A late Rhuddanian (early Llandovery,Silurian) trilobite association from South China and its implications

A recovery trilobite association after the end-Ordovician mass extinction, the ‘Encrinuroides’ Association, is documented from the lower Niuchang Formation (upper Rhuddanian, lower Llandovery, Silurian) of Gaojiang, Meitan, northern Guizhou, South China paleoplate, including 10 genera of eight families. Three subassociations of this association could be differentiated, i.e., ascendingly the ‘Encrinuroides’-Eoleonaspis Subassociation, the ‘Encrinuroides’-Aulacopleura (Paraaulacopleura) Subassociation, and the ‘Encrinuroides’-Meitanillaenus Subassociation. The succession of these subassociations in the section indicates a shallowing upward trend of the environment as a result of regional tectonic activities (i.e., the Qianzhong Uplift) against the background of the Rhuddanian global transgression. A relatively complete trilobite succession in South China across the Ordovician–Silurian transition has been summarized and its implications for the trilobite macroevolution of this particular time interval are discussed in detail.     

Internal supporting structures of some Late Ordovician graptolites from South China revealed by micro-CT

Three-dimensionally preserved pyritic internal moulds of the Late Ordovician graptolites Appendispinograptus leptothecalis and Styracograptus chiai from the Upper Ordovician Wufeng Formation of the Daheba section, Hunan Province, South China, were imaged by micro-CT. The distinctive pea-shaped pits/dimples aligned in a zigzag line arising from the second/third thecal pair on the tubarium in these species are interpreted as the surface expression of the junction between the lateral thecal walls and the thickened internal cross-bars that support the nema. The relations between different internal structures such as the nema, median septum, aboral lists, interthecal septa and cross-bars are summarized, and examples of these supporting systems in different taxa are illustrated. The presence of isolated cross-bars in different diplograptid tubarium may suggest that they have evolved as a result of constructional restrictions.     

A new species of Nelumbo from South China and its palaeoecological implications

A new species of Nelumbo Adanson is described from the Eocene of the Changchang Basin, Hainan Island, South China. This is the first comprehensive report on a fossil Nelumbo species from the Eocene of the Changchang Basin, Hainan Island. The co-occurrence of fruits, tubers, rhizomes with numerous adventitious roots, leaves, and receptacles indicates that the plants grew where they are preserved. These specimens, combined with palynological assemblage and other macrofossils, indicate a tropical or subtropical humid climate in the Changchang Basin from the late early Eocene to early late Eocene.     

Late Miocene Palaeocarya (Engelhardieae: Juglandaceae) from Southwest China and its biogeographic implications

Fossil fruits of Palaeocarya (Juglandaceae) are described from late Miocene sediments of southeastern Yunnan, China. The fruits present a tri-lobed wing consisting of an intact oblong-ovate middle lobe and two lateral lobes. The lobes are apically obovate, and have pinnate venation. The middle lobe is thicker at the base and gradually tapers to the apex. The nutlet, located at the base of the winged fruit, is round and hispid, and is subdivided by a septum into two compartments. Based on extensive morphological comparisons to previously documented fossil fruits, we found that the fossil fruits align most closely with members of the genusPalaeocarya, but have a unique combination of characters. Thus, we describe the fossils as a new species, Palaeocarya hispida sp. nov. This species represents an important range expansion for low-latitude occurrences of Palaeocarya in the late Miocene and therefore substantially improves our understanding of the biogeographic history of the genus. We propose that the wide distribution of Palaeocarya and relatively narrow distributions of close relatives, Engelhardia, Alfaropsis, and Oreomunnea, might be associated with a stepwise cooling and a major ice sheet expansion in the Antarctic and Arctic from the late middle Miocene to early Pliocene. In particular, the climatic oscillations during the Quaternary, such as the last glacial maximum, might have led to a decrease in the geographic distribution of Engelhardieae.     

A latest Ordovician Hirnantia brachiopod fauna from western Yunnan,Southwest China and its paleobiogeographic significance

A new, high-diversity, latest Ordovician brachiopod fauna of nearly 800 brachiopod specimens was collected from the Wanyaoshu Formation (Hirnantian) in the Shaodihe section, Mangshi City, western Yunnan, Southwest China. Altogether 22 genera and two undetermined taxa were identified; dominant are Aegiromena, Anisopleurella, Fardenia, Dalmanella, Hirnantia and Hindella, less common, Paromalomena, Leptaena, Eostropheodonta, Cliftonia, Kinnella, Templeella and Plectothyrella, together with some rare Petrocrania, Xenocrania, Pseudopholidops, Palaeoleptostrophia, Skenidioides, Giraldibella, Draborthis, Dolerorthis and Toxorthis. This is one of the most diverse typical Hirnantia faunas, associated with the Kosov Province. The paleobiogeographic relationships between western Yunnan (Southwest China), Myanmar, Yichang (Central China), Tibet (Southwest China) and Kazakhstan are clarified using Network Analysis and NMDS. The fauna studied is most similar to that of Myanmar; both resided on the Sibumasu terrane. However, the recalculated network diagram, when including the data of Hirnantia fauna from the Prague Basin, indicates that the latter is more closely linked to that of western Yunnan, a testament to the very weak brachiopod provincialism during the Hirnantian, mainly due to the influence of dominant cosmopolitan taxa. Some species of the fauna display significant population variation. Two of the dominant taxa, Aegiromena and Anisopleurella are systematically described, whereas the other two common taxa, Fardenia and Hirnantia are measured and their outlines and internal structures analyzed.     

A new oryctocephalid trilobite from the Balang Formation (Cambrian Stage 4) of northwestern Hunan,South China,with remarks on the classification of oryctocephalids

A new trilobite, Longaspis paiwuensis n. gen. n. sp., from the Balang Formation (Cambrian Stage 4) in northwestern Hunan, South China, is described. This rare trilobite adds to an expanding taxonomic list of organisms recognized from the Balang Lagerstätte, a deposit of exceptional preservation from the Cambrian. Longaspis paiwuensis is an unusually large-sized, micropygous oryctocephalid trilobite that has proparian facial sutures, pit-like lateral glabellar furrows, up to 17 thoracic segments, and a distinct medial notch in the pygidium; it lacks marginal spines.The classification of the family Oryctocephalidae is reviewed. Three subfamilies are recognized, and Longaspis n. gen. is assigned to the subfamily Oryctocarinae.     

晚奥陶世生物大灭绝后第一个腕足动物群:来自滇东北镇雄奥陶系顶部硅化腕足类的证据*

首次记载云南镇雄奥陶系顶部尾坝层(赫南特晚期)硅化腕足类,命名其为幼始准携螺贝群落(Eospirigerina putilla Community)。将它与贵州毕节、仁怀观音桥层的德姆贝-褶窗贝群落(Dalmanella–Plectothyrella Community)[赫南特早中期赫南特贝动物群(Hirnantia Fauna)碳酸盐相浅水代表]比较表明,两者面貌差异显著。确认尾坝层腕足类化石群与中国黔北、赣东北和浙西,以及美国、加拿大、瑞典、挪威、俄罗斯、澳大利亚等地赫南特晚期至鲁丹期的化石同属于晚奥陶世大灭绝后第一个动物群,即埃吉伍德-华夏动物群(Edgewood–Cathay Fauna)。它与Hirnantia动物群在组成、多样性、优势度、生态分异、生物地理及共生生物等区别很大。大灭绝第一幕发生后,显生宙历时最短、源自高纬海域、适应冷凉水环境的“机会主义者”——赫南特贝动物群“如鱼得水”,广布全球,却在第二幕降临后,整体灭绝,成为两幕之间海洋底栖生物群演化的一个鲜明标志。此后,诞生了埃吉伍德-华夏动物群,它以世界性、延限历程长的减缩型(含复活型)属种占优势,新生属很少,体现...     

The late Sandbian – earliest Katian (Ordovician) brachiopod immigration and its influence on the brachiopod fauna in the Oslo Region, Norway

Bulk samples of brachiopods from the middle and upper parts of the Arnestad Formation and the entire Frognerkilen Formation in the Oslo Region demonstrate major vertical changes in the Sandbian to Katian amphicratonic fauna of this part of the Baltic Province. The main influx of new taxa occurred in the upper part of the Arnestad Formation (uppermost Sandbian), and in the succeeding Frognerkilen Formation (basal Katian). Faunal change is initially reflected in an increase in diversity and not as a replacement of the previously incumbent genera, which remain throughout the studied sequence. The majority of the new genera migrated from the East Baltic Region or through this region from Avalonia, while the marginal Laurentian taxa first occupied the Scandinavian part of Baltica. Six taxa migrated from Laurentia, three from Avalonia, two from Gondwana and one apparently originated on the South China palaeoplate. The faunal shifts and immigration of brachiopods appear to have been in response to an initial transgression together with the movement of Baltica into more warm temperate latitudes, but the diverse faunas developed both against a background of subsequent regression and in response to an offshore shift of biofacies. The new amphicratonic fauna was thus generated by faunal shifts from elsewhere in the Baltic Province to the marginal environments of the Oslo Region together with more exotic elements from neighbouring continents and microcontinents.     

Fruit fossils of Rosoideae (Rosaceae) from the late Pliocene of northwestern Yunnan,Southwest China

The subfamily Rosoideae Focke (Rosaceae) has a good fossil record in the Northern Hemisphere, but these fossil records are confined mainly to a few genera, whereas the majority, in particular those with herbaceous members, are still under‐represented. In this study, we describe new fruit fossils of Rosoideae, including Fragaria achenes and Rubus endocarps, from the late Pliocene of northwestern Yunnan, Southwest China. These fossils add new accounts to the fossil archive of Rosoideae and provide the first fossil record of Fragaria in East Asia. The new fossil findings provide a historical backdrop for the modern diversity and distribution of the subfamily in northwestern Yunnan, a topographically complex area accommodating a high diversity for many plant groups. Our Rubus fossils, in combination with other nearby coeval occurrences of the genus, suggest that Rubus was already establishing its modern diversity in northwestern Yunnan during the late Pliocene. This finding enriches our knowledge of the post‐Neogene diversification of flowering plants in northwestern Yunnan, which is thought to be largely driven by dramatic mountain uplifts and environmental complications associated with the southeastern extension of the Tibetan Plateau.     

Late ontogeny of the trilobite Tsinania shanxiensis (Zhang and Wang, 1985) from the Cambrian (Furongian) of Anhui,China and its systematic implications

A new occurrence of Tsinania shanxiensis (Zhang and Wang, 1985) n. comb., is reported herein from northern Anhui, China. This species is here transferred from Guluheia to Tsinania as the differences between the two genera are not enough to support the generic difference. The late ontogeny of T. shanxiensis is documented herein, which provides important evidence for the evolutionary origin of the Family Tsinaniidae. The late ontogeny of the pygidia of T. shanxiensis is divided into four stages: PSS+2 stage, PSS+1 stage, early holaspid stage, and late holaspid stage. The immature morphology of T. shanxiensis retains long pygidial spines, reminicent of Mansuyia. Therefore, the morphology of T. shanxiensis can be regarded as the transitional type between kaolishaniids (e.g., Mansuyia orientalis, M. tani) and tsinaniids (e.g., T. canens, Shergoldia laevigata) during the middle Furongian.     

Middle Miocene Cephalotaxus (Taxaceae) from Yunnan,Southwest China,and its implications to taxonomy and evolution of the genus

The infrageneric taxonomy of the genus Cephalotaxus Siebold et Zuccarini ex Endlicher (Taxaceae) is controversial as morphological characters have been doubted to change coherently among species. Although the epidermal features are considered as important characters in the taxonomy of conifers, they have not been systematically studied in this genus. Neogene foliage shoots with well-preserved epidermal structure not only can provide systematic links to the extant species, but also have implications for the taxonomical definition of the extant species by their epidermal features. In this paper, the morphologies of leafy shoots combining epidermal structure of a Neogene Cephalotaxus, which was collected from the middle Miocene of Southwest China, and ten extant species (variants) were studied. Morphology as well as epidermal structures was compared among species (variants) in Cephalotaxus. Based on the comparisons, the taxonomical definition of extant species in genus Cephalotaxus are discussed. Our results indicate that leaf morphological characters in combination with the features of epidermal structure can distinguish species in genus Cephalotaxus. The fossil foliage shoots from the middle Miocene of Southwest China were assigned to a new species, C. maguanensis n. sp., which shows close affinities to the extant C. hainanensis. On the basis of leaf and epidermal information, we suggest there are seven extant species in Cephalotaxus. With global cooling and the uplift of the Qinghai-Tibet Plateau since the middle Miocene, the cooling climate induced the differentiation and southward expansions of C. maguanensis, and its extant offspring, C. hainanensis, is distributed to Hainan Island and the Indo-Chinese peninsula.     

Early heliolitine tabulate corals from the Sandbian (Upper Ordovician) in the Yunnan-Sichuan border area,Southwest China

Heliolitine tabulate corals are extremely rare in pre-Katian (Upper Ordovician) rocks worldwide, hindering the understanding of their origin and early evolution. Here, we systematically study new heliolitine material from the upper member (Sandbian) of the Huadan Formation in the Yunnan-Sichuan border area, Southwest China, which includes Protaraea ningnanensis, Plasmoporella kiaeri, Wormsipora sp. and Heliolites spinosus. The findings indicate that Protaraea and Heliolites are ancestral forms of coccoserids and typical tubular heliolitines, respectively, and that the absence of trabecular septa is probably a primitive feature of plasmoporellids. The refined Ordovician stratigraphy of this area implies that the coccoserid Protaraea ningnanensis Lin, 1965 from the lower member (late Darriwilian) of the Huadan Formation is the earliest known heliolitine, which further indicates that all heliolitines might have evolved from Protaraea. The comparison of Protaraea with other early tabulate corals also suggests an independent heliolitine origin from auloporinids and sarcinulinids.     

Late Oligocene to Early Miocene foraminifers and ostracods from Karben (Wetterau,Hesse State,Germany): stratigraphic occurrence and palaeoecological implications

Foraminiferal and ostracod assemblages from two upper Oligocene to lower Miocene sections located in the Wetterau region (NE Frankfurt a. M.) were analysed. The lithologie units investigated encompass Cyrenenmergel, Glimmersand, Vilbeler Kies, Cerithiensand, Cerithienkalk andCorbicula-Schichten. Hemicyprideis basiliensis, H. helvetica andCypridopsis? cyclocypriformae are characteristic of Cyrenenmergel deposits, representing the last marine influence around the Rupelian/Chattian boundary. The ostracod assemblages from the Cerithiensand are characterized by variousHemicyprideis species, of whichH. rhenana is the most frequent. The microfossil assemblages of the Cerithiensand indicate normal marine conditions for the basal and upper portions of the unit and slightly brackish to normal marine conditions for the middle portion. Very low salinities are indicated for some levels close to the base (around levels of emergence above sealevel) and for the clay layer around sample 30 in the lower part of the Cerithiensand. The Cerithienkalk represents a transitional facies from normal marine to brackish/freshwater conditions as indicated by successive higher contents of stress tolerant foraminifera and freshwater ostracods. TheCorbicula-Schichten are characterized by largeEucypris-moulds and various proportions ofCypridopsis spp. (freshwater indicators). When foraminifera (Bolivina spp.) are present, the fauna indicates slightly higher but still low salinities for a few levels of theCorbicula-Schichten. Oxygenation levels were generally high (presence of large ostracods and epifaunal foraminifera) and vary from suboxic to high oxic. As in the Mainz Basin, the last occurrence ofHemicyprideis rhenana marks the upper boundary of the Cerithienschichten.Protelphidium nonioninoides can be used to identify the Cyrenenmergel when it occurs in high numbers. Cerithiensand-samples show the highest diversities and contain high portions ofUvigerinella michelsi. Overall, highest diversities and normal marine salinities are reached during deposition of the basal and upper parts of the Cerithiensand, which may correspond to global sealevel rises at the end of the Chattian.     

Upper Ordovician bryozoans from the Xiazhen Formation of Yushan,northeastern Jiangxi,East China

The bryozoan fauna from the Xiazhen Formation (Katian, Upper Ordovician) of northeast Jiangxi Province, southeast China is reported here. Seventeen species of bryozoans belonging to fifteen genera and four orders are identified: Homotrypa yushanensis, Homotrypa sp., Prasopora yushanensis, Trematopora sp., Monotrypella sp., Rhombotrypa sp., Orbignyella sp., Constellaria jiangxiensis, Constellaria sp., Stictopora nicholsoni, Trigonodictya parvula, Ptilodictya ensiformis, Stictoporella sp., Pseudopachydictya sp., Nematopora sp., Arthrostylidae sp. indet., and Chasmatoporidae sp. indet. Four of these genera have been reported previously but nine genera (Trematopora, Monotrypella, Rhombotrypa, Orbignyella, Trigonodictya, Ptilodictya, Stictoporella, Pseudopachydictya, and Nematopora), one rhabdomesine and one fenestrate are found for the first time in the Late Ordovician strata of South China. Our palaeogeographical analysis suggests that the bryozoan association is typical for the Katian, which is mostly widespread in Laurentia, Siberia, Baltica and Mediterranean, and displays palaeobiogeographical relationships to the Laurentia–Siberia Province.     

Sciuridae (Rodentia,Mammalia) from the Late Miocene of Venta del Moro (Cabriel Basin,Spain): biostratigraphical,phylogenetic and palaeoecological inferences

Abstract

Fossil squirrels are relatively abundant in Aragonian and Ramblian assemblages, but very scarce in the Late Turolian and Ruscinian of the Iberian Peninsula. Until now, the locality of Venta del Moro has yielded over 4500 micromammal dental remains, but just nine of them are sciurids. In this assemblage, we have identified the taxa Pliopetaurista pliocaenica, Heteroxerus cf. mariatheresae and Atlantoxerus cf. margaritae. This is the first co-occurrence of the genera Pliopetaurista, Heteroxerus and Atlantoxerus in the same locality, being also one of the oldest records of P. pliocaenica, the oldest record of a form that could be related to A. margaritae, and the youngest record of the genus Heteroxerus. In addition, we hypothesize about the phylogeny of Pliopetaurista in the light of recent discoveries. Finally, the presence of the genera Heteroxerus and Atlantoxerus suggests an open environment and relatively dry environmental conditions, while Pliopetaurista is purportedly a flying squirrel, dweller of forested habitats and wet climates. Based on this, we infer an open environment of grassland type, bordered by forested habitats, agreeing with the conditions supposed for Venta del Moro according to its macromammal and floral assemblages.     

Two new parrot species (Psittaciformes) from the early Pliocene of Langebaanweg,South Africa,and their palaeoecological implications

Two new parrot species (Psittaciformes) are described from the early Pliocene Varswater Formation at Langebaanweg, South Africa, an area where no parrots currently are found. A coracoid, humeri, ulnae, carpometacarpi, tibiotarsi and tarsometatarsi of at least four individuals are assigned to a new species of lovebird Agapornis. Additional tarsometatarsi of at least five individuals including a nestling are referred to a new genus and species of Psittacinae, a taxon endemic to Africa comprising the extant genera Poicephalus and Psittacus. Both species form the as yet earliest geological record of parrots in Africa and document the early diversification of the taxa Agapornis and Psittacinae. Evidence for parrots in general, and a putative graminivorous species of lovebird in particular, indicates that woodlands as well as grasslands were present at Langebaanweg during the early Pliocene, which is consistent with current hypotheses on the palaeoenvironment at and around this site.     

A new assemblage of dabashanellids (Crustacea: Phosphatocopida) from Cambrian Stage 3 of western Hubei Province,South China

Dabashanellids are univalved phosphatocopid crustaceans, and are among the earliest crustaceans known so far. They were reported mainly from the Cambrian Stage 3 of China, but only a single species was established. Here I report a new assemblage of dabashanellids from the Cambrian Stage 3 Shuijingtuo Formation in western Hubei Province, South China. All specimens are sub-millimetre in length, and are preserved as three-dimensional carbonaceous hollow shields. Five species of Dabashanella were recognized, including the previously reported D. hemicyclica, three new species, D. longa n. sp., D. semiorbiculata n. sp., D. unispinata n. sp., and an indeterminate form Dabashanella sp. In addition, five indeterminate forms of arthropods were recovered. These dabashanellids represent part of a meiofauna that lived in the off-shore, deeper environments of mid-northern Yangtze Sea during Cambrian Age 3. This study shows greater morphological diversity and ecologic disparity of dabashanellid crustaceans, and supports the previous suggestion that the invasion of meiofaunal ecdysozoans into the off-shore, deeper environments probably occurred simultaneously with their radiation in the shallow water.     

Evenness and diversity in Upper Cambrian – Lower Ordovician trilobite communities from the Central Andean Basin (Cordillera Oriental,Argentina)

Community evenness has recently received much attention, either because it is related to ecosystem functioning or because it may affect estimation of diversity. Temporal and environmental trends in diversity and evenness of trilobite communities during the Late Cambrian – Early Ordovician of the Cordillera Oriental (north‐western Argentina) are here analysed. Richness and evenness increase through time in both deep subtidal (between fair‐weather and storm wave base) and offshore (below storm wave base) communities. Two significant patterns are superimposed on this general trend: (1) the magnitude of the increase in evenness is much more pronounced in deep than in shallower settings, and (2) richness and evenness trajectories are decoupled (while a significant rise in evenness is recorded in the middle Tremadocian (Tr2), an increase in richness is delayed until the late Tremadocian (Tr3)). In contrast to expectations, a single family (Olenidae) is dominant in samples associated with this earlier rise in evenness relative to richness. Hence, this trend is explained neither by the number of families present in the communities nor by the familial identity of the most abundant taxon. Large‐scale comparisons of the timing and geographical components of these trends are restricted to the patterns recognized in Laurentian North American studies. Results from the Cordillera Oriental mirror those of Laurentia regarding the rise in both metrics in deep marine settings. Nevertheless, the timing of this increase in richness and evenness is delayed in the Cordillera Oriental, supporting the idea that palaeogeographical regions differed in the nature and timing of ecological changes. Finally, the rise in trilobite alpha‐diversity through the Late Cambrian – Early Ordovician of the Cordillera Oriental supports the idea that trilobite alpha‐diversity did not decline worldwide, suggesting that the relative decline in trilobite alpha‐diversity is most probably caused by the dilution effect.     

Fruits and seeds of Ruppia (Potamogetonaceae) from the Pliocene of Yushe Basin, Shanxi, northern China and their ecological implications

Numerous fruits and seeds of Ruppia are reported from the Upper Pliocene (2.3–3.5 Myr ago) Zhangcun Formation in Yushe Basin, Shanxi, northern China. They are the first fossil Ruppia from China and demonstrate the importance of fruit and seed fossils in recording genera not represented by fossil leaves. These Ruppia are characterized by possessing a small oval endocarp, smooth endocarp surface, distinct elliptical external depressions, distinct apical mucro, slightly curved seed shape and conspicuous globose hilum. A new species, R. yushensis Zhao, Collinson and Li, is described from these endocarp and seed features. Comparison with the two European Miocene species, R. palaeomaritima Negru and R. maritime-miocenica Szafer, indicates the existence at that time of three different geographical and stratigraphical species. R. yushensis constitutes the first Pliocene record of Ruppia and extends the range of fossils of this genus from Europe to eastern Asia. R. yushensis is the only aquatic plant in the uppermost middle part of the Zhangcun Formation. This monotypic occurrence indicates a brackish, clear, tranquil and shallow lake in this region in the Late Pliocene. The smooth endocarp surface further suggests a warm temperate or temperate palaeoclimate.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 317–329.     

A Tremadocian (Early Ordovician) palaeoscolecidan worm from graptolitic shales in Hunan Province,South China

A new locality exposing Tremadocian (Early Ordovician) graptolitic shales in Hunan Province, South China, has yielded an exceptionally well‐preserved annulated worm. This palaeoscolecidan is described as Waflascolex changdensis gen. et sp. nov. and reveals extremely fine detail of the cuticle organization and plating array. The new taxon is characterized by three critical characters: incomplete plate rows that occur only on the posterior end of the worm (except in the posterior‐most area) and do not extend over the entire circumference or along the entire trunk; a regular rhomboidal array of platelets around intercalations; and reduced cuticular organization at the posterior termination. The unique cuticular organization and platelet ornamentation in the new taxon offer insight into functional differentiation of plates in the scleritome. Palaeoscolecid distribution through the early Palaeozoic is reviewed, showing that the worms were widespread in the Cambrian and Ordovician, but became more restricted during the Silurian. Ordovician palaeoscolecidans are diverse in scleritome architecture, and strikingly different from Cambrian taxa, indicating that this group diversified as part of the Great Ordovician Biodiversification Event.