Influence of ambient flow around the horseshoe crabLimulus polyphemus on the distribution and orientation of selected epizoans

Cumulatively scored distributions of the slipper shell gastropodCrepidula sp., the sand-building polychaete wormSabellaria vulgaris, the calcareous meandering tubed polychaeteFilograna implexa, the barnacleSemibalanaus balanoides, and the lacy bryozoanSchizoporella sp. were mapped on a cm-square gridded silhouette of the prosoma, opisthosoma, and movable spines, and telson of the horseshoe crabLimulus polyphemus stranded on Reeds Beach, Cape May County, New Jersey in the Delaware Bay. Statistically tested and contoured epizoan frequencies for 81 males and 59 females revealed that slipper shells and agglutinated worm tubes are concentrated near the movable spines on the opisthosoma, but rare on recessed chilarial or opercular pleurites, the prosoma, and telson. Barnacles are disproportionately concentrated in entapophyseal pits on the opisthosoma and in the longitudinal furrows flanking the cardiac lobe of the prosoma. Bryozoans are concentrated near movable marginal spines of the opisthosoma, but also found on all areas of the prosoma, opisthosoma, and hinge area in between, as well as the telson. Different morphologic sites for their respective modal concentrations may result from competitive exclusion of bryozoans by barnacles. Genal angles have lower statistically predicted frequencies of bryozoans, barnacles, and sand worm tubes. Apertural openings of calcareous tubes are predominantly posteriorly oriented whereas the anterior magin of slipper shells are randomly oriented on the opisthosoma. Eschewing differential mortality, nonrandom distributions indicate possible: rugophilic (groove-seeking) larval settlement behavior for barnacles that preferred the recessed hinge area, longitudinal grooves, and pits; rheophilic (current-seeking) larval settlement behavior for the gastropods and tube-building polychaetes that filter feed in the eddies generated near the movable spines of the downcurrent-sloping opisthosoma; and geotactic (gravity-influenced) larval settlement or post-settlement migration to the vaulted crest (cardiac lobe). Deficiencies of protruding epizoic skeletons on the genal angles and anterior margin of the host may be due to removal by abrasion as the horseshoe crab plowed through sediment or shearing of the epizoans from the carapace substratum in strong currents or waves.     

淡水鲎虫一个新科——新疆鲎虫科

文中描述了我国中三叠世淡水鲎虫的1个新科——新疆鲎虫科(Xinjiangiopsi-dae fam.nov.)及陕西铜川中三叠世(铜川组)鲎虫4个新种。     

A giant limulid trackway (<Emphasis Type="Italic">Kouphichnium lithographicum</Emphasis>) from the lithographic limestones of Cerin (Late Kimmeridgian,France): ethological and environmental implications

A well-preserved large limulid trackway related to Kouphichnium lithographicum is described from the Upper Kimmeridgian Lithographic Limestones of Cerin (Ain, France). It is three meters long and exhibits ten successive sets of imprints characterized by a pair of well-preserved tetradactyl external imprints of legs VI (named pushers) and a variable number (mostly three pairs) of straight thin intermediate imprints (legs III–V). The trackway is unusually wide and was probably made by a very large adult approximately 38 cm wide and 80 cm long when related to the proportions of the modern Limulus polyphemus. The straightness of the trackway, the great repeat distance (stride), the length of series of leg imprints, their slight angle to the mid-line, the well-marked pusher (= leg VI) imprints and the absence of groove made by the telson all indicate fast progression of a vigorous animal on the sea-floor. This walking trackway can be easily differentiated from other limulid crawling, ploughing, and dying trackways usually found in lithographic limestones or other deposits of the geological record. Main characteristics of these different limulid trackways are evidenced. The occurrence of a living adult in the Cerin restricted lagoon is probably related to a brief period of connection with the open sea. This occurrence indicates either a passive introduction of the animal into the lagoon or a spawning behaviour.     

An Important Ornithischian Tracksite in the Early Jurassic of the Shenmu Region,Shaanxi, China

A large track site with multiple,well-preserved trackways of an Early Jurassic quadrupedal ornithischian dinosaur is the first report of Moyenisauripus-like trackways from Asia,herein named Shenmuichnu...     

Early Cambrian Ichnofossils from the Mussoorie Syncline and revision of Trace Fossil Biozonation of the Lesser Himalaya,India

A new locality bearing ichnofossils of the Cruziana Assemblage Zone-Ⅲ from the Mussoorie syncline,Lesser Himalaya,is located in rocks of Member-B of the Dhaulagiri Formation,Tal Group,exposed along the Maldewta-Chhimoli fresh road cut section.The site yielded ichnofossils Bergaueria perata,Cochlichnus anguineus,？Diplocraterion isp.,Dimorphichnus obliquus,diplichnitiform Cruziana bonariensis,Diplichnites gouldi,Glockeria isp.,Helminthopsis isp.,Monomorphichnus lineatus,Phycodes palmatum,Palaeophycus striatus,Planolites beverleyensis,Planolites montanus,Treptichnus cf.T.pedum,scratch marks and an undetermined worm impression.An Early Cambrian age （Cambrian Series 2） is assigned to the ichnofossil-bearing strata based on the stratigraphic position between the Drepanuroides and Palaeoolenus trilobite zones.A revised Cambrian ichnofossil zonation is presented for the Tal Group of the Mussoorie syncline.Together with their occurrence on rippled surfaces,and the lateral displacement of some trackways （due to current action）,a sub-aqueous shallow-marine depositional setting is proposed for the rocks of Member-B.     

Yeosu dinosaur track sites of Korea: The youngest dinosaur track records in Asia

Eighty two dinosaur trackways were newly discovered in Upper Cretaceous lacustrine deposits on islands in the vicinity of Yeosu, Korea. Most dinosaur tracks occur in marginal lake deposits with polygonal desiccation cracks. The dinosaur tracks at the Yeosu site include 65 ornithopod trackways, 16 theropod trackways and one sauropod trackway. The prevalence of ornithopod tracks and the limited occurrence of sauropod tracks at the Yeosu site evidently reflect decreased sauropod diversity in the Upper Cretaceous. All ornithopod trackways represent bipeds, and most of the ornithopod tracks are similar to Caririchnium from other sites of the Korean peninsula. All fossil wood specimens collected in the study area represent conifers (three species of cupressaceous and two species of taxodiaceous conifers, and a new species) except for one, which is a discotyledon. It is thus inferred that the southwestern part of the Korean Peninsula was primarily covered with mesic forests with taxodiaceous trees during the Late Cretaceous. The K–Ar age of the Yeosu tracksite is determined as 81–65 Ma (Camapnian to Maastrichtian). It indicates that the Yeosu track site contains the last records of dinosaurs living in Asia. Consequently, semi-arid palaeoclimatic conditions, together with a large lake as a persistent water source and rich vegetation of gymnosperm trees as food, resulted in the preservation of abundant dinosaur tracks in the Upper Cretaceous on the Korean Peninsula.     

宁夏六盘山地区早白垩世恐龙足迹研究

本文记述了发现于宁夏六盘山地区的恐龙足迹化石。共计5个化石点,分别位于宁夏六盘山地区固原市隆德县和泾源县,赋存地层为下白垩统六盘山群和尚铺组、李洼峡组和马东山组,包括蜥脚类和鸟脚类恐龙足迹化石,其中以蜥脚类为主。足迹可识别的有4条行迹,其中,隆德县香水镇行迹所代表的恐龙个体较小,臀高仅1.44m,其余3组行迹代表的恐龙较大,臀高在2.79～3.27m之间;鸟脚类恐龙臀高约为0.48 m,为小型鸟脚类恐龙。采用Alexander的速度计算方法对造迹恐龙的速度进行研究,表明六盘山地区蜥脚类造迹恐龙速度为0.34～1.41m/s之间,复步长与臀高的比值小于2,均为正常行走状态。足迹保存层面岩性主要为粉砂质泥岩、细砂岩,具水平层理、波痕、泥裂、雨雹痕等沉积构造,含植物碎片化石,并发育有大量的无脊椎动物潜穴化石。根据岩性及沉积构造判断,造迹恐龙活动环境为离湖岸较近的浅水地带。     

世界二叠纪生物礁的基本特征及其古地理分布

世界上早二叠世生物礁分布于泛大陆的西北陆棚、乌拉尔山脉的西侧、美国的二叠盆地等地,并集中分布于前两个地区。其早期,以Palaeoaplysina礁或Palaeoaplysina和叶状藻礁占优势,而至晚期则形成以Shamovella (Tubiphytes)和苔藓虫为骨架的礁。中二叠世栖霞期的礁仅发现于北美格拉斯山脉、帕米尔和我国的阿尔格山等地。茅口期的礁是世界上最发育的生物礁之一,北美瓜德罗普山脉的二叠纪礁已成为世界上最典型的礁,其相带分异之清晰堪称为世界之最。北非突尼斯的礁也是研究程度较高的礁之一。中二叠世的礁以海绵、苔藓虫、Shamovella的大量出现为特征,古石孔藻、Shamovella和笛苔藓虫是常见的包覆生物。晚二叠世生物礁分布于欧洲镁灰岩统盆地、特提斯海西缘和最北缘的陆棚以及特提斯海域内的一些地体。我国晚二叠世礁十分发育,成为世界二叠纪礁的一个亮点。礁内的造架生物以珊瑚海绵为主,包括房室海绵和纤维海绵,古石孔藻和Shamovella (Tubiphytes)作为常见的包覆联结生物。     

吐哈盆地及邻区早二叠世沉积特征与构造发育的耦合关系

通过对不同露头剖面和探井资料的分析,在吐哈盆地及邻区区分出三种不同类型沉积相：（1）裂谷型海相火山喷发沉积相;（2）裂谷型陆相火山喷发沉积相;（3）造山带磨拉石相;不同沉积相特征并结合构造分析认为,早二叠世在吐哈及邻区存在二种不同类型的盆地,即前陆盆地和裂谷盆地,同时,还探讨了大陆一大陆碰撞带上两种不同类型盆地的形成机理。     

New dinosaur tracks from Korea, Ornithopodichnus masanensis ichnogen. et ichnosp. nov. (Jindong Formation, Lower Cretaceous): implications for polarities in ornithopod foot morphology

Twelve trackways of ornithopods from Lower Cretaceous lacustrine margin deposits of the Jindong Formation represent new dinosaur trackways described from Korea. The site, discovered during highway construction, was rescued by removing the most important trackways to the Korean Natural Heritage Center in Daejeon, where they are on permanent display. The new ichnotaxon Ornithopodichnus masanensis ichnogen. et ichnosp. nov. is recognized as a distinctive robust tridactyl track, slightly wider than long (l/w ratio = 0.91), with positive (inward) rotation. The toe prints are very thick, broad and U-shaped, resulting in a trefoil outline with a smoothly rounded hind margin. Digit III is short and projects anteriorly much less than digit II and IV (= weak mesaxony). Divarication of digits II-IV is about 70° with interdigital angle II-III larger than III-IV. Trackway width is narrow and the stride length/track length ratio is about 4.2–4.6. The Ornithopodichnus trackways evidently represent gregarious blunt-toed Iguanodon-like bipedal ornithopods, although poorly preserved manus traces are discerned in a few trackways. Ornithopodichnus is distinct from other well known iguanodontid tracks that display much stronger mesaxony and indicates a polarity in ornithopod foot morphology that can be verified by reference to known foot skeletons.     

A new Cenomanian vertebrate tracksite at Tamajón (Guadalajara,Spain): Palaeoichnology and palaeoenvironmental implications

A new Upper Cretaceous vertebrate tracksite has been discovered at Tamajón (Iberian Ranges, Guadalajara, Spain). The track level is a relatively smooth and slightly undulating sandy ferruginous crust, corresponding to an erosive surface at the base of a small meandering channel. It is incised into the underlying planar cross-bedded sandstones of coastal bars located at the middle-upper part of the Utrillas Formation (middle-upper Cenomanian). The site shows an extraordinary concentration of vertebrate tracks, among which numerous sets of two to five isolated digit impressions (“swim tracks”) and, at least, two trackways referred to crocodyliforms, and a single tridactyl footprint probably produced by a theropod dinosaur can be recognized. There are also several long traces (epichnial grooves) revealing sharp direction changes (up to 90°) which seem to be fish fin traces (Undichna unisulca), although crocodyliforms (tail marks) and/or fish invertebrates cannot be rejected as possible tracemakers. Some crocodyliform tracks reveal a thin raised rim, due to the displacement of the sediment by the pressure produced by the feet. Several impressions are moderately deformed by small sediment slides, only preserving their deepest part (claw marks). This is clearly indicative of a soft substrate with a high degree of plasticity and water content at the time of the track registration. Nevertheless, the sediment was hard enough to preserve manus and pes print morphologies and also possible crocodyliform tail and/or fish fin traces. Small rhizoliths can also be recognized and may belong to herbaceous wetland vegetation. The morphology of the palaeochannel, the sedimentary context and the track preservation seem to indicate that the tracks were impressed in a shallow channel located near the coast, under wet conditions and in different moments of time. This discovery represents the first occurrence of vertebrate ichnites in the Utrillas Formation, a stratigraphic unit where osteological and ichnological remains are relatively scarce, and it confirms that some crocodyliforms lived in near coast channels during the deposition of this unit.     

Die Grödener Schichten (Perm,Südalpen). Sedimentologische und geochemische Untersuchungen zur Unterscheidung mariner und kontinentaler Sedimente

Within the sedimentation area of the Gröden formation two facies units have been observed: The attributes of one facies correspond with a deposition in continental environment. Kaolinitic weathering and fresh-water algae support this interpretation. The other facies is deposed in marine environment (supply of Mg for dolomitization and formation of chlorites, scarce forams). The transition facies shows characteristics of a costal region with marginal lagoons. Paleontological data (tetrapod traces, drifted cephalopods and forams) are in accordance with this model. The paleogeographical development during the Middle Permian is as follows: Continental sedimentation at the basis, marine transgression in the Carnic Alps and the western Karawanken during lower Gröden stage, progressiv transgression during the middle Gröden stage to the west with a coast line near Gröden valley and a coast line near River Etsch at the end of the Middle Permian.     

Discussion of “Permian—Triassic vertebrate footprints from South Africa: Ichnotaxonomy,producers and biostratigraphy through two major faunal crises” by Marchetti,L., Klein,H., Buchwitz,M., Ronchi,A., Smith,R.M.H., DeKlerk,W.J., Sciscio,L., and Groenewald,G.H. (2019)

A newly devised tripartite subdivision of the Karoo Basin, South Africa, is based on a number of vertebrate-trackways spanning the Middle Permian (Wordian/Capitanian) to Lower Triassic (reportedly Induan) rocks of the Beaufort Formation. The youngest Footprint Assemblage (FA III) is reported to occur in the lowermost Triassic rocks of the Palingkloof Member. One collection site from which seven trackways are reported, Bethel farm, is a classic South African locality where some authors place the end-Permian extinction event and, coincidentally, the vertebrate-based terrestrial Permian—Triassic Boundary. Published GPS coordinates for the Bethel farm site, though, indicate that these trackways originate from exposures of the Burgersdorp Formation near the town of Aliwal North. Outcrops here are assigned to the Cynognathus Assemblage Zone, and indicate a Middle Triassic age assignment. In addition, we caution the use of any small temporally or spatially restricted dataset, which is of severely limited utility, in making broad generalizations about paleoenvironmental or paleoecological interpretations.     

Stratigraphic and palaeoenvironmental framework of the Early Permian sequence in the Salt Range, Pakistan

The Early Permian Gondwana regime succession of the Nilawahan Group is exposed only in the Salt Range of Pakistan. After a prolonged episode of non-deposition that spanned much of the Palaeozoic, the 350?m thick predominantly clastic sequence of the Nilawahan Group records a late glacial and post-glacial episode in which a range of glacio-fluvial, marine and fluvial environments evolved and accumulated. The Early Permian succession of the Salt Range has been classified into four formations, which together indicates a changing climatic regime during the Early Permian in the Salt Range region. The lower-most, Tobra Formation unconformably overlies a Cambrian sequence and is composed of tillite, diamictite and fresh water facies, which contain a floral assemblage (Gangamopteris and Glossopteris) that confirms an Asselian age. The Tobra Formation is overlain by marginal marine deposits of the Dandot Formation (Sakmarian), which contain an abundant brachiopods assemblage (Eurydesma and Conularia). Accumulation of the Dandot Formation was terminated by a regional sea-level fall and a change to the deposition of the fluvial deposits of the Warchha Sandstone (Artinskian). The Warchha Sandstone was deposited by high sinuosity meandering, avulsion prone river with well developed floodplains. This episode of fluvial sedimentation was terminated by a widespread marine transgression, as represented by the abrupt upward transition to the overlying shallow marine Sardhai Formation (Kungurian). The Early Permian Gondwana sequence represented by the Nilawahan Group is capped by predominantly shallow shelf carbonate deposits of the Tethyan realm. The sedimentologic and stratigraphic relationship of these four lithostratigraphic units in the Salt Range reveals a complex stratigraphic history for the Early Permian, which is mainly controlled by eustatic sea-level change due to climatic variation associated with climatic amelioration at the end of the major Gondwana glacial episode, and the gradual regional northward drift to a lower latitude of the Indian plate.     

Occurrence of faunal coal balls in Gondwana sediments (Permian) of Arunachal Himalaya,India

The nature and genesis of fossiliferous inorganic concretions present within the coal and carbonaceous beds of Garu Formation, Arunachal Pradesh have been studied in detail. These concretionary bodies are described here as “coal balls” due to their close similarity with the true coal balls reported from European and American coals. Further, the presence of coal balls in this area also indicates that the sediments of the Garu Formation were deposited in shallow marginal swamps subjected to marine incursions during the Lower Permian.     

下扬子句容地区海相上组合油气地质条件评价

下扬子句容地区的构造形成演化可以分为海相盆地稳定隆降、逆冲推覆、拉张断陷3个阶段。受盆地形成演化的控制,句容地区海相上组合烃源岩主要为盆地构造稳定阶段沉积的二叠系泥岩及下三叠统青龙群泥岩、碳酸盐岩;储层主要为侏罗系象山组、二叠系龙潭组砂岩储集层和三叠系青龙群碳酸盐岩储集层;区域性盖层为白垩系浦口组泥岩。虽然该区海相地层遭受了多期次不同方式、不同程度的改造作用,对油气成藏条件产生了很大影响和破坏,但研究认为在后期改造相对薄弱地区如该区后缘凹陷,仍具有良好的勘探前景,评价出索墅镇—黄梅桥—陈武地区和句容—行香地区2个有利区块。     

Early Jurassic basal sauropodomorpha dominated tracks from Guizhou,China: Morphology,ethology, and paleoenvironment

The newly discovered large(350 m^2) Yantan dinosaur tracksite, in the Lower Jurassic Ziliujing Formation of Guizhou Province, China, reveals at least 250 footprints of which ~97 can be resolved into trackways of sauropodomorphs. All the trackways are sub parallel likely indicating gregarious behavior. One theropod track(cf. Grallator) was recorded. The sauropodomorph tracks predominantly represent quadrupedal progression(Morphotype A), and footprint morphology is similar to the ichnospecies Liujianpusshunan, characterized by outward pes rotation. Three trackways indicate bipedal progression, and two of these(Morphotype B) indicate inward pes rotation, accompanied by elongate pes digit scratch marks. For the latter phenomenon three possible scenarios are discussed:(1) significant rotation changes accompanying changes in gait,(2) swimming behavior,(3) formation of undertracks.Sedimentological evidence indicates the tracks were made on a linguloid rippled, muddy, immature sandstone substrate characterized by significant differences in substrate consistency across the trackbearing surface. Microbially induced sedimentary structures(MISS) characterized by distinctive wrinkle marks indicate a stressed, probably semi-arid, paleoenvironment that was not conducive to habitation by invertebrate organisms. This is consistent with other evidence that Lower Jurassic sauropodomorph tracks are often associated with semi-arid paleoenvironments.     

Guanling Biota: A Marker of Triassic Biotic Recovery from the end-Permian Extinction in the Ancient Guizhou Sea

After a slow recovery from the end-Permian extinction during the Early Triassic and rapid radiation in the Middle Triassic, evolution of organisms reached a new peak phase in the Late Triassic. The Guanling Biota from the Wayao Member （conodont Paragondolella polygnathiformis Zone）, Falang Formation, Xinpu, Guanling County, Guizhou Province, southwestern China corresponds to this peak that marks the full recovery from the end-Permian extinction of marine ecosystems. The biota is of high diversity, containing well preserved and completely articulated skeletons of vertebrates comprising marine reptiles, fishes, and invertebrates including crinoids, ammonites, bivalves, and other fossils, and is one of the best examples of marine ecosystem records in life history. The fossil marine reptiles and crinoids are most significant in this biota, especially the marine reptiles, which provide an important link between the Triassic Pacific and Tethys, and between Triassic basal forms and the Jurassic-Cretaceous marine top predators. The most remarkable fossils are the large completely articulated ichthyosaur skeletons up to and more than 10 m, and the first recorded thalattosaurs and placodonts in China. Following our review, of the 17 named reptilian taxa the eight listed here are considered to be valid： three ichthyosaurs （Qianichthyosaurus zhoui Li, 1999; Guizhouichthyosaurus tangae Cao and Luo in Yin et al., 2000, Guanlingsaurus liangae Yin in Yin et al., 2000）, three thalattosaurs （Anshunsaurus huangguoshuensis Liu, 1999, Xinpusaurus suni Yin in Yin et al., 2000, Xinpusaurus kohi Jiang et al., 2004）, and two placodonts （Sinocyamodus xinpuensis Li, 2000, Psephochelys polyosteoderma Li and Rieppel, 2002）. Mixosaurus guanlingensis Cao in Yin et al., 2000 might be a junior synonym of Qianichthyosaurus zhoui Li, 1999, and Cymbospondylus asiaticus Li and You, 2002 and Panjiangsaurus epicharis Chen and Cheng, 2003 might be junior synonyms of Guizhouichthyosaurus tangae Cao and Luo in Yin et al., 2000. It needs to re-describe the holotypes after a complete preparation for clarifying the taxonomic status of Typicusichthyosaurus tsaihuae Yu in Yin et al., 2000, Xinpusaurus bamaolinensis Cheng, 2003, Neosinosaurus hoangi （Zhou in Yin et al., 2000）, Wayaosaurus geei Zhou in Yin et al., 2000, Wayaosaurus bellus Zhou in Yin et al., 2000 and Placochelys ？ minutus Yin and Luo in Yin et al., 2000.     

Quartz [0001]-axes preferred orientation, Bluff, New Zealand: Origin elucidated by grain-size measurements

Permian volcanic sediments at Bluff have been strained and thermally metamorphosed by Permian intrusives to metasediments of hornblende—hornfels facies. Quartz, which crystallised as a secondary mineral during metamorphism, has an unusual preferred orientation with c-axes either forming paired maxima in the plane containing the lineation (=maximum principal strain axis = direction of extension) and the perpendicular to schistosity (=minimum principal strain axis = shortening direction) or a broad maximum parallel to the lineation; the paired maxima are approximately 30° either side of the lineation. Some quartz grains are markedly elongate parallel to the lineation, and according to hypotheses of preferred orientation involving crystal plasticity, there should be some correlation between the shape of such grains and their c-axis orientations. Grain-size and shape analysis of Bluff quartz demonstrate that no such correlation exists; the analyses show that the preferred orientation results from oriented nucleation in the residual stress field immediately following the bulk straining of the rocks, with the distribution of c-axes as predicted by Kamb's hypothesis (1959). The time relationships of rock deformation, thermal metamorphism, and nucleation and growth of quartz are discussed.     

滇黔桂盆地早二叠世亚丁斯克期的沉积环境分异与䗴有孔虫响应

亚丁斯克期华南滇黔桂地区沉积环境分异明显,主要发育局限或半局限海台地沉积区、开阔海台地沉积区和台地边缘沉积区,不同沉积区䗴有孔虫动物群面貌也有所区别。本文详细研究了黔南地区的猴场打望剖面和惠水竹子冲剖面阿瑟尔期—亚丁斯克期的地层,在打望剖面识别出4个䗴类化石带,由老至新为Pseudoschwagerina beedei-Sphaeroschwagerina constans共存延限带、Staffella xiushuiensis-Eoparafusulina contracta富集带、Pseudofusulina ellipsoidalis富集带和Misellina subcycloidea间隔带。竹子冲剖面识别出3个化石带,由老至新为Pseudoschwagerina uddeni-Sphaeroschwagerina sphaerica组合带、Eoparafusulina certa富集带和Staffella discoides延限带。对滇黔桂地区不同沉积相区剖面䗴类动物群以及沉积相特征的综合研究显示:开阔台地或台地边缘环境的䗴动物群以Pamirina为特征分子,同时还产出丰富的希瓦格䗴亚科(Schwagerininae)分子;局限或半局限台地环境的䗴类以史塔夫䗴亚科分子(Staffellininae)为主。沉积环境对䗴有孔虫的影响可能主要来自水动力、光照和氧气含量。个体较大、隔壁褶皱强烈的纺锤形希瓦格䗴亚科分子更适应水动力强、透光性好、氧气充足的水体;小个体、结构简单的史塔夫䗴亚科分子可在透光性差、含氧量较低的水体中生存。