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.

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You could visit the Book Centre and look around.

Most of these animals were small to middling sized carnivores, but the later representatives tended to the tiny. Their original niche appears to have been assumed by other groups, including early dinosaurs. Chiniqs could be ancestral to the tritheledontans, (sometimes also assigned to Chiniquodontoidea; see Tritheledonta), which, along with basal mammals, superseded them. It hasn't, and probably can't be proven beyond all reasonable doubt that they were also the direct ancestors of mammals, but they appear damn good candidates. Then again, who knows what further research and discoveries might reveal?

Links: T Mike Keesey, The Dinosauricon, Ages of the Mesozoic http://www.dinosauricon.com/times/index.html Details on funny chronological terms such as Norian etc. Useful for reference. Mikko K Haaramo, Eucynodontia Mikko Haaramo's Eucynodontia These cladogrammes look straightforward, once the work's been done. However, bringing references dating from between 1857 and 2004 together in a coherent form is not easy.

Taxon: CHINIQUODONTOIDEA

Top A. Chiniquodontidae B. Dromatheriidae

Some possible chiniquodontoideans of unclear affiliations. I think of this non-section as a zone of uncertainty. Genera: Aleodon; Eoraetia, Gaumia, Kunminia, other reports Time-Line: Lower Jurassic: Kunminia Upper Triassic: Eoraetia, Gaumia Middle Triassic: Aleodon

Genus: Aleodon Crompton, 1955

Species:	Aleodon brachyrhamphus Crompton, 1955
Place:	Manda Formation, Tanganyika
Country:	Tanzania, also (for the genus) Namibia
Age:	Anisian, Middle Triassic
Remarks:	A skull of this genus, but not necessarily of this species, has also been recovered from the Omingonde Formation of Namibia (Abdale & Smith, 2009*, p.5). A feature the genus shares with Chiniquodon is the long bony palate that reaches to the end of the tooth row. The upper postcanine teeth are widened, and have occlusal outlines ranging from ovoid to elipsoid. This Namibian skull isn't well preserved (p.6) and, with a length of 15.6cm, it's about a centimetre longer than the holotype from Tanzania's Manda Formation. The snout acounts for 6.7cm. The teeth of this individual aren't well represented. The lower jaw is entirely missing. Upper postcanines can at least be counted on the left jaw; 4 incisors, 1 canine and 11 postcanines. (Note: I'm writing this somewhat in advance of the publication from a kind of preview copy, and thanks are due to FA for supplying it. The page numbering of the final edition will differ to those referenced above.) Additional remarks In 1972, this material was referred to the family Chiniquodontidae. Abdala & Giannini, 2002 reject this, (p.1151). It was originally described as a gomphodont, and is known from fragmentary skulls and lower jaw material, (p.1157). The best specimen is a muzzle with teeth, (BMNH 9390). Material is in the collection of The Natural History Museum, London. With thanks to Vince Ward for the species name.
Reference:	Crompton (1955), On some Triassic cynodonts from Tanganyika. Proceedings of the Zoological Society of London, 125, p.617-669.

Link: The Prehistoric Data Files http://www.angellis.net/Web/PDfiles/RepTherapS-al.htm An extensive meta-index of paleonotological genera. I find this site very useful.

Omingonde Formation, Namibia The following is based upon my reading of Abdala & Smith, 2009. My copy, thanks FA, is a pre-publication version, and that means the page numbers will differ to the published paper expected in September, 2009. No new taxa are invovled, so I can't see a necessity to await its official appearance. Namibia hosts a unit of cynodont-yielding rock from the Middle Triassic named the Omingonde Formation (p.1). It's not a new source, but further collecting there has resulted in an extension of the known diversity. A complication is provided by the detail that somewhat different ages appear to be represented. A lower horizon seems to correlate with part of the Cynognathus Assemblage Zone of the Karoo Basin in South Africa. A higher level matches better with somewhat later localities in Zambia and Tanzania. However, one of the newlings, Chiniquodon, packs a surprising punch. As well as being a further cynodont now known from both Africa and South America, its presence in apparently Aniasian aged deposits extends the chronological range of that genus to an abnormal length. The earliest known South American specimens are younger, Ladinian, whereas later ones have been recovered from the Carnian of the Upper Triassic. Its career of spreading mayhem and carnage apparently extended for over nine million years. Old and new Previous cynodont hunters have bagged four of the blighters from the lower area of the upper Omingonde; Cynognathus, Diademodon, Trirachodon and Titanogomphodon. The newer efforts have added four more from the upper reaches; Luangwa, a further traversodont, Aleodon and the aforementioned Chiniquodon. That makes it the most diverse round up in the Middle Triassic world. Half the Omingonde cynodont crew are also now known from South America as well as other African localities; Cynognathus, Diademodon. Luangwa and Chiniquodon. Vertebrate remains from this Formation were first described in 1973. They included amphibians and various therapsids including cynodonts. Subsequent discoveries added more meat to the stew, and the age was thought to correspond with subzones B and C of Karoo's Cynognathus Assemblage Zone, with C being the upper horizon (ie. somewhat more recent). That still seems to apply reasonably well. Sadly, the fossils tend to be in relatively poor condition (p.2). Fortunately, however, comparable specimens from other localities are often better. This tends to restrict Namibian Triassic cynodonts more to supporting roles, when it comes to providing information on the anatomy of taxa. Nevertheless, the cast list itself is of much interest. Compared to elsewhere The cynodonts of the uppermost Omingonde, as opposed to those from the somewhat lower community captained by Cynognathus, clearly contrast to the typical team lists of South Africa's and Antarctica's Cynognathus Assemblage Zone (p.10). Cyno doesn't occur whereas Aleodon, Chiniquodon and traversodontids do (p.12). While this might be explicable in terms of differing environmental factors favouring different critters, a somewhat different phase of time seems more plausible. As far as can be told, neither the climate or depositional conditions contrast strongly between Omingonde and South Africa. The uppermost part of the Cynognathus Assemblage Zone, subzone C, may well be similar in age to Omingonde's younger cynodont fauna (p.14). If so, then the present lack of any Karoo traversodonts could be considered surprising. One possibility is that the geographical situation may not have favoured travies by it then being too far south. However, known fossil yielding outcrops of subzone C are presently restricted to a single valley, and the cast of vertebrates is only a modest one. That apparent travie absence need not reflect the real mix of the Anisian wildlife there. Further Mesozoic site summaries can be found at Localities. Meet the eucynodonts of Omingonde Lower fauna (4 genera, 4 species) Cynognathus sp., Diademodon sp., Trirachodon sp., Titanogomphodon crassus Upper fauna (4 genera, 4 species) Traversodontids Luangwa sp., 1 additional poorly known taxon. Terrifying carnivores Aleodon sp., Chiniquodon sp.

Genus: Eoraetia 'from the dawn Rhaetian'

Species:	Eoraetia siegerti
Place:	Halberstadt
Country:	Germany
Age:	Rhaetian, Upper Triassic
Remarks:	Based on a fragmentary ulna, which was cited as a tritheledontid. With thanks to Vince Ward.
Reference?:	Dietrich WO (1937), Über eine Säugertierelle aus dem Rhät von Halberstadt. Neues Jahrb. Min. Geol. Pal., Beilage-Band 77, p.310-319, 6 figs., 1 pl.

Genus: Gaumia Hahn G, Wild R & Wouters G, 1987 'from Gaume' Remarks: The following is largely based upon my reading of Hahn et al, 1987, and thanks are due to the supplier. This genus is based upon small, isolated teeth (p.11). These are proportionately long and narrow, and the lingual and buccal sides run nearly in parallel. Crowns are descriptively triconodont in style, with a main central cusp and up to two smaller assistants both ahead and behind. All are arranged into a straight line, and there are no cingula. The root is deeper than the height of the crown, and it tapers in a regular manner along its course. It shows no inclinations towards division, and the demarcation between root and crown, although marked, is less pronounced than for its local pal, Lepagia. The most striking differences to Lepagia concern the roots. As well as being comparatively longer and tapered for this genus, the pulp opening at the base is also narrower. The lower teeth would have required a greater depth of jaw for their accommodation. As the root is around 50% longer than the crown, this is less extreme than known from another local cynodont, Pseudotriconodon. With that critter, the root is double the height of the crown, and also near oval in shape. Affinities Few specimens are available, and that limits the amount of information upon the range of individual variability. Gaumia could perhaps have connections with a loosely defined Chiniquodontidae or, then again, perhaps not. The lack of any tendencies towards root division debarred an assignment to Dromatheriidae. Additional notes "The absence of a bipartition of the root suggests that Gaumia is not a member of the family Dromatheriidae" (Godefroit & Battail, 1997), which is why it's not in that section of this directory. This genus easily qualifies as a microvertebrate; ie very small. Crudely put, these teeth have a triangular crown with a triangular root, which makes them look like rough diamonds. The main cusps point proudly upwards. There are actually further cusps, though they're not often very distinct.

Species:	Gaumia longiradiata Hahn G, Wild R & Wouters G, 1987
Place:	Habay-la-Vielle & Saint-Nicolas-de-Point
Country:	Belgium & France
Age:	Norian (late) - Rhaetian, Upper Triassic
Remarks:	The following is based upon my reading of Hahn et al, 1987, and thanks go to the supplier. A grand total of two postcanine teeth were identified for this species at Habay-la-Vielle (p.12), so the provisions allowed but a brief snack rather than a full-blown feast. The crown of the type fossil has a straight line of five cusps, with the central one being much the largest. Those to the fore and aft are the smallest. The central cusp is near to being symmetrical, with its front slope only a little more convex than the rear one, and both were made for cutting. Wear has rounded its tip and done away with the enamel on the front. Both neighbouring cusps have also been worn, and they aren't clearly separated from the centreling by valleys. They're more like should sticking out from its beefier body. Deep and narrow valleys, however, do clearly isolate the front and rear cusps. Division of root and crown is best seen from the front or rear, with the border being fuzzier on the buccal side. The root is one-and-a-half times greater than the crown, and its regular attitude towards tapering rewards it with a truncated triangle sort of profile. Comparatively speaking Compared to both Lepagia and Pseudotriconodon, other members of the local Triassic community, the postcanine is relatively wider (p.13). The level of wear, which is also present on a second specimen, is stronger. That probably indicates these teeth weren't used entirely for cutting food. Some degree of other processing also likely occurred. Holotype R.M.35 works as a postcanine tooth in the Institut royal des Sciences naturelles, Brussels. Its specific name is Latin for 'long rooted' and, remarkably enough, it refers to the length of the root.
Reference:	Hahn, Wild & Wouters (1987), Cynodontier-Zähne aus der Ober-Trias von Gaume (S-Belgien), Mèmoires pour servir à l'Explication des Carte Gèologiques et Minières de la Belgique, Mèmoire 24, p.1-33.

Species:	Gaumia? incisa Hahn G, Wild R & Wouters G, 1987
Place:	Habay-la-Vielle & Hallau
Country:	Belgium & Switzerland
Age:	Upper Triassic
Remarks:	The first part of this entry is also based upon my reading of Hahn et al, 1987 and still more thanks are due to the supplier. An important nomenclatural component is provided by the question mark (p.14). As may be obvious to most it denotes uncertainty about whether this species is part of the genus. Although broadly similar to the teeth of G. longiradicata, there are a couple of rather pronounced differences between the two species. The membership, however, is said to be probable (wahrscheinlich). The postcanine crown is very narrow, three times longer than wide, and the cutting blades are particularly sharp. Valleys clearly separate cusps from each other. While the enamel on the buccal side is flat, a few rills are found at the top of the main cusp on the lingual face. Lengths range from 1.2 to 1.4mm, and referred specimens occur both at Habay-la-Vielle and in deposits near Hallau, Switzerland. Holotype R.M.35 is one of the star exhibits at the Institut royal des Sciences naturelles, Brussels. Its specific name is Latin for eingekerbt, eingeschnitten. 'Notched' seems appropriate. It refers to the clear separation between cusps. Additional notes The postcanines have between three to five cusps, with the middlemost dominant. "The median pair of accessory cusps is set along the lower third of the main cusps. These are not very developed and always very blunt", (Godefroit & Battail 1997, p.592). Furthermore: "when the lateral accessory cusps are present, they are set at the base of the crown, quite anteriorly and/or posteriorly." This pair are small, but the differentiation is more distinct. These teeth lack distinct wear facets. In this way, and in the general arrangement of the crown furnishings, they are similar to Pseudotriconodon. However, the root is triangular, more robust and shows no inclination towards bipartition, which rather counts against membership of Dromatheriidae. Godefroit & Battail 1997, (p.593), go for Cynodontia incertae sedis. The genus has sometimes been referred to Chiniquodontidae.
Reference:	Hahn, Wild & Wouters (1987), Cynodontier-Zähne aus der Ober-Trias von Gaume (S-Belgien), Mèmoires pour servir à l'Explication des Carte Gèologiques et Minières de la Belgique, Mèmoire 24, p.1-33.

Species:	Gaumia sp.
Place:	Hallau
Country:	Switzerland
Age:	Upper Triassic
Remarks:	I suspect this enigmatic entry refers to fossils included within G.? incisa. Some were definitely from Hallau.
Reference:

Of Habay-la-Vielle and Interfacial Methods The following is partly based upon my reading of Hahn, Wild & Wouters, 1987. Thanks are due to both the supplier and his extremely busy photocopier. When last heard from, yet another box was being dragged out of the attic to have the dust blown off. Observant readers may notice the title of Hahn et al is in German. To the best of my knowledge, no translations are available. This article also contains information provided by Eric Freeman. As can be gleaned from either the German, English or French abstracts, this study concerns mostly isolated teeth of four non-mammalian cynodonts from the Upper Triassic (lower Rhaetian) of southern Belgium (p.2). Along with further taxa, a hopefully complete listing should be available at the foot of this article, they all came from the same bonebed of Habay-la-Vielle. As mentioned by Wouters et al, 1984, at least some of the sediment obtained was processed using an interfacial method devised by Eric Freeman. He's actually come up with at least three methods for aiding the discovery of micro-vertebrate fossils by utilizing easily available chemicals, and exploiting particular qualities of fossil bone and teeth. As such techniques could prove useful elsewhere, they certainly merit a mention here. How to find fossils with Fairy Liquid These interfacial methods are aimed at concentrating fossils; sorting them out from the unwanted junk. They exploit a peculiarity. Fossils show a preference for being wetted by oils rather than water. After the usual sort of sieving and filtering, fling the partially interesting remnants into a suitable container holding water and a couple of appropriate chemicals. Stir the broth vigorously and then wander off to do something more appealing. When they feel the time is right, the liquids will separate of their own volition. One will go down and the other up. Of necessity, there'll be a zone between both those faces; an interfacial zone. If the correct chemicals have been used, then that'll happen to be where the fossils concentrate for collection. The great majority of unwanted matter goes elsewhere. This sort of principle has also been exploited by hunters of conodonts and archaeologists chasing Vikings in Jorvik (aka York). The highly sophisticated chemicals involved can often be found in specialist supply centres known as supermarkets. For those favouring conspicuous consumption they have the possible drawback of being cheap as well as easily available. Candidates include petroleum jelly, candle wax, washing-up liquid, kerosene, dry-cleaning fluids and even polystyrene. A brief summary of interfacial methods in accessible English can be found in 1983's New Scientist magazine, 99, p.268, 28th July 1983. A more technical treatment is provided by: Freeman EF (1986), Progress in recovering microvertebrate fossils using Interfacial Methods, Journal of Paleontology, 60 (4), p.957-960 and references therein. None of this is discussed by Hahn et al, 1987, as their task was to describe some of the fossils. Nevertheless, these methods are worth knowing about. In the right circumstances they should be applicable all over the planet. At Habay-la-Vielle, according to Wouters et al, 1984, twenty kilos of sediment was concentrated down to less then two, and that was found to contain 55 grammes of vertebrate fossils (p.352). This challenge was something akin to finding needles in a haystack, but your prospects of success with doing just that are mightily enhanced by using some petrol, a flame thrower and subsequent inspection with a metal detector; aka applied chemistry. Back to the cynodonts Having located your cherished fossils among the more than 99.6% of unwanted rubbish, it'd be even more satisfying if you knew what on Earth they are. In this instance, most of the cynodont material, generally isolated teeth, happened to be from previously unknown genera. That's quite often what happens with remains from previously unsampled localities. The non-mammalian cynodonts required several new genera, and two of those were established in Hahn et al, 1987. Not part of this study were a couple of other members of the ancient wildlife; a 'haramiyidan' and Mojo (affinities disputed). Some advice on orientations of isolated cynodont teeth is provided on page four. These specimens are postcanine teeth of a style termed triconodont. That involves a straight line made by three main cusps, and the middle one is generally the largest. Among later mammals exceptions occurred for this scheme. Cases are known when the first cusp was lost, and others where the sizes became more equal. However, no such complications apply here. Experience relates that the more strongly convex slope of the main cusp is the front one. Additionally, the most strongly convex side is buccal. The reliability of this can naturally be checked on teeth retained in position on jaws, and nothing indicates these differences, when present, can't be trusted. They allow the correct orientation of even isolated teeth to be determined and, in cases with room for doubt, such uncertainties are pointed out. Further Mesozoic site summaries can be found at Localities. Meet the eucynodonts of Habay-la-Vielle (7 genera, 8 species) Non-mammals Plant munching traversodontids (2 genera, 2 species) Habayia halbardieri, Microscalenodon nanus Terrifying carnivores (3 genera, 4 species) Gaumia longiradicata, G.? incisa, Lepagia gaumensis, Pseudotriconodon sp. Mammalia? Haramiyida(ß) (2 genera, 2 species) Mojo usuratus, Thomasia woutersi

Genus: Kunminia Young CC, 1947 'from Kunmin'

Species:	Kunminia minima Young CC, 1947
Place:	Lufeng Formation, Yunnan
Country:	China (& Russia?)
Age:	Sinemurian, Lower Jurassic
Remarks:	Zhang, 1984 (p.3) reports this is based upon a reasonably complete skull with an articulated mandible. It represents a small animal. The known teeth consist of a canine and ten postcanines. It was originally interpreted as an ictidosaur, (tritheledontids and companions). The actual affinities will probably remain unclear without further specimens. The first six postcanines are single-rooted and unicusped. The rear teeth are double-rooted and multicusped. A cast of the holotype is at the Peabody Museum, Yale. Unfortunately, I've completely forgotten where I heard any mention of possibly Russian material, so the question mark is especially valid. Lu and Wu, 1994 mention this as an indeterminate synapsid of some kind or other. In terms of stratigraphy, it's from the upper Dark Red Beds, which are now thought to be Sinemurian.
Reference:	Young (1947), Mammal-like reptiles from Lufeng, Yunnan, China. Proc. Zool. Soc. London 117, p.537-597.

Other reports: Europe Link: Syren, Luxembourg: Centre de Recherches Lorraines: Dominique Delsate http://www.multimania.com/crcrl/articles/syrencrl.htm Diverse teeth are known from this site in Luxembourg. Similar finds are known from France, Belgium and Germany. A shorter, illustrated English print version of this detailed report is available (but not from me!). Reference: Godefroit P, Cuny G, Delsate D & (M.roche?) (1998), Late Triassic Vertebrates from Syren (GD Luxembourg). Neues Jahrbuch Geol. Stake. 210(3), p.305-343. Stuttgart.

Top A. Chiniquodontidae B. Dromatheriidae

A. CHINIQUODONTIDAE

Taxon: Chiniquodontidae von Huene FF, 1936 Chiniquodontids were up to doggish-sized predators. All of them have a characteristic right-angled bend at the back of the upper jaw, (Rubige & Sidor 2001, p.469), but their teeth are rather variable: "For example, Probelesodon has mitten-shaped postcanine teeth that completely lack cingula (i.e., the sides of the teeth are smooth), whereas Aleodon possesses transversely expanded, multicusped postcanines that led to its early classification as a gomphodont (Crompton 1955)." A recent review has been conducted by Abdala & Giannini, 2002. Through means of depriving some species of membership and synonymizing others, these authors restrict the family to just two species of Chiniquodon, (p.1151). I'm following their findings and have relocated a couple of genera to the zone of uncertainty at the head of this page. The aforementioned Probelesodon is now here treated as a synonym of Chiniquodon whilst Aleodon is hanging around above. Other taxa which have been placed within the family at one time or another include: Cromptodon, Cistecynodon parvus (?= Cynognathus), Thrinaxodon brasiliensis (= Prozostrodon) and Probainognathus (Probainognathidae). According to Abdala & Ribeiro 2000 (p.593): "Chiniquodontids are Middle to Late Triassic cynodonts that are restricted to South America". The same authors continue, "The postcanines of chiniquodontids lack cingula and have undivided roots (Abdala 1996). The crowns of the posterior postcanine posses four cusps with the anterior accessory cusp being faintly defined." They have more observations to offer on the dentistry, none of which I'd pretend to fully understand. Accessibly expressed, they had complex teeth for chewing meat. The dating of the Brazilian locations can be a discussion point: "Clearly, the presence of Hyperodapedon and Stagonolepis supports correlation of the Hyperodapedon Assemblage Zone of the Santa Maria Formation with the vertebrates of the Ischigualasto Formation in Argentina, and therefore an Ischigualastian (Adamanian) age assignment", (Lucas & Heckert 2002, p.33-34). I'm attempting to follow the conclusions of Lucas, 2001 and some background is available at: Rio Grande do Sul. Genera: Belesodon (= Chiniquodon), Chiniquodon, Probelesodon (= Chiniquodon), other reports Time-Line: Middle - Upper Triassic: Chiniquodon

Genus: Chiniquodon von Huene FF, 1936 'Chiniquá tooth' Aka: Belesodon von Huene, 1936; Probelesodon Romer, 1969 Remarks: Omingonde Formation of Namibia Abdala & Smith, 2009 report on the arrest of a member of this genus from Namibia (p.7). Its state of preservation makes reference to a particular species presently undo-able. That it turned up in Africa, rather than South America, isn't all that much of a surprise. Four Triassic cynodonts have now been recorded as parts of both the African and South American scenery. However, the Namibian Chiniq is remarkably early; Anisian rather than Ladianian-Carnian. And, if correctly identified, that amounts to a career of about nine million years; unusually long for Middle Triassic cynodonts. The Namibian skull has been squashed, otherwise deformed and left with a length of just over 19cm. Exposure to weather prior to fossilization contributed a generous supply of cracks to the bone, and few of the sutures between individual elements can now be seen.

Species:	Chiniquodon brasilensis Barberena, 1982
Place:	near Chiniquá, Santa Maria Formation, Rio Grande do Sul
Country:	Brazil
Age:	Ladinian, Middle Triassic
Remarks:	A respectably dog-sized predator, with a skull-length of about ten cm. However, Abdala & Giannini, 2002 doesn't mention this species, at least not as member of the genus. The species name makes me suspicious. I wonder if this may somehow be involved with Prozostrodon brasilensis.
Reference:

Links: Projeto Dinossauros Do Brasil http://acd.ufrj.br/mndgp/cinodon.htm A couple of illustrations. Universidade Federal do Rio Grande do Sol http://www.ufrgs.br/geociencias/paleo/cinodontei.html A nice photo of the skull.

Species:	Chiniquodon theotonicus von Huene FF, 1936
Aka:	Belesodon magnificus von Huene FF, 1936; Probelesodon kitchingi Teixeira AMS, 1982; P. lewisi Romer AS, 1969; P. minor Romer AS, 1973
Place:	Santa Maria Formation, Rio Grande do Sul & Los Chanares Formation
Country:	Brazil & Argentina
Age:	Ladinian, Middle Triassic
Remarks:	Some of these possible synonyms are also in use as distinct taxa by various sources. I intend to arrange the links listing for this entry in line with these names, and perhaps include any pertinent info there. I'm presently treating them as synonyms in response to Abdala & Giannini, 2002. Subsequent publications might of course change this. Abdala & Giannini's 2002 paper concerns a study based upon fifteen measurements of the skull, (p.1154), in order to examine detailed growth responses to an overall increase in body size. Their results show: "... that all specimens can be arranged in a continuous growth series independent of their original assignment, indicating the existence of a single form", (p.1167), with the exception of C. sanjuanensis. This means that all the various specimens studied can be accounted for in just two species. Taxonomic history The authors also discuss some of the problems which have beset chiquodontid systematics, (p.1161-1162). Although he described two genera, von Huene didn't actually provide a diagnosis for either of them. The first chiniq to receive such an honour was Probelesodon lewisi in 1969. A major difference given between this and the already named Belesodon was that it's much smaller, but the length of the bony palate of the mouth and shape of teeth were also seen as significant. Part the reason for this unsatisfactory state of affairs, was that previously described material had been either fragmentary, badly preserved or both. The next character to come on stage was P. minor. This taxon was established purely for reasons of size. Then came P. kitchingi. What were held to be distinguishing traits for this species, were generally already known as variations within the larger number of specimens of P. lewisi, though it's somewhat unusual in having a sidewards expansion of the frontal bone in the skull and ten postcanine teeth. Then again, this latter condition is also known from one specimen of the usually nine cheek-toothed lewisis. A few musings on the various skull sizes can be jeered at after the links section. Holotype This taxon was originally based on an incomplete skull and a humerus. The type fossil, according to Abdala 2007 is GPIT 40, and it studies at Tübingen. The holotype of Belesodon magnificus is in the collection of Tübingen University, Germany, where it's affectionately known as GPIT 1050. I thought that was the number of C. theotonicus. However, Abdala, 2007 (p.612) informs me otherwise. Furthermore, checking below strongly indicates I'd gotten it wrong. MCP PV1600 is the type of Probelesondon kitchingi, and it should be interviewable at the Museu de Ciencias e Technologia at Pontificia Universidade, Rio Grande do Sul. Should you prefer discussions with the holotype of P. minor, then visit PULR 12 at the Museo de Antropologia, Universidad Nacional de La Rioja. However, I'd check before hand as it was reportedly stolen. I suppose it could've escaped and returned. Much of the Argentinean material was identified by Romer.
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. NB.: 'Unterkeuper' = Upper Triassic (lower)

Links: Chiniquodon theotonicus Universidae Federal do Rio Grande do Sul http://www.ufrgs.br/geociencias/paleo/cinodontei.html A photo of a skull from a fine gallery of local fossils. 'La Importanica turistica de Ischigualasto' preparado por Dr William Sill http://www.ischigualasto.com/es/ischigualasto.htm Included in the inventory of Annexo VII, (Spanish). M. Alan Kazlev, Late Carnian Age http://www.kheper.auz.com/gaia/Mesozoic/Triassic/Carnian/carnian5.htm More on the fossils of Ischigualasto. Synonyms of Chiniquodon? Belesodon von Huene, 1936 'fine tooth' Belesodon magnificus von Huene, 1936 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. The Mesozoic Eucynodonts' not-terribly-good artist had aquick go at a sketch of the skull. Belosodon (Chiniquodon) I'm informed the thin, pointier end is the snout. The University of Tübingen http://www.uni-tuebingen.de/uni/e16/gpi/sammlung/seiten/zweit12.htm A photo of the skeleton and a brief anatomical description of Belesodon magnificus, (German). The holotype (GPIT 40) is also in the paleontological collection at Tübingen University, where von Huene was a professor for over 60 years. He died in 1969. Lucas, 2001 refers to this genus as a traversondontid, (p.16), whilst Abdala & Giannini regard it as a synonym of Chiniquodon, advice which I've followed. The original description was of an almost complete, poorly preserved skull, and lots of the skeleton, A visit to the Tübingen museum is on my 'things to do' list. In fact, it's been on the list for a couple of years already. Probelesodon Romer AS, 1969 'before fine tooth' Douglas Henderson's Earth History Illustrations http://gallery.in-tch.com/~earthhistory/triassic%20page%201.html Includes an illustration of Probelesodon. Tom Holtz, Dinosaur Mailing List Archive http://www.cmnh.org/dinoarch/1994May/msg00010.html The holotypes of both P. lewisi and P. minor were in the collection of the National University of La Rioja, Argentina. They were stolen in January 1994. P. kitchingi Teixeira, 1982 Remarks: This was based upon a skull without mandibles, (Abdala & Giannini, p.1151). Reference: Teixeira (1982), Um novo cinodonte carnivoro (Probelesodon kitchingi sp. nov.) do Triassico do Rio Grande do Sul, Brasil. Comunicacóes do Museu de Ciencias da PUCRGS 24, p.1-31. Teixeira AMS http://www.unb.br/ig/posg/outros/ufrgs4.htm P. lewisi Romer AS, 1969 Reamrks: The stolen holotype was UPLR 18, (Chure D 2000, p.19). Reference: Romer (1969) The Chanares (Argentina) Triassic reptile fauna. A new chiniquodontid cynodont, Probelesodon lewisi. - Cynodont ancestry. Breviora 333, p.1-24. Valdosta State University's Virtual Museum of Fossils http://gatito.valdosta.edu/fossil_pages/fossils_tri/t66.html This suite of images is accompanied by a Ladinian age, rather than Carnian. Views differ. It also presently cutes a species name of lewskii, rather than lewisi. This is a typo and will be corrected. The specimen pictured was borrowed from Harvard MCZ and a cast was made. The people at Valdosta University, (which is in Georgia), are hoping acquire and upload more specimens, especially tritheledonts and other new South American forms. (With thanks to Dr Edward Chatelain, Associate Professor of Geology). The type fossils of P. lewisi and minor were both stolen from the National University of La Rioja, Argentina in 1994. P. minor Romer AS, 1973. Reamrks: The stolen holotype was UPLR 12, (Chure D 2000, p.19). Reference: Romer (1973), The Chanares Formation (Argentina) Triassic reptile fauna XVIII, Probelesodon minor, a new species of carnivorous cynodont; family Probainognathidae nov. Brevoria, 401, p.1-4. Valdosta State University's Virtual Museum of Fossils http://gatito.valdosta.edu/fossil_pages/fossils_tri/t67.html A further fine collection of photos.

Species:	Chiniquodon sanjuanensis (Martinez RN & Forster CA, 1996) Abdala F & Giannini NP, 2002
Aka:	Probelesodon sanjuanensis Martinez RN & Forster CA, 1996
Place:	Ischigualasto Formation
Country:	Argentina
Age:	Carnian (mid), Upper Triassic
Remarks:	This species is based on a complete skull with articulated lower jaws, and postcranial remains. Many thanks to Darren Naish and the DML, (see link). The morphology is broadly similar to C. theotonicus, as are the proportions of the skull. The differences concern the highly arched cheek bone, and the presence of serrations on the edge of the main postcanine cusp. This feature's certainly present on the rear-most tooth, and probably on its colleagues, (Abdala & Giannini 2002, p.1161). It also has a relatively low number of postcanines, six. Some differences might be accounted for by its being a juvenile, (p.1162). Even so, the authors accept that it represents a separate species, (p.1167). The type fossil, PVSJ 411, works at the Museo de Ciencias Naturales, Universidad Nacional de San Juan, Argentina.
References:	Martinez & Forster (1996), The skull of Probelesodon sanjuanensis, sp. nov., from the late Triassic Ischigualasto Formation of Argentina. Journal of Vertebrate Paleontology, 16, p.285-291.
	Abdala F & Giannini NP (2002), Chiniquodontid Cynodonts: Systematic and Morphometric Considerations. Palaeontology, Vol. 45, Part 6, p.1151-1170.

Links: Journal of Vertebrate Paleontology http://www.vertpaleo.org/jvp/16-285-291.html The abstract of the report by Martinez & Forster, 1996. Darren Naish, Dinosaur Mailing List, 21.11.2002 http://www.cmnh.org/dinoarch/2002Nov/msg00521.html An outline of the findings of Abdala F & Giannini NP (2002), Chiniquodontid cynodonts: systematic and morphometric considerations, Palaeontology 45, p.1151-1170. According to my information Cromptodon mamiferoides Bonaparte, 1972, which the link mentions, is a member of Galesauridae. If correct, this family isn't within Eucynodontia, though it's very close. The fossil comes from the Rio Mendoza Formation of Argentina, and its teeth show resemblances to a juvenile specimen of Aleodon, (Abdala & Giannini 2002, p,1151). The holotype is in the collection of the Universidad Nacional de Tucumán, Argentina, (PVL 3858). This is a tiny lower jaw with postcanine teeth, (p.1157). Reference: Bonaparte (1972), Cromptodon mamiferoides, galesauridae de la Formación Rio Mendoza, Mendoza, Argentina (Therapsida-Cynodontia). Ameghiniana 9, p.343-353. DMG Projects, University of Texas http://digimorph.org/specimens/Probelesodon_sanjuanensis/ A photo of the holotype formerly known as Probelesodon sanjuanensis. "Within Eucynodontia, Probelesodon is the sister taxon to Chiniquodon." The relationship appears even closer.

Species:	Chiniquodon kalanoro Kammerer CF, Flynn JJ, Ranivoharimanana & Wyss, AR, 2010
Place:	Isalo II or Makay Formation
Country:	Madagascar
Age:	Middle (Ladinian) or Upper Triassic (Carnian)
Remarks:	I haven't as yet seen this description. Reportedly, the species is based on a lower jaw known as UA 10607.
Reference:	Kammerer et al (2010), The first record of a probainognathian (Cynodontia, Chiniquodontidae) from the Triassic of Madagascar, Journal of Vertebrate Paleontology, 30(6), p.1889-1894.

Ischigualasto Formation, Upper Triassic, Argentina Hollocher et al, 2005 is a description of a couple of fossils from one location in this Formation. The two specimens were probably provided by early, Upper Triassic dinosaurs, but it's not as obvious as might be thought. In this case, the only bits of bone are remnants of former meals. These fossils are coprolites; dino dung. Perhaps sadly, I don't fell moved to address the main subject of the study. My concern is more with the Formation, the environment it represents and especially the eucynodont fauna. Ischi Ischigualasto is a fossil rich series of rock layers in northwestern Argentina, and the area is not exactly pastoral. The name comes from an indigenous language. The Spanish alternative translates as the Valley of the Moon, and that gives a reasonable approximation of the local plant life. Vegetation is extremely sparse, and the paucity has the advantage of the rocks being well exposed. The Formation occurs in the Ischigualasto- Via Union Basin, which contains continental deposits of up to 4,000 metres from the Middle to Upper Triassic, (p.52). About 340 metres belong to the Ischigualasto section, in which vertebrate remains are both unusually plentiful and excellently preserved. Deposition was mostly the responsibility of river action; both channels and floodplains. Further sediments were supplied by lakes. A radiometric dating from twenty metres above the base of the Formation provided a figure of 228 million years (argon/argon method). The specimens in the study came from approximately 54 metres higher up. According to an estimate, that difference represents about half-a-million years. Neighbouring floors Ischigualasto is on top of Los Rostros Formation, which is therefore somewhat older. It's exposed to the southwest, and has provided few remains of vertebrates. Above can be found Los Colorados Formation, and that's exposed to the northwest. This somewhat younger sequence has provided fossils. I've compiled a list of eucynodonts and included it at the foot of this article. (Additionally, Los Chanares Formation has yielded some fabulous fossils. It predates Los Rostros.) What was the weather like? A three metre section contains a succession of layers, and these include the horizon with the coprolites. The conditions suggest relatively gentle river action in an arid or semi-arid environment. Who left the mess? Most common are remains of two herbivores; a rhynchosaur called Scaphonyx sanjuanensis (the genus is incorrectly spelt on page 62) and the eucynodonts Exaeretodon frenguellis. Neither would have been excreting bits of bone. The size of the droppings suggest a carnivore with a bodyweight of between five and fifteen kilos. Available candidates are: two early dinosaurs (Eoraptor and Herrerasaurus); two cynodonts (Chiniquodon and Ecteninion); a juvenile rauisuchid (Saurosuchus); and a juvenile semiaquatic archosauriform (Chanaresuchus). It was probably Herrerasaurus who did the dirty. Further Mesozoic site summaries can be found at Localities. Meet the eucynodonts of the Ischigualasto Formation (five genera, six species) Traversodontidae (two genera, two species) Exaeretodon frenguelli; Ischignathus sudamericanus Probainognathia (three genera, four species) Probainognathus jenseni; Ecteninion luensis; Chiniquodon sanjuanensis; C. theotonicus. Bonus: Eucynodonts of Los Colorados Formation (upper Norian): (One genus, one species) Chaliminia musteloides.

How long were chiniquodontid skulls? I'm pleased you asked, seeing as Abdala & Giannini, 2002, includes a table of details on page 1164. This is based on 36 specimens of Chiniquodon, Probelesodon and Belesodon, 19 of which are unpublished. Included is a measurement referred to as TL: the length from snout tip to the middle of the back of the skull, the occipital condyle, which is where the skull forms the joint with the uppermost neck vertebra. (With thanks to David Marjanovic.) I'm going to concentrate on the various holotypes and, because I always find measurements difficult to visualise, employ a special bit of equipment known as a piece of A4 paper, (30 x 21cm). Taking into account the bulge on either side of the occipital condyle, Belesodon magnificus would require most the sheet, (TL 26cm). An unpublished specimen would overhang it, (31cm). That's a big animal, especially when clad in muscle and skin. To reach the next of the type fossil lengths, fold the paper and leave a gap of about 1,5cm between the top and the bottom edges. That's about right for Probelesodon kitchingi, (16,4cm). Push the top edge slightly below the bottom and flip the sheet over. The now upper surface gives an impression of the estimated length of Chiniquodon theotonicus's skull, (14,5cm). The easiest way to progress is to next use the sheet in landscape format. Fold it straight down the middle, and we've got an approximation of Probelesodon lewisi, (11cm). To visualise Probelesodon sanjuanensis, recreate the model for P. kitchingi and fold in half, (8cm). As the sheet's now rather crumpled, I'll just mention that the P. minor type fossil is one centimetre shorter, and put the paper in the ironing basket for later. In summary, the holotypes have the following TLs: i. B. magnificus 26cm; ii. P. kitchingi 16,4; iii. C. theotonicus 14,5; iv. P. lewisi 11; v. P. sanjuanensis 8; and vi. P. minor 7. That's a smoothish progression, from ii. down to vi. anyway. But that's restricting the sample to the type fossils. As stated on page 1162, the differentiation between P. minor and P. lewisi was size. A specimen more akin to the holotype of the former is a centimetre longer than the type fossil of the latter. Clearly, size alone cannot a species define in that instance. What these authors did was to measure the specimens whilst: "ignoring their original species assignments", (p.1154). And the result: "shows that all specimens can be arranged in a continuous growth series independent of their original assignment, indicating the existence of a single form", (p.1167). One exception here is P. sanjuanensis which shows genuinely diagnostic distinctions. A second exception might be seen in Belesodon. The difference between this and the next longest holotype is greater than the skull length of the above cited exception. This gap closes partly with the introduction of an incomplete specimen from Chanares known as MCZ 4020. The palate suggests an individual which would possibly measure slightly less than the width of A4, (20,3cm): "This fact is not trivial if we consider that larger sizes in the Chanares fauna are attained by specimens not fossilised in concretions. Meanwhile, nearly all vertebrate fossils in this fauna are recorded within concretions, showing a counter-intuitive bias against the preservation of large-bodied specimens (Rogers et al. 2001)", (p.1165). It certainly runs counter to my intuition. Whether there's a bias against partial specimens of larger individuals is something I don't know, perhaps because I'm unfamiliar with Rogers et al, 2001. I also don't know any reason why this gap shouldn't be closed by fossils from sites other than Chanares, should all these types indeed represent the same species. However, I'm confident that the authors couldn't identify significant, diagnostic differences to distinguish Belesodon from the other specimens studied, other than for size. If anyone else can, please do so.

Other Reports: Vellberg, South-western Germany A number of teeth have been recovered from Kupferzell-Bauersbach. I'm informed these look rather like "traversodont" molars from the Manda Formation, (Zambia and Tanzania), which were described by Broili & Schröder. These were subsequently reidentified by von Huene as being from chiniquodontids. A humerus has also come to light, and is presently being interrogated at Tübingen. (With thanks to Marcel Opitz and Dr Michael Maisch.) References: Schoch R & Wild R (1999), Die Saurier von kupüferzell, der gegenwärtige Forschungsstand. p.409-418, in Hauschke N & Wilde R (eds.), Trias-eine ganz andere Welt, Munich (F. Pfeil). Schoch R (April 2002), STRATIGRAPHY AND TAPHONOMY OF VERTEBRATE-BEARING BEDS IN THE LOWER KEUPER (Middle Triassic) Of VELLBERG (SW GERMANY). (I don't have a more precise citation.)

Top A. Chiniquodontidae B. Dromatheriidae

B. DROMATHERIIDAE

Taxon: Dromatheriidae Gill, 1872 Dromatheriids were small critters of the Upper Triassic, whose anatomical relationship to basal Mammalia is close. Remains are scarce and typically restricted to isolated teeth, though there some pieces of dentary. They were probably insectivores. Most the finds, especially from Europe, are restricted to isolated teeth. Trying to resolve which ones are upper and which lower can clearly be tricky. However: "When wear is developed, it is possible to distinguish the upper teeth from the lower ones; wear develops on the lingual side of the former and on the labial side of the latters", (Godefroit & Battail 1997, p.571). If there's no wear or difference in extent of wear between the two sides of the tooth, then differentiation passes beyond the realms of tricky, and into the mists of impossibility. As a proper taxon of commonly related animals, the family is somewhat iffy. In the language of Abdala & Ribeiro 2000 p.593): "The group, as currently defined, is noticeably heterogenerous, showing different kinds of crown morphologies." The teeth of closely related animals should be reasonably similar in construction. "They are best regarded as a derived " eucynodont" group, but their affinities to mammals remain uncertain due to their incomplete fossil (Sues 2001)", (Luo, Kielan-Jaworowska & Cifelli 2002, p.5). The particular mammal-like qualities known from these fossils are also known from tritheledontids and further cynodont groups. Nevertheless: "The cladogram of the Dromatheriidae proposed by Hahn et al. (1994) shows that this group is clearly paraphyletic, as the ancestry of the mammals lies within it", (Godefroit & Battail 1997, p.571). All of which suggests that this is perhaps better seen as an informal grade, rather than a natural taxon based on common ancestry. Despite the lack of clarity of details, it's clear these animals were close to basal Mammalia. Other nearly-mammals may be encountered at probainognathians. Reference: Simpson GG, (1926), Are Dromatherium and Microconodon mammals? Science 63, p.548-549. Genera: Deccanodon, Dromatherium, Microconodon, Pseudotricondon, Rewaconodon, Tricuspes, other reports Time-Line: Upper Triassic: Deccanodon, Dromatherium, Microconodon, Pseudotriconodon, Rewaconodon, Tricuspes, Saint-Nicolas-de-Point (France), Geissgurglebach (Germany)

Genus: Deccanodon Nath TT & Yadagiri, 2007 'Deccan tooth' Remarks: The authors don't assign the genus, which is based upon one postcanine, to any family. However, as they speicfy similarities with Microconodon, I feel justified in informally placing it within the dromatheriid section of this directory. The generic name honours the Deccan Plateau.

Species:	Deccanodon maleriensis Nath TT & Yadagiri PY, 2007
Place:	Maleri Formation
Country:	India
Age:	Carnian, Upper Triassic
Remarks:	The following is based upon my reading of Nath & Yadagiri, and thanks are due to the supplier. Something both new and ancient is astir in the Maleri Formation of Andrha Pradesh, Central India. This already varied vertebrate collection of the Carnian aged fauna has been enriched by the presence of a formidable killer; a cynodont challenger to outshine the already reported dullard of a depressingly dim-witted coelurosaurian dinosaur (p.57). Admittedly, with a postcanine tooth length of about a millimetre, Deccanodon probably wouldn't have won a duel with the dino using merely muscle and strength. But, had it so chosen, it could doubtlessly have run rings around its rival without being noticed. However, as known remains are presently restricted to those of a dental persuasion, ie. a tooth, some room for uncertainty should be allowed for when regarding the rest of the physique. The authors say these remains are the first for cynodonts from this Formation, although a report of Exaeretodon leads me to doubt that's quite so. I suppose it may have been reinterpreted since 1982, and forgot to tell me. Nevertheless, Decannodon does seem to be the first carnivorous cynodont. And where there's one, then that's usual an excellent place to search for some more. Five postcanine teeth have so far been identified, and only one of them has been referred to the new taxon. Resemblances between them all and Microconodon are cited; crown shape, cusp distinctness, a lack of a constriction between crown and root, and incipient division of the roots. Orientations The five collected specimens all have lines of cusps arranged in straight lines running along the crown, but cusp numbers very from 2 (two specimens), 3 (two sp.) to 6 (one sp.). The possible significance of this isn't addressed, but reasonable possibilities could involve different dental positions, perhaps different taxa or conceivably both and maybe something I haven't thought of. Only the six-cusped postcanine is described. It's a rather odd looking thing and, I confess, simply looking at the overall profile in the graphics left me wondering which end was which. Generally, with this sort of critters, the front slope of a cusp is somewhat more convex than the rear one. That certainly seems to apply for cusps identified as a and c. The situation with b, possibly due to perspectives, seemed less clear cut. However, one photo seems to depict a more convex slope as well. A cingulum, if present, would be expected to be lingual for both uppers and lowers. Presumably, such factors were involved in identifying it as a right canine, but I wouldn't like to guess why they opted for lower. (That isn't to suggest either they had no reason for doing so, or that I think they might be wrong. I simply don't know.) Some justification would have been appreciated by this critic but, in fairness, not much space is available with just four pages. Besides, close looks at graphics amount to good practise. The tooth of 'Deccan tooth' The postcanine is small, a fraction over a millimetre in length, and that obviously indicates a tiny owner. Usually, the tallest cusp on dromi choppers (and the like) are the centrally positioned ones termed a. Most unusually, but not entirely uniquely, its b that's dominant in this instance, and that occurs nearer to the front. There's then a decrease in sizes behind, first for a then c and finally d. A further smaller pair of sentinel cuspules hang out on the very front of the base of the crown, e (lingual) and f (buccal), and their jobs are to interlock with part of the rear of the tooth in front. Such devices, popular with later mammals as well, help add stability to the tooth row. Apart from some of the rear of the single root, the preservation is rather good. It shows the specimen is coated with a uniform covering of enamel. The crown, from the occlusal perspective, is a long, narrow rectangle, and somewhat taller than the depth of the preserved root. Presumably, the use of "preserved" implies the root would've been longer in life, but no speculation is offered upon how much may be missing. In profile, the root has the shape of a semi-eclipse, and a furrow runs vertically along it; incipient bifurcation. A weak cingulum occurs on only one side, and comparative anatomy (took a quick look at the few pointers listed above, should you wish) suggests that's the lingual one. Affinities The authors assign all the 'triconodont'-type teeth (regardless of cusp number) to an undefined "Chinquodontoid group", in their words. At least, I think they do but the phrasing seemed a bit ambiguous. History appears to have conspired against them on one point: "Deccanodon maleriensis gen. et sp. nov. cannot be compared with any Indian specimens, as no Chiniquodontoidea cynodont teeth were reported so far. This wasn't quite correct when written. A find tentatively attributed to Microconodon was reported in the Annual Report of the Ministry of Mines 1999-2000. I happen to know the holotype of Deccanodon was discovered some years ago, as I heard from somebody who tells me they saw it in around 2002, and it could relate to the tooth in question. However, prior to this publication, Rewaconodon from the Tiki Formation saw the light of day in 2004. Still, Nath & Yadagiri don't claim omnipotence, and I should be less mischievous. If a comparison could've been made, then I'm sure they'd have carried one out. Comparisons with Gondwanadon were made and similarities found. Known from a single postcanine from the Tiki Formation, that genus still stands accused of being a very early morganucodontid mammal. However, the pattern of cusp height differential isn't the same. Gondwanadon is more conventional; a>b>c>d. 'Deccan tooth' has b>a>c>d. The lack of a constriction demarcating the root and crown, and the incipient bifurcation of root are more in keeping with Microconodon of North America but, again, cusp a is the largest in that case. A European genus, Meurthodon, does share the enlargement of cusp b. The fact that its tooth (presumably that refers specifically to the holotype) only has four cusps isn't necessarily overly significant. That sort of thing can reflect tooth row positions. Cusp counts frequently vary between teeth from the same set. The root division, however, is much more strongly pronounced for the somewhat later Euro-genus (p.59). With no obvious generic home available for the fossil, the authors established a new one. Given the usual habits of fossil localities, I'd expect it to receive companionship from further local cynodonts at some time. Holotype 22/SR/PAL is a postcanine working in the service of the Geological Survey of India, Southern Region, Hyderabad. The specific name is derived from the Maderi Formation.
Reference:	Nath & Yadagiri (2007), A new mammal-like reptile (Cynodontia) from the Upper Triassic Maleri Formation of Pranhita-Godavari Valley, Andhra Pradesh, Journal of the Geological Society of India, 69, p.57-60.

Genus: Dromatherium 'running beast'

Species:	Dromatherium sylvestre Emmons E, 1857
Place:	Egypt Coal Mine, North Carolina, Newark Supergroup
Country:	USA
Age:	Carnian - Norian, Upper Triassic
Remarks:	An enlarged cast specimen is in the collection of the American Museum of Natural History, New York.
Reference:	Emmons, Fossils of the sandstone and slates of North Carolina. Proc. Amer. Assoc. Adv. Sci., 11th meeting, Montreal 1857, p.76-80. (NB.: This report seems to have been published in 1858.)

Links: ARTFL Project: Webster Dictionary, 1913 http://machaut.uchicago.edu/cgi-bin/WEBSTER.page.sh?page=456 If anyone wants to check my spelling, please feel free. Huber P, Lucas SG & Hunt AP, (1996) http://www.ldeo.colunbia.edu/~polsen/nbcp/huber.html Carnian-Norian Biochronology, Correlation and Biotic Events of the Nonmarine Triassic of North America. This paper reports on attempts to date different locations relatively, according to the vertebrate fossils found; strata with similar fossils are allocated to similar ages. Whilst some people might object to this, they often ignore or are totally ignorant of any other remains which also occur in these strata. It's not so much because of the presence of one particular vertebrate that the sites are found to correlate. It's the whole fossil assemblage. This is only one method of dating. Biochronology often involves demonstrably widespread, short-lived index fossils such as ammonite species, which occur only in marine deposits. With terrestrial remains, pollen can often be a useful indicator, as can other plant material, insects and many freshwater organisms. Radiometric dating techniques are not dependent upon fossils at all. For a short bit of informal and perhaps totally unreliable discussion, see How to Date Dinosaurs by Self MY, (but don't take it too seriously).

Genus: Microconodon Osborn HF, 1886 'small coned tooth' Remarks: 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 this genus, while its friend is a traversodontid.

Species:	Microconodon tenuirostris Osborn HF, 1886
Place:	Egypt Coal Mine, North Carolina & Turkey Branch Formation Virginia, Newark Supergroup
Country:	USA
Age:	Carnian - Norian, Upper Triassic
Remarks:	An enlarged cast specimen is at the AMNH, New York. The original fossil was "an incomplete right dentary with four post-canine teeth", (thanks to Ben Creisler and the DML -9.6.2002.) Several more specimens have been described. Talking of which... The following is based upon my reading of Sues, Olsen & Kroehler, 1994, (see Bibliography for details). Fossils from The Richmond Basin of central Virginia include remains of a traversodontid, Boreogomphodon, and three dentaries and several isolated postcanine teeth from this critter, (p.165). The isolated teeth are very similar to those of Pseudotriconodon from Luxembourg. They've typically got three cusps arranged in a line. The front one (b) is small, the middle one (a) large, and the rear one's (c) about the same size as the first. The shape is broadly like a pointy eared gnome in a hat, (though much smaller). There are no cingula, which means the teeth are smooth. The dentary has a long, low ramus which bears the teeth and lacks a clearly defined angular process. It was home to three distinct types of tooth. As well as the postcanines, there was also room for a canine and three, slightly forward pointing incisors per side. In the smallest jaw from this material, the front postcanines are relatively simple, whilst the posterior are more clearly tricuspid. There's no sign of a diastema, (a gap between teeth, which can be useful for food processing. Ask a rabbit.) However, there is such a thing on the largest specimen. Those simpler front postcanines were lost, and this left a prominent diastema between the canine and the properly tricuspid choppers. The holotype, (no. 10248) is in the collection of the Academy of Natural Sciences, Philadelphia. For some reason I didn't look into, this generic name also finds favour with a number of pornographic pages! If you know why, don't feel compelled to offer further enlightenment.
Reference:	Osborn (1886), Observations on the Triassic mammals Dromatherium and Microconodon. Proceedings of the Acadamy of Natural Sciences, 37, p.359-363.

Links: Biotic Provinciality of the Late Triassic Equatorial Humid Zone http://gsa.confex.com/gsa/2001SE/finalprogram/abstract_4541.htm An item by Olsen PE, Schneider V, Sues H-D, Carter JG & Peyer K. Natural Canvas Fossils http://www.thenaturalcanvas.com/Mammals/pages/4092.html From the catalogue of a fossil trader, (it's nothing to do with me). This is a photo of a rather well preserved 'Microconodon' tooth from Arizona. If you click on the upwards pointing arrow, you may find a couple more images. Whether they actually represent Microconodon is open to question, but that's what Natural Canvas says, (with thanks to David Nathan).

Species:	Microconodon sp.
Place:	Tiki Formation, South Rewa Gondwana basin
Country:	India
Age:	Upper Triassic
Remarks:	This may be the basis of Rewacondon, but I'm not yet sure. Alternatively, should my locality information be incorrect for some reason, it could perhaps pertain to Deccanodon.
Reference:

Link: Ministry of Mines, India http://www.nic.in/mines/archp3gsi.html News from the Annual Report 1999-2000, (Chapter III). The link appears to have died. Should anybody happen across the webpage at a new location, then feel free to let me know of it.

Genus: Pseudotriconodon Hahn G, Lepage J & Wouters G, 1984 'false Triconodon' Remarks: Information supplied by Professor Godefroit suggests that Pseudotriconodon is a dromatheriid. This taxon is largely a dental one; ie. known from gnashers. The postcanine teeth have between three and five triangular cusps on narrow crowns. According to Godefroit and Battail 1997, (p.574), "partial bipartition of the root" is observable in some specimens. This taxon is seen as a rather basal dromatheriid. The reason for the name is that the architecture of the postcanine crowns shares features in common with the molars of Triconodon, a Jurassic mammal. However, these clearly aren't results of a close relationship.

Species:	Pseudotriconodon wildi Hahn G, Lepage J & Wouters G, 1984
Place:	Medernach & Saint-Nicolas-de-Point & Holwell quarries, Somerset
Country:	Luxembourg, France & England
Age:	Norian (late) - Rhaetian, Upper Triassic-?Lower Jurassic
Remarks:	The following is based upon my reading of Hahn, Lepage & Wouters, 1984. Thanks are due to Eric, his photocopier, the postal services of UK and Germany, the manufacturers of ink and paper... A road building scheme in Luxembourg provided a welcome opportunity for fossil hounds hungry for vertebrate remains from the Upper Triassic (p.357). The work exposed a bone bed near Medernach; an unexpected bonus but one with plenty of parallels. As well as a potential for damaging both paleontological and archeological treasures, road building can also make a positive contribution to our appreciation of the past. In this instance, it lead to the recovery of teeth from small fish, crocs, pterosaurs, cynodonts and possible dinos (p.358). 'Three cusps' (or more or less) Various animals of a Norian age had teeth that can be referred to as tricuspid in type, although the term doesn't literally only apply to three-cusped crowns. In essence, there needs to be a centrally located line of cusps, and further small cusps may also be present. In the case of Pseudotriconodon the crowns have a long, narrow outline, with the length being two to three times the width. The sides are relatively straight and run in parallel. The length range is 1 to 1.5mm; smaller than a grain of black pepper. There's a centrally located main cusp which is twice, or more, the height of accompanying cusps situated both ahead and behind. There may be a further smaller cusp to the fore and up to two more aft. Be that as it may, all are arranged in a single line. Absent from these crowns is any trace of cingula. Teeth can have such bits of shelving on their sides, but these ones don't. Another feature is that the tooth enamel is flat and even with no ornamentations. There are also the roots. Each tooth has but one, and it's up to 1.5 times deeper than the crown's height. Even the shorter roots still manage 1.25. Usually, the root is semi-elliptical (the base is unsurprisingly narrower than the top), and there's little or absolutely no tendency towards bifurcation. Sometimes, however, there is some degree of such inclinations near the pulp cavity and, in fewer cases, a groove running down the root to its tip (or even tips). It could be playfully suggested that two roots were kind of thinking of developing but, as with two naked children wanting to play mummy and daddy, their equipment was in no way mature enough to attempt the task they must've heard about somehow or other. This is bifurcation at a prototype stage (if at all). A further fact is the lack of an obvious demarcation between root and crown. There's no indentation or any form of inflation conveniently marking the border. Which way round, up, down and so on A couple of dozen or so postcanine teeth were identified for this genus from Medernach, and working out which are upper and lowers is, in a way, a straightforward. This is because it's presently undoable. No identifying features were apparent. Other bits of orientation could be determined by comparisons with other eucynodonts. One slope of the main cusp is generally somewhat convex along its course while the other is more concave. The second is probably the front. That applies for triconodontid premolars. One flank is somewhat more convex than the other, and that's indicative of the buccal side. Tooth groups Based on the varying numbers of cusps (p.360), the authors came up with five groups of postcanines. Additionally, there were four specimens with two accessory cusps at the rear and an indeterminable number at the front, and two preserving only the roots. There was also a fragment of a crown. The most numerous teeth have five cusps (10 specimens). Four-cusped models came second (5 sp.) and tri-cuspids third (4 sp.). Comparisons with other "dromatheriids" suggest this could be connected with dental positions. For both Dromatherium and Microconodon, the front postcanines are tri-cusped whereas the others have more cusps. Wear There's not much to speak of, and certainly no sign of regular wear facets. Upper and lower teeth couldn't have closely occluded. Some show traces of use on the tip of the main cusps, which became blunted. A strong majority don't even manage that. Affinities At that time of Earth (p.361), four animal groups are known to have had triconodont-style teeth; early mammals, non-mammalian eucynodonts, reptiles called prolacertilians and some pterosaurs. Representatives were lined up for an identity parade. Mammals? No. The crowns are too simple with no traces of cingula and, at most, the roots have weak bifurcation or none. Morganucodon roots, in comparison, are divided into two (although not necessarily entirely completely), and the crowns have cingula with cuspules. Prolacertilia? The most famous prolacertilian is an outrageously bizarre reptile with a ludicrously long neck. This is Tanystropheus; more a neck with an animal stuck on it than the other way round. Adults reach lengths of up to seven metres, and over half of that is contributed by ten to twelve ridiculously long neck vertebrae. More typical adults are three to four metres long. Even so, you can't go around sticking one millimetre teeth into mouths like that. However, their kids were much smaller critters with fairly normal necks, and these happen to be the representatives with triconodont-style teeth. Adult choppers are single-cusped. But the Medernach specimens can't be assigned to junior prolas. Their teeth are only tricuspid, never more, they have oval outlines and the roots don't narrow along their course, or at least hardly. Ptersoaurs? This is trickier. As the original paper is in German, it may be of some use if I attempt to summarize this section more lengthily than usual. However, I'm not doing a direct translation. Several pterosaurs that hang out in the Medernach fauna have triconodont-style teeth. One such is Eudimorphodon. The authors list a number of points meriting attention for comparisons. 1. Enamel Eudimorphodon teeth mostly have ribbing of the enamel, and this isn't the case for Pseudo. Inconveniently, this ribbing is less strong for juveniles, and their enamel can be smooth. 2. occlusal outline The outline is elliptical for Eudi, and the main cusp is wide and fat. Pseudo has more of a stretched rectangular outline with a narrowly built cusp. Inconveniently, juvenile cusps are narrower as well; more like Pseudo than the parents. 3. Side view of crown Pseudo has a long and fairly low main cusp. Eudi has mostly shorter, taller cusps. Inconveniently, some Eudi teeth have main cusps closer to the proportions for Pseudo. 4. Cutting edges The front and rear slopes of Pseudo's cusps are straight and thus sharp. Eudi teeth have more rounded slopes. Inconveniently, this difference doesn't hold true for the kids. 5. Crown-root demarcation Fortunately, there is a clear border between Eudi crowns and roots, whereas the demarcation isn't marked for Pseudo. That difference is useful. At least, it is if that area happens to be preserved. 6. Root depth, crown height Pseudo crowns are an average of 1.25 times higher than root depth. This figure is around 0.94 for Eudi. 7. Root division Some indication of bifurcation can occur for Pseudo, but not always. No such thing has ever been observed for any pterosaur. 8. Frequency (I've got some quibbles with this!) This could help with large enough sample sizes, but not with individual specimens. Both for Medernach's isolated teeth and teeth in place in jaws from elsewhere, Eudi has about the same number of three- and five-cusped teeth, especially in adults. (A juvenile jaw is known with a 12:17 ratio). Four-cusped teeth are rare but can occur. Pseudo, going by the available sample, has more five-cusped specimens (p.362). (My quibble: The authors cite a ratio of nearly 1:4, but I don't follow haw that can be right. Gruppe I (p.360) have three cusps and there are four teeth. Gruppe II have four cusps. Gruppe III and IV are penticuspid, and a single specimen, Gruppe V, has six cusps. The number of front cusps is unknown for Gruppe VI and could, presumably, have been either four- or five-cusped. I also presume those (and the six-cusped tooth) must have been added together to make fifteen specimens; nearly four times Gruppe I. All those seem to have qualified as "five-cusped specimens".) Be that as it may, the point remains valid that penticuspid teeth are more common than tricuspid ones, and tetracuspid specimens are more common than for Eudi. 9. Tooth lengths Eudi adult teeth have an average length of 2mm. Pseudo postcanines can't match that; 1 - 1.5mm. Inconveniently, Eudi kids have teeth of that sort of length as well. If the preservation allows, Eudi and Pseudo teeth can generally be distinguished. However, distinguishing Eudi kids from Pseudo is more difficult and, conceivably, won't always be presently possible. Most significant are points 5 and 7, and those provided grounds for the assignment to Cynodontia. Identity Cynodont. The teeth matched closest with Microconodon and Dromatherium. The unclear demarcation of crown and root and non-divided roots are features held in common. However, incipient bifurcation is stronger in both North American colleagues (p.363). Similarities include the straight line of cusps, the lack of cingula and, of less significance, the size. The new genus was accordingly referred to the family of Dromatheriidae. The text part of the paper is rounded off with an extensive summary, and that's translated into both English and French. Holotype This is the type species of the genus. The holotype, RM1, is in the collection of the Institut Royal des Sciences Naturelles de Belgique in Brussels. The specific name applauds the insights of Rupert Wild, a German paleontologist. He contributed valuable advice on how to differentiate between multi-cusped teeth from diverse vertebrate groups. Think of England The following block is largely based upon my reading of Cluny, 2004, and thanks are due to the kindly supplier. This description concerns a fossil tooth from the Holwell quarries of Somerset, and the age of that locality is subject to dispute. Some chat about that fauna can be found below. The Holwell specimen is named C108 (p.70). It's a lonely, isolated postcanine of 2.0mm in length and 0.8 width. Despite being called 'false three coned tooth', there are actually five cusps in all. There's a big daddy bear-sized cusp in the middle, two much smaller mummy bear-sized cusps (one in front and one behind), and baby bear-sized mini cusps to both extremes. The side walls are almost vertical things, and there's no sign of a constriction to offer elucidation upon the demarcation of the crown and root. All the cusps are arranged into a simple straight line and, for the sake of better clarity, the daddy bear-sized central one is by far the biggest and tallest, by very far. The height of the tooth is 3.1mm, and 1.8 of those have been interpreted as root. What it ain't The absence of any constriction between crown and root rules out Tricuspes, Gaumia and Lepagia, and the lack of any incipient root division means it's not assignable to Microconodon or Dromatherium. The near vertical walls of the tooth contrast to the curvature on one or the other side of Hahnia and Meurthodon. It's from none of that lot then, and the given reasons are comprehensible even to me. The shape corresponds with that of Pseudotriconodon wildi and the size is consistent as well. And that explains why it was assigned to the species. Should the Liassic date be correct, than this would be a fairly radical range extension for a dentally basal "dromatheriid".
Reference:	Hahn et al (1984), Cynodontier-Zähne aus der Ober-Trias von Medernach, Grossherzogtum Luxemburg. Bull. de la Soc. belge de Géologie 93 (4), p.357-373.

Link: Travaux en cours au natur musée http://www.mnhn.etat.lu/colsci/dedo/projets.htm A detailed report on fossil finds from Medernach, (French). A further P. tooth has been reported from the nearby site of Syren. Gilles Cuny http://palaeo.gly.bris.ac.uk/personnel/departed/cuny/Cunygr.html Mentions a probable Pseudotriconodon tooth from the Rhaetian-Liassic of Holwell, Somerset, England, now resident in the museum of the beautiful city of Bath. My psychic powers indicate this has been assigned to P. wildi.

Holwell quarries, Somerset, Rhaetian-Liassic The first part of this article is kind of based upon Cuny, 2004, but not all items mentioned can be found in that study. Remarkably, for a paper on fossils from Somerset, he somehow neglects to mention either cider or the now legendary Glastonbury rock festival. The Holwell quarries are in Somerset, southwest England and, as this is also prime farmyard cider country, things can quite get to look less than clear. Serious students of that subject are inclined to buy the stuff by the gallon and, if mixed with beer, then unclarity can reach an unconscious intensity. This is a discovery some grockles regret making and, after groggily reawakening some time the next day, they begin to appreciate why a cider-beer mix is known as a snake bite. Of course, being Somerset rocks born and bred, the strata in the Holwell quarries know instinctively that some experiments are best avoided. They stick to supping the cider-infused local rain but, what with resultant erosion and general earthly jitteriness, the rocks sometimes have cracks busted into the; fissures. And, over time, such fissures fill up with a jumble of natural junk. Should that happen to include bits of dead or trapped plants and animals, then the collection can be confusing when it comes to trying to date any fossils. Dating can be made difficult. That also applies when drinking snake bite, but that's a different form of the word dating. Fissures featured in the programme of the local rock concert at some time -or may be several times- during the Rhaetian-Liassic; the uppermost Triassic and lowermost Jurassic. And things fell or were dumped in. Some of this stuff were remains from cynodonts and Cuny, at least, considers the fossils he had to deal with were probably Rhaetian (p.69). Although the relevant non-mammalian eucynodont supply is presently limited to the odd tooth and scraps of jaw, what there is happens to overlap with Rhaetian faunas from Western Europe. Other authors -seemingly the majority- regard the locality as being Jurassic. Be that as it may, in terms of the eucynodont fauna, there happen to be overlaps with those European Rhaetian sites; eg Pseudotriconodon, tritylodontids, 'haramiyidans' and morganucodontid mammals. Discovery and neglect The first person to properly scour these fissures for fossils was a geologist in the nineteenth century named Charles Moore, and he published a number of accounts from 1859-1867. His research was one of the earliest deliberate attempts to sieve large quantities of sediment for the purpose of finding micro-fossils, and this was amply rewarded. Following Moore's inevitable extinction, many of his collected treasures from Holwell found sanctuary in the Bath Royal Literary and Scientific Institution. Unfortunately, and despite the impressively regal name, that place closed down in 1940 and hibernated through to 1968. The resultantly inert security perhaps allowed for the mass escape by the entire British supply of Tricuspes, should a single tooth be capable of a mass anything. In any case, that specimen couldn't be found by Cuny. The limestone containing the fissures was deposited far back during the Carboniferous age, and it was exploited from two quarries in the village of Holwell; North and South (p.70). As luck would have it, the cynodont-yielding fissures -as with the Tricupses tooth- are no longer available for interview. The rock's been quarried away. Their content can't be said to offer a typical snap shot of land vertebrates from their time. While Mr Moore amassed a collection of 29 mammalian teeth (assuming haramis are mammals), numerical superiority is ever so slightly achieved by fish. One species, Lissodus minimus, defied minimalism by bequeathing 45,000 teeth. This rock was beneath the sea at the time, and remains of land critters were swept in from elsewhere. Still, even an accumulation of a couple of dozen teeth suffices to suggest this 'elsewhere' can't have been very far away. Age There is, as mentioned, no consensus agreement on this, but the issue isn't as murky as good farmyard cider. (Those familiar with only the transparent fizzy liquid available in pubs knowing nothing worthwhile about cider at all. The drink is for supping and not looking through.) Given their paucity, basing a date on only the eucynodont remains would be more daring than anything the Dan of that name ever got up to, and especially as that part of the fossil record isn't playing a radically different style in music in the Jurassic Liassic, as compared to the Triassic Rhaetian. Tritylodonts, "haramiyidans" and morganucodontids; it was all Rock and Roll fit for both stages. And even should all 'Liassic' haramis turn out to be Rhaetites, the Middle J ones had Lower Jancestors. It's not actually known which of the two quarries yielded the Pseudotriconodon tooth but, as Moore was the finder, that makes the southern quarry more likely. A partial trity jaw came from the North Quarry. Nevertheless, the much more numerous fishes aren't noticeably different in either case, and they've been interpreted as indicating the Rhaetian. Additional notes Benton et al, 2005 (p.34) accuses the quarries of harbouring the trity, Oligokyphus. My information is that this is a filthy lie... Well, not that exactly. A couple of pieces of jaws were tentatively referred to that genus. However, they subsequently confessed to being more indeterminate. Remains aren't good enough to allow them to be referred to any particular trity. Still, they could perhaps be from Olis. The eucynodont fauna, in terms of numbers of taxa, has been experiencing something of a diet during the last ten years. Some have been synonymized (p.35). As had long been suspected, teeth assigned to Haramiyda were indeed the uppers of Thomasia, and only two species have survived interrogation. And then there's the just mentioned down-grading of erstwhile Oli. However, lovers of nomenclatural diversity need not despair. After all, Pseudotricondon is a novelty for this locality. Further Mesozoic site summaries can be found at Localities. Meet the eucynodonts of Holwell quarries, Somerset (five genera, six species) - more information will follow Non-mammalian Eucynodontia> (three genera, three species) 'Dromatheriidae' Pseudotriconodon wildi; "Tricuspes sp." (the specimen has long been lost so I've added quotation marks - see Cuny, 2004). Tritylodontidae. Trity indeterminate. ?Mammalia (one genera, two species) 'Haramiyida' Thomasia moorei, T. antiqua. Mammalia (one genus, one species) Eozostrodon parvus.

Species:	Pseudotriconodon chatterjeei Lucas S & Oakes W, 1988
Place:	Bull Canyon Formation, New Mexico
Country:	USA
Age:	Norian, Upper Triassic
Remarks:	The following is based upon my reading of Lucas & Oakes, 1988. This paper is a relatively short study of a small vertebrate fossil. There's a fragment of dentary a couple of centimetres long, and this contains three alveoli and a solitary postcanine tooth. A further postcanine was found close by and presumably belonged to this set, although it's somewhat damaged and doesn't cleanly fit onto the bone. These teeth closely resemble those of Europe's beloved Pseudotriconodon wildi. However, a couple of differences suggest at least a separate species (p.445). The first is its smaller size (about a third less). On its own, that wouldn't necessarily be of significance. The second factor is provided by striations on both the lingual and labial faces. Such scratches aren't present in the enamel of P. wildi Locality and age A campaign involving screenwashing found traces of various vertebrates in a shale horizon of the Chinle Formation at Bull Canyon, which is in eastern New Mexico. Remains from the locality had been turning up for nearly a century, but the more systematic approach extended the fauna. Dominant are parasuchian reptiles and labyrinthodont amphibians. Also occurring are dinosaurs, pterosaurs and sphenodonts. An 'ictidosaur' was reported ( 'Pachygenelus milleri'), but it's been forcefully accused of being some kind of fish rather than a mammal-like tritheledontan. The fossils suggest a Norian age for the locality. Three coned teeth The postcanine preserved on the jaw has a length of 0.71mm and a width of 0.25 (p.446). The central cusp is the tallest (0.41mm), while the front and rear cusps are near to each other in height (0.23 and 0.21 respectively). The isolated tooth is broadly similar but shorter (0.63). The central cusp is a blunt triangular shape when viewed from either side, and both faces have scratches running down from the top. These fail to reach the base and no opinions are offered on their origins. Their course is generally somewhat diagonal tending towards vertical. The cross-section shape of the tooth is semi-ovoid and there are no cingula. Also lacking is any groove between the crown and the root, and that root shows no inclinations towards division. This is in contrast to various 'dromatheriids' and appears to be relatively primitive. There are no indications of replacement postcanines in the jaw (p.447) so, despite its smaller size, no substantial evidence suggests this might be a juvenile. Family? The reference is actually to 'Dromatheriidae', and the inverted commas are significant. This 'family' is presently used as a handy catch-all for small, broadly similar cynodont teeth, and doesn't denote a natural taxon of one animal and all of its descendants. The two longest established members are Dromatherium and Microconodon, genera which are distinctive enough to probably belong in separate families. However, a robuster classification would require more instructive specimens, and they presently remain elusive. Holotype The type specimen, now known as NMMNH P-4600, is housed at the New Mexico Museum of Natural History, Albuquerque. It was described under the number UNM MV-518, (Morgan & Lucas 1999, p.257). Additional notes Reservations have been expressed about the correctness of this diagnosis, but they weren't supported by clear argumentation, (Godefroit & Battail 1997, p.622).
Reference:	Lucas & Oakes (1988), A Late Triassic cynodont from the American South-West. Paleontology 31, p.445-449.

link: Palaeontology, 31(2) http://palaeontology.palass-pubs.org/pdf/Vol%2031/Pages%20445-449.pdf Spencer & Oakes, 1988 is presently accessible on-line in pdf format.

Species:	Pseudotriconodon ? sp. Hahn G, Wild R & Wouters G, 1987
Place:	Habay-la-Vielle
Country:	Belgium
Age:	lower Rhaetian, Upper Triassic
Remarks:	The following is based upon my reading of Hahn et al, 1987, and thanks are due to the supplier. A single postcanine from Habay-la-Vielle was provisionally assigned to this genus (p.17), but more specimens would be required for confirmation due to the known range of variability; build of crown and the presence of divided root tips in a few cases. While Pseudotriconodon usually exhibits no indications of root division, some exceptions do show minor talents in such a direction. It's a five cusped crown. The tip of the main central cusp has gone due to wear, its front slope is convex and the rear one lightly concave. Valleys between cusps are clearly formed; deeper than for Gaumia longiradicata but shallower than G.? incisa. Demarcation between crown and root is only weakly perceptible at the front and back, but hardly discernable when viewed from the sides. The root has a half-elliptical profile and is 50% deeper than the height of the crown. The form of the root is the most obvious contrast to other local eucynodont insect-baiters. The whole tooth is very similar to fossils from Medernach, Luxembourg, and that's why it's been assigned to the genus. Its length is a majestic 1.24mm.
Reference:	Hahn, Wild & Wouters (1987), Cynodontier-Zähne aus der Ober-Trias von Gaume (S-Belgien), Mèmoires pour servir à l'Explication des Carte Gèologiques et Minières de la Belgique, Mèmoire 24, p.1-33.

Genus: Rewaconodon Datta PM, Das DP & Luo Z-X, 2004 'Rewa coned tooth'

Species:	Rewaconodon tikiensis Datta PM, Das DP & Luo Z-X, 2004
Place:	Tiki Formation, Madhya Pradesh
Country:	India
Age:	Carnian, Upper Triassic
Remarks:	The abstract mentions several isolated three- and four-cusped postcanines, with incipient division of the root. Some plesiomorphic resemblance with Theriherpeton is also cited, but the greatest similarity is with Microconodon. With thanks to Dino Hunter for posting notice of the publication. This very likely involves a fossil I've presently got an entry for under Microconodon, but I'm not certain.
Reference:	Datta, Das & Luo (2004), A Late Triassic dromatheriid (Synapsida : Cynodontia) from India, Annals of the Carnegie Museum, 73(2), p.12-24.

Link: APT On-Line, The Annals of the Carnegie Museum http://apt.allenpress.com/aptonline/?request=get-abstract&issn=0097-4463&volume=073&issue=02&page=0072 The abstract of the publication is available.

Genus: Tricuspes von Huene E,1933 Remarks: This genus is based on more small teeth from a number of European sites, and was also originally interpreted as mammalian. As regards stratigraphy, Rhaetian-Liassic seems to be the most accurate phrase in use, (uppermost Upper Triassic - Lower Jurassic). "Status questionable," (McKenna & Bell, 1997). The genus may belong within Mammaliaformes Rowe, 1988. As the name suggests, the teeth have three main cusps, which are "not perfectly aligned but arranged in a V-like manner," (Godefroit " Battail 1997, p.575). Smaller accessory cusps are sometimes also present. The root's divided, though "both portions remain in close contact with one another", (p.576). Upper and lower dentition can often be identified with the help of wear facets. A single tooth from Holwell quarries, Somerset was referred to the genus, but it managed to get lost. Links: Late Triassic cynodonts from Saint-Nicolas-de-Port, Godefroit P & Battail B. http://cimbad.mnhn.fr/publication/geodiv/g97n3a4.html An abstract on isolated but diverse fossil teeth from north-eastern France. EspacePaléo 3J: Dominique Delsate http://home.pi.be/~nerodj2/Npagehtm/txtvtp3_hbmam2.htm A report on finds from Syren, Luxembourg and Varangéville, France, (French).

Species:	Tricuspes sigogneauae Hahn G, Hahn R & Godefroit P, 1994
Place:	Saint-Nicolas-de-Point & Syren
Country:	France & Luxembourg
Age:	Norian (late) - Rhaetian, Upper Triassic
Remarks:	Upper and lower postcanines. This genus lacks the extra cusp, (m), known from T. tuebingensis. A further difference is that the other accessory cusps, when present, are: "not incorporated in the cutting edge of the crown, but located on its anterior and posterior wall", (Godefroit & Battail 1997, p.577).
Reference:	Hahn et al (1994), Zur Stellung der Dromatheriidae (Ober-Trias) zwischen den Cynodontia und den Mammalia. Geologica et Palaeontologica 28, p.141-159.

Species:	Tricuspes tapeinodon Godefroit P & Battail B, 1997
Place:	Saint-Nicolas-de-Point & Syren
Country:	France & Luxembourg
Age:	Norian (late) - Rhaetian, Upper Triassic
Remarks:	Ten or so isolated teeth. The species name, from the Greek, means 'low tooth'. The molariform teeth have a very low crown, and the central cusp (a) isn't much higher than its two main colleagues (b and c). Also in contrast to T. tuebingensis, there's none of this accessory cusp m stuff. However, when d and e are present, they make up part of the cutting edge, (as in T. tuebingensis but in contrast to T. sigogneauae.
Reference:	Godefroit & Battail (1997), Late Triassic cynodonts from Saint-Nicolas-de-Port (north-eastern France). Geodiversitas 19(3), p.567-631.

Species:	Tricuspes tuebingensis von Huene E, 1933
Place:	Gaisbrunnen, Tübingen & Saint-Nicolas-de-Port
Country:	Germany & France
Age:	Norian (late) - Rhaetian, Upper Triassic
Remarks:	The holotype, (an isolated right molariform), is in the Geologisch-Paläontolgischen Institut der Universität Tübingen. Relatively low-crowned teeth with a couple of reasonable well developed accessory cusps, (d/D and e/E), depending upon whether lower or Upper respectively. It differs from T. sigogneauae: "in the presence of a tiny accessory cusp m on the postero-lingual portion of cusp c", (Godefroit & Battail 1997, p.577).
Reference:	von Huene (1933), Zur Kenntnis des württembergischen Rätbonebeds mit Zahnfunden neuer Säuger und säugerähnlicher Reptilien. Jahresh. Ver. Vaterländ. Naturk. Württemberg 89, p.65-128.

Other Reports: Geissgurgelbach, Germany There are various ways Mammalia can be defined. As I feel a personal attraction for female breasts, (don't worry. I'm housetrained.), I favour a definition involving the ability of babies to suckle, which is presently first-evidenced by Morganucodon. A predilection for breasts may not be part of a scientific approach, but I'm an amateur and find it satisfying. Excuse me. I'm just going to talk to my wife... Later... Now, where was I? Oh yes, breasts. There are a couple of earlier genera which have been considered as mammals in a loose sense, although there's no indication regarding suckling abilities and the inferred presence of proper mammary glands. Fossils which could unbutton the blouse of this matter haven't been found. Adelobasileus is known from the Upper Triassic, (Carnian). In my terms it's a perhaps-mammal. It predates the earliest recognized members of either Tritylodontidae or the family Tritheledontidae, both of which feature strongly in discussions concerning mammal origins. The dromatheriids wish to point out that, scrappy though it may be, their fossil record also predates those families. If better remains, and perhaps a possible transitionary form between Dromatheriidae and Morganucodontidae were found, their case would perhaps strengthen. The following is based upon my reading of Lucas et al, 2001. As judged by the presence of index fossils, the wondrously named Geissgurgelbach location in Germany, (northeast of Stuttgart), seems to be upper Carnian, (p.397). This is where SMNS 80766 was found. It's a fragment of dentary containing one postcanine tooth, (crown length 1,12mm, maximum width 0,58mm). A central cusp (a) is dominant. It's bluntly pointed and has a small wear facet. The front of the crown houses a much smaller cusp (b). A chip is missing from the back of the tooth, (back is used here as meaning the surface nearest the back of the jaw incidentally), at the labial side. If in line with both dromatheriids and morganucodontids, there would have been a further cusp (c) present. Three very small cuspids are also found on the labial side. "The crown base below the cusps is columnar, and the root base is divided into two, highly divergent roots. This root division begins well below the crown base." The details of this morphology are unique. It's probably a new taxon, but the authors declined to name it on the grounds of its incompleteness; thus the classification of indeterminate cynodont. In contrast to dromatheriids: its roots are more strongly divided; the crown isn't as 'trenchant'; and it does possess a labial cingulid. Those three very small cuspids are on that feature. In contrast to the mammalian morganucodontids: there's no evidence that this tooth interlocked with the corresponding upper tooth, as shown by the lack of a system of wear facets; various cuspids and a lingual cingulid at the front aren't present; the root division begins below the base of the crown. (At least some of these 'deficiencies' also apply to Sinoconodon postcanines.) From p.399: "SMNS 80766 can thus be envisioned as a postcanine tooth morphology intermediate between that of a dromatheriid and a morganucodontid. Further division of the root base, addition of an antero-lingual cingulid, and cuspids with development of interlocking wear would transform the SMNS 80766 tooth morphology into a morganucodontid tooth." The specimen resides in the collection of the Staatliches Museum für Naturkunde, Stuttgart. Reference: Lucas SG, Heckert AB, Harris JD, Seegis D & Wild R (2001), Mammal-like tooth from the Upper Triassic of Germany. Journal of Vertebrate Paleontology, 21 (2), p. 397-399.

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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 all the references, so beware. Traditional 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, September 2001. Last update: 12.8.2009 Ktdykes@arcor.de

With further thanks 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/ BIOSIS: The Index to Organism Names http://www.biosis.org.uk/triton/indexfm.htm American Museum of Natural History, New York, Specimen Catalogue Dr Michael Maisch, Tübingen University, for advice on the paleontological collection of his institution and the stratigraphy of South America. M Alan Kazlev, for further considerations on the stratigraphy of South America. Professor Pascal Godefroit for supplying the informative and interesting papers on European microvertebrates. Dr Zhe-Xi Luo for kindly sending further papers.

Bibliography: Abdala F (2007), Redescription of Platycraniellus elegans (Therapsida, Cynodonita) from the Lower Triassic of South Africa, and the cladistic relationships of eutheriodonts, Palaeontology, 50(3), p.591-618. Abdala F & Giannini NP (2002), Chiniquodontid Cynodonts: Systematic and Morphometric Considerations. Palaeontology, Vol. 45, Part 6, p.1151-1170. Abdala F & Ribeiro AM (2000), A new therioherpetid cynodont from the Santa Maria Formation (Middle Triassic), southern Brazil. Geodiversitas. Vol. 22, (4), p.589-596. 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. Benton MJ (1990), The Reign of the Reptiles. Eagle Editions, (printed 1998), ISBN 1-902328-17-5. Benton MJ, Cook E & Hooker JJ (2005), British Mesozoic fossil mammal GCR sites, Chapter 2 of Benton, Cook & Hooker, Mesozoic and Tertiary fossil mammals and birds of Great Britain, Geological Conservation Review Series, No. 32, Joint Nature Conservation Committee, Peterborough, p.27-66. Chure D (2000), New threats to old bones, the theft of fossil vertebrates from museum collections, Cultural Resource Management, 23 (5), p.18-22. Cuny G (2004), A Late Triassic cynodont from Holwell Quarries (Somerset, England), Oryctos, 5, p.69-73. Godefroit P (1997), Reptilian, therapsid and mammalian teeth from the Upper Triassic of Varangéville (northeastern France). Sciences de la Terre 67, p.83-102. 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. Godefroit P & Battail B (1997), Late Triassic cynodonts from Saint-Nicolas-de-Port (north-eastern France). Geodiversitas 19(3), p.567-631. Godefroit P, Cuny G, Delsate D & Roche M (1998), Late Triassic Vertebrates from Syren (Luxembourg). N. Jb. Geol. Paläont. Abh. 210(3), p.305-343, Stuttgart. Hahn G, Lepage JC & Wouters G (1984), Cynodontier-Zähne aus der Ober-Trias von Medernach, Grossherzogtum Luxemburg, Bulletin de la Societe Belge de Géologie, 93(4), p.357-373. Hahn G, Wild R & Wouters G (1987), Cynodontier-Zähne aus der Ober-Trias von Gaume (S-Belgien), Mèmoires pour servir à l'Explication des Carte Gèologiques et Minières de la Belgique, Mèmoire 24, p.1-33. Hollocher KT, Alcober OA, Colombi CE & Hollocher TC (2005), Carnivore coprolites from the Upper Triassic Ischigualasto Formation, Argentina: chemistry, mineralogy, and evidence for rapid initial mineralization, Palaios, 20, p.51-63. Kellner WA & Campos DA (1999), Vertebrate paleontology in Brazil - a review, Episodes, 22(3), p.238-251. Lucas SG (2001), Age and correlation of Triassic tetrapod assemblages from Brazil. Albertiana 26, p.13-20. Lucas SG & Heckert AB (2002), The Hyperodapedon Biochron, Late Triassic of Pangea. Albertiana 27, p.30-38. Lucas SG, Heckert AB, Harris JD, Seegis D & Wild R (2001), Mammal-like tooth from the Upper Triassic of Germany. Journal of Vertebrate Paleontology, 21 (2), p. 397-399. Lucas SG & Oakes W (1988), A Late Triassic cynodont from the American South-West. Paleontology 31, p.445-449. Luo Z-X, Kielan-Jaworowska Z & Cifelli RL (2002), In quest for a phylogeny of Mesozoic mammals. Acta Palaeontologica Polonica 47 (1), p.1-78. Luo Z & Wu X-C (1994), The small tetrapods of the Lower Lufeng Formation, Yunnan, China. p.251-270, (Chapter 14 of) In the Shadow of the Dinosaurs - Early Mesozoic Tetrapods, (eds Fraser NC and Sues H-D), Cambridge University Press. McKenna MC & Bell SK (1997), Classification of Mammals Above the Species Level. Columbia University Press. Morgan GS & Lucas SG (1999), Type specimens of fossil vertebrates in the New Mexico Museum of Natural History and Science. New Mexico Museum of Natural History and Science Bulletin 16, p.253-259. Nath TT & Yadagiri P (2007), A new mammal-like reptile (Cynodontia) from Upper Triassic Maleri Formation of Pranhita-Godavari Valley, Andhra Pradesh, Journal of the Geological Society of India, 69, p.57-60. 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 & 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. Wouters G, Sigogneau-Russell D & Lepage J-C (1984), Decouverte d'une dent d'haramyide (Mammalia) dans le niveaux Rhetiens de la Gaume (en Lorraine Belge), Bulletin de la Société belge de Géologie, p.351-355. Zhang F (1984), The Fossil Record of Mesozoic Mammals in China. Vertebrata PalAsiatica, Vol XXII (1), p. 29-38, (as translated by Will Downs, May 1986 with minor revisions in 1999). This paper was obtained courtesy of the Polyglot Paleontologist.