Siwalik synopsis: A long stratigraphic sequence for the Later Cenozoic of South Asia

The Tertiary sediments of northern Pakistan are an exceptional record of terrestrial sedimentation and represent most of Neogene time. Foremost is the Siwalik Group of the Potwar Plateau, for which multiple, superposed fossil levels span ∼18–6 Ma. Well-developed magnetostratigraphic control provides secure dating so that Siwalik fossil horizons may be interpolated into a time scale with resolution to 100,000 years. We describe the geographic setting of the Potwar, give an overview of the temporal distribution of faunas, and discuss changes in paleohabitat and paleoecology with coinciding faunal change, as seen from the Siwalik viewpoint. The long Siwalik biostratigraphy of many successive assemblages with its resolved time scale may be compared directly with other well-dated sequences. Immigrant arrival and timing of faunal change may be traced. The basins of the Iberian Peninsula show somewhat different timing of introduction of hipparionine horses, and faunal turnover in the Siwaliks clearly precedes the Vallesian crisis in Spain. In contrast to the increasingly seasonal precipitation of the late Miocene Potwar, the paleohabitat of coeval North China appears to have been moist and equable, with high diversity faunas. Continued development and comparison of resolved Neogene records allow increasing resolution of the patterns of faunal change on regional to global levels.     

A revision of fossil Rhizomyid rodents from Northern India and their correlation to a Rhizomyid biochronology of Pakistan

Based on the biochronology of rhizomyid rodents of Northern Pakistan, fossil horizons in the Siwaliks of Northern India can be correlated to the late Neogene sequence of Pakistan with greater precision than previously possible. The indian Siwaliks do not include records of the most primitive Rhizomyidae, but rhizomyid faunas of India display strong affinity with those of Pakistan in the late Miocene. Pliocene differentiation of rhizomyid faunas in India and Pakistan may reflect ecological partitioning within the Indian subcontinent. Use of the term «Nagri for deposits at Haritalyangar suggests an erroneous equivalence in age to the Nagri stratotype in Pakistan.     

A new genus of myrmicine ant (Hymenoptera: Formicidae) from Eocene Baltic amber

Examination of fossil ant specimens from various private and institutional German collections of Baltic amber has revealed a new genus and species of Myrmicinae, described here as Thanacomyrmex hoffeinsorum gen. et sp. nov. The new worker morphotype shares some similarities with the coeval extinct genus Parameranoplus Wheeler, also from Baltic amber, and the extant genera Pristomyrmex Mayr and Acanthomyrmex Emery, suggesting placement of the new fossil in the Myrmecina genus-group within the tribe Crematogastrini. The new genus highlights the tremendous diversity of ants that evolved in the Palaeogene European forests and composed the first major radiation of crown-group ants.     

A new species of Anthracotheriidae, Merycopotamus medioximus nov. sp. from the Late Miocene of the Potwar Plateau, Pakistan

New anthracotheriid remains, discovered by the H-GSP in well-dated localities from the Potwar plateau in the North of Pakistan, between 10.4 and 8.6 Ma, are described and attributed to Merycopotamus medioximus nov. sp. This new species displays an intermediate morphology between the older M. pusillus and the more recent M. dissimilis. These results permit to emend the Merycopotamus diagnosis. To cite this article: F. Lihoreau et al., C. R. Palevol 3 (2004).

Résumé

Une nouvelle espèce d'Anthracotheriidae, Merycopotamus medioximus nov. sp. du Miocène récent du plateau du Potwar, Pakistan. Des restes d'Anthracotheriidae, découverts par le H-GSP dans des localités bien datées du plateau du Potwar, au Nord du Pakistan, entre 10,4 et 8,6 Ma, sont décrits et attribués à Merycopotamus medioximus nov. sp. Cette nouvelle espèce possède une morphologie intermédiaire entre M. pusillus, espèce plus ancienne, et M. dissimilis, plus récente. Ces résultats permettent en outre d'émender la diagnose du genre Merycopotamus. Pour citer cet article : F. Lihoreau et al., C. R. Palevol 3 (2004).     

New Alicornops (Rhinocerotidae) remains from Lower and Middle Siwaliks,Pakistan

Unpublished rhinocerotid remains from the Lower and Middle Siwaliks of Pakistan are described in this paper and recognized as two species of Alicornops. Alicornops complanatum (Heissig) is identified in the Dhok Pathan Formation of the Middle Siwaliks and Alicornops laogouense Deng in the Kamlial Formation of the Lower Siwaliks. The Dhok Pathan Formation levels with A. complanatum are roughly correlated with the late Miocene-Pliocene European mammal zones MN10-15. In turn, levels with A. laogouense of the Kamlial Formation would correlate with the middle–late Aragonian (middle Miocene) European MN5. The recognition of the Chinese species A. laogouense in the Potwar Plateau represents the first discovery of this taxon in Pakistan and increases the geographical and stratigraphic distributions of this species, and adds to the rhinocerotid association from the Siwaliks. In turn, the presence of A. complanatum in the Siwaliks of Potwar Plateau also enlarges its geographic distribution in Pakistan, as it was previously known from the Bugti Hills of Balouchistan. The absence of Alicornops from the Siwaliks in the Chinji and Nagri formations (between late MN5 and MN9 zones) might be due to an inadequate fossil record, as other rhinocerotid species are known from Kamlial to Dhok Pathan formations. However, the two recorded species of Alicornops could also reveal two independent migration waves as supported by the appearance of other taxa in different formations. A summary of fossil Cenozoic rhinocerotids from different areas of Pakistan is also presented.     

Cricetid rodents from the Lower Siwalik Subgroup of Jammu,India: Biochronological significance

We describe the cricetid rodents represented by Megacricetodon daamsi, Megacricetodon sivalensis, and Myocricetodon sivalensis, recovered from two localities, Dehari and Jhajjar Kotli, lying in the upper part of the Lower Siwalik Subgroup of Jammu Province. The cricetid fauna is similar to that reported from the Pakistan Siwaliks. Based on the species identifications and the stratigraphic range of the cricetid taxa in the Siwalik succession of Pakistan, the age of the Dehari locality is correlated to between 13.8 and 13.2 Ma, whereas an age of 13.8–12.5 Ma would be assigned to the Jhajjar Kotli rodent yielding level. The occurrence of similar rodent taxa at the two investigated sites points to the homotaxial nature of the fossiliferous beds.     

A Pelican Tarsometatarsus (Aves: Pelecanidae) from the Latest Pliocene Siwaliks of India

We report a new fossil specimen of a pelican from the Tatrot Formation of the Siwalik Hills, India. It likely represents Pelecanus sivalensis Davies, 1880, the smaller of the two previously published species from the Siwalik Group stratigraphic sequence. This complete tarsometatarsus is the first fossil bone of a pelican collected in India for over 100 years. It is from the latest Pliocene (∼2.6 Ma), and is the youngest pelican fossil from the region. The new specimen exhibits a derived distoplantar ‘slant’ to the plantar margin of the medial crest of the hypotarsus, and a combination of features related to the morphology of the hypotarsus, the distal foramen, trochleae, and overall size that allow further differentiation from known tarsometatarsi of fossil and extant pelicans, including the three species of extant pelicans that occur in India (Pelecanus crispus, P. onocrotalus, and P. philippensis). It is of appropriate size for Pelecanus sivalensis, which to date has been known only by fragments of other skeletal elements of the wing, leg, and shoulder girdle. Thus, the observation that this tarsometatarsus is morphologically distinct from those of known pelicans provides further support for the distinctiveness of at least one extinct species of pelican from the Siwalik Group sediments. While the morphology of the tarsometatarsus allows for separation from other taxa known from tarsometatarsi, we found no clear shared derived states to place this taxon with any confidence in a phylogenetic context relative to any other pelican species, or even determine if it is part of the crown group of Pelecanidae. However, published molecular data are consistent with an origin of the crown clade prior to the Pleistocene, suggesting (along with one morphological character) the possibility that this species belongs to the Old World clade of pelican species.     

First Triassic record of the beetle family Permocupedidae (Insecta: Coleoptera): a peculiar example of a Lazarus taxon

The first Mesozoic representative of the extinct archostematan beetle family Permocupedidae, Frankencupes ultimus, gen. et sp. nov., is described based on two isolated elytra from the Lower Anisian (Middle Triassic) Röt Formation of Lower Franconia, Germany. The new fossil occurrence extends the range of the family from the Lower Wuchiapingian (Upper Permian) up to the Anisian, and represents a fine example of a Lazarus taxon in the fossil record of beetles.     

Taxonomy of the fossil grylloblattid nymphs (Insecta: Grylloblattida)

Nymphs of fossil grylloblattid insects are revised. Newly described taxa are Lemmatonympha gracilissima gen. et sp. nov. and Kaltanympha vorcutensis sp. nov. from the Verkhne-Syr’yaginsk locality (Ufimian, Lek-Vorkuta Formation in the Vorkuta Coal Basin), genera Sylvalitoralis gen. nov. and Tshebardanympha gen. nov. from the Tshekarda locality (Kungurian, Koshelevka Formation, Middle Urals), Tataronymphakamensis gen. et sp. nov. from the Tikhie Gory locality (Lower Kazanian, linguloid beds of the Baitugan Formation, Tatarstan), and Kaltanympha ornata sp. nov. from the Kerbo locality (Upper Tatarian, lower part of the Degali Formation, Evenki Autonomous Region). Liomopterites (?) gracilis Sharov, 1961 from Lower Kazanian deposits of the Kuznetsk Formation in the Kuznetsk Basin is transferred to the genus Kaltanympha Sharov, 1961; Permonympha arcuata Sharov, 1957 from the same locality is synonymized under Permonympha gracile Sharov, 1957; the nymph described from the Karatau locality (Upper Jurassic, Karabastau Formation in southern Kazakhstan) as Blattogryllus karatavicus Rasnitsyn, 1976 is excluded from grylloblattids. Keys to extinct grylloblattid nymphs are provided.     

Miocene hyaenids (Hyaenidae; Carnivora) from Siwaliks of Pakistan

Newly discovered Miocene hyaenid specimens, recently collected from the Siwalik Group, are described and discussed. A careful comparison with the known material reveals that these specimens belong to the early hyaenid species Thalassictis cf. T. proava, Ictitherine indet. and Lepthyaena sivalensis. The stratigraphic range of T. proava extended up to the Dhok Pathan Formation (Middle Miocene to Early Pliocene). The stratigraphic range of T. proava comprises the Middle to Late Miocene, with the youngest record in Hasnot, Potwar Plateau in the Siwalik Group. The material is of great interest because Siwalik carnivoran material is rare.     

A new large barn owl (Aves, Strigiformes, Tytonidae) from the Middle Pleistocene of Sicily, Italy, and its taphonomical significance

A new species of Tytonidae, Tyto mourerchauvireae, is described from the Sicilian cave deposits of Spinagallo, Luparello and Marasà, which have yielded a common vertebrate fossil assemblages referred to the early Middle Pleistocene. T. mourerchauvireae nov. sp. shows a pronounced increase in body size compared to other congeneric taxa. It is larger than the extant Tyto alba and the extinct Tyto balearica and Tyto sanctialbani and comparable in size among the Mediterranean taxa only with the extinct Tyto robusta, which differs in some morphological characteristics of the long bones. The insular adaptations of T. mourerchauvireae nov. sp. and the differences from the other forms of the genus Tyto which spread through the Mediterranean area during Neogene and Pleistocene are discussed. The fossil assemblages of the early Middle Pleistocene of Sicily are dominated by extinct giant Gliridae of the genus Leithia and Maltamys; these taxa are regarded as the primary prey items of T. mourerchauvireae nov. sp. The Gliridae remains were analysed microscopically to detect possible modifications of bones and teeth caused by ingestion and digestion processes. This type of analysis, together with the qualitative study of the fossil assemblages, allows to show the taphonomical importance of T. mourerchauvireae nov. sp. as agent of accumulation in such Sicilian fossil assemblages. Incidentally the taxonomic validity of T. robusta is discussed and confirmed.     

A new genus and species of fossil scorpion (?Euscorpiidae) from the Early–Middle Eocene of Pesciara (Bolca,Italy)

Fossil scorpions are among the oldest terrestrial arthropods known from the fossil record. They have a worldwide distribution and a rich fossil record, especially for the Paleozoic. Fossil scorpions from Mesozoic and Cenozoic deposits are usually rare (except in amber-deposits). Here, we describe the only fossil scorpion from the Early to Middle Eocene Pesciara Lagerstätte in Italy. Eoeuscorpius ceratoi gen. et sp. nov. is probably a genus and species within the family Euscorpiidae. This may be the first fossil record of the Euscorpiidae, which are so far only known from four extant genera. Eoeuscorpius ceratoi gen. et sp. nov. was found in the “Lower Part” of the Pesciara Limestone, which is actually dated Late Ypresian stage (between 49.5 and 49.7 Ma). Besides a possible pseudoscorpion, the here-described fossil scorpion is the second arachnid species known from the Bolca Locality.     

Palaeoisopus,Palaeopantopus andPalaeothea,pycnogonid arthropods from the Lower Devonian Hunsriick Slate,West Germany

Palaeoisopus problematicus andPalaeopantopus maucheri are redescribed and found to be primitive pycnogonids belonging to extinct orders.Palaeoisopus is particularly primitive in retaining a long abdomen with a telson. It differs from other forms also in having flattened limbs indicating a swimming mode of life and in having the ocelli differently arranged.Palaeothea devonica nov. gen. et nov. spec. is a tiny pycnogonid, the only hitherto known fossil representative of the extant order Pantopoda, indicating that entirely modern forms were in existence already in the Early Devonian. A derivation of pycnogonids from early merostomes is regarded as likely.     

New gomphaeschnids and progobiaeshnids from the Yixian Formation in Liaoning Province (China) illustrate the tremendous Upper Mesozoic diversity of the aeshnopteran dragonflies

One new genus and five new species of dragonflies (Odonata) are described and figured from the Yixian Formation of northeastern China, viz. two gomphaeschnids Sinojoria magna nov. sp. and S. cancellosa nov. sp., plus three progobiaeshnids Mongoliaeshna hadrens nov. sp., M. exiguusens nov. sp., and Decoraeshna preciosus nov. gen., nov. sp. These new discoveries confirm the apparently sudden great diversification in China of the clade Aeshnoptera during the Middle-Upper Jurassic, together with the Upper Mesozoic to modern lineages of dragonflies. At the same time, some ‘ancient’ groups of Odonata became extinct.     

The oldest ionoscopiform from China sheds new light on the early evolution of halecomorph fishes

The Halecomorphi are a major subdivision of the ray-finned fishes. Although living halecomorphs are represented solely by the freshwater bowfin, Amia calva, this clade has a rich fossil history, and the resolution of interrelationships among extinct members is central to the problem of understanding the origin of the Teleostei, the largest clade of extant vertebrates. The Ionoscopiformes are extinct marine halecomorphs that were inferred to have originated in the Late Jurassic of Europe, and subsequently dispersed to the Early Cretaceous of the New World. Here, we report the discovery of a new ionoscopiform, Robustichthys luopingensis gen. et sp. nov., based on eight well-preserved specimens from the Anisian (242–247 Ma), Middle Triassic marine deposits of Luoping, eastern Yunnan Province, China. The new species documents the oldest known ionoscopiform, extending the stratigraphic range of this group by approximately 90 Ma, and the geographical distribution of this group into the Middle Triassic of South China, a part of eastern Palaeotethys Ocean. These new data provide a minimum estimate for the split of Ionoscopiformes from its sister clade Amiiformes and shed new light on the origin of ionoscopiform fishes.     

Two new genera of cuckoo wasps (Chrysididae: Amiseginae) from Rovno and Baltic ambers

Cuckoo wasps of the subfamily Amiseginae, Eosega heterotarsata gen. et sp. nov. and Foveorisus kilimniki gen. et sp. nov., from the Late Eocene Baltic (Russia) and Rovno ambers (Ukraine), respectively, are described. Primitive Eosega is close to the extinct genus Palaeobethylus Brues in Kieffer, 1914 and represents the so-called American series of genera of Amiseginae. Foveorisus is found to be morphologically close to a number of extant oriental genera. All known fossil Amiseginae are listed. Stick insects of the family Archipseudophasmatidae, possible hosts of the Eocene amisegines, are recorded in the Rovno amber for the first time.     

A New Late Miocene Odobenid (Mammalia: Carnivora) from Hokkaido,Japan Suggests Rapid Diversification of Basal Miocene Odobenids

The modern walrus, Odobenus rosmarus, is specialized and only extant member of the family Odobenidae. They were much more diversified in the past, and at least 16 genera and 20 species of fossil walruses have been known. Although their diversity increased in the late Miocene and Pliocene (around 8–2 Million years ago), older records are poorly known. A new genus and species of archaic odobenid, Archaeodobenus akamatsui, gen. et sp. nov. from the late Miocene (ca. 10.0–9.5 Ma) top of the Ichibangawa Formation, Hokkaido, northern Japan, suggests rapid diversification of basal Miocene walruses. Archaeodobenus akamatsui is the contemporaneous Pseudotaria muramotoi from the same formation, but they are distinguishable from each other in size and shape of the occipital condyle, foramen magnum and mastoid process of the cranium, and other postcranial features. Based on our phylogenetic analysis, A. akamatsui might have split from P. muramotoi at the late Miocene in the western North Pacific. This rapid diversification of the archaic odobenids occurred with a combination of marine regression and transgression, which provided geological isolation among the common ancestors of extinct odobenids.     

A new species of cormorant (Aves: Phalacrocoracidae) from the Pleistocene of Olduvai Gorge,Tanzania

Phalacrocorax owrei nov. sp. is a small cormorant from the Lower Pleistocene of Olduvai Gorge. Osteological proportions are established for the four subgenera of Phalacrocorax, and P. owrei is assigned to the subgenus Stictocarbo. The species was the most abundant bird in the Bed I deposits and is also represented by a few specimens in the middle part of Bed II. Its last known appearance coincides with a change in the local environment when the climate became more arid and the Olduvai Lake became more saline and more alkaline. At other localities in Bed II the extinct P. tanzanice occurs, as well as P. africanus and P. corbo, which breed on the African lakes and seacoast today.