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Home , Chelidae, Chelodina, Chelodina expansa, Chelus, Compsemys, Elseya

AMERICAN MUSEUM

Noiltates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET NEW YORK, N.Y. 10024 U.S.A. NUMBER 2620 MAY 10, 1977

EUGENE S. GAFFNEY The Side-Necked : A Theory of Relationships Using Shared Derived Characters

NovitatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2620, pp. 1-28, figs. 1-10, tables 1-3 May 10, 1977

The Side-Necked Turtle Family Chelidae: A Theory of Relationships Using Shared Derived Characters

EUGENE S. GAFFNEY'

ABSTRACT The South American and Australian side- b. Dorsal processes of exoccipitals meet medi- necked of the family Chelidae are ana- ally above foramen magnum lyzed using the shared derived character tech- c. First vertebral narrower than second nique of Hennig. The following hypotheses of Group 4. Tribe Chelini (, , Chelo- are tested using the characters in- dina, ) dicated (see fig. 10): a. Lateral margins of parietals distinctly re- Group 1. Family Chelidae (Pseudemydura, Emy- duced dura, , Platemys, Phrynops, Chelus, Group 5. Subtribe Chelina (Chelus, , Chelodina, Hydromedusa) Hydromedusa) a. Unusually developed lateral cheek emargi- a. longer than dorsal ver- nation tebrae b. Loss of quadratojugal b. Medial portions of jugal and postorbital c. Loss of mesoplastra facing more laterally than posteriorly Group 2. Chelinae (, Elseya, Group 6. Infratribe Hydromedusad (Chelodina, Platemys, Phrynops, Chelus, Chelodina, Hydromedusa) Hydromedusa) a. Posterolateral process of parietal absent a. Anterior frontal process at least partially b. Extremely reduced horizontal process of separating nasals parietal Group 3. Infrafamily Chelodd (Platemys, Phry- c. Quadrate-basisphenoid contact nops, Chelus, Chelodina, Hydromedusa) d. Four claws on forefoot a. Symphyseal suture separating lower jaw rami

INTRODUCTION The purpose of the present paper is to develop tles of the family Chelidae. These turtles are a theory of relationships for the side-necked tur- found in , with about 12 living

'Associate Curator, Department of Paleontology, the American Museum of Natural History; Adjunct Assistant Professor, Department of Geological Sciences, Columbia University.

Copyright O The American Museum of Natural History 1977 ISSN 0003-0082 / Price $1.90 2 AMERICAN MUSEUM NOVITATES NO. 2620 usually recognized, and are the dominant ABBREVIATIONS continental turtles of , where about 14 ANATOMICAL living species occur. The chelids are a predom- ang, angular pa, parietal inantly carnivorous, fresh-water aquatic family art, articular pal, palatine that contains some of the strangest turtles (e.g., bo, basioccipital pf, prefrontal Chelus) as well as some of the rarest (e.g., Pseu- bs, basisphenoid pm, premaxilla demydura). There have been few attempts at ex- cor, coronoid po, postorbital plicit phylogeny reconstruction for the chelid den, dentary pr, prearticular genera and little osteologic information is avail- epi, epipterygoid pt, pterygoid able in the literature. My intention here is a ex, exoccipital qi, quadratojugal phylogeny reconstruction relying primarily on fr, frontal qu, quadrate cranial characters. ju,jugal so, supraoccipital mx, maxilla sq, squamosal My method is derived from that of Hennig na, nasal sur, surangular (1966) and others, and is often termed phyloge- op, opisthotic vo, vomer netic systematics or cladism. As I see it, this method is the closest approach in systematics to the logical criteria emphasized by Popper (1968) INSTITUTIONS as characterizing science. Popper has argued that AMNH, the American Museum of Natural His- the best science is developed in terms of hypoth- tory, New York esis formation and test, in which the tests at- FMNH, Field Museum of Natural History, Chi- tempt to falsify rather than confirm the hypoth- cago esis. In phylogeny reconstruction, hypotheses of NMNH, National Museum of Natural History, relationship are tested by character distributions Smithsonian Institution, Washington, D.C. MCZ, Museum of Comparative Zoology, Harvard in which the characters are analyzed in terms of University, Cambridge primitive and derived. The analysis consists of a WAM, Western Australian Museum, Perth further series of testable hypotheses. Testability, i.e., the potential to criticize and falsify a hy- ACKNOWLEDGMENTS pothesis, is the critical feature. The logical as- pects of phylogenetic systematics and a summary Once again I am grateful to Dr. Samuel B. of the method of using shared derived characters McDowell for allowing me free access to his work in phylogeny reconstruction are presented else- on chelids. Dr. Glen Storr of the Western where (Gaffney, In press; Wiley, 1975) and the Australian Museum very kindly lent me reader is referred to these works and more gen- specimens of Pseudemydura. Dr. Richard eral references (Bonde, 1974; Brundin, 1968; Zweifel, the American Museum of Natural Cracraft, 1972, 1974; Eldredge and Tattersall, History, Dr. George Zug, National Museum of 1975; Hennig, 1965). Natural History, Smithsonian Institution, Dr. Although chelids are not discussed here, Hymen Marx, Field Museum of Natural History, this is due more to the lack of skull material than and Dr. Emest Williams, Museum of Comparative lack of interest. I am currently engaged in a Zoology, Harvard University, helped in finding study of fossil chelids from Australia, including and lending me material in their care. Mr. John some good skull material of age. Roger Goode of Frankston, Victoria, obtained Aus- Wood has (in preparation) a study of fossil tralian specimens for me and I appreciate his chelids from South America. Other literature ref- help. erences on fossil chelids may be found in Kuhn (1964) and Mlynarski (1976). PREVIOUS WORK The higher category classification of turtles In the section on Basic Taxa (below) I include used here is developed in Gaffney (1975), and some of the more important literature references the anatomic terminology can be found in Gaff- to particular chelid genera, and here I discuss ney (1972b). some of the more pertinent literature concerning 1977 GAFFNEY: CHELIDAE 3

H

FIG. 1. Lateral views of chelid skulls, measurements are midline condylo-premaxilla length. A. Emydura macquarrii (AMNH 110486; 49 mm.). B. Pseudemydura umbrina (WAM R29341; 35 mm.). C. Platemys platycephala (AMNH 74811; 28 mm.). D. (AMNH 79048; 60 mm.). E. Hydromedusa tectifera (FMNH 31032; 51 mm.). F. Phrynops () gibba (FMNH 45669; 37 mm.). G. (AMNH 108948; 75 mm.). H. Chelus fimbriata (AMNH 108955; 104 mm.). 4 AMERICAN MUSEUM NOVITATES NO. 2620 phylogeny and reviews of the whole family. Werner (1909) provided more limited infor- Attempts at phylogeny reconstruction involving mation. all or most of the genera in this family are The South American chelids are more poorly virtually nonexistent. Boulenger (1889) and Gray known than the Australian forms and there are (1864) presented "natural" keys, which may be no reviews of them. In addition to the works interpreted as phylogenies. Burbidge, Kirsch, and listed above, useful studies dealing with more Main (1974) come closest to a chelid phylogeny than one are Froes (1957), Luederwaldt even though their interests were primarily in the (1926), Siebenrock (1904), and Strauch (1890). Australian forms. On the basis of a phenetic analysis of biochemical data, they conclude that BASIC TAXA Emydura and Elseya have an ancestor in common not found in common with other The theory of chelid relationships developed forms. The three taxa Pseudemydura, here, uses seven generic level taxa as the Emydura-Elseya, and Chelodina are equally fundamental units in the hypothesis. The logical related; that is, they form a trichotomy in their nature of this hypothesis requires that these basic analysis. Furthermore, some South American taxa be strictly monophyletic, but rigorous tests taxa were studied serologically leading to a of monophyly, including studies of species conclusion that the Australian species form one distribution, etc., are beyond the scope of this monophyletic group and the South American paper. I do find it necessary, however, to make a species form another monophyletic group. In partial examination of this problem. general, phenetic studies using morphological My principle difficulty is the small sample of data can be resolved into primitive and derived specimens available for the named genera, and character states (or convergences, etc.) and a the absence of any cranial material objectively phylogenetic study can then deal with the identifiable for many of the species. I have done characters. In the case of serology and related my best with the material at hand, but I suspect methods have been that some of the characters will be subject to techniques numerical more variation than I have indicated. Nonethe- suggested (e.g., Farris, 1972) to analyze phenetic less, I doubt that a more extensive examination data in terms of primitive and advanced of specimens will seriously alter the character dis- characters. Nonetheless, I am not convinced of tributions as I have reported them. In my opin- the usefulness of this approach and, in any case, ion, a more important source of further tests is in Burbidge, Kirsch, and Main (1974) do not pre- other organ systems, such as jaw musculature, sent such an analysis. Although they have pro- hyoid apparatus, limb musculature, and limb duced an interesting phylogeny, albeit distinctly osteology. Only by increasing the number of at variance with mine, I cannot point out areas of areas examined can phylogenetic hypotheses be weakness (or strength) or attempt to resolve it discarded or substantiated. with my character distributions. There are no detailed reviews of the Chelidae, but Boulenger (1889), Gray (1855, 1870), A. Pseudemydura Siebenrock (1909), and Wermuth and Mertens Figures lB, 2B, 4B, 6B (1961) are the best sources for lists of species Specimens Examined. Pseudemydura um- and synonyms. The Australian chelids are brina, WAM R29341, Twin Swamp Reserve, reviewed by Goode (1967) who presented keys, , Australia; Pseudemydura um- figured living specimens and types, and provided brina, WAM R21859, Bullsbrook Reserve, West- a synonymy and bibliography (the last suffers ern Australia, Australia; Pseudemydura umbrina, from errors, however). Other important studies WAM R29338, Ellenbrook Reserve, Western Aus- of the Australian taxa are: Boulenger (1888), tralia, Australia. Burbidge, Kirsch, and Main (1974), Cogger Discussion. Pseudemydura umbrina has had an (1975, distribution maps), de Rooij (1915), interesting taxonomic history, being first named Ogilby (1905), Waite (1929), and Worrell (1963); by Siebenrock (1901; figured and described in whereas Blackmore (1969), Strauch (1890), and 1907; these figures are repeated in Williams, 1977 GAFFNEY: CHELIDAE 5

1958) on the basis of one preserved specimen. differentiate them consistently using cranial char- Subsequently, Glauert (1954) described a form acters. However, this may be due to my ex- he called Emydura inspectata, which Williams tremely small sample of adequately identified (1958) identified as Pseudemydura thereby "re- Elseya skulls. Burbidge, Kirsch, and Main (1974) discovering" what is possibly the rarest Recent and Goode (1967) used features of the intergular turtle. Goode (1967) has photographs of living scute, cervical scute, skull cap, snout, and post- specimens and the Siebenrock type specimen as orbital skin tuberculation, as well as serology in well as sketches of the skull. Burbidge, Kirsch, the case of Burbidge, Kirsch, and Main (ibid.) to and Main (1974) gave good shell and skull figures separate the named species into two genera. (including the lower jaw). Morphologic information substantiates Emy- I interpret the following features of Pseude- dura-Elseya monophyly. Emydura and Elseya mydura as autapomorphies; that is, derived fea- have heavier lower jaws with wider triturating tures found only in this species: areas and slightly developed symphyseal "hooks" in contrast to all other chelids, although there is 1. Quadrate-parietal contact (fig. 1B) some variation in this feature. This would appear 2. Supraoccipital laterally expanded in contrast to be derived within the Chelidae but the com- to other Chelidae (fig. 2B) mon possession of heavy lower jaws among pelo- 3. Parietal laterally expanded (fig. 2B) medusids weakens the use of this feature. Also, 4. Postorbital ventrolaterally expanded (fig. 2B) megacephaly seems to be common in Emydura 5. Anterior extension of squamosal (figs. 1 B, 2B) and Elseya (Goode, 1967) and may occur in 6. Prearticular separating coronoid and splenial other chelids such as Phrynops. Nonetheless, (Burbidge, Kirsch, and Main, 1974) at present it seems best to treat Emydura and 7. Medial approximation of maxillae along labial Elseya as a strictly monophyletic assemblage. ridge or separating nearly separating pre- Other References. Boulenger (1888, 1889; maxlllae into anterior and posterior portions skull figures); Burbidge, Kirsch, and Main (1974, (fig. 4B) skull figures); Gaffney (1975, skull figures); The argument that characters 1 through 6 are Goode (1967, skull figures); Gray (1863a, autapomorphies is developed below in the sec- 1863b, 1872); Hoffmann (1890, skull figures); tion on Group 2. Krefft (1876); Loveridge (1934); Ogilby (1905); Ouwens (1914); Peters and Doria (1878); Sieben- B. Emydura-Elseya rock (1906, 1907, 1912); Vogt (1911). Figures IA, 2A, 4A, 6A, 7 Specimens Examined. Elseya latisternum, C. Platemys AMNH 103700, Bulimba Creek, Brisbane, Figures IC, 2C, 4C, 6C ; Emydura macquarrii, AMNH 77637, Specimens Examined. Platemys platycephala, no data; Emydura macquarrii, AMNH 77648, no AMNH 74811, no data; Platemys platycephala, data; Emydura macquarrii, AMNH 11487, South AMNH 75101, no data; Platemys platycephala, Australia, Australia; Emydura macquarrii, AMNH FMNH 45659, Loreto, . 110486, South Australia, Australia; Emydura Discussion. My sample ofPlatemys skulls con- macquarrii, AMNH 110488, 40 mi. SE Mildura, sists of three specimens of P. platycephala, which Victoria, Australia; Emydura macquarrii, AMNH show the following autapomorphous features: 108962, Patho, Victoria, Australia; Emydura truncated and reduced crista supraoccipitalis, macquarrii, AMNH 103702, Victoria, Australia; lateral edges of parietal parallel and orbits rela- Emydura kreffti, AMNH 72406, no data; Emy- tively large (presumably correlated with small dura kreffti, AMNH 108958, Queensland, Aus- size of adult ). Again, the absence of tralia; , AMNH 108957, Dar- skulls identifiable as P. pallidipectoris, P. spixi, win area, Northern Territory, Australia. and P. radiolata hamper the usefulness of these Discussion. I am treating these two genera as criteria. one basic because I have been unable to One of the principle shell features used to 6 AMERICAN MUSEUM NOVITATES NO. 2620

A B

C

FIG. 2. Dorsal views of chelid skulls. Skull measurements in figure 1. A. Emydura macquarrii (AMNH 110486). B. Pseudemydura umbrina (WAM R29341). C. Platemys platycephala (AMNH 74811). D. Phrynops geoffroanus (AMNH 79048). 1977 GAFFNEY: CHELIDAE 7

A

C D

FIG. 3. Dorsal views of chelid skulls. Skull measurements in figure 1. A. Hydromedusa tectifera (FMNH 31032). B. Phynops (Mesoclemmys) gibba (FMNH 45669). C. Chelodina expansa (AMNH 108948). Chelus fimbriata (AMNH 108955). 8 AMERICAN MUSEUM NOVITATES NO. 2620

pm pm

mx

pal j AX ~~~~Bv

pm pm

CX DX

FIG. 4. Palatal views of chelid skulls. Skull measurements in figure 1. Asterisks (*) show position of foramen posterius canalis carotici interni. A. Emydura macquarrii (AMNH 110486). B. Pseudemy- dura umbrina (WAM R2934 1). C. Platemys platycephala (AMNH 74811). D. Phrynops geoffroanus (AMNH 79048). 1977 GAFFNEY: CHELIDAE 9

pm p

pmpmx

pal ju~~~~~~~~~~~~~~~~~~~~j *~~9 X

C ex sqD

FIG. 5. Palatal views of chelid skulls. Skull measurements in figure 1. Asterisks (*) show position of foramen posterius canalis carotici interni. A. Hydromedusa tectifera (FMNH 31032). B. Phrynops (Mesoclemmys) gibba (FMNH 45669). C. Chelodina expansa (^AMNH 108948). D. Chelus fimbriata (AMNH 108955). 10 AMERICAN MUSEUM NOVITATES NO. 2620 characterize Platemys is the presence of a trough called Hydraspis, and, finding Rhinemys Wagler, along the carapace midline, but this feature is 1830, to be a of Phrynops, erected apparently absent in Platemys radiolata, (Frei- Batrachemys to replace it. Therefore, we find berg, 1945). later works, such as Wermuth and Mertens Other References. Dunn (1945); Freiberg (1961) with the following classification (give or (1945, 1947); Froes (1957); Luederwaldt take a few species): (1926); Medem (1960a); Mertens (1967); Muller (1939); Siebenrock (1897, 1904, skull figures); Batrachemys nasuta Wagler (1830). Batrachemys dahli (erected by Zangerl and Me- dem, 1958) D. Phrynops Figures 1D, 1F, 2D, 3B, 4D, 5B, 6D, 6F Phrynops geoffroanus (with three : ge- offroanus, hilarii, tuberosus) Specimens Examined. Phrynops geoffroanus, Phrynops rufipes AMNH 79048, no data; Phrynops geoffroanus, AMNH 58201, no data; Phrynops geoffroanus, Zangerl and Medem (1958), however, in allu- AMNH 58110, ?Peru; Phrynops (Batrachemys) sion to a study in progress at that time by nasuta, AMNH 108908, no data;Phrynops (Ba- Williams and Vanzolini (unpublished), stated ) sp., AMNH 58123, Iquitos, Peru; that the three genera Batrachemys, Mesoclemmys, Phrynops (Batrachemys) nasuta, MCZ 58099; and Phrynops were closely related and should be Orinoco River, ; Phrynops (Batra- placed in one genus, Phrynops, with the three chemys) nasuta, MCZ 1456, Pernambuco, ; former genera recognized as subgenera. Bour Phrynops (Mesoclemmys) gibba, FMNH 45669, (1973) has argued that even these taxa are not Yarinacocha, Loreto, Peru; Phrynops (Mesoclem- objectively recognizable, but neither he nor mys) gibba, FMNH 45671, Yarinacocha, Loreto, Zangerl and Medem (ibid.) gave a diagnosis of Peru. Phrynops in the larger sense. As Bour (ibid.) has Discussion. The species here included in this suggested, the Zangerl and Medem concept of genus have had a particularly confusing history Phrynops is essentially the same as that of Gray's (1864) Hydraspis, and we appear to have come of generic assignment. Some of the earlier as- full circle. pects may be obtained from Gray (1855, 1864, For the purposes of this study I use Phrynops 1870). In 1909, the taxa involved were classified in Bour's sense, without subgenera, even though I by Siebenrock (closely following Boulenger, cannot rigorously support its strict monophyly. I 1889, and Siebenrock's own work of 1904) as have been unable to find unique derived charac- follow: ters in the skull ofPhrynops but I have also been Rhinemys nasuta unable to find derived characters in common be- Mesoclemmys gibba tween some of the species in Phrynops and Hydraspis hilarii Chelodina, Hydromedusa, or Chelus. Therefore, Hydraspis geoffroyana at present I can falsify neither the hypothesis Hydraspis tuberosa that Phrynops is monophyletic nor the hypoth- Hydraspis rufipes esis that it is paraphyletic. Even if Phrynops is Hydraspis wagleri paraphyletic it will not drastically alter the phylogenetic hypothesis advanced here. Stejneger (1909) showed that the type species Other References. Albrecht (1976, cranial of Hydraspis Bell is longicollis Bell, arteries); Boulenger (1889, skull figures); Dunn 1828, and as this species was earlier made the (1945); Froes (1957); Gray (1873); Kanberg type species of Chelodina Fitzinger, 1826, the (1926); Luederwaldt (1926); Medem (1960a, genus Hydraspis is a junior synonym of Chelo- 1960b, 1960c, 1966, 1973); Mertens (1967, dina. He also argued that Phrynops Wagler, 1830, 1969, 1970); Muller (1939); Siebenrock (1904, is the correct name for the taxon previously 1905); Zangerl and Medem (1958, skull figures). 1977 GAFFNEY: CHELIDAE 11

pt qu foramen jugulare posterius C foramen posterius canalis carotici interni

-NJ''I /Ibo pt \LH I foramen jugulare posterius I foramen jugulare posterius G fenestra postotica H fenestra postotica FIG. 6. Occipital views of chelid skulls. A. Emydura macquarrii (AMNH 76199). B. Pseudemydura umbrina (WAM R29341). C. Platemys platycephala (AMNH 74811). D. Phrynops geoffroanus (AMNH 79048). E. Hydromedusa tectifera (NMNH 15189). F. Phrynops (Mesoclemmys) gibba (FMNH 45669). G. Chelodina expansa (AMNH 108948). H. Chelus fimbriata (AMNH 108955). 12 AMERICAN MUSEUM NOVITATES NO. 2620

E. Chelus 108951, Patho, Victoria, Australia; Chelodina Figures 1H, 3D, SD, 6H longicollis, AMNH 76569, no data; Chelodina longicollis, AMNH 108952, Patho, Victoria, Aus- Specimens Examined. Chelus fimbriata, tralia; Chelodina novaeguineae, AMNH 57589, AMNH 108955, no data; Chelus fimbriata, Mabaduane, Papua, New Guinea; Chelodina no- AMNH 111962, no data; Chelus fimbriata, vaeguineae, AMNH 86547, Armraynald, Queens- AMNH 6596, no data; Chelus fimbriata, AMNH land, Australia; Chelodina expansa (?), AMNH 43298, no data. 103699, Bulimba Creek, Brisbane, Queensland, Discussion. Only one Recent species of this Australia; Chelodina expansa, AMNH 108948, genus is usually recognized, but in any case, Patho, Victoria, Australia; Chelodina expansa, Chelus is riddled with autapomorphies and looks AMNH 108949, Patho, Victoria, Australia; Che- as if it had been run over by a truck. The follow- lodina rugosa (?), AMNH 104338, Mt. Burnett, ing cranial features I interpret as unique derived Western Australia, Australia; Chelodina rugosa, characters for this taxon: AMNH 82532, Cape York Peninsula, Queens- land, Australia; Chelodina rugosa, AMNH 1. Nasals absent (fig. 3D) 108954, Darwin area, Northern Territory, Aus- 2. Prefrontal broadly exposed along dorsal mar- tralia. gin of apertura narium externa [narrowly ex- Discussion. Chelodina has a series of charac- posed in Chelodina] (fig. 3D) ters that I am hypothesizing as autapomorphies 3. Pterygoids extend anteriorly into apertura or unique derived characters. narium interna and often separate vomer from palatines (fig. SD) 1. Nasals usually separated by anterior processes 4. Extreme flattening of skull, particularly in of frontals (fig. 3C) center (fig. 1H) 2. Frontals fused along midline (fig. 3C) 5. Cavum tympani extended laterally to consid- 3. Temporal bar absent (fig. 3c) erable degree (fig. 3D) 4. Extensive quadrate-basisphenoid contact (fig. 6. Medial processes of jugal and postorbital lie 5C) entirely on external surface of skull [see dis- Characters 1 and 2 are unique in turtles, al- cussion under Group 5] (fig. 3D) though the frontal in Chelus reaches the margin 7. Maxilla relatively reduced in exposure on tri- of the apertura narium externa the nasals are turating surface so that palatine bears lingual absent in that form. The temporal bar is absent ridge (fig. 5D) in some cryptodires but the remaining roofing are of distinctly different morphology. An Other References. Boulenger (1889, skull fig- extensive quadrate-basisphenoid contact occurs ures); Dunn (1945); Froes (1957); Fuchs (1931, in pelomedusids (e.g., ) but again, the lower jaw figures); Gregory (1946, skull figures); morphology of the bones involved is inconsistent Hoffmann (1890); Luederwaldt (1926); Medem with the hypothesis that the contacts are homol- (1960a); Muller (1939); Siebenrock (1897, skull ogous. A limited quadrate-basisphenoid contact figures); Wagler (1830, skull figures). occurs in Hydromedusa and I hypothesize that the limited condition is primitive for Hydro- medusa and Chelodina. F. Chelodina These characters are consistent with strict Figures 1G, 3C, 5C, 6G, 8 monophyly of Chelodina. Burbidge, Kirsch, and Specimens Examined. Chelodina steindach- Main (1974) presented serologic data which they neri, AMNH 101978, Woodstock, Western Aus- conclude also argues for Chelodina monophyly. tralia, Australia; Chelodina longicollis, AMNH Rhodin and Mittermeier (1976) gave good 108953, no data; Chelodina longicollis, AMNH descriptions and figures of the skull in Chelodina 108950, no data; Chelodina longicollis, AMNH siebenrocki and their new species, C. parkedi. 108947, no data; Chelodina longicollis, AMNH They provided a glossary of anatomical terms 1977 GAFFNEY: CHELIDAE 13

pm

na mx

fr V a ju ju PO

pa~~~~~~~~~~~~~~~~~~~P

qu

opa A

so

pf sq~~~~s

foramen jugulare posterius FIG. 7. Elseya latisternum (AMNH 103700; 53 mm.). used to describe systematically important fea- (1922); Goode (1967, skull figures; 1968); tures. Goode and Russell (1968); Gray (1856, 1869, Other References. Boulenger (1888); Bur- skull figures); Hoffmann (1890, skull figures); bidge, Kirsch, and Main (1974, skull figures); Fry Loveridge (1934); Ogilby (1890, 1905); Schnee (1915); Gaffney (1975, skull figures); Glauert (1899); Siebenrock (1897, 1905, 1914, skull fig- 14 AMERICAN MUSEUM NOVITATES NO. 2620 ures); Stejneger (1909); Vestjens (1969); Vogt mary of the shared derived characters may be (1911); Waite (1929, skull figures); Werner found in the Abstract. See also table 2 for cranial (1901); Worrell (1961). characters discussed here and in the Basic Taxa section. G. Hydromedusa Figures lE, 3A, 5A, 6E GROUP 1-FAMILY CHELIDAE Specimens Examined. Hydromedusa maxi- Table 1 is a comparison of and miliani, MCZ 2856, Brazil; Hydromedusa tecti- Chelidae using a series of characters that test fera, NMNH 15189; Hydromedusa tectifera, monophyly for both families. I have elsewhere FMNH 31032. (Gaffney, 1975) argued that pleurodires are Discussion. Hydromedusa has the following strictly monophyletic, and that discussion should unique derived features: be consulted. I am including Recent and fossil taxa to the extent that they are available. 1. Relatively large bony apertura narium interna The chelids have the following synapomor- formed by reduced ossification of palatine phies or shared derived characters: (1) Unusually [the fleshy internal narial openings may not developed lateral cheek emargination; (2) loss of be enlarged] (fig. SA) quadratojugal; (3) loss of mesoplastra. 2. Prefrontals meet in midline and may overlap Although Pseudemydura has relatively less anterior processes of frontals so that frontals cheek emargination than the other chelids it still are exposed anterior and posterior to the pre- differs strongly from pelomedusids in the shape frontal contact (fig. 3A) and number of bones bordering the emargina- 3. The cervical (nuchal) scute is relatively large tion. Also, there is reason to think that the ex- and separated from the anterior edge of the tensive in Pseudemydura is derived for shell by medial contact of the first pair of chelids and that the condition in Emydura is the marginal (Boulenger, 1889) primitive one for chelids. The parietals, squamo- The large bony internal nares and the median sals, postorbitals, and supraoccipitals of Pseu- prefrontal contact occur in no other chelids (al- demydura are different in their extension and though they do occur in many other turtles), and shape not only from other chelids but also from it seems most parsimonious to consider them in- pelomedusids. If the Pseudemydura pattern were dependently derived in Hydromedusa rather than primitive, one would expect fo find it in pelome- a primitive retention. The shell feature is unique dusids or cryptodires. among turtles. A further aspect of the emphasis in chelids on Wood and Moody (1976) provide figures for cheek emargination is the loss of the quadra- the shells and descriptions of the shells in Hydro- tojugal in all members of this family. Even Pseu- medusa maximiliani and H. tectifera,' as well as demydura, a form possessing a well-developed shell characters allowing recognition of these two temporal roof, lacks a quadratojugal, further sug- forms. gesting that it evolved from a more emarginate Other References: Froes (1957); Hay (1908, ancestor. Some species of Cuora, Hieremys, skull figures); Luederwaldt (1926); Mertens , and Terrapene also lack a quadrato- (1967); Muller (1939); Peters (1839, skull fig- jugal, and in these forms it appears to be associ- ures); Wagler (1830, skull figures). ated with well developed cheek emargination. Chelids lack mesoplastra, the presence of PHYLOGENETIC HYPOTHESIS which is presumably primitive for pleurodires. The following sections should be read with Although mesoplastra were lost independently the cladogram (fig. 10) in mind. The group num- within the cryptodires, there is no evidence that bers refer to numbers on the cladogram. A sum- this has happened more than once within the pleurodires. ' Note that the recessed nuchal with peripheral bones meeting medially found in Hydromedusa maxi- GROUP 2-SUBFAMILY CHELINAE miliani also occurs in the extinct baenoid (Gaffney, 1972a) and is not unique to Hydromedusa Pseudemydura has a number of unique fea- as stated by Wood and Moody. tures but only one of them would appear to be 1977 GAFFNEY: CHELIDAE is

pm

foramen posterius canalis carotici interni FIG. 8. Chelodina novaeguineae (AMNH 57589; 41 mm.). primitive for the Chelidae. In all other chelids the frontal. No other living turtles possess nasals but frontal has a process that extends anteriorly in fossil turtles (baenoids, toxochelyids, plesio- (with the other frontal) along the median suture chelyids, chelosphargine protostegids, Solnhofia, to partially separate the nasals. In Pseudemydura and ) that do have nasals, the fron- (figs. 2, 3) the nasals slightly separate the frontals tals do not separate the nasals. Therefore, the and there is only a short anterior process on the condition in all chelids except Pseudemydura 16 AMERICAN MUSEUM NOVITATES NO. 2620

TABLE 1 A Comparison of the Pelomedusidae and Chelidae (Including information on Recent and fossil forms.) Character Pelomedusidae Chelidae

1. Nasals absent present (except in Chelus) 2. Prefrontals meet in midline do not meet in midline (except in Hydromedusa) 3. Posterior temporal varies from virtually absent usually poorly developed with emargination (e.g., Dacquemys) to persistent squamosal- extensive (e.g., Pelomedusa) parietal contact (except in Chelodina) 4. Cheek (lateral variable, but never developed to developed to an unusual temporal emargination) the extent seen in Chelidae degree with only a parietal- squamosal bar remaining 5. Quadratojugal present absent 6. Triturating surface usually broader usually narrower 7. Vomer usually absent present 8. Splenial absent present 9. Mesoplastra present absent 10. Cervical (nuchal) scute absent present (except in most Elseya and as an infrequent variation) 11. Cervical vertebraea second biconvex fifth and eighth biconvex a(Wffliams, 1950)

would appear to be derived for pleurodires and a Similarly the second alternative (fig. 9B) also useful test for monophyly of the non- fails from the lack of a derived character in com- Pseudemydura chelids. mon between Pseudemydura and all chelids ex- I am interpreting the other unique features of cept Emydura. Pseudemydura as autapomorphies, that is, de- My contention that Pseudemydura is the sister rived characters found only in this species and, taxon to all other chelids should not be thought therefore, not useful in phylogeny reconstruc- of as an argument that all of its morphology is tion. primitive. The nasal-frontal morphology does This particular hypothesis, namely that Pseu- seem primitive, but other distinctive features of demydura is the sister taxon to the other chelids the skull roof seem to be autapomorphies or ad- is perhaps the weakest aspect of my chelid phy- vanced features unique to this species. The exten- logeny, and I would like to discuss some alterna- sive temporal roof ofPseudemydura is best inter- tives at this point. One alternative (fig. 9A) preted as a unique derived condition for chelids would have Emydura and Pseudemydura as sister rather than a primitive one. In fact, comparison taxa, that is, with an ancestor in common not in with pelomedusids and cryptodires suggests to common with any other turtle. However, Pseu- me that the Emydura type of temporal roof with demydura and Emydura have few unique charac- an extensive lateral or cheek emargination and a ters in common. The cervical vertebrae (particu- shallow posterior emargination is probably primi- larly the anterior ones) of these two genera do tive for chelids. The absence of a quadratojugal have zygapophyses that are more widely sepa- in Pseudemydura is consistent with a hypothesis rated than in other chelid genera but this feature of expansion of the parietals, postorbitals, and is presumably primitive for pleurodires and quite supraoccipital into the emarginated areas. In unsatisfactory for corroborating monophyly. other words, my hypothesis of relationships 1977 GAFFNEY: CHELIDAE 17

FIG. 9. Alternate hypotheses relating Emy- 4 dura, Pseudemydura, and remaining chelids. A third, but preferred alternative, is presented in fig- ure 10. See text for discussion. 2 would have the primitive chelid possessing most of the features of Emydura except the nasal- frontal morphology in which it would resemble Pseudemydura In the lower jaw of Pseudemydura the pre- FIG. 10. A theory of relationships of the chelid turtles. The numbers and associated bars articular extends anteriorly to separate or nearly refer to groups and character distributions dis- separate the coronoid and splenial (Burbidge, cussed in the text (see Abstract for list). The Kirsch, and Main, 1974, p. 389). In all other solid black circles are Australian genera, the chelids the coronoid and splenial have an exten- open circles South American. sive contact. Few turtles retain a splenial, but in those that do (i.e., Solnhofia, baenids, Plesio- on the dorsal margin of the foramen. In Emydura chelys), the coronoid and splenial have a well- and Pseudemydura the exoccipitals do not meet, developed contact. Therefore, the prearticular although they very nearly do so in Pseudemy- extension in Pseudemydura is best interpreted as dura. a unique derived character. As noted by Boulenger (1889), Emydura and Elseya have the first vertebral scute narrower GROUP 3-INFRAFAMILY CHELODD than the second, whereas the other chelids have Platemys, Phrynops, Chelus, Chelodina, and the first vertebral wider than the second. Pseu- Hydromedusa all have a symphyseal suture sepa- demydura (unknown to Boulenger) also has the rating the two lower jaw rami. Pseudemydura first vertebral scute narrower than the second, and Emydura have the rami fused as do all other giving this character the same distribution as the turtles except for , fide Bramble, two characters discussed above. A comparison of [MS], whereas most have the rami sep- this condition with pelomedusids unfortunately arated. The appropriate out-group comparison does not allow a useful test of polarity (primitive must be with cryptodires rather than other vs. derived). Pelomedusids have both conditions reptiles and I conclude that separate rami are a although and Pelomedusa, forms that I derived feature corroborating common ancestry would consider as generally plesiomorphic pelo- of Platemys, Phrynops, Chelus, Chelodina, and medusids, have the narrower first vertebral scute. Hydromedusa. In any case, the character does not falsify the Another feature with this distribution involves hypothesis advanced here. the dorsal processes of the exoccipitals (fig. 6). In the five genera indicated these dorsal processes GROUP 4-TRIBE CHELINI extend dorsomedially to meet each other in a In Phrynops, Chelus, Chelodina, and Hydro- sagittal suture above the foramen magnum and medusa the lateral margins of the parietals are prevent the usual exposure of the supraoccipital distinctly reduced in comparison to the degree of 18 AMERICAN MUSEUM NOVITATES NO. 2620 emargination seen in Emydura and Platemys simonious to accept my hypothesis and reject the (figs. 2, 3). I interpret this more emarginate con- neural bone test because the latter is only one dition as derived for the four taxa indicated and character, whereas my hypothesis suggests sev- suggest that this is consistent with the hypothesis eral. The acceptance of neural absence as being that they are strictly monophyletic. A reduced consistent with strict monophyly of Pseu- squamosal-parietal arch also characterizes this demydura, Emydura-Elseya, and Platemys re- group, except Chelus. quires a number of ad hoc hypotheses to invoke The above four genera also differ from other character convergence. The rejection of the neu- chelids in generally having neurals; the other gen- ral features requires the acceptance of one of the era usually lack them (table 3; see also Boulen- following ad hoc hypotheses: (1) the absence of ger, 1889 for figures). Chelodina has neurals in neurals is a primitive feature for chelids and the only one species, C. oblonga (Burbidge, Kirsch, presence of neurals in the Tribe Chelini (Group and Main, 1974), however. The presence of neu- 4) is an example of convergence, or (2) the three rals would presumably be primitive and their ab- generic level taxa lacking neurals lost them inde- sence derived, making this character distribution pendently. Neither hypothesis is supported by an important contradiction to my hypothesis. other tests and this remains an important prob- Nonetheless, I am concluding that it is more par- lem area. TABLE 2 Cranial Features of Chelid Genera Emydura & Pseudemydura Elseya Platemys Phrynops Chelus Chelodina Hydromedusa

1. Nasals present present present present absent present present 2. Anterior absent present present present present present present process of frontal 3. Nasals no no no no nasals yes no completely absent separated by anterior frontal pro- cess 4. Prefront- no no no no broadly narrowly no als exposed along dorsal margin of apertura narium ex- terna S. Prefront- no no no no no no yes als meet in midline so that front- als are ex- posed an- terior and posterior to prefrontal contact 1977 GAFFNEY: CHELIDAE 19

TABLE 2 - (Continued) Emydura & Pseudemydura Elseya Platemys Phrynops Chelus Chelodina Hydromedusa 6. Frontals no no no no no yes no fused 7. Dorsal por- large small small small small small small tion of postorbital 8. Temporal very exten- moderate; moderate; moderate; moderate; absent very slender; arch sive; formed formed by formed by formed by formed by formed by by parietal, squamosal squamosal squamosal squamosal squamosal squamosal and parie- and parie- and parie- and parie- and supra- and supra- tal tal tal tal occipital occipital 9. Skull no no no no extreme moderate moderate flattened 10. Dorsal (hor- broadly covers cen- covers cen- covers little covers little absent; absent; covers izontal) por- covers ad- tral area of tral area of of adductor of adductor covers none none of ad- tion of ductor fossa adductor adductor fossa but fossa of adductor ductor fossa parietal fossa but fossa but still present fossa not lateral not lateral (although area area greatly re- duced in some) 11. Lateral sub- tapering parallel wasp- wasp- tapering tapering edges of parallel but anteriorly waisted waisted posterior- posteriorly, parietals laterally ex- ly, greatly greatly re- tensive reduced duced 12. Supra- present absent absent absent absent absent present occipital- (temporal parietal bar absent) contact 13. Quadrate- present absent absent absent absent absent absent parietal contact 14. Dorsal hor- broadly not ex- slightly ex- not ex- not ex- not ex- slightly ex- izontal por- expanded panded panded panded panded panded panded tion of supraocci- pital 15. Crista supra- does not ex- extends be- more re- extends be- does not ex- does not ex- does not ex- occipitalis tend be- yond for- duced than yond for- tend be- tend be- tend beyond yond fora- amen mag- in any amen mag- yond fora- yond fora- foramen mag- men magnum num and other chelid, num and men mag- men mag- num condylus does not ex- condylus num num occipitalis tend beyond occipitalis foramen magnum 20 AMERICAN MUSEUM NOVITATES NO. 2620

TABLE 2 - (Continued) Emydura & Pseudemydura Elseya Platemys Phrynops Chelus Chelodina Hydromedusa

16. Medial por- no no no no yes, entire- yes yes tions of ly on exter- jugal and nal surface postorbital of skull facing more laterally than pos- teriorly 17. Dorsal pro- no no yes yes yes yes yes cesses of exoccipi- tals meet above for- amen mag- num 18. Cavum no no no no yes no no tympani extended laterally 19. Maxilla re- no no no no yes no no duced in ex- posure on triturating surface so that palatine bears lingual ridge 20. Medial max- yes no no no no no no illary con- tact divid- ing premax- illae longi- tudinally 21. Vomer- yes yes yes yes no, due to yes no, due to en- palatine anterior larged apertura contact pterygoid narium interna processes reducing reaching palatine vomer 22. Large bony no no no no no no yes apertura narium in- terna formed by reduc- tion of pala- tines 1977 GAFFNEY: CHELIDAE 21

TABLE 2 - (Continued) Emydura & Pseudemydura Elseya Platemys Phrynops Chelus Chelodina Hydromedusa 23. Quadrate- absent absent absent absent absent extensive limited basisphe- noid contact 24. Symphy- no no yes yes yes yes yes seal suture separates lower jaw rami 25. Relatively absent present absent absent absent absent absent massive mandibles with symphy- seal "hooks" 26. Prearticular yes no no no no no no separates or nearly sepa- rates coro- noid splenial

GROUP 5-SUBTRIBE CHELINA the jugal along with the postorbital make up the characteristic postorbital wall found in this group Boulenger (1889, p. 207) divided the "Chely- (Gaffney, 1975). The jugal usually forms the didae" into two sections: "I. Neck longer than more lateral portion of the wall, whereas the the dorsal vertebral column. . . ." and "II. Neck postorbital forms the more medial area. As in shorter than the dorsal vertebral column .. cryptodires the jugal of pleurodires may reach and this distinction, as emphasized by Goode the palatine but there is always a strong contact (1967) and Burbidge, Kirsch, and Main (1974) with the anterior edge of the pterygoid, pre- still has usefulness for systematists. The ex- sumably to aid in support of the uniquely pleu- tremely long cervical vertebrae of Chelus, Chelo- rodiran processus trochlearis pterygoidei. In the dina, and Hydromedusa may be hypothesized as primitive condition both bones consist of two a shared derived character testing the monophy- portions, a medial section exposed behind the letic nature of this group. In all other chelids the fossa orbitalis and forming the front of the fossa cervicals are shorter than the length of the verte- temporalis inferior, and a lateral portion exposed brae attached to the carapace and, as this latter on the external surface of the skull. This situa- condition is found in nearly all other turtles (as tion occurs in pelomedusids, which is the basis far as I know), it is presumably primitive. for considering it primitive. It also occurs in Another feature found only in Chelus, Chelo- Emydura, Pseudemydura, Platemys, and Phry- dina, and Hydromedusa involves the jugal and nops. In Chelus, Chelodina, and Hydromedusa, postorbital bones (figs. 1, 2, 3).1 In pleurodires however, the medial portion, which is covered by musculature (except in Chelus, see below), is 1 Rhodin and Mittermeier (1976) described the turned outward to face somewhat more laterally jugal and postorbital areas in Chelodina and developed rather than more posteriorly as in other chelids a useful terminology for it. Their figures 4 and 6 should (Chelodina novaeguineae, fig. 8, is less like the be consulted as an aid to identification. other forms of Chelodina and more like a) .0 := ,O 4) W 0 +-b CIS CIS .g r. 2? 0 I = t to .; Ei 4) a) bI2 & 0 > !I . :. Ei (.D 02 -0 0

q6) 4) = 4~~~a ca1., U °t E ° E " o C, i ak) --I C >. . - +.

ua)0 a) = E- a) a) 0 a) ,0

0 0 0 3. a) S O^-, 0 a) £E Cd 16 a) 0

co a.- aL) m bD a-) 0 ,0CA E a) 0 2 C. ,0 CD)6g 0 a) U, a) ,W a) a) 0 D4- 5e a) 0 V

0 D0 aL) 0 .0 -, r. Co a) cd a) . ri = Cld b1.0, 0 Cd E A rAc- a) %0~l- 0 a) a)) 0 110 o r., 0 a) Cd 0 = I 0 Cd 9 cne E a) 0

.0

0U *4a a0

a) W CD4 caC .W li sCAD C.) z = a) a) u cd c£ C) 1- l 1977 GAFFNEY: CHELIDAE 23

Emydura, but I interpret this as a secondary con- process of the squamosal reaches the parietal and dition). Chelus, on the other hand, is extreme in the latter bone has no lateral component in this its degree of rotation of the medial portions of region. the jugal and postorbital. The medial areas in Chelodina and Hydromedusa also usually have Chelus lie entirely on the surface of the skull, are a quadrate-basisphenoid contact, usually ventral not covered by musculature, and have no demar- to the prootic, but always leaving some of that cation between medial and external surfaces. bone exposed (figs. 4, 5). The contact is well This condition of Chelus I interpret as uniquely developed in Chelodina but limited in Hydro- derived or autapomorphic, as it does not occur in medusa, barely taking place in some instances. any other turtle. My sample of Hydromedusa (three skulls) is too It is interesting to note that Wood (1976) has limited to determine variation of this feature, but recently described a Miocene Chelus, C. colom- I would not be surprised to see the contact ab- bianus, which differs from C. fimbriatus in hav- sent in some specimens. Nonetheless, there are a ing an intergular enclosed by gulars (one indi- number of unique features in common between vidual apparently has an extra set of gulars) Hydromedusa and Chelodina and I hypothesize much as in Chelodina. This suggests the hypoth- that they are a strictly monophyletic group with esis that an enclosed intergular may be primitive respect to other chelids. for Chelodina, Hydromedusa, and Chelus. In any As noted by Boulenger (1889) Chelodina and case, either the enclosed intergular evolved twice, Hydromedusa have four claws on their forefeet or the open intergular evolved twice. rather than five as in all other chelids and pelomedusids. I regard this as a derived feature GROUP 6-INFRATRIBE HYDROMEDUSAD also. The genera Chelodina and Hydromedusa are CLASSIFICATION very similar in skull morphology. They both have relatively long, thin, and flat skulls, although In my opinion, a classification should be a Chelodina novaeguineae (fig. 8) is deeper than redundant reflection of a phylogenetic hypoth- other Chelodina. Both taxa have extremely re- esis. Further discussion of this point of view may duced temporal roof coverings and a markedly be found in Gaffney (1975, In press) and narrow parietal area between the temporal fossae McKenna (1975). Although stability is often con- (figs. 2, 3). The interorbital distance is narrower sidered an important quality of classifications, I than in other chelids and the orbits face dorsally believe that it is often a spurious and misleading to a greater extent than a flattened form such as indication of the attainment of phylogenetic Chelus. The posterolateral process of the parietal, "truth." All of our notions about phylogeny are seen in other chelids (except Pseudemydura, ap- hypotheses that could be wrong; they can never parently in coincidence with other unique fea- be proved correct. If a classification is to have tures of the temporal roof) are absent in Hydro- wide-ranging biologic usefulness, it must be sus- medusa and Chelodina. Chelodina is the only ceptible to change. The classification presented chelid to entirely lack a temporal bar of some here is as unstable as the phylogenetic hypothesis sort, whereas in Hydromedusa an anteromedial presented here. 24 AMERICAN MUSEUM NOVITATES NO. 2620

CLASSIFICATION OF THE CHELID TURTLES Infraorder (Cope, 1868b) Family Pelomedusidae Cope, 1868a Family Chelidael Gray, 1825 Subfamily Pseudemydurinae, new Pseudemydura Subfamily Chelinae Gray, 1825, new rank Infrafamily Emydurodd,2 new Emydura Elseya Infrafamily Chelodd Gray, 1825, new rank Tribe Platemini, new Platemys Tribe Chelini Gray, 1825, new rank Subtribe Hydraspina3 Bonaparte, 1838, new rank Phrynops Subtribe Chelina Gray, 1825, new rank Infratribe Chelad4 Gray, 1825, new rank Chelus Infratribe Hydromedusad, new Chelodina Hydromedusa

'The family name of this group of turtles has been (1967) gave a number of these spellings and a very use- spelled in a number of ways but most commonly Chelyi- ful guide to the higher categories. dae. However, I am here following Williams (1950) and 2Elsewhere (Gaffney, 1972a) I use the arbitrarily Wermuth and Mertens (1961), among others, in the use chosen ending -odd for infrafamily and continue the of Chelidae. As far as I can see, the oldest valid name for practice here. this family is Chelidina Gray, 1825, which becomes 3Bonaparte (1838) originally named a family level Chelidae with the addition of the appropriate ending. taxon, Hydraspidina, based on the genus Hydraspis, a There could be some question as to whether or not name now considered a synonym of Chelodina (see text Chelides Cuvier, 1817, might not be valid, but I am and Stejneger, 1909). However, when Bonaparte used taking the view that it was not explicitly stated as a Hydraspis it referred to what is now Phrynops and, as I family level taxon. understand the rules on family level taxa, the name The spelling variations are primarily due to the fact Hydraspidina must go with Phrynops. that although Chelus Dumeril, 1806, was the first spell- 4As there are no rules or suggestions dealing with ing of this genus, it was later "corrected" to Chelys and tribal level endings, I arbitrarily choose -ad as the ending many family level taxa were based on the latter spelling, for infratribe. which was popular during the 19th century. Kuhn

LITERATURE CITED Bonaparte, C. E. 1838. Amphibiorum Tabula Analytica. Nuovi .Albrecht, P. W. Annali delle Scienze Naturali, vol. 1, 1976. The cranial arteries of turtles and their pp. 391-393. evolutionary significance. Jour. Morph., Bonde, Niels vol. 149, no. 2, pp. 159-182. 1974. [Review of] Interrelationships of Bell, T. fishes. Greenwood, P. H., R. S. Miles 1828. On Hydraspis, a new genus of fresh- and C. Patterson (eds.), Syst. Zool., vol. water , of the family Emydida. 23, pp. 562-569. Zool. Jour., vol. 3, pp. 511 -513. Boulenger, G. A. Blackmore, E. H. 1888. On the Chelyoid chelonians of New 1969. On the Australian Chelidae (Chelonia). Guinea. Annali de Museo Civico de Victorian Nat., vol. 86, no. 10, pp. Storia Naturale Giacomo Doria, second 280-283. ser., vol. 6, pp. 449-452. 1977 GAFFNEY: CHELIDAE 25

1889. Catalogue of the Chelonians, Rhyncho- Cuvier, Georges cephalians and in the British 1817. Le regne distribue d'apres son Museum (Nat. Hist.). New Edition, organisation. tome 2, 1st ed., Paris, London, Printed by of the Trus- Deterville, Libraire, pp. xviii-532. tees, 311 pp. Dumeril, A. M. Constant Bour, Roger 1806. Zoologie analytique, ou methode natur- 1973. Contribution a la connaissance de elle de classification des animaux. Paris, Phrynops nasutus (Schweigger: 1812) Allais, Libraire, 1806, pp. xxxii-336. et Phrynops tuberculatus (Lueder- Dunn, E. R. waldt: 1926). Description d'une nou- 1945. Los Generos de Anfibios y Reptiles de velle sous-espece originaire du Para- , IV. Cuarta y Ultima Parte: guay, Phrynops tuberculatus vander- Reptiles, Ordens Testudineos y Croco- haegei (-Pleurodira-Cheli- dilinos. Caldasia, vol. III, no. 13, pp. dae). Bull. Soc. Zool. France, tome 98, 239-269. no. 1, pp. 175-190, pl. 1. Eldredge, Niles, and Ian Tattersall Bramble, Dennis Marley 1975. Evolutionary models, phylogenetic re- [MS.] Functional morphology, evolution, and construction, and another look at paleoecology of gopher tortoises. Ph.D. hominid phylogeny. In Szalay, F. (ed.), Diss., 1971, Univ. Calif., Berkeley, 341 Approaches to Primate Paleobiology, PP. Contrib. Primatol., vol. 5, pp. 218-242. Brundin, Lars Farris, James S. 1968. Application of phylogenetic principles 1972. Estimating phylogenetic trees from dis- in systematics and evolutionary theory. tance matrices. American Nat., vol. In Orvig, T. (ed.), Current problems of 106, no. 951, pp. 645-668. lower vertebrate phylogeny. Nobel Fitzinger, L. Symposium 4, New York, Interscience 1826. Neue Classification der Reptilien. Wien, Publishers, John Wiley and Sons, pp. J. G. Heubner Verlag, 66 pp. 473-495. Freiberg, Marcos A. 1945. Una nueva especie de tortuga del gen- Burbidge, Andrew A., John A. W. Kirsch, and A. ero "Platemys" Wagler. Physis, Rev. R. Main Soc. Cien. Nat., tomo 20, pp. 1974. Relationships within the Chelidae 19-23. (Testudines: Pleurodira) of Australia 1947. El alotipo de la tortuga "Platemys pal- and New Guinea. Copeia, no. 2, pp. lidipectoris" Freiberg. Ibid., tomo 20, 392-409. pp. 112-115. Cogger, H. Froes, Oscar Miranda 1975. The reptiles and amphibians of Aus- 1957. Notas quelonologicas. I - Atualizacao tralia. A. H. and A. W. Reed, Sydney, da nomenclatura dos quelonios bra- pp. 1-584. sileiros. Iheringia Zool., no. 2, pp. Cope, Edward D. 1-24. 1868a. An examination of the Reptilia and Fry, D. B. Batrachia obtained by the Orton Expe- 1915. Herpetological Notes, Proceedings of dition to Equador and the Upper the Royal Society of Queensland, vol. Amazon, with notes on other species. 27, pp. 61-95. Proc. Acad. Nat. Sci. Philadelphia, vol. Fuchs, Hugo 20, pp. 96-140. 1931. UCber den Unterkiefer und die Unter- 1868b. On the origin of genera. Ibid., vol. 20, kiefernerven (Ramus tertius nervi trige- pp. 242-300. mini et Chorda tympani) der Ar- Cracraft, Joel rauschildkrote (Podocnemis expansa). 1972. The relationships of the higher taxa of Nebst Bemerkungen zur Kiefer- : problems in phylogenetic reason- gelenksfrage. Zeitschr. Anat. Entwick- ing. Condor, vol. 74, no. 4, pp. lungsgeschichte, vol. 94, pp. 206-274. 379-392. Gaffney, Eugene S. 1974. Phylogenetic models and classification. 1972a. The systematics of the North American Syst. Zool., vol. 23, no. 1, pp. 71-90. family (Reptilia, ). 26 AMERICAN MUSEUM NOVITATES NO. 2620

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