Paleogeography

The geography of the Cambrian world differed greatly from that of the present day. The geographic reconstruction is based on integrated geologic and biological evidence. Fossils in continental-shelf deposits indicate the presence of at least three major faunal provinces (or biogeographical regions) during much of the Cambrian Period.

The most distinct faunal province surrounded the continent of Laurentia. Paleomagnetic evidence indicates that Laurentia was located over the paleoequator during most or all of Cambrian time. This geographic interpretation is supported by the presence of thick, warm-water carbonate-platform deposits that accumulated in a broad belt encircling the continent. These carbonates are commonly flanked on the inner shelf by lagoonal shale and nearshore sandstone deposits. On the outer shelf, the carbonates commonly grade into laminated mudstone and shale that accumulated in deeper water. At times, two almost mutually exclusive ecosystems are separated by temperature and salinity barriers in the shallow water on the carbonate platforms. Inner restricted-shelf deposits were characterized by sparse low-diversity communities that tended to be highly endemic (confined to a particular region). Outer open-shelf deposits are characterized by high-diversity ecosystems that were widely distributed around the continent. Fossils are usually most abundant and most diverse near the outer margins of the carbonate platform. Because Laurentia remained nearly intact structurally, it is ideal for studying the relationships between Cambrian environments and communities of organisms around a low-latitude Cambrian continent.

Another Cambrian faunal province surrounded the small continent of Baltica, which was located in middle to high southern latitudes. Cambrian shelf deposits of Baltica are relatively thin, rarely exceeding 250 metres (820 feet) in thickness, and are composed primarily of sandstone and shale. Seemingly as a consequence of cool-water environments, carbonate deposits are relatively minor and very thin. The wide distribution of many species from the nearshore to deep-shelf environments of Baltica suggests that no significant restrictions in shelf dispersal occurred, unlike the shallow carbonate platforms of Laurentia.

The largest Cambrian faunal province is located around Gondwana, which extended from the low northern latitudes to the high southern latitudes, just short of the South Pole. The rocks and fossil assemblages of Gondwana show major changes that correspond to its great size and wide range of climates and environments. The Antarctic and Australian sectors of Gondwana rested in low latitudes during the Cambrian and have extensive carbonate deposits, although those of Antarctica are poorly exposed through the present-day polar ice cap. Differences in their fossil assemblages in addition to paleomagnetic evidence suggest that present-day North and South China were on separate tectonic plates; however, extensive carbonate deposits in both regions indicate that both plates were found in low latitudes during the Cambrian. The fossil assemblages of South China have strong similarities with those of both Australia and Kazakhstan, but details of the Cambrian geographic relationships remain unclear.

Several terranes (fault-bounded fragments of the Earth’s crust) seem to have been located near or attached to the margin of the northern Africa sector of Gondwana in the high southern latitudes, but many details of their Cambrian geographic relations are unknown. These terranes now make up much of southern Europe and parts of eastern North America. Cambrian deposits in all the terranes are chiefly sandstone and shale and include few or no carbonates. Their faunas closely resemble those of Baltica at generic and higher taxonomic levels, but differences at the species level suggest some geographic separation.

Siberia was a separate continent located in the low latitudes between Laurentia and Gondwana. Fossil similarities between this continent and Gondwana suggest that it was positioned relatively close to the equatorial region of Gondwana. Present-day Kazakhstan seems to be composed of several microcontinental blocks that were in all likelihood separated during the Cambrian. These blocks became amalgamated after the Cambrian, and the subsequent composite continent known as Kazakhstania collided with Siberia during the late Paleozoic Era.

Few Cambrian faunas from continental-slope and deep-ocean environments are known. Limited information from these is important, however, for demonstrating affinities between deep, cool-water ecosystems at all latitudes and shallow, cool-water communities of higher latitudes. There exists a close similarity between the observed distribution patterns of Cambrian and modern marine arthropods. This has been used as persuasive evidence for the existence of thermally stratified Cambrian oceans in lower latitudes with a thermocline separating warm-water from cool-water layers. This inferred thermocline and the wide oceanic separation of biogeographic regions are seen as the likely causes for the high endemism of the biological communities surrounding Laurentia. This interpretation is supported by deposits that date back to the middle of the Cambrian in northern Greenland where, within a few tens of kilometres, normal Laurentian shelf-margin trilobite (distinctive three-lobed marine arthropods) communities grade into deepwater faunas like those in the shallow-shelf deposits of Baltica. Similarly, trilobite species in later Cambrian deepwater faunas found in the western United States also appear in southeastern China, whereas shallow-water communities of the two regions have few genera in common.

As large tectonic plates continued to move during the Phanerozoic Eon, terranes of various sizes were displaced. Endemic Cambrian fossils, in conjunction with other geologic evidence such as physical stratigraphy, have been useful for identifying the geographic origins of some terranes, particularly those that have undergone substantial displacement. Examples of displaced terranes include northern Scotland and west-central Argentina (Precordillera) with Laurentian fossils, eastern Newfoundland with fossils from Baltica, and southern Mexico (Oaxaca) with Gondwanan (South American) fossils.

Another important consequence of continued plate movement has been the formation of large mountain ranges at the sites of plate collision. Pressure and heat generated during collisions since the Cambrian have folded, faulted, and metamorphosed significant volumes of Cambrian rock, especially from the outer margins and slopes of many continental shelves. No crustal rocks found in today’s oceans are older than the Mesozoic Era (252.2 million to 66 million years ago); most pre-Mesozoic deposits that accumulated in the deep ocean basins are thought to have been destroyed after they subducted into Earth’s interior.

Relatively abrupt changes in sea level may have significantly influenced Cambrian environments and life. A global drop in sea level is suggested by extensive unconformities (interruptions in the deposition of sedimentary rock). The time represented by such unconformities in sectors of Laurentia and Baltica bounding the Iapetus Ocean has been called the Hawke Bay event. An apparent absence of a coeval unconformity in western North America seems to be an anomaly. Thick uninterrupted shelf deposits in this sector of Laurentia, however, may have resulted from abnormal shelf subsidence caused by the cooling of crustal rocks following a late Precambrian plate-rifting event. The timing of unconformities found in widely separated continents cannot be correlated with precision; however, it is perhaps significant that a number of characteristic animal groups from the early portions of the Cambrian were either exterminated or severely restricted in their geographic distribution at about the same time in the world’s shallow-shelf environments. Among biostratigraphically important trilobites, the olenellids were exterminated near Laurentia, the holmiids went extinct at the margins of Baltica, and the redlichiids vanished from the shallow-shelf ecosystems near Gondwana. Also, diverse and abundant reef-dwelling archaeocyathans (extinct group of sponges thought to have helped construct the first reefs) disappeared from most low-latitude warm-water continental shelves.

The significant rise in sea level is suggested by rather abrupt and extensive displacements in sedimentary environments and biotas during the middle of the Cambrian (the Ptychagnostus gibbus zone). Lowland areas were flooded, as in parts of Baltica. In warm-water shelf sections of the world, coarse-grained, shallow-water, carbonate rocks were abruptly overlain by fine-grained, deeper-water, laminated limestone or shale. Adaptive radiation of the pelagic agnostoid trilobites was greatly accelerated in open-oceanic environments following this event, perhaps in response to newly expanded habitats.

In Laurentia, a significant drop in sea level near the end of the Cambrian is suggested by both the disappearance of some faunas and a single unconformity that defines the boundary between the Dresbachian and Franconian stages, two regional stages in peripheral areas of North America. However, there is no evidence for a comparably large change in sea level at or near the Cambrian-Ordovician boundary. Associated minor unconformities have made it difficult to select a defining Global Standard Section and Point (GSSP) for the boundary, which ideally should be located in an uninterrupted stratigraphic section.

Several regions of Cambrian volcanism have been identified. Australia was especially active; large areas in the northern and central regions were covered by flood basalts during the early parts of the Cambrian with residual activity extending into the middle of the period. Basalts and mafic intrusives (molten rock derived from basic magmas) in southeastern Australia formed in a volcanic island arc setting during the early and middle parts of the Cambrian. Volcanic suites of similar age are also present in New Zealand and in parts of Antarctica (northern Victoria Land, Ellsworth Mountains, and Pensacola Mountains). Other significant Cambrian volcanic deposits occurring during the early and middle parts of the period are found in southern Siberia and western Mongolia (Altai and Sayan mountains), eastern Kazakhstan and northwestern China (Tian Shan), and northeastern China. Cambrian volcanics are scattered along the easternmost margin of the United States, but most are probably island arc deposits that were accreted to Laurentia after the Cambrian. In the southern United States (Oklahoma), granitic intrusives and basaltic and rhyolitic extrusives are associated with a large tectonic trough that was formed by rifting, or crustal extension, throughout the first half of the period.

Minor volcanic deposits, mainly ash beds and thin flows, are widely known. In general, these have received little study, but some are suitable for determination of isotopic ages. Zircons from a lower Lower Cambrian (pretrilobite) volcanic ash bed in New Brunswick, Can., have a uranium-lead age of 531 million years. Volcanic tuffs near inferred Tommotian-Atdabanian boundaries (Russian designations for the pretrilobite and trilobite portions of the Cambrian explosion) in both Morocco and southwestern China have yielded similar dates of 521 million years.

The tectonic history of the Precambrian is not as well-known as that of the Paleozoic. In general, however, late Precambrian history seems to have been characterized by continental fragmentation, whereas Paleozoic history was characterized by the continental accretion of terranes. The Cambrian was a period of transition between those tectonic modes, and continents were scattered, apparently by fragmentation of Rodinia. Major Cambrian and early Ordovician tectonism affected large areas of Gondwana in what are now Australia, Antarctica, and Argentina. Multiphase tectonism in Antarctica is called the Ross Orogeny, and in Australia it is known as the Delamerian Orogeny. At least some of the volcanic activity noted above, particularly that of volcanic island arcs, is evidence that seafloor spreading and crustal subduction were active geologic processes.

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Paleoclimate

Global climate during Cambrian time was probably warmer and more equable than today. An absence of either land or landlocked seas at the Cambrian poles may have prevented the accumulation of polar ice caps. The general absence of glacial till deposits of Cambrian age is more notable, because these deposits are common and widespread in upper Precambrian strata. They accumulated again during the Ordovician Period in northern Africa as Gondwana began to move over the South Pole. Otherwise, the presence of persistent and widespread limestone deposits found on the margins of a centralized transcontinental arc in North America, for example, indicates that a subtropical climate existed in latitudes between 30° N and 30° S. In addition, arid to semiarid conditions at latitudes around the Tropics of Cancer and Capricorn (approximately 23°27′ N and S latitude, respectively) are suggested by deposits that include sandstone with quartz grains frosted by abrasion through wind transport, ventifacts (wind-polished stones), and evaporites.

More sophisticated research on paleoclimates relies on the detection of changing patterns in the amounts of isotopic oxygen, carbon, and strontium retained in limestone samples to correlate the timing of different geological events. Much remains to be accomplished regarding this promising line of research on Cambrian strata, but certain trends and events are becoming better defined. Strong reductions in isotopic carbon (13C), for example, are correlated from Lower Cambrian strata at localities as distant from one another as the Lena River area of Siberia and the Atlas Mountains of Morocco. Another substantial drop in 13C is believed to mark the transition between the early and middle parts of the Cambrian in the Great Basin of North America. Such decreases may represent a global rise in temperature occurring at the same time as a rise in global sea level. In addition, the delivery of anoxic (oxygen-depleted) ocean waters with reduced amounts of 13C also may have been aided by rising sea level.

Cambrian life

The long history of life on Earth has been punctuated by relatively abrupt changes. Some have argued that the greatest change of all occurred in marine environments near the Precambrian-Cambrian boundary. Fossils from Cambrian rocks include the oldest representatives of most animal phyla having mineralized shells or skeletons. A lack of observed connecting links suggests that processes of biomineralization (specifically, the formation of bones, shells, and teeth) evolved independently in several phyla. Whether or not soft-bodied representatives of some of these phyla originated during the Precambrian but have no preserved record is a debated question. Nevertheless, the hard parts of Cambrian animals had a much greater potential for preservation than the soft parts, and they mark the beginning of a diverse fossil record.

Fossil record of the Precambrian-Cambrian transition

The preservation of the record of the Precambrian-Cambrian transition was significantly affected by global changes in sea level. During latest Precambrian time, the sea level was relatively low, resulting in spatially restricted oceans and expanded continents. Throughout much of the Cambrian, rising seas gradually flooded vast land areas. Sediment was eroded from the continents and deposited in adjacent seas. Because of low sea level, the sedimentary and fossil records of the Precambrian-Cambrian transition are generally most complete toward the outer margins of continental shelves. As a corollary, the time gaps, represented by the boundary surface, generally increase in landward directions. This has led to an absence or serious incompleteness of the transitional record in most areas, particularly in those of classical Cambrian studies. As a result, it is thought that this incompleteness, combined with a general deficiency in knowledge—prior to the mid-1900s—of Precambrian communities, contributed significantly to the long-held notion of an abrupt or sudden appearance of Cambrian fossils.

Considering the biological importance of the Precambrian-Cambrian transition, it is somewhat surprising that the primary impetus for its detailed study came from a project designed to establish a suitable international boundary stratotype (a rock layer that serves as a benchmark of geologic time). Before the project was initiated in 1972, reasonably complete stratigraphic sections across the transition were either largely unrecognized or ignored. Since 1972, information about the transition has accumulated at an accelerating rate. Although many details remain to be learned, the general history of this momentous interval is becoming clear.

The Precambrian-Cambrian biotic transition, once thought to be sudden or abrupt, has been found to include a succession of events spread over many millions of years. It commenced with the appearance of the animal kingdom (i.e., multicelled organisms that ingest food), but the date and details of that event remain obscure. At least three informal phases in the transition can be identified by progressively more diverse and complex biological communities.

The earliest phase of late Precambrian age is characterized by fossils of soft-bodied animals known from many localities around the world. Based on fossils of animal embryos, it is thought that elements of the Ediacaran fauna appeared more than 600 million years ago. The fossils are predominantly the imprints of soft-bodied animals. Their extraordinary preservation, usually in sandstone or shale, was probably the result of rapid burial and protection by smothering sediment. Most of the fossils are relatively simple, and many resemble worms, sea pens, and jellyfish. Dwelling traces (fossilized burrows and other excavations) like those of modern sea anemones are also common. Placing Ediacaran fauna in higher (more generalized) taxonomic levels is controversial, however, because critical diagnostic features are not evident. Some paleontologists have assigned Ediacaran body fossils to the extant phyla Annelida, Coelenterata, and Arthropoda, whereas others have regarded them as members of extinct taxonomic groups of high rank. Some adherents of this latter viewpoint have suggested that the Ediacaran fauna was terminated by a major extinction event, but direct evidence of an abrupt replacement of species has not been found in the geologic record.

Other kinds of fossils also provide valuable clues about life during Ediacaran time. Photosynthetic organisms include unicellular blue-green algae (cyanobacteria) and acritarchs (probable algae), both of low diversity. Individuals of some species were probably abundant, however, and may have been an important source of food for Ediacaran animals. Hard parts of animals, primarily known from Africa and China, are mainly dwelling tubes composed of calcium carbonate and other compounds. Most were probably secreted by sessile, filter-feeding, wormlike animals. Although rare and of low diversity, these forms are significant because they signal the advent of biomineralization. The oldest unequivocal trace fossils, mainly crawling trails, are also of Ediacaran age. The trails suggest that locomotion of the trace makers was accomplished by waves of muscular contraction, like that in annelids and sea slugs, and not by legs. All but the latest Ediacaran trace fossils are relatively simple, suggesting limited and primitive behaviour patterns. Their low diversity further suggests that few kinds of mobile animals lived on the Ediacaran seafloor.

The second phase of the Precambrian-Cambrian biotic transition is characterized by a marked increase in the diversity of its shelly fauna and a lack of trilobites. It is near the lowest stratigraphic occurrence of this fauna that the Precambrian-Cambrian boundary stratotype has been placed. The fauna includes that of the Tommotian Stage, as applied in Russia, and it has often been referred to as the Tommotian fauna. It is known from many localities around the world, but time correlations lack precision. A general acceleration in biotic diversity during this second phase is the beginning of the Cambrian explosion.

Fossils of the second phase, which may be locally abundant, represent several new animal groups of Paleozoic aspect. Calcified archaeocyathans (extinct group of sponges thought to have helped construct the first reefs) diversified rapidly and were the first skeletal metazoans (multicellular animals with differentiated tissues) to develop a modular growth habit. They also evolved a complex symbiotic relationship with reef-building blue-green algae. Mollusks, preserved in both shale and limestone, include at least four classes (Monoplacophora, Gastropoda, Hyolitha, and Rostroconchia). Brachiopods (lamp shells) made their appearance but are low in diversity. Several groups are represented by an astonishing array of small mineralized tubes, scales, and spicules. The presence of arthropods, the first animals to develop legs, is indicated by characteristic trace fossils. The skeletal remains of arthropods are not preserved in the fauna, however, presumably because they were not mineralized. Other trace fossils show a marked increase in abundance and diversity as well as an expansion of behaviour patterns that reflect improvements in locomotion, greater ability to penetrate sediment, and new foraging strategies.

The third phase of the Precambrian-Cambrian biotic transition commenced with the appearance of mineralized trilobite skeletons. The subsequent adaptive radiation of the trilobites was exceptional, and their remains dominate most later Cambrian deposits. For this reason, the Cambrian Period has sometimes been called the Age of Trilobites.

The known Cambrian biota was restricted to marine environments. At least 11 extant animal phyla (Annelida, Arthropoda, Brachiopoda, Chordata, Ctenophora, Echinodermata, Hemichordata, Mollusca, Onychophora [velvet worms], Porifera, and Priapulida), including most of those with a fossil record, first appear in Cambrian rocks. Most of these rapidly diversified as they seemingly adapted to numerous unfilled ecological niches. Another five phyla (Nemertea, Phoronida, Platyhelminthes, Pogonophora, and Sipuncula) may also trace their origin back to the Cambrian fossils, though questions still remain about them. The only extant animal phylum with a good fossil record that is not known from Cambrian rocks are the Bryozoa (moss animals), which first appear in rocks of Early Ordovician age. A summary of the principal biotic groups of the Cambrian is given below.

Photosynthetic organisms

Cambrian photosynthetic organisms, the primary food of animals, are entirely unicellular. These organisms include a variety of prokaryotic (nonnucleated) and eukaryotic (nucleated) bacteria and algae. Their evolution, like that in associated animals, shows a marked acceleration in adaptive radiation and biomineralization near the base of the Cambrian. A new calcareous bottom-dwelling flora dominated by blue-green algae appeared. Some of these organisms formed mounds on the seafloor. Others formed small, concentrically laminated, marble- or biscuit-shaped structures called oncoids, which were locally abundant. Although it was rarely preserved, there existed a noncalcareous benthic flora that also was dominated by blue-green algae. By at least the middle of the Cambrian, some noncalcareous green algae (Chlorophyta) had become common. In North America and Siberia, the axes of one species, Margaretia dorus, exceeded 2 cm (0.8 inch) in diameter and were probably more than 1 metre (3.3 feet) in height. Such large size is attained by modern green algae only in warm, equatorial oceans. The phytoplankton, consisting of acritarchs and blue-green algae, also diversified near the base of the Cambrian. Acritarchs are widespread in many kinds of marine rocks and seem to have potential for an improved zonation of Lower Cambrian rocks. They are difficult to study, however, because of their microscopic size.