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Rock Matrix Surrounding Subfossil Lemur Skull Yields Diverse Collection of Mammalian Subfossils: Implications for Reconstructing Madagascar’S Paleoenvironments

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Rock Matrix Surrounding Subfossil Lemur Skull Yields Diverse Collection of Mammalian Subfossils: Implications for Reconstructing Madagascar’S Paleoenvironments Rock matrix surrounding subfossil lemur skull yields diverse collection of mammalian subfossils: Implications for reconstructing Madagascar’s paleoenvironments Karen E. Samonds1, Sara N. Parent2, Kathleen Species identified includeMicrogale sp. (Afrosoricida: M. Muldoon3, Brooke E. Crowley4 & Laurie R. Tenrecidae), Eliurus myoxinus (Rodentia: Godfrey5 Nesomyidae), Hipposideros sp. cf. H. besaoka and 1Redpath Museum, McGill University, 859 Sherbrooke Triaenops sp. (Chiroptera: Hipposideridae), Galidia St. W., Montreal, Quebec, Canada H3A 2K6 elegans (Carnivora: Eupleridae), Cheirogaleus Email: [email protected] medius and Microcebus murinus (Primates: 2 Department of Anatomical Sciences, School of Cheirogaleidae), and Hippopotamus lemerlei Medicine, Health Sciences Center, Stony Brook (Artiodactyla: Hippopotamidae). Radiocarbon dating University, Stony Brook, New York 11794, USA shows that non-associated surface finds of small Email: [email protected] mammals tend to be younger than extinct larger 3Department of Anatomy, Dartmouth Medical School, mammals at Anjohibe, underscoring the importance and Department of Anthropology, Dartmouth College, of using other methods to establish temporal Hanover, New Hampshire 03755, USA Email: [email protected] associations of small and large mammals. This 4Department of Anthropology, 19 Russell Street, research demonstrates the potential for recovering University of Toronto, Toronto, Ontario, Canada M5S subfossils from matrix that are typically discarded 2S2 during the preparation of larger fossils, and highlights Email: [email protected] the potentially significant loss of information if such 5Department of Anthropology, 240 Hicks Way, University sediments are ignored. of Massachusetts, Amherst, Massachusetts 01003, Keywords: Small mammals, paleontology, acetic USA acid preparation, subfossils, Madagascar Email: [email protected] Résumé détaillé Abstract En raison de l’absence presque complète d’un Due to the near lack of a Cenozoic fossil record, little is registre fossile Cénozoïque à Madagascar, nous ne known about the origin and evolution of Madagascar’s connaissons presque rien des origines et de l’évolution extant fauna. Madagascar’s subfossil record (Late de sa faune actuelle. La plupart des fossiles décrits Pleistocene and Holocene) has been important for de la Cénozoïque sont des « subfossiles », c’est-à- filling in the most recent part of this informational gap, dire seulement quelques milliers d’années d’âge. Ces contributing details on diversity and distribution changes subfossiles ont été très importants pour augmenter in the recent past, but most research has focused nos connaissances des faunes récentes, y compris on larger animals. Less attention has been given to les distributions passées des espèces (actuelles the subfossil record of small mammals, despite the et disparues), et en précisant les dates d’arrivées fact that these groups represent a substantial portion récentes. Toutefois, la plupart de ces recherches of the extant mammalian diversity. To evaluate the était sur les animaux assez grands ; ces animaux potential presence of subfossil microfaunal remains in donc constituent la plupart des espèces décrites du cave breccias (calcite sediment), we used acetic acid registre subfossile. Actuellement, moins est connu sur to dissolve the matrix surrounding a nearly complete le registre subfossile des animaux plus petits, malgré skull and mandible of Archaeolemur sp. cf. A. edwardsi leur dominance numérique parmi les faunes actuelles. from Anjohibe Cave, northwestern Madagascar. The En d’autres pays, les petits mammifères ont été très resulting residue included fossil remains of all five utiles pour mieux comprendre les changements orders represented by Madagascar’s extant mammals climatiques, le paléo-environnement et l’écologie des (Afrosoricida, Carnivora, Chiroptera, Rodentia, and espèces et des écosystèmes disparus. Primates), and one order, Artiodactyla, currently Nous avons évalué la présence des os subfossiles extinct in Madagascar, except for introduced forms. des petits mammifères dans les sédiments calcaires Samonds, K. E., Parent, S. N., Muldoon, K. M., Crowley, B. E. & Godfrey, L. R. 2010. Rock matrix surrounding subfossil lemur skull yields diverse collection of mammalian subfossils: Implications for reconstructing Madagascar’s paleoenvironments. Malagasy Nature, 4: 1-16. 2 Samonds et al. : Rock matrix surrounding subfossil lemur skull consolidés de la grotte d’Anjohibe, au Nord- fossiles récupérés par les tamis, priorités de récolte et ouest de Madagascar. Cette grotte a déjà produit choix des sites) peuvent affecter la taille moyenne des des subfossiles de plusieurs animaux disparus, dont fossiles collectionnés. Par ailleurs, les petits fossiles les lémuriens, hippopotames et crocodiles. Nous avons peuvent être perdus ou détruits en train de préparation utilisé l’acide acétique pour faire fondre une brèche des fossiles et sédiments. Cette préparation peut entourant un crâne et une mandibule presque complète inclure la préparation d’acide pour les sédiments d’Archaeolemur sp. cf. A. edwardsi. Le résidu calcaires (qui, généralement, préservent les petits résultant a inclus des os subfossiles attribuable à fossiles), ou l’enlèvement manuel du sédiment chaque ordre des mammifères actuels de Madagascar entourant un fossile intéressant (qui, généralement, (Afrosoricida, Carnivora, Chiroptera, Rodentia et détruit les petits fossiles ou les laisse enchâssés dans Primates), et un ordre qui était présent, mais est le sédiment). actuellement disparu à Madagascar (Artiodactyla ; les Nous suggérons que les paléontologues hippopotames sont disparus, les espèces restantes doivent considérer les effets de leurs méthodes de sont introduites). Les espèces identifiées incluent reconnaissance, de collecte, et de préparation sur Microgale sp. (Afrosoricida: Tenrecidae), Eliurus leur réussite en récupération des petits fossiles. myoxinus (Rodentia: Nesomyidae), Hipposideros Actuellement, il parait qu’il y a une balance entre sp. cf. H. besaoka et Triaenops sp. (Chiroptera: la récupération rapide des plus grands fossiles Hipposideridae), Galidia elegans (Carnivora: et la récupération d’une faune plus complète, qui Eupleridae), Cheirogaleus medius et Microcebus peut prendre plus de nos efforts et de notre temps. murinus (Primates: Cheirogaleidae) et Hippopotamus Néanmoins, si nous augmenterons la représentation lemerlei (Artiodactyla: Hippopotamidae). La présence des petits fossiles aux études paléontologiques à de Galidia, un carnivore spécifique aux milieux Madagascar, ils nous donneront une reconstruction forestiers, représente la première récolte de ce genre plus complète de l’histoire de la faune malgache, dont d’Anjohibe, et sa présence corrobore la reconstruction la diversité des communautés passées, et l’évolution d’un écosystème forestier (en accord avec la présence des distributions des espèces et écosystèmes. Ces des grands lémuriens adaptés à la locomotion efforts nous donneront une meilleure compréhension arboréale). Deux espèces disparues ont été récoltées de la chronologie de cette histoire, et peuvent aussi (Hipposideros besaoka et Hippopotamus lemerlei), nous donner un cadre pour évaluer les hypothèses ce qui nous donne une reconstruction plus complète sur les causes de ces changements ; alors, on peut de la biodiversité des siècles et millénaires passés mieux comprendre l’histoire environnementale de la dans ce site. Nous avons aussi utilisées la datation faune malgache en même temps qu’évaluer les effets par le carbone-14 pour montrer que les fossiles des changements écologiques présents et futurs. des mammifères actuels d’Anjohibe ne sont pas de Mots clés: Petits mammifères, paléontologie, même âge que les fossiles des espèces disparus ; préparation en acide acétique, subfossiles, l’association des spécimens est alors très importante Madagascar pour les interprétations paléo-écologiques. Cette recherche nous donne un exemple de la présence des petits subfossiles dans les sédiments Introduction des sites subfossiles, et le potentiel pour dévoiler Madagascar’s modern fauna is unique and exhibits l’écologie et l’évolution des espèces passées et one of the highest levels of endemism on the planet leurs environnements. Malheureusement, ces (Goodman & Benstead, 2005). Exploring how, when, sédiments sont souvent perdus (peut-être avec and from where the island’s animals arrived has been des petits fossiles dedans) ou détruits lors de la called “one of the greatest unsolved mysteries of préparation des grands fossiles. Actuellement, les natural history” (Krause et al., 1997). The evolutionary méthodes paléontologiques, sur le terrain et au and biogeographic history of many of the island’s laboratoire, peuvent contribuer à la disparition des groups remains poorly known due to the near-lack of petits subfossiles. Les petites dents et les petits os a Cenozoic fossil record (see Krause et al., 2006 for sont généralement plus fragiles et plus susceptibles discussion). d’être détruits par des processus taphonomiques. En Our only direct window into the evolutionary history plus, leur taille les rendent plus difficile à trouver par of modern Malagasy mammals is based on very des paléontologues sur terrain, et les méthodes de recent subfossil remains, with the deepest glimpse at collecte (techniques de reconnaissance, la taille des a mere ~80,000 years before present (BP) (Samonds, Samonds et al. : Rock matrix surrounding subfossil
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