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Aspects of the Functional Morphology in the Cranial and Cervical Skeleton of the Sabre-Toothed Cat Paramachairodus Ogygia (Kaup, 1832) (Felidae

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Blackwell Science, LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082The

Linnean Society of London, 2005? 2005

1443 363377 Original Article

FUNCTIONAL MORPHOLOGY OF P. OGYGIAM. J. SALESA ET AL.

Zoological Journal of the Linnean Society, 2005, 144, 363–377. With 11 figures

Aspects of the functional morphology in the cranial and cervical skeleton of the sabre-toothed cat

Paramachairodus ogygia (Kaup, 1832) (Felidae,

Machairodontinae) from the Late Miocene of Spain: implications for the origins of the machairodont killing bite

MANUEL J. SALESA1*, MAURICIO ANTÓN2, ALAN TURNER1 and JORGE MORALES2

1School of Biological & Earth Sciences, Byrom Street, Liverpool John Moores University, Liverpool, L3 3A F , U K 2Departamento de Palaeobiología, Museo Nacional de Ciencias Naturales-CSIC, José Gutiérrez Abascal, 2. 28006 Madrid, Spain

Received January 2004; accepted for publication March 2005

The skull and cervical anatomy of the sabre-toothed felid Paramachairodus ogygia (Kaup, 1832) is described in this paper, with special attention paid to its functional morphology. Because of the scarcity of fossil remains, the anatomy of this felid has been very poorly known. However, the recently discovered Miocene carnivore trap of Batallones-1, near Madrid, Spain, has yielded almost complete skeletons of this animal, which is now one of the best known machairodontines. Consequently, the machairodont adaptations of this primitive sabre-toothed felid can be assessed for the first time. Some characters, such as the morphology of the mastoid area, reveal an intermediate state between that of felines and machairodontines, while others, such as the flattened upper canines and verticalized mandibular symphysis, show clear machairodont affinities. © 2005 The Linnean Society of London, Zoological Journal of the

Linnean Society, 2005, 144, 363-377.

ADDITIONAL KEYWORDS: Batallones – Carnivora – Felinae – functional anatomy – Mammalia – Miocene – pantherine – Turolian – Vallesian.

straint in each group, still remain essentially a mystery.

INTRODUCTION

Sabre-toothed predators have evolved several times among different orders of mammals and in somewhat different forms even among nonmammalian synapsids (Turner & Antón, 1997). The similarities in detail between often completely unrelated taxa are so remarkable that the sabre-tooth adaptation has become a text-book example of convergent evolution. However, exactly how it evolved in each case, what the main evolutionary pressures were and where the balance lay between adaptation and phylogenetic con-
One of the reasons for this lack of resolution is the fact that within each convergent group of sabre-tooth predators it is usually the most derived, crown taxa that are the best known anatomically while the basal taxa have much poorer fossil records. Thus, among the Machairodontinae (the sabre-toothed subfamily within the extant family Felidae), the crown taxa such

as Homotherium and Smilodon from the Pliocene and

Pleistocene have been known for many decades thanks to complete skulls and skeletons, while the fossil record of basal genera such as Paramachairodus from the Late Miocene have traditionally consisted of scarce and fragmentary material. The situation is not

*Corresponding author. E-mail: [email protected] © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 363–377

363

364 M. J. SALESA ET AL.

very different across the various families of mammalian sabre-tooths, including the Nimravidae and Barbourofelidae among the Carnivora (Peigné, 2002; Morlo, Peigné & Nagel, 2004) and the marsupial Thylacosmilidae (Argot, 2003, 2004).
Consequently, no studies of the functional morphology of Paramachairodus have yet been published. The discovery of Batallones-1, a new Late Miocene fossil locality near Madrid, Spain, has yielded a large

number of specimens of P . o gygia, with at least 24

individuals represented and including several skull and mandibles (Salesa, 2002). This locality has been interpreted as a carnivore trap based on its special characteristics, such as the presence of an extremely high percentage of carnivores (98%) and the morphology of the site, essentially a hole with semivertical walls (Antón & Morales, 2000; Morales et al., 2000). The carnivore guild of Batallones-1 also includes the lion-sized sabre-toothed felid Machairodus aphanis-

tus, the bear-dog Amphicyon sp. aff. A. castellanus, the primitive hyaenid Protictitherium crassum and other

carnivores such as mustelids and Simocyon batalleri, a medium-sized carnivore related to the extant red panda (Antón & Morales, 2000; Morales et al., 2000; Salesa & Fraile, 2000; Salesa, 2002; Antón et al., 2004a; Peigné et al., 2005).
The result of this unbalanced fossil record is that emphasis on a series of highly convergent crown taxa has led to an exaggerated perception of homogeneity in adaptation and to suggestions that the so-called ‘sabre-tooth complex’ would be under ‘strong pleiotropic control’ (Dawson et al., 1986). A better fossil record and a detailed study of basal taxa would greatly help to clarify the origins of this predatory adaptation, in particular of the ‘canine shear-bite’, the derived killing bite modality that has been proposed as a functional explanation of the morphology of crown sabre-toothed

taxa such as Smilodon and Homotherium (Akersten,

1985; Antón & Galobart, 1999) The genus Paramachairodus Pilgrim, 1913 includes primitive, leopard-sized sabre-toothed cats known from the Late Miocene faunas of Eurasia (Beaumont, 1975; Morales & Soria, 1977; Montoya, 1994; Morlo, 1997; Salesa et al., 2003). Two species have been tra-

ditionally referred to this genus: P . o gygia  (Kaup,

1832), of Vallesian-Early Turolian age (MN 9–11 of

Mein, 1975) and P . o rientalis (Kittl, 1887) of Turolian

age (MN 11–13), distinguished by several dental traits. The more primitive species, P . o gygia, had noncrenulated upper canines, a P4 without ectostyle and with a strong protocone, and a P3 with a posterointer-

nal expansion. P . o rientalis  had crenulated upper

canines, a P4 with a well marked ectostyle as well as a reduced, backwardly displaced protocone and a P3 reduced in size and without posterointernal expansion (Salesa et al., 2003).

In 1924, Zdansky created the species P . m aximiliani

for a damaged skull from the Late Miocene of China. The differences between this third species and

P. o rientalis  are largely confined to the more curved upper canines of P . m aximiliani, where there are

crenulations on both keels (Zdansky, 1924). These characters are now considered to be of little systematic value, because crenulations are not equally evident in

both keels. Thus for most authors, P . m aximiliani is a synonym of P . o rientalis  (Pilgrim, 1931; Beaumont,

1978; Salesa, 2002; Salesa et al., 2003). The anatomy of Paramachairodus has been largely unknown, because it is present in only a few fossil localities such as Pikermi (Greece, MN 12), Maragha (Iran, MN 11) or Eppelsheim (Germany, MN 9) and represented by relatively scarce material. Most finds have been of dentitions. While it has long been clear that Paramachairodus was a sabre-toothed cat, the absence of more extensive anatomical information has meant that its machairodont adaptations and wider aspects of its palaeobiology have remained unknown.
In this paper we present a functional analysis of the cranial, mandibular and cervical anatomy of P . o gygia, concentrating on aspects directly related to the canine shear-bite adaptation. This analysis has revealed an intermediate morphology between the extant felines and the more derived sabre-toothed cats such as

Smilodon or Homotherium.

MATERIAL AND METHODS

Functional study of the cranial and cervical anatomy

of Paramachairodus ogygia has been possible because

of the great number of newly available fossils of this species: a total of 16 skulls, 12 mandibles and several vertebrae belonging to the Batallones-1 assemblage. All the material is housed at the Museo Nacional de Ciencias Naturales-CSIC in Madrid, Spain. Comparisons with other Felidae have been made using the

extant felines Panthera leo, Panthera tigris, Panthera pardus, Panthera onca, Uncia uncia, Neofelis nebu- losa, Caracal caracal, Lynx pardina and Puma con-

color, belonging to the collections of the Museo Anatómico de la Universidad de Valladolid and Museo Nacional de Ciencias Naturales (Madrid). We also used published information on modern felid anatomy (Barone, 2000; Reinhard & Jennings, 1935; Sisson & Grossman, 1962) and on the morphology of other, more derived sabre-toothed cats such as Smilo-

don fatalis and Homotherium latidens. The latter taxa

were chosen as the ideal reference for comparison with

P. o gygia  because they exemplify the most derived

state for the sabre-toothed adaptations within the

Machairodontinae, whilst P . o gygia  occupies a near

basal position in the same subfamily. The degree of machairodont adaptations appears to be essentially

© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 363–377

FUNCTIONAL MORPHOLOGY OF P . O GYGIA  365

independent of such body size differences as those

observed between P . o gygia and the referred taxa.

Differences in the height of the crown of the upper

canines between P . o gygia  and reference taxa were

assessed using an ANOVA on the index between crown height and basal skull length. All the measured upper canines were unworn specimens. Differences in relative elongation of the neck were assessed by calculating an index between the corpus length of the atlas and the third cervical vertebra. The lack of associated

C3 and atlas of P . o gygia in Batallones-1 was overcome

by taking an average measurement between several isolated C3 and atlas vertebrae and comparing the resultant index with similar indices of reference taxa using a Student’s t-test. These results provided the most reliable indication of elongation of the neck in the absence of complete, associated cervical series of

P. o gygia.

In the more derived members of this group, the paraoccipital process becomes very reduced, almost vestigial, and the mastoid process is hyper-developed (Emerson & Radinsky, 1980), overlapping the auditory bulla in its growth in some genera such as Smilodon

(Fig. 1). Thus, the mastoid region of P . o gygia  can be

considered primitive for the Machairodontinae, but it is clearly within the machairodont lineage because its morphology is derived in relation to the earliest felids

of the genera Proailurus and Pseudaelurus. There are

few known basicranial remains of these genera, but in the Proailurus-like skull from Ginn Quarry (Hunt, 1998: fig. 19A), and in the holotype of Pseudaelurus validus (Rothwell, 2001: fig. 3), the paroccipital process can be seen to be well developed ventrally, surpassing the level of the mastoid process; this condition is also found in fossil and extant species of the Felinae. The mastoid process is the attachment area for some important muscles directly involved in headdepression movements, such as the brachiocephalicus (which has its origin on the humerus), part of the fibres of the sternocephalicus (extending from the

manubrium of the sternum) and the obliquus capitis

anterior (with its origin on the ventrolateral surface of the atlas wings) (Reinhard & Jennings, 1935; Barone, 2000). The inferred position of these attachments on

the cranium of P . o gygia is shown in Figure 2.

The anteroventral displacement of these attachment areas in the Machairodontinae in relation to the Felinae caused by the modification of the mastoid process has an important consequence. The distance between the muscular attachment areas and the atlanto-cranial articulation enlarges, resulting in an increase in the length of the lever arm of the flexor muscles of the head (Antón & Galobart, 1999) and thus producing a more powerful contraction. Moreover, there is another interesting implication of this displacement of muscle attachment areas. In the primitive model, exemplified by the felines, the main

function of the obliquus capitis anterior is the exten-

sion of the head on the atlas (Sisson & Grossman, 1962; Barone, 2000). This is because a great number of the fibres of this muscle are disposed above the point of rotation of the head over the atlas (Fig. 3A) and thus their contraction produces the extension of the head. In the sabre-toothed cats, most of the fibres of the

obliquus capitis anterior are below the point of rota-

tion of the head (Fig. 3B) because of the anteroventral displacement of the mastoid process. The function of this muscle is thus ventral flexion of the head, a motion directly involved in the canine shear-bite (Antón et al., 2004b), as discussed further in the final section.

FUNCTIONAL MORPHOLOGY OF THE SKULL
AND MANDIBLE

GENERAL CRANIAL MORPHOLOGY

The skull of P . o gygia  shows an overall morphology

similar to that of a pantherine cat (Figs 1, 6, 11), although there are some differences. The shape of the nasals is neither completely rectangular, as in Smilo- don or Homotherium, nor triangular, as in most of the pantherines, but shows an intermediate morphology. All nuchal, lambdoid and sagittal crests are well developed, and the join between sagittal and frontal crests is situated at the level of the postorbital processes of the frontal bone, as in pantherines and

felines in general. The zygomatic arch of P . o gygia is

broadly similar to that of a pantherine, but with a postorbital process that is almost vestigial, unlike the pantherines in which this process is large and pointed. The tympanic bullae of P . o gygia are similar to those of the pantherines. They are rounded, but less inflated than in the latter, and extend from the anterior margin of the mastoid process to the posterior margin of the postglenoid process.

MASTOID AREA

The morphology of the mastoid region in P . o gygia

shows an intermediate condition between that of the more derived Pleistocene sabre-toothed cats, such as

Smilodon or Homotherium, and that of the fossil and extant Felinae. The paraoccipital process of P . o gygia

is relatively smaller than in the latter, whereas the mastoid process has a similar size to that in the felines, although it is displaced in an anteroventral direction (Fig. 1). This morphology is plesiomorphic for the sabre-toothed cats.
The reduction of the paroccipital process in the sabre-toothed cats, including P . o gygia, can be related to the need to increase the gape of the mandible

© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 363–377

366 M. J. SALESA ET AL.

p.p. p.p. p.p. m.p.

m.p. m.p.

  • A
  • B
  • C

5 cm
D

Figure 1. Left mastoid morphology of some species of Felidae showing the different development of the mastoid process (m.p.) and paraoccipital process (p.p.). A, Panthera leo. B, Paramachairodus ogygia from Batallones-1, B-1377. C, Smilodon fatalis from Rancho La Brea. D, B-1377, skull of P . o gygia from Batallones-1 in left lateral view with mastoid area circled.

because of the enlargement of the upper canines. The function of the digastricus muscle, whose insertion areas are the paroccipital process and the ventral flange of the dentary, is to open the mandible (Sisson & Grossman, 1962; Barone, 2000). If the ventral projection and size of the paramastoid process is reduced, which is the condition shown by sabre-toothed felids, the attachment of this muscle is displaced dorsally, increasing fibre lengths and thus allowing efficient contraction at larger gapes (Emerson & Radinsky, 1980; Antón & Galobart, 1999).

The morphology of the mastoid region in P . o gygia

shows an early stage of the skull modifications that will reach their maximum development in the later

Machairodontinae, such as Smilodon or Homothe- rium. The remodelling of this area in P . o gygia  pro-

duces an increase in the force generated by the flexor muscles of the head and an increase in the maximum gape of the mandible. These changes suggest that the development of the canine shear-bite, the machairodont type of killing bite, had already started in this Late Miocene species (Salesa, 2002). This kind of

© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 363–377

FUNCTIONAL MORPHOLOGY OF P . O GYGIA  367

A

Ob. Cap. Post.

brachiocephalicus

Br.
Ob. Cap. Ant.

rectus capitis lateralis

digastricus sterno-

Ob. Cap. Post.

cephalicus obliquus capitis

anterior

B

Figure 2. Inferred areas of muscular attachment on the

skull of Paramachairodus ogygia.

Ob. Cap. Ant.
Br.

attack consisted of a well-aimed bite to the throat of the prey, which damaged its blood vessels and trachea and resulted in almost instantaneous death. This is in contrast with the feline killing method where the prey has to be suffocated with a bite in the throat lasting several minutes (Turner & Antón, 1997). These two different hunting methods explain the significant differences in the cranial and cervical morphology of the Felinae and Machairodontinae, and they are already present in one of the most primitive

machairodonts, P . o gygia  (Martin, 1980; Biknevicius

& van Valkenburgh, 1996; Turner & Antón, 1997; Antón & Galobart, 1999). This kind of attack was probably developed in parallel by other ‘sabre-toothed’ carnivores, such as the Barbourofelines and Thylacosmilinae, and their mastoid anatomy shows some analogies with that of the true sabre-toothed felids (Turner & Antón, 1997; Morales et al., 2001).

Figure 3. Anatomical disposition of the muscles brachio-

cephalicus (Br.), obliquus capitis anterior (Ob. Cap. Ant.) and obliquus capitis posterior (Ob. Cap. Post.) in Felinae and Machairodontinae. A, Panthera leo. B, Homotherium

latidens (artwork by M. Antón).

correlated with the existence of a high vertical stress in that zone during the canine shear-bite, due in part to the use of the mandible as an anchor. The curved mandibular symphysis of felines does not suffer such stress during the attack, because this is achieved with the maxilla and mandible biting at the same time (Biknevicius, van Valkenburgh & Walker, 1996). Another possible interpretation, compatible with the one outlined above, is that an increase in the vertical height of the symphysis is an efficient way to counter symphyseal bending due to axial twisting of the mandibular corpora, an argument that has been put forward to explain the vertically high symphyses of sabre-toothed therapsids (Jenkins, Thomasson & Norman, 2002). It is also possible that the verticalized mandibular

symphysis of P . o gygia  and other sabre-toothed cats

can be explained as a consequence of reorganization in the alveolar zone of the lower canines. Because of the smaller palate width of the sabre-toothed cats relative to the felines, the lower canines had to become verticalized to allow occlusion. Nevertheless, it is probable that both mechanisms acted together, creating the specialized symphysis model of the Machairodontinae. Some sabre-toothed cat taxa, such as Homotherium and Megantereon, developed a mandibular flange, a kind of ventral projection in the symphysis that could

MANDIBULAR SYMPHYSIS

The mandibular symphysis of P . o gygia  is strongly

verticalized, forming an almost square angle with the ventral flange of the mandibular corpus (Fig. 4B). Owing to this verticalization, its anterior surface is flat and describes a distinctive plane. This morphology, typical of the sabre-toothed cats (Fig. 4C), is clearly different from the feline model, in which the anteroventral surface of the mandible is gently curved upwards (Fig. 4A). The morphology of the mandibular

symphysis of P . o gygia  can be related to the canine

shear-bite, in which the mandible acts as an anchor while the head is flexed downwards, sinking the upper canines into the flesh of prey. This morphological change in the mandibular symphysis of P . o gygia relative to the feline model can be

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    Hyaenodontidae (Creodonta, Mammalia) and the Position of Systematics in Evolutionary Biology by Paul David Polly B.A. (University of Texas at Austin) 1987 A dissertation submitted in partial satisfaction of the requirements for the degree of Doctor of Philosophy in Paleontology in the GRADUATE DIVISION of the UNIVERSITY of CALIFORNIA at BERKELEY Committee in charge: Professor William A. Clemens, Chair Professor Kevin Padian Professor James L. Patton Professor F. Clark Howell 1993 Hyaenodontidae (Creodonta, Mammalia) and the Position of Systematics in Evolutionary Biology © 1993 by Paul David Polly To P. Reid Hamilton, in memory. iii TABLE OF CONTENTS Introduction ix Acknowledgments xi Chapter One--Revolution and Evolution in Taxonomy: Mammalian Classification Before and After Darwin 1 Introduction 2 The Beginning of Modern Taxonomy: Linnaeus and his Predecessors 5 Cuvier's Classification 10 Owen's Classification 18 Post-Darwinian Taxonomy: Revolution and Evolution in Classification 24 Kovalevskii's Classification 25 Huxley's Classification 28 Cope's Classification 33 Early 20th Century Taxonomy 42 Simpson and the Evolutionary Synthesis 46 A Box Model of Classification 48 The Content of Simpson's 1945 Classification 50 Conclusion 52 Acknowledgments 56 Bibliography 56 Figures 69 Chapter Two: Hyaenodontidae (Creodonta, Mammalia) from the Early Eocene Four Mile Fauna and Their Biostratigraphic Implications 78 Abstract 79 Introduction 79 Materials and Methods 80 iv Systematic Paleontology 80 The Four Mile Fauna and Wasatchian Biostratigraphic Zonation 84 Conclusion 86 Acknowledgments 86 Bibliography 86 Figures 87 Chapter Three: A New Genus Eurotherium (Creodonta, Mammalia) in Reference to Taxonomic Problems with Some Eocene Hyaenodontids from Eurasia (With B. Lange-Badré) 89 Résumé 90 Abstract 90 Version française abrégéé 90 Introduction 93 Acknowledgments 96 Bibliography 96 Table 3.1: Original and Current Usages of Genera and Species 99 Table 3.2: Species Currently Included in Genera Discussed in Text 101 Chapter Four: The skeleton of Gazinocyon vulpeculus n.
  • 7 X 11 Long.P65

    7 X 11 Long.P65

    Cambridge University Press 978-0-521-73586-5 - Carnivoran Evolution: New Views on Phylogeny, Form, and Function Edited by Anjali Goswami and Anthony Friscia Index More information Index Page numbers in bold type refers to figures. Acinonyx jubatus, 226, 304 Arctoidea, 12, 27, 30, 31, 40, 94, 104 Actiocyon, 115, 125 Artiodactyla, 312 active replacement, 311 Atilix paludinosis, 9 Adelphailurus, 411 Ailuridae, 11, 30, 92–126, 231, 304 badgers, see Mustelidae diagnosis, 116 bamboo, 12, 94 Ailurinae, 92, 116–20 Barbourofelinae, 10, 34, 295, 301 Ailuropoda, 15, 20, 31, 94, 104, 107, 226 basicranium, 37, 65, 107, 155, 157, 158, 159 Ailurus, 117, 126 Bassaricyon, 11 Ailurus fulgens, 3, 30, 93, 126 Bassaricyon lasius, 377 alisphenoid canal, 82, 107, 110 Bassariscus, 231, 238 allometry, 43, 165, 168 Bassariscus astutus, 388 multiphasic, 172–6, 178–85 Bathygale, 32 postcranial, 411–59 bear-dogs, see Amphicyonidae Alopecocyon, 112, 113, 115, 122, 125 behaviour, 411, 454 Alopex lagopus, 392 Bergmann’s Rule, 396 American lion, see Panthera leo cf. atrox biogeography, 225–39, 247–65, 361 Amphictis, 30, 92, 109, 112, 115, 116, biomechanics 123–124 cranial, 466–81 Amphicynodon, 109 postcranial, 450–9 Amphicynopsis, 297 bite force, 466–81 Amphicyon major, 299 body size, 39–43, 226, 248, 249, 269, Amphicyonidae, 17, 34, 41, 142, 193, 295 270, 301, 314, 325, 330, 412, 413 Ancient DNA, 25 bone cracking, 8, 16, 19, 289, 304 Andrewsarchus, 304 brain size, 39, 43–7 Aonyx, 236 lions, 168 aquatic species, 4, see Pinnipedia Buxolestes, 271 ArcGIS, 379 Arctictis,
  • Fossil Calibration Schemes the Soft Explosive Model Of

    Fossil Calibration Schemes the Soft Explosive Model Of

    Supplemental File 2: Fossil calibration schemes The soft explosive model of placental mammal evolution Matthew J. Phillips*,1 and Carmelo Fruciano1 1School of Earth, Environmental and Biological Sciences, Queensland University of Technology, Brisbane, Australia *Corresponding author: E-mail: [email protected] Contents Fossil Calibration schemes ......................................................................................................... 1 Table S2 .................................................................................................................................. 1 New calibrations ..................................................................................................................... 3 Figure S2 ................................................................................................................................ 7 References ............................................................................................................................ 10 Calibration schemes Table S2. Soft-bound calibrations employed for the MCMCtree analyses of the 122-taxon, 128- taxon, and 57-taxon empirical datasets. Calibrations among placental mammals are largely based on dos Reis et al. [1], hence the designations dR32 and dR40 (numbers indicating the number of calibrations). Several new calibrations, including some inspired by Springer et al. [2], are described below the table. Also note that the 122-taxon dR40, 128-taxon and 57-taxon analyses employ bounds as listed below, whereas the dR32 analyses,
  • University of Florida Thesis Or Dissertation Formatting

    University of Florida Thesis Or Dissertation Formatting

    UNDERSTANDING CARNIVORAN ECOMORPHOLOGY THROUGH DEEP TIME, WITH A CASE STUDY DURING THE CAT-GAP OF FLORIDA By SHARON ELIZABETH HOLTE A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2018 © 2018 Sharon Elizabeth Holte To Dr. Larry, thank you ACKNOWLEDGMENTS I would like to thank my family for encouraging me to pursue my interests. They have always believed in me and never doubted that I would reach my goals. I am eternally grateful to my mentors, Dr. Jim Mead and the late Dr. Larry Agenbroad, who have shaped me as a paleontologist and have provided me to the strength and knowledge to continue to grow as a scientist. I would like to thank my colleagues from the Florida Museum of Natural History who provided insight and open discussion on my research. In particular, I would like to thank Dr. Aldo Rincon for his help in researching procyonids. I am so grateful to Dr. Anne-Claire Fabre; without her understanding of R and knowledge of 3D morphometrics this project would have been an immense struggle. I would also to thank Rachel Short for the late-night work sessions and discussions. I am extremely grateful to my advisor Dr. David Steadman for his comments, feedback, and guidance through my time here at the University of Florida. I also thank my committee, Dr. Bruce MacFadden, Dr. Jon Bloch, Dr. Elizabeth Screaton, for their feedback and encouragement. I am grateful to the geosciences department at East Tennessee State University, the American Museum of Natural History, and the Museum of Comparative Zoology at Harvard for the loans of specimens.
  • Raymond M. Alf Museum of Paleontology EDUCATOR's GUIDE

    Raymond M. Alf Museum of Paleontology EDUCATOR's GUIDE

    Raymond M. Alf Museum of Paleontology EDUCATOR’S GUIDE Dear Educator: This guide is recommended for educators of grades K-4 and is designed to help you prepare students for their Alf Museum visit, as well as to provide resources to enhance your classroom curriculum. This packet includes background information about the Alf Museum and the science of paleontology, a summary of our museum guidelines and what to expect, a pre-visit checklist, a series of content standard-aligned activities/exercises for classroom use before and/or after your visit, and a list of relevant terms and additional resources. Please complete and return the enclosed evaluation form to help us improve this guide to better serve your needs. Thank you! Paleontology: The Study of Ancient Life Paleontology is the study of ancient life. The history of past life on Earth is interpreted by scientists through the examination of fossils, the preserved remains of organisms which lived in the geologic past (more than 10,000 years ago). There are two main types of fossils: body fossils, the preserved remains of actual organisms (e.g. shells/hard parts, teeth, bones, leaves, etc.) and trace fossils, the preserved evidence of activity by organisms (e.g. footprints, burrows, fossil dung). Chances for fossil preservation are enhanced by (1) the presence of hard parts (since soft parts generally rot or are eaten, preventing preservation) and (2) rapid burial (preventing disturbance by bio- logical or physical actions). Many body fossils are skeletal remains (e.g. bones, teeth, shells, exoskel- etons). Most form when an animal or plant dies and then is buried by sediment (e.g.
  • Mammals from the Earliest Uintan (Middle Eocene) Turtle Bluff Member, Bridger Formation, Southwestern Wyoming, USA, Part 1: Primates and Rodentia

    Mammals from the Earliest Uintan (Middle Eocene) Turtle Bluff Member, Bridger Formation, Southwestern Wyoming, USA, Part 1: Primates and Rodentia

    Palaeontologia Electronica palaeo-electronica.org Mammals from the earliest Uintan (middle Eocene) Turtle Bluff Member, Bridger Formation, southwestern Wyoming, USA, Part 1: Primates and Rodentia Thomas S. Kelly and Paul C. Murphey ABSTRACT The Turtle Bluff Member (TBM) is the stratotype section for the earliest Uintan bio- chron, Ui1a, of the middle Eocene Uintan North American Land Mammal age. For more than a century, the TBM had yielded only a few fragmentary specimens. As the result of many years of field work, numerous mammal fossils have now been recov- ered and provide an unprecedented opportunity to better define this poorly known interval. This is the first in a series of papers that provide detailed descriptions and tax- onomic revisions of the fauna of the TBM. Here we document the occurrence of the fol- lowing taxa in the TBM: Uintasorex parvulus; Microsyops annectans; Notharctus robustior; Omomys carteri; Trogolemur myodes; Washakius insignis; Thisbemys corru- gatus; Microparamys minutus; Microparamys sp.; Sciuravus nitidus; Tillomys senex; Tillomys? parvidens; Taxymys lucaris; Pauromys sp., cf. P. perdit us ; three informal sci- uravid species (sp. A, B and C); cf. Pareumys sp.; Metanoiamys sp.; and Elymys? emryi new species. Except for the previously described Hemiacodon engardae, all of the primates from the TBM are holdover taxa from the earlier Bridgerian Land Mammal age, whereas the rodents exhibit a modest diversification during the earliest Uintan. Elymys? emryi and four additional informal rodent species (Microparamys sp., sci- uravid sp. A, cf. Pareumys sp., and Metanoiamys sp.) make their appearances in the TBM and, as such, can be added to the list of index species characterizing biochron Ui1a.
  • Optimizing Phylogenetic Supertrees Using Answer Set Programming Laura Koponen1, Emilia Oikarinen1, Tomi Janhunen1, and Laura Säilä2 1 HIIT / Dept

    Optimizing Phylogenetic Supertrees Using Answer Set Programming Laura Koponen1, Emilia Oikarinen1, Tomi Janhunen1, and Laura Säilä2 1 HIIT / Dept

    Optimizing Phylogenetic Supertrees Using Answer Set Programming Laura Koponen1, Emilia Oikarinen1, Tomi Janhunen1, and Laura Säilä2 1 HIIT / Dept. Computer Science, Aalto University 2 Dept. Geosciences and Geography, University of Helsinki Computational logic day 2015 — Aalto, Finland Outline Introduction — the supertree problem ASP Encodings — trees, quartets and projections Experiments — Felidae data Conclusions Koponen et al., Optimizing Phylogenetic Supertrees Using ASP Computational logic day 2015 2/31 I Several measures can be used used I Optimal tree not necessarily unique I Output: a phylogenetic tree that covers all taxa from input and reflects the relationships in input as well as possible The supertree problem I Input: a set of overlapping, possibly conflicting phylogenetic trees (rooted, leaf-labeled) Koponen et al., Optimizing Phylogenetic Supertrees Using ASP Computational logic day 2015 3/31 The supertree problem I Input: a set of overlapping, possibly conflicting phylogenetic trees (rooted, leaf-labeled) I Output: a phylogenetic tree that covers all taxa from input and reflects the relationships in input as well as possible I Several measures can be used used I Optimal tree not necessarily unique Koponen et al., Optimizing Phylogenetic Supertrees Using ASP Computational logic day 2015 4/31 Solving the supertree problem I Typically heuristic methods are used, e.g. matrix representation with Parsimony (MRP) [Baum, 1992; Ragan,1992] I input trees encoded into a binary matrix, and maximum parsimony analysis is then used to construct
  • A New, Fast Method to Search for Morphological Convergence with Shape Data

    A New, Fast Method to Search for Morphological Convergence with Shape Data

    This is a repository copy of A new, fast method to search for morphological convergence with shape data. White Rose Research Online URL for this paper: https://eprints.whiterose.ac.uk/163919/ Version: Published Version Article: Castiglione, Silvia, Serio, Carmela, Tamagnini, Davide et al. (7 more authors) (2019) A new, fast method to search for morphological convergence with shape data. PLoS ONE. e0226949. ISSN 1932-6203 https://doi.org/10.1371/journal.pone.0226949 Reuse This article is distributed under the terms of the Creative Commons Attribution (CC BY) licence. This licence allows you to distribute, remix, tweak, and build upon the work, even commercially, as long as you credit the authors for the original work. More information and the full terms of the licence here: https://creativecommons.org/licenses/ Takedown If you consider content in White Rose Research Online to be in breach of UK law, please notify us by emailing [email protected] including the URL of the record and the reason for the withdrawal request. [email protected] https://eprints.whiterose.ac.uk/ RESEARCH ARTICLE A new, fast method to search for morphological convergence with shape data Silvia Castiglione1, Carmela Serio1, Davide Tamagnini2, Marina Melchionna1, Alessandro Mondanaro1,3, Mirko Di Febbraro4, Antonio Profico2, Paolo Piras5,6, 1 1 Filippo Barattolo , Pasquale RaiaID * 1 Dipartimento di Scienze della Terra, dell’Ambiente e delle Risorse, University of Naples Federico II, Napoli, Italy, 2 Dipartimento di Biologia Ambientale, Sapienza Università