Additional trilobite fauna from the basal part of the Cambrian Series 3, Stage 5, Kunzam La (=Parahio) Formation,Parahio Valley,Spiti (Northwest Himalaya), India and its biostratigraphic significance

Additional Cambrian Series 3, Stage 5, trilobite fauna comprising Peronopsis sp., Eosoptychoparia cf. Spinosa Gaotanaspis cf. pingzhaiensis and Gaotanaspis cf. transversa have been recorded just immediately above the known Oryctocephalus indicus biozone (Cambrian Series 3, Stage 5) in the Parahio valley section (Spiti region). The FAD of Peronopsis and LAD of Gaotanaspis are used to establish a Peronopsis–Gaotanaspis concurrent biozone immediately above the Oryctocephalus indicus biozone. The first records of Gaotanaspis cf. pingzhaiensis and G. transversa from the Cambrian of Spiti region and the other faunal elements are correlated with the Peronopsis taijiangensis biozone of the Kaili Formation (South China). The stratigraphic thickness from the base of the O. indicus biozone to the top of the Peronopsis–Gaotanaspis concurrent biozone in the Kunzam La (=Parahio) Formation and its comparison to the Kaili Formation (South China) indicate a possible stratigraphic condensation in the basal part of the Cambrian Series 3, Stage 5 of the Parahio valley (Spiti). Sedimentological and sequence stratigraphic analysis supports this contention.     

Cambrian trilobite fauna of the Haydenaspis parvatya level (Series 2, Stage 4): Its stratigraphic position with respect to the Oryctocephalus indicus Biozone (Miaolingian Series,Wuliuan Stage) in the Spiti Himalaya,India

Refinement of the Cambrian biozonation in the Parahio valley, Spiti region (Himalaya) leads to the demarcation of the stratigraphic distance between the Haydenaspis parvatya level (Series 2, Stage 4) and the Oryctocephalus indicus Biozone (base of the Miaolingian Series, Wuliuan Stage), which has been a subject of debate. The present work suggests that these two trilobite beds in the Kaltarbo section are separated by 183.4 m of strata. Whether either taxon ranges beyond the beds they are found in this section or others in the Parahio Valley is yet unknown.     

Ontogeny of the redlichiid trilobite Eoredlichia intermediata from the Chengjiang Lagerstätte,lower Cambrian,southwest China

A detailed investigation of the morphology and ontogeny of the redlichioid trilobite Eoredlichia intermediata (Bulletin of the Geological Society of China, 3–4, 1940, 333) from the lower Cambrian Yu'anshan Member of the Heilinpu Formation, in Kunming, Yunnan Province, southwest China, is presented. The new material comprises a relatively complete ontogenetic series ranging from the early meraspid to the holaspid period, which reveals more details on morphological variation such as the appearance of bacculae in some holaspid specimens, contraction and disappearance of fixigenal spines, and macropleural spines of the first and second thoracic segments, which are all documented for the first time and can also be used as developmental markers defining ontogenetic phases. Two distinct morphotypes, possibly an expression of intraspecific variation or sexual dimorphism, are distinguished by the morphology of pleural spines of the second thoracic segment in meraspid degree 14 and holaspides. The trunk segmentation schedule of E. intermediata is also discussed and conforms to the protarthrous developmental mode. The distinction of the thoracic region into two portions can be observed during late meraspid and early holaspid periods, which might be considered as a reference for a better understanding on the relationship of tagmosis and growth segmentation among the Cambrian redlichiid trilobites.     

A new stratigraphic occurrence of the taxon Pagetia (Trilobita) from the Spiti region and its biostratigraphic significance in correlation of the Wuliuan Stage (Miaolingian Series) in the Kashmir and the Spiti regions (Tethyan Himalaya), India

Abundant, though moderately well-preserved, specimens of Pagetia sp. are recorded along with the ptychopariid Xingrenaspis dardapurensis from a new stratigraphic level which lies above the Oryctocephalus salteri biozone in the Spiti region (Himalaya). This occurrence of Pagetia in a higher stratigraphic level (higher than the Oryctocephalus salteri biozone) from the Spiti region helps in understanding the distribution of this taxon in the Cambrian of the Kashmir and Spiti regions of the Himalaya. A Pagetia-Xingrenaspis association from the Kashmir region is already known, the present discovery of a similar association in the Spiti region enables the Wuliuan (Miaolingian) biostratigraphic correlation between the Kashmir and the Spiti regions. The record of the taxon Pagetia from a higher stratigraphic level in the Spiti region contradicts the previous assumption that the Pagetia bearing level in Kashmir is equivalent to the Pagetia-Oryctocephalus indicus (O. indicus biozone, Hayden horizon 2) in the lowest part of the Wuliuan in the Spiti region.     

Ontogeny of the Upper Cambrian (Furongian) Olenid trilobite Protopeltura aciculate (Angelin, 1854) from Skåne and Västergötland,Sweden

Abstract: The ontogeny of the trilobite Protopeltura aciculata (Angelin, 1854) is described on the basis of material from the upper Cambrian (Furongian) of Andrarum (Skåne) and Hjelmsäter (Västergötland), Sweden. P. aciculata is present in the Parabolina brevispina and Parabolina spinulosa zones. Protopeltura aciculata is represented by all stages of growth, from early protaspides to holaspides, although most of the specimens are disarticulated and precise degrees are unknown. The cranidia have therefore been allocated to five morphological groups. Cuticular sculpture of the cranidia changes throughout ontogeny. Large tubercles are present in earlier stages, disappear gradually in middle meraspid stages and are replaced with a very faint granulation. The transitory pygidium, relatively large and shield‐shaped with upwardly and backwardly directed marginal spines in early meraspides, later becomes very small, triangular‐shaped and lacking spines as a late meraspid and holaspid. The development of hypostomes and librigenae is also described. Protopeltura aciculata shows major intraspecific variations throughout development, especially regarding the pygidium where variation is much less constrained than in many other olenids. This high developmental plasticity may be a survival strategy for a trilobite living in a stressed environment. Protopeltura inhabited a dysoxic environment, possibly unusually prone to localised spreading of anoxic or toxic water. Some morphs may have been less vulnerable than others to such stresses, surviving by chance and thus enabling the species to continue.     

Intraspecific variation of trunk segmentation in the oryctocephalid trilobite Duyunaspis duyunensis from the Cambrian (Stage 4, Series 2) of South China

A detailed exploration of growth and trunk segmentation of the oryctocephalid trilobite, Duyunaspis duyunensis Chang & Chien in Zhou et al. 1977, from the lower Cambrian (Stage 4, Series 2) Balang Formation in western Hunan Province, South China, is presented. Because of the excellent preservation, the complete post‐protaspid ontogenetic series from merapsid degree 0 to the holaspid phase is described. The ontogenetic series reveals new information on morphological changes such as the migration of the posterior branch of the facial sutures (from proparian to opisthoparian) and contraction of the posteromedial notch in the pygidium. The abundance of articulated specimens available from a narrow stratigraphical interval makes this material singularly useful for studying the morphogenesis and post‐embryonic growth of D. duyunensis in comparison with other oryctocephalids. Strong evidence that multiple numbers of pygidial segment are recognized in each meraspid degree as well as in the holaspid period showed unusual intraspecific variability in the rate of trunk segmentation, providing insights into how Cambrian subisopygous trilobites controlled their body patterning, including size, shape and trunk segment number in both thorax and caudal plate during growth.     

Lower-Middle Cambrian boundary in open shelf facies of the Siberian Platform

In the Early and Middle Cambrian (of traditional usage), three broad facies belts were developed over the Siberian Platform. The lithofacies belts comprise evaporitic-carbonate, reef-shoal, and open-marine environments.Trilobites assemblages are most diverse in the open-marine lithofacies (Judoma-Olenek facies), represented especially by the Kuonamka Formation, in the eastern part of the platform. Species, genera and families of trilobites gradually increased through the Middle-Cambrian boundary interval in the Kuonamka Formation. Protolenidae dominated in the time represented by the Lermontovia Zone. In strata assigned to the Anabaraspis Zone, the protolenids disappear, the paradoxidids appear, and the diversity of dorypygids increases. The number of trilobite families increases in the Oryctocara and Kounamkites zones.Well represented in the Kuonamka formation of the eastern Siberian Platform are three levels of trilobites that have potential for global correlation. They are the Oryctocephalus reticulatus (=O. indicus latus), Oryctocephalus indicus, and Ovatoryctocara granulatus levels. The Lower-Middle Cambrian boundary in the South China correlates to the lower part of the Kounamkites Zone of the Siberian Platform.     

Phylogeny and taxonomic reassessment of pikas Ochotona pallasii and O. argentata (Mammalia,Lagomorpha)

We examined a cranial morphometric data set consisting of 186 specimens from the entire distribution range of Ochotona pallasii sensu lato and O. argentata, as well as 67 complete sequences of the COI gene and 28 sequences of the MGF and PRKCI introns from these and closely allied species. Our results show that the two allopatric morphologically similar taxa composing O. pallasii sensu lato – from Mongolia and adjacent territories and Kazakhstan – are paraphyletic relative to O. argentata. Genetic distances between these three taxa are larger than the intraspecific variation known for the subgenus Pika, in which the species under consideration belong; these distances are even larger than the interspecific differences among closely related species such as O. hyperborea, O. mantchurica and O. hoffmanni. Thus, the three focal taxa are recognized here as distinct species. Inspection of the type specimen of O. pallasii indicated that this specimen was not collected in Kazakhstan, has previously been theorized. The most probable place of the holotype's origin is Russian south‐eastern Altai (Chuyskaya Steppe); whatever its exact origin, it definitively originates from the ‘Mongolian’ taxon. Based on this evidence, the senior synonym for the Kazakh pika is O. opaca Argyropulo, 1930. Thus, we propose to recognize three separate species in the O. pallasii species group: O. pallasii (Mongolia and adjacent territories), O. opaca (eastern Kazakhstan) and O. argentata (Helan Shan Range, China).     

Moulting,ontogeny and sexual dimorphism in the Cambrian ptychopariid trilobite Strenuaeva inflata from the northern Swedish Caledonides

Abstract: Three thousand seven hundred disarticulated remains together with several articulated specimens of the Cambrian Series 2 ptychopariid trilobite Strenuaeva inflata Ahlberg and Bergström, 1978 have been collected from the Torneträsk area, northern Sweden. The material provides significant new data on the morphology, ontogeny, moulting and enrolment of the species. Two distinct morphotypes, possibly an expression of sexual dimorphism, are recognized. The morph with a pair of bulbs in the frontal area, interpreted as brood pouches, is considered to represent females. Statistical treatment of the length/width ratio in cranidia reveals isometric growth during ontogeny for both morphotypes. The transition from the meraspid to holaspid ontogenetic period has been established through recognition of the successive development of the number of thoracic segments in articulated late meraspides. Throughout its life cycle, S. inflata went through 11 meraspid degrees and at least 17 holaspid growth stages. Inferred moult ensembles and exuviae reveal the successive opening of cephalic sutures and the function of the rostral plate during exuviation. As in other ellipsocephalid trilobites in which enrolment is known, the pygidium and two or three thoracic segments of S. inflata are concealed beneath the cephalon (spiral enrolment) during complete enrolment.     

Morphology and ontogeny of the eodiscoid trilobite Sinodiscus changyangensis from the lower Cambrian of South China

Abstract: A large number of complete specimens together with numerous disarticulated sclerites of the eodiscinid trilobite Sinodiscus changyangensis Zhang in Zhou et al., 1977 have been collected from the lower Cambrian Shuijingtuo Formation in Changyang, Hubei Province, South China. An ontogenetic series is established based on the immature and mature exoskeletons including the previously unknown protaspides and meraspides, in particular. No further substages can be differentiated in the protaspid specimens herein. Changes that took place during the meraspid period include the addition of postcephalic segments and prominent pygidial larval notches in early meraspid development which became progressively less distinct and disappeared in degree 2. Two holaspid stages are recognized based on the addition of a new pygidial segment, indicating that the start of the holaspid phase preceded the onset of the epimorphic phase and accordingly, its developmental mode is attributed to the protarthrous pattern. The trunk segmentation schedule of S. changyangensis is discussed, which is similar to other primitive eodiscoid trilobites, that is, as the boundary between the thorax and pygidium migrated posteriorly, there is no change in the number of the trunk segments. The processes of liberation of the thoracic segment and segment insertion into the pygidium are separated from one another, and the two different mechanisms, somitogenesis and tagmosis, progress independently during the ontogenetic development of the postcephalic region of these primitive eodiscinids.     

Mobergellans (Problematica) from the Cambrian of Greenland, Siberia and Kazakhstan

Mobergellans are disc-shaped problematic fossils of phosphatic composition bearing prominent muscle attachment structures on the presumed internal surface. They are found exclusively in sediments of Early Cambrian age, mainly from Laurentia, Baltica and Siberia. Type material ofDiscinella brastadi Poulsen, 1932 from North-East Greenland is re-studied and synonomised withDiscinella micans (Billings, 1871). Specimens from Siberia described asD. brastadi are re-assigned toMobergella hexactina n. sp., and specimens from Siberia described asMobergella radiolata Bengtson, 1968 are re-assigned toM. sibirica n. sp.Aktugaia triangula Mis-Sarzhevsky, 1976 from Kazakhstan is redescribed, and its relationship to other mobergellans is discussed.      

Palynomorphs from the Cambrian Series 3, Parahio valley (Spiti), Northwest Himalaya

We present low-diversity acritarch assemblages, including Alliumella baltica, Bavlinella faveolata, Brocholaminaria nigrita, Brocholaminaria sp., Eliasum sp., Leiosphaeridia minutissima, Leiosphaeridia spp., Lophosphaeridium sp., Pterospermella solida, Satka colonialica, Siphonophycus sp., and Synsphaeridium sp., as well as filamentous algae and cryptospore-like microfossils recorded from 45 rock samples around the Oryctocephalus indicus Zone (Cambrian Series 3) in the Cambrian Kunzam La (Parahio) Formation at the Kaltarbo locality in the Parahio valley, Northwest Himalaya. One new species Synsphaeridium parahioense was established. The acritarch assemblages at the basal part of the Cambrian Series 3 in the Kunzam La (Parahio) Formation were dependent on the local marine environment. The new discovery of cryptospore-like microfossils from the Cambrian Kunzam La (Parahio) Formation supports the extensive distribution of possible cryptospores in the Cambrian stratigraphic sequences.     

Precambrian–Cambrian boundary interval occurrence and form of the enigmatic tubular body fossil Shaanxilithes ningqiangensis from the Lesser Himalaya of India

The affinity of the Ediacaran fossil Shaanxilithes ningqiangensis and putatively related forms has long been enigmatic; over the past few decades, interpretations ranging from trace fossils to algae to metazoans of uncertain phylogenetic placement have been proposed. Combined morphological and geochemical evidence from a new occurrence of S. ningqiangensis in the Krol and Tal groups of the Lesser Himalaya of India indicates that S. ningqiangensis is not a trace fossil, but rather an organic‐walled tubular body fossil of unknown taxonomic affinity. Specimens consist of compressed organic cylindrical structures, characterized by extended, overlapping or fragmented iterated units. Where specimens intersect, overlapping rather than branching or intraplanar crossing is observed. Lithologic comparisons and sequence stratigraphic data all suggest a late Ediacaran age for the uppermost Krol Group and basalmost Tal Group. By extending the biogeographical distribution of S. ningqiangensis, hitherto confined to the Ediacaran of China and potentially Siberia, to the Precambrian–Cambrian boundary interval of India, this new occurrence of S. ningqiangensis expands the biostratigraphic utility of this enigmatic fossil to the inter‐regional and intercontinental scale. Moreover, study of these new and exceptionally preserved samples may help to significantly constrain the long‐debated problem of Shaanxilithes' affinity, elucidating its ‘problematic’ status and shedding new light upon the ecology and taphonomy of one of the most significant intervals in early life history.     

Potential global stratotype sections and points in China for defining Cambrian stages and series

Three stratigraphic sections in the Cambrian of China that contain complete successions across important biohorizons having chronostratigraphic value, and that may be useful for developing Cambrian stage or series boundaries, are reviewed. The Wuliu section (Guizhou, South China), contains the FAD of Oryctocephalus indicus at 58.2 m above the base of the Kaili Formation. The Wangcun North section (Hunan, South China), contains the FADs of Ptychagnostus (or Acidusus) atavus, Ptychagnostus punctuosus, and Lejopyge laevigata at 1.2 m, 56.7 m, and 111.3 m, respectively, above the base of the Huaqiao Formation. The Xiaoyangqiao section (Jilin, Northeast China), contains the FAD of Cordylodus proavus in the Fengshan Formation at 10.89 m above the zero point of the section. A fourth potential stratotype section, the Wangcun South section (Hunan, South China), which contains the P. atavus, P. punctuosus, and L. laevigata zones, is discussed briefly.     

The ontogeny of cinctans (stem‐group Echinodermata) as revealed by a new genus,GraciaCystis, from the middle Cambrian of Spain

Abstract:  A new cinctan echinoderm, Graciacystis ambigua gen. et sp. nov. from Cambrian Series 3 rocks of Spain, is described based on more than 100 articulated specimens that range from 6 to 14.5 mm in thecal length. This material shows that Graciacystis ambigua, while plastic in thecal shape, is highly conservative in its thecal construction, with a fixed number of marginal plates and very limited addition of plates in the stele and ventral membrane through ontogeny. Ventral swellings on marginal elements are absent from the smallest specimens and become gradually more marked during growth. A cladistic analysis shows Graciacystis to be a basal cinctan, more derived than Sotocinctus and the Trochocystitidae and as sister group to a large clade formed by Sucocystidae + Gyrocystidae. The determinate growth pattern seen in Graciacystis seems to be the general pattern for all cinctans.     

Environment and space as drivers of variation in skull shape in two widely distributed South‐American Tayassuidae,Pecari tajacu and Tayassu pecari (Mammalia: Cetartiodactyla)

The influence of the environment on the geographical variation of morphological traits has been recognized in a number of taxa. Pecari tajacu and Tayassu pecari are ideal models to investigate intraspecific geographic variation in skull because of their wide and heterogeneous geographical distribution in South America. We used geometric morphometric procedures to examine the geographical variation in skull shape of 294 adult specimens of these species from 134 localities. We quantified to what extent skull shape variation was explained by environment, skull size and geographical space using variation partitioning analysis. We detected a strong pattern of geographic variation for P. tajacu skull shape, but not for T. pecari. The environment seems to be the major selective force that drives skull shape variation in both species. Nevertheless, other spatially structured processes (e.g. genetic drift, gene flow) might also have affected variation in the skull shape of the more widespread species P. tajacu. Allometric relationships might reflect the biomechanical constraints that are thought to be strong enough to limit size‐related changes in T. pecari skull shape.     

Upper Cambrian agnostoid trilobites from the Machari Formation, Yongwol, Korea

The Middle to Upper Cambrian Machari Formation in Korea is well known for abundant and diverse trilobites along with other invertebrate fossils. Based on recent reappraisal on the trilobites of the Machari Formation, eight trilobite zones have been proposed for the Upper Cambrian sequence: i.e., in ascending order the Glyptagnostus stolidotus, Glyptagnostus reticulatus, Proceratopyge tenuis, Hancrania brevilimbata, Eugonocare longifrons, Eochuangia hana, Agnostotes orientalis and Pseudoyuepingia asaphoides zones. Trilobites from the lower five zones and part of the Pseudoyuepingia asaphoides Zone have been already published elsewhere. In this paper, we describe the agnostoid trilobites of the upper three zones, Eochuangia hana, Agnostotes orientalis and Pseudoyuepingia asaphoides zones, of the Machari Formation. These comprise 24 species belonging to 13 genera, including one new genus (Yongwolagnostus) and ten new species (Homagnostus? sulcatus, Ivshinagnostus alatus, Ivshinagnostus quadratus, Yongwolagnostus stenorhachis, Yongwolagnostus dubius, Nahannagnostus pratti, Pseudagnostus? dividuus, Pseudagnostus medius, Pseudorhaptagnostus? urceus and Ammagnostus serus). These biozones correlate well with middle Upper Cambrian zones established in South China, Australia, Kazakhstan, Siberia, and Canadian Rocky Mountains.     

Geographic variation of body morphology of the Atlantic bluefin tuna, (Thunnus thynnus,Linnaeus, 1758)

Geometric morphometric methods were used to explore body shape morphology in 260 Atlantic bluefin tuna, Thunnus thynnus, collected in Sardinia (Western Mediterranean) during the breeding phase and in the Bay of Biscay (North Eastern Atlantic) during the feeding phase. The shape of each specimen was captured by high resolution digital images and recording the 2‐D coordinates of seven morphological landmarks. A general procruste analysis (GPA) was applied in order to eliminate any morphological variations resulting from size, position or orientation of specimens. A thin plate‐spline (TPS) method was then used to provide a graphical representation of the shape conformation between two sets of data. Results of the regression model between the direct and indirect measurements accounted for a R² = 0.98. The Principal Components Analysis shows differences linked to the two sampling areas, accounting for 37% and 19.97% of the body shape variation in the first (PC1) and second (PC2) principal component, respectively. Specifically, the deformation grid projection highlights the major differences regarding the anterior‐ventral part of the body (landmark 5‐6‐7). These differences might not necessarily be linked to an actual population substructure. Instead, it was hypothesized that such body shape differences were due to the diverse life phases during which specimens were collected, since the reproductive specimens show a ‘pot‐bellied’ shape, which was larger than for the feeding specimens that showed a ‘slimmer’ shape. Analyses of likely sexual dimorphism conducted on Sardinian specimens did not reveal any significant differences; whereas body shape differences related to the pre‐ and post‐reproductive sizes were detected.     

Oxytropis sobolevskajae sp. nov. (Fabaceae: Papilionoideae,Galegeae) from Tuva Republic (south Siberia,Russia)

Oxytropis sobolevskajae Pjak, a new species endemic to Tuva Republic (south Siberia, Russia), is described and illustrated. The new species belongs to O. sect. Xerobia Bunge subsect. Ampulla Vass. and is confined to the central Tuva depression. This species is probably most closely related to O. leptophylla (Pall.) DC., from which it is easily distinguished by the number of leaflet pairs, the length and type of inflorescence, and the length of the keel cusp.