A New Extinct Genus of Cavioidea (Rodentia,Hystricognathi) from the Miocene of Patagonia (Argentina) and the Evolution of Cavioid Mandibular Morphology

The family Caviidae is represented in modern faunas by cavies and maras, whereas the family Hydrochoeridae is represented by capybaras. The evolutionary origin of these families has been related to a diversity of plesiomorphic fossil forms (recorded from the late Oligocene up to the middle Miocene) traditionally grouped in the family “Eocardiidae”. These fossil forms were included, together with Caviidae and Hydrochoeridae, within the Cavioidea s.s. (sensu stricto), because they share high crowned cheek teeth, double-hearted occlusal surface, short lower incisors, and moderate hystricognathy. Within Cavioidea s.s., caviids and hydrochoerids were interpreted as forming its crown group, because they have unique craniomandibular and dental features. In this contribution, a new taxon of Cavioidea s.s. from the middle Miocene of central Patagonia, Argentina, is described, and its phylogenetic position is determined on the basis of a morphological cladistic analysis in which “eocardiids” were included. The study permits the understanding of the sequence of appearance of characters that originated the highly divergent morphology of crown-group cavioids. The analysis of the sequence of appearance of the characters that traditionally diagnosed the crown group indicates that these changes did not occur at the same time. On the contrary, many of these features seem to have appeared at different nodes of the evolutionary history of Cavioidea s.s. The remarkably derived morphology of modern cavioids is the result of a stepwise appearance of a mosaic of evolutionary innovations that originated gradually along the history of Cavioidea during the late-middle Miocene.     

Chromosome studies in some members of the family Caviidae (Mammalia: Rodentia)

A study of the chromosomes of Cavia aperea and Galea musteloides has been made following the introduction into captivity of these two cavies from Argentina. The evolutionary relationships of the two genera have been considered, and the possible ancestry of C. porcellus from C. aperea was investigated in hybrids of the two species.     

Lung morphology in rodents (Mammalia, Rodentia) and its implications for systematics

A new nomenclature of the lung lobes and of the bronchial tree is presented, with which the lungs in 40 species of 11 rodent families are described. Whole, fixed lungs and silicone casts of the bronchial tree are tested for 23 characters, based on the distribution of lung lobes, the number and geometry of first order bronchi, the pulmonary blood supply, and lung symmetry. Ten lung morphotypes are recognized, seven of them representing one or more families: Castor type (Castoridae), Cryptomys type (Bathyergidae), Ctenodactylus type (Ctenodactylidae), Eliomys type (Gliridae), Myocastor type (Myocastoridae), Octodon type (Octodontidae and Echimyidae) and Rattus type (Sciuridae, Muridae pt. and Dipodidae). The Hydromys type is found only in Hydromys chrysogaster (Muridae), while Galea type A and B both appear in Galea musteloides (Caviidae). The data are phylogenetically analyzed by the program PAUP 4.0 using as outgroup Lagomorpha or Insectivora. On the species level, there are no well-resolved cladograms. On the family level, the cladograms do not contradict traditional rodent systematics with one exception: the Caviidae do not fall within Caviomorpha or even within the Hystricomorpha, but form a sister group to Dipodidae (Myomorpha). This appears to be a result of convergence. The lungs of Gliridae are more similar to those of Muridae than to those of Sciuridae. Included in the ingroup, Oryctolagus (Lagomorpha) forms a clade with Caviidae + Dipodidae. Thus, the "Glires hypothesis" is neither supported nor refuted.     

Major Radiations in the Evolution of Caviid Rodents: Reconciling Fossils,Ghost Lineages,and Relaxed Molecular Clocks

Background

Caviidae is a diverse group of caviomorph rodents that is broadly distributed in South America and is divided into three highly divergent extant lineages: Caviinae (cavies), Dolichotinae (maras), and Hydrochoerinae (capybaras). The fossil record of Caviidae is only abundant and diverse since the late Miocene. Caviids belongs to Cavioidea sensu stricto (Cavioidea s.s.) that also includes a diverse assemblage of extinct taxa recorded from the late Oligocene to the middle Miocene of South America (“eocardiids”).

Results

A phylogenetic analysis combining morphological and molecular data is presented here, evaluating the time of diversification of selected nodes based on the calibration of phylogenetic trees with fossil taxa and the use of relaxed molecular clocks. This analysis reveals three major phases of diversification in the evolutionary history of Cavioidea s.s. The first two phases involve two successive radiations of extinct lineages that occurred during the late Oligocene and the early Miocene. The third phase consists of the diversification of Caviidae. The initial split of caviids is dated as middle Miocene by the fossil record. This date falls within the 95% higher probability distribution estimated by the relaxed Bayesian molecular clock, although the mean age estimate ages are 3.5 to 7 Myr older. The initial split of caviids is followed by an obscure period of poor fossil record (refered here as the Mayoan gap) and then by the appearance of highly differentiated modern lineages of caviids, which evidentially occurred at the late Miocene as indicated by both the fossil record and molecular clock estimates.

Conclusions

The integrated approach used here allowed us identifying the agreements and discrepancies of the fossil record and molecular clock estimates on the timing of the major events in cavioid evolution, revealing evolutionary patterns that would not have been possible to gather using only molecular or paleontological data alone.     

Phylogenetic relationships, ecological correlates, and molecular evolution within the cavioidea (mammalia, rodentia)

A molecular phylogeny of the rodent superfamily Cavioidea was derived using two nuclear sequences (exon #10 of the growth hormone receptor gene and intron #1 of the transthyretin gene) and one mitochondrial gene (12S rRNA). A combined analysis produced a highly derived and well-supported phylogenetic hypothesis that differed from traditional taxonomy primarily in the placement of two taxa. Kerodon, traditionally included within the subfamily Caviinae with guinea pigs and its relatives, is placed sister to the family Hydrochaeridae and closely aligned with the subfamily Dolichotinae. Inclusion of Hydrochaeris within the Caviidae renders the familial classification paraphyletic. Our data further support the taxonomic separation of the families Agoutidae and Dasyproctidae. Both the molecular and traditional morphological interpretations are assessed in testing an ecological constraints hypothesis regarding social behaviors. Whereas traditional taxonomy is consistent with an environmental constraints explanation for social behavior, the molecular data suggest that phylogenetic effects may be a more important factor in the evolution of social behavior in this group. Although lineage-specific rate heterogeneity was identified in all three molecular data sets, no significant support was obtained for the metabolic rate hypothesis. However, both nuclear genes displayed patterns in accordance with the generation time hypothesis.     

The most ancient caviine rodent (Hystricognathi,Cavioidea) comes from the late Miocene of Northwest Argentina (South America)

The family Caviidae is one of the most diverse groups among South American hystricognath rodents and is represented by three main living lineages: Caviinae (cavies), Dolichotinae (maras) and Hydrochoerinae (capybaras). Caviinae includes the smaller forms of caviids represented by the extant Microcavia, Cavia and Galea. They are distributed in a wide range of environments throughout South America. In addition, three other genera from the late Miocene–Pliocene (Dolicavia, Palaeocavia and Neocavia) are recognised in high latitudes. In northwestern Argentina, the fossil forms of Caviinae have been poorly studied and for most of them there is no precise stratigraphic information. We describe and evaluate the phylogenetic affinities of the most ancient caviine from the Chiquimil Formation, Catamarca province, northwestern Argentina (9.14–7.14 ma). According to the morphological analysis of the mandibular and dental morphology and the results of the phylogenetic analysis, we assigned the new species tentatively to genus Palaeocavia. The phylogenetic position of the new species suggests an earlier origin for the lineage Palaeocavia + Cavia and for the entire clade Caviinae.     

Postnatal allometry of the skeleton in Tupaia glis (Scandentia: Tupaiidae) and Galea musteloides (Rodentia: Caviidae)--a test of the three-segment limb hypothesis

During the evolution of therian mammals, the two-segmented, sprawled tetrapod limbs were transformed into three-segmented limbs in parasagittal zig-zag configuration (three-segment limb hypothesis). As a consequence, the functional correspondence of limb segments has changed (now: scapula to thigh, upper arm to shank, fore arm plus hand to foot). Therefore, the scapula was taken into account in the current study of the postnatal growth of the postcranial skeleton in two small mammalian species (Tupaia glis, Galea musteloides). Comparisons were made between the functionally equivalent elements and not in the traditional way between serially homologous segments. This study presents a test of the three-segment limb hypothesis which predicts a greater ontogenetic congruence in the functionally equivalent elements in fore and hind limbs than in the serially homologous elements. A growth sequence, with decreasing regression coefficients from proximal to distal, was observed in both species under study. This proximo-distal growth sequence is assumed to be ancestral in the ontogeny of eutherian mammals. Different reproductive modes have evolved within eutherian mammals. To test the influence of different life histories on ontogenetic scaling during postnatal growth, one species with altricial juveniles (Tupaia glis) assumed to be the ancestral mode of development for eutherians and one species with derived, precocial young (Galea musteloides) were selected. The growth series covered postnatal development from the first successive steps with a lifted belly to the adult locomotory pattern; thus, functionally equivalent developmental stages were compared. The higher number of allometrically positive or isometrically growing segments in the altricial mammalian species was interpreted as a remnant of the fast growth period in the nest without great locomotor demands, and the clearly negative allometry in nearly all segments in the precocial young was interpreted as a response to the demand on early locomotor activity. Different life histories seem to have a strong influence on postnatal ontogenetic scaling; the effects of the developmental differences are still observable when comparing adults of the two species.     

Cranial features of dendrobatid larvae (Amphibia: Anura: Dendrobatidae)

The larval neurocranium and visceral arches of seven dendrobatid species representing four genera are described, based on cleared-and-stained and serially sectioned specimens. A variety of characters is shared by all seven species. Larval features do not substantiate the assumption of close ranoid affinities of the Dendrobatidae. Instead dendrobatid larvae share features such as the special quadripartite cartilago suprarostralis, the lack of the larval processus oticus, the presence of three foramina acustica, and the lack of a foramen perilymphaticum accessorius with many bufonoid larvae. The first of these characters is unique to bufonids, hylids, dendrobatids, and some New World leptodactylids; the other characters also occur in pelobatids and are presumably plesiomorphic for the Neobatrachia. The free proximal ends of Ceratobranchialia II and III are an autapomorphy of the Dendrobatidae supporting the monophyly of the family. Some features of the cranium are paedomorphic: low cartilago orbitalis, lack of connection between cartilage orbitalis and otic capsule (most species), and vestigal taeniae tecti. New anatomical terms are introduced. © 1995 Wiley-Liss, Inc.     

A new karyotype for <Emphasis Type="Italic">Cavia magna</Emphasis> (Rodentia: Caviidae) from an estuarine island and <Emphasis Type="Italic">C. aperea</Emphasis> from adjacent mainland

We describe a new karyotype for Cavia magna Ximenez, 1980 from an estuarine island and the karyotype of Cavia aperea Erxleben, 1777 from an adjacent mainland. The species have differences in diploid number (2n), autosomal fundamental number, quantity, and distribution of heterochromatin as dissimilar distributions of the nucleolus-organizing regions (Ag-NORs). The C. aperea karyotype has a diploid number of 64 as previously reported for C. aperea and most other Cavia species. In contrast, this new C. magna karyotype exhibits a variant diploid number of 2n = 62, considering that previous work reported a karyotype of 2n = 64 for C. magna. The discovery of a distinct diploid number within C. magna represents the first record of intra-specific chromosomal variation in a species of Caviidae. The diploid number of 2n = 62, heterochromatin quantity, Ag-NOR distributions, and inversed X chromosome from this population of C. magna are as seen in the geographically proximate (Cavia intermedia Cherem Olimpio and Ximenez; intermediate Cavy). These data provide further evidence supporting C. magna as the sister species of C. intermedia.     

A New Extinct Species of the Genus Microcavia (Rodentia,Caviidae) from the Upper Pleistocene of the Northern Basin of Uruguay,with Paleobiogeographic and Paleoenvironmental Comments

A new species, Microcavia criolloensis (Rodentia, Caviidae), from the Upper Pleistocene (Sopas Formation) of the northern basin of Uruguay (South America) is described. This species is characterised by an exclusive association of skull and dental state characters. An analysis of similarity including other species of the genus and the related genera of Caviidae was performed. The new species of Microcavia behaved in the phenogram obtained as a discrete unit closely associated to Microcavia chapalmalensis and Microcavia niata. A mosaic of habitats, ranging from open and arid or semiarid microhabitats to fluvial and riparian forests, is suggested here, mainly based on the current adaptations of the genus Microcavia and the records of several taxa of mammals in the Sopas Formation. The geographic area occupied by Microcavia in the Pleistocene was different, with fossil records located more than 500 km eastward with respect to its present distribution. This variation could have occurred as a response to environmental changes in the last interglacial and glacial cycles.     

Ontogeny and cranial morphology of the tympanic region of the Tupaiidae, with special reference to Ptilocercus

The structure of the tympanic region of the skull of Ptilocercus lowii was studied in an embryo of 30 mm crown-rump length and in 5 osteocrania. As in Tupaia, the anterior wall of the bulla of Ptilocercus is not completed by a tympanic process of the alisphenoid, contrary to earlier reports. Ptilocercus resembles Tupaia in the following derived characters. The ventral wall of the tympanic cavity is formed by a rostral entotympanic and by a caudal tympanic process of the petrosal. The entotympanic develops in primary connection with the tubal cartilage. The tympanic aperture of the auditory tube is bordered by the entotympanic. The ring-shaped tympanicum is covered by the entotympanicum and is aphaneric. The musculus tensor tympani is lacking. Among mammals, these characters can be regarded as synapomorphic for the Tupaiidae, that is, to have been present in the common ancestor of the two subfamilies. From the evidence of the tympanic region, the Tupaiidae, therefore, form a monophyletic group. Besides these synapomorphies, there are remarkable differences between Ptilocercus and Tupaia in the structure of the bulla. In Ptilocercus the bulla is smaller and less pneumatized than in Tupaia. An anterior intrabullar septum, present in Tupaia, is lacking in Ptilocercus. The epitympanic wing of the alisphenoid is smaller in Ptilocercus than in Tupaia. A lateral prefacial commissure of the tegmen tympani is present in Ptilocercus, but absent in Tupaia. The caudal tympanic process of the petrosal is larger in Ptilocercus than in Tupaia. These characters are autapomorphic for the Ptilocercinae and for the Tupaiinae, respectively. They demonstrate that the auditory bulla of Ptilocercus and that of Tupaia have evolved independently to a considerable extent. An early phylogenetic separation of their respective ancestors seems likely. The tympanic region of the skull provides no evidence for close relationships of the tree shrews to the primates or to any other eutherians. The classification of the Tupaiidae in a separate order, Scandentia, is supported.     

Respiration at high altitudes, phosphate-protein interaction: the sequence of hemoglobins from guinea pig and dromedary (author's transl)

The sequence of the main hemoglobin component of the guinea pig (Cavia aerea f. porcellus, Caviidae) and that of the hemoglobin of the dromedary (Camelus dromedarius, Camelidae) is given. The sequence is obtained automatically by the sequenator using the quadrol and the propyne programme. The sequence of the alpha-and beta-chains of the guinea pig is compared with that of the human hemoglobin; the sequence of the dromedary in comparison to the Ilama shows in the alpha-chains five amino acid exchanges, in the beta-chains there are only two exchanges in beta 2 and beta 76. Beta 2 in dromedary is the P2-glycerate contact histidine. This sustains the interpretation of the high altitude respiration of the Ilama as mutation beta2His leads to Asn.     

Glaresidae, archaeopteryx of the Scarabaeoidea (Coleoptera)

Abstract. Evidence is presented that Glaresidae are the most primitive livinj scarabaeoid group and as such represent the sister group of the rest of the Scarabaeoidea. This is based on a review of the states of seventy-two morphologica characters. The plesiomorphic states of many characters are unique to the Glaresidae or are shared with other primitive scarabaeoids; seven may be synapomorphic with other groups and two are autapomorphic for the family.     

Reexamination of the morphological evidence for the cohort Epitheria (Mammalia,Eutheria)

Novacek and co-workers recognized a monophyletic clade Epitheria, comprising all eutherians except edentates and the extinct palaeoryctoids, on the basis of two synapomorphies: a stirrupshaped stapes and a foramen ovale enclosed within the alisphenoid. To evaluate this phylogenetic hypothesis, we reexamined the distributions of stapedial morphologies and positions of the foramen ovale across Recent and extinct mammals and nonmammalian cynodonts. The states and distributions of the stapes and forament ovale characters used by Novacek and coworkers were modified by recognizing two stapedial characters (one relating to shape of the crura, the other to the nature of the foramen) and a single, multistate foramen ovale character (within, behind, and lateral to the alisphenoid). The taxon-character matrix used by Novacek (1989, 1992b), substituting our amended stapedial and foramen ovale characters and adding several previously unscored extinct taxa and three new characters, was subjected to a series of PAUP manipulations. Identified among the most parsimonious trees were three major topologies for the base of Eutheria: (1) a polytomy including an Edentata/Ungulata clade, (2) a polytomy with Edentata and Ungulata as separate clades, and (3) Edentata and (when included) Palaeoryctoidea as the successive outgroups to a monophyletic Epitheria. We conclude that topology 2 best reflects the current state of knowledge. An edentate/ungulate clade is supported by three characters (from the mastoid region and subarcuate fossa); however, other morphological studies require modification of the distributions of these characters in xenarthrans and bassal ungulates, thereby eliminating support for this clade. In nearly all manipulations, obtaining a monophyletic Epitheria required that one or two steps be added to the most parsimonious trees. When a monophyletic Epitheria was obtained, it was supported by a triangular stapes and, in some trees, the reappearance of a stapedial artery (lost earlier at the level of Recent therians) and a transpromontorial internal carotid artery. In the most parsimonious trees, a foramen ovale within the alisphenoid was an equivocal synapomorphy of Recent therians or cutherians, and a stapes with strongly convex crura (our state closest to the stirrup-shaped state of Novacek and co-workers) appeared independently within various eutherian lineages. The reduction or loss of the stapedial foramen was identified as an independent event in monotremes and within marsupials and various eutherian lineages.To whom correspondence should be addressed.     

The location of eggs retained in the oviducts of mares

The oviducts of 24 mares were examined to determine the site of retention of unfertilized eggs. The ampullary-isthmic junction regions of 42 of the 48 oviducts were serially sectioned and examined histologically. The remaining parts of the oviducts were flushed and the flushings searched microscopically. Of 45 eggs located, 40 were in the sectioned segments of 24 oviducts and only 5 were in the flushings. All but one of the sectioned segments contained prominent masses of material obstructing the lumen, but these were apparently not the direct cause of egg retention since eggs were found on both sides of them.     

The occurrence of Cavia tschudi (Rodentia,Caviidae) in the Southwest of Salta Province,Argentina

The first record of Cavia tschudi Fitzinger, 1857 for the subandean area of Salta Province, Argentina, is herein presented.

This species presently inhabits the subandean area of the provinces of Tucumán and Jujuy. The materials are composed of two mandibles coming from an archaeological site, which had been occupied ca. 1480 AD, at 2300 m a.s.l.