Cecilia C. Morgan

Phylogeny and Evolutionary Patterns of South American Octodontoid Rodents

Octodontoidea is the most diverse clade of hystricognath rodents, and is richly recorded in South America since at least the Oligocene. A parsimony-based morphological phylogenetic analysis of a wide range of extant and extinct octodontoids recovered three major clades, here recognised as Echimyidae, Octodontidae, and Abrocomidae. Taxa previously assigned to Echimyidae or Octodontoidea incertae sedis are here interpreted for the first time as early representatives of Ctenomyinae (Octodontidae), Octodontinae or Abrocomidae. Based on our results, we estimate the divergence of octodontoid families and subfamilies to have occurred during the Late Oligocene, which is consistent with molecular estimates, but older than previous inferences based on the fossil record. Contrary to previous suggestions, we show the first appearances of modern members of Abrocomidae, Octodontinae and Ctenomyinae to be distinctly decoupled from the origin of these clades, with different stages in the evolutionary history of octodontoids seemingly following distinct phases of palaeoenvironmental change. Depending on the phylogenetic pattern, fossils from the stage of differentiation bear evolutionary information that may not be provided by crown groups, thus highlighting the unique and important contribution of fossils to our understanding of macroevolutionary patterns.

Morphological Diversity of the Humerus of the South American Subterranean Rodent Ctenomys (Rodentia, Ctenomyidae)

Abstract Humeral variation associated with digging ability in the subterranean rodent Ctenomys was analyzed through 6 functionally significant indexes. The humerus of some extinct and living species was slightly more specialized than that of fossorial octodontoids †Actenomys and Octodon, whereas it was highly specialized in some living species. The constant occurrence of greater epicondyles suggests a hierarchical pattern in the acquisition of scratch-digging specializations. A possible relationship between humeral morphological diversity and environments is preliminarily discussed.

Carpal-metacarpal specializations for burrowing in South American octodontoid rodents

Among the ecomorphologically diverse Octodontoidea rodents, fossorial habits are prevalent in Ctenomyidae and Octodontidae and occur in some members of Echimyidae. To detect traits linked to scratch‐digging, we analyzed morpho‐structural variation in the carpus and metacarpus of 27 species of extinct and living octodontoids with epigean, fossorial and subterranean habits. Within a context of relative morphological uniformity, we detected the following specialized traits in the burrowing Clyomys (Echimyidae), Spalacopus (Octodontidae), Ctenomys and †Eucelophorus (Ctenomyidae): broad shortened carpus, robust metacarpals, markedly broad and short metacarpal V, and predominance of ray III (mesaxony, incipient in Spalacopus). In addition, the specialized subterranean Ctenomys presented an enlarged scapholunar in extensive contact with the unciform, and with a complex‐shaped proximal articular surface. These features are interpreted as responses to mechanical requirements of scratch‐digging, providing greater carpal rigidity and resistance to direct forces exerted during the digging stroke. In Ctenomys, the radius‐scapholunar joint restricts movement at wrist level. The phylogenetic distribution of traits shows that the most derived carpal and metacarpal morphologies occur among subterranean octodontoids, also possessing important craniodental adaptations, and supports the hypothesis that the acquisition of digging specializations would have been linked to increasing burrowing frequency in some lineages. Nevertheless, octodontoids with less morphological specializations have metacarpal modifications advantageous for digging, suggesting that scratch‐digging specialization preceded the acquisition of tooth‐digging traits, in agreement with the general claim that scratch‐digging is the primary digging strategy in burrowing mammals.

Contrasting Phylogenetic and Diversity Patterns in Octodontoid Rodents and a New Definition of the Family Abrocomidae

Octodontoidea is the most species-rich clade among hystricomorph rodents. Based on a combined parsimony analysis of morphological and molecular data of extinct and extant species, we analyze the history of South American octodontoids and propose ages of divergence older than interpreted so far. Early Abrocomidae are recognized for the first time, and a new definition of the family is provided. Traditionally accepted fossil-based times of origin for the southern clades are reinterpreted as later stages of differentiation markedly uncoupled from the origin, differentiation implying specializations for open environments as shown in a morphospace of skull variation. Origin of crown groups is also strongly uncoupled from origin of clades as a consequence of extinction of deep lineages. In the resulting diversity pattern of modern southern clades of octodontoids, the combination of greater disparity, less content of evolutionary history, and lower taxonomic diversity, compared to their northern counterparts, appears at first counterintuitive. We propose that primary components of diversity derived from evolutionary transformation or anagenesis, on the one hand, and from cladogenesis and extinction, on the other, should not be considered associated, or at least not necessarily. Certain patterns of relationships between these distinct components could be driven by environmental dynamics. Like environments, octodontoid diversity would have been more stable in northern South America, whereas in the south, both strong adaptive change and extinction would have been triggered by emerging derived environments.

Adaptive diversity of incisor enamel microstructure in South American burrowing rodents (family Ctenomyidae, Caviomorpha)

The aim of this study was to analyse the morphofunctional and adaptive significance of variation in the upper incisor enamel microstructure of South American burrowing ctenomyids and other octodontoid taxa. We studied the specialized subterranean tooth‐digger †Eucelophorus chapalmalensis (Pliocene – Middle Pleistocene), and compared it with other fossil and living ctenomyids with disparate digging adaptations, two fossorial octodontids and one arboreal echimyid. Morphofunctionally significant enamel traits were quite similar among the species studied despite their marked differences in habits, digging behaviour and substrates occupied, suggesting a possible phylogenetic constraint for the Octodontoidea. In this context of relative similarity, the inclination of Hunter–Schreger bands, relative thickness of external index (EI) and prismless enamel zone were highest in †Eucelophorus, in agreement with its outstanding craniomandibular tooth‐digging specialization. Higher inclination of Hunter–Schreger bands reinforces enamel to withstand high tension forces, while high external index provides greater resistance to wear. Results suggest increased frequency of incisor use for digging in †Eucelophorus, which could be related to a more extreme tooth‐digging strategy and/or occupancy of hard soils. Higher external index values as recurring patterns in distant clades of tooth‐digging rodents support an adaptive significance of this enamel trait.

Systematics and evolutionary significance of the small Abrocomidae from the early Miocene of southern South America

Abstract Octodontoidea is the most species-rich clade among hystricomorph rodents, and has a fossil record going back to at least the late Oligocene. Affinities of fossils previous to the late Miocene differentiation of the extant families Abrocomidae, Echimyidae and Octodontidae are controversial, essentially because these fossils may share few apomorphies with modern species. In fact, pre-late Miocene representatives of Abrocomidae had not been recognised until very recently. Here we revise the early Miocene genus Acarechimys, originally assigned to Echimyidae, and alternatively to stem Octodontoidea or to Octodontidae. A systematic and parsimony-based phylogenetic analysis of the species traditionally included in Acarechimys showed that this genus is part of stem Abrocomidae. These results are primarily supported by morphology of the mandible and lower molars. Acarechimys is here restricted to three species, A. minutus, A. pulchellus and Acarechimys pascuali sp. nov., while another species, A. constans, is here transferred to a new abrocomid genus. The remaining species were nested within Octodontidae. According to these results, Abrocomidae might have been as diverse as its sister clade Octodontidae-Echimyidae during the late Oligocene–early Miocene. Extinction of this diversity would have resulted in marked loss of evolutionary history, with extant abrocomids being currently restricted to late-diverged euhypsodont representatives.

Morphology of the lower deciduous premolars of South American hystricomorph rodents and age of the Octodontoidea

Abstract We analyse the crest homologies of lower deciduous premolars (Dp4) of South American caviomorphs in a comparative context including other hystricomorphs and under a dynamic topological criterion. An hexalophodont pattern in which the three anteriormost crests are assumed to be the anterolophid, metalophulid I and metalophulid II, respectively, is proposed as ancestral morphology. Simplified pentalophodont morphologies would have resulted from loss of the metalophulid I, and distinct transformation pathways are recognised especially for Octodontoidea and Cavioidea. A basal octodontoid morphological pattern can be recognised in the fossil record from as early as the middle Eocene, supporting that an initial divergence among major clades of caviomorphs had already occurred by this time. Programs for identifying potentially useful dental characters in phylogeny and taxonomy of caviomorphs need to be revised. Interpreting lophids as dynamic components in the morpho-functional variation of molars, rather than as static landmarks, is central for understanding dental evolution in these rodents.