PLEASE NOTE: THIS PROJECT IS NOT SCIENTIFIC. IT IS A HOBBY.
"I was looking for information on an old animal and found this lot. What is this project?"
It's got lots of information on old animals. For a short bit of background information, see here.

Traversodontidae is composed of fairly advanced, plant-eating, non-mammalian therapsids, and is part of a line called Cynognathia. The name of the family gives a clue as to the construction of their postcanine teeth, which are of an expanded width. This is in contrast to the sectorial (blade-like) condition seen in the mouths of their meat-eating colleagues of Probainognathia. Better equipped for dealing with plants It's also a contrast to herbivore forerunners such as Trirachodon, which used to maintain some sectorial teeth in adulthood, (at least in earlier forms). The postcanines of these later animals were more complex, (Kemp 2005, p.67), as there was an impressive crest running transversely across the crown. This is positioned towards the rear in uppers but to the front in lowers, and this resulted in a form of cutting action, (p.68). The lower jaw then went gently backwards and that produced a measure of grinding ability. Reasonably close occlusion between the tooth rows had developed. Such a festure is generally only found in mammals. The benefits were somewhat limited, as the teeth tended to erode to stumps. Decent enamel would've helped, but none was available. Other mammalianosities Traversodonts generally had relatively wide snouts, and the maxilla extends sidewards beyond the teeth. This may indicat the presence of useful bags known as cheeks. Such devices allow for food to be more thoroughly chewed. The ribs are also more mammal-like than for earlier cynodonts. Previous models had spines fitted with costal plates. These bits of bone overlapped with each other where the ribs met the back bone. These strengthed the spine, but what advantage that provided is unclear, (p.111). Such features are much reduced in more basal traversodonts. Derived models, (eg. Exaeretodon), did away with them entirely. This gives the spine a less clanky, more mammalian appearance. Traversodontids foraged through the world of the Middle and Upper Triassic, and so some made the acquaintance of early dinosaurs. Although they seemed destined to become megastars in their own right, they actually reacted by going extinct. Typically, the upper postcanine teeth of these animals are roughly rectangular in outline, when viewed from above (occlusal). The corresponding lowers tend to be squarer, (Abdala & Ribeiro 2003, p.538). Along with the even more mammal-like Tritylodontidae, (see Jurassic cynodonts), they have been placed in the taxon of Gomphodontia, (or alternatively Tritylodontoidea). It's possible that the generally later tritys are descended from traversodontids, but this is increasingly seen as doubtful. Presently, I'm swayed by argmentation placing the tritys within Probainognathia. This page has been rebuilt. It originally contained Diademodontidae, Trirachodontidae and Traversodontidae. Those other two families are more basal and may be visited at Cynognathia.

TRAVERSODONTIDAE

Taxon: Traversodontidae von Huene, 1936 Traversodontids were the most widespread, diverse and derived of the Triassic gomphodonts. They were herbivores of the later Middle and early Upper Triassic. The first representatives were relatively small. By the Ladinian-Carnian, most members were large to very large. Strangely enough, the last of the travies would have been bullied by mice, had rodents been invented. They were tiny, shrew-sized creatures; about as big as the first basal mammals. Our ancestors appeared on the stage just as the traversodontids were bowing out. Their departure was possibly connected with the radiation of non-mammalian tritylodontids. The Upper Triassic representatives of the northern and southern hemispheres show distinguishing characteristics in the postcanine teeth. The northerners have more resemblance to older southern forms, (Abdala & Ribeiro 2003, p.540). Boreogomphodon and Plinthogomphodon are reminiscent of Luangwa and Traversodon. Whilst Arctotraversodon is most similar to Massetognathus, this is specifically the Ladinian version. Of the tiny European forms, Maubeugia seems closest to Boreogomphodon, and Rosieria has features in common with Scalenodon angustifrons and Traversodon. This suggests that the northerners diverged and developed in their own sweet way, presumably during the Ladinian. Their teeth remained relatively conservative. A non-traversodont At least one critter in the following collection isn't a travie. Rather, it's an Upper Triassic gomphodont of unclear affinities. Gomphodonts presently divide into three families; Diademodontidae, Trirachodontidae and Traversodontidae. Italy's first gomphodont genus, Gorngomphodon, has declined invitations to fit into any of those. Still, it's chronologically closer to the travies so I've accommodated on this directory.

Links: Mikko K. Haaramo, Traversodontidae Mikko Haaramo's Traversodontidae Mikko Haaramo saves me a great deal of time. He's also a very helpful chap. M Alan Kazlev, The Kannemeyeriid-Rhynchosaur-Traversodontid Empire M Alan Kazlev's The Kannemeyeriid-Rhynchosaur-Traversondontid Empire A good paleo-ecological overview of terrestrial goings on during the Triassic. T Mike Keesey, The Ages of the Mesozoic http://dinosauricon.com/times/index.html The times they were a changing, and this shows when.

Ischigualato, Argentina Traversodontids are a significant element of the local fossil fauna. Links: 'La Importanica turistica de Ischigualasto' preparado por Dr William Sill http://www.ischigualasto.com/es/ischigualasto.htm Annexo VII 3. Ischichuca & Chanares Formations: Megagomphodon Annexo VII 5. Ischigualasto Formation: Exaeretodon, Ischignathus, Proexaeretodon Raymond Rogers, Research in Argentina http://www.macalstr.edu/~geology/rogers/Argentina.html Reports and photos from Ischigualasto. Genera: Andescynodon, Arctotraversodon, Boreogomphodon, Colbertosaurus, Dadadon, Exaeretodon, Gomphodontosuchus, Gornogomphodon (not a travie). Habayia, Ischignathus, Luangwa, Massetognathus, Maubeugia, Megagomphodon (=Massetognathus), Menadon, Microscalenodon, Nanogomphodon, Pascualgnathus, Plinthogomphodon, Proexaetetodon (=Exaeretodon), Protuberum, Rosieria, Rusconiodon (perhaps = Andescyndon), Santacruzodon, Scalenodon (and partly = Luangwa), Scalenodontoides, Theropsis (=Exaeretodon), Theropsodon, Trirachodon (partly = Scalenodon), Traversodon, other reports Time-Line: Upper Triassic: Arctotraversodon, Boreogomphodon, Exaeretodon, Gomphodontosuchus, Gornogomphodon (not a travie), Habayia, Ischignathus, Maubeugia, Microscalenodon, Plinthogomphodon, Rosieria, Scalenodontoides, Saint-Nicholas-de-Pont (France), Syren (Luxembourg), North Carolina Middle Triassic: Colbertosaurus, Dadadon, Exaeretodon, Luangwa, Massetognathus, Menadon, Nanogomphodon, Protuberum Santacruzodon, Scalenodon, Theropsodon, Traversodon Lower Triassic: Andescynodon, Pascualgnathus, Rusconiodon

Genus: Andescynodon Bonaparte JF, 1969 'Andes dogtooth'

Species:	Andescynodon mendozensis Bonaparte JF, 1969
Place:	Cerro de las Cabras Formation, (?Rio Mendoza Formation)
Country:	Argentina
Age:	probably late Scythian - early Anisian, Lower - Middle Triassic
Remarks:	A small and basal form. A number of specimens are in the collection of the Universidad Nacional de Tucumán, Argentina. Formation One source, I forget which, informed me the locality was in the Rio Mendoza Formation. A more recent one, Abdala & Smith, 2009, cites the Cerro de las Cabras Formation (p. 12). I can't explain the significance, if any, of that apparent descrepency.
Reference:	Bonaparte (1969), Dus nuevas 'faunas' de reptiles triásicos de Argentina, Gondwana Stratigraphy, IUGS Symposium, Mar del Plata, p.283-306.

Link: Clarin digital / Sociedad: Un nuevo tramo de ruta destruira yacimientos unicos de fosiles http://www.clarin.com/diario/2000-10-aa/s-05401.htm A report in Spanish, which gives an age of ca. 240 Ma. It also refers to Rusconiodon.

Genus: Arctotraversodon Sues HD, Hopson JA & Shubin NH, 1992 'northern Traversodon' Aka: Scalenodontoides (partly)

Species:	Arctotraversodon plemmyridon (Hopson JA, 1984) Sues HD, Hopson JA & Shubin NH, 1992
Aka:	?Scalenodontoides plemmyridon Hopson JA, 1984
Place:	Annapolis Formation & Wolfville Formation, Fundy Group, Nova Scotia
Country:	Canada
Age:	Carnian, Upper Triassic
Remarks:	Specimens held at the Peabody Museum, Yale, under the name of S. plemmyridon.
References:	Sues, Hopson & Shubin (1992), Affinities of ?Scalenodontoides plemmyridon Hopson, 1984 (Synapsida: Cynodontia) from the Upper Triassic of Nova Scotia. J. of Vertebrate Paleontology, Vol. 12(2), p.133-141.
	Hopson (1984), Late Triassic traversodont cynodonts from Nova Scotia and southern Africa. Palaeont. afr. 25, p.181-201.

Links: Centre Consolidated School, Lunenburg, Nova Scotia http://www.ednet.ns.ca/educ/schoolpages/ccs/geology/TriassicPeriod.html Two years of hard work went into this project. The result is a magnificent introduction to geology and paleontology. About the Triassic: George, Terri and Sandy Hrynewich http://www3.ns.sympatico.ca/gphrynew/triassic.htm An interesting review of Nova Scotian fossils by a family of enthusiastic and knowledgeable fossilers. Journal of Vertebrate Paleontology, Vol. 12(2): 133-141 http://www.vertpaleo.org/jvp/contents-12-2.html A synopsis by Hues HD, Hopson JA and Shubin NH.

Genus: Boreogomphodon Sues HD & Olsen PE, 1990 'northern Gomphodon' Remarks: Abundant finds have been reported of this small, late genus, though they are thought mostly to represent juvenile animals. Typically, the upper postcanine teeth are less than 5mm wide.

Species:	Boreogomphodon jeffersoni Sues HD & Olsen PE, 1990
Place:	Richmond Basin, Virginia & Pennsylvania-Newark Supergroup
Country:	USA
Age:	Carnian, Upper Triassic
Remarks:	In the case of one location, Zions View, vertebrate coprolites are commonly found, (ie. fossilized dung). Whether this includes boreogomph droppings is something I don't know. Tomahawk Locality, Turkey Branch Formation The following is based mainly upon my reading of Sues, Olsen & Kroehler, 1994, (see Bibliography for details). The Richmond Basin, central Virginia, is coal country. Reports of fossils date back to the early nineteenth century, (p.163), when fish remains turned up in the pits. These are mostly incomplete and disarticulated remains of the same genus, Dictyopyge. In 1981, Olsen found that a 15-20cm thick layer of the Turkey Branch Formation yielded vertebrate remains. This is at a site called the Tomahawk Locality, named after Tomahawk Creek, (p.161). There were plenty of fishy bits in that there mudstone, but there were also land critters; four-legged ones. They're generally represented by isolated bones and teeth. These black fossils tend to be well preserved, though fragile, (p.164). There are often tiny fractures and, as the surrounding mudstone matrix dries out, there's a goodly chance of disintegration. Glues are applied on site to get around this problem. The eucynodonts in the fauna comprise Boreogomphodon, (this entry), and Microconodon, a poorly known animal with very mammal-like teeth. Further Mesozoic site summaries can be found at Localities. Boreogomphodon The most abundant remains of identifiable tetrapod belong to Boreogomphodon. "It is represented by three excellently preserved partial skulls (including one with a natural endocast), scattered remains of a fragmentary skeleton, a number of isolated dentaries, maxillae, and premaxillae, and many isolated teeth. In addition, a few isolated cynodont limb-bones (humerus, femur) may prove referable to this taxon." The authors observe that the structure of the upper postcanines is closest to Traversodon stahleckeri and Luangwa drysdalli. They've got widened crowns, three main cusps at the back and a large basin at the front. There's one accessory cusp too, as well as a posterior cingulum, (a kind of shelf). The lower postcanines have three cusps at the front whereas most traversodontids make do with two. This is a feature in common with Arctotrversodon from Nova Scotia, (and an undescribed colleague from southern). Details of the upper and lower jaws, and the fact that the tooth rows are somewhat inset: "provide suggestive evidence for a flexible cheek and a buccal oral vestibule, as inferred for other traversodont cynodonts (Hopson, 1984)." The young ones Because of the small size, the relative paucity of postcanines (four or five), the shortness of the snout and the depth of the dentary, most jaws are probably from juveniles, (p.164-165). This interpretation is further supported by the presence of apparently 'sectorial' teeth, (slicers), which several upper jaws have at the rear of the tooth row. This is a feature known from juvenile gomphodonts; eg. Diademodon and Massetognathus. Holotype The type fossil is a left maxilla with three postcanine teeth in situ, (Sues & Olsen 1990, p.1022). This is affectionately named USNM 437632, and resides in the collection of The National Museum of Natural History. The species name honours the first person to describe fossil vertebrates from Virginia. This was a fellow called Thomas Jefferson, who also had interests outside of paleontology.
Reference:	Sues & Olsen (1990), Triassic vertebrates of Gondwanan aspect from the Richmond basin of Virginia. Science 249, p.1020-1023.

Links: Newark Photo Album http://rainbow.ldgo.columbia.edu/courses/v1001/newarkalb.html Photos of a number of fossils, including Boreogomphodon. Biotic Provinciality of the Late Triassic Equatorial Humid Zone http://gsa.confex.com/gsa/2001SE/finalprogram/abstract_4542.htm Some observations on the well preserved tetrapod assemblages of Virginia and North Carolina.

Genus: Colbertosaurus 'Colbert's reptile'

Species:	Colbertosaurus muralis
Place:	Mendoza
Country:	Argentina
Age:	Middle Triassic
Remarks:	A cast specimen is included in the AMNH collection, New York. Further information welcome.
Reference:

Genus: Dadadon Flynn JJ et al, 2000 'Dada's tooth' Remarks: Dada is a nickname of Francois d'Assise Randriamanendika. It was selected in recognition of his help in the field, (Flynn et al 2000, p.423). For significant reasons which I nevertheless don't feel compelled to elaborate upon, the authors didn't select a 'generic' name for either Dadadon or Menadon. Rather, they authored a praenomen in each case. The nomenclature appears to be traditionally binomial in line with the publishing requirements, but appearances can be deceptive.

Species:	Dadadon isaloi Flynn JJ, Parrish JM, Rakotosamimanana B, Ranivoharimanana L, Simpson WF & Wyss AR, 2000
Place:	Isalo II fauna, Morondava Basin
Country:	Madagascar
Age:	Ladinian or Carnian, Middle or Upper Triassic
Remarks:	The following is based upon my reading of Flynn et al. This paternal sounding genus is known from two specimens, with the type fossil being a partial skull preserving the complete set of right upper teeth, (excepting for the first incisor), and left postcanines, (p.423). The total length would have been about 13 centimetres. The presence of four incisors is an old-fashioned touch shared with Gomphodontosuchus. The number had been reduced to three in several roughly contemporary forms, (Menadon and Exaeretodon). An unusual feature of the skull is a rounded, suborbital process, (p.424), which is also a characteristic of the subsequently described Santacruzodon. Dental formula The skull is both weathered and somewhat crushed but the fossil took care of its teeth. They're in good condition. (The animal was neglectful of its lower jaw.) The known formula per side is: (uppers): four incisors, one canine and nine to eleven postcanines. Teeth The incisors are simple cones with faint scratches, (p.425). Canines are fairly small and have no serrations on the front or rear edges. There are vertical scratches and a small diastema between this tooth and the first postcanine. This is mainly the result of the canine being in a more lateral position. Postcanines grow progressivley larger and wider along the row from front to back. The first three are considerably smaller and less complex than the following ones. The first is triangular from the occlusal view, while the second and third have more similarity with the rest. There's a strong crest crossing the crown at the back, two internal cusps and some basining. A well developed cingulum is present on the lateral side. A weak crest goes across the crowns at the front. Looking for relatives Subsequent research suggests Santacruzodon is reasonably closely related, but that genus wasn't established when Flynn & Co published in 2000. The authors found some close similarities with Massetognathus, especially specimens originally assigned to M. teruggii. One clear difference is that Dada was less keen on postcanines, (nine to eleven). The skulls were generally of a similar size but the only Argentinian individual with as few as eleven upper postcanines was a bit of a tiddler; nine centimetres. Skulls with a length of thirteen centimetres favour twelve to fifteen of these teeth. Holotype The holotype, Ua-10605, and a second specimen are in the collection of the University of Antananarivo. The 'specific' name refers to the informal Isalo "Group" of rock strata. The authors offer a definition of Traversodontidae as being: "the clade consisting of all eucynodont species sharing a more recent common ancestor with Exaeretodon than with Probainognathus or Mammalia", (p.765). This definition is not generally followed by other researchers, and nor is it followed in this directory.
Reference:	Flynn et al (2000), New traversodontids (Synapsida: Eucynodontia) from the Triassic of Madagascar. Journal of Vertebrate Paleontology: 20 (3), p.422-427.

Link: The Field Museum, Chicago http://www.fmnh.org/research_collections/research_feature.htm Various finds from Madagascar.

Isalo II, Madagascar, ?Carnian The following is based upon my reading of Flynn et al, 2000. Much of western Madagascar is made up of twwo basins; the Mahajanga to the north and the Morondava in the south. These contain rock exposures of fossil bearing rock from various ages, some of which form the Malagasy "Karroo"; a comparison to the vertebrate rich strata of southern Africa. In terms of fossils, Madagascar has a lot of catching up to do. However, it wins hands down for the Middle Jurassic. The real Karoo stops playing during the Lower Jurassic. Triassic eucynodonts Prior to the year 2000, the Malagasy inventory for published Triassic cynodonts amounted to less than one. Flynn and colleagues raised the count to two and promised more. At least four are present in the fauna, and three are traversodonts, (p.422). Isalo II The rock exposures continue into the southeast of the island in the neighbouring Morondava Basin. An informal term, the Isalo "Group", refers to stata in both Basins. The ages are presently not well understood, but the picture is becoming clearer. The sequence is widespread and divided into three rough subunits: Isalo I, II and III. The upper reaches of Isalo III have been pinned down to the Bathonian, Middle Jurassic. Vertebrates include a small sauropod called Lapparentosaurus and the much, much smaller mammal, Ambondro. The traversodonts come from near the foot of Isalo II, and must be older. They were found to the east of Sakaraha in the Morondava Basin. Previously, the only fossils found in the area were wood and fragments of a rhynchosaur called Isalorhynchus. The new locality is more generous. Uncertain times Several factors contribute to the lack of clarity concerning ages. Marine sediments with convenient index taxa are lacking, as are deposits of datable volcanic debris. Additionally, the vertevrates which have been found aren't shared with faunas from further afield. This may indicate a high degree of endemic evolution. Alternatively, the fossils could date from an otherwise poorly represented slice of two. Both possibilities might even apply. Age of Isala II The vertebrates so far recovered suggest an age of Ladinian or Carnian; either side of the Middle - Upper Triassic transition. Remains include sphenodontids and prosauropods, and these groups haven't been recorded elsewhere prior to the Carnian, (p.426). They coexisted with rhynchosaurs and dicynodonts. With one possible and outrageous exception, (there appears to have been a ludicrously late dicynodont in the Lower Cretaceous of Australia), those lines aren't known beyond the Carnian. Consequently, that's the most probable age. The travies are a bit more basal than Carnian forms from South America. This could be because they're somewhat earlier, (possibly upper Ladinian). However, that's not necessarily the case. Basal lineages can survive for longer than more derived ones. The euro-travies, such as Habayia, are more primitive and later than their Malagasy relatives. With regard to the age, from Flynn et al, 1999, p.764: "Collectively, this evidence (Fig. 2) suggests that either a Middle Triassic or early Carnian (rather than middle or late Carnian) age for the fauna is likely, and the basal Isalo II beds may represent a stratigraphic interval not well sampled previously by a continental tetrapod assemblage." Further Mesozoic site summaries can be found at Localities. These include Triassic traces, the Morondava Basin - an even more informal approach to the same locations. Meet the eucynodonts of Isalo II (at least four taxa) Eucynodont: one undescribed taxon. Traversodontidae (three taxa) Dadadon isaloi; Menadon besairei; one undescribed taxon.

Genus: Exaeretodon Cabrera, 1943 Aka: Proexaeretodon Bonaparte JF, 1963 'before Exaeretodon', Theropsis Remarks: This genus grew to a length of up to 1.8m. (A specimen pictured in Kemp, 2005 has a head and body length of 1.35m, p.69.) Various institutions have fossils; eg. Museo Argentino de Ciencias Naturales, Buenos Aires; Harvard University; Museu de Ciênces e Technologia, Porto Alegre; Universidad Nacional de Tucumán; and the Universidad Nacional de San Juan. Abundant fossils have been recovered from Argentina. These include complete skulls, postcranial material and even a mould of the internal skull, (Abdala, Barberena & Dornelles, 2002). This genus had a particularly well-developed method of grinding. This was achieved through the presence of a diastema between the canine and the postcanines and specialisations of the tooth rows, (Kemp 2005, p.69). Theropsis lanceolata is a perennial garden plant better known as the false lupine. The flowers are yellow. (NB: The same name can be used legitimately for a plant and an animal; eg. Cannabis Blyth, 1850 is a bird. If offered one to smoke, just say no.)

Links: Valdosta State University's Virtual Museum of Fossils http://gatito.valdosta.edu/fossil_pages/fossils_tri/t60.html Fine photos of an Exaeretodon skull from Argentina, (cast specimen). Lou Tremblay's Dino Adventures http://www.pinkyweb.com/tremblay/argentina2001.html A fine gallery of images from Argentina, October 2001 http://www.pinkyweb.com/tremblay/cynodontinfo.gif Lou Tremblay gives us the lowdown on Exaeretodon. A concise summary of the genus.

Species:	Exaeretodon frenguelli Cabrera, 1943
Aka:	Proexaeretodon vincei Bonaparte, 1963
Place:	Ischigualasto Formation & Rio Grande do Sul
Country:	Argentina & Brazil
Age:	Carnian (early), Upper Triassic
Remarks:	Abdala et al, 2002 report comparisons involving E. frenguelli fossils. One of the specimens concerned, (PVL 2565), is the holotype of P. vincei, (p.322). I've taken this as an indication that it's a junior synonym. This specimen is in the collection of the Universidad Nacional de Tucumán, Argentina. The accompanying Table 2 gives the basal length of the skull for 23 specimens, (excluding E. riograndensis). This figure ranges from 15 - to an estimated 40cm. The longest non-estimated example is 38cm. Ten individuals are 30cm or more, whilst the average length is about 28cm.
Reference:	Cabrera (1943), El primer hallazgo de terapsidos en la Argentine, Notas del Museo de La Plata, Paleontology, 55, p.317-331.

Links: Adrian Giacchino & Yamila Gurovich, Net Paleo: Vertebrados triasicos http://www.paleonet.com.ar/divulga/ischigualasto/ischigualasto.html An attractively illustrated item from Argentina, (Spanish). The University of Texas CT Facility The University of Texas CT Facility: Exaeretodon Photographic images of a juvenile skull. "Most researchers accept an alternative view, that Exaeretodon is a close relative of mammals." Interesting.

Species:	Exaeretodon argentinus Bonaparte JF, 1962
Place:	Ischigualasto Formation, San Juan Province
Country:	Argentina
Age:	Carnian (early), Upper Triassic
Remarks:	It's taken about seven years but, thanks to Neal, some information has arrived on this species. It gets a mention in an abstract by Jun Liu; 2007, The taxonomy of the traversodontid cynodonts Exaeretodon and Ischignathus, Rev. bras. paleontol., 10(2), p.133-136. The abstract is all I happen to have. There it says that this specific name is considered to be valid for Argentinian fossils. That's not much information, I'll grant you, but it's good to know this long extinct travie is indeed still alive, if you see what I mean. I was afraid the species might have been extincted into synonymity. Tain't so. Actually, there's a bit more. Originally, the material involved was referred to E. frenguelli. Liu reaches the conclusion that Ischignathus is a junior synonym of E. argentinus. Before reacting to that suggestion by rearranging the bookkeeping of this directory, I think best wait on the off-chance that the relevant paper might turn up.
Reference:

Species:	Exaeretodon major (von Huene, 1935-42) Barberena MC, 1974
Aka:	Traversodon? major von Huene, 1935-42
Place:	Santa Maria Formation, Rio Grande do Sul
Country:	Brazil
Age:	Ladinian, Middle Triassic
Remarks:	This species is represented by a toothless and fragmentary maxilla, (Abdala, Barberena & Dornelles, 2002). Despite the absence of dentition, (p.320) seven alveoli are angled similarly to the condition in E. frenguelli. In times of size, the only known specimen is close to the adult representatives of E. riograndensis. Although such characters are not definitely diagnostic for traversodontid species, the clear difference in dimensions between the front and rear alveoli lead the authors to favour the retention of this as a distinct species for the while. However, confirmation of its status can only come from more specimens. I wonder if this fossil might be connected with Massetognathus major.
Reference:	von Huene (1935-42), Die fosilen Reptilien des südamerikanischen Gondwanalandes, CH Beck, München, 332pp.
	Barberena (1974), Contribuicao des cinodontes gonfodontes (Cynodontia-Tritylodontoidea) do Brasil. Ministério de Educacao e Cultura, Universidade Federal do Rio Grande do Sol. Tese Livre Docente, 194pp.

Species:	Exaeretodon riograndensis Abdala F, Barberena MC & Dornelles J, 2002
Place:	Santa Maria Formation, Rio Grande do Sul
Country:	Brazil
Age:	Carnian or Norian (lower), Upper Triassic
Remarks:	The following is based upon my reading of Abdala et al, 2002. The description involves three specimens which were recovered in 1985 in the Candelaria District, (p.313). The holotype is particularly well preserved, and provides previously unknown information for the genus. The authors conclude that Tritylodontidae is more closely related with Mammaliaformes, than it is with Traversodontidae. In comparison to E. frenguelli, some of the skull bones of this species are extended: "the posterodorsal projection of the jugal behind of the posteroventral; prefrontal relatively wide", (p.320). A feature called the prootic is equipped with a series of crests. These may have provided attachment surfaces for the posterior pterygoid muscle. It also exhibits less variability in the number of postcanine teeth. The skulls of the three specimens range in length from 17cm, (probably a juvenile), up to 26cm. The holotype is 22cm, (p.322). Teeth The largest individual has an upper dental formula of three incisors, one canine and six postcanines, one of which was in the process of erupting, (5 + 1). As is to be expected, these animals didn't have diphyodont tooth replacement. The lower formula is only known for the juvenile. It's the same as with its upper dentition; 3, 1 and 6 (plus one erupting, ie. 6 + 1), (p.320). The upper incisors are directed somewhat forwards. The largest are the first two, and there are wear facets on the lingual edges. The smaller third incisor has no such wear facets. The upper canine is large and forward pointing. Enamel is restricted to its labial surface. At the margin of the enamel is a wear facet. In the juvenile, the lower canine is relatively small. When the mouth was closed, it fitted in behind of its upper colleague. Regarding the upper postcanines: "In unworn teeth, the labial posterior accessory cusp is completely isolated from the crest that descends from the main labial cusp... Thus, two basins characterize the unworn postcanine: the principal one formed by the anterior accessory labial cusp and the main labial cusp, and a posterior small one formed only by the posterior accessory labial cusp", (p.320) (I've omitted a reference from the quotation. Should the language have left anyone somewhat disorientated, it concerns one main and two accessory cusps on the external part of the crown -labial. The language of teeth can be a bit of a mouthful.) Anyway, after slight wear, that posterior accessory cusp becomes incorporated into the principal basin. Taking into account both these specimens and a larger number of E. frenguellii (plural), the authors are able to reach some conclusions regarding postcanine replacement. Smaller (and presumably younger) individuals are generally somewhat toothier. In the Argentinien species, probable juveniles exhibit eight or nine postcanines per side, (p.322). Larger specimens usually have six. However, some bigger individuals actually have more, (10 + 1 erupting in one case). This probably shows unusual instances of tooth retention. For example, the front postcanines of MCP 1522 PV are very worn, and seem to have passed their lose-by date. Tooth replacement and an eccentricity This tendency to lose front postcanines in favour of replacements at the rear, is the general pattern for both traversodontids and Sinoconodon. However, the number of teeth decreasing with age is more eccentric. In both Scalenodon and Massetognathus, larger individuals have usually invested in more teeth, (p.323). Exaeretodon exhibits a trend not previously recorded in any non-mammalian eucynodont. Holotype The hololtype, MCP PV 1522, is a skull in the collection of the Museu de Ciências e Tecnologia, Pôrto Alegre, Brazil. The species name refers to Rio Grande do Sul, (p.314). The suggested age is based upon the presence of Proterochampsa fossils.
Reference:	Abdala et al (2002), A new species of the traversodontid cynodont Exaeretodon from the Santa Maria Formation (Middle/Late Triassic) of Southern Brazil: J. of Vert. Paleont., 22 (2), p.313-325.

Link: Journal of Vertebrate Paleontology, 22(2) http://www.vertpaleo.org/jvp/22-313-325.html The abstract.

Species:	Exaeretodon statisticae Chatterjee, 1982
Place:	lower Maleri Formation, Andhra Pradesh
Country:	India
Age:	Carnian, Upper Triassic
Remarks:	This may be a junior synonym of E. frenguelli.
Reference:	Chatterjee (1982), A new cynodont reptile from the Triassic of India, Journal of Paleontology, 56 (19), p.203-214.

Genus: Gomphodontosuchus von Huene FF, 1928 'Gomphodont crocodile'

Species:	Gomphodontosuchus brasiliensis von Huene FF, 1928
Place:	Hyperodapedon Assemblage Zone, Santa Maria Formation, Rio Grande do Sul
Country:	Brazil
Age:	Carnian, Upper Triassic
Remarks:	A specimen of this creature is housed in the paleontological collection of the University of Tübingen. Somewhat unusually, it doesn't seem to have a catalogue number, (Abdala & Ribeiro 2003, p.543). This was the first traversodontid ever described.
Reference:	von Huene (1928), Ein Cynodontier aus der Trias Brasiliens. Centralblatt für Minerologie, Geologie und Paläontologie (B), p.251-270.

Genus: Gornogomphodon Renesto S & Lucas SG, 2009 'Gorno gomphodon' Ramrks: This genus isn't a traversodontid. Rather, it's perhaps a gomphodont that refuses to enter into any established family, and I can't think of anywhere in particular to place it. Here seems as good as anywhere. The generic name refers to the Formation that yielded Italy's first reported gomphodont eucynodont remains.

Species:	Gornogomphodon caffii Renesto S & Lucas SG, 2009
Place:	Zambla Alta, Gorno Formation
Country:	Italy
Age:	middle Carnian, Upper Triassic
Remarks:	The following is based upon my reading of Renesto & Lucas, 2009. I'm going to commence with a confession that may outrage the legions of Italian fans of Triassic cynodonts. Personally, I'm left unconvinced that Gornogomphodon, Italy's first offered contribution to Upper Triassic eucynodonts, actually is a cynodont. I haven't got any better explanation to offer for the identity of this small fragment of upper jaw with three teeth, accept the authors know far more than me, and respect the hard work and reasoning that's behind their conclusions. But I'm unconvinced. Hopefully, one day, a more revealing specimen will turn up. The difficulty I have is that this fossil certainly merits the term 'aberrant', if it be a cynodont. Actually, it seems presently aberrant whatever its identity! Perhaps it's an aberrant something else with broad, unusually built teeth and strange roots. The snag is I've got no idea as to what else. A gomphodont cynodont is the best supported explanation I presently know of. The fragment in question has a length of a bit over 1.5cm, and came from Lombardy's Gorno Formation about 50km south of the Swiss border (p.357). The teeth are notably wide, can be termed tri-cusped and have cingula both at the front and the rear. That, say the authors: "... supports their identification as postcanines of a "eucynodontian"" and, in that instance, they place explanation remarks around the term. I'm not sure why, and suspect they have no particular significance. However, the morphology on display rules out an assignment to any established family of gomphodonts. The age is middle Carnian, an impoverished sector of time for Euro-cynodonts, and the deposits built up in shallow lagoons. Informally seen The postcanine teeth vaguely remind me of a baby lying in a crib or a drunk left paralytic in a canoe. The buccal side hosts a large blunt cusp, and the lingual side has a much lower cusp. The third cusp, if that word is appropriate, is the low bulge of the sleeping body stretched out between both those points. Cingula run round the entire crown so as to form the walls of the crib or canoe, and a thin trench runs between them and the snoozing body inside. It's not a traversodontid. A travie upper postcanine needs to be equipped with a basin to complement the build of the lower partner, and no such thing happens to be present. It's also not a trirachodontid. Its buccal cusp is too tall, and the cingula lack cuspules. Indeed, the single buccal cusp is presently singular for a gomphodont, should Gorno have been one. That cusp 'should' adjoin with a second cusp there, but it has no such partner. The complete encirclement of the crown by cingula is a further singularity for such critters, and the roots are described as being: "... similar to the protothecodont condition", something that's presently sadly beyond my appreciation. The authors assure the reader that the roots are, however divided. The most complete of the three teeth, the rear one, has height of 5.6mm, a width of 9.3 and a length of 4.5 (p.358). To the root of the matter The roots are described as: "... deep and seem to be divided and open, so that the inner alveolar bone that surrounds the tooth seems to bulge into the middle of the pulp cavity. It is difficult to interpret this structure..." I find the discussion on this hard to appreciate due to my insufficient knowledge. As there's no "alveolar space" involved, this apparently doesn't appear to be a: "... mammal-like multiple rooted tooth..." Anyway, they consider the roots to be more likely "protothecodont", a situation common for early reptiles and, probably, basal therapsids. If so, as cynodonts certainly can't be considered as anything like basal therapsids, this apparent condition for a gomphodont eucynodont would greatly puzzles me. What is Gorno? The authors naturally considered other possibilities for the affinities of this fossil, but felt able to rule them out. They were initially unsure as to whether it was from a tetrapod or part of a fish. The crown shape and the arrangement of the teeth isn't unlike pycnodont fish. However, enquiries revealed the walls of the "roots" (the authors use quotation marks at that point for some reason) are too thick. Their width should be less than, and not about the same as that of the crown itself. The teeth in this jaw are arranged in a somewhat curved row, and that's unknown for pycnodonts. Another point is that pycnodont teeth with distinct cusps, as opposed to rills and other forms of ornamentation, happen to be unknown (p.359). Consequently, the fish were found innocent of producing this jaw fragment. They then switch their enquiries to tetrapods with relatively complex postcanine crowns that could be termed molar-like. Procolophonids, a group of small reptiles that appear to have been largely plant-eaters during the Triassic, have teeth that don't sport three cusp or cingular ridges, and they lack distinct necks. Another group of herbivores, trilophosaurids, have wide, tri-cusped crowns. However, those cusps are found on a raised loph, and the central cusp is the tallest. "Not guilty!" is the verdict for both cases. Gomphodont? The width is obviously in line with gomphodont postcanines although, as stated, I'd personally have problems should the same be claimed for the roots. (No such claim is explicitly stated.) Similarities are apparent with trirachodontids, diademodontids and traversodontids. In the first instance, there's a line of three main cusps running across the crown, cingula at both the front and rear, and no shearing surfaces on the cusp sides. However, in order to be qualified as a trira, the cingula should have cuspules, and the central cusp ought to be the tallest, not the buccal one. Comparable diademodontid postcanines are near round in outline, and have a wide basin on the lingual side. Gornogomphodon uppers aren't and don't respectively. By no stretch of the imagination, not even after a plentiful intake of LSD and magic mushrooms, can Gorno be successfully accused of having an occlusal basin on the central area of its upper postcanine. There are very narrow gullies between the central cusp and the cingula, but nothing that could be remotely termed a basin. That's enough to rule out traversodontids as well. Odd bod Those of you who'd like to qualify as aberrations will, as Gorno demonstrates, have to do some of you own things in unforeseen ways. For example, Gorno insists on having but a solitary buccal cusp that happens to be the tallest on the crown. Odd. The cingula succeed in completely encompassing the crown. Odd. The ratio of width to length is extreme even by the standards of extremist gomphodont upper postcanines. Somewhat odd. Although I very possibly wouldn't have understood all too much, I can't help but feel that the roots merited discussion during considerations on features not found on "other" gomphodonts. Be that as it m, the new genus receives a referral to Eucynodontia family uncertain. Holotype MCSNB 5863 is a fragment of upper jaw with three teeth residing in the collection of the Museo Civico di Scienze Naturali 'E. Caffi', Bergamo. The specific name honours Enrico Caffi, who was the first director of that institution.
Reference:	Renesto & Lucas (2009), Cynodont teeth from the Carnian (Late Triassic) of northern Italy, Acta Palaeontologica Polonica, 54(2), p.357-360.

Link: APP 54(2), Renesto S & Lucas SO http://www.app.pan.pl/archive/published/app54/app54-357.pdf The original description in pdf format is presently freely accessible on-line.

Genus: Habayia Godefroit P, 1999 'from Habay' Remarks: The generic name is derived from Habay-la-Vieille in southern Belgium.

Species:	Habayia halbardieri Godefroit P, 1999
Place:	Habay-la-Vielle
Country:	Belgium
Age:	Rhaetian, Upper Triassic
Remarks:	The following is based upon my reading of Godefroit, 1999. The Upper Triassic opened with an impressive diversity of middle to large herbivores. Among them were armoured aetosaurs, rhynchosaurs, dicynodonts and traversodonts. Remains of the latter have been found in most continents. The family was radiating nicely, and bodysizes ranged up to a couple of metres. They could've gone on for fame and fortune. However, things worked out differently during the Carnian. All four of these groups were totally or nearly swept away. From the last stage of the Triassic, the Rhaetian, the only known traversodont fossils are a few teeth from Europe, (p.385). The global supply of identified Habayia remains consists of one tooth, (unless anything else has turned up in the last few years). It's tiny. In terms of quantity and size, this is typical enough of these presumably last traversodonts. The mini-herbivore eucynodont niches were dominated, (if that's an appropriate word), by haramiyids. A further group, the trtiylodontids, were more substantial animals; lengths of up to perhaps a metre. Dinosaurs called prosauropods were starring as megaherbivores, and ornithischians were coming on-line. A motorway to the past The sparse remains of Habayia and its cohabitees of ancient Belgium came to light courtesy of a road construction project. This work revealed outcrops of sediments dating from the Upper Triassic and Lower Jurassic. A group of amateur paleontologists took the chance to systematically sample Rhaetian bonebeds between Habay-la-Vieille and Fouches. Numerous remains were obtained from one site called HLV. A sedimentological survey revealed four sequences in the strata. The earliest deposition was at a stage of oceanic regression. During the other three, the tide was very much in. HLV has three main bonebeds, and they were laid down by tidal action. Palynological research dates then as Rhaetian. Of particular interest were the eucynodont fossils. As well as this genus, teeth have been found for: Thomasia woutersi, Mojo usuratus, Lepagia gaumensis, Gaumia longiradicata, G. ? incisa, Pseudotriconodon and Microscalenodon nanus. Although all these animals were small, seven genera is a fair level of diversity. The body of Habayia It was presumably very small. The tooth It's a right upper postcanine. The length of the crown is 1.05 and the width 1.3mm, (p.387). When seen from the occlusal perspective, the outline in vaguely pentangular. There are two main cusps; a lingual and a labial one, and they're connected by a ridge running across the middle of the tooth. This produces a large basin at the front and a smaller one behind. Also present is a mesiolabial accessory cusp. Apart from the minute size, it's typically traversodont. However, there are distinctions in detail which justified a new genus. Affinities While the morphology of Microscalenodon may be simiplified, it's primitive for this genus, Maubeugia and Rosieria: "the mesiolabial cusp is not displaced medially and there is no trace of a mesiolingual accessory cusp", (p.392). While perhaps sounding superficially paradoxical, these most recent of traversodonts were old-fashioned when alive. Upper Triassic Europe The paper contains a genus count for non-mammalian cynodonts, (so it's not limited to traversodonts), from Norian-Rhaetian locations in Europe: Saint-Nicolas-de-Port, France 11 genera; 2. Habay-la-Vieille, Belgium 5; Württemburg, Germany at least 4; Hallau, Switzerland more than 3; Medernach, Luxembourg 3; Syren, Luxembourg 2. Less determinate cynodont teeth have also been found at two further French localities; Varangéville and Saint-Germain-les-Arlay. Traces of early mammals are found in many of these locations as well. There were small herbivores, (should haramiyids be considered as mammals), and mini-predators. The environment of Western Europe clearly supported a wide variety of small eucynodonts. However, comparing this record with faunas from elsewhere necessitates caution. The European sites have been systematically scoured for microvertebrate remains. Less intensive efforts would've missed at least most of the fossils. "Recent discoveries of abundant macroscopic plant material in southern Belgium indicates a dense vegetation in the vicinity of Habay-la-Vieille, providing the basic trophic support for such a variety of primary and secondary consumers." Dense plant growth provided a diversity of opportunities, but there was a further environmental peculiarity at the time. The Rhaetian was characterized by oceanic transgressions. This may be reflected in the fossil record of tetrapods, (four-legged land critters and their descendants). Medernach (Luxembourg) is relatively early for the locations mentioned above, (middle Norian). Large amphibian tetrapods are plentiful. More recent are Saint-Nicholas-de-Port and Varangéville. Those fossils were laid down during periods of transgression. While large animals are still well represented, their micro-cohabitees appear to have been far more diverse than previously. Habay-la-Vieille and Syren are typically Rhaetian. Large vertebrates are scarce, and small animals were relatively varied. More sea meant less terrestrial and freshwater habitats, and this added up to bad news for big amphibians and crocodile-style phytosaurs. However, it didn't inhibit small vertebrates. As the available land was also chopped up into an archipelago of islands, isolation may have fuelled diversification. Holotype The holotype, IRSNB R203, resides at the Royal Belgian Institute of Natural Sciences, Brussels. The specific name honours Mr B Halbardier, the mayor of Habay, who helped smooth the way for new excavations. Salut. A further traversodont tooth from the same site was also described, and referred to as genus and species indeterminate.
Reference:	Godefroit P (1999) New traversodontid (Therapsida: Cynodontia) teeth from the Upper Triassic of Habay-la-Vieille (southern Belgium). Paläontologische Zeitschrift 73 (3/4), p.385-394.

Genus: Ischignathus Bonaparte JF, 1963 'Ischi jaw' Remarks: Along with Exaeretodon, this was one of the largest herbivores of the time.

Species:	Ischignathus sudamericanus Bonparte JF, 1963
Place:	Ischigualasto Formation
Country:	Argentina
Age:	Carnian, Upper Triassic
Remarks:	The genus is based on one jaw, which is similar to Exaerotodon, but larger. There is the possibility that it might be from a bigger representative of that taxon.
Reference:	Bonaparte (1963), Descripción de Ischignathus sudamericanus n.gen. n.sp., nuevo cinodonte gonfodonte del Triásuci Medio superior de San Juan, Argentina. Acta geológica lilloana 4, p.111-128, Tucuman.

Links: INTELRADIO, Formación Ischigualasto http://www.intelradio.com.ar/dinosaurios/ischigualasto.htm An Argentinean report on fossils, including Ischignathus, (Spanish). FAUNA OF ISCHIGUALASTO 230 MILLION YEARS AGO http://www.ischigualasto.org/ENGLISH/30fosiles/index.htm A handy and informative review of fossils from Ischigualasto, courtesy of del Museo de Ciencias Naturales de la Universidad Nacional de San Juan. It's fairly reader friendly.

Genus: Luangwa Brink, 1963 'Luangwa' (a geographical term) Aka: Scalenodon (partly) Reamarks: Abdala & Smith, 2009, report on the presence of this genus in the Omingonde Formation of Namibia. Several specimens were identified, but not referred to any particular species (p.2). The deposits concerned date from the Anisian, and so are much the same age as the Luangwan Luangwa. The fossils were part of a snout, three skulls and some bits of skeleton. Distortion and poor preservation imposed limitations on what could be usefully described, and presumably precluded a specific allocation. The skull lengths ranged from 14-14.5cm (estimated) to 20.4cm. That latter individual, previously mis-informed that it was called Scalenodon angustifrons, is the largest member of Luangwa so far found. As could be expected for a larger and, therefore, older animal, the proportional ratios of some skull parts differ from those of other specimens of the genus. Additionally, another fossil from there was referred to as some traversodontid or other. It probably doesn't belong to Luangwa. Although it evaded further determination, it nevertheless indicates two travies were parts of the local wildlife.

Species:	Luangwa drysdalli Brink AS, 1963
Aka:	Scalenodon drysdalli
Place:	upper Ntawere Formation, Luangwa Valley
Country:	Zambia
Age:	Anisian, Middle Triassic
Remarks:	Specimens reside at The Bernard Price Institute, Johannesburg and in the collection of The Oxford University Museum of Natural History. A distinct genus Battail referred this species to Scalenodon in 1991, (Abdala & Sa-Teixeira 2004, p.17), but subsequent studies have maintained a seperate genus. Examples include that paper and Rubidge & Sidor, 2001. The snout and temporal area on the skull of L. are short, and the animal was blessed with large orbits. The lower jaw had a high coronoid process. Jaws of Scalenodon aren't well-preserved, but the snout and temporal region are proportionately longer and the eye holes smaller. Luangwa was also less keen on postcanine teeth. Although similarly sized to S. angustifrons, the L. type fossil was content with seven as opposed to nine. A larger individual did have nine. Seven to nine is pretty much the range for L. sudamericana as well, although the two known specimens of that species are somewhat smaller. There are enough distinctions to justify a separate genus. Size One Oxford specimen is known as TSK 121, and this includes at least a pelvis and the femur. Blob, 2001 offers a bodymass estimate of 8.8 kilos for this individual, (p.20). That's something like equivalent to a cat. The basal length of the skull from the holotype is 13.6cm (Abdala & Sa-Teixeira 2005, p.18). Holotype The holotype, BP/1/3731, is a reasonably complete skull in the Bernard Price Institute, Johannesburg. The specific name honours geologist Alan R Drysdall. (With thanks to Steve for the confirmation.)
Reference:	Brink (1963), Two cynodonts from the Ntawere Formation in the Luangwa valley of Northern Rhodesia, Palaeontologia africana 8, p.77-96.

Link: http://www.users.bigpond.com/delacroix/data35.html This creature was featured on a Zambian postage stamp in 1973.

Species:	Luangwa sudamericana Abdala F & Sa-Teixeira AM, 2004
Place:	Santa Maria Formation, Rio Grande do Sul
Country:	Brazil
Age:	Middle Triassic
Remarks:	The following is based upon my reading of Abdala & Sa-Teixeira AM, 2004. (Thanks are due to Fernando for sending the paper.) An African far south of the Amazon Luangwa sudamericana has more similarities with relatives from Africa than it does with others from its own continent, and this may reflect the age (p.11). Anatomically, the species is allied with L. drysdalli from Zambia and Scalenodon angustifrons from Tanzania; animals from the Anisian stage of the Triassic, rather than the later Ladinian. Upper postcanine teeth are oval in outline and have a cingulum towards the front of the labial side. A cingulum at the back is a generic feature of Luangwa, as is an anterior cingulum on the lowers. The skull has a short snout, very large orbits and the temporal area is also short. These are further generic characteristics. What's the time? The Santa Maria Formation spans the Ladinian-Carnian transition of the Middle - Upper Triassic. However, unless this animal was a living fossil, some earlier, Anisian deposits could also be involved. The formation is mischievous as exposures occur in outcrops. The German connection Research in this formation dates back to 1928 when a German paleontologist, Friedrich von Huene, spent six months working in a number of localities. As a result of his efforts, a significant collection of local fossils ended up in Tübingen, southwest Germany. His discoveries included the first traversodont ever described, Gomphodontosuchus. Further Tübingen Santa Marian eucynodonts are Traversodon and Chiniquodon. The African connection Transatlantic travel wasn't an option in the Triassic, as the Atlantic Ocean didn't exist. The similarities among the fossil wildlife reflect the fact that South America and Africa were neighbouring parts of Pangaea. Although Luangwa is the first traversodont common to both continents, a globetrotting eucynodont carnivore called Cynognathus had already achieved this distinction. A specimen was arrested in the Puesto Viejo Formation of Argentina in the 1960s. Further African influences aren't surprising. Luangwa of Argentina The new species is based on two specimens, and these were lurking in the collections of two institutions in Porto Alegre. Precisely where they originally came from is unknown, although UFRGS 267PV was obtained somewhere on the district border between Candelaria and Vera Cruz, (p.12). This is a partial skull with a fragment of lower left jaw. The holotype, MCP 3167PV, is a reasonably complete snout with part of the front cheek area, (including the rim of the right orbit). Specific distinctions In contrast to its African relatives, (L. drysdalli and Scalenodon angustifrons), the upper postcanines of the new species have an extra, well-defined cusp on the labial crest (p.13). A further accessory cusp occurs on the front of the labial crest, but this has been identified in at least one specimen of L. drysdalli, in contrast to the type fossil. Getting a head As one individual provided a snoutless skull and a second chipped in with a snout, the partnership allows a reasonably complete picture of the face. L. sudamericana was a short-nosed traversodont with large eye holes, (p.14). The temporal area is unusually short as well, and it features a pineal foramen in the broad parietal crest (p.15). Teeth (p.16) Dental formulae in non-mammalian eucynodonts vary in different individuals, and even in the jaws of one animal. The type fossil of the species has per side: (uppers): four incisors, one canine and nine - ten postcanines; (lowers): three, one and eight - nine respectively. (The second specimen has seven to eight upper postcanines). Uppers The incisors are mostly broken, but what remains there are suggest they were rounded to ellipsoid in the occlusal perspective. Part of the third crown is available and canine-like. The labial face has thick enamel, while the internal side seems to be free of the stuff. The leading edge is smooth but its posterior counterpart carries faint denticulations. A short diastema separates the incisors and canine. That tooth is ellipsoid and enamelled on both faces. Strong serrations are present both fore and aft. Upper postcanines are oval in outline and they became bigger along the row. Most are heavily eroded, although the rear ones are less damaged. A main cusp occurs towards the back on the labial side. This forms a crest with smaller cusps in front and behind. A further ridge runs across the crown featuring a large cusp (labial), a wide central one and a small, lingual companion. This crest is situated towards the back of the tooth. Lowers The incisors are roughly circular in occlusal outline, and thick enamel is probably limited to the lateral face. A short diastema comes between these teeth and the ovoid, somewhat bigger canine. Both faces are enamelled. Whether denticulations were present is unclear. No diastema is found between the canine and postcanines. As with the uppers, most crowns are heavily eroded. The final ones have crests on the front of them. These involve a wide cusp on the lingual side and a taller, narrower labial cusp. Old gomphodont fashions When seen from the occlusal perspective, the upper postcanines of both species of Luangwa and Scalenodon angustifrons are roughly ovoid in outline, (p.18). This resembles the shape in old timers such as Diademodon and Cricodon. Derived traversodonts, such as Massetognathus and Exaeretodon, favoured a rectangular or trapezoid shape. How big were these animals? Table 1 presents some measurements of Luangwa specimens, with two representatives of both species. This is the holotype and a friend from each. The full data set contains eleven various aspects and is in millimetres. I'm going to restrict myself to three characteristics and use centimetres. An asterix donates an estimate. L. sudamericana (type) L. sudamericana (friend) L. drysdalli (type) L. drysdalli (friend) Basal length of the skull __ *9.4 13.6 __ Snout length 5.02 *3.2 6.0 __ Length of the row of upper postcanines 3.36 3.37 3.86 4.11 Source: Abdala & Sa-Teixeira 2004, Table 1, p.18. Taken at face value, that suggests the African species was larger. However, as only four faces are involved, the small sample size might not necessarily be an accurate reflection. A question of age (p.20) For various reasons, the age of the Santa Maria Formation has been difficult to pin down. In general, it ranges from the Ladinian to the upper Carnian. The presence of Luangwa suggests the possibility of the base extending further back to the Anisian stage. This has been suspected before. A basal rhynchosaur turned up at a location called Mariante, and this prompted a Late Anisian / Early Ladinian assignment. That fauna has subsequently been referred to as Ladinian, (p.21). Another possibility would be a chronological extension for the genus. Clarity would come with further knowledge about the components of the faunas which yielded these Brazilian specimens. Unfortunately, as the exact locations aren't known, that's presently trickier than tricky. However, as prospecting work is very much continuing, should clearly Anisian sites be out there, they'll be found sooner or later. Holotype The type fossil, MCP 3167PV, is a resident of the Museo de Ciências e Technologia in Porto Alegre. The reason behind the specific name is as blindingly obvious as it appears.
Reference:	Abdala & Sa-Teixeira (2004), A traversodontid cynodont of African affinity in the South American Triassic, Palaeontologia africana 40, p.11-22.

Genus: Massetognathus Romer AS, 1967 'chewer jaw' Aka: Megagomphodon 'large gomphodon' Romer, 1972 Remarks: The canines of this genus were relatively small, (Abdala & Ribeiro 2003, p.530). Institutions with specimens include, Harvard University, the Universidad Nacional de La Rioja and the Universidade Federal do Rio Grande do Sol.

Reassigned species: M. major Romer, 1972 see M. pascuali; M. teruggii Romer, 1967 see M. pascuali

Link: Valdosta State University's Virtual Museum of Fossils http://gatito.valdosta.edu/fossil_pages/fossils_tri/t61.html They don't only offer pictures of skulls in Georgia. Here's a well preserved left leg.

Species:	Massetognathus pascuali Romer AS, 1967
Aka:	M. major Romer 1972; M. teruggi; M. teruggii Romer 1967; Megagomphodon oligodens Romer, 1972
Place:	Los Chanares Formation
Country:	Argentina
Age:	Ladinian, Middle Triassic
Remarks:	The creature reached about 50cm long and weighed perhaps 10 kg when alive. It's the most common land vertebrate in this fauna. The following is based upon my reading of Abdala & Giannini, 2000. In 1967, Romer AS established Massetognathus and recognized two species from the Chanares Formation. M. pascuali was described as a smaller animal with a skull length of about 8-9.5 cm. Some differences in detail of the skull were also cited, and emphasis was placed upon the presence of 12 upper and 11 lower postcanines per side, (p.501). M. teruggii was established for individuals with skull lengths of around 11.5-14cm. The opposite differentiations of construction applied as compared to M. pascuali, and the postcanine count was reported as 13-15 uppers, and 11-13 lowers. At the time, Romer had five specimens to work with. In 1972 he added M. major. This consisted of a single skull without lower jaw. It was a bit over 20cm in length. The orbits extended further forward; the antorbital region was narrow; and there were 13 postcanines with less variability amongst the rear ones. It also exhibited a more weakly developed labial platform lateral to these teeth which produced a narrower snout. For good measure, a separate genus, Megagomphodon, was proposed for a pair of skulls held to be closely related; skull length 18cm, 17-18 smallish postcanines, and a more slender skull. One genus, different ages Given that all this material comes from one limited area, he suspected this apparent abundance of similar species might have been overdoing things, and all specimens could belong to a single taxon in varying stages of growth. Subsequently, other authors proposed synonymity; a possibility which Abdala & Giannini set out to test. They had 31 skulls available and took 17 different measurements into account, (p.502). The results, (p.503), suggest all the specimens are representatives of the same species. For example: "Thus, the facial, orbital, and braincase components grow slower than, or at the same rate as the skull as a whole, whereas the temporal region grows faster both longitudinally and transversely." Different bits of the skull grow at differing speeds. If you've got a newly born baby available, have a look. The eyes are relatively large, as are the orbits. Advantages of hindsight Of course Abdala & Giannini enjoyed several advantages over Romer. They had more carefully researched literature to refer to, (eg by Romer AS), and many more specimens. That undoubtedly helped because: "The existence of intermediate specimens filling the proposed size gaps erected by Romer...", (p.504). Size is an unreliable characteristic for classification. Their results indicate that the snout in juveniles is already relatively large. It then develops more slowly than the rest of the skull. This is in contrast to the temporal area. As individuals grow older, the number of postcanines generally increases. The teeth are also proportionately smaller, seeing as more have to be accommodated in the row. These kinds of details lead to the conclusion, (p.505); "Therefore, we recognize Massetognathus pascuali as the only valid species for Chanares gomphodonts, and consider M. teruggi... [not my typo!] ...M. major, and Megagomphodon oligodens as junior synonyms. This is coincident with Battail's (1991) opinion, but we have now provided a clear argument to justify this viewpoint." Holotypes All holotypes are in the collection of the Universidad Nacional de la Rioja, (p.502). They are: Massetognathus pascuali PULR 10; M. major PULR 11; M. teruggii PULR 13; and Megagomphodon Oligodens PULP without/number.
References:	Romer (1967), The Chanares (Argentina) Triassic reptile fauna. III. Two new gomphodonts, Massetognathus pascuali and M. terugii. Breviora 264, p.1-25.
	Romer (1972), The Chanares (Argentina) Triassic reptile fauna. XVII. The Chanares gomphodonts. Breviora 396, p.1-9.

Link: Dinosaurios: Massetognathus http://www.conce.plaza.cl/~dinos/links/dinos/masseto.htm An illustrated file card, (Spanish).

Species:	Massetognathus ochagaviae Barberena MC, 1981
Place:	Dinodontosaurus Assemblage Zone, Santa Maria Formation, Rio Grande do Sul
Country:	Brazil
Age:	Ladinian, Middle Triassic
Remarks:
Reference:	Uma nova espécie de Massetognathus (Massetognathus ochagaviae, sp. nov.) de Formacao Santa Maria, Triássico do Rio Grande do Sul, Pesquisas 14, p.181-195.

Los Chanares Formation, Middle Triassic, Northwest Argentina The following is based upon my reading of Rogers et al, 2001. This Formation formed in a basin which was fed by sediment from the surrounding highlands. Rivers brought and dumped material on the floodplain, (p.461). At some stages, deposits also built up on the beds of lakes. This sort of process is usual enough. The special aspect of Chanares is the quality of the terrestrial vertebrate fossils, and much of the responsibility has been blamed on volcanic activity. Large amounts of ash were strewn across the landscape. Some was presumably delivered by the rivers, but a more simply rained down from the sky. The generally hospitable environment occasionally had ferocious temper tantrums. The star locality is the name-giving Los Chanares. Remains of over a hundred bodies of land critters have so far been identified. Most are eucynodonts, but they're accompanied by dicynodont cousins and archosaurs. Some of these animals have been preserved in direct association with one another, and this applies for individuals who wouldn't generally have socialised; carnivores and their foodstuff. One poorly represented element is large-sized animals, which are found in other contemporary assemblages. There seems to have been a preservation bias involved, as there's little reason to suppose none lived here. The arrangement of bones suggests the possibility that many of the corpses were dumped on a shoreline. By changing water courses, volcanic activity can result in dramatic floods. In any case, the remains are of such quality that they can't have been lying around in the open for all too long. After burial in a tomb rich in volcanic ash, carbonate started to accumulate around them, and they ended up girdled in nodules of stone known as concretions. A brief look at the wildlife The Chanares Formation provides one of the best glimpses of land animals available from the Ladinian, though Brazil would probably claim the world championship. The latter stages of the Middle Triassic are of particular interest, (as can validly be said of any time)! This was just prior to some dramatic radiations ensured the spread of dinosaurs, extremely mammal-like non-mammalian synapsids and basal mammals. In terms of diversity, the local winners are archosauriforms of one kind or another, (p.462). These include proterochampids. Although crocodile-like, their lineage emerged before the divergence of croc and dino lines. Also represented are early crocodile relatives, (eg. the bipedal Gracilisuchus), and close dino relations such as Lagerpeton. In numbers of specimens, the assemblage is dominated by us eucynodonts. Herbivores are most frequent, but meat-eaters of relevance to mammalian origins are present as well. Vegetarian dicynodonts kept them company. The Formation A belt of rock exposures are located south of the town of La Rioja, and the classic Los Chanares location is one of the most northerly of them. Below can be found the Tarjados Formation while the Los Rastros is higher. Vertebrates have also been recovered a few kilometres to the south near Canon del Gualo, but not in great numbers. Other exposures have so far been even less generous. Remains are heavily concentrated in just the one main cemetery. Agua de la Pena Group This is a collection of rock formations, and not a local band. Collectively, it witnesses the time from the lower Ladinian, (about 234 million years ago), until fairly late in the Norian, (older than 210 million years ago). The Chanares forms the base and Los Rastros separates it from the Ischigualastro Formation. At the apex comes Los Colorados, (Fig. 2, p.463). A radiometric date is available from low in the Ischigualastro, and that states the underlying strata must predate 228 million years. While rock samples will sometimes be kind enough to supply information on their absolute age, it's also possible to ask the fossils. In this case, the Chanares fauna has yielded no dinosaurs, aetosaurs or ornithosuchids. It was presumably a bit too old. Comparing the inhabitants with Middle Triassic Europeans doesn't reveal much usable information, as the two areas had distinctive populations. The dominant land vertebrates in Europe were rhynchosaurs, lepidosauromorphs and temnospondyl amphibians. Similarities do occur with the Dinodontosaurus Assemblage of the Santa Maria Formation in Rio Grande do Sul, (p.464). Shared genera are Dinodontosaurus, Massetognathus, Probelesodon (aka Chiniquodon) and Chanaresuchus. As the Brazilian assemblage is probably Ladinian, then so is the Chanares. Los Chanares locality Back in the 1960s, the Formation was recognised as having two informal units at the main cemetery. The lower one has a tinge of blue and provides the fossils. The upper is nearer to a virginal white and fossil free. The lower unit can be followed through much of the area, and varies in thickness from 20 to 35 metres. Starting at about ten metres from the base, the fine-grained sediments also contain brown nodules with diameters of up to two metres. Some of these formed around a core provided by vertebrate remains, and that's the source of most fossils. Origins of the Formations The underlying Tarjados was predominantly deposited by relatively fast flowing water action, whereas Los Rastros is more associated with lakes, (p.468), and at least six episodes were involved, (p.469). These are separated by narrower layers of fluvial deposit. There are also many strata containing volcanic ash, (p.470). The Chanares Formation is the middle of the sandwich, and its ingredients have a less clear recipe. There's plenty of volcanic debris around, but it's possible that much was transported in by rather grubby streams. This would better account for the similarly delivered silt and sand. The strata with fossils were probably built up on a floodplain, and that explains the presence of a healthy variety of terrestrial vertebrates. Original interpretations favoured much more direct volcanic excitement. A further aspect is that at least some deposition may have been on the shoreline of a lake, (p.471). Counting animals At the time of publication, sixteen terrestrial vertebrate taxa had been identified; ten archosauriforms, three eucynodonts and three dicynodonts, (p.472). That could give a false impression as to which group dominated. A headcount of individuals in three collections presents an entirely different picture; eucynodonts 81%, dicynodonts 7%, archosauriforms 12%. About 58% if all identified specimens are provided by the genus Massetognathus. Aquatic and amphibious animals haven't been found. Even the crocs are landlubbers. Quality of preservation Remains of tetrapods are restricted to the lower unit, and a large majority have come from within nodules. The specimens occur in concentrations separated by expanses of fossil-free rock. This contains nodules as well, but they lack animals as the cores. Los Chanares main cemetery covers only about 75 square metres. Outside of this very restricted area, less than 5% of the exposed concretions contain fossils. The preservation quality is generally excellent, with bones showing minimal evidence of weathering. Fossils are found independently of nodules, but these are rare and not concentrated together, (p.473). These remains are often from relatively large animals; skulls of big archosaurs, dicynodonts and Massetognathus (including the holotype of "Megagomphodon oligodens"). Such isolated material is usually more distorted and weathered. They also have a different colour, which reflects differing processes of fossilisation; pinky grey rather than dark brown. Finds from nodules These range from bits of bone to groups of partially articulated skeletons. Bones are frequently intact or, if broken, cleanly snapped. Lighter elements, such as ribs and vertebrae, are reasonably common. These body parts are more liable to be transported off by natural forces, than is the case for substantial skulls and long bones. A sample of 23 prepared nodules were categorised into three groups. Eight contained remains of one individual, generally Massetognathus. These ran from a few bones to near complete skeletons. Eight more contained parts of more than one individual of the same species. All bar one were again provided by the usual traversodont, while the exception consisted of partial Chanaresuchus skeletons. In on case, there was a collection of four Massetognathus skulls along with various bones. Some force of nature had left the skulls strongly aligned, as if their owners had all been gazing in the same direction. However, being dead, they wouldn't have been in a fit state to spectate anything. The other seven nodules contained representatives of at least two taxa. These included various combinations of archosaurs, dicynodonts and herbivorous and meat-eating eucynodonts. Four more nodules had something else as their core; droppings. In three instances, these coprolites were in association with skeletal material, so they didn't come out the rear ends of vegetarians. In one case, the remains included ribs, vertebrae and armour from a small archosaur. Sensitive scientific instruments - sledge hammers After patient searching, extracting fossils can be delicate work. That doesn't apply much to this location, where sledge hammers are wielded with enthusiasm for opening nodules. (It's not destructive as long as you can recognise where to aim.) The restricted nature of the area is also useful to researchers. They know what they're potentially looking for and where it might be. The processes which produced these fossils were generally very helpful. However, there are bound to be some disadvantages. Exposed nodules of rock often fall down and gravity can send them skipping along slopes. Getting information on precise stratigraphic positions and original associations may be rendered impossible. While the concentration makes the search easier, the preserved picture could be distorted. It might be a highly selective insight into the past, rather than a well-balanced cross-section of the former fauna, (p.474). Single deaths and collective die-ins There appear to have been several ways of getting fossilized at Los Chanares. Isolated specimens were probable products of what's termed 'attritional mortality'; victims of everyday deaths -illness, starvation, murder. The aforementioned small archosaur had that route chosen on its behalf. These remains generally lay around until natural processes provided a burial. Some were barely disturbed while others were broken to pieces. So it goes. But this normal state of affairs can't readily account for the collections of corpses. Four Massetognathus wouldn't routinely die in a close huddle. This would be better explained by an unusual, mass-mortality, killer mechanism. That could also account for the peculiar concentrations of remains in compact groups. Apart from the absence or presence of fossils, there's no obvious difference in the composition of the rock, so the depositional history must have been broadly similar, (p.476). Evidence for mass slaughter The fauna is rich in individuals as well as diverse in taxa. The tally from the three collections runs to over one hundred animals. As these are only the prepared fossils, this is a minimum figure. Two of the collections have unprepared material, and it would be very strange if many more individuals aren't found in that. The minimum inventory involves 94 eucynodonts: 68 Massetognathus, 18 Probainognathus and 8 Probelesodon (aka Chiniquodon). The presence of a number of individuals in single nodules is suspicious. In four cases, carnivorous archosaurs have been found alongside with edible eucynodonts. They wouldn't have been hanging out together in normal circumstances. A third suspicious factor is the lack of weathering, and the fact that so much skeletal material is still articulated. These characters rule out prolonged expose to the elements and scavengers, and anything beyond gentle transportation. A psychological profile of a killer The killer selected members of various taxa and animals of differing ages. It killed carnivores and prey, and young and old. What's not known is whether the bodies came together shortly before or following death, though the second option strikes me as more probable. Larger animals may have been less at risk, as their remains are more common elsewhere in the Formation. However, it's also possible this 'immunity' is an illusion caused by post-mortem sorting. The alignment of remains in nodules offers further evidence. This pattern is what could result from carcases being washed ashore. It's also been noted that one fairly complete Massetognathus has a "passive" death stance, which is consistent with a period of involuntary floatation, (p.477). A relevant parallel might have been provided by events at Smithers Lake in southeastern Texas, which formed the basis of a study published in 1989. A severe storm played havoc with the local wildlife, and dead alligators, fish, birds and plant material ended up assembled together along a 1,300 metre band of the southern shore. As the water delivered the remains from much the same direction, many were aligned in a similar fashion. Versatile volcanoes Volcanic activity can kill in various ways. A place which comes immediately to mind is Pompeii, where many deaths have been blamed on suffocation under blankets of ash. Another possibility may be demonstrated by the Eocene fossil site of Messel, Germany, involved releases of poisonous gas. Volcanoes can also kill with lava flows, but they have less direct means in the arsenal as well. Rivers can be forced into changes of direction by blockages. Long established lakes can be breached or new ones formed. This can lead to flooding, mudslides and rockfalls. Such occurrences can be sudden and catastrophic. As there's plenty of evidence of volcanic activity in Los Chanares Formation, it's an obvious suspect. Being suspicious isn't necessarily a sign of guilt In order to avoid Lynch law, other potential suspects have been interviewed; eg. drought, climate and burial alive. Most are untenable when checked against the available evidence. The Mongolian Upper Cretaceous offers an example of death by vast quantities of sand at Ukhaa Tolgod. This resulted in lots of articulated skeletons from a great variety of animals, and the entombing sandstone shows precious little order. Virtually all individuals from the Los Chanares locality are disarticulated to some extent, and that suggests some degree of exposure after death. Furthermore, the similar alignment of individuals isn't what would be expected from living animals, should they have suddenly been hit by, for example, a mudslide. Many would've tried to struggle free. Another suggestion involved entrapment in burrows. Creatures of a similar size to Massetognathus have been recovered from Karoo, South Africa, and they date from the Lower Triassic and even Upper Permian. Such a cause would explain the lack of large victims. As the ground featured plenty of ash, it was presumably soft enough for tunnelling. One unsatisfactory aspect is the lack of burrow casts but, as such structures require freaky circumstances in order to be preserved, this absence isn't necessarily significant. It would've been very odd if none of the local population had indulged in plenty of digging. The tunnellers of Karoo included both dicynodonts and cynodonts; groups represented here. However, Los Chanares fossils are relatively flat, and this suggests they formed while lying on the ground rather than below it in chambers. And again, the similar alignment of individuals isn't suggestive of such an entombment. There may not be a smoking gun, but the finger of guilt points strongly at death by volcanic activity. Further Mesozoic site summaries can be found at Localities. Meet the eucynodonts of Los Chanares Formation (three genera, three species) Traversodontidae (one genus, one species) Massetognathus passcuali Probainognathia (two genera, two species) Probainognathus jenseni; Probelesodon (Chiniquodon) lewisi. Los Chanares Formation support cast featured: Dicynodonts: Dinodontosaurus platiceps, D. platygnathus, D. brevirostris. Archosauriforms: Luperosuchus fractus, Lagerpeton chanarensis, Chanaresuchus bonapartei, Gualosuchus reigi, Gracilisuchus stipanicicorum, Lewisuchus admixtus, Marasuchus lilloenses, Pseudolagosuchus major, Tropidosuchus romeri, Tarjadia ruthae.

Genus: Maubeugia Godefroit P & Battail B, 1997 'for Maubeug' (as in Dr P-L Maubeug)

Species:	Maubeugia lotharingica Godefroit P & Battail B, 1997
Place:	Saint-Nicolas-de-Point
Country:	France
Age:	Norian (late)-Rhaetian (early), Upper Triassic
Remarks:	One of the late European midgets, based on a left upper postcanine, (<1mm wide). The species name is Latin, 'from Lorraine'. The authors note similarities to the dentition of Boreogomphodon and Scalenodon, (though they don't specify which S. species). The holotype is affectionately known as IRSNB R172. It resides in the Institut Royal des Sciences Naturelles de Belgique, Brussels, (Godefroit & Battail 1997, p.604).
Reference:	Godefroit P & Battail B (1997): Late Triassic cynodonts from Saint-Nicolas-de- Port (north-eastern France). Geodiversitas 19(3), p.567-631.

Genus: Menadon Flynn JJ, Parrish JM, Rakotosamimanana B, Ranivoharimanana L, Simpson WF & Wyss AR, 2000 'red tooth' Remarks: 'Mena' is Malagasy for 'red'. This refers to both Madagascar, (the red island), and a helpful villager who led the authors to the site and assisted with the field work. He doesn't have a surname, (p.422). As for Dadadon the paper doesn't propose a generic name. This is referred to as a praenomen. However, for convenience and uniformity I'm choosing to disregard that, which doesn't mean it's not significant.

Species:	Menadon besairei Flynn JJ, Parrish JM, Rakotosamimanana B, Ranivoharimanana L, Simpson WF & Wyss AR, 2000
Place:	Isalo II fauna, Morondava Basin
Country:	Madagascar
Age:	Ladinian or Carnian, Middle or Upper Triassic
Remarks:	The following is based upon my reading of Flynn et al, 2000. The authors found that Menadon has similarities with Exaeretodon, Scalenodontoides and Gomphodontosuchus (p.422), which are relatively derived members of the family. The cheek bone (jugal) has a suborbital process, the zygomatic arch is deep, (that's also a reference to the cheek area and not a landmark in Athens), and there are a number of dental specializations characteristic of traversodontids. Dental formula The right side of the skull, (which is nearly 16cm in length), is fairly complete along with the lower jaw, (p.423). Consequently, both the upper and lower probable tooth formulae could be assessed. There are three incisors and a canine per side in both cases. The traditional traversodontid condition was four incisors. As for the postcanines, the lower count is given as six to seven, whilst the upper situation is eight. This is less than for most family members. Lower teeth The lower jaw is both relatively deep and short with robust incisors, which are angled forwards at about 45°, (with the gradient estimated by me from the photo). Wear has made some details of the lower postcanine crowns unclear. Uppers The third upper incisor (I3) is unusually large and canine-like. The next tooth in the series is probably the canine, and it seems to be accompanied by an older colleague in the process of being replaced. Both upper and lower canines feature five serrations on the front and rear edges. A wide diastema is between the anterior teeth and the postcanine brigade. The first three postcanines are rather pathetic. Number one is triangular and separated from the rest by a short gap. Two and three are highly worn, oval pegs. The other five postcanines are wider than long. They're roughly rectangular from the occlusal perspective, with curvature of the front lingual corner and deep basins; similar to the choppers of Gomphodontosuchus. There's a high labial crest and a traditional travie posterior ridge. This has only two cusps. A central one isn't present. Relationships A phylogenetic analysis suggests Menadon is a relatively primitive member of a clade which led to Exaeretodon and Scalenodontoides, (p.425). Along with Gomphodontosuchus, these animals have a common ancestor not shared with Luangwa or Massetognathus. Holotype The holotype is a skull and lower jaw known as Ua-10601. It's in the collection of the University of Antananarivo, Madagascar. The 'specific' name honours the geologist Henri Besairies, (p.422).
Reference:	Flynn et al (2000), New traversodontids (Synapsida: Eucynodontia) from the Triassic of Madagascar. Journal of Vertebrate Paleontology: 20 (3), p.422-427.

Link: Santa Barbara Museum of Nat Hist. Bulletin, (July / August 2002) http://www.sbnature.org/members/bulletin/bull222.pdf This edition contains a photo of what seems to be a reasonably complete skull, (p. 3).

Genus: Microscalenodon Hahn, Lepage & Wouters, 1988 'small Scalenodon'

Species:	Microscalenodon nanus Hahn, Lepage & Wouters, 1988
Place:	Habay-la-Vielle
Country:	Belgium
Age:	Rhaetian, Upper Triassic
Remarks:	Based on two teeth, an upper and a lower postcanine. May also be known from Saint-Nicolas-de-Point in France. This seems to be the smallest and most recent representative of this family. Seriously micro.
Reference:	Traversodontiden-Zähne (Cynodontia) aus der Ober-Trias von Gaume (Süd Belgien). Bull. Inst. R. Sci. Nat. Belb. Sci. Terre. 58, p.177-186.

Genus: Nanogomphodon Hopson JA & Sues HD, 2006 'dwarf molar tooth' Remarks: The single known small tooth is somewhat molar-like rather than literally a molar. The crown construction is relatively complex.

Species:	Nanogomphodon wildi Hopson JA & Sues HD, 2006
Place:	Baden-Württemberg
Country:	Germany
Age:	Ladinian, Middle Triassic
Remarks:	Helpful paleo pixies have managed to smuggle a copy of the description into my paws and, and thanks are due. My understanding of that paper naturally provides the basis of this article. Nanogomphodon is the first traversodontid to be described from the Middle Triassic of Europe (p.124). As with later possible European travies, assuming they've been correctly assigned to the family, this genus is also presently known only from isolated teeth or, more accurately here, a tooth. In any case, it's from a small member of the increasing tribe of northern hemisphere travies. These authors express doubts about the travie credentials of Upper Triassic Euro-fossils previously referred to the family from France, Belgium and Luxembourg; eg. Maubeugia. The resemblance, they say, is vague. However, they don't suggest an alternative interpretation, and don't go as far as stating such taxa aren't travies. 'dwarf molar tooth' A single lower travie tooth was found by Hans Hagdorn in 1980. He's an amateur fossil hunter, and took the opportunity to collect some sediment that was conveniently exposed by a house construction project in Michelbach an der Bilz. This was undertaken under the auspices of Stuttgart's Museum für Naturkunde. These efforts added remains for a new tetrapod fauna of this age to the not overly rich German record. This postcanine features a tri-cusped ridge running across the front of the crown and, to the rear, a long basined area with a further ridge behind it. That supports three small cusps, and the basin is closed lingually by a further clear ridge. The crown also has a frontal cingulum featuring five small cusps. This enormous relic of travie glories has a maximum length of 2.02mm and a maximum width of 1.54. Clear is that the otherwise absent owner would have faced difficulties with any application for joining the Megaherbivore Association. Dentists will be delighted (or upset, in the case of greedy ones) to learn the tooth is in good condition, with only the end of the root missing. Somewhat atypical is the tri-cusped ridge at the front. More usual for travies would be only two cusps (p.126). The buccalmost of the trio is in all ways the largest, while the other two cusps are near similar with one another. There's a slight hollow apparent on the cingulum at the front of the tooth, and this presumably was an area of contact with the preceding tooth. Wear facets on the crown are in line with travie expectations, and that subject gets more intimately discussed in the paper. (It's somewhat too complex for me to attempt to chew over here!) Affinities This fossil shares similarities with Boreogomphodon from Virginia. Boreo also has a tri-cusped ridge at the front of lower cheek teeth. The size is modest for travie standards, but it's similar to that known from isolated teeth of the NAm genus; presumably juvenile ones. Whether this Nano was also a kid or simply a small animal, packed with lustful adult interests, can be answered with a definite perhaps. There's presently no way of knowing. A feature not held in common with Broeo is the proportionate length and width. This crown is longer than the norm, but the same can be said of South America's Santacruzodon. Holotype SMNS 51962 resides in the Staatliches Museum für Naturkunde, Stuttgart. The specific name is for Rupert Wild, a further Stuttgart cynodont specimen who has contributed much to the growth of knowledge of Triassic tetrapods. A further testament towards that happens to be the last most recently updated entry on these directions; an insect-killing cynodont named Pseudotriconodon wildi. More thanks are due to DinoTracker for posting notice of the publication on-line.
Reference:	Hopson JA & Sues HD (2006), A traversodont cynodont from the Middle Triassic (Ladinian) of Baden-Württemberg (Germany), Paläontologische Zeitschrift, 80 (2), p.124-129.

Genus: Pascualgnathus Bonaparte JF, 1966 'Pascual's jaw'

Species:	Pascualgnathus polanski Bonaparte JF, 1966
Place:	Puesto Viejo Formation
Country:	Argentina
Age:	Anisian, Middle Triassic
Remarks:	This genus could belong to Diademodontidae. If not, it's a small and early representative of Traversodontidae, which had an unusually large canine, (Abdala & Ribeiro 2002, p.530). Some material is in the collection of the Museo de La Plata, whilst other specimens can be contacted at the Universidad Nacional de Tucumán. Both institutions are in Argentina. (The species name was forwarded by Vince Ward.)
Reference:	Bonaparte (1966), Una nueva 'fauna' triásica de Argentina (Therapside: Cynodontia Dicynodontia). Consideraciones filgenéticas y paleobiogeográficas. Ameginiana 4, p.243-296.

Genus: Plinthogomphodon Sues HD, Olsen PE & Carter JG, 1999 'Brick tooth' Remarks: As the first page of the publication has just become visible on-line, I now know the meaning of the name! Apparently, the mudstone from the locality has been a popular choice for making bricks. That, rather than any particular bricky characteristic of the tooth, is the reasoning.

Species:	Plinthogomphodon hepetairus Sues HD, Olsen PE & Carter JG, 1999
Place:	Deep River Basin, North Carolina, Newark Supergroup
Country:	USA
Age:	Carnian - Norian, Upper Triassic
Remarks:	The only part of the description I've seen is page 351. That contains little information on the specimen itself, although there is mention of the obliteration of most of the formerly functional teeth. A full copy would be welcome. Based on a partial snout which appears to have been swallowed by a large, rauisuchian predator about 220 Ma. Abdala & Ribeiro, 2003 (p.540) mentions this material is from a small, juvenile individual. Holotype (UNC) no. 15576 is two pieces of a snout housed in the collection of the University of North Carolina, Chapel Hill. The specific name translates as 'companion of reptiles'. This is because it was find lying around close to bits of archosaurs; namely a nasty rauischian and a sphenosuchian.
Reference:	Sues et al (1999), A Late Triassic traversodontid cynodont from the Newark Supergroup of North Carolina, Journal of Vertebrate Paleontology, 19, p.351-354.

Links: The Journal of Vertebrate Paleontology 19(2): 351 http://www.vertpaleo.org/jvp/19-351-354.html The synopsis. Science Daily, (University of North Carolina at Chapel Hill) http://www.sciencedaily.com/releases/1999/08/990818071044.htm How the fossil came to light.

Genus: Protuberum Reichel M, Schultz CL & Soares MB, 2009 'protuberances' Remarks: The ribs and illia of this somewhat eccecentric travie have lots of protuberances on them, thus the generic name.

Species:	Protuberum cabralensis Reichel et al, 2009
Place:	Dinodontosaurus Assemblage Zone, Santa Maria Formation, Rio Grande do Sul
Country:	Brazil
Age:	Ladinian, Middle Triassic
Remarks:	As I've got a pdf of the paper, more information will be added at some time or other. Holotype MGB 368/100 is composed of a skull (minus some teeth and the lower jaw), and a lucky dip selection of skeletal bones. It struts its stuff in the Museu Guido Borgomanero, Mata. The specific name refers to the municipality of Novo Cabrais, home of the type fossil's erstwhile grave.
Reference:	Reichel et al (2009), A new traversodontid cynodont (Therapsida, Eucynodontia) from the Middle Triassic Santa Maria Formation of Rio Grande do Sul, Brazil, Palaeontology, 52(1), p.229-250.

Genus: Rosieria Godefroit P & Battail B, 1997 'from Rosières' (Rosières-aux-Salines) Remarks: "... the upper postcanines of Rosieria can easily be derived from those of Traversodon..." (Godefroit & Battail, 1997, p. 612.)

Species:	Rosieria delsatei Godefroit P & Battail B, 1997
Place:	Saint-Nicolas-de-Point
Country:	France
Age:	Norian (late)-Rhaetian (early), Upper Triassic
Remarks:	Another isolated tooth of less than a millimetre. The species is named in honour of Dr D Delsate. A further possible specimen has also been identified. The holotype is IRSNB R173. It works at the Institut Royal des Sciences Naturelles de Belgique, Brussels, (Godefroit & Battail 1997, p.610).
Reference:	Godefroit P & Battail B (1997): Late Triassic cynodonts from Saint-Nicolas-de-Port (north-eastern France). Geodiversitas 19(3), p.567-631.

Link: Geodiversitas 19 (3), p.567-631 http://cimbad.mnhn.fr/publication/geodiv/g97n3a4.html The abstract of the author's article.

Genus: Rusconiodon Bonaparte JF, 1970 Remarks: In a 2009 paper I haven't yet read, Liu & Powell concluded that this genus is a junior synonym of Andescynodon.

Species:	Rusconiodon mignonei Bonaparte JF, 1970
Place:	Cerro de las Cabras Formation (?Rio Mendoza Formation)
Country:	Argentina
Age:	Spathian, Lower Triassic
Remarks:	The same site as Andescynodon, and a similar animal too. Basal and small. According to Kemp, 2005, the transverse ridge of the postcanines is not much more than developed than in Diademodon, which is primitive. The same applies for postcanines, (p.69). Formation One source, I forget which, informed me the locality was in the Rio Mendoza Formation. A more recent one, Abdala & Smith, 2009, cites the Cerro de las Cabras Formation (p. 12). I can't explain the significance, if any, of that apparent descrepency. (This note is identical to part of the entry for the just mentioned Andescynodon. The same situation pertains.)
Reference:	Bonaparte (1970), Annotated list of the South American Triassic tetrapods, Proceedings of the Second Gondwana Symposium, International Union of Geological Sciences Commission on Stratigraphy, CSIR Pretoria, p.665-682.

Genus: Santacruzodon Abdala F & Ribeiro AM, 2003 'Santa Cruz tooth' Remarks: The generic name is derived from the city where the discovery was made, and the Greek for 'tooth'.

Species:	Santacruzodon hopsoni Abdala F & Ribeiro AM, 2003
Place:	Santa Cruz do Sul, Santa Maria Formation, Rio Grande do Sul
Country:	Brazil
Age:	Ladinian, Middle Triassic or Carnian, Upper Triassic
Remarks:	The following is based upon my understanding of Abdala & Ribeiro, 2003. The locality of Santa Cruz do Sul was a recent discovery, (p.529), and the vertebrates found were all eucynodonts. Traversodontids are strongly represented, with the most common being Santacruzodon. A speciality of this animal, an autapomorphy, is a very large cusp on the rear labial side of upper postcanines. It contributes over half the length of the labial crest. A phylogenetic analysis suggested the new genus is most closely allied with Dadadon from Madagascar. Occlusion, the key to travie success Traversodontids were active and agile herbivores, and their lifestyles required fuel. Partly paralleling a trick subsequently used by mammals, they found a way of making their teeth more efficient food processors. Upper and lower postcanines became properly aligned with each other, (p.530). Occlusion and complimentary morphology meant that plants could be exploited both more quickly and thoroughly. As this was the most efficient dentition the world had ever known, the global radiation of travies is none too surprising. A homage and a lament Armed with this sophisticated dental technology, it's easy enough to understand how traversodontids continued to flourish in the Carnian, with some obtaining bodysizes of towards two metres. The group was poised for even greater success. In a fair world, their excellent dentitions would have ensured a starring role among middle to large-sized terrestrial herbivores, and perhaps some would've developed a taste for meat. However, the world ain't fair. Aircraft engineers will assure you that concord was far and away the most sophisticated passenger plane ever to have left the runway, but the future turned out to be jumbo. After the Carnian, jumbo descendants of carnivores were the head gardeners; plateosaur dinos with pathetically simple, disposable teeth. Why travies had little future is a mystery. The reasons for the commercial failure of concord included high fuel consumption, a vast increase in oil prices and relatively low capacity. The Santa Cruz do Sul fauna Four traversodontids were hanging out in this Ladinian locality, three of which were previously unknown. That presumably means a further two taxa await description. The main differences concern the structure of the postcanines. Rare remains of chiniquodontids have also been found. The Santa Cruz tooth The signature of this genus is the unusually large cusp at the back of the labial crest on the upper postcanine. Otherwise, it has a mixture of characteristics shared with various travies. The jugal bone has a ball-shaped suborbital process as known from Dadadon. Flattened incisors are found in Massetognathus pascuali. However, in Santacruzodon, they carry 7-9 marginal cuspules, and that's a feature found in Arctotraversodon. A series of small cusps form a crest at the front of the upper postcanines, (p.533), and this has been observed for 'Scalenodon' attridgi. Skull The head isn't well preserved, and individual bones are indistinct. There's also some crushing. The estimated length is about eight centimetres; a small rabbity size. (I reached that figure with the precision of a workbench measure, and was pleased to find the authors agree.) The length of the upper postcanine row is 2.9cm while the lowers manage less than 2.8. Upper teeth There are four incisors and they're shaped like arrowheads. The outer face is flat while the inner is somewhat convex. A diastema comes between them and a poorly preserved, unimpressive canine. There's no diastema between that tooth and the postcanines. Those teeth vary in structure and numbers. There are seven to ten of them in the available specimens. When seen from the occlusal perspective the front postcanines are generally triangular in outline. Rear ones are wider with a tri-cusped labial crest. A further tri-cusped crest crosses at the back of the crown. The lingual and middle positioned cusps are close with one another, and separated from the labial one by a basin. Eight or nine cuspules form a lower crest at the front of the tooth. Lowers The three incisors are larger than the uppers and something like spoon-shaped, (p.534). The front face is convex while the lingual one is strongly concave. There's a fairly puny canine, with a small diastema between that and the postcanines. These teeth number between nine and eleven. A crest runs across the front of their crowns, and it's composed of a pair of cusps. The taller, labial one is positioned slightly further forwards than its internal colleague. There's a further crest on the lingual side and a cingular crest at the back. This is sometimes of a similar height to the anterior one. Shared traits As mentioned, the ball-shaped process on the jugal is a similarity with Dadadon. The same applies to the differentiation evident in the front and rear upper postcanines, with their triangular or roughly rectangular outlines. (This feature is also known from Gomphodontosuchus.) Both Dada and Santa have large numbers of postcanines. They have further shared features too but, as these occur in earlier travies, they're uninformative, primitive traits; plesiomorphies. The three cusps on the labial side of the upper postcanines is a characteristic in common with Massetognathus. Only two occur in Dadadon. The incisors also have similarities with Masseto, but even more so with Arctotraversodon. An analysis with a proviso A data matrix of 28 characteristics and 15 genera was analysed, ( Diademodon, Trirachodon and 13 traversodontids). As 21 of the traits concerned teeth, the authors advised some caution. The state of preservation of Santacruzodon means that potentially informative skull and jaw characters aren't known for this genus, and couldn't be taken into account. Results According to this analysis, the most basal travies in the sample were Scalenodon angustifrons and a taxon combining Andescynodon and Pascualgnathus, (p.536). Next came a monophyletic clade containing Luangwa and 'Scalenodon' hirschoni. (Note: Abdala & Sa-Teixeiri, 2004 supports a relationship between S. angustifrons and Luangwa with distinctions from 'S.' hirschoni.) A clade of Ladinian and Carnian travies has Traversodon at its base and Dadadon and Santacruzodon in a monophyletic pairing. The remaining sample travies form a more derived clade, (Massetognathus, Gomphodontosuchus, Exaeretodon, Menadon and Scalenodontoides). Holotype The holotype, a fragmentary skull with lower jaws, is known as MCN PV 2768, (p.530). In the company of further specimens, it's in the collection of Museu de Ciências Naturais, Porto Alegre. The specific name is in honour of Dr James Hopson, who has contributed greatly to the knowledge of therapsids. Thanks are due to Stephan Pickering for the notification and abstract.
Reference:	Abdala & Ribeiro (2003), A new traversodontid cynodont from the Santa Maria Formation (Ladinian-Carnian) of southern Brazil, with a phylogenetic analysis of Gondwanan traversodontids. Zoological Journal, 139 (4), p.529-545.

Genus: Scalenodon Crompton, 1955 'uneven tooth' (With thanks to C.V. Vick) Aka: Trirachodon (partly) Remarks: This genus is known from Africa, though there may also be a Russian species. However, at least some of the proposed species do not belong in this taxon. Abdala & Ribeiro, 2003 (p.540) also raise doubts about the chronological uniformity of the Manda Formation. The fauna contains both basal traversodontids (S. angustifrons and others with apparently derived dental features otherwise known from Ladinian-Carnian representatives, ("S." hirschsoni and "S." charigi). In combination with wider evidence from the locality, it's possible that material from more than one age is represented. Blob, 2001 offers a bodymass estimate for one ?Scalenodon sp. specimen in the Natural History Museum, London, (p.20). BMNH R.9391 weighed something like 4.2 kilos, which is a bit less than a middling hare. Remains of this specimen consist of at least a pelvis, femur, tibia and foot.

Reassigned species: S. drysdalli see Luangwa drysdalli

Species:	Scalenodon angustifrons (Parrington FR, 1946) Crompton, 1955
Aka:	Trirachodon angustifrons Parrington, 1946
Place:	Manda Formation
Country:	Tanzania
Age:	Anisian, Middle Triassic
Remarks:	Abdala & Ribeiro, 2003 (p.536) found that this species fits into a clade of basal traversodontids along with Pascualgnathus and Andescynodon. However: "Some caution should be introduced to our analysis and its resulting cladograms," (p.534). The set of comparisons they made focused strongly on teeth. A different or wider set of data could possibly produce differing results. Although not necessarily incompatible with that conclusion, Abdala & Sa-Teixeira, 2004 focused on allegiances with Luangwa. (The two studies had differing objectives.) Postcanines The type fossil preserves alveoli for nine postcanine teeth. Some larger specimens could have found room for eleven. As with the holotype of L. drysdalli, the incisors and canine feature serrated edges, (p.18). The equivalents in 'S.' hirschoni and 'S.' attridgi don't. Holotype The holotype, UMZC T.907, is in the collection of the University Museum of Zoology, Cambridge, (with thanks to Dr Fernando Abdala). This is a partial skull with lower jaw (Abdala & Sa-Teixeira 2004, p.12). The length of the skull is about 13cm. The collection houses at least a dozen specimens, and a cast specimen is at the Peabody Museum, Yale.
References:	Parrington (1946), On the cranial anatomy of cynodonts. Proceedings of the Zoological Society of London, 116, p.181-197.
	Crompton (1955), On some Triassic cynodonts from Tanganyika. Proceedings of the Zoological Society of London, 125, p.617-669.

Species:	"Scalenodon attridgei" Crompton, 1972
Place:	Manda Formation
Country:	Zambia
Age:	Anisian, Middle Triassic
Remarks:	Possibly not part of this genus. At least one specimen is enjoying the hospitality of The Natural History Museum, London. It might be synonymous with "S. charigi", (Abdala & Ribeiro 2003, p.540).
Reference:	Crompton (1972), Bulletin of the British Museum (Natural History) Geology, 21, p.29-71.

Species:	Scalenodon boreus Tatarinov LP, 1973
Place:	Karagachka, Orenburg Province, Southern Urals
Country:	Russia
Age:	Anisian - Ladinian, Middle Triassic
Remarks:	Battail & Surkov, 2000 contains some information on page 115. Remains found so far are not exactly extensive. Actually, they consist of two upper postcanine teeth. The structure's similar to S. angustifrons, but these are smaller and differ in some details. Holotype The type tooth, PIN 2973/1, may be consulted at the Paleontological Institute in Moscow.
Reference:	Tatarinov (1973), Cynodonts of Gondwanan habit in the Middle Triassic of the USSR. Paleontological Journal 2, p.200-205.

Link: Mathematical http://mathematical.com/dinotatarinov.html A list of species described by Dr Tatarinov.

Species:	"Scalenodon charigi" Crompton AW, 1972
Place:	Manda Formation
Country:	Zambia
Age:	Anisian, Middle Triassic
Remarks:	Possibly not part of this genus, (Abdala & Ribeiro 2003, p.540).
Reference:	Crompton (1972), Bulletin of the British Museum (Natural History) Geology, 21, p.29-71.

Species:	"Scalenodon hirschoni" Crompton, 1972
Place:	Manda Formation
Country:	Zambia
Age:	Anisian, Middle Triassic
Remarks:	Abdala & Ribeiro, 2003 (p.536) find this species is more closely related to Luangwa. However, it also has features in common with Gomphodontosuchus and a possible clade containing Menadon, Scalenodontoides and Exaeretodon. In terms of traversodontid systematics, it displays both basal and derived features. (Abdala & Sa-Teixeira, 2004 complicates things by pointing to similarities between S. angustifrons and Luangwa as well.) A study by Hopson & Kitching in 2001 found support for sister-group affinities with Tritylodontidae. In either case, a close relationship with S. augustifons was not recognized. At least one specimen is in The Natural History Museum, London (BMNH 8577).
Reference:	Crompton (1972), Bulletin of the British Museum (Natural History) Geology, 21, p.29-71.

Genus: Scalenodontoides Crompton & Ellenberger, 1957 'Scalenodon form'

Reassigned species: ?S. plemmyridon Hopson, 1984 see Arctotraversodon plemmyridon

Species:	Scalenodontoides macrodontes Crompton & Ellenberger, 1957
Place:	Lower Elliot Formation & ?Molteno Formation
Country:	Lesotho & South Africa
Age:	Norian, Upper Triassic
Remarks:	This seems to be the latest of the large traversodontids. Lucas & Hancox, 2000, (p.6) assigns the Lower Elliot to the Norian. The paper makes no mention of remains in the underlying Molteno Formation, which seems to be Carnian. A complete skull was reported in 1993 by Gow CE & Hancox PJ. Material is in the collection of the Bernard Price Institute, Johannesburg.
Reference:	Crompton & Ellenberger (1957), On a new cynodont from the Molteno Beds and the origin of the tritylodontids. Annals of the South African Museum 44 (1), p.1-13, Cape Town.

Genus: Theropsodon von Huene FF, 1950

Species:	Theropsodon njalilus von Huene FF, 1950
Place:	Ansian, Manda Formation, Ruhuhu Basin
Country:	Tanzania
Age:	Middle Triassic
Remarks:	Based on a complete skull with lower jaw, this fossil is not very well preserved. The holotype is at Tübingen, Germany. (With thanks to Vince Ward and Dr Michael Maisch.)
Reference:	von Huene, FF (1950), Die Theriodontier des ostafrikanischen Ruhuhu-Gebietes in der Tübinger Sammlung. Neues Jahrb. Geol. Pal., Abh. 92, p.47-136, 68 figs, 2 pls. (The theriodonts of the east African Ruhuhu area in the Tübingen collection).

Genus: Traversodon von Huene, 1936

Reassigned species?: T. major von Huene, see Exaeretodon major

Species:	Traversodon stahleckeri von Huene, 1936
Place:	Dinodontosaurus Assemblage Zone, Santa Maria Formation, Rio Grande do Sul
Country:	Brazil & Argentina
Age:	Ladinian, Middle Triassic
Remarks:	The AMNH, New York has a fair few specimens. These were collected by Colbert in Brazil, 1959. Material is also in the collection of the Univeridade Federal do Rio Grande do Sul and the University of Tübingen. Based on guesswork, the holotype is most probably at the last cited institution. Grew to a length of about 50cm.
Reference:	von Huene (1936), Die Fossilien Reptilien des südamerikanischen Gondwanalandes an der Zeitenwende (Denwa-Molteno-Unterkeuper = Ober-Karnisch). Ergebnisse der Sauriergrabungen in Südbrasilien 1928/29. Lieferung 2, p.93-159.

Other Reports: Syren, Luxembourg Link: Dominique Delsate, Espace Paleo 3J http://home.planetinternet.be/~nerodj2/RecapWEB/pagehtm/rhetien.htm A description of Upper Triassic finds in Luxembourg and Lorraine, including traversodontid material, (French). Arizona, USA Indeterminate gomphodont material was collected from the Chinle Formation, Apache County by Hazen in 1946. This is also housed at the Peabody Museum in Yale. A large gomphodont has also been struck at the Dinosaur Hill locality in Petrified Forest National Park, which is the same formation. It gets a mention in Therrien & Fastovsky, 2000 (p.203). Other vertebrates from the site include: Coelophys, a crocodylomorph, a rauischian predator, phytosaurs, aetosaurs, further saurischian and ornithischian dinos and bits of metoposaurs. North Carolina, USA Heckert A, Camp J, Schneider V, Olsen P & Nesbitt S have written a 2006 Society of Vertebrate Paleontology Abstract, on p.74A. It concerns a new microvertebrate locality in the Norian Cumnock Formation of North Carolina. Remains are diverse and include at least two eucynodonts. One is tentatively referred to Traversodontidae, while its friend is thought to be the ferocious Microconodon.

Help: Should anybody have any further information, I'd be pleased to hear of it. Regarding references and Bibliography: I haven't and can't verify the references, so beware. Traditionally papers used in constructing this page are in the bibliography. If you feel these are too few, then send some more. With thanks to all the featured sources. Trevor Dykes, June 2001, Latest update: 4.12.2009. ktdykes@arcor.de

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With further thanks for due to: The Society of Vertebrate Paleontology's Bibliography of Fossil Vertebrates (John Damuth) http://www.bfvol.org/ Yale Peabody Museum, Collection Search (VP) http://george.peabody.yale.edu/vp/ Mark Isaak, Curiosities of Biological Nomenclature http://home.earthlink.net/~misaak/taxonomy.html who supplied the Cannabis Blyth, 1850. BIOSIS: The Index to Organism Names http://www.biosis.org.uk/triton/indexfm.htm Professor Pascal Godefroit for kindly supplying the informative papers.

Bibliography: Abdala F, Barberena MC & Dornelles J (2002), A new species of the traversodontid cynodont Exaeretodon from the Santa Maria Formation (Middle/Late Triassic) of Southern Brazil. Journal of Vertebrate Paleontology, 22 (2), p.313-325. Abdala F & Giannini NP (2000), Gomphodont cynodonts of the Chanares Formation: the analysis of an ontogenetic sequence. Journal of Vertebrate Paleontology, 20 (3), p.501-506. Abdala F & Ribeiro AM (2003), A new traversodontid cynodont from the Santa Maria Formation (Ladinian-Carnian) of southern Brazil, with a phylogenetic analysis of Gondwanan traversodontids. Zoological Journal, 139 (4), p.529-545. Abdala F & Smith RMH (2009 -'Proof only'-copy), A Middle Triassic cynodont fauna from Namibia and its implications for the biogeography of Gondwana, In press, Journal of Vertebrate Paleontology. Abdala F & Sa-Teixeira (2004), A traversodontid cynodont of African affinity in the South American Triassic, Palaentologia africana, 40, p.11-22. Battail B & Surkov MV (2000), Mammal-like reptiles from Russia, Chapter 6 in Benton MJ, Shishkin MA, Unwin AM & Kurochkin EN (Eds.), The age of dinosaurs in Russian and Mongolia, Cambridge University Press. Benton MJ (1990), The Reign of the Reptiles. Eagle Editions, (printed 1998), ISBN 1-902328-17-5. Blob, RW (2001), Evolution of hindlimb posture in nonmammalian therapsids: biomechanical tests of plaeontological hypotheses, Paleobiology, 27(1), p.14-38. Flynn JJ, Parrish JM, Rakotosamimanana B, Ranivoharimanana L, Simpson WF & Wyss AR (2000), New traversodontids (Synapsida: Eucynodontia) from the Triassic of Madagascar. Journal of Vertebrate Paleontology, 20 (3), p.422-427. Flynn JJ, Parrish JM, Rakotosamimanana B, Simpson WF, Whatley RL & Wyss AR (1999), A Triassic fauna from Madagascar, including early dinosaurs. Science, 286, p.763-765. Godefroit P & Battail B (1997), Late Triassic cynodonts from Saint-Nicolas-de- Port (north-eastern France). Geodiversitas 19(3), p.567-631. Godefroit P (1999), New traversodontid (Therapsida: Cynodontia) teeth from the Upper Triassic of Habay-la-Vieille (southern Belgium). Paläontologische Zeitschrift 73 (3/4), p.385-394, 6 Abb., 1 Tab. Hopson JA & Sues HD (2006), A traversodont cynodont from the Middle Triassic (Ladinian) of Baden-Württemberg (Germany), Paläontologische Zeitschrift, 80 (2), p.124-129. Hurum JH (1998), The inner ear of two late Cretaceous Multituberculate mammals, and its implications for multituberculate hearing. Journal of Mammalian Evolution, 5 (1), p.65-93. Kemp TS (2005), The Origin and Evolution of Mammals, Oxford University Press, pp.331. Lucas SG (2001), Age and correlation of Triassic tetrapod assemblages from Brazil. Albertiana 26, p.13-20. Lucas SG & Hancox PJ (2000), Tetrapod-based correlation of the nonmarine Upper Triassic of Southern Africa. Albertiana 25, p.5-9. Lucas SG & Heckert AB (2002), The Hyperodapedon Biochron, Late Triassic of Pangea. Albertiana 27, p.30-38. Renesto S & Lucas SG (2009), Cynodont teeth from the Carnian (Late Triassic) of northern Italy, Acta Palaeontologica Polonica, 54(2), p.357-360. Rogers RR, Arcucci AB, Abdala F, Sereno PC, Forster CA & May CL (2001), Paleoenvironment and taphonomy of the Chanares Formation tetrapod assemblage (Middle Triassic), Northwestern Argentina: spectacular preservation in volcanogenic concretions, Palaios, 16, p.461.481. Rubidge BS & Sidor CA (2001), Evolutionary patterns among Permo-Triassic therapsids. Annual Reviews of Ecology and Systematics 32, p.449-480. Sues H-D & Olsen PE (1990), Triassic vertebrates of Gondwanan aspect from the Richmond basin of Virginia. Science 249, p.1020-1023. Sues H-D, Olsen PE & Kroehler PA (1994), Small tetrapods from the Upper Triassic of the Richmond basin, (Newark Supergroup), Virginia, p.161-170, in (eds) Fraser NC & Sues H-D, In the Shadow of the Dinosaurs. Cambridge University Press. Therrien F & Fastovsky DE (2000), Paleoenvironments of early theropods, Chinle Formation (Late Triassic), Petrified Forest National Park, Arizon, Palaios, 15, p.194-211.