Papers by Analia M. Forasiepi

We report two fossil procyonids, Cyonasua sp. and Chapalmalania sp., from the late Pliocene of Ve... more We report two fossil procyonids, Cyonasua sp. and Chapalmalania sp., from the late Pliocene of Venezuela (Vergel Member, San Gregorio Formation) and Colombia (Ware Formation), respectively. The occurrence of these pre Holocene procyonids outside Argentina and in the north of South America provides further information about the Great American Biotic Interchange (GABI). The new specimens are recognized in the same monophyletic group as procyonids found in the southern part of the continent, the “Cyonasua group,” formed by species of Cyonasua and Chapalmalania. The phylogenetic analysis that includes the two new findings support the view that procyonids dispersed from North America in two separate events (initially, previous to the first major migration wave—GABI 1—and then within the last major migration wave—GABI 4—). This involved reciprocal lineage migrations from North to South America, and included the evolution of South American endemic forms.
Palaontol Z, Jun 2014
We describe a new toxodontid species, Nesodon taweretus sp. nov.

Journal of Systematic Palaeontology, Jun 2014
An almost complete skull of Australohyaena antiqua (Ameghino), from the Late Oligocene (Deseadan ... more An almost complete skull of Australohyaena antiqua (Ameghino), from the Late Oligocene (Deseadan SALMA) of Cabeza Blanca, Chubut Province, Argentina is described and analysed. For more than a century, this species was represented by isolated teeth. The genus Australohyaena gen. nov. is proposed based on a phylogenetic reconstruction that demonstrates that A. antiqua is a Borhyaenidae (Mammalia, Sparassodonta), grouped with Arctodictis and Borhyaena, but not with Pharsophorus lacerans, the genus to which antiqua was formerly assigned. A. antiqua is recognized by several features on the skull, dentary and dentition. In addition, a short snout, large canines, deep jaw, reduced protocone and talonid determine A. antiqua as hypercarnivorous. A vaulted skull, well-developed temporal fossa and little difference on the jaw depth at p3 and m4, are suggestive of bone-cracker specializations. A. antiqua is within the largest Deseadan sparassodonts with a body mass of about 70 kg. Homoplasies are detected within borhyaenoids on lower molar cusps. The metaconid is lost within Sparassodonta, although Pharsophorus and borhyaenids retained the metaconid on m2Àm4 or m2Àm3.

Swiss Journal of Palaeontology, May 2014
During the Cenozoic paucituberculatans were much more diverse taxonomically and ecomorphologicall... more During the Cenozoic paucituberculatans were much more diverse taxonomically and ecomorphologically than the three extant genera of shrew-like marsupials. Among paucituberculatans, palaeothentids were abundant during the Early Miocene, although most of the fossil remains consist of isolated teeth or fragmentary jaws. We describe a new and exceptional partial skeleton of Palaeothentes lemoinei (Palaeothentidae), collected from the Santa Cruz Formation (Santacrucian age, Early Miocene) in Patagonia. Whereas the skull of P. lemoinei has more plesiomorphic traits in the face, palate, and cranial vault than that of living paucituberculatans, the dental morphology is more derived. The osseous inner ear was examined using micro-CT scanning, revealing a cochlea with 1.9 turns, the presence of a ''second crus commune'', an anterior semicircular canal (SC) projecting slightly dorsally from the dorsal-most point of the posterior SC, and lateral and posterior SCs projecting laterally to the same level. On the basis of postcranial anatomy, previous studies have demonstrated that P. lemoinei was an agile cursorial form, an inference supported by study of the new postcranial elements.

Journal of Mammalian Evolution, 2013
The Santa Cruz Formation (late early Miocene, Santacrucian age) registers 11 species of mammalian... more The Santa Cruz Formation (late early Miocene, Santacrucian age) registers 11 species of mammalian predators (Metatheria, Sparassodonta). Together with large carnivorous flightless birds, they comprised the terrestrial predator guild. The Santacrucian sparassodonts were diverse in body size, had different locomotory habits, and were primarily hypercarnivores. The objective of this work is to analyze the guild structure of the sparassodonts of the Santa Cruz Formation, using the variables of body mass, diet, and locomotion as proxies. Furthermore, we analyze the interaction with other predators and potential prey. The univariated test V of Poole and Rathcke and the multivariated test of Clark-Evans were used to construct the models. In the multivariate test, we made a Principal Component Analysis to resume and standardize the variables. With body mass and locomotion we obtained an evenly spaced pattern of segregation for the sparassodont species, being non-significant and significant, respectively. The pattern was aggregated and significant only with diet. The analysis of all variables together resulted in an evenly spaced and significant pattern, which is consistent with character displacements (segregation of species throughout the morphospace) that would help to diminish interspecific competition during the Santacrucian age and would allow selection of prey species of different sizes and substrate specializations. When the body size pattern of predator birds and sparassodonts were plotted together, the pattern is evenly spaced and nonsignificant. Other factors, including locomotion, would differentiate these species and their ecological niches.

Nature Communications, 2014
The multiple skeletal components of the skull originate asynchronously and their developmental sc... more The multiple skeletal components of the skull originate asynchronously and their developmental schedule varies across amniotes. Here we present the embryonic ossification sequence of 134 species, covering all major groups of mammals and their close relatives. This comprehensive data set allows reconstruction of the heterochronic and modular evolution of the skull and the condition of the last common ancestor of mammals. We show that the mode of ossification (dermal or endochondral) unites bones into integrated evolutionary modules of heterochronic changes and imposes evolutionary constraints on cranial heterochrony. However, some skull-roof bones, such as the supraoccipital, exhibit evolutionary degrees of freedom in these constraints. Ossification timing of the neurocranium was considerably accelerated during the origin of mammals. Furthermore, association between developmental timing of the supraoccipital and brain size was identified among amniotes. We argue that cranial heterochrony in mammals has occurred in concert with encephalization but within a conserved modular organization.

Journal of Morphology, 2014
Used as markers of postnatal growth closure sequences of 22 ectocranial sutures and synchondroses... more Used as markers of postnatal growth closure sequences of 22 ectocranial sutures and synchondroses were recorded in a sample of 1161 skulls belonging to 38 species from all major placental clades: Afrotheria, Xenarthra, Laurasiatheria and Euarchontoglires (Boreoeutheria). The maximum closure level, which is not significantly correlated to body mass, is higher in Afrotheria and Xenarthra than in Boreoeutheria. Only the basioccipito-basisphenoid and the basioccipito-exoccipital synchondroses close in all species sampled, the supraoccipito-exoccipital and the inter-parietal sutures do in most species. Parsimov retrieved more heterochronic shifts for Afrotheria and Xenarthra than for Boreoeutheria. The amount of intraspecific variation differs among the species sampled being high among xenarthran species and low among afrotherians. Specimens (162) representing 12 marsupial genera were also analysed. Placentals exhibit a larger number of suture closures than marsupials and in both groups the sutures at the base of the skull are the first to fuse starting with the basioccipitoexoccipital. J. Morphol. 275:125-140, 2014. V C 2013 Wiley Periodicals, Inc.

Journal of Evolutionary Biology, 2014
Mammals display a broad spectrum of limb specializations coupled with different locomotor strateg... more Mammals display a broad spectrum of limb specializations coupled with different locomotor strategies and habitat occupation. This anatomical diversity reflects different patterns of development and growth, including the timing of epiphyseal growth plate closure in the long bones of the skeleton. We investigated the sequence of union in 15 growth plates in the limbs of about 400 specimens, representing 58 mammalian species: 34 placentals, 23 marsupials and one monotreme. We found a common general pattern of growth plate closure sequence, but one that is universal neither between species nor in higher-order taxa. Locomotor habitat has no detectable correlation with the growth plate closure sequence, but observed patterns indicate that growth plate closure sequence is determined more strongly through phylogenetic factors. For example, the girdle elements (acetabulum and coracoid process) always ossify first in marsupials, whereas the distal humerus is fused before the girdle elements in some placentals. We also found that heterochronic shifts (changes in timing) in the growth plate closure sequence of marsupials occur with a higher rate than in placentals. This presents a contrast with the more limited variation in timing and morphospace occupation typical for marsupial development. Moreover, unlike placentals, marsupials maintain many epiphyses separated throughout life. However, as complete union of all epiphyseal growth plates is recorded in monotremes, the marsupial condition might represent the derived state.

Journal of Systematic Palaeontology, 2014
We describe a new metatherian mammal, Fieratherium sorex gen. et sp. nov., found in western Argen... more We describe a new metatherian mammal, Fieratherium sorex gen. et sp. nov., found in western Argentina (Quebrada Fiera locality, southern Mendoza Province), in Late Oligocene deposits (Agua de la Piedra Formation, Deseadan age). The only known specimen is a juvenile with fragments of both dentaries, the right maxilla and a fragment of the left premaxilla with dentition. The loci and the number teeth preserved suggest a dental formula of I?3/i3, C1/c1, P3/p3, M?3/m?3. Fieratherium sorex has a convergent shrew-like appearance and a unique combination of features among metatherians and other South American mammals of Palaeogene age, including the well-known faunas of Patagonia. An analysis of its phylogenetic affinities suggests that Fieratherium is the sister-group of the Paucituberculata. As already described by other authors for several mammalian taxa, the mainly Patagonian South American Palaeogene fossil record offers little information to understanding the evolution of northern lineages. Fieratherium may represent a taxon belonging to a lineage that had its origin in Neotropical regions, so far unrecorded in the southern region of South America.

Papéis Avulsos de Zoologia, 2013
New records of Lestodelphys sp. (Marsupialia: Didelphimorphia: Didelphinae: Thylamyini) were foun... more New records of Lestodelphys sp. (Marsupialia: Didelphimorphia: Didelphinae: Thylamyini) were found at the base of Eloisa Member, Río Luján Formation (late Pleistocene) from three sites located on the right bank of the Río Reconquista, Merlo County, northeast Buenos Aires Province (Argentina). The records of these specimens are among the northernmost for Lestodelphys from the Pleistocene of Buenos Aires Province. The material consists of four lower hemimandibles. They have a combination of features suggesting either: (1) a high morphological variability in the fossil species L. juga, in which case the current diagnosis of this taxon has to be reviewed; or (2) that the validity of L. juga must no longer be considered, as already stated by previous researchers. The occurrence of Lestodelphys is used as an environmental indicator because the current species is confined to areas of arid to semiarid conditions. The presence of the new specimens in the Pleistocene of Merlo reinforces previous paleo-environmental inferences for the levels of provenance.

PlosOne, 2013
Bone microstructure reflects physiological characteristics and has been shown to contain phylogen... more Bone microstructure reflects physiological characteristics and has been shown to contain phylogenetic and ecological signals. Although mammalian long bone histology is receiving increasing attention, systematic examination of the main clades has not yet been performed. Here we describe the long bone microstructure of Xenarthra based on thin sections representing twenty-two species. Additionally, patterns in bone compactness of humeri and femora are investigated. The primary bone tissue of xenarthran long bones is composed of a mixture of woven, parallel-fibered and lamellar bone. The vascular canals have a longitudinal, reticular or radial orientation and are mostly arranged in an irregular manner. Concentric rows of vascular canals and laminar organization of the tissue are only found in anteater bones. The long bones of adult specimens are marked by dense Haversian bone, a feature that has been noted for most groups of mammals. In the long bones of armadillos, secondary osteons have an oblique orientation within the three-dimensional bone tissue, thus resulting in their irregular shape when the bones are sectioned transversely. Secondary remodeling is generally more extensive in large taxa than in small taxa, and this could be caused by increased loading. Lines of arrested growth are assumed to be present in all specimens, but they are restricted to the outermost layer in bones of armadillos and are often masked by secondary remodeling in large taxa. Parameters of bone compactness show a pattern in the femur that separates Cingulata and Pilosa (Folivora and Vermilingua), with cingulates having a lower compactness than pilosans. In addition, cingulates show an allometric relationship between humeral and femoral bone compactness.
Journal of Mammalian Evolution, 2013
South America was isolated from other continents during most of the Cenozoic, developing a singul... more South America was isolated from other continents during most of the Cenozoic, developing a singular mammalian fauna. In contrast to North America, Europe, Asia, and Africa, up to the late Neogene, the carnivore adaptive zone in South America was populated by crocodiles (Sebecidae), large snakes (Madtsoiidae), large birds (Phorusrhacidae), and metatherian mammals (Sparassodonta). Sparassodonta were varied and comprised a wide range of body masses (≈ 2-50 kg) and food habits. Their diversity decreased towards the late Miocene (Huayquerian Stage/Age) and the group became extinct in the

Ameghiniana, 2012
We report on a new dryolestoid (Mammalia, Dryolestoidea, Meridiolestida) from the Los Bastos Form... more We report on a new dryolestoid (Mammalia, Dryolestoidea, Meridiolestida) from the Los Bastos Formation (Coniacian), Neuquén Province, Patagonia, Argentina, consisting of an edentulous left dentary . The alveoli preserved suggest the presence of three incisors, one double-rooted canine, and six double-rooted postcanines (probably three premolars and three molars). Based on comparisons with previously known dentaries and isolated teeth, MCF-PVPH 412 would have been about the same size as Reigitherium Bonaparte. Among Dryolestoidea, MCF-PVPH 412 is assigned to Meridiolestida because there were probably three molars, the roots of the posterior molars are anteroposteriorly compressed, and there is no Meckelian groove. In addition, the penultimate lower premolar would be the largest in the tooth series, which is also true in other meridiolestidans. The position of the mandibular foramen, the probable presence of three premolars, and the outline of the posteroventral part of the jaw suggest affinities with the Mesungulatoidea (e.g., Coloniatherium Rougier, Forasiepi, Hill and Novaceck; Peligrotherium Bonaparte, Van Valen and Kramarz; and Reigitherium). The Coniacian specimen represents the oldest Mesungulatoidea and fills the gap in the record between the oldest South American dryolestoid (i.e., Cenomanian) and the better known Campanian-Maastrichtian taxa. The discovery of MCF-PVPH 412 in the Coniacian of Patagonia is consistent with the dryolestoid diversification during the Late Cretaceous that makes them the most abundant mammals during that period in South America.
Mastozoología Neotropical , 2012
<adolfogil@arqueologiamendoza. org>]. 3 Universidad Nacional de Cuyo.

Andean Geology, 2012
Se presentan los recientes avances en el conocimiento de la estratigrafía y la edad de las unidad... more Se presentan los recientes avances en el conocimiento de la estratigrafía y la edad de las unidades volcanosedimentarias cenozoicas expuestas en el área de sierra de Huantraico-sierra Negra y cerro Villegas (Departamento Pehuenches, provincia del Neuquén, Argentina). Los análisis estratigráficos y de los restos fósiles hallados permiten reconocer y diferenciar seis unidades litoestratigráficas: a. ʻRodados Lustrosos' (Eoceno Tardío-Oligoceno Temprano?), conformados por conglomerados gruesos texturalmente maduros (hasta 18 m de espesor) dispuestos sobre un marcado paleorrelieve labrado en sedimentitas preeocenas (fase diastrófica Incaica); b. Formación Rincón Escondido (nom. nov.) (Oligoceno Temprano-Oligoceno Tardío bajo), compuesta por depósitos fluviales de dominio psamítico asociado a un fuerte aporte piroclástico (31 m), se dispone y rellena el mismo paleorrelieve sobre el cual se sitúan los ʻRodados Lustrosos'; c. Formación Sierra Negra (nom. nov.) (Aquitaniano-Burdigaliano, Mioceno Temprano), suprayace a un nuevo paleorrelieve que llega a erosionar hasta depósitos del Cretácico Temprano (fase diastrófica Pehuenche); esta unidad puede subdividirse en el Miembro Filo Morado (hasta 250 m de basaltos alcalinos olivínicos, pirocalstitas, depósitos volcanoclásticos y sedimentarios volcanogénicos) y el Miembro Puesto Tillerías (nom. nov.) (hasta 600 m de coladas basálticas y andesíticas con intercalaciones de tobas y tufitas); d. Formación Pichi Tril (Burdigaliano tardío-Langhiano), conformada por cuerpos intrusivos de composición andesítica, cuyo emplazamiento se vincula a una deformación acaecida a fines del Mioceno Temprano (fase diastrófica Quechua); e. Formación Desfiladero Negro (Tortoniano Temprano), que conforma un conjunto de diques básicos que atraviesan al resto de la sucesión-una intrusión relacionada con deformación del Mioceno Tardío bajo; f. Formación La Tiza (Tortoniano tardío?-Mesiniano temprano, Mioceno Tardío), compuesta por hasta 85 m de depósitos conglomerádicos oligomícticos gruesos de baja madurez textural, originados a partir de la erosión de los cuerpos volcánicos y las coladas de la Formación Sierra Negra. La sucesión completa fue nuevamente deformada hacia finales del Mioceno (Mesiniano tardío). El hallazgo de restos de mamíferos fósiles en las formaciones Rincón Escondido (nom. nov.) y Sierra Negra (nom. nov.), sumadas a las dataciones radiométricas efectuadas por otros autores, han permitido calibrar geocronológicamente a las unidades volcanosedimentarias de esta región.
Journal of Human Evolution, 2012

Ameghiniana, 2011
A new Cardiomyinae rodent (Hydrochoeridae) from Arroyo Seco de la Frazada, Mendoza Province, Arge... more A new Cardiomyinae rodent (Hydrochoeridae) from Arroyo Seco de la Frazada, Mendoza Province, Argentina, is described. The new species, Caviodon cuyano sp. nov., is distinguished by its greater robustness, deeper fissures, and greater number of prisms in M3, which are derived characters within the context of Hydrochoeridae evolution. An assessment of the species of Caviodon Ameghino showed that tooth growth is allometric and occlusal morphology becomes more complex during ontogeny, as in hydrochoerines. The true diversity of this genus, when tooth postnatal ontogeny is considered, may be less than heretofore estimated. The fossil-bearing levels, the middle section of the Aisol Formation, have been considered no older than Huayquerian in age (late Miocene). The evolutionary stage of the new species suggests that the age of these sediments could be even younger, no older than early Pliocene and possibly up to late Pliocene.

Palaeontologische Zeitschrift, 2011
Astraponotus Ameghino, 1901, the only valid Mustersan (late Eocene) astrapothere, typifies the Am... more Astraponotus Ameghino, 1901, the only valid Mustersan (late Eocene) astrapothere, typifies the Ameghino's ''Capas Astraponotenses''. This taxon is traditionally interpreted as structurally ancestral to all the Oligocene-Miocene astrapotheriids. However, it was imperfectly known: only isolated teeth and very partial mandibles have hitherto been described. In this contribution we provide the first description of the skull, mandible, and complete dentition of Astraponotus based on new materials from the Gran Hondonada and other Mustersan localities in central Patagonia, Argentina. The features observed in the dentition of Astraponotus are intermediate between the Casamayoran (middle Eocene) and the Oligocene-Miocene astrapotheres in the degree of hypsodonty, reduction of the dental formula, and development of accessory occlusal elements. Concordantly, the skull retains some plesiomorphies, also observed in Trigonostylops, whereas the auditory region and the basicranium are much closer to those of Parastrapotherium, Astrapotherium, and Granastrapotherium. On the other hand, the skull of Astraponotus differs from all known astrapotheres by the disproportioned height and narrowness of the braincase, the extreme reduction of the nasals and the premaxillaries, the absence of anteorbital rim, and the reduction of the frontal region. Some of these features represent cranial specializations exactly opposite to that of Astrapotherium. These characters look astonishingly derived for an Eocene astrapothere, suggesting that extreme cranial specializations occurred independently during the evolution of the order, and that Astraponotus represents a distinctive lineage from that of Astrapotherium and other Miocene forms.

Zootaxa, 2010
A new genus and species, Patagosmilus goini, of the family Thylacosmilidae (Mammalia, Metatheria,... more A new genus and species, Patagosmilus goini, of the family Thylacosmilidae (Mammalia, Metatheria, Sparassodonta) is described here. The new taxon is based on a single specimen collected from the west margin of the Río Chico, in Río Negro Province, Argentina, from the middle Miocene Colloncuran SALMA. Until now, two formally recognized species were encompassed in the family Thylacosmilidae: Thylacosmilus atrox, from the late Miocene-late Pliocene Huayquerian to Chapadmalalan SALMAof Argentina and probably Uruguay; and Anachlysictis gracilis, from the middle Miocene Laventan SALMA of Colombia. Recognition of the Patagonian taxon, Patagosmilus, provides new anatomical data, likely to be included in future phylogenetic analyses. The overall morphology of Patagosmilus suggests that it has a more generalized anatomy than Thylacosmilus. The dental morphology suggests the new Patagonian taxon was probably closer to Thylacosmilus than Anachlysictis. Saber-tooth thylacosmilids have several autapomorphic features in the skull that differentiate them from other sparassodonts, including the delayed replacement or non-replacement of the deciduous last premolar.
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Papers by Analia M. Forasiepi