Un nuevo genero para Habrothrix angustidens y Akodon serrensis (Rodentia, Cricetidae): de nuevo palentologia y neontologia se encuentran en el legado de Lund.
During the last 2 decades, several species of sigmodontine rodents named by Winge (1887) based on material from the rich paleontological deposits of Lagoa Santa area (Minas Gerais, Brazil) were recognized as senior synonyms of living entities (e.g., Voss, 1993; Voss and Carleton, 1993; Musser et al., 1998; Pardinas and Teta, 2011, 2013). These findings, partly based on earlier observations (e.g., Avila-Pires, 1960; Massoia and Fornes, 1965; Hershkovitz, 1966) helped identify redundancies in sigmodontine classification. In addition, these studies provided new data for understanding the tempo and mode of evolution of this rodent group and its past diversity (Voss, 1988; Voss and Myers, 1991). In fact, of 25 species recorded by Winge (1887) in Quaternary deposits (summarized in Voss and Myers, 1991:table 4), only Habrothrix angustidens, Habrothrix clivigenis, Oxymycterus cosmodus, Oxymycterus talpinus, and Calomys anoblepas, still remain as presumptively extinct forms (Pardinas and Teta, 2013).
During the course of our studies of the collections of Lund housed in the Danish city of Copenhagen, once again paleontology meets neontology. Habrothrix angustidens Winge, 1887, a putative fossil species recorded from the area of Lagoa Santa, Minas Gerais, Brazil, appears morphologically undistinguishable from Akodon serrensis Thomas, 1902, a living form from the Atlantic Forest of Argentina and Brazil. The present paper documents this synonymy--first found and generously communicated to us by Robert Voss--and proposes a new genus for both forms. The need for a new genus was uncovered by the phylogenetic analyses of D'Elia (2003) and D'Elia et al. (2003), who employed mitochondrial and nuclear DNA sequences and broad taxonomic coverage and recovered A. serrensis as sister to Thaptomys and not as part of the genus Akodon. That hypothesis is here morphologically and karyologically supported and expanded through descriptions and comparisons against related genera that conform the Akodon Division (sensu D'Elia, 2003) of the tribe Akodontini (see also D'Elia and Pardinas, 2015).
MATERIALS AND METHODS
In this report we studied fossil and recent specimens (see Appendix 1) housed at the following institutions: American Museum of Natural History, New York, USA (AMNH); The Natural History Museum, London, UK (BMNH); Centro Nacional Patagonico, Puerto Madryn, Argentina (CNP); Field Museum of Natural History, Chicago, USA (FMNH); Coleccion Nacional de Mastozoologia, Museo Argentino de Ciencias Naturales "Bernardino Rivadavia," Buenos Aires, Argentina (MACN); Museu Nacional, Rio de Janeiro, Brazil (MN); Museu de Zoologia, Departamento de Biologia Animal, Universidade Federal de Vinosa, Vinosa, Minas Gerais, Brazil (MZUFV); Universitets Zoologisk Museum, Copenhagen, Denmark (ZMK); and voucher specimens CIT and BIO that will be deposited at Museu de Zoologia, Universidade de Sao Paulo, Sao Paulo, Brazil. Fossil specimens are represented by craniodental fragments; recent specimens are preserved as dry skins, skulls, skeletons and individuals stored in fluid. Most of the studied material came from several localities in southeastern Brazil, while a few are from Misiones Province in northeastern Argentina (see an annotated gazetteer of recording localities in Supplementary Material S1).
Anatomical descriptions and terminology follow Reig (1977), Carleton (1980), Voss (1988), and Carleton and Musser (1989). Standard external measurements (length of head and body; length of tail; length of hind foot with and without claw; length of ear from notch; all in millimeters) and weight (in grams) were transcribed from the specimen labels or museum catalogs as originally recorded by collectors. Craniodental dimensions were taken using a digital caliper and recorded to the nearest 0.01 mm and follow definitions provided by Weksler and Geise (1996) and Voss et al. (2002). They were, arranged in alphabetical order: breadth across M1s, breadth across occipital condyles, breadth of braincase, breadth of incisive foramina, breadth of M1, breadth of rostrum, breadth of zygomatic plate, condylo-incisive length, depth of incisor, height of skull, height of mandible, least interorbital breadth, length of bulla, length of diastema, length of incisive foramina, length of mandible, length of orbital fossa, length of palatal bridge, length of rostrum, length of maxillary molar row, and zygomatic breadth. Measured specimens were sorted into age categories following toothwear classes defined for Zygodontomys by Voss (1991). Because our aim is to provide a general characterization of a new genus, we adopt a univariate statistics approach based on mean values for large samples of adult individuals (toothwear classes 3-5).
Karyotypes from selected specimens (see Appendix 1) were prepared in the field. Chromosomes at metaphases were obtained via in vitro culture (culture of bone marrow grown in Dulbecco's MEM with 10% fetal bovine serum and colchicine [10.sup.-6] M), following incubation in KCl 0.075M solution at 37[grados]C by 40 minutes, centrifuged, fixed in Carnoy solution (methanol: acetic acid, 3:1), dropped onto clean slides and air-dried (Geise, 2014). Conventional coloration with Giemsa 5% was used to observe diploid (2n) and fundamental (FN, excluding sexual chromosomes) numbers and chromosome morphology variation; 50 metaphases of each specimen were analyzed with a Nikon Eclipse 50i photomicroscope under 1000 x magnification. Karyotypes were compared with literature reports. For intergeneric chromosomal comparisons, Deltamys sp. (CIT 944, male, Esmeralda, Rio Grande do Sul, Brazil), Necromys lasiurus (CIT 952, male, Rio Formoso, Pernambuco, Brazil), Thalpomys lasiotis (BIO 288, male, Brasilia, Distrito Federal, Brazil), and Thaptomys nigrita (CIT 331, male, Iguape, Sao Paulo, Brazil), metaphases were also obtained from fibroblast cell cultures derived from ear biopsies grown in Dulbecco's modified Eagle's medium supplemented with 20% fetal calf serum. For all species, GTG-banding was based on routine cytogenetic procedures, and FISH with whole chromosome probes was conducted following Ventura et al. (2009) using the Akodon paranaensis chromosome set as paints. FISH with telomeric probes was also performed only on T. lasiotis using Telomere PNA FISH Kit/FITC (K 5325-Dako) following the manufacturers' protocol for the purpose of cytogenetic comparisons. After FISH, the slides were counterstained with DAPI diluted in Vecta-shield (Vector) and analyzed using a fluorescence microscope (Zeiss Axiophot) equipped with software for image capture (MetaSystems).
RESULTS AND DISCUSSION
Habrothrix angustidens was described by Winge (1887:28-29) based on fossil remains recovered by the Danish naturalist Peter Lund in Lagoa Santa cave deposits in the first half of the XIX century. Winge (1887) also recognized or named 5 other forms in Habrothrix. One of these, H. clivigenis was referred to Akodon by Trouessart (1898) and remains a putative fossil form. The other 4, H. cursor, H. orycter, H. lasiotis, and H. lasiurus are now allocated, respectively, to the recent genera Akodon, Thaptomys, Thalpomys, and Necromys. Interestingly, all these genera belong to the tribe Akodontini. This highlights the accurate systematic perception of Winge, despite working mostly with fragmentary fossil material representing a large number of taxonomic groups.
Winge (1887:28) emphasized the similarities between H. angustidens with H. cursor but also was impressed by the absence of a capsular projection of the incisors and the smaller size of the former. He specifically highlighted the peculiar condition of the dentition of H. angustidens, and most remarkably, mentioned the difficulty of distinguishing the enamel from the dentine on the occlusal surfaces. Winge (1887:pl. II, figure 9) illustrated 2 remains belonging to H. angustidens, both from Lapa da Serra das Abelhas; an anterior skull fragment and a right mandible. Shortly after, Trouessart (1898:536) listed H. angustidens under Akodon; this action must be viewed as a taxonomic act without major implications because this author transferred to this genus all the Habrothrix species of Winge plus a wide array of sigmodontine forms.
In 1902 Oldfield Thomas received from Brazil a large mammal collection made by Alphonse Robert in the Serra do Mar of the Brazilian state of Parana. Thomas (1902:61) recognized one sigmodontine as new and described it as Akodon serrensis with type locality at "Roca Nova ... at an altitude of about 1000 meters" (Thomas, 1902:59). Thomas also received in the same collection specimens of A. cursor, and he directly compared both species. Akodon serrensis was characterized basically on coloration traits and said to be intermediate in size between A. azarae (= "arenicola" in the original work) and A. cursor. Thomas (1902:61) also noted "molars very large in proportion, as large as those of the much bigger species A. cursor; " however, he did not make any mention about their morphology.
Miranda Ribeiro (1905:188) added a new name to the nominal forms allied to A. serrensis. Based on a single female from Retiro de Ramos (also known as Retiro do Ramos), Rio de Janeiro, Brazil, he erected [Hesperomys] (A.) serrensis var. leucogula, a subspecies characterized by a more olive dorsal coloration, a yellowish-ochraceous belly, and a narrow pure white band in the lower part between the chin and the anterior limbs.
The XXth century passed without major systematic or nomenclatural modifications. While H. angustidens was basically forgotten, A. serrensis--including H. (A.) serrensis leucogula--was maintained under Akodon from the time of the catalog of Gyldenstolpe (1932) up to the more recent treatments (e.g., Musser and Carleton, 2005; Patton et al., 2015). Hershkovitz (1990a:14) ranked leucogula as a valid and different species from serrensis, including both forms in an informal group that he named Akodon mollis size class. Other researchers addressed morphological and chromosomal considerations of A. serrensis populations, especially in southern Brazil. In part, these studies were triggered by a misunderstanding involving several species of the A. cursor group (sensu Smith and Patton, 2007) and A. serrensis. This contorted history, which started when Liascovich and Reig (1989) erroneously referred one Argentinean Akodon specimen to A. serrensis and reported for it a diploid complement of 44, was fully resolved by Christoff et al. (2000; see also Pardinas et al., 2003; Gonsalves et al., 2007). It is of interest to transcribe a paragraph of Liascovich and Reig (1989:394) in order to reflect the understanding of A. serrensis at that time: "Akodon serrensis is similar to A. cursor in size and color pattern, and the two species are difficult to distinguish based on external characters. Fresh skins of A. serrensis have a conspicuously more ochraceous venter, especially evident in the anal region; however, even this distinction must be confirmed by examination of a large series of specimens. The skulls of both species also are similar in having long, narrow rostra, moderately inflated frontal regions, short zygomatic plates, a reduced interparietal, and elongated palatal foramina that reaches the level of the protocone of M1. The main differences between the two species are: the nasals are longer and project farther behind the maxillo-frontal suture in A. cursor; the zygomatic plate is much deeper in A. serrensis; the palate is shorter in A. cursor, its posterior border terminating anterior to the level of the alveoli of M3; the molars are narrow, elongated, and larger in A. serrensis; and the mandible in A. cursor is slender and elongated with a more posteriorly projecting condylar process and a more inflated capsular projection (root of incisor)." It is difficult to know how influential it was that one of the specimens used by Liascovich and Reig (1989) was, in reality, an Akodon paranaensis (see Pardinas et al., 2003; Gonsalves et al., 2007). Despite having the holotype of A. serrensis in hand, these authorities failed to distinguish these species. This necessitates that we call attention to the similarity of A. serrensis and other species of the cursor group of Akodon (e.g., cursor, montensis, paranaensis).
However, a new scenario emerged from molecular studies. D'Elia et al. (2003:358) explicitly called attention to the generic position of A. serrensis as follows: "A novel finding of our study with respect to akodontine systematics is the placement of Akodon serrensis well outside the Akodon clade ... A. serrensis appears as sister to the monotypic genus Thaptomys, and this clade is sister to the genus Necromys ... The phylogenetic position of Akodon serrensis is surprising because to the best of our knowledge nobody has cast doubt upon the inclusion of A. serrensis within the genus Akodon ... We will not make any taxonomic judgment about this topology until further studies ... are carried out. However, a combined analysis of nuclear and mitochondrial DNA sequences points in the same direction" In fact, D'Elia (2003:319) revisited this topic in a combined cytb and IRBP analysis "Ten extra steps are needed to force A. serrensis within the Akodon clade. Akodon serrensis appears to be sister to the monotypic genus Thaptomys (node 18), although this relationship is weakly supported (JK<50%, BS = 1). This clade is sister to the clade Thalpomys-Necromys. Further studies, including morphological comparisons and the sequencing of additional specimens, are needed to determine if A. serrensis should be placed on its own genus or in Thaptomys" Despite the relevance, these findings were basically ignored in the chromosomal research developed with Akodon species in southeastern Brazil (e.g., Ventura et al., 2006; Hass et al., 2008; Bonvicino, 2011). More recently, analyzing cytb over a taxonomically diverse sample of Akodon, Smith and Patton (2007:848) corroborated the separate position of serrensis although they highlighted its alternative relationship to Akodon + Deltamys rather than Thaptomys.
The taxonomic hypothesis that H. angustidens and A. serrensis were congeners was achieved by R. Voss after a restudy of the type material of both forms. According to him "The type series of Habrothrix angustidens consists of subfossil cranial fragments and disassociated hemimandibles from several caves in the vicinity of Lagoa Santa ... All of this material qualifies as syntypes because Winge did not designate a holotype ... Altogether, this material ... preserves the entire upper and lower dentition, the left zygomatic plate, diastema, palate, interorbital region, and mandibles. We compared this material side-by-side with a complete topotypical cranium of Akodon serrensis (ZMUC 1257) and found these two taxa to be phenotypically indistinguishable. In addition to qualitative resemblances, the few measurements that we were able to take from the lectotype of angustidens all fall within the range of variation seen in the topotypic series of serrensis. The only possible taxonomic conclusion is that angustidens and serrensis are synonyms" (R. Voss, in litt. November 4, 2008; Supplementary Material S2).
Family Cricetidae Fischer, 1817
Subfamily Sigmodontinae Wagner, 1843
Tribe Akodontini Vorontsov, 1959
Castoria, gen. nov.
Figs. 2-5; Tables 1-3
Habrothrix sensu Winge (1887:28), part, not Habrothrix Wagner, 1843
Akodon sensu Thomas (1902:61), part, not Akodon Meyen, 1833
Hesperomys sensu Miranda Ribeiro (1905:188), part, not Hesperomys Waterhouse, 1839
Type species.--Habrothrix angustidens Winge, 1887.
Etymology.--In homage to Castor Cartelle, a contemporany paleontologist (born in Ourense, Spain in 1938) studying Quaternary Brazilian mammalian fauna who effectively resurrected in Brazil the genius and figure of Peter W. Lund (see Cartelle, 2002). Continuous efforts of Cartelle over the last 4 decades have enlarged our knowledge of biotic evolution in tropical areas through the excavation, cleaning, sorting, and study of a huge quantity of osteological material recovered from karstic caves in northeastern Brazil.
Geographic distribution.--Known from southeastern Brazil and the northeastern tip of Argentina, roughly between 20[grados] to 30[grados]S and mostly restricted to an elevational range above 800 m (Fig. 1; Supplementary Material S1).
Chronological distribution.--Late Pleistocene? -Recent. Fossils were found in 5 cave deposits Lapa do Capao Secco, da Cerca Grande, a chamber near Escrivania, Quebra Chavelha, and Serra das Abelhas- in the area of Lagoa Santa, Minas Gerais, Brazil (Winge, 1887). Absolute radiometric dates performed on fossils and breccias excavated by Lund from Lagoa Santa indicate an array of ages covering at least the last 0.5 MA (e.g., Auler et al., 2006).
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Contents.--Tentatively, the genus is considered to be composed by one species, Castoria angustidens (Winge, 1887), nov. comb. Neither morphology nor metrics (Table 1) allow us to distinguish, at the specific level, the anterior skull portion that represents H. angustidens from those of living populations traditionally ascribed to A. serrensis. In fact, an exploratory multivariate analysis based on principal components placed the lectotype of H. angustidens within the morphospace of recent Brazilian specimens of A. serrensis (Supplementary Material S3). Minor differences between the fossil H. angustidens fragment--particularly the narrow condition of the molar toothrow--and recent A. serrensis individuals are here attributed to age variation. Given the broad geographic range of this genus and modest morphometric segregation (results not shown; see also Kosloski, 1997), the variation of Castoria is a topic that deserves further study. The names serrensis Thomas, 1902 and leucogula Miranda Ribeiro, 1905 are available.
[FIGURE 3 OMITTED]
Morphological diagnosis.--Rodents of the subfamily Sigmodontinae (sensu Reig, 1980) with the following combination of characters: small-medium size (head-and-body length ~ 95 mm; condylo-incisive length ~ 25 mm); tail shorter than head-and-body (~ 85% of head-and-body length); manual and pedal digits with short claws covered by long and sparse ungual tufts; pelage soft and long; skull robust with pointed rostrum, broad and somewhat flat interorbital region with frontal borders moderately divergent backwards, and enlarged braincase without patent crests; broad, anteriorly rounded, and slanting zygomatic plate; zygomatic arches with their lower border parallel with respect to the alveolar plane; long incisive foramina, anteriorly constricted, with expanded palatal process of premaxillary; short, broad, and flat bony palate; broad mesopterygoid fossa with anterior margin rounded; alisphenoid strut present; tegmen tympani overlapping squamosal with the participation of a distinct posterior suspensory process of the latter bone; carotid arterial circulation pattern primitive; bullae short and rounded; upper incisors opisthodont; molars noticeable large in relation to the cranium proportions, terraced, enlarged, and moderately hypsodont; main molar cusps arranged in opposite pairs; M1 without anteromedian flexus with large procingulum and marked trilophodont pattern; M3 with cylindrical outline; m1 with anteromedian flexid weakly present and procingulum anterior-posteriorly compressed; m1-2 with small ectolophids present and mesolophids probably fussed to metaconids; length of m2 slightly shorter than those of m1; length of m3 subequal than those of m2; lower incisor alveolus without distinct capsular process on lateral dentary surface; gall bladder present; stomach unilocular and hemiglandular; diploid number (2n) = 46 and fundamental number (FN) = 46.
Morphological description.--The following description is based on representative samples from localities in Minas Gerais and Rio de Janeiro states in Brazil and 2 individuals from Argentina. It aims to provide a general characterization of the genus. Additional morphological details and measurements can be found in Hershkovitz (1998:215-218).
[FIGURE 4 OMITTED]
Castoria includes small-medium sized sigmodontine rodents (Tables 1 and 2) characterized by a darker general coloration above and below, short mystacial vibrissae, short claws, and a moderately long tail although shorter than the combined length of head and body. The dorsal and ventral colors are subtly delimited with weak countershading (Supplementary Material S4). The skins exhibit thick, soft fur (guard hairs ~ 9 to 12 mm); the cuticle pattern of guard hairs was described as narrow and leaf-shaped; medullar pattern of these same hairs were a mixture of alveolar and striped types, with three to four layers of cells of irregular shape (Silveira et al., 2013). The dorsal fur is dark yellowish- to orangish-brown; the ventral surface is washed with bright buff or tawny over dark-gray hair bases. The eyes are small. The mystacial vibrissae are short, not extending posteriorly much beyond the base of the pinnae; genal vibrissae are present. The ears are medium in size, rounded, and internally and externally covered by brown hairs. The tail is dark brown, unicolored, and sparsely haired, covered by sets of 3 somewhat hispid hairs per scale, each dorsal hair covering 2-3 scale-rows; apical tuft absent. The manus and pes in Castoria are covered with short hairs, with their basal portion brown and apical portion whitish. The claws, basally open on the pes, are inconspicuous and dorsally covered by several whitish or darker ungual hairs slightly surpassing their apices. The plantar surface is naked, pigmented, and slightly squamated (sensu Pacheco, 2003). The pes has 6 pads, 4 interdigital and 2 plantar, the thenar and hypothenar; the thenar pad is enlarged and narrow; the hypothenar pad is subequal in size to the 1st interdigital pad; dIII and dIV are the largest, subequal in size and showing a slightly syndactylous condition (Fig. 2).
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The skull in dorsal view is characterized by a flat, broad appearance with a pointed rostral region and a rounded braincase (Fig. 3). The nasals are pointed and usually slightly projecting beyond the anterior margin of the premaxillary (Fig. 4A). The rostrum is flanked by shallow but distinct zygomatic notches (Fig. 4B); the frontal sinuses are inflated and the interorbital region is broad, with rounded supraorbital margins in young and adult specimens (older animals develop small, sharp, convergent postorbital edges). The zygomatic arches are delicate, widest across the squamosal roots; they converge anteriorly and their midportions run parallel to the alveolar plane; the jugal is well developed (Fig. 4C). The braincase is large, smooth, and globose; the lambdoid crests are developed only in older individuals. The interparietal varies in size and form, being typically rhomboidal and compressed. The zygomatic plate in lateral view has a straight anterior margin that aligns vertically or slopes slightly backward; the origin of the superficial masseter is marked by a scar (not a tubercle) that is just below and slightly behind the anteroventral margin of the zygomatic plate. The foramina of the carotid circulation consist of pattern 1 (sensu Voss, 1988); a thick alisphenoid strut is present, and the subsquamosal fenestra is large (Figs. 4G and 4H). The incisive foramina are long (extending between the M1 anterocones) and characterized by a constricted anterior portion (at or near the premaxillary-maxillary sutures) and a markedly inflated premaxillary process (Fig. 4E); the palate is broad and uncomplicated by keels or deep lateral grooves; and the mesopterygoid fossa extends anteriorly to or between the M3s (Fig. 4F). There are no large posterolateral palatal pits, just tiny foramina flanking the mesopterygoid fossa. The mesopterygoid roof is perforated by very narrow but long sphenopalatine vacuities (Fig. 4F). The auditory bullae are small and flask-shaped, with broad bony Eustachian tubes.
The mandible is relatively robust, moderately low and elongated; the lower border of the ramus bends upward and backward behind the level of the m2 and descends again behind the m3. The mental foramen opens on the dorsolateral surface of the diastema. The alveolus of the lower incisor is contained in a ridge, not a distinct process, on the lateral surface of the mandible behind the base of the coronoid process. The angular process is pointed and bent upwards (Fig. 4I).
The upper incisors are opisthodont in orientation, with yellow-orange enamel bands, straight or very slightly inclined in frontal view and sharp, not inclined medially in ventral view (cf. Voss, 1988:376-377). The exterior edge of the upper incisors shows a thick enamel band; the dentine lake is straight (sensu Steppan, 1995:figure 10). The molars are very narrow in proportion to their length, with a marked unilateral tubercular hypsodonty (sensu Hershkovitz, 1962:89; Fig. 4D). They are also characterized by the opposite pattern of the main cusps, strongly bi-level (subadults) to terraced (adults) coronal surface (sensu Hershkovitz, 1962:86), simplification, and a tendency to retain little occlusal detail even in young animals; the cusps and lophs wear rapidly away, exposing large dentine basins (Fig. 5). The latter feature, shared with other akodontines (e.g., Akodon, Oxymycterus), makes it necessary to study a series of individuals of different ages in order to adequately assess dental variation. The upper molar tooth-rows are parallel or slightly convergent backwards in adult and old-adult specimens. The M1 is typically composed by 3 subequal "lobules" characterized by central basins and abrupt posterior walls inclined backwards. The procingulum lacks any trace of an anteromedian flexus and shows a well-developed surface occlusal area. The paracone "lobule" is the largest because it likely involves the mesoloph plus the paralophule; a trace of the latter is present as an acute posterior edge. The proto- and hypoflexus are subequals and scarcely developed, while para- and metaflexus are moderately directed backwards, with their internal margins reaching the middle line of the molar; the mures flanked by these flexus are particularly enlarged, lingually disposed, and longitudinally oriented. No evidence of a free posteroloph was detected in any studied specimen. The M2 shows a tetralophodont pattern with anteroloph and posteroloph obsolescent in young individuals to bilophodont in adults. The metacone "lobule" is smaller than the paracone one. The M3 is certainly complex in juveniles, with an internal enamel ring of equivocal identity (mesofossette?) and reduced hypoflexus and posterior lobe. In subadult to adult individuals, M3 acquires a cylindrical outline without internal accidents. We documented a deformation of the occlusal figure of the upper molars by the differential wear of labial and lingual sides as a consequence of age; whereas the labial portion is partially eroded to the coronal base, the lingual remains with persistent striae and thicker enamel. Lower molars are characterized by occlusal simplicity in subadult and adult individuals, a compressed procingulum mostly fused with the remainder portion of the m1, and a large m3 subequal to or even exceeding the m2 in length.
On the basis of 9 individuals, Castoria has 13 thoracic ribs, 13 thoracic vertebrae, 6 lumbar vertebrae, and 29-30 caudal vertebrae. The sacrum involves 3 subequal and squared sacral vertebrae. The neural spine of the axis is enlarged anteriorly-posteriorly and is fan-shaped. The neural spine of the 2nd thoracic vertebra is enlarged, twice or more as long as nearby spines. The caudal vertebrae lack hemal arches but hemal processes are present starting between the 3rd and 4th caudal vertebrae, becoming most pronounced between the 5th and 6th, and then decreasing in size. Gross stomach morphology in Castoria conforms with the unilocular-hemiglandular type (sensu Carleton, 1973:figure 2), a widespread condition in sigmodontine rodents. The bordering fold bisects the stomach on a line from the incisura angularis to a point opposite it on the greater curvature (Supplementary Material S5). Animals from Rio de Janeiro, Sao Paulo, and Misiones indicate that a gall bladder is present (cf. Geise et al., 2004; this paper).
Cytogenetics.--The specimens of Castoria angustidens cytogenetically analyzed (Appendix 1) presented the same karyotype, with 2n = 46 and FN = 46 (Supplementary Material S6); the autosomes comprising 21 acrocentric pairs and 1 small metacentric pair (the smallest of the complement). The X chromosome is a medium-sized acrocentric, and the Y one of the smallest of the complement, also acrocentric. This karyotype was previously recorded for specimens collected in states of Rio de Janeiro (Geise et al., 1998; Olifiers et al., 2007), Parana (Barros-Battesti et al., 1998; Hass et al., 2008), Santa Catarina (Testoni et al., 2012), Minas Gerais (Moreira et al., 2009), and Espirito Santo (Bonvicino et al., 2002). Cytogenetic information is lacking for Argentinean populations; the animal referred to A. serrensis by Liascovich and Reig (1989) and characterized as having 2n = 44 (FN = 44) was correctly reidentified as Akodon paranaensis by Christoff et al. (2000).
Phylogeny.--Phylogenetic analyses of combined molecular markers (e.g., D'Elia, 2003; D'Elia et al., 2003; Smith and Patton, 2007) showed that Castoria belongs to the Akodon Division of the tribe Akodontini, a well-supported clade composed by the recent genera Akodon, Deltamys, Necromys, Podoxymys, Thalpomys, and Thaptomys (D'Elia, 2003; Leite et al., 2015). Within the Akodon Division, its position varies depending upon the molecular markers used, taxonomic sampling, and the type of analysis conducted. In fact, in those studies with moderate taxonomic coverage (e.g., excluding Vilela et al., 2014), Castoria was recovered as sister to a clade including Akodon + Deltamys (Smith and Patton, 2007; Ventura et al., 2013; Abreu et al., 2014; Pardinas et al., 2014b), but elsewhere, it appears as sister to Thaptomys (e.g. D'Elia, 2003; D'Elia et al., 2003), or as sister to Deltamys (Fabre et al., 2012; Leite et al., 2015). Despite its uncertain position, no study has recovered Castoria within or as sister to the genus Akodon. This finding supports the hypothesis that this lineage deserves generic recognition.
Comparisons.--The fact that A. serrensis remained allocated as a species of Akodon for more than a century is a clear signal of the strong morphological similarity between these 2 genera. In addition, some character traits that were used as evidence of the distinctiveness of some genera of the Akodon Division, such as the morphology of the male accessory glands (see Reig, 1987), were--and remain--unknown for A. serrensis. Castoria is smaller than Akodon cursor or Akodon montensis and can be externally distinguished by its shorter manual claws and overall darker coloration. The relative size of molar toothrows, noticeably larger in Castoria than in Akodon (as Gyldenstolpe, 1932:104 highlighted), the general morphology of the interorbital region--flat and divergent posteriorly in Castoria and smooth and hourglass-shaped in Akodon, and the marked differences in the development of the premaxillary process as seen through the openings of the incisive foramina--more inflated and extended in Castoria than in Akodon, are the main cranial features separating these genera (Fig. 6). However, there are more trenchant differences between Castoria and Akodon in molar morphology. Castoria displays a conspicuous pattern of opposite main cusps in contrast to Akodon where these elements are moderately alternating; in addition, the former lacks the anteromedian flexus that typically characterizes the M1 procingulum in Akodon species (Table 3; Fig. 7). An additional important trait to distinguish Castoria from species of the cursor group of Akodon, which are sympatric with Castoria, is the morphology of the incisive foramina. In Castoria, the foramina are constricted anteriorly, anterior to the premaxillary-maxillary suture. In Akodon, the margins are not constricted, and the lateral margins of the foramina are convex and wider medially, resembling parentheses and showing no changes in curvature (Fig. 6; see also Supplementary Material S7). However, this degree of morphological differentiation is considered as interspecific variation within the genera Handleyomys (cf. Voss et al., 2002:22) and Drymoreomys (Percequillo et al., 2011:371).
Thaptomys appears as the sister taxon for Castoria in some molecule-based phylogenies (e.g., D'Elia, 2003). Both share weak pelage countershaping and inflation of the premaxillary process. However, morphological differences are strong. Castoria has a moderately long tail and rostrum, contrasting with the shorter condition of both in Thaptomys. Castoria has a well-developed upper free margin of the zygomatic plate whereas Thaptomys does not. Castoria lacks the defined capsular projection of the lower incisor root characterizing the mandible of Thaptomys. A gall bladder is present in Castoria and absent in Thaptomys. Molar morphology in Thaptomys is very similar to that in Akodon, so that Castoria differs from Thaptomys in all the respects noted earlier regarding Akodon.
[FIGURE 6 OMITTED]
Although Castoria differs in numerous aspects with respect to Deltamys, both share weak countershading, a moderately long tail, and the general "soricine" external aspect. Their skulls are easily distinguished by the pointed rostrum and broad interorbital region of Castoria versus the comparatively shortened rostrum and narrow interorbital region of Deltamys. In addition, Castoria has a broad zygomatic plate and an alisphenoid strut whereas Deltamys has a narrow plate without free upper margin and lacks the alisphenoid strut. In keeping with the molecular phylogenetic hypothesis of Smith and Patton (2007), which recovered Castoria close to Delta mys, these genera show many similarities in molar occlusal morphology (Fig. 7). In particular, both have the main cusps arranged in opposite pairs, the tendency to a trilophodont M1 in adults, and an anteriorly-posteriorly compressed procingulum that coupled with its coalescence to the metaconid produce on the m1 a nearly bilophodont pattern. The GTG-banding patterns of Castoria chromosomes allowed the identification and pairing of all homologues (Supplementary Material S6), and therefore, comparisons with the GTG-banded chromosomes of Akodon paranaensis (2n = 44 and FN = 44; reported in Ventura et al., 2009). Chromosome painting using A. paranaensis (APA) revealed that a tandem rearrangement, in which chromosome 7 of A. paranaensis is homologous to two chromosome pairs (16 and 21) of Castoria angustidens, explains their differences in 2n and FN. Meanwhile, analyzed species of Akodon share the sequences of APA 7 conserved as one chromosome (see Ventura et al., 2009). Comparisons of karyotypes among C. angustidens and other closely related akodontines (Deltamys, Necromys, Thalpomys and Thaptomys), showed that two groups besides Akodon can also be delimited with respect to APA 7. The first group unites Castoria and Thaptomys, both with homologous sequences of chromosome 7 of A. paranaensis reorganized into two acrocentric chromosomes (data not shown).
The second group comprises Deltamys sp., Necromys lasiurus and Thalpomys lasiotis, which present the 2 chromosome pairs homologous to APA 7 (as Castoria and Thaptomys), but also presents an exclusive association of chromosomes APA 7 and APA 8. Castoria presents a distinct pattern and nature of heterochromatin, distributed in large and positive pericentromeric blocks after telomeric FISH with (TTAGGG) n sequences as probes (Ventura et al., 2006, as Akodon serrensis). Thalpomys lasiotis also exhibits a pericentromeric region rich in (TTAGGG)n-like sequences (for CBG-banding patterns see Yonenaga-Yassuda et al., 1987, as Akodon reinhardti, and Andrade et al., 2004); this homoplasic feature in light of currently understood phylogenetic relationships, suggests that accumulation of the (TTAGGG) n-like repetitive sequences could lead to generic-level differentiation in the Akodontini.
[FIGURE 7 OMITTED]
Natural History.--Castoria has been recorded in at least 44 localities, almost all in the Atlantic Forest of Brazil (41) and Argentina (3), encompassing 7 ecoregions (Supplementary Material S1). Little is known about its natural history. The principal vegetation where Castoria is recorded is humid mountain forest, including those with Araucaria augustifolia (Supplementary Material S8). The elevational range for Castoria collection localities is from 300 to 2100 m (correlation between latitude and elevation was r = 0.5; Supplementary Material S9). Usually Castoria is interpreted as an element of montane fauna (Gonsalves et al., 2007; Abreu et al., 2014). Paglia et al. (2012) reported the diet of this rodent as insectivorous/omnivorous, although the basis for this assessment is unknown. Morphometric analysis of the appendicular skeleton suggests that this species uses terrestrial locomotion (Coutinho et al., 2013). Vieira and Monteiro-Filho (2003) captured this mouse exclusively on the ground. Miranda Ribeiro (1905) indicates that Castoria inhabits "galerias subterraneas," that according to Voss (1993:35) possibly correspond to "shallow tunnels under matted roots and moss" and piles of litter in thickets of Chusquea. The species was the most frequently trapped small mammal in the Parque Estadual do Desengano, a disturbed montane forest (~1000-1425 m elevation) in the northern portion of Rio de Janeiro (Modesto et al., 2008). Castoria angustidens was also the dominant species among recorded sigmodontines in an elevational transect of Parque Estadual da Serra do Brigadeiro from 1200 to 1800 m, particularly at middle to upper elevations (1450-1800 m; Moreira et al., 2009). In Parque Nacional de Itatiaia, it was trapped in montane forests and "campos de altitude" up to ~2100 m (Geise et al., 2004). Captive animals fed mainly on pulp of fruits and also on small to medium-sized seeds of a wide array of plant species from the Atlantic Forest (Vieira et al., 2003). A rich assemblage of parasites, including Acari, Anoplura, and Siphonaptera has been recorded from Brazilian individuals (Barros-Battesti et al., 1998; Linardi and Guimaraes, 2000; Moraes et al., 2003). Teixeira et al. (2014) recorded natural spillover infection between this species and the Jabora strain of hantavirus, whose primary host is Akodon montensis. According to these authors, spillover infection in secondary hosts, including C. angustidens and A. paranaensis, may play an important role in maintaining the hantavirus sylvatic cycle during periods of low prevalence in primary hosts.
Biogeography.--Castoria is distributed in southeastern Brazil, from Espirito Santo to Rio Grande do Sul, and in Misiones, northeastern Argentina. Its distributional range overlaps those of other akodontine rodents, such as Blarinomys breviceps, Brucepattersonius spp., and Thaptomys nigrita. Unlike those taxa and other Atlantic Forest sigmodontines, Castoria is much restricted to a narrow coastal fringe at relatively high elevations (500-1600 m) in well conserved forest remnants. Most of the recorded localities are located within Serra do Mar coastal forests and Araucaria Moist forests ecoregions (Olson et al., 2001; Pardinas et al., 2015). The Serra do Mar is recognized as one of the more remarkable centers of endemism in South America, with evidence from plants (especially of the families Myrtaceae, Lauraceae, Melastomataceae, and Orchidaceae), birds, butterflies, amphibians, and mammals (Da Silva, 2000a and references therein). Araucaria Moist forests spread from middle-level plateaus of ~500 m to the high slopes (1600 m) of the Serra da Mantiqueira. Among 352 plant species recorded in this region, 47 (13.3%) are endemic; this moist forest is also recognized as an important endemic bird area (Da Silva, 2000b). Castoria has been recorded from the Bahia Coastal and Interior Forests and at ecotones with the Alto Parana Atlantic forests. At a regional scale, at least other 2 genera, Delomys and Juliomys, have similar distributions, with populations mostly restricted to these same ecoregions (see Pavan and Leite, 2011; Gonqalves and Oliveira, 2014). The genera Delomys and Juliomys comprise both widely distributed species as well as more elevationally restricted taxa. This pattern highlights the importance of montane topographies in the evolution of sigmodontines of the Atlantic Forest (Gonqalves and Oliveira, 2014) and identifies this area as a major center of diversification of this subfamily (Smith and Patton, 1999).
Conservation.--C. angustidens was categorized as "Least Concern" by the IUCN. Several arguments were used to justify this placement, including its wide distribution, presumed large populations, its presence in a number of protected areas (e.g., Parque Estadual do Desengano, Parque Estadual da Serra do Brigadeiro, Parque Nacional de Itatiaia, Parque Nacional da Serra dos Orgaos, Parque Nacional da Bocaina), and because "it is unlikely to be declining at nearly the rate required to qualify for listing in a threatened category" (http://www. iucnredlist.org/details/full/756/0). However, it is likely that some populations of this species have been negatively affected by habitat destruction and forest fragmentation.
Remarks.--Our systematic hypothesis involves fossil (H. angustidens) and recent materials (A. serrensis). This kind of taxonomic work is obviously limited owing to the anatomical incompleteness and absence of genetic information for the fossils. However, as discussed several times revisiting the Lund collections (Voss, 1993; Voss and Carleton, 1993; Pardinas et al., 2008), fossils of Lagoa Santa often preserve enough information to make confident inferences regarding both genus and species (Supplementary Material S2 and S10).
Winge's description of H. angustidens was explicitly based on multiple specimens, none of which was designated as type (Winge, 1887:28-29). Therefore, all of those specimens must be considered syntypes in accordance with Article 72.1.1 of the Code (ICZN). We designate as lectotype of H. angustidens--according to Article 74.1--the cranial anterior fragment illustrated by Winge (1887:II, figure 2); this fossil was recently numbered in ZMK collections as 1/1845:13246 and is associated with Winge's handwritten label saying "Habrothrix angustidens Tegnet [figured] ?Lapa da Serra das Abelhas" (Supplementary Material S10). In addition, a photograph of this specimen was published in the catalog of Hansen (2012:58-59).
The lectotype of H. angustidens comes from "Lapa da Serra das Abelhas," and like other fossils of the same provenience (e.g., the types of H. orycter or H. clivigenis) is fragmentary and stained yellowish by superficial concretions. Several of the caves worked by Lund--including Serra das Abelhas--were destroyed by human activities over the last 170 years; however, a few, such as Cerca Grande located about 16 km NW Lagoa Santa city, where H. angustidens was also recorded, still remain. It is noteworthy that, despite extensive surveys of modern mammal fauna made by Lund around Lagoa Santa, Castoria was not detected (cf. Voss and Myers, 1991). In fact, the nearest record for the species is from Fazenda da Neblina in the Parque Estadual da Serra do Brigadeiro (Moreira et al., 2009), ca. 200 km SE of Lagoa Santa. The Lapa da Serra das Abelhas sigmodontine assemblage includes 8 taxa without recent records of occurrence near Lagoa Santa, including Castoria, Delomys, Euryoryzomys, Juliomys, Sooretamys, and Thaptomys (Winge, 1887; Pardinas and Teta, 2013). The taxonomic structure of this assemblage is characterized by elements presently restricted to Atlantic Forest in mixture with Cerrado species, a non-analogue fauna probably triggered by cooling episodes during the Late Pleistocene (Voss and Carleton, 1993; Cartelle, 1999; Pardinas and Teta, 2013).
The cursory indication by Hershkovitz (1990a:14) concerning the synonymy of A. serrensis and Akodon pervalens Thomas, 1925, a poorly known form with type locality at Carapari (Tarija, Bolivia), must be dismissed. A. pervalens--judging from examination of its holotype and paratype--looks like a true Akodon, phenotypically similar to species of the A. cursor species group as earlier suggested by Myers (1989). Although the holotype of A. pervalens is old (see Jayat et al., 2007:210) and some of its cranial and dental features (e.g., broad interorbital region, nasals tapering forward, absence of anteromedian flexus on M1) recall those of older A. serrensis, these are only general resemblances produced by aging (Supplementary Material S7).
Castoria belongs to a clade of akodontine genera historically understood as a part of Akodon (cf. Reig, 1987), which has recently been reorganized into several genera (for a review, see D'Elia and Pardinas, 2015). Morphological differences among Akodon, Deltamys, Thalpomys, Thaptomys, Necromys, and Castoria, as well as more controversial forms such as Microxus, are typically subtle (Table 3). This seems to be a signature of the Akodon Division, one that prevented the recognition of these different genera for most of the XXth century.
Since the 1970s, several distinctive lineages formerly subsumed in Akodon--directly or under subgeneric combinations--have been progressively elevated to full generic rank. Paradigmatic cases were Abrothrix (now included in a different tribe) and Thaptomys, but also more recently Thalpomys and Deltamys. Reig (1987:358) anticipated these changes when he wrote "I tentatively recognize a central genus Akodon, subdivided into five subgenera ... I do not deny, however, that more detailed and comprehensive further studies could eventually elevate some of these taxa to generic level" These generic partitions within Akodon were possible because of the accumulation of morphological and karyological information and were then consolidated by DNA-based phylogenetic analyses (e.g., Hershkovitz, 1990b; Smith and Patton, 1993, 1999, 2007; Geise et al., 2001; D'Elia, 2003; D'Elia et al., 2003; Pardinas et al., 2005; Jayat et al., 2010). It is interesting to point out that at least two putative genus-rank groupings remain subsumed under Akodon despite clear morphological indications about their singularities; these are Microxus and Hypsimys (see Reig, 1987:359-360; Ventura et al., 2000:497; Voss, 2003:20-21). Perhaps their retention in Akodon is more the result of historical inertia than of unequivocal phylogenetic support (see Patton et al., 1989; Smith and Patton, 2007). An example of such progress is the recent finding that Akodon bogotensis and Akodon latebricola are neither Akodon nor Akodontini (Alvarado-Serrano and D'Elia, 2013). The naming of Neomicroxus demonstrates that the content of Akodon, as presently understood containing about 38 living species (Pardinas et al., 2015), requires detailed morphological and molecular scrutiny.
Castoria adds a new piece to Akodontini historical biogeography. In fact, the exclusion of the Abrotrichini (see Smith and Patton, 1999), the inclusion of Scapteromyini (see Smith and Patton, 1999; D'Elia et al., 2005), and the elevation to genera of several erstwhile subgenera or species (Deltamys, Gyldenstolpia; D'Elia et al., 2003; Pardinas et al., 2008) have reshaped our vision about this diverse tribe. Of the 15 genera currently recognized (see Patton et al., 2015), three are Atlantic Forest endemic, Blarinomys, Brucepattersonius, and Thaptomys. Castoria offers further evidence for a potential area of endemism in southeastern Brazil, a picture also supported by several genera of non-akodontine sigmodontine rodents (Smith and Patton, 1999; Pardinas et al., 2014a; Prado et al., 2014) and echimyid rodents (Upham and Patterson, 2015:Fig. 1).
Recibido 30 setiembre 2015. Aceptado 13 abril 2016. Editor asociado: B Patterson
Robert Voss (AMNH) deserves credit for recognizing the congeneric status between H. angustidens and A. serrensis. He helped us not only by providing this information but also the original notes of his study and provided casts of H. angustidens that are now housed in CNP collections. Rob's selfless attitude is an example that better communication among researchers is possible. J. Oliveira (MN) kindly provided on long-term loan a large sample of specimens collected at key localities in Brazil. C. Galliari, J. Notarnicola, M. Lareschi, and R. Robles shared field efforts in order to obtain Argentinean material for study. A. Bezerra photographed for us the holotype of A. serrensis. M. Lareschi provided parasitological data. Valuable specimens were freely accessed thanks to the curators D. Flores (MACN), J. Patton (MVZ), and B. Patterson (FMNH); K. L. Hansen (ZMK) kindly answered our request about Lund's collections in Copenhagen. This paper was partially funded by Agencia Nacional de Promocion Cientifica y Tecnologica (PICT 2008-547 and 2014-1039 to UFJP) and Consejo Nacional de Investigaciones Cientificas y Tecnicas PIP 6179 (to UFJP), Centro Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq), Coordenado de Aperfeicoamento de Pessoal de Nivel Superior (CAPES), Fundado Carlos Chagas Filho de Amparo a Pesquisa do Estado do Rio de Janeiro (FAPERJ), World Wildlife Fund (WWF) and UERJ/ Prociencia (to LG), and Fundado de Amparo a Pesquisa do Estado de Sao Paulo, FAPESP (BP.PD 2009/54300-0 to KV). Specimens from Rio de Janeiro state were collected under collecting license of IBAMA and ICMBio (license # 598633). This contribution was originally triggered by a suggestion of G. D'Elia who, after a decade of shared efforts and contributions of phylogenetic tips, declined to be an author; this manuscript also received significant input from P. Teta, who participated in several aspects of museum study and also in the masterful production of the figures. We are deeply grateful to all these individuals and institutions. Finally, it is important to mention that the manuscript benefited from three reviews, reflecting the zeal of the editors to ensure its quality; and also, the associate editor was deeply committed to ensure the linguistic correctness employed in the text. This is GEMA's (Grupo de Estudios de Mamiferos Australes) contribution #10.
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Studied specimens belong to the following mammal collections: AMNH, American Museum of Natural History, New York, USA; BMNH, Natural History Museum, London, UK; CNP, Coleccion de Mamiferos del Centro Nacional Patagonico, Chubut, Argentina; FMNH, Field Museum of Natural History, Chicago, USA; MACN, Coleccion Nacional de Mastozoologia, Museo Argentino de Ciencias Naturales "Bernardino Rivadavia," Buenos Aires, Argentina; MN, Museu Nacional, Rio de Janeiro, Brazil; MZUFV, Museu de Zoologia, Departamento de Biologia Animal, Universidade Federal de Vinosa, Vinosa, Minas Gerais, Brazil; ZMK, Universitets Zoologisk Museum, Copenhagen, Denmark. Symbols denote specific studies carried out on each specimen (* = measured; k = karyotyped).
Akodon montensis (n = 46): Argentina, Misiones, Balneario Municipal de Aristobulo del Valle sobre el arroyo Cuna Piru, 10 km NN W Aristobulo del Valle (CNP 1785, 1786, 1789, 1790, 1794, 1795, 1796, 1797, 1798, 1799, 1800, 1802, 1804, 1805, 1807, 1808, 1809, 1810, 1811, 1812, 1813, 1814, 1815, 1818, 1819, 1821, 1825, 1829, 1831, 1833, 1835, 1838, 1844, 1845, 1847), Reserva Privada de Usos Multiples Guarani (CNP 4243, 4244, 4245, 4247, 4248, 4249, 4250, 4251, 4252, 4253, 4254). Castoria angustidens nov. comb. (n = 123): Argentina, Misiones, Reserva Privada de Vida Silvestre Urugua-i (MACN 22246), RP 2, 6 km NE Arroyo Paraiso (CNP 449 *). Brazil, Minas Gerais, Parque Estadual da Serra do Brigadeiro, Fazenda da Neblina (MZUFV 520 *, 551 *, 1076, 1148 *, 1150 *, 1151 *, 1153 *, 1154 *, 1155 *, 1156 *, 1215 *, 1220 *, 1234, 1242, 1266 *, 1281 *, 1298 *, 1630, 1635, 1653, 1655, 2555; UFMG 1858 *, 1859 *, 1860 *, 1861 *, 1862 *), Parque Estadual da Serra do Brigadeiro, Serra das Caberas (MZUFV 1243 *, 1248 *, 1250 *, 1267 *, 1280 *, 1286 *, 1287 *, 1294 *, 1295 *), Lapa da Serra das Abelhas (ZMK 1/1845:13246 * [lectotype of H. angustidens]; CNP 4267 [lectotype: cast of upper molar row], CNP 4268 [paralectotype: cast of lower molar row]); Parana, Roca Nova (BMNH 22.214.171.124 *-126.96.36.199 *, ZMK 1257 *); Rio de Janeiro, Parque Nacional da Serra dos Orgaos, Vale das Antas (MN 69807 * (k), 69808 *, 69809 *, 69810 *, 69811 *, 69812 * k, 69813 *, 77085 *, 77099 * (k), 77100 *, 77101 *, 77102 *, 77104 *, 77105 *, 77108 *, 77111 *, 77112 *, 77113 *, 77116 *, 77117 * (k), 77119 *, 77121 *, 77122 *, 77123 *, 77078 * (k), 77079 * (k), 77081 *, 77082 *, 77084 * (k), 81130 *, 81131 *, 81133 *, 81134 *, 81135 *, 81136 *, 81137 *, 81138 *, 77086 *, 77087 * (k), 77088 *, 77090 *, 77091 *, 81144 *, 77096 *, 81147 *, 81148 *, 81150 *, 81151 *, 81152 *, 81153 *, 81154 *, 81155 *, 81156 *, 81157 *, 81158 *, 81159 *, 81160 *, 81161, 81162 *, 81163 *, 81164 *, 81165 *, 81166 *, 81168 *, 81169 *, 81170 *, 81171 *, 81172 *, 77097 * (k), 81175 *), Sitio Xitaca, Debossan, Nova Friburgo (MN 35912 *, 35920 *, 35921 *, 35923 *, 35924 *, 35926 *), Parque Nacional do Itatiaia (MN 69803 *), Retiro de Ramos (MN 2513 * [holotype of H. serrensis leucogula]), Visconde de Maua (MN 69804 *); Sao Paulo, Fazenda Intervales (FMNH 143275).
Deltamys kempi (n = 6): Argentina, Buenos Aires, La Balandra (CNP 581, 893, 3086, 3087, 4141), General Lavalle (CNP 2377).
Necromys lactens (n = 5): Argentina, Jujuy, Barcena, ca. 3 km S sobre RN 9 (CNP 1486), Abra de la Cienaga Negra (CNP 1487), San Francisco (CNP 3032, 3037, 4124).
Necromys lasiurus (n = 8): Argentina, Santiago del Estero, Estacion Experimental INTA "La Maria" (CNP 2358); Chaco, 5 km NW Puerto Las Palmas (CNP 3036, 3049); Misiones, Estancia Santa Ines (CNP 3038, 3041, 3043, 3045, 3048).
Necromys obscurus (n = 5): Argentina, Buenos Aires, Arroyo de las Brusquitas (CNP 891, 2380, 3039, 3047), Estacion Experimental INTA "Balcarce" (CNP 3053).
Thaptomys nigrita (n = 23): Argentina, Misiones, Balneario Municipal de Aristobulo del Valle sobre el arroyo Cuna Piru, 10 km NN W Aristobulo del Valle (CNP 1788, 1791, 1816, 1826, 1827, 1828, 1832, 1839, 1846, 1929, 1931), Refugio Mocona (CNP 1938, 3008, 3010, 3011), Asentamiento Aborigen Kaaguy Poty, 1 km NNW interseccion RP 7 y arroyo Cuna Piru (CNP 1979, 2370, 2371, 2372), Posada Las Malvinas (CNP 3526), Reserva Privada de Vida Silvestre Urugua-i (CNP 3875). Brazil: Minas Gerais, Parque Estadual da Serra do Brigadeiro, Fazenda da Neblina (MZUFV 1110, 1112)
SUPPLEMENTARY MATERIAL ONLINE
S1. Gazetteer of the recording localities for the genus Castoria. https://www.sarem.org.ar/wp-content/uploads/2016/06/SAREM_MastNeotrop_23-1_Pardinas-sup1.doc
S2. Study notes on Habrothrix angustidens and Akodon serrensis made by R. S. Voss in November, 2008. https://www.sarem.org.ar/wp-content/uploads/2016/06/SAREM_MastNeotrop_23-1_Pardinas-sup2.doc
S3. Principal component analysis of all age classes of Brazilian specimens of Castoria, including the lectotype of Habrothrix angustidens. https://www.sarem.org.ar/wp-content/uploads/2016/06/SAREM_MastNeotrop_23-1_Pardinas-sup3.doc
S4. External appereance of Castoria angustidens, nov. comb. https://www.sarem.org.ar/wp-content/uploads/2016/06/SAREM_MastNeotrop_23-1_Pardinas-sup4.doc
S5. Gastric morphology of Castoria angustidens, nov. comb. https://www.sarem.org.ar/wp-content/uploads/2016/06/SAREM_MastNeotrop_23-1_Pardinas-sup5.doc
S6. Chromosomes of Castoria angustidens, nov. comb. https://www.sarem.org.ar/wp-content/uploads/2016/06/SAREM_MastNeotrop_23-1_Pardinas-sup6.doc
S7. Comparison of the holotypes of Akodon pervalens and A. serrensis. https://www.sarem.org.ar/wp-content/uploads/2016/06/SAREM_MastNeotrop_23-1_Pardinas-sup7.doc
S8. Habitat of Castoria angustidens, nov. comb. at Itamonte, Brazil. https://www.sarem.org.ar/wp-content/uploads/2016/06/SAREM_MastNeotrop_23-1_Pardinas-sup8.doc
S9. Joint distribution of latitude and elevation for Castoria angustidens, nov. comb. collection localities. https://www.sarem.org.ar/wp-content/uploads/2016/06/SAREM_MastNeotrop_23-1_Pardinas-sup9.doc
S10. Lectotype of Castoria angustidens, nov. comb. (ZMK 1/1845:13246). https://www.sarem.org.ar/wp-content/uploads/2016/06/SAREM_MastNeotrop_23-1_Pardinas-sup10.doc
S1.-Gazetteer of locality records for the genus Castoria.
This gazetteer summarizes geographical and ecological information about Castoria collection localities -arranged by increasing latitude- and habitats. Except where indicated, voucher specimens have been identified by direct examination and/or karyotype.
CJ--Field catalogue of Erika Hingst-Zaher.
CNP--Coleccion de Mamiferos, Centro Nacional Patagonico, Puerto Madryn, Chubut, Argentina.
MACN--Coleccion Nacional de Mastozoologia, Museo Argentino de Ciencias Naturales "B. Rivadavia," Buenos Aires, Argentina.
MN--Museu Nacional, Rio de Janeiro, Brazil.
MZUFV--Museu de Zoologia, Departamento de Biologia Animal, Universidade Federal de Vicosa, Vicosa, Minas Gerais, Brazil.
MZUSP--Museu de Zoologia da Universidade de Sao Paulo.
P--Field catalogue of Yves Sbalqueiro.
UFMG--Colecao de Mamiferos, Universidade Federal de Minas Gerais.
UFRGS--Colecao de Mamiferos, Universidade Federal de Rio Grande do Sul.
1. Brazil, Minas Gerais, Lagoa Santa, Lapa da Serra das Abelhas (~19[degrees]37'S, 43[degrees]53'W; exact locality coordinates unknown); Cerrado (type locality of Habrothrix angustidens, see Winge 1887).
2. Brazil, Minas Gerais, Simonesia, Mata do Sossego (20[degrees]7'S, 42[degrees]1'W, 575 m); Atlantic Forest--humid mountain forest, secondary forest (MN 35910).
3. Brazil, Espirito Santo, Ibitirama, Parque Nacional do Caparao (20[degrees]29'S, 41[degrees]43'W, 970 to 2890 m), including Pedra Rocha trapping station listed by Hershkovitz (1998) and Bonvicino et al. (2002) and Arrozal, Segredo, and Terreirao included by us (this paper); specimens housed at MN and FMNH; Atlantic Forest--mountain field, mountain scrub, humid mountain forest, secondary forest.
4. Brazil, Minas Gerais, Alto do Caparao, Parque Nacional do Caparao (20[degrees]26'10.4"S, 41[degrees]48'2.8"W, 970 to 2890 m) including Arrozal, Cachoeira Bonita, Centro de Visitantes, Pico da Bandeira, Segredo, Terreirao, Tronqueira, Vale Encantado, and Vale Verde trapping stations listed by Hershkovitz (1998) and Bonvicino et al. (2002) and Alfa, Beta, and Casa de Julio included by us (this paper); specimens housed at MN and FMNH; Atlantic Forest--mountain field, mountain scrub, humid mountain forest, secondary forest.
5. Brazil, Espirito Santo, Castelinho (20[degrees]31'S, 40[degrees]59'W, 1200 m); Atlantic Forest--humid mountain forest, secondary forest (MN 32631).
6. Brazil, Minas Gerais, Fervedouro, Parque Estadual da Serra do Brigadeiro, Fazenda da Neblina (20[degrees]40'0"S, 42[degrees]31'15"W, 1400 m); Atlantic Forest--mountain field, mountain scrub, humid mountain forest, secondary forest (MZUFV 1220).
7. Brazil, Minas Gerais, Tombos, Mata do Banco (20[degrees]56'28"S, 42[degrees]06'28"W, 467 m); Atlantic Forest--humid mountain forest (MZUFV 1900).
8. Brazil, Minas Gerais, Posses (21[degrees]41'S, 46[degrees]10'W, 1117 m); Atlantic Forest--humid mountain forest (UFMG 1855).
9. Brazil, Rio de Janeiro, Santa Maria Magdalena, Parque Estadual do Desengano (21[degrees]45'S, 41[degrees]41'W, 1425 m); Atlantic Forest--mountain field, mountain scrub, humid mountain forest (Modesto et al. 2008).
10. Brazil, Sao Paulo, Serra da Fartura, Sao Joao da Boa Vista (21[degrees]55'S, 46[degrees]43'W, 1142 m); Atlantic Forest--humid mountain forest (Moraes et al. 2003).
11. Brazil, Minas Gerais, Itamonte, Parque Nacional de Itatiaia, Brejo da Lapa (22[degrees]20'54.4"S, 44[degrees]41'38.2"W, 2100 m); Atlantic Forest--humid mountain forest with Araucaria (MN 48027).
12. Brazil, Rio de Janeiro, Visconde de Maua, Fazenda Marimbondo (22[degrees]21'38.5"S, 44[degrees]34'14.6"W, 1570 m); Atlantic Forest--humid mountain forest (MN 69804).
13. Brazil, Rio de Janeiro, Nova Friburgo, Tres Picos, Salinas (22[degrees]22'12"S, 42[degrees]30'32"W, 1380 m); Atlantic Forest--humid mountain forest (MN 80379).
14. Brazil, Rio de Janeiro, Itatiaia, Parque Nacional de Itatiaia: several localities including Abrigo Rebocas (22[degrees]23'07.5"S, 44[degrees]40'43.93"W, 2395 m; MN 69680), Piscina Maromba (22[degrees]26'S, 44[degrees]37'W, 1170 m; MN 69803), and Retiro de Ramos, 19 km from Parque Nacional de Itatiaia main house (exact locality coordinates unknown; type locality of Akodon serrensis leucogula); Atlantic Forest--humid mountain forest. The following sigmodontines also taken here: A. montensis, Akodonparanaensis, Brucepattersonius sp., Delomys altimontanus, D. dorsalis, D. sublineatus, O. nigripes, and Thaptomys nigrita.
15. Brazil, Rio de Janeiro, Nova Friburgo, Debossan, Sitio Xitaca (22[degrees]26'S, 42[degrees]32'W, 1010 m); Atlantic Forest--humid mountain forest (MN 35912). C. angustidens was obtained together with the following sigmodontines: Akodon montensis, Delomys sublineatus, Oligoryzomys nigripes, and Sooretamys angouya.
16. Brazil, Rio de Janeiro, Teresopolis, Parque Nacional da Serra dos Orgaos, Base da Pedra do Sino (22[degrees]27'34"S, 43[degrees]01'40"W, 2100 m); Atlantic Forest--Mountain field, mountain scrub, humid mountain forest (MN 69806).
17. Brazil, Rio de Janeiro, Cachoeiras de Macacu, Parque Nacional dos Tres Picos (22[degrees]27'45"S, 42[degrees]39'11"W, 1300 m); Atlantic Forest--humid mountain forest (MN 71942).
18. Brazil, Rio de Janeiro, Teresopolis, Parque Nacional da Serra dos Orgaos, Vale das Antas (22[degrees]27'50"S, 43[degrees]02'33"W, 1950 m); Atlantic Forest--Mountain field, mountain scrub (MN 69807). The following sigmodontines also taken here: Brucepattersonius sp., Delomys dorsalis, O. nigripes, Oxymycterus judex, and Thaptomys nigrita.
19. Brazil, Minas Gerais, Passa Quatro, Fazenda do Itaguare (22[degrees]28'S, 45[degrees]05'W, 1500 m) Atlantic Forest--humid mountain forest (UFMG 1854).
20. Brazil, Sao Paulo, Campos do Jordao, Parque Estadual de Campos do Jordao (22[degrees]44'22"S, 45[degrees]35'29"W, 1628 m); Atlantic Forest--humid mountain forest with Araucaria (CJ 5).
21. Brazil, Sao Paulo, Sao Jose do Barreiro, Parque Nacional da Bocaina (22[degrees]50'S, 44[degrees]41'W, 1400 m); Atlantic Forest--humid mountain forest (MN 69805).
22. Brazil, Rio de Janeiro, Paraty, Parque Nacional da Bocaina, RJ-165 road (23[degrees]11'54"S, 44[degrees]50'38"W, 1120 m); Atlantic Forest--humid mountain forest (MN 77791).
23. Brazil, Parana, Morretes, Represa de Guaricana (23[degrees]36'S, 49[degrees]04'W, 941 m) Atlantic Forest--humid mountain forest with Araucaria (MZUSP 29128).
24. Brazil, Sao Paulo, Salesopolis, Estacao Ecologica Boraceia (23[degrees]38'00"S, 45[degrees]52'00"W, 870 m); Atlantic Forest--humid mountain forest (MZUSP 9469).
25. Brazil, Sao Paulo, Sao Paulo (23[degrees]43'S, 46[degrees]43'W, 780 m); Atlantic Forest--humid mountain forest (MZUSP 6410).
26. Brazil, Sao Paulo, Paranapiacaba (23[degrees]46'41"S, 46[degrees]18'16"W, 820 m) Atlantic Forest --humid mountain forest (MZUSP 1796).
27. Brazil, Sao Paulo, Capao Bonito, Fazenda Intervales (24[degrees]20'00"S, 48[degrees]25'00"W, 700 m) Atlantic Forest--humid mountain forest (MZUSP 27226).
28. Brazil, Parana, Curitiba, Quatro Barras (25[degrees]21'56"S, 49[degrees]04'37"W, 936 m); Atlantic Forest--humid mountain forest with Araucaria (Rabelo 2007; P 828).
29. Brazil, Parana, Curitiba, Sao Jose dos Pinhais (25[degrees]26'S, 49[degrees]00'W, 1024 m); Atlantic Forest--humid mountain forest with Araucaria (Rabelo 2009).
30. Brazil, Parana, Piraquara, Roca Nova (25[degrees]26'30"S, 49[degrees]00'51"W, 905 m); Atlantic Forest--humid mountain forest (type locality of Akodon serrensis, see Thomas 1902; P 888).
31. Brazil, Parana, Piraquara, Mananciais da Serra (25[degrees]29'16.35"S, 45[degrees]58'32.2"W, 1117 m); Atlantic Forest--humid mountain forest with Araucaria (MN 78416).
32. Brazil, Parana, Sao Jose dos Pinhais, Guaricana (25[degrees]32'05"S, 49[degrees]12'23"W, 906 m); Atlantic Forest--humid mountain forest with Araucaria (P 108).
33. Brazil, Parana, Tijucas do Sul, Fazenda Panagro (25[degrees]57'14"S, 49[degrees]16'13"W, 1117 m); Atlantic Forest--originally sub-tropical forests of Auracaria augustifolia; today forests are reduced to small patches forming mosaics with agriculture and stockbreeding areas (Barros-Battesti et al. 1998; P 566).
34. Argentina, Misiones, "rio San Antonio, 32 km al norte de Bernardo de Irigoyen": from this locality [ca. 25[degrees]58'S, 53[degrees]10'W, 400 m], was collected 1 specimen by Harold Cleveland on 4 August 1971 and housed at the Coleccion de Mamiferos del IADIZA with the number M-00677 (Justo and De Santis 1977:47). This material was the base for the first reference of the species in Argentina, but subsequently criticized and discarded by Galliari et al. (1996:46), Pardinas et al. (2003:139), and Pereira et al. (2005:84) arguing measurement inconsistencies or a misidentification in favor to Thaptomys or the species of the cursor group of Akodon. Since the voucher specimen is loss from IADIZA collections (R. Ojeda, pers. comm. 2010), any inference about their identity is weak. However, is interesting to note that the information offered by Justo and De Santis (1977) regarding coloration and cranial morphology totally agrees with which is known for C. angustidens Argentinean specimens. More indeed, it was obtained in the Araucaria district, a typical habitat of this rodent in Brazil. In this context we consider here to resurrect this record as valid.
35. Argentina, Misiones, Reserva Privada de Vida Silvestre Urugua-i (25[degrees]59'S, 54[degrees]05'W; Pereira et al. 2005:83): a single specimen (MACN 22246) was recorded after an effort of 1,565 trap-nights in primary forest (ca. 300 m) belonging to the districts of "Laurel" and "Palo Rosa" (sensu Martinez-Crovetto 1963). The following sigmodontines also taken here: Akodon sp. (including A. montensis), Brucepattersonius sp., Delomys dorsalis, Nectomys squamipes, Oligoryzomys nigripes, Oxymycterus misionalis, Euryoryzomys russatus, and Sooretamys angouya.
36. Brazil, Parana, General Carneiro (26[degrees]25'S, 51[degrees]19'W, 1101 m); Atlantic Forest (Raboni et al. 2012).
37. Brazil, Santa Catarina, Reserva Biologica de Sassafras, Doutor Pedrinho (26[degrees]42'S, 49[degrees]40'W, 950 m); Atlantic Forest--"Floresta Ombofila Mista" (Testoni et al. 2012).
38. Argentina, Misiones, Provincial Route 2, 6 km NE Arroyo Paraiso (27[degrees]12'47.7"S, 54[degrees]01'59.9"W, 312 m): a single specimen (CNP 449) was obtained in secondary Atlantic Forest. The animal is an adult female and was trapped by Rosario Robles on April 24, 2007 (original number LTU 372).
39. Brazil, Santa Catarina, Rancho Queimado, Alto da Boa Vista (27[degrees]42'S, 49[degrees]09'W, 1200 m); Atlantic Forest--"Floresta Ombrofila Mista," edges of forest fragments, near high-altitude grasslands (Testoni et al. 2012).
40. Brazil, Santa Catarina, Campo Belo do Sul, Fazenda Gateados (27[degrees]58'S, 50[degrees]49'W, 1000 m); Atlantic Forest--"Floresta Ombrofila Mista" (Testoni et al. 2012).
41. Brazil, Santa Catarina, Urubici, Parque Nacional de Sao Joaquim, Fazenda Caiambora (28[degrees]01'S, 49[degrees]36'W, 1136 m); Atlantic Forest--humid mountain forest with Araucaria ((Goncalves et al. 2007).
42. Brazil, Rio Grande do Sul, Cambara do Sul, Parque Nacional da Serra Geral, Campo Limpo do Arroio do Segredo, Canion Fortaleza (29[degrees]04'17.4"S, 49[degrees]59'24.5"W, 1000 m); Atlantic Forest--humid mountain forest with Araucaria (MN 78550).
43. Brazil, Rio Grande do Sul, Sao Francisco de Paula, Centro de Pesquisas e Conservacao da Natureza Pro-Mata (29[degrees]26'53"S, 50[degrees]35'01"W, 900 m); Atlantic Forest--ecotone between "Campos de Altitude e Floresta Ombrofila Mista no Planalto das Araucarias" (UFRGS 2550-UFRGS 2558).
44. Brazil, Rio Grande do Sul, Sao Francisco de Paula, Centro de Pesquisas e Conservacao da Natureza Pro-Mata--CPCN Pro-Mata (29[degrees]29'08.9"S, 50[degrees]12'38.7"W, 600 to 900 m), Ombrophilous Mixed Forest (Araucaria Forest; Abreu et al. 2014).
Abreu, M., A. U. Christoff, V. H. Valiati, and L. R. de Oliveira. 2014. New distribution records of Serra do Mar Grass Mouse Akodon serrensis Thomas, 1902 (Mammalia: Rodentia: Sigmodontinae) in the southernmost Brazil. Check List 10:655-659.
Barros-Battesti, D. M., M. Arzua, P. M. Linardi, J. R. Botelho, and I. J.
Sbalqueiro. 1998. Interrelationship between ectoparasites and wild rodents from Tijucas do Sul, State of Parana, Brazil. Memorias do Instituto Oswaldo Cruz 93:719-725.
Bonvicino, C. R., S. M. Lindbergh, and L. S. Maroja. 2002. Small non-flying mammals from conserved and altered areas of Atlantic Forest and Cerrado: comments on their potential use for monitoring environment. Brazilian Journal of Biology 62:765-774.
Galliari, C., U. F. J. Pardinas, and F. J. Goin. 1996. Lista comentada de los mamiferos argentinos. Mastozoologia Neotropical 3:39-61.
Gonsalves, P. R., P. Myers, J. Vilela, J., and J. A. Oliveira. 2007. Systematics of species of the genus Akodon (Rodentia: Sigmodontinae) in southeastern Brazil and implications for the biogeography of the campos de altitude. Miscellaneous Publications, Museum of Zoology, University of Michigan 197:1-24.
Hershkovitz P. 1998. Report on some sigmodontine rodents collected in southeastern Brazil with descriptions of a new genus and six new species. Bonner Zoologische Beitrage 47:193-256.
Justo E. and L. de Santis. 1977. Akodon serrensis serrensis Thomas en la Argentina (Rodentia, Cricetidae). Neotropica 23:47-48.
Martinez-Crovetto, R. 1963. Esquema fitogeografico de la Provincia de Misiones (Argentina). Bonplandia 1:171-223.
Modesto, T. C., F. S. Pessoa, M. C. Enrici, N. Attias, T. Jordao-Nogueira, L. M. Costa, H. G. Albuquerque, and H. G. Bergallo. 2008. Mamiferos do Parque Estadual do Desengano, Rio de Janeiro, Brasil. Biota Neotropica 8:153-159.
Moraes, L. B., D. A. P. Bossi, and A. X. Linhares. 2003. Siphonaptera parasites of wild rodents and marsupials trapped in three mountain ranges of the Atlantic Forest in Southeastern Brazil. Memorias do Instituto Oswaldo Cruz 98:1071-1076.
Pardinas U. F. J., G. D'Elia, and S. Cirignoli. 2003. The genus Akodon (Muroidea: Sigmodontinae) in Misiones, Argentina. Mammalian Biology 68:129-143.
Pereira, J., P. Teta, N. Fracassi, A. Johnson, and P. Moreyra. 2005. Sigmodontinos (Rodentia, Cricetidae) de la reserve de vida silvestre Uurugua-i (Provincia de Misiones, Argentina), Con la confirmacion de la presencia de "Akodon" serrensis en la Argentina. Mastozoologia Neotropical 12:83-89.
Rabelo, G. P. 2007. O genero Akodon (Rodentia, Cricetidae)--principais mecanismos responsaveis pela variabilidade cariotipica e sua possivel implicado na distribuido das especies ocorrentes no estado do Parana. Monografia de Bacharelado em Ciencias Biologicas, Universidade Federal do Parana, Parana, Brazil.
Raboni, S.M., A. Delfraro, L. Borba, B. R. Teixeira, V. Stella, M. R. Araujo, S. Carstensen, G. Rubio, A. Maron, E. R. S. Lemos, P. S. D'Andrea, and C. N. D. Santos. 2012. Hantavirus infection prevalence in wild rodents and human antihantavirus serological profiles from different geographic areas of south Brazil. American Journal of Tropical Medicine and Hygiene 87:371-378.
Testoni, A.F., J. Fumis, S. L. Althoff, F. R. Tortato, and J. J. Cherem. 2012. Akodon serrensis Thomas, 1902 (Mammalia: Rodentia: Sigmodontinae): Records in Santa Catarina state, southern Brazil. Check List 8:1344-1346.
Thomas, O. 1902. On mammals from the Serra do Mar, Parana, collected by Mr. Alphonse Robert. Annals and Magazine of Natural History Series 7, 9:59-64.
Winge, H. 1887. Jordfundne og nulevende Gnavere (Rodentia) fra Lagoa Santa, Minas Geraes, Brasilien. E Museo Lundii, 1(3):1-178 + 18 plates.
S2. Study notes on Habrothrix angustidens and Akodon serrensis made by R. S. Voss in November, 2008.
Notes on Akodon serrensis Thomas, 1902
I examined the type (BMNH ???) and eight topotypes (BMNH ???, ZMUC 1257) of Akodon serrensis, consisting of skins and skulls collected at Ro9a Nova, Serra do Mar, between 930 and 1150 m above sea level by A. Robert in August and September of 1901. The skins are dark yellowish- or orangish-brown dorsally, with thick, soft fur; the ventral surface is washed with bright buff or tawny over dark-gray hair bases. Tails are dark, unicolored, and sparsely haired. The eyes are small, and the mystacial vibrissae are short, not extending posteriorly much beyond the base of the pinnae. Genal vibrissae are present.
The skull in dorsal view is characterized by a long, pointed rostrum flanked by shallow zygomatic notches; the frontal sinuses are inflated and the interorbital region is broad, with rounded supraorbital margins in young specimens (older animals develop small, sharp, convergent postorbital edges). The zygomatic arches are delicate, widest across the squamosal roots, and convergent anteriorly. The braincase is large, smooth, and globose. The zygomatic plate in lateral view has a straight anterior margin that is aligned vertically or slopes slightly backward; the origin of the superficial masseter is marked by a scar (not a tubercle) that is just below and slightly behind the anteroventral margin of the zygomatic plate. The carotid circulation is primitive, an alisphenoid strut is present, and the subsquamosal fenestra is large (but smaller than the postglenoid foramen). The incisive foramina are long (extending deeply between the M1 anterocones); the palate is broad and uncomplicated by keels or deep lateral grooves; and the mesopterygoid fossa extends anteriorly to or between the M3s. There are no large posterolateral palatal pits, just tiny foramina flanking the mesopterygoid fossa.
The mesopterygoid roof is completely bony or perforated by very narrow sphenopalatine openings. The auditory bullae are small and flask-shaped with broad bony eustacian tubes.
The upper incisors are narrow and opisthodont, with yellow-orange enamel bands. The molars are very narrow in proportion to their length, and retain little occlusal detail even in young animals, the cusps and lophs wearing rapidly away to dentine. The alveolus of the lower incisor is contained in a ridge, not a distinct process, on the lateral surface of the mandible behind the base of the coronoid process.
Notes on the type series of Habrothrix angustidens Winge, 1887
The type series of Habrothrix angustidens consists of subfossil cranial fragments and disassociated hemimandibles from several caves in the vicinity of Lagoa Santa; Winge's handwritten labels suggest that all were recovered from mineral sediments, not from Recent owl pellets. The caves from which specimens were collected include those that Lund called "Lapa Serra das Abelhas", "Lapa da Escrivania", "Lapa de Cerca Grande", and "Lapa da Quebra Chavelha." All of this material qualifies as syntypes because Winge did not designate a holotype. In order to more securely fix the application of the name angustidens, we designate the large cranial fragment that Winge illustrated (op. cit. plate II, fig. 9) as the lectotype, by which action all the remaining material become paralectotypes.
Altogether, this material (lectotype plus paralectotypes) preserves the entire upper and lower dentition, the left zygomatic plate, diastema, palate, interorbital region, and mandibles. We compared this material side-by-side with a complete topotypical cranium of Akodon serrensis (ZMUC 1257) and found these two taxa to be phenotypically indistinguishable. In addition to qualitative resemblances, the few measurements that we were able to take from the lectotype of angustidens all fall within the range of variation seen in the topotypic series of serrensis:
Length of Diastema: 6.3 mm Length of Molars: 4.9 mm Breadth of M1: 1.2 mm Length of Incisive Foramina: 5.9 mm Breadth of Rostrum: 4.4 mm Breadth of Palatal Bridge: 2.6 mm Breadth of Zygomatic Plate: 2.2 mm Least Interorbital Breadth: 5.2 mm Depth of Incisors: 1.2 mm
The only possible taxonomic conclusion is that angustidens and serrensis are synonyms.
Robert S. Voss
4 November 2008
Measurements (mm) of the type series of Akodon serrensis (a) Minimum Maximum Mean N Length of Head and Body (b) 85 105 92 8 Length of Tail (b) 70 85 78 8 Hind Foot (including claws) (c) 23 25 24 8 Ear (b) 17 18 17 8 Condylo-incisive Length 22.5 24.8 23.7 8 Length of Diastema 6.3 7.5 6.8 9 Length of Molars 4.8 5.2 4.9 9 Breadth of M1 1.1 1.2 1.2 9 Length of Incisive Foramina 5.5 6.1 5.8 9 Breadth of Rostrum 4.2 4.7 4.5 9 Breadth of Palatal Bridge 2.5 3.3 2.8 9 Breadth of Zygomatic Plate 2.2 2.6 2.4 9 Least Interorbital Breadth 5.2 5.6 5.4 9 Breadth of Braincase 11.7 12.7 12.2 9 Depth of Incisor 1.1 1.3 1.2 9 Length of Orbital Fossa 7.7 8.4 8.0 9 (a) BMNH 188.8.131.52-184.108.40.206, ZMUC 1257. (b) Measured by collector. (c) Measured from dry skin.
S3.-Individual specimen scores projected onto principal components I and II extracted from the analysis of log-transformed values of five cranial and two dental variables of all age classes of Castoria angustidens nov. comb. Gray circles represent recent individuals; black circle corresponds to the lectotype of Habrothrix angustidens Winge (ZMK 1/1845:13246).
Loadings of five cranial and two dental variables onto the first two principal components of Castoria angustidens nov. comb.
Principal components Variables First Second Length of diastema 0.52 -0.19 Length of incisive foramina 0.44 -0.36 Crown length of maxillary toothrow 0.22 0.38 Breadth of first upper molar 0.36 -0.21 Breadth of rostrum 0.37 0.35 Interorbital breadth 0.25 0.70 Breadth of zygomatic plate 0.40 -0.14
S4.-External appearance of Castoria angustidens nov. comb. based on the skin of the holotype of Akodon serrensis Thomas, 1902 (BM 220.127.116.11) in dorsal (top), ventral (middle), and left lateral (bottom) view.
S5.-Gastric morphology of Castoria angustidens nov. comb. based on CNP 449 (Misiones, Argentina): diagram (above), field-photograph of the bisected specimen (below). Anatomical features indicated are: a, antrum; bf, bordering fold; c, corpus; e, posterior end of esophagus; ia, incisura angularis.
S6.-A) GTG-banding patterns of Castoria angustidens female karyotype, 2n = 46 and FN = 46. Inset, X and Y chromosomes from the male individual. B) Male C. angustidens metaphase after hybridization using double probe (Akodon paranaensis 6 and 7). Whole painted chromosomes corresponding to A. paranaensis 6 and 7 on C. angustidens metaphase are indicated on the left. The painted chromosomes of C. angustidens are indicated on the DAPI-stained metaphase on the right. Asterisks indicate the 2 chromosomes of C. angustidens painted by A. paranaensis 7. C) Male Necromys lasiurus metaphase, 2n = 34 and FN = 34 after hybridization using double probe (A. paranaensis 6 and 7). Painted segments and chromosomes corresponding to A. paranaensis genome on N. lasiurus metaphase are indicated on the left. The painted chromosome and segments of N. lasiurus are indicated on the DAPI-stained metaphase on the right. Asterisks indicate the 2 chromosomes of N. lasiurus painted by A. paranaensis 7. The identification of chromosomes was based on GTG-banding patterning. Painted segments are pink.
S7.-Ventral view of the skulls of the holotypes of Akodon pervalens Thomas, 1925 (to the left; BM 18.104.22.168; photograph thanks to British Museum of Natural History, [C] The Natural History Museum, London) and Akodon serrensis Thomas, 1902 (to the right; BM 22.214.171.124; photograph taken by A. Bezerra). Main differences are, among others, the broad rostrum (pervalens) vs. the acute rostrum (serrensis); the foraminal margins resembling parentheses (pervalens) vs. the constricted anterior portion of the incisive foramina (serrensis); the smaller braincase (pervalens) vs. the inflated and larger braincase (serrensis).
S8.-Montane forest with Araucaria at about 2100 m, Itamonte, Brazil inhabited by Castoria angustidens nov. comb. Photograph taken by L. Geise.
S9.-Latitude and elevation for collection localities of Castoria angustidens nov. comb. The plotted points are listed in the gazetteer (Supporting Information S1).
S10.-Lectotype of Castoria angustidens nov. comb. (ZMK 1/1845:13246) from Lapa da Escrivania Nr. 5: right lateral (a), left lateral (b), ventral (c) and dorsal (d) view of the skull (scale = 5 mm) and upper and lower right toothrows (e) from epoxy casts. The figure in (f) depicts the original label handwritten by H. Winge with the indication that the skull fragment was illustrated -"tegnet" in Danish; the original illustration of Habrothrix angustidens from Winge (1887:II, fig. 2) is shown on (g).
Ulyses F. J. Pardinas (1), Lena Geise (2), Karen Ventura (3), and Gisele Lessa (4)
(1) Instituto de Diversidad y Evolucion Austral (IDEAus, CONICET), CC 128, 9120 Puerto Madryn, Chubut, Argentina. [Correspondence: <email@example.com>].
(2) Laboratorio de Mastozoologia, Departamento de Zoologia, Instituto de Biologia, Universidade do Estado do Rio de Janeiro, Rua Sao Francisco Xavier, 524, Maracana, 20550-900 Rio de Janeiro, Brazil.
(3) Departamento de Genetica e Biologia Evolutiva, Instituto de Biociencias, Universidade de Sao Paulo, Rua do Matao 277, 05508-090 Sao Paulo-SP and Instituto de Recursos Naturais, Universidade Federal de Itajuba, Avenida BPS 1303, 37500-903 Itajuba, Minas Gerais, Brazil.
(4) Museu de Zoologia, Departamento de Biologia Animal, Universidade Federal de Vinosa, 36571-000 Vinosa, Minas Gerais, Brazil.
Table 1 Selected measurements (in mm) of the lectotype of Habrothrix angustidens (ZMK 1-1845:13246), the type series of Akodon serrensis (BMNH 126.96.36.199-BMNH 188.8.131.52, ZMK 1257; mean, minimum--maximun and sample size in parenthesis), and the holotype of Akodon serrensis leucogula (MN 2513). H. angustidens A. serrensis Length of head and body (a) -- 92, 85 - 105 (8) Length of tail (a) -- 78, 70 - 85 (8) Hind foot with claws (b) -- 24, 23 - 25 (8) Eara -- 17, 17 - 18 (8) Condylo-incisive length -- 23.7, 22.5 - 24.8 (8) Length of diastema 6.3 6.8, 6.3 - 7.5 (9) Length of upper molars 4.9 4.9, 4.8 - 5.2 (9) Breadth of M1 1.2 1.2, 1.1 - 1.2 (9) Length of incisive foramina 5.9 5.8, 5.5 - 6.1 (9) Breadth of rostrum 4.4 4.5, 4.2 - 4.7 (9) Breadth of palatal bridge 2.6 2.8, 2.5 - 3.3 (9) Breadth of zygomatic plate 2.2 2.4, 2.2 - 2.6 (9) Least interorbital breadth 5.2 5.4, 5.2 - 5.6 (9) Breadth of braincase -- 12.2, 11.7 - 12.7 (9) Depth of incisor 1.2 1.2, 1.1 - 1.3 (9) Length of orbital fossa -- 8.0, 7.7 - 8.4 (9) A. s. leucogula Length of head and body (a) 105 Length of tail (a) 73 Hind foot with claws (b) 24 (a) Eara -- Condylo-incisive length 25.8 Length of diastema 7.1 Length of upper molars 4.6 Breadth of M1 1.6 Length of incisive foramina 5.6 Breadth of rostrum 4.6 Breadth of palatal bridge -- Breadth of zygomatic plate 2.7 Least interorbital breadth 5.3 Breadth of braincase 11.1 Depth of incisor -- Length of orbital fossa 8.2 (a) Measured by collector. (b) Measured from dry skin. Table 2 Descriptive statistics (in mm; mean [+ or -] standard deviation, range, and sample size in parenthesis) for 2 samples of adult individuals of Castoria angustidens nov.comb., from Brazil (see Supplementary Material S1 for additional data about these localities); PE = Parque Estadual; PN = Parque Nacional. Fazenda da Neblina, PE da Serra do Brigadeiro females Length of head and body 98.70 [+ or -] 26.90 Length of tail 89.80 [+ or -] 23.38 Length of hind foot with claw 25.57 [+ or -] 6.31 Length of hind foot without claw 23.29 [+ or -] 5.51 Length of ear 17.20 [+ or -] 2.58 Weight 29.67 [+ or -] 10.42 Breadth across M1s 5.67 [+ or -] 1.26 Breadth across occipital condyles 6.55 [+ or -] 1.01 Breadth of braincase 11.68 [+ or -] 0.58 Breadth of incisive foramina 2.13 [+ or -] 2.27 Breadth of M1 1.25 [+ or -] 2.53 Breadth of rostrum 5.09 [+ or -] 1.44 Breadth of zygomatic plate 2.82 [+ or -] 2.09 Condylo-incisive length 25.39 [+ or -] 4.53 Height of skull 8.26 [+ or -] 0.54 Height of mandible 6.28 [+ or -] 1.11 Least interorbital breadth 5.54 [+ or -] 1.30 Length of bulla 4.77 [+ or -] 1.52 Length of diastema 7.26 [+ or -] 0.87 Length of incisive foramina 5.96 [+ or -] 1.21 Length of mandible 14.02 [+ or -] 1.22 Length of orbital fossa 8.52 [+ or -] 0.54 Length of palatal bridge 4.32 [+ or -] 1.66 Length of rostrum 11.06 [+ or -] 0.60 Length of maxillary molar row 4.96 [+ or -] 1.47 Zygomatic breadth 14.57 [+ or -] 2.14 Fazenda da Neblina, PE da Serra do Brigadeiro females Length of head and body 84.00 - 113.00 (10) Length of tail 83.00 - 95.00 (10) Length of hind foot with claw 24.00 - 28.00 (7) Length of hind foot without claw 22.00 - 25.00 (7) Length of ear 14.00 - 20.00 (10) Weight 19.00 - 38.00 (6) Breadth across M1s 5.36 - 5.99 (10) Breadth across occipital condyles 6.23 - 6.82 (10) Breadth of braincase 11.11 - 12.15 (10) Breadth of incisive foramina 1.95 - 2.39 (10) Breadth of M1 1.14 - 1.37 (10) Breadth of rostrum 4.69 - 5.62 (10) Breadth of zygomatic plate 2.45 - 3.30 (10) Condylo-incisive length 23.51 - 27.24 (10) Height of skull 7.94 - 8.69 (10) Height of mandible 5.67 - 6.66 (10) Least interorbital breadth 5.21 - 5.86 (10) Length of bulla 4.55 - 5.06 (10) Length of diastema 6.76 - 7.98 (10) Length of incisive foramina 5.45 - 6.66 (10) Length of mandible 13.34 - 14.86 (10) Length of orbital fossa 8.07 - 9.29 (10) Length of palatal bridge 3.65 - 4.55 (10) Length of rostrum 10.00 - 12.13 (10) Length of maxillary molar row 4.41 - 5.16 (10) Zygomatic breadth 13.60 - 15.35 (10) Fazenda da Neblina, PE da Serra do Brigadeiro males Length of head and body 96.67 [+ or -] 27.88 Length of tail 83.33 [+ or -] 23.15 Length of hind foot with claw 25.50 [+ or -] 7.07 Length of hind foot without claw 23.33 [+ or -] 6.26 Length of ear 16.71 [+ or -] 3.30 Weight 28.50 [+ or -] 9.82 Breadth across M1s 5.58 [+ or -] 1.06 Breadth across occipital condyles 6.49 [+ or -] 0.78 Breadth of braincase 11.67 [+ or -] 1.19 Breadth of incisive foramina 2.09 [+ or -] 2.09 Breadth of M1 1.27 [+ or -] 2.33 Breadth of rostrum 4.99 [+ or -] 1.25 Breadth of zygomatic plate 2.56 [+ or -] 1.97 Condylo-incisive length 24.74 [+ or -] 4.83 Height of skull 8.22 [+ or -] 0.27 Height of mandible 6.28 [+ or -] 0.86 Least interorbital breadth 5.47 [+ or -] 1.08 Length of bulla 4.79 [+ or -] 1.03 Length of diastema 7.02 [+ or -] 0.67 Length of incisive foramina 5.75 [+ or -] 1.03 Length of mandible 13.49 [+ or -] 1.45 Length of orbital fossa 8.26 [+ or -] 0.30 Length of palatal bridge 4.39 [+ or -] 1.40 Length of rostrum 10.71 [+ or -] 0.78 Length of maxillary molar row 5.02 [+ or -] 1.21 Zygomatic breadth 14.21 [+ or -] 1.99 Fazenda da Neblina, PE da Serra do Brigadeiro males Length of head and body 83.00 - 116.00 (9) Length of tail 74.00 - 95.00 (9) Length of hind foot with claw 24.00 - 29.00 (6) Length of hind foot without claw 22.00 - 26.00 (6) Length of ear 16.00 - 18.00 (7) Weight 21.00 - 39.00 (6) Breadth across M1s 5.24 - 5.97 (9) Breadth across occipital condyles 6.16 - 6.84 (9) Breadth of braincase 11.31 - 12.32 (8) Breadth of incisive foramina 1.77 - 2.25 (9) Breadth of M1 1.12 - 1.42 (9) Breadth of rostrum 4.50 - 5.55 (9) Breadth of zygomatic plate 1.76 - 2.97 (9) Condylo-incisive length 23.28 - 26.36 (9) Height of skull 8.00 - 8.40 (9) Height of mandible 5.80 - 6.67 (9) Least interorbital breadth 5.24 - 5.70 (9) Length of bulla 4.45 - 5.07 (8) Length of diastema 6.56 - 7.52 (9) Length of incisive foramina 5.39 - 6.22 (9) Length of mandible 12.47 - 14.06 (9) Length of orbital fossa 7.92 - 8.58 (9) Length of palatal bridge 4.02 - 4.72 (9) Length of rostrum 9.54 - 11.69 (9) Length of maxillary molar row 4.69 - 5.32 (9) Zygomatic breadth 13.52 - 14.69 (8) Vale das Antas, PN da Serra dos Orgaos females Length of head and body 100.03 [+ or -] 14.09 Length of tail 84.10 [+ or -] 12.30 Length of hind foot with claw 25.77 [+ or -] 3.30 Length of hind foot without claw 23.87 [+ or -] 3.46 Length of ear 15.80 [+ or -] 3.90 Weight 24.48 [+ or -] 4.13 Breadth across M1s 5.74 [+ or -] 3.95 Breadth across occipital condyles 6.55 [+ or -] 3.81 Breadth of braincase 11.48 [+ or -] 2.89 Breadth of incisive foramina 2.17 [+ or -] 4.61 Breadth of M1 1.30 [+ or -] 4.77 Breadth of rostrum 4.99 [+ or -] 4.10 Breadth of zygomatic plate 2.76 [+ or -] 4.51 Condylo-incisive length 25.46 [+ or -] 0.81 Height of skull 8.25 [+ or -] 3.49 Height of mandible 6.40 [+ or -] 3.59 Least interorbital breadth 5.38 [+ or -] 4.02 Length of bulla 4.86 [+ or -] 4.12 Length of diastema 7.39 [+ or -] 3.66 Length of incisive foramina 5.87 [+ or -] 3.93 Length of mandible 13.85 [+ or -] 2.18 Length of orbital fossa 8.24 [+ or -] 3.49 Length of palatal bridge 4.53 [+ or -] 4.18 Length of rostrum 11.19 [+ or -] 2.09 Length of maxillary molar row 4.99 [+ or -] 4.09 Zygomatic breadth 14.30 [+ or -] 1.80 Vale das Antas, PN da Serra dos Orgaos females Length of head and body 84.00 - 116.00 (31) Length of tail 60.00 - 96.00 (29) Length of hind foot with claw 16.00 - 31.50 (30) Length of hind foot without claw 14.00 - 29.00 (30) Length of ear 11.00 - 22.00 (30) Weight 17.00 - 36.00 (30) Breadth across M1s 5.42 - 6.09 (27) Breadth across occipital condyles 5.95 - 7.00 (27) Breadth of braincase 11.03 - 12.01 (27) Breadth of incisive foramina 1.85 - 2.49 (27) Breadth of M1 1.18 - 1.46 (27) Breadth of rostrum 4.12 - 5.58 (27) Breadth of zygomatic plate 2.28 - 3.29 (27) Condylo-incisive length 23.47 - 27.47 (27) Height of skull 7.84 - 8.59 (27) Height of mandible 5.78 - 6.94 (25) Least interorbital breadth 5.05 - 5.94 (27) Length of bulla 4.47 - 5.42 (27) Length of diastema 6.60 - 8.20 (27) Length of incisive foramina 5.34 - 6.53 (27) Length of mandible 13.11 - 14.77 (25) Length of orbital fossa 7.73 - 8.78 (27) Length of palatal bridge 3.74 - 5.07 (27) Length of rostrum 10.23 - 12.16 (21) Length of maxillary molar row 4.70 - 5.28 (27) Zygomatic breadth 13.60 - 15.06 (23) Vale das Antas, PN da Serra dos Orgaos males Length of head and body 102.20 [+ or -] 12.20 Length of tail 85.10 [+ or -] 10.18 Length of hind foot with claw 25.36 [+ or -] 2.60 Length of hind foot without claw 23.18 [+ or -] 3.05 Length of ear 16.81 [+ or -] 4.42 Weight 25.47 [+ or -] 7.54 Breadth across M1s 5.66 [+ or -] 4.73 Breadth across occipital condyles 6.49 [+ or -] 4.49 Breadth of braincase 11.42 [+ or -] 3.66 Breadth of incisive foramina 2.09 [+ or -] 5.32 Breadth of M1 1.30 [+ or -] 5.45 Breadth of rostrum 5.03 [+ or -] 4.84 Breadth of zygomatic plate 2.75 [+ or -] 5.21 Condylo-incisive length 25.47 [+ or -] 1.54 Height of skull 8.24 [+ or -] 4.30 Height of mandible 6.38 [+ or -] 4.40 Least interorbital breadth 5.34 [+ or -] 4.78 Length of bulla 4.79 [+ or -] 4.88 Length of diastema 7.39 [+ or -] 4.45 Length of incisive foramina 5.87 [+ or -] 4.70 Length of mandible 13.90 [+ or -] 3.13 Length of orbital fossa 8.31 [+ or -] 4.29 Length of palatal bridge 4.58 [+ or -] 4.92 Length of rostrum 10.98 [+ or -] 3.54 Length of maxillary molar row 4.98 [+ or -] 4.84 Zygomatic breadth 14.28 [+ or -] 2.81 Vale das Antas, PN da Serra dos Orgaos males Length of head and body 87.00 - 119.00 (40) Length of tail 70.00 - 101.00 (39) Length of hind foot with claw 22.00 - 28.00 (39) Length of hind foot without claw 19.00 - 25.00 (40) Length of ear 11.00 - 23.00 (40) Weight 15.00 - 60.00 (38) Breadth across M1s 5.19 - 6.27 (34) Breadth across occipital condyles 6.07 - 7.41 (33) Breadth of braincase 10.90 - 11.91 (33) Breadth of incisive foramina 1.76 - 2.33 (34) Breadth of M1 1.14 - 1.43 (34) Breadth of rostrum 4.40 - 5.66 (34) Breadth of zygomatic plate 2.40 - 3.06 (34) Condylo-incisive length 23.26 - 26.76 (33) Height of skull 7.85 - 8.61 (34) Height of mandible 5.77 - 6.94 (32) Least interorbital breadth 4.81 - 5.76 (34) Length of bulla 4.23 - 5.27 (34) Length of diastema 6.61 - 8.00 (34) Length of incisive foramina 4.97 - 6.44 (34) Length of mandible 13.14 - 14.79 (32) Length of orbital fossa 7.74 - 8.92 (34) Length of palatal bridge 4.15 - 6.25 (34) Length of rostrum 9.17 - 11.91 (31) Length of maxillary molar row 4.73 - 5.28 (34) Zygomatic breadth 13.54 - 15.18 (30) Table 3 Morphological comparisons of selected traits among Castoria gen. nov. and other related sigmodontines of the Akodon Division. Characters Castoria Head and body length as % total length ~ 85% Countershading weak Rostrum long and narrow Interorbital region broad Frontals convergent with sharp margins Free upper border of zygomatic plate yes Premaxillary process > 1/2 of the incisive foramen length Alisphenoid strut present Middle lacerate foramen open Subsquamosal [ssf] / postglenoid [pgf] ssf ~ pgf fenestrae size Capsular process absent Molar design main cusps opposite Anteromedian flexus absent Procingulum m1 compressed m2 / m3 in length m2 ~ m3 Gall bladder present Characters Akodon (cursor group) Head and body length as % total length ~ 80% Countershading moderate Rostrum long and narrow Interorbital region moderate Frontals "hourglass," with rounded margins Free upper border of zygomatic plate yes Premaxillary process < 1/2 of the incisive foramen length Alisphenoid strut present Middle lacerate foramen open Subsquamosal [ssf] / postglenoid [pgf] ssf ~ pgf fenestrae size Capsular process absent Molar design main cusps alternate Anteromedian flexus present Procingulum m1 not compressed m2 / m3 in length m2 > m3 Gall bladder present or absent Characters Deltamys Head and body length as % total length ~ 85% Countershading weak Rostrum short and narrow Interorbital region narrow Frontals "hourglass," with rounded margins Free upper border of zygomatic plate no Premaxillary process > 1/2 of the incisive foramen length Alisphenoid strut absent Middle lacerate foramen close Subsquamosal [ssf] / postglenoid [pgf] ssf ~ pgf fenestrae size Capsular process absent Molar design main cusps opposite Anteromedian flexus present Procingulum m1 compressed m2 / m3 in length m2 > m3 Gall bladder present Characters Necromys (excluding amoenus and urichi) Head and body length as % total length ~ 55-60% Countershading moderate Rostrum short and broad Interorbital region broad Frontals convergent with beaded margins Free upper border of zygomatic plate yes Premaxillary process < 1/2 of the incisive foramen length Alisphenoid strut present Middle lacerate foramen open Subsquamosal [ssf] / postglenoid [pgf] ssf < pgf fenestrae size Capsular process present Molar design main cusps opposite Anteromedian flexus absent Procingulum m1 not compressed m2 / m3 in length m2 [much greater than] m3 Gall bladder present or absent Characters Thalpomys Head and body length as % total length ~ 60% Countershading strong Rostrum short and broad Interorbital region broad Frontals convergent with ledged margins Free upper border of zygomatic plate yes Premaxillary process < 1/2 of the incisive foramen length Alisphenoid strut ? Middle lacerate foramen open Subsquamosal [ssf] / postglenoid [pgf] ssf ~ pgf fenestrae size Capsular process present Molar design main cusps alternate Anteromedian flexus present Procingulum m1 not compressed m2 / m3 in length m2 [much greater than] m3 Gall bladder present Characters Thaptomys Head and body length as % total length ~ 35% Countershading weak Rostrum short and broad Interorbital region broad Frontals convergent with rounded margins Free upper border of zygomatic plate no Premaxillary process > 1/2 of the incisive foramen length Alisphenoid strut present Middle lacerate foramen close Subsquamosal [ssf] / postglenoid [pgf] ssf [much less than] pgf fenestrae size Capsular process present Molar design main cusps alternate Anteromedian flexus present Procingulum m1 not compressed m2 / m3 in length m2 > m3 Gall bladder absent
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|Author:||Pardinas, Ulyses F.J.; Geise, Lena; Ventura, Karen; Lessa, Gisele|
|Date:||Jun 1, 2016|
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