| Pustula Temporal range: | |
|---|---|
Scientific classification  | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Brachiopoda |
| Class: | † Strophomenata |
| Order: | † Productida |
| Family: | † Echinoconchidae |
| Subfamily: | † Pustulinae |
| Genus: | † Pustula Thomas, 1914 |
Pustula is an extinct genus of brachiopods which lived during the Carboniferous period. It is the type genus of the subfamily Pustulinae. Its fossils have been found in Eurasia, North America and northern Africa. [1]
Members of the genus Pustula had a subrectangular outline with a weak ventral sulcus, minimal trails and dorsal median folds. Like other echinoconchids, they had recumbent spines arranged in irregular concentric bands, with Pustula having elongated spine bases and weak spine differentiation. The dorsal face of this brachiopod was trifid, and the cardinal process was narrow. Marginal ridges and cardinal process pits are absent in this genus, while the cardinal ridges diminished towards the ears. [1]
Brunton et al. (1995) deemed Pustula to be a primitive member of the Echinoconchidae family, a placement also supported by Muir-Wood & Cooper (1960). [2] [3] Leighton & Maples (2002) conducted multiple phylogenetic analyses which are strongly in agreement that the four subfamilies Buxtoniinae, Echinoconchinae, Pustulinae and Juresaniinae form the family Echinoconchidae. The cladogram results of their phylogenetic analyses are displayed in the cladogram below: [4]
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Brachiopods, phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection. Two major categories are traditionally recognized, articulate and inarticulate brachiopods. The word "articulate" is used to describe the tooth-and-groove structures of the valve-hinge which is present in the articulate group, and absent from the inarticulate group. This is the leading diagnostic skeletal feature, by which the two main groups can be readily distinguished as fossils. Articulate brachiopods have toothed hinges and simple, vertically-oriented opening and closing muscles. Conversely, inarticulate brachiopods have weak, untoothed hinges and a more complex system of vertical and oblique (diagonal) muscles used to keep the two valves aligned. In many brachiopods, a stalk-like pedicle projects from an opening near the hinge of one of the valves, known as the pedicle or ventral valve. The pedicle, when present, keeps the animal anchored to the seabed but clear of sediment which would obstruct the opening.
Rhynchonelliformea is a major subphylum and clade of brachiopods. It is roughly equivalent to the former class Articulata, which was used previously in brachiopod taxonomy up until the 1990s. These so-called articulated brachiopods have many anatomical differences relative to "inarticulate" brachiopods of the subphyla Linguliformea and Craniformea. Articulates have hard calcium carbonate shells with tongue-and-groove hinge articulations and separate sets of simple opening and closing muscles.
Paterinata is an extinct class of linguliform brachiopods which lived from the lower Cambrian ("Tommotian") to the Upper Ordovician (Hirnantian). It contains the single order Paterinida and the subfamily Paterinoidea. Despite being some of the earliest brachiopods to appear in the fossil record, paterinides stayed as a relatively subdued and low-diversity group even as other brachiopods diversified later in the Cambrian and Ordovician. Paterinides are notable for their high degree of convergent evolution with rhynchonelliform (articulate) brachiopods, which have a similar set of muscles and hinge-adjacent structures.
Dictyonellidae is a family of sponges in the order Bubarida, based on the genus DictyonellaSchmidt, 1868.
Helen Marguerite Muir-Wood was a British paleontologist and historian of paleontology who spent her career at London's Natural History Museum. She is a recipient of the prestigious Lyell Medal for her contributions to the field.
Productida is an extinct order of brachiopods in the extinct class Strophomenata. Members of Productida first appeared during the Silurian. They represented the most abundant group of brachiopods during the Permian period, accounting for 45-70% of all species. The vast majority of species went extinct during the Permian-Triassic extinction event, though a handful survived into the Early Triassic. Many productids are covered in hollow tubular spines, which are characteristic of the group. A number of functions for the spines have been proposed, including as a defensive mechanism against predators.
Kutorginates (Kutorginata) are an extinct class of early rhynchonelliform ("articulate") brachiopods. The class contains only a single order, Kutorginida (kutorginides). Kutorginides were among the earliest rhynchonelliforms, restricted to the lower-middle part of the Cambrian Period.
The orthotetides (Orthotetida) are an extinct order of brachiopods in the class Strophomenata. Though not particularly diverse or abundant relative to strophomenides (Strophomenida) or productides (Productida), orthotetides were nevertheless the longest-lasting order of strophomenates, surviving from the Middle Ordovician (“Llanvirn”) up until the Late Permian. Externally, many orthotetides are difficult to distinguish from strophomenides. Most fundamental differences between the two orders are internal: orthotetides have more elaborate cardinal processes and a greater diversity of shell microstructure.
The following is a taxonomy of extant (living) Brachiopoda by Emig, Bitner & Álvarez (2019). There are over 400 living species and over 120 living genera of brachiopods classified within 3 classes and 5 orders, listed below. Extinct groups are not listed.
Schizophoria is an extinct genus of brachiopod belonging to the superfamily Enteletoidea. Specimens have been found in Devonian through Permian beds in North America, Australia, central and southeast Asia, and eastern Europe.
Diaphragmus is an extinct genus of brachiopod belonging to the order Productida and family Linoproductidae. Specimens have been found in Carboniferous beds in North America.
Productidae is an extinct family of brachiopods which lived from the Upper Devonian to Upper Permian periods in marine environments. It is the most diversified family in the suborder Productidina, with some 100 genera.
Echinoconchidae is an extinct family of brachiopods which lived from the Lower Carboniferous to Upper Permian periods in marine habitats. Currently, four subfamilies are assigned to it, though the evolutionary relationships between them and the family Productidae have been heavily debated for the better part of the 20th Century.
Echinoconchinae is an extinct subfamily of brachiopods which lived during the Carboniferous and Permian periods in marine habitats. The family had a cosmopolitan distribution.
Echinoconchus is an extinct genus of brachiopods which lived during the Lower Carboniferous period. The genus was abundant and had a cosmopolitan distribution.
Echinaria is an extinct genus of brachiopods which lived during the Upper Carboniferous and Lower Permian periods. Its fossils have been found in Eurasia, North America and northern South America.
Juresaniinae is an extinct subfamily of brachiopods which lived during the Carboniferous and Permian periods in marine habitats.
Pustulinae is an extinct subfamily of medium to large-sized brachiopods which lived during the Early Carboniferous period in marine habitats. It is named after the type genus Pustula.
Antiquatonia is an extinct genus of brachiopod belonging to the order Productida and family Productidae. Specimens have been found in Carboniferous beds across many continents, suggesting the genus had a cosmopolitan distribution. Species level taxonomy of Antiquatonia is in need of revision.
Pulchratia is an extinct genus of brachiopods which lived in marine habitats during the Upper Carboniferous period. Its fossils have been found in North America.
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