| 1. |
- Chen, Feiyang, et al.
(författare)
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Cambrian ecological complexities: Perspectives from the earliest brachiopod – supported benthic communities in the early Cambrian Guanshan Lagerstätte
- 2022
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Ingår i: Gondwana Research. - Amsterdam : Elsevier. - 1342-937X .- 1878-0571. ; 107, s. 30-41
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Tidskriftsartikel (refereegranskat)abstract
- The Cambrian radiation is characterized by the emergence of diverse bilaterian animal phyla and theestablishment of complex marine ecosystems. The Guanshan Biota records an unusual ecological transition from trilobite- to brachiopod-dominated communities during Cambrian Stage 4. This community transition is accompanied by direct evidence of in situ biological interactions such as durophagous pre-dation and kleptoparasitism. Here we describe new material from the Guanshan biota, focusing on an association of palaeoscolecidomorphs and brachiopods with parasitic tube worms that occur on micro-bedding planes. The bedding plane assemblages are dominated by the organophosphatic brachiopod Neobolus wulongqingensis encrusted with kleptoparasitic tube-dwelling worms, along with infaunal palaeoscolecidans. Taphonomic and sedimentological evidence indicates that these specimens are com-monly preserved in life position, and thus the association between individuals represent potential biological interactions. This case study reveals that ecosystems during the early Cambrian exhibited a well-developed system of tiering and a complex trophic network, easily distinguished from the simple communities typical of precursor deposits in the Ediacaran. Brachiopods forming extremely dense concentrations on the sea floor are effectively acting as ecosystem engineers, not only to stabilize the soft-substrate seafloor, but also act as an alternative substrate for the oldest empirically demonstrated kleptoparasites.The in situ biological interactions preserved in the Guanshan Biota are critical for filling gaps in ourknowledge of ecosystem complexity in the Cambrian.
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| 2. |
- Hu, Yazhou, et al.
(författare)
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Distinguishing borings and burrows in intraclasts : Evidence from the Cambrian (Furongian) of North China
- 2023
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Ingår i: Sedimentary Geology. - : Elsevier. - 0037-0738 .- 1879-0968. ; 443
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Tidskriftsartikel (refereegranskat)abstract
- Hardgrounds represent synsedimentary cemented stratigraphic beds that form at or near the seafloor. Borings represent a key line of evidence for investigations of hardground development and record the evolution of bioerosion and boring organisms. The unequivocal identification of borings is done through identification of the crosscutting relationship between the proposed boring and a hard substrate, such as lithoclasts and/or shells, with morphological criteria able to be used when dealing with a homogeneous substrate, such as micritic hardgrounds. Bioeroded hardgrounds and burrows with a micrite halo/lining are subject to fracturing and reworking, resulting in accumulations of intraclasts in flat-pebble conglomerates (FPC). The recognition of borings and broken burrows with a halo can be challenging in FPC. Using trace fossils preserved in situ and in FPC from late Cambrian carbonates of North China, we establish a set of criteria for distinguishing borings from burrows with a halo in FPC. Features such as the relative volume of burrows and borings versus the host pebble and the number of traces per pebble, the cross-cutting relationship with different colored laminae, and the presence of pyrite or glauconite encrustations can all be invoked to aid recognition of borings. Examination of the cross-cutting relationship and encrustation of trace fossils are not sufficient on their own. Our results suggest caution is necessary in defining borings in FPC, particularly as synsedimentary deformation of burrows with a halo in late Cambrian FPC can create structures that resemble borings.
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| 3. |
- Liang, Yue, et al.
(författare)
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Evolutionary contingency in lingulid brachiopods across mass extinctions
- 2023
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Ingår i: Current Biology. - : Elsevier BV. - 0960-9822 .- 1879-0445. ; 33:8, s. 1565-1572.e3
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Tidskriftsartikel (refereegranskat)abstract
- Morphology usually serves as an effective proxy for functional ecology,1,2,3,4,5 and evaluating morphological, anatomical, and ecological changes permits a deeper understanding of the nature of diversification and macroevolution.5,6,7,8,9,10,11,12 Lingulid (order Lingulida) brachiopods are both diverse and abundant during the early Palaeozoic but decrease in diversity over time, with only a few genera of linguloids and discinoids present in modern marine ecosystems, resulting in them frequently being referred to as “living fossils.”13,14,15 The dynamics that drove this decline remain uncertain, and it has not been determined if there is an associated decline in morphological and ecological diversity. Here, we apply geometric morphometrics to reconstruct global morphospace occupation for lingulid brachiopods through the Phanerozoic, with results showing that maximum morphospace occupation was reached by the Early Ordovician. At this time of peak diversity, linguloids with a sub-rectangular shell shape already possessed several evolutionary features, such as the rearrangement of mantle canals and reduction of the pseudointerarea, common to all modern infaunal forms. The end Ordovician mass extinction has a differential effect on linguloids, disproportionally wiping out those forms with a rounded shell shape, while forms with sub-rectangular shells survived both the end Ordovician and the Permian-Triassic mass extinctions, leaving a fauna predominantly composed of infaunal forms. For discinoids, both morphospace occupation and epibenthic life strategies remain consistent through the Phanerozoic. Morphospace occupation over time, when considered using anatomical and ecological analyses, suggests that the limited morphological and ecological diversity of modern lingulid brachiopods reflects evolutionary contingency rather than deterministic processes.
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| 4. |
- Liang, Yue, et al.
(författare)
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When lingulid brachiopods became infaunal(?) : Perspectives from the morphological and anatomical information
- 2023
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Ingår i: Estonian journal of earth sciences. - : Estonian Academy Publishers. - 1736-4728 .- 1736-7557. ; 72:1, s. 140-140
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Tidskriftsartikel (refereegranskat)abstract
- Morphology usually serves as an effective proxy for functional ecology, and the evaluation of morphological, anatomical, and ecological changes allows for a deeper understanding of the nature of diversification and macroevolution. Lingulid (Order Lingulida) brachiopods were diverse and abundant during the early Palaeozoic, but decreased in diversity over time, with only a few genera of linguloids and discinoids present in modern marine ecosystems, frequently referred to as 'living fossils'. The dynamics that drove this decline remain unclear and it has not been determined if there is an associated decline in morphological and ecological diversity. We applied geometric morphometrics to reconstruct global morphospace occupied by lingulid brachiopods through the Phanerozoic, with results showing that maximum morphospace occupation was reached in the Early Ordovician. At this time of peak diversity, linguloids with sub-rectangular shells already possessed several evolutionary features common to all modern infaunal forms such as the rearrangement of mantle canals and reduction of the pseudointerarea. The endOrdovician mass extinction had a differential effect on linguloids, disproportionally wiping out those with rounded shells whilst forms with sub-rectangular shells survived both the endOrdovician and the Permian-Triassic mass extinctions, with post-extinction faunas predomi nantly composed of infaunal forms. For discinoids, both morphospace occupation and epi-benthic life strategies remain consistent through the Phanerozoic. Analysis of the morphospace occupation of lingulids over time, taking into account their body size, anatomical features and ecological changes, suggests that the reduced morphological and ecological diversity observed in modern lingulid brachiopods reflects evolutionary contingency rather than deterministic processes.
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| 5. |
- Luo, Mei, et al.
(författare)
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First Report of Small Skeletal Fossils from the Upper Guojiaba Formation (Series 2, Cambrian), Southern Shaanxi, South China
- 2023
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Ingår i: Biology. - : MDPI. - 2079-7737. ; 12:7
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Tidskriftsartikel (refereegranskat)abstract
- A small skeletal fossil assemblage is described for the first time from the bioclastic limestone interbeds of the siltstone-dominated Guojiaba Formation, southern Shaanxi, China. The carbonate-hosted fossils include brachiopods (Eohadrotreta zhujiahensis, Eohadrotreta zhenbaensis, Spinobolus sp., Kuangshanotreta malungensis, Kyrshabaktella sp., Lingulellotreta yuanshanensis, Eoobolus incipiens, and Eoobolus sp.), sphenothallids (Sphenothallus sp.), archaeocyaths (Robustocyathus sp. and Yukonocyathus sp.), bradoriids (Kunmingella douvillei), chancelloriids sclerites (Onychia sp., Allonnia sp., Diminia sp., Archiasterella pentactina, and Chancelloria cf. eros), echinoderm plates, fragments of trilobites (Eoredlichia sp.), and hyolithelminths. The discovery of archaeocyaths in the Guojiaba Formation significantly extends their stratigraphic range in South China from the early Tsanglangpuian at least to the late Chiungchussuan. Thus, the Guojiaba Formation now represents the lowest known stratigraphic horizon where archaeocyath fossils have been found in the southern Shaanxi area. The overall assemblage is most comparable, in terms of composition, to Small skeletal fossil (SSF) assemblages from the early Cambrian Chengjiang fauna recovered from the Yu’anshan Formation in eastern Yunnan Province. The existing position that the Guojiaba Formation is correlated with Stage 3 in Cambrian Series 2 is strongly upheld based on the fossil assemblage recovered in this study.Simple SummarySmall skeletal fossils are reported for the first time from the early Cambrian Guojiaba Formation, southern Shaanxi, China. All specimens were recovered from bioclastic limestone interbeds and encompass a wide variety of skeletal clades, including brachiopods, sphenothallids, archaeocyaths, bradoriids, sponge spicules, echinoderm plates, and trilobite spines. The archaeocyaths described herein are considerably older than those described from the Xiannvdong Formation, which was previously assumed to contain the lowest archaeocyath-bearing horizons in South China. The brachiopod Lingulellotreta yuanshanensis is recorded for the first time from the Fucheng area, with previous records confined mainly to the Chengjiang Fauna-bearing Yu’anshan Formation in the lower Cambrian, the eastern Yunnan Province. Micro-morphological and elemental analyses of the small skeletal fossil assemblages were carried out using SEM, BSEM, and Micro X–ray fluorescence. The skeletal fauna in the Guojiaba Formation resembles the assemblages recovered from the upper Yu’anshan Formation (Chengjiang Fauna) in eastern Yunnan Province. Based on the recovered assemblage, the biostratigraphic age of the Guojiaba Formation correlates with the Chiungchussuan Stage (Stage 3 of Cambrian Series 2).
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| 6. |
- Topper, Timothy P, et al.
(författare)
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Competition and mimicry : the curious case of chaetae in brachiopods from the middle Cambrian Burgess Shale
- 2015
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Ingår i: BMC Evolutionary Biology. - : Springer Science and Business Media LLC. - 1471-2148. ; 15:1
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Tidskriftsartikel (refereegranskat)abstract
- Background: One of the first phyla to acquire biomineralized skeletal elements in the Cambrian, brachiopods represent a vital component in unraveling the early evolution and relationships of the Lophotrochozoa. Critical to improving our understanding of lophotrochozoans is the origin, evolution and function of unbiomineralized morphological features, in particular features such as chaetae that are shared between brachiopods and other lophotrochozoans but are poorly understood and rarely preserved. Micromitra burgessensis and Paterina zenobia from the middle Cambrian Burgess Shale are among the most remarkable examples of fossilized chaetae-bearing brachiopods. The form, functional morphology, evolutionary and ecological significance of their chaetae are studied herein.Results: Like in Recent forms, the moveable but semi-rigid chaetae fringe both the dorsal and ventral mantle margins, but in terms of length, the chaetae of Burgess Shale taxa can exceed twice the maximum length of the shell from which it projects. This is unique amongst Recent and fossil brachiopod taxa and given their size, prominence and energy investment to the organism certainly had an important functional significance. Micromitra burgessensis individuals are preserved on hard skeletal elements, including conspecific shells, Tubulella and frequently on the spicules of the sponge Pirania muricata, providing direct evidence of an ecological association between two species. Morphological analysis and comparisons with fossil and extant brachiopod chaetae point to a number of potential functions, including sensory, defence, feeding, defouling, mimicry and spatial competition.Conclusions: Our study indicates that it is feasible to link chaetae length to the lack of suitable substrate in the Burgess Shale environment and the increased intraspecific competition associated with this. Our results however, also lend support to the elongated chaetae as an example of Batesian mimicry, of the unpalatable sponge Pirania muricata. We also cannot discount brachiopod chaetae acting as a sensory grille, extending the tactile sensitivity of the mantle into the environment, as an early warning system to approaching predators.
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| 7. |
- Topper, Timothy, P., et al.
(författare)
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DO BRACHIOPODS SHOW SUBSTRATE-RELATEDPHENOTYPIC VARIATION? A CASE STUDY FROM THE BURGESS SHALE
- 2017
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Ingår i: Palaeontology. - London : Wiley. - 0031-0239 .- 1475-4983. ; 60:2, s. 269-279
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Tidskriftsartikel (refereegranskat)abstract
- As sessile, benthic filter feeders, brachiopods share an intimate relationship with their chosen substrate. Individuals of Micromitra burgessensis in the Burgess Shale Formation are preserved in life position, attached to a range of hard substrates, including skeletal debris, conspecific brachiopods, sponges and enigmatic tubes. Here we investigate the phenotypic variability of M. burgessensis associated with differing substrate attachments. We apply geometric morphometrics to test for variation by plotting landmarks on the exterior of ventral and dorsal valves of M. burgessensis specimens that are preserved attached to different substrates. Using principal component, canonical variate analyses and ANOVA, we determine that there is some variation in shape related to substrate. Canonical variate analyses, for ventral valves and dorsal valves, indicate that specimens attached to the same substrate are recognizable in shape from specimens attached to other substrate types. The strength of differentiation however, is not robust and combined with our discriminate analysis of separate populations suggests that there is the potential for substrates to exercise only weak control over the morphology of Brachiopoda.
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| 8. |
- Topper, Timothy P., et al.
(författare)
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Survival on a soft seafloor : life strategies of brachiopods from the Cambrian Burgess Shale
- 2015
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Ingår i: Earth-Science Reviews. - : Elsevier BV. - 0012-8252 .- 1872-6828. ; 151, s. 266-287
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Forskningsöversikt (refereegranskat)abstract
- Abstract Understanding the structure of benthic communities in the Cambrian remains a major challenge. Direct evidence for species interrelationships is rare and therefore past ecological interactions typically cannot be reconstructed with great accuracy. Here we reveal the community patterns and modes of life of brachiopods – one of the most important filter-feeding groups of Cambrian ecosystems – from the Cambrian Burgess Shale LagerstÀtte. Burgess Shale brachiopods attached to a range of hard substrates, including skeletal debris, conspecific brachiopods and enigmatic tubes, with an overwhelming preference for attachment on the demosponge Pirania muricata. The dominance of P. muricata as a substrate choice – even in bedding assemblages where P. muricata individuals are rare – and similarities to the gregarious attachment strategies of extant brachiopod species suggests that brachiopods larvae in the Burgess Shale community selected their attachment substrates. The distribution of brachiopod taxa is also intricately linked with the presence of suitable hard substrates, with species declining in bedding assemblages where their preferred hard substrates are absent. In addition, brachiopods in the Burgess Shale community are predominantly low epifaunal tierers and do not exploit niches high in the water column, despite the presence of suitable attachment sites. Our analysis of tiering height versus host height indicates that there is no selection by brachiopod larvae in regard to the height of attachment and individuals attached at the first point of contact with the selected substrate. Through comparisons with the ‘early’ Cambrian Chengjiang Biota, we confirm that by the ‘middle’ Cambrian (Series 3, Stage 5) brachiopods had already developed a range of attachment strategies similar to some modern brachiopod populations. Our results provide significant insight into the ecological constraints and adaptability of brachiopods in the earliest animal communities of the Cambrian.
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| 9. |
- Zhang, Zhifei, et al.
(författare)
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An encrusting kleptoparasite-host interaction from the early Cambrian
- 2020
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Ingår i: Nature Communications. - : Springer Science and Business Media LLC. - 2041-1723. ; 11
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Tidskriftsartikel (refereegranskat)abstract
- Parasite–host systems are pervasive in nature but are extremely difficult to convincingly identify in the fossil record. Here we report quantitative evidence of parasitism in the form of a unique, enduring life association between tube-dwelling organisms encrusted to densely clustered shells of a monospecific organophosphatic brachiopod assemblage from the lower Cambrian (Stage 4) of South China. Brachiopods with encrusting tubes have decreased biomass (indicating reduced fitness) compared to individuals without tubes. The encrusting tubes orient tightly in vectors matching the laminar feeding currents of the host, suggesting kleptoparasitism. With no convincing parasite–host interactions known from the Ediacaran, this widespread sessile association reveals intimate parasite–host animal systems arose in early Cambrian benthic communities and their emergence may have played a key role in driving the evolutionary and ecological innovations associated with the Cambrian radiation.
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| 10. |
- Zhang, Zhiliang, et al.
(författare)
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Fossil evidence unveils an early Cambrian origin for Bryozoa
- 2021
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Ingår i: Nature. - : Springer Science and Business Media LLC. - 0028-0836 .- 1476-4687. ; 599:7884, s. 251-255
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Tidskriftsartikel (refereegranskat)abstract
- Bryozoans (also known as ectoprocts or moss animals) are aquatic, dominantly sessile, filter-feeding lophophorates that construct an organic or calcareous modular colonial (clonal) exoskeleton. The presence of six major orders of bryozoans with advanced polymorphisms in lower Ordovician rocks strongly suggests a Cambrian origin for the largest and most diverse lophophorate phylum. However, a lack of convincing bryozoan fossils from the Cambrian period has hampered resolution of the true origins and character assembly of the earliest members of the group. Here we interpret the millimetric, erect, bilaminate, secondarily phosphatized fossil Protomelission gatehousei from the early Cambrian of Australia and South China as a potential stem-group bryozoan. The monomorphic zooid capsules, modular construction, organic composition and simple linear budding growth geometry represent a mixture of organic Gymnolaemata and biomineralized Stenolaemata character traits, with phylogenetic analyses identifying P. gatehousei as a stem-group bryozoan. This aligns the origin of phylum Bryozoa with all other skeletonized phyla in Cambrian Age 3, pushing back its first occurrence by approximately 35 million years. It also reconciles the fossil record with molecular clock estimations of an early Cambrian origination and subsequent Ordovician radiation of Bryozoa following the acquisition of a carbonate skeleton.
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