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On the Phylogenetic Arrangement of the Sauropsida

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On the Phylogenetic Arrangement of the Sauropsida ON THE PHYLOGENETIC ARRANGEMENT OF THE SAUROPSIDA. DR. G. BAUR, YALECOLLEGE MUSEUM, NEW HAVEN,Cam. Cope' has given the following classification of the Reptilia :- A. Extremities not differentiated in form beyond proximal segment. I. 0s quadratum immovably articulated to squamosal, etc. Tubercular and capitular rib-articulations present and distinct. I. Ichthyopterygia. A.A. Elements of extremities differentiated. 11. 0s quadratum immovably articulated, capitular and tubercular rib-articulations distinct. ARCHOSAURIA. Pubis and ischium united, and with little or no obturater foramen ; one posterior cranial arch ; limbs ambulatory ; a procoracoid. 2. Theromovpha. Ischium and pubis distinct, the latter directed forwards, back- wards, or downwards ; two posterior cranial arches ; limbs am- bulatory ; no procoracoid. 3. Binosauria (+Crocodilia) . Ischium and pubis united ; two postcranial arches ; anterior limbs volant. 4. Omithosauria. 111. 0s quadratum closely united to cranial arches ; but one rib-articulation. SYNAPTOSAURIA. Distinct hyposternal and postabdominal bones ; ribs joining each two vertebrae, and generally forming a carapace ; one pos- terior cranial arch. 5. Testudinata. Hyposternal and postabdominal bones not distinct ; two pos- terior cranial arches ; ribs attached to one vertebra ; a sternum ; ? no procoracoid. 6. Rhynckocephalia. Cape, E. D. On the evolution of the vertebrata, progressive and retrogressive. Amer. Naturalist, March, 1885, pp. 245-246. 94 BAUR. POL. I. Hyposternal and postabdominal bones not distinct ; two pos- terior cranial arches; ribs attached to one centrum; no ster- num ; a procoracoid. 7. Sauropterygia. IV. 0s quadratum attached only at the proximal extremity, and more or less immovable; ribs with one head. STREPTO- STYLICA. Brain case membranous in front of proijtic bone; trabecula not persistent. 8. Lacevtilia. Brain case with osseous walls anterior to prootic ; a scapular arch and sternum. 9. Pythonomorpka. Brain case with osseous walls anterior to proijtic ; no scapular arch nor sternum ; trabecular grooves of sphenoid and pre- sphenoid bones. 10. Ophidia. The phylogeny of the Reptilia is expressed by Cope in the following way :- Dinosauria Testudinata. Rhynchocephalia. Lacertilia. opbidia. (Crocodilia) . \ I / I \IPythonomorpha. Pterosauria. \ \ I / Theromorpha. No. I.] ARRANGEMENT OF THE SAUROPSfDA. 95 CRITICISM OF THIS CLASSIFICATION. Ichthyopterygia. The Ichthyopterygia are separated by Cope from the rest of the Reptilia because of the non-differentiation of the extremities in form beyond proximal segment. That the Ichthyopterygia stand in the same relation to one or the other group of Reptiles as the Cetaceans do to the Ungulata and Carnivora seems to me unquestionable. Prof. Haeckel expressed this idea twenty years ago in his celebrated work: ‘I Generelle Morphologie der Organismen.” Vol. I., pp. I 84- 185. Berlin, 1866. Of the Halisauria (= Enaliosauria = Ichthyopterygia + Sau- ropterygia), he says, - Ihre vielfachen Ahnlichkeiten mit den Fischen und insbeson- dere mit den Ganoiden haben zu der Annahme gefiihrt, dass sie diesen niher als den iibrigen Reptilien venvandt seien, und man hat selbst neuerdings versucht, die Ganoiden, Ganocephalen, Labyrinthodonten, Ichthyosaurier und Sauropterygier als fort- laufende Glieder einer einzigen Entwicklungsreihe darzustellen. Indessen ist es viel wahrscheinlicher, hss diese k’knZichkeiten mehr Anpassungs ahnlichkeiten sind, und dassdie Halisaurier sich zu den iibrigen Reptilien verhalten, wie die Cetaceen zu den Saugethieren.” In 1881 Prof. C. Vogt’ expressed the same opinion. Quite lately I have brought forward new proofs for the cor- rectness of this view? Therefore it is impossible to separate the Ichthyopterygia from the rest of the Reptiles ; it would be the same as to sepa- rate the Cetaceans from the Mammals. Weber3 has shown, in a very important memoir, that the Cetaceans descended from a group of Mammals, probably ancestral to both the Ungulata and the Carnivora. ’Revue scientifique. 12. Mlirz, 1881; Kosmos, Vol. 9, 1881,pp. 318-319. Baur, G. Bemerkungen iiber SIuropterygia und Ichthyopterygia. Zool. Anz., NO. 221. 1886. 3Weber, Max. Studien iiber Singethiere. Ein Beitrag ZUT Frage nach dem Ursprnng da Cetaceen. Jena, 1886. 96 BAUR. [VOL. I. The question now is, from which group of Reptiles descended the Ichthyopterygia ? The skull shows characters of the Rhynchocephalia, the oldest Crocodiles (Belodon) and the Dinosaurs; but it is still more generalized than in these groups. This is proved espe- cially by two bones, - the opisthotic and the supratemporal. The opisthotic is separate as in the Testudinata.’ Huxley a speaks of a flattened bone between the postorbital, postfrontal, and the squamosal; this bone (temporal, Cuvier - prosquamosal, Owen), according to Huxley, does not appear to have any precise homologue among other Reptilia. I shall show, in another paper, that this peculiar bone is nothing else than the supratemporal I’ of the Lacertilia, and the ‘I squamosal ” of the Stegocephali ; that the (( squamosal ” of the Stegocephali is really the supatemporaZ, the supra- temporal ” of the Stegocephali, the squnmosal, of the Reptilia. The Ichthyopterygia, therefore, are the only Reptiles, so far as now known, which have a supratemporal, like that of the old Stegocephali. Another character common to the Rhynchocephalia, a few Lizards and Dinosauria, the oldest Crocodilia (Belodon), and Sa~ropterygin,~is the presence of the postorbital and the post- frontal in a separate condition. The scapular arch of The Ichthyopterygia is Lacertilian or Rhynchocephalian. The ribs are two-headed, like those of the Crocodilia, Dinosauria, etc. They are different from the ribs of all other known Reptiles, because they are never connected with the neurapophyses; they never leave the body of the vertebra (according to Owen’s figures). Abdominal ribs are developed, as in the Rhynchocephalia and Sauropterygia. Therefore we have combined characters of the Rhynchoce- phalia, the oldest Crocodilia and Sauropterygia. To-day we do not know a group of Reptiles showing such characters; 1 Cope, E. 0. On the homologies of some of the cranial bones of Reptilia, and on the systematic arrangement of the class. Proc. Amer. Assoc. Adv. Sc., Vol. XIX., p. 199. 1871. ”uley, T. H. A manual of the anatomy of vertebrsted animals. London, 1871, p. 246. 3 In Sirnorourus these two bones have already united. No. 1.3 ARRANGEMENT OF THE SAUROPSIDA. 97 but that the Ichthyopterygia must have taken their origin from such a one I have no doubt.' The Sauropterygia and Testudinata. This group of Reptiles is classified by Cope with the Synapto- sauyia, which have but one rib articulation. This is not correct; the oldest Sauropterygia, the Lariosauridae and Nothosauridae, and even Plesiosaums, have two-headed ribs in the cervicals and the former families even in the anterior dorsals. The posterior dorsals have the capitulum and tuberculum united, and therefore are morphologically two-headed. It always seemed difficult to determine the systematic position of the Sauropterygia. Huxley considered this group allied to the Crocodilia, especially to the Teleosauride ; Owen cbmpared it with the Testudinata. That they descended from land-living reptiles is certain? The Sauropterygia begin in the Triassic with the Lario- sauridae and Nothosauridae. They are in no direct relation to the Ichthyopterygia. The skull and shoulder-girdle are entirely different in both. The skull is very characteristic. It resembles the Rhyncho- cephalia and old Crocodilia (Belodon, Teleosaurus) ; the parietal foramen is present. The postorbital is free or united with the postfrontal. The whole shape (at least of some forms) resembles very much the skull of the Crocodilia; but one character shows at once the specialization of the Sauroptery- gian skull; it is the absence of the lower temporal arch, as in the Lacertilia; a quadratojugal seems never to be developed. Another resemblance to the Rhynchocephalia consists in the structure of the abdominal ribs. If we compare these elements in the Lariosauridae and Nothosauridae with those in Sphenodon we find exactly the same condition. In Lariosaurus and Sphenodon they are entirely identical, as the figures of Deecke prove.3 'The Baptanodontia (Sauranodontia), Marsh, are specialized forms of the Ichthy- opterygia, like the Mystacoceti among the Cetacea, and the Pteranodontidse among the Ornithosauria. 'Barn, G. Bemerkungen iiber Sauroptetlrgia und IchthpptetYgia. Zool. Am., No. 221, 1886. a Deecke, W. uber Lariosaurus und einige andere Saurier der 1ombardischenTrias. Zcitschr. Deutsch. Geol. Ges. Bd. 38. P1. 1x1. Fig. I. BAUR. POL. I. There seem to be the larger number of connections with the Testudinata. The cervical ribs of the Testudinata are entirely rudimentary. Hoffmann ' has shown that they are developed in the embryos. In the adult Testudinata there are often well-developed dia- pophyses and parapophyses in the anterior cervicals. The former are connected with the neurapophyses, the latter with the body of the vertebra; both touch each other. Therefore we have a real para-diapophysis, and, consequently, if a rib should be developed, a rib with a capitulo-tuberculum. That the an- cestors of the Testudinata had well-developed ribs on the cervicals is shown, not only by Hoffmann's researches, but also by the still developed para-diapophyses. The pelvic arch of the Nothosauridz' is only comparable with that of the Testudinata. The shape of the pubis is very
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