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Lower Cambrian Vendobionts from China and Early Diploblast Evolution

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Lower Cambrian Vendobionts from China and Early Diploblast Evolution REPORTS provide exceptional morphological detail. Pre- Lower Cambrian Vendobionts from sumably they were rapidly buried by storm events, and most are oriented at a shallow angle China and Early Diploblast Evolution to the bedding plane. The split between part and counterpart is therefore oblique, necessitating D.-G. Shu,1,2* S. Conway Morris,3* J. Han,1 Y. Li,4 X.-L. Zhang,1 H. Hua,1 Z.-F. Zhang,1 composite reconstructions of each specimen J.-N. Liu,1 J.-F. Guo,1 Y. Yao,1 K. Yasui5 (Figs. 1, C to E, and 2, B and C). The body is foliate, with a bluntly terminating Ediacaran assemblages immediately predate the Cambrian explosion of metazoans and should stalk that lacks obvious attachment structures have played a crucial role in this radiation. Their wider relationships, however, have remained (Figs. 1; A, C, D, and F; and 2; A, B, and D). refractory and difficult to integrate with early metazoan phylogeny. Here, we describe a frondlike Body length ranges between 2.5 and 7.5 cm. fossil, Stromatoveris (S. psygmoglena sp. nov.), from the Lower Cambrian Chengjiang Lagersta¨tte Orientations were adopted for convenience and (Yunnan, China) that is strikingly similar to Ediacaran vendobionts. The exquisite preservation imply no direct homologies with other orga- reveals closely spaced branches, probably ciliated, that appear to represent precursors of the nisms. Upper and lower surfaces are markedly diagnostic comb rows of ctenophores. Therefore, this finding has important implications for the different. The former bears prominent branches early evolution of this phylum and related diploblasts, some of which independently evolved a with quite pronounced relief. Most specimens frondose habit. are too incomplete to count the precise number of branches Etypical width was circa (ca.) 1 mm^, diacaran assemblages represent Earth_s and 2). Five of them, including the holotype, but in the holotype about 15 are visible on earliest complex macroscopic organisms were collected near Meishucun, whereas the either side of the midline. The latter is defined Ein the history of life (1). They lack skeletal other three are from Jianshan, near Haikou. Spec- by a relatively narrow groove (Figs. 1A and 2, hard parts but are relatively diverse (1, 2), have a imens show preservation typical of other soft- A and E). The most proximal branches arise defined community structure (3), and show a bodied fossils from Chengjiang and therefore along an adaxially inclined line, but all branches global distribution (2). Yet their phylogenetic interpretation remains highly controversial. Rad- ical reassignments to lichens (4) and fungi (5) have won little support (1, 6), but the traditional assignments to animals (7) remain problematic. A few taxa can be compared, with varying degrees of reliability, to known animal groups, including sponges (8), cnidarians (9), mollusks (10), and arthropods (11). Most, however, remain in phylogenetic limbo because comparisons to either extant phyla or putative stem groups are frustratingly imprecise. The provocative vendo- biont hypothesis (12–14) seeks to unify disparate taxa, including forms otherwise assigned to groups as separate as cnidarians and arthropods, on the basis of a distinctive body plan with a modular quilted construction and possibly syn- cytial tissue. If correct, this hypothesis has two major implications. First, did at least some vendobionts derive independently from protis- tans, or are they a sister group of either animals (15) or even diploblasts? Crucial in this respect are various frondlike fossils, some of which have been compared to the cnidarians, specifi- cally the pennatulaceans (7, 16), whereas others are clearly akin to other vendobionts (12, 17). Here, we describe eight specimens of Stromatoveris psygmoglena gen. sp. nov., (18) a frondose fossil from the Lower Cambrian Chengjiang Lagerst.tte near Kunming, Yunnan, and interpret them as a new vendobiont (Figs. 1 1Early Life Institute and Key Laboratory of Continental Dynamics, Northwest University, Xi’an 710069, China. 2School of Earth Sciences and Resources, China University of Geosciences, Beijing 100083, China. 3Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge 4 CB2 3EQ, UK. College of Earth Science and Land Resources, Fig. 1. The Cambrian vendobiont S. psygmoglena,gen.sp.nov.(A, B,andG) Holotype, ELI-Vend-05-001. Chang’an University, Xi’an 710054, China. 5Marine Biological Laboratory, Graduate School of Science, Hiroshima University, (A) Upper surface, (B) fragment (counterpart) of lower surface, and (G) enlargement of the boxed area in 2445 Mukaishima-cho, Onomichi, Hiroshima 722-0073, (A). (C and D) ELI-Vend-05-002. (C) Composite photograph of upper and lower surfaces and (D) distal part Japan. of upper surface removed to reveal lower surface. (E) ELI-Vend-05-003, composite photo of part and *To whom correspondence should be addressed. E-mail: counterpart to show both upper and lower surfaces. (F) ELI-Vend-05-004, composite photograph of part [email protected] (D.-G.S.); [email protected] (S.C.M.) and counterpart to show upper surface and axial rod located between upper and lower surfaces. www.sciencemag.org SCIENCE VOL 312 5 MAY 2006 731 REPORTS are parallel and fan distally at a shallow angle to The lower region broadly appears to be feeding is conjectural and depends in part on the midline. Branches are mostly single, but oc- divisible into two regions. More distally an ovoid phylogenetic comparisons. Given the absence of casionally they bifurcate or fuse in an irregular central smooth area is strongly concave (Figs. 1D definitive zooids, one possibility is that the fashion (Figs. 1A and 2, A and E). In some and 2B), but adjacent there are subdued ridges branches were ciliated and served to transport food specimens, branches were filled with sediment, that run slightly oblique to the body margin and in particles via the narrow grooves between the suggesting that during life they were hollow the opposite direction to the branches on the up- branches. Assuming a density of cilia comparable tubes. The lateral margins of the branches show per surface. In addition, there is a diffuse orna- to typical suspension feeders, this would provide a occasional short prongs or spurs. These may mentation, roughly ovoid. More basally there is highly effective trap. What appear to be inter- represent interconnections either between the subdued ribbing and a prominent leopard-skin connections between the branches (Figs. 1; A, E, branches or possibly into the interior of the ornamentation, at least near the margin (Figs. 1B and G; and 2; A and D) suggest an exchange frond. Branch numbers appear to have shown and 2A). The lower surface becomes smooth system that presumably also connected to the only a modest increase with body size, and dif- toward the basal region. One specimen (fig. S1, interior. During the organism_s life, the interior ferences in relative width in larger specimens C and E) shows striking arcuate structures on the was presumably filled with fluid or gelatinous suggest that the individual branches continued to stem. Although these might be another type of tissue. The central axis (Figs. 1F and 2D), grow. Distally, however, new branches can be surface ornamentation, they are interpreted as analogous to the axial rod of pennatulaceans, seen to differentiate from the surface of the body-wall support, possibly collagenous. presumably provided additional support, but no blade (Figs. 1A and 2, A and E). The interior contains a substantial sediment fill other internal organs are discernible. The branches were evidently firmly attached (Figs. 1, C to E, and 2, C and D, and fig. S1B), Although it cannot be placed in any known to the body, but some branches show slight im- more pronounced in the stalk, suggesting its cross genus, in overall form Stromatoveris is similar to brication. The branches bear transverse, closely section was approximately circular, whereas the a number of Ediacaran frondlike fossils. It is most spaced striations (Figs. 1, A and G, and 2, A and blade was somewhat compressed. As with the similar to the otherwise poorly documented B, and fig. S1, A and D), but there is no evidence branches, this sediment infill suggests the interior Khatyspytia (19), but the latter is more slender for zooids. The regions between the branches was largely hollow and possibly fluid-filled during and has a larger number of shorter branches. were narrow and recessed, and in some speci- the organism_s life. An axial structure (ca. 0.4 mm General resemblances also exist with the fronds mens there are associated dark strands (Fig. 1; A, across), with apparently ferruginous preservation, Vaizitsinia (19), Charniodiscus (16, 20), Glaess- C,andEtoG;andfig.S1;A,B,F,andG). is located nearer to the upper surface. In one nerina (16), and, more remotely, such forms as These may have been discrete structures, possibly specimen (Figs. 1F and 2D), it has been partially Charnia (19, 20). The phylogenetic position of canal-like and presumably located in the body excavated, and toward the base a similar structure these Ediacaran fronds (and by implication wall. They have irregular margins (Fig. 1E), which occurs, but it runs transversely and possibly was Stromatoveris) is controversial (21), with oppos- may have been extensions connecting to other rotated to its present position as a result of decay. ing views favoring cnidarians, especially pen- regions of the body wall. Distal to the branches The organism was benthic and embedded in natulaceans (7, 16), or vendobionts (12, 14). Key the surface of the frond is smooth, although a the seafloor by the stalk. Whether it lived upright points in these differing interpretations include continuation of the midline occurs as a narrow or recumbent is less clear. The apparent rigidity of attachment of the branches to the frond and meandering groove (Figs.
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