Diadectidae is an extinct family of early tetrapods that lived in what is now North America and Europe during the Late Carboniferous and Early Permian, and in Asia during the Late Permian. They were the first herbivorous tetrapods, and also the first fully terrestrial animals to attain large sizes. Footprints indicate that diadectids walked with an erect posture.
The best known and largest representative of the family is Diadectes, a heavily built animal that attained a maximum length of several metres. Several other genera and various fragmentary fossil remains are also known. Although well known genera like Diadectes first appear in the Late Pennsylvanian, fragmentary remains of possible diadectids are known from much earlier deposits, including a piece of lower jaw found in Mississippian strata from Tennessee.
Description
thumb|left|Life restoration of [[Diadectes, a well-known Early Permian diadectid]]
Diadectids have large bodies with relatively short limbs. The rib cage is barrel-shaped to accommodate a large digestive tract necessary for the digestion of cellulose in plants. The skulls of diadectids are wide and deep with blunt snouts. The internal nares (holes for the nostrils) are also short. Paleontologist E.C. Case compared diadectids to turtles in 1907, noting their large pectoral girdles, short, strong limbs, and robust skulls. Case described them as "lowly, sluggish, inoffensive herbivorous reptiles, clad in an armor of plate to protect them from the fiercely carnivorous pelycosaurs."
Diadectids have a heterodont dentition, meaning that their teeth vary in shape along the length of the jaws. The teeth are wide and bear many cusps or projections, an indication that diadectids ate tough plants. Some teeth are leaf-shaped and laterally compressed, another indication that diadectids were able to shred plant material. The procumbent front teeth of the lower jaw project forward. Diadectids likely had strong jaw muscles for processing plant material; the placement of the jaw joints above or below the level of the occlusal plane (the plane at which the teeth come together) would have given diadectid jaws mechanical advantage. The joints themselves give the jaws a complex range of movement suitable for consuming plants. Large holes and cavities in the skull called adductor chambers and temporal openings would have provided room for large jaw-closing muscles.
History of study
The first diadectid to be described was Diadectes. American paleontologist Edward Drinker Cope named the genus in 1878 on the basis of several vertebrae and teeth from the Early Permian of Texas. Cope erected the family Diadectidae in 1880 to include Diadectes and Empedocles, a genus he named two years earlier. Nothodon, named by Cope's rival Othniel Charles Marsh in 1878, was soon placed in the family.
Cope named several other diadectids, including Helodectes in 1880, Chilonyx and Empedias in 1883, and Bolbodon in 1896. Paleontologist E.C. Case named four other diadectids: Desmatodon in 1908, Diasparactus in 1910, Diadectoides in 1911, and Animasaurus along with paleontologist Samuel Wendell Williston in 1912. Case and Williston considered Marsh's Nothodon and Cope's Bolbodon to be synonymous with Diadectes. Marsh named Nothodon in the American Journal of Science only five days before Cope described Diadectes in Proceedings of the American Philosophical Society. Under rules of the International Code of Zoological Nomenclature, the name Nothodon would have priority over Diadectes, but because the name Diadectes has been in use since Case and Williston first synonymized the genera, Diadectes remains the accepted name. A fourth genus, Ambedus, was named in 2004 from the Early Permian of Ohio.
thumb|left|[[Orobates, a diadectid from Germany]]
Diadectids are also known from Germany. Phanerosaurus was described from several vertebrae near Zwickau by German paleontologist Christian Erich Hermann von Meyer in 1860, but was not recognized as a diadectid until 1925. A second species of Phanerosaurus was identified from some vertebrae and a fragmentary skull in 1882, and was given its own genus, Stephanospondylus, in 1905. In 1998, a new species of Diadectes, D. absitus, was described from the Bromacker sandstone quarry of the Tambach Formation in the Thuringian Forest of central Germany. A new genus of diadectid called Orobates was also named from the Bromacker Quarry in 2004.
In 2015, the known geographic range of diadectids was expanded with the description of a new genus and species of diadectid from China, Alveusdectes fenestralis. Alveusdectes is also the youngest known diadectid by 16 million years, coming from a unit of the Late Permian Shangshihezi Formation that dates to about 256 million years. However, the assignment of Alveusdectes to Diadectidae has been questioned. A phylogenetic analysis from 2024 recovers Alveusdectes as a non-diadectomorph synapsid, more closely related to the eothyridids Eothyris and Oedaleops.
Classification
Diadectids have long been considered close relatives of the amniotes, tetrapods that lay eggs on land or retain the fertilized egg within the mother. In 1987, the paleontologist D.M.S. Watson placed the family in the larger group Diadectomorpha, which includes another family of large-bodied diadectomorphs, the Limnoscelidae, as well as the monotypic diadectomorph family Tseajaiidae, represented by the genus Tseajaia. Throughout the twentieth century, amniotes and diadectomorphs were often grouped together using the old name Cotylosauria, a name originally used for the most basal grade of what was then thought to be reptiles. In the early part of the century, many paleontologists regarded diadectids, along with other cotylosaurs (such as placodonts), to be close relatives of turtles. In most recent studies of early tetrapod phylogeny, Cotylosauria is no longer recognized and Diadectomorpha is placed as the sister taxon of Amniota. However, while the majority of analyses now place diadectids outside Amniota, some have found them to be true amniotes.
Most phylogenetic studies of the three diadectomorph families – Diadectidae, Limnoscelidae, and Tseajaiidae – have found diadectids and limnoscelids to be more closely related to each other than either is to Tseajaia. In other words, Diadectidae and Limnoscelidae form a clade within Diadectomorpha and Tseajaia is excluded from the clade. In a 2010 phylogenetic analysis, Diadectidae formed a clade that was characterized by wide cheek teeth with cusps on either side. Unlike previous studies, it was found to be more closely related to Tseajaiidae than Limnoscelidae. The family was defined as Diadectes and all taxa sharing a more recent common ancestor with Diadectes than with Tseajaia. Below is a cladogram modified from the 2010 analysis: The same results were found in the 2010 analysis. Two new genera were erected in the study to include D. absitus and D. sanmiguelensis. D. sanmiguelensis, the more basal of the two forms, was placed in the new genus Oradectes. D. absitus was renamed Silvadectes.
Evolutionary history
Diadectids were some of the first tetrapods, or four-legged vertebrates, to attain large sizes. Diadectids first appear in the Late Carboniferous with the genus Desmatodon, although recently described bones from Tennessee suggest that they may have appeared even earlier in the Early Carboniferous. They underwent a small evolutionary radiation in the Late Carboniferous and Early Permian, diversifying into thirteen species and outnumbering other diadectomorphs, such as the limnoscelids. This radiation was likely the result of diadectids' expansion into a new herbivorous ecological niche that was previously unfilled.
Diadectids had a much wider geographic distribution than their relatives; while the distribution of limnoscelids is limited to parts of North America and Tseajaia is restricted to just the southwestern United States, diadectids are present in North America, Europe, and Asia.
Although they bear similarities to those of amniotes, the tarsal bones of diadectids are poorly ossified and loosely connected. The digits of the foot connect only to the fourth distal tarsal, providing a wide range of movement in the foot. This flexibility enabled diadectids to rotate their feet in a forward position while walking, providing greater force when pushing off. The feet could also be placed closer to the midline of the body to give diadectids an erect stance. The close positioning of the footprints attributed to the more advanced diadectides suggests that the animals held their feet almost underneath their bodies, giving them a more efficient gait and to some degree paralleling the stance of mammals more than that of the sprawling amphibians and most reptiles.
References
General references
- Carroll, R. L. (1988). Vertebrate Paleontology and Evolution. WH Freeman and Company, New York.