New trace fossils of termites (Insecta: Isoptera) from the late Eocene‐early Miocene of Egypt,and the reconstruction of ancient isopteran social behavior

Three new ichnogenera and five new ichnospecies are described for new trace fossils of termitaria (including associated gallery systems) of subterranean termites from upper Eocene through lower Miocene rocks of northern Egypt. All but two ichnospecies (Krausichnus trompitus, ichnogen, and ichnosp. nov., and A’, altus, ichnosp. nov.) show varying degrees of affinity to nests of the extant subterranean termite species Sphaerotermes sphaerothorax (Termitidae, Macro‐termitinae).

Termitichnus qatranii (Bown, 1982) is divided into two ichnospecies, the previously named T. qatranii, and a more generalized form, T. simplicidens, ichnosp. nov. Vondrichnus obovatus, ichnogen. and ichnosp. nov., is named for simple, possibly macrotermitine nests with oblate form, and Fleaglellius pagodus, ichnogen. and ichnosp. nov., records a nest form similar to V. obovatus, but one in which successive vertical growth by chamber apposition has produced a subterranean, tower‐like structure, reminiscent of that built by extant, epigeous Cubitermes. Krausichnus trompitus, ichnogen. and ichnosp. nov., and A’, altus, ichnosp. nov., record two unique nest architectures probably produced by unknown but related species of humivorous termites. The nest architecture expressed by Krausichnus is of wholly unknown affinity and is only distantly related in form to the ichnofossils of other termite nests known from the Tertiary of Egypt.

Study of the architecture of the nests of these ancient termites reveals details important in reconstructing the phylogeny of termite nests and documents two novel blueprints for chamber expansion and society budding in what were probably primitive Macrotermi‐tinae. It also indicates that at least two constructed edifices, earlier known only as epigeous manifestations of extant termite species, were almost certainly first developed by species living underground.

The extant Macrotermitinae are a subfamily of termites originally believed to have evolved in the post‐Eocene of the Ethiopian biogeo‐graphic region. We offer fossil evidence of four distinct, possibly macrotermitine structures from upper Eocene rocks. This evidence suggests that this termite subfamily (or at least their peculiar mode of nest construction) might have evolved considerably earlier, as it is already well established and exhibits several variants by the late Eocene in Egypt.     

EARLY DEVONIAN BIOEROSION IN THE RADE DE BREST, ARMORICAN MASSIF, FRANCE

Abstract:  Abundant and diverse bioerosion trace fossils in the Lower Devonian of the Rade de Brest, France, were produced by boring ctenostomate bryozoans, polychaete annelids and algae. Two unidentifiable traces of encrusting sclerobionts are also considered. Two new taxa are described: Pinaceocladichnus cristatus ichnosp. nov. and Ichnogutta erectus ichnogen. et ichnosp. nov. Both are attributed to the work of boring bryozoans. Sedimentary condensation during intervals of sea-level rise seems to have had an important influence in promoting infestation by organisms, the activities of which are now preserved as trace fossils. A zonation of bioerosion trace fossils within the offshore zone, in the Lower Devonian of the Armorican Massif, is also suggested.     

南极利文期顿岛晚三叠世复理石相遗迹化石

南极利文斯顿岛赫德半岛中的迈尔斯陡崖组主要是浅变质的石英长石杂砂岩、粉砂岩、泥岩和粉砂岩与泥岩互层等组成的复理石,厚度可能超过３０００m,形成于浊积扇的上扇和下扇部分,时代可能为晚三叠世。在复理石相岩石中发育一套深水的遗迹化石,共有１５个遗迹属、１６人遗变种,其中有１０个可以鉴定到遗迹种,两个比较种,４个只鉴定到遗迹属,未鉴定到遗迹种,１建立１个新遗迹属及新遗迹种,除新遗迹属种外,其余１４个遗迹属,１５个遗迹种都曾经在深海复理石相浊积岩内发现过,其中Ｂelorhaphe,Glockerichnus,Lophoctenium,Rhabdo-glyphus,Paleodictyon,Suborenzinia,Spirophycus,Strobilorhaphe,Tuberculicnus,Cochlichnus等,属于浊流前产生在复理石相泥岩内的高度分异的雕画迹（Ｇraphoglyptida),它们产于泥岩却保存为上覆砂岩底面的铸型凸起。别外,Ｆucusopsis和Neonereites却产生在砂岩内代表浊流后形成的沉积后遗迹组合,文内还描述３种浊流常见的无机构造,并上上述遗迹化石进行比较。     

再论假裸枝叠层石科*

内蒙古桌子山地区中奥陶统公乌素组遗迹化石十分丰富.本文共鉴定、描述14个遗迹属、17个遗迹种,其中1新遗迹属——Biscopulatichnus、4新遗种和6未定种.并根据不同遗迹属在不同层位中的相对丰度,建立了4个遗迹组合,自下而上的顺序是:1. Taenidium-Phycodes 组合;2. Volkichnium 组合;3. Helminthopsis 组合;4. Granularia-Circulichnus 组合,大致相当于 Seilacher (1967) 的 Zoophycos 遗迹相,反映出公乌素组沉积期,水体较深且宁静、缺氧、有机质含量高,推测应为大陆斜坡环境.     

Review of Japanese Dinosaur Track Localities: Implications for Ichnotaxonomy,Paleogeography and Stratigraphic Correlation

Nine dinosaur ichnospecies from the Lower Jurassic to Upper Cretaceous of Japan, including two that are new, are described herein. The new ichnotaxa are Asianopodus pulvinicalx ichnogen. et ichnosp. nov. and Schizograllator otariensis ichnosp. nov. The Japanese ichnotaxa are allied to Lower Jurassic ichnospecies in South China, North America, Western Europe and South Africa, and Upper Jurassic to Lower Cretaceous ichnospecies from Southeast and East Asia. This suggests they were part of a global ichnofauna before continental drift began in the Middle Jurassic, leading to the development of a more endemic dinosaur fauna in the Cretaceous. At least two assemblages, an ornithopod-gracile-toed theropod-dominated community, in northeastern Asia, and a robust theropod- and sauropod-dominated community in the southern part of the continent, existed in the Cretaceous. This parallels North American dinosaur distribution patterns in the Cretaceous and seems to be a reflection of paleolatitudinal controls.     

a new ichnogenus for etchings made by cheilostome bryozoans into calcareous substrates

Some encrusting cheilostome bryozoans etch a pattern of small pits into hard calcareous substrates, especially calcitic and aragonitic shells of molluscs. These patterns, herein described as Leptichnus ichnogen. nov., comprise pits which are sub-circular to elongate in cross section and are found in either uniserial ( L. dromeus isp. nov.) or multiserial arrangements ( L. peristroma isp. nov., the type species). Each pit corresponds to the location of a single zooid in the bryozoan colony. The oldest known Leptichnus is Late Cretaceous (Maastrichtian), the trace fossil first becomes common in the Cenozoic, and at least nine modern cheilostome genera produce incipient Leptichnus. Leptichnus can be the only evidence remaining of encrusting cheilostomes following taphonomic or diagenetic loss of their calcareous skeletons. The mechanism by which bryozoans etch into their calcareous substrates is unknown but is almost certain to be chemical and necessitates having windows in the basal walls of the zooids which permit contact with the substratum beneath. Etching may result in better adherence to the substrate, giving protection from abrasion and bioerosion.     

中国湘南早侏罗世双壳类壳体内的遗迹化石--兼论遗迹化石属Talpina Hagenow, 1840

论文描述的双壳类壳体内的钻孔遗迹组合发现于湖南省东南部宜章县下坪附近的心田门组上部(早侏罗世,早辛涅缪尔期)浅海沉积地层中。这个遗迹化石群中,帚虫类动物制造的钻孔Talpinahunanensisichnosp.nov.(新遗迹化石种)明显地占据了大部分,偶尔共生的其它钻孔包括:Rogerellaichnosp.(蔓足亚纲尖胸类的遗迹)、Calcideletrixichnosp.(可能是藻类的遗迹)、藻或菌造的遗迹(暂定为"Mycelites"ichnosp.)、以及一种可能被海绵动物制造的未定钻孔。生物侵蚀者侵害了海底上较大的双壳类壳体。钻孔保存为天然印模(石核),它们是文石质壳体溶解后留下的空间中的充填物。此外,文中还讨论了遗迹化石属TalpinaHagenow,1840,尤其讨论了有关遗迹种颇为复杂的鉴别问题,同时校正了该遗迹属的属征。     

Microtaphonomy of bioclasts and paleoecology of microencrusters from Upper Jurassic spongiolithic limestones (External Prebetic,Southern Spain)

Microencrusters and microtaphonomic features of the Oxfordian spongiolithic limestones of the External Prebetic were studied using thin-section analysis. The spongiolithic limestone is a bioclastic-rich packstone with common echinoderm, mollusc and brachiopod remains. The bioclasts show a high fragmentation index and frequent microborings. The encrustation index (E _i) is higher for fragments of serpulids, ammonoids and bivalves, and increases with the initial grain-size of bioclasts. The main microencrusters consist of benthic microbial communities (BMC) and nubeculariids, as well as subordinate calcareous and siliceous agglutinated foraminifera, serpulids and bryozoans. BMC are usually the first colonizers, and encrusting foraminifera mainly appear on bioclasts larger than 2 mm. BMC dominate in well-developed encrustations on upward facing surfaces of larger bioclasts that are also colonized by foraminifera (nubeculariids and Subdelloidina). Bullopora, serpulids and bryozoans are more common on lower surfaces. The fact that the values of encrustation index, encrustation thickness and diversity of the microencrusters increase with the size of bioclasts is related to a higher stability and exposure time of the available bioclastic substrate. The microencruster distribution on upper and lower surfaces of large bioclasts may be related to photic control, space competition and/or predation avoidance.     

Lower Jurassic Arthropod Resting Trace from the Hartford Basin of Massachusetts,USA

Cheliceratichnus lockleyi ichnogen. nov. et ichnosp. nov. is a new ichnotaxon of arthropod resting trace (cubichnium) from the Lower Jurassic (Hettangian) East Berlin Formation in Holyoke, Massachusetts, USA. The trace fossil is preserved as showing many of the external anatomical features of the exoskeleton, which resemble those of some chelicerates, notably sun spiders (Solifugae). The resting trace is directly associated with a trackway of the ichnospecies Acanthichnus cursorius Hitchcock. This is the first described fossil resting trace of a solifugan-like arthropod, and the first direct evidence of a trackmaker of A. cursorius.     

The infaunal echinoid Micraster: Taphonomic pathways indicated by sclerozoan trace and body fossils from the Upper Cretaceous of northern Spain

Micraster echinoid fossils are common in the Upper Cretaceous Olazagutía Formation of northern Spain. Tests frequently record sclerobiont signatures (including bioerosion and encrustation), left by reaction and/or coaction phenomena. Among bioerosion structures, Oichnus simplex, O. paraboloides, O. ichnosp. A, Trypanites solitarius, Rogerella ichnosp. indet., Centrichnus cf. eccentricus, Maeandropolydora ichnosp. indet. and fungal microborings are found, as well as pits and fractures. As for sclerozoan body fossils, bivalves (Dimyidae, Anomiidae, Plicatulidae and other Pectinacea), polychaete annelids (serpulids and spirorbids), lituolid foraminiferans (Haddoniidae and Coscinophragmatidae) and bryozoans (cheilostomate ones and others undetermined), as well as other less common groups, have been identified. Taphonomic paths followed by Micraster tests are analysed, based on conservation state and degree of colonisation and sedimentary filling. This allows to distinguish between accumulated fossils and non accumulated ones (including resedimented and reelaborated ones).     

Late Cretaceous Mammal Tracks from North America

Mammal tracks from the latest Cretaceous (Campanian-Maastrichtian) of Colorado are described as Schadipes crypticus ichnogen. et ichnosp. nov., on the basis of material from the Laramie Formation in Golden, eastern Colorado. This ichnospecies, and a closely related form (Schadipes sp.) from the Mesaverde Group of western Colorado, represent the only mammal tracks so far identified from the Upper Cretaceous. A possible mammal track from North Africa (Agadirichnus elegans) was originally attributed to a lizard/lacertilian. Other purported Lower Cretaceous mammal tracks are based on isolated specimens of materials that are dubious or as yet undescribed in detail. Morphologically, Late Cretaceous mammal tracks resemble those of some modern rodents. However, based on the dominant mammalian elements of faunas at that time they are probably of marsupial or multituberculate affinity.     

Faunal distribution and colonization strategy in a Middle Ordovician hardground community

The attached fauna of one of the many hardgrounds from the Galena Group (Trentonian Substage) of the Upper Mississippi Valley is described. The fauna is composed of three principal elements, viz. (1) borers, including Cicatricula retiformis ichnogen. et ichnosp. nov., (2) pelmatozoans with encrusting holdfasts, and (3) bryozoans. Analysis of the distribution of members of each population on the hardground shows that most are strongly aggregated. The nature of, and reasons for, such aggregations are considered in the light of comparable Recent shallow-water marine populations. The community on this hardground, and those on other Galena Group hardgrounds, are immature. This is a consequence of frequent and damaging scour, which these organisms were poorly adapted to resist.     

Large gryphaeid oysters as habitats for numerous sclerobionts: a case study from the northern Red Sea

The shell of a living specimen of the Indo-Pacific gryphaeid giant oyster Hyotissa hyotis was colonized by numerous encrusting, boring, nestling and baffling taxa which show characteristic distribution patterns. On the upper valve, sponge-induced bioerosion predominates. On the lower valve intergrowth of chamid bivalves and thick encrusting associations—consisting mostly of squamariacean and corallinacean red algae, acervulinid foraminifera, and scleractinian corals—provides numerous microhabitats for nestling arcid and mytilid bivalves as well as for encrusting bryozoans and serpulids. Such differences between exposed and cryptic surfaces are typical for many marine hard substrata and result from the long-term stable position of the oyster on the seafloor. The cryptic habitats support a species assemblage of crustose algae and foraminifera that, on exposed surfaces, would occur in much deeper water.     

Halorina cryptica nov. ichnogen., nov. ichnosp., mass-occurrence of Upper Triassic crustacean microcoprolites from neptunian dikes and sills cutting the Dachstein-type carbonate platform and their paleoenvironmental significance (Northern Apuseni Mountains,Romania)

The new crustacean microcoprolite Halorina cryptica nov. ichnogen., nov. ichnosp., is reported from cryptic cavities cutting the Upper Triassic (Carnian to lower Rhaetian) Dachstein-type limestones from the Northern Apuseni Mountains, Romania. The new ichnotaxon is extremely abundant in cavities, neptunian dikes and sills filled with red ferruginous carbonate sediment. The associated microfauna consists of ostracods and rare foraminifera. The microfacies is represented by bioclastic coprolite-bearing wackestone-packstone to grainstone. The red ferruginous carbonate fillings are strongly bioturbated. The neptunian sill located at the top of the studied section contains a rich ichnofauna associated with brachiopod accumulations. It is dominated by the dimerelloid rhynchonellid Halorella, indicating a late Norian-early Rhaetian age. Although neptunian dikes and sills are rather common in the Dachstein-type carbonate platform that extended on the northwestern Tethyan margin, we report here the first record of mass-occurrence of crustacean microcoprolites in neptunian dikes and sills filled with red carbonate sediments of marine origin. They are interpreted here as cryptic cavities with specific palaeoenvironmental features (e.g., lack of light, abundant nutrients, intensive microbial activity) where crustacean arthropods thrived.     

A NEW WIDE-GAUGE SAUROPOD TRACK SITE FROM THE LATE CRETACEOUS OF MENDOZA, NEUQUÉN BASIN, ARGENTINA

Abstract:  Agua del Choique is a new Late Cretaceous sauropod track site from Mendoza Province, Neuquén Basin, Argentina. It is situated in the Loncoche Formation, late Campanian – early Maastrichthian in age, and is one of the youngest sauropod tracks site recorded in the world. Agua del Choique represents a lake setting and river-dominated delta deposits, and comprises at least 160 well-preserved tracks, located on a calcareous sandstone bed. A new ichnotaxon, Titanopodus  mendozensis ichnogen. et ichnosp. nov., is erected for the footprints of this track site. Titanopodus  mendozensis exhibits the following association of features: (1) wide-gauge trackway (manus and pes trackway ratios of 18–22 and 26–31 per cent respectively), (2) pronounced heteropody (manus-pes area ratio of 1:3), (3) outer limits of trackway defined, in some cases, by the manus tracks, and (4) manus impression with an asymmetrical crescent contour and acuminate external border. Titanopodus  mendozensis is an excellent case study of the wide-gauge style of locomotion produced by Late Cretaceous derived titanosaurs that have no impression of manual phalanges. These features, and the fossil record from the Loncoche Formation, suggest that the trackmakers were, probably, middle size saltasaurine or aeolosaurine titanosaurs (14–16 m long).     

A new sauropodomorph ichnogenus from the Lower Jurassic of Sichuan,China fills a gap in the track record

A dinosaur tracksite in the Lower Jurassic Ziliujing Formation of Sichuan Province, China consists of a spectacular sub-vertical exposure, with multiple track-bearing levels and trackways showing parallel and bimodal orientations. Based on well-preserved material, the new ichnogenus and ichnospecies, Liujianpus shunan ichnogen. nov. ichnosp. nov. is erected to accommodate distinctive sauropodomorph trackways occurring in this assemblage. Liujianpus has a unique combination of features, some relating to the early Jurassic basal sauropodomorph (prosauropod in traditional usage) ichnogenus Otozoum, others to the sauropod ichnogenus Brontopodus. Despite such a mix of basal sauropodomorph- and sauropod-like features, the trackmaker of Liujianpus is likely a basal sauropodomorph. This identification is consistent with the occurrence of basal sauropodomorph skeletons from geographically and chronologically close localities. The other distinct morphotype from the tracksite is linked to a sauropod trackmaker. As such, the ichnofauna consisting of two distinct foot morphotypes reflects the diversity of sauropodomorph dinosaurs in the Early Jurassic of Asia.     

陕西子洲中侏罗世早期恐龙足迹初步研究

陕西子洲县中侏罗统延安组砂岩中,同一地点共厚约1.7 m的5个层面发现4种食肉类恐龙足迹,自上而下分别为:1)第5层面大型三趾型足迹,实雷龙足迹科王氏子洲足迹(新遗迹属、新遗迹种)Zizhoupus wangi ichnogen.et ichnosp,nov.;2)第4和第3层面中型三趾型足迹,实雷龙足迹科龙尾峁张北足迹(新遗迹种)Changpeipus longweimaoensis inchnosp.nov.;3)第2层面小型三趾或四趾型足迹,虚骨龙类小理河陕西足迹(新遗迹种)Shensipus xiaoliheensis inchnosp.nov.;4)第1层面小型三趾型足迹,虚骨龙类的铜川陕西足迹Shensipus tungchuanensis.     

内蒙古查布地区下白垩统巨齿龙足印化石

内蒙古鄂尔多斯市鄂托克旗查布地区下白垩统中产有大量的蜥脚类和兽脚类恐龙足迹及鸟类足迹化石,作者对这个地区的足迹化石进行全面普查,发现许多新暴露出来的恐龙足迹化石,包括兽脚类恐龙足迹一新属新种Chapuslockleyiichnogen.etichnosp.nov.属于巨齿龙类(megalosaurid)所留。与兽脚类足迹保存在一起的蜥脚类足迹Brontopodusbirdi分布凌乱,反映了造迹恐龙搏斗的场面。通过测量连续的兽脚类足迹计算,Chapuslockleyi的造迹恐龙的行走速度为4.33km/h。通过足迹化石还识别出造迹恐龙的病态特征。     

河北侏罗-白垩纪界线最古老的恐爪龙类足迹

记述中国河北省赤城县倪家沟化石点一组恐龙行迹,并命名一新属新种--中国猛龙足迹(Menglongipus sinensis ichnogen.et ichnosp.nov.).足迹来自土城子组,位于侏罗-白孚纪界线.相邻的义县组曾发现最古老的恐爪龙类骨骼化石,而中国猛龙足迹的发现表明恐爪龙类早在义县组之前便出现在该地区.中国猛龙足迹的造迹者体长约65cm,非常接近于基十的近鸟类.此外,为四川伶盗龙足迹(Velociraptorichnus sichuanensis)的模式标本提供更多细节,并讨论驰龙类足迹型与其他兽脚类足迹之间的联系.     

Cenomanian cheilostome bryozoans from Devon,England

The Cenomanian witnessed a spectacular evolutionary radiation of cheilostome bryozoans, both in terms of species diversity and morphological disparity. However, Cenomanian cheilostome faunas are inadequately known. Twelve species of cheilostome bryozoans are here described from the Cenomanian Beer Head Limestone Formation of SE Devon, England, a nearshore facies of limestones and sands. Two of the cheilostome species are new – Wilbertopora manubriformis sp. nov. and Foratella cervisia sp. nov. – and five cannot be identified beyond genus level. Syntypes of Foratella forata (d’Orbigny, 1853), the Senonian type species of Foratella Canu, 1900, are illustrated using SEM and a lectotype is chosen. All of the species present are neocheilostomes, which were larval brooders. Compared with the non-brooding malacostegans that dominate pre-Cenomanian faunas, most species have avicularia and extensive frontal walls, features probably adaptive against small predators.