Cavity-dwelling organisms in Lower Cambrian patch reefs from southern Labrador

An intact and well-preserved biota of cavity-dwelling organisms (coelobionts) of early Cambrian age is found within growth-framework cavities in archaeocyathid patch reefs of the lower Forteau Formation (upper Bonnia-Olenellus Zone) of southern Labrador. The biota was diverse, and consisted of encrusting and vagrant organisms. Renalcis , a Renalcis-like form, Epiphyton, Girvanella , and Serligia , all lived attached to walls and roofs of the cavities; other attached forms were Bija, Archaeotrypa , and two types of Wetheredella , an agglutinated foraminifer. Other organisms, including trilobites, probable calcareous ostracodes, brachiopods, echinoderms, and worms, as well as fungi, are also found preserved within the cavities. The structure of the coelobiontic community is surprisingly modern; it indicates that as early as the Lower Cambrian coelobiontic organisms were well-established in reefs, and were an important part of the reef community as a whole.     

Archaeocyaths: alternatively explained as consortia of siphonous algae and cyanobacteria-like microbes in shallow Cambrian seas

In order to test the systematic position of archaeocyaths, a group of Cambrian, exclusively marine fossils of unclear biological affinity, currently ascribed mostly to sponges (Porifera), we present an alternative explanation supporting the early idea of siphonous green algal affinity of archaeocyaths. The new arguments are based on a study of the morphology, microstructure, and microtaphonomy of early Cambrian (Tommotian and Atdabanian) archaeocyathan specimens from northeastern Siberia. Light microscopic and scanning electron microscopic (SEM) observations revealed that in many specimens the outer, often microlaminated, skeletal fabric grades continuously into the microfabric similar to that of microbial bush-like and dendritic microfossils such as Angulocellularia and Renalcis. The polyhedral to subglobular units characterizing the skeletal microfabric of the studied archaeocyaths and their microbial outgrowths are most similar to recent and fossil mineralized (calcified) colonies of benthic coccoidal cyanobacteria. This means that what had heretofore been interpreted as skeletons secreted by archaeocyathan organisms represent calcareous crusts precipitated by epibiontic cyanobacteria-like microbes, probably associated with a variety of other bacteria, colonizing surfaces and internal spaces of soft-bodied archaeocyathan organisms. Our study shows that a number of the morphological traits of archaeocyaths conserved by the mineralized microbial cover are similar to those of of modern siphonous green algae, like the marine genus Codium Stackhouse (Bryopsidales). Identifiable in well-preserved archaeocyaths are remains of such traits characteristic for Codium species as utricles, medullary threads, juvenile stages (primordia), and coalescence structures. The taxonomic and palaeoenvironmental implications of the recognition of archaeocyaths as possible consortia of siphonous green seaweeds (Chlorophyta) and cyanobacteria-like microbes are briefly discussed.     

Cambrian Series 3 lithistid sponge–microbial reefs in Shandong Province,North China: reef development after the disappearance of archaeocyaths

The Cambrian Series 3 Zhangxia Formation in Shandong Province, North China, includes small‐scale lithistid sponge–microbial reefs. The lithistid sponges grew on oolitic and bioclastic sediments, which were stabilized by microbial activities. The relative abundances of microbial components (e.g. calcimicrobe Epiphyton and stromatolites) vary among the reefs. However, the microbial components commonly encrusted or bound the lithistid sponges, formed remarkable encrustations on the surfaces of the sponges. Epiphyton especially grew upward and downward. The lithistid sponges thus provided substrates for the attachment and development of microbes, and the microbes played essential roles as consolidators, by encrusting reef‐building sponges. Additionally, the lithistid sponges were prone to degradation via microbial activities and diagenetic processes, and were thus preserved as micritic bodies, showing faint spicular networks or abundant spicules. Such low preservation potential within the reef environment obscured the presence of the sponges and their widespread contribution as reef‐building organisms during the Cambrian. During the prolonged interval after the demise of archaeocyaths, purely microbial reefs, such as stromatolites and thrombolites have been considered to be the principal reef builders, in association with rare lithistid sponge–microbial associations. However, recent findings, including those from Shandong Province and Korea, suggest that the lithistid sponge‐bearing reefs were more extensive during the Epoch 3 to the Furongian than previously thought. These lithistid sponge–microbial reefs were precursors of the sponge–microbial reefs that dominated worldwide in the Early Ordovician.     

Stromatoporoid-grade and other sponge fossils from the upper Krol Formation of the Lesser Himalaya (India) Implications for the biotic evolution around the Precambrian-Cambrian boundary interval

Summary A study of fossils in thin sections of a sample from the uppermost Krol E Member in the Mussoorie Hills of the Lesser Himalaya, India, proves the existence of morphologically differentiated calcified sponges within the Precambrian-Cambrian boundary time interval. The sponges, described as Mussooriella kroli n.g., n.sp. and Maldeotaina composita n.g., n.sp. indicate the presence of different organization grades at the Precambrian-Cambrian interval. Mussooriella had a calcareous skeleton consisting of skeletal elements composed of an inner laminated part and a distinct peripheral layer with knobs. Maldeotaina is characterized by a stromatoporoid-grade growth pattern following a thalamid-grade pattern. The stromatoporoidgrade part of the skeletons reminds strongly on skeletal elements common in labechiid Ordovician and younger stromatoporoids. Maldeotaina also shows criteria of Early Cambrian fossils, originally described as stromatoporoids and later excluded from this group and transferred to archaeocyaths. These similarities point to an Early Cambrian age of the fossil-bearing horizon in the topmost Krol E Member. Growth cavities with crypts indicate that the sponges might have contributed to the formation of small metazoan reefs-like structures. Although the study is based on limited material and many interpretations are still tentative, a thorough documentation of the preliminary results seems reliable considering the high potential of the fossils of the upper Krol Formation important source in understanding of early metazoan differentiation.     

贵州寒武纪海绵动物化石组合特征

贵州寒武纪海绵化石丰富。分布广泛,主要产于早寒武世梅树村期的戈仲伍组或牛蹄塘组底部、筇竹寺期的牛蹄塘组和早、中寒武世凯里组。梅树村期的海绵属种单调,主要为六射海绵类麦粒海绵（Triticispongia sp．）及一些海绵骨针,代表寒武纪海绵首次爆发或辐射的前奏和先驱。牛蹄塘组海绵多达13个属,是寒武纪海绵辐射的重要窗口,以普通海绵略占优势,Leptomitus为优势属。凯里组的海绵共计7属,以普通海绵为主。贵州寒武纪海绵动物群的发展具有明显的阶段性和组成的差异性,其演变过程受到环境的制约。贵州早期海绵动物的发现和研究不仅改变了寒武纪生物多样性的认识,而且还为探讨海绵动物本身的起源、辐射提供了重要材料。     

Anatomy and lifestyles of Early Cambrian priapulid worms exemplified by Corynetis and Anningvermis from the Maotianshan Shale (SW China)

Accurate information on the anatomy and ecology of worms from the Cambrian Lagerstätten of SW China is sparse. The present study of two priapulid worms Anningvermis n. gen. and Corynetis Luo & Hu, 1999 from the Lower Cambrian Maotianshan Shale biota brings new information concerning the anatomical complexity, functional morphology and lifestyles of the Early Cambrian priapulids. Comparisons are made with Recent priapulids from Sweden (live observations, SEM). The cuspidate pharyngeal teeth of Anningvermis (circumoral pentagons) and the most peculiar radiating oral crown of Corynetis added to the very elongate pharynx of these two forms are interpreted as two different types of grasping apparatus possibly involved in the capture of small prey. Corynetis and Anningvermis are two representative examples of the Early Cambrian endobenthic communities largely dominated by priapulid worms (more than ten species in the Maotianshan Shale biota) and to a much lesser extent by brachiopods. Corynetis and Anningvermis were probably active mud-burrowers and predators of small meiobenthic animals. Likewise predator priapulid worms exploited the interface layer between the seawater and bottom sediment, where meiobenthic organisms were abundant and functioned as prey. This implies that complex prey-predator relationship between communities already existed in the Early Cambrian. This study also shows that the circumoral pentagonal teeth and caudal appendage were present in the early stages of the evolutionary history of the group and were important features of the priapulid body plan already in the Early Cambrian. Two new families, one new genus and new species are introduced and described in the appendix.     

A new look at the Precambrian and Cambrian event chronostratigraphic correlation of Mongolia

The Mongolian Precambrian and Cambrian event chronostratigraphic classification of sedimentary, sedimentary—volcanogenic, volcanogenic, and metamorphic rock sediments spread over the territory of Mongolia according to the new chronostratigraphic classification approved by the International Stratigraphic Commission is discussed. These ancient rock units are readily divided into two independent complexes. The lower complex (1000–3500 m) is represented by the Archean and Early Proterozoic crystalline basement and the upper complex is the latest Precambrian–Cambrian sediments (220–7500 m). A detailed study of abundant Neoproterozoic and Early Cambrian faunal (ichnofossils, hexactinellid sponges, archaeocyaths, trilobites, etc.) and floral (stromatolites, microphytolites, cyanobacterial mats, microfossils, etc.) fossils provides the first regional chronostratigraphic subdivision of different facies sediments. Every stage is characterized by distinctive geohistorical and biological events.     

贵州早、中寒武世斗篷海绵科(Choiidae)的发现

本文描述了贵州遵义下寒武统牛蹄塘组生物群和台江中寒武世凯里生物群中保存完整的斗篷海绵科(Choiidae)中斗篷海绵(Choia)和小斗篷海绵(Choiaella)化石标本,共计2属2种2未定种：Choia zunyiensis sp.nov.,Choia sp.,Choiaella avata sp.nov.和Choiaella sp.。Choia分布很广．在我国下寒武统澄江生物群及北美中寒武统的四大生物群中均有产出。Choiaella原仅产于我国澄江生物群,凯里生物群中Choiaella的发现使其时代上延至中寒武世。贵州早、中寒武世斗篷海绵科的发现及研究为海绵动物的早期演化及古地理学等方面的研究提供了重要的新材料。     

Facies of Liassic sponge mounds, central High Atlas, Morocco

Summary Liassic sponge mounds of the central High Atlas (Rich area, northern Morocco) have a stratigraphic range from the Lower/Upper Sinemurian boundary interval up to the lower parts of the Lower Pliensbachian (Carixian). The base of Liassic sponge mounds consists of a transgressive discontinuity, i.e., a condensed section of microbioclastic wackestones with firm- and hardgrounds, ferruginous stromatolites, sponge spicules and ammonites. The top of Liassic sponge mounds is an irregular palaeorelief covered by cherty marl-limestone rhythmites, namely hemipelagic spicular wackestones with radiolaria. In the Rich area, section Foum Tillicht, the sponge mound succession has a total thickness of about 250 meters. Within this succession we distinguished between three mound intervals. The lower mound interval shows only small, meter-scale sponge mounds consisting of boundstones with lyssakine sponges, commensalicTerebella and the problematicumRadiomura. This interval forms a shallowing-upward sequence culminating in a bedded facies withTubiphytes, calcareous algae (Palaeodasycladus), sponge lithoclasts, coated grains, and thin rims of marine cement. The middle mound interval is aggradational with decametric mounds and distinct thrombolitic textures and reefal cavities. The mound assemblage here consists of hexactinellid sponges, lithistid demosponges, non-rigid demosponges,Radiomura, Serpula (Dorsoserpula), Terebella, encrusting bryozoa, and minor contributions by calcareous sponges, and excavating sponges (typeAka). Thrombolites are dendrolitic and may reach sizes of several tens of centimeters, similar to the maximum size of siliceous sponges. The upper mound interval appears retrogradational and geometries change upsection from mound shapes to flat lenses and level-bottom, biostromal sponge banks. The biotic assemblage is similar to that of the middle mound interval and there is no difference between mound and bank communities. The demise of sponge mounds is successive from regional spread in the Sinemurian to more localised spots in the Lower Pliensbachian. This reduction correlates with an increasing influence of pelagic conditions. At Foum Tillicht, sponge mounds lack any photic contribution and there is virtually no differentiation into subcommunities between mound surface and cavity dwelling organisms. There is some evidence that the heterotrophic food web of mound communities was sourced by oxygen minimum zone edge effects, namely microbial recycling of essential elements such as N and P. Basin geometry suggests a waterdepth of several 100's of meters, well below the photic zone and possibly only controlled by the depth range of the oxygen minimum zone. Palaeoceanographic conditions of well-stratified deeper water masses diminished gradually during widespread transgression across the Sinemurian to Pliensbachian boundary culminating in the Lower Pliensbachianibex ammonite zone.     

Cambrian anemones with preserved soft tissue from the Chengjiang biota, China

The group Cnidaria includes 'jellyfish', soft-bodied anemone and anemone-like forms and calcified corals. These diploblastic organisms have a fossil record extending back to the earliest metazoans of the Neoproterozoic; however certain cnidarians of the subclass Zoantharia, characterized by soft-bodied anemone-like forms, are absent or poorly represented in the fossil record. Despite the paucity of fossils, it is thought that calcification by soft anemone-like animals was responsible for producing the skeleton that allowed the preservation of the first corals. We report discovery of an abundant assemblage of in situ soft-bodied polyps with tissues. They are preserved in exquisite detail and come from the well-known Lower Cambrian Chengjiang biota of Yunnan, China. The soft-bodied polyps display a simple anatomy that is comparable to some extant anemones of the order Actinaria. The new fossils are assigned to Archisaccophyllia kunmingensis n. gen. et n. sp. Their simple and conservative form suggests that these fossils may represent some kind of ancestral rootstock. The preserved life assemblage provides a unique snapshot of Lower Cambrian anemone life and provides clues for relationships with extant actiniarians as well as calcified corals.     

Quantitative paleoecology of Cambrian (Series 2–Miaolingian) communities from central Sonora,Mexico

This work presents a paleoecological approach to study benthic communities preserved in Cambrian strata exposed at the Sahuaral locality in central Sonora, northwestern Mexico. Based on species richness and abundance, this work analyzed 627 individuals corresponding to 33 species from five distinctive Cambrian formations: the lowermost Proveedora Formation characterized by bioturbation horizons that record the occurrence of a benthic-infaunal biota from coastal shallow seas, followed by the Buelna Formation which contains reef deposits dominated by autotrophic epibenthic communities. The Cerro Prieto Formation was deposited in a high-energy shallow marine environment, dominated by primary consumers-epifaunal and suspensions feeders. The uppermost unit, the El Gavilán Formation, is an intercalation of shale and fossiliferous limestone characterized by the fossil remains of suspension feeders, detritus feeders and nektobenthic organisms of low energy deep water environments. The paleobiota consist of ichnofossils, microbial oncolites, chancellorids, brachiopods, hyolithids, and trilobites in a sequence of increasing diversity and complexity. Paleoecological indices (Simpson Dominance, Shannon-Weaver Index, Evenness Index, and Total Richness) were determined and compared in the study area. In addition, the effective number of species and the true diversities were obtained, which are similar between each station distributed in each formation. However, it is possible that the high degree of dominance is a factor to indicate an inequality between the true diversity and the species richness identified. These results provide new analytical and paleoecological criteria for better understanding depositional environments and fossil associations where the spatial distribution of benthic organisms, richness and abundance provide a valued window to Cambrian biotas. The structure of the marine community of the El Sahuaral area has a great affinity with other Cambrian communities, such as those from the Stephen Formation (Burgess Shale, British Columbia) and the Chengjiang biota of the Yangtze Shelf deposits in South China.     

Upper Triassic (Carnian) reef biota from the Sambosan Accretionary Complex, Kyushu, Japan

Calcified sponges, algae, and reef problematica are abundant yet poorly known from the Triassic of Japan. They are abundant in shallow-water carbonate, redeposited blocks of the Sambosan Accretionary Complex, Konosé Group, and southern Kyushu. Based on study of thin-sections from reef limestone exposed along the Kuma River, some important organisms and reef microfacies are described, which seem typical of Upper Triassic reef complexes. The most abundant reef organisms are hypercalcified sponges, including sphinctozoans, inozoans and chaetetids, followed by cyanophycean algae (including “Tubiphytes”-like organisms), and solenoporacean red algae. Loose sponge spicules in one thin-section also indicate the occurrence of rare hexactinellid sponges. Chambered demosponges described from the Konosé carbonate rocks include Solenolmia manon manon (Münster), Colospongia sp., Jablonskyia andrusovi (Jablonsky), several unidentified chambered sponges as well as the inozoid Permocorynella sp. 1 and Permocorynella sp. 2. Also present are chaetetid sponges and solenoporacean red algae belonging to Parachaetetes cassianus (Flügel) and Parachaetetes? sp. or Solenopora? sp. Especially abundant in thin-sections are cyanophyceans and “Tubiphytes”-like organisms. Among the organisms is Cladogirvanella Ott and Hedstroemia sp. The composition of the biota and presence of typical problematic organisms increases our knowledge of shallow-water Upper Triassic carbonate rocks in a remote setting in western Panthalassa. The composition of the biota indicates a mostly Carnian age. Most comparable organisms are known from both the northeastern and southern Tethys.     

AN ORDOVICIAN LOBOPODIAN FROM THE SOOM SHALE LAGERSTÄTTE, SOUTH AFRICA

Abstract:  The first lobopodian known from the Ordovician is described from the Soom Shale Lagerstätte, South Africa. The organism shows features homologous to Palaeozoic marine lobopodians described from the Middle Cambrian Burgess Shale, the Lower Cambrian Chengjiang biota, the Lower Cambrian Sirius Passet Lagerstätte and the Lower Cambrian of the Baltic. The discovery provides a link between marine Cambrian lobopodians and younger forms from the Silurian and Carboniferous. The new fossil preserves an annulated trunk, lobopods with clear annulations, and curved claws. It represents a rare record of a benthic organism from the Soom Shale, and demonstrates intermittent water oxygenation during the deposition of the unit.     

Branching archaeocyaths as ecosystem engineers during the Cambrian radiation

The rapid origination and diversification of major animal body plans during the early Cambrian coincide with the rise of Earth's first animal-built framework reefs. Given the importance of scleractinian coral reefs as ecological facilitators in modern oceans, we investigate the impact of archaeocyathan (Class Archaeocyatha) reefs as engineered ecosystems during the Cambrian radiation. In this study, we present the first high-resolution, three-dimensional (3D) reconstructions of branching archaeocyathide (Order Archaeocyathida) individuals from three localities on the Laurentian paleocontinent. Because branched forms in sponges and corals display phenotypic plasticity that preserve the characteristics of the surrounding growth environment, we compare morphological measurements from our fossil specimens to those of modern corals to infer the surface conditions of Earth's first reefs. These data demonstrate that archaeocyaths could withstand and influence the flow of water, accommodate photosymbionts, and build topographically complex and stable structures much like corals today. We also recognize a stepwise increase in the roughness of reef environments in the lower Cambrian, which would have laid a foundation for more abundant and diverse coevolving fauna.     

Paleoecology of Middle Ordovician stromatoporoid mounds in Vermont

Fossiliferous mounds of carbonate mud are a distinctive facies in the middle Chazy Group (Crown Point Formation) at Isle La Motte, Lake Champlain. The mounds are surrounded by bedded calcarenite of spar-cemented pelmatozoan debris. Channels which cut into the mounds during mound growth are filled with the same calcarenite. The mud-free intermound rocks and the mound biota suggest agitated, normal marine shallow-water environments. The principal lime-secreting organisms within the mounds are stromatoporoids, calcareous algae, tabulate corals, sponges, and bryozoans. Each mound is dominated in terms of biomass by one of three groups: stromatoporoids, calcareous algae, and bryozoans. Most of the mound biota first appear at the base of the Crown Point Formation. In the lower Crown Point Formation the organisms increase in number and species. Both changes in the biota are related to periods of shallowing of the Chazy sea which are also reflected in the character of the carbonate sands.     

Non-rigid cryptic sponges in oyster patch reefs (Lower Kimmeridgian, Langenberg/Oker, Germany)

Summary Two patch reefs which predominately consist of the oysterNanogyra nana (Sowerby 1822) are exposed in Lower Kimmeridigian strata of the Langenberg hillrange, central Germany. Left oyster valves making up the frame-work of the reefs formed small abundant cavities that were inhabited by a unique sponge community. The excellent preservation of non-rigid sponges was related to early organomineralization within the decaying sponge tissue. As a process of sponge taphonomy, different types of microbially induced carbonates precipitated preserving spicule aggregates. Organomineralization within sponge soft tissues is especially favored with the Langenberg patch reefs due to the closed or semi-closed system conditions with the cavities. The δ¹³C values ofin situ formed microbialities reveal that carbonate precipitation was in equilibrium with Jurassic seawater. The carbon of the microbialites does not derive from the bacterial remineralization of organic matter, but is of a marine source. Likewise, organomineralization is probably related to bacterial EPS or decaying sponge tissues providing an organic matrix for initial carbonate precipitation. Biomarker analyses revealed, that the patch reef microbialites contain terminally branched fatty acids (iso-andanteiso-pentadecanoic acid) in significant concentrations. These fatty acids, like hopanoid hydrocarbons, are most likely of a bacterial source. This is in agreement with sulfate-reducing bacteria remineralizing the decaying sponges as further indicated by the occurrence of framboidal pyrite in sponge microbialites.     

寒武纪早期澄江生物群中受创伤的中间型始莱德利基三叶虫(Eoredlichia intermedia)

三叶虫是寒武纪海洋中的常见动物,其受伤愈合现象的化石记录可以反映生态系统内物种之间竞争关系。中国华南寒武纪早期澄江化石生物群因保存软躯体构造化石而成为研究寒武纪大爆发时期古海洋生态群落的绝佳窗口,然而,澄江生物群中三叶虫受伤愈合现象的化石至今未见报道。中间型始莱德利基虫(Eoredlichia intermedia Lu,1940)是澄江生物群的常见物种,也是我国寒武系第二个三叶虫化石带的标准带化石。在2 000余枚中间型始莱德利基虫标本中,发现两枚成体标本的头甲侧边缘、颊刺和胸甲肋节保存了不同程度的残缺。据此推测,Eoredlichia intermedia在生命活动过程中曾受到碎壳型捕食动物的攻击或其他伤害而形成伤口。伤口部位边缘加厚且光滑,显示了明显的愈合痕迹。这些特征表明始莱德利基虫在受到非致死的伤害后,具备自身修复损伤的能力。这是迄今所知最为古老的,也是在澄江生物群中首次发现的三叶虫受伤并且愈合的标本,反映了寒武纪第二世第三期底栖生态系统中碎壳型捕食者和三叶虫猎物之间的军备竞赛生态关系。     

Calcification processes of siliceous sponges in Viséan Limestones (Counties Sligo and Leitrim,Northwestern Ireland)

Summary In the Lower Carboniferous limestones and shales of the Benbulben Range, Counties Sligo and Leitrim, northwestern Ireland, a suite of carbonate nodules, about 1 to 4 cm in diameter, has been sampled and investigated by thin sectioning. The nodules consist of micritic, peloidal and fenestral fabrics. Many of them contain relics of desma bearing demosponges and hexactinellid sponge skeletons. The nodules are interpreted as calcified siliceous sponges. Micrite and peloids have been formed via microbial activity during the decay of the soft sponge tissue. The actual processes are deduced from Recent examples investigated at Lizard Island, Autralia, byReitner (1993). The skeletal opal was dissolved very early. In places where the skeleton was already embedded in micrite the spicules are preserved as molds cemented by granular ferroan calcite. The nodules were extensively inhabited by agglutinating polychaetes and bored by sponges. Micrite clasts have been exported to the surrounding seafloor before the sponges were completely covered by sediments. Fenestral fabrics represent primary sponge cavities, that may be enlarged due to volume reduction of the soft tissue during calcification. Some originated from non-calcification of decaying tissue. The granular calcite cement, filling the fenestral fabrics, contains relics of spicules and faintly visible peloids floating unsupportedly in the cement. These peloids were probably produced in situ by calcification of organic mucilages that filled the cavities almost entirely. It is evident that most diagenetic processes occurring within the sponges happened on the seafloor, most likely within the still living individuals. Possibly the nodules represent a precursor stage of mud mound development.     

贵州寒武纪杷榔动物群的古生态学初探*

贵州剑河辣子寨村辣子寨剖面杷榔组的杷榔动物群是近期发现的一个化石丰度较大的产地。目前已发现了海绵动物、刺细胞动物、曳鳃动物、腕足动物、软舌螺动物、节肢动物、棘皮动物、藻类等8大类群的化石代表,计有63属80种。本文依据辣子寨剖面杷榔化石库下化石层的生物组合及各类群的生态特性、埋藏特征,探讨杷榔动物群的生态系及生活环境。杷榔动物群的生物群落,包括了底栖固着、内栖钻孔、表栖游移、漂游和游泳的生物类群,且生物群生活在赤道地区以北开阔海盆的透光区至弱透光区、水域含氧、海底有氧—弱氧、盐分正常、快速沉积事件频发的陆棚较深水环境。其古生态特点为:生物种类丰富,具有滤食、食草、食泥、食肉和食腐的动物;而捕食-被捕食的食物网的关系较为复杂,具有明显的金字塔型的营养结构。