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A Middle Cambrian possible cnidarian from the Murrawong Creek Formation, NE New South Wales
Abstract
A coralomorph, Tretocylichne perplexa gen. et sp. nov., tentatively interpreted as a cnidarian with mineralized skeleton and unusually preserved in the form of epidote coatings, is described from richly fossilliferous limestone clasts collected from the Murrawong Creek Fmn near Woolomin, NE New South Wales. The trilobite fauna including Pagetia pollosta Jell, 1975, P. whitehousei Jell, 1975, Hypagnostus parvifrons (Linnarsson, 1869) and Utagnostus cf. trispinulus Robison, 1964, permits a middle Middle Cambrian dating. -Author
... Recently, cloudinids were also discovered at the basal Terreneuvian (ca 539-521 Ma) [15,16], indicating that they were also present during the ongoing biotic replacement at the Ediacaran-Cambrian boundary. Despite the fact that Cloudina is one of the most studied Ediacaran organisms, its phylogenetic affinity has remained debated; suggestions including being a diploblastic metazoan such as a stem-or crown-group cnidarian [6,17] or an animal with annelid/bilaterian affinity [10,18]. ...
... In the type species, Cl. hartmannae, there are transverse annulations on the outer surface of the funnel-shaped elements (figure 1a,c). Each funnel-like element is nested deeply within the preceding element, and they frequently flare aperturally to form flanges that may fuse to neighbouring flanges [17,18]. Dichotomous branching commonly occurs deep within royalsocietypublishing.org/journal/rsos R. Soc. ...
... 8: 210829 4 the parental funnel forming two daughter funnels of equal size, but recently, budding from the outside of the parental funnel has also been reported [25]. In Cloudina riemkeae the funnel-like elements are small and nested only shallowly [17]. The type specimen of Cloudina carinata were originally thought to occur in the terminal Ediacaran [26], but is now known to be basal Terreneuvian in age [27]. ...
The Ediacaran–Cambrian transition and the following Cambrian
Explosion are among the most fundamental events in the
evolutionary history of animals. Understanding these events is
enhanced when phylogenetic linkages can be established among
animal fossils across this interval and their trait evolution
monitored. Doing this is challenging because the fossil record of
animal lineages that span this transition is sparse, preserved
morphologies generally simple and lifestyles in the Ediacaran
and Cambrian commonly quite different. Here, we identify
derived characters linking some members of an enigmatic
animal group, the cloudinids, which first appeared in the
Late Ediacaran, to animals with cnidarian affinity from the
Cambrian Series 2 and the Miaolingian. Accordingly, we present
the first case of an animal lineage represented in the Ediacaran
that endured and diversified successfully throughout the
Cambrian Explosion by embellishing its overall robustness and
structural complexity. Among other features, dichotomous
branching, present in some early cloudinids, compares closely
with a cnidarian asexual reproduction mode. Tracking this
morphological change from Late Ediacaran to the Miaolingian
provides a unique glimpse into how a primeval animal group
responded during the Cambrian Explosion.
... Peel (2011) described Cothonion sympomatum from the Paralleldal Formation (Cambrian Series 2, Stage 4) of southern Peary Land, North Greenland. Coralla of Tretocylichne perplexa Engelbretsen, 1993 from the Murrawong Creek Formation (Cambrian Series 3, Drumian) of New South Wales often display an octagonal form and weak septation, while the holdfast is often perforated into the calice (Engelbretsen, 1993). ...
... Journal of Paleontology 91 (5) in consequence of which, Cambroctoconida is tentatively placed within the anthozoan Class Octocorallia. While the presence of pores is not established in Tretocylichne on account of its relatively coarse preservation (Engelbretsen, 1993), the octagonal form and eight-fold septation support its assignment to Cambroctoconida. Material of Lipopora is coarsely silicified, but the cylindrical coralla display eight or sixteen septa, which seem to confirm their affinity with Cambroctoconus. ...
The problematic calcified cnidarian Cambroctoconus is described from the Henson Gletscher Formation (Cambrian Series 2, Stage 4–Series 3, Stage 5) of North Greenland, representing the first record from Laurentia of a genus otherwise recently described from China, Kyrgyzstan, and Korea. Internal molds produced by penetrative phosphatization mirror the pervasive pore system of the calice walls and septa. The pore system is compared to the network of gastrodermal solenia that distributes nutrients between polyps and surrounding stolon tissues in present day octocorals. In conjunction with the octagonal form of the individual coralla and eight-fold symmetry of septa, the pore system promotes assignment of Cambroctoconus to the Octocorallia, a basal clade in cnidarian phylogeny. Octocorals (‘soft corals’) are diverse in present day seas, but have a poor fossil record despite the general development of distinctive calcareous spicules. New taxa: Order Cambroctoconida new; Cambroctoconus koori new species.
... The uncertainty of "coralomorph" higher-level taxonomy, as noted in a number of reports (Savarese et al., 1993;Wood, 1999;Rowland and Shapiro, 2002), includes the sheer strangeness of many of these early Cambrian fossils (particularly forms like Lipopora, Harklessia, and the Cothoniida) (see also Engelbretsen, 1993). This makes it difficult to relate them to the subsequent great Palaeozoic coral groups, the Tabulata and the Rugosa, that make their first appearance in the Ordovician. ...
Re-evaluation of eumetazoan modular coloniality gives a new perspective to Ediacaran–Ordovician animal diversification. Highly integrated eumetazoan colonies (porpitids [“chondrophorines”], pennatulacean octocorals, anthozoans) prove to be unknown in the Ediacaran. Ediacaran Evolutionary Radiation (EER, new term) fossils include macroscopic and multicellular remains that cannot be compellingly related to any modern group. Claims of eumetazoan coloniality in the Ediacaran are questionable. The subsequent Cambrian Evolutionary Radiation (CER, terminal Ediacaran–late early Cambrian) records appearance and diversification of deep burrowers and a relatively abrupt development of biomineralization. The CER began in a transition zone that spans the Ediacaran–Cambrian boundary and includes the final few million years of the Ediacaran. The early CER has pseudocolonial(?) Corumbella that may be related to some Phanerozoic taxa (conulariids) and records appearance of the first macroscopic biomineralised organisms (Cloudina, Namacalathus, Namapoikea), which may not be eumetazoans. Modular eumetazoans dominate and define many Ordovician and younger habitats (coral, bryozoan, sabellitid reefs; pelagic larvaceans, salps, early–middle Palaeozoic graptolites), but eumetazoan coloniality largely “missed” the EER and CER. All purported Ediacaran–Ordovician porpitids (“chondophorines”) and pennatulaceans are not colonial eumetazoans. Only in the late early Cambrian (late CER) or early middle Cambrian do a few modular colonial eumetazoans first occur as fossils. These include Sphenothallus (available evidence precludes Torellella coloniality), some corals (colonial “coralomorphs”), and lower middle Cambrian graptolithoids. Modular eumetazoan colonies (corals, graptolithoids) in the late early and early middle Cambrian (late Epoch 2–early Epoch 3) and appearance of mid-water predators (cephalopods, euconodonts) and bryozoans in the late Cambrian–earliest Ordovician (late Furongian–early Tremadocian) are the root for the Great Ordovician Biodiversification Interval (GOBI, new term) and diverse later Phanerozoic communities.
... The lowermost of these, designated unit 1, is 65 m thick and is dominated by coarse-grained allochthonous limestone blocks yielding a diverse shelly fauna of late 'Middle' Cambrian age (Undillan–Boomerangian, P. punctuosus to L. laevigata trilobite zones). Fossils described from this level include a possible cnidarian (Engelbretsen 1993), lingulate brachiopods (Engelbretsen 1996), molluscs (Brock 1998a) possible rhynchonelliformean brachiopods (Brock 1998bBrock , 1999) and trilobites (Cawood 1976, Sloan & Laurie 2004 now known to be of 'Middle' Cambrian to early Late Cambrian age based on the presence of paraconodonts and absence of euconodonts (Stewart 1995). Hence the age of the Pipeclay Creek Formation effectively constrains the age of deposition of the underlying Murrawong Creek Formation to that of the limestone clasts found in conglomerates of unit 1. Lithologies in the Pipeclay Creek Formation consist predominantly of siltstone and mudstone with subsidiary chert, tuff, sandstone and fine-grained conglomerate, more than 900 m in total thickness (Bradley, in Pickett 1982; Cawood 1983; Leitch & Cawood 1987). ...
We present a comprehensive review of a significant interval spanning 100million years in the geological history of New South Wales, listing all currently accepted groups, formations and constituent members of Cambrian and Ordovician age. These units are briefly described and placed in their tectonic context, with the most up-to-date biostratigraphic and isotopic age dating assembled to constrain correlations (depicted in 25representative stratigraphic columns) across orogenic belts and terranes. Rock units previously assigned a Cambrian or Ordovician age, whose names are now obsolete, redundant or are known to be younger, are also discussed or listed in an appendix. The increasingly diverse literature on the Cambrian and Ordovician stratigraphy of the state is reflected in an extensive bibliography. This review is intended to benefit the mineral exploration industry, research workers both locally and overseas, and geological mapping generally by providing a ready reference to Cambrian and Ordovician rocks in NSW. It also indicates where current data are insufficient to resolve precise age determinations and correlations, thereby highlighting those areas that require further work before a complete synthesis of the early Palaeozoic geological history of NSW can be undertaken.
... The relationship of the Cambrian forms to the Ordovician and younger corals is problematic, and Jell (1984) proposed Coralomorpha as an informal embracive term for taxa of uncertain affinity, including several species not considered by Hicks (2006). Cambrian corals/coralomorphs are generally considered to represent calcification events within early cnidarian evolution that are unrelated to the later rugose and tabulate corals (Scrutton 1997, 1999, Engelbretsen 1993, Fuller & Jenkins 2007. 'True corals' are well represented in the early Cambrian (Botoman) and have been referred to Tabuliconida Scrutton, 1997, whereas Cothonion sympomatum Jell & Jell, 1976 from the 'first discovery limestone' of the Coonigan Formation of New South Wales, Australia (31810 0 30 00 S, 142819 0 E), was placed in the Cothoniida of uncertain position (Scrutton 1997(Scrutton , 1999; see also Oliver & Coates 1987). ...
Originally described from the Coonigan Formation of New South Wales, Australia, a second occurrence of the operculate coral Cothonion sympomatum is here described from the Paralledal Formation of North Greenland. Both finds are of late early Cambrian age, Series 2, Stage 4 in the emerging fourfold classification of the Cambrian. The new find supports widespread distribution patterns seen in early Cambrian Small Shelly Fossils, although associated trilobites belong to traditional redlichiid (Australia) and olenellid (Greenland) realms, respectively.
The Cambrian cnidarian Cambroctoconus orientalis occurs in clusters exhibiting pendent growth in crypts within thrombolite frameworks. Here we successfully reconstruct its three‐dimensional (3D) in situ mode of life. The 3D reconstruction shows the initial individuals of colonies and attachment structures, budding sites and growth modes of coralla in response to the extent of sheltered spaces. The earliest individuals were attached to the framework by a suction‐cup‐like structure and were basically orthogonal to the walls, growing downward. Their calicular bases were pierced by a hole. Individuals were initially cylindrical and gradually formed an octagonal column during growth. Budding occurred everywhere, and toroidal budding traces are abundant upon the corallite surfaces. The attachment structures were secondarily reinforced by continuous secretion from the coenenchymal tissue. Offsets continued to grow only when sufficient growth space was available, while producing as many individuals as possible. Growth of individuals was commonly halted by the surrounding framework. Growth directions were in part modified by corallite curvature to avoid collisions between corallites and to make full use of the intricate void spaces. In deteriorating conditions, offset corallites appeared through regeneration rather than budding from the remaining soft tissues. Active exploitation of the sheltered spaces within thrombolites, which remained dominant during the Cambrian, was thus conducted by C. orientalis . Notably, the cryptobionts treated herein adopted subtle growth strategies that have not previously been fully understood and are not easily interpreted with reference to organisms living in open spaces.
Five species, Lipopora lissa Jell and Jell, 1976, Lipopora daseia Jell and Jell, 1976, Tretocylichne perplexa Engelbretsen, 1993 from Australia, Cambroctoconus orientalis Park, Woo, Lee, Lee, Lee, Han and Chough, 2011 from China, and Cambroctoconus kyrgyzstanicus Peel, 2014 from Kyrgyzstan, belonging to the Cambrian stem-group cnidarians have been documented in the fossil record. Cambroctoconus coreaensis sp. nov., interpreted here as a stem-group cnidarian, from the Seokgaejae section in the Daegi Formation, Taebaek Group (Cambrian Series 3), Taebaeksan Basin, central-eastern Korean Peninsula, has a slender cup-shaped skeleton. A cladistic analysis produced 21 most parsimonious trees, which invariably placed the six stem-group cnidarians below the crown-group, but their relationships within the stem-group are unresolved. Nine out of the 21 trees suggest a monophyletic relationship for the Cambrian stem-group cnidarians, whereas in six other trees a monophyly of Cambroctoconus and Tretocylichne appeared as the sister-group to the crown-group cnidarians with Lipopora at the most basal branch. This result may reflect the fact that crown-group cnidarians evolved in the Precambrian, and suggests that the diversity of stem-group cnidarians was a result of an independent radiation in the Cambrian.
Tomteluva perturbata gen. et sp. nov. and Nasakia thulensis gen. et sp. nov., two new rhynchonelliformean brachiopod taxa, are described from carbonate beds from the lower middle Cambrian (Series 3, Stage 5) basinal Stephen Formation, Canada, and the upper lower Cambrian (Series 2, Stage 4) Henson Gletscher Formation, North Greenland, respectively. The two taxa are characterized by an unusual coral-like morphology typified by a high conical ventral valve with an anteriorly curved umbo and a tube-like structure inside the ventral valve, interpreted as pedicle tube. Both resemble the problematic late middle Cambrian (Drumian) species Anomalocalyx cawoodi Brock from Australia, whose systematic affiliation is controversial. Together, the three genera are interpreted as representatives of a new family of rhynchonelliformean brachiopods, the Tomteluvidae fam. nov., which is interpreted as an aberrant or derived taxon within the Order Naukatida. Convergence between the Tomteluvidae and the coralla of small solitary Cambrian coralimorphs, as well as the late Palaeozoic reef-building richthofenioid brachiopods, might indicate adaptation to a similar life habits and environments. However, their small size (length 4 mm), well-developed pedicle and perfect morphological symmetry make it more likely that tomteluvids lived attached to frondose algae or sponges, above the seafloor, in a similar fashion to the acrotretoid brachiopods with which they show a high degree of morphological convergence. Morphological features of the pedicle tube of N. thulensis suggest that the tomteluvid pedicle is homologous to that in modern rhynchonelliformean brachiopods. This is the first evidence of the pedicle type within the Naukatida and represents the oldest confirmation of a rhynchonellate pedicle.
Early Middle Cambrian bituminous coquinoid limestones from a tectonically isolated outcrop in southwestern Kyrgyzstan yield a remarkably diverse fauna, with stem-group cnidarians, trilobites, rhynchonelliformean brachiopods, and other shelly fossils. The fossil site is in the northern foothills of the Turkestan Range and thus forms part of the westernmost extension of the South Tien Shan. The fauna includes two fairly well known trilobite species, Glabrella ventrosa Lermontova, 1940 and Dorypyge richthofeniformis Lermontova, 1940, that provide confident support for an Amgan age of the rocks. New described taxa include the stem-group cnidarian Cambroctoconus kyrgyzstanicus Peel sp. nov., the trilobite Olenoides sagittatus Geyer sp. nov., and the helcionelloid Manasoconus bifrons Peel gen. et sp. nov. Additional fossils within the samples include the trilobites Olenoides sp. A, Kootenia sp., and Pseudoeteraspis? sp.; the rhynchonelliform brachiopods Narynella cf. ferganensis (Andreeva, 1962), Narynella? sp., Austrohedra? sp. nov., and two species of uncertain generic affinity; the tommotiid Tesella sp.; the hyolithelminth Hyolithellus sp.; and the palaeoscolecid Hadimopanella oezgueli Gedik, 1977. Of particular interest is Cambroctoconus kyrgyzstanicus with an octagonal corallum and a sparsely septate calyx.
Early Middle Cambrian bituminous coquinoid limestones from a tectonically isolated outcrop in southwestern Kyrgyzstan yield a remarkably diverse fauna, with stem-group cnidarians, trilobites, rhynchonelliformean brachiopods, and other shelly fossils. The fossil site is in the northern foothills of the Turkestan Range and thus forms part of the wes- ternmost extension of the South Tien Shan. The fauna includes two fairly well known trilobite species, Glabrella ventrosa Lermontova, 1940 and Dorypyge richthofeniformis Lermontova, 1940, that provide confident support for an Amgan age of the rocks. New described taxa include the stem-group cnidarian Cambroctoconus kyrgyzstanicus Peel sp. nov., the trilobite Olenoides sagittatus Geyer sp. nov., and the helcionelloid Manasoconus bifrons Peel gen. et sp. nov. Additional fossils within the samples include the trilobites Olenoides sp. A, Kootenia sp., and Pseudoeteraspis? sp.; the rhynchonelliform brachiopods Narynella cf. ferganensis (Andreeva, 1962), Narynella? sp., Austrohedra? sp. nov., and two species of uncertain generic affinity; the tommotiid Tesella sp.; the hyolithelminth Hyolithellus sp.; and the palaeoscolecid Hadimopanella oezgueli Gedik, 1977. Of particular interest is Cambroctoconus kyrgyzstanicus with an octagonal corallum and a sparsely septate calyx.
Numerous Early Cambrian corals or "coralomorphs," as they are often classified, are recorded from North America, Australia. and Siberia. A new Early Cambrian coral, Harklessia yuenglingensis n. gen. and sp., is found in conjunction with archaeo-cyathan-microbial reefs in Esmeralda County, southwestern Nevada. The coral-bearing reefs are within quartzic, trilobite-rich packstone beds in the upper portion of the Harkless Formation (Bonnia-Olenellus Zone). Coralla are constructed by subpolygonal to polygonal, cerioid, close-packed corallite tubes. Coralla average 12 cm in height by 18 cm in diameter with individual corallite tubes ranging from 1.2 to 3.2 mm in diameter. Corallites are greater than 25 mm in length. Septa and tabulae are not present. Many of the Early Cambrian corals previously described have attributes of the class Anthozoa and subclass Zoantharia, with some specific similarities to tabulate corals. Harklessia yuenglingensis is placed confidently within the class Anthozoa, subclass Zoantharia because its morphological characteristics indicate an affinity to true corals, but whether H. yuenglingensis is a tabulate coral remains uncertain.
Middle Cambrian (Floran-Undillan) allochthonous limestone clasts from the Murrawong Creek Formation, southern New England Fold Belt, northeastern N.S.W., Australia, yield a well preserved silicified molluscan fauna consisting of the helcionelloids Latouchella accordionataRunnegar & Jell,, Latouchella aliciae nov. sp., Yochelcionella dalekiRunnegar & Jell,, Anabarella simesiMacKinnon,, and the paragastropod Pelagiella sp. cf. P. deltoidesRunnegar & Jell.. On the basis of a single bellerophontiform-like muscle scar fortuitously preserved in one specimen, Protowenella is tentatively re-interpreted as an archaeogastropod, though a tergomyan affinity cannot be ruled out. The fauna displays strong regional affinities with the “first discovery limestone” member of the Coonigan Formation, western N.S.W., sequences in the Georgina Basin, northern Australia, and northwest Nelson in New Zealand.
Calcareous articulate brachiopods are rare components of the high diversity, phosphatic, silicified, and epidote coated shelly fauna derived from Middle Cambrian (Floran-Undillan) allochthonous limestone clasts from the Murrawong Creek Formation, southern New England Fold Belt, northeastern New South Wales, Australia. Three taxa are described, the kutorginids
Nisusia metula
n. sp., and
Yorkia
sp. indet., and the protorthid
Arctohedra austrina
n. sp.
Yorkia
is documented from Australia for the first time. An unusual valve (possibly a brachial valve) of enigmatic affinity is also reported and illustrated. Generically, the taxa provide broad regional paleobiogeographic links with the “first discovery limestone” Member of the Coonigan Formation, western New South Wales, and the Current Bush Limestone in the Georgina Basin, northern Australia, and globally, with broadly contemporaneous sequences in western North America, Siberia, and South China.
The first partially articulated scleritome of a tommotiid, Eccentrotheca sp., is described from the Lower Cambrian of South Australia. The Eccentrotheca scleritome consists of individual sclerites; fused in a spiral arrangement, forming a tapering tube-shaped skeleton with an inclined apical aperture and a circular to subcircular cross section. Traditionally, tommotiid sclerites have been assumed to form a dorsal armor of imbricating phosphatic plates in slug-like bilaterians, analogous to the calcareous sclerites of halkieriids. The structure of the Eceentrotheca scleritome is here reinterpreted as a tube composed of independent, irregularly shaped sclerites growing by basal-marginal accretion that were successively fused to form a rigid, protective tubular structure. The asymmetrical shape and sometimes acute inclination of the apical aperture suggests that the apical part of the scleritome was cemented to a hard surface via a basal disc, from which it projected vertically. Rather than being a vagrant member of the benthos, Eccentrotheca most likely represented a sessile, vermiform filter feeder. The new data suggest that the affinities of Eccentrotheca, and possibly some other problematic tommotiids, lie with the lophophorates (i.e., the phoronids and brachiopods), a clade that also possesses a phosphatic shell chemistry and a sessile life habit.
Fourteen species of lingulate brachiopods are documented from allochthonous limestone blocks of the Murrawong Creek Formation in the southern New England Fold Belt, northeastern New South Wales, Australia. The fauna includes Treptotreta jucunda Henderson and MacKinnon 1981, Treptotreta sp. cf. T. sp. nov. Henderson 1992, Amictocracens teres Henderson and MacKinnon 1981, Stilpnotreta magna Henderson and MacKinnon 1981, Anabolotreta tegula Rowell and Henderson 1978, Neotreta orbiculata Koneva 1990, Linnarssonia sp., Linnarssonia sp. cf. L. ophirensis (Walcott 1912), Pegmatreta clavigera sp. nov., Acrothele subsidua (White 1874), Micromitra sp. cf. M. modesta (Lochman 1940), Micromitra sp. Henderson 1992, Lingulella sp. A Henderson 1992, and Kyrshabaktella certa Koneva 1986.The associated trilobite assemblages indicate a medial Middle Cambrian age for the blocks, and the stratigraphic ranges of several of the lingulate species have been extended. The fauna displays biogeographic links at the specific level with northeastern and southeastern Australia, New Zealand, Antarctica, North America, Kazakhstan, Siberia, and Britain; the strongest links (four species in common) are with the Georgina Basin in northeastern Australia and the Tasman Formation in New Zealand.
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