overlapped by pars facialis of maxilla. Parasphenoid robust, 2/3 width of frontoparietals; cultriform process rounded terminally, extending anteriorly to level posterior to palatines. Parasphenoid alae broad, horizontal. Pterygoid well developed; anterior ramus extending to level of palatine, ventrally overlapped by pars dentalis of maxilla; posterior ramus acuminate; medial ramus articulating with crista parotica via cartilage. Anterior and posterior rami forming a substantial brace; inner margins of rami forming a crescentic curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal, oriented approximately horizontal to midline. Quadratojugal robust, anteriorly articulating with maxilla, posteriorly fused with squamosal. Squamosal well developed. Otic ramus distinct, longer than zygomatic ramus, widely separated from crista parotica. Zygomatic ramus slender. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Premaxilla broad; alary processes long, widely separated. Palatal shelf of premaxilla broad, expanded laterally; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla distinctly V-shaped. Vomers reduced medially, articulating with ventral surface of sphenethmoid; anterior process reduced; posterior process long, spine-like, converging medially; lateral alae forming anterior and medial margins of choanae; dentigerous processes absent.

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a b

Fig. 4.4. Skull of Batrachylodes trossulus. (a) SAM47122, female, dorsal (b) SAM8209D, female, ventral. Scale bar = 1 mm

Lateral margin of hyoid plate indented; plate cartilaginous, as long as wide. Anterior cornua slender, cartilaginous, curving anteromedially; anterior processes absent. Alary processes broad, blade-like, cartilaginous lateral extensions directly arising from above anterior margin of hyoid plate. Posterolateral processes long (approximately 55% length of posterior cornua), straight with distal notch. Posterior cornua bony, except distally. Posterior cornua abuts hyoid plate, narrowly separated proximally, separated at angle of 22°. Both proximal and distal ends of posterior cornua undilated.

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Fig. 4.5. Hyoid apparatus of Batrachylodes trossulus, SAM47122, female, ventral. Scale bar = 1 mm.

Firmisternal pectoral girdle robust. Epicoracoids ossified but overlain with cartilage. Omosternum present, unforked. Sternum ossified, well developed, short, approximately as long as broad, with large cartilaginous xiphoid process. Clavicles large, slender, straight. Coracoids well developed, anterior margin concave, posterior margin straight, expanded medially and well separated medially; oriented horizontally. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

Fig. 4.6. Pectoral girdle of Batrachylodes trossulus, SAM47122, female, ventral. Scale bar = 5 mm.

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Eight presacral vertebrae. Neural spines present on Presacrals II, III, decreasing in size anterior to posterior. Transverse processes of Presacrals III and IV moderately expanded distally. Lengths of transverse processes: III > IV > II = V = VI = VII = VIII. Orientation of transverse processes: II, III, VI, VII, VIII horizontal, IV, V angled posteriorly. Sacral diapophyses undilated, cylindrical, with anterior edge oriented horizontally. Urostyle bearing dorsal crest along entire length.

Fig. 4.7. Vertebral column of Batrachylodes trossulus, SAM47122, female, dorsal. Scale bar = 1 mm

Ilia cylindrical, shaft slightly curved comprising approximately 70% of total length of pelvic girdle with distinct crest on shaft (approximately 55% width of shaft, 90% length of shaft). Prominent oval, dorsal protuberance proximally, arising anteriorly to acetabulum. Dorsal acetabular expansion concave, angled approximately 30° to ilial shaft. Ischium well developed, forming posterior and posterodorsal acetabulum. Ventral portion of acetabulum moderately developed with slightly ossified pubis fused with ilium and ischium. Acetabulum large, width approximately 63% of ilium (minus shaft) and ischium; anterior edge of rim approximately 1/2 above the base of ilial shaft.

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Fig. 4.8. Pelvic girdle of Batrachylodes trossulus, SAM47122, female, lateral. Scale bar = 1 mm.

Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements slightly T-shaped. Prepollical elements moderately developed, mostly cartilaginous. Sesamoid bones ventrally at 1st metacarpophalangeal joint. Phalangeal formula of foot: 2-2-3-4-3. Distal tips of phalangeal elements T-shaped larger than fingers. Prehallux well developed, elements cartilaginous. Sesamoid bone at junction of tibiotarsus and ventrally between interphalangeal joints.

Fig. 4.9. Hand of Batrachylodes trossulus, SAM47122, female, dorsal. Scale bar =1 mm

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4.3.1.4 Batrachylodes vertebralis Boulenger, 1887 Figs 4.10 - 4.15 Skull robust, with well-ossified neurocranium. Nasal plate found in males covering sphenethmoid dorsally, merging with it ventrally. Nasal plate absent in females, entire sphenethmoid ossified to level of anterior edge of nasals forming an acuminate apex anteriorly; ossified portion extending 1/2 length of orbit in ventral view. Prootic fused with exoccipital dorsally and ventrally to form otoccipital. Crista parotica well developed but not articulated with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, posteriorly fused with otoccipital, no flanges posterolaterally. Nasals large, broad, juxtaposed medially, overlying bony sphenethmoid. Maxillary process of nasal short, broad, acuminate, lying adjacent to preorbital process of pars facialis of maxilla. Palatines well ossified, long, broad, slightly expanded, rounded medially, terminating on sphenethmoid at a level between anterior extremity of cultriform process of parasphenoid and vomers; lateral expansion overlapped by pars facialis of maxilla. Parasphenoid robust, 2/3 width of frontoparietals; cultriform process truncate terminally, extending anteriorly to level posterior to palatines. Parasphenoid alae broad, horizontal. Pterygoid well developed; anterior ramus extending to level of palatine, ventrally overlapped by pars dentalis of maxilla. Anterior and posterior rami forming a substantial brace; inner margins of rami forming a crescentic curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal, oriented approximately horizontal to midline. Quadratojugal robust, fully articulated. Squamosal well developed. Otic ramus distinct, longer than zygomatic ramus, not articulating with crista parotica. Zygomatic ramus slender. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Premaxilla broad; alary processes long, widely separated. Palatal shelf of premaxilla broad, expanded laterally; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla distinctly V-shaped. Vomers reduced medially, articulating with ventral surface of sphenethmoid; anterior process reduced; posterior process long, spine-like, converging medially; lateral alae reduced, forming anterior and medial margins of choanae; dentigerous processes absent.

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a

b

Fig. 4.10. Skull of Batrachylodes vertebralis, MCZ41848, female. (a) dorsal (b) ventral. Scale bar = 1 mm.

Lateral margin of hyoid plate indented; plate cartilaginous, longer than wide. Anterior cornua slender, cartilaginous, curving anteromedially, expanded at

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midpoint; anterior processes absent. Alary processes curved anteromedially, cartilaginous lateral extensions directly arising from above anterior margin of hyoid plate. Posterolateral processes long (approximately 65% length of posterior cornua), straight, elongate. Posterior cornua bony, except distally. Posterior cornua abuts hyoid plate, narrowly separated proximally, separated at angle of 25°. Both proximal and distal ends of posterior cornua undilated.

Fig. 4.11. Hyoid apparatus of Batrachylodes vertebralis, SAM47110, male, ventral. Scale bar = 1 mm

Firmisternal pectoral girdle robust. Omosternum present, unforked. Sternum ossified, well developed, short, with large cartilaginous xiphoid process. Clavicles large, slender, straight. Coracoids well developed, expanded medially and well separated medially; oriented horizontally or slightly posterolaterally. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

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Fig. 4.12. Pectoral girdle of Batrachylodes vertebralis, MCZ41834, male, ventral. Scale bar = 1 mm.

Eight presacral vertebrae. Neural spines present on Presacrals II, III and IV, decreasing in size anterior to posterior. Transverse processes of Presacrals III and IV moderately expanded distally. Lengths of transverse processes: III > IV > II = V = VI = VII = VIII. Orientation of transverse processes: II angled anteriorly, III, IV, V, VI angled posteriorly, VII and VIII horizontal. Sacral diapophyses undilated, cylindrical, with anterior edge oriented slightly posterolaterally. Urostyle bearing dorsal crest on 2/3 or entire of length.

Fig. 4.13. Vertebral column of Batrachylodes vertebralis, SAM47110, male, dorsal. Scale bar = 1 mm.

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Ilia cylindrical, shaft slightly curved, approximately 70% of total length of pelvic girdle with distinct crest on shaft (approximately 50% width of shaft, 85% length of shaft). Prominent oval, dorsal protuberance proximally, arising approximately 2/3 above acetabulum. Dorsal acetabular expansion concave, angled approximately 30° to ilial shaft. Ischium well developed, bearing prominent crest, forming posterior and posterodorsal acetabulum. Ventral portion of acetabulum well developed with ossified pubis fused with ilium and ischium. Acetabulum large, width approximately 50% of ilium (minus shaft) and ischium; anterior edge of rim approximately 1/3 above the base of ilial shaft.

Fig. 4.14. Pelvic girdle of Batrachylodes vertebralis, SAM47110, male, lateral. Scale bar = 1 mm.

Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements T- shaped. Prepollical elements well developed, ossified. Sesamoid bones ventrally at interphalangeal joints. Phalangeal formula of foot: 2-2-3-4-3. Distal tips of phalangeal elements T-shaped although not as extensive as hand. Prehallux well developed, ossified elements overlain by cartilage.

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a b

Fig. 4.15. Extremities of Batrachylodes vertebralis, SAM47110, male. (a) hand, dorsal (b) foot, dorsal. Scale bar = 1 mm.

4.3.1.5. Batrachylodes wolfi (Sternfeld, 1918) Figs 4.16 - 4.19 Skull robust, with well-ossified neurocranium. Nasal plate found in males covering sphenethmoid dorsally, merging with it ventrally. Nasal plate absent in females, entire sphenethmoid ossified to level of posterior edge of nasals, just below anterior margin of frontoparietals, forming a pointed apex. Ossified portion extending 1/2 length of orbit in ventral view. Prootic fused with exoccipital dorsally and ventrally to form otoccipital. Crista parotica well developed but not articulated with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, anteriorly extend to level of posterior margin of nasals, posteriorly fused with otoccipital, no flanges posterolaterally. Nasals large, broad, juxtaposed medially,

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overlying bony sphenethmoid, nasal plate (in males). Maxillary process of nasal short, broad, acuminate, lying adjacent to preorbital process of pars facialis of maxilla, overlying palatines posteriorly. Palatines well ossified, long, slightly expanded medially, terminating on sphenethmoid at a level between anterior extremity of cultriform process of parasphenoid and vomers; lateral expansion overlapped by pars facialis of maxilla. Parasphenoid robust, 2/3 width of frontoparietals; cultriform process rounded terminally, extending anteriorly to level posterior to palatines. Parasphenoid alae broad, horizontal. Pterygoid well developed; anterior ramus extending to level of palatine. Posterior ramus acuminate; medial ramus articulating with crista parotica via cartilage. Anterior and posterior rami forming a substantial brace; inner margins of rami forming a crescentic curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal, oriented approximately horizontal to midline. Quadratojugal robust and fully articulated. Squamosal well developed. Otic ramus distinct, longer than zygomatic ramus, not articulating with crista parotica. Zygomatic ramus slender, tapering to point. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Premaxilla broad; alary processes long, widely separated. Palatal shelf of premaxilla broad, expanded laterally; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla distinctly V-shaped. Vomers reduced medially, articulating with ventral surface of sphenethmoid; anterior process reduced; posterior process long, spine-like, converging medially; lateral alae reduced, forming anterior and medial margins of choanae; dentigerous processes absent.

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a

b

Fig. 4.16. Skull of Batrachylodes wolfi, SAM8195c, female. (a) dorsal (b) ventral. Scale bar = 1 mm.

Lateral margin of hyoid plate indented; plate cartilaginous, longer than wide. Anterior cornua slender, cartilaginous, curving anteromedially, expanded at

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midpoint; anterior processes absent. Alary processes broad, blade-like, cartilaginous lateral extensions arising from a broad stalk level with anterior margin of hyoid plate. Posterolateral processes moderate (approximately 50% length of posterior cornua), straight, rounded distally. Posterior cornua bony, except distally. Posterior cornua abuts hyoid plate, narrowly separated proximally, separated at angle between 25°- 30°. Proximal end of posterior cornua undilated, distal end dilated.

Fig. 4.17. Hyoid apparatus of Batrachylodes wolfi, SAM4927A, male, ventral. Scale bar =1 mm.

Firmisternal pectoral girdle robust. Omosternum present, unforked. Sternum ossified, well developed, short, with large cartilaginous xiphoid process. Clavicles large, slender, straight. Coracoids well developed, anterior margin concave, posterior margin straight; expanded medially and well separated medially; oriented horizontally or slightly posterolaterally. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

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Fig. 4.18. Pectoral girdle of Batrachylodes wolfi, SAM4927A, male, dorsal. Scale bar = 1 mm.

Eight presacral vertebrae. Neural spines present on Presacral II, III and IV, decreasing in size anterior to posterior. Lengths of transverse processes: III > IV = V = VI = VII = VIII > II. Orientation of transverse processes: III, IV, V, VI angled posteriorly, II, VII, VIII horizontal. Sacral diapophyses undilated, cylindrical, with anterior edge oriented horizontally. Urostyle bearing dorsal crest on 2/3 or entire length. Ilia cylindrical, shaft curved, approximately 70% of total length of pelvic girdle with distinct crest on shaft (approximately 50% width of shaft, 100% length of shaft). Prominent round, dorsal protuberance proximally, arising above acetabulum. Dorsal acetabular expansion concave, angled approximately 30° to ilial shaft. Ischium well developed, and forming posterior and posterodorsal acetabulum. Ventral portion of acetabulum well developed with ossified pubis fused with ilium and ischium. Acetabulum large, width approximately 50% of ilium (minus shaft) and ischium; anterior edge of rim level with base of ilial shaft.

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Fig. 4.19. Pelvic girdle of Batrachylodes wolfi, SAM5164c, male, lateral. Sutures not identifiable. Scale bar = 1 mm.

Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements T- shaped. Prepollical elements well developed, ossified. Sesamoid bones ventrally at metacarpophalangeal joints. Phalangeal formula of foot: 2-2-3-4-3. Distal tips of phalangeal elements T-shaped although not as extensive as hand. Prehallux well developed, ossified elements cartilaginous. Sesamoid bone at junction of tibiotarsus.

4.3.2 Ceratobatrachus Boulenger, 1884

4.3.2.1 Ceratobatrachus guentheri Boulenger, 1884 Figs 4.20 - 4.25 Skull broad, robust, fused, with well-ossified neurocranium. Sphenethmoid ossified to level just below anterior process of vomers, with ossified part extending 1/2 length of orbit in ventral view. Prootic fused with exoccipital dorsally and ventrally to form otoccipital. Crista parotica well developed, long and slender, articulated with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, anteriorly fused with nasals, posteriorly fused with otoccipital. Lateral margins with upturned flange. Nasals large, broad, triangular, fused medially, overlying bony sphenethmoid. Maxillary process of nasal long, narrow, acuminate, overlying preorbital process of pars facialis of maxilla and palatines. Palatines long, broad, expanded laterally, fused to sphenethmoid at a level above anterior extremity of cultriform process of parasphenoid becoming confluent with vomers. Parasphenoid robust, spade

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shaped, constricted posteriorly; cultriform process acuminate terminally, extending anteriorly to level with palatines. Parasphenoid alae tapered, fused to medial rami of pterygoids laterally, horizontal. Pterygoid well developed; anterior ramus extending to level of palatine, fused with maxilla; posterior ramus expanded distally, indented, club-shaped, fused to ventral arm of squamosal and quadratojugal. Anterior and posterior rami forming a substantial brace; inner margins of rami curved between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal, oriented approximately horizontal to midline. Quadratojugal robust, fully articulated becoming confluent with maxilla, spine present on lateral edge. Squamosal well developed. Otic ramus large distinct but shorter than zygomatic ramus. Zygomatic ramus slender, blunt, fused to maxilla. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Maxilla expanded posteriorly. Premaxilla broad; alary processes long, widely separated. Palatal shelf of premaxilla broad, expanded laterally; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla distinctly V-shaped. Vomers large, articulating with ventral surface of sphenethmoid and palatines posteriorly, premaxillaries anteriorly; posterior process large converging medially; lateral alae acuminate. Dentigerous processes long with transverse orientation, bearing large fang-like teeth. Maxillary and premaxillary teeth fang-like, with each tooth borne on a pedestal of bone, curved posteriorly. Mandible dentate.

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a

b

Fig. 4.20. Skull of Ceratobatrachus guentheri, MCZ62226, female. (a) dorsal (b) ventral. Scale bars = 10 mm

Lateral margin of hyoid plate indented; plate mostly cartilaginous, some calcification; longer than wide. Anterior cornua slender, cartilaginous, curving anteromedially; anterior processes present, curved laterally. Alary processes broad, blade-like, cartilaginous lateral extensions arising from a broad stalk level with anterior margin of hyoid plate. Posterolateral processes moderate

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(approximately 50% length of posterior cornua), straight, distally acuminate. Posterior cornua bony. Posterior cornua abuts hyoid plate, almost touching proximally, separated at angle approximately 30°. Proximal end of posterior cornua dilated, distal end undilated.

Fig. 4.21. Hyoid apparatus of Ceratobatrachus guentheri, MCA42118, subadult, ventral. Scale bar = 1 mm.

Firmisternal pectoral girdle very robust. Omosternum present, forked. Sternum ossified, well developed, longer than wide, with large cartilaginous xiphoid process. Epicoracoid cartilage ossified. Clavicles large, oriented horizontally. Coracoids well developed, anterior and posterior margins concave; greatly expanded medially and well separated medially, oriented horizontally. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

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Fig. 4.22. Pectoral girdle of Ceratobatrachus guentheri, MCZ62226, female, ventral. Scale bar = 5 mm.

Eight presacral vertebrae. Neural spines present on Presacral II, III and IV, decreasing in size anterior to posterior. Lengths of transverse processes: III > IV > V = VI = VII = VIII > II. Orientation of transverse processes: III, IV, V, VI angled posteriorly, II, VII, VIII horizontal. Sacral diapophyses undilated, rounded, with anterior edge oriented posterolaterally. Urostyle bearing dorsal crest along entire length.

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Fig. 4.23. Vertebral column of Ceratobatrachus guentheri, MCZ62226, female, dorsal. Scale bar = 10 mm

Ilia tapered, shaft curved, approximately 75% of total length of pelvic girdle with crest on shaft (approximately 30% width of shaft, 90% length of shaft). Prominent round, dorsal protuberance proximally, level with anterior rim of acetabulum. Dorsal acetabular expansion straight, angled approximately 30° to ilial shaft. Ischium well developed, forming posterior and posterodorsal acetabulum. Ventral portion of acetabulum well developed with ossified pubis fused with ilium and ischium. Acetabulum large, width approximately 70% of ilium (minus shaft) and ischium; anterior edge of rim below the base of ilial shaft.

Fig. 4.24. Pelvic girdle of Ceratobatrachus guentheri, MCZ62226, female, lateral. Sutures between ischium and pubis not identifiable. Scale bar = 10 mm

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Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements obtuse. Prepollical elements well developed, ossified. Sesamoid bones ventrally at metacarpophalangeal joints, at proximal and middle interphalangeal joints on digits III, IV. Phalangeal formula of foot: 2-2-3-4-3. Distal tips of phalangeal elements obtuse. Prehallux well developed, ossified. Sesamoid bones at junction of tibiotarsus, ventrally at metacarpophalangeal joint of toes II, III, IV; proximal and middle interphalangeal joints on digits III, IV, V; middle and distal interphalangeal joints on digits IV.

a b

Fig 4.25. Extremities of Ceratobatrachus guentheri. MCZ62226, female. (a) hand, dorsal. Scale bar = 5 mm (b) foot, dorsal. Scale bar = 10 mm.

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4.3.3 Discodeles Boulenger, 1918

4.3.3.1 Discodeles bufoniformis (Boulenger, 1884) Figs 4.26 - 4.31 Skull robust, broad, with well-ossified neurocranium. Sphenethmoid ossified to level of anterior margins of palatines. Ossified part extending 1/2 length of orbit in ventral view. Prootic fused with exoccipital dorsally and ventrally to form otoccipital. Crista parotica well developed articulating with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, anteriorly extend to posterior margin of nasals, posteriorly fused with otoccipital, no flanges posterolaterally. Nasals large, broad, triangular, juxtaposed medially, overlying anterior margin of bony sphenethmoid. Maxillary process of nasal long, broad, acuminate, lying adjacent to preorbital process of pars facialis of maxilla, overlying palatines. Palatines well ossified, long, broad, expanded distally, tapering to a point medially, terminating on sphenethmoid at a level between anterior extremity of cultriform process of parasphenoid, fused to vomers, especially dentigerous process. Lateral expansion overlapped by preorbital process of maxilla. Parasphenoid robust, 2/3 width of frontoparietals; cultriform process usually rounded terminally, extending anteriorly to level posterior to palatines. Parasphenoid alae broad but tapering laterally, oriented horizontally. Pterygoid well developed. Flange occurring on medial ramus. Medial ramus juxtaposed with crista parotica. Anterior ramus extending 1/2 length of orbit, ventrally overlapped by pars dentalis of maxilla. Anterior and posterior rami forming a substantial brace; inner margins of rami forming a crescentic curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal, oriented approximately horizontally to midline. Quadratojugal robust, fully articulated. Squamosal well developed. Otic ramus distinct, equal in length to zygomatic ramus, articulating with crista parotica. Zygomatic ramus broad, tapering to point. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Premaxilla broad; alary processes long, widely separated. Palatal shelf of premaxilla narrow; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla U-shaped. Vomers large, articulating with ventral surface of sphenethmoid and palatines; anterior process acuminate; posterior process broad

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triangular converging medially; lateral alae large, forming anterior and medial margins of choanae; dentigerous processes present, crescent shaped, orientated obliquely, short bearing 6-7 large fang-like teeth.

a

b

Fig. 4.26. Skull of Discodeles bufoniformis, CAS109895. (a) dorsal (b) ventral. Scale bars = 10 mm. Lateral margin of hyoid plate indented; plate mostly cartilaginous but calcification present; longer than wide. Anterior cornua slender, cartilaginous,

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curving anteromedially; anterior processes present, curved with a distinct lateral branch. Alary processes club-shaped, cartilaginous lateral extensions arising from a broad stalk level with anterior margin of hyoid plate. Posterolateral processes long (approximately 65% length of posterior cornua), curved, tapering to a point. Posterior cornua bony, except distally. Posterior cornua abuts hyoid plate, narrowly separated proximally, separated at angle between 18-23°. Proximal end of posterior cornua expanded, distal end undilated.

Fig. 4.27. Hyoid apparatus of Discodeles bufoniformis, MCZ38581, female, ventral. Scale bar = 5 mm

Firmisternal pectoral girdle robust. Epicoracoids bony, fused. Omosternum present, forked. Sternum ossified, well developed, longer than wide, with large cartilaginous xiphoid process containing some calcification. Clavicles large, slender, straight, tapering sharply. Coracoids well developed, expanded medially and well separated medially, hourglass shaped, oriented horizontally. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

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Fig. 4.28. Pectoral girdle of Discodeles bufoniformis, CAS109895, ventral. Scale bar = 5 mm

Eight presacral vertebrae. Neural spines present on all presacrals, decreasing in size anterior to posterior. Transverse processes of Presacrals II, III and IV moderately expanded distally. Lengths of transverse processes: III > IV > V = VI = VII = VIII > II. Orientation of transverse processes: II angled anteriorly, III, IV, V, VI angled posteriorly, VII and VIII horizontal. Sacral diapophyses undilated, cylindrical, with anterior edge oriented posterolaterally. Urostyle bearing dorsal crest along entire length.

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Fig. 4.29. Vertebral column of Discodeles bufoniformis, CAS109985, dorsal. Scale bar = 10 mm

Ilia cylindrical, shaft straight comprising approximately 60% of total length of pelvic girdle with distinct crest on shaft (approximately 55% width of shaft, 100% length of shaft). Prominent large round, dorsal protuberance proximally, arising approximately 1/2 above acetabulum. Dorsal acetabular expansion concave, angled approximately 32° to ilial shaft. Ischium well developed, forming posterior and posterodorsal acetabulum. Ventral portion of acetabulum well developed with ossified pubis fused with ilium and ischium. Acetabulum large, width approximately 45% of ilium (minus shaft) and ischium; anterior edge of rim below the base of ilial shaft.

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Fig. 4.30. Pelvic girdle of Discodeles bufoniformis, MCZ38578, male, lateral. Scale bar = 5 mm.

Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements obtuse. Prepollical elements well developed, cartilaginous. Phalangeal formula of foot: 2-2-3-4-3. Distal tips of phalangeal elements obtuse. Prehallux well developed, cartilaginous.

Fig. 4.31. Hand of Discodeles bufoniformis, CAS109985, dorsal. Scale bar = 10 mm.

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4.3.3.2 Discodeles guppyi (Boulenger, 1884) Figs 4.32 - 4.37 Skull robust, broad, with well-ossified neurocranium. Sphenethmoid ossified to level of posterior edge of nasals fused to palatines dorsally; ossified portion extending 1/2 length of orbit in ventral view. Prootic fused with exoccipital dorsally but not ventrally to form otoccipital. Crista parotica well developed articulating with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, anteriorly reaching posterior margin on nasals, posteriorly fused with otoccipital; no flanges posterolaterally. Nasals large, broad, triangular, juxtaposed medially, overlying bony sphenethmoid. Maxillary process of nasal long, broad, acuminate, lying adjacent to preorbital process of pars facialis of maxilla and overlapping palatines. Palatines well ossified, long, broad, become confluent with sphenethmoid at medial margin; terminating on sphenethmoid at a level between anterior extremity of cultriform process of parasphenoid; fused with vomers. Parasphenoid robust, 1/2 width of frontoparietals; cultriform process usually acuminate terminally, extending anteriorly to level posterior to palatines. Constricted posteriorly forming a distinct diamond process overlying centre of alae. Parasphenoid alae broad, horizontal. Pterygoid well developed; anterior ramus extending to level of palatine, ventrally overlapped by pars dentalis of maxilla. Medial ramus articulating with crista parotica. Posterior ramus tapering to point. Anterior and posterior rami forming a substantial brace; inner margins of rami forming a crescentic curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal, oriented approximately horizontal to midline. Quadratojugal robust, fully articulated. Squamosal well developed. Otic ramus distinct, equal in length to zygomatic ramus, overlying and fused with crista parotica. Zygomatic ramus broad, elongate, ovoid. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Premaxilla broad; alary processes long, widely separated. Palatal shelf of premaxilla broad; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla U-shaped. Vomers large, articulating with ventral surface of sphenethmoid, fused to palatines; anterior process large, acuminate; posterior process large, broad converging medially bearing large dentigerous process. Lateral alae large, Posterolateral alae acuminate. Dentigerous processes present, large, curved, short bearing approximately seven teeth.

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a

b

Fig. 4.32. Skull of Discodeles guppyi, SAM47092, male. (a) dorsal (b) ventral. Scale bar = 10 mm Lateral margin of hyoid plate indented; plate cartilaginous, longer than wide. Anterior cornua slender, cartilaginous, curving anteromedially; anterior processes

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present, curved laterally. Alary processes broad, club-shaped, cartilaginous lateral extensions directly arising level with anterior margin of hyoid plate. Posterolateral processes moderate (approximately 35% length of posterior cornua), curved, acuminate. Posterior cornua bony, except distally. Posterior cornua abut hyoid plate, narrowly separated proximally, separated at angle of 20°. Proximal ends of posterior cornua expanded, distal ends undilated.

Fig. 4.33. Hyoid apparatus of Discodeles guppyi, SAM47091, female, ventral. Scale bar = 10 mm.

Firmisternal pectoral girdle robust. Epicoracoids ossified. Omosternum present, forked. Sternum ossified, well developed, longer than wide, with cartilaginous xiphoid process. Clavicles large, straight, tapering to a point. Coracoids well developed, expanded medially and well separated medially, hourglass shape, oriented posterolaterally. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

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Fig. 4.34. Pectoral girdle of Discodeles guppyi, SAM47092, male, ventral. Scale bar = 5 mm.

Eight presacral vertebrae. Neural spines present on all presacrals, decreasing in size anterior to posterior. Transverse processes of Presacrals III and IV expanded distally. Lengths of transverse processes: III > IV > V = VI = VII = VIII > II. Orientation of transverse processes: III, IV, V, VI, VII angled posteriorly, II, VIII horizontal. Sacral diapophyses undilated, cylindrical, with anterior edge oriented slightly posterolaterally. Urostyle bearing dorsal crest along entire length.

118

Fig. 4.35. Vertebral column of Discodeles guppyi, SAM47092, male, dorsal. Scale bar = 10 mm.

Ilia cylindrical, shaft straight comprising approximately 68% of total length of pelvic girdle with distinct crest on shaft (approximately 55% width of shaft, 100% length of shaft). Prominent round, dorsal protuberance proximally, arising approximately 2/3 above acetabulum. Dorsal acetabular expansion sigmoidal, angled approximately 27° to ilial shaft. Ischium well developed, forming posterior and posterodorsal acetabulum. Ventral portion of acetabulum well developed with ossified pubis fused with ilium and ischium. Acetabulum moderate, width approximately 42% of ilium (minus shaft) and ischium; anterior edge of rim level with base of ilial shaft.

Fig. 4.36. Pelvic girdle of Discodeles guppyi, SAM47092, male, lateral. Scale bar = 10 mm

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Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements obtuse. Prepollical elements well developed, mostly cartilaginous, some ossification. Phalangeal formula of foot: 2-2-3-4-3. Distal tips of phalangeal elements obtuse. Prehallux well developed, slightly ossified. Very small sesamoid bone at junction of tibiotarsus.

Fig. 4.37. Hand of Discodeles guppyi, SAM47092, male, dorsal. Scale bar = 10 mm.

4.3.4. Palmatorappia Ahl, 1927

4.3.4.1. Palmatorappia solomonis (Sternfeld, 1920) Figs 4.38 - 4.43 Skull moderate, broad, with poorly ossified neurocranium. Sphenethmoid poorly ossified but reaching a level to posterior process of vomers; widely separated into two halves. Ossified part extending 1/3 length of orbit in ventral view. Crista parotica not well developed, prootic and exoccipital not confluent, not articulated with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, medially separated; anteriorly extend to well below nasals, posteriorly fused with otoccipital, no flanges posterolaterally. Nasals moderate, broad, bulbous medially, well separated medially, not overlying sphenethmoid. Maxillary process of nasal long, slender,

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acuminate, reaching preorbital process of pars facialis of maxilla, just overlapping palatines. Palatines well ossified, long, slender tapering medially, terminating and fused with sphenethmoid at a level between anterior extremity of cultriform process of parasphenoid and vomers; lateral end expanded. Parasphenoid robust, 2/3 width of frontoparietals; cultriform process truncate terminally, extending anteriorly to midway between sphenethmoid, well below palatines. Parasphenoid alae broad, cylindrical, oriented slightly posterolaterally. Pterygoid moderately developed; anterior ramus extending just posteriorly to palatine, acuminate, ventrally overlapped by pars dentalis of maxilla for a considerable distance. Posterior ramus tapering to a fine point. Medial ramus juxtaposed with or separate from crista parotica. Anterior and posterior rami forming a brace; inner margins of rami forming a crescentic curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal. Quadratojugal slender, anteriorly not articulated with maxilla, posteriorly articulating via a spur to squamosal. Squamosal well developed. Otic ramus distinct, longer than zygomatic ramus, tapering sharply, not articulating with crista parotica. Zygomatic ramus slender, ending in a rounded knob. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla; articulating via cartilage to palatine. Premaxilla broad; alary processes long, widely separated. Palatal shelf of premaxilla broad; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla V-shaped. Vomers reduced medially, articulating with anterior margin of ventral surface of sphenethmoid; anterior process reduced; posterior process long, broad converging medially; lateral alae acuminate, forming anterior and medial margins of choanae; dentigerous processes absent.

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a

b

Fig. 4.38. Skull of Palmatorappia solomonis, MCZ60864, male. (a) dorsal (b) ventral. Scale bar = 1 mm.

Lateral margin of hyoid plate is straight; plate cartilaginous, slightly longer than wide. Anterior cornua slender, cartilaginous; anterior processes absent. Alary processes short, acuminate, flat, cartilaginous anterior extensions directly arising from above anterior margin of hyoid plate. Posterolateral processes absent.

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Posterior cornua bony, except distally. Posterior cornua abuts hyoid plate, narrowly separated proximally, separated at angle of 20°. Both proximal and distal ends of posterior cornua undilated.

Fig. 4.39. Hyoid apparatus of Palmatorappia solomonis, MCZ60858, female, ventral. Scale bar = 1 mm.

Firmisternal pectoral girdle moderate. Epicoracoids mostly cartilaginous, slight ossification. Omosternum present, forked. Sternum ossified, slender, short, longer than wide, constricted in centre, with cartilaginous xiphoid process. Clavicles slender, oriented posterolaterally, poorly ossified, tapering sharply. Coracoids well developed, expanded medially, well separated medially, slightly hourglass shaped, oriented horizontally. Scapula fused with clavicle (via cartilage), coracoid and suprascapula.

123

Fig. 4.40. Pectoral girdle of Palmatorappia solomonis, MCZ60863, male, ventral. Scale bar = 1 mm.

Eight presacral vertebrae. Neural spines absent. Transverse processes of Presacrals II, III moderately expanded distally. Lengths of transverse processes: III > IV = V = VI = VII = VIII > II. Orientation of transverse processes: II, III angled anteriorly, IV, V, VI angled posteriorly, VII and VIII horizontal. Sacral diapophyses undilated, cylindrical, with anterior edge oriented posterolaterally. Urostyle bearing dorsal crest 2/3 of length.

Fig. 4.41. Vertebral column of Palmatorappia solomonis, MCZ60864, male, dorsal. Scale bar = 1 mm.

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Ilia cylindrical, shaft slightly curved, approximately 75% of total length of pelvic girdle; crest absent. Prominent oval, dorsal protuberance proximally, arising approximately 2/3 above acetabulum. Dorsal acetabular expansion straight, angled approximately 35° to ilial shaft. Ischium well developed, forming posterior and posterodorsal acetabulum. Ventral portion of acetabulum well developed with ossified pubis fused with ilium and ischium. Acetabulum moderate, width approximately 38% of ilium (minus shaft) and ischium; anterior edge of rim approximately 2/3 above the base of ilial shaft.

Fig. 4.42. Pelvic girdle of Palmatorappia solomonis, MCZ60864, male, lateral. Scale bar = 1 mm.

Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements Y- shaped. Prepollical elements well developed, mostly cartilage, slight ossification. Phalangeal formula of foot: 2-2-3-4-3. Distal tips of phalangeal elements T- shaped, not as extensive as hand. Prehallux cartilaginous. Sesamoid bone at junction of tibiotarsus.

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a b

Fig. 4.43. Extremities of Palmatorappia solomonis, MCZ60864, male. (a) hand, dorsal (b) foot, dorsal. Scale bars = 1 mm.

4.3.5 Platymantis Günther, 1859 "1858"

4.3.5.1 Platymantis guppyi (Boulenger, 1884)

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4.3.5 Platymantis Günther, 1859 "1858"

4.3.5.1 Platymantis guppyi (Boulenger, 1884) Figs 4.44 - 4.49 Skull robust, broad, more so in females, with well-ossified neurocranium. Sphenethmoid ossified to level of centre of nasals forming an acuminate apex anteriorly; ossified portion extending half to 2/3 length of orbit in ventral view. Prootic fused with exoccipital dorsally but not ventrally to form otoccipital. Crista parotica well developed articulating with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, separated anteriorly, expanded and diverging to level of posterior margin of nasals, posteriorly fused with otoccipital, no flanges posterolaterally. Nasals large, broad, triangular, juxtaposed medially, overlying bony sphenethmoid. Maxillary process of nasal long, slender, acuminate, lying adjacent to or slightly overlapping preorbital process of pars facialis of maxilla, overlapping palatines that reach

posterior margin. Palatines well ossified, long, expanded laterally, tapering medially, terminating on sphenethmoid at a level between anterior extremity of cultriform process of parasphenoid and posterior to vomers. Parasphenoid robust, constricted posteriorly, half width of frontoparietals; cultriform process truncate terminally, extending anteriorly to almost reach palatines. Parasphenoid alae broad, horizontal. Pterygoid well developed; anterior ramus bifurcated, extending to level of palatine, ventrally overlapped by palatal shelf of maxilla. Medial ramus broad, fused to crista parotica. Anterior and posterior rami forming a substantial brace; inner margins of rami forming a crescentic curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal oriented approximately parallel to midline. Quadratojugal robust, fully articulated. Squamosal well developed. Otic ramus distinct with distinct shelf (flange), longer than zygomatic ramus, fused with crista parotica. Zygomatic ramus slender, curved. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Palatal shelf broad. Maxilla with cartilaginous connection with palatine. Maxilla articulating with pars dentalis of premaxilla. Premaxilla broad; alary processes long, separated medially. Palatal shelf of premaxilla broad; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla U-shaped. Vomers unreduced, overlying ventral surface of sphenethmoid; anterior process large, acuminate; posterior process large, broad converging medially; lateral alae large, broad, forming anterior and medial margins of choanae; dentigerous processes present, orientation transverse or slightly oblique, short with approximately six teeth.

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a

b

Fig. 4.44. Skull of Platymantis guppyi, SAM47128, female. (a) dorsal (b) ventral. Scale bar = 10 mm.

Lateral margins of hyoid plate straight; plate cartilaginous, longer than wide. Anterior cornua slender, cartilaginous, curving anteromedially; anterior processes curved, slender. Alary processes T-shaped, flat, cartilaginous lateral extensions arising via a stalk from above anterior margin of hyoid plate. Posterolateral

128

processes short (approximately 36% length of posterior cornua), acuminate. Posterior cornua bony, except distally. Posterior cornua abuts hyoid plate, narrowly separated proximally, separated at angle of 33°. Both proximal and distal ends of posterior cornua undilated.

Fig. 4.45. Hyoid apparatus of Platymantis guppyi, SAM47128, female, ventral. Scale bar = 10 mm.

Firmisternal pectoral girdle robust. Epicoracoid cartilages ossified. Omosternum present, forked. Sternum ossified, well developed, longer than wide, with large cartilaginous xiphoid process. Clavicles large, slender, tapering sharply, oriented horizontally. Coracoids well developed, expanded medially and well separated medially, hourglass shaped, oriented horizontally. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

129

Fig. 4.46. Pectoral girdle of Platymantis guppyi, SAM47128, female, ventral. Scale bar = 5 mm.

Eight presacral vertebrae. Neural spines present on Presacrals II to VII, decreasing in size, anterior to posterior. Transverse processes of Presacrals III and IV moderately expanded distally. Lengths of transverse processes: III > IV > V = VI = VII = VIII > II. Orientation of transverse processes: III, IV, V, VI, VII angled posteriorly, II and VIII horizontal. Sacral diapophyses undilated, cylindrical, with anterior edge oriented posterolaterally. Urostyle bearing dorsal crest along entire length.

Fig. 4.47. Vertebral column of Platymantis guppyi, SAM47128, female, dorsal. Scale bar = 10 mm.

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Ilia cylindrical, shaft straight comprising approximately 73% of total length of pelvic girdle with distinct crest on shaft (approximately 40% width of shaft, 94% length of shaft). Prominent round, dorsal protuberance proximally, arising approximately half above acetabulum. Dorsal acetabular expansion concave, angled approximately 15° to ilial shaft. Ischium well developed, although not fused to ilium, bearing a slight crest anteriorly, forming posterior acetabulum. Ventral portion of acetabulum well developed with ossified pubis fused with ilium and ischium. Acetabulum large, width approximately 46% of ilium (minus shaft) and ischium; anterior edge of rim approximately level with base of ilial shaft.

Fig. 4.48. Pelvic girdle of Platymantis guppyi, SAM47128, female, lateral. Scale bar = 10 mm.

Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements T- shaped, almost Y-shaped on digits 3, 4. Prepollical elements slightly ossified. Sesamoid bones ventrally at interphalangeal joints. Phalangeal formula of foot: 2- 2-3-4-3. Distal tips of phalangeal elements T-shaped on digits 1,2; Y-shaped on 3,4,5. Prehallux well developed, cartilaginous.

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a b

Fig. 4.49. Extremities of Platymantis guppyi, SAM47128, female. (a) hand, dorsal (b) foot, dorsal. Scale bars = 10 mm.

4.3.5.2 Platymantis myersi Brown, 1949 Figs 4.50 - 4.55 Skull robust, longer than wide, with moderately ossified neurocranium. Sphenethmoid ossified to level of posterior margin of vomers. Ossified portion extending less than 1/3 length of orbit in ventral view. Prootic not fused with exoccipital dorsally or ventrally to form otoccipital. Crista parotica well developed but not articulated with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, anteriorly extending to anterior margin of palatines, posteriorly fused with otoccipital, no flanges posterolaterally. Nasals large, broad, triangular, juxtaposed medially, overlying bony sphenethmoid. Maxillary process of nasal long, broad, acuminate, lying adjacent to preorbital process of pars facialis of maxilla, overlapping palatines reaching posterior margin. Palatines well ossified, long, broad, truncate medially, terminating on sphenethmoid, fused to vomers. Parasphenoid robust, half width of frontoparietals; cultriform process acuminate terminally, extending anteriorly to level anterior to palatines. Parasphenoid alae moderate, horizontal. Pterygoid

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moderately developed; anterior ramus tapering to a fine point, extending just posteriorly to level of palatine, ventrally overlapped by pars dentalis of maxilla. Medial ramus short articulating with crista parotica. Anterior and posterior rami forming a brace; inner margins of rami forming a curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal oriented approximately parallel to midline. Quadratojugal robust, fully articulated. Squamosal moderately developed. Otic ramus distinct, longer than zygomatic ramus, not articulating with crista parotica. Zygomatic ramus slender, tapering to point. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Alary processes of premaxilla short, separated. Palatal shelf of premaxilla broad; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla U-shaped. Vomers large, articulating with ventral surface of sphenethmoid, fused to anterior margin of medial aspect of palatine; anterior process slender, acuminate; posterior process large, broad converging medially; lateral alae reduced, acuminate, forming anterior and medial margins of choanae; dentigerous processes present, obliquely oriented, short with approximately 5-6 teeth.

a

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b

Fig. 4.50. Skull of Platymantis myersi, SAM47157, female. (a) dorsal (b) ventral. Scale bar = 5 mm.

Lateral margins of hyoid plate straight; plate cartilaginous, longer than wide. Anterior cornua slender, cartilaginous, curving anteromedially. anterior processes present, curve anterolaterally. Alary processes blade or shield-like, cartilaginous lateral extensions arising via a stalk from above anterior margin of hyoid plate. Posterolateral processes short, acuminate. Posterior cornua bony, except distally. Posterior cornua abuts hyoid plate, narrowly separated proximally almost touching, separated at angle of 32°. Both proximal and distal ends of posterior cornua undilated.

134

Fig. 4.51. Hyoid apparatus of Platymantis myersi, SAM47157, female, ventral. Scale bar = 5 mm.

Firmisternal pectoral girdle robust. Epicoracoids ossified. Cleithrum present, ossified. Omosternum present, forked. Sternum ossified, well developed, slender tapering distally, much longer than wide, with large cartilaginous xiphoid process. Clavicles large, slender, oriented horizontally. Coracoids well developed, hourglass-shaped, expanded medially and separated medially; oriented horizontal. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

Fig. 4.52. Pectoral girdle of Platymantis myersi, SAM47157, female, ventral. Scale bar = 1 mm.

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Eight presacral vertebrae. Neural spines present on Presacrals II to VII, decreasing in size anterior to posterior, very large on II. Transverse processes of Presacral III moderately expanded distally. Lengths of transverse processes: III > IV > V > II = VI = VII > VIII. Orientation of transverse processes: IV, V, VI, VII angled posteriorly, II, III and VIII horizontal. Sacral diapophyses undilated, cylindrical, with anterior edge oriented posterolaterally. Urostyle bearing dorsal crest along entire length.

Fig. 4.53. Vertebral column of Platymantis myersi, SAM47157, female, dorsal. Scale bar = 5 mm.

Ilia cylindrical, shaft straight comprising approximately 67% of total length of pelvic girdle with distinct crest on shaft (approximately 72% width of shaft, 90% length of shaft). Prominent round, dorsal protuberance proximally, arising approximately half above acetabulum. Dorsal acetabular expansion concave, angled approximately 26° to ilial shaft. Ischium well developed, although not fused to ilium, forming posterior acetabulum. Ventral portion of acetabulum well developed with ossified pubis fused with ilium and ischium. Acetabulum large, width approximately 50% of ilium (minus shaft) and ischium; anterior edge of rim approximately 1/3 above base of ilial shaft.

136

Fig. 4.54. Pelvic girdle of Platymantis myersi, SAM47157, female, lateral. Scale bar = 5 mm.

Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements obtuse. Prepollical elements slightly ossified. Phalangeal formula of foot: 2-2-3-4- 3. Distal tips of phalangeal elements obtuse. Prehallux well developed, cartilaginous.

a b Fig. 4.55. Extremities of Platymantis myersi, SAM47157, female. (a) hand, dorsal. Scale bar = 1 mm (b) foot, dorsal. Scale bar = 5 mm.

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4.3.5.3 Platymantis neckeri (Brown and Myers, 1949) Figs 4.56 - 4.61 Skull robust, longer than wide, with well-ossified neurocranium. Sphenethmoid ossified to level of approximately 1/3 above posterior margin of vomers. Ossified part extending half length of orbit in ventral view. Prootic fused with exoccipital dorsally but not ventrally to form otoccipital. Crista parotica well developed juxtaposed with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, anteriorly truncate, extending to posterior margin of palatines; posteriorly fused with otoccipital, no flanges posterolaterally. Nasals large, broad, narrowly separated medially, not overlying bony sphenethmoid. Maxillary process of nasal long, broad, acuminate, lying adjacent to preorbital process of pars facialis of maxilla. Palatines well ossified, long, broad, overlying sphenethmoid, fused to vomers; lateral end slightly expanded. Parasphenoid robust, half width of frontoparietals; cultriform process acuminate terminally, extending anteriorly to level anterior to palatines. Parasphenoid alae moderately lying horizontal. Pterygoid moderately developed; anterior ramus tapering to a point, extending just posteriorly to level of palatine, ventrally overlapped by pars dentalis of maxilla. Medial ramus slender articulating with crista parotica. Posterior ramus acuminate. Anterior and posterior rami forming a brace; inner margins of rami forming a curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal oriented approximately parallel to midline. Quadratojugal robust, fully articulated. Squamosal well developed. Otic ramus distinct, longer than zygomatic ramus, not articulating with but juxtaposed with crista parotica. Zygomatic ramus broad, tapering to point. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Premaxilla broad; alary processes long, separated medially. Palatal shelf of premaxilla broad; palatine process acuminate. Vomers large, articulating with ventral surface of sphenethmoid, fused to anterior margin of medial aspect of palatine; anterior process broad, acuminate; posterior process large, broad converging medially; lateral alae acuminate, anterior lateral alae broad, larger than posterior, forming anterior and medial margins of choanae; dentigerous processes present, obliquely oriented, short with approximately 7-8 teeth.

138

a

b

Fig. 4.56. Skull of Platymantis neckeri, SAM4926D, male. (a) dorsal (b) ventral. Scale bars = 5 mm.

Lateral margins of hyoid plate straight; plate cartilaginous, longer than wide. Anterior cornua slender, cartilaginous, curving anteromedially; anterior processes present, curved laterally, almost forming a complete arch. Alary processes curved,

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acuminate, flat, cartilaginous lateral extensions arising directly from and above anterior margin of hyoid plate. Posterolateral processes short (approximately 35% length of posterior cornua), triangular, acuminate. Posterior cornua bony, except distally. Posterior cornua directly abuts hyoid plate, narrowly separated proximally almost touching, separated at angle of 43°. Proximal ends of posterior cornua expanded, distal ends undilated.

Fig. 4.57. Hyoid apparatus of Platymantis neckeri, SAM4928B, male, ventral. Scale bar = 5 mm.

Firmisternal pectoral girdle robust. Epicoracoids ossified. Cleithrum present, ossified. Omosternum present, forked. Sternum ossified, well developed, slender tapering distally, much longer than wide, with large cartilaginous xiphoid process. Clavicles large, slender, oriented posterolaterally. Coracoids well developed, hourglass shaped, expanded medially and separated medially; oriented posterolaterally. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

140

Fig. 4.58. Pectoral girdle of Platymantis neckeri, SAM4926D, male, ventral. Scale bar = 1 mm.

Eight presacral vertebrae. Neural spines present on Presacrals II, III, IV, decreasing in size anterior to posterior. Transverse processes of Presacral III, IV moderately expanded distally. Lengths of transverse processes: III > IV > II> V = VI = VII = VIII. Orientation of transverse processes: II, III slightly angled anteriorly; IV, V, VI, VII, VIII angled posteriorly. Sacral diapophyses undilated, cylindrical, with anterior edge oriented posterolaterally. Urostyle bearing dorsal crest along entire length.

141

Fig. 4.59. Vertebral column of Platymantis neckeri, SAM4926D, male, dorsal. Scale bar = 5 mm.

Ilia cylindrical, shaft straight comprising approximately 72% of total length of pelvic girdle with distinct crest on shaft (approximately 50% width of shaft, 95% length of shaft). Prominent round, dorsal protuberance proximally, arising approximately half above acetabulum. Dorsal acetabular expansion straight, angled approximately 20° to ilial shaft. Ischium well developed, forming posterior acetabulum. Ventral portion of acetabulum well developed with ossified pubis fused with ilium and ischium. Acetabulum large, width approximately 53% of ilium (minus shaft) and ischium; anterior edge of rim approximately level with base of ilial shaft.

Fig. 4.60. Pelvic girdle of Platymantis neckeri, SAM4926D, male, lateral. Scale bar = 5 mm.

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Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements T- shaped. Prepollical elements ossified. Sesamoid bones present in pairs ventrally at metacarpophalangeal joints. Phalangeal formula of foot: 2-2-3-4-3. Distal tips of phalangeal elements T-shaped although not as extensive as hands. Prehallux well developed, three elements ossified, two cartilaginous. Sesamoid bones absent from toes but present at junction of tibiotarsus.

a b

Fig. 4.61. Extremities of Platymantis neckeri, SAM4926D, male. (a) hand, dorsal. Scale bar = 1 mm (b) foot, dorsal. Scale bar = 5 mm.

4.3.5.4 Platymantis parkeri (Brown, 1965) Figs 4.62 - 4.67 Skull moderate, broad, with moderately ossified neurocranium. Sphenethmoid ossified to level of posterior margin of vomers. Ossified part extending 1/3 length of orbit in ventral view. Prootic not fused with exoccipital dorsally or ventrally to form otoccipital. Crista parotica short, broad developed but not articulating with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, anteriorly acuminate, extending to posterior margin of nasals; posteriorly fused with otoccipital, no flanges posterolaterally. Nasals large, broad, narrowly separated medially, not overlying

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bony sphenethmoid. Maxillary process of nasal long, acuminate, lying adjacent to preorbital process of pars facialis of maxilla, overlapping palatines reaching posterior margin. Palatines well ossified, long, curved, tapering medially, terminating on sphenethmoid. Parasphenoid robust, 1/3 to half width of frontoparietals; cultriform process acuminate terminally, extending anteriorly to level posterior to palatines. Parasphenoid alae broad, horizontal. Pterygoid moderately developed; anterior ramus tapering to a point, extending 2/3 length of orbit, well below level of palatine, ventrally overlapped by pars dentalis of maxilla. Medial ramus short articulating with and fused with crista parotica. Posterior ramus acuminate. Anterior and posterior rami forming a brace; inner margins of rami forming a curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal oriented approximately parallel to midline. Quadratojugal slender and fully articulated. Squamosal well developed. Otic ramus distinct, longer than zygomatic ramus, not articulating with crista parotica. Zygomatic ramus slender, tapering to point. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Premaxilla broad; alary processes long, separated medially. Palatal shelf of premaxilla broad, expanded laterally; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla forming a broad V-shaped. Vomers moderate, overlying the sphenethmoid; anterior process reduced, triangular; posterior process moderate, triangular converging medially; lateral alae acuminate. Dentigerous processes present, very obliquely oriented, very short with approximately 2-3 teeth.

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a

b

Fig. 4.62. Skull of Platymantis parkeri, SAM5169B, female. (a) dorsal (b) ventral. Scale bars = 1 mm.

Lateral margins of hyoid plate indented; plate cartilaginous, longer than wide. Anterior cornua slender, cartilaginous, curving anteromedially; anterior processes present, curved posterolaterally, almost forming a complete arch. Alary processes

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broad, blade-like, cartilaginous lateral extensions directly arising from and below anterior margin of hyoid plate. Posterolateral processes short (approximately 32% length of posterior cornua), triangular, acuminate. Posterior cornua bony, except distally. Posterior cornua directly abuts hyoid plate, narrowly separated proximally, separated at angle of 34°. Proximal ends of posterior cornua expanded, distal ends undilated.

Fig. 4.63. Hyoid apparatus of Platymantis parkeri, SAM5169B, female, ventral. Scale bar = 1 mm.

Firmisternal pectoral girdle robust. Epicoracoids ossified. Cleithrum present, ossified. Omosternum present, forked. Sternum ossified, well developed, short, proximal end expanded, tapering distally, with large cartilaginous xiphoid process. Clavicles large, slender, oriented posterolaterally. Coracoids well developed, hourglass shaped, expanded medially and separated medially; oriented posteromedially. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

146

Fig. 4.64. Pectoral girdle of Platymantis parkeri, SAM5169B, female, ventral. Scale bar = 1 mm.

Eight presacral vertebrae. Neural spines present on Presacrals II, III, IV, decreasing in size anterior to posterior. Transverse processes of Presacral III, IV moderately expanded distally. Lengths of transverse processes: III > II > IV > V = VI = VII = VIII. Orientation of transverse processes: II, angled anteriorly; IV, V, VI angled posteriorly, III, VII, VIII horizontal. Sacral diapophyses undilated, cylindrical, with anterior edge oriented posterolaterally. Urostyle bearing dorsal crest along 2/3 of its length.

Fig. 4.65. Vertebral column of Platymantis parkeri, SAM5169B, female, dorsal. Scale bar = 1 mm.

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Ilia cylindrical, shaft almost straight comprising approximately 75% of total length of pelvic girdle with distinct crest on shaft (approximately 60% width of shaft, 90% length of shaft). Prominent oval, dorsal protuberance proximally, arising approximately 2/3 above acetabulum. Dorsal acetabular expansion concave, angled approximately 36° to ilial shaft. Ischium well developed, forming posterior acetabulum. Ventral portion of acetabulum well developed with ossified pubis fused with ilium and ischium. Acetabulum large, width approximately 60% of ilium (minus shaft) and ischium; anterior edge of rim approximately half above base of ilial shaft.

Fig. 4.66. Pelvic girdle of Platymantis parkeri, SAM5169B, female, lateral. Scale bar = 1 mm.

Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements obtuse. Sesamoid bones present ventrally at metacarpophalangeal joints of all digits and between proximal interphalangeal joint on digits 3,4. Phalangeal formula of foot:: 2-2-3-4-3. Distal tips of phalangeal elements obtuse. Prehallux well developed, two elements ossified. Sesamoids bones present ventrally at metatarsophalangeal joints of all digits and between proximal interphalangeal joint on digits 2,3,4,5; and between distal interphalangeal joint on digits 4. One large sesamoid at junction of tibiotarsus.

148

Fig. 4.67. Extremities of Platymantis parkeri, SAM5169B, female. (a) hand, dorsal (b) foot, dorsal. Scale bars = 1 mm.

4.3.5.5 Platymantis solomonis (Boulenger, 1884) Figs 4.68 - 4.74 Skull robust, as long as wide, with a well-ossified neurocranium. Sphenethmoid ossified fused to and reaching anterior margin of palatines, fused to vomers. Ossified portion extending half the length of the orbit in ventral view. Prootic fused with exoccipital dorsally but not ventrally to form otoccipital. Crista parotica well developed but not articulating with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, anteriorly truncate, extending to posterior margin of nasals; posteriorly fused with otoccipital, no flanges posterolaterally. Nasals large, broad, triangular, juxtaposed medially, overlying bony sphenethmoid. Maxillary process of nasal long, slender, acuminate, lying adjacent to preorbital process of pars facialis of maxilla, overlapping palatines extending beyond posterior margin. Palatines well ossified, long, expanded laterally, terminating on and fused with sphenethmoid. Parasphenoid robust, half width of frontoparietals, constricted posteriorly;

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cultriform process acuminate terminally, extending anteriorly to level anterior to palatines. Parasphenoid alae moderate, horizontal. Pterygoid well developed; anterior ramus tapering to a blunt point, extending to palatines, ventrally overlapped by pars dentalis of maxilla; medial ramus tapering, articulating with crista parotica. Posterior ramus acuminate. Anterior and posterior rami forming a brace; inner margins of rami forming a curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal oriented approximately parallel to midline. Quadratojugal robust, fully articulated. Squamosal well developed. Otic ramus distinct, equal in length to zygomatic ramus, not articulating with crista parotica. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Premaxilla broad; alary processes long, separated medially. Palatal shelf of premaxilla broad, expanded laterally; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla forming a broad V-shaped. Vomers large, articulating with ventral surface of sphenethmoid, fused to palatines; anterior process reduced, triangular; posterior process broad, triangular converging medially; lateral alae large, posterior lateral alae confluent with posterior process of vomer. Anterior lateral alae notched. Dentigerous processes present, transversely oriented, elongate with approximately 10 teeth.

a

150

b

Fig. 4.68. Skull of Platymantis solomonis, MCZ58565, female. (a) dorsal (b) ventral. Scale bars = 10 mm.

Lateral margins of hyoid plate slightly curved; plate cartilaginous, longer than wide. Anterior cornua slender, cartilaginous, curving anteromedially; anterior processes present, curved posterolaterally, almost forming a complete arch. Alary processes broad, club-like, flat, cartilaginous lateral extensions arising via a stalk, 2/3 above anterior margin of hyoid plate. Posterolateral processes short (approximately 23% length of posterior cornua), slender, acuminate. Posterior cornua bony, except distally. Posterior cornua directly abuts hyoid plate, narrowly separated proximally, almost touching; separated at angle between 20-30°. Proximal ends of posterior cornua expanded, distal ends undilated.

151

Fig. 4.70. Hyoid apparatus of Platymantis solomonis, SAM47134, female, ventral. Scale bar = 5 mm.

Firmisternal pectoral girdle robust. Epicoracoids ossified. Cleithrum present, ossified. Omosternum present, forked. Sternum ossified, well developed, long, proximal end expanded, tapering distally, with large cartilaginous xiphoid process. Clavicles large, moderate, oriented horizontally. Coracoids well developed, hourglass shaped, expanded medially and separated medially; oriented posteromedially. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

Fig. 4.71. Pectoral girdle of Platymantis solomonis, SAM47134, female, ventral. Scale bar = 5 mm.

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Eight presacral vertebrae. Neural spines present on all presacrals but very large on Presacrals II, III decreasing in size anterior to posterior. Transverse processes of Presacral III, IV moderately expanded distally. Lengths of transverse processes: III > IV > V = VI = VII = VIII >II. Orientation of transverse processes: III, IV, V, VI, VII angled posteriorly, II, VIII horizontal. Sacral diapophyses undilated, cylindrical, with anterior edge oriented posterolaterally. Urostyle bearing dorsal crest along entire length.

Fig. 4.72. Vertebral column of Platymantis solomonis, MCZ58565, female, dorsal. Scale bar = 10 mm.

Ilia cylindrical, shaft straight comprising approximately 72% of total length of pelvic girdle with distinct crest on shaft (approximately 56% width of shaft, 95- 100% length of shaft). Prominent oval, dorsal protuberance proximally, arising approximately half above acetabulum. Dorsal acetabular expansion straight, angled approximately 38° to ilial shaft. Ischium well developed, forming posterior and posteroventral acetabulum. Ventral portion of acetabulum well developed with ossified pubis fused with ilium and ischium. Acetabulum moderate, width approximately 51% of ilium (minus shaft) and ischium; anterior edge of rim approximately 1/3 above base of ilial shaft.

153

Fig. 4.73. Pelvic girdle of Platymantis solomonis, MCZ58565, female, lateral. Scale bar = 5 mm.

Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements rounded. Prepollical elements well developed, ossified. Sesamoids bones present ventrally at metacarpophalangeal joint of all digits, between proximal interphalangeal joint on digits 3,4 and between distal interphalangeal joint on digit 4. Phalangeal formula of foot: 2-2-3-4-3. Distal tips of phalangeal elements pointed. Prehallux well developed, elements ossified. Sesamoid bones present ventrally at metatarsophalangeal joint on all digits and between proximal interphalangeal joint on digits 3,4,5. One large sesamoid at junction of tibiotarsus.

a b

Fig. 4.74. Extremities of Platymantis solomonis, MCZ58565, female. (a) hand, dorsal hand (b) foot, dorsal. Scale bars = 5 mm.

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4.3.5.6 Platymantis weberi Schmidt, 1932 Figs 4.75 - 4.80 Skull robust, as long as wide, with a well-ossified neurocranium. Sphenethmoid ossified extending beyond to level approximately 1/3 above of posterior margin of nasals. Ossified part extending half length of orbit in ventral view, forming a point anteriorly. Prootic fused with exoccipital dorsally but not ventrally to form otoccipital. Crista parotica well developed articulating with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, anteriorly truncate or slightly acuminate, extending to posterior margin of nasals; posteriorly fused with otoccipital, no flanges posterolaterally. Nasals large, broad, triangular, juxtaposed medially, overlying bony sphenethmoid. Maxillary process of nasal long, slender, acuminate, lying adjacent to preorbital process of pars facialis of maxilla, overlapping palatines extending beyond posterior margin. Palatines well ossified, long, terminating on sphenethmoid. Parasphenoid robust, half width of frontoparietals, constricted posteriorly; cultriform process acuminate terminally, extending anteriorly to level anterior to palatines. Parasphenoid alae moderate, horizontal. Pterygoid well developed; anterior ramus tapering to a point, extending to palatines, ventrally overlapped by and fused with pars dentalis of maxilla. Medial ramus overlapping crista parotica. Posterior ramus acuminate. Anterior and posterior rami forming a brace; inner margins of rami forming a curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal oriented approximately parallel to midline. Quadratojugal robust, fully articulated. Squamosal well developed. Otic ramus distinct, equal in length to zygomatic ramus, juxtaposed with crista parotica. Zygomatic ramus broad, tapering to point. Maxilla and premaxilla dentate. Preorbital process present on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Premaxilla broad; alary processes long, separated medially. Palatal shelf of premaxilla broad, expanded laterally; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla forming a broad V- shape. Vomers large, articulating with ventral surface of sphenethmoid; anterior process moderate, triangular; posterior process broad, triangular converging medially; lateral alae large, anterior alae larger than posterior; posterior alae confluent with posterior process of vomer. Dentigerous processes present, obliquely oriented, moderate in length with approximately 8 teeth.

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a

b

Fig. 4.75. Skull of Platymantis weberi, SAM4412, male. (a) dorsal (b) ventral. Scale bar = 5 mm.

Lateral margins of hyoid plate straight; plate cartilaginous, longer than wide. Anterior cornua slender, cartilaginous, curving anteromedially. Alary processes

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broad, club-like, flat, cartilaginous lateral extensions arising via a stalk, level with anterior margin of hyoid plate. Anterior processes present, curved posterolaterally, almost forming a complete arch. Posterolateral processes short (approximately 20% length of posterior cornua), slender, acuminate. Posterior cornua bony, except distally. Posterior cornua directly abuts hyoid plate, narrowly separated proximally, almost touching; separated at angle approximately 30°. Proximal ends of posterior cornua expanded, distal ends undilated.

Fig. 4.76. Hyoid apparatus of Platymantis weberi, SAM4412, male, ventral. Scale bar = 5 mm.

Firmisternal pectoral girdle robust. Epicoracoids ossified. Cleithrum present, ossified. Omosternum present, forked. Sternum ossified, well developed, long, proximal end expanded, tapering distally, with large cartilaginous xiphoid process. Clavicles large, moderate, fused with epicoracoids, oriented horizontally. Coracoids well developed, hourglass shaped, expanded medially, narrowly separated medially. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

157

Fig. 4.77. Pectoral girdle of Platymantis weberi, SAM4412, male, ventral. Scale bar = 1 mm.

Eight presacral vertebrae. Neural spines present on all presacrals but very large on Presacrals II, III decreasing in size anterior to posterior. Transverse processes of Presacral III, IV moderately expanded distally. Lengths of transverse processes: III > IV > V = VI = VII = VIII >II. Orientation of transverse processes: III, IV, V, VI, VII angled posteriorly, II, VIII horizontal. Sacral diapophyses undilated, cylindrical, with anterior edge oriented posterolaterally. Urostyle bearing dorsal crest along entire length.

Fig. 4.78. Vertebral column of Platymantis weberi, SAM4412, male, dorsal. Scale bar = 5 mm.

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Ilia cylindrical, shaft slightly curved, approximately 70% of total length of pelvic girdle with distinct crest on shaft (approximately 70% width of shaft, 95% length of shaft). Prominent round, dorsal protuberance proximally. Dorsal acetabular expansion concave, angled approximately 40° to ilial shaft. Ischium well developed, forming posterior portion of acetabulum. Ventral portion of acetabulum well developed with ossified pubis fused with ilium and ischium. Acetabulum moderate, width approximately 45% of ilium (minus shaft) and ischium; anterior edge of rim approximately level with base of ilial shaft.

Fig. 4.79. Pelvic girdle of Platymantis weberi, SAM4412, male, lateral. Scale bar = 5 mm.

Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements obtuse. Prepollical elements well developed, ossified. Sesamoid bones present ventrally at metacarpophalangeal joints on all digits and between proximal interphalangeal joint on digits 3, 4. Phalangeal formula of foot: 2-2-3-4-3. Distal tips of phalangeal elements obtuse. Prehallux well developed, elements ossified. Sesamoids bones present ventrally at metatarsophalangeal and proximal interphalangeal joints on all digits. One large sesamoid at junction of tibiotarsus.

159

Fig. 4.80. Extremities of Platymantis weberi, SAM4412, male. (a) hand, dorsal (b) foot, dorsal. Scale bar = 5 mm.

4.3.6 Rana Linnaeus, 1758

4.3.6.1 Rana kreffti Boulenger, 1882 Figs 4.81 - 4.86 Skull robust, as long than wide, with a well-ossified neurocranium. Sphenethmoid ossified into large nasal plate extending to anterior margin of nasals. Ossified part extending 1/2 length of orbit in ventral view. Prootic fused with exoccipital dorsally but not ventrally to form otoccipital. Crista parotica well developed articulating with otic ramus of squamosal. Frontoparietal fontanelle roofed. Frontoparietals extending entire length of orbit, anteriorly truncate, extending to posterior margin of nasal plate and palatines; posteriorly fused with otoccipital, no flanges posterolaterally. Nasals narrow, widely separated, not overlying bony sphenethmoid. Maxillary process of nasal long, broad, acuminate, lying adjacent to preorbital process of pars facialis of maxilla, overlapping

160

palatines extending to posterior margin. Palatines well ossified, long, tapering to a fine point medially, terminating on sphenethmoid. Parasphenoid robust, 1/2 width of frontoparietals, cultriform process acuminate terminally with jagged edge, extending to level posterior to palatines. Parasphenoid alae broad, oriented posterolaterally. Pterygoid well developed; anterior ramus tapering to a point, extending to palatines, ventrally overlapped by pars dentalis of maxilla. Medial ramus broad, articulating with crista parotica. Posterior ramus ending in a blunt point. Anterior and posterior rami forming a brace; inner margins of rami forming a curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming distinctly curved profile between maxilla and squamosal, oriented approximately horizontal to midline. Quadratojugal slender, juxtaposed anteriorly with maxilla fused posterior via a spur with squamosal. Squamosal well developed. Otic ramus distinct, slightly longer than zygomatic ramus, articulating with crista parotica. Zygomatic ramus broad, tapering to point. Maxilla and premaxilla dentate. Well developed preorbital process on pars facialis of maxilla. Maxilla articulating with pars dentalis of premaxilla. Premaxilla broad; alary processes long, separated medially. Palatal shelf of premaxilla broad; palatine process acuminate. Area between palatal shelf and palatine process of premaxilla forming a broad U-shaped. Vomers moderate articulating with ventral surface of sphenethmoid; anterior process reduced to a small triangular point; posterior process broad, triangular converging medially; lateral alae moderate, posterior alae extending as a long sharp spine. Anterior alae small, triangular. Dentigerous processes present, obliquely orientated, short with approximately four teeth.

161

a

b

Fig. 4.81. Skull of Rana kreffti, MCZ38087, male. (a) dorsal (b) ventral. Scale bar = 5 mm.

162

Lateral margins of hyoid plate straight; plate cartilaginous, longer than wide. Anterior cornua slender, cartilaginous, curving anteromedially; anterior processes present, almost straight, slightly curved. Alary processes T-shaped, cartilaginous lateral extensions arising via a stalk, below level of anterior margin of hyoid plate. Posterolateral processes short, triangular, acuminate. Posterior cornua bony, except distally. Posterior cornua directly abuts hyoid plate, narrowly separated proximally; separated at angle of approximately 30°. Proximal ends of posterior cornua expanded, distal ends undilated.

Fig. 4.82. Hyoid apparatus of Rana kreffti, SAM5161A, female, ventral. Scale bar = 5 mm.

Firmisternal pectoral girdle robust. Epicoracoids cartilaginous with some ossification. Cleithrum present, ossified. Omosternum present, unforked or very slightly forked at base. Sternum ossified, well developed, short, proximal end greatly expanded, tapering distally, with large cartilaginous xiphoid process. Clavicles large, moderate, oriented horizontally. Coracoids well developed, hourglass shaped, expanded medially and separated medially; oriented horizontally. Scapula fused with clavicle, coracoid and suprascapula. Suprascapulae ossified.

163

Fig. 4.83. Pectoral girdle of Rana kreffti, MCZ38087, male, ventral. Scale bar = 5 mm. Crossed lines represent cartilage.

Eight presacral vertebrae. Neural spines present on all presacrals but very prominent on Presacrals II, III, IV, V decreasing in size anterior to posterior. Transverse processes of Presacral III, IV moderately expanded distally. Lengths of transverse processes: III > IV > II = V = VI = VII = VIII. Orientation of transverse processes: II angled anteriorly, III, IV, V, VI, angled posteriorly, VII, VIII horizontal. Sacral diapophyses slightly dilated, cylindrical, with anterior edge oriented posterolaterally. Urostyle bearing dorsal crest along entire length.

Fig. 4.84. Vertebral column of Rana kreffti, MCZ38087, male, dorsal. Scale bar = 5 mm.

164

Ilia cylindrical, shaft slightly curved, approximately 74% of total length of pelvic girdle with distinct crest on shaft (approximately 70% width of shaft, 90% length of shaft). Prominent round, dorsal protuberance proximally, arising approximately 1/2 above acetabulum. Dorsal acetabular expansion straight, angled approximately 44° to ilial shaft. Ischium well developed, bearing a small crest, forming posterior and posteroventral acetabulum. Ventral portion of acetabulum well developed with ossified pubis fused with ilium and ischium. Acetabulum moderate, width approximately 44% of ilium (minus shaft) and ischium; anterior edge of rim approximately level with base of ilial shaft.

Fig. 4.85. Pelvic girdle of Rana kreffti, MCZ38087, male, lateral. Sutures not identifiable. Scale bar = 5 mm.

Phalangeal formula of hand: 2-2-3-3. Distal tips of phalangeal elements rounded. Prepollical elements well developed, ossified. Phalangeal formula of foot: 2-2-3-4-3. Distal tips of phalangeal elements rounded. Prehallux well developed, elements ossified. Sesamoid bones present only at junction of tibiotarsus.

165

Fig. 4.86. Extremities of Rana kreffti, MCZ38087, male. (a) hand, dorsal (b) foot, dorsal. Scale bars = 5 mm.

166 Chapter 5. Karyology of the Solomon Island Ranids

5.1 Introduction Karyotypes can provide information for reconstructing phylogenetic relationships (Bogart and Tandy, 1981; Green, 1986). For instance, among the myobatrachid frogs of Australia, chromosome morphology proved to be valuable for inferring inter- and intra-generic relationships (Mahony and Robinson, 1980). However, Hillis (1991) and Green and Borkin (1993) point out that traditional methods of gross karyotype analysis may be insufficient for phylogenetic analysis. The aim of this chapter is to examine the karyotypes of the ranid frogs of the Solomon Islands. Comparisons with published data on a small number of related species from Australia and New Guinea are also given, to determine the extent of karyotype variability observed in Solomon Island ranids and its suitability for phylogenetic reconstruction.

5.2 Methods

5.2.1 Material examined Material is deposited in the following institutions: AM = Australian Museum, Sydney, Australia. SAM = South Australian Museum, Adelaide, Australia.

Batrachylodes vertebralis, SAM R37000-3, 1f, 3m, Pavora River, Choiseul Is., Solomon Is. (6°46'S 156°32'E); Ceratobatrachus guentheri, SAM R37004-6, 2f, 1m, Pavora River, Choiseul Is., Solomon Is. (6°46'S 156°32'E); Discodeles bufoniformis, SAM R37007-8, 2f, Pavora River, Choiseul Is., Solomon Is. (6°46'S 156°32'E); D. guppyi, SAM R37009, 1m, Pavora River, Choiseul Is., Solomon Is. (6°46'S 156°32'E); AM R129684-6, 3m, 4km NNE Amelei, New Britain (6°03'S 150°40'E); AM R129589, 1m, Amelei, New Britain (6°06'S 150°40'E); Platymantis myersi, SAM R37010, 1f, Pavora River, Choiseul Is., Solomon Is. (6°46'S 156°32'E); P. neckeri, SAM R37011, 1f, Pavora River, Choiseul Is., Solomon Is. (6°46'S 156°32'E); P. solomonis, SAM R37012-13, 1f, 1m, Pavora River, Choiseul Is., Solomon Is. (6°46'S 156°32'E); P. weberi, SAM R37014, 1f, Pavora River, Choiseul Is., Solomon Is. (6°46'S 156°32'E). see Appendix 1 - specimens identified by a letter c

167 5.2.2 Preparation of Chromosome spreads Chromosomes were prepared in the field by Dr Steve Donnellan, South Australian Museum, from duodenal epithelium by the method of Mahony and Robinson (1980). Minor modifications were necessary because of remote field conditions. Colchicine solution (0.01% in distilled water) was injected intraperitoneally three hours before the frogs were killed. The duodenum was hypotonically treated in distilled water for three minutes then fixed in a freshly prepared solution of 3 parts methanol, 1 part acetic acid. Epithelial cells were scraped from the lining of the duodenum, and the resulting cell suspension was pipetted with 5 ml of fixative into a small plastic tube. The suspension was centrifuged with a hand centrifuge for five minutes, the supernatant pipetted off and the cells resuspended in fixative. This step was performed at least twice. The lid of the tube was then firmly secured with masking tape and the tube placed in a cylinder of liquid nitrogen. Upon return to the laboratory the tubes were removed from liquid nitrogen and brought to ambient temperature. The cell suspension was transferred to a centrifuge tube and spun at 1000 rpm for 5 minutes. The supernatant was removed and the cell button resuspended in 5 ml of fresh fixative, and air-dried chromosome spreads prepared and stained with buffered Giemsa.

5.2.3 Giemsa Banding (G-banding) G-banding produces differential staining in the chromosome arms. All methods involve treating the chromosomes with either a proteolytic enzyme such as trypsin or mild denaturation of chromosomes with an alkali such as NaOH or heating the slides with saline citrate solution at 65°C. These act to remove partially the protein component of the chromosomes (Vosa, 1973). With G-banding, heterochromatic regions of the chromosomes stain darker. G-banding is thought to operate by modifying chromosomal proteins or by selective denaturation and renaturation of chromosomal DNA, or both (Harrison et al. 1981; McKay, 1973). The advantages of using such a technique are that (i) the staining is permanent, (ii) no fluorescent microscopy is needed and (iii) the results are easily photographed.

168 5.2.3.1 Procedure Giemsa stock stain was prepared by grinding 1 g of powdered Gurr’s Giemsa R66 stain and dissolving it in 66 ml of glycerol. This solution was then heated at 56°C for two hours. Methanol (66 ml) was then added and the solution mixed thoroughly, and then allowed to stand at 37°C. The solution was mixed every day for seven days and then filtered. To make the staining solution: 10 ml of the

Giemsa stock stain was added to 90 ml of Sorenson's PO4 Buffer (pH 6.8)/Giemsa buffer (working solution). Two forms of Giemsa buffer were made: a stock solution consisting of 750 ml 0.2N KH2PO4 (27.2 g/l) and 335 ml of 0.2N NaOH (8 g/l); and a working solution consisting of 21 ml of the Giemsa stock solution and 1029 ml of de-ionised water, adjusted to a pH of 6.8 - 7.0. A Trypsin solution was made by adding 1 ml of trypsin stock (1.25 g of trypsin (Difco 1:250) in 200 ml of distilled water that has been acidified to a pH of 2.5 with concentrated HCl to 49 ml of Phosphate Buffered Saline (PBS). PBS was made by dissolving 8 g of NaCl, 0.2 g of KH2HPO4 and 1.16 g of Na2HPO4 in 1 L of distilled water. Prepared air-dried slides were immersed into the trypsin solution for 5-25 seconds at room temperature. The slides were then rinsed in PBS for 30 seconds, for another minute in fresh PBS and then stained with the working Giemsa solution for 4-5 minutes. Slides were rinsed well in distilled water for 5-10 seconds and allow to air dry. The slides were then cleaned with xylene and mounted with a coverslip, using Depex mountant, for long term storage.

5.2.4 Constitutive Heterochromatin Banding (C-banding) C-banding was first discovered by Pardue and Gall (1970) who noticed that the centromeric regions of the mouse chromosome complement stained more darkly with Giemsa than the remainder of the chromosome. Constitutive heterochromatin is present in varying amounts in virtually all vertebrate genomes. This heterochromatin consists of satellite DNA of highly repetitive short DNA sequences (Miklos and John, 1983). The regions that stain darkly by this method are called C-bands. More commonly C-bands are seen at the centromeric position, however they do occur on the distal part of the long arm of the Y chromosome. C- banding has been widely used for studying the distribution of centromeric heterochromatin and also for identifying chromosomes, studying the

169 rearrangements on abnormal cells and distinguishing chromosomes of one species from the other in somatic cell hybrids. The mechanism involved in C-band production is thought to be selective removal of various proteins from the chromosome in hot Saline Sodium Citrate (SSC) (Elspeth et al., 1985; Holmquist, 1979). Upon staining with Giemsa, the regions from which the proteins have been removed stain darkly. The regions are not necessarily constitutive heterochromatin; they may, for example, be regions that are inactive at different stages of the cell cycle. These regions are known as facultative heterochromatin.

5.2.4.1 Procedure Prepared air dried slides were place in 0.2 N HCl for 1 hour. They were then placed in barium hydroxide (BaOH, 3 g dissolved in 60 ml of water) for 1 hour. Slides were transferred to a solution of Saline Sodium Citrate (SSC) (17.52 g of NaCl and 8.82 g of trisodium citrate dissolved into 1000 ml of distilled water) and incubated in a water bath at 65°C for 1 hour. Slides were cooled in running water and stained with 10% Giemsa (refer to G-banding section for method) for 20 minutes. Slides were rinsed with distilled water, air dried and mounted with a coverslip using Depex mountant.

5.2.5 Silver Staining - Staining for active nucleolus organiser regions The silver-staining method of Bloom and Goodpasture (1976) was used to localise the nucleolar organiser regions (NORs). Two drops of silver nitrate were placed on to the air-dried slide preparations and overlain with a coverslip. The coverslip was sealed with nail varnish. Slides were then placed in a sealed container and incubated in a water bath for 10-12 hours at 56-60°C. The coverslip was then removed, the slide rinsed with distilled water and allowed to air dry.

5.2.6 Photomicroscopy After staining, suitable spreads were selected for photography using criteria such as the number of overlapping chromosomes, clarity of banding and the degree of chromatids. Chromosome pairs were cut from photographic prints and numbered according to size, and arranged in parallel rows so that conformation of the

170 banding pattern could be observed. Arm lengths of chromosomes were measured on at least three metaphase spreads using vernier calipers reading to 0.05 mm and mean relative lengths (RL) and arm ratios calculated. Terminology for centromere position follows Levan et al. (1964). For convenience and not phylogenetic inference, results were compared and contrasted with the 2n=26, fundamental number (FN)=52 karyotype in species of Rana. This karyotype format is common among the species from the subfamily Raninae examined by King (1990) and listed by Kuramoto (1990). Results were also compared with published accounts of other species of Platymantis (Kuramoto, 1985; Ota and Matsui, 1995; Mahony et al., 1996) and four species of Rana from New Guinea and surrounding regions (Mahony et al., 1996).

5.3 Results Differential staining methods (C-banding and G-banding) were attempted, but banded metaphases of sufficient quality to permit interspecific comparisons were not obtained.

5.3.1 Solomon Island species Results are summarized in Table 5.1.

Batrachylodes vertebralis has 2n=20, FN=40; pairs 2, 3, 4, and 5 are submetacentric, the remainder are metacentric (Fig. 5.1). The NOR is located proximally on the pair 3.

Idiogram for Batrachylodes vertebralis

18 16 14 12 10 8 6 4

Relative Length Relative 2 0 12345678910 Chromosome Number

Fig. 5.1. Idiogram of the karyoptype of Batrachylodes vertebralis. White circles represent centromeres.

171 Ceratobatrachus guentheri has 2n=30, FN=38; pairs 1 and 7 are subtelocentric; 11 and 12 are metacentric and the remaining 11 pairs are telocentric (Fig. 5.2). The NOR is located proximally on the pair 6.

Idiogram for Ceratobatrachus guentheri

16 14 12 10 8 6 4

Relative Length Relative 2 0 1 2 3 4 5 6 7 8 9 101112131415 Chromosome Number

Fig. 5.2. Idiogram of the karyoptype of Ceratobatrachus guentheri. White circles represent centromeres.

Discodeles bufoniformis has 2n=24, FN=48, consisting of five large and seven small pairs. Pairs 1, 5, 6, 10 and 12 are metacentric; 2, 3, 7 and 11 are submetacentric and 4, 8 and 9 are subtelocentric (Fig. 5.3). The NOR is located proximally on pair 9.

Idiogram for Discodeles bufoniformis

20

15

10

5 Relative Length Relative 0 123456789101112 Chromosome Number

Fig. 5.3. Idiogram of the karyoptype of Discodeles bufoniformis. White circles represent centromeres.

Discodeles guppyi has 2n=24, FN=46, consisting of five large and seven small pairs. Pairs 1, 5, 10 and 12 are metacentric; 2, 7 and 11 are submetacentric; and pairs 3, 6, 8 and 9 are subtelocentric and pair 4 is telocentric (Fig. 5.4). The

172 NOR is located proximally on pair 6. Although the karyotype was similar to that seen in D. bufoniformis, the following differences were observed: the position of the NOR is located proximally on pair 9 in D. bufoniformis and pair 6 in D. guppyi; pair 3 is submetacentric in D. bufoniformis but subtelocentric in D. guppyi and pair 4 is subtelocentric in D. bufoniformis and telocentric in D. guppyi.

Idiogram for Discodeles guppyi

20 18 16 14 12 10 8 6 4 Relative Length Relative 2 0 123456789101112 Chromosome Number

Fig. 5.4. Idiogram of the karyoptype of Discodeles guppyi. White circles represent centromeres.

Platymantis myersi has 2n=22, FN=44; pairs 1, 5, 10 and 11 are metacentric, and the remainder submetacentric or subtelocentric (Fig. 5.5). The NOR is located proximally on the pair 7.

Idiogram for Platymantis myersi

18 16 14 12 10 8 6 4

Relative Length Relative 2 0 1234567891011 Chromosome Number

Fig. 5.5. Idiogram of the karyoptype of Platymantis myersi. White circles represent centromeres.

Platymantis neckeri has 2n=26, FN=42; pairs 1, 2, 4 and 13 are metacentric; 7, 8, 11 and 12 are submetacentric and five pairs: 3, 5, 6, 9 and 10 are telocentric

173 (Fig. 5.6). The NOR is located proximally on the pair 4.

Idiogram for Platymantis neckeri

16 14 12 10 8 6 4

Relative Length Relative 2 0 12345678910111213 Chromosome Number

Fig. 5.6. Idiogram of the karyoptype of Platymantis neckeri. White circles represent centromeres.

Platymantis solomonis has 2n=22, FN=42; pairs 5 and 9 are metacentric; pair 6 is telocentric and the remainder are either submetacentric or subtelocentric (Fig. 5.7). The NOR is located proximally on the pair 7.

Idiogram for Platymantis solomonis

18 16 14 12 10 8 6 4

Relative Length Relative 2 0 1234567891011 Chromosome Number

Fig. 5.7. Idiogram of the karyoptype of Platymantis solomonis. White circles represent centromeres.

Platymantis weberi has 2n=22, FN=40; pairs 1, 5 and 11 are metacentric; 2, 3, 4, 6, 7 and 8 are submetacentric or subtelocentric and two pairs: 9 and 10 are telocentric (Fig. 5.8). The NOR is located proximally on the pair 7.

174 Idiogram for Platymantis weberi

20 18 16 14 12 10 8 6 4 Relative Length Relative 2 0 1234567891011 Chromosome Number

Fig. 5.8. Idiogram of the karyoptype of Platymantis weberi. White circles represent centromeres.

Karyotype morphology was different among the species of Platymantis. However, with the exception of Platymantis neckeri, the Solomon Island species examined has 2n=22. Platymantis myersi and P. neckeri also shared the same NOR position on pair 7.

175 Table 5.1. Relative Length and Arm Ratio calculation of chromosomes of Solomon Island ranids. Chromosome Batrachylodes vertebralis Ceratobatrachus guentheri Discodeles bufoniformis Discodeles guppyi Number 2n=20,FN=40 2n=30,FN=38 2n=24,FN=48 2n=24,Fn=26 RLsd Arm RLsd Arm RLsd Arm RLsd Arm Ratiosd Ratiosd Ratiosd Ratiosd 1 16.810.82 1.710.85 m 14.740.88 2.972.43 st 16.890.32 1.720.74 m 18.620.71 1.300.06 m 2 13.390.75 2.220.30 sm 14.311.58 ∞ t 14.810.42 1.960.43 sm 15.530.78 2.160.40 sm 3 12.591.68 2.880.89 sm 12.860.89 ∞ t 12.400.71 2.210.05 sm 11.970.55 3.120.47 st 4 11.310.63 2.080.39 sm 11.700.51 ∞ t 12.640.23 5.211.33 st 11.960.35 ∞ t 5 11.380.77 2.010.17 sm 9.660.67 ∞ t 11.530.12 1.240.08 m 11.410.58 1.470.12 m 6 9.790.51 1.430.47 m 8.740.55 ∞ t 7.970.22 1.690.02 m 7.850.35 3.270.41 st 7 9.610.87 1.570.14 m 8.280.44 2.792.21 st 6.350.57 1.760.17 sm 6.020.21 1.760.13 sm 8 5.850.24 1.700.55 m 7.410.80 ∞ t 6.590.30 2.960.40 st 6.220.45 2.911.36 st 9 4.840.61 1.240.13 m 6.760.56 ∞ t 5.470.26 1.920.56 st 5.260.31 2.040.38 st 10 4.420.57 1.270.18 m 5.532.74 ∞ t 5.370.14 1.730.42 m 5.160.40 1.380.13 m 11 7.030.95 1.200.16 m 5.020.46 1.820.79 sm 4.690.42 1.240.20 sm 12 6.120.63 1.120.19 m 5.170.39 1.940.99 m 4.610.35 1.660.09 m 13 5.391.75 ∞ t 14 5.171.75 ∞ t 15 3.971.36 ∞ t

176 Table 5.1. continued Chromosome Platymantis myersi Platymantis neckeri Platymantis solomonis Platymantis weberi Number 2n=22,FN=44 2n=26,FN=42 2n=22,FN=42 2n=22,FN=40 RL±sd Arm RL±sd Arm Ratiosd RL±sd Arm RL±sd Arm Ratiosd Ratiosd Ratiosd 1 16.360.53 1.310.08 m 13.660.77 1.690.25 m 16.200.11 2.580.22 sm 17.410.93 1.300.12 m 2 14.290.33 1.860.16 sm 12.950.35 1.330.33 m 13.720.68 2.750.06 sm 14.170.81 2.170.20 sm 3 12.450.55 2.740.40 st 13.091.11 ∞ t 12.160.28 3.180.28 st 12.030.56 2.420.58 sm 4 11.480.34 1.920.08 sm 11.620.94 1.650.11 m 11.090.38 2.710.25 sm 11.451.16 9.012.73 st 5 10.430.21 1.430.27 m 8.850.69 ∞ t 10.390.21 1.360.03 m 10.530.90 1.440.18 m 6 9.600.42 2.150.23 sm 10.030.99 ∞ t 10.401.04 ∞ t 9.660.75 2.590.95 sm 7 8.350.38 2.870.44 sm 8.330.28 2.820.53 sm 7.540.08 2.810.66 sm 7.811.11 3.621.82 st 8 6.160.10 1.900.22 sm 7.360.41 1.680.17 sm 6.430.56 2.631.03 sm 6.520.29 3.500.61 st 9 5.760.34 1.770.31 sm 7.571.22 ∞ t 6.410.13 1.450.04 m 5.600.36 ∞ t 10 5.130.59 1.390.26 m 6.530.66 ∞ t 5.670.10 1.990.02 sm 4.820.49 ∞ t 11 4.500.34 1.290.43 m 5.900.18 1.950.25 sm 4.980.01 1.550.13 sm 4.450.44 1.450.16 m 12 5.890.57 2.150.74 sm 13 5.230.55 1.490.42 m

177 5.3.2 Comparison Species from literature

The following species taken from literature : Rana daemeli1, R. garritor1, R. jimiensis1, R. supragrisea1, Platymantis boulengeri1, P. dorsalis2, P. papuensis2, P. hazelae2 and P. pelewensis3 are summarised in Table 5.2. 1Mahony et al., 1996; 2Kuramoto, 1985; 3Ota and Matsui, 1995.

Table 5.2. Relative Length and Arm Ratio calculations for South-East Asian ranids. Chromosome Rana daemeli1 Rana garritor1 Rana jimiensis1 Rana supragrisea1 Number 2n=26,FN=52 2n=26,FN=52 2n=26,FN=52 2n=26,FN=52 RL Arm RL Arm RL Arm RL Arm Ratio Ratio Ratio Ratio 1 15.95 1.07 m 15.66 1.28 m 14.51 1.37 m 15.52 1.13 m 2 12.64 1.94 sm 13.82 2.42 sm 12.38 1.84 sm 13.95 2.03 sm 3 11.19 2.34 sm 11.19 1.68 m 11.34 1.61 m 11.63 1.69 m 4 10.96 1.49 m 9.75 2.07 sm 10.70 1.56 m 10.39 2.03 sm 5 10.05 3.27 st 9.58 1.49 m 9.88 1.66 sm 9.72 1.40 m 6 5.40 1.67 m 6.01 1.35 m 5.78 1.49 m 5.50 1.46 m 7 5.29 1.89 sm 5.80 2.07 sm 6.04 2.08 sm 5.91 1.99 sm 8 5.39 1.84 sm 5.88 1.78 m 5.60 1.46 m 5.47 1.83 sm 9 4.61 1.32 sm 4.96 2.53 m 4.80 1.45 m 4.94 1.69 m 10 4.69 2.09 m 4.71 1.13 m 5.45 1.66 m 4.49 1.53 m 11 4.79 2.06 m 4.79 1.05 m 4.96 1.64 m 4.59 1.50 m 12 4.60 1.02 sm 3.99 1.81 m 4.73 1.20 m 4.51 1.55 m 13 4.10 1.61 m 3.87 1.28 m 3.88 1.55 m 3.67 1.64 m

178 Table 5.2. continued. Chromosome Platymantis boulengeri1 Platymantis dorsalis2 Platymantis hazelae2 Platymantis papuensis2 Number 2n=24,FN=48 2n=20,FN=40 2n=26,FN=52 2n=22,FN=44 RL Arm RL Arm RL Arm RL Arm Ratio Ratio Ratio Ratio 1 17.48 3.11 m 17.40 1.08 m 15.60 1.27 m 16.00 1.26 m 2 14.46 3.64 sm 13.00 4.19 st 12.90 1.70 sm 13.20 1.89 sm 3 14.06 3.71 sm 12.30 1.27 m 11.20 2.17 sm 11.60 2.27 sm 4 12.25 3.61 st 11.60 2.77 sm 10.80 1.83 sm 11.30 1.90 sm 5 12.21 2.08 m 10.10 1.25 m 9.90 1.33 m 10.20 1.38 m 6 10.75 2.57 sm 8.60 1.12 m 5.80 1.33 m 9.10 1.60 m 7 10.60 2.83 sm 7.80 1.15 m 5.70 2.40 sm 7.60 1.89 sm 8 8.15 1.88 sm 7.30 1.12 m 5.30 1.33 m 5.90 1.61 m 9 7.30 1.90 m 6.80 1.22 m 5.00 2.31 sm 5.30 2.08 sm 10 6.06 1.83 m 5.10 1.14 m 4.90 1.26 m 5.20 1.42 m 11 5.39 1.58 sm 4.70 1.20 m 4.50 1.26 m 12 5.27 1.74 m 4.50 1.24 m 13 3.80 2.99 sm

Table 5.2. continued Chromosome Platymantis pelewensis3 Number 2n=22,FN=44 RL Arm Ratio 1 16.10 1.27 m 2 12.80 1.90 sm 3 11.70 2.25 sm 4 11.30 1.95 sm 5 10.40 1.35 m 6 9.60 1.79 sm 7 7.10 2.44 sm 8 5.80 1.56 sm 9 5.50 1.96 sm 10 5.10 1.48 m 11 4.50 1.28 m

179 Mahony et al. (1996) examined four species of Rana from New Guinea: R. daemeli, R. garritor, R. jimiensis and R. supragrisea. All had 2n=26, FN=52. The karyotypes are very similar to one another and to other Rana karyotypes (Haertel et al., 1974; Green et al., 1981; Green, 1985; Green and Delisle, 1985; Nishioka et al., 1987a, 1987b). All chromosomes were biarmed and shared the following karyotype arrangement: 1, 6, 10, 11 and 13 metacentric, 2 and 7 submetacentric. Differences were observed in pairs 3, 9 and 12, which are metacentric in R. garritor, R. jimiensis and R. supragrisea but submetacentric in R. daemeli. Pairs 4 and 5 are submetacentric or subtelocentric in R. daemeli and R. jimiensis and metacentric in R. garritor and R. supragrisea, pair 8 is submetacentric in R. daemeli and R. supragrisea but metacentric in R. garritor and R. jimiensis. RLs of chromosomes were the same as those seen in other 2n=26, FN=52 Rana karyotype. Mahony et al. (1996) also examined Platymantis boulengeri from New Guinea. The species has 2n=24, FN=48; pairs 1, 5, 9, 10 and 12 are metacentric; pair 4 subtelocentric and the remainder are submetacentric. RLs of pairs 6 and 7 are greater than those found in species of Rana. Kuramoto (1985) examined three species of Platymantis. He found that Platymantis dorsalis had 10 pairs of biarmed chromosomes. Pair 2 was subtelocentric; pair 4 was submetacentric, and the other eight metacentric. The NOR was located on pair 9. Platymantis papuensis had 11 pairs of biarmed chromosomes of pairs 2, 3, 4, 7, and 9 were submetacentric the other six pairs metacentric. No NOR was observed. Platymantis hazelae had 13 pairs of biarmed chromosomes consisting of five large and eight small pairs. Pairs 2, 3, 4, 7, 9 and 13 were submetacentric and the other seven pairs metacentric. Again no NOR was recorded. Platymantis pelewensis was karyotyped by Ota and Matsui (1995). They found that the karyotype had 2n=22 homologous chromosomes. Pairs 1, 5, 8, 10 and 11 were metacentric. The remainder were submetacentric. The NOR was located on the short arm of pair 7.

180 5.4 Discussion The data presented here and elsewhere (Kuramoto, 1985; Mahony et al., 1996; Ota and Matsui, 1995) demonstrate that reduction and/or increases in diploid number have occurred in the evolution of the ranid genera Batrachylodes, Ceratobatrachus, Discodeles and Platymantis. Reduction in diploid number and FN in ranids is not uncommon. Species of the genera Platymantis, Ptychadena and Rana have been reported as differing from the 2n=26, FN=52 karyotype of Rana (Birstein, 1984; Bogart and Tandy 1981; Green, 1983; Green and Borkin, 1993; Kuramoto, 1972, 1980, 1985; Schmid, 1980; Seto, 1965). King (1990) proposed that the reductions in diploid number occurred independently in each of the three genera. Two mechanisms have been proposed for the reduction in diploid number. It could have been accomplished by numerous small translocations (Morescalchi, 1973), or from a two stage process involving the production of telocentrics by inversion, and subsequent chromosome fusion of the derived telocentrics (Morescalchi, 1981), which King (1990) termed the "inversion-fusion cycle". Evidence for this cycle can be seen in Rana mucosa (2n=26, FN=46) which has three of the smaller chromosome pairs telocentric, and Pyxicephalus adspersus (2n=26, FN=48), which has two pairs of small telocentrics (Morescalchi, 1981; Green, 1986). Evidence for the subsequent fusion of the telocentric chromosomes is found in the Rana species which have 2n=24, FN=48 or 2n=22, FN=44. These species have a larger pair 6 and larger pairs 6 and 7 respectively, than in the 2n=26, FN=52 karyotype of Rana. The centric fusion of the smaller telocentrics, could account for the increase in chromosome size and reduction in diploid number and FN. The species of Solomon Island ranids examined all had relatively larger pairs 6 and 7 when compared to karyotypes of either Rana species. Evidence that centric fissions have been involved in other anurans is found in the Neotropical leptodactylid genus Eleutherodactylus which has diploid numbers ranging from 18 to 34 (Bogart, 1974; Cole, 1974; de Weese, 1975). King (1990) has argued that the major factor initiating chromosomal change in this genus has been fission of metacentric chromosomes. Kuramoto (1985) postulated that the karyotype of the New Guinean species Platymantis papuensis (2n=22, FN=44) could be derived from the karyotype of P. hazelae (2n=26, FN=52) by the "inversion-fusion cycle", but was unable to derive the karyotype of the Philippine species P. dorsalis (2n=20, FN=40), by the same

181 mechanism. Amongst the Solomon Island ranid frogs the "inversion-fusion cycle" could be used to explain some observed reductions and increases in diploid number and FN. Pericentric inversions may explain the occurrence of telocentric chromosomes among the smaller pairs in Ceratobatrachus guentheri, Platymantis neckeri and P. weberi, differences in centromeric positions between P. myersi and P. solomonis, and the change in NOR location between P. myersi and P. weberi. However, banding evidence is required to test this hypothesis. The karyotype of Batrachylodes vertebralis which had the lowest diploid number and FN, may support the occurrence of fusions. This species has chromosomes with increased RLs which could result from the fusion of the smaller chromosome elements to form a larger pair. However, the evolution of some of the karyotypes of Solomon Island ranids cannot be readily explained by the "inversion-fusion cycle". Evidence for centric fissions comes from a number of species. The karyotype of Platymantis neckeri can most readily be related to that of P. boulengeri by centric fission of the largest chromosome to produce the telocentrics 5 and 6 (Mahony et al., 1996). The karyotypes of Discodeles bufoniformis and D. guppyi can be achieved from a 2n=26 karyotype by considering the centric fissioning of a small chromosome and subsequent telomeric fusion of these chromosomes onto larger chromosomes. The location of the NOR gives supportive evidence of the role of fissions and fusions in the Solomon Island taxa examined. Although the NOR was found on a number of different chromosome pairs between species, it was always found proximally. The NORs of most specimens were dimorphic, a feature which is common in the karyotypes of frogs (Schmid, 1982). It is possible to relate the different locations observed by the process of centric fission or fusion. The frequency of chromosomal rearrangements in Batrachylodes, Ceratobatrachus, Discodeles and Platymantis with diploid numbers ranging between 20 to 30 is relatively high when compared with the conservative nature of karyotypes of most families of Amphibia (Morescalchi, 1973, 1979). King (1990) documented that 53 of the 60 species of karyotyped Rana have 2n=26, FN=52, five have 2n=24, FN=48, and two have 2n=22, FN=44, whilst Kuramoto (1990) reported that 43 of another 49 species of Rana have 2n=26. In over 60 species of Australian hylids examined, only one species has been found with a change in diploid number (King, 1980, 1981; Menzies and Tippet, 1976). An exception to this general pattern of karyotypic conservatism in the Anura is found in the

182 leptodactylid genus Eleutherodactylus where the diploid number ranges from 18 to 34 (Bogart, 1974; Cole, 1974; de Weese, 1975). Morescalchi (1979) has noted the association between rapid chromosomal evolution and speciation in tropical habitats, most evident in Eleutherodactylus, while Bogart (1981) proposed a correlation between chromosome variability and terrestrial habit (which involves direct development, parental behaviour, increased territoriality, and small clutch size). The level of karyotypic variation in the Solomon Island ranids is high, when it is considered that only a small number of species have been examined. It is tempting to draw some parallels between these frogs and Eleutherodactylus. Both show direct development, occur in the tropics, and have distributions which include a significant island component. Further study of the chromosomes of the Solomon Island ranids offers the opportunity to determine whether chromosomal rearrangements have any relationship with patterns of speciation, biogeography (dispersal versus vicariance) or shifts in habitat utilisation or specialisation. Considerable chromosome repatterning has occurred in the Solomon Island ranids examined in this study. Unfortunately, phylogenetic analysis of the karyotype at this level of resolution is uninformative because, with the exception of three species, Platymantis myersi, P. papuensis and P. pelewensis, each species has a different karyotype format. However, if it is possible to identify the chromosomal segments involved in these changes by the use of chromosome banding techniques, then a reassessment of the phylogenetic significance of these changes would be warranted. It is unfortunate that the various attempts at both G- banding and C-banding were unsuccessful. Only very faint bands were observed for some species examined. Reasons for the failure of bands to appear may have been due to the initial preparation of material in remote field conditions or it may be that there is little heterochromatin in chromosomes of these frogs. In order to test the identity of the various chromosomal rearrangements already observed, banding techniques will have to be applied. Also, karyotypes of more species of Batrachylodes and Platymantis, the majority of whose species have not been examined, should be determined.

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Chapter 6. Phylogenetic Analysis

6.1 Introduction The Solomon Island ranid frogs are well known taxonomically, yet, their phylogenetic relationships have not been investigated. They are endemic to the region and are highly diverse in both habit and habitat. This chapter explores the phylogenetic relationships between the ranid frogs of the Solomon Islands. The null hypothesis to be tested is that "the ranid frogs of the Solomon Islands do not form a monophyletic clade". Acceptance or rejection of this hypothesis provides information regarding invasion events into the Solomon Islands. If the null hypothesis is rejected then there has only been one invasion event into the Solomon Islands, i.e. the Solomon Island ranids are each other closest relatives. If the null hypothesis is accepted then there are two possible explanations either: a) There have been two invasion events into the Solomon Islands by ranid frogs. or b) There have been multiple invasions into the Solomon Islands by ranid frogs, i.e. the ranid frogs from the Solomon Islands are not each others closest relatives. If there have been two invasion events then there would be two clearly defined clades containing species of Solomon Island ranids with outgroup species forming sister taxa. If there have been multiple invasion events then there would be multiple clades inclusive of outgroup taxa. Furthermore if there are only two clades and hence two invasion events this would provide some evidence for Brown's (1952) assertion that there is an older element and a more recent element of frogs within the islands so that Batrachylodes, Ceratobatrachus, Discodeles, Palmatorappia and some members of Platymantis form a monophyletic group whilst Rana and some members of Platymantis (P. myersi, P. weberi) form a second monophyletic group

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Closely allied to these possible explanations are questions arising regarding the origin of the frogs within the zoogeographic region: is the most recent common ancestor to the Solomon Island ranids found in New Guinea, New Britain, the Philippines, Borneo or a combination of the these land masses? Lastly, the question of whether all or some of the Solomon Island genera should be placed in separate clades or not (sensu Noble 1931, Savage, 1973, or Dubois, 1992) will be addressed.

6.2 Methods To test the above hypotheses, 341 character states distributed across 136 morphological characters from osteology, external morphology and morphometrics from 27 taxa listed in Table 6.1 were analysed using the parsimony program Nona (v0.99, v2.00 Goloboff, 1999). Cladograms were constructed using the Ratchet command (Nixon, 1999) with the following settings: 200 iterations/rep, 10 trees in hold/iteration, 36 characters sampled, ambi= poly-. The random constraint level was set at 10. Descriptions of all morphological characters and their states are given in Appendix 3.

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Table 6.1 Character coding for 27 taxa by 136 characters Species 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 Outgroup Rana temporaria 1 0 2 0 2 0 1 1 1 0 1 0 2 1 0 1 1 1 1 2 1 1 1 1 0 0 0 0 1 0 Staurois natator 0 0 2 0 2 1 0 2 0 0 0 0 2 1 1 1 1 1 0 0 0 1 0 1 0 ? 1 0 ? ? Meristogenys orphnocnemis 0 0 2 1 2 1 2 2 1 1 1 0 1 1 1 1 2 1 0 1 1 1 1 1 0 0 0 0 0 0 Ingerana baluensis 1 1 2 1 0 1 0 2 1 0 0 1 2 1 0 1 1 1 0 2 1 1 1 1 1 1 1 1 1 0 Occidozyga laevis 1 0 1 1 0 1 1 2 0 1 1 0 0 0 1 1 1 2 0 1 0 0 1 0 0 ? 0 0 ? ? Limnonectes magnus 1 1 2 1 0 1 1 2 1 0 2 0 1 1 0 0 1 1 1 2 1 1 1 1 0 0 0 1 1 0 Rana grisea 0 0 2 0 2 1 1 2 1 0 0 1 1 1 1 1 1 1 0 1 1 1 1 1 0 0 1 0 0 0 Platymantis boulengeri 1 1 1 1 0 1 1 2 1 1 0 0 2 1 1 0 1 0 1 2 1 1 0 0 0 0 0 1 1 0 Platymantis corrugata 1 0 2 1 1 1 0 2 0 0 0 1 2 0 1 0 1 0 0 2 1 1 1 0 0 0 0 1 1 0 Platymantis papuensis 1 0 2 1 1 1 1 1 0 0 0 0 2 0 1 1 1 0 1 1 1 1 1 0 0 0 0 1 1 0 Platymantis schmidti 1 0 1 1 1 1 2 0 0 1 0 0 1 0 1 0 1 1 0 2 1 1 1 0 0 0 0 1 1 1 Ingroup Batrachylodes elegans 1 0 0 1 0 1 2 0 0 0 0 0 2 0 1 1 0 0 0 ? 0 1 1 1 1 ? ? ? ? ? Batrachylodes mediodiscus 1 1 0 0 0 1 0 0 0 0 0 0 2 0 1 1 0 0 0 1 0 1 1 1 1 ? 0 0 ? ? Batrachylodes trossulus 1 1 0 0 0 0 0 0 0 0 0 0 2 0 1 1 0 0 0 2 0 1 1 1 1 ? 0 0 ? ? Batrachylodes vertebralis 1 0 0 1 0 1 0 0 0 0 0 0 2 0 1 1 0 0 0 2 0 1 1 1 1 ? 0 0 ? ? Batrachylodes wolfi 1 0 0 1 0 1 0 0 0 0 0 0 2 0 1 1 0 0 1 2 0 1 1 1 1 ? 0 0 ? ? Ceratobatrachus guentheri 1 1 2 1 0 1 2 2 1 1 2 1 2 0 0 1 1 1 1 1 1 1 1 1 0 0 1 1 1 0 Discodeles bufoniformis 1 0 1 1 1 0 1 1 1 0 1 1 1 0 0 0 0 0 0 2 1 0 1 1 0 0 1 1 1 1 Discodeles guppyi 1 0 1 1 2 1 1 2 1 0 0 1 1 0 0 0 1 0 1 2 1 1 1 1 0 0 1 1 1 1 Palmatorappia solomonis 1 0 2 0 0 0 0 0 0 0 0 1 1 1 1 1 0 0 0 1 0 0 1 1 1 ? 0 0 ? ? Platymantis guppyi 1 0 2 1 2 1 1 1 1 0 0 0 1 0 1 1 1 1 1 ? 1 1 1 0 0 0 0 1 1 0 Platymantis myersi 1 0 1 1 1 1 1 0 0 0 0 0 1 0 1 0 1 1 0 1 1 1 1 0 0 0 0 1 1 0 Platymantis neckeri 1 0 1 1 1 1 1 1 1 0 0 0 1 0 1 1 1 1 1 1 1 1 1 0 0 0 0 1 1 0 Platymantis parkeri 1 0 1 0 0 1 0 0 0 0 0 0 2 0 1 1 1 1 0 1 1 0 1 0 1 1 0 0 1 0 Platymantis solomonis 1 1 2 1 1 1 1 1 1 0 0 0 1 1 0 0 1 1 0 2 1 1 1 0 0 0 0 1 1 0 Platymantis weberi 1 0 2 1 1 1 1 1 0 0 0 1 2 0 1 1 1 1 1 2 1 1 1 0 0 0 0 1 1 0 Rana kreffti 0 0 2 0 2 1 1 2 1 0 0 1 1 0 1 1 1 1 0 1 1 1 1 1 0 0 1 0 0 0

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Table 6.1 continued Species 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Outgroup Rana temporaria 0 1 0 1 0 1 0 ? ? 0 ? ? 1 0 0 4 0 ? ? ? 0 0 0 0 0 1 1 1 1 0 0 Staurois natator 1 0 1 0 0 1 1 1 2 1 1 2 0 0 0 4 1 2 1 0 0 0 2 0 0 1 0 1 1 0 0 Meristogenys orphnocnemis 1 0 0 0 0 1 1 1 0 1 1 0 1 0 0 4 1 0 1 1 1 0 0 0 0 1 1 1 2 1 0 Ingerana baluensis 1 1 1 1 1 1 1 1 2 1 1 2 0 0 0 4 1 2 1 0 ? 0 ? 0 0 1 1 0 ? ? 0 Occidozyga laevis 1 1 0 1 1 0 0 ? ? 0 ? ? 1 0 0 4 0 ? 1 1 ? ? 0 1 0 1 1 0 ? ? 0 Limnonectes magnus 1 0 0 0 0 0 0 ? ? 0 ? ? 1 0 1 4 0 ? 0 1 2 0 0 0 0 1 1 0 ? ? 0 Rana grisea 1 0 0 0 0 1 1 0 0 1 0 0 1 0 1 4 0 ? 1 1 1 1 0 0 0 1 1 1 1 0 0 Platymantis boulengeri 1 1 0 1 1 1 1 0 1 1 1 0 1 0 0 4 0 ? 1 1 0 1 2 1 0 1 1 1 1 1 0 Platymantis corrugata 1 0 0 1 1 1 0 ? ? 1 0 0 1 1 1 4 0 ? 1 1 3 1 1 1 0 1 1 1 1 1 0 Platymantis papuensis 1 0 0 1 1 1 0 ? ? 1 0 1 1 0 1 4 0 ? 1 1 1 1 2 1 0 1 1 1 1 1 2 Platymantis schmidti 1 0 0 1 1 1 1 0 1 1 1 0 1 0 0 4 1 1 3 1 1 1 0 1 0 1 1 1 2 1 2 Ingroup Batrachylodes elegans 0 0 1 0 1 1 1 1 1 1 1 0 0 0 0 4 1 1 1 0 0 0 0 0 1 0 0 1 1 1 0 Batrachylodes mediodiscus 0 0 1 1 1 1 1 1 0 1 1 0 0 0 0 2 1 0 1 0 0 0 0 0 1 1 0 1 1 1 0 Batrachylodes trossulus 0 0 1 0 1 1 1 1 0 1 1 0 0 0 0 1 1 0 1 0 1 0 0 0 1 1 0 1 1 1 0 Batrachylodes vertebralis 0 0 1 1 1 1 1 1 1 1 1 0 0 0 0 3 1 1 0 0 0 0 0 0 1 1 0 1 1 1 0 Batrachylodes wolfi 0 0 1 1 1 1 1 1 0 1 1 0 0 0 0 4 1 0 0 0 0 0 0 0 1 1 1 1 1 1 0 Ceratobatrachus guentheri 2 1 0 1 1 1 0 ? ? 1 0 0 1 0 1 4 0 ? 2 1 1 1 2 1 0 2 1 1 1 1 1 Discodeles bufoniformis 1 0 0 0 0 1 0 ? ? 1 1 2 1 0 1 4 0 ? 1 1 1 1 0 1 0 1 1 1 1 1 2 Discodeles guppyi 1 0 0 0 0 1 1 0 1 1 1 2 1 0 0 4 1 1 1 1 1 1 0 0 0 1 1 1 1 1 0 Palmatorappia solomonis 0 0 2 0 0 1 1 1 2 1 1 2 0 0 0 4 1 2 1 0 1 0 1 0 0 1 0 0 ? ? 0 Platymantis guppyi 1 0 2 0 0 0 1 1 0 1 1 0 0 0 0 4 1 1 1 0 1 0 0 0 0 1 0 1 0 0 0 Platymantis myersi 1 0 0 0 0 0 1 0 1 1 1 1 1 0 0 4 1 1 3 1 1 1 0 1 0 1 1 1 2 1 0 Platymantis neckeri 1 0 1 1 0 1 1 1 0 1 1 1 0 0 0 4 1 1 1 1 0 1 0 1 0 1 1 1 1 0 0 Platymantis parkeri 1 0 0 1 1 1 1 0 1 1 0 2 0 0 0 0 0 ? 1 1 1 1 0 1 0 1 1 1 1 1 0 Platymantis solomonis 1 1 0 1 1 1 0 ? ? 1 1 0 1 0 1 4 0 ? 3 1 1 1 0 1 0 1 1 1 2 1 2 Platymantis weberi 1 0 0 1 1 1 1 0 1 1 0 0 1 0 0 4 1 1 3 1 1 1 0 1 0 1 1 1 2 1 0 Rana kreffti 1 1 0 0 0 1 1 0 1 1 0 2 1 0 1 4 1 0 1 1 1 1 0 0 0 1 1 1 1 1 0

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Table 6.1 continued Species 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 Outgroup Rana temporaria 1 1 0 1 1 1 3 0 2 0 0 2 0 2 1 1 1 1 1 1 3 1 0 1 1 2 0 1 1 0 1 Staurois natator 0 1 0 0 0 0 2 1 2 0 1 0 0 1 1 0 1 0 1 1 2 0 ? 2 0 0 ? 2 1 0 1 Meristogenys orphnocnemis 0 1 0 1 2 0 3 0 0 1 1 3 0 2 1 0 0 ? ? 1 1 1 0 ? 1 0 0 1 1 1 1 Ingerana baluensis 0 0 0 0 0 1 2 2 1 2 0 1 0 1 1 0 0 ? ? 0 ? 1 0 ? 1 1 0 1 1 1 1 Occidozyga laevis 1 1 0 1 2 1 2 1 2 2 0 3 0 1 1 1 1 1 1 0 ? 1 0 1 1 1 0 0 1 1 1 Limnonectes magnus 0 1 2 1 2 1 3 1 2 2 0 3 0 0 1 1 1 0 1 1 0 1 0 0 2 1 0 1 1 0 1 Rana grisea 0 1 0 1 2 1 3 0 2 1 1 3 0 1 0 0 1 1 1 1 3 1 0 1 1 0 0 0 1 0 2 Platymantis boulengeri 1 1 2 1 1 1 3 1 0 2 1 3 0 0 1 0 1 0 1 1 3 1 1 1 0 ? ? 0 1 1 2 Platymantis corrugata 1 1 2 1 1 1 3 1 0 2 1 3 0 2 0 0 1 1 1 1 3 1 1 1 0 0 0 0 1 1 1 Platymantis papuensis 1 1 0 1 1 1 3 1 0 2 1 2 0 0 1 0 1 1 1 1 3 1 0 2 0 0 0 1 1 1 1 Platymantis schmidti 1 1 1 1 1 1 3 1 0 2 0 2 0 0 0 0 1 1 1 1 3 1 0 1 0 ? 0 0 1 0 1 Ingroup Batrachylodes elegans 0 0 0 0 0 1 1 1 0 0 0 2 1 0 0 0 1 1 1 0 ? 1 2 0 1 2 1 ? 1 ? ? Batrachylodes mediodiscus 0 0 0 0 0 1 1 1 0 0 0 2 1 0 0 0 1 1 1 0 ? 1 2 1 1 2 1 0 1 0 1 Batrachylodes trossulus 0 0 0 0 0 1 1 1 0 0 0 2 1 0 0 0 1 0 1 0 ? 1 2 1 1 2 1 0 1 0 1 Batrachylodes vertebralis 0 1 0 0 0 1 1 1 0 0 0 2 1 0 0 0 1 0 0 0 ? 1 2 0 0 2 1 0 1 0 1 Batrachylodes wolfi 0 0 0 0 0 1 1 1 0 0 0 2 1 0 0 0 1 1 1 0 ? 1 2 1 0 2 1 0 1 1 1 Ceratobatrachus guentheri 1 1 0 1 1 1 2 0 0 2 1 2 0 2 1 0 1 1 1 1 3 1 0 1 1 2 1 1 1 0 0 Discodeles bufoniformis 1 1 0 1 1 1 3 0 2 2 0 3 0 1 1 0 1 1 1 1 3 1 1 1 0 0 1 0 1 1 1 Discodeles guppyi 0 1 0 1 0 1 3 1 2 2 0 3 0 1 1 0 1 0 1 1 3 1 1 1 0 0 0 2 1 1 1 Palmatorappia solomonis 0 0 1 0 0 0 1 1 2 2 1 0 2 2 0 0 1 0 0 0 ? 0 ? 0 0 1 ? 0 0 ? ? Platymantis guppyi 1 1 0 0 2 1 3 1 0 2 1 3 0 0 0 0 1 1 1 1 3 1 0 1 0 0 0 0 1 1 0 Platymantis myersi 1 1 0 1 1 1 3 1 0 2 0 3 0 2 0 0 1 1 1 1 3 1 0 1 0 0 0 0 1 0 2 Platymantis neckeri 0 1 1 1 1 1 3 1 0 2 1 2 1 0 0 0 1 0 1 1 3 1 0 0 0 0 0 0 1 1 1 Platymantis parkeri 0 1 0 1 1 1 0 0 0 2 0 2 1 0 0 0 1 0 1 1 3 1 0 2 0 0 0 0 1 0 1 Platymantis solomonis 1 1 2 1 1 1 3 1 0 2 0 3 0 0 1 0 1 1 1 1 3 1 1 0 0 0 0 0 1 1 1 Platymantis weberi 1 1 2 1 1 1 3 1 0 2 1 3 0 0 1 0 1 1 1 1 3 1 0 1 0 1 0 1 1 1 2 Rana kreffti 0 1 2 1 1 1 3 0 2 1 1 3 0 1 1 0 1 1 1 1 3 1 0 2 0 0 0 1 1 0 2

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Table 6.1 continued Species 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 Outgroup Rana temporaria 0 1 1 1 1 3 0 2 0 1 0 0 0 0 2 0 0 1 1 0 0 2 0 2 0 1 0 Staurois natator 0 2 1 0 1 2 0 2 0 1 1 1 0 0 0 0 2 1 1 0 1 1 1 2 0 1 0 Meristogenys orphnocnemis 1 1 1 1 0 3 0 2 0 1 1 1 0 0 0 0 1 1 1 0 0 1 1 2 0 0 0 Ingerana baluensis 0 0 1 1 0 4 0 2 0 1 0 0 0 0 0 0 0 1 1 0 0 1 0 2 0 0 0 Occidozyga laevis 0 2 0 2 0 3 0 0 0 1 0 0 1 1 2 1 0 0 0 ? 0 1 0 ? 0 0 3 Limnonectes magnus 1 0 0 1 0 3 0 2 1 2 1 0 1 0 0 0 1 1 1 0 0 1 1 2 1 1 ? Rana grisea 0 0 1 1 0 1 0 1 0 1 1 0 0 0 0 0 1 1 1 0 0 1 1 3 0 1 0 Platymantis boulengeri 0 0 1 1 1 3 0 2 1 1 1 0 1 0 0 0 1 1 1 1 0 1 1 2 0 0 1 Platymantis corrugata 0 1 1 1 0 2 0 1 0 1 1 0 1 0 0 1 1 1 1 0 0 1 1 2 0 0 3 Platymantis papuensis 0 1 1 1 0 2 0 1 0 1 1 0 1 0 0 0 1 1 1 0 0 1 1 2 1 0 0 Platymantis schmidti 0 0 1 1 0 3 0 2 0 2 1 1 1 0 0 0 1 1 1 0 0 1 1 2 1 0 1 Ingroup Batrachylodes elegans ? ? ? ? ? ? ? ? ? ? 0 0 0 0 0 1 1 1 1 0 0 1 0 2 2 1 0 Batrachylodes mediodiscus 1 1 1 1 0 4 0 2 0 1 0 0 0 0 1 1 0 1 1 0 0 2 0 2 2 1 0 Batrachylodes trossulus 1 1 1 1 0 4 1 2 0 1 0 0 0 0 1 1 0 1 1 0 0 2 0 2 0 1 3 Batrachylodes vertebralis 1 1 1 1 0 4 0 2 0 1 0 0 0 1 1 1 0 1 1 0 0 1 0 2 2 1 0 Batrachylodes wolfi 0 1 1 1 0 4 0 1 0 1 0 0 0 1 1 1 0 1 1 0 0 1 0 2 2 1 0 Ceratobatrachus guentheri 0 0 1 1 0 0 1 0 1 1 1 1 1 0 0 0 1 0 1 1 0 0 1 2 0 1 0 Discodeles bufoniformis 0 1 0 1 0 2 0 0 0 1 0 0 1 0 0 0 0 1 1 0 0 1 1 2 0 1 2 Discodeles guppyi 0 2 1 1 0 3 0 1 0 1 0 0 1 0 0 0 1 1 1 0 0 1 1 2 0 1 2 Palmatorappia solomonis 1 0 ? 1 1 3 ? 2 0 1 0 1 0 1 0 0 0 0 1 0 0 1 1 1 0 0 0 Platymantis guppyi 0 1 1 1 0 2 0 1 0 0 1 1 1 0 0 0 0 0 1 1 0 0 1 2 2 0 0 Platymantis myersi 0 1 1 1 0 3 0 2 0 1 1 0 1 0 0 0 0 1 1 0 0 1 1 2 1 0 1 Platymantis neckeri 0 0 1 1 1 2 0 1 0 1 1 0 0 0 0 0 1 1 1 0 0 1 1 2 2 0 3 Platymantis parkeri 1 1 1 1 0 3 1 2 0 1 0 0 1 0 0 0 0 0 1 1 0 1 0 0 0 0 3 Platymantis solomonis 1 0 1 1 0 2 0 2 0 1 0 0 1 0 0 1 0 1 1 0 0 1 1 2 0 0 2 Platymantis weberi 0 1 1 1 0 2 0 1 0 1 0 0 1 0 0 0 1 0 1 0 0 1 1 2 0 0 0 Rana kreffti 0 0 1 2 0 1 0 1 0 2 1 0 0 1 0 0 1 1 1 0 1 1 1 3 0 0 0

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Table 6.1 continued Species 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 Outgroup Rana temporaria 0 0 1 1 1 0 0 1 1 1 0 0 0 1 1 1 1 Staurois natator 1 2 1 1 1 0 0 1 1 0 0 0 0 1 0 0 2 Meristogenys orphnocnemis 1 2 1 1 0 0 0 1 1 1 0 0 1 1 0 2 4 Ingerana baluensis 1 2 1 1 0 1 1 0 0 0 0 0 1 1 1 0 1 Occidozyga laevis 0 0 0 ? ? ? 0 1 1 0 0 1 1 0 0 0 1 Limnonectes magnus 0 2 1 1 1 0 0 1 1 0 1 1 1 1 0 0 1 Rana grisea 0 2 1 0 1 0 0 1 1 1 0 1 1 1 0 0 3 Platymantis boulengeri 0 2 1 0 1 0 1 0 0 0 1 1 1 1 1 2 1 Platymantis corrugata 2 1 1 0 1 0 1 0 0 1 2 0 1 1 1 1 1 Platymantis papuensis 0 2 1 1 1 0 1 0 0 0 1 1 1 1 1 2 1 Platymantis schmidti 2 2 1 0 1 0 1 0 0 0 1 0 1 1 0 2 1 Ingroup Batrachylodes elegans 0 0 1 0 0 1 1 0 0 0 0 0 0 0 0 2 1 Batrachylodes mediodiscus 0 0 0 ? ? ? 1 0 0 0 0 0 0 0 0 2 1 Batrachylodes trossulus 0 0 0 ? ? ? 1 0 0 0 0 0 0 0 0 0 1 Batrachylodes vertebralis 0 0 0 ? ? ? 1 0 0 0 1 0 0 0 0 2 1 Batrachylodes wolfi 0 0 0 ? ? ? 1 0 0 0 0 0 0 0 0 2 1 Ceratobatrachus guentheri 1 1 1 0 1 0 1 0 0 1 2 1 1 0 1 1 1 Discodeles bufoniformis 0 2 1 1 0 0 1 0 0 1 1 0 1 2 1 1 1 Discodeles guppyi 0 2 1 1 0 0 1 0 0 0 1 0 1 1 0 2 1 Palmatorappia solomonis 1 2 1 0 1 0 1 0 0 0 0 0 0 0 1 1 1 Platymantis guppyi 0 2 1 0 1 0 1 0 0 0 0 0 1 0 2 1 1 Platymantis myersi 2 2 1 1 0 0 1 0 0 0 1 0 1 2 2 2 1 Platymantis neckeri 0 2 1 0 1 0 1 0 0 0 0 0 1 0 1 2 1 Platymantis parkeri 0 2 1 1 0 0 1 0 0 0 1 0 1 2 1 1 0 Platymantis solomonis 2 2 1 1 1 0 1 0 0 0 1 0 1 1 2 2 1 Platymantis weberi 2 2 1 1 1 1 1 0 0 0 1 0 1 1 0 2 1 Rana kreffti 1 1 1 1 1 0 0 1 1 1 0 0 1 0 0 2 1

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6.2.1 Character Selection Characters were selected based primarily on three factors (1) my own observation of relevant differences across taxa, (2) diagnostic characters from literature, (3) ability to be accurately placed into discrete states. There may be more morphological characters not included here that may illuminate Solomon Island ranid phylogeny, e.g. myological characters. These characters were investigated, albeit briefly, yet were found to be not variable amongst Solomon Island ranids. A broader study needs to be undertaken for their inclusion into a phylogenetic analysis. Many of the characters listed in Appendix 3 are adapted from many publications (see Table 6.2). However, all character state definitions were scored only after personal observation. Hence differences that occur between my character descriptions and those in the literature are based on a more thorough analysis, including multiple specimens examined. Specimens examined are listed in Appendix 1 and are identified by a lower case p. All character states identified in these samples were included in the data matrix.

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Table 6.2 The references used to assist in character identification for each species. Species Cited Literature Batrachylodes elegans Brown and Parker, 1970 B. mediodiscus Brown and Parker 1970 B. trossulus Brown and Myers, 1949a; Brown 1952; Brown and Parker 1970 B. vertebralis Boulenger, 1887; Van Kampen, 1923; Kinghorn, 1928; Schmidt, 1932; Tanner, 1951; Brown 1952; Brown and Parker 1970 B. wolfi Brown and Parker 1970 Ceratobatrachus guentheri Boulenger, 1884; Boulenger, 1886a, Barbour, 1921; Van Kampen, 1923; Kinghorn, 1928; Tanner, 1951; Brown, 1952 Discodeles bufoniformis Boulenger 1884, 1886a, 1918a, 1920; Van Kampen 1923; Kinghorn 1928; Brown 1952 D. guppyi Boulenger 1884, 1886a, 1918a, 1920; Van Kampen 1923; Kinghorn 1928; Brown 1952 Ingerana baluensis Boulenger 1886; Van Kampen 1923, Inger 1954 Limnonectes magnus Boulenger 1920; Inger, 1954 Meristogenys orphnocnemis Matsui, 1986 Occidozyga laevis Inger 1954 Palmatorappia solomonis Ahl, 1927; Kinghorn, 1928; Brown, 1952 Platymantis boulengeri Loveridge, 1948; Brown and Tyler, 1968 P. corrugata Boulenger, 1882; Alcala, 1986; Inger 1954 P. guppyi Boulenger, 1884, 1886a; Van Kampen 1923; Kinghorn 1928; Brown 1952 P. myersi Brown, 1949; Brown 1952 P. neckeri Brown and Myers, 1949b; Brown 1952 P. papuensis Brown and Tyler 1968; Zweifel, 1969 P. parkeri Brown, 1965 P. schmidti Brown and Tyler 1968 P. solomonis Boulenger 1884, 1886a; Kinghorn 1928; Brown 1952 P. weberi Tanner 1951, Brown 1952, Brown and Tyler 1968 Rana kreffti Boulenger 1882, 1886a, 1920; Van Kampen 1923; Kinghorn 1928; Brown 1952 R. grisea Van Kampen 1913, Van Kampen 1923 R.temporaria Ecker 1889 Staurois natator Inger 1954

6.2.2 Outgroup selection Representative ranids (New Guinea: Platymantis papuensis, Rana grisea, New Britain: Platymantis schmidti, P. boulengeri, Philippines: Platymantis corrugata, Occidozyga laevis, Limnonectes magnus, Staurois natator, Borneo: Meristogenys orphnocnemis, Ingerana baluensis, Staurois natator) were included in this analysis based on their likely status as sister taxa to some or all of the Solomon Island ranids. These taxa were initially assigned as the outgroup based on the assumption that the ingroup was monophyletic (Swofford and Olsen, 1990). However, this assumption proved incorrect as initial runs of the phylogenetic analysis showed lack of resolution of the relationships between the Solomon Island ranids with other south-east Asian ranids. The European Rana temporaria was ultimately used as the outgroup.

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All genera within the Solomon Island ranids were represented and as many species as possible within these genera were included.

6.2.3 Assumptions for phylogeny reconstruction One problem with reconstructing phylogenies is controlling for the multitude of parameters that cannot be determined without first knowing the actual phylogeny (Asher, 1999). Such parameters need a priori estimating. These include character weighting, character ordering, treatment of missing data and choice of outgroup. Asher (1999) identified these aspects (as well as others) and created “assumption sets” to enable the processing of his data. This format has been followed throughout this chapter. Character weighting, missing data and ordering of multistate characters were all manipulated using the methods of Asher (1999) and are summarised in Table 6.3.

6.2.3.1 Character weighting Like Asher (1999) the methods of Farris (1969, 1989) and Carpenter (1988) were modified to a posteriori weight characters. In this study R*C scaled to base 100 was used. Character weighting was calculated using the trees derived from analysis 1 (Figs 6.1 - 6.3) i.e. the original data set. As three most parsimonious trees (MPTs) were found from analysis 1, the tree with the best support (ascertained by bootstrapping) was used (tree 1, Fig. 6.1), to obtain weights for each character. The results can be seen in analysis 2.

6.2.3.2 Missing data Missing data were of two types, states that were not scorable due to an incompleteness of data e.g. damaged specimen, so the state was unknown and those states that were inapplicable because they were absent in a taxon. It has been suggested that for the latter type of missing data the absent character can be recorded as “not present” (Maddison, 1993), but the result of having more multistate characters in the data set may not resolve the tree (Nixon and Davis, 1991; Platnick et al., 1991). In this analysis I used two data sets: one with all missing characters (either unknown or inapplicable) scored as ? and one where

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the unknown missing states were scored as ? and inapplicable missing states were scored as an extra state “not present”. The results can be seen in analysis 3.

6.2.3.3 Character ordering Following Asher (1999) and using the methods of Frost et al. (1991), Mabee (1989) and Wilkinson (1992) this study also examined the effect of unordered and ordered multistate characters. Firstly all characters were given equal weight and secondly those character state changes that represented an obvious morphocline were constrained.

Table 6.3. Combinations of different parameters analysed (see text for details) Weighting (a) all characters weighted (b) a posteriori weighting by equally RC x 100 Missing data (a) all characters included (b) inapplicable states recorded as “not present” Ordering (a) all characters unordered (b) multistate characters with distinct morphoclines ordered

6.2.4 Support and Confidence The size of the original data matrix 27 x 136 prevented exact searching techniques. Therefore the Ratchet analysis (Nixon, 1999) was performed to search for cladograms. This study used Bremer values as an indicator of support for trees found from the analyses (Bremer 1988, 1994), i.e. the number of extra steps needed to lose a branch in the strict consensus of the most parsimonious cladograms (Bremer, 1994). A branch present in one of the most parsimonious cladograms is more strongly supported by the data if a large increase in length of additional cladograms is required before that branch is lost in the consensus (Kallersjo et al., 1992). Using this index, strength of evidence for the monophyly of a group is usually equated to the number of synapomorphies. Total support index (t) (the sum of all group support divided by the length of the most parsimonious tree) was also calculated (Kallersjo et al., 1992; Bremer, 1994). Because exact searching could not be used, branch support indices are only approximations. Measures of confidence i.e. bootstrapping, were not employed in this thesis, due to the multitude of assumptions that cannot be met when using this statistical analysis (Kluge and Wolf, 1993; Bremer, 1994). Thus it was decided not to

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present arbitrary statistical measures of confidence based on simulated data. In addition, Bremer support is preferable over bootstrap methods as the former is calculated on an unchanged data matrix whilst the latter uses random resampling of characters (Gustaffson and Bremer, 1995).

6.3 Results For consistency and comparison the description of trees has been formalised. When writing out clades in the text, open parentheses are used to group species, for example,

becomes (M. orphnocnemis(R. grisea R. kreffti)). When detailing synapomorphies the character and character state are written in open parentheses followed by a description, for example, (115(1)) large tympanum. Such descriptions can be cross referenced to section 6.6 character descriptions. For all cladograms the white circles represent unique and unreversed changes and the black squares homoplasies (convergences, reversals and parallelisms). For all cladograms the species are identified by abbreviated generic names but full species names. For clarity Pasolomonis is Palmatorappia solomonis whereas Psolomonis is Platymantis solomonis.

6.3.1 Analysis 1 This analysis comprised the data set aaa (Table 6.3). Three MPTs were found of length 639 with consistency index (CI) 31 and a retention index (RI) of 50.

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Rtemporaria 2 8 9 13 17 20 21 23 34 53 67 80 88 90 93 95 99 105 107 109 110 118 121 132 Olaevis 1 0 1 0 2 0 0 0 1 1 2 0 0 1 2 2 0 1 1 0 0 3 0 0 1 15 27 44 48 63 78 81 84 85 93 100 101 116 Lmagnus 1 0 1 1 0 2 0 0 0 2 0 1 2 1 6 9 16 37 58 66 77 81 90 103 123 135 Morphnocnemis 0 4 28 68 70 71 104 128 2 1 2 1 2 0 0 1 1 1 0 4 46 59 75 88 117 122 130 134 135 0 2 0 0 1 1 0 1 3 11 26 40 44 91 93 97 99 115 Rgrisea 0 0 0 0 1 0 1 0 3 13 31 41 63 65 84 85 95 101 105 112 120 132 31 61 102 106 108 114 120 0 1 1 0 1 2 0 1 1 3 Rkreffti 0 0 1 0 1 1 2 0 1 2 2 1 2 0 2 2 1 1 1 0 2 3 21 40 45 46 68 98 109 111 115 118 132 134 135 Pparkeri 1 0 0 0 0 0 0 1 0 1 0 3 2 1 0 1 8 14 27 31 68 77 85 88 90 Ibaluensis 6 12 24 25 42 102 108 114 123 13 38 47 74 93 119 134 1 1 0 1 1 2 0 1 1 1 0 4 22 24 62 70 80 88 93 95 102 108 112 117 125 126 127 135 36 39 46 76 86 134 0 2 1 1 0 0 0 0 0 1 2 2 1 0 1 0 3 33 34 66 71 82 96 103 114 123 Snatator 0 2 0 0 1 0 1 2 2 0 1 2 1 1 0 1 1 2 1 1 1 0 0 2 5 12 16 17 21 30 32 63 73 74 79 89 105 109 115 122 132 134 0 0 0 0 1 0 1 1 1 1 32 37 49 62 64 65 80 104 Pasolomonis 0 1 0 0 0 0 2 1 2 2 0 0 1 0 1 0 0 1 1 1 0 0 0 0 0 0 6 33 55 85 108 Belegans 2 0 0 1 1 2 16 17 30 54 70 83 87 107 116 117 120 132 45 62 79 129 Bvertebralis 0 0 0 0 1 0 2 1 1 2 1 0 0 3 1 0 1 106 121 18 56 90 99

7 8 13 65 75 85 88 125 126 127 1 0 Bwolfi 38 47 84 1 1 1 1 19 0 0 0 1 0 0 0 1 0 0 0 0 1 1 3 85 113 Bmediodiscus 1 5 33 50 98 116 118 134 1 0 1 2 Btrossulus 0 0 1 1 0 3 0 6 9 19 29 60 63 93 101 103 122 2 15 48 58 116 118 119 Pschmidti 2 1 2 1 2 1 0 2 1 0 33 35 41 74 91 108 123 132 1 0 3 2 1 1 2 Pmyersi 0 0 1 2 2 0 0 2 4 33 35 49 51 53 56 58 64 65 91 101 103 108 109 111 113 133 134 4 27 61 8 37 38 42 59 116 122 132 Pguppyi 2 0 0 0 0 0 0 0 0 2 0 0 1 0 0 1 0 2 1 2 41 50 61 63 73 78 84 93 96 104 118 1 1 1 1 1 0 0 0 2 0 0 Pneckeri 1 1 0 0 1 1 0 0 0 1 0 3 41 60 84 116 7 18 71 90 97 99 Ppapuensis 52 130 1 2 2 1 1 1 1 1 2 1 2 13 15 19 22 40 50 63 78 83 96 118 2 1 1 4 7 8 9 31 93 100 111 122 Pboulengeri 1 1 0 2 0 1 0 2 0 1 1 1 6 10 11 14 23 26 30 48 55 60 67 68 74 85 90 98 103 109 113 117 119 120 128 129 132 134 1 0 2 1 1 1 0 1 1 0 12 40 75 Cguentheri 2 2 1 0 1 1 2 2 2 1 2 0 2 1 0 1 1 0 0 1 1 1 1 2 0 1 2 0 1 48 58 86 91 109 124 133 Pweberi 3 2 1 2 0 1 0 11 19 63 119 6 7 43 50 52 74 75 120 122 128 129 134 Pcorrugata 1 2 2 2 0 2 1 3 1 2 0 1 0 1 2 1 15 18 83 1 11 13 31 48 58 84 92 93 99 133

0 0 1 Psolomonis 8 12 14 40 71 1 0 1 1 3 2 0 1 0 2 2 5 10 16 21 68 87 94 99 128 132 134 1 1 0 1 0 2 23 26 29 33 34 41 63 69 74 117 119 123 Dbufoniformis 0 1 0 0 0 1 0 0 1 2 1 4 7 18 36 44 46 53 61 65 78 88 93 97 133 1 1 1 1 0 0 2 0 2 1 1 0 0 Dguppyi 2 2 1 1 0 1 0 0 0 0 2 2 3 0 Fig. 6.1. Tree 1 of Analysis 1. Length = 639, CI = 31 RI = 50. The Solomon Island ranids are distributed amongst three large clades.

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Rtemporaria 2 8 9 13 17 20 21 23 34 53 67 80 88 90 93 95 99 105 107 109 110 118 121 132 Olaevis 1 0 1 0 2 0 0 0 1 1 2 0 0 1 2 2 0 1 1 0 0 3 0 0 1 15 27 44 48 63 78 81 84 85 93 100 101 116 Lmagnus 1 0 1 1 0 2 0 0 0 2 0 1 2 1 6 9 16 37 58 66 77 81 90 103 123 135 Morphnocnemis 0 4 28 68 70 71 104 128 2 1 2 1 2 0 0 1 1 1 0 4 46 59 75 88 117 122 130 134 135 0 2 0 0 1 1 0 1 3 11 26 40 44 91 93 97 99 115 Rgrisea 0 0 0 0 1 0 1 0 3 31 61 102 106 108 114 120 13 31 41 63 65 84 85 95 101 105 112 120 132 0 1 1 0 1 2 0 1 1 3 Rkreffti 0 0 1 0 1 1 2 0 1 2 2 1 2 0 2 2 1 1 1 0 2 3 21 40 45 46 68 98 109 111 115 118 132 134 135 Pparkeri 1 0 0 0 0 0 0 1 0 1 0 3 2 1 0 1 8 14 27 31 68 77 85 88 90 Ibaluensis 6 12 24 25 42 72 102 108 114 123 13 38 47 74 93 119 134 1 1 0 1 1 2 0 1 1 1 36 39 46 76 86 134 0 4 22 24 62 70 80 88 93 95 102 108 112 117 125 126 127 135 0 2 1 1 0 2 0 0 0 0 1 2 2 1 0 1 0 3 33 34 66 71 82 96 103 114 123 Snatator 1 1 1 0 0 2 0 2 0 0 1 0 1 2 2 0 1 2 1 1 0 1 1 2 5 12 16 17 21 30 32 63 73 74 79 89 105 109 115 122 132 134 0 0 0 0 1 0 1 1 1 1 32 37 49 62 64 65 80 104 Pasolomonis 0 1 0 0 0 0 2 1 2 2 0 0 1 0 1 0 0 1 1 1 0 0 0 0 0 0 6 33 55 85 108 Belegans 2 0 0 1 1 2 16 17 30 54 70 83 87 107 116 117 120 132 45 62 79 129 Bvertebralis 0 0 0 0 1 0 2 1 1 2 1 0 0 3 1 0 1 106 121 18 56 90 99

7 8 13 65 75 85 88 125 126 127 1 0 Bwolfi 38 47 84 1 1 1 1 19 0 0 0 1 0 0 0 1 0 0 0 0 1 1 3 85 113 Bmediodiscus 1 5 33 50 98 116 118 134 1 0 1 2 Btrossulus 0 0 1 1 0 3 0 6 9 29 60 63 72 93 101 103 122 2 15 48 58 116 118 Pschmidti 2 1 1 2 1 2 0 2 1 0 33 35 41 74 91 108 123 132 1 0 3 2 1 1 Pmyersi 0 0 1 2 2 0 0 2 4 33 35 49 51 53 56 58 64 65 91 101 103 108 109 111 113 134 4 27 61 8 37 38 42 59 116 122 132 Pguppyi 2 0 0 0 0 0 0 0 0 2 0 0 1 0 0 1 0 1 1 1 1 2 41 50 61 63 72 73 78 84 93 96 104 118 1 1 0 0 0 2 0 0 Pneckeri 1 1 0 0 1 2 1 0 0 0 1 0 3 7 18 71 90 97 99 48 58 86 91 102 109 124 Pweberi 1 1 1 1 2 1 3 2 1 2 0 0 1 41 60 84 116 Ppapuensis 12 40 75 52 130 1 2 2 1 2 13 15 22 36 40 44 50 78 83 96 118 2 0 1 2 1 1 4 7 8 9 31 93 100 111 122 Pboulengeri 1 1 0 0 1 1 0 0 0 1 1 1 6 10 14 17 23 26 30 48 55 60 67 68 74 85 90 98 103 109 113 117 119 120 128 129 132 134 1 0 2 1 1 1 0 1 1 0 17 36 44 46 Cguentheri 2 2 0 1 1 1 2 2 2 1 2 0 2 1 0 1 1 0 0 1 1 1 1 2 0 1 0 0 1 0 6 7 43 50 52 74 75 120 122 128 129 134 Pcorrugata 0 2 1 3 1 2 0 1 0 1 2 1 15 18 83 1 13 17 31 48 58 84 92 93 99 133

0 0 1 Psolomonis 8 12 14 40 71 102 1 1 1 1 3 2 0 1 0 2 2 5 10 16 21 68 87 94 99 128 132 134 1 1 0 1 0 0 2 23 26 29 33 34 41 69 74 117 123 Dbufoniformis 0 1 0 0 0 1 0 0 1 2 1 4 7 18 36 44 46 53 61 65 78 88 93 97 133 1 1 1 1 0 0 2 2 1 1 0 Dguppyi 2 2 1 1 0 1 0 0 0 0 2 2 3 0 Fig. 6.2. Tree 2 of Analysis 1. Length = 639, CI = 31, RI = 50. The Solomon Island species are distributed amongst three clades. This tree differs from Tree 1 in the placement of Platymantis weberi.

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Rtemporaria 2 8 9 13 17 18 20 21 23 34 53 67 80 88 93 95 99 105 107 109 110 118 121 132 Olaevis 1 0 1 0 2 0 0 0 0 1 1 2 0 0 2 2 0 1 1 0 0 3 0 0 1 15 44 48 63 78 81 84 85 92 93 100 101 116 129 Lmagnus 1 0 1 0 2 0 0 0 2 1 0 1 2 1 1 6 9 16 58 66 77 81 92 103 123 135 Morphnocnemis 0 18 28 68 70 104 128 2 1 2 2 0 0 1 1 1 0 4 31 33 102 106 108 114 120 46 75 88 117 122 130 134 135 0 0 0 0 1 0 1 3 11 26 37 40 44 51 91 93 97 115 Rgrisea 0 0 1 0 1 1 2 0 0 0 1 0 1 0 3 13 31 38 41 63 65 84 85 95 101 105 112 120 132 0 1 1 0 0 1 1 2 0 1 3 Rkreffti 0 1 1 2 2 1 2 0 2 2 1 1 1 0 36 39 46 71 76 86 134 32 49 56 58 64 91 101 103 108 109 111 113 134 Pguppyi 1 1 1 1 0 0 2 2 0 0 0 0 0 0 1 0 0 1 0 1 32 41 50 63 73 78 84 93 96 104 118 Pneckeri 1 1 0 1 1 0 0 0 1 0 3 41 60 84 116 7 13 23 75 85 88 97 125 126 127 Ppapuensis 52 130 1 2 2 1 1 0 0 0 0 0 2 1 0 0 2 13 15 19 22 40 50 63 72 78 83 96 118 2 1 1 4 7 8 9 31 93 100 111 122 Pboulengeri 1 1 0 2 0 1 0 2 3 0 1 1 1 6 10 11 14 23 26 30 48 55 60 67 68 74 85 90 98 103 109 113 117 119 120 128 129 132 134 1 0 2 1 1 1 0 1 1 0 40 75 Cguentheri 4 33 51 53 65 72 2 2 1 0 1 1 2 2 2 1 2 0 2 1 0 1 1 0 0 1 1 1 1 2 0 1 0 1 48 58 86 91 109 124 133 1 1 1 1 1 2 Pweberi 3 2 1 2 0 1 0 11 19 63 72 119 6 7 43 50 52 74 75 120 122 128 129 134 Pcorrugata 1 2 2 3 2 0 2 1 3 1 2 0 1 0 1 2 1 15 18 83 1 11 13 31 48 58 84 92 93 99 133

0 0 1 Psolomonis 8 12 14 40 71 1 0 1 1 3 2 0 1 0 2 2 5 10 16 21 68 87 94 99 128 132 134 8 34 37 38 59 129 1 1 0 1 0 2 23 26 29 33 34 41 63 69 74 117 119 123 Dbufoniformis 0 1 0 0 0 1 0 0 1 2 1 0 1 0 1 1 1 4 7 18 36 44 46 53 61 65 78 88 93 97 133 1 1 1 1 0 0 2 0 2 1 1 0 0 Dguppyi 2 2 1 1 0 1 0 0 0 0 2 2 3 0 6 9 19 29 60 63 93 101 103 122 15 48 58 116 118 119 Pschmidti 2 1 2 1 2 1 0 2 1 0 33 34 35 41 72 74 91 132 0 3 2 1 1 2 Pmyersi 0 0 0 1 3 2 2 2 7 18 71 90 97 99 3 21 40 45 46 68 98 109 111 115 118 132 134 135 Pparkeri 0 0 0 0 3 2 0 0 0 0 0 0 1 0 1 0 3 2 1 0 1 8 14 27 31 68 77 85 88 90 Ibaluensis 4 6 24 25 27 102 114 13 38 47 74 93 119 134 1 1 0 1 1 2 0 1 1 1 0 4 22 24 62 70 80 88 93 95 102 108 112 117 125 126 127 133 135 0 0 1 1 0 0 0 1 2 2 1 0 1 0 3 33 34 66 71 82 96 103 114 123 Snatator 0 2 0 0 1 0 1 2 2 0 1 2 1 1 0 1 1 0 2 5 12 16 17 21 30 32 63 73 74 79 89 105 109 115 122 132 134 0 0 0 0 1 0 1 1 1 1 23 32 37 49 51 53 62 64 65 80 104 129 Pasolomonis 0 1 0 0 0 0 2 1 2 2 0 0 1 0 1 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 6 33 55 85 108 Belegans 2 0 0 1 1 16 17 30 54 70 83 87 107 116 117 120 132 133 45 62 79 129 Bvertebralis 0 0 0 1 0 2 1 1 2 1 0 0 0 3 1 0 1 106 121 18 56 90 99

1 0 Bwolfi 38 47 84 1 1 1 1 19 0 0 1 1 3 85 113 Bmediodiscus 1 5 33 50 98 116 118 134 1 0 1 2 Btrossulus 0 0 1 1 0 3 0 Fig. 6.3. Tree 3 of Analysis 1. L = 639, CI = 31, RI = 50. The Solomon Island species are distributed amongst five clades. This tree differs from Trees 1 and 2 by the placement of the clade containing species of Batrachylodes and the relationships of the species Platymantis guppyi and P. neckeri.

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Fig. 6.4. Strict Consensus Tree for 3MPTs from analysis 6. Tree Length = 663, CI = 30, RI = 48, t = 0.15. Bremer support values are shown below the line.

In terms of Bremer values, the most strongly supported taxa in analysis 1 is the clade containing the species of Batrachylodes, with greater than 10 steps needed before resolution is lost (Fig. 6.4). Support for monophyly at terminal clades is strong ranging from 9 steps for the Rana clade (R. grisea R. kreffti); 8 steps for the Discodeles clade (D. bufoniformis D. guppyi); 6 steps for (P. schmidti P. myersi) and 5 steps for (S. natator Pa. solomonis) (Fig. 6.4). However, overall the support for most clades is strong with most ranging between 2 and 4 steps (Fig. 6.4). When the three MPTs are presented as a strict consensus tree (Fig. 6.4) the relationships of the Solomon Island ranid frogs can be seen more clearly. Consistent topologies occurring in the three trees can be seen in clades (M. orphnocnemis(R. grisea R. kreffti) suggesting a Bornean/ New Guinea origin for R. kreffti. The clade is supported by Bremer value of 6. Other resolved areas on the tree are the relationships within the Solomon Island genus Batrachylodes and their relationship with the clade (I. baluensis(S. natator Pa. solomonis) indicating again

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a Bornean origin for these frogs. The sister group to these clades is P. parkeri (Figs 6.1 - 6.4), a species from the Solomon Islands, with a Bremer support value of 6 steps, again indicating a strong relationship. The Solomon Island frog Platymantis solomonis and the two species of Discodeles form a clade that has a Philippine ranid frog, Platymantis corrugata as its sister group. This suggests a different origin and hence invasion event into the Solomon Islands. Yet support for this clade is not as strong as others with a Bremer support value of 2 steps. In addition the clade appears to be related to other Solomon Islands/New Guinea species and is unresolved at is base (Fig. 6.4). The entire Solomon Island ingroup (except R. kreffti) is not resolved at its basal clade and the tree collapses splitting into five groups. This may be due to a possible paraphyly within the genus Platymantis, as the tree does indicate that some of its members are more closely related to other species than to each other, i.e. Platymantis appears not to be monophyletic. Alternatively, character selection may be dubious. This latter aspect is investigated is section 6.4. In examining the three MPTs (Figs 6.1 - 6.3) it can be seen that the Solomon Island ranid frogs do not form a monophyletic group and hence the null hypothesis is accepted. Data indicate there may have been at least six entries into the Solomon Islands. Firstly (and more probably, the most recent invasion) is the clade, common to all three MPTs, consisting of (M. orphnocnemis(R. grisea R. kreffti)) (Figs 6.1 - 6.3). The one member of the genus Rana recognised from the Solomon Islands (in this study) is the sister taxon of the New Guinean Rana grisea and together they form a clade with the Bornean Meristogenys orphnocnemis (Fig. 6.5). The clade comprising only R. grisea and R. kreffti is supported by 10 synapomorphies in trees 1 and 2 (Fig. 6.5).

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Fig. 6.5. Subsection of trees 1, 2, 3, (Fig. 6.1, 6.2, 6.3) from analysis 1 highlighting the clade containing the Solomon Island ranid frog, Rana kreffti.

Two of these, (97(1)) the position of the dorsal protuberance on the ilium is 1/3 anterior to the upper rim of the acetabulum and (115(3)) a large tympanum (greater than 53% of the head width) are unequivocal. Other synapomorphies supporting this monophyly are: (11(1)) a curved, posteriorly sloping alary process of the premaxilla, (26(1)) the posterior alary process of the vomers and the palatines are in contact, (40(0)) a shallow circummarginal groove on the toes, (44(1)) length of the 1st finger greater than 4th, (91(2)) a high dorsal crest (> 65% of ilium height), (93(0)) a straight dorsal acetabular expansion on the ilium, and (99(1)) the superior margin of the acetabulum is level with the inferior margin of the ilial shaft, all of which are parallelisms. The remaining synapomorphy, (3(0)) nasals widely separated is a reversal. Tree 3 conflicts with trees 1 and 2 in having 11 synapomorphies supporting this clade (Fig. 6.5). Support for the tree is the same as in trees 1 and 2 except for the addition of synapomorphies: (37(0)) a shallow almost indistinct circummarginal groove on the fingers and (51(1)) a raised, protruding middle palmar tubercle, both parallelisms. Tree 3 deletes character 99 at this clade (Fig. 6.5). The clade which encompasses the Bornean species, Meristogenys orphnocnemis as the sister taxon to (R. grisea R. kreffti) is supported by eight synapomorphies in trees 1 and 2 (Fig. 6.5). Two characters: (28(0)) vomerine teeth positioned between the choanae and (70(1)) the omosternum is slightly forked at its base are unequivocal. The clade is also supported by (4(2)) the frontoparietal is notched anteriorly, (68(0)) pointed toes discs, (104(0)) the absence of conical tubercles on upper eyelids and (128(1)) the presence of a dorsolateral fold. All of these characters are reversals whereas (0(0)) narrow

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nasals and (71(1)) a long and thin bony sternum are parallelisms. Tree 3 conflicts with trees 1 and 2 by having seven synapomorphies (Fig. 6.5), differing in the addition of the synapomorphy (18(0)) no inward curve at the base of cultriform process, a parallelism and the deletion of characters 4 and 76. Another clade represented in all three MPTs (Fig. 6.1, 6.2, 6.3) is the one consisting of species from the Solomon Island genus Batrachylodes: (B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus)))) (Fig. 6.6).

Fig. 6.6. The Batrachylodes clade, an subset from Fig. 6.1, 6.2 and 6.3

In all three trees, the sister taxa B. mediodiscus and B. trossulus are supported by four synapomorphies, however none of these is unequivocal (Fig. 6.6). Synapomorphy (1(1)) the nasals are closely applied or fused to the frontoparietal is a parallelism. Reversals include (85(1)) a square hyoid plate, (3(0)) nasals are widely separated from each other and (113(2)) a broad snout. The clade (B. wolfi(B. mediodiscus B. trossulus)) is supported by the same three synapomorphies in all three trees (Fig. 6.6). Again none are unequivocal and the support can be described at best, as weak (Bremer support = 1, Fig. 6.4). Two of the characters, 38 and 47, are based on features not present in the outgroups, so therefore (38(0)) a ventrally positioned circummarginal groove on fingers and (47(0)) round finger discs are parallelisms. The other synapomorphy, (84(1)) the alary process of hyoid is level with anterior margin of hyoid plate is a reversal. The clade (B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))) is supported by two synapomorphies, again common to all three trees (Fig. 6.6). One

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synapomorphy, (106(1)) a straight loreal region is unequivocal. The other, (121(0)) absence of a tongue notch is a parallelism. Finally the clade including all the species of Batrachylodes (examined in this study) i.e. (B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus)))) is supported by 13 synapomorphies in all three MPTs. However, the designation of these synapomorphies differs in tree 3 from trees 1 and 2 (Fig. 6.6). In trees 1 and 2 there are three unequivocal synapomorphies supporting the Batrachylodes clade. These are (2(0)) a short maxillary process of the nasal, (54(1)) the presence of a 4th palmar tubercle and (83(2))the posterolateral horn of the hyoid is elongate with a broad, rounded tip. Of the remaining characters: (16(0)) the cultriform process of the parasphenoid does not reach the palatines, (17(0)) an obtuse cultriform process, (87(1)) the posterolateral horn of hyoid is longer than the length of hyoid plate, (107(1)) small white tubercles present on chin of males, (116(2)) the choanae are larger than the eustachian openings and (132(0)) the texture of the dorsum is smooth, are parallelisms. Whereas (30(0)) the medial ramus of pterygoid does not reach the prootic, (70(0)) an unforked omosternum, (117(1)) oval shaped choanae and (120(0)) tongue broad posteriorly and tapering to a point anteriorly are all reversals. Tree 3 differs in the deletion of character 2 (Fig. 6.6) and therefore the clade is supported by only two unequivocal synapomorphies, and the addition of 133(0) texture of venter is smooth, which is a parallelism. The Batrachylodes clade is one of two clades supported by eight synapomorphies (Figs 6.1 - 6.3). The other clade being that of (Ibaluensis(Snatator Pasolomonis) which contains two species from Borneo and one from the Solomon Islands. This clade is common to all three MPTs. The Solomon Island species, Palmatorappia solomonis is the sister species to Staurois natator, which is found in Borneo and the Philippines, and this clade is supported by ten synapomorphies in all three MPTs (Fig. 6.7). One is unequivocal (82(0)) the absence of a posterolateral horn on the hyoid. Of the remaining: (3(0)) nasals widely separated from each other, (34(0)) no sesamoid bones at interphalangeal joints of feet, (96(1)) the inferior margin of dorsal protuberance of the ilium is below the anterior rim of acetabulum and (123(1)) narrow tongue notch are reversals. Whilst: (33(0)) no sesamoid bones at interphalangeal joints of hand, (66(0)) length of 3rd toe is shorter than 5th, (71(1)) a long, slender bony sternum,

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(103(1)) the canthus rostralis viewed dorsally is curved and (114(1)) a long snout are parallelisms.

Fig. 6.7. Clade containing Palmatorappia solomonis

The larger clade encompassing Ingerana baluensis from Borneo as the sister species to the Palmatorappia, Staurois clade is supported by seven synapomorphies in all three MPTs (Fig. 6.7). Two are unequivocal: (38(2)) a dorsally located circummarginal groove on the fingers and (47(2)) pointed finger discs. Of the five remaining synapomorphies, (13(1)) a quadratojugal spur present is a reversal and (74(1)) the width of the transverse process of vertebrae 2 being equal to width of vertebra 4, is a convergence (except Pa. solomonis), although there is lack of resolution at the base of the tree. (93(0)) the dorsal acetabular expansion of the ilium is straight (except Pa. solomonis), (119(1)) tongue free along half its length and (134(0)) texture of thighs is smooth (except Pa. solomonis) are parallelisms. As mention previously the clades ((I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))) are supported by eight synapomorphies in trees 1 and 2 and by 12 synapomorphies in tree 3 (Figs 6.1 - 6.3). In all, (62(0)) indistinct subarticular tubercles on the fingers is

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unequivocal. In trees 1 and 2: (32(1)) a T-shaped distal phalanx (except Pa. solomonis), (37(1)) a deep circummarginal groove on fingers, (49(0)) a low, flattened inner palmar tubercle, (64(0)) indistinct subarticular tubercles on toes, (65(0)) the anterior margin of the subarticular tubercles on the toes when viewed laterally is round and (80(0)) an absence of an anteromedial process on the anterior cornua of the hyoid (not seen in S. natator) are parallelisms, whilst the synapomorphy (104(0)) no conical tubercles on upper eyelid, is a reversal. Tree 3 conflicts with trees 1 and 2 in the addition of synapomorphies: (23(1)) the posterior alary process of vomers are not reduced, (51(0)) a low, flattened middle palmar tubercle, (53(0)) a low flattened outer palmar tubercle and (129(0)) absences of folds on the dorsum [except in B. vertebralis] all reversals. The sister group to the clades ((I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))) is Platymantis parkeri, a ranid from the Solomon Islands. This suggests not only immigration but emigration from the islands. The clade (P. parkeri((I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus)))))) is common to all three MPTs but each differ in their synapomorphy support. There are nine synapomorphies supporting the clade in tree 1 (Fig. 6.8): (24(1)) a narrow dentigerous process of vomers and (25(1)) vomerine teeth arising from the posterior margin of the dentigerous process are unequivocal. Synapomorphies (6(0)) zygomatic ramus shorter than otic ramus of squamosal (except B. elegans), (42(0)) length of 1st finger shorter than or equal to length of 2nd and (123(0)) a narrow tongue notch [except the clade (S. natator Pa. solomonis) and (B. vertebralis(B. wolfi(B. mediodiscus B. trossulus)))] are parallelisms whereas (12(2)) the lateral process of premaxilla is 2x the width of palatine process (except Pa. solomonis), (102(0)) a rounded canthus rostralis (except S. natator), (108(0)) an obtuse snout (except S. natator, B. elegans) and (114(0)) a short snout (except clade (S. natator Pa. solomonis)) are reversals. Tree 2 conflicts with Tree 1 in having ten synapomorphies supporting the clade with the addition of (72(2)) neural spines present only on vertebrae 2-4 [except clade (I. baluensis(S. natator Pa. solomonis))] a reversal (Fig. 6.8).

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Fig. 6.8. The relationship of Platymantis parkeri to two clades.

Tree 3 differs from trees 1 and 2 in only having seven synapomorphies supporting this clade (Fig. 6.8). There is an addition of synapomorphies (4(0)) anterior margin of frontoparietal acuminate (except S. natator), a convergence and (27(0)) no articulation between the vomers and the palatines (except M. baluensis) a reversal. Tree 3 differs from trees 1 and 2 in the deletion of characters 12, 42, 108, and 123, and from Tree 2 in the deletion of character 72. In trees 1 and 2 the clade (P. parkeri((I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus)))))) is one of two large clades supported by ten synapomorphies (Figs 6.1 and 6.2), the other being ((P. schmidti P. myersi)((P. guppyi P. neckeri)((P. papuensis(P. boulengeri C. guentheri))(P. weberi(P. corrugata(P. solomonis(D. bufoniformis D.

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guppyi))))))) in tree 1 and ((P. schmidti P. myersi)((P. guppyi P. neckeri)(P. weberi((P. papuensis(P. boulengeri C. guentheri))(P. corrugata(P. solomonis(D. bufoniformis D. guppyi))))))) in tree 2. The only difference between these clades is the placement of the Solomon Island species Platymantis weberi, as either the sister species to (P. corrugata(P. solomonis(D. bufoniformis D. guppyi))) or the sister species to ((P. papuensis(P. boulengeri C. guentheri))(P. corrugata(P. solomonis(D. bufoniformis D. guppyi)))). In both trees support is weak for either scenario as there are no unequivocal synapomorphies. Tree 3 differs in topology for this clade and will be discussed separately. The Solomon Island species of Discodeles: D. bufoniformis and D. guppyi are each other's closest relatives being supported by 13 synapomorphies in tree 1 and 11 in tree 2 (Fig. 6.9). However, none are unequivocal.

Fig. 6.9. The Discodeles clade.

In tree 1, (2(1)) a broad and long maxillary process of the nasal, (26(1)) an articulation between the posterior alary process of the vomers and the palatines, (29(1)) curved vomerine teeth, (33(0)) no sesamoids at interphalangeal joints in hand, (41(2)) a dorsally located circummarginal groove on toes, (117(1)) oval shaped choanae and (123(0)) a narrow tongue notch are parallelisms. Whereas (23(1)) the posterior alary process of the vomer is not reduced, (34(0)) no sesamoids on the interphalangeal joints of the foot, (63(0)) subarticular tubercles on the fingers have a rounded anterior margin in lateral view, (69(0)) rudimentary webbing on feet and (119(0)) tongue 1/3rd free posteriorly are reversals. The synapomorphy (74(1)) the width of the transverse processes of vertebra 2 equal to that of vertebra 4 is a convergence although there is little resolution at the base of the tree. Tree 2 differs in the deletion of the characters 63 and 119. Platymantis solomonis is the sister species of the clade (D. bufoniformis D. guppyi) and this clade is supported by five synapomorphies in tree 1 and six in tree 2, again none of which is unequivocal (Fig. 6.9). Synapomorphies include:

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(8(1)) a curved ramus of the squamosal, (14(0)) the sphenethmoid overlain entirely by nasals dorsally and (71(0)) a short and thick bony sternum all of which are reversals whereas (12(1)) the lateral process of the premaxilla being less than or equal to the palatine process of the premaxilla and (40(1)) a deep circummarginal groove on the toes are parallelisms. Tree 2 differs from tree 1 by the addition of 120(0) a rounded lateral margin of canthus rostralis, which is a reversal. It appears that the Solomon Island clade (P. solomonis(D. bufoniformis D. guppyi) has an origin in the Philippines since Platymantis corrugata is identified as its sister species in trees 1 and 2 (Fig. 6.10). This clade is held by three synapomorphies, none are unequivocal. In both trees 1 and 2 the synapomorphies (15(0)) sphenethmoid not projecting beyond palatines when viewed ventrally, (18(0)) base of cultriform process does not curve inwards (except D. guppyi) and (83(1)) a cuspidate posterolateral horn of the hyoid are all parallelisms.

Fig. 6.10. The relationship of the Philippine frog Platymantis corrugata to Solomon Island ranids.

The clade (P. weberi(P. corrugata(P. solomonis(D. bufoniformis D. guppyi)))) is supported by four synapomorphies in tree 1 only (Fig. 6.11). Of these, two are parallelisms: (11(1)) a curved, posterior sloping alary process of the premaxilla (except P. solomonis) and (119(2)) the tongue free for 2/3 of its length (except clade (D. bufoniformis D. guppyi)) and two are reversals: (19(2)) a V-shaped projection on the posteromedial margin of the parasphenoid and (63(2)) anterior margin of subarticular tubercles truncated in lateral view (except the Discodeles

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clade). If Platymantis weberi, a Solomon Island species, is the sister species to the clade (P. corrugata(P. solomonis(D. bufoniformis D. guppyi))) then movement has occurred from the Solomons to the Philippines and back again!

Fig. 6.11. The relationship of Platymantis weberi to the clade P. corrugata(P. solomonis(D. bufoniformis D. guppyi)) from Tree I Analysis 1.

The clade of (P. weberi(P. corrugata(P. solomonis(D. bufoniformis D. guppyi)))) is one of two supported by three synapomorphies in tree 1 (Fig. 6.1). The other clade comprises (P. papuensis(P. boulengeri C. guentheri)). Ceratobatrachus guentheri is a Solomon Island ranid and it appears to be closely related to Platymantis boulengeri from the neighbouring islands of New Britain. These two species form a clade of which the New Guinean Platymantis papuensis is the sister species (Fig. 6.12). The clade (P. boulengeri C. guentheri) is supported by ten synapomorphies in both trees 1 and 2 (Fig. 6.12.).

Fig. 6.12. The clade containing the Solomon Island ranid Ceratobatrachus guentheri

None are unequivocal. Those synapomorphies that are parallelisms are: (1(1)) nasals are closely applied or fused with the frontoparietal, (7(2)) the otic

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ramus of the squamosal overlaps the crista parotica, (9(1)) the width of the otic ramus of the squamosal is 2x that of the zygomatic, (93(0)) the dorsal acetabular expansion on the ilium is straight, (100(1)) a markedly curved ilial shaft, (111(1)) a vertically elliptical tympanum and (122(0)) a shallow tongue notch. Two synapomorphies are reversals: (8(1)) curved rami of the squamosal and (31(1)) a bony articulation between the pterygoid and quadratojugal. The synapomorphy, (4(0)) anterior margin of the frontoparietal is acuminate is a convergence. The clade (P. papuensis(P. boulengeri C. guentheri)) is supported by two synapomorphies in trees 1 and 2. They are: (52(2)) an obovate outer palmar tubercle and (130(1)) the presence of a tarsal fold, both parallelisms. In tree 1, three synapomorphies support the clades ((P. papuensis(P. boulengeri C. guentheri))(P. weberi(P. corrugata(P. solomonis(D. bufoniformis D. guppyi))))). Reversals are (12(2)) the width of the lateral process of the premaxilla is 2x that of the palatine process (except clade (P. solomonis(D. bufoniformis D. guppyi)) and (75(1)) proximal end of the posterior cornua of the hyoid expanded (except P. corrugata). Whereas (40(0)) a shallow circummarginal groove on the toes (except P. boulengeri and the clade (P. solomonis(D. bufoniformis D. guppyi))) is a parallelism. This arrangement of clades is not present in tree 2 (Fig. 6.1, 6.2). The clade of the Solomon Island species (P. guppyi P. neckeri) is supported by eight synapomorphies in both trees 1 and 2 (Fig. 6.13).

Fig. 6.13. The clade Platymantis guppyi and P. neckeri

None are unequivocal. Six are parallelisms: (37(1)) a deep circummarginal groove on the fingers, (38(0)) a ventral positioned circummarginal groove on fingers, (42(0)) the length of the 1st finger is less than or equal to the 2nd, (116(2)) the choanae are larger than the eustachian tube openings, (122(0)) a shallow tongue notch and (132(0)) a smoothly textured dorsum, whilst two are reversals: (8(1)) curved ramus of the squamosal and (59(0)) a low, flattened outer plantar tubercle. This clade is one of two supported by six synapomorphies, the other

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clade being ((P. papuensis(P. boulengeri C. guentheri))(P. weberi(P. corrugata(P. solomonis(D. bufoniformis D. guppyi))))) (Figs 6.1 and 6.2). Synapomorphies (7(1)) the otic ramus of the squamosal lies alongside the crista parotica (except P. boulengeri, C. guentheri, P. corrugata, D. guppyi) and (18(1)) the base of the cultriform process curves inwards (except P. corrugata, P. solomonis, D. bufoniformis) are reversals, whereas (71(1)) a long, thin bony sternum (except clade (P. solomonis(D. bufoniformis D. guppyi))), (90(1)) a long dorsal crest on ilium (except C. guentheri), (97(2)) half of the dorsal protuberance of the ilium is anterior to the anterior rim of the acetabulum (except P. boulengeri, C. guentheri, D. guppyi) and (99(1)) the superior margin of the acetabulum is level with the inferior margin of the ilial shaft (except (P. boulengeri C. guentheri), P. solomonis, D. bufoniformis) are parallelisms. The closest relative of the Solomon Islands species, Platymantis myersi, is the New Britain species, Platymantis schmidti. This relationship is seen in both trees 1 and 2 (Fig. 6.14).

Fig. 6.14. The clade containing Platymantis schmidti and P. myersi

The clade is supported by seven synapomorphies in tree 1, all of which are parallelisms: (2(1)) a broad, long maxillary process of the nasal, (15(0)) sphenethmoid does not project beyond the palatines seen ventrally, (48(3)) an obovate inner palmar tubercle, (58(2)) a conical outer palmar tubercle, (116(1)) size of the choanae equal to the eustachian tube openings, (118(1)) a triangular shaped eustachian tube opening and (119(2)) the tongue free 2/3 of its length. Tree 2 only identifies six synapomorphies, deleting character 119 (Fig. 6.14). This clade is one of two clades supported by three synapomorphies in both trees 1 and 2 (Figs 6.1 and 6.2). In tree 1, the other clade is ((P. guppyi P. neckeri)((P. papuensis(P. boulengeri C. guentheri))(P. weberi(P. corrugata(P. solomonis(D. bufoniformis D. guppyi)))))), whereas in tree 2 it is ((P. guppyi P. neckeri)(P. weberi((P. papuensis((P. boulengeri C. guentheri))(P. corrugata(P. solomonis(D. bufoniformis D. guppyi)))))) (Fig. 6.1, 6.2). One of the three supporting

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synapomorphies, (4(1)) the anterior margin of the frontoparietal truncated is unequivocal. Of the other two: (27(1)) the presence of an articulation between the vomers and the palatines is a parallelism and (61(1)) a rugose plantar surface (except P. neckeri, D. guppyi) is a reversal. These large clades: ((P. schmidti P. myersi)((P. guppyi P. neckeri)((P. papuensis(P. boulengeri C. guentheri))(P. weberi(P. corrugata(P. solomonis(D. bufoniformis D. guppyi))))))) in tree 1 and ((P. schmidti P. myersi)((P. guppyi P. neckeri)(P. weberi((P. papuensis(P. boulengeri C. guentheri)))(P. corrugata(P. solomonis(D. bufoniformis D. guppyi))))))) in tree 2 are supported by ten synapomorphies. Four of the ten synapomorphies are unequivocal: (7(0)) the otic ramus of the squamosal is well separated from the crista parotica, (125(1)) large eggs, (126(0)) unpigmented eggs, and (127(0)) number of eggs is few <50. There is one reversal: (65(1)) anterior margin of the subarticular tubercles of the toes pointed, as viewed laterally (except clade (B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus)))), P. guppyi, D. guppyi) and five parallelisms: (8(0)) rami of squamosal straight (except I. baluensis, (P. guppyi P. neckeri), (P. boulengeri C. guentheri) and (P. solomonis(D. bufoniformis D. guppyi))), (13(0)) no quadratojugal spur (except (I. baluensis(S. natator Pa. solomonis), P. boulengeri, P. solomonis), (75(0)) proximal end of the posterior cornua of the hyoid unexpanded (except M. orphnocnemis, S. natator, P. papuensis(P. boulengeri C. guentheri), P. weberi, P. solomonis(D. bufoniformis D. guppyi)), (85(0)) a narrow hyoid plate (except I. baluensis, B. elegans, (B. mediodiscus B. trossulus), C. guentheri) and (88(0)) the ventral acetabular expansion of the ilium forms a straight line (except I. baluensis, S. natator, P. papuensis, C. guentheri, P. weberi, D. guppyi). Six synapomorphies, four of which are unequivocal support the large clade ((M. orphnocnemis(R. grisea R. kreffti))((P. parkeri((I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))))((P. schmidti P. myersi)((P. guppyi P. neckeri)((P. papuensis(P. boulengeri C. guentheri))(P. weberi(P. corrugata(P. solomonis(D. bufoniformis D. guppyi))))))))) in tree 1 (Fig. 6.1) and clade ((M. orphnocnemis(R. grisea R. kreffti))(P. parkeri((I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))))((P. schmidti P. myersi)((P. guppyi P. neckeri)(P. weberi((P. papuensis(P. boulengeri C. guentheri))(P. corrugata(P. solomonis(D.

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bufoniformis D. guppyi)))))))) in tree 2 (Fig. 6.2). The unequivocal synapomorphies are (39(1)) circummarginal groove on toes present, (76(0)) the distal end of the posterior cornua of the hyoid unexpanded, (86(0)) the length of the posterior cornua is shorter than the hyoid plate and (134(2)) the skin texture of the ventromedial aspect of the thigh is warty. Synapomorphies: (36(1)) presence of a circummarginal groove on the fingers (except P. papuensis, C. guentheri, P. corrugata, P. solomonis, D. bufoniformis) and (46(1)) presence of discs on fingers (except R. grisea, P. parkeri, clade (P. papuensis(P. boulengeri C. guentheri)), P. corrugata, P. solomonis, D. bufoniformis) are parallelisms. The sister species to the aforementioned large clade in both trees 1 and 2 is the Philippine ranid frog Limnonectes magnus (Figs 6.1 and 6.2). This clade is supported by seven synapomorphies with (31(0)) a cartilaginous articulation between the posterior ramus of the pterygoid and the quadratojugal, (106(0)) a concave loreal region and (120(2)) the tongue is broad but narrow anteriorly being unequivocal. The remainder: (61(0)) a smooth plantar surface (except P. schmidti, P. myersi, P. guppyi, P. weberi, P. papuensis, P. boulengeri, C. guentheri, P. corrugata, P. solomonis, D. bufoniformis), (102(1)) an angular canthus rostralis (except P. weberi, P. solomonis, D. bufoniformis, D. guppyi, P. parkeri, I. baluensis, Pa. solomonis, B. elegans, B. vertebralis, B. wolfi, B. mediodiscus, B. trossulus), (108(1)) a pointed snout (except P. myersi, P. guppyi, P. solomonis, D. bufoniformis, P. parkeri, I. baluensis, Pa. solomonis, B. vertebralis, B. wolfi, B. mediodiscus, B. trossulus) and (114(1)) a long snout (except P. parkeri, I. baluensis, B. elegans, B. vertebralis, B. wolfi, B. mediodiscus, B. trossulus) are all parallelisms. Although Tree 3 is similar to trees 1 and 2 in the topology of its terminal clades, the topology of the whole tree does differ (Fig. 6.3). The clade (P. parkeri((I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus)))))) is present and similar to that seen in trees 1 and 2. However the sister group to this clade is the clade (P. schmidti P. myersi) (Fig. 6.15). This latter clade (P. schmidti P. myersi) is supported by six synapomorphies, not seven as seen in trees 1 and 2, differing in the deletion of character 2. The clade ((P. schmidti P. myersi)(P. parkeri((I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))))) is supported by six synapomorphies one of which, (7(0)) no

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articulation between the otic ramus of the squamosal and the crista parotica is unequivocal (Fig. 6.14). Synapomorphy (18(0)) the base to the cultriform process does not curve inwards (except B. wolfi) is a parallelism, whilst (71(0)) a short, thick bony sternum (except clade (S. natator Pa. solomonis), (90(0)) a short dorsal crest on ilium (except I. baluensis, Pa. solomonis, B. wolfi), (97(3)) dorsal protuberance of ilium positioned 2/3 above acetabulum (except I. baluensis, S. natator, the Batrachylodes clade) and (99(2)) the position of the superior margin of the acetabulum is well above the inferior margin of the ilial shaft (except B. wolfi) are reversals.

Fig. 6.15. Clade extracted from Fig. 6.3.

In tree 3 the clade ((P. papuensis(P. boulengeri C. guentheri))(P. weberi(P. corrugata(P. solomonis(D. bufoniformis D. guppyi))))) is the same as that seen in tree 1 (Fig. 6.1, 6.3). The only exceptions being two synapomorphies supporting the entire clade instead of three (character 12 is deleted in tree 3) and five synapomorphies not four supporting the clade (P. weberi(P. corrugata(P. solomonis(D. bufoniformis D. guppyi)))). Synapomorphy (72(3)) neural spines present on all vertebrae is a parallelism in tree 3. The clade ((P. papuensis(P. boulengeri C. guentheri))(P. weberi(P. corrugata(P. solomonis(D. bufoniformis D. guppyi))))) together with the already mentioned larger clade of ((P. schmidti P. myersi)(P. parkeri((I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))))) forms an even larger clade supported by six synapomorphies, although none are unequivocal (Fig. 6.3). All

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synapomorphies are parallelisms, they are: (8(0)) rami of the squamosal are straight (except clades (P. boulengeri C. guentheri), (P. solomonis(D. bufoniformis D. guppyi)), and I. baluensis), (34(1)) sesamoid bones at the interphalangeal joints of the foot (except (D. bufoniformis D. guppyi), P. myersi, (S. natator Pa. solomonis)), (37(0)) a shallow circummarginal groove on fingers (except clade (I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))), (38(1)) a terminally positioned circummarginal groove on fingers (except clades (I. baluensis(S. natator Pa. solomonis)) and (B. wolfi(B. mediodiscus B. trossulus)), (59(1)) a protrudent, raised outer plantar tubercle (except S. natator) and (129(1)) longitudinal ridges on dorsum (except C. guentheri, P. corrugata, clade (I. baluensis(S. natator Pa. solomonis)), B. elegans, (B. wolfi(B. mediodiscus B. trossulus))). The sister species to this large clade is Platymantis neckeri and this is supported by one unequivocal synapomorphy, (4(1)) a truncated anterior margin of the frontoparietal and three reversals: (33(1)) sesamoids at interphalangeal joints on hand (except (D. bufoniformis D. guppyi), P. myersi, (S. natator Pa. solomonis), B. elegans, B. trossulus), (65(1)) a pointed anterior margin to subarticular tubercles on toes (except D. guppyi, clade (I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))), (72(2)) neural spines present only on vertebrae 2-4 (except P. boulengeri, clade (P. weberi(P. corrugata(P. solomonis(D. bufoniformis D. guppyi)))), P. myersi and clade (I. baluensis(S. natator Pa. solomonis))) and two parallelisms: (51(1)) a protruding, raised middle palmar tubercle (except clade (I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))) and (53(1)) a protrudent, raised outer palmar tubercle (except D. guppyi and clade (I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))) (Fig. 6.3). Platymantis guppyi completes the clade as the sister species to all the above (Fig. 6.3). There are ten synapomorphies supporting this clade, three of which are unequivocal. They are: (125(1)) large eggs, (126(0)) unpigmented eggs and (127(0)) eggs few in number. The remaining seven synapomorphies are: (7(1)) otic ramus of squamosal lies alongside crista parotica (except (P. boulengeri C. guentheri), P. corrugata, D. guppyi, clade ((P. schmidti P. myersi)(P. parkeri((I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B.

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mediodiscus B. trossulus)))))), a reversal, (13(0)) no quadratojugal spur (except P. boulengeri, P. solomonis, (I. baluensis(S. natator Pa. solomonis))), (23(0)) a reduced posterior alary process of the vomer (except C. guentheri, (D. bufoniformis D. guppyi), clade (I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))), (75(0)) the proximal end of the posterior cornua of the hyoid is unexpanded (except I. baluensis, S. natator, P. papuensis, P. boulengeri, C. guentheri, P. weberi, P. solomonis, D. bufoniformis, D. guppyi), (85(0)) a narrow hyoid plate (except C. guentheri, M .baluensis, B. elegans, B. mediodiscus, B. trossulus) and (88(0)) a straight ventral acetabular expansion on the ilium are parallelisms. The only convergence is (97(2)) the dorsal protuberance of ilium is positioned 1/2 above anterior rim of acetabulum. The already discussed clade of (M. orphnocnemis(R. grisea R. kreffti) (Fig. 6.4) joins with this large clade and the resulting clade is supported by seven synapomorphies, unlike the six seen in trees 1 and 2 (Fig. 6.1, 6.2, 6.3). The difference being the addition of synapomorphy (71(1)) a long, thin bony sternum (except clades (P. solomonis(D. bufoniformis D. guppyi)), ((P. schmidti P. myersi)(P. parkeri((I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus)))))), (S. natator Pa. solomonis)) is a parallelism. Completing the tree, Limnonectes magnus is the sister species to the clade ((M. orphnocnemis(R. grisea R. kreffti))(P. guppyi(P. neckeri(((P. papuensis(P. boulengeri, C. guentheri))(P. weberi(P. corrugata(P. solomonis(D. bufoniformis D. guppyi)))))((P. schmidti P. myersi)(P. parkeri((I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))))))))). This is supported by seven synapomorphies, the only difference between tree 3 and trees 1 and 2 being the replacement of character 61 for 33, a parallelism. Like trees 1 and 2, Occidozyga laevis is the sister species to all the clades with Rana temporaria being set as the outgroup (Figs 6.1 - 6.3).

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6.4 Character Reassessment In cases where assumption sets produced more than one maximum parsimonious tree, a strict consensus tree is reported.

6.4.1 Analysis 2 Data set baa (Table 6.3). One MPT, L=8420 CI=44, RI=70

Using a base weight of 100, the reweighted length of the cladogram in Fig. 6.16, obtained after four successive approximation runs, is 8420. This cladogram was constructed using an heuristic analysis and not a ratchet analysis as the latter resets the data set to equal weights (Nixon, 1999). The reweighted branch support values range between 33 and 263; thus cladograms 8520 reweighted steps long are necessary to lose all groups in the consensus. Due to the base weight of 100 assigned to this analysis, these reweighted branch support values are difficult to compare with the equal-weighted values found in analysis 1. So to counteract this, the reweighted values were rescaled so that the cladogram in Fig. 6.16 and 6.17 become 639 steps long (Gustafsson and Bremer, 1995). If this is done the rescaled and reweighted support values ranges between (33[8420/639]) = 2.5 and (263[8420/639]) = 19.95 = 20.

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Fig. 6.16. Most parsimonious tree for analysis 2 after four successive weighting runs.

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Fig. 6.17. Bremer support values for analysis 2. N.B. Bremer support values have been rescaled (see text for details).

After weighting characters, one most parsimonious tree was found with a ci of 44 and a ri of 70. There are some similarities in the tree to those found from analysis 1. The clade (M. orphnocnemis(R. grisea R. kreffti) is present and strongly supported with a Bremer value of 9 steps (Fig. 6.17). The species of Batrachylodes form an identical clade as seen in the first analysis (Fig. 6.17 compared with 6.1, 6.2, 6.3) but with stronger support. The Philippine origin of the (P. solomonis(D. bufoniformis D. guppyi) is still evident and well supported (Fig. 6.17). The relationship between C. guentheri and P. boulengeri from New Britain is again present. The major difference in the tree reconstructed using successive weighting is of the collapsing of the clade (I. baluensis(S. natator Pa. solomonis) (Fig. 6.16). In this analysis these species do not form a clade but become sister species to the Batrachylodes clade (Fig. 6.18).

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Fig. 6.18. Different clade arrangement for Bornean species and Solomon Island species due to successive weighting.

Nine synapomorphies support the clade (Pa. solomonis(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus(B. trossulus)))))): (67(1)) medium leg length as measured against snout vent is unequivocal. The synapomorphy (7(0)) otic ramus of the squamosal well separated from the crista parotica is a convergence; whereas (16(0)) a short cultriform process, (17(0)) an obtuse cultriform process, (75(0)) an unexpanded proximal end of the posterior cornua of the hyoid, (92(1)) an oval shaped dorsal protuberance on the ilium (except B. wolfi), (122(0)) a shallow tongue notch and (132(0)) a smooth dorsum are parallelisms. The only reversal being (30(0)) a reduced medial ramus of the pterygoid. Ingerana baluensis is the sister species to this clade being supported by seven synapomorphies (Fig. 6.18). Two are unequivocal: (62(0)) indistinct subarticular tubercles on fingers (except B. vertebralis) and (97(4)) the dorsal protuberance of the ilium is entirely anterior to the acetabulum (except Pa. solomonis). An acuminate anterior margin of the frontoparietal (4(0)), is a convergence. Parallelisms are: 24(1) long, acuminate vomers, (80(0)) no anteromedial process on the anterior cornua of the hyoid, (86(1)) the length of the posterior cornua equals that of the hyoid plate (except Batrachylodes clade). Whilst (102(0)) a rounded canthus rostralis is a reversal. Finally the Bornean ranid, Staurois natator is the sister species to the entire clade (Fig. 6.18) for which there are seven synapomorphies two of which are unequivocal: (38(2)) a dorsally positioned circummarginal groove on the fingers (except Batrachylodes clade) and (72(0)) no neural spines on vertebra (except Batrachylodes clade and I. baluensis). Of the remaining synapomorphies: (6(0)) a shorter zygomatic ramus compared to otic ramus of the squamosal (except B. elegans), (41(2)) a dorsally positioned circummarginal groove on the toes (except Batrachylodes clade),

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(65(0)) subarticular tubercles on toes are rounded distally and (67(2)) a long leg length relative to snout-vent (except Batrachylodes clade and I. baluensis) are parallelisms. The synapomorphy (12(2)) a wide lateral process of the premaxilla (except Pa. solomonis) is a reversal. Support for this arrangement is high with Bremer values of 11, 14 and 20 respectively (Fig. 6.17). The tree (Fig. 6.16) identifies two large clades for the Solomon Island ranid frogs. One being (S. natator(I. baluensis(Pa. solomonis(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))))) and the other (P. guppyi(P. neckeri((P. schmidti P. myersi)(P. parkeri(P. boulengeri C. guentheri)))((P. weberi(P. papuensis(P. corrugata(P. solomonis(D. bufoniformis D. guppyi)))))))). The former clade suggests a Bornean origin for the Batrachylodes species and Palmatorappia solomonis as does the latter clade if you follow the tree back to the sister clade of (M. orphnocnemis(R. grisea R. kreffti)). But in the latter clade there has been movement from the Solomon Islands to New Britain and New Guinea and then re-entry into the Islands. The only anomaly is the Philippine species of Platymantis (P. corrugata) nested within the New Guinean/Solomon Island clade (Fig. 6.16). The latter clade splits into two clades with one ((P. schmidti P. myersi)(P. parkeri(P. boulengeri C. guentheri))) indicating a New Britain origin. The tree does resolve the relationships of P. neckeri, P. guppyi and P. weberi unlike that seen in analysis 1 (compare Figs 6.16 and 6.17 with 6.1, 6.2, 6.3).

6.4.2 Analysis 3

Data set aba (Table 6.3). 3 MPTs, L= 688, CI = 32, RI = 51

Inapplicable character states were recoded as an extra state “not present” Strict Consensus Tree L = 713, CI = 31, RI = 49 (Fig. 6.19).

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Fig. 6.19. Strict Consensus tree from analysis 3

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Fig. 6.20. Bremer support values for Analysis 3.

When inapplicable missing states are manipulated, longer trees are found. This is probably due to there being more changes due to extra character states. Again the monophyly of the Rana clade (M. orphnocnemis(R. grisea R. kreffti)) is well supported and the large clade comprising (P. parkeri((I. baluensis(S. natator Pa. solomonis))(B. elegans(B. vertebralis(B. wolfi(B. mediodiscus B. trossulus))))) is also present (Fig. 6.19). In addition the clade (P. corrugata(P. solomonis(D. bufoniformis D. guppyi))) and the clades (P. guppyi P. neckeri); (P. schmidti P. myersi); (P. boulengeri C. guentheri) are all present with support (Fig. 6.20). However the tree collapses in consensus with seven clades becoming unresolved.

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6.4.3 Analysis 4 Data set aab (Table 6.3). 20 multistate characters with a defined morphocline were ordered (NONA: additive FARRIS). 11 MPTs, L = 666, CI = 30, RI = 50

Fig. 6.21. Strict Consensus tree with Bremer support for Analysis 4.

When multistate characters are ordered the resolution of the tree is lost. The tree is longer than that found in analysis 1 and there are many more parsimonious trees found (11). There is no resolution for the majority of the species within the analysis, only support for the strong Batrachylodes clade.

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6.5 Discussion When the data are examined without character manipulation (analysis 1) three MPTs were found. Examination of these trees show no support for the hypothesis that "the Solomon Island ranid frogs are a monophyletic group". When the typology of the strict consensus tree (analysis 1) was forced to achieve a monophyletic origin for the Solomon Island ranids an extra 135 steps are needed (Fig. 6.22).

Rtemporaria

Olaevis

Morphnocnemis

Snatator

Ibaluensis

Lmagnus

Pcorrugata

Pboulengeri

Pschmidti

Ppapuensis

Rgrisea

Rkreffti

Pguppyi

Pmyersi

Pneckeri

Pparkeri

Psolomonis

Pweberi

Cguentheri

Dbufoniformis

Dguppyi

Pasolomonis

Belegans

Bvertebralis

Bwolfi

Bmediodiscus

Btrossulus Fig. 6.22. Forced topology for monophyly of Solomon Island ranids. Tree length = 774, ci = 26. ri = 35.

Such an arrangement, which still retains the clades for Batrachylodes and Discodeles, and allows for only one invasion event into the Solomon Islands from a New Guinean ancestor, is unattainable from this data set. Therefore the Solomon Island ranid frogs do not share a common ancestor and are not each others closest relatives. The null hypothesis is accepted in support of multiple

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origins for these frogs. Multiple invasions into the Solomon Islands by ranid frogs is given the most support from the character set of analyses 1 and 2. From analysis 1, there are at least two or three major clades (depending on the tree topology) all having an admixture of Solomon Island ranids and other South East Asian ranids (Figs 6.1 – 6.3). Results from analysis 1 also did not support Brown's (1952) predictions of two major radiations into the Solomon Islands. Admittedly there was one consistent Rana grouping (perhaps the recent element of Brown (1952)) but at no stage does this clade contain species from Platymantis, namely P. myersi and P. weberi, although Brown's (1952) assumptions that these species of Platymantis are closely related to New Guinean forms might be supported with further analyses (see below). Generalisations of the origins and relationships of the ranids from the Solomon Islands are as follows: species of the genus Batrachylodes form a well supported clade in all trees. Their origin lies in Borneo and/or the Philippines as this clade is closely allied with the clade containing the Bornean species Ingerana baluensis and the Philippine and Bornean species Staurois natator as well as another Solomon Island ranid, Palmatorappia solomonis. The ultimate sister taxon to these clades is the Solomon Island species Platymantis parkeri. Such an arrangement is well supported (Bremer support of 6, Fig. 6.4) and suggests emigration as well as invasion. Another possibility is that this tree topology may be a causal effect of miniaturisation rather than true relationships. All ranids in this area of the tree have snout-vent lengths less than 35 mm (except S. natator mean = 40.1 mm). However characters supporting the clade are not those that result from heterochrony as they are not size related, nor are they known states on ontogenetic trajectories in other frogs (M. Davies, pers comm.). The only species of Rana, R. kreffti from the Solomon Islands used in this study forms a strong clade with the New Guinean species, R. grisea. Completing the larger clade is the Bornean species, Meristogenys orphnocnemis. Brown (1952, 1996) suggested that the Rana species of the Solomon Islands was a recent arrival to the islands and results shown here do support this notion as all trees suggest that Rana kreffti and its allies differentiated prior to the Solomon Island lineage. As for the remainder of the ingroup, data suggest a New Britain or Solomon Island origin with multiple emigrations and invasion events. The ancestor to

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species within the genera Ceratobatrachus, Discodeles, Platymantis may be some form of "platymantid" frog. This may be Brown's (1952) "older element". It also supports Barbour's (1921) assumption that the fauna of the Solomon Islands "is obviously Papuan not Australian in origin" (Barbour, 1921:92). However, when the three trees from analysis 1 are combined in strict consensus, resolution of the entire Solomon Island ingroup (except R. kreffti) is lost at its basal clade. The tree collapses splitting into five groups indicating that Platymantis may be paraphyletic as many of its species are grouped with other species from different genera. For the suggestion of a "platymantid" ancestor to the Solomon Island ranids to be confirmed further examination of character selection is necessary. Analysis 1 suggests at least three major evolutionary events for the Solomon Island frogs, one involving Rana kreffti, another for Batrachylodes and Palmatorappia and a third (although with less support) involving Platymantis, Discodeles and Ceratobatrachus. The analyses that involved ordering multistate characters and rescoring missing values as an extra state gave trees that provided no resolution to the relationships of the Solomon Island ranids. For this reason analyses 3 and 4 were not considered informative and will not be discussed further. However to elucidate the possibility of multiple origins of ranids into the Solomon Islands suggested by analysis 1, results from analysis 2 (successive weighting) deserve consideration. Analysis 2 involved reanalysing the character set of analysis 1 by successive reweighting using the methods of Farris (1969, 1989). Successive weighting has been criticised by Swofford and Olsen (1990) for being circular however this is refuted by Carpenter (1994) and Farris (2001). After four weighting procedures one parsimonious tree was found whose tree length was rescaled to the length found in analysis 1 for comparative purposes (section 6.5.1). The tree produced three major clades containing the ingroup species (Fig. 6.16) and all were strongly supported (Fig. 6.17). Like analysis 1, one clade contains Rana kreffti which is closely related to the New Guinean R. grisea with a Bornean origin. A clade containing Batrachylodes (all species examined) and Palmatorappia solomonis also occurs. This clade differs from that in analysis 1 in that Palmatorappia is the sister species to Batrachylodes with their origins being in Borneo, as evidenced by Ingerana baluensis being the sister species to the clade. The Philippine and

227

Bornean frog, Staurois natator completes the clade. The other major clade contains all species of Platymantis, Discodeles and Ceratobatrachus. The species of Discodeles have previously been considered to be closely related to Platymantis (Boulenger, 1918b). This clade is strongly supported (Fig. 6.17). However if this topology is accepted as providing the most likely relationships, clarification of origins by the tree indicates that these ranids have their ancestral origin from within the Solomon Islands because two endemic species, P. guppyi and P. neckeri form the basal part of the clade, i.e. they are sister species to the entire clade. This may also corroborate findings that the Solomon Island ranids had undergone a long period of isolation (Barbour, 1921; Brown, 1996; Noble, 1931; Burt and Burt, 1932, Myers 1954, Tyler, 1972) as indicated by their endemicity and reflecting a possible centre of differentiation (Burt and Burt, 1932). Brown (1952) also suggested that his "older element" (Batrachylodes, Ceratobatrachus, Discodeles, Palmatorappia and some Platymantis) of Solomon Island ranids might have its centre of origin in the Solomons/Bismarck Archipelago. Indeed, Brown (1965) when recognising Platymantis (then Cornufer) as one genus commented that "Cornufer appears to represent a relict, peripheral group of ranid frogs…" (Brown, 1965:3). Within this clade, relationships suggest patterns of movement of species entering New Britain and New Guinea from the Solomon Islands, a concept not unlike that suggested by Brown and Tyler (1968) and Tyler (1979). The placement of the Philippine frog Platymantis corrugata nested with this clade however, may be an artefact, merely showing that it is a Platymantis with perhaps stronger relationships with other Philippine platymantid frogs. As other platymantid frogs from the Philippines were not included in the outgroup this cannot be tested. It is hard to support the ideas of Dubois (1981, 1986, 1992) for separate subfamilial status for the Solomon Island frogs because Palmatorappia and Batrachylodes are shown as closely related in every analysis whereas Dubois consistently places them in separate tribes. There is some support for Noble's (1931) and Savage's (1973) separation of the Solomon Island ranids into the subfamily Platymantinae, as well as Laurent's (1986) concept of the tribe, Platymantini. Unlike the subfamilial composition of Dubois (1992) these other subfamilial groupings do include Batrachylodes with the other Solomon Island genera as well as other southeast Asian ranids. Dubois (1986, 1992) had based

228

his relationships on characters that reflected the mode of reproduction whereas Savage (1973), Laurent (1986) and to a lesser degree Noble (1931) did not. In this study such characters do form the basal part of the ingroup clade in analysis 1 and Batrachylodes is included in all the clades. Hence, Dubois' (1986, 1992) placement of Batrachylodes in a separate subfamily to the remaining Solomon Island genera (except Rana) must ultimately be based on characters of the omosternum. This is not supported in the current analysis. The subfamilial rank of Platymantinae as proposed by Brown (1996), Tyler (1979) and Zweifel and Tyler (1982) (i.e. restricted to the genera Ceratobatrachus, Discodeles, Palmatorappia and Platymantis) cannot be supported if the phylogenies generated by analysis 1 are accepted. In all trees, southeast Asian ranids (e.g. Staurois, Ingerana) are nested within terminal clades, as is the Solomon Island genus Batrachylodes. However, some consideration may be given to this subfamilial structure when examining the tree generated from analysis 2. The southeast Asian ranid genera, Staurois and Ingerana form a distinct clade containing Batrachylodes and Palmatorappia whereas the genera Ceratobatrachus, Discodeles and Platymantis also form a distinct clade, perhaps this latter grouping is the Platymantinae of Brown (1996), Tyler (1979) and Zweifel and Tyler (1982). As to the basal part of the clade, in all analyses the origin is an Oriental one. The Philippine ranids Occidozyga laevis and Limnonectes magnus complete the basal part of the tree. Tyler (1979) proposed an Oriental origin, in particular the Philippines, for the Solomon Island ranids and data from this analysis support this hypothesis. However, the relationships of the Solomon Island ranids will remain contentious until the phylogeny of the speciose and widely distributed genus Platymantis is resolved.

229 References

Ahl, E. (1927) Ueber neue oder seltene Froschlurche aus den Zoologischen Museum Berlin. Sitzungsberichte Gesselschaft Naturforschenden Freunde, 111-117. Alcala, A. C. (1986) Guide to Philippine Flora and Fauna: Amphibians and Reptiles. National Resources Management Center, Ministry of Natural Resources and University of the Philippines, Manila. Allison, A. (1996) Zoogeography of amphibians and reptiles of New Guinea and the Pacific region. In The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: patterns and processes. (Eds Keast, A. and MIller, S.E.) Academic Publishing, Amsterdam, pp 407-436. Asher, R. J. (1999) A morphological basis for assessing the phylogeny of the "Tenrecoidea" (Mammalia, Lipotyphla). Cladistics, 15, 231-252. Atoda, K. (1950) Metamorphosis of the "Non-aquatic Frog" of the Palau Islands, Western Carolines. Pacific Science, 4, 202-207. Barbour, T. (1921) Reptiles and amphibians from the British Solomon Islands. Proceedings of the New England Zoölogical Club, 7, 91-112. Birstein, V. J. (1984) Localization of NORs in karyotypes of four Rana species. Genetica, 64, 149-154. Bloom, S. E. and Goodpasture, C. (1976) An improved technique for selective silver staining of nucleolar organiser regions in human chromosomes. Human Genetics, 34, 199-206. Bogart, J.P. (1974) A karyosystematic study of frogs in the genus Leptodactylus (Anura: Leptodacylidae). Copeia, 1974, 728-737. Bogart, J. P. (1981) How many times has terrestrial breeding evolved in Anuran amphibians? Monitore zoologico Italiano NS Supplemento, 15, 29-40. Bogart, J. P. and Tandy, M. (1981) Chromosome lineages in African ranoid frogs. Monitore zoologico italiano NS Supplemento, xv, 55-91. Boulenger, G. A. (1882) Catalogue of the Batrachia Salientia S. Ecaudata in the collection of the British Museum. London.

230 Boulenger, G. A. (1884) Diagnoses of new reptiles and batrachians from the Solomon Islands collected and presented to the British Museum by H.B. Guppy Esq., M.B., HMS 'Lark'. Proceedings of the Zoological Society of London, 1884, 210-213. Boulenger, G. A. (1886a) III. On the reptiles and batrachians of the Solomon Islands. Transactions of the Zoological Society of London, 12(1), 35-62. Boulenger, G. A. (1886b) Remarks in connexion with the preceding note. Annals and Magazine of Natural History, 17, 463-464. Boulenger, G. A. (1887) Second contribution to the herpetology of the Solomon Islands. Proceedings of the Zoological Society of London, 23, 333-338. Boulenger, G. A. (1888) Third contribution to the herpetology of the Solomon Islands. Proceedings of the Zoological Society of London, 1888, 88-90. Boulenger, G. A. (1890) Fourth contribution to the herpetology of the Solomon Islands. Proceedings of the Zoological Society of London, 1890, 30-31. Boulenger, G. A. (1918a) On the Papuan, Melanesian and North-Australian species of the genus Rana. Annals and Magazine of Natural History, 1, 236-242. Boulenger, G. A. (1918b) Remarks on the Batrachian genera Cornufer , Tschudi, Platymantis , Gthr., Simomantis, g.n., and Staurois, Cope. Annals and Magazine of Natural History, 1, 372-375. Boulenger, G. A. (1920) A monograph of the South Asian, Papuan, Melanesian and Australian frogs of the genus Rana. Records of the Indian Museum, 20, 1-226. Bremer, K. (1988) The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution, 42, 795-803. Bremer, K. (1994) Branch support and tree stability. Cladistics, 10, 295-304. Brown, W. C. (1949) A new frog of the genus Platymantis from the Solomon Islands. American Museum Novitates, 1387, 1-4. Brown, W. C. (1952) The amphibians of the Solomon Islands. Bulletin of the Museum of Comparative Zoology, 107, 1-81. Brown, W. C. (1965) New frogs of the genus Cornufer (Ranidae) from the Solomon Islands. Breviora, 218, 1-16. Brown, W. C. (1996) Biogeography of amphibians in the islands on the southwest Pacific. Proceedings of the California Academy of Sciences, 50(2), 21-38.

231 Brown, W. C. and Menzies, J. I. (1978) A new Platymantis (Amphibia: Ranidae) from New Ireland, with notes on the amphibians of the Bismark Archipelago. Proceedings of the Biological Society of Washington, 91, 965- 971. Brown, W. C. and Myers, G. S. (1949a) A new frog of the genus Cornufer from the Solomon Islands, with notes on the endemic nature of the Fijian frog fauna. American Museum Novitates, 1418, 1-10. Brown, W. C. and Myers, G. S. (1949b) A new frog of the genus Batrachylodes from the Solomon Islands. Journal of the Washington Academy of Sciences, 39, 379-380. Brown, W. C. and Parker, F. (1970) New frogs of the genus Batrachylodes (Ranidae) from the Solomon Islands. Breviora, 346, 1-31. Brown, W. C. and Tyler, M. J. (1968) Frogs of the genus Platymantis (Ranidae) from New Britain with descriptions of new species. Proceedings of the Biological Society of Washington, 81, 69-86. Brown, W. C. and Webster, T. P. (1969) A new frog of the genus Discodeles Ranidae from Guadalcanal Island. Breviora, 338, 1-9. Burt, C. E. and Burt, M. D. (1932) Herpetological results of the Whitney South Sea Expedition. VI. Pacific Island amphibians and reptiles in the collection of the American Natural History. Bulletin of the American Natural History Museum, 63, 461-597. Carpenter, J. M. (1988) Choosing among multiple equally parsimonious cladograms. Cladistics, 4, 291-296. Carpenter, J. M. (1994) Successive weighting, reliability and evidence. Cladistics, 10, 215-220. Cole, C. J. (1974) Chromosome evolution in selected treefrogs, including casque- headed species (Pternohyla, Triprion, Hyla and Smilsca). American Museum Novitates, 2541, 1-10. Cope, E. D. (1865) Sketch of the primary groups of Batrachia Salientia. Natural History Review, 5, 97-120. Davis, D. D. and Gore, V. R. (1947) Clearing and staining skeletons of small vertebrates. Fieldiana: Technique, 4, 3-16. David, T. W. E. (1950) The Geology of the Commonwealth of Australia, vol. 1. Edward Arnold and Co., London [cited not seen].

232 Deckert, K. (1938) Beiträge zur osteologie und systematik ranider froschlurche. Sitzungsberichte Gesselschaft Naturforschenden Freunde, Berlin, 1938, 127-184. [cited not seen] de Weese, J. (1975) Chromosomes in Eleutherodactylus (Anura, Leptodactylidae). Mammalian Chromosome Newsletter, Houston, 16, 121-123. Dingerkus, G. and Uhler, L. D. (1977) Enzyme clearing of alcian blue stained whole small vertebrates for demonstration of cartilage. Stain Technology, 52, 229-232. Dubois, A. (1975) Un nouveau sous-genre (Paa) et trois nouvelles espèces du genre Rana. Remarques sur la phylogénie des Ranidés (Amphibiens, Anoures). Bulletin du Museum National d'Histoire Naturelle, Paris, 3, 1093- 1115. Dubois, A. (1981) Liste des genres et sous-genres nominaux de Ranoidea (Amphibiens Anoures) du monde, avec identification de leurs espèces- types: conséquences nomenclaturales. Monitore zoologico Italiano NS Supplemento, 15, 225-284. Dubois, A. (1983) Classification et nomenclature supragenerique des amphibiens anoures. Bulletin Mensuel de la Societe Linnenne de Lyon, 52, 270-276. Dubois, A. (1984) La nomenclature supragénérique des amphibiens anoures. Mémoires du Museum National d'Histoire Naturelle, Paris, 131, 1-56. Dubois, A. (1986) Miscellanea taxinomica batrachologica (I). Alytes, 5, 7-95. Dubois, A. (1988) Some comments of the genus concept in zoology. Monitore zoologico Italiano Supplemento, 22, 27-44. Dubois, A. (1992) Notes sur la classification des Ranidae (Amphibiens Anoures). Bulletin Mensuel de la Société Linnéenne de Lyon, 61, 305-352. Duellman, W. E. (1985) Reproductive modes in anuran amphibians: phylogenetic significance of adaptive strategies. South African Journal of Science, 81, 174-178. Dumeril, A. (1853) Mémoire sur les Batraciens anoures, de la famille des Hylæformes ou Rainettes, comprenant la description d'un genre nouveau et onze espèces nouvelles. Annales des Sciences Naturelles Paris, (3)19, 135-179. Dunteman, G.H. (1989) Principal components analysis. Sage Publications, Newbury Park.

233 Ecker, A. (1889) The Anatomy of the Frog. Clarendon Press, Oxford. Elspeth, M. J., Harrison, C. J., Allen, T. D. and Harris, R. (1985) The structural basis for C-banding: a scanning electron microscopy study. Chromosoma, 91, 363-368. Farris, J. S. (1969) A successive approximations approach to character weighting. Systematic Zoology, 18, 374-385. Farris, J. S. (1989) The retention index and the rescaled consistency index. Cladistics, 5, 417-419. Farris, J. S. (2001) Support weighting. Cladistics, 17(4), 389-394. Frost, D. R. (Ed.) (1985) Amphibian species of the world: a taxonomic and geographic reference. Association of Systematics Collections, Lawrence, Kansas, Allen Press. Frost, D. R. (2002) Amphibian species of the world: an online reference. V2.21 (15 July 2002). Electronic database available at http://research.amnh.org/herpetology/amphibia/index.html Frost, D. R., Wozencraft, W. C. and Hoffman, R. S. (1991) Phylogenetic relationships of hedgehogs and gymnures (Mammalia: Insectivora: Erinaceidae). Smithsonian Contributions to Zoology, 518, 1-69. Goloboff, P. A. (1999) Analyzing large data sets in reasonable times: Solutions for composite optima. Cladistics-The International Journal of the Willi Hennig Society, 15, 415-428. Green, D. M. (1983) Evidence for chromosome number reduction and chromosomal homosequentiality in the 24-chromosome Korean frog Rana dybowskii and related species. Chromosoma, 88, 222-226. Green, D. M. (1985) Differentiation in amount of centromeric heterochromatin between subspecies of the red-legged frog, Rana aurora. Copeia, 1985, 1071-1074. Green, D. M. (1986) Systematics and evolution of western North American frogs allied to Rana aurora and Rana boylii: Karyological evidence. Systematic Zoology, 35, 273-282. Green, D. M. and Borkin, L. J. (1993) Evolutionary relationships of Eastern Paleartic brown frogs, genus Rana: paraphyly of the 24-chromosome species group and the significance of chromosome number change. Zoological Journal of the Linnean Society, 109, 1-25.

234 Green, D. M. and Delisle, D. M. (1985) Allotriploidy in natural hybrid frogs, Rana chiricahuensis x R. pipiens, from Arizona: chromosomes and electrophoretic evidence. Journal of Herpetology, 19, 385-390. Green, D. M., Wasserman, A. O. and Bogart, J. P. (1981) Karyotypes of the frogs, Rana septentrionalis and R. virgatipes. Copeia, 1981, 879-882. Greenslade, P. J. M. (1969) Land fauna: Insect distribution patterns in the Solomon Islands. Philosophical Transactions of the Royal Society B, 255, 271-284. Günther, G. C. L. G. (1859) Catalogue of the Batrachia, Salientia in the collection of the British Museum, Taylor and Francis, London. Gustafsson, M. and Bremer, K. (1995) Morphology and phylogenetic interrelationships of the Asteraceae, Calyceraceae, Campanulaceae, Goodeniaceae, and related families (Asterales). American Journal of Botany, 82(2), 250-265. Haertel, J. D., Owczarzak, A. and Storm, R. M. (1974) A comparative study of the chromosomes from five species of the genus Rana (Amphibia: Salientia). Copeia, 1974, 109-114. Harrison, C. J., Britch, M., Allen, T. D. and Harris, R. (1981) Scanning electron microscopy of human metaphase chromosomes. Journal of Cell Science, 56, 409-422. Hediger, H. (1934) Beitrag zur Herpetologie und Zoogeographie Neu Britanniens und einiger umliegender Gebiete. Zoologische Jarbuecher, 65, 441-582. Hillis, D. M. (1991) The phylogeny of amphibians: Current knowledge and the role of cytogenetics. In Amphibian Cytogenetics and Evolution(Eds, Green, D. M. and Sessions, S. K.) Academic Press, San Diego, pp. 7-31. Holmquist, G. (1979) The mechanism of C-banding: depurination and B- elimination. Chromosoma, 72, 203-224. Inger, R. F. (1954) Systematics and zoogeography of Philippine Amphibia. Fieldiana: Zoology, 33, 181-531. Jolliffe, I. T. (1986) Principal Components Analysis. Springer-Verlag, New York. Kallersjo, M., Farris, J. S., Kluge, A. G. and Bult, C. (1992) Skewness and permutation. Cladistics, 8, 275-287.

235 King, M. (1980) C-banding studies on Australian hylid frogs : Secondary constriction structure and the concept of euchromatin transformation. Chromosoma, 80, 191-217. King, M. (1981) A cytotaxonomic analysis of Australian hylid frogs of the genus Litoria. In Proceedings of the Melbourne Herpetological Symposium (Eds Bansk, C. B. and Martin, A. A.), Melbourne. King, M. (1990) Amphibia. Animal Cytogenetics, 4, 1-242. Kinghorn, J. R. (1928) Herpetology of the Solomon Islands. Records of the Australian Museum, 16, 123-178. Kluge, A. G. and Wolf, A. J. (1993) Cladistic: What's in a word? Cladistics, 9, 183- 199. Kuramoto, M. (1972) Karyotypes of six species of frog (genus Rana) endemic to the Ryukyu Islands. Caryologia, 25, 547-549. Kuramoto, M. (1980) Karyotypes of several frogs from Korea, Taiwan and the Philippines. Experientia, 36, 826-828. Kuramoto, M. (1985) Karyological divergence in three platymantine frogs, Family Ranidae. Amphibia-Reptilia, 6, 355-361. Kuramoto, M. (1990) A list of chromosome numbers of anuran amphibians. Bulletin of Fukuoka University of Education, 39, 83-127. Laurent, R. F. (1943) Sur l'ostéologie de deux Ranides exotiques. Bulletin du Musée Royal d'Histoire Naturelle de Belgique, 19, 1-4. Laurent, R.F. (1951) Sur la nécessité de supprimer la famille des Rhacophoridae, mais de créer celle des Hyperoliidae. Revue de Zoologie et Botanique de Africaines, 45, 116-122. Laurent, R. F. (1981) Systématique et répartition géographique. In: P.-P. Grassé, Ed. Traité de Zoologie. Tome XIV, Amphibiens, Fasc. A. Pris: Masson. Laurent, R. F. (1986) Sous-classe des Lissamphibiens (Lissamphibia). Systématique. In Traité de Zoologie, Vol. 14 Batraciens, Fasc 1-B (Eds, Grassé, P. P. and Delsol, M.) Paris, pp. 594-797. Levan, A., Fredga, K. and Sandberg, A. A. (1964) Nomenclature for centromeric position of chromosomes. Hereditas, 52, 201-220. Loveridge, A. (1948) New Guinean reptiles and amphibians in the Museum of Comparative Zoology and United States National Museum. Bulletin of the Museum of Comparative Zoology, 101, 305-430.

236 Mabee, P. M. (1989) Assumptions underlying the use of ontogenetic sequences for determining character state order. Transactions of the American Fisheries Society, 118, 151-158. Maddison, W. P. (1993) Missing data versus missing characters in phylogenetic analysis. Systematic Biology, 42, 576-581. Mahony, M. J., Norris, R. M. and Donnellan, S. C. (1996) Karyotypes of south- west Pacific ranid frogs (Anura: Ranidae). Australian Journal of Zoology, 44, 119-128. Mahony, M. J. and Robinson, E. S. (1980) Polyploidy in the Australian leptodactylid frog genus Neobatrachus. Chromosoma, 81, 199-212. Matsui, M. (1986) Three new species of Amolops from Borneo (Amphibia, Anura, Ranidae). Copeia, 1986, 623-630. McKay, R. G. (1973) The mechanism of G and C-banding in mammalian metaphase chromosomes. Chromosoma (Berl), 44, 1-14. Menzies, J. I. (1982a) Systematics of Platymantis papuensis (Amphibia: Ranidae) and related species of the New Guinea region. British Journal of Herpetology, 6, 236-240. Menzies, J. I. (1982b) The voices of some male Platymantis species of the New Guinean region. British Journal of Herpetology, 6, 241-245. Menzies, J. I. (1987) A taxonomic revision of Papuan Rana (Amphibia: Ranidae). Australian Journal of Zoology, 35, 373-418. Menzies, J. I. and Tippet, J. (1976) Chromosome numbers of Papuan hylid frogs and the karyotype of Litoria infrafrenata (Amphibia, Anura, Hylidae). Journal of Herpetology, 10, 167-173. Mertens, R. (1929) Zur synonymie der Froschgattung Batrachylodes Boulenger. Zoologischer Anzeiger, 80, 266-268. Miklos, G. L. G. and John, B. (1983) Heterochromatin and satellite DNA in man: properties and prospects. American Journal of Human Genetics, 31, 264- 280. Morescalchi, A. (1973) Amphibia. In Cytotaxonomy and Vertebrate Evolution (Ed, Chiarelli, A. B. and Capanna, E.) Academic Press, London, pp. 233-248. Morescalchi, A. (1979) New developments in vertebrate taxonomy. I. Cytotaxonomy of the amphibians. Genetics, 50(3), 179-193.

237 Morescalchi, A. (1981) Karyology of the main groups of African frogs. Monitore zoologico Italiano NS Supplemento, 15, 41-53. Myers, G. S. (1954) Ability of amphibians to cross sea barriers with especial reference to Pacific zooogeography. Proceedings of the 7th Pacific Science Congress, 4, 19-27. Nishioka, M., Okumoto, H. and Ryuzaki, M. (1987a) A comparative study on the karyotypes of pond frogs distributed in Japan, Korea, Taiwan, Europe and North America. Scientific Report from the Laboratory of Amphibian Biology, Hiroshima University, 9, 135-163. Nishioka, M., Okumoto, H., Ueda, H. and Ryuzaki, M. (1987b) Karyotypes of brown frogs distributed in Japan, Korea, Europe and North America. Scientific Report from the Laboratory of Amphibian Biology, Hiroshima University, 9, 165-212. Nixon, K. C. (1999) The Parsimony Ratchet, a new method for rapid parsimony analysis. Cladistics, 15, 407-414. Nixon, K. C. and Davis, J. I. (1991) Polymorphic taxa, missing values and cladistic analysis. Cladistics, 7(3), 233-241. Noble, G. K. (1925) An outline of the relation of ontogeny to phylogeny within the Amphibia I. American Museum Novitates, 165, 1-17. Noble, G. K. (1927) The value of life history data in the study of the evolution of the Amphibia. Annals of the New York Academy of Science, 30, 31-128. Noble, G. K. (1931) The Biology of the Amphibia. McGraw-Hill Book Company, Inc., New York. Ogilby, J. D. (1890) Report on a zoological collection from the Solomon Islands Part II. Records of the Australian Museum, 1, 5-7. Ohler, A. and Dubois, A. (1989) Démonstration de l'origine indépendante des ventouses digitales dans deux lignées phylogénétiques de Ranidae (Amphibiens, Anoures). Comptes rendus de l'Academie des sciences. Serie III, Sciences de la vie, 309, 419-422. Ota, H. and Matsui, M. (1995) Karyotype of a ranid frog, Platymantis pelewensis, from Belau, Micronesia, with comments on its systematic implications. Pacific Science, 49, 296-300. Pardue, M. J. and Gall, J. G. (1970) Chromosomal localization of mouse satellite DNA. Science, 168, 1356-1358.

238 Platnick, N. I., Griswold, C. E. and Coddington, J. A. (1991) On missing entries in cladistic analysis. Cladistics, 7, 337-344. Roux, J. (1905) La famille des Ranidae. Zoologischer Anzeiger, 28, 777-785. Roux, J. (1918) Note sur quelques espèces d'Amphibiens de l'Archipel Indo- Australien. Revue Suisse de Zoologie, 26, 409-415. Savage, J. M. (1973) The geographic distribution of frogs: patterns and predictions. In Evolutionary Biology of the Anurans: Contemporary Research on Major Problems(Ed, Vial, J. D.) University of Missouri, Columbia, pp. 351-445. Schmid, M. (1980) Chromosome banding in Amphibia IV. Differentiation of GC- and AT-rich chromosome regions in Anura. Chromosoma, 77, 83-103. Schmid, M. (1982) Chromosome banding in Amphibia VII. Analysis of the structure and variability of NORs in Anura. Chromosoma, 87, 327-344. Schmidt, A. A. (1992) Breeding of the Solomon Island frog, Ceratobatrachus guentheri Boulenger, 1884. Salamandra, 1, 3-8. Schmidt, K. P. (1932) Reptiles and amphibians from the Solomon Islands. Publication of the Field Museum of Natural History, Zoology Series, 18, 175-190. Seto, T. (1965) Cytogenetic studies in lower vertebrates, II Karyological studies of several species of frogs (Ranidae). Cytologia, 30, 437-446. SPSS (1993) SPSS Professional Statistics 6.1. SPSS Inc, Chicago, Il. Sternfeld, R. (1918) Zur Tiergeographie Papuasiens und der pazifischen Inselwelt. Abhandelungen Senckenbergia Naturforsch Gessel, 36, 375-436. Swofford, D. L. and Olsen, G. J. (1990) Phylogeny reconstruction. In Molecular Systematics (Eds, Hillis, D.M and Moritz, C.) Sinauer Associates, Inc., Sunderland, Massachusetts, pp. 411-501. Tanner, V. M. (1951) Pacific islands herpetology No V. Guadalcanal, Solomon Islands: A check list of species. The Great Basin Naturalist, 11, 53-86. Thompson, R. B. and Hackman, B.D. (1969) Some geological notes on areas visited by the Royal Society Expedition to the British Solomon Islands 1965. Philosophical Transactions of the Royal Society B, 255, 189-202. Trueb, L. (1966) Morphology and development of the skull in the frog, Hyla septentrionalis. Copeia, 3, 562-573.

239 Trueb, L. (1973) Bones, frogs and evolution. In Evolutionary Biology of the Anurans: Contemporary Research on Major Problems Evolutionary Biology of the Amphibians (Ed, Vial, J. D.) University of Missouri, Columbia, pp. 65- 132. Trueb, L. (1979) Leptodactylid frogs of the genus Telmatobius in Ecuador with a description of a new species. Copeia, 1979, 714-733. Tschudi, J. J. (1838) Classification der Batrachia. Neuchatel. Tyler, M. J. (1972) An analysis of the lower vertebrate faunal relationships of Australia and New Guinea. In Bridge and Barrier: The Natural and Cultural History of Torres Strait., Vol. Publ. BG/3 (Ed, Walker, D.) Department of Biogeography and Geomorphology, Canberra, pp. 231-256. Tyler, M. J. (1976) Comparative osteology of the pelvic girdle of Australian frogs and description of a new fossil genus. Transactions of the Royal Society of South Australia, 100, 3-14. Tyler, M. J. (1979) Herpetofaunal relationships of South America with Australia. In The South American Herpetofauna: its origins, evolution and dispersal., Vol. 7 (Ed, Duellman, W. E.) University of Kansas, Lawrence, Kansas, pp. 1- 485. Van Kampen, P. N. (1919) Die amphibienfauna von Neu-Guinea; Fest-nummer. Bijdragen Tot De Dierkunde, Koninklijk Zoölogisch Genootschap "Naturia Artis Magistra" (21), 51 [cited not seen]. Van Kampen, P. N. (1923) The Amphibia of the Indo-Australian Archipelago. E.J. Brill Ltd., Leiden, Holland. Vogt, T. (1912) Beitrag zur reptilien- und amphibienfauna der Südseeinseln. Sonder-Abdruck aus den Sitzungsberichte Gessellschaft naturforschenden Freunde zu Berlin, 1912, 1-13. Vosa, C. G. (1973) Heterochromatin and chromosome structure. In Chromosome Identification - Techniques and Applications in Biology and Medicine.(Eds Caspersson, T. and Zeck, L.) Academic Press, New York, pp. 152-155. Werner, F. (1894) Über einige Novitäten der herpetologischen Sammlung des Wiener zoolog. vergl. anatom. Instituts. Zoologischer Anzeiger, 17, 155- 159. Werner, F. (1900) Die reptilien- und Batrachierfauna des Bismark Archipels. Mitteilungen Zoologische Museum, Berlin, 1, 8-11, 111-117.

240 Wilkinson, M. (1992) Ordered versus unordered characters. Cladistics, 8, 375-385. Zweifel, R. G. (1960) Results of the 1958-1959 Gilliard New Britain Expedition. 3. Notes on the frogs of New Britain. American Museum Novitates, 2023, 1- 27. Zweifel, R. G. (1966) Cornufer unicolor Tschudi 1838 (Amphibia Salientia); request for suppression under the plenary powers Z.N.(S.) 1749. Bulletin of Zoological Nomenclature, 23, 167-168. Zweifel, R. G. (1967) Identity of the frog Cornufer unicolor and application of the generic name Cornufer. Copeia, 1967, 117-121. Zweifel, R. G. (1975) Two new frogs of the genus Platymantis (Ranidae) from New Britain. American Museum Novitates, 2582, 1-7. Zweifel, R. G. and Tyler, M. J. (Eds.) (1982) Amphibia of New Guinea. Dr W. Junk Publishers, The Hague.

241 APPENDIX 1. Specimen List

Genus Species Registration Number Location Date Collected Sex Character Dataset

Batrachylodes elegans AMNH71778 "Kunua, montane region, Bougainville" M "m, p" Batrachylodes elegans AMNH79540 "Bougainville Id., Mutahi" F "m, p" Batrachylodes elegans AMNH79541 "Bougainville Id., Mutahi" M "m, p" Batrachylodes elegans AMNH79542 "Bougainville Id., Mutahi" M "m, p" Batrachylodes elegans AMNH79543 "Bougainville Id., Mutahi" F "m, p" Batrachylodes elegans CAS117444 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 F "m, p" 3600 ft " Batrachylodes elegans CAS117445 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 F "m, p" 3600 ft " Batrachylodes elegans CAS117446 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 F "m, p" 3600 ft " Batrachylodes elegans CAS117447 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 M "m, p" 3600 ft " Batrachylodes elegans CAS117448 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 F "m, p" 3600 ft " Batrachylodes elegans CAS117449 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 F "m, p" 3600 ft " Batrachylodes elegans CAS117450 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 M "m, p" 3600 ft " Batrachylodes elegans CAS117451 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 M "m, p" 3600 ft " Batrachylodes elegans CAS117452 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 . o 3600 ft " Batrachylodes elegans CAS117453 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 M "m, p" 3600 ft " Batrachylodes elegans CAS117454 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 F "m, p" 3600 ft " Batrachylodes elegans CAS117455 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 F "m, p" 3600 ft " Batrachylodes elegans CAS117456 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 F "m, p" 3600 ft " Batrachylodes elegans CAS117457 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 S 3600 ft " Batrachylodes elegans CAS117458 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 F "m, p" 3600 ft " Batrachylodes elegans CAS117459 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 M "m, p" 3600 ft " Batrachylodes elegans CAS117460 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 M "m, p" 3600 ft " Batrachylodes elegans CAS117462 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 F "m, p" 3600 ft "

242 Batrachylodes elegans CAS117463 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 M "m, p" 3600 ft " Batrachylodes elegans CAS117464 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 F "m, p" 3600 ft " Batrachylodes elegans CAS117465 (paratype) "Solomon Islands: N end Bougainville Id. Mutahi, 3000- 1-May-66 S 3600 ft " Batrachylodes elegans SAMR8210A "Mutahi, Bougainville" 11-May-66 M "m, p, o" Batrachylodes elegans SAMR8210B "Mutahi, Bougainville" 11-May-66 M "m, p" Batrachylodes gigas CAS117442 "Solomon Islands: S end Bougainville Id. Lake Loloru area, 19-May-05 F ca. 4300 ft" Batrachylodes mediodiscus AMNH79524 "Bougainville Is., Mutahi" M "m, p" Batrachylodes mediodiscus AMNH79525 "Bougainville Is., Mutahi, 3000-3600'" M "m, p" Batrachylodes mediodiscus AMNH79526 "Bougainville Is., Mutahi, 3000-3600'" M "m, p" Batrachylodes mediodiscus AMNH79527 "Bougainville Is., Mutahi, 2700'" M "m, p" Batrachylodes mediodiscus CAS101780 (paratype) "Solomon Islands, Bougainville Id., Kunua" 22-Jan-63 M "m, p" Batrachylodes mediodiscus CAS21811 "Solomon Islands, Bougainville Id., Kunua" F "m, p" Batrachylodes mediodiscus CAS21812 (paratype) "Solomon Islands, Bougainville Id., Aresi area" 1-May-63 M "m, p" Batrachylodes mediodiscus CAS21813 "Solomon Islands, Bougainville Id., Aresi area" 1-May-63 M "m, p" Batrachylodes mediodiscus CAS21815 (paratype) "Solomon Islands, Bougainville Id., Pipekei" 1-Dec-62 M "m, p" Batrachylodes mediodiscus SAMR8204A "Mutahi, Bougainville" 10-May-66 M "m, p, o" Batrachylodes mediodiscus SAMR8204B "Mutahi, Bougainville" 10-May-66 M "m, p, o" Batrachylodes mediodiscus SAMR8205 "Mutahi, Bougainville" 10-May-66 M "m, p" Batrachylodes mediodiscus SAMR8259A "Mutahi, Bougainville" 9-May-66 M "m, p" Batrachylodes mediodiscus SAMR8259B "Mutahi, Bougainville" 9-May-66 F "m, p" Batrachylodes minutus AMNH35425 Choiseul Island F m Batrachylodes minutus AMNH79258 "Bougainville, Mutahi, Solomon Islands" F m Batrachylodes minutus CAS101781 (paratype) "Solomon Islands, Bougainville Id., Aresi" 14-Apr-63 F m Batrachylodes minutus CAS21852 "Solomon Islands, Bougainville Id., Pipekei" 15-May-05 F m Batrachylodes minutus CAS23989 (paratype) "Solomon Islands, Bougainville Id., S end Ororoi area, 2000 1-Jun-66 M m ft" Batrachylodes minutus CAS23990 "Solomon Islands, Bougainville Id., Mutahi area, 2700-5000 10-May-66 F m ft" Batrachylodes minutus CAS23991 (paratype) "Solomon Islands, Bougainville Id., Mutahi area, 2200-3200 17-May-66 M m ft" Batrachylodes minutus CAS23992 (paratype) "Solomon Islands, Bougainville Id., Mutahi area, 2200-2300 17-May-66 M m ft" Batrachylodes montanus CAS117443 (paratype) "Solomon Islands: N end Bougainville Id. Melilup area, 1-May-66 M 3500-4000 ft" Batrachylodes trossulus A27/SAMR47121 "Pavora River, Choiseul" 1-Mar-90 M "m, p" Batrachylodes trossulus A43/SAMR47122 "Pavora River, Choiseul" 1-Mar-90 F "m, p, o" Batrachylodes trossulus AMNH69270 "Bougainville Is., Buka Island" F "m, p" Batrachylodes trossulus AMNH70021 "Bougainville Is., Kunua" M "m, p" Batrachylodes trossulus AMNH71673 "Bougainville Is., Kunua" M "m, p" Batrachylodes trossulus AMNH71674 "Bougainville Is., Kunua" F "m, p" Batrachylodes trossulus AMNH71675 "Bougainville Is., Kunua" M "m, p" Batrachylodes trossulus AMNH71676 "Bougainville Is., Kunua" M "m, p" Batrachylodes trossulus AMNH71677 "Bougainville Is., Kunua" M "m, p"

243 Batrachylodes trossulus AMNH79529 "Bougainville Is., Mutahi" F "m, p" Batrachylodes trossulus AMNH79530 "Bougainville Is., Mutahi" F "m, p" Batrachylodes trossulus AMNH79532 "Bougainville Is., Mutahi" F "m, p" Batrachylodes trossulus K771/SAMR47120 "Torakina, Bougainville" 1987 M "m, p" Batrachylodes trossulus SAMR5163 "Kunua, Bougainville" 28-Jun-63 F "m, p" Batrachylodes trossulus SAMR5167A "Kunua, Bougainville" 12-Jul-63 F "m, p" Batrachylodes trossulus SAMR5167B "Kunua, Bougainville" 12-Jul-63 F "m, p" Batrachylodes trossulus SAMR5167C "Kunua, Bougainville" 12-Jul-63 M "m, p, o" Batrachylodes trossulus SAMR8209A "Mutahi, Bougainville" 11-May-66 F "m, p, o" Batrachylodes trossulus SAMR8209B "Mutahi, Bougainville" 11-May-66 F "m, p" Batrachylodes trossulus SAMR8209C "Mutahi, Bougainville" 11-May-66 F "m, p" Batrachylodes trossulus SAMR8209D "Mutahi, Bougainville" 11-May-66 F "m, p, o" Batrachylodes vertebralis CAS117473 "Solomon Ids., N end of Bougainville, Mutahi, 3000-3600 ft" 1-May-66 M o Batrachylodes vertebralis A22/SAMR47108 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p, o" Batrachylodes vertebralis A23/SAMR47109 "Pavora River, Choiseul" 6/3/90 - 12/3/90 M "m, p" Batrachylodes vertebralis A24/SAMR47110 "Pavora River, Choiseul" 6/3/90 - 12/3/90 M "m, p, o" Batrachylodes vertebralis A25/SAMR47111 "Pavora River, Choiseul" 6/3/90 - 12/3/90 M "m, p" Batrachylodes vertebralis A26/SAMR47112 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Batrachylodes vertebralis A28/SAMR47113 "Pavora River, Choiseul" 6/3/90 - 12/3/90 S Batrachylodes vertebralis A42/SAMR47114 "Pavora River, Choiseul" 6/3/90 - 12/3/90 M "m, p" Batrachylodes vertebralis A60/SAMR37000 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "c, m, p" Batrachylodes vertebralis A67/SAMR47115 "Pavora River, Choiseul" 6/3/90 - 12/3/90 S Batrachylodes vertebralis A82/SAMR37002 "Pavora River, Choiseul" 6/3/90 - 12/3/90 M "c, m, p" Batrachylodes vertebralis A83/SAMR37003 "Pavora River, Choiseul" 6/3/90 - 12/3/90 M "c, m, p" Batrachylodes vertebralis B04/SAMR37001 "Pavora River, Choiseul" 6/3/90 - 12/3/90 M "c, m, p" Batrachylodes vertebralis B92/SAMR47116 "Mt Austen, Guadalcanal" 13-Mar-90 F "m, p" Batrachylodes vertebralis B93/SAMR47117 "Mt Austen, Guadalcanal" 13-Mar-90 M "m, p" Batrachylodes vertebralis B94/SAMR47118 "Mt Austen, Guadalcanal" 13-Mar-90 M "m, p" Batrachylodes vertebralis B95/SAMR47119 "Mt Austen, Guadalcanal" 13-Mar-90 M "m, p" Batrachylodes vertebralis MCZ41832 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41833 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41834 "Popoheirarai, Bougainville" 17-May-66 M "m, p, o" Batrachylodes vertebralis MCZ41835 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41836 "Popoheirarai, Bougainville" 17-May-66 M "m, p, o" Batrachylodes vertebralis MCZ41837 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41838 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41839 "Popoheirarai, Bougainville" 17-May-66 M "m, p, o" Batrachylodes vertebralis MCZ41840 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41841 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41842 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41843 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41844 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41845 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41846 "Popoheirarai, Bougainville" 17-May-66 M "m, p, o" Batrachylodes vertebralis MCZ41847 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41848 "Popoheirarai, Bougainville" 17-May-66 F "m, p, o" Batrachylodes vertebralis MCZ41849 "Popoheirarai, Bougainville" 17-May-66 M "m, p"

244 Batrachylodes vertebralis MCZ41850 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41851 "Popoheirarai, Bougainville" 17-May-66 F "m, p" Batrachylodes vertebralis MCZ41852 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41853 "Popoheirarai, Bougainville" 17-May-66 F "m, p" Batrachylodes vertebralis MCZ41854 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41855 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41856 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis MCZ41857 "Popoheirarai, Bougainville" 17-May-66 F "m, p" Batrachylodes vertebralis MCZ41858 "Popoheirarai, Bougainville" 17-May-66 M "m, p" Batrachylodes vertebralis SAMR4243 "Kunua, Bougainville" 1-Mar-90 M "m, p, o" Batrachylodes vertebralis SAMR4422 "Kunua, Bougainville" 2-Jul-62 F "m, p" Batrachylodes vertebralis SAMR4423 "Kunua, Bougainville" 2-Jul-62 F "m, p" Batrachylodes vertebralis SAMR4767 "Kunua, Bougainville" - F "m, p" Batrachylodes vertebralis SAMR4924A "Kunua, Bougainville" jan-feb 63 F "m, p" Batrachylodes vertebralis SAMR4924B "Kunua, Bougainville" jan-feb 63 F "m, p" Batrachylodes vertebralis SAMR4924C "Kunua, Bougainville" jan-feb 63 M "m, p" Batrachylodes vertebralis SAMR4924D "Kunua, Bougainville" jan-feb 63 M "m, p" Batrachylodes vertebralis SAMR4924E "Kunua, Bougainville" jan-feb 63 F "m, p" Batrachylodes vertebralis SAMR4924F "Kunua, Bougainville" jan-feb 63 M "m, p" Batrachylodes vertebralis SAMR5162A "Kunua, Bougainville" 28-Jun-63 M "m, p" Batrachylodes vertebralis SAMR5162B "Kunua, Bougainville" 28-Jun-63 M "m, p, o" Batrachylodes vertebralis SAMR5162C "Kunua, Bougainville" 28-Jun-63 M "m, p" Batrachylodes vertebralis SAMR5162D "Kunua, Bougainville" 28-Jun-63 M "m, p" Batrachylodes vertebralis SAMR5162E "Kunua, Bougainville" 28-Jun-63 M "m, p" Batrachylodes vertebralis SAMR5162F "Kunua, Bougainville" 28-Jun-63 M "m, p, o" Batrachylodes vertebralis SAMR5162G "Kunua, Bougainville" 28-Jun-63 M "m, p" Batrachylodes vertebralis SAMR5162H "Kunua, Bougainville" 28-Jun-63 M "m, p" Batrachylodes vertebralis SAMR5162I "Kunua, Bougainville" 28-Jun-63 F "m, p" Batrachylodes vertebralis SAMR5162J "Kunua, Bougainville" 28-Jun-63 M "m, p" Batrachylodes vertebralis SAMR5162K "Kunua, Bougainville" 28-Jun-63 M "m, p" Batrachylodes vertebralis SAMR5162L "Kunua, Bougainville" 28-Jun-63 M "m, p" Batrachylodes vertebralis SAMR8351 "Mutahi, Bougainville" 10-May-65 F "m, p" Batrachylodes vertebralis UM60303 "Highlands interior of Malaita, Solomon Islands" 1923-24 M "m, p" Batrachylodes wolfi AMNH71752 "Bougainville, Kunua" M "m, p" Batrachylodes wolfi AMNH71753 "Bougainville, Kunua" M "m, p" Batrachylodes wolfi AMNH71757 "Bougainville, Kunua" M "m, p" Batrachylodes wolfi AMNH71758 "Bougainville, Kunua" F "m, p" Batrachylodes wolfi AMNH71762 "Bougainville, Kunua" M "m, p" Batrachylodes wolfi AMNH71764 "Bougainville, Kunua" M "m, p" Batrachylodes wolfi AMNH71768 "Bougainville, Kunua" M "m, p" Batrachylodes wolfi AMNH71774 "Bougainville, Kunua" M "m, p" Batrachylodes wolfi AMNH71777 "Bougainville, Kunua" M "m, p" Batrachylodes wolfi SAMR4927A "Kunua, Bougainville" feb '63 M "m, p, o" Batrachylodes wolfi SAMR4927B "Kunua, Bougainville" feb '63 M "m, p" Batrachylodes wolfi SAMR4927C "Kunua, Bougainville" feb '63 M "m, p" Batrachylodes wolfi SAMR4927D "Kunua, Bougainville" feb '63 M "m, p" Batrachylodes wolfi SAMR4927E "Kunua, Bougainville" feb '63 F "m, p"

245 Batrachylodes wolfi SAMR5164A "Kunua, Bougainville" 28-Jun-63 M "m, p" Batrachylodes wolfi SAMR5164B "Kunua, Bougainville" 28-Jun-63 M "m, p, o" Batrachylodes wolfi SAMR5164C "Kunua, Bougainville" 28-Jun-63 M "m, p, o" Batrachylodes wolfi SAMR5164D "Kunua, Bougainville" 28-Jun-63 M "m, p" Batrachylodes wolfi SAMR8195A "Mutahi, Bougainville" 5-May-66 F "m, p" Batrachylodes wolfi SAMR8195B "Mutahi, Bougainville" 5-May-66 M "m, p" Batrachylodes wolfi SAMR8195C "Mutahi, Bougainville" 5-May-66 F "m, p, o" Batrachylodes wolfi SAMR8196A "Mutahi, Bougainville" 9-May-66 F "m, p" Batrachylodes wolfi SAMR8196B "Mutahi, Bougainville" 9-May-66 F "m, p" Batrachylodes wolfi SAMR8778A "Mutahi, Bougainville" 3-Mar-66 F "m, p" Batrachylodes wolfi SAMR8778B "Mutahi, Bougainville" 3-Mar-66 F "m, p" Batrachylodes wolfi SAMR8778C "Mutahi, Bougainville" 3-Mar-66 M "m, p" Ceratobatrachus guentheri A53/SAMR47123 "Pavora River, Choiseul" 6/3/90 - 23/3/90 F p Ceratobatrachus guentheri A61/SAMR47124 "Pavora River, Choiseul" 6/3/90 - 23/3/90 F p Ceratobatrachus guentheri A65/SAMR37004 "Pavora River, Choiseul" 6/3/90 - 23/3/90 F "c, p" Ceratobatrachus guentheri A66/SAMR37003 "Pavora River, Choiseul" 6/3/90 - 23/3/90 M p Ceratobatrachus guentheri A97/SAMR47125 "Pavora River, Choiseul" 6/3/90 - 23/3/90 M p Ceratobatrachus guentheri B02/R37005 "Pavora River, Choiseul" 6/3/90 - 23/3/90 F "c, p" Ceratobatrachus guentheri SAMR37006 "Pavora River, Choiseul" 6/3/90 - 23/3/90 F "c, p" Ceratobatrachus guentheri MCZ42113 "Kunua, Bougainville" . M p Ceratobatrachus guentheri MCZ42114 "Kunua, Bougainville" . S Ceratobatrachus guentheri MCZ42115 "Kunua, Bougainville" . M p Ceratobatrachus guentheri MCZ42116 "Kunua, Bougainville" . M p Ceratobatrachus guentheri MCZ42117 "Kunua, Bougainville" . S Ceratobatrachus guentheri MCZ42118 "Kunua, Bougainville" . S Ceratobatrachus guentheri MCZ42119 "Kunua, Bougainville" . S Ceratobatrachus guentheri MCZ42120 "Kunua, Bougainville" . S Ceratobatrachus guentheri MCZ42121 "Kunua, Bougainville" . S Ceratobatrachus guentheri MCZ62219 "Mutahi, Bougainville" 17-May-66 F p Ceratobatrachus guentheri MCZ62220 "Mutahi, Bougainville" 17-May-66 F p Ceratobatrachus guentheri MCZ62221 "Mutahi, Bougainville" 17-May-66 M p Ceratobatrachus guentheri MCZ62222 "Mutahi, Bougainville" 17-May-66 F "p, o" Ceratobatrachus guentheri MCZ62223 "Mutahi, Bougainville" 17-May-66 M p Ceratobatrachus guentheri MCZ62224 "Mutahi, Bougainville" 17-May-66 F p Ceratobatrachus guentheri MCZ62225 "Mutahi, Bougainville" 17-May-66 F "p, o" Ceratobatrachus guentheri MCZ62226 "Mutahi, Bougainville" 17-May-66 F "p, o" Ceratobatrachus guentheri MCZ62227 "Mutahi, Bougainville" 17-May-66 M p Ceratobatrachus guentheri MCZ62228 "Mutahi, Bougainville" 17-May-66 F p Ceratobatrachus guentheri MCZ62229 "Mutahi, Bougainville" 17-May-66 F p Ceratobatrachus guentheri MCZ62230 "Mutahi, Bougainville" 17-May-66 F p Ceratobatrachus guentheri SAMR4221 "Kunua, Bougainville" 13-Mar-62 M p Ceratobatrachus guentheri SAMR4407 "Kunua, Bougainville" 1-Jul-62 F p Ceratobatrachus guentheri SAMR4408 "Kunua, Bougainville" 1-Jul-62 F p Ceratobatrachus guentheri SAMR4409 "Kunua, Bougainville" 1-Jul-62 F p Ceratobatrachus guentheri SAMR8057 "Turiboiru, Bougainville" 14-Mar-62 F p Ceratobatrachus guentheri SAMR8058 "Turiboiru, Bougainville" 14-Mar-62 F p Ceratobatrachus guentheri SAMR8192 "Turiboiru, Bougainville" 9-Mar-62 F p

246 Ceratobatrachus guentheri SAMR8193 "Turiboiru, Bougainville" 3-Mar-62 M p Ceratobatrachus guentheri SAMR8194 "Turiboiru, Bougainville" 3-Mar-62 F p Ceratobatrachus guentheri SAMR8203A "Matsiogu, Bougainville" 30-Mar-66 F p Ceratobatrachus guentheri SAMR8203B "Matsiogu, Bougainville" 30-Mar-66 M p Ceratobatrachus guentheri SAMR8207A Malai/Shortlands 22-Apr-66 F p Ceratobatrachus guentheri SAMR8207B Malai/Shortlands 22-Apr-66 M p Ceratobatrachus guentheri SAMR8207C Malai/Shortlands 22-Apr-66 M p Ceratobatrachus guentheri SAMR8776 "Malai, New Guinea" 15-Apr-66 M p Discodeles bufoniformis CAS109895 "Solomon Ids., Bougainville Id, Matsioga 2100 ft" 24-Mar-66 o Discodeles bufoniformis A05/SAMR45938 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles bufoniformis A07/SAMR45940 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles bufoniformis A44/SAMR47100 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles bufoniformis A47/SAMR37007 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "c, m, p" Discodeles bufoniformis A48/SAMR37008 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "c, m, p" Discodeles bufoniformis A49/SAMR37009 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles bufoniformis A54/SAMR47102 "Pavora River, Choiseul" 6/3/90 - 12/3/90 M "m, p" Discodeles bufoniformis A62/SAMR45942 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles bufoniformis A72/SAMR47103 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles bufoniformis A86/SAMR47104 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles bufoniformis A87/SAMR47105 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles bufoniformis A88/SAMR47106 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles bufoniformis A89/SAMR47107 "Pavora River, Choiseul" 6/3/90 - 12/3/90 M "m, p" Discodeles bufoniformis AMNH35438 Vella Lavella F "m, p" Discodeles bufoniformis AMNH35441 Ronongo F "m, p" Discodeles bufoniformis AMNH35468 Ronongo F "m, p" Discodeles bufoniformis MCZ38551 "Kunua, Bougainville" 17-May-66 F "m, p, o" Discodeles bufoniformis MCZ38553 "Kunua, Bougainville" 1963 F "m, p" Discodeles bufoniformis MCZ38554 "Kunua, Bougainville" 1963 F "m, p" Discodeles bufoniformis MCZ38556 "Kunua, Bougainville" 17-May-66 F "m, p, o" Discodeles bufoniformis MCZ38557 "Kunua, Bougainville" 1963 F "m, p, o" Discodeles bufoniformis MCZ38561 "Kunua, Bougainville" 1963 F "m, p" Discodeles bufoniformis MCZ38562 "Kunua, Bougainville" 1963 F "m, p" Discodeles bufoniformis MCZ38565 "Kunua, Bougainville" 1963 M "m, p" Discodeles bufoniformis MCZ38567 "Kunua, Bougainville" 1963 F "m, p" Discodeles bufoniformis MCZ38568 "Kunua, Bougainville" 1963 M "m, p" Discodeles bufoniformis MCZ38569 "Kunua, Bougainville" 1963 F "m, p" Discodeles bufoniformis MCZ38570 "Kunua, Bougainville" 1963 M "m, p" Discodeles bufoniformis MCZ38578 "Kunua, Bougainville" 17-May-66 M "m, p, o" Discodeles bufoniformis MCZ38580 "Kunua, Bougainville" 1963 F "m, p" Discodeles bufoniformis MCZ38581 "Kunua, Bougainville" 1963 F "m, p" Discodeles bufoniformis MCZ38582 "Kunua, Bougainville" 1963 M "m, p" Discodeles bufoniformis MCZ38584 "Kunua, Bougainville" 1963 F "m, p" Discodeles bufoniformis MCZ38586 "Kunua, Bougainville" 1963 F "m, p" Discodeles bufoniformis MCZ38587 "Kunua, Bougainville" 1963 M "m, p, o" Discodeles bufoniformis SAMR4272A "Kunua, Bougainville" 1963 M "m, p" Discodeles bufoniformis SAMR4272B "Kunua, Bougainville" 1963 F "m, p" Discodeles bufoniformis SAMR4417 "Kunua, Bougainville" 1963 M "m, p"

247 Discodeles bufoniformis SAMR4428A Bougainville . F "m, p" Discodeles bufoniformis SAMR4428B Bougainville . F "m, p" Discodeles bufoniformis SAMR4920B "Kunua, Bougainville" apr-may 63 F "m, p" Discodeles bufoniformis SAMR4930A "Kunua, Bougainville" apr-may 63 F "m, p" Discodeles bufoniformis SAMR4930C "Kunua, Bougainville" apr-may 63 S Discodeles bufoniformis SAMR4930D "Kunua, Bougainville" apr-may 63 S Discodeles bufoniformis SAMR5159 "Kunua, Bougainville" 16-Jun-63 M "m, p" Discodeles bufoniformis SAMR7001 "Pomugu/Kandrian, New Britain" 30-Jan-66 F "m, p" Discodeles bufoniformis SAMR8050 "Turiboiru, Bougainville" 10-Mar-66 M "m, p" Discodeles bufoniformis SAMR8178A "Turiboiru, Bougainville" 13-Mar-66 F "m, p" Discodeles bufoniformis SAMR8178B "Turiboiru, Bougainville" 13-Mar-66 M "m, p" Discodeles bufoniformis SAMR8180 "Turiboiru, Bougainville" 12-Mar-66 F "m, p" Discodeles bufoniformis SAMR8319 "Kandrian, New Britain" Feb'66 F "m, p" Discodeles bufoniformis SAMR8320 "Walindi Ptn.,New Britain" 18-Jan-66 F "m, p" Discodeles bufoniformis SAMR8321 "Cape Gloucester, New Guinea" 18-Jan-66 F "m, p" Discodeles bufoniformis SAMR8322 "Kandrian, New Britain" Jan'66 F "m, p" Discodeles bufoniformis SAMR8323 "Pomugu/Kandrian, New Britain" 30-Jan-66 F "m, p" Discodeles bufoniformis SAMR8324 "Pomugu/Kandrian, New Britain" 1-Jan-66 F "m, p" Discodeles bufoniformis SAMR8352 "Kandrian, New Britain" 1-Jan-66 F "m, p" Discodeles bufoniformis SAMR8353 "Kandrian, New Britain" 1-Jan-66 F "m, p" Discodeles bufoniformis SAMR8354 "Kandrian, New Britain" 1-Jan-66 F "m, p" Discodeles bufoniformis UM71403 "Ysabel Id., Solomon Islands" 1931-32 F "m, p" Discodeles guppyi A06/SAMR45939 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles guppyi A14/SAMR47091 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p, o" Discodeles guppyi A33/SAMR45941 "Pavora River, Choiseul" 6/3/90 - 12/3/90 M "m, p" Discodeles guppyi A34/SAMR47093 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles guppyi A35/SAMR47094 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles guppyi A36/SAMR47095 "Pavora River, Choiseul" 6/3/90 - 12/3/90 M "m, p" Discodeles guppyi A37/SAMR47096 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles guppyi A38/SAMR47097 "Pavora River, Choiseul" 6/3/90 - 12/3/90 S Discodeles guppyi A46/SAMR37009 "Pavora River, Choiseul" 6/3/90 - 12/3/90 M "c, m, p" Discodeles guppyi A51/SAMR47092 "Pavora River, Choiseul" 6/3/90 - 12/3/90 M "m, p, o" Discodeles guppyi A56/SAMR47098 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles guppyi A93/SAMR47154 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles guppyi A94/SAMR47099 "Pavora River, Choiseul" 6/3/90 - 12/3/90 F "m, p" Discodeles guppyi AMR129589 "Amelei, New Britain" . c Discodeles guppyi AMR129684 "Amelei, New Britain" . c Discodeles guppyi AMR129685 "Amelei, New Britain" . c Discodeles guppyi AMR129686 "Amelei, New Britain" . c Discodeles guppyi AMR137174 (AMH 36661) "Bsurata village, Malaita Id., Solomon Ids., (08 49 S 160 49 23-Mar-91 M "m, p" E)" Discodeles guppyi AMR137175 (AMH 36662) "Bsurata village, Malaita Id., Solomon Ids., (08 49 S 160 49 23-Mar-91 M "m, p" E)" Discodeles guppyi AMR137176 (AMH 36663) "Bsurata village, Malaita Id., Solomon Ids., (08 49 S 160 49 23-Mar-91 M "m, p" E)" Discodeles guppyi AMR137178 (AMH 36665) "Bsurata village, Malaita Id., Solomon Ids., (08 49 S 160 49 23-Mar-91 M "m, p" E)"

248 Discodeles guppyi FMNH41375 "Solomon Islands, New Georgia, Munda" 1-Nov-63 F "m, p" Discodeles guppyi FMNH44295 New Georgia 1945 F "m, p" Discodeles guppyi FMNH44296(Bent) New Georgia 1945 F "m, p" Discodeles guppyi FMNH44533 . . F "m, p" Discodeles guppyi SAMR4416 "Kunua, Bougainville" 7-Sep-62 M "m, p" Discodeles guppyi SAMR4931A "Kunua, Bougainville" 8-Apr-63 S Discodeles guppyi SAMR4931B "Kunua, Bougainville" 8-Apr-63 S Discodeles guppyi SAMR4931C "Kunua, Bougainville" 8-Apr-63 S Discodeles guppyi SAMR7060 "Pomugu/Kandrian, New Britain" 30-Jan-66 S Discodeles guppyi SAMR7077 "Pomugu/Kandrian, New Britain" 30-Jan-66 S Discodeles guppyi SAMR8182 "Turiboiru, Bougainville" 12-Mar-66 F "m, p" Discodeles guppyi UM100000 "Florida Is., Halavo, Solomon Islands" 1-Oct-44 F "m, p" Discodeles opisthodon CAS9345 "Bougainville, Solomon Islands" S Discodeles opisthodon AMNH34274 "Bougainville, Solomon Islands" F m Discodeles opisthodon AMNH35354 "Bougainville, Solomon Islands" F m Discodeles opisthodon AMNH35356 "Bougainville, Solomon Islands" F m Discodeles opisthodon AMNH35357 "Bougainville, Solomon Islands" F m Discodeles opisthodon AMNH35436 "Fauro, Solomon Islands" F m Discodeles opisthodon UM55770 "Ugi Islands, Solomon Islands" 1920-25 M m Discodeles opisthodon UM55771 "Ugi Islands, Solomon Islands" 1920-25 M m Palmatorappia solomonis MCZ60844 "Mutahi, Bougainville" 18-May-66 S Palmatorappia solomonis MCZ60845 "Mutahi, Bougainville" 18-May-66 M p Palmatorappia solomonis MCZ60846 "Mutahi, Bougainville" 18-May-66 M p Palmatorappia solomonis MCZ60847 "Mutahi, Bougainville" 18-May-66 M p Palmatorappia solomonis MCZ60848 "Mutahi, Bougainville" 18-May-66 F p Palmatorappia solomonis MCZ60849 "Mutahi, Bougainville" 18-May-66 M p Palmatorappia solomonis MCZ60850 "Mutahi, Bougainville" 18-May-66 M p Palmatorappia solomonis MCZ60851 "Mutahi, Bougainville" 18-May-66 M p Palmatorappia solomonis MCZ60852 "Mutahi, Bougainville" 18-May-66 F "p,o" Palmatorappia solomonis MCZ60853 "Mutahi, Bougainville" 18-May-66 M p Palmatorappia solomonis MCZ60854 "Mutahi, Bougainville" 18-May-66 M p Palmatorappia solomonis MCZ60855 "Mutahi, Bougainville" 18-May-66 M p Palmatorappia solomonis MCZ60856 "Mutahi, Bougainville" 18-May-66 S Palmatorappia solomonis MCZ60857 "Mutahi, Bougainville" 18-May-66 M p Palmatorappia solomonis MCZ60858 "Mutahi, Bougainville" 18-May-66 F "p,o" Palmatorappia solomonis MCZ60859 "Mutahi, Bougainville" 18-May-66 M "p,o" Palmatorappia solomonis MCZ60860 "Mutahi, Bougainville" 18-May-66 F p Palmatorappia solomonis MCZ60861 "Mutahi, Bougainville" 18-May-66 F p Palmatorappia solomonis MCZ60862 "Mutahi, Bougainville" 18-May-66 F p Palmatorappia solomonis MCZ60863 "Mutahi, Bougainville" 18-May-66 M "p,o" Palmatorappia solomonis MCZ60864 "Mutahi, Bougainville" 18-May-66 M "p,o" Palmatorappia solomonis MCZ60865 "Mutahi, Bougainville" 18-May-66 M p Palmatorappia solomonis MCZ60866 "Mutahi, Bougainville" 18-May-66 M p Palmatorappia solomonis SAMR4426 "Kunua, Bougainville" 27-Aug-62 M "p,o" Palmatorappia solomonis SAMR4427 Bougainville 27-Apr-32 F p Palmatorappia solomonis SAMR4766 Bougainville 27-Apr-32 M p Palmatorappia solomonis SAMR4925A "Kunua, Bougainville" jan-feb 63 M p

249 Palmatorappia solomonis SAMR4925B "Kunua, Bougainville" jan-feb 63 M p Palmatorappia solomonis SAMR4925C "Kunua, Bougainville" jan-feb 63 M "p,o" Palmatorappia solomonis SAMR4925D "Kunua, Bougainville" jan-feb 63 M p Palmatorappia solomonis SAMR5168A "Kunua, Bougainville" 12-Jun-63 M p Palmatorappia solomonis SAMR5168B "Kunua, Bougainville" 12-Jun-63 F p Palmatorappia solomonis SAMR5168C "Kunua, Bougainville" 12-Jun-63 M p Palmatorappia solomonis SAMR8198 "Mutahi, Bougainville" 9-May-66 F p Palmatorappia solomonis SAMR8208 T.P.N.G??? . M p Platymantis acrochorda (=acrochordus) CAS101786 (paratype) "Solomon Islands, Bougainville Id., Kunua, Aresi Mt region, 6-Aug-63 F m 2000-4000 ft" Platymantis acrochorda (=acrochordus) CAS21832 (paratype) "Solomon Islands, Bougainville Id., Kunua area" F m Platymantis acrochorda (=acrochordus) FMNH169948 "Solomon Islands, Bougainville Id., Kunua area" . F m Platymantis acrochorda (=acrochordus) FMNH169949 "Solomon Islands, Bougainville Id., Kunua area" . F m Platymantis aculeodactylus AMNH34733 Choiseul Is. F m Platymantis aculeodactylus AMNH70065 "Kunua, Bougainville Is.," F m Platymantis aculeodactylus AMNH71899 "Kunua, Bougainville Is.," F m Platymantis aculeodactylus AMNH79458 "Mutahi, Bougainville Is.," F m Platymantis aculeodactylus CAS109796 "Solomon Islands, Bougainville Id., Mutahi, 2600-2900 ft" 14-Apr-66 M m Platymantis aculeodactylus CAS109797 "Solomon Islands, Bougainville Id., Mutahi, 1800-2700 ft" 17-May-66 F m Platymantis aculeodactylus CAS109798 "Solomon Islands, Bougainville Id., Mutahi, 2700 +/- 500 ft" 11-May-66 F m Platymantis aculeodactylus CAS109799 "Solomon Islands, Bougainville Id., Mutahi, 2700 +/- 500 ft" 11-May-66 F m Platymantis aculeodactylus CAS109801 "Solomon Islands, Bougainville Id., Mutahi, 2200-3200 ft" 17-May-66 F m Platymantis aculeodactylus CAS109802 "Solomon Islands, Bougainville Id., Mutahi, 2200-3200 ft" 17-May-66 F m Platymantis aculeodactylus CAS109803 "Solomon Islands, Bougainville Id., Mutahi, 2200-3200 ft" 17-May-66 F m Platymantis aculeodactylus CAS109804 "Solomon Islands, Bougainville Id., Mutahi, 2200-3200 ft" 17-May-66 F m Platymantis aculeodactylus CAS110811 "Solomon Islands, Bougainville Id., Mutahi, " 11-May-66 F m Platymantis aculeodactylus CAS110812 "Solomon Islands, Bougainville Id., Mutahi, " 13-May-66 F m Platymantis aculeodactylus CAS110813 "Solomon Islands, Bougainville Id., Mutahi, " 13-May-66 F m Platymantis aculeodactylus CAS110814 "Solomon Islands, Bougainville Id., Mutahi, " 13-May-66 F m Platymantis aculeodactylus CAS110815 "Solomon Islands, Bougainville Id., Mutahi, " 13-May-66 F m Platymantis aculeodactylus CAS110816 "Solomon Islands, Bougainville Id., Mutahi, " 13-May-66 F m Platymantis aculeodactylus CAS21830 "Solomon Islands, Bougainville Id., Kunua" M m Platymantis aculeodactylus CAS21831 "Solomon Islands, Bougainville Id., Kunua" F m Platymantis aculeodactylus CAS93988 "Solomon Islands, Bougainville Id., Kunua" 5-Aug-63 F m Platymantis aculeodactylus CAS93989 "Solomon Islands, Bougainville Id., Kunua" F m Platymantis aculeodactylus CAS93990 "Solomon Islands, Bougainville Id., Kunua" F m Platymantis aculeodactylus FMNH141976 "Solomon Islands, Bougainville Id., Kunua" F m Platymantis akarithymus CAS22875 "Bismarck Islands, New Britain Id., south coast, Ponugu, 7 1-Jan-66 M miles NW Kandrian" Platymantis batantae AMNH74192 (holotype) "West Irian: Batanta Id.,; Mt. Besar, above Wailebet, 1500'" 19-Jun-64 F Platymantis batantae AMNH74193 (paratype) "West Irian: Batanta Id.,; Mt. Besar, above Wailebet, 1500'" M Platymantis boulengeri AMNH64254 "New Britain: Iambon, camp no. 6, 1500 ft, S. slope F p Whiteman Range" Platymantis boulengeri AMNH64627 "New Britain: Iambon, camp no. 6, 1500 ft, S. slope F p Whiteman Range" Platymantis boulengeri AMNH70074 New Britain: Rabaul F p Platymantis boulengeri CAS22876 "Bismarck Islands, New Britain Id., Talasea area on 1-Jan-66 F p

250 Willaumez Peninsula" Platymantis boulengeri SAMR6162 "Kerevat, New Britain" May'65 F p Platymantis boulengeri SAMR7067 "Kandrian, New Britain" 30-Jan-66 F p Platymantis boulengeri SAMR7090 "Malasait/Baining Rgs., New Britain" 11-Jan-66 F p Platymantis boulengeri SAMR7108 "Pomugu/Kandrian, New Britain" 30-Jan-66 F p Platymantis boulengeri SAMR7118 "Raunsegna, N. Bainings, New Britain" 27-Dec-65 F p Platymantis boulengeri SAMR7706 "Kerevat, New Britain" 10-Feb-66 S Platymantis boulengeri SAMR8124 "Kerevat, New Britain" July'66 M p Platymantis boulengeri SAMR8848A "Kerevat, New Britain" July'66 F p Platymantis boulengeri SAMR8848B "Kerevat, New Britain" July'66 F p Platymantis boulengeri SAMR8848C "Kerevat, New Britain" July'66 F p Platymantis boulengeri SAMR8848D "Kerevat, New Britain" July'66 F p Platymantis boulengeri SAMR8848E "Kerevat, New Britain" July'66 F p Platymantis boulengeri SAMR8848F "Kerevat, New Britain" July'66 F p Platymantis boulengeri SAMR8848G "Kerevat, New Britain" July'66 F p Platymantis boulengeri SAMR8848H "Kerevat, New Britain" July'66 F p Platymantis boulengeri SAMR8848I "Kerevat, New Britain" July'66 S Platymantis parkeri AMNH69314 "S end of Buka Is., Solomon Islands" M "m, p" Platymantis parkeri AMNH69315 "S end of Buka Is., Solomon Islands" M "m, p" Platymantis cheesmanae AMNH75101 (paratype) "Cyclops Range, 3000-4000', West Irian, New Guinea" F Platymantis cheesmanae AMNH75102 (paratype) "Cyclops Range, 3000-4000', West Irian, New Guinea" F Platymantis cheesmanae FMNH43319 "New Guinea, Dutch New Guinea, Doromena" 1-May-45 M Platymantis corrugatus CAS129284 "Philippine Ids., Negros Id., Negros Oriental Prov. E slope 17-Dec-61 M p Cuernos de Negros, 1200 ft" Platymantis corrugata (=corrugatus) CAS145690 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, Sierra 31 Mar- 1 Apr M p Bullones" 1964 Platymantis corrugata (=corrugatus) CAS145691 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, Sierra 2-3 Apr 1964 M p Bullones" Platymantis corrugata (=corrugatus) CAS145692 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, Sierra 2-3 Apr 1964 M p Bullones" Platymantis corrugata (=corrugatus) CAS145693 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, Sierra 4-5 Apr 1964 M p Bullones" Platymantis corrugata (=corrugatus) CAS22242 "Philippine Ids., Mindoro Id., Mindoro Or. Prov., SE slope of 4-Apr-63 F p Barawanan Pk., E side of Mt Halcon, 1700 ft" Platymantis corrugata (=corrugatus) CAS22243 "Philippine Ids., Mindoro Id., Mindoro Or. Prov., SE slope of 4-Apr-63 F p Barawanan Pk., E side of Mt Halcon, 1700 ft" Platymantis corrugata (=corrugatus) CAS22817 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 26-Apr-62 M p km SE of Sierra Bullones" Platymantis corrugata (=corrugatus) CAS23025 "Philippine Ids., Camiguin Id., Catibawassan Falls area, 4.5 28-29 June 1966 M p Km S Mambajao town, secondary forest, 1200 ft" Platymantis corrugata (=corrugatus) CAS23026 "Philippine Ids., Camiguin Id., Catibawassan Falls area, 4.5 28-29 June 1966 F p Km S Mambajao town, secondary forest, 1200 ft" Platymantis corrugata (=corrugatus) CAS23701 "Philippine Ids., Negros Id., Negros Oriental Prov. 5 km NW 8-Apr-62 F p of Biaknabato, La Castellana area, S slope Mt Canlaon, 3100 ft" Platymantis corrugata (=corrugatus) CAS62227 "Polillo Id., Philippine Ids." 15 Jul - 16 Aug F p 1920

251 Platymantis corrugata (=corrugatus) FMNH172815 "Philippine Ids., Mindanao, Davao" F p Platymantis corrugata (=corrugatus) FMNH173174 "Philippine Ids., Negros Id., Cuernos de Negros" F p Platymantis corrugata (=corrugatus) FMNH247745 "Philippine Ids., Negros Id., Negros Oriental Prov, Mt. 4-Jul-90 M p Talinis" Platymantis corrugata (=corrugatus) FMNH50536 "Philippine Ids., Mindanao, Davao Prov., E slope of Mt 31-Aug-46 M p McKinley" Platymantis corrugata (=corrugatus) FMNH50539 "Philippine Ids., Mindanao, Davao Prov., E slope of Mt 1-Sep-46 M p McKinley" Platymantis corrugata (=corrugatus) FMNH50541 "Philippine Ids., Mindanao, Davao Prov., E slope of Mt 7-Sep-46 F p McKinley" Platymantis corrugata (=corrugatus) FMNH50545 "Philippine Ids., Mindanao, Davao Prov., Tagum, Sitio 11-Oct-46 F p Taglawig" Platymantis corrugata (=corrugatus) FMNH50547 "Philippine Ids., Mindanao, Davao Prov., Upi, Burungkot" 7-Jan-47 F p Platymantis corrugata (=corrugatus) FMNH50548 "Philippine Ids., Mindanao, Davao Prov., Mt Apo, Todaya" 1-Nov-46 M p Platymantis corrugata (=corrugatus) SAMR13601 "Bohol Is., Philippines" 29-Apr-64 F p Platymantis corrugata (=corrugatus) SAMR13602A "Bohol Is., Philippines" 1-May-62 F p Platymantis corrugata (=corrugatus) SAMR13602B "Bohol Is., Philippines" 1-May-62 S Platymantis corrugata (=corrugatus) SAMR8856 "Negos Is., Philippines" M p Platymantis corrugata (=corrugatus) SAMR8857 "16 Kms S Canayan Town, Philippines" 29-May-56 M p Platymantis dorsalis CAS124785 "Philippine Ids., Mindanao Id., Kibwalam, Malalag, Davao 13-May-67 . del Sur, 2000-2800 ft" Platymantis dorsalis CAS124786 "Philippine Ids., Mindanao Id., Kibwalam, Malalag, Davao 18 dec 1967-1 F del Sur, 1800-2200 ft" jan 1968 Platymantis dorsalis CAS124787 "Philippine Ids., Mindanao Id., Kibwalam, Malalag, Davao 18 dec 1967-1 F del Sur, 1800-2200 ft" jan 1968 Platymantis dorsalis CAS124788 "Philippine Ids., Mindanao Id., Kibwalam, Malalag, Davao 18 dec 1967-1 F del Sur, 1800-2200 ft" jan 1968 Platymantis dorsalis CAS124789 "Philippine Ids., Mindanao Id., Kibwalam, Malalag, Davao 18 dec 1967-1 F del Sur, 1800-2200 ft" jan 1968 Platymantis dorsalis CAS133991 "Philippine Ids., Negros Id., Negros Oriental Prov., 1-Apr-58 M Dayungan,N slope Talinis, 3500 ft" Platymantis dorsalis CAS133992 "Philippine Ids., Negros Id., Negros Oriental Prov., 1-Apr-58 M Dayungan,N slope Talinis, 3500 ft" Platymantis dorsalis CAS133994 "Philippine Ids., Negros Id., Negros Oriental Prov., 11-Apr-58 F Dayungan,E slope Talinis, 3200 ft" Platymantis dorsalis CAS16610 "Philippine Ids., Negros Id., Negros Oriental Prov., Upper 29-Dec-54 F Mabuya Creek area, S side Mayaposi Hill, 23 km W of Bais" Platymantis dorsalis CAS22871 "Philippine Ids., Danjugan Id., Negros Occ. Prov., (sm. Id. 11-May-66 F Near Camayan on W side of Negros)" Platymantis dorsalis CAS22872 "Philippine Ids., Danjugan Id., Negros Occ. Prov., (sm. Id. 11-May-66 M Near Camayan on W side of Negros)" Platymantis dorsalis FMNH110364 "Philippine Ids., Luzon, Mt. Makiling" F Platymantis dorsalis FMNH110368 "Philippine Ids., Luzon, Mt. Makiling" M Platymantis dorsalis FMNH173166 "Philippine Ids., Luzon, Ifagao, Banane" F Platymantis dorsalis FMNH173168 "Philippine Ids., Luzon, Ifagao, Banane" F Platymantis dorsalis FMNH173173 "Philippine Ids., Luzon, Ifagao, Banane" M

252 Platymantis dorsalis FMNH50552 "Philippine Ids., Mindanao, Davao Prov. E slope Mt 20-Aug-46 F McKinley" Platymantis dorsalis FMNH50562 "Philippine Ids., Mindanao, Davao Prov. Caburan, " 20-Jan-47 F Platymantis dorsalis FMNH61601 "Philippine Ids., Negros Id., Bais, Pagyabunan" 8-May-49 F Platymantis dorsalis FMNH61602 "Philippine Ids., Negros Id., Bais, Pagyabunan" 8-May-49 F Platymantis dorsalis FMNH77890 "Philippine Ids., Negros Id., Cuernos de Negros" 1-Dec-55 F Platymantis dorsalis FMNH77891 "Philippine Ids., Negros Id., Cuernos de Negros" 1-Dec-55 F Platymantis dorsalis FMNH96080 "Philippine Ids., Mindanao, Zamboanga Prov., Mt 1-Apr-56 F Malindang" Platymantis dorsalis SAMR13598A "Philippine, Negros Ids." 12-May-58 F Platymantis dorsalis SAMR13598B "Philippine, Negros Ids." 12-May-58 F Platymantis gilliardi AMNH23545 "Admiralty Islands, Pak (Tack?) Isd." . Platymantis gilliardi AMNH23546 "Admiralty Islands, Pak (Tack?) Isd." . Platymantis gilliardi AMNH23547 "Admiralty Islands, Lambussu Isd. (Jesu Maria Isd.)" F Platymantis gilliardi AMNH64253 (type) "Iambon, camp 6, 1500', s. slope Whiteman Range, New F Britain" Platymantis gilliardi CAS22877 "Bismarck Ids., New Britain Id., south coast 7 miles NW of 1-Jan-66 M Pomugu, Kandrian" Platymantis gilliardi CAS22878 "Bismarck Ids., New Britain Id., south coast 7 miles NW of 1-Jan-66 F Pomugu, Kandrian" Platymantis gilliardi SAMR129626 F Platymantis gilliardi SAMR129688 F Platymantis gilliardi SAMR7754 F Platymantis guentheri CAS23152 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 2-May-62 F km SE Sierra Bullones" Platymantis guentheri CAS23154 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 2-May-62 F km SE Sierra Bullones" Platymantis guentheri CAS23155 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 2-May-62 F km SE Sierra Bullones" Platymantis guentheri CAS23156 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 2-May-62 F km SE Sierra Bullones" Platymantis guentheri CAS23157 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 2-May-62 F km SE Sierra Bullones" Platymantis guentheri CAS23158 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 3-May-62 M km SE Sierra Bullones" Platymantis guentheri CAS23159 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 3-May-62 M km SE Sierra Bullones" Platymantis guentheri CAS23160 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 3-May-62 F km SE Sierra Bullones" Platymantis guentheri CAS23162 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 3-May-62 F km SE Sierra Bullones" Platymantis guentheri CAS23163 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 3-May-62 F km SE Sierra Bullones" Platymantis guentheri CAS23169 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 8-May-62 F km SE Sierra Bullones" Platymantis guentheri CAS23170 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 7-May-62 F km SE Sierra Bullones"

253 Platymantis guentheri CAS23172 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 7-May-62 F km SE Sierra Bullones" Platymantis guentheri CAS23174 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 8-May-62 M km SE Sierra Bullones" Platymantis guentheri CAS23175 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 8-May-62 M km SE Sierra Bullones" Platymantis guentheri CAS23176 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 7-May-62 M km SE Sierra Bullones" Platymantis guentheri CAS23177 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 8-May-62 M km SE Sierra Bullones" Platymantis guentheri CAS23179 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 8-May-62 F km SE Sierra Bullones" Platymantis guentheri CAS23181 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 8-May-62 F km SE Sierra Bullones" Platymantis guentheri CAS23182 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 20-Apr-62 M km SE Sierra Bullones" Platymantis guentheri FMNH134987 "Philippine Ids., Bohol Id., Dusita area, 10 km SE Sierra 17-Apr-61 F Bullones" Platymantis guentheri FMNH134988 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 8-May-61 M km SE Sierra Bullones" Platymantis guentheri FMNH50573 "Philippine Ids., Mindanao Id., Davao Prov., Tagum, Sitio 12-Oct-46 F Taglawig" Platymantis guentheri UM175608 "Philippine Is., S. Negros Oriental, Lake Balinsasayao, 14 May 18 1983 F Km W + 3km N of Dumaguete (123 11'E; 9 22'N) found in leaf axil of coconut" Platymantis guentheri UM175609 "Philippine Is., S. Negros Oriental, Lake Balinsasayao, 14 F Km W + 3km N of Dumaguete (123 11'E; 9 22'N) found near Tagat farm at 900 M alt" Platymantis guppyi A20/SAMR47131 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis guppyi A50/SAMR47132 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis guppyi A52/SAMR47133 "Pavora River, Choiseul" 6/3-12/3'90 F "m, p" Platymantis guppyi BIG/SAMR47128 "Isabel, Solomons" 1989 F "m, p, o" Platymantis guppyi MED/SAMR47129 "Isabel, Solomons" 1989 F "m, p" Platymantis guppyi SAMR4242 "Kunua, Bougainville" 28-Feb-62 M "m, p" Platymantis guppyi SAMR4273A Bougainville . F "m, p" Platymantis guppyi SAMR4273B Bougainville . M "m, p" Platymantis guppyi SAMR4414 "Kunua, Bougainville" 16-Jul-62 F "m, p" Platymantis guppyi SAMR4415 "Kunua, Bougainville" 16-Jul-62 F "m, p" Platymantis guppyi SAMR4917A "Kunua, Bougainville" feb 63 F "m, p" Platymantis guppyi SAMR4917B "Kunua, Bougainville" feb 63 M "m, p, o" Platymantis guppyi SAMR4917C "Kunua, Bougainville" feb 63 M "m, p" Platymantis guppyi SAMR4917D "Kunua, Bougainville" feb 63 M "m, p" Platymantis guppyi SAMR4917E "Kunua, Bougainville" feb 63 M "m, p" Platymantis guppyi SAMR4917F "Kunua, Bougainville" feb-may 63 F "m, p" Platymantis guppyi SAMR4922A "Kunua, Bougainville" feb-may 63 F "m, p" Platymantis guppyi SAMR4922B "Kunua, Bougainville" feb-may 63 F "m, p" Platymantis guppyi SAMR4922C "Kunua, Bougainville" feb-may 63 M "m, p, o"

254 Platymantis guppyi SAMR4922D "Kunua, Bougainville" feb-may 63 F "m, p" Platymantis guppyi SAMR4922E "Kunua, Bougainville" feb-may 63 F "m, p" Platymantis guppyi SAMR5158 "Kunua, Bougainville" 16-Jun-63 F "m, p" Platymantis guppyi SAMR8197 "Kunua, Bougainville" 10-May-66 F "m, p" Platymantis guppyi SMA/SAMR47130 "Isabel, Solomons" 1989 F "m, p" Platymantis hazelae AMNH88040 "Negros Id., Negros Oriental Prov., slope on N. side F Bunyawi ravine about 1 km NW of N. peak of Cuernos de Negros, 3600-3700'" Platymantis hazelae AMNH88041 "Negros Id., Negros Oriental Prov., slope on N. side F Bunyawi ravine about 1 km NW of N. peak of Cuernos de Negros, 3600-3700'" Platymantis hazelae CAS133946 "Philippine Ids., Negros Id., Negros Oriental Prov., 24-Dec-57 M Dayungan, Talinis, 3600 ft" Platymantis hazelae CAS133948 "Philippine Ids., Negros Id., Negros Oriental Prov., 26-Dec-57 M Dayungan, Talinis, 3500 ft" Platymantis hazelae CAS133949 "Philippine Ids., Negros Id., Negros Oriental Prov., 26-Dec-57 M Dayungan, Talinis, 3500 ft" Platymantis hazelae CAS133950 "Philippine Ids., Negros Id., Negros Oriental Prov., 27-Dec-57 M Dayungan, Talinis, 4200 ft" Platymantis hazelae CAS133951 "Philippine Ids., Negros Id., Negros Oriental Prov., 27-Dec-57 F Dayungan, Talinis, 3200 ft" Platymantis hazelae CAS133952 "Philippine Ids., Negros Id., Negros Oriental Prov., 28-Dec-57 M Dayungan, Talinis, 3600 ft" Platymantis hazelae CAS133953 "Philippine Ids., Negros Id., Negros Oriental Prov., 8-Feb-58 M Dayungan, Talinis, 4000-5000 ft" Platymantis hazelae CAS133954 "Philippine Ids., Negros Id., Negros Oriental Prov., 8-Feb-58 F Dayungan, Talinis, 4000-5000 ft" Platymantis hazelae CAS134232 "Philippine Ids., Negros Id., Negros Oriental Prov., 22-Dec-60 F Palaypay, 7km SW Pamlona, 1000 ft" Platymantis hazelae CAS134233 "Philippine Ids., Negros Id., Negros Oriental Prov., 22-Dec-60 F Palaypay, 7km SW Pamlona, 1000 ft" Platymantis hazelae CAS134234 "Philippine Ids., Negros Id., Negros Oriental Prov., 22-Dec-60 M Palaypay, 7km SW Pamlona, 1000 ft" Platymantis hazelae CAS137641 "Philippine Ids., Panay Id., Aklan Prov., Laserna Barrio, 14-May-73 F 2200-2300 ft" Platymantis hazelae CAS137642 "Philippine Ids., Panay Id., Aklan Prov., Laserna Barrio, 14-May-73 M 2200-2300 ft" Platymantis hazelae CAS61478 (paratype) "Balbalan, Kalinga subprov. Mt Prov., Luzon Id., Philippines 25-Apr-20 M " Platymantis ingeri AMNH88049 (paratype) "Bohol Id., Cantaub area, 15 km SE of Sierra Bullones, F 2500'" Platymantis ingeri CAS133548 "Philippine Ids., Mindanao Id., Agusan del Norte Prov. S 21-May-71 F side Mt Hilonghilong, 5000-5100 ft" Platymantis ingeri CAS21198 (paratype) "Philippine Ids., Bohol Id., Bohol Prov., Disuta area, 11 km 22-Apr-62 F SE Sierra Bullones, 2300 ft" Platymantis ingeri CAS21208 (paratype) "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 15 6-May-62 F km SE Sierra Bullones, 2500 ft"

255 Platymantis ingeri CAS21212 "Philippine Ids., Bohol Id., Bohol Prov., Disuta area, 11 km 7-May-62 F SE Sierra Bullones, 2300 ft" Platymantis ingeri CAS21216 "Philippine Ids., Bohol Id., Bohol Prov., Disuta area, 12 km 12-May-62 F SE Sierra Bullones, 2300 ft" Platymantis ingeri CAS21217 "Philippine Ids., Bohol Id., Bohol Prov., Disuta area, 12 km 12-May-62 M SE Sierra Bullones, 2300 ft" Platymantis ingeri CAS21717 "Philippine Ids., Bohol Id., Bohol Prov., Disuta area, 11 km 6-May-62 F SE Sierra Bullones, 2300 ft" Platymantis ingeri CAS21718 "Philippine Ids., Bohol Id., Bohol Prov., Disuta area, 11 km 6-May-62 F SE Sierra Bullones, 2300 ft" Platymantis ingeri CAS23165 "Philippine Ids., Bohol Id., Bohol Prov., Cantaub area, 14 8-May-62 F km SE Sierra Bullones" Platymantis ingeri CAS23434 "Philippine Ids., Bohol Id., Bohol Prov., Disuta area, Sierra 31 mar - 1 apr F Bullones, 1200 ft" 1964 Platymantis ingeri CAS93354 (paratype) "Philippine Ids., Bohol Id., Bohol Prov., Disuta area, 10 km 25-Apr-61 F SE Sierra Bullones, 1600 ft" Platymantis ingeri CAS93355 (paratype) "Philippine Ids., Bohol Id., Bohol Prov., Disuta area, 10 km 25-Apr-61 M SE Sierra Bullones, 1600 ft" Platymantis insulatus CAS117440 "Philippine Ids., Iloilo subprov., South Gigante Id., off coast 11-Jun-68 . of Panay Id., 100-150 ft" Platymantis insulatus CAS119967 (paratype) "Philippine Ids., Iloilo subprov., South Gigante Id., off coast 11-Jun-68 F of Panay Id." Platymantis insulatus CAS119968 (paratype) "Philippine Ids., Iloilo subprov., South Gigante Id., off coast 11-Jun-68 . of Panay Id." Platymantis insulatus CAS119969 (paratype) "Philippine Ids., Iloilo subprov., South Gigante Id., off coast 11-Jun-68 F of Panay Id." Platymantis insulatus CAS157235 "Philippine Ids., Iloilo Prov., Gigante North Id." 7-8 Jul 1984 . Platymantis insulatus CAS157236 "Philippine Ids., Iloilo Prov., Gigante North Id." 7-8 Jul 1984 . Platymantis insulatus CAS157237 "Philippine Ids., Iloilo Prov., Gigante North Id." 7-8 Jul 1984 F Platymantis insulatus CAS157238 "Philippine Ids., Iloilo Prov., Gigante North Id." 7-8 Jul 1984 . Platymantis lawtoni CAS135733 (paratype) "Philippine Ids., Romblon Prov., Tablas Id., Busay, 27-May-72 . Dubduban Barrio: San Agustin area, 800-1800 ft" Platymantis levigatus CAS136098 (paratype) "Philippine Ids., Tablas Id., San Agustin, Dubduban, 800 ft" 26-May-72 M Platymantis levigatus CAS137258 "Philippine Ids., Romblon Prov., Tablas Id., Busay, 22-May-72 F Dubduban Barrio: San Agustin area, 50-150 ft" Platymantis levigatus CAS137259 "Philippine Ids., Romblon Prov., Tablas Id., Busay, 25-May-72 M Dubduban Barrio: San Agustin area, 800 ft" Platymantis levigatus CAS137260 "Philippine Ids., Romblon Prov., Tablas Id., Busay, 25-May-72 M Dubduban Barrio: San Agustin area, 800 ft" Platymantis levigatus CAS137261 "Philippine Ids., Romblon Prov., Tablas Id., Busay, 25-May-72 F Dubduban Barrio: San Agustin area, 800 ft" Platymantis levigatus CAS137262 "Philippine Ids., Romblon Prov., Tablas Id., Busay, 25-May-72 M Dubduban Barrio: San Agustin area, 800 ft" Platymantis levigatus CAS137263 "Philippine Ids., Romblon Prov., Tablas Id., Busay, 26-May-72 F Dubduban Barrio: San Agustin area, 800 ft" Platymantis levigatus CAS137264 "Philippine Ids., Romblon Prov., Tablas Id., Busay, 26-May-72 F Dubduban Barrio: San Agustin area, 800 ft"

256 Platymantis levigatus CAS137295 "Philippine Ids., Romblon Prov., Tablas Id., Mt Progresso, 1-May-72 S San Agustin area, 1200-1700 ft" Platymantis levigatus CAS139208 "Philippine Ids., Romblon Prov., Tablas Id., Busay, 26-May-72 S Dubduban Barrio: San Agustin area, 800 ft" Platymantis macrops CAS110810 "Solomon Islands., Bougainville Id., Mutahi, 2700 ft" 20-May-66 M Platymantis macrops CAS21795 (paratype) "Kieta area, Bougainville, Solomons" 15-May-05 M Platymantis magnus AMNH98079 (paratype) "New Ireland, ca. 3 mi S Kavieng" F Platymantis magnus CAS143639 (paratype) "Bismarck Archipelago: New Britain Id., Madina High 25-Sep-76 M School area, 80kms SE Kavieng" Platymantis mimicus CAS22874 (paratype) "Bismarck Ids., New Britain Id., Numundo plantation 1-Jan-66 M Willaumez Peninsula, 18 miles S Talasea" Platymantis mimicus SAMR18083 M Platymantis mimicus SAMR18084 F Platymantis mimicus SAMR18085 M Platymantis mimicus SAMR18086 M Platymantis mimicus SAMR18087 M Platymantis mimicus SAMR18088 S Platymantis mimicus SAMR7750 M Platymantis mimicus SAMR7751 M Platymantis myersi A19/SAMR47158 "Pavora River, Choiseul" 6/3-12/3'90 F "m, p" Platymantis myersi A39/SAMR47159 "Pavora River, Choiseul" 6/3-12/3'90 F "m, p" Platymantis myersi A81/SAMR37010 "Pavora River, Choiseul" 6/3-12/3'90 F "c, m, p" Platymantis myersi A90/SAMR47157 "Pavora River, Choiseul" 6/3-12/3'90 F "m, p, o" Platymantis myersi AMNH35348 (type) "Bougainville, Solomon Islands" F "m, p" Platymantis myersi AMNH70066 "Bougainville Id., Kunua" S Platymantis myersi AMNH70067 "Bougainville Id., Kunua" M "m, p" Platymantis myersi CAS106742 "Solomon Islands, Bougainville, Melilup, 3000-4000 ft" 19-May-66 M "m, p" Platymantis myersi CAS106743 "Solomon Islands, Bougainville, Melilup, 3000-4000 ft" 19-May-66 S Platymantis myersi CAS106744 "Solomon Islands, Bougainville, Melilup, 3000-4000 ft" 19-May-66 F "m, p" Platymantis myersi CAS106745 "Solomon Islands, Bougainville, Melilup, 3000-4000 ft" 19-May-66 F "m, p" Platymantis myersi CAS106746 "Solomon Islands, Bougainville, Melilup, 3000-4000 ft" 19-May-66 F "m, p" Platymantis myersi CAS106747 "Solomon Islands, Bougainville, Mutahi, 2700 ft" 20-May-66 S Platymantis myersi CAS106748 "Solomon Islands, Bougainville, Mutahi, 2700 ft" 20-May-66 S Platymantis myersi CAS9335 (paratype) "Bougainville, Solomon Islands" M "m, p" Platymantis myersi CAS93993 "Solomon Islands., Bougainville Id., Kunua" 6-Aug-63 F "m, p" Platymantis myersi CAS93994 "Solomon Islands., Bougainville Id., Kunua" 6-Aug-63 F "m, p" Platymantis myersi FMNH141977 "Solomon Islands., Bougainville Id., Kunua" F "m, p" Platymantis myersi FMNH141978 "Solomon Islands., Bougainville Id., Kunua" F "m, p" Platymantis myersi FMNH141979 "Solomon Islands., Bougainville Id., Kunua" F "m, p" Platymantis neckeri A17/SAMR47155 "Pavora River, Choiseul" 6/3-12/3'90 F "m, p" Platymantis neckeri A41/SAMR47156 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis neckeri A57/SAMR37011 "Pavora River, Choiseul" 6/3-12/3'90 F "c, m, p" Platymantis neckeri AMNH34317 "Bougainville, Solomon Islands" M "m, p" Platymantis neckeri AMNH34318 "Bougainville, Solomon Islands" M "m, p" Platymantis neckeri AMNH34325 "Kunua,montane region, Bougainville" F "m, p" Platymantis neckeri AMNH34329 (type) "Kunua, Bougainville" F "m, p" Platymantis neckeri AMNH35334 (paratype) "Bougainville, Solomon Islands" M "m, p"

257 Platymantis neckeri AMNH79560 "Mutahi, Bougainville, 2700'" F "m, p" Platymantis neckeri SAMR4926A "Kunua, Bougainville" 6-Feb-63 F "m, p" Platymantis neckeri SAMR4926B "Kunua, Bougainville" 6-Feb-63 F "m, p" Platymantis neckeri SAMR4926C "Kunua, Bougainville" 6-Feb-63 M "m, p" Platymantis neckeri SAMR4926D "Kunua, Bougainville" 6-Feb-63 M "m, p, o" Platymantis neckeri SAMR4928A "Kunua, Bougainville" 28-Feb-63 M "m, p" Platymantis neckeri SAMR4928B "Kunua, Bougainville" 28-Feb-63 M "m, p, o" Platymantis neckeri SAMR4928C "Kunua, Bougainville" 28-Feb-63 M "m, p" Platymantis neckeri SAMR8211 "Mutahi, Bougainville" 10-May-66 F "m, p" Platymantis neckeri SAMR8261 "Mutahi, Bougainville" 9-May-66 M "m, p" Platymantis papuensis AMNH59970 "Fergusson Is., Iamelele no. 1, 0-50m PNG" F p Platymantis papuensis AMNH59972 "Fergusson Is., Iamelele no. 1, 0-50m PNG" F p Platymantis papuensis AMNH59974 "Fergusson Is., Iamelele no. 1, 0-50m PNG" F p Platymantis papuensis AMNH59975 "Fergusson Is., Iamelele no. 1, 0-50m PNG" F p Platymantis papuensis AMNH59976 "Fergusson Is., Iamelele no. 1, 0-50m PNG" F p Platymantis papuensis AMNH60144 "Normanby Is., Waikaiuna, 15m, PNG" F p Platymantis papuensis AMNH60145 "Normanby Is., Waikaiuna, 15m, PNG" F p Platymantis papuensis AMNH60148 "Normanby Is., Waikaiuna, 15m, PNG" M p Platymantis papuensis CAS107537 "New Guinea, Lae" 17-Feb-66 F p Platymantis papuensis SAMR15255A "4km N Lae, Morobe Provence, TPNG" 7-Mar-74 M p Platymantis papuensis SAMR15255B "4km N Lae, Morobe Provence, TPNG" 7-Mar-74 M p Platymantis papuensis SAMR15257 "4km N Lae, Morobe Provence, TPNG" 7-Mar-74 F p Platymantis papuensis SAMR3606A "Dutch New Guinea, nr Hollandia" Dec'44-Jan'45 F p Platymantis papuensis SAMR4397 F p Platymantis papuensis SAMR5800 "Lovengan, Manus Is. , New Guinea" 1-Aug-64 M p Platymantis papuensis SAMR5882A "Popondetta, Papua New Guinea" Nov-Dec 1964 F p Platymantis papuensis SAMR5882B "Popondetta, Papua New Guinea" Nov-Dec 1964 M p Platymantis papuensis SAMR5889A "Pukago, nr Maprik, New Guinea" 31 Oct - 1 Nov F p '64 Platymantis papuensis SAMR5889B "Pukago, nr Maprik, New Guinea" 31 Oct - 1 Nov F p '64 Platymantis papuensis SAMR8972 "Lorengau, Manus Is., Admiralty Is." 1-Aug-64 F p Platymantis papuensis SAMR8973 "Lorengau, Manus Is., Admiralty Is." 1-Aug-64 F p Platymantis papuensis SAMR9440 "Agenehambo/Popondetta, New Guinea" 15-Feb-68 M p Platymantis papuensis SAMR9441 "Agenehambo/Popondetta, New Guinea" 15-Feb-68 M p Platymantis parkeri A30/SAMR47126 "Pavora River, Choiseul" 4 -21 mar 1990 F "m, p" Platymantis parkeri A84/SAMR47127 "Pavora River, Choiseul" 4 -21 mar 1990 F "m, p" Platymantis parkeri AMNH70069 (paratype) "Bougainville Island, Kunua area" F "m, p" Platymantis parkeri AMNH70070 (paratype) "Bougainville Island, Kunua area" F "m, p" Platymantis parkeri AMNH70071 (paratype) "Bougainville Island, Kunua area" F "m, p" Platymantis parkeri AMNH79563 "Bougainville Island, Turiboiru" F "m, p" Platymantis parkeri CAS101787 (paratype) "Solomon Islands, Bougainville, Kunua area" 17-Jun-63 M "m, p" Platymantis parkeri CAS109805 "Solomon Islands, Bougainville, Mutahi, 2700 ft" 11-May-66 F "m, p" Platymantis parkeri CAS109806 "Solomon Islands, Bougainville, Mutahi, 2700 ft" 20-May-66 M "m, p" Platymantis parkeri CAS109807 "Solomon Islands, Bougainville, Mutahi, 2700 ft" 20-May-66 M "m, p" Platymantis parkeri CAS109808 "Solomon Islands, Bougainville, Mutahi, 2700 ft" 20-May-66 F "m, p" Platymantis parkeri CAS109809 "Solomon Islands, Bougainville, Mutahi, 2700 ft" 20-May-66 F "m, p"

258 Platymantis parkeri CAS109810 "Solomon Islands, Bougainville, Mutahi, 2700 ft" 20-May-66 M "m, p" Platymantis parkeri CAS21773 (paratype) "Solomon Islands, Bougainville, Kunua area" 1-Jun-62 F "m, p" Platymantis parkeri CAS21774 (paratype) "Solomon Islands, Bougainville, Kunua area" 1-Jun-62 M "m, p" Platymantis parkeri CAS93987 "Solomon Islands, Bougainville Id., Kunua" 23-Jul-63 F "m, p" Platymantis parkeri SAMR4424 "Kunua, Bougainville" 2-Jul-62 F "m, p" Platymantis parkeri SAMR5169A "Kunua, Bougainville" 12-Jun-63 M "m, p, o" Platymantis parkeri SAMR5169B "Kunua, Bougainville" 12-Jun-63 F "m, p, o" Platymantis parkeri SAMR8199 "Mutahi, Bougainville" 13-May-66 M "m, p, o" Platymantis pelewensis AMNH55040 "Koror, Palau Island, N. Pacific" F Platymantis pelewensis FMNH121586 "Palau Id., Pelelin Id" jan-feb 1948 M Platymantis pelewensis FMNH121587 "Palau Id., Pelelin Id" jan-feb 1948 M Platymantis pelewensis FMNH121588 "Palau Id., Pelelin Id" jan-feb 1948 M Platymantis pelewensis FMNH121590 "Palau Id., Pelelin Id" jan-feb 1948 M Platymantis pelewensis FMNH121591 "Palau Id., Pelelin Id" jan-feb 1948 M Platymantis pelewensis FMNH121592 "Palau Id., Pelelin Id" jan-feb 1948 M Platymantis pelewensis FMNH121593 "Palau Id., Pelelin Id" jan-feb 1948 M Platymantis pelewensis FMNH121594 "Palau Id., Pelelin Id" jan-feb 1948 M Platymantis pelewensis FMNH121595 "Palau Id., Pelelin Id" jan-feb 1948 M Platymantis pelewensis FMNH121596 "Palau Id., Pelelin Id" jan-feb 1948 F Platymantis pelewensis FMNH121597 "Palau Id., Pelelin Id" jan-feb 1948 M Platymantis pelewensis FMNH69781 "Palau Id., Angaur Id" F Platymantis pelewensis FMNH69782 "Palau Id., Angaur Id" F Platymantis pelewensis FMNH69783 "Palau Id., Angaur Id" F Platymantis pelewensis FMNH69784 "Palau Id., Angaur Id" M Platymantis pelewensis FMNH69785 "Palau Id., Angaur Id" M Platymantis pelewensis FMNH69787 "Palau Id., Angaur Id" M Platymantis pelewensis FMNH69789 "Palau Id., Angaur Id" M Platymantis pelewensis FMNH69790 "Palau Id., Angaur Id" M Platymantis pelewensis FMNH69791 "Palau Id., Angaur Id" M Platymantis pelewensis FMNH69792 "Palau Id., Angaur Id" M Platymantis pelewensis FMNH69793 "Palau Id., Angaur Id" M Platymantis pelewensis FMNH69796 "Palau Id., Angaur Id" M Platymantis pelewensis FMNH69798 "Palau Id., Angaur Id" M Platymantis pelewensis FMNH69800 "Palau Id., Angaur Id" M Platymantis pelewensis FMNH69801 "Palau Id., Angaur Id" M Platymantis pelewensis FMNH69803 "Palau Id., Angaur Id" M Platymantis polillensis CAS62251 (syntype) "Polillo Id., Philippine Ids." 4-16 Aug 1920 M Platymantis polillensis CAS62252 (syntype) "Polillo Id., Philippine Ids." 4-16 Aug 1920 S Platymantis punctata AMNH74185 "West Irian, Batanta Island, Mt. Besar above Wailebet, F 1500-1600 ft" Platymantis punctata AMNH74186 "West Irian, Batanta Island, Mt. Besar above Wailebet, M 1500-1600 ft" Platymantis punctata AMNH74188 "West Irian, Batanta Island, Mt. Besar above Wailebet, M 1500-1600 ft" Platymantis rhipiphalcus CAS22873 (paratype) "Bismarck Ids., New Britain Id., San Remo plantation 1-Jan-66 F Willaumez Peninsula, 20 mile Talasea" Platymantis rubristriatus FMNH14192 "Indonesia, Waigei Id." 6-Jun-29 M

259 Platymantis rubristriatus FMNH14193 "Indonesia, Waigei Id." 6-Jun-29 M Platymantis rubristriatus FMNH14194 "Indonesia, Waigei Id." 6-Jun-29 F Platymantis schmidti AMNH64258 (paratype) "Whiteman Range, Camp 6, Iambon, S. slope, 1500' New F p Britain" Platymantis schmidti AMNH64259 (paratype) "Whiteman Range, Camp 6, Iambon, S. slope, 1500' New F p Britain" Platymantis schmidti AMNH64261 (paratype) "Whiteman Range, Camp 6, Iambon, S. slope, 1500' New F p Britain" Platymantis schmidti AMNH64264 (paratype) "Whiteman Range, Camp 6, Iambon, S. slope, 1500' New F p Britain" Platymantis schmidti AMNH83083 "E. New Britain, Rabaul" M p Platymantis schmidti AMNH83088 (tototype) "E. New Britain, Talasea" M p Platymantis schmidti AMNH83089 (tototype) "E. New Britain, Talasea" M p Platymantis schmidti AMNH83091 (tototype) "E. New Britain, Talasea" F p Platymantis schmidti AMNH83092 (tototype) "E. New Britain, Talasea" F p Platymantis schmidti AMNH83093 (tototype) "E. New Britain, Talasea" F p Platymantis schmidti AMNH83094 (tototype) "E. New Britain, Talasea" M p Platymantis schmidti AMNH83095 (tototype) "E. New Britain, Talasea" M p Platymantis schmidti AMNH83098 (tototype) "E. New Britain, Talasea" M p Platymantis schmidti AMNH83101 (tototype) "E. New Britain, Talasea" M p Platymantis schmidti CAS139651 New Britain Id: Kerevat: Kereba plantation 4-Jan-69 F p Platymantis schmidti CAS139652 New Britain Id: Kerevat: Kereba plantation 4-Jan-69 F p Platymantis schmidti CAS22879 (paratype) "Bismarck Ids., New Britain Id., Talasea area, Willaumez 1-Jan-66 F p Peninsula" Platymantis schmidti CAS22880 (paratype) "Bismarck Ids., New Britain, Baining Ranges Gazelle 1-Jan-66 M p peninsula" Platymantis schmidti CAS22881 (paratype) "Bismarck Ids., New Britain, Baining Ranges Gazelle 1-Jan-66 M p peninsula" Platymantis schmidti CAS22882 (paratype) "Bismarck Ids., New Britain, Baining Ranges Gazelle 1-Jan-66 F p peninsula" Platymantis schmidti SAMR129601 F p Platymantis schmidti SAMR129602 F p Platymantis schmidti SAMR5650A "Rangoulit, Bairing Rgs, New Britain" 26-Jul-64 F p Platymantis schmidti SAMR5650B "Rangoulit, Bairing Rgs, New Britain" 26-Jul-64 M p Platymantis schmidti SAMR5650C "Rangoulit, Bairing Rgs, New Britain" 26-Jul-64 F p Platymantis schmidti SAMR6163 "Kerevat, New Britain" 1-May-65 F p Platymantis schmidti SAMR6447A "Kerevat, New Britain" 1-Jul-65 F p Platymantis schmidti SAMR6447B "Kerevat, New Britain" 1-Jul-65 S Platymantis schmidti SAMR7764 "Talasea, New Britain" 16-Jan-66 F p Platymantis schmidti SAMR7765 "Kerevat, New Britain" feb-mar 1966 F p Platymantis schmidti SAMR7766 "Walindi Ptn., Talasea, New Britain" 18-Jan-66 F p Platymantis schmidti SAMR7767 "Numondo Ptn., Willaumez Pen., New Britain" 19-Jan-66 F p Platymantis schmidti SAMR8023 "Kerevat, New Britain" 1-Apr-66 F p Platymantis schmidti SAMR8024 "Kerevat, New Britain" 1-Apr-66 F p Platymantis schmidti SAMR8025 "Kerevat, New Britain" 1-Apr-66 F p Platymantis schmidti SAMR8026 "Kerevat, New Britain" 1-Apr-66 F p Platymantis schmidti SAMR8027 "Kerevat, New Britain" 1-Apr-66 F p

260 Platymantis schmidti SAMR8028 "Kerevat, New Britain" 1-Apr-66 F p Platymantis schmidti SAMR8029 "Kerevat, New Britain" 1-Apr-66 F p Platymantis schmidti SAMR8030 "Kerevat, New Britain" 1-Apr-66 F p Platymantis schmidti SAMR8308A "Kerevat, New Britain" 13-Feb-66 F p Platymantis schmidti SAMR8308B "Kerevat, New Britain" 13-Feb-66 M p Platymantis schmidti SAMR8308C "Kerevat, New Britain" 13-Feb-66 S Platymantis solomonis A01/SAMR47136 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis solomonis A02/SAMR47137 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis solomonis A03/SAMR47138 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis solomonis A04/SAMR47139 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis solomonis A12/SAMR47140 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis solomonis A13/SAMR47141 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis solomonis A15/SAMR47142 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis solomonis A16/SAMR47143 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis solomonis A21/SAMR47144 "Pavora River, Choiseul" 6/3-12/3'90 F "m, p" Platymantis solomonis A59/SAMR37013 "Pavora River, Choiseul" 6/3-12/3'90 M "c, m, p" Platymantis solomonis A63/SAMR47135 "Pavora River, Choiseul" 6/3-12/3'90 F "m, p, o" Platymantis solomonis A69/SAMR47145 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis solomonis A70/SAMR47146 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis solomonis A73/SAMR47147 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis solomonis A74/SAMR47148 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis solomonis A77/SAMR47149 "Pavora River, Choiseul" 6/3-12/3'90 M "m, p" Platymantis solomonis A80/SAMR37012 "Pavora River, Choiseul" 6/3-12/3'90 F "c, m, p" Platymantis solomonis A91/SAMR47150 "Pavora River, Choiseul" 6/3-12/3'90 F "m, p" Platymantis solomonis AMR137087 (AMH 36491) "Mono Id., Shortland Ids., Solomon Islands" 14-Nov-90 M "m, p" Platymantis solomonis AMR137091 (AMH 36496) "Mt Moabe, Mono Id., Shortland Ids., Solomon Ids." 15-Nov-90 F "m, p" Platymantis solomonis AMR137095 (AMH 36500) "Mt Moabe, Mono Id., Shortland Ids., Solomon Ids." 15-Nov-90 F "m, p" Platymantis solomonis AMR137096 (AMH 36501) "Mt Moabe, Mono Id., Shortland Ids., Solomon Ids." 15-Nov-90 F "m, p" Platymantis solomonis AMR137103 (AMH 36509) "Mt Moabe, Mono Id., Shortland Ids., Solomon Ids." 15-Nov-90 F "m, p" Platymantis solomonis AMR137126 (AMH 36560) "Sulu Bay,Malaita Id., Solomon Ids." 15-Mar-91 F "m, p" Platymantis solomonis B06/R47134 "Pavora River, Choiseul" 6/3-12/3'90 F "m, p, o" Platymantis solomonis B69/SAMR47151 "Mt Austen, Gaudalcanal" 19-Mar-90 F "m, p" Platymantis solomonis B91/47152 "Mt Austen, Gaudalcanal" 19-Mar-90 F "m, p" Platymantis solomonis MCZ58549 "Turiboiru, Bougainville" 15-Mar-66 M "m, p" Platymantis solomonis MCZ58550 "Turiboiru, Bougainville" 15-Mar-66 S Platymantis solomonis MCZ58551 "Turiboiru, Bougainville" 15-Mar-66 M "m, p" Platymantis solomonis MCZ58552 "Turiboiru, Bougainville" 15-Mar-66 F "m, p" Platymantis solomonis MCZ58553 "Turiboiru, Bougainville" 15-Mar-66 F "m, p" Platymantis solomonis MCZ58554 "Turiboiru, Bougainville" 15-Mar-66 M "m, p" Platymantis solomonis MCZ58555 "Turiboiru, Bougainville" 15-Mar-66 F "m, p" Platymantis solomonis MCZ58556 "Turiboiru, Bougainville" 15-Mar-66 F "m, p, o" Platymantis solomonis MCZ58557 "Turiboiru, Bougainville" 15-Mar-66 S Platymantis solomonis MCZ58558 "Turiboiru, Bougainville" 15-Mar-66 S Platymantis solomonis MCZ58559 "Turiboiru, Bougainville" 15-Mar-66 F "m, p, o" Platymantis solomonis MCZ58560 "Turiboiru, Bougainville" 15-Mar-66 F "m, p" Platymantis solomonis MCZ58561 "Turiboiru, Bougainville" 15-Mar-66 S Platymantis solomonis MCZ58562 "Turiboiru, Bougainville" 15-Mar-66 F "m, p"

261 Platymantis solomonis MCZ58563 "Turiboiru, Bougainville" 15-Mar-66 M "m, p" Platymantis solomonis MCZ58564 "Turiboiru, Bougainville" 15-Mar-66 M "m, p" Platymantis solomonis MCZ58565 "Turiboiru, Bougainville" 15-Mar-66 F "m, p, o" Platymantis solomonis MCZ58566 "Turiboiru, Bougainville" 15-Mar-66 M "m, p" Platymantis solomonis MCZ58567 "Turiboiru, Bougainville" 15-Mar-66 F "m, p" Platymantis solomonis MCZ58568 "Turiboiru, Bougainville" 15-Mar-66 F "m, p, o" Platymantis solomonis MCZ58569 "Turiboiru, Bougainville" 15-Mar-66 F "m, p, o" Platymantis solomonis SAMR4413 "Kunua, Bougainville" 29-Jun-62 F "m, p" Platymantis solomonis SAMR4929 "Kunua, Bougainville" 29-Jun-62 F "m, p" Platymantis solomonis SAMR8051 "Turiboiru,, Bougainville" 14-Mar-66 F "m, p" Platymantis solomonis SAMR8052 "Turiboiru,, Bougainville" 14-Mar-66 M "m, p" Platymantis solomonis SAMR8053 "Turiboiru,, Bougainville" 14-Mar-66 M "m, p" Platymantis solomonis SAMR8054 "Turiboiru,, Bougainville" 14-Mar-66 M "m, p" Platymantis solomonis SAMR8060 "Turiboiru,, Bougainville" 10-Mar-66 F "m, p" Platymantis solomonis SAMR8061 "Turiboiru,, Bougainville" 10-Mar-66 F "m, p" Platymantis solomonis SAMR8190B "Mutahi, Bougainville" 11-May-66 M "m, p" Platymantis solomonis SAMR8191 "Turiboiru, Bougainville" 8-Mar-66 F "m, p" Platymantis solomonis SAMR8202 "Mutahi, Bougainville" 10-May-55 M "m, p" Platymantis solomonis SAMR8206 Malai/Shortlands 22-Apr-66 F "m, p" Platymantis sp. SAMR8125 "Kerevat, New Britain" 1-Jul-66 M Platymantis sp. SAMR8127 "Kerevat, New Britain" 1-Jul-66 M Platymantis spelaeus AMNH109454 (paratype) "Negros Id., Negros Oriental Prov., Basay, Tiyabanan F Banio" Platymantis spelaeus CAS153477 (paratype) "Philippine Ids., Negros Id., Negros Oriental: Basay, Actin 18-Mar-81 F Barrio" Platymantis spelaeus CAS153458 (paratype) "Philippine Ids., Negros Id., Negros Oriental: Basay, Actin 18-Mar-81 M Barrio" Platymantis spelaeus CAS153470 (paratype) "Philippine Ids., Negros Id., Negros Oriental: Basay, 18-Mar-81 M Tiyabanan Barrio" Platymantis spelaeus CAS153471 (paratype) "Philippine Ids., Negros Id., Negros Oriental: Basay, 18-Mar-81 M Tiyabanan Barrio" Platymantis spelaeus CAS153472 (paratype) "Philippine Ids., Negros Id., Negros Oriental: Basay, 18-Mar-81 M Tiyabanan Barrio" Platymantis spelaeus CAS153473 (paratype) "Philippine Ids., Negros Id., Negros Oriental: Basay, 18-Mar-81 S Tiyabanan Barrio" Platymantis spelaeus CAS153476 (paratype) "Philippine Ids., Negros Id., Negros Oriental: Basay, 18-Mar-81 S Tiyabanan Barrio" Platymantis spelaeus CAS153475 (paratype) "Philippine Ids., Negros Id., Negros Oriental: Basay, 18-Mar-81 M Tiyabanan Barrio" Platymantis spelaeus CAS153478 (paratype) "Philippine Ids., Negros Id., Negros Oriental: Basay, Actin 18-Mar-81 F Barrio" Platymantis spelaeus CAS153479 (paratype) "Philippine Ids., Negros Id., Negros Oriental: Basay, Actin 18-Mar-81 F Barrio" Platymantis spelaeus CAS153480 (paratype) "Philippine Ids., Negros Id., Negros Oriental: Basay, Actin 18-Mar-81 M Barrio" Platymantis spelaeus CAS153481 (paratype) "Philippine Ids., Negros Id., Negros Oriental: Basay, Actin 18-Mar-81 M Barrio"

262 Platymantis spelaeus CAS153482 (paratype) "Philippine Ids., Negros Id., Negros Oriental: Basay, Actin 18-Mar-81 F Barrio" Platymantis subterrestris FMNH142603 "Philippine Ids., Luzon Id., Camarines Sur Prov., Mt. Isaroo, 12-26 apr 1961 F Curry, Pili, 5600 ft" Platymantis subterrestris FMNH172392 "Philippine Ids., Luzon Id., Banane, Ifagao" F Platymantis subterrestris FMNH173165 "Philippine Ids., Luzon Id., Banane, Ifagao" F Platymantis vitianus CAS147574 "Fiji, Ovalau: Lovoni" 19-Feb-76 . Platymantis vitianus CAS159433 "Fiji Ids., Viwa Id.," 1-Jul-65 F Platymantis vitianus CAS172511 "Fiji Ids., Viwa Id., Viwa village" 5-Oct-88 F Platymantis vitianus CAS172512 "Fiji Ids., Viwa Id., Viwa village" 5-Oct-88 F Platymantis vitianus CAS172526 "Fiji, Ovalau Id., 0.5 mi N Navuloa village" 8-Oct-88 F Platymantis vitianus CAS172531 "Fiji, Ovalau Id., St John's college, 10 km N Levuka" 9-Oct-88 M Platymantis vitianus CAS172532 "Fiji, Ovalau Id., St John's college, 10 km N Levuka" 9-Oct-88 M Platymantis vitianus FMNH13634 "Fiji, Ovalau Id." 20-Mar-29 F Platymantis vitianus FMNH13635 "Fiji, Ovalau Id." 20-Mar-29 F Platymantis vitianus FMNH13637 "Fiji, Ovalau Id." 20-Mar-29 M Platymantis vitiensis FMNH23000 Fiji Id. M Platymantis vitiensis AMNH23614 "Viti Levu, Fiji" F Platymantis vitiensis CAS172436 "Fiji, Viti Levu, Savura Creek road" 26-Sep-88 M Platymantis vitiensis CAS172437 "Fiji, Viti Levu, Savura Creek road" 26-Sep-88 F Platymantis vitiensis CAS172438 "Fiji, Viti Levu, Savura Creek road" 26-Sep-88 M Platymantis vitiensis CAS172439 "Fiji, Viti Levu, Savura Creek road" 26-Sep-88 M Platymantis vitiensis CAS172440 "Fiji, Viti Levu, Savura Creek road" 26-Sep-88 F Platymantis vitiensis CAS172441 "Fiji, Viti Levu, Savura Creek road" 26-Sep-88 M Platymantis vitiensis CAS172442 "Fiji, Viti Levu, Savura Creek road" 26-Sep-88 M Platymantis vitiensis CAS172447 "Fiji, Viti Levu, Savura Creek road" 26-Sep-88 F Platymantis vitiensis CAS172449 "Fiji, Viti Levu, Savura Creek road" 26-Sep-88 M Platymantis vitiensis CAS172450 "Fiji, Viti Levu, Savura Creek road" 26-Sep-88 F Platymantis vitiensis CAS172451 "Fiji, Viti Levu, Savura Creek road" 26-Sep-88 M Platymantis vitiensis CAS172452 "Fiji, Viti Levu, Savura Creek road" 18-Dec-88 F Platymantis vitiensis CAS172453 "Fiji, Viti Levu, Savura Creek road" 18-Dec-88 M Platymantis vitiensis CAS172454 "Fiji, Viti Levu, Savura Creek road" 18-Dec-88 M Platymantis vitiensis CAS172455 "Fiji, Viti Levu, Savura Creek road" 18-Dec-88 M Platymantis vitiensis CAS172457 "Fiji, Viti Levu, Savura Creek road" 18-Dec-88 M Platymantis vitiensis CAS172456 "Fiji, Viti Levu, Savura Creek road" 18-Dec-88 M Platymantis vitiensis CAS172462 "Fiji, Viti Levu, Savura Creek road" 30-Dec-88 M Platymantis vitiensis CAS172614 "Fiji, Viti Levu, Naraiyawa village" 10-Dec-88 M Platymantis vitiensis SAMR9460 Fiji F Platymantis vitiensis SAMR9461 Fiji F Platymantis vitiensis SAMR9462 Fiji F Platymantis vitiensis SAMR9463 Fiji F Platymantis weberi A18/SAMR47160 "Pavora River, Choiseul" 4-21 mar 1990 F "m, p" Platymantis weberi A40/SAMR47161 "Pavora River, Choiseul" 4-21 mar 1990 F "m, p" Platymantis weberi A55/SAMR47153 "Pavora River, Choiseul" 4-21 mar 1990 F "m, p" Platymantis weberi A71/SAMR47162 "Pavora River, Choiseul" 4-21 mar 1990 F "m, p" Platymantis weberi A75/SAMR47163 "Pavora River, Choiseul" 4-21 mar 1990 F "m, p" Platymantis weberi A76/SAMR47164 "Pavora River, Choiseul" 4-21 mar 1990 F "m, p"

263 Platymantis weberi A78/SAMR47165 "Pavora River, Choiseul" 4-21 mar 1990 F "m, p" Platymantis weberi A85/SAMR47166 "Pavora River, Choiseul" 4-21 mar 1990 M "m, p" Platymantis weberi A92/SAMR47167 "Pavora River, Choiseul" 4-21 mar 1990 F "m, p" Platymantis weberi AMNH71905 "Bougainville I., Kunua, mts, 2000'" F "m, p" Platymantis weberi AMNH71906 "Bougainville I., Kunua, mts, 2000'" F "m, p" Platymantis weberi AMNH71907 "Bougainville I., Kunua, mts, 2000'" F "m, p" Platymantis weberi AMR137000 "Gold Ridge, Guadalcanal, Solomon Ids." 1-Apr-90 F "m, p" Platymantis weberi AMR137001 "Makarakomburu (S slope), Guadalcanal, Solomon Ids." 1-May-90 F "m, p" Platymantis weberi AMR137004 "Makarakomburu (S slope), Guadalcanal, Solomon Ids." 1-May-90 F "m, p" Platymantis weberi AMR137005 "Makarakomburu (S slope), Guadalcanal, Solomon Ids." 1-May-90 F "m, p" Platymantis weberi AMR137009 "Makarakomburu (S slope), Guadalcanal, Solomon Ids." 1-May-90 F "m, p" Platymantis weberi AMR137013 "Makarakomburu (S slope), Guadalcanal, Solomon Ids." 1-May-90 F "m, p" Platymantis weberi AMR137018 "Makarakomburu (S slope), Guadalcanal, Solomon Ids." 1-May-90 F "m, p" Platymantis weberi AMR137019 "Makarakomburu (S slope), Guadalcanal, Solomon Ids." 1-May-90 M "m, p" Platymantis weberi AMR137020 "Makarakomburu (S slope), Guadalcanal, Solomon Ids." 1-May-90 M "m, p" Platymantis weberi AMR137021 "Makarakomburu (S slope), Guadalcanal, Solomon Ids." 1-May-90 M "m, p" Platymantis weberi AMR137024 "Makarakomburu (S slope), Guadalcanal, Solomon Ids." 1-May-90 F "m, p" Platymantis weberi AMR137025 "Makarakomburu (S slope), Guadalcanal, Solomon Ids." 1-May-90 M "m, p" Platymantis weberi AMR137026 "Makarakomburu (S slope), Guadalcanal, Solomon Ids." 1-May-90 F "m, p" Platymantis weberi AMR137092 "Mt Moabe, Mono Id., Shortland Ids., Solomon Ids." 15-Nov-90 M "m, p" Platymantis weberi AMR137107 "Mt Moabe, Mono Id., Shortland Ids., Solomon Ids." 15-Nov-90 M "m, p" Platymantis weberi B03/SAMR37014 "Pavora River, Choiseul" 6/3-12/3'90 F "c, m, p" Platymantis weberi B90/SAMR47168 "Mt Austen, Gaudalcanal" 19-Mar-90 M "m, p" Platymantis weberi SAMR10888 "Rabaul, New Britain" 4-Jan-69 F "m, p" Platymantis weberi SAMR4410 "Kunua, Bougainville" 7-Jul-62 F "m, p" Platymantis weberi SAMR4411 "Kunua, Bougainville" 7-Jul-62 F "m, p" Platymantis weberi SAMR4412 "Kunua, Bougainville" 7-Jul-62 M "m, p, o" Platymantis weberi SAMR8212 "Mutahi, Bougainville" 13-May-66 F "m, p" Rana kreffti AMR137127 "Su'u Bay Malaita Id., Solomon Ids." 16-Mar-91 F p Rana kreffti AMR137128 "Su'u Bay Malaita Id., Solomon Ids." 18-Mar-91 F p Rana kreffti AMR137129 "Su'u Bay Malaita Id., Solomon Ids." 18-Mar-91 M p Rana kreffti AMR137171 "Bsurata village, Malaita Id., Solomon Ids., (08 49 S 160 49 23-Mar-91 F p E)" Rana kreffti CAS54666 "San Cristoval Id., Solomon Ids." F p Rana kreffti CAS72076 "Malaita Id., Solomon Ids." 30-May-33 F p Rana kreffti CAS72077 "Malaita Id., Solomon Ids." 30-May-33 M p Rana kreffti CAS72078 "Malaita Id., Solomon Ids." 30-May-33 M p Rana kreffti CAS72079 "Malaita Id., Solomon Ids." 30-May-33 F p Rana kreffti CAS72166 "Guadalcanar Id., Solomon Ids.," 21-May-33 F p Rana kreffti MCZ38079 "Kunua, Bougainville" 1962-63 M p Rana kreffti MCZ38080 "Kunua, Bougainville" 1962-63 M p Rana kreffti MCZ38081 "Kunua, Bougainville" 1962-63 M p Rana kreffti MCZ38082 "Kunua, Bougainville" 1962-63 M p Rana kreffti MCZ38083 "Kunua, Bougainville" 1962-63 F "p, o" Rana kreffti MCZ38084 "Kunua, Bougainville" 1962-63 F "p, o" Rana kreffti MCZ38085 "Kunua, Bougainville" 1962-63 M p Rana kreffti MCZ38086 "Kunua, Bougainville" 1962-63 F p

264 Rana kreffti MCZ38087 "Kunua, Bougainville" 1962-63 M "p, o" Rana kreffti MCZ38088 "Kunua, Bougainville" 1962-63 F p Rana kreffti MCZ38089 "Kunua, Bougainville" 1962-63 M "p, o" Rana kreffti MCZ38090 "Kunua, Bougainville" 1962-63 M p Rana kreffti MCZ38206 "Kunua, Bougainville" 1962-63 M p Rana kreffti MCZ38207 "Kunua, Bougainville" 1962-63 F "p, o" Rana kreffti MCZ38208 "Kunua, Bougainville" 1962-63 F p Rana kreffti MCZ38209 "Kunua, Bougainville" 1962-63 F p Rana kreffti MCZ38210 "Kunua, Bougainville" 1962-63 M p Rana kreffti MCZ38211 "Kunua, Bougainville" 1962-63 M p Rana kreffti MCZ38212 "Kunua, Bougainville" 1962-63 F p Rana kreffti MCZ38213 "Kunua, Bougainville" 1962-63 F p Rana kreffti SAMR16777 "Maraone, Santa Ana" 6-Mar-78 F p Rana kreffti SAMR4236 "Kunua, Bougainville" 1962-63 M p Rana kreffti SAMR4237 "Kunua, Bougainville" 28-May-62 M p Rana kreffti SAMR4238 "Kunua, Bougainville" 28-May-62 F p Rana kreffti SAMR4239 "Kunua, Bougainville" 28-May-62 M p Rana kreffti SAMR4240 "Kunua, Bougainville" 28-May-62 F "p, o" Rana kreffti SAMR4241 "Kunua, Bougainville" 28-May-62 M p Rana kreffti SAMR4921B "Kunua, Bougainville" 8-Mar-63 F p Rana kreffti SAMR5157 "Kunua, Bougainville" 7-Jun-63 F "p, o" Rana kreffti SAMR5161A "Kunua, Bougainville" 16-Jun-63 F "p, o" Rana kreffti SAMR5161B "Kunua, Bougainville" 16-Jun-63 F p Rana kreffti SAMR8055 "Turiboiru, Bougainville" 3-Mar-66 F p Rana kreffti SAMR8056 "Turiboiru, Bougainville" 3-Mar-66 F p Rana kreffti SAMR8179 "Turiboiru, Bougainville" 13-Mar-66 F p Rana kreffti SAMR8775A "Matsiogu, Bougainville" 30-Mar-66 F p Rana kreffti SAMR8775B "Matsiogu, Bougainville" 30-Mar-66 M p Rana kreffti UM60258(1) "Bio, Solomon Islands" 1923-24 M p Rana kreffti UM60258(2) "Bio, Solomon Islands" 1923-24 F p Rana kreffti (=novaebrittanae) AMNH64280 New Britain: south slope of Whiteman range S Rana kreffti (=novaebrittanae) AMNH69340 Buka Island M Rana kreffti (=novaebrittanae) AMNH69343 Kunua M Rana kreffti (=novaebrittanae) AMNH69344 Kunua M Rana kreffti (=novaebrittanae) AMNH69346 Kunua M Rana kreffti (=novaebrittanae) AMNH71924 Kunua M Rana kreffti (=novaebrittanae) AMNH71928 Kunua F Rana kreffti (=novaebrittanae) AMNH71932 Kunua F Rana kreffti (=novaebrittanae) AMNH71933 Kunua F Rana kreffti (=novaebrittanae) CAS106178 "Bismarck Archipelago: New Britain Id., Talasea" 1-Jun-63 M Rana kreffti (=novaebrittanae) CAS110830 "Solomon Islands, Bougainville Id., Mutahi, 1800-2700 ft" 17-May-66 F Rana kreffti (=novaebrittanae) CAS110831 "Solomon Islands, Bougainville Id., Mutahi, 1800-2700 ft" 17-May-66 M Rana kreffti (=novaebrittanae) CAS110832 "Solomon Islands, Bougainville Id., Mutahi, 1800-2700 ft" 17-May-66 F Rana kreffti (=novaebrittanae) CAS110833 "Solomon Islands, Bougainville Id., Mutahi, 1800-2700 ft" 17-May-66 F Rana kreffti (=novaebrittanae) CAS110834 "Solomon Islands, Bougainville Id., Mutahi, 1800-2700 ft" 13-May-66 F Rana kreffti (=novaebrittanae) CAS110836 "Solomon Islands, Bougainville Id., Mutahi, 1800-2700 ft" 19-May-66 F Amolops afghanus CAS172759 "Thailand, Doi Ithanon National Park, 1300 ft" 21 0ct 1988 F

265 Amolops jerboa CAS105983 "Bornea, Sarawak, 3rd div. Kapit Dist. Mengiong River S 11-Sep-63 F Serbong" Amolops jerboa CAS105984 "Bornea, Sarawak, 3rd div. Kapit Dist. Mengiong River S 1-Aug-67 F Serbong" Amolops jerboa CAS23022 "Kiau, Mt Kina Balu, N Bornea, 3000 ft" 1-Apr-29 M Amolops jerboa CAS7287 "Tanah Rata, Cameroon Highlands, Pahang, Malay 30-Oct-40 F Peninsula" Amolops jerboa CAS7288 "Tanah Rata, Cameroon Highlands, Pahang, Malay 30-Oct-40 F Peninsula" Amolops jerboa CAS7289 "Tanah Rata, Cameroon Highlands, Pahang, Malay 30-Oct-40 F Peninsula" Amolops larutensis CAS7274 "Perak, Malaysia" 25-Oct-40 M Amolops larutensis CAS7275 "Perak, Malaysia" 25-Oct-40 M Amolops larutensis CAS7280 "Cameroon Highlands, Ringlet, Pahang, Malaysia" 29-Oct-40 F Amolops larutensis CAS7281 "Cameroon Highlands, Ringlet, Pahang, Malaysia" 29-Oct-40 F Amolops larutensis CAS7282 "Cameroon Highlands, Ringlet, Pahang, Malaysia" 29-Oct-40 F Amolops larutensis CAS7283 "Cameroon Highlands, Ringlet, Pahang, Malaysia" 29-Oct-40 F Meristogenys (=Amolops) orphnocnemis FMNH235887 "Malaysia, Sabah, Tenom Dist., Sungai Malutut" 24-Nov-88 F p Meristogenys (=Amolops) orphnocnemis FMNH235889 "Malaysia, Sabah, Tenom Dist., Sungai Malutut" 24-Nov-88 F p Meristogenys (=Amolops) orphnocnemis FMNH235890 "Malaysia, Sabah, Tenom Dist., Sungai Malutut" 24-Nov-88 F p Meristogenys (=Amolops) orphnocnemis FMNH237304 "Malaysia, Sabah, Sipitang Dist., Mendolong" 26-Sep-89 F p Meristogenys (=Amolops) orphnocnemis FMNH237317 "Malaysia, Sabah, Tambunam Sunsuron, Mirad-Ahad" 15-Sep-89 F p Meristogenys (=Amolops) orphnocnemis FMNH237331 "Malaysia, Sabah, Sipitang Dist., Mendolong" 22-Sep-89 F p Meristogenys (=Amolops) orphnocnemis FMNH237337 "Malaysia, Sabah, Tambunam Sunsuron, Mirad-Ahad" 18-Sep-89 F p Meristogenys (=Amolops) orphnocnemis FMNH237354 "Malaysia, Sabah, Tambunam Sunsuron, Mirad-Ahad" 9-Sep-89 F p Meristogenys (=Amolops) orphnocnemis FMNH237362 "Malaysia, Sabah, Sipitang Dist., Mendolong" 22-Sep-89 F p Meristogenys (=Amolops) orphnocnemis FMNH235884 "Malaysia, Sabah, Tenom Dist., Sungai Malutut" 24-Nov-88 F p Meristogenys (=Amolops) whiteheadi CAS3272 "Kiau, Mt Kina Balu, N Bornea, 3000 ft" 1-Apr-29 F Meristogenys (=Amolops) whiteheadi CAS3274 "Kadamaian River, Mt. Kinabalu, N Borneo" . M Meristogenys (=Amolops) whiteheadi CAS3275 "Kadamaian River, Mt. Kinabalu, N Borneo" . F Ingerana (=Micrixalus) baluensis AMNH90504 "4th Div., Bintulu Dist., Tubua camp on Sungei Penisula, F p Sarawak" Ingerana (=Micrixalus) baluensis AMNH90505 "4th Div., Bintulu Dist., Tubua camp on Sungei Penisula, S Sarawak" Ingerana (=Micrixalus) baluensis FMNH223126 F p Ingerana (=Micrixalus) baluensis FMNH223127 M p Ingerana (=Micrixalus) baluensis FMNH223128 M p Ingerana (=Micrixalus) baluensis FMNH223135 M p Ingerana (=Micrixalus) baluensis FMNH223136 F p Ingerana (=Micrixalus) baluensis FMNH223137 F p Ingerana (=Micrixalus) baluensis FMNH223141 F p Ingerana (=Micrixalus) baluensis FMNH223142 F p Ingerana (=Micrixalus) baluensis FMNH223143 F p Ingerana (=Micrixalus) baluensis FMNH223145 M p Ingerana (=Micrixalus) baluensis FMNH238301 Ingerana (=Micrixalus) baluensis FMNH238307 Ingerana (=Micrixalus) baluensis FMNH223147 F p

266 Ingerana (=Micrixalus) baluensis FMNH235002 "Malaysia, Sabah, Sipitang Dist., Mendolong" 29-Nov-87 F p Ingerana (=Micrixalus) baluensis FMNH235603 F p Ingerana (=Micrixalus) baluensis FMNH235604 F p Ingerana (=Micrixalus) baluensis FMNH238311 "Malaysia, Sabah, Sipitang Dist., Mendolong" F p Ingerana (=Micrixalus) baluensis FMNH238315 "Malaysia, Sabah, Sipitang Dist., Mendolong" F p Ingerana (=Micrixalus) baluensis FMNH238317 "Malaysia, Sabah, Tenom Purulon, Kilampun" 12-Jul-89 F p Ingerana (=Micrixalus) baluensis FMNH240958 "Malaysia, Sabah, Danum Valley Field Centre" 19-Nov-89 F p Ingerana (=Micrixalus) baluensis FMNH244552 F p Ingerana (=Micrixalus) baluensis FMNH244848 "Malaysia, Sabah, Lahad Datu Dist., Danum Valley Field 15-Oct-90 F p Centre" Ingerana (=Micrixalus) baluensis FMNH244849 "Malaysia, Sabah, Lahad Datu Dist., Danum Valley Field 2-Nov-90 F p Centre" Ingerana (=Micrixalus) baluensis FMNH244850 "Malaysia, Sabah, Lahad Datu Dist., Danum Valley Field 15-Nov-90 F p Centre" Ingerana (=Micrixalus) baluensis FMNH244851 "Malaysia, Sabah, Lahad Datu Dist., Danum Valley Field 24-Nov-90 F p Centre" Micrixalus tenasserimensis AMNH97869 Salween River (17 57'N) Burma M Micrixalus tenasserimensis AMNH97870 Salween River (17 57'N) Burma F Micrixalus tenasserimensis AMNH97871 Salween River (17 57'N) Burma F Micrixalus tenasserimensis AMNH97872 Salween River (17 57'N) Burma M Micrixalus tenasserimensis AMNH97873 Salween River (17 57'N) Burma S Occidozyga laevis CAS16250 "Philippine Ids., Negros Id., Negros Oriental Prov., Ocoy 19-Oct-54 F p River Valley, 3 km W Palimpinon" Occidozyga laevis CAS16256 "Philippine Ids., Negros Id., Negros Oriental Prov., Ocoy 19-Oct-54 F p River Valley, 3 km W Palimpinon" Occidozyga laevis CAS16259 "Philippine Ids., Negros Id., Negros Oriental Prov., Ocoy 19-Oct-54 F p River Valley, 3 km W Palimpinon" Occidozyga laevis CAS16260 "Philippine Ids., Negros Id., Negros Oriental Prov., Ocoy 19-Oct-54 F p River Valley, 3 km W Palimpinon" Occidozyga laevis CAS16269 "Philippine Ids., Negros Id., Negros Oriental Prov., Ocoy 19-Oct-54 F p River Valley, 3 km W Palimpinon" Occidozyga laevis CAS16272 "Philippine Ids., Negros Id., Negros Oriental Prov., Ocoy 19-Oct-54 F p River Valley, 3 km W Palimpinon" Occidozyga laevis CAS16275 "Philippine Ids., Negros Id., Negros Oriental Prov., Ocoy 19-Oct-54 F p River Valley, 3 km W Palimpinon" Occidozyga laevis CAS16278 "Philippine Ids., Negros Id., Negros Oriental Prov., Ocoy 19-Oct-54 M p River Valley, 3 km W Palimpinon" Occidozyga laevis CAS16282 "Philippine Ids., Negros Id., Negros Oriental Prov., Ocoy 19-Oct-54 F p River Valley, 3 km W Palimpinon" Occidozyga laevis CAS16283 "Philippine Ids., Negros Id., Negros Oriental Prov., Ocoy 19-Oct-54 F p River Valley, 3 km W Palimpinon" Occidozyga laevis CAS16290 "Philippine Ids., Negros Id., Negros Oriental Prov., Ocoy F p River Valley, 3 km W Palimpinon" Occidozyga laevis CAS16279 "Philippine Ids., Negros Id., Negros Oriental Prov., Ocoy p River Valley, 3 km W Palimpinon" Occidozyga laevis AMNH72679 "Philippine Ids., Marinduque Id., Santa Cruz" 10-Aug-64 p Occidozyga laevis AMNH72680 "Philippine Ids., Marinduque Id., Santa Cruz" 10-Aug-64 p

267 Occidozyga laevis AMNH102937 "Philippine Ids., Marinduque Id., Santa Cruz" 10-Aug-64 p Occidozyga laevis AMNH102938 "Philippine Ids., Marinduque Id., Santa Cruz" 10-Aug-64 p Occidozyga laevis AMNH102939 "Philippine Ids., Marinduque Id., Santa Cruz" 10-Aug-64 p Occidzyga baluensis CAS105980 "Borneo, Sarawak, 4th div. Bintulu Dist., Sugnei Pesi" . . Occidzyga baluensis CAS105981 "Borneo, Sarawak, 4th div. Bintulu Dist., Sugnei Pesi" . . Rana arfarki SAMR9439 "Ining River, Soliabeda, New Guinea, S Highlands" 23-Sep-67 F Rana daemeli BG1 F Rana daemeli BG2 F Rana daemeli J791 . Rana daemeli J792 . Rana daemeli J804 . Rana daemeli J805 . Rana daemeli NHMV16282.2 M Rana daemeli NHMV16283.1 M Rana daemeli R121929 F Rana daemeli R121930 . Rana daemeli R121931 F Rana daemeli R122006 F Rana daemeli R122007 F Rana daemeli R122008 F Rana daemeli R122009 F Rana daemeli R122010 F Rana daemeli R122011 F Rana daemeli R122012 F Rana daemeli R122013 F Rana daemeli R122014 F Rana daemeli R122015 F Rana daemeli R129518 M Rana daemeli R129519 M Rana daemeli R129554 M Rana daemeli R129555 M Rana daemeli R129556 M Rana daemeli R129569 S Rana daemeli R129570 S Rana daemeli R129571 F Rana daemeli R129572 F Rana daemeli R129576 F Rana daemeli R129584 F Rana daemeli R129590 M Rana daemeli R129591 M Rana daemeli R129592 M Rana daemeli R129593 M Rana daemeli R129594 M Rana daemeli R129595 M Rana daemeli R129596 F Rana daemeli R129597 M Rana daemeli R129599 F

268 Rana daemeli R129600 M Rana daemeli R129603 F Rana daemeli R129604 M Rana daemeli R129605 M Rana daemeli R129619 S Rana daemeli R129631 M Rana daemeli R129639? . Rana daemeli R129690 M Rana daemeli R129691 S Rana daemeli R35091 . Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 F p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 M p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 M p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 F p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 M p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 M p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 F p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 M p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 F p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 F p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 F p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 F p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 F p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 F p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 M p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 F p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 M p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 M p Rana grisea SAMR5213 "Okapa, New Guinea" 6-Nov-63 M p Rana grisea SAMR5408A "Aiyura, E. New Guinea" 24-Oct-59 F p Rana grisea SAMR5408B "Aiyura, E. New Guinea" 24-Oct-59 F p Rana grisea SAMR5687A "Yaveyuta, W. New Guinea" 7-Jun-64 F p Rana grisea SAMR5687B "Yaveyuta, W. New Guinea" 7-Jun-64 M p Rana grisea SAMR5687C "Yaveyuta, W. New Guinea" 7-Jun-64 M p Rana grisea SAMR5687D "Yaveyuta, W. New Guinea" 7-Jun-64 M p Rana grisea SAMR5687E "Yaveyuta, W. New Guinea" 7-Jun-64 M p Rana grisea SAMR5687F "Yaveyuta, W. New Guinea" 7-Jun-64 M p Rana grisea SAMR5687G "Yaveyuta, W. New Guinea" 7-Jun-64 S Rana grisea SAMR5694A "Nivi, Yandime, W. New Guinea" 21-Jun-64 F p Rana grisea SAMR5694B "Nivi, Yandime, W. New Guinea" 21-Jun-64 F p Rana grisea SAMR5694C "Nivi, Yandime, W. New Guinea" 21-Jun-64 M p Rana grisea SAMR5694D "Nivi, Yandime, W. New Guinea" 21-Jun-64 F p Rana grisea SAMR5694E "Nivi, Yandime, W. New Guinea" 21-Jun-64 S Rana grisea SAMR5694F "Nivi, Yandime, W. New Guinea" 21-Jun-64 S Rana grisea SAMR5694G "Nivi, Yandime, W. New Guinea" 21-Jun-64 S Rana grisea SAMR5838A "Suwaira, W. New Guinea" 10-Aug-64 M p Rana grisea SAMR5838B "Suwaira, W. New Guinea" 10-Aug-64 F p

269 Rana grisea SAMR6165A "Koge, W. New Guinea" 28-Apr-65 F p Rana grisea SAMR6165B "Koge, W. New Guinea" 28-Apr-65 F p Rana grisea SAMR6178A "Wahgi R., (Ninmul) W. New Guinea" 8 - 10-Apr-65 F p Rana grisea SAMR6178B "Wahgi R., (Ninmul) W. New Guinea" 8 - 10-Apr-65 F p Rana grisea SAMR6178C "Wahgi R., (Ninmul) W. New Guinea" 8 - 10-Apr-65 M p Rana grisea SAMR8296A "Mt Michael Lufa, New Guinea" 15-Mar-66 F p Rana grisea SAMR8296B "Mt Michael Lufa, New Guinea" 15-Mar-66 F p Rana grisea SAMR8296C "Mt Michael Lufa, New Guinea" 15-Mar-66 F p Limnonectes (=Rana) magnus (=magna) CAS23405 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, 10 km SE 30-Apr-62 F p Sierra Bullones, 1300 ft" Limnonectes (=Rana) magnus (=magna) CAS23406 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, 10 km SE 30-Apr-62 F p Sierra Bullones, 1300 ft" Limnonectes (=Rana) magnus (=magna) CAS23407 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, 10 km SE 30-Apr-62 F p Sierra Bullones, 1300 ft" Limnonectes (=Rana) magnus (=magna) CAS23408 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, 10 km SE 30-Apr-62 F p Sierra Bullones, 1300 ft" Limnonectes (=Rana) magnus (=magna) CAS23409 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, 10 km SE 30-Apr-62 F p Sierra Bullones, 1300 ft" Limnonectes (=Rana) magnus (=magna) CAS23410 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, 10 km SE 30-Apr-62 F p Sierra Bullones, 1300 ft" Limnonectes (=Rana) magnus (=magna) CAS23411 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, 10 km SE 30-Apr-62 M p Sierra Bullones, 1300 ft" Limnonectes (=Rana) magnus (=magna) CAS23421 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, 10 km SE 30-Apr-62 M p Sierra Bullones, 1300 ft" Limnonectes (=Rana) magnus (=magna) CAS23423 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, 10 km SE 30-Apr-62 F p Sierra Bullones, 1300 ft" Limnonectes (=Rana) magnus (=magna) CAS23427 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, 10 km SE 30-Apr-62 F p Sierra Bullones, 1300 ft" Limnonectes (=Rana) magnus (=magna) CAS23428 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, 10 km SE 30-Apr-62 M p Sierra Bullones, 1300 ft" Limnonectes (=Rana) magnus (=magna) CAS23431 "Philippine Ids., Bohol Id., Bohol Prov., Dusita, 10 km SE 30-Apr-62 F p Sierra Bullones, 1300 ft" Limnonectes (=Rana) magnus (=magna) AMNH69972 "Philippine Ids., Mindanao, Cotabato, Mt. Tatatum, 5100 ft" 7-Mar-62 p Limnonectes (=Rana) magnus (=magna) AMNH69973 "Philippine Ids., Mindanao, Cotabato, Mt. Tatatum, 5100 ft" 7-Mar-62 p Limnonectes (=Rana) magnus (=magna) AMNH52613 "Umingan, Pangasinan, Luzon, Philippine Id." M p Limnonectes (=Rana) magnus (=magna) AMNH68176 "Philippines, Luzon, Isabela Prov. San Mariano Bo. M p Disusuan" Limnonectes (=Rana) magnus (=magna) AMNH68179 "Philippines, Luzon, Isabela Prov. San Mariano Bo. F p Disulap" Limnonectes (=Rana) magnus (=magna) AMNH68180 "Philippines, Luzon, Isabela Prov. San Mariano Bo. F p Disulap" Limnonectes (=Rana) magnus (=magna) AMNH68181 "Philippines, Luzon, Isabela Prov. San Mariano Bo. F p Disulap" Limnonectes (=Rana) magnus (=magna) AMNH68183 "Philippines, Luzon, Isabela Prov. San Mariano Bo. F p Dimalasud" Rana papua AMNH98992 "PNG, East Sepik Prov., Maprik" 20-Nov-76 F Rana papua AMNH98993 "PNG, East Sepik Prov., Maprik" 20-Nov-76 M

270 Rana sanguinea FMNH51287 . Rana sanguinea FMNH51288 F Rana temporaria SAMR1654 "Butzow, Germany" 21-Mar-31 M p Rana temporaria SAMR1655 "Butzow, Germany" 4-Apr-31 F p Rana temporaria SAMR1656A "Butzow, Germany" 7-Apr-31 F p Rana temporaria SAMR1656B "Butzow, Germany" 7-Apr-31 F p Rana temporaria SAMR1656C "Butzow, Germany" 7-Apr-31 F p Rana temporaria SAMR1657(10) "Butzow, Germany" 4-Apr -7-Apr -31 M p Rana temporaria SAMR1657(11) "Butzow, Germany" 4-Apr -7-Apr -31 M p Rana temporaria SAMR1657(12) "Butzow, Germany" 4-Apr -7-Apr -31 M p Rana temporaria SAMR1657(13) "Butzow, Germany" 4-Apr -7-Apr -31 M p Rana temporaria SAMR1657(14) "Butzow, Germany" 4-Apr -7-Apr -31 M p Rana temporaria SAMR1657(3) "Butzow, Germany" 4-Apr -7-Apr -31 M p Rana temporaria SAMR1657(4) "Butzow, Germany" 4-Apr -7-Apr -31 M p Rana temporaria SAMR1657(5) "Butzow, Germany" 4-Apr -7-Apr -31 M p Rana temporaria SAMR1657(7) "Butzow, Germany" 4-Apr -7-Apr -31 M p Rana temporaria SAMR43145 M p Rana temporaria SAMR43146 F p Staurois natator CAS20036 F p Staurois natator CAS19989 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 M p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator CAS19990 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 F p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator CAS19992 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 F p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator CAS19993 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 F p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator CAS19994 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 F p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator CAS19995 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 M p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator CAS20004 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 F p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator CAS20029 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 M p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator CAS20031 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 F p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator CAS20033 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 F p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator CAS20041 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 F p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator CAS20051 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 F p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator CAS23364 "Philippine Ids., Bohol Id., Bohol Prov., Dusita area, 11 km 17-Apr-62 p SE Sierra Bullones" Staurois natator AMNH70008 "Mindanao, Cotabatc Prov., Mt. Matatum, 5000', 7-Mar-62 M p Philippines"

271 Staurois natator AMNH70009 "Mindanao, Cotabatc Prov., Mt. Matatum, 5000', 7-Mar-62 M p Philippines" Staurois natator AMNH70011 "Palawan, Aborlan, Philippines" 1-May-62 p Staurois natator AMNH70012 "Palawan, Aborlan, Philippines" 1-May-62 F p Staurois natator FMNH196600 F p Staurois natator FMNH196614 F p Staurois natator CAS20037 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 F p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator CAS20044 "Philippine Ids., Mindanao Id., Zamboanga del Norte Prov., 30-Apr-59 F p Dapitan River area, 1 km SE Gumay, 2300 ft" Staurois natator FMNH234689 F p Staurois natator FMNH234690 F p Staurois natator FMNH234691 F p Staurois natator FMNH234692 F p Staurois natator FMNH234695 F p Staurois natator FMNH234696 M p Staurois natator FMNH234697 M p Staurois natator FMNH234699 M p Staurois natator FMNH234701 M p Staurois natator FMNH234702 M p Staurois natator FMNH234704 M p Staurois natator FMNH234707 M p Staurois natator FMNH234708 F p Staurois natator FMNH234710 M p Staurois natator FMNH234711 M p Staurois natator SAMR13600A "Philippine, Mindanao Ids, Zamboanga del Norte Prov" 30-Apr-59 M p Staurois natator SAMR13600B "Philippine, Mindanao Ids, Zamboanga del Norte Prov" 30-Apr-59 F p

272 APPENDIX 2. Discriminant Function Analysis

DISCRIMINANT ANALYSIS based on 14 Principal Components generated from six species of Batrachylodes.

Stepwise variable selection Selection rule: minimize Wilks' Lambda

Maximum number of steps ...... 26 Minimum tolerance level ...... 0.001 Minimum F to enter ...... 3.84 Maximum F to remove ...... 2.71

Canonical Discriminant Functions

Maximum number of functions ...... 5 Minimum cumulative percent of variance ...... 100.00 Maximum significance of Wilks' Lambda ...... 1.00

Prior probability for each group is 0.17

Table A2.1 Standardized canonical discriminant function coefficients for Batrachylodes Func 1 Func 2 Func 3 Func 4 Func 5

PRIN1 -1.56 -.17 .18 .04 .12 PRIN10 -.11 .48 .31 -.07 -.32 PRIN14 -.11 -.35 -.23 -.42 .32 PRIN2 1.08 .07 -.20 .32 .44 PRIN3 .48 -.94 -.08 .12 -.30 PRIN4 .90 -.13 .84 -.16 .21 PRIN5 .06 .24 .27 .29 -.39 PRIN6 .02 .30 -.09 .43 .46 PRIN7 -.11 .55 -.02 -.23 -.19 PRIN8 -.20 .12 -.00 .58 -.17 PRIN9 .00 .22 -.29 -.56 -.05

Table A2.2 Pooled within-groups correlations between discriminating variables and canonical discriminant functions (Variables ordered by size of correlation within function) for six species of Batrachylodes Func 1 Func 2 Func 3 Func 4 Func 5

PRIN1 -.47* -.24 .38 .10 .37 PRIN3 .07 -.62* -.08 .14 -.43 PRIN7 -.01 .23* -.01 -.16 -.17 PRIN13 -.09 -.15* -.07 .05 -.04 PRIN4 .13 -.10 .85* -.20 .31 PRIN9 .00 .10 -.18 -.43* -.05 PRIN8 -.02 .05 -.00 .43* -.15 PRIN14 -.01 -.15 -.14 -.32* .29 PRIN12 -.07 -.11 -.07 .14* -.05 PRIN2 .10 .03 -.13 .26 .43* PRIN6 .00 .13 -.06 .33 .42* PRIN5 .01 .10 .16 .22 -.35* PRIN10 -.01 .20 .19 -.06 -.29* * denotes largest absolute correlation between each variable and any discriminant function.

273 Table A2.3 Canonical discriminant functions evaluated at group means (group centroids) for six species of Batrachylodes Group Func 1 Func 2 Func 3 Func 4 Func 5

elegans -2.56 -.58 1.81 .22 -.27 mediodiscus 2.63 -.63 -.03 1.47 1.03 minutus 6.75 .76 -.71 1.42 -1.45 trossulus 6.22 -.95 .30 -1.06 .11 vertebralis -1.10 1.34 -.24 -.20 .09 wolfi -3.31 -1.92 -1.15 -.08 -.17

DISCRIMINANT ANALYSIS based on 14 Principal Components generated from three species of Discodeles.

Stepwise variable selection Selection rule: minimize Wilks' Lambda

Maximum number of steps ...... 28 Minimum tolerance level ...... 0.001 Minimum F to enter ...... 3.84 Maximum F to remove ...... 2.71

Canonical Discriminant Functions

Maximum number of functions ...... 2 Minimum cumulative percent of variance ...... 100.00 Maximum significance of Wilks' Lambda ...... 1.00

Prior probability for each group is .33

Table A2.4 Standardized canonical discriminant function coefficients for Discodeles Func 1 Func 2

PRIN2 1.06 .18 PRIN3 -.54 .86 PRIN7 -.33 -.51

274 Table A2.5 Pooled within-groups correlations between discriminating variables and canonical discriminant functions (Variables ordered by size of correlation within function) for three species of Discodeles Func 1 Func 2

PRIN2 .80* .32 PRIN13 .21* -.04 PRIN4 .17* -.12 PRIN12 -.14* -.04 PRIN11 .14* .03 PRIN1 -.12* -.03 PRIN9 .11* .03 PRIN5 -.08* .06 PRIN10 -.04* .01 PRIN3 -.22 .87* PRIN7 -.10 -.37* PRIN6 -.13 .14* PRIN8 .07 .11* PRIN14 -.00 .11* * denotes largest absolute correlation between each variable and any discriminant function.

DISCRIMINANT ANALYSIS based on 14 Principal Components generated from eight species of Platymantis.

Stepwise variable selection Selection rule: minimize Wilks' Lambda

Maximum number of steps ...... 28 Minimum tolerance level ...... 0.001 Minimum F to enter ...... 3.84 Maximum F to remove ...... 2.71

Canonical Discriminant Functions

Maximum number of functions ...... 7 Minimum cumulative percent of variance ...... 100.00 Maximum significance of Wilks' Lambda ...... 1.00

Prior probability for each group is .125

275 Table A2.6 Standardized canonical discriminant function coefficients for eight species of Platymantis Func 1 Func 2 Func 3 Func 4 Func 5 Func 6 Func 7

PRIN1 1.49 .45 -.16 .02 -.04 -.05 .08 PRIN11 .40 .39 -.11 .16 .05 -.02 -.14 PRIN12 -.09 .06 -.16 -.06 -.65 -.07 -.12 PRIN13 -.23 .10 -.04 .25 .09 -.25 .53 PRIN14 .13 .08 -.23 -.13 .23 .40 .34 PRIN2 .51 -1.07 .00 .04 -.01 .03 -.01 PRIN3 .80 .16 .88 -.41 -.07 -.10 -.10 PRIN4 .09 .38 .33 .86 -.32 .14 -.13 PRIN5 .42 -.15 -.31 .38 .38 -.28 -.43 PRIN6 -.30 -.34 .56 .39 .13 -.43 .46 PRIN7 .27 .11 .34 .14 .03 .75 .19 PRIN8 .33 .30 .05 .03 .50 .10 -.06 PRIN9 .46 .37 .43 .23 .48 .09 -.26

Table A2.7 Pooled within-groups correlations between discriminating variables and canonical discriminant functions (Variables ordered by size of correlation within function) for Platymantis Func 1 Func 2 Func 3 Func 4 Func 5 Func 6 Func 7

PRIN1 .50* .22 -.35 .05 -.20 -.23 .48 PRIN2 .25 -.78* .01 .18 -.08 .22 -.05 PRIN3 .09 .03 .65* -.41 -.11 -.16 -.20 PRIN10 .13 .11 .16* -.03 -.08 .06 .05 PRIN4 .01 .05 .19 .66* -.38 .18 -.19 PRIN12 -.01 .01 -.06 -.03 -.50* -.06 -.11 PRIN9 .03 .03 .16 .12 .37* .07 -.26 PRIN8 .02 .02 .01 .02 .36* .08 -.05 PRIN7 .02 .01 .13 .08 .03 .67* .20 PRIN14 .01 .01 -.01 -.06 .17 .32* .31 PRIN6 -.02 -.03 .24 .23 .11 -.42 .52* PRIN13 -.01 .01 -.01 .12 .07 -.20 .48* PRIN5 .02 -.01 -.11 .20 .30 -.24 -.43* PRIN11 .02 .03 -.04 .07 .03 -.02 -.12* * denotes largest absolute correlation between each variable and any discriminant function.

Table A2.8 Canonical discriminant functions evaluated at group means (group centroids) for eight species of Platymantis Group Func 1 Func 2 Func 3 Func 4 Func 5 Func6 Func7 acrochordus -3.94 -1.56 -.83 -2.41 -3.44 -2.50 -.20 aculeodactylus -5.59 -1.59 -2.11 1.83 -.35 .31 -.03 guppyi 6.63 -5.34 -1.84 -.94 .35 .05 .02 myersi 1.16 2.88 .51 -1.10 -1.10 1.31 .62 neckeri 4.51 -4.07 2.89 1.89 -.39 -.23 .17 parkeri -7.90 -4.05 1.53 -1.31 .62 .10 -.03 solomonis .22 3.49 -.17 .07 .47 -.45 .30 weberi 1.54 2.75 .40 -.09 -.05 .22 -.83

276 Batrachylodes – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.9 3.2 2.6 2.9 2.9 1.8 1.50 1.5 1.0 1.8 2.5 2.8 3.5 0.50

3.8 3.1 2.5 2.8 2.8 1.7 1.4 0.9 1.0 2.7 3.0 0.5 3.7 2.7 1.6 1.25 1.6 0.25 3.0 2.4 2.7 1.3 0.8 2.6 3.6 2.6 1.5 2.0 2.5 2.9 2.6 0.7 0.5 2.3 1.2 2.5 3.5 2.5 1.4 1.00 1.4 0.00 2.8 2.5 0.6 2.0 0.0 3.4 2.2 2.4 1.3 1.1 2.4 2.7 2.4 0.5 0.0 3.3 2.1 2.3 1.2 0.75 1.0 1.2 1.5 2.3 1.5 -0.25 2.6 2.3 0.4 3.2 2.0 2.2 1.1 0.9 2.2 2.5 2.2 0.3 -0.5 1.0 -0.5 3.1 2.1 1.0 0.50 1.0 -0.50 1.9 0.8 2.1 3.0 2.4 2.1 2.0 0.9 0.2 1.0 0.5 1.8 0.7 2.0 2.9 2.3 2.0 1.9 0.8 0.25 0.1 0.8 -1.0 -0.75 0.0 -1.0 2.8 2.2 1.7 1.9 1.8 0.7 0.6 0.0 1.9 2.7 2.1 1.6 1.8 1.7 0.6 0.00 0.5 -0.1 0.6 0.5 -1.5 1.8 -0.5 -1.00 median median median median median median median median median median median median median median median - median - 3.1553 2.4527 1.9803 2.1541 2.1547 1.1490 0.7952 0.8961 0.43823 0.9914 1.7317 0.2390 2.1552 2.3906 0.0834 0.0943 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean - 3.16243 2.48019 1.99404 2.16930 2.16483 1.12412 0.73718 0.89060 0.43067 1.01159 1.71095 0.04423 2.16137 2.37202 0.09435 0.10773 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.22011 0.20321 0.16852 0.23205 0.25577 0.23230 0.27105 0.18006 0.21843 0.18323 0.29592 0.60174 0.17657 0.39400 0.34478 0.31282

Ceratobatrachus – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.4 3.6 3.1 3.7 3.8 2.3 2.4 1.8 2.25 2.4 3.1 0.5 3.3 3.7 0.7 0.7

3.0 0.4 0.6 0.6 3.6 3.7 2.2 2.3 1.7 2.2 3.0 3.2 3.6 4.3 3.5 2.00 0.3 0.5 2.9 0.5 3.5 3.6 2.1 2.2 1.6 2.0 2.9 3.1 3.5 0.2 0.4 4.2 3.4 1.75 2.8 0.4 3.4 3.5 2.0 2.1 1.5 1.8 2.8 0.1 3.0 3.4 0.3 2.7 0.3 4.1 3.3 1.50 -0.0 0.2 3.3 3.4 1.9 2.0 1.4 1.6 2.7 2.9 3.3 2.6 0.2 -0.1 0.1 4.0 3.2 1.25 3.2 3.3 1.8 1.9 1.3 1.4 2.6 2.8 3.2 2.5 -0.2 0.0 0.1 3.9 3.1 2.4 3.1 3.2 1.7 1.8 1.2 1.00 1.2 2.5 -0.3 2.7 3.1 -0.1 0.0 median median median median median median median median median median median median median median median median 4.2485 3.4920 2.8987 3.5041 3.6085 2.0360 2.2176 1.5507 1.4702 1.7155 2.8633 0.20293 3.1385 3.4690 0.39541 0.48841 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 4.22380 3.44936 2.84989 3.46234 3.54985 2.00411 2.18042 1.52874 1.49321 1.78038 2.83484 0.20902 3.10011 3.45149 0.36712 0.43526 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.13570 0.12767 0.15835 0.13483 0.14660 0.15112 0.12330 0.13518 0.21718 0.22824 0.14577 0.17510 0.14915 0.14227 0.18426 0.17951

277 Discodeles – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

5.5 4.4 4.5 3.2 2.2 4.0 4.5

3.0 1.5 4.5 4.0 4.2 3.0 3.0 2.5 2.0 4.5 1.5 1.5

5.0 4.0 1.8 4.0 4.0 2.8 3.5 2.5 1.0 4.0 3.5 3.8 1.6 4.0 1.0 2.6 2.5 2.0 1.0 4.5 3.6 1.4 3.5 3.5 2.4 2.0 3.0 0.5 3.5 3.0 3.4 1.2 3.5 0.5 2.2 2.0 1.5 0.5 4.0 3.2 1.0 3.0 1.5 0.0

3.0 2.5 3.0 3.0 2.0 0.8 2.5 3.0 0.0 3.5 2.8 1.8 1.5 1.0 0.6 1.0 -0.5 2.5 0.0 median median median median median median median median median median median median median median median median 4.7337 3.9152 3.1130 3.8214 3.9192 2.6027 2.1798 1.7596 1.6094 2.0605 3.0910 0.5563 3.4657 3.7612 0.7792 0.7724 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 4.57764 3.85805 3.19701 3.65230 3.74537 2.52380 2.29055 1.90901 1.56722 2.09865 3.10774 0.62126 3.53599 3.80585 0.82169 0.80381 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.49029 0.52187 0.52609 0.43111 0.48874 0.35407 0.44385 0.46018 0.36762 0.61066 0.43724 0.51310 0.45889 0.46551 0.39236 0.41104

Palmatorappia – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.50 2.90 2.45 2.425 2.55 1.6 1.4 1.1 0.8 1.4 2.2 1.0 2.45 2.75 0.75 1.00

2.85 2.70 2.40 2.400 1.5 2.40 3.45 1.3 1.0 0.7 2.1 0.75 2.50 1.2 0.50 2.80 0.5 2.65 2.35 2.375 1.4 2.35 3.40 1.2 0.9 0.6 2.0 0.50 2.75 2.60 2.30 2.350 2.45 1.3 1.0 2.30 0.25 2.70 0.0 2.55 3.35 1.1 0.8 0.5 1.9 0.25 2.25 2.325 1.2 2.25 2.65 2.50 2.40 0.8 0.00 3.30 1.0 0.7 0.4 1.8 0.00 2.20 2.300 1.1 2.20 2.60 -0.5 2.45 3.25 2.55 2.15 2.275 2.35 1.0 0.9 0.6 0.3 0.6 1.7 2.15 2.40 -0.25 -0.25 median median median median median median median median median median median median median median median median 3.3690 2.7948 2.3321 2.3627 2.4891 1.3762 1.2149 0.9708 0.43825 1.2920 2.1187 0.73716 2.2565 2.5494 0.50682 0.74194 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.37814 2.77658 2.30909 2.35860 2.46642 1.31816 1.21387 0.92230 0.46977 1.24525 2.05586 0.59366 2.27671 2.55715 0.40961 0.62843 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.059105 0.082221 0.074616 0.041941 0.059899 0.152484 0.123513 0.141187 0.117101 0.189770 0.123685 0.438179 0.072454 0.092951 0.249514 0.264745

278 Platymantis – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4 3.5 4 2.0 3.5 4 2.0 2.0 1.5 2.5 2 1.5 3.0 1.5 3.0 2 1.0 2.0 1.0 2 1.0 3 4 3 2.5 0.5 2.5 3 1.0 3 0.5 3 1.5

2.0 0.0 0.0 0.0 2.0 1 0.5 1.0 1 -0.5 -0.5 1.5 -1.0 2 3 1.5 2 1 0.5 0.0 -1.0 2 2 -1.0 2 1.0 -1.5 1.0 0.0 -0.5 -2.0 -1.5 median median median median median median median median median median median median median median median median 3.7693 3.1942 2.6487 2.8565 2.8588 1.7842 1.6074 1.3403 1.0852 1.3558 2.4493 0.1398 2.8217 3.1561 0.1740 0.2624 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.7236 3.1176 2.5897 2.8003 2.8261 1.7302 1.5191 1.2967 1.0211 1.3690 2.3834 0.1722 2.7663 3.0975 0.1645 0.2389 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.3727 0.3780 0.3605 0.3756 0.3850 0.3556 0.4437 0.3279 0.4118 0.3644 0.4315 0.6581 0.3888 0.3879 0.4642 0.4843

Rana – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.5 4.0 3.4 3.6 3.5 2.4 2.3 2.1 2.1 2.2 3.4 1.0 3.6 3.9 1.0 1.0

4.4 3.9 3.3 3.5 3.4 2.2 2.0 3.5 3.8 0.9 2.3 2.0 3.2 0.8 3.8 2.0 0.8 3.4 0.8 4.3 3.2 3.4 3.3 2.1 1.9 3.7 2.2 1.9 3.7 3.3 0.7 3.1 3.2 2.0 3.0 3.6 0.6 4.2 3.3 1.8 1.8 0.6 3.6 2.1 1.8 3.2 0.6 3.0 3.1 1.9 3.5 2.8 0.4 4.1 3.5 3.2 1.7 3.1 0.5 2.9 3.0 2.0 1.8 1.7 1.6 0.4 3.4 3.4 3.0 0.4 4.0 3.1 1.6 2.6 0.2 2.8 2.9 1.7 3.3 3.3 1.9 1.6 2.9 0.3 3.9 3.0 1.5 1.4 0.2 2.7 2.8 1.6 2.4 3.2 0.0 3.2 1.8 1.5 2.8 0.2 3.8 2.6 2.9 2.7 1.5 1.4 3.1 3.1 1.2 0.0 2.7 0.1 1.7 1.4 2.2 -0.2 3.7 3.0 2.5 2.8 2.6 1.4 1.3 2.6 3.0 0.0 3.6 2.9 2.4 2.7 2.5 1.6 1.3 1.2 1.3 1.0 2.0 -0.2 2.5 2.9 -0.1 -0.4 median median median median median median median median median median median median median median median median 3.9190 3.3109 2.6803 3.0084 2.8622 1.8679 1.6084 1.6094 1.6074 1.4586 2.6373 0.30010 2.9143 3.2498 0.40544 0.40212 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.97615 3.35533 2.73025 3.06030 2.91280 1.93437 1.64393 1.62676 1.61252 1.48459 2.67060 0.34865 2.95042 3.28376 0.43448 0.41341 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.21016 0.18665 0.19490 0.18834 0.21538 0.19096 0.19541 0.17576 0.14371 0.21088 0.22514 0.26739 0.21450 0.19612 0.22820 0.26002

279 Batrachylodes – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.5 2.6 2.6 1.5 1.2 0.9 1.5 2.3 2.5 2.8 0.3 0.4 2.7 2.3 1.2 1.1 0.8 2.2 0.2 0.3 3.4 2.5 2.5 1.4 1.4 0.5 2.4 2.7 2.1 1.0 0.7 0.1 0.2 2.6 2.2 2.4 2.4 1.3 1.1 1.3 2.0 3.3 0.9 2.3 2.6 -0.0 0.1 0.6 2.3 2.3 1.2 1.2 1.9 0.0 2.5 2.1 0.8 -0.1 -0.0 3.2 1.0 2.2 2.5 0.5 1.8 2.2 2.2 1.1 0.7 1.1 -0.2 -0.1 2.4 2.0 1.7 3.1 0.4 2.1 2.4 0.6 0.9 -0.3 -0.2 2.1 2.1 1.0 1.0 1.6 -0.5 0.3 3.0 2.3 1.9 0.5 2.0 2.3 -0.4 -0.3 2.0 2.0 0.9 0.8 0.9 1.5 0.2 0.4 1.4 -0.5 -0.4 2.9 2.2 1.8 1.9 2.2 1.9 1.9 0.8 0.1 0.8 -1.0 0.3 0.7 1.3 -0.6 -0.5 2.8 2.1 1.7 1.8 1.8 0.7 0.2 0.0 0.7 1.2 1.8 2.1 -0.7 -0.6 median median median median median median median median median median median median median median median - median 3.2328 2.5345 2.0268 2.3016 2.2343 1.2000 0.8544 0.9708 0.55962 0.9933 1.8656 0.3716 2.2322 2.4965 0.0305 0.03922 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean - 3.2105 2.5132 2.0273 2.2314 2.2128 1.1692 0.7985 0.9683 0.5351 1.0046 1.7888 0.1998 2.2051 2.4616 0.0584 0.0441 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.1379 0.1267 0.1200 0.1542 0.1549 0.1684 0.2130 0.1291 0.1971 0.1324 0.2151 0.4263 0.1340 0.1408 0.2090 0.2190

Ceratobatrachus – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.3 3.4 2.8 3.6 3.8 2.2 2.2 1.6 1.7 2.2 2.8 0.3 3.1 3.5 0.5 0.5

3.5 2.1 0.2 4.2 3.3 2.1 1.5 1.6 2.7 3.0 3.4 0.4 0.4 2.7 3.4 3.6 2.0 2.0 0.1 4.1 1.4 2.6 2.9 3.3 0.3 3.2 2.0 1.5 0.3 2.6 3.3 1.9 -0.0 4.0 1.3 2.5 2.8 3.2 0.2 3.4 1.8 3.1 3.2 1.8 1.9 1.4 -0.1 0.2 3.9 2.5 1.2 2.4 2.7 3.1 0.1 3.0 3.1 1.7 1.8 1.3 -0.2 0.1 3.2 1.6 3.8 1.1 2.3 2.6 3.0 -0.0 2.4 3.0 1.6 -0.3 2.9 1.7 1.2 0.0 3.7 1.0 2.2 2.5 2.9 -0.1 2.9 3.0 1.5 1.4 -0.4 2.3 2.8 1.6 1.1 -0.1 3.6 2.8 1.4 0.9 2.1 -0.5 2.4 2.8 -0.2 3.5 2.7 2.2 2.7 2.8 1.3 1.5 0.8 1.0 1.2 2.0 -0.6 2.3 2.7 -0.3 -0.2 median median median median median median median median median median median median median median median median 4.0490 3.3078 2.7100 3.2834 3.3673 1.9022 2.0189 1.3661 1.4207 1.5354 2.6848 0.05353 2.9447 3.2949 0.18647 0.28507 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean Mean 3.98053 3.22120 2.64068 3.25446 3.32212 1.84588 1.97794 1.29645 1.36323 1.59509 2.57000 0.01064 2.84625 3.20613 0.17485 0.22708 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.20733 0.20675 0.19108 0.20778 0.23412 0.24490 0.14735 0.20800 0.18859 0.26921 0.22842 0.19548 0.23232 0.22056 0.22495 0.23168

280 Discodeles – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.6 4.25 3.75 3.8 3.7 2.7 2.6 2.2 1.6 2.2 3.4 3.8 4.25 1.25 1.2

3.7 2.1 3.6 2.6 1.0 3.50 2.4 2.0 3.2 1.0 4.4 4.00 3.6 2.0 1.4 3.6 4.00 1.00 3.5 2.5 3.25 3.5 2.2 1.9 1.8 3.0 0.8 3.4 2.4 4.2 3.75 1.2 3.4 3.75 0.75 3.4 1.8 0.5 3.00 3.3 2.3 2.0 1.6 2.8 0.6 3.3 1.7 4.0 3.50 1.0 3.2 3.50 0.50 3.2 2.2 2.75 3.2 1.8 1.6 1.4 2.6 0.4 0.0 3.1 2.1 3.8 3.25 3.1 1.5 0.8 3.0 3.25 0.25 2.50 1.6 1.2 2.4 0.2 3.0 3.0 2.0 1.4 3.6 3.00 2.25 2.9 2.9 1.9 1.4 1.3 0.6 1.0 2.2 -0.5 2.8 3.00 0.00 0.0 median median median median median median median median median median median median median median median median 4.1989 3.4391 2.8616 3.2745 3.2635 2.1691 1.9755 1.5974 1.2809 1.6312 2.7948 0.3436 3.2546 3.4657 0.6259 0.6206 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 4.14049 3.44163 2.87333 3.26622 3.28448 2.21527 1.97108 1.60002 1.25510 1.58736 2.78742 0.38175 3.21762 3.46073 0.61139 0.58882 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.24156 0.31094 0.29267 0.21397 0.21107 0.21483 0.24390 0.21449 0.20258 0.24732 0.23216 0.43903 0.27193 0.27589 0.28301 0.31236

Palmatorappia – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.5 2.85 2.40 2.45 2.55 1.50 1.4 1.0 0.7 1.40 2.2 0.9 2.35 2.65 0.8 1.0

2.35 2.40 2.50 1.45 1.3 0.6 0.8 2.30 2.60 0.7 0.9 3.4 2.80 1.35 2.1 0.9 2.35 1.40 0.7 2.30 2.45 1.2 0.5 2.25 2.55 0.6 0.8 3.3 2.75 1.30 2.0 2.30 1.35 0.6 0.8 2.25 2.40 1.1 0.4 2.20 2.50 0.5 0.7 3.2 2.70 2.25 1.30 1.25 1.9 0.5 2.20 2.35 1.0 0.3 2.15 2.45 0.4 0.6 0.7 2.20 1.25 0.4 3.1 2.65 1.20 1.8 2.15 2.30 0.9 0.2 2.10 2.40 0.3 0.5 2.15 1.20 0.3 0.6 3.0 2.60 1.15 1.7 2.10 2.10 2.25 1.15 0.8 0.1 0.2 2.05 2.35 0.2 0.4 2.9 2.55 2.05 2.05 2.20 1.10 0.7 0.5 0.0 1.10 1.6 0.1 2.00 2.30 0.1 0.3 median median median median median median median median median median median median median median median median 3.3700 2.7828 2.2945 2.3263 2.4643 1.3737 1.1506 0.87963 0.38526 1.2542 2.0788 0.70802 2.2370 2.5221 0.38861 0.63656 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.33675 2.76310 2.28604 2.30592 2.43824 1.34168 1.15225 0.85279 0.39988 1.26304 2.04231 0.61951 2.22637 2.51229 0.39635 0.61438 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.10772 0.06227 0.06696 0.08061 0.08012 0.09251 0.12894 0.10758 0.13013 0.06534 0.12055 0.18405 0.08333 0.08134 0.13837 0.13656

281 Platymantis – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.6 1.8 1.5 1.5 1.5 3.4 3 2.0 1.6 2 3.0 1.0 4 3 2 1.0 3.2 1.4 1.0 3 3 2.5 0.5 3.0 1.5 1.2 0.5 0.5 1 3 2.8 1.0 0.0 2.0 0.0 0.0 2.6 1.0 0.8 2 1 -0.5 -0.5 2.4 0.6 -0.5 1.5 2 3 1 -1.0 2.2 2 0.5 0.4 -1.0 2 -1.0 2.0 0.2 0 1.0 2 -1.5 -1.5 1.8 0.0 0.0 -1.5 median median median median median median median median median median median median - median median median - median 3.5444 2.9549 2.3832 2.6218 2.6152 1.5518 1.3350 1.1314 0.8713 1.1909 2.2628 0.1278 2.5726 2.9014 0.0101 0.0583 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean 3.5374 2.9154 2.3798 2.6288 2.6202 1.5839 1.3312 1.1262 0.8806 1.1908 2.2079 0.0156 2.5618 2.8950 0.0144 0.0590 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.3177 0.3278 0.3082 0.3224 0.3228 0.3086 0.3826 0.2970 0.3918 0.2888 0.3659 0.6210 0.3428 0.3443 0.4198 0.4379

Rana – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.2 3.7 3.1 3.3 3.1 2.2 2.0 1.9 1.80 1.8 2.9 0.6 3.2 3.6 0.6 0.7

1.75 3.6 3.0 2.1 1.9 1.8 1.7 0.5 3.1 3.5 0.5 0.6 4.1 3.2 3.0 1.70 2.8 0.4 3.0 0.5 3.5 2.9 2.0 1.8 1.7 1.6 3.4 0.4 4.0 1.65 0.3 2.9 0.4 3.1 2.9 2.7 3.4 2.8 1.9 1.7 1.6 1.60 1.5 3.3 0.3 3.9 0.2 2.8 0.3 3.3 2.7 1.8 1.6 1.5 1.55 1.4 3.2 0.2 3.0 2.8 2.6 0.1 2.7 0.2 3.8 1.50 3.2 2.6 1.7 1.5 1.4 1.3 3.1 0.1 0.0 2.6 0.1 2.9 2.7 1.45 2.5 3.7 3.1 2.5 1.6 1.4 1.3 1.2 3.0 0.0 1.40 -0.1 2.5 0.0 3.6 3.0 2.4 2.8 2.6 1.5 1.3 1.2 1.35 1.1 2.4 -0.2 2.4 2.9 -0.1 -0.1 median median median median median median median median median median median median median median median median 3.7793 3.1508 2.5696 2.9231 2.7317 1.7976 1.4827 1.4805 1.5602 1.3673 2.5388 0.23899 2.8109 3.1357 0.25464 0.26236 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.82667 3.19415 2.62617 2.95618 2.77580 1.81859 1.53745 1.52314 1.58401 1.39998 2.57534 0.22074 2.83181 3.17315 0.28111 0.26648 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.14200 0.14045 0.14107 0.11096 0.13004 0.11861 0.13158 0.12326 0.09946 0.12844 0.13752 0.19330 0.16162 0.13846 0.16760 0.17289

282 Batrachylodes elegans – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.45 2.75 2.25 2.50 2.5 1.40 1.2 1.20 0.8 1.4 2.2 0.8 2.45 2.75 0.4 0.4

3.40 2.45 1.35 1.15 0.7 2.40 2.70 0.3 2.70 2.20 1.1 1.3 2.1 0.7 0.3 2.4 3.35 2.40 1.30 1.10 0.6 2.35 2.65 0.2 2.65 2.15 1.0 1.2 2.0 0.6 0.2

3.30 2.35 2.3 1.25 1.05 0.5 2.30 2.60 0.1

2.60 2.10 0.9 1.1 1.9 0.5 0.1 3.25 2.30 1.20 1.00 0.4 2.25 2.55 0.0 2.2 2.55 2.05 0.8 1.0 1.8 0.4 0.0 3.20 2.25 1.15 0.95 0.3 2.20 2.50 -0.1

3.15 2.50 2.00 2.20 2.1 1.10 0.7 0.90 0.2 0.9 1.7 0.3 2.15 2.45 -0.2 -0.1 median median median median median median median median median median median median median median median median 3.2962 2.6333 2.1702 2.3599 2.3321 1.2669 0.9478 1.0852 0.43178 1.1019 1.9615 0.47623 2.3360 2.5696 0.11333 0.15700 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.29835 2.64420 2.15901 2.35104 2.34585 1.26206 0.94042 1.06929 0.48092 1.12848 1.96001 0.49359 2.31741 2.58623 0.09426 0.15524 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.07957 0.05244 0.06908 0.08246 0.09703 0.07703 0.09980 0.07470 0.13177 0.10251 0.11223 0.12813 0.08987 0.08885 0.13724 0.13447

Batrachylodes minutus – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.00 2.24 1.80 1.92 1.95 1.0 0.5 0.75 0.5 0.85 -0.7 1.90 2.175 -0.4 -0.5

1.35 -0.8 2.95 2.22 1.91 1.90 0.4 0.70 0.4 2.150 0.9 0.80 1.88 -0.6 -0.6 1.75 -0.9 2.90 2.20 1.90 1.85 0.3 0.65 0.3 2.125 1.30 -1.0 2.85 0.8 0.75 1.86 -0.8 -0.7 1.70 -1.1 2.18 1.89 1.80 0.2 0.60 0.2 2.100 2.80 1.25 -1.2 0.7 0.70 1.84 -1.0 -0.8 2.16 1.88 1.75 0.1 0.55 0.1 2.075 2.75 1.65 -1.3 2.70 2.14 1.87 1.70 0.6 0.0 0.50 0.0 0.65 1.20 -1.4 1.82 2.050 -1.2 -0.9 median median median median median median median median median median median median - median median median - median - 2.8965 2.1622 1.7210 1.8946 1.8579 0.73716 0.32208 0.66783 0.23111 0.76081 1.2920 0.9943 1.8469 2.0857 0.7550 0.6931 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 2.87643 2.17255 1.71335 1.89361 1.83624 0.75717 0.27204 0.64760 0.25835 0.75930 1.29204 1.02450 1.85094 2.09964 0.73301 0.70172 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.103594 0.030643 0.053830 0.016136 0.077958 0.115613 0.172255 0.081550 0.202327 0.061609 0.063087 0.189036 0.029021 0.037993 0.284841 0.146233

283 Batrachylodes trossulus – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnTL lnT4 lnTD3 lnTD4

3.10 2.00 2.10 2.05 1.1 0.9 0.90 0.6 1.05 1.7 -0.4 2.15 2.8 -0.1 -0.0

2.40 0.8 0.85 -0.1 2.05 2.00 0.5 1.00 2.7 3.05 1.0 1.6 -0.2 1.95 -0.5 2.10 0.7 0.80 -0.2 2.35 2.00 1.95 0.4 0.95 2.6 3.00 0.9 1.5 -0.3 0.6 0.75 -0.3 1.90 1.95 1.90 0.3 0.90 -0.6 2.05 2.5 2.30 0.5 0.70 -0.4 2.95 0.8 1.4 -0.4 1.90 1.85 0.2 0.85 2.4 0.4 0.65 -0.5 1.85 -0.7 2.00 2.90 2.25 0.7 1.3 -0.5 1.85 1.80 0.1 0.80 2.3 0.3 0.60 -0.6 2.85 2.20 1.80 1.80 1.75 0.6 0.2 0.55 0.0 0.75 1.2 -0.8 1.95 2.2 -0.6 -0.7 median median median median median median median median median median median median - median median median - median - 2.9513 2.3360 1.8764 1.9351 1.8990 0.8774 0.53063 0.74194 0.30008 0.9400 1.4679 0.6733 2.0308 2.2966 0.2748 0.3285 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 2.96949 2.33191 1.87650 1.94220 1.90750 0.90812 0.52793 0.76163 0.32459 0.92781 1.46857 0.65164 2.03876 2.32426 0.31885 0.31328 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.07910 0.04652 0.04270 0.06994 0.07753 0.12390 0.13453 0.09558 0.15313 0.07898 0.09843 0.08830 0.05621 0.13250 0.11983 0.14168

Batrachylodes vertebralis – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.5 2.70 2.15 2.5 2.5 1.5 1.1 1.2 1.0 1.05 2.0 0.7 2.40 2.6 0.2 0.1

2.65 2.35 2.10 2.4 1.9 0.6 0.1 3.4 2.4 1.4 1.0 1.1 0.8 -0.0 2.60 1.00 2.30 2.5 2.05 -0.0 2.3 1.8 0.5 3.3 2.55 2.3 1.3 0.9 1.0 0.6 2.25 -0.1 2.00 -0.1 0.95 2.4 2.50 2.2 1.7 0.4 2.20 3.2 1.95 2.2 1.2 0.8 0.9 0.4 -0.2 -0.2 2.45 2.1 1.6 0.3 2.15 0.90 2.3 1.90 -0.3 3.1 2.40 2.1 1.1 0.7 0.8 0.2 2.10 -0.3 2.0 1.5 0.2 1.85 -0.4 2.35 0.85 2.05 2.2 3.0 2.0 1.0 0.6 0.7 0.0 -0.4 1.9 1.4 0.1 2.30 1.80 2.00 -0.5 2.9 2.25 1.75 1.9 1.8 0.9 0.5 0.6 -0.2 0.80 1.3 0.0 1.95 2.1 -0.6 -0.5 median median median median median median median median median median median median median median median - median - 3.1355 2.4467 1.9762 2.1175 2.1459 1.1474 0.6716 0.9062 0.41211 0.9381 1.7263 0.21498 2.1453 2.3475 0.1866 0.1166 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean - 3.13801 2.44442 1.96739 2.12846 2.14362 1.14501 0.72565 0.89810 0.36783 0.93770 1.70951 0.22026 2.13692 2.35629 0.17818 0.13454 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.11809 0.09824 0.10472 0.12384 0.11305 0.12893 0.19495 0.12663 0.24629 0.06645 0.18158 0.15815 0.10015 0.11935 0.19601 0.18636

284 Batrachylodes wolfi – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.40 2.65 2.15 2.40 2.45 1.40 1.10 1.05 0.80 1.4 2.05 0.7 2.40 2.65 0.5 0.6

2.40 1.05 1.00 0.75 1.3 0.6 2.60 0.4 3.35 2.60 1.35 2.00 2.35 0.4 2.10 2.35 2.35 1.00 0.95 0.70 1.2 0.5 2.55 0.3 3.30 2.55 1.30 1.95 2.30 0.2

2.05 2.30 2.30 0.95 0.90 0.65 1.1 0.4 2.50 0.2

3.25 2.50 1.25 1.90 2.25 0.0 2.25 0.90 0.85 0.60 1.0 0.3 2.45 0.1 2.00 2.25 3.20 2.45 1.20 1.85 2.20 -0.2 2.20 0.85 0.80 0.55 0.9 0.2 2.40 0.0

3.15 2.40 1.95 2.20 2.15 1.15 0.80 0.75 0.50 0.8 1.80 0.1 2.15 2.35 -0.1 -0.4 median median median median median median median median median median median median median median median median 3.3359 2.5553 2.0699 2.3403 2.3626 1.3066 1.0188 0.9103 0.60140 1.2204 1.9747 0.56246 2.2986 2.5802 0.25461 0.21087 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.30117 2.55733 2.07329 2.32727 2.34353 1.29410 1.00077 0.90948 0.60958 1.17847 1.95543 0.52002 2.29348 2.54284 0.23129 0.16511 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.087765 0.061477 0.054724 0.058297 0.072660 0.083273 0.078609 0.087362 0.080423 0.143694 0.071324 0.180169 0.062554 0.085187 0.143311 0.218551

Ceratobatrachus guentheri – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.4 3.6 3.1 3.7 3.8 2.3 2.4 1.8 2.25 2.4 3.1 0.5 3.3 3.7 0.7 0.7

3.0 0.4 0.6 0.6 3.6 3.7 2.2 2.3 1.7 2.2 3.0 3.2 3.6 4.3 3.5 2.00 0.3 0.5 2.9 0.5 3.5 3.6 2.1 2.2 1.6 2.0 2.9 3.1 3.5 0.2 0.4 4.2 3.4 1.75 2.8 0.4 3.4 3.5 2.0 2.1 1.5 1.8 2.8 0.1 3.0 3.4 0.3 2.7 0.3 4.1 3.3 1.50 -0.0 0.2 3.3 3.4 1.9 2.0 1.4 1.6 2.7 2.9 3.3 2.6 0.2 -0.1 0.1 4.0 3.2 1.25 3.2 3.3 1.8 1.9 1.3 1.4 2.6 2.8 3.2 2.5 -0.2 0.0 0.1 3.9 3.1 2.4 3.1 3.2 1.7 1.8 1.2 1.00 1.2 2.5 -0.3 2.7 3.1 -0.1 0.0 median median median median median median median median median median median median median median median median 4.2485 3.4920 2.8987 3.5041 3.6085 2.0360 2.2176 1.5507 1.4702 1.7155 2.8633 0.20293 3.1385 3.4690 0.39541 0.48841 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 4.22380 3.44936 2.84989 3.46234 3.54985 2.00411 2.18042 1.52874 1.49321 1.78038 2.83484 0.20902 3.10011 3.45149 0.36712 0.43526 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.13570 0.12767 0.15835 0.13483 0.14660 0.15112 0.12330 0.13518 0.21718 0.22824 0.14577 0.17510 0.14915 0.14227 0.18426 0.17951

285 Discodeles bufoniformis – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.5 4.0 4.25 4.5 3.0 3.00 2.2 3.0 4.0 1.5 1.50 1.5

5.0 2.5 4.0 4.5 4.00 2.8 2.75 2.0 1.25 2.5 1.0 4.0 3.5 4.0 3.5 1.0 3.75 2.6 2.50 1.8 1.00 4.0 4.5 2.0 3.5 3.50 2.4 2.25 1.6 2.0 0.5 0.75 3.5 3.0 3.5 3.0 0.5 3.5 3.25 2.2 2.00 1.4 0.50 4.0 1.5 3.0 1.5 0.0 3.00 2.0 1.75 1.2 0.25 3.0 2.5 3.0 2.5 3.0 0.0 3.5 2.75 1.8 1.50 1.0 1.0 1.0 -0.5 2.5 0.00 median median median median median median median median median median median median median median median median 4.4172 3.6363 3.1130 3.5565 3.6750 2.5088 2.1798 1.7596 1.5810 1.9906 3.0129 0.4383 3.4444 3.7116 0.7792 0.7605 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 4.49736 3.72869 3.15594 3.59345 3.69526 2.47414 2.26274 1.88268 1.55577 2.02712 3.07466 0.53905 3.49384 3.74842 0.78544 0.76711 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.41039 0.45338 0.47769 0.37162 0.41723 0.32066 0.39462 0.42372 0.28595 0.53284 0.40817 0.48534 0.43503 0.43664 0.36267 0.37948

Discodeles guppyi – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

5.5 4.5 4.5 3.25 2.25 3.5 4.0 4.5 1.75

4.5 4.0 3.0 2.5 1.5 4.5 1.5 3.00 2.00 1.50 3.0 5.0 4.0 4.0 3.5 4.0 2.75 1.75 1.25 3.5 2.5 4.0 2.5 2.0 1.0 4.0 1.0 4.5 2.50 1.50 1.00 3.5 3.5 3.0 3.5 2.0 3.0 2.25 1.25 0.75 3.5 2.0 1.5 0.5 3.5 0.5 4.0 1.5 2.00 1.00 0.50 2.5 3.0 3.0 2.5 3.0 3.5 3.0 1.75 1.5 1.0 0.75 1.0 0.0 3.0 0.25 0.0 median median median median median median median median median median median median median median median median 5.0084 4.4212 3.3669 3.9700 4.1096 2.7551 2.1497 2.0062 1.7850 2.5603 3.2581 0.8700 3.6889 4.0775 0.8448 0.8582 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 4.74155 4.12216 3.33193 3.77245 3.84770 2.62518 2.38194 1.99551 1.59060 2.24792 3.17677 0.89138 3.62395 3.92571 0.94083 0.92441 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.59939 0.56137 0.66382 0.52073 0.60679 0.40235 0.58592 0.57367 0.50157 0.73859 0.49497 0.52627 0.50373 0.50990 0.47269 0.49787

286 Palmatorappia solomonis – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.50 2.90 2.45 2.425 2.55 1.6 1.4 1.1 0.8 1.4 2.2 1.0 2.45 2.75 0.75 1.00

2.85 2.70 2.40 2.400 1.5 2.40 3.45 1.3 1.0 0.7 2.1 0.75 2.50 1.2 0.50 2.80 0.5 2.65 2.35 2.375 1.4 2.35 3.40 1.2 0.9 0.6 2.0 0.50 2.75 2.60 2.30 2.350 2.45 1.3 1.0 2.30 0.25 2.70 0.0 2.55 3.35 1.1 0.8 0.5 1.9 0.25 2.25 2.325 1.2 2.25 2.65 2.50 2.40 0.8 0.00 3.30 1.0 0.7 0.4 1.8 0.00 2.20 2.300 1.1 2.20 2.60 -0.5 2.45 3.25 2.55 2.15 2.275 2.35 1.0 0.9 0.6 0.3 0.6 1.7 2.15 2.40 -0.25 -0.25 median median median median median median median median median median median median median median median median 3.3690 2.7948 2.3321 2.3627 2.4891 1.3762 1.2149 0.9708 0.43825 1.2920 2.1187 0.73716 2.2565 2.5494 0.50682 0.74194 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.37814 2.77658 2.30909 2.35860 2.46642 1.31816 1.21387 0.92230 0.46977 1.24525 2.05586 0.59366 2.27671 2.55715 0.40961 0.62843 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.059105 0.082221 0.074616 0.041941 0.059899 0.152484 0.123513 0.141187 0.117101 0.189770 0.123685 0.438179 0.072454 0.092951 0.249514 0.264745

Platymantis acrochorda – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.65 2.95 2.5 2.80 2.90 1.85 1.4 1.2 1.2 1.3 2.2 -0.3 2.65 2.95 -0.0 0.2

2.75 2.85 1.1 2.1 3.60 1.80 2.60 2.90 2.4 1.2 -0.4 2.90 2.70 2.80 1.3 1.0 1.0 2.0 -0.2 0.0 3.55 1.75 2.55

2.85 2.3 2.65 2.75 0.9 1.1 1.9 -0.5 2.85

3.50 1.70 2.50 2.60 2.70 1.2 0.8 0.8 1.8 -0.4 -0.2 2.80 2.2 1.0 -0.6 2.80 3.45 1.65 2.45 2.55 2.65 0.7 1.7

3.40 2.75 2.1 2.50 2.60 1.60 1.1 0.6 0.6 0.9 1.6 -0.7 2.40 2.75 -0.6 -0.4 median median median median median median median median median median median median - median median median - median - 3.6014 2.9167 2.3780 2.7147 2.8034 1.7228 1.3321 0.9619 0.9123 1.1639 2.1163 0.3784 2.5551 2.8875 0.1587 0.0408 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 3.56648St 2.89148St 2.33607St 2.67815 2.78425 1.71442 1.30183 0.94567 0.87972 1.14771 2.00004 0.43239 2.53677 2.86609 0.19093 0.10097 d Dev d Dev d Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.091996 0.077342 0.102333 0.095282 0.108498 0.068844 0.101822 0.205078 0.219438 0.135562 0.232266 0.123096 0.084848 0.070081 0.193248 0.184945

287 Platymantis aculeodactylus – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.50 2.30 2.55 2.60 1.40 1.20 1.20 0.80 1.10 1.85 -0.4 2.50 2.80 -0.0 0.1

1.35 1.15 3.45 2.70 1.15 1.05 -0.1 -0.0 2.25 2.50 2.55 0.75 1.80 -0.5 2.45 1.30 1.10 2.75 3.40 -0.1 1.10 1.00 -0.2 2.20 2.45 2.50 1.25 1.05 0.70 1.75 -0.6 2.40 3.35 2.65 -0.2 2.70 1.20 1.00 1.05 0.95 -0.3 3.30 2.15 2.40 2.45 0.65 1.70 -0.7 2.35 -0.3 1.15 0.95 1.00 0.90 -0.4 2.65 3.25 2.60 -0.4 2.10 2.35 2.40 1.10 0.90 0.60 1.65 -0.8 2.30 0.95 0.85 -0.5 3.20 -0.5 1.05 0.85 2.60 2.05 2.30 2.35 0.55 1.60 -0.9 2.25 0.90 0.80 -0.6 3.15 2.55 1.00 0.80 -0.6 3.10 2.00 2.25 2.30 0.95 0.75 0.85 0.50 0.75 1.55 -1.0 2.20 2.55 -0.7 -0.7 median median median median median median median median median median median median - median median median - median - 3.3604 2.6383 2.1994 2.4078 2.4689 1.2528 0.9183 1.0332 0.63658 0.9783 1.7639 0.664 2.3910 2.6939 0.1803 0.1454 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 3.33640 2.62768 2.18532 2.39841 2.45812 1.22328 0.91087 1.02425 0.66087 0.96964 1.74800 0.69180 2.37428 2.68310 0.24846 0.20611 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.07444 0.05274 0.07296 0.05829 0.07053 0.10819 0.09042 0.08043 0.08100 0.07568 0.07497 0.17732 0.08063 0.07167 0.15311 0.19613

Platymantis boulengeri – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.4 3.6 3.2 3.4 3.6 2.1 2.1 2.0 1.7 2.50 2.9 1.00 3.3 3.7 0.7 0.8

4.3 0.7 3.5 3.1 3.3 2.0 1.9 1.6 2.8 3.2 3.6 0.6 2.0 0.75 4.2 3.4 2.25 0.6 3.0 3.2 1.5 2.7 3.1 3.5 0.5 3.4 1.9 1.8 4.1 1.9 0.50 0.5 2.9 3.1 1.4 2.6 3.0 3.4 0.4 3.2 2.00 4.0 3.3 1.8 1.7 0.4 2.8 3.0 1.8 1.3 2.5 0.25 2.9 3.3 0.3 3.9 3.2 1.7 1.6 0.3 3.0 1.75 2.7 2.9 1.2 2.4 2.8 3.2 0.2 3.8 1.7 0.00 0.2 3.1 1.6 1.5 2.6 2.8 1.1 2.3 2.7 3.1 0.1 3.7 2.8 1.50 0.1 1.6 -0.25 3.0 1.5 1.4 3.6 2.5 2.7 1.0 2.2 2.6 3.0 0.0 0.0 3.5 2.9 2.4 2.6 2.6 1.5 1.4 1.3 0.9 1.25 2.1 -0.50 2.5 2.9 -0.1 -0.1 median median median median median median median median median median median median median median median median 4.0070 3.3969 2.8769 3.0732 3.2121 1.9402 1.7579 1.7138 1.5063 1.9036 2.6568 0.33647 3.0057 3.3149 0.20701 0.41211 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.99767 3.34954 2.85105 3.07816 3.21287 1.89048 1.77446 1.70177 1.38885 1.89759 2.60182 0.33228 2.98427 3.30531 0.30847 0.39671 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.24354 0.20290 0.22284 0.22621 0.26716 0.17572 0.19561 0.20916 0.23232 0.31590 0.24202 0.32693 0.22403 0.23370 0.22300 0.21829

288 Platymantis corrugata – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.8 3.4 2.8 2.9 3.1 1.9 1.7 1.7 1.2 1.7 2.6 0.75 2.9 3.3 0.2 0.4

3.3 2.7 3.0 1.8 1.6 1.6 0.50 2.8 3.2 3.7 2.8 1.6 0.3 1.0 2.4 0.1 3.2 2.6 2.9 1.7 1.5 1.5 0.25 2.7 3.1 3.6 2.7 1.5 0.2

3.1 2.5 2.8 1.6 1.4 0.8 1.4 2.2 0.00 2.6 3.0 0.0

3.5 2.6 1.4 0.1 3.0 2.4 2.7 1.5 1.3 1.3 -0.25 2.5 2.9 0.6 2.0 -0.1 3.4 2.5 1.3 0.0 2.9 2.3 2.6 1.4 1.2 1.2 -0.50 2.4 2.8

3.3 2.8 2.2 2.4 2.5 1.3 1.1 1.2 0.4 1.1 1.8 -0.75 2.3 2.7 -0.2 -0.1 median median median median median median median median median median median median - median median median median 3.6819 3.2276 2.6472 2.7828 2.8614 1.5890 1.4609 1.5006 1.0282 1.4525 2.3399 0.1985 2.7584 3.0999 0.05827 0.17814 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean Mean 3.64564 3.16171 2.59293 2.74584 2.82337 1.58887 1.43645 1.48291 0.92534 1.41629 2.23922 0.16708 2.69861 3.04846 0.04314 0.14988 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.14156 0.16159 0.16020 0.14217 0.14844 0.14157 0.15195 0.12824 0.23759 0.13612 0.21205 0.27450 0.15563 0.16783 0.11940 0.12004

Platymantis dorsalis – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 LnT4 lnTD3 lnTD4

3.8 3.3 3.0 2.9 2.9 1.8 1.6 1.5 1.3 1.4 2.5 0.1 2.9 3.2 0.2 0.3

0.2 2.9 2.8 1.7 1.4 1.2 2.4 -0.0 0.1 3.7 3.2 2.8 1.5 1.3 2.8 3.1 0.1 2.8 -0.1 -0.0 2.7 1.6 1.3 1.1 2.3 3.6 3.1 2.7 1.4 1.2 2.7 3.0 -0.0 2.7 -0.2 -0.1 2.6 1.5 1.2 1.0 2.2 -0.1 3.5 3.0 2.6 2.6 1.3 1.1 -0.3 2.6 2.9 -0.2 2.5 1.4 1.1 0.9 2.1 -0.2 2.5 -0.4 -0.3 3.4 2.9 2.5 1.2 1.0 2.5 2.8 -0.3 2.4 1.3 1.0 0.8 2.0 2.4 -0.5 -0.4 -0.4 3.3 2.8 2.4 1.1 0.9 2.4 2.7 2.3 1.2 0.9 0.7 1.9 2.3 -0.6 -0.5 -0.5 3.2 2.7 2.2 2.2 2.3 1.1 1.0 0.8 0.6 0.8 1.8 -0.7 2.3 2.6 -0.6 -0.6 median median median median median median median median median median median median - median median median median 3.5791 3.1272 2.5369 2.6844 2.7543 1.6174 1.3610 1.2837 0.9746 1.1632 2.2407 0.1054 2.6700 2.9932 0.06766 0.04879 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean 3.55692 3.08549 2.54585 2.63907 2.69810 1.56414 1.30079 1.25026 0.94285 1.14659 2.24959 0.17278 2.65924 3.00684 0.02424 0.01268 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.13953 0.12241 0.15027 0.13646 0.14198 0.15966 0.14521 0.14238 0.17110 0.12535 0.15639 0.17753 0.13711 0.14968 0.20676 0.21650

289 Platymantis gilliardi – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.9 3.4 2.8 3.0 3.2 1.9 1.6 1.5 1.3 1.6 2.5 0.2 3.0 3.3 0.2 0.2

1.4 1.5 3.2 0.1 3.8 3.3 2.7 2.9 1.8 1.5 1.2 2.4 3.0 -0.0 2.8 -0.0 1.3 1.4 3.1 -0.0 3.7 3.2 2.6 2.8 1.7 1.4 1.1 2.3 2.8 1.2 1.3 3.0 -0.1 3.6 3.1 2.5 2.7 1.6 1.3 1.0 2.2 -0.2 2.6 -0.2 1.1 1.2 2.9 -0.2 2.6 3.5 3.0 2.4 2.6 1.5 1.2 0.9 2.1 1.0 1.1 2.8 -0.3 -0.4 2.4 -0.4 2.4 3.4 2.9 2.3 2.5 1.4 1.1 0.8 2.0 0.9 1.0 2.7 -0.4 3.3 2.8 2.2 2.4 2.2 1.3 1.0 0.8 0.7 0.9 1.9 -0.6 2.2 2.6 -0.6 -0.5 median median median median median median median median median median median median - median median median - median 3.7504 3.1482 2.6001 2.8244 2.8558 1.8107 1.4363 1.3925 1.1553 1.4598 2.3000 0.2899 2.7596 3.1134 0.1111 0.05353 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 3.68848 3.13157 2.57640 2.76651 2.78895 1.72479 1.40874 1.31161 1.07678 1.37312 2.25046 0.26765 2.71094 3.06109 0.13717 0.09232 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.181911 0.144736 0.163884 0.183301 0.258754 0.184448 0.162676 0.206617 0.160286 0.212946 0.179468 0.226173 0.213354 0.207485 0.243788 0.27055

Platymantis guentheri – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.0 3.4 2.8 2.9 3.2 1.9 1.7 1.4 1.2 1.8 3.0 1.50 3.0 3.4 0.8 1.0

2.7 1.6 2.8 1.8 1.3 3.0 1.0 2.8 0.6 0.8 3.8 3.2 2.6 1.5 1.6 1.25 2.8 3.2 2.7 1.7 1.2 2.5 2.8 1.4 0.8 2.6 0.4 0.6 2.6 3.6 3.0 1.4 1.00 2.6 3.0 2.4 1.6 1.3 1.1 2.5 2.6 0.6 2.4 0.2 0.4 2.3 1.5 1.2 1.0 3.4 2.8 1.2 0.75 2.4 2.8 2.4 2.2 2.4 1.1 0.4 2.2 0.0 0.2 1.4 0.9 2.3 3.2 2.6 2.1 1.0 1.0 0.50 2.2 2.6 2.2 0.2 2.0 -0.2 0.0 1.3 0.8 2.0 2.2 0.9 3.0 2.4 1.9 2.1 2.0 1.2 0.8 0.7 0.0 0.8 1.8 0.25 2.0 2.4 -0.4 -0.2 median median median median median median median median median median median median median median median median 3.7386 3.1162 2.6056 2.7819 2.9014 1.7047 1.4279 1.2267 0.8796 1.4279 2.6858 1.2090 2.8273 3.1219 0.43178 0.51879 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.62966 3.04344 2.51612 2.69703 2.78892 1.63573 1.37455 1.16908 0.77290 1.37363 2.58289 1.08704 2.70929 3.01871 0.39414 0.44880 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.24450 0.23248 0.21655 0.22304 0.26478 0.19132 0.23741 0.18619 0.28576 0.24720 0.27597 0.35373 0.26318 0.25583 0.28563 0.28836

290 Platymantis guppyi – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.6 4.0 3.4 3.6 4.00 2.4 2.5 2.0 1.8 2.50 3.6 2.25 3.6 4.0 1.8 2.0

3.5 2.4 2.3 1.6 3.4 2.00 1.6 1.8 4.4 3.8 3.2 3.4 3.75 2.3 1.8 2.25 3.4 3.8 2.2 3.3 2.2 1.4 3.2 1.75 1.4 1.6 4.2 3.6 3.0 3.50 1.6 2.00 3.2 3.6 3.2 2.1 2.1 1.2 3.0 1.50 1.2 1.4 3.1 2.0 2.0 4.0 3.4 2.8 3.25 1.4 1.75 3.0 3.4 3.0 1.9 1.0 2.8 1.25 1.0 1.2 1.9 3.8 3.2 2.6 2.9 3.00 1.8 1.2 1.50 2.8 3.2 0.8 2.6 1.00 0.8 1.0 1.8 2.8 1.7 3.6 3.0 2.4 2.7 2.75 1.7 1.6 1.0 0.6 1.25 2.4 0.75 2.6 3.0 0.6 0.8 median median median median median median median median median median median median median median median median 4.2648 3.6079 3.0194 3.3157 3.4653 2.0222 2.1459 1.6174 1.4375 2.0412 3.0929 1.6658 3.2966 3.6339 1.3610 1.5602 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 4.23211 3.57131 2.95879 3.26539 3.40142 2.00909 2.08687 1.54262 1.36917 2.00788 2.99488 1.56058 3.18596 3.53515 1.27841 1.41852 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.27193 0.24887 0.29722 0.23928 0.30347 0.18564 0.24761 0.26912 0.22284 0.32526 0.30842 0.41353 0.29779 0.28921 0.38601 0.40066

Platymantis hazelae – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.50 2.85 2.40 2.55 2.6 1.6 1.3 0.95 0.6 1.25 2.25 1.0 2.45 2.9 0.40 0.65

0.5 3.45 2.50 1.20 0.9 2.40 0.60 2.80 2.35 1.2 0.90 2.20 2.8 0.35 2.5 1.5 0.4 3.40 2.45 1.15 0.8 2.35 0.55 2.75 2.30 1.1 0.85 2.15 2.7 0.30 0.3 3.35 2.40 2.4 1.4 1.10 0.7 2.30 0.50 0.2 2.70 2.25 1.0 0.80 2.10 2.6 0.25 3.30 2.35 1.05 0.6 2.25 0.45 0.1 2.3 1.3 2.65 2.20 0.9 0.75 2.05 2.5 0.20 3.25 2.30 1.00 0.5 2.20 0.40 0.0 3.20 2.60 2.15 2.25 2.2 1.2 0.8 0.70 -0.1 0.95 2.00 0.4 2.15 2.4 0.15 0.35 median median median median median median median median median median median median median median median median 3.3861 2.7402 2.2354 2.3571 2.4024 1.4351 1.1314 0.89200 0.30010 1.0986 2.1330 0.66269 2.3135 2.6261 0.26236 0.43825 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.36085 2.73748 2.23535 2.37556 2.41580 1.40786 1.08881 0.85017 0.28606 1.12224 2.13980 0.70253 2.31591 2.63858 0.26767 0.46362 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.091446 0.070894 0.07970 0.092380 0.109411 0.114255 0.141170 0.075862 0.221987 0.092187 0.083405 0.174806 0.096734 0.105966 0.089325 0.091910

291 Platymantis ingeri – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.6 3.1 2.5 2.6 2.6 1.5 1.2 1.2 0.9 1.3 2.3 0.6 2.6 2.9 0.3 0.4

3.5 3.0 2.5 1.2 2.2 0.3 2.5 1.1 0.8 2.5 2.8 0.2 2.4 1.4 1.1 0.4 3.4 2.9 2.4 1.1 2.1 0.2 2.4 1.0 0.7 2.4 2.7 0.1

3.3 2.8 2.3 2.3 1.3 1.0 1.0 2.0 0.2 0.1

2.3 0.9 0.6 2.3 2.6 0.0 3.2 2.7 2.2 0.9 1.9 0.0 2.2 1.2 0.9 0.0 2.2 0.8 0.5 2.2 2.5 -0.1 3.1 2.6 2.1 0.8 1.8 -0.1

3.0 2.5 2.1 2.1 2.0 1.1 0.8 0.7 0.4 0.7 1.7 -0.2 2.1 2.4 -0.2 -0.2 median median median median median median median median median median median median median median median median 3.4075 2.9277 2.4204 2.4583 2.5076 1.3686 1.1442 0.9969 0.59333 1.1442 2.1395 0.38861 2.4732 2.8232 0.18232 0.25464 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.38138 2.90796 2.37523 2.43746 2.46649 1.36963 1.10008 0.97728 0.60148 1.08301 2.09867 0.39127 2.44752 2.78945 0.12705 0.21730 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.11939 0.10700 0.10223 0.09642 0.12917 0.11627 0.09602 0.10499 0.11263 0.12769 0.14109 0.17206 0.10913 0.12548 0.12864 0.13649

Platymantis levigatus – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.7 3.0 2.5 2.7 2.9 1.7 1.2 1.3 0.7 1.3 2.3 0.1 2.8 3.0 0.4 0.4

2.8 3.6 1.2 2.9 2.4 2.6 1.6 1.1 1.2 2.2 -0.0 2.9 0.6 0.2 2.7 3.5 1.1 2.6 0.2 2.8 2.3 2.5 1.5 1.0 1.1 2.1 -0.1 2.8 2.6 3.4 0.5 1.0 0.0 2.5 2.7 2.2 2.4 1.4 0.9 1.0 2.0 -0.2 2.7 3.3 0.9 2.4 0.0 2.4 0.4 -0.2 2.6 2.1 2.3 1.3 0.8 0.9 1.9 -0.3 2.6 3.2 0.8 2.3 3.1 2.5 2.0 2.2 2.2 1.2 0.7 0.8 0.3 0.7 1.8 -0.4 2.2 2.5 -0.4 -0.2 median median median median median median median median median median median median - median median median median 3.5219 2.8848 2.3274 2.5502 2.6290 1.5686 1.0762 1.1652 0.57291 1.0613 2.1759 0.0101 2.5556 2.8547 0.09428 0.17427 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean Mean 3.474584 2.845652 2.276044 2.492125 2.590862 1.540383 1.006806 1.107090 0.542605 1.024478 2.066179 0.059963 2.490218 2.792769 0.049130 0.143121 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.211762 0.164286 0.161656 0.172658 0.193875 0.155286 0.177120 0.164349 0.151746 0.174058 0.219590 0.184016 0.187487 0.165812 0.196184 0.194823

292 Platymantis myersi – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.2 3.6 3.1 3.4 3.4 2.4 2.2 1.9 1.6 1.8 3.0 0.50 3.4 3.7 0.75 0.75

3.3 3.3 2.1 1.8 1.5 2.9 3.3 3.6 4.1 3.5 3.0 1.7 2.2 0.50 0.50 1.4 0.25 3.2 3.2 2.0 1.7 2.8 3.2 3.5 4.0 3.4 2.9 1.6 1.3 2.0 0.25 0.25 3.1 3.1 1.9 1.6 2.7 3.1 3.4 3.9 3.3 2.8 1.2 1.5 0.00 3.0 3.0 1.8 1.5 2.6 3.0 3.3 1.8 0.00 0.00 1.1 3.8 3.2 2.7 1.4 2.9 2.9 1.7 1.4 2.5 2.9 3.2 1.0 -0.25 1.6 -0.25 -0.25 3.7 3.1 2.6 1.3 2.8 2.8 1.6 1.3 0.9 2.4 2.8 3.1 3.6 3.0 2.5 2.7 2.7 1.4 1.5 1.2 0.8 1.2 2.3 -0.50 2.7 3.0 -0.50 -0.50 median median median median median median median median median median median median median median median median 3.8890 3.3289 2.7154 2.9821 2.9861 2.0169 1.8042 1.3863 1.0835 1.4071 2.6423 0.13976 3.0301 3.2992 0.03922 0.00995 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.92271 3.33474 2.77421 3.03606 3.02150 2.00094 1.80982 1.42842 1.16610 1.43914 2.64599 0.07997 3.04846 3.34100 0.07432 0.10677 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.16376 0.16159 0.15226 0.18024 0.17573 0.22917 0.16059 0.17375 0.21849 0.15726 0.19361 0.26087 0.18839 0.17401 0.25490 0.25533

Platymantis neckeri – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.2 3.5 2.9 3.1 3.4 2.05 1.8 1.7 1.4 1.7 3.0 1.4 3.2 3.6 1.25 1.4

1.6 3.1 3.4 2.00 1.6 1.3 2.8 3.0 3.2 1.2 4.0 1.6 1.2 2.8 3.4 1.00 1.5 3.0 3.3 1.95 1.5 1.2 2.7 2.9 3.0 1.0 1.4 2.9 3.8 3.2 1.90 1.4 1.0 1.4 2.6 1.1 3.2 0.75 1.3 2.8 2.6 2.8 2.8 0.8 3.1 1.85 1.3 1.0 1.2 2.7 3.6 1.2 0.8 2.4 3.0 0.50 2.5 2.7 2.6 0.6 3.0 1.80 1.2 0.9 1.1 2.6 3.4 2.9 2.4 2.6 2.4 1.75 1.0 1.0 0.6 1.1 2.2 0.8 2.5 2.8 0.25 0.4 median median median median median median median median median median median median median median median median 3.8504 3.1843 2.6532 2.8893 2.9339 1.8840 1.6657 1.4303 0.9969 1.5096 2.6627 1.0626 2.8809 3.2186 0.7518 0.8788 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.84322 3.20454 2.66966 2.87842 2.92099 1.89568 1.53510 1.38940 0.99054 1.46674 2.65819 1.09429 2.86369 3.21047 0.73705 0.88279 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.203524 0.155477 0.157089 0.140776 0.212619 0.091720 0.296471 0.192021 0.221413 0.159851 0.229494 0.215648 0.202308 0.216760 0.261184 0.251006

293 Platymantis papuensis – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.1 3.45 2.85 3.2 3.2 2.2 1.9 1.8 1.5 1.7 2.7 0.3 3.1 3.4 0.4 0.5

4.0 3.40 2.80 1.8 1.7 1.4 1.6 0.2 0.4 3.1 3.1 2.1 0.3 2.6 3.0 3.3 2.75 1.6 1.3 1.5 0.3 3.9 3.35 1.7 0.1 3.0 3.0 2.0 0.2 2.70 1.5 1.2 1.4 0.2 2.5 2.9 3.2 3.8 3.30 1.6 -0.0 2.65 2.9 2.9 1.9 1.4 1.1 1.3 0.1 0.1 3.7 3.25 1.5 -0.1 2.4 2.8 3.1 2.60 1.3 1.0 1.2 -0.0 2.8 2.8 1.8 0.0 3.6 3.20 1.4 -0.2 2.55 1.2 0.9 1.1 -0.1 2.3 2.7 3.0 2.7 2.7 1.7 -0.1 3.5 3.15 2.50 1.3 1.1 0.8 1.0 -0.3 -0.2 3.4 3.10 2.45 2.6 2.6 1.6 1.2 1.0 0.7 0.9 2.2 -0.4 2.6 2.9 -0.2 -0.3 median median median median median median median median median median median median median median median median 3.8778 3.2910 2.7280 2.9837 2.9674 1.8924 1.7290 1.4350 1.2801 1.5085 2.5297 0.05353 2.9148 3.2751 0.20672 0.28123 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.86336 3.27723 2.70353 2.97696 2.95316 1.90640 1.68236 1.43141 1.24550 1.45219 2.51799 0.00525 2.89487 3.23882 0.17094 0.24983 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.14824 0.09256 0.10664 0.13049 0.12705 0.14164 0.15853 0.16669 0.18043 0.16605 0.11771 0.15516 0.09968 0.10183 0.14799 0.17443

Platymantis parkeri – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

2.95 2.10 1.7 1.95 1.00 0.50 0.55 0.6 0.8 1.4 -0.75 1.75 2.10 -0.6 -0.8

1.90 0.50 2.05 2.90 1.6 1.90 0.95 0.45 1.70 0.4 0.7 1.3 -1.00 -0.8 -1.0 2.05 0.45 2.00 2.85 1.5 1.85 0.90 0.40 1.65 1.85 0.2 0.6 1.2 -1.25 0.40 1.95 2.80 1.4 1.80 0.85 0.35 1.60 -1.0 -1.2 2.00 0.35 1.90 0.0 0.5 1.1 -1.50 1.80 2.75 1.3 1.75 0.80 0.30 1.55 0.30 1.85 -1.2 -1.4 -0.2 0.4 1.0 -1.75 2.70 1.2 1.70 0.75 0.25 1.50 1.95 1.75 0.25 1.80 2.65 1.1 1.65 0.70 0.20 0.20 -0.4 0.3 0.9 -2.00 1.45 1.75 -1.4 -1.6 median median median median median median median median median median median median - median median median - median - 2.7584 2.0470 1.5497 1.8602 1.8358 0.94585 0.37156 0.39204 0.02937 0.55389 1.1969 1.3471 1.5994 1.9516 1.2040 1.0498 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 2.76347 2.04068 1.53478 1.83809 1.82564 0.91216 0.35891 0.39465 0.01540 0.56101 1.16863 1.29284 1.61548 1.95574 1.13987 1.08094 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.07018 0.04215 0.11547 0.06021 0.07697 0.07693 0.06572 0.08177 0.22191 0.09674 0.12565 0.25943 0.07432 0.07886 0.18835 0.19024

294 Platymantis pelewensis – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.80 3.05 2.55 2.90 2.9 1.85 1.6 1.4 1.2 1.275 2.4 -0.0 2.75 3.10 0.10 0.4

2.85 1.80 0.05 3.75 2.50 2.70 3.05 0.3 3.00 1.5 1.250 2.8 1.3 1.1 2.3 -0.1 2.80 1.75 -0.00 3.70 2.45 2.65 3.00 0.2 2.95 1.4 1.225 2.75 1.70 -0.05 3.65 2.40 2.7 1.2 1.0 2.2 -0.2 2.60 2.95 0.1 2.70 1.65 -0.10 2.90 1.3 1.200 3.60 2.35 2.55 2.90 0.0 2.65 1.60 -0.15 2.6 1.1 0.9 2.1 -0.3 2.85 1.2 1.175 3.55 2.30 2.50 2.85 -0.1 2.60 1.55 -0.20 3.50 2.80 2.25 2.55 2.5 1.50 1.1 1.0 0.8 1.150 2.0 -0.4 2.45 2.80 -0.25 -0.2 median median median median median median median median median median median median - median median median - median - 3.5599 2.9418 2.3428 2.6603 2.6071 1.6174 1.3863 1.1600 0.9083 1.2030 2.3046 0.2357 2.6707 2.9982 0.1054 0.0619 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean 3.59045 2.92731 2.37610 2.69061 2.64274 1.66042 1.38741 1.19554 0.96616 1.22122 2.25953 0.19125 2.61778 2.96678 0.06099 0.03293 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.112776 0.088807 0.087174 0.106589 0.150946 0.115254 0.144939 0.123787 0.135425 0.044392 0.123615 0.134486 0.106478 0.099830 0.117217 0.170094

Platymantis schmidti – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.10 3.55 3.00 3.20 3.2 2.10 2.00 1.70 1.5 1.8 2.8 0.6 3.20 3.55 0.7 0.7

3.15 1.65 4.05 3.50 2.95 2.05 1.95 0.5 3.15 3.50 0.6 0.6 1.4 1.7 3.10 3.1 1.60 2.7 4.00 2.90 2.00 1.90 0.4 3.10 3.45 0.5 3.45 0.5 3.05 1.55 1.3 1.6 3.95 2.85 1.95 1.85 0.3 3.05 3.40 0.4 3.0 2.6 3.40 3.00 1.50 0.4 3.90 2.80 1.90 1.80 1.2 1.5 0.2 3.00 3.35 0.3 3.35 2.95 1.45 0.3 2.9 2.5 3.85 2.75 1.85 1.75 0.1 2.95 3.30 0.2 2.90 1.40 1.1 1.4 3.30 0.2 3.80 2.70 1.80 1.70 0.0 2.90 3.25 0.1 2.85 2.8 1.35 2.4 1.0 1.3 3.25 0.1 3.75 2.65 2.80 1.75 1.65 1.30 -0.1 2.85 3.20 0.0 3.70 3.20 2.60 2.75 2.7 1.70 1.60 1.25 0.9 1.2 2.3 -0.2 2.80 3.15 -0.1 0.0 median median median median median median median median median median median median median median median median 3.8877 3.3457 2.7915 2.9852 2.9104 1.8802 1.8009 1.4011 1.2541 1.4024 2.5568 0.29267 2.9696 3.3370 0.30010 0.37841 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.89398 3.35531 2.78934 2.97997 2.92729 1.89410 1.79606 1.41938 1.25805 1.42469 2.56304 0.28079 2.98219 3.33746 0.33034 0.39431 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.09802 0.09105 0.09468 0.09532 0.11463 0.08925 0.08609 0.09669 0.12853 0.13415 0.10322 0.14786 0.09717 0.09501 0.15615 0.15343

295 Platymantis solomonis – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.4 3.7 3.4 3.5 3.5 2.5 2.3 1.9 1.8 2.0 3.0 0.3 3.5 3.8 0.5 0.6

4.3 3.6 3.4 3.4 2.4 2.2 1.8 1.7 2.9 0.2 3.4 3.7 0.4

3.2 3.3 1.8 3.3 0.4 4.2 3.5 3.3 2.3 2.1 1.7 1.6 2.8 0.1 3.6 0.3 3.2 3.2 4.1 3.4 3.2 2.2 2.0 1.6 1.5 2.7 -0.0 3.5 0.2 3.0 1.6 0.2 3.1 3.1 4.0 3.3 3.1 2.1 1.9 1.5 1.4 2.6 -0.1 3.4 0.1 3.0 3.0 2.8 1.4 0.0 3.9 3.2 3.0 2.0 1.8 1.4 1.3 2.5 -0.2 3.3 -0.0 2.9 2.9 3.8 3.1 2.9 1.9 1.7 1.3 1.2 2.4 -0.3 3.2 -0.1 2.6 2.8 1.2 2.8 -0.2 3.7 3.0 2.8 2.7 1.8 1.6 1.2 1.1 2.3 -0.4 2.7 3.1 -0.2 3.6 2.9 2.4 2.7 2.6 1.7 1.5 1.1 1.0 1.0 2.2 -0.5 2.6 3.0 -0.3 -0.4 median median median median median median median median median median median median - median median median median 3.9923 3.3604 2.8416 3.1224 3.0426 2.0719 1.8878 1.5217 1.3863 1.5215 2.6333 0.1054 3.0922 3.4142 0.13976 0.24275 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean Mean 4.00082 3.34263 2.82525 3.09992 3.06137 2.06020 1.87815 1.51854 1.35434 1.49302 2.59755 0.10149 3.06780 3.38257 0.13146 0.20637 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.23167 0.20127 0.22622 0.21773 0.23613 0.22097 0.20462 0.20473 0.19661 0.26638 0.21344 0.22505 0.21007 0.19949 0.21672 0.24444

Platymantis spelaeus – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.15 3.50 2.950 3.125 3.200 2.10 1.775 1.600 1.8 1.75 2.9 1.0 3.07 3.43 1.0 0.70

1.70 1.575 2.8 0.9 0.9 4.10 3.48 3.100 3.175 1.7 3.06 3.42 0.65 2.925 1.750 2.05 1.65 1.550 2.7 0.8 0.8 4.05 3.46 3.075 3.150 1.6 3.05 3.41 0.60 1.60 2.900 1.725 1.525 2.6 0.7 0.7 2.00 1.55 4.00 3.44 3.050 3.125 1.5 3.04 3.40 0.55 1.500 2.5 0.6 0.6 1.50 2.875 1.700 3.95 3.42 3.025 3.100 1.4 3.03 3.39 0.50 1.475 2.4 0.5 0.5 1.95 1.45 3.90 3.40 2.850 3.000 3.075 1.675 1.450 1.3 1.40 2.3 0.4 3.02 3.38 0.4 0.45 median median median median median median median median median median median median median median median median 3.9972 3.4500 2.9020 3.0489 3.1224 1.9912 1.7579 1.5581 1.3788 1.5194 2.7609 0.53937 3.0325 3.3932 0.57647 0.53063 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 4.004225 3.452854 2.909920 3.054778 3.128093 2.001665 1.737902 1.543529 1.432661 1.544954 2.710830 0.626543 3.037468 3.397684 0.621286 0.543688 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.055251 0.024485 0.030164 0.041371 0.036650 0.052789 0.033902 0.051635 0.137135 0.097051 0.165276 0.189356 0.013928 0.016385 0.157028 0.071817

296 Platymantis vitianus – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.8 4.0 3.5 3.8 4.0 2.6 2.7 2.2 2.0 2.50 3.4 0.8 3.8 4.0 0.6 0.8

3.4 2.6 3.9 3.7 1.9 3.3 3.9 0.7 2.5 0.7 3.6 0.5 4.6 3.8 2.0 2.25 3.3 2.5 3.8 3.6 1.8 3.2 3.8 0.6 2.4 0.6 3.4 0.4 3.2 2.4 4.4 3.7 3.5 3.6 1.8 1.7 2.00 3.1 3.7 0.5 3.1 2.3 2.3 0.5 3.2 0.3 3.6 3.4 1.6 3.0 3.6 0.4 3.0 2.2 4.2 3.4 1.6 1.75 2.2 0.4 3.0 0.2 3.5 3.3 1.5 2.9 3.5 0.3 2.9 2.1 4.0 3.4 2.8 3.2 3.2 2.1 2.0 1.4 1.4 1.50 2.8 0.3 2.8 3.4 0.1 0.2 median median median median median median median median median median median median median median median median 4.4272 3.7241 3.2267 3.4861 3.5403 2.3105 2.2511 1.7182 1.5811 2.1333 3.0544 0.57170 3.4068 3.6338 0.48241 0.48586 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 4.397720 3.671815 3.181465 3.485410 3.519063 2.336220 2.306880 1.730833 1.612286 2.108796 3.024106 0.557581 3.332507 3.649221 0.429925 0.501782 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.227040 0.185254 0.208990 0.198030 0.225374 0.169261 0.188798 0.217526 0.190280 0.267790 0.176841 0.167363 0.256906 0.220731 0.152993 0.189279

Platymantis vitiensis – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.0 3.2 2.7 2.9 2.9 1.80 1.7 1.3 1.1 1.6 2.70 1.2 2.9 3.2 0.55 0.7

2.65 0.50 0.6 2.6 1.75 3.9 2.8 1.6 1.0 1.5 1.1 2.8 3.1 2.8 1.2 3.1 2.60 0.45 0.5 2.5 1.70 3.8 2.7 1.5 0.9 1.4 1.0 2.7 2.55 0.40 0.4 3.0 2.4 2.7 1.65 1.1 3.0 2.50 0.35 0.3 3.7 2.6 1.4 0.8 1.3 0.9 2.6 2.3 1.60 2.45 0.30 0.2 2.9 2.6 1.0 2.9 3.6 2.5 1.3 0.7 1.2 0.8 2.5 2.2 1.55 2.40 0.25 0.1 3.5 2.8 2.1 2.5 2.4 1.50 1.2 0.9 0.6 1.1 2.35 0.7 2.4 2.8 0.20 0.0 median median median median median median median median median median median median median median median median 3.6098 2.8881 2.3798 2.6358 2.6583 1.6184 1.4109 1.1019 0.8755 1.4279 2.4344 0.9183 2.5388 2.8626 0.41862 0.36111 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.64227 2.93909 2.39249 2.66057 2.66251 1.63090 1.42552 1.093978 0.87120 1.38695 2.47145 0.91851 2.57905 2.91184 0.36198 0.36251 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.12842 0.12040 0.14954 0.11520 0.12313 0.08423 0.11654 0.117585 0.15132 0.13898 0.09585 0.13383 0.12712 0.11844 0.11729 0.18058

297 Platymantis weberi – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.4 3.6 3.2 3.3 3.4 2.3 2.2 1.7 1.5 1.7 3.0 0.3 3.4 3.6 0.6 0.6

3.5 3.2 2.2 1.6 1.6 0.2 4.2 1.4 2.8 0.4 0.4 3.4 3.0 3.1 3.2 2.1 2.0 1.5 3.2 3.4 1.5 0.1 4.0 3.3 3.0 2.0 1.3 1.4 2.6 0.2 0.2 1.4 2.8 3.0 1.8 3.0 3.2 3.2 2.9 1.9 1.3 -0.0 3.8 1.3 1.2 2.4 -0.0 -0.0 3.1 2.8 1.8 1.2 -0.1 2.6 2.8 1.6 2.8 3.0 1.2 3.6 3.0 2.7 1.7 1.1 1.1 2.2 -0.2 -0.2 -0.2 1.1 2.9 2.4 2.6 2.6 1.6 1.4 1.0 2.6 2.8 3.4 1.0 2.0 -0.4 -0.4 -0.3 2.8 2.5 1.5 1.0 0.9 3.2 2.7 2.2 2.4 2.4 1.4 1.2 0.9 0.9 0.8 1.8 -0.4 2.4 2.6 -0.6 -0.6 median median median median median median median median median median median median median median median median 3.8982 3.2910 2.7259 2.9968 2.9712 1.9706 1.8625 1.4134 1.3015 1.3737 2.5757 0.13976 2.9339 3.2600 0.13976 0.23902 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.88992 3.29646 2.74617 3.00418 2.96638 1.98114 1.80378 1.41020 1.27408 1.37152 2.56324 0.06973 2.95945 3.28111 0.14973 0.23726 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.17152 0.16527 0.17129 0.16691 0.17178 0.17956 0.18468 0.17278 0.15967 0.18609 0.16669 0.16438 0.17080 0.17814 0.18286 0.22076

Rana kreffti – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.5 4.0 3.4 3.6 3.5 2.4 2.4 2.1 2.1 2.2 3.4 1.00 3.6 3.9 1.0 1.00

4.4 3.9 3.5 3.4 2.0 3.5 3.8 2.3 2.0 3.2 3.8 2.2 2.0 3.4 0.8 4.3 3.2 3.4 3.3 1.9 0.75 3.7 0.75 2.2 1.9 3.7 3.3 3.2 3.0 3.6 4.2 3.3 2.0 1.8 1.8 0.6 3.6 2.1 1.8 3.2 3.0 3.1 0.50 3.5 0.50 2.8 4.1 3.5 3.2 1.7 3.1 3.0 2.0 1.8 1.7 1.6 3.4 0.4 3.4 3.0 4.0 3.1 1.6 2.6 2.8 2.9 0.25 3.3 0.25 3.3 1.9 1.6 2.9 3.9 3.0 1.6 1.5 1.4 0.2 2.8 2.4 3.2 3.2 1.8 1.5 2.8 3.8 2.6 2.9 2.7 1.4 0.00 3.1 0.00 3.1 1.4 1.2 2.7 0.0 1.7 1.4 2.2 3.7 3.0 2.8 2.6 1.3 2.6 3.0 3.6 2.9 2.4 2.7 2.5 1.6 1.2 1.2 1.3 1.0 2.0 -0.25 2.5 2.9 -0.2 -0.25 median median median median median median median median median median median median median median median median 3.9200 3.3329 2.6899 3.0282 2.8662 1.8563 1.6371 1.6174 1.6074 1.4586 2.6254 0.28518 2.9014 3.2347 0.36426 0.36464 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.98873 3.36920 2.74641 3.07114 2.92778 1.94701 1.66559 1.64066 1.61340 1.49248 2.66372 0.34557 2.94689 3.28540 0.42253 0.41170 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.22819 0.20483 0.21739 0.20479 0.23572 0.20408 0.20393 0.19848 0.15667 0.23299 0.24572 0.29917 0.23676 0.21436 0.25211 0.29187

298 Rana novaebritanniae – females lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.3 3.55 2.80 3.4 3.2 2.3 1.9 1.650 1.8 1.65 3.0 0.6 3.25 3.6 0.6 0.6

3.50 3.20 4.2 3.3 2.2 1.625 1.60 3.1 1.8 2.9 0.5 3.5 2.75 1.7 0.5 0.5 3.45 3.15 4.1 3.2 2.1 1.600 1.55 3.0 1.7 2.8 0.4 3.4 3.40 3.10 4.0 2.70 3.1 2.0 1.575 1.6 1.50 0.4 0.4 3.35 3.05 2.9 1.6 2.7 0.3 3.3 3.9 3.0 1.9 1.550 1.45 3.30 3.00 2.65 1.5 0.3 0.3 2.8 1.5 2.6 0.2 3.2 3.8 2.9 1.8 1.525 1.40 3.25 2.95 3.7 3.20 2.60 2.8 2.7 1.7 1.4 1.500 1.4 1.35 2.5 0.1 2.90 3.1 0.2 0.2 median median median median median median median median median median median median median median median median 3.9154 3.2697 2.6497 3.0017 2.8344 1.8749 1.5581 1.5892 1.6054 1.4563 2.6658 0.34359 2.9544 3.2542 0.52466 0.44469 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.93087 3.30537 2.673694 3.02127 2.85887 1.88889 1.574152 1.581965 1.60932 1.453881 2.697343 0.358595 2.964125 3.277372 0.476293 0.418933 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.12415 0.08651 0.054357 0.10999 0.10832 0.13200 0.154465 0.045088 0.08825 0.083549 0.121784 0.131449 0.093857 0.106736 0.112456 0.120413

299

Batrachylodes elegans – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.50 2.700 2.35 2.55 2.6 1.5 1.10 1.25 0.8 1.5 2.15 0.7 2.45 2.80 0.3 0.4

3.45 2.30 1.05 2.10 2.75 0.3 2.675 2.50 1.4 1.20 1.4 0.2 2.5 0.7 0.6 2.40 3.40 2.25 1.00 2.05 2.70 0.2 2.650 2.45 1.3 1.15 1.3 0.1

3.35 2.20 2.4 0.95 0.6 2.00 0.5 2.35 2.65 0.1

2.625 2.40 1.2 1.10 1.2 0.0 3.30 2.15 0.90 1.95 2.60 0.0

2.3 0.5 0.4 2.30 2.600 2.35 1.05 -0.1 3.25 2.10 1.1 0.85 1.1 1.90 2.55 -0.1

3.20 2.575 2.05 2.30 0.80 1.00 1.85 2.25 2.50 2.2 1.0 0.4 1.0 0.3 -0.2 -0.2 median median median median median median median median median median median median median median median median 3.3463 2.6308 2.1535 2.3983 2.3659 1.2850 0.9419 1.1167 0.61218 1.2075 1.9865 0.53642 2.3471 2.6090 0.18232 0.13972 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.35087 2.63908 2.15702 2.39884 2.36246 1.27714 0.95252 1.12560 0.62069 1.19529 1.99144 0.52943 2.34586 2.61425 0.12084 0.15640 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.07263 0.02940 0.05651 0.04704 0.08355 0.08670 0.07156 0.07067 0.10738 0.10007 0.07119 0.09006 0.05309 0.06012 0.12790 0.13045

Batrachylodes mediodiscus – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.200 2.550 2.050 2.250 2.25 1.3 0.9 1.05 0.8 1.10 1.75 0.1 2.25 2.8 -0.0 0.2

2.525 2.025 2.225 1.05 1.70 0.1 1.2 0.8 1.00 0.7 -0.0 2.7 -0.1 3.175 2.20 2.20 2.500 2.000 2.200 1.00 1.65 -0.0 1.1 0.7 0.95 0.6 -0.1 2.6 -0.2

3.150 2.475 1.975 2.175 2.15 0.95 1.60 -0.1 2.15 1.0 0.6 0.90 0.5 -0.2 2.5 -0.3 2.450 1.950 2.150 0.90 1.55 -0.2 3.125 2.10 0.9 0.5 0.85 0.4 -0.3 2.4 -0.4 0.85 1.50 2.425 1.925 2.125 2.10 -0.3

3.100 2.400 1.900 2.100 2.05 0.8 0.4 0.80 0.3 0.80 1.45 -0.4 2.3 -0.5 -0.4 median median median median median median median median median median median median - median median median - median - 3.1241 2.4427 1.9727 2.1650 2.1616 1.1003 0.61771 0.9243 0.61248 0.9477 1.6292 0.1166 2.1436 2.4305 0.1425 0.0256 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.13172 2.45909 1.97064 2.17160 2.15640 1.08700 0.63253 0.93127 0.60241 0.94701 1.63570 0.11267 2.15098 2.45786 0.17442 0.04951 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.02813 0.04658 0.03795 0.03861 0.04623 0.11075 0.11994 0.05819 0.12894 0.08551 0.06633 0.13446 0.04113 0.10492 0.15615 0.14030

300

Batrachylodes minutus – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

2.92 2.200 1.80 1.9425 1.950 0.82 0.6 0.72 0.50 1.0 1.450 -0.6 1.925 2.175 -0.45 -0.300

-0.325 2.175 1.78 1.9400 1.925 0.5 1.425 1.900 -0.50 2.90 0.80 0.71 0.45 0.9 -0.8 2.150 -0.350

2.150 1.76 1.9375 1.900 0.4 1.400 1.875 -0.55

-0.375 2.88 0.78 0.70 0.40 0.8 -1.0 2.125 2.125 1.74 1.9350 1.875 0.3 1.375 1.850 -0.60 -0.400

2.86 2.100 1.72 1.9325 1.850 0.76 0.2 0.69 0.35 0.7 1.350 -1.2 1.825 2.100 -0.65 -0.425 median median median median median median median median median median median median - median median median - median - 2.8987 2.1541 1.7422 1.9402 1.8779 0.78390 0.36464 0.71295 0.36464 0.91228 1.4422 0.8916 1.9036 2.1353 0.6162 0.3567 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 2.89588 2.15373 1.75301 1.93825 1.89232 0.78831 0.36486 0.70797 0.39510 0.87073 1.41676 0.87214 1.88360 2.13622 0.57555 0.35728 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.025011 0.032495 0.026675 0.003328 0.028999 0.020766 0.141837 0.013075 0.065243 0.136932 0.046114 0.246814 0.035923 0.030107 0.070373 0.042887

Batrachylodes trossulus – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.025 2.40 2.00 2.000 2.00 0.900 0.7 0.90 0.6 0.925 1.50 -0.6 2.025 -0.25 -0.15

2.30 -0.20 3.000 1.975 0.900 -0.30 1.95 0.875 0.5 1.45 -0.7 2.000 1.95 0.6 0.85 2.35 -0.25 2.975 1.950 0.875 -0.35 1.90 0.850 0.4 1.40 -0.8 1.975 2.25 -0.30 2.950 1.925 1.90 0.5 0.80 0.850 -0.40 2.30 -0.35 1.85 0.825 0.3 1.35 -0.9 1.950 2.925 1.900 0.825 -0.45 2.20 -0.40 1.85 0.4 0.75 1.80 0.800 0.2 1.30 -1.0 1.925 1.875 0.800 -0.50 2.900 2.25 -0.45 2.875 1.75 1.850 1.80 0.775 0.3 0.70 0.1 0.775 1.25 -1.1 1.900 2.15 -0.55 -0.50 median median median median median median median median median median median median - median median median - median - 2.9574 2.3081 1.8254 1.9206 1.8863 0.85866 0.44776 0.77011 0.36116 0.83291 1.3623 0.7995 1.9579 2.2332 0.3285 0.3156 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 2.95024 2.31160 1.84094 1.92239 1.87794 0.85154 0.46706 0.78064 0.35096 0.84041 1.37194 0.83574 1.96276 2.23274 0.34465 0.33434 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.048170 0.039295 0.059513 0.041473 0.055155 0.041179 0.108100 0.046836 0.096283 0.037330 0.078750 0.146163 0.038871 0.044997 0.071721 0.115962

301

Batrachylodes vertebralis – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.4 2.70 2.25 2.45 2.35 1.2 1.20 0.9 1.2 2.1 0.6 2.40 2.65 0.3 0.2

2.20 2.40 1.4 1.15 0.8 2.35 2.60 2.65 2.30 1.1 2.0 0.5 0.2 0.1 3.3 2.15 2.35 1.10 0.7 1.1 2.30 2.55 2.60 1.0 1.9 0.1 -0.0 2.25 1.3 0.4 3.2 2.10 2.30 1.05 0.6 2.25 2.50 2.55 0.9 1.8 -0.0 -0.1 1.0 2.20 0.3 2.05 2.25 1.2 1.00 0.5 2.20 2.45 3.1 2.50 0.8 1.7 -0.1 -0.2 2.00 2.20 2.15 0.95 0.4 0.2 2.15 2.40 0.9 2.45 1.1 0.7 1.6 -0.2 -0.3 3.0 1.95 2.15 0.90 0.3 2.10 2.35 2.10 0.1 2.40 0.6 1.5 -0.3 -0.4 1.90 2.10 0.85 0.2 0.8 2.05 2.30 2.9 1.0 2.05 0.0 2.35 1.85 2.05 0.5 0.80 0.1 1.4 2.00 2.25 -0.4 -0.5 2.8 2.30 1.80 2.00 2.00 0.9 0.4 0.75 0.0 0.7 1.3 -0.1 1.95 2.20 -0.5 -0.6 median median median median median median median median median median median median median median median - median - 3.2268 2.4857 2.0149 2.1939 2.2138 1.1817 0.8329 0.9746 0.50078 0.9594 1.8374 0.35066 2.2072 2.4204 0.0202 0.0408 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean - 3.21564 2.51958 2.03943 2.22974 2.20507 1.18540 0.80857 0.98227 0.49899 0.95827 1.80518 0.31745 2.20059 2.44946 0.05229 0.08205 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.09472 0.10078 0.10708 0.12028 0.08326 0.12872 0.17356 0.11801 0.23512 0.08568 0.16966 0.15405 0.09722 0.11602 0.16201 0.21291

Batrachylodes wolfi – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

2.150 2.38 2.425 1.15 1.10 0.80 2.05 0.60 2.400 2.600 0.3 0.3

1.20 3.35 2.65 1.35 1.10 0.75 0.2 2.125 2.36 2.400 1.05 2.375 2.575 0.55 0.2 2.00 1.05 0.70 0.1 2.100 2.34 2.375 1.00 1.15 2.350 2.550 1.30 0.1 3.30 2.60 1.00 0.65 0.50 -0.0 2.075 2.32 2.350 0.95 1.95 2.325 2.525 0.95 0.60 1.10 -0.1 1.25 0.0 2.050 2.30 2.325 0.90 0.45 2.300 2.500 3.25 2.55 0.90 0.55 -0.2 1.90 1.05 -0.1 2.025 2.28 2.300 0.85 2.275 2.475 0.85 0.50 -0.3 1.20 0.40 3.20 2.50 2.000 2.26 2.275 0.80 0.80 0.45 1.00 1.85 2.250 2.450 -0.2 -0.4 median median median median median median median median median median median median median median median median 3.3164 2.5859 2.0567 2.3389 2.3599 1.3083 0.9987 0.9708 0.60701 1.1184 1.9344 0.47933 2.3046 2.5537 0.09527 0.11777 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.30863 2.58671 2.06135 2.33599 2.35872 1.30149 0.98317 0.96614 0.62225 1.12712 1.93534 0.48664 2.30950 2.53981 0.08563 0.09706 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.03915 0.04012 0.03268 0.02929 0.03132 0.04570 0.07229 0.06721 0.10444 0.05110 0.04589 0.05168 0.03745 0.03722 0.12910 0.12847

302

Ceratobatrachus guentheri – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.3 3.4 2.8 3.6 3.8 2.2 2.2 1.6 1.7 2.2 2.8 0.3 3.1 3.5 0.5 0.5

3.5 2.1 0.2 4.2 3.3 2.1 1.5 1.6 2.7 3.0 3.4 0.4 0.4 2.7 3.4 3.6 2.0 2.0 0.1 4.1 1.4 2.6 2.9 3.3 0.3 3.2 2.0 1.5 0.3 2.6 3.3 1.9 -0.0 4.0 1.3 2.5 2.8 3.2 0.2 3.4 1.8 3.1 3.2 1.8 1.9 1.4 -0.1 0.2 3.9 2.5 1.2 2.4 2.7 3.1 0.1 3.0 3.1 1.7 1.8 1.3 -0.2 0.1 3.2 1.6 3.8 1.1 2.3 2.6 3.0 -0.0 2.4 3.0 1.6 -0.3 2.9 1.7 1.2 0.0 3.7 1.0 2.2 2.5 2.9 -0.1 2.9 3.0 1.5 1.4 -0.4 2.3 2.8 1.6 1.1 -0.1 3.6 2.8 1.4 0.9 2.1 -0.5 2.4 2.8 -0.2 3.5 2.7 2.2 2.7 2.8 1.3 1.5 0.8 1.0 1.2 2.0 -0.6 2.3 2.7 -0.3 -0.2 median median median median median median median median median median median median median median median median 4.0490 3.3078 2.7100 3.2834 3.3673 1.9022 2.0189 1.3661 1.4207 1.5354 2.6848 0.05353 2.9447 3.2949 0.18647 0.28507 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean Mean 3.98053 3.22120 2.64068 3.25446 3.32212 1.84588 1.97794 1.29645 1.36323 1.59509 2.57000 0.01064 2.84625 3.20613 0.17485 0.22708 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.20733 0.20675 0.19108 0.20778 0.23412 0.24490 0.14735 0.20800 0.18859 0.26921 0.22842 0.19548 0.23232 0.22056 0.22495 0.23168

Discodeles bufoniformis – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.4 3.6 3.0 3.5 3.6 2.5 2.1 1.8 1.5 1.8 3.0 0.6 3.4 3.6 1.00 0.8

4.3 3.5 2.9 3.5 2.9 3.4 2.4 2.0 1.7 3.3 3.5 1.7 1.4 0.4 0.75 0.6 4.2 3.4 2.8 3.4 2.8 3.3 2.3 1.9 1.6 3.2 3.4 1.6 1.3 0.2 4.1 3.3 2.7 3.3 2.7 3.2 2.2 1.8 1.5 3.1 3.3 0.50 0.4 4.0 3.2 2.6 1.5 1.2 0.0 3.2 2.6 3.1 2.1 1.7 1.4 3.0 3.2 3.9 3.1 2.5 0.25 0.2 1.4 1.1 -0.2 3.1 2.5 3.0 2.0 1.6 1.3 2.9 3.1 3.8 3.0 2.4

3.7 3.0 2.4 2.9 2.9 1.9 1.5 1.3 1.0 1.2 2.3 -0.4 2.8 3.0 0.00 0.0 median median median median median median median median median median median median median median median median 4.0411 3.2316 2.6783 3.2015 3.2321 2.1558 1.8611 1.4748 1.3686 1.5251 2.6550 0.00498 3.0532 3.2905 0.46648 0.35758 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 4.05941 3.24994 2.72037 3.21683 3.27968 2.15790 1.85409 1.50815 1.30932 1.50313 2.67425 0.05431 3.07885 3.31272 0.45498 0.38717 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.19987 0.17792 0.18534 0.17070 0.19472 0.14728 0.15876 0.12726 0.12950 0.20835 0.17570 0.21687 0.18326 0.16966 0.22620 0.21277

303

Discodeles guppyi – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.75 4.25 3.75 3.75 3.75 2.75 2.50 2.25 1.50 2.25 3.4 1.2 4.00 4.25 1.2 1.2

3.50 2.00 3.2 1.0 3.75 4.00 1.0 1.0 4.50 4.00 3.50 3.50 2.50 2.25 2.00 1.25

3.25 1.75 3.0 0.8 3.50 3.75 0.8 0.8

4.25 3.75 3.25 3.25 2.25 2.00 1.75 1.00

3.00 1.50 2.8 0.6 3.25 3.50 0.6 0.6

4.00 3.50 3.00 3.00 2.00 1.75 1.50 0.75 2.75 1.25 2.6 0.4 3.00 3.25 0.4 0.4

3.75 3.25 2.50 2.75 2.75 1.75 1.50 1.25 0.50 1.00 2.4 0.2 2.75 3.00 0.2 0.2 median median median median median median median median median median median median median median median median 4.2243 3.6507 3.0745 3.3109 3.2711 2.3115 2.0844 1.6563 1.1969 1.7210 2.9444 0.8544 3.3868 3.6454 0.9517 0.9322 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 4.248591 3.697225 3.077279 3.332078 3.290866 2.291767 2.127069 1.722509 1.182796 1.699664 2.938303 0.818345 3.402650 3.658088 0.819930 0.857678 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.260651 0.263747 0.291584 0.256620 0.243235 0.272098 0.257211 0.251400 0.263156 0.261532 0.217880 0.204761 0.266976 0.272086 0.210389 0.198400

Palmatorappia solomonis – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.5 2.85 2.40 2.45 2.55 1.50 1.4 1.0 0.7 1.40 2.2 0.9 2.35 2.65 0.8 1.0

2.35 2.40 2.50 1.45 1.3 0.6 0.8 2.30 2.60 0.7 0.9 3.4 2.80 1.35 2.1 0.9 2.35 1.40 0.7 2.30 2.45 1.2 0.5 2.25 2.55 0.6 0.8 3.3 2.75 1.30 2.0 2.30 1.35 0.6 0.8 2.25 2.40 1.1 0.4 2.20 2.50 0.5 0.7 3.2 2.70 2.25 1.30 1.25 1.9 0.5 2.20 2.35 1.0 0.3 2.15 2.45 0.4 0.6 0.7 2.20 1.25 0.4 3.1 2.65 1.20 1.8 2.15 2.30 0.9 0.2 2.10 2.40 0.3 0.5 2.15 1.20 0.3 0.6 3.0 2.60 1.15 1.7 2.10 2.10 2.25 1.15 0.8 0.1 0.2 2.05 2.35 0.2 0.4 2.9 2.55 2.05 2.05 2.20 1.10 0.7 0.5 0.0 1.10 1.6 0.1 2.00 2.30 0.1 0.3 median median median median median median median median median median median median median median median median 3.3700 2.7828 2.2945 2.3263 2.4643 1.3737 1.1506 0.87963 0.38526 1.2542 2.0788 0.70802 2.2370 2.5221 0.38861 0.63656 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.33675 2.76310 2.28604 2.30592 2.43824 1.34168 1.15225 0.85279 0.39988 1.26304 2.04231 0.61951 2.22637 2.51229 0.39635 0.61438 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.10772 0.06227 0.06696 0.08061 0.08012 0.09251 0.12894 0.10758 0.13013 0.06534 0.12055 0.18405 0.08333 0.08134 0.13837 0.13656

304

Platymantis corrugata – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.8 3.2 2.7 2.8 2.9 1.6 1.5 1.6 1.2 1.5 2.4 0.00 2.8 3.1 0.1 0.2

3.7 1.4 2.3 -0.0 0.1 3.1 2.6 2.7 2.8 1.5 1.5 1.4 2.7 3.0 1.0 -0.25 3.6 1.3 2.2 -0.1 -0.0 3.0 2.5 2.6 2.7 1.4 1.4 1.3 2.6 2.9

3.5 1.2 0.8 2.1 -0.50 -0.2 -0.1

2.9 2.4 2.5 2.6 1.3 1.3 1.2 2.5 2.8 3.4 1.1 2.0 -0.3 -0.2 0.6 -0.75 2.8 2.3 2.4 2.5 1.2 1.2 1.1 2.4 2.7 3.3 1.0 1.9 -0.4 -0.3

3.2 2.7 2.2 2.3 2.4 1.1 0.9 1.1 0.4 1.0 1.8 -1.00 2.3 2.6 -0.5 -0.4 median median median median median median median median median median median median - median median median - median - 3.4642 2.9413 2.3805 2.5734 2.6575 1.4586 1.2413 1.3324 0.8286 1.2809 2.0656 0.3857 2.4899 2.8220 0.1165 0.0305 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 3.45524 2.95977 2.40634 2.56981 2.63151 1.40212 1.22346 1.31988 0.79738 1.25975 2.08930 0.38717 2.49908 2.83267 0.13237 0.04040 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.11048 0.08990 0.10635 0.09872 0.11124 0.11456 0.13760 0.10989 0.19358 0.13459 0.11245 0.18113 0.09917 0.10738 0.13375 0.14213

Platymantis dorsalis – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.45 2.95 2.400 2.55 2.60 1.6 1.3 1.25 1.0 1.10 2.20 -0.1 2.55 2.90 -0.10 -0.05

2.375 -0.2 -0.10 2.90 2.50 1.5 1.2 0.9 2.15 -0.15 3.40 2.55 1.20 1.05 2.50 2.85 2.350 -0.3 -0.15 2.85 2.45 1.4 1.1 0.8 2.10 -0.20

3.35 2.325 2.50 1.15 1.00 -0.4 2.45 -0.20 2.80 2.80 2.40 1.3 1.0 0.7 2.05 -0.25 2.300 -0.5 -0.25 3.30 2.45 1.10 0.95 2.40 2.75 2.35 1.2 0.9 0.6 2.00 -0.30 -0.30 2.275 -0.6 2.75

3.25 2.70 2.250 2.30 2.40 1.1 0.8 1.05 0.5 0.90 1.95 -0.7 2.35 -0.35 -0.35 median median median median median median median median median median median median - median median median - median - 3.3690 2.8954 2.3504 2.4553 2.5088 1.4159 1.1019 1.1663 0.87963 1.0080 2.0807 0.4155 2.4940 2.8449 0.1985 0.2231 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 3.36124 2.86317 2.33907 2.45197 2.49003 1.39862 1.09404 1.15332 0.82923 1.01490 2.07960 0.40103 2.47802 2.82845 0.20999 0.18667 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.062925 0.069547 0.044999 0.073739 0.061820 0.128797 0.136826 0.064711 0.161257 0.066841 0.067875 0.198322 0.066019 0.058286 0.102667 0.113053

305

Platymantis guentheri – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.60 2.925 2.400 2.65 2.70 1.7 1.40 1.15 0.8 1.3 2.50 1.05 2.600 0.35 0.45

1.35 2.90 0.40 2.900 1.00 2.575 3.55 2.375 2.60 2.65 0.30 1.6 1.10 0.7 1.2 2.45 1.30 0.35 2.875 0.95 2.550 3.50 2.350 2.55 2.60 0.25 1.25 2.85 0.30 2.850 1.5 1.05 0.6 1.1 2.40 0.90 2.525 1.20 0.25 3.45 2.325 2.50 2.55 0.20 2.825 0.85 2.500 1.15 2.80 0.20 1.4 1.00 0.5 1.0 2.35 3.40 2.300 2.45 2.50 0.15 2.800 0.80 2.475 1.10 0.15 3.35 2.775 2.275 2.40 2.45 1.3 1.05 0.95 0.4 0.9 2.30 0.75 2.450 2.75 0.10 0.10 median median median median median median median median median median median median median median median median 3.4826 2.8696 2.3665 2.5467 2.5900 1.5138 1.2323 1.0346 0.59056 1.2119 2.4014 0.8431 2.5369 2.8469 0.18145 0.19474 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.47210 2.86455 2.35683 2.53486 2.58652 1.51272 1.22299 1.04124 0.58527 1.17131 2.40269 0.87338 2.53659 2.84582 0.20106 0.22984 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.064283 0.031489 0.033256 0.064509 0.067537 0.102073 0.100702 0.061849 0.095196 0.104776 0.049887 0.096697 0.039200 0.041634 0.080864 0.107870

Platymantis guppyi – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.3 3.6 3.1 3.3 3.5 2.2 2.2 1.6 1.75 2.1 3.1 1.75 3.3 3.6 1.4 1.6

3.4 2.0 4.2 3.5 3.0 2.1 3.0 3.2 3.5 3.2 1.5 1.4 2.0 1.50 1.50 1.2 3.3 1.9 4.1 3.4 2.9 2.0 2.9 3.1 3.4 3.1 1.4 1.2 3.2 1.8 4.0 3.3 2.8 1.9 1.8 1.25 2.8 1.25 3.0 3.3 1.0 3.1 1.7 3.0 1.3 1.0 3.9 3.2 2.7 1.8 2.7 2.9 3.2 3.0 1.6 1.6 1.00 1.00 0.8 2.9 1.2 0.8 3.8 3.1 2.6 1.7 2.6 2.8 3.1 2.9 1.5 3.7 3.0 2.5 2.8 2.8 1.6 1.4 1.1 0.75 1.4 2.5 0.75 2.7 3.0 0.6 0.6 median median median median median median median median median median median median median median median median 4.0108 3.3182 2.7383 3.1028 3.1381 1.8946 1.9516 1.2920 1.3610 1.8342 2.7020 1.2837 2.9528 3.2944 0.9400 1.0852 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 4.00100St 3.33357 2.76084 3.07105 3.13994 1.87445 1.87414 1.31628 1.28514 1.77695 2.74178 1.27637 2.96651 3.31025 0.97453 1.08264 d Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.170905 0.174094 0.173502 0.149988 0.188006 0.132950 0.205887 0.125430 0.234252 0.216920 0.189926 0.256282 0.191860 0.187371 0.234785 0.250377

306

Platymantis hazelae – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.35 2.8 2.3 2.35 2.45 1.5 1.2 0.90 0.6 1.3 2.15 0.7 2.4 2.7 0.25 0.50

0.5 3.30 2.40 2.10 2.2 1.4 0.85 1.2 0.6 0.20 0.45 2.7 2.30 1.1 2.3 2.6 0.4 3.25 2.35 2.05 2.1 1.3 0.80 1.1 0.5 0.15 0.40 0.3 3.20 2.6 2.25 2.30 1.0 2.00 2.2 2.5 0.2 2.0 1.2 0.75 1.0 0.4 0.10 0.35 3.15 2.25 1.95 0.1 2.5 2.20 0.9 2.1 2.4 1.9 1.1 0.70 0.9 0.3 0.05 0.30 3.10 2.20 1.90 0.0

3.05 2.4 1.8 2.15 2.15 1.0 0.8 0.65 -0.1 0.8 1.85 0.2 2.0 2.3 0.00 0.25 median median median median median median median median median median median median median median median median 3.2981 2.6348 2.1587 2.2885 2.3466 1.3297 0.9746 0.79299 0.17395 0.9969 2.0516 0.50682 2.2246 2.5533 0.19062 0.41211 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.27584 2.62385 2.12691 2.28345 2.34321 1.31939 0.98905 0.78403 0.20445 1.01266 2.03700 0.50358 2.20773 2.53124 0.16422 0.40809 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.082394 0.078700 0.113003 0.047166 0.076649 0.108420 0.101798 0.044686 0.188025 0.138655 0.069380 0.117313 0.083039 0.084828 0.071429 0.064347

Platymantis mimicus – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.7 3.1 2.6 2.9 2.8 1.8 1.6 1.20 1.4 1.3 2.5 -0.1 2.8 3.2 -0.05 -0.05

1.7 1.5 1.15 2.4 3.6 2.8 2.7 3.1 3.0 2.5 1.2 2.7 -0.10 1.2 -0.2 -0.10 1.6 1.4 1.10 2.3 3.5 2.7 2.6 3.0 2.9 2.4 1.1 2.6 -0.15 1.5 1.3 1.05 2.2 3.4 2.6 2.5 1.0 -0.3 2.9 -0.15 1.4 1.2 1.00 2.1 2.8 2.3 1.0 2.5 -0.20 3.3 2.5 2.4 2.8 1.3 1.1 0.95 2.0 0.8 -0.4 -0.20 2.7 2.2 0.9 2.4 -0.25 3.2 2.4 2.3 2.7 1.2 1.0 0.90 1.9 3.1 2.6 2.1 2.3 2.2 1.1 0.9 0.85 0.6 0.8 1.8 -0.5 2.3 2.6 -0.30 -0.25 median median median median median median median median median median median median - median median median - median - 3.5193 2.9887 2.4327 2.6218 2.5565 1.4586 1.3686 1.0852 0.8920 1.0852 2.2628 0.2614 2.6624 2.9740 0.1863 0.1054 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 3.44269 2.89232 2.35587 2.55751 2.49147 1.44072 1.31161 1.04297 0.94540 1.04763 2.13065 0.28728 2.55261 2.88330 0.17180 0.11533 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.188140 0.158972 0.155831 0.203121 0.186493 0.214184 0.207386 0.127197 0.263192 0.179461 0.238492 0.111435 0.206780 0.190415 0.076912 0.068862

307

Platymantis myersi – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.1 3.5 3.00 3.3 3.2 2.25 1.9 1.60 1.5 1.46 3.0 0.50 3.3 3.6 0.3 0.3

1.55 1.4 3.2 3.5 0.2 4.0 3.4 2.75 3.2 3.1 0.2 2.00 1.8 1.44 2.8 0.25 1.50 1.3 3.1 3.4 0.1

3.9 3.3 2.50 3.1 3.0 0.1

1.45 1.2 3.0 3.3 0.0 1.75 1.7 1.42 2.6 0.00 3.8 3.2 2.25 3.0 2.9 0.0 1.40 1.1 2.9 3.2 -0.1

3.7 3.1 2.00 2.9 2.8 1.50 1.6 1.35 1.0 1.40 2.4 -0.25 2.8 3.1 -0.2 -0.1 median median median median median median median median median median median median median median median median - 3.9984 3.4102 2.8058 3.0634 3.0277 1.9601 1.8165 1.4110 1.3218 1.4446 2.7955 0.10436 3.1050 3.4171 0.05827 0.0202 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.93891 3.32690 2.63177 3.06640 3.02167 1.95954 1.77445 1.44437 1.25204 1.43809 2.68373 0.09983 3.05760 3.36035 0.03695 0.06711 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.148187 0.171361 0.393243 0.135675 0.119553 0.287800 0.137009 0.099276 0.193651 0.022153 0.237081 0.260126 0.183231 0.193327 0.189715 0.169406

Platymantis neckeri – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.85 3.20 2.65 2.9 2.95 2.1 1.7 1.5 1.2 1.50 2.7 1.2 2.85 3.3 1.0 0.9

3.80 1.45 1.1 2.80 3.15 2.90 1.6 1.4 0.8 0.8 2.60 2.8 2.0 1.1 2.6 3.2 3.75 1.40 1.0 2.75 3.10 2.85 1.5 1.3 0.6 0.7

3.70 2.55 2.7 1.9 1.0 1.35 2.5 0.9 2.70 3.1

3.05 2.80 1.4 1.2 0.4 0.6 3.65 1.30 0.8 2.65 2.50 2.6 1.8 0.9 2.4 3.0 3.00 2.75 1.3 1.1 0.2 0.5 3.60 1.25 0.7 2.60

3.55 2.95 2.45 2.5 2.70 1.7 1.2 1.0 0.8 1.20 2.3 0.6 2.55 2.9 0.0 0.4 median median median median median median median median median median median median median median median median 3.7129 3.0711 2.5265 2.8139 2.8015 1.8833 1.4303 1.2494 0.9744 1.3489 2.5131 0.9730 2.7473 3.0845 0.54519 0.68733 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.70032 3.06800 2.52260 2.78586 2.80924 1.86543 1.42474 1.27004 0.97694 1.34826 2.50210 0.97092 2.70967 3.05373 0.52375 0.66031 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.10302 0.07407 0.05161 0.11555 0.07640 0.11431 0.12706 0.14068 0.09493 0.09976 0.09977 0.15838 0.09605 0.10323 0.21818 0.19059

308

Platymantis papuensis – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.75 3.100 2.600 2.90 2.85 1.85 1.65 1.5 1.4 1.45 2.40 0.1 3.15 0.2 0.3

2.75 2.575 1.80 1.60 1.40 -0.0 3.075 2.80 1.4 0.2 3.70 2.85 1.3 2.35 3.10 0.1 2.550 1.75 1.55 1.35 -0.1 3.050 2.75 1.3 0.1 2.70 3.65 2.525 2.80 1.70 1.50 1.2 1.30 2.30 -0.2 3.05 0.0

3.025 2.70 1.2 0.0 2.500 1.65 1.45 1.25 -0.3 2.65 3.60 2.75 1.1 2.25 3.00 -0.1 3.000 2.65 1.1 -0.1 2.475 1.60 1.40 1.20 -0.4

3.55 2.975 2.450 2.70 2.60 1.55 1.35 1.0 1.0 1.15 2.20 -0.5 2.60 2.95 -0.2 -0.2 median median median median median median median median median median median median - median median median - median 3.6576 3.0727 2.5249 2.7562 2.7575 1.7492 1.5282 1.2179 1.1346 1.2413 2.3016 0.1744 2.6912 3.0397 0.0408 0.09531 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean 3.64880 3.06343 2.52970 2.77943 2.74114 1.72295 1.51801 1.22857 1.15489 1.25571 2.29958 0.18552 2.69749 3.04565 0.03403 0.09600 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.062637 0.036551 0.049321 0.060958 0.074374 0.082270 0.07595 0.130412 0.097892 0.089972 0.048360 0.160752 0.045706 0.055173 0.107924 0.103725

Platymantis parkeri – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

2.90 2.050 1.70 1.90 2.00 0.95 0.5 0.5 0.6 0.65 1.4 -1.0 1.75 2.05 -0.8 -0.8

1.65 0.60 2.025 0.90 -1.1 1.70 -0.9 2.85 1.95 0.4 0.4 2.00 -1.0 1.85 0.4 1.3 1.60 0.55 2.000 0.85 -1.2 1.65 -1.0 2.80 1.90 0.3 0.3 1.95 -1.2 1.55 0.50 1.975 1.80 0.80 0.2 1.2 -1.3 1.60 -1.1 1.50 0.45 2.75 1.85 0.2 0.2 1.90 -1.4 1.950 0.75 -1.4 1.55 -1.2 1.45 0.40 1.75 0.0 1.1 2.70 1.80 0.1 0.1 1.85 -1.6 0.70 1.50 1.925 1.40 0.35 -1.5 -1.3

2.65 1.900 1.35 1.70 1.75 0.65 0.0 0.0 -0.2 0.30 1.0 -1.6 1.45 1.80 -1.4 -1.8 median median median median median median median median median median median median - median median median - median - 2.7441 1.9892 1.5347 1.8485 1.8421 0.90016 0.30010 0.37844 0.03922 0.56531 1.2000 1.3471 1.6014 1.9502 1.1087 1.0217 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 2.74314 1.97515 1.53954 1.82297 1.84253 0.86783 0.31816 0.32156 0.12158 0.51986 1.21524 1.33585 1.60334 1.94887 1.12250 1.12839 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.068228 0.046231 0.085907 0.065418 0.073315 0.099366 0.119998 0.144460 0.219815 0.098487 0.104384 0.155521 0.069850 0.070823 0.150014 0.248081

309

Platymantis pelewensis – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.45 2.75 2.20 1.55 1.25 1.10 1.00 1.20 2.10 -0.1 2.45 -0.1 -0.0

0.95 2.75 2.50 2.40 1.05 1.15 1.50 2.05 3.40 2.70 0.90 -0.2 2.40 -0.2 -0.1 1.20 2.15 1.00 1.10 1.45 0.85 2.00 2.70 2.45 2.35 3.35 2.65 -0.3 2.35 -0.3 -0.2 0.95 0.80 1.05 1.40 1.15 1.95 2.10 0.90 0.75 1.00 2.65 3.30 2.60 2.30 2.40 2.30 -0.4 -0.4 -0.3 1.35 0.70 1.90 0.85 0.95 1.10 0.65 2.60 3.25 2.55 -0.5 2.25 -0.5 -0.4 1.30 1.85 2.05 0.80 0.90 2.35 2.25 0.60 3.20 2.50 1.25 1.05 0.75 0.55 0.85 1.80 -0.6 2.20 2.55 -0.6 -0.5 median median median median median median median median median median median median - median median median - median - 3.3254 2.6476 2.1223 2.4587 2.3504 1.4398 1.1346 0.9969 0.72271 1.0080 1.9755 0.4308 2.3360 2.6769 0.2877 0.2357 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 3.32009 2.64776 2.11639 2.45027 2.34431 1.42556 1.13956 0.97652 0.72298 1.01415 1.97272 0.41740 2.33122 2.66643 0.32138 0.24546 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.04830 0.05266 0.03697 0.04140 0.05050 0.06743 0.04990 0.07928 0.09942 0.07176 0.06054 0.10308 0.05157 0.04733 0.12363 0.12097

Platymantis schmidti – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.9 3.4 2.9 3.1 3.1 1.9 1.9 1.6 1.3 1.6 2.7 0.4 3.1 3.5 0.4 0.4

3.0 1.8 1.5 0.3 3.0 3.4 0.3 3.3 2.8 3.0 1.5 2.6 0.3 3.8 1.8 1.2 2.9 1.7 1.4 0.2 2.9 3.3 0.2 3.2 2.7 2.9 1.4 2.5 0.2

3.7 2.8 1.7 1.6 1.3 1.1 0.1 2.8 3.2 0.1

3.1 2.6 2.8 1.3 2.4 0.1 2.7 1.5 1.2 0.0 2.7 3.1 0.0 3.6 1.6 1.0 3.0 2.5 2.7 1.2 2.3 0.0 2.6 1.4 1.1 -0.1 2.6 3.0 -0.1

3.5 2.9 2.4 2.6 2.5 1.5 1.3 1.0 0.9 1.1 2.2 -0.2 2.5 2.9 -0.2 -0.1 median median median median median median median median median median median median median median median median 3.6911 3.1028 2.5185 2.7453 2.6803 1.6790 1.4609 1.2090 1.1474 1.2556 2.2986 0.10436 2.7180 3.0540 0.04879 0.13976 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.67474 3.10914 2.53694 2.76590 2.70633 1.68015 1.48628 1.23422 1.09823 1.26622 2.31646 0.07187 2.73142 3.10556 0.08272 0.14325 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.10016 0.12067 0.10827 0.10841 0.12353 0.10751 0.13834 0.12215 0.10838 0.11548 0.11389 0.15663 0.11574 0.16259 0.15992 0.15775

310

Platymantis solomonis – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.2 3.4 2.9 3.2 3.2 2.3 2.1 1.7 1.8 1.7 2.7 0.2 3.2 3.5 0.3 0.4

2.0 4.1 2.8 3.1 3.1 2.2 1.6 1.6 2.6 0.1 3.1 3.4 0.2 0.3 3.3 1.6 1.9 4.0 3.0 2.1 1.5 -0.0 2.7 3.0 1.5 2.5 3.0 3.3 0.1 0.2 3.2 1.8 1.4 3.9 2.9 2.0 1.4 -0.1 2.6 2.9 1.7 1.4 2.4 2.9 3.2 -0.0 0.1 3.8 3.1 2.8 1.9 1.2 1.3 -0.2 2.5 2.8 1.6 1.3 2.3 2.8 3.1 -0.1 -0.0 3.7 2.7 1.8 1.2 -0.3 3.0 1.5 1.0 2.4 2.7 1.2 2.2 2.7 3.0 -0.2 -0.1 3.6 2.6 1.7 1.1 -0.4 1.4 2.9 0.8 2.3 2.6 1.1 2.1 2.6 2.9 -0.3 -0.2 3.5 2.5 1.6 1.3 1.0 -0.5 3.4 2.8 2.2 2.5 2.4 1.5 1.2 1.0 0.6 0.9 2.0 -0.6 2.5 2.8 -0.4 -0.3 median median median median median median median median median median median median - median median median median 3.8828 3.1928 2.6965 3.0228 2.9429 1.9932 1.6926 1.4552 1.4182 1.3229 2.4164 0.1625 2.9403 3.2508 0.04879 0.11769 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean Mean 3.83806 3.15987 2.62863 2.94951 2.90232 1.94657 1.71866 1.40787 1.30764 1.31855 2.40241 0.21198 2.88066 3.19818 0.00824 0.06806 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.23293 0.17637 0.18276 0.22006 0.23560 0.23561 0.23333 0.20665 0.29677 0.19656 0.20066 0.18905 0.19238 0.19157 0.16762 0.18009

Platymantis speleaus – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.8 3.25 2.75 2.9 3.0 1.9 1.6 1.5 1.4 1.5 2.60 0.4 2.85 3.20 0.4 0.5

2.55 3.20 2.70 1.3 3.15 2.9 1.4 1.4 2.80 0.3 0.4 3.7 2.8 1.8 1.5 0.3 2.50 3.15 2.65 1.2 3.10 2.8 1.3 1.3 2.75 0.2 0.3 2.45 3.6 3.10 2.60 2.7 1.7 1.4 1.1 0.2 3.05 2.40 2.7 1.2 1.2 2.70 0.1 0.2 3.05 2.55 1.0 3.00 2.35 3.5 2.6 1.6 1.3 0.1 2.6 1.1 1.1 2.65 0.0 0.1 3.00 2.50 2.95 0.9 2.30

3.4 2.95 2.45 2.5 2.5 1.5 1.2 1.0 0.8 1.0 2.25 0.0 2.60 2.90 -0.1 0.0 median median median median median median median median median median median median median median median median 3.7551 3.1987 2.6497 2.8362 2.8820 1.8083 1.5347 1.3635 1.1756 1.3083 2.5281 0.26236 2.7801 3.1268 0.29267 0.32930 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.69448 3.16036 2.63396 2.76795 2.81621 1.74038 1.47680 1.30371 1.14526 1.28342 2.46354 0.22110 2.74348 3.08564 0.23584 0.27266 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.126602 0.091990 0.083523 0.131961 0.133337 0.135729 0.134542 0.133596 0.158891 0.147135 0.104085 0.125464 0.090054 0.106638 0.145918 0.156760

311

Platymantis vitianus – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.25 3.75 3.25 3.50 3.50 2.25 2.25 1.75 1.3 2.25 3.00 0.75 3.50 3.75 0.75 0.50

2.00 3.50 3.00 3.25 3.25 2.00 3.25 3.50 0.50 4.00 2.00 1.50 1.2 2.75 0.50 0.25 1.75

3.25 2.75 3.00 3.00 1.75 3.00 3.25 0.25

1.50 3.75 1.75 1.25 1.1 2.50 0.25 0.00 3.00 2.50 2.75 2.75 1.50 2.75 3.00 0.00 1.25

3.50 2.75 2.25 2.50 2.50 1.25 1.50 1.00 1.0 1.00 2.25 0.00 2.50 2.75 -0.25 -0.25 median median median median median median median median median median median median median median median median 3.8969 3.2699 2.7403 3.0015 2.9869 1.7238 1.8146 1.3916 1.1134 1.6121 2.6796 0.16104 2.9995 3.3368 0.21511 0.16321 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.88907 3.25776 2.72200 3.00157 2.99559 1.73017 1.83088 1.359944 1.13961 1.62612 2.63998 0.24572 2.97149 3.296491 0.22207 0.17781 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.25153 0.244238 0.300581 0.227442 0.25929 0.247351 0.235794 0.266482 0.088763 0.333790 0.271864 0.264967 0.285459 0.291520 0.319798 0.223996

Platymantis vitiensis – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.8 3.2 2.7 3.0 3.0 1.9 1.7 1.3 0.9 1.6 2.8 1.2 2.9 3.2 0.6 0.6

3.1 2.6 2.9 2.9 1.6 2.7 1.1 2.8 3.1 0.5 3.7 1.8 0.8 1.5 0.5 1.2 2.8 2.8 2.6 1.0 2.7 3.0 2.5 1.5 3.0 0.4 3.6 1.7 0.7 1.4 0.4 2.7 2.7 2.5 0.9 2.6 1.1 2.9 2.4 1.4 2.9 0.3 3.5 2.6 2.6 1.6 0.6 1.3 2.4 0.8 2.5 0.3 2.8 2.3 1.3 2.8 0.2 1.0 2.5 2.5 2.3 0.7 2.4 3.4 1.5 0.5 1.2 0.2 2.7 2.2 1.2 2.7 0.1 2.4 2.4 2.2 0.6 2.3 0.9 3.3 1.4 0.4 1.1 0.1 2.6 2.1 2.3 2.3 1.1 2.1 0.5 2.2 2.6 0.0 3.2 2.5 2.0 2.2 2.2 1.3 1.0 0.8 0.3 1.0 2.0 0.4 2.1 2.5 -0.1 0.0 median median median median median median median median median median median median median median median median 3.5785 2.8510 2.2821 2.5798 2.5382 1.5697 1.3052 0.9554 0.71784 1.2794 2.3480 0.7907 2.5236 2.8347 0.21106 0.25039 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.56306 2.87850 2.29865 2.56494 2.54114 1.56363 1.31916 0.97152 0.69235 1.29766 2.33535 0.78005 2.50294 2.83252 0.20269 0.23817 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.11850 0.17946 0.14301 0.15705 0.16261 0.13008 0.14664 0.11688 0.14944 0.12221 0.17125 0.16893 0.16668 0.15399 0.15508 0.13964

312

Platymantis weberi – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

3.9 3.3 2.8 3.0 3.0 2.0 1.8 1.40 1.5 1.35 2.50 0.50 2.95 3.25 0.3 0.2

1.35 2.45 2.90 0.2 1.30 3.20 2.7 1.4 0.1 3.8 3.2 2.9 2.9 1.9 1.7 0.25 1.30 2.40 2.85 0.1 1.25 3.15 2.6 1.3 -0.0 1.25 2.35 2.80 -0.0 3.7 3.1 2.8 2.8 1.8 1.6 1.20 0.00 3.10 1.20 2.30 2.75 -0.1 2.5 1.2 -0.1 1.15 3.05 1.15 2.25 2.70 -0.2 3.6 3.0 2.7 2.7 1.7 1.5 -0.25 2.4 1.1 -0.2 1.10 3.00 1.10 2.20 2.65 -0.3 3.5 2.9 2.3 2.6 2.6 1.6 1.4 1.05 1.0 1.05 2.15 -0.50 2.60 2.95 -0.4 -0.3 median median median median median median median median median median median median - median median median - median - 3.6587 3.0824 2.5185 2.7259 2.7120 1.6790 1.5644 1.2528 1.1474 1.1632 2.3656 0.1863 2.7770 3.1553 0.1054 0.0943 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean - Mean Mean Mean - Mean - 3.69355 3.09333 2.53657 2.80330 2.77130 1.74858 1.58283 1.25919 1.17024 1.17380 2.34484 0.16683 2.76663 3.11261 0.10721 0.07224 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.105092 0.099370 0.101685 0.132261 0.125323 0.145956 0.118005 0.099080 0.148892 0.096245 0.091521 0.217768 0.110294 0.092249 0.176185 0.142568

Rana kreffti – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.2 3.6 3.1 3.20 3.05 2.2 1.9 1.8 1.80 1.8 2.9 0.6 3.2 3.5 0.6 0.7

3.00 1.75 3.0 3.15 1.7 0.5 3.1 0.5 0.6 4.1 3.5 2.1 1.8 1.7 3.4 2.95 1.70 2.8 3.10 0.4 3.0 0.5 2.9 1.6 0.4 4.0 3.4 2.90 2.0 1.7 1.6 1.65 3.3 3.05 0.3 2.9 0.4 2.7 2.8 2.85 1.60 1.5 0.3 3.9 3.3 3.00 1.9 1.6 1.5 0.2 2.8 3.2 0.3 2.7 2.80 1.55 1.4 0.2 2.6 2.95 0.1 2.7 0.2 3.8 3.2 2.75 1.8 1.5 1.4 1.50 3.1 2.6 1.3 0.1 2.90 0.0 2.6 0.1 2.70 1.45 2.5 3.7 3.1 1.7 1.4 1.3 3.0 2.5 1.2 0.0 2.85 2.65 1.40 -0.1 2.5 0.0 3.6 3.0 2.4 2.80 2.60 1.6 1.3 1.2 1.35 1.1 2.4 -0.2 2.4 2.9 -0.1 -0.1 median median median median median median median median median median median median median median median median 3.8083 3.1679 2.5802 2.9434 2.7754 1.7976 1.5560 1.5195 1.5993 1.3750 2.5573 0.28123 2.8457 3.1591 0.30748 0.30379 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.84179 3.20409 2.62491 2.95908 2.78255 1.82644 1.56415 1.53136 1.60660 1.38657 2.58633 0.25485 2.84359 3.17976 0.29670 0.30771 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.14259 0.14263 0.13469 0.11423 0.13102 0.12805 0.12065 0.12844 0.10336 0.13741 0.15256 0.21491 0.17523 0.14811 0.19632 0.19662

313

Rana novaebritanniae – males lnSV lnTIB lnTAR lnHL lnHW lnED lnEN lnIN lnTY lnIOW lnF3 lnFD3 lnT3 lnT4 lnTD3 lnTD4

4.2 3.7 3.0 3.3 3.1 2.1 2.0 1.9 1.75 1.8 2.9 0.4 3.2 3.6 0.6 0.5

1.70 4.1 3.6 2.9 2.0 1.9 1.8 3.5 0.5 3.2 3.0 1.7 2.8 0.3 0.4 3.0 1.65 4.0 3.5 2.8 1.9 1.8 1.7 3.4 0.4 3.1 2.9 1.6 2.7 0.2 0.3 1.60 3.9 3.4 2.7 1.8 1.7 1.6 2.8 3.3 0.3 1.55 3.0 2.8 1.5 2.6 0.1 0.2 3.8 3.3 2.6 1.7 1.6 1.5 3.2 0.2 1.50 2.6 2.9 2.7 1.4 2.5 0.0 0.1 3.7 3.2 2.5 1.6 1.5 1.4 3.1 0.1 1.45

3.6 3.1 2.4 2.8 2.6 1.5 1.4 1.3 1.40 1.3 2.4 -0.1 2.4 3.0 0.0 0.0 median median median median median median median median median median median median median median median median 3.7492 3.1389 2.5649 2.9083 2.7104 1.8031 1.4702 1.4725 1.5304 1.3558 2.5388 0.18578 2.8021 3.1335 0.24294 0.20698 Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean Mean 3.80651 3.18089 2.62774 2.95231 2.76679 1.80811 1.50184 1.51219 1.55390 1.41787 2.56069 0.17525 2.81610 3.16433 0.26162 0.21152 Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev Std Dev 0.14489 0.14263 0.15474 0.11134 0.13394 0.10937 0.14221 0.12070 0.08933 0.11888 0.11945 0.15743 0.14755 0.13037 0.12883 0.12201

314

APPENDIX 3. Character Descriptions

NASALS

Character 0: Shape of Nasals

a b

(a) Platymantis guppyi (x3) SAM47128 (b) Rana kreffti (x6) MCZ38087

State 0: nasals narrow. maximum width (excluding maxillary process) is less than 30% of the maximum length (including maxillary process) (fig. b) State 1: nasals broad, triangular. maximum width (excluding maxillary process) is greater than 30% (invariably 50%) of the maximum length (including maxillary process) (fig. a)

Character 1: Articulation of the nasal with the frontoparietal

a b c

(a) Palmatorappia solomonis (x12) MCZ60864 (b) Platymantis solomonis (x6) MCZ58569 (c) Ceratobatrachus guentheri (x3) MCZ62226

State 0: separate (fig. a) State 1: edges closely applied or fused along entire margins (figs b,c)

315

Character 2: The shape of the maxillary process of the nasal.

a b c

(a) Batrachylodes vertebralis (x12) MCZ41848 (b) Discodeles guppyi (x3) SAM47092 (c) Platymantis solomonis (x6) MCZ58569

State 0: short (fig. a) State 1: broad and long (fig. b) State 2: narrow and long (fig. c)

Character 3: Relationships of the medial edge of the nasals to each other.

a

c a’

b

(a) Rana grisea (x6) SAM5697B (a') Palmatorappia solomonis (x12) MCZ60864 (b) Discodeles guppyi (x3) SAM47092 (c) Ceratobatrachus guentheri (x3) MCZ62226

State 0: widely separated (figs a, a’) State 1: closely applied (figs b, c)

316

FRONTOPARIETAL

Character 4: Shape of anterior edge of frontoparietal

a b c

(a) Batrachylodes wolfi (x12) SAMR5164c (b) Discodeles bufoniformis (x6) MCZ38578 (c) Rana grisea (x6) SAMR5687B

State 0: anterior margin of frontoparietal acuminate or forming a broad arc (antero-medial margin anterior to the antero-lateral margin) (fig. a) State 1: anterior margin of frontoparietal truncated (antero-medial margin and antero-lateral margin nearly horizontal) (fig. b) State 2: anterior margin of frontoparietal notched (antero-medial margin posterior to antero-lateral margin) (fig. c)

Character 5: Degree of fusion of frontoparietal elements

a b c

(a) Palmatorappia solomonis (x12) MCZ60864 (b) Batrachylodes vertebralis (x12) MCZ41836 (c) Ceratobatrachus guentheri (x3) MCZ62226

State 0: not fused (fig. a)

317

State 1: fused (figs b, c)

SQUAMOSAL

Character 6: Ratio of the length of the zygomatic ramus to the otic ramus of the squamosal. Measured from the centre of the shaft.

a b c

(a) Palmatorappia solomonis (x12) MCZ60864 (b) Platymantis solomonis (x6) MCZ38569 (c) Meristogenys orphnocnemis (x6) CAS235890

State 0: zygomatic ramus is shorter than otic (fig. a) State 1: zygomatic ramus is equal in length to the otic ramus (fig. b) State 2: zygomatic ramus is longer than otic (fig. c)

318

Character 7: Articulation of otic ramus of squamosal and the crista parotica

a b c

(a) Batrachylodes wolfi (x12) SAMR5164C (b) Platymantis solomonis (x6) MCZ58568 (c) Platymantis boulengeri (x6) SAMR7090

State 0: otic ramus well separated from crista parotica (fig. a) State 1: otic ramus lying alongside crista parotica (fig. b) State 2: otic ramus overlapping crista parotica (fig. c)

319

Shapes of squamosal

a b c d

f e g

i h

Staurois natator, Platymantis papuensis, P. weberi, P. corrugata, P. myersi, P. parkeri, Batrachylodes vertebralis, B. wolfi, B. mediodiscus, B. trossulus (fig. a) Palmatorappia solomonis (fig. b) Occidozyga laevis, Platymantis schmidti (fig. c) Discodeles bufoniformis, D. guppyi, Platymantis guppyi (fig. d) Platymantis solomonis, P. neckeri, Rana kreffti, Ingerana baluensis (fig. e) Limnonectes magna, Rana grisea (fig. f) Meristogenys orphnocnemis (fig. g) Ceratobatrachus guentheri (fig. h) Platymantis boulengeri (fig. i)

Character 8: Shape of the rami (zygomatic + otic) of the squamosal

State 0: straight State 1: curved

320

Character 9: Width of otic ramus

State 0: not expanded: less than or equal to zygomatic ramus State 1: expanded: 2x or more zygomatic ramus

MAXILLA

Character 10: Relationship of the lateral process of the premaxilla to the palatal shelf of the maxilla

b c (a) Batrachylodesa wolfi (x12) SAMR5164c (b) Platymantis solomonis (x6) MCZ85868 (c) Limnonectes magnus (x6) CAS23405

State 0: widely separated (fig. a) State 1: lying alongside (fig.b) State 2: wrapped around (fig. c)

PREMAXILLA

Character 11: Orientation of alary process of the premaxilla in lateral view.

a b

(a) Limnonectes magnus (x6) CAS23405 (b) Ingerana baluensis (x12) FMNH238301

o State 0: alary process nearly vertical (~80-90 ) (fig. a) State 1: alary process slopes posteriorly, usually curved (fig. b)

321

Character 12: Width of lateral process of premaxilla.

a b c

(a) Occidozyga laevis (x6) CAS16269 (b) Palmatorappia solomonis (x12) MCZ60864 (c) Batrachylodes vertebralis (x25) MCZ41834

State 0: lateral process acuminate projecting extensively towards the maxilla (fig. a) State 1: lateral process less than or equal to the palatine process of the premaxilla (fig. b) State 2: lateral process wider (approx. 2x) than the palatine process of the premaxilla (fig. c)

QUADRATOJUGAL

Character 13: Quadratojugal spur

a b

(a) Platymantis myersi (x6) SAM47157 (b) Staurois natator (x6)

State 0: absent (fig. a) State 1: present (fig. b)

322

SPHENETHMOID

Character 14: Projection of sphenethmoid in dorsal view

a b

(a) Limnonectes magnus (b) Platymantis weberi

State 0: overlain entirely by nasals dorsally (fig. a) State 1: not overlain or overlain by posterior margin of nasals dorsally (fig. b)

Character 15: Projection of sphenethmoid in ventral view

a b (a) Limnonectes magnus (b) Platymantis weberi

State 0: sphenethmoid not projecting anteriorly beyond palatines (fig. a) State 1: sphenethmoid projects anteriorly beyond palatines (fig. b)

323

PARASPHENOID

Character 16: Extension of cultriform process of the parasphenoid

b a

c d

(a) Palmatorappia solomonis (x12) MCZ60864 (b) Rana kreffti (x6) MCZ38087 (c) Platymantis weberi (x6) SAMR4412 (d) Meristogenys orphnocnemis (x6) CAS235890

State 0: ending well below to the palatines (fig. a) State 1: reaching the palatines (figs b, c) State 2: projecting beyond the palatines (fig. d)

324

Character 17: Shape of cultriform process. Terminology taken from Lawrence (1955)

a b c d e

(a) Batrachylodes vertebralis (x12) MCZ41848 (b) Platymantis corrugatus (x6) SAM13602A (c) Platymantis schmidti (x6) SAMR7766 (d) Ceratobatrachus guentheri (x3) MCZ62226 (e) Occidozyga laevis (x12?) CAS16269

State 0: obtuse (with rounded apex) (figs a, b), may appear truncate State 1: lanceolate (shaped as the head of a lance, broadened at base and tapering toward the apex) (figs c, d) State 2: caudate (bearing a tail-like process) (fig. e)

Character 18: Base of cultriform process curving inward.

a b (a) Palmatorappia solomonis (x6) MCZ60864 (b) Ceratobatrachus guentheri (x3) MCZ62226

State 0: absent (fig. a) State 1: present (fig. b) Character 19: Shape of posteromedial margin of parasphenoid

325

b

c

a

d e

(a) Staurois natator (b) Batrachylodes mediodiscus (x12) (c) Meristogenys orphnocnemis (x6) FMNH235890 (d) Ceratobatrachus guentheri (x3) MCZ6226 (e) Platymantis solomonis (x6) MCZ85868

State 0: margin has no projections (fig. a) State 1: projection is curved, not extending to base of otoccipital (figs b, c) State 2: projection is V-shaped, extending towards the base of the otoccipital(figs d, e)

326

VOMERS

Character 20: Vomerine teeth

a b

(a) Batrachylodes vertebralis (x12) MCZ41848 (b) Platymantis guppyi (x3) SAM47128

State 0: absent (fig. a) State 1: present (fig. b)

Shapes of the vomer

a b c d e

f g h i j

k l m

Staurois natator (fig. a) Amolops orphnocnemis, Rana grisea (fig. b) Occidozyga laevis (fig. c) Batrachylodes vertebralis (all Batrachylodes) (fig. d) Palmatorappia solomonis (fig. e)

327

Ceratobatrachus guentheri (fig. f) Discodeles bufoniformis (fig. g) Discodeles guppyi (fig. h) Platymantis solomonis (corrugatus, schmidti, weberi, myersi, papuensis, neckeri, guppyi) (fig. i) Platymantis boulengeri (fig. j) Platymantis parkeri (fig. k) Rana kreffti, Limnonectes magnus (fig l) Ingerana baluensis (fig. m)

Character 21: Anterior process of vomer

State 0: reduced State 1: not reduced

Character 22: Anterior alary (lateral) process of vomer

State 0: reduced State 1: not reduced

Character 23: Posterior alary (lateral) process of vomer

State 0: reduced State 1: not reduced

Character 24: Dentigerous process of vomer

State 0: broad State 1: narrow, acuminate, long

Character 25: Position of vomerine teeth on dentigerous process

State 0: arising from the central area of the dentigerous process of the vomer (fig. i from character 25) State 1: arising from the posterior margin of the dentigerous process of the vomer, this may form an isthmus (fig, m from character 25)

328

Character 26: Articulation of the posterior alary process of vomers with the palatines

a b c d

(a) Batrachylodes vertebralis (x12) MCZ41848 (b) Rana kreffti (x6) MCZ38027 (c) Platymantis schmidti (x6) SAMR7766 (d) Discodeles guppyi (x3) SAM47092

State 0: separate (figs a, c) State 1: present; touching or overlying (fig. b)

Character 27: Articulation of the dentigerous process of vomers with the palatines

State 0: separate (fig. b) State 1: present; touching and/or fused and/or overlapping (figs c, d)

Character 28: Position of the vomerine teeth in relation to the choanae

State 0: between State 1: between and extending posterior

Character 29: Shape of vomerine teeth

State 0: straight (fig. i from character 25) State 1: curved (fig. h from character 25)

329

PTERYGOID

Character 30: Extent of medial ramus of the pterygoid

a b b

c

(a) Palmatorappia solomonis (x12) MCZ60864 (b) Discodeles bufoniformis (x3) MCZ38557 (c) Ceratobatrachus guentheri (x3) MCZ62226

State 0: not reaching prootic (fig. a) State 1: overlying prootic (fig. b) State 2: reaching alae of parasphenoid (fig. c)

Character 31: Articulation of the posterior ramus of the pterygoid with the quadratojugal

State 0: present, cartilaginous State 1: present, bony

330

EXTREMITIES

Character 32: Shape of the distal phalanx, this character is size dependant, with digit 1 occasionally exhibiting marginal features

a b c

(a) Ceratobatrachus guentheri (x6) MCZ62226 (b) Batrachylodes vertebralis (x12) MCZ41846 (c) Palmatorappia solomonis (x12) MCZ60863

State 0: pointed or rounded (fig. a) State 1: T-shaped (fig. b) State 2: Y-shaped (fig. c)

Character 33: Sesamoid bones between the joints on the hand

State 0: absent State 1: present

Character 34: Sesamoid bones between the joints on the foot

State 0: absent State 1: present

Character 35: Sesamoid bones at the base of the tibio-tarsus

State 0: absent State 1: present

331

Character 36: Circummarginal groove on fingers (2,3,4)

a b

State 0: absent (fig. a) State 1: present (fig. b)

Character 37: Depth of circummarginal groove on fingers

State 0: shallow (indistinct) State 1: deep (distinct)

Character 38: Position of circummarginal groove on fingers

State 0: ventral (inferior) State 1: terminal State 2: dorsal (superior)

Character 39: Circummarginal groove on toes (2,3,4,5)

State 0: absent State 1: present

332

Character 40: Depth of circummarginal groove on toes

a b c

State 0: shallow (figs a, b) State 1: deep (fig. c)

Character 41: Position of circummarginal groove on toes

State 0: ventral State 1: terminal State 2: dorsal

Character 42: Relative length of finger 1 to finger 2

State 0: F1  F2 State 1: F1 > F2

Character 43: Relative length of finger 2 to finger 4

State 0: F2  F4 State 1: F2 > F4

Character 44: Relative length of finger 1 to finger 4

State 0: F1  F4 State 1: F1 > F4

333

Character 45: Length of 3rd finger (FL3) as a ratio of snout-vent length (SV)

State 0: (lnFL3 +1):(lnSV +1) less than or equal to 0.530 State 1: (lnFL3 +1):(lnSV +1) between 0.531 and 0.550 State 2: (lnFL3 +1):(lnSV +1) between 0.551 and 0.580 State 3: (lnFL3 +1):(lnSV +1) between 0.581 and 0.600 State 4: (lnFL3 +1):(lnSV +1) greater than 0.610

Character 46: Discs on fingers

a b

State 0: absent (fig. a) State 1: present (fig. b)

334

Character 47: Shape of finger discs

a b c

State 0: round (fig. a) State 1: truncate (fig. b) State 2: pointed (fig. c)

Character 48: Shape of the Inner Palmar Tubercle. Terminology taken from Lawrence (1955)

State 0: ovate (egg-shaped, broadest below the middle, usually but not necessarily rounded at each end) State 1: elliptical (broadest point midway between ends and the width about one- half the length, maybe be classified as narrowly elliptical or broadly elliptical depending on the ratio of width to length) State 2: round State 3: obovate (broadest part above the middle and narrower end toward the base)

Character 49: Height of the Inner Palmar Tubercle

State 0: low, flattened State 1: protrudent, raised

Character 50: Shape of the Middle Palmar Tubercle. Terminology taken from Lawrence (1955)

State 0: ovate (egg-shaped, broadest below the middle, usually but not necessarily rounded at each end) State 1: elliptical (broadest point midway between ends and the width about one- half the length, maybe be classified as narrowly elliptical or broadly elliptical depending on the ratio of width to length) State 2: round State 3: obovate (broadest part above the middle and narrower end toward the base)

335

Character 51: Height of the Middle Palmar Tubercle

State 0: low, flattened State 1: protrudent, raised

Character 52: Shape of the Outer Palmar Tubercle. Terminology taken from Lawrence (1955)

State 0: elliptical (broadest point midway between ends and the width about one- half the length, maybe be classified as narrowly elliptical or broadly elliptical depending on the ratio of width to length) State 1: round State 2: obovate (broadest part above the middle and narrower end toward the base)

Character 53: Height of the Outer Palmar Tubercle

State 0: low, flattened State 1: protrudent, raised

Character 54: 4th Palmar Tubercle

State 0: absent State 1: present

Character 55: Shape of the Inner Plantar Tubercle. Terminology taken from Lawrence (1955)

State 0: ovate (egg-shaped, broadest below the middle, usually but not necessarily rounded at each end) State 1: elliptical (broadest point midway between ends and the width about one- half the length, maybe be classified as narrowly elliptical or broadly elliptical depending on the ratio of width to length) State 2: round

Character 56: Height of the Inner Plantar Tubercle

State 0: low, flattened State 1: protrudent, raised

Character 57: Outer Plantar Tubercle

State 0: absent State 1: present

Character 58: Shape of the Outer Plantar Tubercle

State 0: elliptical State 1: round State 2: conical Character 59: Height of the Outer Plantar Tubercle

336

State 0: low, flattened State 1: protrudent, raised

Character 60: Appearance of Palmar Surface of Hand

State 0: smooth State 1: rugose. small tubercles State 2: warty, large tubercles

Character 61: Appearance of Plantar Surface of Foot

State 0: smooth State 1: rugose, small tubercles

Character 62: Appearance of Subarticular Tubercles on the fingers

State 0: indistinct State 1: distinct

Character 63: Anterior Margin of Subarticular Tubercles on the fingers (seen laterally)

State 0: round State 1: pointed State 2: truncate

Character 64: Height of Subarticular Tubercles on the Toes

State 0: indistinct: low, flattened State 1: distinct: prominent, raised

Character 65: Anterior Margin of Subarticular Tubercles on the Toes (viewed laterally)

State 0: round State 1: pointed State 2: truncate

Character 66: Relative Length of Toe 3 to Toe 5

State 0: T3 < T5 State 1: T3 > T5

337

Character 67: Length of 4th toe (TL4) as a ratio of snout-vent length (SV)

State 0: (lnTL4 +1):(lnSV +1) less than or equal to 0.750 State 1: (lnTL4 +1):(lnSV +1) between 0.760 and 0.800 State 2: (lnTL4 +1):(lnSV +1) between 0.801 and 0.820 State 3: (lnTL4 +1):(lnSV +1) greater than 0.821

Character 68: Shape of toe discs

a c b

(a) (b) (c)

State 0: pointed (fig. a) State 1: round (fig. b) State 2: truncate (fig. c)

Character 69: Extent of Webbing on Feet

State 0: rudimentary; minute at base, below or reaching proximal subarticular tubercles on all digits State 1: 2/3, reaching distal sat but not reaching disc on all State 2: entire, reaching disc on all toes

338

PECTORAL GIRDLE

Character 70: Shape of Omosternum

a b c d

(a) Batrachylodes wolfi (x12) SAMR5164c (b) Staurois natator (x12) CAS23364 (c) Rana kreffti (x6) MCZ38087 (d) Discodeles bufoniformis (x6) MCZ38581

State 0: unforked, no indentation at base (figs a, b) State 1: slightly forked, slight indent at base (fig. c) State 2: forked, marked indentation forming two posterior struts that attach to clavicles (fig. d)

Character 71: Length of the sternum (bony body only)

State 0: Bony part of sternum short and thick: length is less than 30% of the total length of the pectoral girdle (anterior/posterior plane) State 1: Bony part of sternum long and thin: l 2x w; length is greater than 30% of the total length of the pectoral girdle (anterior/posterior plane)

VERTEBRAL COLUMN

Character 72: Neural Spines, all spines decrease in size anterior to posterior

State 0: absent State 1: present on vertebrae 2-3 only State 2: present on vertebrae 2-4 only State 3: present on vertebrae 2-7, may or may not be on 8

Character 73: Width of vertebra 2 to vertebra 4, including transverse processes, in the medio-lateral plane.

State 0: 2 < 4 State 1: 2 = 4 State 2: 2 > 4

339

Character 74: Width of transverse processes of vertebra 2 to vertebra 4, in the antero-posterior plane.

State 0: 2 < 4 State 1: 2 = 4 State 2: 2 > 4

HYOID

Character 75: Expansion of proximal end of posterior cornua of the hyoid

a b

(a) Platymantis solomonis (x6) SAM47134 (b) Batrachylodes wolfi (x12) SAMR4297A

State 0: little or no expansion. Width of proximal end the same as the shaft (fig. a) State 1: proximal end expanded. Width of proximal end is greater than that of the shaft (fig. b)

Character 76: Expansion of distal end of posterior cornua of the hyoid

State 0: little or no expansion. Width of distal end the same as the shaft (fig. b from character 88) State 1: distal end expanded. Width of distal end is greater than that of the shaft (fig. a from character 88)

Character 77: Alary process of the hyoid

State 0: absent State 1: present

340

Various shapes of the alary process of the hyoid

b c d e e’

a b’

f f’ g g’

(a) Ceratobatrachus guentheri (x6) MCZ62226 (b) Discodeles bufoniformis (x6) MCZ38581 (b’) Platymantis weberi (x6) SAMR4412 (c) Staurois natator (x12) CAS23364 (d) Rana kreffti (x6) MCZ38084 (e) Limnonectes magnus (x6) CAS23405 (e’) Discodeles guppyi (x6) SAM47091 (f) Platymantis neckeri (x6) SAMR4928B (f’) Platymantis boulengeri (x6) SAMR7067 (g) Batrachylodes vertebralis (x12) SAM47110 (g’) Palmatorappia solomonis (x12) MCZ60858

Character 78: Alary process

State 0: non-pedunculate (fig. e) State 1: pedunculate (fig. b)

Character 79: Alary process

State 0: unexpanded State 1: expanded

Character 80: Antero-medial process of the anterior cornua of the hyoid

State 0: absent State 1: present

341

Character 81: Shape of anterior horn of the anterior cornua of the hyoid

d e a b

c

f

(a) Limnonectes magnus (x6) CAS23405 (b) Meristogenys orphnocnemis (6) FMNH235889 (c) Staurois natator (x12) CAS23364 (d) Rana kreffti (x6) SAMR5161A (e) Platymantis schmidti (x6) SAM8030 (f) Platymantis solomonis (x6) SAM47134

State 0: anterior horn represented by a small bump anteriorly (fig. a) State 1: anterior horn projecting anteriorly from hyoid, may be slightly curved at tip (fig. b) State 2: anterior horn projecting anteriorly and slightly curving medially (fig. c) State 3: anterior horn curving laterally, can form an arch (figs d, e, f)

Character 82: Postero-lateral horn of the hyoid

State 0: absent State 1: present

342

Character 83: Shape of postero-lateral horn of the hyoid

a b

(a) Limnonectes magnus (x6) CAS23405 (b) Ceratobatrachus guentheri (x6) MCZ62226

State 0: triangular (fig. a) State 1: cuspidate (somewhat abruptly and sharply concave and constricted into an elongated sharp-pointed tip) (fig. b)

Character 84: Position of alary process of the hyoid in relation to the anterior margin of the hyoid plate.

a b c

(a) Palmatorappia solomonis (x12) MCZ60858 (b) Platymantis weberi (x6) SAMR4412 (c) Rana kreffti (6) SAMR5161A

State 0: above (fig. a) State 1: level (fig. b) State 2: below (fig. c)

343

Measurements taken from the hyoid

PL = Plate Length PL PW = Plate Width PLH = Postero-lateral Horn Length PCL – posterior cornua length PW

PLH

PCL

Character 85: Shape of the hyoid plate expressed as a ratio of length : width (PL:PW)

State 0: narrow (length > width; ratio > 1.1) State 1: square (length approximates width; ratio within 10% of 1) State 2: broad (length < width; ratio < .9)

Character 86: Length of the posterior cornua (PCL) as expressed as a ratio of hyoid plate length (PL)

State 0: short (posterior cornua length < hyoid plate length; ratio < 0.9) State 1: equal (posterior cornua length < hyoid plate length; ratio 0.9-1.1) State 2: long (posterior cornua length < hyoid plate length; ratio > 1.1)

344

Character 87: Length of the posterior-lateral horn (PLH) as expressed as a percentage of plate length (PL)

State 0: short: PPL is less than 50% of PL State 1: long: PPL is greater than 50% of PL

PELVIC GIRDLE

Character 88: Shape of the ventral acetabular expansion if the ilium. Defined from Tyler, 1976:4.

a b c

(a) Batrachylodes vertebralis (x12) SAM47110 (b) Amolops orphnocnemis (x6) FMNH235890 (c) Staurois natator (x12) CAS23364

State 0: "form of a straight line extending ventrally and posteriorly" (fig 2A, Tyler 1976) (fig. a) State 1: "a gradual concavity" (fig 2B, Tyler 1976) (fig. b) State 2: "a concavity of the preacetabular zone and a convexity underneath" (2C, Tyler 1976) (fig. c)

Measurements taken from the pelvic girdle

PGL ISL DAEA DCL

DCH

ISH

PGL = pelvic girdle length AW ISL = ilial shaft length DCH = dorsal crest height ISH = ilial shaft height AW = acetabular width DCL = dorsal crest length IIW = ischium + ilium length (- shaft) IIW Character 89: Dorsal Crest DAEA = dorsal acetabular expansion angle

345

State 0: absent State 1: present

Character 90: Length of dorsal crest (DCL) expressed as a ratio of ilial shaft length (ISL)

State 0: dorsal crest 80.00-92.99% of ilial shaft State 1: dorsal crest 93.00-100.00% of ilial shaft

Character 91: Height of dorsal crest (DCH) expressed as a ratio of total ilium height (ISH + DCH)

State 0: low (dorsal crest <45.00% of total ilial shaft) State 1: medium (dorsal crest 45.01-64.99% of total ilial shaft) State 2: high (dorsal crest >65.00% of total ilial shaft)

Character 92: Shape of the dorsal protuberance

a b

(a) Discodeles guppyi (x3) SAM47092 (b) Platymantis parkeri (x25) SAMR15159B

State 0: round (fig. a) State 1: oval (fig. b)

346

Character 93: Shape of the dorsal acetabular expansion

a b c

(a) Platymantis boulengeri (x6) (b) Batrachylodes vertebralis (x12) SAM47110 (c) Occidozyga laevis (x3) CAS16269

State 0: straight (fig. a) State 1: concave (fig. b) State 2: sigmoidal (fig. c)

Character 94: Length of the ilial shaft (ISL) expressed as a percentage of pelvic girdle length (PGL)

State 0: short (ilial shaft < 65.00% of pelvic girdle) State 1: long (ilial shaft >65.01% of pelvic girdle

Character 95: Length of the pelvic girdle (PGL) expressed as a percentage of snout-vent length (SV)

State 0: short (pelvic girdle <30.00% of snout-vent) State 1: medium (pelvic girdle between 30.01-59.99% of snout-vent) State 2: long (pelvic girdle >60.00% of snout-vent)

347

Character 96: The position of the inferior margin of the dorsal protuberance to the anterior rim of the acetabulum

b a

(a) Palmatorappia solomonis (x12) MCZ60684 (see fig.51a) (b) Discodeles guppyi (x3) SAM47092

State 0: above (fig. a) State 1: below (fig. b)

Character 97: Position of the dorsal protuberance to the anterior rim of the acetabulum

State 0: level State 1: 1/3 anterior State 2: 1/2 anterior State 3: 2/3 anterior State 4: entirely anterior

Character 98: Width of the acetabulum (AW) expressed as a ratio of the width of the ilium (minus the shaft) and ischium (IIW)

State 0: small (acetabulum < 55.00% of ischium & ilium width) State 1: large (acetabulum > 55.01% of ischium & ilium width)

348

Character 99: The position of the superior margin of the acetabulum to the inferior margin of the ilial shaft

a

b

(a) Ceratobatrachus guentheri (x3) MCZ62225 (b) Platymantis neckeri (x6) SAMR4296D

State 0: well below (fig. a) State 1: level (fig. b) State 2: well above (see measurements from pelvic girdle figure)

349

Character 100: Angle of the ilial shaft

a

b

c

(a) Discodeles guppyi (b) Rana kreffti (c) Limnonectes magnus

State 0: straight or slightly curved (figs a, b) State 1: markedly curved (fig. c)

Character 101: Angle of the dorsal acetabular expansion (DAEA) to the ilial shaft

State 0: narrow (angle < 19.990) State 1: medium (angle 20.000-49.990) State 2: wide (angle > 50.000)

HEAD

Character 102: Shape of the lateral margin of the snout (canthus rostralis) viewed dorsally. Terminology from Duellman (1979)

State 0: rounded State 1: angular

350

Character 103: Anterior projection of the lateral margin of the snout (canthus rostralis) from the eye to the nostril, viewed dorsally

b a

(a) Rana kreffti (x6) MCZ38209 (b) Palmatorappia solomonis (x6) MCZ60860

State 0: straight (fig. a) State 1: curved (fig. b)

Character 104: Conical tubercles on the posterior region of the upper eyelid

State 0: absent State 1: present

Character 105: Angle of loreal region (side of snout)

State 0: oblique State 1: vertical

Character 106: Shape of loreal region

State 0: concave State 1: straight State 2: convex

Character 107: Small, white tubercles on venter surface of chin (snout) in males

State 0: absent State 1: present

351

Character 108: Shape of snout viewed laterally

State 0: obtuse State 1: pointed State 2: truncate

Character 109: Projection of the upper lip

State 0: absent State 1: present

Character 110: Tympanum

State 0: absent State 1: present

Character 111: Shape of the tympanum

State 0: round State 1: vertically elliptical

Character 112: Head Shape expressed as a percentage of the length (HL) to width (HW)

State 0: broad: (lnHL +1):(lnHW +1) less than or equal to 1.05 State 1: narrow: (lnHL +1):(lnHW +1) greater than 1.051

Character 113: Shape of the snout expressed as a ratio of internarial distance to eye-naris distance

State 0: narrow: (lnIN +1):(lnEN +1) less than or equal to 0.800 State 1: equal: (lnIN +1):(lnEN +1) between 0.801 and 1.100 State 2: broad: (lnIN +1):(lnEN +1) greater than 1.101

Character 114: Length of the snout : eye-naris distance expressed as a percentage of head length

State 0: short: (lnEN +1):(lnHL +1) less than or equal to 0.500 State 1: long: (lnEN +1):(lnHL +1) greater than to 0.501

Character 115: Size of tympanum expressed as a percentage of the length of head

State 0: (lnTYMP +1):(lnHL +1) less than or equal to 0.370 State 1: (lnTYMP +1):(lnHL +1) between 0.371 and 0.390 State 2: (lnTYMP +1):(lnHL +1) between 0.391 and 0.530 State 3: (lnTYMP +1):(lnHL +1) greater than 0.531

352

MOUTH

Character 116: Size of choanae compared to Eustachian tube opening

State 0: less than State 1: equal State 2: greater than

Character 117: Shape of the choanae

State 0: round State 1: oval

Character 118: Shape of Eustachian tube opening

State 0: oval, slit-like State 1: triangular State 2: round

Character 119: Attachment of the Tongue

State 0: 1/3 of tongue is free posteriorly State 1: 1/2 of tongue is free posteriorly State 2: 2/3 of tongue is free posteriorly

Character 120: Shape of the tongue

b c a d (a) Occidozyga laevis (b) Batrachylodes trossulus (c) Ceratobatrachus guentheri (d) Platymantis boulengeri

State 0: broadest posteriorly (near notch) tapering to a point anteriorly (figs a, b) State 1: heart-shaped, broadest in middle, narrowing anteriorly at attachment (fig. c) State 2: broad in middle but not as narrow anteriorly as seen in State 1 (fig. d)

353

Character 121: Tongue notch

State 0: absent State 1: present

Character 122: Depth of tongue notch

b c a

(a) Ceratobatrachus guentheri (b) Ingerana baluensis (c) Platymantis solomonis

State 0: shallow (fig. a) State 1: deep (figs b, c)

Character 123: Width of Tongue notch

State 0: narrow (fig. b) State 1: broad (fig. c)

354

Character 124: Shape of tongue notch

b a

(a) Platymantis boulengeri (b) Ingerana baluensis

State 0: U-shaped (fig. a) State 1: V-shaped (fig. b)

EGGS

Character 125: Egg size

State 0: small (less than 2mm) State 1: large (greater than 2.01mm)

Character 126: Egg pigmentation

State 0: absent State 1 present

Character 127: Number of eggs

State 0: Few State 1: Numerous

SKIN

Character 128: Dorsolateral fold

State 0: absent State 1: present

Character 129: folds on dorsum

State 0: absent State 1: longitudinal, linear ridges State 2: urn-shaped in scapular (axillary) region

355

Character 130: Tarsal fold

State 0: absent State 1: present

Character 131: Supratympanic fold

State 0: absent State 1: present

Character 132: Texture of skin on dorsum

State 0: smooth State 1: rugose : fine granules State 2: warty : low, flat tubercles

Character 133: Texture of skin on venter

State 0: smooth State 1: rugose : fine granules State 2: warty : low, flat tubercles

Character 134: Texture of skin ventromedially of thigh

State 0: smooth State 1: rugose : fine granules State 2: warty : low, flat granules

LEGS

Character 135: Tibia length (TIB) expressed as a percentage of the snout vent length (SV)

State 0: (lnTIB +1):(lnSV +1) less than or equal to 0.770 State 1: (lnTIB +1):(lnSV +1) between 0.771 and 0.880 State 2: (lnTIB +1):(lnSV +1) between 0.881 and 0.891 State 3: (lnTIB +1):(lnSV +1) between 0.892 and 0.910 State 4: (lnTIB +1):(lnSV +1) greater than 0.911

356

Mahony, M.J., Norris, R.M. and Donnellan, S.C. (1996) Karyotypes of South-west Pacific Ranid Frogs (Anura:Ranidae) Australian Journal of Zoology, v.44 (2), pp. 119-128, 1996

NOTE: This publication is included in the print copy of the thesis held in the University of Adelaide Library.

It is also available online to authorised users at:

http://dx.doi.org/10.1071/ZO9960119

Norris, R.M. (1999) Testing multiple species hypothesis on frogs. Perspectives in Human Biology, v. 4 (1), pp.51-64, 1999

NOTE: This publication is included in the print copy of the thesis held in the University of Adelaide Library.