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Measurements of the molars of Huerzelerimys minor sp. nov.
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In this paper the various species and populations attributed to the genus Progonomys Schaub, 1938 are revised. Valerymys Michaux, 1969 is considered to be synonymous with Occitanomys Michaux, 1969, since the type-species V. ellenbergeri (Thaler, 1966) was included in Occitanomys by Aguilar et al. (1986a). Other species from Western Europe that had...
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... In addition, the Yihachi Fauna includes taxa with the FAD of Albanensia, Paralactaga, Dipus, Nannocricetus, Sinocricetus, Abudhabia, Pararhizomys, Prosiphneus, Ochotona and Ochotonoma, the LAD of Mioechinus?, Heterosminthus and Protalactaga in Bahean LMS/A of late Miocene Qiu and Li, 2016), and the FAD of Huezelerimys in MN 10 (Mein et al., 1993). Accordingly, the age of the Yihachi Fauna is likely early Late Miocene, late Bahean Chinese land mammal stage/age, roughly equivalent to MN 10-11 of the European land mammal zonation (Figs. ...
... The earliest record of Muridae in North Africa is represented by Progonomys cathalai from the early Vallesian locality of Farafra (Egypt; Heissig 1982), although Mein et al. (1993) considered that this attribution is contingent on a revision. Progonomys cathalai has been identified in other several northern Africa localities such as Oued Zra (Morocco; Jaeger 1977a), Bou Hanifia 1, 2 and 5 (Algeria; Ameur-Chehbeur 1988), Oued Tabia (Morocco; Benammi et al. 1996), Afoud 6 (Morocco; Benammi et al. 1995;Benammi 2001) and Ma 486 (Morocco; Görler and Zucht 1986). ...
... In contrast, this feature is shown in half of specimens of Progonomys mauretanicus (see Coiffait-Martin 1991). The European Progonomys hispanicus Michaux, 1971 was transferred to the genus Occitanomys by Mein et al. (1993). The t1 of the studied M1 is somewhat more displaced anteriorly than that of Occitanomys hispanicus. ...
... Therefore, the ascription to this species can be discarded. Other species excluded from the genus Progonomys by Mein et al. (1993) are Progonomys clauzoni Aguilar, Calvet and Michaux, 1986; Progonomys debruijni Jacobs, 1978, and Progonomys yunannensis Qiu and Storch, 1990. Anyhow, the studied molars are somewhat smaller than those of Progonomys clauzoni and larger than those of Progonomys debruijni and Progonomys yunannensis. ...
The continental Late Miocene is relatively well represented in North Africa, but stratigraphic gaps are present in the Vallesian due to the scarcity of localities from this time. In this paper, we present the small mammal association from the section of Guefaït 1, in the Aïn Beni Mathar Basin (Jerada, eastern Morocco). This sequence contains three successive fossiliferous levels: Guefaït 1A, Guefaït 1B and Guefaït 1C. The small mammal association of the richest level, Guefaït 1A, includes insectivores (Schizogalerix sp., Lartetium cf. africanum), bats (cf. Hypsugo sp.) and rodents of the families Gerbillidae (Zramys salemi, Myocricetodon seboui, Myocricetodon aff. trerki, Myocricetodon sp.), Muridae (Progonomys cathalai), Gliridae (Microdyromys cf. chaabi) and Sciuridae (Atlantoxerus sp.). Large mammals are scarcely represented by Gazella sp. and Hipparion sp. The rodent association, characterised by the occurrence of Progonomys cathalai and Zramys salemi, indicates an early Late Miocene, latest Vallesian age for the sites of Guefaït 1. The novel small mammal combination from this site constitutes a significant finding that improves our understanding of evolution of terrestrial Late Miocene faunas from northern Africa.
... tubercles; t1bis and t2bis. appendices to t1 and t2 (After Qiu and Li, 2016) 1 Systematic paleontology Muridae Gray, 1821 Murinae Gray, 1821 Huerzelerimys Mein et al., 1993 Type species Parapodemys vireti Schaub, 1938. Diagnosis (adopted from Mein et al., 1993) "Molars smaller than or similar in size to those of extant Rattus rattus, and with a poor development of the longitudinal connection between tubercles. ...
... appendices to t1 and t2 (After Qiu and Li, 2016) 1 Systematic paleontology Muridae Gray, 1821 Murinae Gray, 1821 Huerzelerimys Mein et al., 1993 Type species Parapodemys vireti Schaub, 1938. Diagnosis (adopted from Mein et al., 1993) "Molars smaller than or similar in size to those of extant Rattus rattus, and with a poor development of the longitudinal connection between tubercles. Upper molars without t7, but with t4 and t8 connected by a weak crest. ...
A skull with mandible and several cervicals of a new species of Huerzelerimys, H. asiaticus, collected from the Late Miocene Liushu Formation in Linxia Basin, Gansu Province, is described in this paper. The skull is the first one ever found for the genus Huerzelerimys. Its main characters are: size small; skull broad and short with stout rostrum; interorbital roof narrow; premaxillary laterodorsal crest well developed; frontal crests weak and subparallel; incisive foramina long, with their posterior ends lined up with anterior root of M1; posterior palatal foramina located mesial to M2; caudal border of hard palate lying posterior to M3; interpterygoid foramen absent; alisphenoid canal bony; bulla large and inflated; internal carotid foramen located near the basilar tubercle; mandible having low horizontal ramus and deeply concave diastema; anterior end of masseteric ridge lined up with anterior margin of m1; mental foramen situated slightly anterior to masseteric ridge and anteroventral to m1, near to mandibular diastema; upper incisors orthodont; M1 having slightly anteriorly located t1; in M1 and M2 t6 and t9 connected by distinct crest and t12 crest-like; t1 and t3 of M2 and t1 of M3 connected to t5; M3 having t3 and a large isolated t8; small Acc of m1 connected with both Alc and Abc; m1 and m2 having distinct buccal cingula, larger c2 attached to protoconid, and low crest-like posterior heel; m2 and m3 having isolated Abc; c1 absent in m3. The evolutionary level of the described skull tends to show that the age of the upper part of the Liushu Formation yielding H. asiaticus may belong to late Bahean, corresponding to European upper MN11 or lower MN12.
... Remarks-The Udabno specimens share some features with the genus Progonomys Schaub, 1938. However, all the M1s have a t6-t9 connection, so the assignment to the genus Progonomys is excluded (Schaub, 1938;Mein et al., 1993). The genus Hansdebruijnia was created by Storch and Dahlmann (1995) based on a unique combination of characters. ...
The locality of Udabno, Georgia, preserves material referred to the hominoid primate Dryopithecus. In this study, we describe the microvertebrate fauna from the site, which includes fishes (Scardinius sp., Carassius sp.), anurans (Pelophylax sp.), squamates (Lacertidae indet., Ophidia indet.), insectivores (Schizogalerix sinapensis, Turiasorex cf. pierremeini), and rodents (Byzantinia pikermiensis, Hansdebruijnia erksinae, Vasseuromys pannonicus). The association of two rodents, the cricetid B. pikermiensis and the murid H. erksinae, constrains the age of the Udabno fauna more precisely than previous studies. Byzantinia pikermiensis was first recognized at the site of Pikermi, Chomateri, and later recovered in other sites from Turkey, including the hominid-bearing site of Çorakyerler. The murid H. erksinae was also described from Çorakyerler in Turkey. Therefore, the presence of these rodents at Udabno indicates that the fauna postdates late Vallesian faunas such as Biodrak, Greece, and Bayraktepe II, Turkey, which are characterized by the association of the more archaic species Byzantinia nikosi and Progonomys cathalai. This study demonstrates that Udabno is temporally equivalent to typical Turolian sites from the Mediterranean area, such as Pikermi, Çorakyerler, and Crevillente 2.
... cathalai, but later Mein et al. (1993) noticed that the specimen closely follows the taxonomic definition of †Parapodemus (also see Martín-Suárez and . The single specimen possesses a complete stephanodonty (the cusps enterostyle (t4), protocone (t5), paracone (t6), metacone (t9) and hypocone (t8) united in a continuous loop) and lacks posterostyle (t7) although there is a continuous high crest between enterostyle and hypocone (Mein et al., 1993; Figure S5). ...
... cathalai, but later Mein et al. (1993) noticed that the specimen closely follows the taxonomic definition of †Parapodemus (also see Martín-Suárez and . The single specimen possesses a complete stephanodonty (the cusps enterostyle (t4), protocone (t5), paracone (t6), metacone (t9) and hypocone (t8) united in a continuous loop) and lacks posterostyle (t7) although there is a continuous high crest between enterostyle and hypocone (Mein et al., 1993; Figure S5). ...
... The descendant of †?Karnimata is considered †Karnimata darwini ( Figure S6) from Siwalik Group, Pakistan (Jacobs, 1978;Kimura et al., 2015). Mein et al. (1993) and Wöger (2011) synonymized †Karnimata darwini from Siwalik to be †Progonomys woelferi Bachmayer and Wilson, which is otherwise distributed only in Europe. Storch and Ni (2002) pointed out that †Karnimata darwini differs from †Progonomys woelferi in having enterostyle (t4) on M1 in a more anterior, symmetrical position relative to paracone (t6), and anterostyle (t1) more closely approaching lingual antercone (t2). ...
Murid rodents (Rodentia: Muridae) represent the most diverse and abundant mammalian family. In this study, we provide a refined set of fossil calibrations which is used to reconstruct a dated phylogeny of the family using a multilocus dataset (six nuclear and nine mitochondrial gene fragments) encompassing 161 species representing 82 murid genera from four extant subfamilies (Deomyinae, Gerbillinae, Lophiomyinae, and Murinae). In comparison with previous studies on murid or muroid rodents, our work stands out for the implementation of nine robust fossil constraints within the Muridae thanks to a thorough review of the fossil record. Before being assigned to specific nodes of the phylogeny, all potential fossil constraints were carefully assessed; they were also subjected to several cross-validation analyses. The resulting phylogeny is consistent with previous phylogenetic studies on murids, and recovers the monophyly of all sampled murid subfamilies and tribes. Based on nine controlled fossil calibrations, our inferred temporal timeframe indicates that the murid family likely originated in the course of the Early Miocene, 22.0-17.0 million years ago (Ma), and that most major lineages (i.e. tribes) started diversifying ca. 10 Ma. Historical biogeography analyses support the tropical origin for the family, with an initial internal split (vicariance event) between Afrotropical and Oriental (Indomalaya and Philippines) lineages. During the course of their diversification, the biogeographic pattern of murids is marked by several dispersal events toward the Australasian and the Palearctic regions. The Afrotropical region was also secondarily colonized at least three times from the Indomalaya, indicating that the latter region has acted as a major centre of diversification for the family.
... In this work, the authors provide a detailed, preliminary list of the fauna, making a first estimation of its age between units MN10 and MN 11. Subsequently, Mein et al., (1993) in a more general and detailed study of the rodent (Murids) determined the presence of Huerzelerimys minor in the site and considered it explicitly as a Karstic fisure. ...
Corral de Lobato, a karstic site in the area of Molina de Aragón has been studied in a preliminary way. Even though there are not many Neogene karstic sites in the Iberian Chain, they occur in four clusters, with ages ranging from latest middle Miocene (MN7/8) to early Pleistocene (MN17). Correlations between these clusters and the reference stratigraphical units of the Tagus Basin, as well as with local and global events, are proposed. These karstic sites provide a complementary source of fossil vertebrate remains to that of the stratified sites formed lowland. The Heritage significance of such sites arises from the enhanced preservation of rare taxa or associations, and the operation of biotic concentrative processes.
... The genus Karnimata was erected by Jacobs (1978) for a late Miocene lineage of murines from the Siwaliks of Pakistan. Brandy (1979) recognized several species from the late Miocene and Pliocene of Afghanistan, and Storch (1987) and Mein et al. (1993) attributed species from China and Spain to the genus. Some of these species are quite distinctive in morphology and are now recognized as being derived from separate lineages. ...
... Both Storch (1987) and Cai and Qiu (1993) recognized distinct discrepancies between the Siwalik and the Chinese forms of Karnimata, particularly the stephanodonty of the M1 of the latter. Subsequently, from the viewpoint of fossils from Spain, Mein et al. (1993) proposed that the type species Karnimata darwini is a synonym of Progonomys woelferi Bachmayer and Wilson, 1970, and that all Siwalik Karnimata darwini should be transferred to Progonomys. ...
... The primitive murid Tedfordomys is morphologically close to Progonomys, as revised by Mein et al. (1993) and documented by Wessels (2009). The similarities between the new genus and Progonomys include slender molars with weak longitudinal crests, t1 not greatly posteriorly positioned on M1, t6 isolated from t9, and t7 absent on M1 and M2; the tma on m1 is absent or weak and the anterior mure is undeveloped. ...
We review and expand the systematics and biostratigraphic record of murine rodents in Yushe Basin. Several new taxa were established for the Pliocene fossil record of North China
following our initial field work of the 1980s. The rodent collection was expanded and developed by complementary field work in 1991, and subsequent study. Herein we present a new Late Miocene genus and review all occurrences of Yushe Murinae. Relatively primitive taxa characterize the Late Miocene of Yushe, and these appear to be related to early lineages that diversified in the Indian Subcontinent. The interval of about 6–5 Ma in Yushe records several murines that occur at Ertemte, Inner Mongolia: Karnimatoides hipparionus, Apodemus orientalis, and Micromys chalceus.
Chardinomys
is an additional genus that distinguishes Yushe microfaunas from other Asian assemblages, and the common C. yusheensis ranges from latest Miocene through early Pliocene. With Chardinomys, the genus Huaxiamys characterizes Late Neogene Yushe murine assemblages. The Pliocene of Yushe Basin records successive species of Chardinomys, Micromys, Huaxiamys, and Apodemus. Derived species of Chardinomys, Micromys, and Apodemus persist into the Pleistocene.
... It shows a similar conservative stage of evolution as P. hussaini. All early Progonomys across Eurasia show similar molar structures despite some variation in size, and relative sizes of cusps (see Mein et al. 1993). Reasonable conclusions on dental morphology and metrics led Wessels (2009) to consider P. hussaini and P. sinensis as junior synonyms of the somewhat younger P. cathalai schaub, 1938, type locality of which is Montredon in southern France. ...
... It is likely that the populations over this vast area were distinct, if difficult to define morphologically. Current age estimates for Progonomys place its dispersal to Europe ( Mein et al. 1993) in late mammal zone MN 9, around 10 Ma. Our geohistorical view of Siwalik murine evolution sees P. hussaini as the Potwar successor of a somewhat older successful disperser that was the progenitor of P. cathalai and P. sinensis. ...
... Karnimata has been thought to occur in the late Miocene of Shanxi and Inner Mongolia, China, but Qiu and Li (2016) distinguish this material as new genus Karnimatoides. Mein et al. (1993) considered the possibility that the holotype of Siwalik Karnimata darwini might represent the European species, Progonomys woelferi bachMeyer et WiLson, 1979, possibly rendering the genus name a junior synonym. However, Storch and Ni (2002) ruled this out, noting the anterior position of the anterostyle, which unlike Progonomys is characteristic of Karnimata. ...
Abstract: The early late Miocene is an interval of increased diversification for murine rodents. Whereas the genus Progonomys
became widespread throughout Eurasia by 10 Ma, it appears from the known paleontological record that southern Asia is the
arena of evolution and diversification at the base of the Murinae. The Siwalik fossil record of the Potwar Plateau in northern
Pakistan preserves fossil evidence relevant to unraveling this history. Murine rodents are recorded there throughout the middle
Miocene, and diversification began in Siwalik assemblages before 11 Ma. The well-established Progonomys and Karnimata lineages were already present between 11 and 10 Ma, and these represent extant murine crown groups. Here we document diversity in Siwalik murines dating to 10.5 to 10.1 Ma, and clarify their recognition by naming a new species of Karnimata and referring specimens of Progonomys from this interval to P. hussaini. In addition, we define at least two other uncommon murine
species that coexist with them. One of these is an early record of Parapodemus, a fossil genus of Tribe Apodemurini, which
constitutes a calibration point for the Apodemus/Tokudaia split. Together, these fossil taxa provide further evidence bearing on the major split among murines leading to the clades Murini and Arvicanthini.
... Family Muridae Illiger, 1811 Genus Huerzelerimys Mein, Martín Suárez and Agustí, 1993 Huerzelerimys oreopitheci (Engesser, 1989) Figs 8D-G; Tab. 3 ...
... It further differs from H. vireti by the higher number of roots (five) in the M1 and M2. In H. vireti, the M1 shows three roots, although a minute fourth one may be present in some specimens, and the M2 may show three or four roots (Van de Weerd 1976;Martín-Suárez & Freudenthal 1993;Mein et al. 1993). Finally, the molars are more hypsodont and all the specimens show a t6-t9 connection, while this is variable in H. vireti (Van de Weerd 1976;Martin Suárez & Freudenthal 1993;Van Dam 1997). ...
The Late Miocene continental successions of the Baccinello-Cinigiano basin (Grosseto), one of the longest and most continuous vertebrate-bearing continental successions in the Neogene Italian record, yielded at least four superimposed vertebrate assemblages bracketed in the time span 8.3-6.4 Ma. The Baccinello-Cinigiano basin is famous for recording endemic vertebrate assemblages that include the youngest European Miocene hominoid, Oreopithecus bambolii. The late Miocene endemic vertebrate fauna known as the Baccinello V0 assemblage is the oldest vertebrate fauna within the Baccinello-Cinigiano basin succession, being correlated to the European mammal Neogene unit MN11. Recent field surveys along the Trasubbie river allowed studying in detail the basal Baccinello-Cinigiano sedimentary succession, and sampling fossiliferous level bearing microvertebrates along the small creek Fosso della Fittaia. The sample "Fosso della Fittaia 2013" yielded about 170 fossil remains improving our documentation of the oldest vertebrate assemblages from the Baccinello-Cinigiano basin. As far as rodents are concerned, in addition to the already recognized murid Huerzelerimys and glirid Anthracoglis, a few dental remains are assigned to a new genus and species of giant dormouse. It is further worth noting the occurrence in the sample of shrew remains (the first described from the Baccinello-Cinigiano basin) identified as cf. Lartetium. The latter attests the presence of a crocidosoricine in the Fosso della Fittaia 2013 assemblage, postdating the youngest known occurrences of the subfamily by at least 1 my. The vertebrate assemblage is completed by a diverse herpetofauna and the first fish remains reported from the basin.
... This approach can accommodate wide variations of tooth morphology in a single fossil species or genus (e.g. van de Weerd 1976;Mein et al. 1993;van Dam 1997;Wessels 2009). In these studies, when an apomorphic character state reaches a threshold of 50% frequency of occurrence, those specimens are recognised as belonging to a separate species or genus in a chronoclinal lineage. ...
A general lineage concept is widely adopted in studies of species delimitation for extant taxa with DNA sequence data. In the general lineage concept, species are separately evolving metapopulations that acquire sets of properties during the process of speciation. Murine rodents from the Miocene Siwalik Group of Pakistan have an excellent fossil record showing temporal trends of morphological change, and are therefore ideal to observe the process of lineage divergence as fossil evidence. Here, we review the evolution of Siwalik murine rodents in the sense of the general lineage concept and draw lines of evidence for lineage separation. For currently available datasets, we applied the initial split criterion as well as three operational criteria (phenetic, diagnosable and ecological properties). In early Siwalik murine rodents, the acquisition of distinct phenetic properties emerges subsequent to the event of the evolutionary split. The phenetic evidence occurs closer in time to ‘true’ divergence, while distinct diagnosable and ecological properties fully arise later. Within a sequence of time-controlled samples, these observations explicitly show that phenetic properties (here, shape data and a combination of linear measurements) more precisely delimit the divergence of fossil species than does the fixation of diagnostic properties.
