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Condylura (Mammalia, Talpidae) reloaded: New insights about the fossil representatives of the genus

Gabriele Sansalone, Tassos Kotsakis, and Paolo Piras

Plain Language Abstract

The star nosed mole, Condylura cristata, possesses a unique mixture of apomorphic and plesiomorphic characters such as the star-like appendages surrounding the nostrils and a forelimb arm bone with slight adaptation to digging. Despite the presence of an abundant literature about the life history of the extant star nosed mole, very little is known about the extinct species belonging to the genus. In this survey we review the fossil record of Condylura in order to provide new insights about the systematics, palaeoecology and palaeobiogeography of this genus. Furthermore, we re-evaluate a fossil Condylura from the middle Miocene of Kazakhstan. The re-description of this specimen allows to hypothesize that evolutionary lineage of star nosed moles probably originated in Eurasia rather than in North America.

Resumen en Español

Condylura (Mammalia, Talpidae) revisitado: nuevas ideas sobre los representantes fósiles del género

El topo de nariz estrellada, Condylura cristata, ha sido profundamente investigado por diferentes autores debido a sus peculiaridades morfológicas y de comportamiento. Por el contrario, se sabe muy poco sobre las relaciones filogenéticas, evolución y diversidad de los miembros fósiles de este género. En el presente estudio se aportan nuevos conocimientos sobre los especímenes fósiles atribuidos a Condylura teniendo en cuenta aspectos sistemáticos, paleobiogeográficos y paleoecológicos. Además, proporcionamos una re-descripción de un Condylura fósil del Mioceno medio de Kazakstán. Confirmamos que el fósil kazajo pertenece al género Condylura, de acuerdo con las características morfológicas del húmero, y discutimos sus implicaciones y su impacto en el escenario filogenético y la ecología de este peculiar género de tálpido. Este espécimen representa el primer registro del género, lo que sugiere un origen eurasiático en lugar del escenario más comúnmente aceptado de un origen norteamericano. La presencia tanto de caracteres plesiomórficos como apomórficos en Condylura apoya firmemente la hipótesis de que este género podría ser considerado como un clado hermano de Talpinae.

Palabras clave: Talpidae, húmero, sistemática, Mioceno, paleobiogeografía, paleoecología

Traducción: Enrique Peñalver (Sociedad Española de Paleontología)

Résumé en Français

Révision des représentants fossiles du genre Condylura (Mammalia, Talpidae)

Le condylure étoilé, Condylura cristata, a été étudié en détail par différents auteurs en raison de ses particularités morphologiques et comportementales. En revanche, les relations phylogénétiques, l'évolution, et la diversité des membres fossiles de ce genre sont très peu connues. Nous présentons ici une révision des spécimens fossiles attribués à Condylura, en abordant les aspects systématiques, paléobiogéographiques, et paléoécologiques. De plus, nous redécrivons un fossile de Condylura du Miocène moyen du Kazakhstan. Nous confirmons que le fossile kazakh appartient au genre Condylura d'après des caractères morphologiques de l'humérus, et nous discutons les implications et l'impact sur le scénario phylogénétique et l'écologie de ce genre particulier de talpidé. Ce spécimen représente la plus ancienne occurrence de Condylura, suggérant ainsi une origine eurasiatique de ce genre plutôt qu'une origine nord-américaine (le scénario généralement accepté). La présence de caractères à la fois plésiomorphes et apomorphes chez Condylura soutient fortement l'hypothèse que ce genre pourrait être le groupe-frère des Talpinae.

Mots-clés : Talpidae ; humérus ; systématique ; Miocène ; paléobiogéographie ; paléoécologie

Translator: Antoine Souron

Deutsche Zusammenfassung

Condylura (Mammalia, Talpidae) neu aufgelegt: neue Erkenntnisse über die fossilen Vertreter der Gattung

Der Sternmull, Condylura cristata, wurde wegen seiner morphologischen Besonderheiten und wegen seines Verhaltens von verschiedenen Autoren eingehend untersucht. Demgegenüber ist sehr wenig über die phylogenetischen Beziehungen, Evolution und Diversität der fossilen Vertreter dieser Gattung bekannt. In der vorliegenden Untersuchung geben wir neue Einblicke in fossile Stücke die Condylura zugeschrieben werden unter Berücksichtigung systematischer, paläobiogeografischer und paläoökologischer Aspekte. Des Weiteren offerieren wir eine Neubeschreibung eines fossilen Condylura aus dem mittleren Miozän von Kasachstan. Wir bestätigen, dass das kasachische Fossil zur Gattung Condylura gehört, basierend auf morphologischen Merkmalen im Humerus, und wir diskutieren seine Auswirkungen und seinen Einfluss auf das phylogenetische Szenario und auf die Ökologie dieser bestimmten Maulwurfsgattung. Dieses Stück stellt den ersten Nachweis dieser Gattung dar und legt somit einen eurasischen Ursprung nahe, entgegen des allgemein akzeptierten Szenarios eines nordamerikanischen Ursprungs. Das Vorkommen von sowohl plesiomorphen als auch apomorphen Merkmalen unterstützt die Hypothese, dass diese Gattung als Schwestertaxon der Klade Talpidae angesehen werden könnte.

Schlüsselwörter: Talpidae, Humerus, Systematik, Miozän, Paläobiogeographie, Paläoökologie

Translator: Eva Gebauer

Arabic

Translator: Ashraf M.T. Elewa

 

 

FIGURE 1. A detail of the star-like appendages from a preserved specimen. Image modified from Catania (2012b).

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FIGURE 2. Drawing of a left humerus of Condylura cristata showing the anatomical terms used in this study (modified from Hutchinson, 1984). Abbreviations: bg, bicipital groove; gt, greater tuberosity; gs, greater sulcus; hh, humeral head; le, lateral epicondyle; lt, lesser tuberosity; me, medial epicondyle; met, medial edge of the trochlea; ms, minor sulcus; of, olecranon fossa; pc, pectoral crest; tr, trochlea; tt, teres tubercle.

 

FIGURE 3. Drawings ( 1-5, 8) and photographs ( 6 and 7) showing fossil and extant representatives of genus Condylura. 1 and 2, Condylura sp., left humerus in caudal ( 1) and frontal ( 2) views, modified from Lychev (1963). 3-5, cf. Condylura sp., right humerus (YPM 20699, YPM) in caudal ( 3) and lateral ( 4) views and close-up of the greater tuberosity in medial view ( 5), modified from Hutchinson (1984, figure 1B). 6 and 7, Condylura kowalskii, left humerus (MF/1006/16, ISEZ-PAN) in frontal ( 6) and caudal ( 7 ) views, modified from Rzebik-Kowalska (2014, figure 2.1, 2.2). 8, Condylura cristata, right humerus in caudal view, modified from Hutchinson (1984, figure 1A). Scale bars represent 1 mm.

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FIGURE 4. 1, Condylura kowalskii, mandible fragment bearing p1 and p2 in lingual view (MF/1006/4, ISEZ-PAN); from Rzebik-Kowalska (2014, figure 1.3). 2, Line drawing of a left mandible of Condylura cristata in labial view, modified from Laerm et al. (2007).

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FIGURE 5. Drawings ( 1-4) and photographs ( 5-6) of talpid species humeri in caudal view. 1, Yanshuella primaeva, left humerus (IVPP-6455, IVPP), from Storch and Qiu (1983, figure 39b). 2, Yunoscaptor scalprum, left humerus (IVPP-V-9741.38, IVPP), from Storch and Qiu (1991, figure 48b). 3, Scapanulus oweni, left humerus, from Hutchinson (1968). 4, Wilsonius ripafodiator, left humerus (UO-22366, UO), from Hutchinson (1968, figure 55). 5, Leptoscaptor robustior, right humerus (P35-58.6, NHMA), from Ziegler (2003, figure 3B2). 6, Leptoscaptor bavaricum, left humerus (P10-610.2, NHMA), from Ziegler (2003, figure 2I).

 figure5a

FIGURE 6. 1, Talpidae phylogeny resulting from a parsimony analysis based on 157 discrete morphological characters, modified from Sánchez-Villagra et al. (2006, figure 1). 2, Unconstrained Talpidae phylogeny based on a parsimony analysis (DELTRAN optimization) based on 176 discrete morphological characters, modified from Schwermann and Thompson (2015, figure 16A).

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FIGURE 7. Map showing Condylura fossil bearing localities and a possible palaeobiogeographical scenario. The orange striped area represents the present distribution of the extant C. cristata. The orange star represents Condylura sp. (middle Miocene locality of Kalkaman Lake, Kazakhstan). The red square represents cf. Condylura sp. (late Miocene locality of Malheur River, Oregon, USA). The green triangles represent C. kowalskii and C. izabellae (Pliocene Polish localities). The blue circles represent the late Pleistocene-Holocene localities of C. cristata.

figure7

 

 

sansaloneGabriele Sansalone. Roma Tre University of Rome, Dept. of Sciences, L.S. Murialdo, 1 - 00146 Rome, Italy/Center for evolutionary ecology, C.da Fonte Lappone, Pesche, Italy/Form, Evolution and Anatomy Research Laboratory, Zoology, School of Environmental and Rural Sciences, University of New England, Armidale, NSW 2351, Australia This email address is being protected from spambots. You need JavaScript enabled to view it.

Gabriele Sansalone had is master degree in evolutionary biology. In 2012 he started his PhD at the Roma Tre University of Rome, Italy in vertebrate palaeontology, focusing on Talpidae systematics and evolution. He's investigating the evolution of fossoriality in moles via Geometric Morphometrics and 3D biomechanical simulation.

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kotsakisTassos Kotsakis. Roma Tre University of Rome, Dept. of Sciences, L.S. Murialdo, 1 - 00146 Rome, Italy/Center for evolutionary ecology, C.da Fonte Lappone, Pesche, Italy This email address is being protected from spambots. You need JavaScript enabled to view it.

Born in 1945. Graduate in 1971. Researcher of CNR from 1976, Full Professor from 1990, visiting professor at Université Pierre et Marie Curie (Paris VI; 2008), during several years was Dean of the Faculty of Sciences, Head of Department, President of the Teaching Council of Geological Sciences, Coordinator of the Council of PhD students, Coordinator of the Vertebrate Palaeontology Group of CNR, several times member of the Academic Senate. Founder member of the Roman Theriological Association, of the Societas Herpetologica Italica and of the Center for Evolutionary Ecology. Member of editorial board of several scientific journals. He has been and is leading researcher and valuer of both national and international projects. He has been member of several panels for the evaluation of Departments of Geology and Mining Engineering in many countries of EU. Tutor of fourteen PhDs in Vertebrate Palaeontology. He works on Vertebrate Palaeontology, particularly on insular fossil faunas, both endemic and non-endemic, on the Cenozoic continental palaeobiogeography of the Mediterranean area, and on the systematics of Neogene and Quaternary small mammals of the Mediterranean area. The results of his work are reported in 175 papers published or in press, in both national and international reviews and volumes. His participation on Congresses and Workshops is testified by more than 100 abstracts.

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pirasPaolo Piras. Center for evolutionary ecology, C.da Fonte Lappone, Pesche, Italy/Dipartimento di Scienze Cardiovascolari,Respiratorie, Nefrologiche, Anestesiologiche e Geriatriche, “Sapienza” University of Rome, Rome, Italy/Dipartimento di Ingegneria Strutturale e Geotecnica, “Sapienza” University of Rome, Via Eudossiana 18, 00100, Rome, Italy This email address is being protected from spambots. You need JavaScript enabled to view it.

Paolo Piras is an evolutionary paleobiologist. After the PhD in paleontology at Roma Tre University, Italy; he was postdoctoral research fellow in biology (2008) at Pierre et Marie Curie University (Paris, France) and at Roma Tre University (2010-2012) in paleontology. Currently is postdoctoral fellow at Sapienza, University of Roma, Italy, for studying shape changes of the human left ventricle via Geometric Morphometrics using data coming from 3D echocardiography. He applies methods of modern shape analysis to biological and biomechanical problems spanning from systematics of extinct and extant vertebrates to human cardiac mechanics. 

 

List of material used for comparisons.

Condylura cristata. NHM-3.11.5, humerus and mandible. NHM-3.11.6, humerus and mandible. NHM-1.7.4, mandible. NHM-12.12.1, mandible. NHM-1.25.2, mandible. NHM-7.7.2659, mandible. NHM-7.7.132, mandible. NHM-7.7.2792, mandible. NHM-12.5.3, mandible. NHM-1.7.2, mandible. NHM-1.7.3, mandible. Natural History Museum NHM, London, UK. 62567, mandible. NHMW.

Condylura izabellae. MF/1007/01, humerus. ISEZ-PAN.

Condylura kowalskii. MF/1006/4, mandibular fragment bearing p1-p2. MF/1006/16, humerus. MF/1006/18, humerus. MF/1006/21, humerus. MF/1006/22, humerus. MF/1006/23, humerus. ISEZ-PAN.

Leptoscaptor bavaricum. P10-610.2, humerus. Ptb68_1962, humerus. Ptb68_1962b, humerus. Ptb68_1962c, humerus. NHMA.

Leptoscaptor robustior. P35-58.6, humerus. NHMA.

Scapanulus oweni. Drawings from Hutchinson, 1968; Storch and Qiu, 1983.

Mioscalops ripafodiator. UCMP-70140b, humerus. UCMP-V70140a, humerus. UCMP-V70140d, humerus. University of California Museum of Palaeontology, UCMP, Berkeley, California, USA.

Yanshuella primaeva. IVPP-6455, humerus. IVPP-6456, humerus. IVPP.

Yunoscaptor scalprum. IVPP-V-9741.38, humerus. IVPP.