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( Ozotoceros Bezoarticus Celer: Cervidae) Cranial Morphology

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( Ozotoceros Bezoarticus Celer: Cervidae) Cranial Morphology DOI 10.1515/mammalia-2013-0051 Mammalia 2014; aop Guillermo H. Cassini * , David A. Flores and Sergio F. Vizca í no Postnatal ontogenetic scaling of Pampas Deer ( Ozotoceros bezoarticus celer : Cervidae) cranial morphology Abstract: The pampas deer Ozotoceros bezoarticus is a Introduction medium-sized, elegant, and lightly built cervid that was once a characteristic inhabitant of open grasslands across Cervidae account for the larger diversity of South Ameri- a wide geographical distribution in South America. In can extant ungulates with five genera and eight species Argentina, the subspecies Ozotoceros bezoarticus celer is recognized in Argentina ( Merino 2006 ). The pampas the most southern and abundant populations and rela- deer Ozotoceros bezoarticus (Linnaeus, 1758) was once tively well represented in osteological collections per- a characteristic inhabitant of open grasslands across a mitting a morphometric study of cranial ontogeny. We wide geographical distribution, from 5 ° to 41 ° S includ- measured 17 cranial variables on an ontogenetic series ing regions of Brazil, Bolivia, Paraguay, Argentina, and in order to evaluate the multivariate allometry of neu- the whole Uruguay ( Jackson 1987 , Jackson and Giulietti rocranial and splanchnocranial components through 1988 , Pautasso et al. 2002 , Merino 2006 , González et al. developmental stages. The cranial ontogenetic pattern 2010 , Merino and Vieira Rossi 2010 ). Only few small iso- in O. b. celer exhibits a conservative plan in which both lated populations remain at present, and the pampas deer sexes share most of the allometric trends except in three is considered by the UICN as near threatened in South variables related to the rostrum. Isometry was detected America ( González and Merino 2008 ). Four separated in 9 out of 17 variables, whereas four grows with negative populations of two subspecies are present in Argentina: allometry, and only nasal length showed positive allom- Ozotoceros bezoarticus leucogaster Goldfuss, 1817, in the etry. As for most mammals, those variables related to the provinces of Santa Fe and Corrientes ( Parera and Moreno neurocranial components and sensorial capsules exhibit 2000 , Pautasso et al. 2002 ), and Ozotoceros bezoarti- negative allometry. The growth of the rostrum indicates cus celer Cabrera, 1943 , in the provinces of San Luis and that both sexes have longer face than younger. However, Buenos Aires ( Dellafiore et al. 2003 , P é rez Carusi et al. the lack of allometry in most of the variables seems to be 2009 ). The southern and more abundant population of O . supporting the idea of a younger with the general appear- b . celer is relatively well represented in osteological collec- ance of an adult but smaller size. tions and pertinent for morphological studies. The pampas deer is a medium-sized (30 – 40 kg), Keywords: Cervidae; early development; multivariate elegant, and lightly built cervid ( Jackson 1987 ). As most allometry; skull; South America. cervids, it presents sexual dimorphism, with males having three pointed small antlers (∼ 30 cm) and females lacking *Corresponding author: Guillermo H. Cassini, Divisi ó n antlers and showing instead a white twisted shape tuft Mastozoolog í a, Museo Argentino de Ciencias Naturales “ Bernardino on the frontal zone ( Cabrera 1943 , Cosse Larghero 2002 ). Rivadavia ” , CONICET, Av. Angel Gallardo 470, C1405DJR, Ciudad The offspring weigh 1.5 – 2 kg at birth, and at the age of Aut ó noma de Buenos Aires, Argentina; and Departamento de 7 months, they show almost complete adult aspect, except Ciencias B á sicas, Universidad Nacional de Luj á n, Ruta 5 y Av. Constituci ó n s/n, Luj á n (6700), Buenos Aires, Argentina, e-mail: for the smaller body size ( Fr ä derich 1981 , Gimenez-Dixon [email protected] 1991 ). The pampas deers have broad feeding strategies David A. Flores: Divisi ó n Mastozoolog í a, Museo Argentino de throughout their distribution in relation to phytogeo- Ciencias Naturales “ Bernardino Rivadavia ”, CONICET, Av. Angel graphical variation ( Merino 2003 ) and, consequently, Gallardo 470, C1405DJR, Ciudad Aut ó noma de Buenos Aires, are described as having an opportunistic foraging strat- Argentina Sergio F. Vizca í no: Divisi ó n Paleontolog í a Vertebrados, Museo egy typical of intermediate or mixed feeders ( Cosse et al. de La Plata, CONICET, Paseo del Bosque s/n, B1900FWA La Plata, 2009 ). Different populations of the subspecies studied Argentina here, Ozotoceros bezoarticus celer , presents slightly dif- ferent feeding behaviors, as those from San Luis Province Brought to you by | National Dong Hwa University Authenticated | 134.208.103.160 Download Date | 3/25/14 10:29 PM 2 G.H. Cassini et al.: Multivariate skull allometry of Ozotoceros feed primarily on new green growth grass ( Jackson and of the mandibular corpus) growth is mainly prenatal, Giulietti 1988 , but see Merino et al. 2009 ), whereas those whereas the posterior section of the jaw (i.e., molar part of from Campos del Tuy ú Wildlife Reserve (Buenos Aires the mandibular corpus and ramus of the mandible) devel- Province) were described as mixed grass feeders with ops in parallel with the transition in main food source preference for grasses ( Jackson and Giulietti 1988 ). from milk to herbal forage ( Todd and Wharton 1934 , Pond Although some authors carried out cranial mor- 1977 , Herring 1985 , H ø ye and Forchhammer 2006 ). phometric studies on South American cervids including In this contribution, we investigate the way in which Ozotoceros bezoarticus (e.g., Wemmer and Wilson 1987 , the cranial configuration of both populations of Ozoto ceros Delupi and Bianchini 1995 , Beade et al. 2000 , González bezoarticus celer changes from juveniles to adults, apply- et al. 2002 ), they are focused primarily on taxonomic ing a multivariate quantitative approach on a well-rep- variations. More recently, Merino et al. (2005) correlated resented ontogenetic series of skulls of different inferred diet and morphology in this species and made prelimi- age stages. This statistical approach is possible thanks to nary observations on ontogeny. Perhaps the first contri- recent efforts to collect dead carcasses in San Luis prov- bution to the skull ontogeny of ungulates was the Allen ince and “ Campos del Tuy ú Wildlife Reserve ” yielded to (1913) paper on Ovibos moschatus (Zimmermann 1780). enlarge the number of O. b. celer skulls in museum col- However, with some exceptions as horses ( Radinsky 1984 ), lections of Argentina ( Figure 1 A – F). Together with the col- camels (Al-Sagair and ElMougy 2002 ), hippos ( Weston lections made in the late 1960s, an excellent ontogenetic 2003 ), and the extinct South American toxodontids (order series for the two localities of this endangered species is Notoungulata) ( Cassini et al. 2012 ), little is known about available. The ontogenetic series studied here allows the the ontogenetic development of ungulate cranium and its evaluation of the morphological changes on functional relationships with the transition to a herbivorous feeding grounds, during a period of development in which critical style in an allometric framework. To date, some studies changes occur, as interactions between growth of neuro- on ontogeny in ungulates suggest that the medioanterior cranial and splanchnocranial components through devel- regions of the mandible (i.e., diastema and premolar parts opmental stages causes modifications in the function of A D E B F C 100 nm Figure 1 Three ontogenetical states of Ozotoceros bezoarticus . (A) Female juvenile MLP 18.VIII.92.4 (TSL = 168 mm mirror image); (B) female subadult MLP 31.VIII.98.1 (TSL = 197 mm); (C) female adult MLM 388 (TSL = 218 mm); (D) male juvenile MLP 23.VIII.96.1 (TSL = 150 mm mirror image); (E) male subadult MLM 401 (TSL = 188 mm); (F) male adult CDT 096 (TSL = 223 mm). Brought to you by | National Dong Hwa University Authenticated | 134.208.103.160 Download Date | 3/25/14 10:29 PM G.H. Cassini et al.: Multivariate skull allometry of Ozotoceros 3 the skull linked to the transition from juvenile to adult 255.2 mm in total skull length (MLM-391), with highly worn diets ( Abdala et al. 2001 , Giannini et al. 2004 , Flores et al. adult dentition. We chose the specimens coming from the 2006 ). In addition, we compared the allometric trends two aforementioned populations ( O. b. celer from Buenos of O. b. celer with other larger-size ungulates as hippos Aires and San Luis Provinces) in order to avoid the ecogeo- ( Weston 2003 ) and the fossil toxodontids Adinotherium graphic effects. and Nesodon ( Cassini et al. 2012 ), in order to evaluate the differences and similarities of the ontogenetic pattern (in allometric terms) of the skull in three nonrelated and mor- Study of growth and measurements phologically divergent ungulates. Allometry of growth explicitly considers timing of changes throughout the life of an individual ( Kunz and Robson Materials and methods 1995 , Prestrud and Nilssen 1995 , Maunz and German 1996 , Stern and Kunz 1998 ). By contrast, allometry of size com- pares changes against overall size along a growth series, Study specimens and the time frame is implicitly incorporated but not specified. Allometry of size may, in turn, be interspecific We analyzed an ontogenetic series of 70 specimens (36 with the purpose of studying functional changes from females and 34 males) ( Tables 1 and 2 ) of Ozotoceros an evolutionary perspective within a lineage ( Davis 1962 , bezoarticus celer housed at the mammal collections of Radinsky 1984 ,
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