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![Head anatomy of Fuxianhuia. (a, c–f) Specimens of Fuxianhuia protensa from the Yu'anshan Formation (Chengjiang biota). (a) Full specimen; NIGPAS166941. (c) Inset of head area showing successive anterior sclerites and mandibles; area of Figure 2d, e, k indicated; NIGPAS166941. (d) Head area showing anterior sclerites and mandibles with palps. The pair of post-oral plate-like structures (en) are interpreted as the compressed basal endites of the post-oral limbs; area of Figure 2f, g indicated; see also Supplementary Fig. 1; NIGPAS166940. (e) Head area with three-dimensional preservation showing the overlap of the labrum on top of the mandibles (see also Fig. 3); NIGPAS166939. (f) Head area showing the three-podomerous morphology of the mandibles; area of Figure 2j indicated; NIGPAS166937. Asterisks in (c) and (d) mark the location of the atrium oris. (b) Specimen of Fuxianhuia xiaoshibaensis from the Hongjingshao Formation (Xiaoshiba biota). Note midline separation between lateral halves of the carapace (solid arrowheads). Symmetrical median bulges are possibly caused by the mandibles underneath the carapace; NIGPAS166943. Abbreviations: an, antennule; en, endites; hy, hypostome; lb, labrum; le, left eye; ma, mandible; mg, mandibular gnathal lobe; os, ocular sclerite; pn, podomere n; pd, peduncle of eye stalk; re, right eye. Scale bars: (a) 6 mm; (b–f) 2 mm.]()
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![Details of the head anatomy of Fuxianhuia. (a, b, h, i, k) Specimens of Fuxianhuia xiaoshibaensis from the Hongjingshao Formation (Xiaoshiba biota). (a) Anteriormost section of the head in ventral view; NIGPAS166945. (b) Close-up of the ocular sclerite, hypostome and proximal portion of mandibles shown in (a). Asterisks mark the two lobes of the ocular sclerite in ventral view; NIGPAS166945. (h) Close-up of isolated mandible; NIGPAS166944. (i) Close-up of sclerotized tip of mandibular palp; NIGPAS166942. (k) Detail of articulation between stalk and eye (see Fig. 1c); NIGPAS166941. (c–g, j, l) Specimens of Fuxianhuia protensa from the Yu'anshan Formation (Chengjiang biota). (c) Carbonaceous remains inside ocular lobe (open arrowhead); NIGPAS166938. (d) Anterior section of the head showing arrangement of the three anterior sclerites; NIGPAS166941 (see Fig. 1c for location). (e) Close-up of right mandible; dotted line replicates proximal margin of masticatory surface, suggesting incomplete separation between anterior and posterior faces; NIGPAS166941 (see Fig. 1c for location). (f) Close-up of post-oral area, showing possible endites preserved as single blocks; NIGPAS166940 (see Fig. 1d and Supplementary Fig. 1). (g) Close-up on mandible with two-segmented palp, with possible differentiated proximal element partially covering the labrum; NIGPAS166940 (see Fig. 1d and Supplementary Fig. 1). (j) Close-up of sclerotized tip of mandibular palp (see Fig. 1f); NIGPAS166937. (l) Post-mandibular area; note rounded elements near the base of the appendages (solid arrowheads), possibly representing enditic outgrowths (see also Fig. 3); NIGPAS166939. Abbreviations: ca, carapace; can, cephalic appendage n; en, endites; hy, hypostome; lb, labrum; le, left eye; ma, mandible; mg, mandibular gnathal lobe; mp, mandibular palp; os, ocular sclerite; pn, podomere n; pd, peduncle of eye stalk; pe, putative proximal element of gnathal mandibular lobe; re, right eye. Scale bars: (a, d, g, h, l) 1 mm; (b, c, e, f, i, k) 0.5 mm; (j) 0.25 mm.]()
![Stacked tomographic images of Fuxianhuia protensa NIGPAS166939 (see Fig. 1e). Only the left eye, hypostome, labrum, mandibles and carapace are originally visible on the specimen. The asterisk represents the location of the hypostome, rendered here as a depression, perhaps due to the relative lack of pyritization. This demonstrates the distinct anatomical identity of the hypostome compared to the labrum. Note also that only the distal portions of the three posterior cephalic limbs are preserved on the left side of the animal, suggesting the first two pairs are reduced. Abbreviations: an, antennule; ca, carapace; cn, cephalic segment n; can, cephalic appendage n; lb, labrum; le, left eye; m, mouth; mp, mandibular palp; os, ocular sclerite; pd, peduncle of eye stalk; re, right eye; tn, trunk segment n.]()
![Reconstructions summarizing the anatomy of Fuxianhuia's head. (a) Drawing of the mandibles with putative proximal element and multi-segmented, enditic basis of a small post-oral appendage. Note slightly diverging faces on masticatory margin of mandibles (straight lines); solid arrowheads point to coarse teeth on these margins. The presence of a fifth, proximal endite on the post-oral basipod is conjectural. (b, c) Stylized diagrams. For clarity, the carapace and putative proximal gnathal element have not been drawn. Features of the protocerebral (ocular) somite are in blue and of the deutocerebral somite in green. The labrum (yellow) and antennules in (b) have been made transparent to show underlying features. Mandibles are ochre coloured. The central black circle represents the mouth. The mouth depression (atrium oris) is surrounded by endites (light yellow). (b) Dorsal view. The mandibles come in contact in front of the mouth (closest element to the viewer), below the labrum. (c) Ventral view. Abbreviations: an, antennule; ds, distal sclerotized tip; en, endites; hy, hypostome; lb, labrum; le, left eye; ma, mandible; mg, mandibular gnathal lobe; mp, mandibular palp; os, ocular sclerite; pe, putative proximal element of the mandibular gnathal lobe; pd, peduncle of eye stalk; pn, podomere n.]()
![Representative fossil taxa from Chengjiang background mudstone beds (BGB): (A–D) external skeletons of euarthropods (A—Fuxianhuia protensa, B—Yunnanocephalus yunnanensis, C—bivalved arthropod, D—Acanthomeridion serratum); (E) distal claw of frontal appendage of Amplectobelua symbrachiata; (F) potential hexactinellid sponge Triticispoingia diagonata; (G) vetulicolian Beidazoon venustum; (H) undetermined taxon; (I–M) fossils with soft internal cellular organ/tissue preserved (I—potential mollusc; J—Naraoia spinosa; K—Ovalicephalus mirabilis; L—Urokodia aequalis; M—Isoxys auritus).]()
![Comparison of head anatomy between euthycarcinoids and fuxianhuiids. (a, c) Apankura machu JUY-P24 from the Santa Rosita Formation, Argentina (image courtesy of G. Edgecombe): (a) head area, ventral view. (b, d) Fuxianhuia protensa NIGPAS166941, dorsal view. (c, d) Interpretative drawings. In comparison with other euthycarcinoids, the mandibles of Apankura are remarkably large, whereas the wide labrum known in other taxa is interpreted here as incompletely preserved. The central H-shaped feature is impossible to identify with absolute confidence, but its shape and location are consistent with the hypopharynx reported in Euthycarcinus and Heterocrania. Apankura displays an ‘eye bar' akin to those of fuxianhuiids, but it is interpreted as the doublure of the large ocular sclerite present in other euthycarcinoids and preserved similarly in ventral view. Abbreviations: ca, carapace; hx, hypopharynx; hy, hypostome; lb, labrum; le, left eye; ma, mandible; os, ocular sclerite; re, right eye. Scale bars: (a) 0.25 mm; (b) 2 mm.]()
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![FIGURE 18—Diversity of prey-predator relationships in the lower Cambrian Maotianshan Shale biota inferred from fossil evidence (e.g., coprolite, gut contents, and functional morphology). Exact role of phytoplankton and possible zooplankton undocumented. 1 = ctenophores (Chen and Zhou, 1997; Dzik, 2002); 2 = eldoniids (Zhu et al., 2002); 3 = chaetognaths (Chen and Huang, 2002); 4 = anomalocaridids (Chen et al., 1994; Collins, 1996); 5 = Fuxianhuia and related forms (Chen et al., 1995a; Zhu et al., 2004); 6 = naraoiids (Vannier and Chen 2002); 7 = priapulids (Huang et al., 2004); 8 = Isoxys (Vannier and Chen 2001); 9 = vetulicolids (Shu et al. 2001); 10 = waptiids; 11 = Haikouella (early chordate; Chen et al., 1999); 12 = trilobites; 13 = bradoriids (Shu et al., 1999); 14 = hyoliths; 15 = Ercaia (Chen et al., 2001). Not to scale]()
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![FIGURE 3—Fossils from the Lower Cambrian Yu'anshan Formation at Mafang (Figs. A–K) and Ercaicun (Fig. L) near Haikou, Kunming. All bar scales 10 mm except: C, I, = 5 mm; L, 1 mm. Abbreviations: an, antenna; ap, appendage; as, axial spine; ex, exopodite; eye, compound eye; fc, free cheek; hs, head shield; leg, tentacle-like leg; te, trunk end. A) Three lobopodians (arrowed separately in different colors) of Microdictyon sinicum coexisting with their host Eldonia eumorpha (RCCBYU 10414). B) Fuxianhuia protensa (RCCBYU 10192), an adult with three juveniles (arrows). C) Eoredlichia intermedia with one free cheek detached (RCCBYU 10199). D) Four laterally compressed Leanchoilia illecebrosa (RCCBYU 10193). E) Concentration of exoskeletons of Yunnanocephalus yunnanensis (RCCBYU 10366). F) Dense accumulation of Yunnanozoon lividum (RCCBYU 10418). G) Eoredlichia intermedia (RCCBYU 10190). H) Xandarella spectaculum (RCCBYU 10191), with head shield, antennae and appendages preserved in separate laminae. I) Naraoia longicaudata (RCCBYN 10367) with nonsymmetry gut fillings (arrow). J) Dorsoventrally compressed Leanchoilia illecebrosa (RCCBYU 10194). K) Paucipodia inermis (RCCBYU 10186) preserved across two thin layers marked by the dark shadow. L) Kunmingella douvillei (RCCBYU 10196) with outstretched appendages and trunk]()
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![FIGURE 6. Fossils from the lower Cambrian Yu'anshan Formation at Mafang showing the preservational characteristics of the two taphofacies. All scale bars = 5 mm, except G–H = 2 mm. Comparison of the preservation of Fuxianhuia protensaHou, 1987 (arthropod) within A) event (BA7-047) and B) background (BA12-108) mudstones. Comparison of the preservation of Onychodictyon feroxHou, Ramsköld, & Bergström, 1991 (lobopod) within C) event (XBA7-C3-020) and D) background (XBA6-E3-011) mudstones. Comparison of the preservation of Mafangscolex sinensis (Hou & Sun, 1988) (priapulid) within E) event (BA6-57) and F) background mudstones (BA11-103). Comparison of the preservation of Kunmingella douvilleiMansuy, 1912 (arthropod) within G) event (BA5-142) and H) background mudstones (BA10-049). General preservational characteristics of event mudstones. I) Lingulella chengjiangensisJin, Hou, & Wang, 1993 (brachiopod) (BA5-123). J) Undescribed soft-shelled lingulid brachiopod (BA8-081). K) Cindarella eucalla Chen, Ramsköld, Edgecombe, & Zhou in Chen et al., 1996 (arthropod) (BA4-057). L) Waptia ovataLee, 1975 (arthropod) (XBA5-A1-003). M) Archisaccophyllia kunmingensisHou et al., 2005 (cnidarian) (BA9-139). General preservational characteristics of background mudstones. N) Retifacies abnormalisHou, Chen, & Lu, 1989 (BA5-044). O) Eoredlichia intermediaLu, 1941 (arthropod) (BA8-206). P) Diandongia pistaRong, 1974 (brachiopod) (BA11-69). Q) Beidazoon venustumShu, 2005 (vetulicolid) (BA11-084). R) Fossil in a event mudstone, showing the background–event bed boundary, burrowing in the lower part of the background mudstone bed, a graded silt band within the event mudstone, and the position and posture of Eldonia eumorpha Sun & Hou, 1987 (enigmatic animal) (BA9-264). S) Mafangscolex sinensis (Hou & Sun, 1988) (priapulid) preserved while burrowing through an event mudstone layer (XBA6-E3-24).]()
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Fuxianhuia
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Head anatomy of Fuxianhuia . ( a , c – f ) Specimens of Fuxianhuia prote... Available to Purchase
in Fuxianhuiids are mandibulates and share affinities with total-group Myriapoda
> Journal of the Geological Society
Published: 22 March 2021
Fig. 1. Head anatomy of Fuxianhuia . ( a , c – f ) Specimens of Fuxianhuia protensa from the Yu'anshan Formation (Chengjiang biota). ( a ) Full specimen; NIGPAS166941. ( c ) Inset of head area showing successive anterior sclerites and mandibles; area of Figure 2d, e, k indicated
Journal Article
Fuxianhuiids are mandibulates and share affinities with total-group Myriapoda Available to Purchase
Journal: Journal of the Geological Society
Publisher: Geological Society of London
Published: 22 March 2021
Journal of the Geological Society (2021) 178 (5): jgs2020-246.
...Fig. 1. Head anatomy of Fuxianhuia . ( a , c – f ) Specimens of Fuxianhuia protensa from the Yu'anshan Formation (Chengjiang biota). ( a ) Full specimen; NIGPAS166941. ( c ) Inset of head area showing successive anterior sclerites and mandibles; area of Figure 2d, e, k indicated...
FIGURES
Image
Details of the head anatomy of Fuxianhuia . ( a , b , h , i , k ) Spec... Available to Purchase
in Fuxianhuiids are mandibulates and share affinities with total-group Myriapoda
> Journal of the Geological Society
Published: 22 March 2021
Fig. 2. Details of the head anatomy of Fuxianhuia . ( a , b , h , i , k ) Specimens of Fuxianhuia xiaoshibaensis from the Hongjingshao Formation (Xiaoshiba biota). ( a ) Anteriormost section of the head in ventral view; NIGPAS166945. ( b ) Close-up of the ocular sclerite, hypostome
Image
Stacked tomographic images of Fuxianhuia protensa NIGPAS166939 (see Fig.... Available to Purchase
in Fuxianhuiids are mandibulates and share affinities with total-group Myriapoda
> Journal of the Geological Society
Published: 22 March 2021
Fig. 3. Stacked tomographic images of Fuxianhuia protensa NIGPAS166939 (see Fig. 1e ). Only the left eye, hypostome, labrum, mandibles and carapace are originally visible on the specimen. The asterisk represents the location of the hypostome, rendered here as a depression, perhaps due
Image
Reconstructions summarizing the anatomy of Fuxianhuia 's head. ( a ) Drawi... Available to Purchase
in Fuxianhuiids are mandibulates and share affinities with total-group Myriapoda
> Journal of the Geological Society
Published: 22 March 2021
Fig. 5. Reconstructions summarizing the anatomy of Fuxianhuia 's head. ( a ) Drawing of the mandibles with putative proximal element and multi-segmented, enditic basis of a small post-oral appendage. Note slightly diverging faces on masticatory margin of mandibles (straight lines); solid
Image
Representative fossil taxa from Chengjiang background mudstone beds (BGB): ... Available to Purchase
Published: 12 July 2024
Figure 2. Representative fossil taxa from Chengjiang background mudstone beds (BGB): (A–D) external skeletons of euarthropods (A— Fuxianhuia protensa , B— Yunnanocephalus yunnanensis , C—bivalved arthropod, D— Acanthomeridion serratum ); (E) distal claw of frontal appendage of Amplectobelua
Image
Comparison of head anatomy between euthycarcinoids and fuxianhuiids. ( a , ... Available to Purchase
in Fuxianhuiids are mandibulates and share affinities with total-group Myriapoda
> Journal of the Geological Society
Published: 22 March 2021
Fig. 4. Comparison of head anatomy between euthycarcinoids and fuxianhuiids. ( a , c ) Apankura machu JUY-P24 from the Santa Rosita Formation, Argentina (image courtesy of G. Edgecombe): ( a ) head area, ventral view. ( b , d ) Fuxianhuia protensa NIGPAS166941, dorsal view. ( c , d
Journal Article
Unveiling an ignored taphonomic window in the early Cambrian Chengjiang Biota Available to Purchase
Journal: Geology
Publisher: Geological Society of America
Published: 12 July 2024
Geology (2024) 52 (10): 753–758.
...Figure 2. Representative fossil taxa from Chengjiang background mudstone beds (BGB): (A–D) external skeletons of euarthropods (A— Fuxianhuia protensa , B— Yunnanocephalus yunnanensis , C—bivalved arthropod, D— Acanthomeridion serratum ); (E) distal claw of frontal appendage of Amplectobelua...
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FIGURE 18 —Diversity of prey-predator relationships in the lower Cambrian M... Available to Purchase
in Early Cambrian Food Chain: New Evidence from Fossil Aggregates in the Maotianshan Shale Biota, SW China
> PALAIOS
Published: 01 February 2005
; Dzik, 2002 ); 2 = eldoniids ( Zhu et al., 2002 ); 3 = chaetognaths ( Chen and Huang, 2002 ); 4 = anomalocaridids ( Chen et al., 1994 ; Collins, 1996 ); 5 = Fuxianhuia and related forms ( Chen et al., 1995a ; Zhu et al., 2004 ); 6 = naraoiids ( Vannier and Chen 2002 ); 7 = priapulids (Huang et al
Journal Article
A New Arthropod Jugatacaris agilis n. gen. n. sp. from the Early Cambrian Chengjiang Biota, South China Available to Purchase
Journal: Journal of Paleontology
Publisher: Paleontological Society
Published: 01 May 2011
Journal of Paleontology (2011) 85 (3): 567–586.
... of Fuxianhuia ( Hou, 1987b ) is reinterpreted as bivalved carapace ( Budd, 2002 ). Superficial resemblances in “bivalved” carapace are, however, probably due to convergent evolution as recognized by a number of authors ( Briggs, 1978 ; Hou and Bergström, 1997 ; Taylor, 2002 ). An anus, preserved...
FIGURES
Journal Article
GRAVITATIONAL CONSTRAINTS ON THE BURIAL OF CHENGJIANG FOSSILS Available to Purchase
Journal: PALAIOS
Publisher: SEPM Society for Sedimentary Geology
Published: 01 July 2007
PALAIOS (2007) 22 (4): 448–453.
...). Chengjiang fossils are thought to have experienced only gentle transport prior to burial ( Hou et al., 2004 ). For example, some adults of Fuxianhuia —a common Chengjiang arthropod—co-occur with their small-sized juveniles ( Fig. 3B ). This mixed-sized association suggests limited transport prior to burial...
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FIGURE 3 —Fossils from the Lower Cambrian Yu'anshan Formation at Mafang (Fi... Available to Purchase
Published: 01 July 2007
shield; leg, tentacle-like leg; te, trunk end. A) Three lobopodians (arrowed separately in different colors) of Microdictyon sinicum coexisting with their host Eldonia eumorpha (RCCBYU 10414). B) Fuxianhuia protensa (RCCBYU 10192), an adult with three juveniles (arrows). C) Eoredlichia intermedia
Journal Article
THE LOWER CAMBRIAN CRUSTACEAN PECTOCARIS FROM THE CHENGJIANG BIOTA, YUNNAN, CHINA Available to Purchase
Journal: Journal of Paleontology
Publisher: Paleontological Society
Published: 01 July 2004
Journal of Paleontology (2004) 78 (4): 700–708.
... Perspicaris and Canadaspis and the very different Fuxianhuia Hou, 1987 (placed closest to Pectocaris ) in a clade supposed to share a subchelate frontal appendage and a straight hinge line. There is no sign of a subchelate appendage in most of these disparate genera, including Pectocaris...
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Journal Article
Information landscapes and sensory ecology of the Cambrian Radiation Available to Purchase
Journal: Paleobiology
Publisher: Paleontological Society
Published: 01 May 2010
Paleobiology (2010) 36 (2): 303–317.
... ; Chen 2009 ). Tardipolypods, such as Hallucigenia , might have had a simple type of visual system, such as eyespots, but lacked image-forming eyes ( Chen and Zhou 1997 ). Compound eyes first appeared in proarthropods such as Fuxianhuia and Shankouia . The primitive condition of the image-forming...
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Journal Article
Early Cambrian Food Chain: New Evidence from Fossil Aggregates in the Maotianshan Shale Biota, SW China Available to Purchase
Journal: PALAIOS
Publisher: SEPM Society for Sedimentary Geology
Published: 01 February 2005
PALAIOS (2005) 20 (1): 3–26.
... ; Dzik, 2002 ); 2 = eldoniids ( Zhu et al., 2002 ); 3 = chaetognaths ( Chen and Huang, 2002 ); 4 = anomalocaridids ( Chen et al., 1994 ; Collins, 1996 ); 5 = Fuxianhuia and related forms ( Chen et al., 1995a ; Zhu et al., 2004 ); 6 = naraoiids ( Vannier and Chen 2002 ); 7 = priapulids (Huang et al...
FIGURES
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FIGURE 6. Fossils from the lower Cambrian Yu'anshan Formation at Mafang sho... Available to Purchase
in QUANTITATIVE ANALYSIS OF TAPHOFACIES AND PALEOCOMMUNITIES IN THE EARLY CAMBRIAN CHENGJIANG LAGERSTÄTTE
> PALAIOS
Published: 01 January 2009
FIGURE 6. Fossils from the lower Cambrian Yu'anshan Formation at Mafang showing the preservational characteristics of the two taphofacies. All scale bars = 5 mm, except G–H = 2 mm. Comparison of the preservation of Fuxianhuia protensa Hou, 1987 (arthropod) within A) event (BA7-047) and B
Journal Article
Taphonomy of Naraoiids (Arthropoda) from the Middle Cambrian Kaili Biota, Guizhou Province, South China Available to Purchase
Journal: PALAIOS
Publisher: SEPM Society for Sedimentary Geology
Published: 01 February 2006
PALAIOS (2006) 21 (1): 15–25.
... as digestive organs (e.g., Burgessia , Canadaspis , Leanchoilia , Molaria , Naraoia , Odaraia , Perspicaris , Plenocaris , Skania , and Tegopelte from the Burgess Shale, and Almenia , Canadaspis , Fortiforceps , Fuxianhuia , Kuamaia , Leanchoilia , Naraoia , Retifacies , Sinoburius...
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Journal Article
EVIDENCE OF PRIAPULID SCAVENGING FROM THE EARLY CAMBRIAN CHENGJIANG DEPOSITS, SOUTHERN CHINA Available to Purchase
Journal: PALAIOS
Publisher: SEPM Society for Sedimentary Geology
Published: 01 December 2007
PALAIOS (2007) 22 (6): 691–694.
... the Chengjiang ( Babcock, 2003 ). Fragments of a eodiscoid trilobite have been found in a Fuxianhuia -like arthropod from the Middle Cambrian Kaili fauna ( Zhu et al., 2004 ), suggesting that arthropods by this time had used gnathobases and large appendages to break down the exoskeleton of some prey...
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Journal Article
A new Burgess Shale–type assemblage from the “thin” Stephen Formation of the southern Canadian Rockies Available to Purchase
Journal: Geology
Publisher: Geological Society of America
Published: 01 September 2010
Geology (2010) 38 (9): 811–814.
.... Great appendage arthropod A is broadly similar to Fuxianhuia from the Chengjiang biota ( Hou, 1987 ) and has a pair of short frontal appendages, a truncated head shield bearing one pair of large pedunculate eyes, and at least 32 trunk somites associated with presumably biramous limbs ( Fig. 3F ; Fig...
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Journal Article
The early Cambrian Kuamaia lata , an artiopodan euarthropod with a raptorial frontal appendage Open Access
Journal: Journal of Paleontology
Publisher: Paleontological Society
Published: 01 September 2024
Journal of Paleontology (2024) 98 (5): 808–820.
..., Systematics Association Special Volume Series : London , Chapman & Hall , p. 125 – 138 .
Chen , A.
, 2005 , A new Fuxianhuia -like arthropod of the early Cambrian Chengjiang fauna in Yunnan : Yunnan Geology , v. 24 , p. 108 – 113...
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