Paleontology: Diversity and Extinction, Taphonomy, Functional Morphology, Morphometrics, Vertebrates and Paleoecology

This chapter answers the heading “Paleontology” of Part B1 of the Geology and Geophysics (GG) paper: “Diversity of life through time, mass extinctions- causes and effects; Taphonomy - processes of fossilization. functional morphology of invertebrates (bivalves, brachiopods, gastropods, echinoids, ammonites) and microfossils - (foraminifera, Ostracoda, conodonts, bryozoa); Statistical analysis of morphology; Vertebrate paleontology. Basic concepts of paleoecology; Fossils and paleoenvironments.” It teaches diversity and its indices, the Big Five extinctions, taphonomy, the functional reading of nine fossil groups, allometry, vertebrate transitions and paleoenvironments. The numericals are the Shannon index, rarefaction, per-capita extinction rate, allometric prediction and a confidence interval.

1. Diversity of life through time and how it is measured

The fossil record reads as a series of radiations and collapses. Soft-bodied Ediacaran organisms (about 570 to 539 Ma) give way to the Cambrian explosion of skeletonised animals. Marine diversity then climbs steeply in the Great Ordovician Biodiversification Event, holds a long Paleozoic plateau, is cut hard at the end of the Permian, and rises through the Mesozoic and Cenozoic to its highest level. Sepkoski’s three evolutionary faunas summarise the pattern: the Cambrian fauna (trilobites, inarticulate brachiopods), the Paleozoic fauna (articulate brachiopods, crinoids, rugose and tabulate corals, nautiloids) and the Modern fauna (bivalves, gastropods, echinoids, bony fishes, cheilostome bryozoans), each rising to dominance largely because mass extinctions removed the one before.

Raw counts of taxa depend on sample size, so richness must be compared after rarefaction: the expected number of species in a random subsample of n individuals out of N is E(S_n) = Σ [1 − C(N − N_i, n) / C(N, n)], where N_i is the count of species i. Indices that use abundance are Simpson’s D = Σ p_i² (the chance that two individuals drawn at random belong to the same species; 1 − D rises with diversity) and Shannon’s H = −Σ p_i ln p_i, with p_i = N_i / N. For S species H peaks at ln S when all are equally abundant; evenness is J = H / ln S. Example: a bed of 60 shells holding 30, 20 and 10 individuals of three species has p = 0.50, 0.33, 0.17, so H = 1.01 against a maximum ln 3 = 1.10, and J = 0.92.

Fossil-record diversity is counted from stratigraphic ranges. The range-through method counts a taxon in every interval between its first and last appearance, which repairs sampling gaps; the better sampling of young rocks still inflates recent diversity (the pull of the Recent). Percentage extinction (extinct ÷ standing diversity × 100) is not a rate. A per-capita rate that lets intervals of unequal length be compared is q = −ln(N_bt / N_b) / Δt, where N_b is the number of taxa crossing the bottom boundary, N_bt the number crossing both boundaries and Δt the interval length in Myr; taxa confined to the interval are not counted.

2. Mass extinctions: causes and effects

A mass extinction is a geologically brief interval (usually under a few Myr) in which a large share of species worldwide disappears, far above the background rate, across unrelated groups and habitats.

The Big Five; the percentages are estimates, but the end-Permian is the largest
EventAge (Ma, approx.)Marine species lost (approx.; estimates vary)Main proposed causes
End-Ordovician445about 85%Gondwanan glaciation with sea-level fall, then warming and anoxia
Late Devonian372about 75%Repeated anoxic pulses, climate change, collapse of reef ecosystems
End-Permian252about 90% or moreSiberian Traps volcanism, warming, ocean anoxia and acidification
End-Triassic201about 75 to 80%Central Atlantic Magmatic Province volcanism, CO₂ warming, acidification
End-Cretaceous66about 75%Chicxulub impact, with Deccan Traps volcanism (relative roles debated)

At the end of the Cretaceous the evidence for impact is an iridium-enriched clay layer, shocked quartz, spherules and the Chicxulub crater (Yucatán, about 180 km across). Volcanism acts through CO₂ (long-term warming and acidification), SO₂ aerosols (short-term cooling) and warm, stagnant, oxygen-poor oceans. The end-Permian removed the trilobites and rugose and tabulate corals, so the Paleozoic fauna never regained dominance.

Effects. Extinction is selective: geographically restricted, narrowly adapted, tropical reef-building and large-bodied taxa suffer most. Survivors radiate into the emptied ecospace (mammals after the end-Cretaceous, archosaurs after the end-Permian), but marine recovery takes several million years, and disaster taxa (the bivalve Claraia and the therapsid Lystrosaurus after the end-Permian) briefly dominate. A Lazarus taxon vanishes from the record and reappears later. Because a genus survives if any one of its species does, the percentage of species lost normally exceeds the percentage of genera lost in the same event.

⚠️ Signor–Lipps effect: a sudden event can look gradual
The last fossil of a taxon is almost never from the moment it died, so ranges appear to end below the true extinction horizon. With sparse sampling a single sudden event therefore looks like a gradual decline spread over several metres of rock.

3. Taphonomy: processes of fossilisation

Taphonomy studies everything that happens to remains between death and discovery. Biostratinomy covers death to final burial (scavenging, decay, disarticulation, transport, abrasion, bioerosion); fossil diagenesis covers what follows burial (compaction, cementation, dissolution, recrystallisation, replacement). Preservation needs durable hard parts, rapid burial and often oxygen-poor bottom water that keeps out scavengers.

  • Permineralisation: pores fill with silica, calcite or pyrite while the original tissue stays (petrified wood, bone).
  • Replacement: the original substance dissolves and another mineral takes its place molecule by molecule (silicified or pyritised shells).
  • Recrystallisation or inversion: the composition is kept but the crystal form changes (aragonite to calcite), destroying fine structure.
  • Carbonisation (distillation): volatiles escape and a thin carbon film remains (graptolites, leaves, soft tissue).
  • Moulds and casts: a dissolved shell leaves an external mould (impression of the outside) and an internal mould or steinkern (the infilled interior); a void filled by cement gives a cast.
  • Unaltered remains: amber (insects), freezing (mammoths), desiccation, tar seeps.

Mineralogy decides what survives. Aragonite (many bivalves and gastropods, scleractinian corals, ammonite shells) dissolves or inverts easily, so these groups often survive only as moulds, whereas low-Mg calcite (articulate brachiopods, trilobites), high-Mg calcite (echinoderm plates), apatite (bone, conodonts, Lingula) and silica (radiolarians, sponge spicules) are more durable. An assemblage may be autochthonous (in life position), parautochthonous or allochthonous (transported). Time-averaging mixes generations spanning hundreds to thousands of years into one bed, so a death assemblage is a long-term average, not a snapshot.

Lagerstätten are deposits of exceptional fossil content. Konzentrat deposits concentrate hard parts (bone beds, shell beds); Konservat deposits preserve soft tissue: the Burgess Shale (Cambrian, British Columbia), Chengjiang (early Cambrian, China) and the Solnhofen Limestone (Late Jurassic, Germany). The record is therefore biased towards hard-shelled, shallow-marine, rapidly buried organisms, and diversity counts inherit this bias.

4. Functional morphology of invertebrates

Functional morphology reads a skeleton as an engineering solution. The method is to compare with living relatives, model the structure, and test the model against the sediment and associated fossils.

Standard functional readings of the syllabus groups
GroupSkeleton and symmetryFunctional features and what they tell
BivalvesTwo valves, mirror images of each other across the commissure plane; one valve alone is usually not symmetrical (inequilateral)A deep pallial sinus (notch in the pallial line where siphon muscles attach) means long siphons and a deep burrower; a byssal notch means anchoring by threads; one valve cemented down means an oyster-type cementer
BrachiopodsTwo unequal valves (pedicle and brachial); the symmetry plane cuts through both valves, and each valve is itself bilaterally symmetrical. Articulate forms have low-Mg calcite shells with teeth and sockets; inarticulate forms (Lingula) have phosphatic shells held by muscles onlyThe pedicle emerges through an opening in the pedicle valve to anchor the animal; internal brachidia (spiral in spiriferids) support the feeding lophophore; a wide hinge line stabilises the shell on soft mud
GastropodsOne univalve shell, usually coiled: trochospiral (helical) or planispiral. Torsion, a 180° twist of the body in the larva that brings the mantle cavity forward, is not the same as shell coilingCoiling is dextral (aperture on the right when the apex is up and the aperture faces the viewer) or sinistral; a siphonal canal at the aperture is typical of carnivorous snails, while a rounded entire aperture is common in grazers
EchinoidsCalcite test of plates arranged in five ambulacral (tube-foot pores) and five interambulacral areas; radial in regular urchins, secondarily bilateral in irregular onesRegular urchins live on the surface, with the mouth below, the anus (periproct) at the apex and a chewing Aristotle’s lantern. Irregular urchins (heart urchins, sand dollars) burrow: periproct shifted to the posterior, short dense spines, flat or heart-shaped test; heart urchins have lost the lantern
AmmonitesExternal chambered shell, usually planispiral; septa divide the chambers and a siphuncle runs along the outer (ventral) margin in ammonoidsThe suture, where a septum meets the shell wall, is typically goniatitic (simple zigzag) in the Devonian–Permian, ceratitic in the Triassic and ammonitic (finely frilled) in the Jurassic–Cretaceous. Compressed, streamlined, involute shells swim efficiently, open evolute shells less so. Rapid evolution makes them Mesozoic zone fossils
⚠️ Two different symmetry rules
In a bivalve the plane of symmetry lies between the valves, so the two valves are mirror images and each is normally asymmetrical. In a brachiopod the plane cuts through both valves, so each valve is symmetrical but the two valves differ. A bilaterally symmetrical single valve therefore points to a brachiopod, because most bivalve valves are inequilateral.

5. Microfossils: foraminifera, ostracoda, conodonts and bryozoa

Microfossils, mostly under a millimetre long, are recovered by sieving and acid residues; a small drill sample holds hundreds, so they are the working tools of subsurface biostratigraphy.

The four microfossil groups of the syllabus
GroupTest and rangeMain uses
ForaminiferaSingle-celled protists with a chambered test: agglutinated, porcellaneous or hyaline calcareous. Benthic forms from the Cambrian, planktic forms from the JurassicPlanktic species zone the Jurassic to Recent; benthic assemblages and the planktic : benthic ratio (higher offshore) show depth; Neogloboquadrina pachyderma coils mostly sinistral in cold water and dextral in warm; δ¹⁸O of calcite tests gives temperature and ice volume
OstracodaBivalved crustaceans with a low-Mg calcite carapace; valves unequal and hinged, with muscle scars; Ordovician to RecentOrnament, thickness and assemblage composition indicate salinity, temperature and depth; Mg/Ca and δ¹⁸O of the shell give water temperature
ConodontsTooth-like phosphatic (apatite) elements of an extinct eel-like chordate, forming apparatuses of several element types; late Cambrian to end-TriassicFast evolving, so the main zonation of Ordovician to Triassic marine rocks. The colour alteration index (CAI 1, pale, to CAI 5, black) records irreversibly the maximum heating, from about 50 to 80 °C (CAI 1) to above 300 °C (CAI 5)
BryozoaColonial animals; each zooid lives in a calcareous box or tube (zooecium) and feeds with a lophophore; Ordovician to Recent; mostly marineColony form tends to track water energy: encrusting sheets in turbulent water, erect branching or fan-like (fenestrate) colonies in calmer water; cheilostomes dominate Cenozoic shelves

6. Statistical analysis of morphology

Allometry describes growth of one part relative to another by Huxley’s power law y = a x^b, which is a straight line of slope b on a log–log plot: log y = log a + b log x. Isometry (shape unchanged with size) means b equals the value geometry demands: 1 for two lengths, 2 for an area against a length, 3 for a volume or mass against a length. A larger b is positive allometry, a smaller b negative. The slope is ontogenetic within one growth series, static among individuals of one age and evolutionary among species. Example: a shell with x = 10 and y = 4 and another with x = 40 and y = 32 give b = ln 8 / ln 4 = 1.5, so at x = 90 the predicted y is 4 × 9^1.5 = 108.

For n measurements the mean is x̄, the standard deviation s, the standard error SE = s / √n and the coefficient of variation CV = 100 s / x̄. A 95% confidence interval for the mean is x̄ ± t s / √n, with t from Student’s distribution at n − 1 degrees of freedom: for n = 16, x̄ = 42.0 mm, s = 6.0 mm and t = 2.131, the half-width is 2.131 × 6 / 4 = 3.20 mm. Quadrupling the sample only halves the interval. A bimodal size histogram suggests two cohorts, sexual dimorphism or two species, which must be tested, not assumed.

Shape is compared with multivariate methods: principal components (the first axis usually carries size), discriminant analysis, and landmark-based geometric morphometrics, which removes position, scale and rotation before shape is compared. Raup’s theoretical morphology describes a coiled shell by W (whorl expansion rate), D (distance of the generating curve from the axis) and T (translation along the axis, zero in planispiral shells).

⚠️ The slope is not the correlation
A correlation coefficient r measures scatter about the line, not the slope. Two allometric lines with the same b can have very different r. Growth is judged from b and its confidence interval, compared with the isometric value for those dimensions.

7. Vertebrate paleontology

The vertebrate story is a series of transitions, each recognised from a few key anatomical changes.

Major vertebrate transitions
TransitionKey evidence
Fish to tetrapods (Late Devonian, about 375 to 360 Ma)Lobe-finned fishes with limb-like fin bones; Tiktaalik (neck, wrist-like fin) then Acanthostega and Ichthyostega with digits
AmniotesThe amniotic egg frees reproduction from water (Carboniferous). Skull openings classify them: anapsid none, synapsid one lower temporal opening (therapsids, mammals), diapsid two (lizards, snakes, crocodiles, dinosaurs, birds)
Therapsids to mammals (Triassic)Cynodont ancestors; a single lower jaw bone (dentary) joined to the squamosal; the old jaw-joint bones (articular, quadrate) become the middle-ear malleus and incus; teeth differentiate into incisors, canines, premolars and molars
Dinosaurs and birdsSaurischians are lizard-hipped (theropods, sauropodomorphs); ornithischians are bird-hipped (pubis swung back beside the ischium). Birds descend from theropod saurischians; Archaeopteryx (Late Jurassic, Solnhofen) has feathers, teeth and a long bony tail. Pterosaurs, ichthyosaurs and plesiosaurs are not dinosaurs
WhalesEocene Pakicetus and related forms from the Indo-Pakistan region show the land-to-sea transition

Teeth and limbs read diet and movement: sharp shearing teeth in carnivores; flat, high-crowned (hypsodont) cheek teeth in grazers, which spread with Miocene grasslands. India has a rich record: the Late Triassic Maleri Formation (rhynchosaurs, phytosaurs), the Early Jurassic Kota Formation (the sauropod Barapasaurus), the Late Cretaceous Lameta Formation (titanosaurs, Rajasaurus, eggs) and the Siwalik Group (Miocene to Pleistocene mammals, the ape Sivapithecus).

8. Paleoecology, fossils and paleoenvironments

Paleoecology reconstructs how ancient organisms related to their surroundings. Its basic terms are life habit (benthic epifauna or infauna, nekton, plankton), feeding mode (suspension, deposit, grazing, carnivore), community (organisms that lived together) against assemblage (what was buried together, which may mix communities through transport and time-averaging). The method is uniformitarian: use nearest living relatives, functional morphology and the enclosing sediment, and prefer several independent indicators.

Fossil evidence and the environments it indicates
EvidenceWhat it indicates
Hermatypic (zooxanthellate) reef coralsWarm, clear, shallow water of normal salinity within the photic zone
Stenohaline groups (echinoderms, brachiopods, cephalopods, corals)Normal marine salinity; their absence, with a few abundant euryhaline species such as some bivalves, gastropods and ostracods, points to brackish or hypersaline water
Laminated black shale, no bioturbation, no benthic shellsOxygen-depleted bottom water
Mostly entire-margined leaves in a woody dicot floraWarm, typically tropical climate; more toothed leaves mean cooler
Trace fossils: Skolithos (vertical dwelling burrows), Cruziana (horizontal feeding and resting traces), Zoophycos, Nereites (patterned grazing traces)Ichnofacies: Skolithos, clean mobile sands of a high-energy shoreface; Cruziana, quieter lower shoreface to shelf; Zoophycos, quiet deeper muddy water; Nereites, deep-sea turbidite floors

Index fossils date rocks. They must be widespread, abundant, fast-evolving (short range), easy to identify and not restricted to one facies. Facies fossils are the reverse: long-ranging and tied to a habitat, so they show environment rather than age. A good paleoenvironmental indicator is therefore usually a poor index fossil, and the reverse.

Key takeaways

  • Diversity moves through three evolutionary faunas (Cambrian, Paleozoic, Modern). Compare richness only after rarefaction; Shannon H = −Σ p ln p peaks at ln S, Simpson D = Σ p²; an extinction rate is per capita, q = −ln(N_bt/N_b)/Δt, not a bare percentage.
  • The Big Five (about 445, 372, 252, 201 and 66 Ma) are the end-Ordovician, Late Devonian, end-Permian (largest), end-Triassic and end-Cretaceous. Extinction is selective, species loss normally exceeds genus loss, and the Signor–Lipps effect makes sudden events look gradual.
  • Taphonomy runs from biostratinomy (death to burial) to diagenesis. Aragonite shells dissolve and leave moulds, time-averaging blurs assemblages, and Konservat Lagerstätten (Burgess Shale, Chengjiang, Solnhofen) preserve soft tissue while Konzentrat deposits only concentrate hard parts.
  • Bivalve valves are mirror images across the commissure and each is normally asymmetrical; brachiopod valves are each symmetrical and unequal. A deep pallial sinus means a deep burrower, torsion is not coiling, and ammonoid sutures run goniatitic, ceratitic, ammonitic.
  • Allometry is y = a x^b, slope b on log–log axes, compared with the isometric 1, 2 or 3; r is not the slope. A mean’s 95% interval is x̄ ± t s/√n, so quadrupling n halves it.
  • Vertebrate history runs fish, tetrapod, amniote, therapsid, mammal, with birds nested in theropod (saurischian) dinosaurs, not the bird-hipped ornithischians. Index fossils are widespread, short-ranging and facies-free; facies fossils and ichnofacies (Skolithos, Cruziana, Zoophycos, Nereites) read environment.

Practice questions (18)

Attempt each one before opening the answer. Every explanation names the tempting wrong option as well as the right one, because that is where marks are lost.

  1. A clay layer at a stratigraphic boundary contains an iridium anomaly, shocked quartz grains and spherules, and a buried crater about 180 km across has been dated to the same time. Which mass extinction is this evidence most directly associated with?

    1. The end-Ordovician extinction, attributed to Gondwanan glaciation and a fall in sea level
    2. The end-Permian extinction, attributed to Siberian Traps volcanism and ocean anoxia
    3. The end-Triassic extinction, attributed to volcanism of the Central Atlantic Magmatic Province
    4. The end-Cretaceous extinction, attributed mainly to a large impact at Chicxulub
    Show answer

    Answer: D — The end-Cretaceous extinction, attributed mainly to a large impact at Chicxulub

    Iridium is rare in crustal rock but abundant in meteorites, and shocked quartz, spherules and a crater of about 180 km are the signature of a large impact; this set of evidence belongs to the end-Cretaceous boundary at about 66 Ma. The end-Permian is the tempting wrong answer because it is the largest extinction, but its record points to flood-basalt volcanism and ocean chemistry, not shock features. The end-Triassic and end-Ordovician events are likewise tied to volcanism and glaciation.
  2. In a measured section the last occurrences of 12 ammonite species are spread through the 4 m of rock below a boundary clay, although independent evidence shows that all 12 were eliminated in one brief event at the boundary. Which effect best explains the apparent gradual disappearance?

    1. The Signor–Lipps effect: with incomplete sampling the last preserved fossil of a taxon lies below its true last occurrence, so ranges seem to end early
    2. The Lazarus effect: taxa vanish from the record during the crisis interval and then reappear in younger rocks once the crisis has passed and conditions recover
    3. The pull of the Recent: younger rocks are better sampled and preserve more taxa, so ranges appear to extend upward towards the top of a section
    4. Time-averaging: shells of different ages are mixed within single beds, so older species are found reworked into the younger beds above them
    Show answer

    Answer: A — The Signor–Lipps effect: with incomplete sampling the last preserved fossil of a taxon lies below its true last occurrence, so ranges seem to end early

    Incomplete preservation means the last fossil found lies below the true last occurrence, so a single event is smeared into an apparent gradual decline: this is the Signor–Lipps effect. A Lazarus taxon needs a reappearance above the gap, which is not described here. The pull of the Recent concerns the better sampling of young rocks and exaggerates diversity near the present. Time-averaging would push apparent last occurrences upward (older shells in younger beds), the opposite of the smearing below the boundary.
  3. A mass extinction removes 60% of the genera of a marine group. Species were lost at random, with the same probability in every genus, and most genera of the group contain several species. Which statement about species loss is correct?

    1. Species loss was about 60%, because species and genera are lost in equal proportions when extinction is random
    2. Species loss was below 60%, because a genus survives if any one of its species survives
    3. Species loss was well above 60%, because a genus disappears only when all of its species have disappeared
    4. Species loss cannot be related to genus loss without knowing the number of individuals per species
    Show answer

    Answer: C — Species loss was well above 60%, because a genus disappears only when all of its species have disappeared

    A genus is lost only if all k of its species are lost. With a random species loss fraction p the genus loss fraction is p^k, which for k > 1 is smaller than p. A genus loss of 60% therefore needs p^k = 0.6, so p = 0.6^(1/k) is well above 0.6 (for k = 3, p is about 0.84). The tempting answer, species loss below 60%, reverses this logic: surviving genera are the ones that retained at least one species, but extinct genera lost all of theirs, which makes the species percentage the larger.
  4. A sandstone bed contains many cavities shaped like gastropod and bivalve shells but no shell material, while the associated articulate brachiopod shells (low-Mg calcite) are intact. Which explanation is most likely?

    1. Carbonisation drove off the volatile components of the molluscan shells and left only thin films of carbon on the bedding planes of the sandstone
    2. The aragonitic molluscan shells dissolved after burial and left moulds, while the low-Mg calcite brachiopod shells resisted dissolution
    3. Permineralisation filled the pores of the molluscan shells with silica, so that they weathered out later as solid casts
    4. Time-averaging mixed shells of several ages, and the molluscan shells were destroyed during the repeated reworking of the bed
    Show answer

    Answer: B — The aragonitic molluscan shells dissolved after burial and left moulds, while the low-Mg calcite brachiopod shells resisted dissolution

    Aragonite is the less stable CaCO3 polymorph and is dissolved or inverted early in diagenesis, whereas low-Mg calcite is stable. Mollusc shells (aragonite) therefore survive as external and internal moulds beside intact calcitic brachiopods. Carbonisation leaves carbon films, not shaped cavities. Permineralisation preserves the shell material itself, so the cavities would not be empty. Time-averaging blends ages but does not selectively delete one mineralogy.
  5. A fossil shell has two unequal valves. A plane of bilateral symmetry cuts through the middle of each valve, and the larger valve has a small opening near its beak for a fleshy stalk. Which identification is best supported?

    1. A bivalve mollusc, because bivalves have unequal valves and each valve has its own plane of symmetry
    2. A gastropod, because the small opening near the beak is the siphonal canal of a snail with a bivalved shell
    3. An oyster-type bivalve, because one valve is larger than the other and cemented to the substrate by a stalk
    4. A brachiopod, because each valve is symmetrical on its own and the pedicle emerges through the larger valve
    Show answer

    Answer: D — A brachiopod, because each valve is symmetrical on its own and the pedicle emerges through the larger valve

    In brachiopods the plane of symmetry passes through both valves, so each valve is symmetrical, the valves differ in size, and the pedicle leaves through an opening in the pedicle valve. Bivalve molluscs have a symmetry plane between the two valves, so a valve alone is normally asymmetrical. Oysters are inequivalve, which makes them the tempting answer, but their valves are irregular and they have no pedicle opening. A gastropod has one shell, not two valves.
  6. In a deep-sea core from the North Atlantic, the share of sinistrally coiled Neogloboquadrina pachyderma falls steadily from 90% at the base to 10% at the top, while the planktic : benthic foraminiferal ratio stays constant. Which interpretation is best?

    1. Surface water cooled upwards, because sinistral coiling in this species marks warm surface water rather than cold water
    2. Surface water warmed upwards, because this species coils mainly sinistrally in cold water and dextrally in warm water
    3. Water depth increased upwards, because sinistral individuals dominate in the shallow shelf waters inhabited by this species
    4. Salinity decreased upwards, because dextral coiling is restricted to brackish water in this species
    Show answer

    Answer: B — Surface water warmed upwards, because this species coils mainly sinistrally in cold water and dextrally in warm water

    In Neogloboquadrina pachyderma the dominant coiling direction tracks surface temperature: sinistral in cold water, dextral in warm. A fall from 90% to 10% sinistral therefore records warming upcore. A depth change would normally shift the planktic : benthic ratio, which stays constant here, and coiling in this species is not a salinity or depth indicator. The option claiming that sinistral coiling marks warm water reverses the temperature relationship.
  7. Dinosaurs are divided into lizard-hipped and bird-hipped groups. Which statement about hip structure and the ancestry of birds is correct?

    1. Birds descend from theropod dinosaurs, which are lizard-hipped saurischians; the bird-hipped dinosaurs are the ornithischians
    2. Birds descend from the ornithischians, which is why that group is called bird-hipped and why it includes Archaeopteryx
    3. Lizard-hipped saurischians have the pubis swung back beside the ischium, as birds do, and are therefore the ancestors of birds
    4. Birds descend from pterosaurs, the flying members of the bird-hipped dinosaur group, which survived the end-Cretaceous event
    Show answer

    Answer: A — Birds descend from theropod dinosaurs, which are lizard-hipped saurischians; the bird-hipped dinosaurs are the ornithischians

    Birds evolved from small theropods, a lizard-hipped saurischian lineage; Archaeopteryx is a theropod-like bird of the Late Jurassic. The ornithischians are called bird-hipped because their pubis rotated backwards beside the ischium, a condition birds evolved independently, so the name misleads. Pterosaurs are not dinosaurs and became extinct at the end-Cretaceous.
  8. A mudstone unit contains millions of individuals of only three species, all thin-shelled bivalves, gastropods and ostracods. Echinoderms, brachiopods, corals and cephalopods are absent from this unit although they occur in the beds above and below it. Which environment is most likely?

    1. A warm, clear, shallow reef flat with normal marine salinity and a rich, diverse fauna
    2. An open outer shelf with normal marine salinity, where all the stenohaline groups of the beds above and below would have flourished
    3. A brackish lagoon or estuary, where only euryhaline taxa tolerate the fluctuating salinity and can reach high abundance
    4. A cold, deep, well-oxygenated basin floor, where the stenohaline groups are excluded by low temperature alone
    Show answer

    Answer: C — A brackish lagoon or estuary, where only euryhaline taxa tolerate the fluctuating salinity and can reach high abundance

    Echinoderms, brachiopods, corals and cephalopods are stenohaline and need normal marine salinity. Their absence between beds that contain them, with a low-diversity, very abundant assemblage of tolerant bivalves, gastropods and ostracods, indicates fluctuating, brackish conditions such as a lagoon or estuary. A reef flat or an open shelf would carry the stenohaline groups, and the bed lies between marine beds, not a cold basin floor, so temperature alone does not explain it.
  9. Which description fits a Konservat-Lagerstätte?

    1. A bed in which the hard parts of many generations are concentrated by currents, as in a bone bed or a shell bed
    2. A sandstone in which the shells have dissolved and survive only as external moulds and internal moulds
    3. A deposit in which fossils from several different habitats were mixed together by transport, so that the whole assemblage is allochthonous
    4. A deposit, such as the Burgess Shale or the Solnhofen Limestone, that preserves soft tissues or whole bodies in exceptional detail
    Show answer

    Answer: D — A deposit, such as the Burgess Shale or the Solnhofen Limestone, that preserves soft tissues or whole bodies in exceptional detail

    A Konservat-Lagerstätte is defined by exceptional preservation of soft tissue or whole bodies, as in the Burgess Shale, Chengjiang and Solnhofen. A concentration of hard parts, such as a bone bed, is a Konzentrat-Lagerstätte, which is the tempting confusion. Moulds and mixed transported assemblages describe other taphonomic outcomes and carry no claim of exceptional soft-tissue preservation.
  10. Which of the following statements about mass extinctions are correct?

    1. Extinction is often selective: narrowly adapted, geographically restricted and reef-building taxa tend to suffer more than widespread generalists
    2. A Lazarus taxon is one that survives an extinction because its individuals are unusually long-lived
    3. Disaster taxa, such as the bivalve Claraia and the therapsid Lystrosaurus after the end-Permian event, briefly become abundant in the aftermath
    4. The percentage of genera lost in an extinction event is normally larger than the percentage of species lost in the same event
    Show answer

    Answer: A — Extinction is often selective: narrowly adapted, geographically restricted and reef-building taxa tend to suffer more than widespread generalists; C — Disaster taxa, such as the bivalve Claraia and the therapsid Lystrosaurus after the end-Permian event, briefly become abundant in the aftermath

    Extinction is selective, and disaster taxa bloom after the end-Permian crisis, so those two statements are correct. A Lazarus taxon is one that disappears from the fossil record and later reappears, not one that is long-lived. Because a genus is lost only when all its species are, the percentage of genera lost is normally smaller than the percentage of species lost, so the genus-versus-species statement is reversed.
  11. Which of the following statements about invertebrate functional morphology are correct?

    1. Gastropod torsion is a 180° twist of the body mass during larval development and is distinct from the coiling of the shell
    2. Regular echinoids are bilaterally symmetrical, with the anus displaced from the apex to the posterior margin of the test
    3. A deep pallial sinus in a bivalve indicates long siphons and a deep-burrowing habit
    4. Ammonoid sutures are typically goniatitic in the Devonian to Permian, ceratitic in the Triassic and ammonitic in the Jurassic and Cretaceous
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    Answer: A — Gastropod torsion is a 180° twist of the body mass during larval development and is distinct from the coiling of the shell; C — A deep pallial sinus in a bivalve indicates long siphons and a deep-burrowing habit; D — Ammonoid sutures are typically goniatitic in the Devonian to Permian, ceratitic in the Triassic and ammonitic in the Jurassic and Cretaceous

    Torsion is a larval twist unrelated to shell coiling; the pallial sinus marks the attachment of siphon retractors, so a deep sinus means long siphons and deep burial; and the goniatitic, ceratitic, ammonitic sequence is the usual pattern of increasing suture complexity. Regular echinoids are radially symmetrical with the anus at the apex; it is the irregular echinoids that have a displaced anus and secondary bilateral symmetry, so that statement is wrong.
  12. Which of the following statements about vertebrate evolution are correct?

    1. Tiktaalik, a Late Devonian lobe-finned fish with a neck and wrist-like fin bones, is a transitional form between fishes and tetrapods
    2. Synapsids, the group containing the therapsids and mammals, have a single opening in the skull behind the eye on each side
    3. Pterosaurs and ichthyosaurs are dinosaurs that returned to the air and the sea respectively
    4. In the evolution of the mammalian ear, the articular and quadrate bones of the reptilian jaw joint became the malleus and incus
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    Answer: A — Tiktaalik, a Late Devonian lobe-finned fish with a neck and wrist-like fin bones, is a transitional form between fishes and tetrapods; B — Synapsids, the group containing the therapsids and mammals, have a single opening in the skull behind the eye on each side; D — In the evolution of the mammalian ear, the articular and quadrate bones of the reptilian jaw joint became the malleus and incus

    Tiktaalik is the classic fish-tetrapod transitional form; synapsids are defined by one lower temporal opening; and the reptilian articular and quadrate became the malleus and incus as the mammalian jaw joint shifted to the dentary and squamosal. Pterosaurs (flying archosaurs) and ichthyosaurs (marine reptiles) are not dinosaurs, so that statement is wrong.
  13. Which of the following statements about fossils and paleoenvironments are correct?

    1. A death assemblage is a snapshot of a single living community, because time-averaging is negligible in most shell beds
    2. A good index fossil is widespread, abundant, quickly evolving and not restricted to a single facies
    3. A woody dicot flora in which most leaves have toothed margins indicates a warmer climate than one with mostly entire margins
    4. Vertical dwelling burrows of the Skolithos type in clean sand typify a high-energy shoreface environment
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    Answer: B — A good index fossil is widespread, abundant, quickly evolving and not restricted to a single facies; D — Vertical dwelling burrows of the Skolithos type in clean sand typify a high-energy shoreface environment

    An index fossil must be widespread, abundant, fast-evolving and independent of facies, and the Skolithos ichnofacies of vertical dwelling burrows marks high-energy sandy shorefaces. A death assemblage is usually time-averaged and may mix communities, so it is not a snapshot. Entire-margined leaves, not toothed ones, dominate warm climates, so the leaf statement is reversed.
  14. A bed contains four species with 40, 30, 20 and 10 individuals. Using Shannon’s H = −Σ p_i ln p_i, with p_i the proportion of individuals of species i and the natural logarithm, the value of H is ____ (give the answer to two decimal places).

    Numerical answer — type the value.

    Show answer

    Answer: 1.28

    p = 0.40, 0.30, 0.20, 0.10. The terms −p ln p are 0.4 × 0.9163 = 0.3665, 0.3 × 1.2040 = 0.3612, 0.2 × 1.6094 = 0.3219 and 0.1 × 2.3026 = 0.2303. Their sum is 1.2799, which is 1.28. The maximum for four species would be ln 4 = 1.39, so the bed is fairly even.
  15. In a stage lasting 4 Myr, 120 genera cross its bottom boundary. Of these, 30 also cross its top boundary and the other 90 become extinct within the stage. A further 40 genera originate and become extinct entirely inside the stage. Use q = −ln(N_bt / N_b) / Δt, where N_b is the number of taxa crossing the bottom boundary, N_bt the number crossing both boundaries and Δt the length in Myr. The per-capita extinction rate q, in per Myr, is ____ (give the answer to three decimal places).

    Numerical answer — type the value.

    Show answer

    Answer: 0.347

    N_b = 120 and N_bt = 30, so N_bt / N_b = 0.25 and −ln 0.25 = 1.3863. Dividing by Δt = 4 Myr gives q = 0.3466, which is 0.347 per Myr. The 40 genera confined to the stage cross neither boundary and are not counted, and using 90 (the number that went extinct) instead of the survivors is the usual slip.
  16. A sample of N = 10 fossil individuals contains three species with 6, 3 and 1 individuals. Two individuals are drawn at random without replacement. Using E(S_n) = Σ [1 − C(N − N_i, n) / C(N, n)], where N_i is the count of species i, the expected number of species in the subsample of n = 2 is ____ (give the answer to one decimal place).

    Numerical answer — type the value.

    Show answer

    Answer: 1.6

    C(10, 2) = 45. For the species with 6 individuals, C(4, 2) = 6, so 1 − 6/45 = 0.8667. For 3 individuals, C(7, 2) = 21, so 1 − 21/45 = 0.5333. For 1 individual, C(9, 2) = 36, so 1 − 36/45 = 0.2. The sum is 1.6. Check: the chance that both drawn individuals share a species is (15 + 3 + 0)/45 = 0.4, so E(S_2) = 1 × 0.4 + 2 × 0.6 = 1.6.
  17. In a brachiopod population two specimens are measured: x = 4 mm and y = 10 mm for the smaller, and x = 64 mm and y = 40 mm for the larger. Assume y = a x^b holds exactly. The predicted value of y, in mm, for a specimen with x = 100 mm is ____.

    Numerical answer — type the value.

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    Answer: 50

    The slope on a log–log plot is b = ln(40/10) / ln(64/4) = ln 4 / ln 16 = 0.5, which is below the isometric value of 1 for two lengths, so this is negative allometry. Then y(100) = 10 × (100/4)^0.5 = 10 × 5 = 50 mm. Using a straight-line (linear) extrapolation instead of the power law is the usual slip.
  18. The lengths of n = 25 brachiopod shells have a mean of 30.0 mm and a standard deviation of 5.0 mm. For a 95% confidence interval of the mean use x̄ ± t s / √n with t = 2.064 (24 degrees of freedom). The full width of the interval, from lower to upper limit, in mm, is ____ (give the answer to two decimal places).

    Numerical answer — type the value.

    Show answer

    Answer: 4.13

    The standard error is s / √n = 5.0 / 5 = 1.0 mm. The half-width is t × SE = 2.064 mm, so the interval is 27.94 to 32.06 mm and its full width is 2 × 2.064 = 4.128, which is 4.13 mm. Reporting only the half-width (2.06) is the usual slip.