Phanerozoic changes in the high-rank suprageneric diversity structure of brachiopods: Linear and non-linear effects

Phanerozoic evolution of brachiopods produced many linear (established by a comparison of successive geologic time units) and non-linear (established by a comparison of non-successive geologic time units) effects, which can be examined quantitatively by using the similarity coefficients (Czekanowski's Quantified Coefficient and Gower Index) and correlation tools. The high-rank suprageneric diversity structure accounts for a number of superfamilies in each of 26 orders for every epoch of geological time. The intensity of turnovers in this structure was generally low during the entire Phanerozoic. It was slightly stronger during the Early Paleozoic, but close to zero during the Cenozoic, when the high-rank suprageneric diversity structure of brachiopods stabilized finally. Significant turnovers took place at the Middle Cambrian–Early Ordovician, the Late Ordovician–Early Silurian, the Late Silurian–Early Devonian, the Middle Devonian–Mississippian, and the Permian–Triassic transitions. Influences of mass extinctions, both major like those End Ordovician or Permian/Triassic and minor like Early Jurassic or Jurassic/Cretaceous, on the high-rank suprageneric diversity structure of brachiopods is registered. The strongest was the consequences of the Permian/Triassic catastrophe, which perhaps even reset the brachiopod evolution. No evident direct relationships are established between intensity of turnovers and eustatic fluctuations. However, the changes in the diversity structure recorded with the Gower Index provide evidence that eustatic lowstands were more favorable for intensification in these changes.     

Commissural asymmetry in brachiopods

Consistent asymmetric folding of the commissure is a characteristic feature of a small but significant number of brachiopod species. The feature may be obligate or facultative and is almost entirely confined to rhynchonellids, most of which are Mesozoic. The detailed nature of the asymmetry is very variable, but does not extend to internal hard parts such as crura. Taken as a whole, asymmetric brachiopod species show no preference for any particular environment or geographic region, and in no circumstances seem to have been markedly more or less successful than symmetric species. We are thus led to suggest that asymmetry was a genetically based condition which cropped up periodically in brachiopod evolution, and which possibly was selected neither particularly for nor against.     

'Flapping valves' in brachiopods

M. J. S. Rudwick and others postulate 'rhythmic-flow' feeding for the Permian richtho-feniacean bi-achiopods, whereas R. E. Grant claims that they fed by normal ciliary action. Suspension-feeding has two components, current generation and food capture; normal brachiopod lophophores do both, but this is neither universal nor compulsory among animals. Opening and closing the richthofeniid shell generated a 'tidal-flow' current precisely analogous to respiratory currents in mammals; this is neither inefficient nor 'self-defeating', as Grant claims. Grant's analysis fails because he chose the wrong mechanical analogy (a pump). Morphological features of richthofeniids are better explained on a tidal-flow hypothesis than on a ciliary-flow model, and all the data adduced by Grant in rejecting the former is compatible with it or favorable to it. It explains morphological features that are bizarre mysteries on the ciliaiy-current model, and is therefore superior even though it implies that these Permian brachiopods were radically innovative.     

Ecophenotypic strategies of dalmanellid brachiopods

General ecophenotypic patterns, of particular interest when they apply to all, or most, taxa of the group concerned, can never be demonstrated until after monophyletic taxa have been recognized, that is, until after the initial stages of phylogeny construction have been carried out. In criticizing certain dalmanellid phylogenies, and based in large part on a study of five 'species subgroups'. Hurst & Watkins (1978; Geologica et Palaeontologica 12 ) postulate ecophenotypic patterns for Isorthis , and Hurst (1978; Palaeontology 21 ) postulates general patterns of ecophenotypic variation for dalmanellid brachiopods. These patterns may be invalid for four reasons: (1) Univariate and 'bivariate' statistical analysis of the samples used to define the five subgroups reveals no significant differences between subgroups, or vertical trends, for the very morphological characters claimed to exhibit the ecophenotypic patterns; (2) Hurst & Watkins' discriminant function analysis contains procedural errors and its results are ambiguous; (3) several of the five subgroups represent mixtures of unrelated taxa; (4) in recognizing the alleged patterns, Hurst & Watkins ignored contrary evidence from many taxa (and from many dalmanellid studies). □ Brachiopoda, Dalmanellidae, Silurian, ecology, evolution, systematics.     

Allometric studies of fossil brachiopods

In different groups of fossil brachiopods new adaptations were formed by changes in size, shape, and proportions of morphological structures in ontogenetic and phylogenetic development.     

Biogeography of the recent brachiopods

The vertical, latitudinal, and circumcontinental zonality of the distribution of the species, genera, and families of recent brachiopods is considered. The distortions of the latitudinal and meridional symmetry of the biogeographic structure of the ocean are analyzed in view of the patterns of the global circulation of the surface and intermediate waters. Thus ancient faunas may be reconstructed based on data on the structural characteristics of the taxocene of recent brachiopods. The features of the paedomorphic evolution of brachiopods from the different families in extreme habitats (interstitial, underwater caverns, submarine rises, abyssal depths, hydrothermal areas, and margins of habitats) are discussed. The biogeographic structure of bottom dwellers is shown to simplify with depth as well as with simplification of the hydrological structure of the ocean. The important role of the bathyal oceanic zone (slopes of continents, islands, submarine mountains, ridges, and rises) in the preservation of faunal relicts is shown. The historical change from brachiopods to bivalves that occurred from the Paleozoic to the Mesozoic and Cenozoic is shown to have resulted not from competitive exclusion, but from complex and global changes in the plankton composition, which were unfavorable for articulate brachiopods, which had already developed specialized feeding habits, feeding on food that led to the production of almost no metabolic waste products; they had even partly lost their alimentary canal. The development of shelly plankton and, especially, of diatoms hampered the post-Paleozoic revival of large assemblages of articulate brachiopods in shallow-water habitats. The unfilled ecological niches were colonized by bivalves, which were widely adapted to feeding on live phyto-and zooplankton. Recent articulate brachiopods, which are adapted to feeding on the products of decay of dead plankton, form a belt of densely populated settlements of the organic biofilter outside the photic zone on the seaward edge of shelves and on the upper parts of the slopes of continents, islands, and submarine rises throughout the world.     

Origin and dispersal of the earliest brachiopods

Brachiopods first appeared at the very beginning of the Phanerozoic together with the first skeletal organisms. Most brachiopod taxa that arose in the first half of the Cambrian had a short temporal range and became completely extinct by the middle of the Middle Cambrian. Rigid articulation of the valves of brachiopods was provided by various structures, which also appeared in the Early Cambrian. This fact points to the importance of this feature for the formation of the whole group and at the same time testifies to the high variability of rigid articulation at the early stages of brachiopod evolution. This is a typical manifestation of archaic diversity in this animal phylum, which appeared very early in the Phanerozoic. Another important property of the archaic diversity of the early brachiopods was the large number of centers of diversification. As for the majority of groups, climatic zonality was the main factor determining the distribution of brachiopods at the beginning of the Phanerozoic. The main ecological types of brachiopods also appeared in the Early Cambrian.     

Gene-history correlation and population structure

Correlation of gene histories in the human genome determines the patterns of genetic variation (haplotype structure) and is crucial to understanding genetic factors in common diseases. We derive closed analytical expressions for the correlation of gene histories in established demographic models for genetic evolution and show how to extend the analysis to more realistic (but more complicated) models of demographic structure. We identify two contributions to the correlation of gene histories in divergent populations: linkage disequilibrium, and differences in the demographic history of individuals in the sample. These two factors contribute to correlations at different length scales: the former at small, and the latter at large scales. We show that recent mixing events in divergent populations limit the range of correlations and compare our findings to empirical results on the correlation of gene histories in the human genome.     

Symbolic trephinations and population structure

The sample examined consists of 19 skulls with symbolic trephinations and 86 skulls without trepanations dated from the X century. Skulls were all excavated in the Great Hungarian Plain in the Carpathian Basin, which was occupied by the Hungarian conquerors at the end of the IX century. The variations of 12 cranial dimensions of the trephined skulls were investigated and compared to the skulls without trepanations after performing a discriminant analysis. The classification results evince that the variability of non-trephined skulls shows a more homogeneous and a more characteristic picture of their own group than the trephined samples, which corresponds to the notion, formed by archaeological evidence and written historical sources, of a both ethnically and socially differing population of the Hungarian conquerors. According to historical research, a part of the population was of Finno-Ugric origin, while the military leading layer of society can be brought into connection with Turkic ethnic groups. All the same, individuals dug up with rich grave furniture and supposed to belong to this upper stratum of society are primarily characterized by the custom of symbolic trephination, and, as our results demonstrate, craniologically they seem to be more heterogeneous.     

Size-frequency distributions along a latitudinal gradient in Middle Permian fusulinoideans

Geographic gradients in body size within and among living species are commonly used to identify controls on the long-term evolution of organism size. However, the persistence of these gradients over evolutionary time remains largely unknown because ancient biogeographic variation in organism size is poorly documented. Middle Permian fusulinoidean foraminifera are ideal for investigating the temporal persistence of geographic gradients in organism size because they were diverse and abundant along a broad range of paleo-latitudes during this interval (～275-260 million years ago). In this study, we determined the sizes of Middle Permian fusulinoidean fossils from three different paleo-latitudinal zones in order to examine the relationship between the size of foraminifers and regional environment. We recovered the following results: keriothecal fusulinoideans are substantially larger than nonkeriothecal fusulinoideans; fusulinoideans from the equatorial zone are typically larger than those from the north and south transitional zones; neoschwagerinid specimens within a single species are generally larger in the equatorial zone than those in both transitional zones; and the nonkeriothecal fusulinoideans Staffellidae and Schubertellidae have smaller size in the north transitional zone. Fusulinoidean foraminifers differ from most other marine taxa in exhibiting larger sizes closer to the equator, contrary to Bergmann's rule. Meridional variation in seasonality, water temperature, nutrient availability, and carbonate saturation level are all likely to have favored or enabled larger sizes in equatorial regions. Temporal variation in atmospheric oxygen concentrations have been shown to account for temporal variation in fusulinoidean size during Carboniferous and Permian time, but oxygen availability appears unlikely to explain biogeographic variation in fusulinoidean sizes, because dissolved oxygen concentrations in seawater typically increase away from the equator due to declining seawater temperatures. Consequently, our findings highlight the fact that spatial gradients in organism size are not always controlled by the same factors that govern temporal trends within the same clade.     

Lines of phylogenetic studies of brachiopods

The main techniques used in phylogenetic studies of brachiopods are reviewed, onto-phylogenetic and morphobiological. The advantages and shortcomings of each method are discussed.     

The origin of the spiriferidine brachiopods

Wright, Anthony D. 197901 15: The origin of the spiriferidine brachiopods. Lethaia . Vol. 12. pp. 29–33. Oslo. ISSN 0024–1164.
In recent years doubts have been expressed as to whether the strophic spiriferidines should continue to be grouped with the non-strophic spiriferids in the order Spiriferida or not. The solution to this problem will only be found by establishing the origin of the spiriferidines. A characteristic feature of the spiriferidines is their well developed micro-ornament; this is shared also by the orthid Platystrophiinae, and together with the striking similarity in overall morphology (well recognized in the early nineteenth century) it suggests that here, in the Platystrophia plexus, is the ancestral stock of the strophic spirebearers.
The Platystrophünae share with the non-strophic Rhynchonellida the potential to develop a spire. It is here considered that the former gave rise to the spiriferidines and the latter to the non-strophic spire-bearers. These quite separate developments justify the separation of the spire-bearers into the two orders Spiriferida and Atrypida.     

Ultrastructure of the loop of terebratulide brachiopods

The folded and twisted calcareous ribbon, forming both the ascending and descending lamellae of the loop of Waltonia inconspicua (Sowerby), is a two-layered structure consisting of a wedge of regularly stacked secondary layer fibres that overlie a thin layer of non-fibrous calcite (herein termed brachiotest). On one surface, that facing into the mantle cavity, secondary fibrous mosaic predominates, but smooth, finely banded brachiotest occurs as a narrow marginal lip upon which secondary layer fibres proliferate and progressively overlap. This growing edge of the ribbon is secreted by long, folded epithelial cells with digitate extensions to their apical plasmalemmas, which are distinguishable from the cuboidal epithelium-secreting fibres and their membranous sheaths. The other surface, facing the body cavity and the brachial coelom, consists entirely of roughened brachiotest exhibiting prominent banding that is aligned parallel to the growing loop edge. This surface is overlain by microfilamentar epithelium acting as a holdfast for the connective tissue frame of the lophophore. The other edge of the ribbon consists of truncated sections of both secondary-layer fibres and brachiotest which bear signs of resorption consistent with the degenerated state of the associated epithelium. Growth of the Waltonia loop is controlled by these localized processes of secretion and resorption of the fibrous and brachiotest layers and is typical of all terebratulides so far studied. The brachiotest is not homologous with the non-fibrous primary shell secreted at the valve margin. □ Brachiopoda, Articulata, Terebratulida, ultrastructure, lophophore, loop.     

Predatory asteroids and the decline of the articulate brachiopods

Various causes, such as increased predation pressure, the lack of planktotrophic larvae, a 'resetting' of diversity, increased competition from benthic molluscs and the decline of the Palaeozoic fauna, have been suggested to explain the failure of the brachiopods to reradiate following the Permo-Triassic mass extinction. Increased predation pressure has hitherto appeared improbable, because typical predators of brachiopods, such as teleostean fish, brachyuran crabs and predatory gastropods, did not undergo major radiation until the late Mesozoic and early Cenozoic. However, new evidence strongly suggests that one important group of predators of shelly benthic organisms, the asteroids, underwent a major radiation at the beginning of the Mesozoic. Although asteroids appeared in the early Ordovician, they remained a minor element of the marine benthos during the Palaeozoic acme of the brachiopods. However, these early asteroids lacked four important requirements for active predation on a bivalved epifauna: muscular arms (evolved in the early Carboniferous); suckered tube feet, a flexible mouth frame and an eversible stomach (all evolved in the early Triassic). Thus radiation of the Subclass Neoasteroidea coincided with both their improved feeding capability and the decline of the articulates. The asteroids were the only group of predators of brachiopods that underwent a major adaptive radiation in the earliest Mesozoic. The asteroids may therefore have contributed to inhibiting a Mesozoic reradiation of the brachiopods. Epifaunal species lacking a muscular pedicle may have been particularly vulnerable. Unlike bivalve molluscs, modern brachiopods show only a limited range of adaptations to discourage asteroid predation. □ Asteroidea, Brachiopoda, evolution, predation, functional morphology.     

Habitat-dependent population regulation and community structure

Summary Density-dependence provides a causal link between processes acting at different levels of ecological organization. The linkage between density-dependent habitat use, population regulation and community organization is examined on the basis of qualitative and quantitative differences between habitats. These differences are expressed as characteristic shapes on isodars which are lines of equal fitness, and are plotted in density space as lines at every point of which the fitness of individuals in one habitat is equal to that of individuals in another. Isodars can be constructed for single species or modified to include the effects of interacting species. Isodars are easily analyzed by linear regression to differentiate between alternative modes of population regulation and to suggest patterns of community structure. Different isodars are causally related to different kinds of community structure, and suggest the existence of four new forms of community organization; equal, differential, switched and mixed preferences. A preliminary isodar analysis on a common rodent species demonstrates that population regulation depends upon habitat, and that mixed preferences probably organize the rodent community. Habitat-dependent population regulation has farreaching implications to studies of temporal and spatial scale, and to all ecological processes that are density-dependent.