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A juvenile-rich palaeocommunity of the lower Cambrian Chengjiang biota sheds light on palaeo-boom or palaeo-bust environments


The fossil record, including the record of Burgess Shale-type deposits, is biased towards late ontogenetic stages. Larval stages, juvenile and subadult specimens exist but are very rare and often preserved as phosphatic fossils, resulting in biased population structures. Here, we report a new Burgess Shale-type Lagerstätte from Haiyan, China. The Haiyan palaeocommunity is extraordinary in that it is rich in fossils of early and middle ontogenetic stages of various phyla, with eggs also commonly found in the studied interval. This Lagerstätte also hosts a considerable number of new taxa—many related to later biotas of Gondwana and Laurentia. We propose that the deposit may either preserve one of the earliest nurseries in the fossil record or, alternatively, records several attempted invasions. Our study highlights the complexity of biotas and their interactions in the lower Cambrian ocean and calls for a better understanding of the mechanisms responsible for the observed spatial variation of fossil community composition in the Cambrian.

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Fig. 1: Location map and early Cambrian stratigraphy of the Haiyan Lagerstätte.
Fig. 2: Exceptionally preserved juvenile specimens from the Haiyan Lagerstätte.
Fig. 3: Larvae and eggs from the Haiyan Lagerstätte.
Fig. 4: Body sizes and taxa from the Haiyan Lagerstätte.
Fig. 5: Comparison of dominant ecological groupings within three different Chengjiang biota localities and the Walcott Quarry of the Burgess Shale. A complete list of ecological groupings for each of the localities can be found in Supplementary Table 4.

Data availability

All elements necessary to allow interpretation and replication of the results, including full datasets, are provided in Supplementary Tables 14. All specimens are kept at YKLP at Yunnan University. Photographic material of the studied material is available from the corresponding authors on request.


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We thank M. Wilson and R. LaVine for comments on earlier versions of the manuscript. We thank N. Hughes and J. Schiffbauer for their reviews that improved the manuscript. This paper is a contribution to IGCP668, Equatorial Gondwanan History and Early Palaeozoic Evolutionary Dynamics. X.Y. was supported by the National Natural Science Foundation of China (grant nos. 41062001, 41562001) and the State Key Laboratory of Palaeobiology and Stratigraphy (Nanjing Institute of Geology and Palaeontology, CAS) (grant no. 103113). S.P. was supported by the State Key Laboratory of Palaeobiology and Stratigraphy (Nanjing Institute of Geology and Palaeontology, CAS) (grant no. 20191101). D.Z. was supported by Key Research Program of the Institute of Geology & Geophysics, Chinese Academy of Sciences (grant no. IGGCAS-201905).

Author information

Authors and Affiliations



X.Y. designed and led the study. X.Y. and S.P. completed fieldwork and collected all specimens used in the study. X.Y. photographed the specimens and completed EDS analyses. X.Y. completed all the measurements. X.Y. and J.K. completed all analyses. X.Y., J.K. and S.R.K. created the figures. X.Y., J.K., S.R.K., S.P., D.Y. and Y.L. wrote the manuscript. All authors discussed and agreed in interpretation of the data.

Corresponding authors

Correspondence to Xianfeng Yang or Julien Kimmig.

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The authors declare no competing interests.

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Peer review information Nature Ecology & Evolution thanks Nigel Hughes, James Schiffbauer and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. Peer reviewer reports are available.

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Extended data

Extended Data Fig. 1 Detailed stratigraphy of the Haiyan Lagerstätte.

Stratigraphic succession of the Cambrian Stage 3 Haiyan section with ranges of major fossil lineages preserved in the Haiyan Lagerstätte and the upper soft-bodied interval.

Extended Data Fig. 2 Taphonomy of the Haiyan Lagerstätte.

a–g, Soft-bodied tissue preservation of a new hemichordate, YKLP14443. a, Microprobe EDS spectrum of the matrix. b, Stereo microscope image. c, Backscatter electron (BSE) image showing pyrite on the surface of fossil, black arrow indicates the boundary of the fossil and the matrix. d, Elemental map overlay, including silicon (Si), aluminium (Al), oxygen (O), potassium (K), iron (Fe). e–j, individual elemental maps e, Iron. f, Silicon. g, Aluminium. h, Carbon (C). i, Sulfur (S). j, Oxygen. k, Unpolished mudstone from the lower level of the Haiyan Lagerstätte. l, Polished mudstone from the lower level of the Haiyan Lagerstätte. Scale bars, 200 μm (c–i), 1 mm (b),10 mm (k, l).

Extended Data Fig. 3 Exceptionally preserved new specimens of rare Chengjiang biota taxa in the Haiyan Lagerstätte.

a, Larva of the deuterostome Yunnanozoon sp., YKLP14449. b, New ctenophore, YKLP14450. c, A juvenile of the problematicum Archotuba elongata, YKLP14451. d, A juvenile specimen of the sipunculan worm Cambrosipunculus sp., YKLP14452. e, Priapulid worm Anningvermis multispinosa, YKLP14453. f, Vetulicolian Heteromorphus confusus, YKLP14454. g, Priapulid worm Acosmia maotiania, YKLP14455. h, Phoronid Iotuba chengjiangensis, YKLP14456. i, Arthropod Urokodia aequalis, YKLP14457. j, The problematicum Eldonia eumorpha, YKLP14458, preserved with a priapulid worm Maotianshania cylindrica and the brachiopod Longtancunella chengjiangensis. k, The problematicum Archotuba elongata, YKLP14459, preserved together with specimens of the palaeoscolecid worm Maotianshania cylindrica, the entoproct Cotyledion tylodes and the priapulid worm Paraselkirkia sinica. Scale bars, 500 μm (c,), 1 mm (a, b, d), 5 mm (e, g, i), 10 mm (f, h, k), 20 mm (j). df = dorsal fin.

Extended Data Fig. 4 Further elements of the Haiyan Lagerstätte biota.

a, A probable juvenile specimen of the mollusk Wiwaxia sp., YKLP14473. b, A juvenile specimen of the problematicum Eldonia eumorpha, YKLP14474. c, A juvenile specimen of the chancelloriid Allonnia phrixothrix, YKLP14475. d, The cnidarian Xianguangia sinica, YKLP14476. e, A new ctenophoran, YKLP14477. f, The brachiopod Diandongia pista, YKLP14478. g, The brachiopod Lingulellotreta malongensis, YKLP14479. h, An indeterminate juvenile sponge, YKLP14480. i–k, Hyolithids. i, Ambrolinevitus ventricosus, YKLP14481. j, Burithes yunnanensis, YKLP14482, k, Burithes yunnanensis, YKLP14483. l, The cnidarian Archisaccophyllia kunmingensis, YKLP14484. m–p, Sponges. m, A new sponge, YKLP14485. n, Crumillospongia biporosa, YKLP144486. o, Choiaella radiata, YKLP14487. p, A new sponge, YKLP14488. q, The brachiopod Heliomedusa orienta, YKLP14489. r, The alga Fuxianospira gyrata, YKLP14490. Scale bars, 500 μm (e), 1 mm (c, n), 2 mm (a), 5 mm (b, d–m, o, q), 10 mm (p, r).

Extended Data Fig. 5 Further elements of the Haiyan Lagerstätte biota.

a, Molt of an indeterminate vetulicolian, YKLP14523. b, Molt of the deuterostome Yunnanozoon lividum, YKLP14524. c, Cluster of juvenile bivalved arthropods, recording possible gregarious behaviour, YKLP14525. d, Cluster of hemichordate fossils, recording possible gregarious behaviour, YKLP14526. e, Circular aggregate of trilobite and hyolith shell fragments, possibly a coprolite, YKLP14527. f, Burrows under the carapace of an indeterminate bivalved arthropod, YKLP14528. Scale bars, 5 mm (f), 10 mm (a, b), 20 mm (c–e).

Extended Data Fig. 6 Further arthropods from the Haiyan Lagerstätte.

a, Missizhouia longicaudata, YKLP14491. b, An uncertain juvenile arthropod, YKLP14492. c, A juvenile specimen of Tanglangia sp., YKLP14493. d, Kunmingella douvillei, YKLP14494. e, Primicaris larvaformis, YKLP 14495. f, Chuandianella ovata, YKLP14496. g, Fortiforceps foliosa, YKLP14497. h, Sunella sp., YKLP14498. i, Pectocaris sp., YKLP14499. j, Eoredlichia intermedia, YKLP14500. k, Naraoia spinosa, YKLP14501. l, Acanthomeridion sp., YKLP14502. m, Yunnanocephalus yunnanensis., YKLP14503. n, Isoxys auritus, YKLP14504. o, Chengjiangocaris sp., YKLP14505. p, Combinivalvula chengjiangensis, YKLP14506. q, A juvenile specimen of Chengjiangocaris sp., YKLP14507. Scale bars, 500 μm (b), 1 mm (e), 2 mm (a, d, n, p, q), 5 mm (b, c, f–l, o), 10 mm (m).

Extended Data Fig. 7 Vermiform fossils from the Haiyan Lagerstätte.

a–d, Priapulid worms. a, Palaeopriapulites parvus, YKLP14508. b, Sicyophorus rara, YKLP14509. c, Indeterminate priapulid worm, YKLP14510. d, Cricocosmia jinningensis, YKLP14511. e, Indeterminate vermiform fossil, YKLP14512. f–i, Priapulid worms f, Anningvermis multispinosa, YKLP14513. g, Xiaoheiqingella peculiaris, YKLP14514. h, Maotianshania cylindrica, YKLP14515. i, Yunnanpriapulus sp., YKLP14516. j, The lobopod Onychodictyon ferox, YKLP14517. k, The lobopod Hallucigenia fortis, YKLP14518. l, The priapulid worm Eximipriapulus sp., YKLP14519. m, Indeterminate vermiform fossil, YKLP14520. n, The priapulid worm Sicyophorus rara, YKLP14521. o, The lobopod Cardiodictyon catenulum, YKLP14522. Scale bars, 500 μm (k), 1 mm (a, m), 2 mm (b, c, i, j, l, n), 5 mm (e–h, o), 10 mm (d).

Extended Data Fig. 8 Exceptionally preserved new species from the Haiyan Lagerstätte.

a, A new Amiskwia sagittiformis-like animal, YKLP14435. b, close-up of the fin of (a). c, New chancelloriid sp. A., YKLP14436. d, New chancelloriid, sp. B., YKLP14437. e, The holotype of the deuterostome Herpetogaster haiyanensis, YKLP14404. f, New frond-like taxon, YKLP14438. g, New chaetognath preserving the gut with some gut content, YKLP14439. h, A new colonial hydroid, YKLP14440. i, Early ontogenetic stage of the new chaetognath, YKLP14441 j, k, New hemichordate, extending the range of the range of the Enteropneusta into the lower Cambrian. j, YKLP14442. k, YKLP14443. l, New wiwaxiid, preserving detailed sclerites, YKLP14444. m, New arthropod, YKLP14445. n, A new problematic taxon preserving fine spines (arrows), YKLP14446. o, New sponge, preserving detailed spicules, YKLP14447. p, New lophophorate preserving internal soft tissues, YKLP14448. Scale bars, 5 mm (a, e–i), 2 mm (b–d, l, k, m–p), 500 μm (j).

Extended Data Fig. 9 Adult arthropods from the Haiyan Lagerstätte.

a, A juvenile specimen of Saperion glumaceum, YKLP14460. b, Indeterminate bivalved arthropod, YKLP14461. c, Missizhouia longicaudata, YKLP14462. d, Pectocaris sp., YKLP14463. e, Parapaleomerus sp., YKLP14464. f, Posterior portion of thorax of an indeterminate arthropod, YKLP14465. g, Leanchoilia illecebrosa, YKLP14466. h, Isoxys paradoxus, YKLP14467. i, Indeterminate arthropod, YKLP14468. j, Amplectobelua symbrachiata, YKLP14469. k, Naraoia spinosa, YKLP14470. l, Haifengella corona, YKLP14471. m, Branchiocaris yunnanensis, YKLP14472. Scale bars, 1 mm (a), 5 mm (c, f), 10 mm (b, e, g, i, k, m), 20 mm (d, h, j, l).

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Yang, X., Kimmig, J., Zhai, D. et al. A juvenile-rich palaeocommunity of the lower Cambrian Chengjiang biota sheds light on palaeo-boom or palaeo-bust environments. Nat Ecol Evol 5, 1082–1090 (2021).

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