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1.
It has long been assumed that the extant bilaterian phyla generally have their origin in the Cambrian explosion, when they appear in an essentially modern form. Both these assumptions are questionable. A strict application of stem- and crown-group concepts to phyla shows that although the branching points of many clades may have occurred in the Early Cambrian or before, the appearance of the modern body plans was in most cases later: very few bilaterian phyla sensu stricto have demonstrable representatives in the earliest Cambrian. Given that the early branching points of major clades is an inevitable result of the geometry of clade diversification, the alleged phenomenon of phyla appearing early and remaining morphologically static is seen not to require particular explanation. Confusion in the definition of a phylum has thus led to attempts to explain (especially from a developmental perspective) a feature that is partly inevitable, partly illusory. We critically discuss models for Proterozoic diversification based on small body size, limited developmental capacity and poor preservation and cryptic habits, and show that the prospect of lineage diversification occurring early in the Proterozoic can be seen to be unlikely on grounds of both parsimony and functional morphology. Indeed, the combination of the body and trace fossil record demonstrates a progressive diversification through the end of the Proterozoic well into the Cambrian and beyond, a picture consistent with body plans being assembled during this time. Body-plan characters are likely to have been acquired monophyletically in the history of the bilaterians, and a model explaining the diversity in just one of them, the coelom, is presented. This analysis points to the requirement for a careful application of systematic methodology before explanations are sought for alleged patterns of constraint and flexibility. 相似文献
2.
In this attempt to synthesize present knowledge into a coherent story, the Cambrian explosion is interpreted to represent a true adaptive radiation, an event similar to Phanerozoic radiation events in principle but unique in its possibilities. A model of bilaterian evolution helps explain how this particular event involved the sudden initiation of major coelomate phyla. In many of these groups, preservable skeletons are part of the basic body plan. Biochemical‐sequence analyses indicate that acoelomates and pseudocoelomates branched off long before the coelomate radiation. The great differences between Vendian and Cambrian ichnofaunas, therefore, may be the result of a major shift in composition from Vendian acoelomate‐pseudocoelomate‐procoelomate faunas to Phanerozoic faunas dominated by new coelomate groups. 相似文献
3.
The Lower Cambrian Poleta Formation in the White-Inyo Mountains of eastern California contains well-preserved and laterally extensive exposures of the large looping and meandering trace fossil Taphrhelminthopsis nelsoni n.isp. Such traces are typical features on upper bed surfaces of Lower Cambrian shallow marine sandstones and occur with Ediacaran fossils at other localities. Morphologic, sedimentologic and goniogram analyses suggest that the inferred tracemaker was a large soft-bodied echinozoan- or mollusc-grade animal with a volume greater than 14 cm 3 that actively grazed or ingested sediment at the sediment-water interface. Although portions of these traces appear to reflect relatively 'complex' behavior, looping patterns are not periodic as expected for a systematic foraging strategy. T. nelsoni traces are patchy in distribution and commonly associated with suspect-microbial features, suggesting that tracemakers may have been targeting microbial-based or related concentrations of food resources. Such behavioral patterns are typical of shallow late Neoproterozoic-early Cambrian settings, and like suspect-microbial structures are later restricted to deep marine or stressed settings. 相似文献
4.
A new trace fossil, Raaschichnus gundersoni , from the Upper Cambrian St. Lawrence Formation of Wisconsin, was produced by an aglaspidid arthropod. The rusophyciform trace, which occurs singly and in series, is distinctive in showing marks left by the tail-spine. Other trace fossils previously considered to have been made by aglaspidids were probably excavated by animals lacking a tail-spine and with appendage morphologies very different from the aglaspidids of Wisconsin. 相似文献
5.
A set of criteria is suggested for associating a trace fossil with a plausible fossil: (1) close association in the field; (2) concurrent stratigraphic range; (3) a minimal choice of candidates available that could have made the trace; (4) consistent size range between trace fossil population and that of the maker; and (5) consistent biogeographic ranges for both. In the Upper Cambrian Andam Formation of Oman, there is a rare, direct association between trilobite body fossils and the trace fossil Cruziana in the same rock succession. An association of the trilobite Maladioidella cf. colcheni with the trace fossil Cruziana semiplicata is supported by the criteria listed. Maladioidella is associated with C. semiplicata over a wide geographical area in western Gondwana. It is therefore likely that some Cruziana species were made by trilobites. 相似文献
6.
Priapulids and their extinct relatives, the archaeopriapulids and palaeoscolecids, are vermiform, carnivorous ecdysozoans with an armoured, extensible proboscis. These worms were an important component of marine communities during the Palaeozoic, but were especially abundant and diverse in the Cambrian. Today, they comprise just seven genera in four families. Priapulids were among the first groups used to test hypotheses concerning the morphological disparity of Cambrian fossils relative to the extant fauna. A previous study sampled at the generic level, concluding that Cambrian genera embodied marginally less morphological diversity than their extant counterparts. Here, we sample predominantly at the species level and include numerous fossils and some extant forms described in the last fifteen years. Empirical morphospaces for priapulids, archaeopriapulids and palaeoscolecids are relatively insensitive to changes in the taxon or character sample: their overall form has altered little, despite the markedly improved sampling. Cambrian and post-Cambrian genera occupy adjacent rather than broadly overlapping regions of these spaces, and Cambrian species still show lower morphological disparity than their post-Cambrian counterparts. Crucially, the significance of this difference has increased with improved taxon sampling over research time. In contrast with empirical morphospaces, the phylogeny of priapulids, archaeopriapulids and palaeoscolecids derived from morphological characters is extremely sensitive to details of taxon sampling and the manner in which characters are weighted. However, the extant Priapulidae and Halicryptidae invariably resolve as sister families, with this entire clade subsequently being sister group to the Maccabeidae. In our most inclusive trees, the extant Tubiluchidae are separated from these other living taxa by a number of small, intervening fossil clades. 相似文献
7.
Oldhamia trace fossils represent a complex horizontal burrow system usually associated with microbial mats, occurring in the upper part of the Cambrian Stage 2 of the Terreneuvian Series, with a peak in abundance during Stage 3 and 4 of Series 2, and then rapidly disappearing in the Wuliuan Stage of the Miaolingian Series. We here report Oldhamia radiata from the Cambrian Series 2 Xinji Formation in western Henan of North China, associated with microbial mats, as evidenced by the presence of wrinkle structures. The distinctive patterns indicate the exploitation of microbial mats by the tracemakers of O. radiata, featuring a specific feeding strategy under the mats where possibly anoxic and sulfidic conditions prevailed during the early Cambrian. 相似文献
8.
磷酸盐化球状化石是近 2 0年来才相继发现的一类微体化石 ,主要产于下寒武统梅树村阶和震旦系陡山沱组的含磷地层。研究初期其生物亲缘就存在动植物之争 ,但多数专家将其作为分类位置未定的球状化石。近年来 ,由于一些学者研究发现部分磷酸盐化球状化石具有后生动物胚胎发育序列的特征 ,认为它们可能是刺细胞动物或两侧对称动物的胚胎化石 ,从而引起了国内外众多学者 ,尤其是藻类和疑源类专家的关注和热烈讨论。本文在总结球状化石研究进程的基础上 ,对其当前的研究焦点 ,如归属问题、后生动物胚胎化石、保存机制及分类命名 ,进行了简略的评述 ;同时 ,简要介绍了其研究意义。 相似文献
10.
The Cambrian fauna can now reasonably be seen as containing many taxa that lie in the stem-groups of the extant phyla. As such, these fossils suggest how both the ‘body plans’ of extant phyla were assembled, and also how various ‘minor’ phyla relate to the larger groupings of today such as the arthropods and annelids. The various arthropod and lobopod taxa of the Cambrian faunas have been controversial and have generally been considered either as lying in the crown or (occasionally) stem groups of the euarthropods, onychophorans and tardigrades. However, phylogenetic analysis strongly suggests that many of even the most euarthropod-like taxa do not lie within the euarthropod crown-group but are more basal. Further, the commonly expressed view that Cambrian lobopods are in effect stem- or crown-group onychophorans also seems not to be well supported. Lobopods in the Cambrian appear to be diverse and not particularly closely related to one another, and certainly cannot be combined in a monophyletic clade. Both these advances offer hope that the tardigrades (placed as the sister group to the euarthropods in many analyses of extant taxa, here collectively named the Tactopoda) may be more closely related to some of these Cambrian taxa than others. The challenge for both neontologists and palaeontologists is to refine the systematic analysis of both living and fossil taxa in order to maximise the usefulness of the (admittedly few) characters that unite tardigrades to their Cambrian forbears. 相似文献
11.
We describe a new assemblage of small carbonaceous fossils (SCFs) from diagenetically minimally altered clays and siltstones of Terreneuvian age from the Lontova and Voosi formations of Estonia, Lithuania and Russia. This is the first detailed account of an SCF assemblage from the Terreneuvian and includes a number of previously undocumented Cambrian organisms. Recognizably bilaterian‐derived SCFs include abundant protoconodonts (total‐group Chaetognatha), and distinctive cuticular spines of scalidophoran worms. Alongside these metazoan remains are a range of protistan‐grade fossils, including Retiranus balticus gen. et sp. nov., a distinctive funnel‐shaped or sheet‐like problematicum characterized by terminal or marginal vesicles, and Lontohystrichosphaera grandis gen. et sp. nov., a large (100–550 μm) ornamented vesicular microfossil. Together these data offer a fundamentally enriched view of Terreneuvian life in the epicratonic seas of Baltica, from an episode where records of non‐biomineralized life are currently sparse. Even so, the recovered assemblages contain a lower diversity of metazoans than SCF biotas from younger (Stage 4) Baltic successions that represent broadly equivalent environments, echoing the diversification signal recorded in the coeval shelly and trace‐fossil records. Close comparison to the biostratigraphical signal from Fortunian small shelly fossils supports a late Fortunian age for most of the Lontova/Voosi succession, rather than a younger (wholly Stage 2) range. 相似文献
12.
Abstract: The Cambrian index fossil Treptichnus pedum is reported from the Puerto Blanco Formation near Pitiquito, Sonora, Mexico, and new occurrences of the Neoproterozoic index fossil Cloudina are reported from the underlying La Ciénega Formation. Considered together, these fossils constrain the location of the Ediacaran/Cambrian boundary in Mexico. The suite of fossils in this sequence is important because it provides an independent biostratigraphic datum for constraining the position of Laurentia during the onset of metazoan diversification, for testing proposed lithostratigraphic correlations among western North American Neoproterozoic–Cambrian successions, and for testing regional tectonic paradigms such as the Mojave-Sonora Megashear hypothesis. 相似文献
13.
The genomes of taxa whose stem lineages branched early in metazoan history, and of allied protistan groups, provide a tantalizing outline of the morphological and genomic changes that accompanied the origin and early diversifications of animals. Genome comparisons show that the early clades increasingly contain genes that mediate development of complex features only seen in later metazoan branches. Peak additions of protein‐coding regulatory genes occurred deep in the metazoan tree, evidently within stem groups of metazoans and eumetazoans. However, the bodyplans of these early‐branching clades are relatively simple. The existence of major elements of the bilaterian developmental toolkit in these simpler organisms implies that these components evolved for functions other than the production of complex morphology, preadapting the genome for the morphological differentiation that occurred higher in metazoan phylogeny. Stem lineages of the bilaterian phyla apparently required few additional genes beyond their diploblastic ancestors. As disparate bodyplans appeared and diversified during the Cambrian explosion, increasing complexity was accommodated largely through changes in cis‐regulatory networks, accompanied by some additional gene novelties. Subsequently, protein‐coding genic richness appears to have essentially plateaued. Some genomic evidence suggests that similar stages of genomic evolution may have accompanied the rise of land plants. 相似文献
14.
Shell decollation is a growth strategy that has been adopted by a number of invertebrate taxa to offset the metabolic and ecological disadvantages of shell growth. However, little is known about the origin and evolution of this process. We here describe well-preserved specimens of the hyolith Cupitheca decollata sp. nov. preserving the decollation process, from the early Cambrian Yu'anshan Formation ( c. 518 Ma) of South China. Based on a large number of specimens collectively representing different developmental stages, we use high-resolution X-ray microtomography and scanning electronic microscopy to reconstruct the process of decollation in this taxon. Cupitheca is among the earliest known small shelly fossils, and thus our discovery confirms that periodic decollation had evolved by the onset of the Cambrian explosion, reflecting the high intensity of the predator–prey arms race in early Cambrian ecosystems. A comparison between the decollation processes of Cupitheca and other shelly invertebrates suggests that periodic decollation and the associated molecular mechanisms of calcium dissolution, uptake, allocation and deposition may have had multiple independent origins. 相似文献
15.
The interrelationships and evolutionary history of molluscs have seen great advances in the last decade. Recent phylogenetic studies have allowed alternative morphology‐based evolutionary scenarios to be tested and, most significantly, shown that the aplacophorans are sister group to polyplacophorans (chitons), corroborating palaeontological and embryological evolutionary scenarios in which aplacophorans are secondarily simplified from a chiton‐like ancestor. Aplacophoran morphology therefore does not represent the plesiomorphic condition for molluscs as a whole. The mollusc crown group radiated in the Early Cambrian, and rapidly thereafter, stem lineages to the major molluscan classes emerged: cephalopods, gastropods, bivalves (= pelecypods), monoplacophorans, rostroconchs (inferred stem scaphopods) and aculiferans. This attests to the fast, adaptive radiation of the crown group during the Cambrian explosion. Kimberella from the latest Ediacaran exhibits several molluscan traits, which justifies its position as a molluscan stem‐group member, rather than as a more basal Lophotrochozoan. The interrelationships among the conchiferan molluscs are still a matter of contention and require further palaeontological and molecular phylogenetic scrutiny. 相似文献
16.
We describe an assemblage of small carbonaceous fossils (SCFs) and acritarchs from cored siltstones of the Lappajärvi impact structure, west‐central Finland. Previous studies had detected a depauperate acritarch biota ascribed to a deep Proterozoic origin—this age, however, was based on recovery of long‐ranging poorly age‐diagnostic sphaeromorphs. To resolve the age and provenance of these crater sediments, we applied low‐manipulation processing techniques optimized for retrieval of larger organic‐walled microfossils. Our study revealed a previously undetected assemblage containing numerous metazoan SCFs consisting of flattened ‘protoconodonts’ (grasping spines assignable to total group Chaetognatha) and a distinctive fossilised chaeta, possibly representing the oldest known annelid remains. Phylogenetically problematic fossils include various acritarchs (large Leiosphaeridia sp., Tasmanites tenellus, smaller sphaeromorphs, Synsphaeridium, Archaeodiscina and Granomarginata) and filamentous forms ( Palaeolyngbya‐ and Rugosoopsis‐like filaments, Siphonophycus), likely representing prokaryotic or protistan grades of organisation. As well as adding new diversity to an emerging SCFs record, these data substantially refine the age of these sediments by more than half a billion years, to an early Cambrian Terreneuvian age. More specifically, the assemblage is equivalent to that of the Lontova Formation from the Baltic States and northwest Russia, but is previously unreported from Finland. Identification of Lontova‐type SCFs/organic‐walled microfossils at Lappajärvi further constrains the poorly resolved extent of maximum flooding during the early Cambrian in Baltica. Renewed attention should be directed to strata that have thus far produced only biostratigraphically long‐ranging or ambiguous palynological assemblages—‘SCF‐style’ processing can reveal hitherto undetected, age‐informative microfossils that are otherwise selectively removed in conventional palynological studies. 相似文献
17.
The Cambrian explosion is a grand challenge to science today and involves multidisciplinary study. This event is generally believed as a result of genetic innovations, environmental factors and ecological interactions, even though there are many conflicts on nature and timing of metazoan origins. The crux of the matter is that an entire roadmap of the evolution is missing to discern the biological complexity transition and to evaluate the critical role of the Cambrian explosion in the overall evolutionary context. Here, we calculate the time of the Cambrian explosion by a “C-value clock”; our result quite fits the fossil records. We clarify that the intrinsic reason of genome evolution determined the Cambrian explosion. A general formula for evaluating genome size of different species has been found, by which the genome size evolution can be illustrated. The Cambrian explosion, as a major transition of biological complexity, essentially corresponds to a critical turning point in genome size evolution. 相似文献
18.
This study investigates whether the recently described Cambrian fossil Haikouella (and the very similar Yunnanozoon) throws light on the longstanding problem of the origin of craniates. In the first rigorous cladistic analysis of the relations of this animal, we took 40 anatomical characters from Haikouella and other taxa (hemichordates, tunicates, cephalochordates, conodont craniates and other craniates, plus protostomes as the outgroup) and subjected these characters to parsimony analysis. The characters included several previously unrecognized traits of Haikouella, such as upper lips resembling those of larval lampreys, the thick nature of the branchial bars, a mandibular branchial artery but no mandibular branchial bar, muscle fibers defining the myomeres, a dark fibrous sheath that defines the notochord, conclusive evidence for paired eyes, and a large hindbrain and diencephalon in the same positions as in the craniate brain. The cladistic analysis produced this tree: (protostomes, hemichordates (tunicates, (cephalochordates, (Haikouella, (conodonts + other craniates))))), with the \"Haikouella + craniate\" clade supported by bootstrap values that ranged from 81-96%, depending on how the analysis was structured. Thus, Haikouella is concluded to be the sister group of the craniates. Alternate hypotheses that unite Haikouella with hemichordates or cephalochordates, or consider it a basal deuterostome, received little or no support. Although it is the sister group of craniates, Haikouella is skull-less and lacks an ear, but it does have neural-crest derivatives in its branchial bars. Its craniate characters occur mostly in the head and pharynx; its widely spaced, robust branchial bars indicate it ventilated with branchiomeric muscles, not cilia. Despite its craniate mode of ventilation, Haikouella was not a predator but a suspension feeder, as shown by its cephalochordate-like endostyle, and tentacles forming a screen across the mouth. Haikouella was compared to pre-craniates predicted by recent models of craniate evolution and was found to fit these predictions closely. Specifically, it fits Northcutt and Gans' prediction that the change from ciliary to muscular ventilation preceded the change from suspension feeding to predatory feeding; it fits Butler's claim that vision was the first craniate sense to start elaborating; it is consistent with the ideas of Donoghue and others about the ancestor of conodont craniates; and, most strikingly, it resembles Mallatt's prediction of the external appearance of the ancestral craniate head. By contrast, Haikouella does not fit the widespread belief that ancestral craniates resembled hagfishes, because it has no special hagfish characters. Overall, Haikouella agrees so closely with recent predictions about pre-craniates that we conclude that the difficult problem of craniate origins is nearly solved. 相似文献
19.
Trilobites, key components of early Palaeozoic communities, are considered to have been invariably fully marine. Through the integration of ichnological, palaeobiological, and sedimentological datasets within a sequence-stratigraphical framework, we challenge this assumption. Here, we report uncontroversial trace and body fossil evidence of their presence in brackish-water settings. Our approach allows tracking of some trilobite groups foraying into tide-dominated estuaries. These trilobites were tolerant to salinity stress and able to make use of the ecological advantages offered by marginal-marine environments migrating up-estuary, following salt wedges either reflecting amphidromy or as euryhaline marine wanderers. Our data indicate two attempts of landward exploration via brackish water: phase 1 in which the outer portion of estuaries were colonized by olenids (Furongian–early late Tremadocian) and phase 2 involving exploration of the inner to middle estuarine zones by asaphids (Dapingian–Darriwilian). This study indicates that tolerance to salinity stress arose independently among different trilobite groups. 相似文献
20.
Echinoderms are unique among animal phyla in having a pentaradial body plan, and their fossil record provides critical data on how this novel organization came about by revealing intermediate stages. Here, we report a spiral-plated animal from the early Cambrian of Morocco that is the most primitive pentaradial echinoderm yet discovered. It is intermediate between helicoplacoids (a bizarre group of spiral-bodied echinoderms) and crown-group pentaradiate echinoderms. By filling an important gap, this fossil reveals the common pattern that underpins the body plans of the two major echinoderm clades (pelmatozoans and eleutherozoans), showing that differential growth played an important role in their divergence. It also adds to the striking disparity of novel body plans appearing in the Cambrian explosion. 相似文献
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