Signor-Lipps Research Articles

Demise of the middle Paleozoic crinoid fauna: gradual or mass extinction?

There are essentially three major crinoid faunas of the Paleozoic: 1) the lower Paleozoic (Ordovician) fauna dominated by disparids and diplobathrid camerates; 2) the middle Paleozoic (Silurian-middle Mississippian) fauna dominated by monobathrid camerates, cladids, and flexibles; and 3) the upper Paleozoic (middle Mississippian-Permian) fauna dominated by cladids. Change from the middle Paleozoic fauna to the late Paleozoic fauna, at or near the Osagean-Meramecian boundary, was characterized by Laudon (1948) as “one of the most remarkable faunal breaks in the entire Paleozoic era”. The monobathrids that had reached their zenith in the Osagean (Tournaisian-Visean) became a very minor component of late Paleozoic faunas. Conventional thinking has implied that a mass extinction of crinoids occurred at the Osagean-Meramecian boundary.We have biostratigraphically subdivided the late Osagean and early Meramecian into four zones (times A-D, oldest to youngest, all within the Gnathodustexanus zone) in order to study the origination and extinction of all crinoid species during the changeover from the middle to upper Paleozoic faunas. Rather than a mass extinction, a monotonic turnover of species fits a pattern of gradual extinction as extinctions outpaced originations. Data are based on 216 species (taxonomically updated) from 69 localities and are as follows:The pattern of originations and extinctions for monobathrids is very similar to the pattern for all crinoid species.The above data have maximum diversity during Time B of the late Osagean followed by declining diversity in times C and D into the early Meramecian. Late Meramecian crinoid faunas (after time D) were less diverse with a maximum of about 45 species. The gradual decline of crinoids across the Osagean-Meramecian boundary cannot be explained as sampling bias (Signor-Lipps effect), because ranges of crinoids thought to be extinct at the boundary are extended into the early Meramecian as well as disappearing at various tinles prior to the boundary.The decline in diversity and the monotonic turnover of species are hypothesized to be the result of habitat reduction as the Eastern Interior Basin of North America was gradually infilled with clastic sediments at the end of the Acadian Orogeny. Most notable in this regard was the smothering of the Keokuk Limestone carbonate bank. Changes in sea level also contributed to habitat reduction. Lowering of sea level at the end of the Osagean caused restriction of open marine environments. A transgression in the early Meramecian was followed by shoaling and restriction during deposition of the late Meramecian Salem and St. Louis limestones.

Sampling, species abundance, and the stratigraphic signature, of mass extinction: A test using Holocene tidal flat molluscs

Assume that a catastrophe caused the sudden extinction of the molluscan biota of a Holocene tidal flat. Would this event be recorded accurately in the stratigraphic record? How accurately would the extinction of rare species be recorded? How would sudden, gradual, and stepwise patterns of extinction compare? Bivalve and gastropod species occurrences were recorded from eight tidal channel sediment cores (up to 70 cm long) from Bahia la Choya, northern Gulf of California. The distribution of last occurrences of 45 species suggests that a sudden extinction would appear to be gradual in the stratigraphic record, due to error in biostratigraphic range end-points (Signor-Lipps effect). Extinction is typically not accurately recorded for species with less than 15% stratigraphic abundance (i.e., occurring in less than 15% of the sample intervals; 71% of the species). The data were subjected to simulations of stepwise and gradual extinction. Differences between the sudden, stepwise, and gradual scenarios are apparent in (1) frequency distributions of biostratigraphic last occurrences, (2) graphic relations between stratigraphic abundance and last occurrence, and (3) patterns of apparent diversity decline below the extinction boundary. Extinction studies in outcrops and cores could utilize last occurrence and stratigraphic abundance data (readily available in the fossil record) to distinguish among sudden, stepwise, and gradual patterns of extinction.

Delayed recovery and the spacing of major extinctions

Approximate periodicity for peak rates of global extinction during the past 250 m.y. may have resulted from delayed recovery following major extinction events. Two components can be envisioned for such delays: persistence of inimical environmental conditions for some time after the onset of the crisis, and slow restoration of vulnerable taxa. This general hypothesis is consistent with statistical evidence of linkage between measured rates of extinction of marine invertebrate genera for contiguous stages and substages of the geologic column. The nine broad valleys between the “periodic” peak rates for the past 250 m.y. exhibit only three trivial secondary peaks, indicating that, if the pattern is not artifactual, trends in global rates of extinction have not readily been abruptly reversed. Moreover, the smooth observed trends reflect the fact that regional crises tend to remove many species but few genera. To some degree, high rates of extinction that precede peak rates must represent bias of the incomplete fossil record (the Signor-Lipps effect). High rates that immediately follow peak rates also may, to a degree, reflect biological legacy: (1) final extinction of weakened genera or (2) extinction of new genera that contain few species or represent failed evolutionary “experiments.” Nonetheless, there is evidence that protracted intervals of stressful environmental conditions contributed to high rates of extinction preceding or following certain peak intervals, including the Scythian, Cenomanian, Early Paleocene, and Early Oligocene. The reef-building rudists, for example, suffered heavy extinction during both Cenomanian and Turonian time and then failed to recover quickly.The late Neogene record of bivalve molluscs in the Western Atlantic offers a more detailed picture of delayed recovery. Here early intervals of glacial expansion caused heavy extinction, leaving an impoverished, eurythermal fauna that was virtually unaffected by late Pleistocene glacial episodes. The episode of heavy extinction in Late Eocene time exhibits a similar phenomenon on a worldwide scale. Among the planktonic foraminifera, warm-adapted stenothermal species died out, and eurythermal forms predominated throughout Oligocene time; restoration of vulnerable, stenothermal species proceeded gradually during the Miocene Epoch. This example of delayed recovery and others like it following earlier global crises may have prevented such crises from following one another in rapid succession, yielding an appearance of periodicity.

A Substantial Bias in Nonparametric Tests for Periodicity in Geophysical Data

A nonparametric test that has been used to conclude that extinction rates are periodic with a period of 26 million years is shown to be substantially biased toward this conclusion, regardless of whether or not the data are periodic in origin (and, indeed, regardless of the actual period if they are in fact periodic). The test is shown to be sensitive to measurement error of a type expected with these data (early recording of extinctions due to missing fossil specimens, the "Signor-Lipps effect"), and it is shown that because of the unequal spacing in time, such models may be expected to produce statistically significant but artifactual periods of (in this case) exactly 26 million years over the span of time actually used.