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The genus Helicoprion (Chondricthyes, Euchondrocephali) is preserved primarily by a continuous spiral root that forms the base for more than 130 teeth. Helicoprion is found globally in Lower Permian marine rocks and at least 100 specimens exist in public collections worldwide. Ten species of the genus are reviewed in the context of new morphometric analyses conducted on dozens of specimens. Helicoprion whorls share a common inner spiral geometry that results in exponential growth of tooth and root dimensions. Novel growth equations permit calculation of whorl diameter, volution number and tooth count from incomplete specimens. The results of meristic and geometric analyses identify taxobases that distinguish three emended species concepts. Differentiation of form is evident among specimens only after the second volution or roughly the 85th tooth. Helicoprion davisii has widely spaced, stout teeth with tall cutting surfaces and is distinguished from H. bessonowi, which has narrow, closely spaced teeth with short cutting surfaces. Helicoprion ergassaminon is an intermediate form, having narrow, closely spaced teeth with tall cutting surfaces. Several large specimens in the study are too dissimilar to place in the new emended species concepts.
Pywackia baileyi Landing inLanding et al., 2010, from the upper Cambrian Yudachica Member of Oaxaca State, southern Mexico, consists of small, phosphatic, proximally tapering cylindrical rods covered by shallow polygonal calices. The bryozoan-like morphology of this fossil prompted its interpretation as the first bryozoan known from the Cambrian. However, restudy of some of the original material, employing scanning electron microscopy for the first time, questions the assignment of Pywackia to the Bryozoa. Striking similarities between Pywackia and the modern pennatulacean octocoral Lituaria lead to an alternative hypothesis interpreting Pywackia an early fossil octocoral. While Pywackia is probably not a true pennatulacean, a group with a definitive fossil record stretching back only to the Late Cretaceous, it can be envisaged as having had a similar skeletal structure and ecology to Lituaria, the rods representing mineralized axes of tiny colonies that lived with their proximal ends buried in the sediment and distal ends covered by feeding polyps. Landing et al. (2010) considered the phosphatic composition of Pywackia specimens to be the result of diagenetic replacement, but the evidence is equivocal. If Pywackia had a primary phosphatic skeleton, this would support the hypothesized existence of phosphatic biomineralization early in the evolutionary history of Cnidaria, as well as providing further evidence that Pywackia is not a bryozoan.
The Sesong Formation is a member of the Taebaek Group, Korea, which extends from late Cambrian Series 3 to middle Furongian in age. Recent studies on the trilobites of the Sesong Formation have contributed significantly to the revision of the biostratigraphy. However, trilobites in the lower part of the formation, which may include the “Stephanocare Zone”, have remained essentially overlooked since the establishment of the biozone, making it difficult to correlate with the equivalent biozones of North China. Here we report trilobite faunas from the lower part of the Sesong Formation in two different sections, the Seokgaejae and the Jikdong sections, which yield two species of Jiulongshania among other species. Species of Jiulongshania have been known to occur successively in North China, so are useful for detailed correlation. Specimens of Stephanocare richthofeni are fragmentary and rarely occur in association with Jiulongshania regularis, while Jiulongshania species occur throughout the studied intervals. Accordingly, it is reasonable to extend the previously established Jiulongshania Zone of the uppermost part of the underlying Daegi Formation into the lower part of the Sesong Formation. By doing so, the Jiulongshania Zone is correlated with the Blackwelderia Zone of North China with confidence. The lowermost part of the Sesong Formation in the Jikdong section yields a fauna including J. regularis, which implies that the boundary between the Daegi and Sesong formations is diachronous within the Taebaeksan Basin. The Daegi/Sesong formation boundary in Korea is comparable to the Zhangxia/Gushan boundary in North China in that it displays an abrupt change from a carbonate-dominant facies to a shale-dominant facies. The correlation employing the Jiulongshania species indicates that the facies shift occurred significantly earlier in Shandong, North China than in the Taebaeksan Basin, Korea.
A paleoecological study of benthic foraminifera through the lower Hettangian in the Doniford Bay section (west Somerset, U.K.) is presented. The sudden and brief appearance of Oberhauserellidae in the aftermath of the Late Triassic extinction is defined as a proxy for environmental perturbations indicating severe biotic stress conditions. Oberhauserellidae, associated with the genus Reinholdella are distinguished from other species by a high abundance, low diversity, high dominance and an abnormally small size. This suite of characters mimics an opportunistic behavior where these r-strategists and grazer feeders maximize their full ecological potential at a time of low-oxygen conditions on the sea-floor and a high food supply: both of which appear to be the main triggers of this paleoecological change. The disappearance of these opportunistic benthic foraminifera coincides with the appearance of infaunal, low-oxygen-tolerant generalists, and the restoration of stable environmental conditions (e.g., well-stratified water mass and oligotrophic conditions), characterizing the initial stages of recovery following the Late Triassic extinction event.
Quaternary raised marine terraces containing the remains of diverse, shallow water marine invertebrate faunas are widespread across the coast of Angola. These deposits and faunas have not been studied in the same detail as contemporaneous features in northwest and southernmost Africa. We analyzed the fossil assemblages and sedimentology of two closely spaced middle Pleistocene marine terrace deposits in Baía das Pipas, southwest Angola. This revealed 46 gastropod and 29 bivalve species, along with scleractinian corals, encrusting bryozoans, polychaete tubes, barnacles, and echinoids. The fauna is characteristic of intertidal and nearshore rocky substrates and sandy soft-bottom habitats. Sedimentological analysis is consistent with faunal data and indicates an upper shoreface paleoenvironment along a gravel coast. This diverse fauna stands out as a rare example of a marine Pleistocene assemblage from over 6,000 km of the West African coast. The assemblage is dominated by extant tropical West African molluscs, including species from the “Senegalese fauna” that colonized northern Africa and beyond during Pleistocene interstadials. Additionally, as along the modern coast of the Namibe Desert, the influence of the cool-water Benguela Current is apparent in the paleofauna by the occurrence of a few temperate species. The distribution and thermal tolerances of extant species identified in the Pipas fauna indicate that this region experienced similar climatic and oceanographic conditions as that of the present during this interstadial. Seasonal temperature varied between ∼20 and 28°C and resulted from upwelling in this tropical setting.
Rare scales of agnathan thelodonts Paralogania ludlowiensis and Thelodus sp. cf. T. parvidens, or alternatively Thelodus macintoshi, and acanthodian fishes Nostolepis striata, Gomphonchus sp., Gomphonchoporus sp. aff. G. hoppei, and Machaeraporus stonehousensis (Legault) n. gen., plus acanthodian fin spines, teeth and tooth whorls have been identified from the upper Moydart and Stonehouse formations near Arisaig, Nova Scotia. The assemblage agrees well with the late Silurian (uppermost Ludlow–Přídolí) age assigned to these strata based on invertebrate assemblages, and the vertebrate taxa show affinity with British and Baltic faunas of this age. All vertebrate-bearing strata were deposited in various positions off the western shores of the Avalonia terrane during or after its collision with Laurentia.
A new fossil ant genus, Bilobomyrma new genus, and two new species are described based on males from the late Eocene Rovno (B. ukrainica n. sp.) and Baltic (B. baltica n. sp.) ambers. We tentatively place this genus in the myrmicine tribe Formicoxenini. Bilobomyrma is characterized by its 13-segmented antennae without an apical club; by the short scape, which is subequal to the length of the first and second funicular segments together; by the shape of the second funicular segment, which is distinctly longer than the any other funicular segment except for the apical one; by the presence of notauli on the scutum; by the absence of spurs on the middle and hind tibiae. At the same time, Bilobomyrma differs from other myrmicine genera by the peculiar shape of its clypeus, having a strongly incised medially, bilobed anterior margin, and its forewing venation: the wings have three closed cells—mcu, 12r and 3r; the cell 3r is very short, only twice as long as its width; the distal section of veins RS and M diverge from the cell 1 2r separately.
Burgess Shale-type faunas provide unique insights into the Cambrian “explosion”. Their degree of representativeness of Cambrian marine life in general is, however, less easy to establish. One line of evidence is to consider only the skeletal component of a Burgess Shale-type fauna and compare that with a typical Cambrian assemblage. This paper describes a new species of helcionelloid mollusk (Totoralia reticulata n. sp.) from the middle Cambrian Burgess Shale of British Columbia. Whilst much rarer than the co-occurring smooth shelled helcionelloid Scenella amii, the strongly costate morphology of Totoralia reinforces comparisons with Cambrian shelly faunas. The extension of the range of Totoralia from Argentina to Canada adds support to the proposed derivation of the Precordillera terrane of Mendoza from Laurentia.
Permian rocks from the Indonesian island of Timor contain an abundant and diverse pelmatozoan echinoderm fauna that was extensively monographed in the first half of the twentieth century. Recent collections have produced specimens of Corrugatoblastus savilli, n. gen. n. sp., the first new genus of blastoid described from Timor in 70 years. Corrugatoblastus savilli is a ridged and furrowed, conical, fissiculate blastoid with a highly unusual thecal morphology mimicking a small, solitary, rugose coral. We have placed C. savilli in the Family Codasteridae, although it has several morphological features that are unique when compared to other genera in the family and to other blastoid genera regardless of familial assignment.
Extraordinarily well-preserved pithonellid microfossils (calcitarchs, “calcispheres”) from the Turonian (upper Cretaceous) of Tanzania reveal previously unknown morphological traits, crystallographic patterns, and chemical signatures, providing new insight to this enigmatic group of microfossils. Using combined transmitted-reflected light microscopy, scanning electron microscope imagery, electron microprobe elemental analysis and stable isotope geochemistry, the present study reveals four new aspects of the genus Pithonella, notably, the following. An affinity with cyst-forming organisms, potentially the dinoflagellates, is indicated by the presence of a hatch opening and corresponding operculum. The pristine outer wall architecture consists of thin, smooth shingle-shaped plates with regular rows of slit-shaped pores and an apical sub-angular or circular pore. This primary surface pattern is significantly different from previous descriptions of an outer wall consisting of “parquet-shaped” prismatic crystal rows; this latter surface pattern is formed by secondary overgrowth. The crystallographic pattern of the inner wall is crypto-crystalline. Unaltered pithonellids reveal a calcite chemistry characterized by comparably high Mg-contents, relatively enriched stable carbon isotope values, and stable oxygen values indicating a surface water habitat. Based on these previously unseen traits, the diagnosis of the genus Pithonella is emended. A new species, Pithonella diconica, is described from the lower-middle Turonian sediments of Tanzania.
The orthidine brachiopod genera Plaesiomys and Hebertella are significant constituents of Late Ordovician benthic marine communities throughout Laurentia. Species-level phylogenetic analyses were conducted on both genera to inform systematic revisions and document evolutionary relationships. Phylogenetic analyses combined discrete and continuous characters, from which character states were determined using a statistical approach, and utilized both cladistic and Bayesian methodologies. Plaesiomys cutterensis, P. idahoensis, and P. occidentalis are herein recognized as distinct species rather than subspecies of P. subquadratus. Similarly, Hebertella montoyensis and H. prestonensis are recognized as distinct species separate from H. occidentalis, and H. richmondensis is recognized as a distinct species rather than a geographical variant of H. alveata. Hebertella subjugata is removed from its tentative synonymy with H. occidentalis and revalidated.
The development of species-level evolutionary hypotheses for Plaesiomys and Hebertella provides a detailed framework for assessing evolutionary and paleobiogeographic patterns of Late Ordovician brachiopods from Laurentia. The geographic range of Hebertella expanded throughout Laurentia during the Richmondian into both intracratonic and marginal basins. Plaesiomys subquadratus participated in the Late Ordovician Richmondian Invasion. The recovered phylogenetic topology for Plaesiomys suggests that P. subquadratus may have migrated into the Cincinnati region from a basin situated to the paleo-northeast.
Discovery of well-preserved specimens from the Nanba section in Yiyang, Hunan Province, in combination with a literature review, enabled us to re-evaluate and revise the graptolite genus Ancoragraptus. This is a genus of biradiate, multiramous anisograptids with horizontal to reclined rhabdosomes and free lower part of the metasicula and slightly isolated autothecal apertures. According to the revised definition, two species are included in Ancoragraptus, i.e., Ancoragraptus bulmani (Spjeldnæs, 1963) and Ancoragraptus psigraptoidesCho, Kim, and Jin, 2009. It is the first time that A. bulmani has been reported from China. The occurrences of Ancoragraptus reported worldwide are reviewed in the present study and found to be restricted to the lower upper Tremadocian. The restriction of Ancoragraptus in stratigraphical distribution makes it a taxon with a high potential for precise biostratigraphical correlation at both regional and global scale.
Isolated teeth from the Middle Permian (early Guadalupian) Kaibab Formation of Arizona are described as a new species of the xenacanth shark genus Bransonella. Bransonella tribula n. sp. is a small tooth in which the intermediate cusp is 65% of the length of the principal cusps and the cristae on the labial face extend down over the base, covering it, and bifurcating to form distinctive double crested ridges. Fin spines from the same localities in the Kaibab Formation show the characteristic xenacanth feature of a double row of large thorn-like denticles along the posterior margin. Bransonella tribula n. sp. is the only xenacanth shark known from the Kaibab Formation at present, however, due to the lack of articulated material the fin spines are attributed to ?Bransonella tribula n. sp. The ecomorphology of Bransonella suggests a primitive, small, gracile, marine xenacanth that fed near the sea floor like the modern catsharks (Scyliorhinidae).
Representatives of the family Inaniguttidae dominate a diverse and well-preserved radiolarian assemblage from Kazakhstan. The fauna was extracted from a limestone sample of the Shundy Formation, a limestone sequence that accumulated on the slope of a carbonate platform and which crops out in the Aksuran Mountains (North Balkhash Region). The family Inaniguttidae is represented in the studied assemblage by five genera and 14 species (including two new species, Triplococcus aksuranensis and Inanibigutta maletzi). The genus Triplococcus is particularly abundant, representing half of the Inaniguttids in the studied assemblage. Based on the presence of species Haplentactinia juncta, the fauna can be correlated with the upper Darriwilian Haplentactinia juncta–Inanigutta unica assemblage. Identified trilobites (Endymionia semielliptica and Porterfieldia sp. aff. P. delicata) found in the same sample also suggest a late Darriwilian age, which agrees with the age suggested previously by graptolites.
We report on the faunal transition of benthic foraminifera during the middle Eocene at Site U1333 (4862 m water depth, 3,560–3,720 m paleo-water depth) of Integrated Ocean Drilling Program Expedition 320 in the eastern equatorial Pacific Ocean. During the period ∼41.5–40.7 Ma, which includes carbonate accumulation event 3 (CAE-3), the benthic foraminiferal accumulation rate (BFAR) increased gradually and then it declined rapidly. In contrast, BFAR was considerably lower during ∼40.7–39.4 Ma, corresponding to the middle Eocene climatic optimum (MECO), and then it increased during ∼39.3–38.4 Ma, including CAE-4. Diversity (E [S200]) was slightly lower in the upper part of the study interval than in the lower part. The most common benthic foraminifera were Nuttallides truempyi, Oridorsalis umbonatus, and Gyroidinoides spp. in association with Globocassidulina globosa and Cibicidoides grimsdalei during the period studied. Quadrimorphina profunda occurred abundantly with N. truempyi, O. umbonatus, and G. globosa during ∼39.4–38.4 Ma, including CAE-4, although this species was also relatively common in the lower part of the study interval. Virgulinopsis navarroanus and Fursenkoina sp. A, morphologically infaunal taxa, were common during ∼38.8–38.4 Ma, corresponding to the late stage of CAE-4. Based on Q-mode cluster analysis, four sample clusters were recognized and their stratigraphic distributions were generally discriminated in the lower and upper parts of the study interval. Thus, there was only a small faunal transition in the abyssal eastern equatorial Pacific during the middle to late-middle Eocene. The faunal transition recognized in this study may be related to recovery processes following intense carbonate corrosiveness in the eastern equatorial Pacific during MECO.
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As one of the highly diversified radiolarian groups in the Mesozoic, Vallupinae occurred widely in the tuffaceous claystone from the Mariana Trench of the upper Tithonian–Berriasian where the specimens are extremely well-preserved. The occurrence of diversified Vallupinae presented here is further evidence that the range of Vallupinae extends to Berriasian. The stratigraphic range and proposed possible phylogenetic relationships within the Subfamily Vallupinae are provided and 17 radiolarian species of this subfamily are presented, including three new species: Vallupus gracilis, Mesovallupus brevispina, and Neovallupus mediforma.