A new insight into pycnodontiform fishes Francisco José POYATO-ARIZA Unidad de Paleontología, Departamento de Biología, Universidad Autónoma de Madrid, Cantoblanco, E-28049 Madrid (Spain) [email protected] Sylvie WENZ Laboratoire de Paléontologie, UMR 8569, Muséum national d’Histoire naturelle, 8 rue Buffon, F-75231 Paris cedex 05 (France) [email protected] Poyato-Ariza F. J. & Wenz S. 2002. — A new insight into pycnodontiform fishes. Geodiversitas 24 (1) : 139-248. ABSTRACT The present paper provides a revision of the pycnodontiform fish genera that are based on articulated material. The results of the first cladistic analysis on the interrelationships of the order Pycnodontiformes Berg, 1937 are also pre- sented: it is based on 105 characters for 33 taxa. The monophyly of the order, of the suborder Pycnodontoidei Nursall, 1996, and of the family Pycnodontidae Agassiz, 1833 sensu Nursall 1996b (in large sense) are confirmed; the “suborder Gyrodontoidei” appears, in contrast, as a para- phyletic group. The revision of the historic nomenclatural problems and the hypothesis of phylogenetic relationships show that the genera Eomesodon Woodward, 1918, Coelodus Heckel, 1854, and Palaeobalistum Blainville, 1818, as previously recognized, were not natural assemblages. The new genera Apomesodon n. gen., Ocloedus n. gen., Oropycnodus n. gen., and Abdobalistum n. gen., together with two new species (Apomesodon surgens n. gen., n. sp. and KEY WORDS Abdobalistum thyrsus n. gen., n. sp.), are erected to locate former species of Actinopterygii, Pycnodontiformes, those non-monophyletic genera. The subfamilies Proscinetinae n. rank, revision, Pycnodontinae n. rank, and Nursalliinae n. rank, are also proposed, together cladistics, with the superfamily Pycnodontoidea n. rank, which gathers the sister-groups phylogeny, new genera, Coccodontidae and Pycnodontidae. A completely revised systematic palaeon- new species. tology of the Pycnodontiformes is provided. GEODIVERSITAS • 2002 • 24 (1) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.mnhn.fr/publication/ 139 Poyato-Ariza F. J. & Wenz S. RÉSUMÉ Un nouveau regard sur les poissons pycnodontiformes. Ce travail présente une révision des poissons pycnodontiformes, fondée sur des spécimens en connexion sans tenir compte des dentures ou des dents iso- lées. Il a pour but d’établir les relations phylogénétiques au sein du groupe des Pycnodontiformes Berg, 1937 à l’aide d’une analyse cladistique prenant en compte 33 taxons et 105 caractères. La monophylie de l’ordre, du sous-ordre Pycnodontoidei Nursall, 1996 et de la famille Pycnodontidae Agassiz, 1833 sensu Nursall 1996b (au sens large) a été confirmée, ce qui n’est pas le cas du « sous-ordre Gyrodontoidei ». Les sous-familles Proscinetinae n. rank, Pycnodontinae n. rank et Nursalliinae n. rank ont été proposées ainsi que que la super-famille Pycnodontoidea n. rank, qui réunit les groupes-frères Coccodontidae et Pycnodontidae. Quatre genres (Apomesodon n. gen., Ocloedus n. gen., Oropycnodus n. gen. et Abdobalistum n. gen.) et deux espèces MOTS CLÉS (Apomesodon surgens n. gen., n. sp. et Abdobalistum thyrsus n. gen., n. sp.) ont Actinopterygii, Pycnodontiformes, été créés pour les taxons exclus des genres Eomesodon Woodward, 1918, révision, Coelodus Heckel, 1854 et Palaeobalistum Blainville, 1818, qui apparaissent analyse cladistique, comme non-monophylétiques. Ce travail s’achève par une mise à jour de la phylogénie, nouveaux genres, systématique paléontologique des Pycnodontiformes à la lumière des données nouvelles espèces. nouvelles présentées ici. INTRODUCTION Blainville (1818), Agassiz (1833-43), Pictet (1854), Heckel (1856), Wagner (1862), One of the most characteristic and popular fossils Woodward (1895a), and Hennig (1906). of many preservational Lagerstätten are the pyc- The order Pycnodontiformes was erected by Berg nodontiform fishes. They have been found in in 1937 (p. 1278) by giving the name only. Later Triassic to Eocene strata all over the world. In on, a small diagnosis and a familiar arrangement general lines, a pycnodontiform is easily recogniz- are given by the same author in 1940 (p. 208- able by its laterally compressed body, resulting in 210, 413, 414 in the English translation of a more or less rounded outline, and by its well 1947). A most exhaustive account, with all gen- developed durophagous dentition, so that these era known at the time, including their geographic fishes have always called the attention of both and temporal distribution, was presented by amateurs and professional palaeontologists. Lehman (1966: 170-181). Some papers have Pycnodontiforms have been known for a long extensively dealt with certain pyncodontiforms in time. The oldest plate that we have seen is Knorr the last decades (e.g., Blot 1987; Lambers 1991), (1768: pls 1755), containing a long philosophic- but the first recent look at the ensemble of these al dissertation on the nature of this “petrified ani- fishes was not given until Nursall (1996a on their mal”. The first descriptions with scientific palaeoecology and distribution, 1996b on a characters on what would later be pycnodont detailed phylogenetic hypothesis, 1999a on the fishes are about two centuries old (e.g., Volta Mesturidae, 1999b on the pycnodontiform 1809, usually cited as 1796, but a note on the last bauplan). page of the book explains that it was not pub- The aims of the present paper are to update their lished until 1809, which should be the valid nomenclatural problems, and to accomplish the date). Pycnodonts are a relevant part of many key first cladistic analysis on the interrelatiohships of classic palaeoichthyological works, such as the order Pycnodontiformes. The phylogenetic 140 GEODIVERSITAS • 2002 • 24 (1) A new insight into pycnodontiform fishes hypothesis by Nursall (1996b) is based in a dis- MGSB Museu Geològic del Seminari cussion of the distribution of his proposed char- Conciliar de Barcelona; ML Muséum d’Histoire naturelle, Lyon; acters, rather than on a cladistic analysis sensu MNHN Muséum national d’Histoire stricto. As acknowledged by this author, his naturelle, Paris; HDJ, HAK, MSE, hypothesized relationships “should provide direc- SLN, BCE, CRN, CNJ: abbreviations tions for further analysis” (Nursall 1999b: 149). of localities in this collection. MNHUB Museum für Naturkunde, der Facing works on certain pycnodontiforms from Humboldt Universität, Berlin; Spain and the Lebanon, both new and revised NHML Natural History Museum, London; forms (Kriwet et al. 1999; Poyato-Ariza & Wenz NMW Naturhistorisches Museum in Wien; 1995, 2000, work in progress), we found that UAM Universidad Autónoma de Madrid. they do not fit neatly into Nursall’s phylogenetic hypothesis. This is due not only to the relatively high number of his incertae sedis forms, but also METHODS to incongruences in the distribution and in the polarity of the characters used by him when Our characters are based, on the first place, on applied to the forms involved in our studies. We those presented by Nursall (1996b, re-taken in therefore realized that we needed to perform a 1999b). We have not used those characters cladistic analysis if we wanted to present coherent regarding the phylogenetic relationships of the phylogenetic hypotheses for the taxa that we are Pycnodontiformes with other high taxa (1996b: describing and revising. Both the cladistic analy- characters 72-93), because the present paper does sis and its taxonomic implications, including a not address this issue. Regarding Nursall’s sets of new arrangement of the Pycnodontiformes, with characters, many of them actually turn out to be several new taxa, are presented herein. different states of a same given character. For example, characters 25 (“The prehensile teeth INSTITUTIONAL ABBREVIATIONS are styliform”; Nursall 1996b: 133) and 40 ADR private collection of Mr. Armando (“Incisiform prehensile teeth”; Nursall 1996b: Díaz-Romeral, Cuenca; BMM Burgmeister Müller Museum, 135) are actually different states of a single char- Solnhofen; acter on the morphology of the prehensile teeth. DGM-DNPM Divisão de Geologia e Mineralogia, We therefore gathered all the different states into Departamento Nacional da produção discrete characters. Some of the characters, Mineral, Rio de Janeiro; FSL Faculté des Sciences de Lyon; notably those concerning the contour scales and IEI Institut d’Estudis Ilerdencs, Lleida; the cloaca, were substantially developed into IGM Instituto de geología, Universidad additional characters, and we added some charac- Nacional Autónoma de México ters of our own as well. The result is the character (Departamento de Paleontología), México D. F.; data set discussed below and listed on JM Jura Museum, Eichstätt; Appendix 1. The correspondence of our charac- IRSNB Institut royal des Sciences naturelles ters with those of Nursall (1996b) is noted in the de Belgique, Brussels; section below for each case wherever pertinent. MB Museum Bergér, Harthof; MCCM Museo de las Ciencias de Castilla-La The characters were polarised by using the Mancha, Cuenca, Spain. Provisionally compound outgroup methodology by Maddison housed at the UAM for study; et al. (1984). The outgroup was formed by MCSNB Museo Civico di Scienze Naturale “E. Caffi” di Bergamo; Pteronisculus (Nielsen 1942; Lehman 1952); MCSNM Museo Civico di Storia Naturale di Semionotus (Olsen & McCune 1991); Dapedium Milano; (Wenz 1968; Thies 1988; Thies & Herzog MCSNV Museo Civico di Storia Naturale di 1999); Amia (Allis 1897, 1898; Grande 1996;