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Figure 4.4 Schematic size-frequency histograms: (a) ideal (positively) skewed, common of numerous invertebrate populaces with high infant mortality; (b) regular (Gaussian) distribution, typical of steady-state or transported assemblages; (c) left (negatively) skewed, common of high senile mortality; (d) multimodal distribution, typical of populaces with seasonal spawning patterns; and (e) multimodal distribution, v decreasing amplitude, usual of populations farming by molting (ecdysis).

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Size of individuals (age units) size of individuals (age units)

Figure 4.4 Schematic size-frequency histograms: (a) ideal (positively) skewed, typical of numerous invertebrate populaces with high infant mortality; (b) typical (Gaussian) distribution, typical of steady-state or transported assemblages; (c) left (negatively) skewed, typical of high senile mortality; (d) multimodal distribution, typical of populaces with seasonal spawning patterns; and (e) multimodal distribution, with decreasing amplitude, typical of populations farming by molting (ecdysis).

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Size of people (age units)

Figure 4.5 Schematic survivorship curves: form I tracks, increasing mortality through age; form II, consistent mortality through age; form III, diminish mortality v age.

Size of individuals (age units)

Figure 4.5 Schematic survivorship curves: form I tracks, enhancing mortality with age; form II, continuous mortality through age; form III, diminish mortality through age.

where K is the carrying capacity of the population or top limit of populace size, N is the actual populace size, r is the intrinsic price of population increase and also t is the unit that time.

Thus, when N philosophies K the price of population growth slows appropriate down and the population will approach a stable equilibrium. Such populations are common of an ext stable environments dominated by equilibrium species (K strategists). By contrast opportunistic types thrive in an ext adverse, unstable environments, where high expansion rates are usual (r strategists).

Habitats and also niches: addresses and occupations

All contemporary and fossil organisms deserve to be classified in regards to their habitat, where they live

Box 4.1 The terebratulide brachiopod Dielasma indigenous the Permian that the Tunstall Hills

The smooth terebratulide brachiopod Dielasma is typical in the limestones and dolomites linked with the Permian reefs the the Sunderland area in northeast England. Is it possible to usage data from simple length dimensions of the brachiopod shell to recognize the expansion strategies of these animals? One sample mirrors a bimodal pattern arguing two successive cohorts are present in the population; as whole the survivorship curve argues increasing mortality v age, in maybe a stable, equilibrium atmosphere (Fig. 4.6). But this was no the just environment around these Permian reefs; various other samples show different-shaped curves, part demonstrating high infant mortality in possibly much less stable environments, conversely, a populace with a bell-shaped curve suggests that the shells have been transported and sorted prior to burial. A choice of datasets is accessible by adhering to this link, http://www.blackwellpublishing.com/paleobiology/.

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Figure 4.6 Size-frequency histogram (a), polygon (b), cumulative frequency polygon (c) and survivorship curve (d) for a sample of 102 conjoined valves that Dielasma from the Permian reef basic deposit that the Tunstall Hills, Sunderland. (From Hammer & Harper 2005.)

(their address) or with recommendation to their niche, their way of life (their occupation). Modern-day organisms accounting a variety of atmospheres from the peak of mount Everest in ~ heights of nearly 9 km to depth of over 10 km in the Marianas Trench in the Pacific Ocean. Recognition of extremophiles (see p. 205), living in even an ext bizarre habitats, has significantly extended our knowledge of the environmental selection of life on Earth. A huge number the physical, chemical and biological components may characterize an organism"s environment; unfortunately, few can be recognized in the fossil record.

Some that the many abundant and also diverse areas inhabit the littoral zone, whereby rocky shores organize some that the most varied and also extensively learned faunas. Because that example, almost 2000 separation, personal, instance organisms have actually been recorded from a 250 mm2 quadrat ~ above an

Approximate

Approximate

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Figure 4.7 review of contemporary marine environments and their depth ranges, together with the approximate positions of the key benthic zones. (Based on Ager 1963.)

exposed wave-battered platform about the Scottish island the Oronsay. Unfortunately, few rocky coastlines have been recorded from the geology record; where they occur, often linked with paleo-islands, there are exciting and also unusual biotas and sediments (Johnson & Baarli 1999).

The bulk of fossil pets have been uncovered in maritime sediments, occupying a wide variety of depths and conditions. The distribution of the maritime benthos is managed principally by depth of water, oxygenation and temperature. The main depth zones and also pelagic atmospheres are depicted on Fig. 4.7. In addition, the photic ar is the depth the water penetrated by light; this have the right to vary according to water purity and also salinity but in optimum conditions it can expand down to about 100 m. Terrestrial atmospheres are largely governed through humidity and temperature, and also organisms inhabit a wide selection of continent environments, varying from the Arctic tundras come the lush woodlands of the tropics.

Marine environments host a selection of lifestyles (Fig. 4.8). The upper surface waters space rich in floating plankton, and also nektonic biology swim at miscellaneous levels in the water column. In ~ the benthos - the beasts the live in or ~ above the seabed - mobile nektoben-thos scuttle throughout the seafloor and the solved or sessile benthos are fixed by a variety of structures. Infaunal biology live beneath the sediment-water interface, when epifauna live above it.

Members of most areas are involved in some type of competition because that food, irradiate and space resources. For example, the stratification that tropical rain forests reflects vain in the top canopy because that light, if vegetation adapted for damp, darker conditions is occurred at lower levels. Similar stratification or tiering is a function of most maritime communities, becoming greater and more sophisticated v geological time (Fig. 4.9), rather choose the high-rise buildings in Manhattan seeking to optimize space on a densely inhabited island. Low-level tiers were typically occupied by brachiopods and corals throughout the Paleozoic, if the greater tiers were populated by crinoids. The Mesozoic and also Cenozoic faunas, however, are an ext mollus-kan-based through the lower tiers inhabited by epifaunal bivalves and brachiopods and also the top tiers populated by bryozoans and also crinoids.

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Figure 4.8 an option of naval lifestyles above, in ~ the surface, within and also at the basic of the water column. (Based top top Ager 1963.)

Seaweeds (brown, eco-friendly algae)

Seaweeds (brown, environment-friendly algae)

25 cm

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Giant bivalves, corals, sponges, " giant brachiopods

Most brachiopods, " bivalves, bryozoans

Figure 4.9 Epifaunal tiering of naval benthic communities; infaunal tiering recorded in map fossil assemblages is debated on p. 205. (From Copper 1988.)

25 cm

Giant bivalves, corals, sponges, " large brachiopods

Most brachiopods, " bivalves, bryozoans

Figure 4.9 Epifaunal tiering of naval benthic communities; infaunal tiering recorded in trace fossil assemblages is debated on p. 205. (From Copper 1988.)

Trace fossil associations display that burrows may be organized in an infaunal, tiered power structure (see chapter 19). Ausich and also Bottjer (1982) identified three levels with enhancing depth indigenous the sediment-water interface: 0 come -60 mm, -60 come -120 mm and also -120 mm to -1 m. During the faster Paleozoic, just the first tier was repetitively occupied, the second tier was lived in from the so late Silurian and, finally, the third tier was populated in the Carboniferous. Tiering was also selectively impacted by die out events, and also tiers deeper 보다 500 mm space rare ~ the so late Cretaceous since of predation by bony fishes.

Trophic structures: bottom or peak of the food chain?

Food pyramids form the basis of most ecological systems, defining the energy flow v a chain of different organisms from very abundant primary producers to relatively few predators. A number of basic trophic or feeding methods are recognized (Fig. 4.10). Several marine food chains (basically, that eats what) have actually been documented including those dominated by suspension feeders such as brachiopods, bryozoans and sponges. This fed mainly on phytoplankton and also other organic detritus. Suspension feeding was particularly common in Paleozoic benthos; the Mesozoic and Cenozoic faunas were an ext dominated by detritus feeders, such together echi-noids, and also food chains were normally longer and also more facility (Fig. 4.11).

It could seem fairly easy at an initial sight to reconstruct a food chain for a fossil assemblage, offering you deserve to work out who ate what. However that is much easier said 보다 done. One of the many spectacular fossil lake deposits, conquered by amphibians, has actually been documented from the top Carboniferous of

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Figure 4.10 Trophic groups, task of members and their life sites. 1, primary producers: phytoplankton in surface waters v (a) cyanobacteria and (b) benthic algae. 2, Herbivores: browsing and also grazing gastropods. 3, Deposit feeders: (a) deposit-feeding gastropod and (b) shallow infaunal bivalve. 4, Suspension feeders: (a) semi-infaunal, byssally-attached bivalve, (b) shallow infaunal bivalve, (c) crinoid, (d) epifaunal bivalve, and (e) deep infaunal bivalve. 5, Carnivores: (a) nektonic fishes, (b) nekton-benthic fishes, (c) epifaunal gastropod, and (d) infaunal gastropod. (From Brenchley & Harper 1998.)

Figure 4.10 Trophic groups, activity of members and their life sites. 1, major producers: phytoplankton in surface ar waters with (a) cyanobacteria and (b) benthic algae. 2, Herbivores: browsing and also grazing gastropods. 3, Deposit feeders: (a) deposit-feeding gastropod and also (b) shallow infaunal bivalve. 4, Suspension feeders: (a) semi-infaunal, byssally-attached bivalve, (b) shallow infaunal bivalve, (c) crinoid, (d) epifaunal bivalve, and (e) deep infaunal bivalve. 5, Carnivores: (a) nektonic fishes, (b) nekton-benthic fishes, (c) epifaunal gastropod, and (d) infaunal gastropod. (From Brenchley & Harper 1998.)

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zoo / phytoplankton-suspension feeders -carnivores
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Figure 4.11 Reconstructions of two various food chain communities. (a) A neighborhood with a suspension-feeding food chain, displaying a range of suspension feeders, collecting food in various ways (bivalves with a mucous catch or setae, bryozoans and brachiopods with lophophores, foraminiferans v cilia, corals through tentacles, and also sponges with flagellae). (b) A neighborhood with a detritus-feeding food chain conquered by various species of bottom-dwelling deposit feeders and nektonic carnivores represented by a cephalopod and also placoderm. (From Copper 1988.)

Figure 4.11 Reconstructions of two various food chain communities. (a) A ar with a suspension-feeding food chain, displaying a range of suspension feeders, collecting food in different ways (bivalves v a mucous trap or setae, bryozoans and brachiopods with lophophores, foraminiferans through cilia, corals with tentacles, and also sponges through flagellae). (b) A community with a detritus-feeding food chain conquered by various types of bottom-dwelling deposit feeders and nektonic carnivores stood for by a cephalopod and also placoderm. (From Copper 1988.)

lake margin upland lake margin upland

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Figure 4.12 (a) Trophic structures in and also around a so late Carboniferous lake complex, Nyrany, Czechoslavakia. (b) Trophic structures in a late Permian reef complex, northeast England. (a, based on Benton 1990; b, native Hollingworth & Pettigrew 1998.)

Figure 4.12 (a) Trophic structures in and also around a so late Carboniferous lake complex, Nyrany, Czechoslavakia. (b) Trophic structures in a late Permian reef complex, northeast England. (a, based upon Benton 1990; b, from Hollingworth & Pettigrew 1998.)

Czechoslovakia (Fig. 4.12a). The lake ecosystem recreated because that the citizens of the Nyrany Lake facility has three main ecological communities: an open up water and lake association, dominated by fishes in addition to various larger amphibians; a shallow water and also swamp/lake association through amphibians, small fishes, land plants and also other tree debris; and also finally a terrestrial-marginal association with microsaur (small, primitive) amphibians and also primitive reptiles. Food chains have been worked out for each of these associations by mindful study the the teeth (was that a herbivore with grinding teeth or a carnivore v slashing teeth?) of each beast, and also comparisons with contemporary relatives. Because that example, in the open-water environments fishes, such together the spiny acanthodians, fed top top plankton however were themselves attacked by the amphibians, presumably in ~ the height of the food chain. In the linked terrestrial environments, plant material was consumed by a range of invertebrates, consisting of insects, millipedes, spiders, snails and worms; these listed food and also nutrients for a range of little amphibians, themselves food for bigger amphibians and also reptiles.

A great example of a marine food web originates from the Zechstein Reef facies of northern Europe, date from the late Permian

(Fig. 4.12b). The Zechstein benthos was conquered by varied associations the brachiopods, overshadowed in the greater tiers by fan- and vase-shaped bryozoans (Hollingworth & Pet-tigrew 1988). Both groups were sessile filter feeders. Stalked echinoderms to be rarer and also occupied the highest possible tiers. Mollusks such together bivalves and also gastropods were important deposit feeders and also grazers. Among the largest predators was Janassa, a benthic ray, equipped v a formidable battery that teeth capable of crushing the shells the the sedentary benthos.

Megaguilds

Assignment of organisms to megaguilds gives another way to classify and understand the materials of a fossil community. Guilds are groups of functionally comparable organisms developing together in a community. Megagu-ilds are simply a selection of adaptive strategies based on a combination of life place (e.g. Shallow, active, infaunal burrower) and feeding type (e.g. Suspension feeder). Part paleontologists have used the term "guild" for these categories; however, these were most likely finer ecological divisions within the so-called Bambachian megaguilds, called after the American paleontologist Richard Bambach, who first used the ide (Bambach 1983). Megaguilds have additionally become an effective tool in assessing irreversible ecological adjust (see p. 105).

Controlling factors

The environmental niche of an biology is established by a huge selection of limiting factors, many of which are not tape-recorded in the rock document (Fig. 4.13). Crucial limiting factors for naval organisms room light, oxygen levels, temperature, salinity, depth and substrate (Pickerill & Brenchley 1991).

Light is the main energy source for primary producers, therefore diatoms, dinoflagellates, coc-coliths and cyanobacteria are dependent ~ above light and usually accounting the photic zone. Most biological productivity occurs in the peak 10-20 m of the water column. Virtually all eukaryotic bio organisms require oxygen for your metabolic processes, absorbing oxygen through diffusion, in the situation of small-bodied organisms, or with gills or lung in the situation of the bigger metazoans. There is a well-developed

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Figure 4.13 coastline to basin transect showing the relative prestige of different components on the distribution of organisms. (From Brenchley & Harper 1998.)

oxygen-depth profile in the world"s seas and oceans. Oxygen levels generally decrease under to 100-500 m, whereby the quantity of oxygen absorbed by organic matter exceeds primary oxygen production. Below in the oxygen minimum zone (OMZ), the lowest oxygen values are reached. The number of numerous organisms, such together corals, echinoderms, mollusks, polychaetes and sponges autumn off dramatically in the OMZ.

Levels that oxygen in marine atmospheres are essential in determining who resides where. Aerobic (normoxic) environments have >1.0 ml L-1 concentrations of oxygen, dys-aerobic (hypoxic) atmospheres have 0.11.0 ml L-1 and anaerobic (hypoxic-anoxic) have ostracodes and small benthic forams, vice versa, hypersaline assemblages room of really low diversity with just a couple of bivalves, gastropods and also ostracodes.

Depth is just one of the most often quoted controls ~ above the circulation of marine organisms (Fig. 4.14). Although the direct affects that depth are related to hydrostatic pressure, numerous other factors, both chemical and also physical, are related to depth; for example, in basic terms, the grain dimension of sediment and water temperature decreases with depth. Back hydrostatic press does not normally distort the shells and also soft organization of biology it deserve to dramatically influence organisms with pockets that gas in their bodies, such as fishes and nautiloids. Except the effects of hydrostatic pressure, depth can additionally control the solubility that calcium carbonate; cold water contains much more dissolved carbon dioxide (CO2) giving a means to corrode carbonate. At offered depths in the world"s oceans, lead carbonate material starts to dissolve at so-called compensations depths. Listed below the carbonate compensation depth (CCD) the dissolution of calcium lead carbonate exceeds supply and at about 4-5 km calcite is no preserved. The depth is shallower for arago-nite, v the aragonite compensation depth (ACD) put at 1-2 km. Both the CCD and also ACD vary through latitude, gift shallower at greater latitudes, and also both parameters have actually varied throughout geological time. Nevertheless, depth alone more than likely has tiny effect on biotic distribution, fairly the numerous depth-related determinants can be used to reconstruct the water depths of ancient marine communities.

Finally, the state of the substrate, rates of sedimentation and turbidity dramatically impact the distribution of benthic organisms (Brenchley & Pickerill 1993). Biology have complex ecological requirements, part preferring a particular grain size, a certain type of organic product or they even respond come chemical signals (chemotaxic). There room also complicated taphonomic feedback processes, wherein biogenic substrates such as covering pavements can form attachment web page for new communities. In general terms, within near-shore environments, over there is a wide correlation between community distribution and also grain size. Diversity often tends to be greatest in muddy sands, center in sandy muds, short in pure sands and virtually zero in soft muds. Moreover whether the sediments kind soupy muds, loose sands, firmgrounds or hardgrounds will influence faunal distributions.

Paleocommunities

Paleocommunities space recurrent teams of organisms concerned some specific set of environmental problems or limiting factors. Numerous of the concepts and also techniques used to marine fossil communities are based upon the work-related of biologist such together the Danish scientist Carl Petersen, researching in the late 1800s and also early 1900s. Petersen known a collection of level-bottom benthic communities approximately the Scandinavian coasts; the major control on neighborhood distribution was water depth, back other determinants such as the substrate were likewise influential.

Paleontologists were sluggish to embrace these insights from modern-day marine biology. There were a few pioneer studies on Carboniferous assemblages in the 1930s, however it to be the standard work by Alfred Ziegler in the 1960s the really brought these methods to the attention of paleontologists. He established five depth-related, brachiopod-dominated communities in the reduced Silurian rocks that Wales and also the Welsh borderlands (see thing 12). These neighborhoods stretched native the intertidal zone in the east to the deep shelf and also continental slope in the direction of the west, matching perfectly the ancient paleogeography. This totality system has been revamped and is now an ext widely well-known as the benthic assemblage zones (Fig. 4.15). These zones are characterized on a wide range of faunal and also sedimentological criteria and also may be subdivided, internally, on the basis of, for example, substrate type and the level of turbulence (Brett et al. 1993).

Describing fossil communities

Sometimes the simplest work are the hardest to do properly. For over a hundred years, paleontologists have provided lists of species from specific localities, yet these room not advantageous for eco-friendly work uneven the loved one abundances the the different species are recorded as well. We need to understand which varieties dominate (sometimes one types makes up more than 50% of the sample) and which are rare (i.e. Much less than 5% of the collection). Currently it is an ext common to paper the absolute and also relative variety of every organism, depicted graphically through frequency histograms, and based ~ above data obtained from heat transects, quadrats or more commonly currently from bed-by-bed collecting or bulk samples.

Counting conventions remain a problem. With many organisms the is fairly simple to calculation how plenty of individuals to be actually represented in a given assemblage: univalved types (e.g. Gastropods) count as one, whereas twin-valved species (e.g. Bivalves and also brachio-pods) may be assessed by adding the most common valve (right or left, dorsal or ventral) to the number of articulated or conjoined shells. Animals that molt, such together ostracods and trilobites, early american organisms and also those

Figure 4.15 Silurian maritime benthic assemblage zones and identifying criteria. (From Brenchley & Harper 1998.)

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Figure 4.15 Silurian marine benthic assemblage zones and identifying criteria. (From Brenchley & Harper 1998.)

that easily fragment (e.g. Bryozoans) and those v multi-element skeletons (e.g. Cri-noids and also vertebrates) need more devoted counting techniques. These straightforward data space then transformed into a more realistic snapshot of old communities populating previous landscapes and also seascapes, through histograms and pie charts. Raw number data are incredibly useful, but these can also be convert to diversity, dominance and evenness parameters, and also parameters because that taxonomic distinc-tiveness (Box 4.2). These with each other can give us a rich outline of the composition and structure that the paleocommunity and enable numerical comparisons with other similar assemblages. Such ideologies have come to be routine in researches of invertebrate paleoecology yet it is lot more an overwhelming to apply these methods to vertebrate assemblages where sample size are generally much smaller.

Detailed evaluation of paleocommunity structures has actually permitted acknowledgment of a variety of specific neighborhood types. Pioneer communities are those that have just entered brand-new ecospace, and they may be dominated by one or two an extremely abundant opportunistic species, in comparison to long-established and rather stable equilibrium neighborhoods where relatively high diversities of much more or less equally abundant animals are present. The eco-friendly relationships in between organisms is also an important aspect the community development (Box 4.3).

Paleocommunity advance through time

Communities undoubtedly adjust with time. Components such as eco-friendly fluctuation, immigration and emigration that animals and also plants, evolution and also extinction of types and coevolutionary transforms will alter the composition and also structure the a community. But are the components of communities tightly linked and also thus evolve together or is the a fairly haphazard random process? life communities, when very first established, show initial high rates of replacement and instability, whereas later stages are more stable with little change,

Box 4.2 eco-friendly statistics and sampling sufficiency: room you obtaining enough?

It is often complicated to evaluate the adequacy of a paleoecological sample. Part authorities have said that samples of about 300 give a fairly accurate census that a fossil assemblage. Commonly, investigators plot rarefaction curve (Fig. 4.16). This are developed simply by collecting samples that 10 and also identifying the number of species in each. Because that each sample the 10 plotted follow me the x-axis, the cumulative number of species is plotted along the y-axis. The curve might level off at the point where no additional varieties are figured out with additional collecting and also this fixes the sample dimension that is sufficient to count the majority of species present (Fig. 4.16).

A variety of statistics has actually been provided to describe aspects of fossil communities. Although the number of species accumulated from one assemblage gives a rough overview to the diversity that the association, obviously in most cases the bigger the sample, the higher the diversity. Diversity steps are normally standardized versus the sample size. Supremacy measures have actually high worths for communities with a few abundant elements and low values where types are more or less evenly represented; procedures of evenness are usually the train station of dominance.

Margalef diversity = S - 1/log N

where S is the number of species, N is the number of specimens, ni is the variety of the ith species, and also p, is the family member frequency of ,th species.

Many number techniques have been provided to analysis paleocommunities and their distributions. Phenetic methods (see chapter 2) are based on the investigation of a similarity or street matrix

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Figure 4.16 building of a rarefaction curve based upon data collected from a mid-Devonian brachiopod-dominated fauna, north France. The main species of brachiopod space illustrated: (a) Schizophoria, (b) Douvillina, (c) Productella, (d) Cyrtospirifer, (e) Rhipidiorhynchus, and also (f) Athyris. The curve levels turn off at around 300 specimens, suggesting this sample size is a enough census the the fauna. Magnification about x0.5 because that all.

Figure 4.16 building and construction of a rarefaction curve based upon data accumulated from a mid-Devonian brachiopod-dominated fauna, north France. The main species of brachiopod room illustrated: (a) Schizophoria, (b) Douvillina, (c) Productella, (d) Cyrtospirifer, (e) Rhipidiorhynchus, and (f) Athyris. The curve levels off at about 300 specimens, suggesting this sample size is a adequate census the the fauna. Magnification about x0.5 because that all.

Continued acquired from a raw data procession of the visibility or absence or number abundance the fossils at each site. Cluster analysis is most typically used in environmental studies and there is a wide variety of both distance and similarity measures, together with clustering techniques, to pick from. R-mode analysis clusters the variables, in most paleoecological researches the taxa, conversely, Q-mode analysis clusters the cases, generally the localities or assemblages (Fig. 4.17).

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For example, late Ordovician brachiopod-dominated assemblages from southern China have been investigated by swarm analyses (Hammer & Harper 2005) and fall into a number of ecogroups. This data are accessible at http://www.blackwellpublishing.com/paleobiology/.