B. CIMOLOMYIDAE & BOFFIIDAE

Taxon: Cimolomyidae Marsh, 1889 Boffiidae Hahn G & Hahn R, 1983 References: Marsh (1889), Discovery of Cretaceous Mammalia. Part II. Am. J. of Sci., 3, 38, p.81-92. Hahn & Hahn (1983), Multituberculata. in Westphal F (ed.), Fossilium Catalogus, I: Animalia, Pars 127. Kugler Publications, Amsterdam, p.409. Remarks: The affinities of these two families are uncertain. Both were previously placed within Taeniolabidoidea. What's a cimolomyid, mister? The following is based upon my reading of Sahni, 1972 (p.370-371). Cimolomyids were multituberculate mammals, and were rather popular in North America during the latter days of the Upper Cretaceous; small critters, which terrified tyrannosaurids predominantly by eating plants at them. When Marsh established the family in 1889, he never got round to confessing what it included (p.370). As, however, Meniscoessus is a close relative of Cimolomys, Sahni allowed both genera to join, and specified a total of six species: C. clarki, C. gracilis, M. major, M. robustus, M. conquistus and M. borealis. (Somewhat curiously, on page 373 of the paper, Sahni recognizes Cimolomys as having three valid species rather than the two just mentioned; C. trochus. I don't know why it wasn't included in the above list as well, but it could perhaps be connected with its then relatively recent date of description, 1969. If it's accepted as a valid species, then it must be seen as one of the family. And now we return to page 370). Family familiarities So seen a cimolomyid was a middling-sized to large multi for the time of Earth (which means a small animal), but still larger ones developed later. The lower p4 premolar is high and 'accurate' (a word I don't claim to understand as yet), but it bears less serrations than do the equivalents of most contemporary cousins and cousins' cousins. The rear two to three serrations are more distinct than the others. Grooves, caused by usage, occur on the labial face of the tooth leading back to a ledge at the rear, and this houses a number of cuspules (between one and three). Generally, but not always, later cimolomyids preferred to be somewhat larger than earlier versions. The upper P4 premolar plays a prominent role in distinguishing those two genera. The Cimolomys model is a long but low tooth. Meniscoessus is more modest in terms of length, but exuberant when it comes to height (p.371). The molars are harder to tell apart, but their owners presumably managed expertly and, in contrast to strangely derived primates with microscopes and puzzled expressions, consistently placed them in the appropriate jaws. The upper M1 has a well developed lingual cusp row, which ends close to the second cusp of the middle row. These teeth do become easier to distinguish in members after the Campanian age, as evolution's busy little elves rearranged the furniture into useful adaptations. Since Sahni's study, further discoveries and research have expanded the family. It may also iclude Buginbaatar of Asia and various further North American taxa. Outcasts

Link: Mikko Haaramo's Taeniolabidoidea Mikko Haaramo's Taeniolabidoidea Taeniolabidoidea, before it was redefined by Kielan-Jaworowska & Hurum, 2001.

Genera: Allacodon (?= Cimolomys), Boffius, Buginbaatar, Cimolomys (partly = Meniscoessus), Dripriodon (= Meniscoessus), Essonodon, Halodon (= Meniscoessus), Meniscoessus, Meniscoessus (partly = Cimolomys), Ptilodus (partly = Cimolomys), Selenacodon (= Cimolomys / Meniscoessus), Tripriodon (= Meniscoessus), other reports Time-Line: Paleocene: Boffius Upper Cretaceous: Buginbaatar, Cimolomys, Essonodon, Meniscoessus

Genus: Boffius Vianey-Liaud M, 1979 Family: Boffiidae Hahn G & Hahn R, 1983 Aka: Boffia?

Species:	Boffius splendidus Vianey-Liaud M, 1979
Place:	Hainin
Country:	Belgium
Age:	Lower Paleocene
Remarks:	Three specimens are at the AMNH, New York. This was a relatively large multi and is the only known member of its family.
Reference:	Vianey-Liaud (1979), Les Mammifères montiens de Hainin (Paléocene moyen de Belgique). Part I, Multituberculés. Palaeovertebrata, 9, p.117-131.

Genus: Buginbaatar Kielan-Jaworowska Z & Sochava AV, 1969 'Bügiin hero'

Species:	Buginbaatar transaltaiensis Kielan-Jaworowska Z & Sochava AV, 1969
Place:	Khaichin Uul-I (Bügiin Tsav)
Country:	Mongolia
Age:	Maastrichtian, Upper Cretaceous
Remarks:	The only known Upper K Mongolian multi not to belong to Djadochtatherioidea. Remains are incomplete and the assignment of B. to the Cimolomyidae is tentative, (Kielan-Jaworowska & Hurum 2001, p.408). This is presently the only mammalian taxon known from this location, to which it's also restricted. Kielan-Jaworowska et al, 2000 mentions a skull discovered by a Soviet-Mongolian Geological Excavation in 1968 (p.573). As well as being phylogenticly distinct from other Mongolian Cretaceous multis, the critter also lived somewhat later. That study also provides further information on pages 595-596. Affinities and characters It's clear this critter isn't a djado multi. Only a single premolar is preserved and there's no obvious sign any others were originally present. Additionally, the molars are all proportionately larger and have an excessive fondness for cusps. Other than for that, its affinities are presently unclear. However, it does seem to have more in common with North American fossils rather than other known Mongolian ones. There are similarities with Meniscoessus, eucosmodontids and Taeniolabididae. However, eucosmodontids retained front upper incisors and had arcuate lower p4 premolars. Buginbaatar seems to possess neither. That same p4, and the long and cuspy upper P4, aren't characters known from taeniolabids. As for its presence in Central Asia, it was presumably involved in Upper Cretaceous faunal exchanges with North America along various other terrestrial vertebrates. It could indicate Asiatic origins for some North American lineages, or the opposite configuration. Presently, a lack of non-djadoch Asian multis from the Upper Cretaceous leaves Buginbaatar a lonely heart in search of local company. Multi remains from Maastrichtian deposits were reported from Northern China a couple of years ago, and new mammal sites are likely the only way of providing closer information on affinities within the context of Asia. Dental and cusp formulae The dental formula per side isn't completely certain: (uppers): ?1 incisor, 0 canine, ?1 premolar, 2 molars; (lowers): 1, 0, 1 and 2 respectively. The cusp formulae (buccal to lingual): Uppers: P4 3:6:1; M1 7:8:6; M2 2:3:2. Lowers: p4 3 cusps with ridge; m1 5:6; m2 3:3. Holotype The type fossil has two catalogue numbers, PIN 3487-1 and 3487-2. These refer to partial upper and lower jaws in the collection of the Paleontological Institute, Moscow.
Reference:	Kielan-Jaworowska & Sochava (1969), The first multituberculate from the uppermost Cretaceous of the Gobi Desert (Mongolia). Acta Palaeontologica Polonica, 14, p.355-367.

Link: Acta Palaeontologica Polonica 42(2), 1997 http://www.paleo.pan.pl/acta/acta42-2.htm Kielan-Jaworowska & Hurum (1997). An abstract on Djadochtatheria, (since found by the same authors (2001) to be a superfamily; Djadochtatherioidea.

Genus: Cimolomys Marsh OC, 1889 Aka: Allacodon Marsh, 1889 (partly); Meniscoessus (partly); Ptilodus (partly); Selenacodon Marsh, 1889 (partly) Remarks: The following is based upon my reading of Osborn, 1893. Henry Osborn reviewed Upper Cretaceous material from Wyoming collected by (or for) Marsh and Cope, and this was supplemented by 400 additional mammal fossils obtained in 1892 by Dr Wortman and OA Peterson (p.331). Without attempting to assign any of the fossils to a particular species, he identified three genera of multituberculates. He termed them Ptilodus, Meniscoessus and Allacodon. As presently understood, Ptilodus is restricted to the Paleocene, so none of the specimens actually belonged to that taxon. They've been spread around to Cimolomys, Cimolodon and perhaps elsewhere. Osborn's 'Ptilodus' He concluded that Cimolomys, Cimolodon, Nanomys and (in part) Allacodon were all synonyms of Ptilododus (p.314). The actual situtation is more complex, but he was hampered by having little other than isolated teeth available. A few jaw fragments where able to help sort out the identiy and placement of premolars and molars. Three bits of dentary contained alveoli which showed a further, small lower premolar had been part of the dentition. However, it was small enough to evade capture. Even the helpful ants had been unimpressed. Pardon? Didn't you just write helpful ants? That's right. One good place to look for fossils like this is in an appropriate anthill, as some insects collect the things. A contact of mine is a rancher in eastern Wyoming, and the land contains lots of cow dung and outcrops of Hell Creek Formation rock. Some of these sites yield Cretaceous mammal fossils. One successful method of obtaining more involves raking the ground in a likely area and scattering grains of wild rice. This encourageous the ants to concentrate their activities. They collect up the cereal and other similarly sized stuff. That happens to include isolated Cretaceous teeth. Allacodon Marsh had based this genus on upper premolars (not P4s). They were broadly similar to teeth sported by Ctenacodon during the Upper Jurassic. Allocating them to a particular genus was nigh on impossible in 1893, so Osborn didn't attempt to do so (p.315). They seem to pertain to either (or both) Cimolomys and Cimolodon. Additional notes A. pumilus presently seems to be classed as Cimolomys sp. , according to the Peabody. Simpson referred it to a redefined Cimolomys gracilis (sensu Simpson) (Marsh, 1889) Simpson, 1929. A species is questionably present in the Straight Cliffs Formation of Utah, (Santonian). References: Marsh (1889), Discovery of Cretaceous Mammalia. Am. J. Sci, 3, xxxviii: p.177-180. Simpson (1929), American Mesozoic Mammalia. Mem. Peabody Mus. Nat. Hist. iii (i), p.1-235.

Reassigned species: C. bellus Marsh, 1889 see Cimolodon nitidus; C. digona see C. gracilis and Cimolodon nitidus; C. formosus Osborn, 1891 see Mesodma formosa; C. gracilus (Marsh, 1889) see (partly) Cimolodon nitidus, Mesodma formosa and Mesodma thompsoni; C. major Russell, 1936 see Meniscoessus major; C. minutus Marsh, 1889 see Cimolodon nitidus; C. nitidus see Cimolodon nitidus and Mesodoma thompsoni; C. primaevus Simpson, 1929 see Mesodoma primaeva; C. sculptus see Meniscoessus robustus

Species:	Cimolomys gracilis Marsh OC, 1889
Aka:	Cimolomys digona Marsh, 1889; Meniscoessus brevis; Ptilodus gracilus Osborn HF, 1893; Selenacodon brevis Marsh, 1889
Place:	Montana, South Dakota, Wyoming & Alberta, Sasakatchewan
Country:	USA & Canada
Age:	Maastrichtian, Upper Cretaceous
Remarks:	The type fossil’s from Wyoming. Rat sized, ca. 415g. Began its career as C., spent four decades as P. and was then brought back into C., courtesy of Simpson. A number of specimens are in the Peabody collection, Yale, with further specimens at the University of Wyoming. Much material has been subsequently referred to Cimolodon nitidus Marsh, 1889 and Mesodma (Marsh, 1889). This is rather messy and Marsh was clearly filled with nomenclatural enthusiasm in 1889. Ptilodus gracilis is considered to be valid by at least some authors. USNM 6076 at the Smithsonian Institute apparently goes by that name, (Wible & Rougier 2000, p.76).
Reference:	Marsh (1889), Discovery of Cretaceous Mammalia. Part II. Am. J. Sci. (3) 18, p.81-92.

Species:	Cimolomys trochuus Lillegraven JA, 1969
Place:	Alberta
Country:	Canada
Age:	Maastrichtian, Upper Cretaceous
Remarks:	The holotype is in the collection of the University of Alberta.
Reference:	Lillegraven (1969), Latest Cretaceous mammals of the upper part of Edmonton Formation of Alberta, Canada, and review of Marsupial-Placental dichotomy in mammalian evolution. The University of Kansas Paleontological Contributions, 50, p.1-122.

Links: University of Wyoming, Jason A Lillegraven http://home.gg.uwyo.edu/people/faculty/pubsArticles.asp?PersonID=136 Details on the interests and pursuits of a paleontologist. There’s also a link to the University’s fossil database, which can be searched. This contains a reference to Cimolomys nitidus. I treat this as a further synonym of Cimolodon nitidus. The University of Alberta http://www.biology.ualberta.ca/wilson.hp/UALVP/UALVPtypelist.html Type specimens in the UALVP Collections.

Species:	Cimolomys clarki Sahni A, 1972
Place:	Montana, Texas, Wyoming, ?Utah
Country:	USA
Age:	Campanian-Maastrichtian, Upper Cretaceous
Remarks:	The following is based largely upon my reading of Sanhi, 1972, which concerned fossils from the Judith River Formation of Montana. From what was then known to the author, the key to the door of this species is size (p.371). C. clarki is much like the later C. gracilis, but it's a smaller edition. Lower premolars The type fossil is such a tooth -a p4- but it, and all other available specimens were incomplete. However, assisted by a friend supplying otherwise absent information, a composite picture was possible. The tooth roof had something like eight or nine serrations sending ridges down the sides of the blade. These are short for the first two or three, lengthen for the rest, and the serrations are strongest for the rear couple (or trio). These rear serrations have grooves on their external surfaces, and a ledge to the back of the crown (posterolabial) is a strong feature. This contains the worn remnants of three cuspules. It's a double-rooted tooth, with the first partner being dominant. There's a natural cave towards the base of the front, and this would've been a hiding place for the shy and pathetic p3. All walls of this cavity, which is relatively small, are decorated with enamel with the exception of the rear one. Lower molars As is the multi way, there were two molars per mandible. Mischievously, number one shares both its size and basic cusp layout with Mesodma primaeva, and this restricts its diagnostic potentials. That indicates a cusp formula of 5-6 (labial) : 4 lingual and a length of 3.4mm plus or minus a bit. The m2 has the formula of 4:2. Apart from being a bit smaller, this is the same as for C. gracilis. The labial cusps tend towards being crescents, without quite attaining the quality, and deep valleys divide its four members. A ridge links the last of these with the lingual row, and it sometimes has a small cuspule on it. The two lingual cusps are large in size and achieve more height than the external crew. A further sidewards ridge often joins the front members of both rows. Upper incisors Perhaps the singular would've been better. A double cusped incisor is from a multi of an appropriate size, and was referred to this species. Upper premolars A pair of P4s were arrested for loitering at Clambank Hollow. The cusp formula (labial - lingual) is 2:5-6:2. The cusps of the middle row increase gradually in height from front to back, and have scratches on both sides. This tooth is low crowned, and thickened concentrations of enamel produce bulges at the base on both the external and internal sides. A ridge connects the highest middle cusp with further cusps on the rear labial corner of the tooth, a trait also favoured by C. gracilis. The occlusal outline of the whole tooth is something akin to a pinched rectangle. There were two tubular roots, but they've broken off. Upper molars A dozen or more M1s surrendered after the struggles at Clambank Hollow and, apart from their smaller size, they're not much different from C. gracilis. The middle cusp row, however, does have less members. The formula modelled by one specimen is 5:6:6. Labial cusps are shaped like cones and pyramids, and a ridge connects the first one to the leading cusp of the middle parade. Clear valleys lie between the other members of this labial brigade. The middle cusps are crescents with their open ends directed to the rear. The six lingual cusps become smaller from front to back. The front root of the tooth is long in terms of cross section (p.373), while its rear partner takes a broader approach, and it also has a couple of little accessory roots helping out with anchoring duties. An M2 has the following cusp formula: 2:3:4. The middle cusps are distinct from one another, with the final one being a crescent. A ridge connects it with the labial row. All four lingual cusps are pyramids. A further runs across the front of the crown connecting the leaders of all three groups, and this snuggles up intimately with the rear of the proceeding molar. This tooth has only the two main roots, and these are similar in size. The attitudes of premolars, a matter of taste While the upper P4 premolars are about 70% of the size for C. clarki (compared to C. gracilis), the lower p4 took a entirely different inclination. The two known for C. clarki average 4mm in length as opposed to about 3mm. All else being equal, this would be odd. So something wasn't equal. It may be recalled and, if so, then I have no need to mention it again, that a p4 for this species has eight or nine serrations. This number is low for multis of this time and place. Such serrations delivered enhanced shearing abilities and, whatever food tempted the attentions of Cimolomys most strongly, it presumably didn't require all too much shearing. Although the form of the tooth remained similar, its diminishment in size would've benefited these particular multis, in that this left more space for increasing the other food processing equipment. Holotype AMNH 77179 is a lower right premolar (p4) employed by the American Museum of Natural History, New York. It hails from Clayball Hill, Montana, and the specific name honours Captain William Clark, an early explorer of the River Missouri. Early, that is, for a white man. Plenty of people already know much about the subject. Additional notes Some specimens are in the AMNH, New York, whilst the OMNH, Oklahoma also has material. Probably weighed about the same as a malnourished rat, 300g.
Reference:	Sahni (1972), The vertebrate fauna of the Judith River Formation, Montana. Bulletin of the American Museum of Natural History, 147, p.321-412.

Link: NAMPFD, 12 mi NE of Worland http://flatpebble.nceas.ucsb.edu/nam/listfiles/12_mi_NE_of_Worland.html One of Dr John Alroy’s many carefully crafted mammal fossil inventories.

Species:	Cimolomys milliensis Eaton JG, 1993a
Place:	Kaiparowits Formation, Utah
Country:	USA
Age:	Campanian, Upper Cretaceous
Remarks:	The locality was apparently Mill Creek.
Reference:

Genus: Essonodon Simpson GG, 1927 Aka: Cimolodon (partly) Remarks: The inclusion of this taxon within the family is tentative, (Kielan-Jaworowska & Hurum 2001, p.408). Unfortunately, Simpson didn't provide the meaning of the name in his description, so a translation would be welcome. It's something-tooth.

Species:	Essonodon browni Simpson GG, 1927
Aka:	Cimolodon nitidus Marsh, 1889
Place:	Hell Creek, Montana; San Juan Basin, New Mexico; Wyoming; & Frenchman Formation, Saskatchewan
Country:	USA & Canada
Age:	Maastrichtian, Upper Cretaceous
Remarks:	The following is based upon my reading of Simpson, 1927. The holotype is a single lower molar; 2.4mm long and 3.6 wide (p.2). It's vaguely rectangular-ish in outline, but the lingual margin is shorter than the buccal. (As rectangles go, this one shouldn't be taken overly literally. There's not a straight line to be seen.) As traditional for lower multi molars, there are two rows of cusps running along the crown. Three form the buccal row with two on the opposing side. That pair are much larger, with the front one being the biggest cusp of all. This is usual for ptilodontids (as then used by Simpson). The tooth is triple-rooted, with two of them supporting the buccal side. The truth about the tooth A single tooth wasn't much to work with, but it was all that was available. It happened to contrast with other multi m2s, and the author had no doubts about establishing a new genus. It couldn't be accommodated in an established one. The length was proportionately less than for any known multi, and a total of only five cusps is low (p.3). Six is typical for an m2 excepting for artefacts caused by wear and tear. Some have more, but this was the first with five. Nevertheless, the shape, arrangement and ridged ornamentation of the cusps are broadly in keeping with ptilodontids (as then used). This is a specialised variation on that general theme. Holotype The type fossil, AMNH 14410, is an isolated lower molar (m2) in the collection of the American Natural History Museum, New York. It came from Crooked Creek in Montana. I presume the specific names honours the collector, Barnum Brown, but that's not stated. Additional notes Lofgren, 1995 reports on a single fossil, an m2 lower molar, collected from a McGuire Creek, Montana locality (p.94). It was a bit worn, had a length of 2.49mm and a cusp formula of 3:2 (buccal-lingual). It differed little from either the holotype or other known specimens (p.95). A mighty multi, which weighed in at over 2,5 kilos according, at least, to one molar-based estimate. Further material has been reported from the Frenchman Formation of Canada. Several specimens are at the Peabody, Yale, which were originally collected for Marsh. Wyoming University also has some material. One was originally diagnosed as Cimolodon.
Reference:	Simpson GG (1927), Mammalian fauna of the Hell Creek Formation of Montana, American Museum Novitates, 267, p.1-7.

LocalitiesThe first part of this essay has been written with reference to Norrell MA, Gaffney ES & Dingus L (1995), Discovering Dinosaurs, Little Brown and Company, 204pp. A relaxing trip to Hell It all started in 1901, when the director of the New York zoo went on holiday. As befitting for his job, Dr WT Hornaday loved animals, so he headed west to Montana in order to shoot some deer. Whether the luck went more with the primate or the ungulates is something I don't know, but he did happen upon an accumulation of fossilized bones. They were too large to do much with, so Hornaday took some photos. Back in New York he showed these to staff at the American Museum of Natural History, and interest was high. Barnum Brown had seen something similar previously. These bones were the last will and testament of a Triceratops, and he felt himself selected as the grateful heir. Accompanied by Richard Lull, Brown went west in 1902 and they found the dearly departed. They also met another playmate; Tyrannosaurus. Taking photos was far easier than carting the fossils off. For one thing, the nearest railway station was about 150 miles away. Nevertheless, they managed somehow or other. The location was part of the Hell Creek Formation; one of the most generous of fossil mines. Over a century has since rolled by and there's still plenty of Upper Cretaceous paydirt in them there hills (and valleys). The next section is based upon my reading of Simpson, 1927. 1907 Brown continued collecting at Hell Creek and elsewhere. In 1907 he reported his finds weren't restricted to obscure animals called dinosaurs. He'd also struck Mesozoic mammals (p.1). He had two lots of fossil teeth. One had been gathered from near Crooked Creek in Dawson County while the second, the Cameron Collection, is of less certain provenance; somewhere near Forsyth and Snow Creek. Although Brown produced a provisional list of the specimens, twenty years elapsed before descriptions appeared. They were in Simpson's 1927 paper. Since then many more taxa have been identified from Hell Creek locations. The Formation is widespread in parts of Montana, and sneaks over the state line into Wyoming. Similarly aged formations with fossils occur in a grand sweep of the North American Midwest, extending from New Mexico up to Alberta and Saskatchewan, and even into Alaska. The fossils Simpson had available were a pleasing appetiser for the continuing feast. Those interested in current excavations might dig the prospecting efforts of Wyoming rancher Frank Bliss. A gallery of photos is among the delights. This growing collection is presently undescribed, but studies have commenced. Further Mesozoic site summaries can be found at Localities. Links: Dinosaurs at a working cattle ranch (Bliss Ranch) http://www.cattleranch.org/pages/567495/index.htm Welcome to the Wyoming/Montana border where dinosaurs hide under the grass the cattle eat! Cretaceous "Hell Creek Faunal Facies"; Late Maastrichtian, Phillip Bigelow http://www.dinosauria.com/jdp/misc/hellcreek.html Essonodon features in this listing, which has a short commentary on Hell Creek. AMNH Novitates, Simpson, 1927 http://digitallibrary.amnh.org/dspace/bitstream/2246/3143/1/N0267.pdf You've read the webpage so now see the film, er, original paper. It's presently freely available in pdf format.

Genus: Meniscoessus Cope ED, 1882 Aka: Cimolomys (partly); Dipriodon Marsh, 1889; Halodon Marsh, 1889; Oracodon Marsh, 1889; Moeniscoessus; Selenacodon (partly); Tripriodon Marsh, 1889 Remarks: A quick introduction to the molars As is generally the case for multis, upper molars have three rows of cusps while lowers get by with two. A less usual characteristic is the shape of the cusps, as these form crescents in this genus. The uppers have the open end at the front while the situation is reversed in the lowers. This resulted in an excellent grinding system, and the broad principle is also favoured by some specialist chewers among placental ungulates. (That brief summary owes thanks to Osborn 1893, p.319). Nomenclatural spaghetti The following has been greatly aided by my reading of Simpson, 1936a. Meniscoessus is a nomenclatural minefield for the uninitiated. It could be thought of as chaotic, but that would be wrong. Early efforts resulted in far more names then desirable, but provided an orderly array of sorted specimens. This was essential for further work. Besides, the only people who can gain no benefit from hindsight, are those tackling something new. They can't learn from the mistakes of their predecessors and must either: a. get everything right (highly unlikely); or b. make mistakes for the enlightenment of themselves and others. Extra hurdles In the case of Meniscoessus, the fossils available did their best to make life difficult. They all insisted on being found as isolated teeth. You couldn't have checked them against more complete specimens, as there weren't any. An extra layer of potential confusion was added, when Othniel Marsh coincidentally used the same generic name for some dinosaur remains. This was invalid, but he wasn't aware ED Cope had already claimed title deed on Meniscoessus for some mammal molars from New Mexico. This certainly has its implications for anybody trying an internet search. Puzzling puzzles Marsh and Cope developed a serious mutual hatred, and quite why is unclear. It's therefore somewhat ironic, that various multi genera established by Marsh were soon synonymised with Cope's earlier published taxon of Meniscoessus. Pure speculation suggests this may have been both highly amusing and very annoying, depending upon to whom. This was thanks to an 1893 study by Osborn (p.1). Osborn reviewed the material described by Marsh in 1889, and reached the conclusion that various tooth forms were parts of a coherent series. Dipriodon robustus, interpreted originally as a left, rear upper molar (but actually a right lower); D. lunatus (a right m1 rather than an M1 or M2); and Halodon sculptus (that really is a p4 premolar although Osborn, 1893 appears to state it's a lower incisor!); all these teeth (and three further genera for other uppers and lowers) belonged together. His case was very persuasive despite not being able to point to a single fossil with associated choppers. At last Simpson credited Osborn as showing "remarkable insight" in his 1936 study, and he was well placed to judge. He'd come across a piece of jaw in the collection of the American Museum of Natural History, which had been harvested at some time by Miss Idella Kennedy. It had found its way to New York via the Carter County Geological Society of Montana. This convenient bit of Meniscoessus preserved the p4, both molars and the root of the third premolar as well. These teeth corresponded with both of Marsh's Dipriodon 'species' and his 'genus' of Halodon. "Hello!" Simpson might have said. It told him Osborn was right about those. Still a tangle Marsh had established nine species, all of which were later referred to Meniscoessus (p.2). Different species are probably involved. However, as the material available wasn't informative enough to be clearly distinguished, Simpson bundled them all together into M. robustus in 1929. Unfortunately, the jaw didn't shed much light on that problem. The m2 is about the same length as the holotype of M. robustus, but it's not as wide (p.3). There were no clear distinctions from the other two aforementioned holotypes, but the relevant teeth of the jaw weren't in good enough condition to reveal whether they represent a different species. Nevertheless, as the new jaw clearly belonged to the genus, AMNH 22708 was diagnosed as M. sp. (meaning some species or other). Limited novelty and down with concrescence The fossil didn't add much new to the knowledge about individual teeth, but it was kind enough to give an illustration of the roots. This picture was already known to some, but hadn't burrowed deeply into the literature. The postcanines have supernumerary roots. There's a main one at both the front and back, and these start at their tops with widths similar to those of the crowns. However, each tooth has at least one extra little helper. This is in the middle of the lingual side for the premolar (p4). Of the two molars, m1 has a pair of extra roots on both sides, while m2 is a bit less endowed; one lingual and two buccal. Of that latter duo, one isn't fully separated from the main front root (p.4). (At this juncture, Simpson alerts the reader to a foot note. This briefly addresses ideas postulated by: "Adherents of the moribund concrescence theory of molar evolution..." This 'moribund theory' doesn't particularly awake my curiosity. There seems to have been a view that molars began their career as multi-rooted, and the number declined over time. Suffice it to add: "... just two roots on lower cheek teeth is primary..." (The sentence is rather long.) This statement was subsequently supported by the expert testimony of Morganucodon, a genus supplying evidence from the Upper Triassic and Lower Jurassic.) Gaining a sense of proportions As all previous Meniscoessus specimens were isolated, there was a lack of certainty regarding the proportional dimensions involved, as nothing could be reasonably assigned to one individual animal. Most multis then known as ptilodontids had p4s which are longer than either molar. It's shorter than either in this genus. The m1 is a bit longer than m2. Generally, the first molar is much longer; frequently double the length of the second. Relatives It had earlier been suggested that this species was closely related with, and even perhaps ancestral to Taeniolabis. The new jaw reconfirmed this isn't so. It spoke of no particular taeniolabidid traits. Various details, including the relative proportions, failed to reveal an intimate relationship with any of the then accepted ptilodontids. "It appears to be a specialized, aberrant Cretaceous ptilodontid." A connection with something like the later Eucosmodon couldn't then be ruled out, but the author deemed it 'improbable'. The present position is in line with that. Additions from other sources The history of this genus is complicated and confusing. It’s attributed to Cope, 1882. Later, this was joined by "Meniscoessus" Marsh, 1889. The second usage apparently related to teeth described as belonging to small, carnivorous dinosaurs. Further names included Tripriodon and lord knows what else, including Triprotodon. Close similarities were then noticed with an already established dinosaur genus, Paronychodon Cope, 1876, also based on teeth from the Laramie Formation. Over time, an impressive school of names was synonymized under P. However, this is now considered a nomina dubia, and a non-dinosaurian dubious name to boot. In 1929, Simpson published American Mesozoic Mammalia, (Mem. of the Peabody Museum, 3 pt. 1; i-xv). The name Tripriodon, (‘three saw tooth’), was resurrected. These ‘theropod’ teeth were actually mammalian. The mammal T. since seems to have also gone the way of the dubious dino. At least, that’s my best shot at interpreting what I’ve read. It doesn’t equip me to make sense of the Peabody specimen YPM 11852. Its history is given as Tripriodon caperatus Marsh, 1889 holotype; Meniscoessus caperatus holotype; Meniscoessus robustus (sensu Simpson) (Marsh, 1889) Simpson, 1929; Paronychodon lacustris Cope, 1876. Strange. Be that as it may, the holotype of the mammalian M. robustus definitely began its career as Dipriodon robustus. Hunter & Archibald, 2002 say so, (p.196). Paronychodon lacustus Matthew, 1916 (p.477) adds either clarity or more mud. Cope described this as reptilian in 1876. However, in a review of North American Cretaceous mammals from 1893, Osborn noted similarities with incisors from Meniscoessus, and suggested that could be possibly be its true identity. However, as subsequent dino researchers have carried on using the name for dino teeth, it appears that a provisional referral to a multituberculate was incorrect. Osborn, 1893 Osborn synonymised the following names with this genus on page 317; Tripriodon, Selenacodon, part of Halodon and questionably Oracodon. The final one mentioned was based on upper premolars, which hadn't then been identified for Meniscoessus. Marsh had named two species; O. anceps and O. conulus (p.319). This assignment was tentative as neither matched well enough with any of the available lower p4s. Be that all as it may, Meniscoessus Cope is a valid multituberculate, and is known from some quite good remains, as well as a great many teeth. Possible specimens have also been reported from Utah. Reference: Cope (1882), Mammalia in the Laramie Formation. American Naturalist, 16, p.830-831. Link: American Musuem of Natural History Archives http://digitallibrary.amnh.org/dspace/bitstream/2246/2143/1/N0825.pdf The study I've termed Simpson, 1936a is presently freely accessible in pdf format.

Reassigned species: M. brevis see Cimolomys gracilus; M. coelatus see M. robustus; M. fragilis see M. robustus; M. greeni Wilson, 1987 see M. robustus; M. lunatus see M. robustus; M. sculptus see M. robustus

Link: Anne Weil http://www.ucmp.berkeley.edu/mammal/multis/meniscoessus.jpg A photo of a reconstructed lower jaw. Triebold Paleontology, Inc. http://www.trieboldpaleontology.com/retail/bonesnteeth/smtms.htm Another free and entirely voluntary advert. A cast of a fairly complete skull, 7cm long. Yours for $74,95. If anyone feels like buying me one, I wouldn’t mind.

Species:	Meniscoessus conquistus Cope ED, 1882
Place:	Wyoming, Colorado, South Dakota & St Mary River Formation
Country:	USA & Canada
Age:	Maastrichtian, Upper Cretaceous
Remarks:	Matthew, 1916 (p.447) provides some insights. Dr JL Wortman collected three 'mammal' fossils from the Lance Formation of Wyoming, and Cope referred all of them to this new species. There were two teeth and part of a humerus. One of the teeth turned out not to be a multi premolar after all. It inconveniently belonged to an anylosaur dinosaur. The partial arm bone is mammalian, but rather small for Meniscoessus, at least in the way the genus was used ninety years ago. Holotype According to Osborn, 1893, the type fossil was an upper molar (M2). Additional notes This was the first North American Cretaceous mammal to be named and described, (Kermack et al 1965, p.536). The species has been referred to the 'M. robustus group', (Hunter & Pearson, 1996, p.636).
Reference:	Cope (1882), Mammalia in the Laramie formation, American Naturalist xvi, p.830-831.

Link: The AMNH, Catalogue number 5735 http://paleo.amnh.org/fossil/FRC.image_card?img_id=5735-01 Reportedly, these Triceratops bones were found with the type of Meniscoessus in 1882 in South Dakota. Cope’s material was from the Laramie Formation of Colorado, whilst Marsh’s seems to be from Wyoming. Perhaps it moved. Puzzling.

Species:	Meniscoessus caperatus (Marsh, 1889)
Aka:	Tripriodon caperatus Marsh, 1889
Place:	Wyoming
Country:	USA
Age:	Maastrichtian, Upper Cretaceous
Remarks:	This is supposed to be Marsh’s 'theropod' and an example resides at Yale. However, Osborn, 1893 appears to list it as both lower incisors (p.318) and an upper molar for this genus (an M2 (p.319)), although all under the name of Tripriodon caperatus. I don't pretend to understand how it can be both of those very different teeth.
Reference:

Species:	Meniscoessus robustus (Marsh OC, 1889)
Aka:	Cimolomys sculptus; Dipriodon lacunatus; D. lunatus Marsh, 1889; D. robustus Marsh, 1889; Halodon sculptus Marsh, 1889; M. coelatus; M. fragilis; M. greeni Wilson RW, 1987; M. lunatus; M. sculptus; Moeniscoessus robustus; Oracodon anceps Marsh, 1889; O. conulus Marsh, 1892; Selenacodon fragilis Marsh, 1889; Tripriodon coelatus Marsh, 1889
Place:	Lancian Formation, Wyoming; Hell Creek F., Montana, North Dakota, South Dakota; & St Mary's River Fm., Alberta; Frenchman Fm. & possibly Ravenscrag Fm., Saskatchewan
Country:	USA & Canada
Age:	Maastrichtian, Upper Cretaceous - Paleocene?
Remarks:	The first part of this rather episodic entry is based upon my reading of Lofgren, 1995 (p.93-94). Lofgren's study concerns over thirty postcanines rescued from both Lancian and Puercan localities at McGuire Creek, Montana. Accepted at face value that would suggest this species survived the K-T extinction event(s). However, it seems more likely some of these specimens were reworked by mischievous river action cutting down during the Paleoncene into fossil-bearing Cretaceous deposits, and that lead to things getting somewhat mixed. All donors probably dropped dead during the Upper Cretaceous. I've combined figures from two tables into the following list of lengths, and they only concern McGuire Creek localities. Postcanine teeth lengths Uppers: P4 (2 specimens) 4.25-4.42mm; M1 (3 sp.) 9.05-9.82mm; M2 (9 sp.) 6.75-8.39mm. Lowers: p4 (1 sp.) 7.09mm; m1 (7 sp.) 7.54-8.72mm; m2 (4 sp.) 6.40-7.70mm. On average McGuire Creek specimens from apparently Paleocene sites are a bit larger (p.94) and somewhat more numerous than those from the single relevant Upper Cretaceous locality. However, the sample sizes involved aren't large and, in terms of size and morphology, all seem to be from the same species. While it may have been a survivor of the K-T party, all the apparently Paleocene fossils occur in river channel deposits. None were found in other Paleocene localities in Montana or, for that matter, in the whole of North America. This pattern was reflected among various other typically Cretaceous taxa from McGuire Creek. That factor, along with analysis of pollen remains from these sites (overwhelmingly Paleocene), indicates reworking is the most likely explanation. Holotype The holotype of M. robustus, YPM 11234, is a right lower molar, (m2). It came from the Lance Formation of Wyoming and presently resides in the Peabody Museum, Yale, (Hunter & Archibald 2002, p.196). Additional notes Marsh authored a swamp of names. At least one source of confusion was the referral to teeth from differing positions in the mouth to different genera. A body weight guestimate is 3.3kg. The Peabody is home to a pile of specimens, though what they all actually are lies beyond the scope of my interest. More are in the Wyoming collection. The first usage of M. robustus seems to go back to Osborn, 1891. Getting ahead This is the only species for which skull material has so far been found, (Weil 1999, p.67). It's also the largest and most derived, though the specimens may belong to a number of closely related species, (Hunter & Pearson 1996, p.636-637). This paper refers to the 'M. robusts group' which includes M. conquistus, M. robustus and M. borealis. (Both those possible species presently occupy separate entries on this page.) The first mentioned is the smallest in size, whilst borealestes is the largest. About half of the lower p4 premolars assigned to this taxon have nine serrations. Clemens et al, 2003 report that the length of molars can reach nearly 10mm. Amongst Mesozoic multis, this is only exceeded by Bubodens magnus (Taeniolabididae?), which seems to be awaiting publication very patiently. (What I originally read led me to believe that'd be in about 2004!) Synonym corner According to Osborn, 1893, Oracodon anceps and O. conulus were upper premolars, and he tenatively assigned them to this genus (p.319). Selenacodon fragilis was established on the basis of an M1 upper molar. M2s were supplied by M. conquistus and Tripriodon coaelatus (p.319). On page 318 he seems to state Halodon sculptus is based on lower incisors, whereas Simpson, 1936a has this as a lower premolar (p4). Relevant lower molars had been termed Dipriodon lunatus (m1). These generally had fove cusps in the buccal row and four in the lingual one. D. robustus was established for the shorter m2s, and the cusp rows contained four and two cusps respectively.
References:	Marsh (1889), Discovery of Cretaceous Mammalia part 1, American Journal of Science, 38, p.81-96.
	Osborn (1891), A review of the Cretaceous Mammalia. Proc. Acad. Nat. Sci., Phila., p.124-135.
	Wilson (1987), Late Cretaceous (Fox Hills) multituberculates from the Red Owl local fauna of western South Dakota. Dakoterra 3, p.118-122.

Species:	Meniscoessus borealestes Simpson, 1927
Place:	Hell Creek, Montana
Country:	USA
Age:	Maastrichtian, Upper Cretaceous
Remarks:	It's possible this is synomynous with M. robustus, but I've decided to provide it with its own enclosure. The systematics of this genus are far from settled, as more cooperative specimens would be required to resolve a number of difficulties. The following is based on my reading of Simpson, 1927 and I'm going to stick to the terminology in that paper. Simpson had a number of isolated teeth he referred to this genus, with the holotype being an upper M2 molar (p.3). While well preserved it had suffered some wear. The three rows of cusps beloved by upper multi molars, (except in basal taxa), number 4, 4 and 5, counting from the buccal to lingual sides. Crown measurements are: length 8.3mm, width 7.0. In frightningly technical terminology, this is big. The quartet of buccal cusps differ a bit in size and the external face of each is rounded. Each is blessed with sharp ridges running from the summit, which travel inwards and forwards. The front cusp has branching of the forward directed ridge. The cusps of the middle row are the largest, while those in the lingual brigade are smaller but more similarly sized. Relatives and further specimens As the tooth is consistent with the already established genus, (known originally from New Mexico (p.4)), Simpson assigned this species to it. However, it's 20% larger than M. conquistus, and the anterior lingual cusp is more clearly separated from the second. This justified a different species. A small number of other teeth were somewhat tentatively referred to the genus; a badly broken M1, an upper and a lower incisor, and a nearly seven millimetre long premolar. This p4 was in reasonable condition and bore eight serrations. Holotype The holotype of M. borealis is AMNH 14411, an isolated upper molar (M2). It lives in New York and was collected from Crooked Creek, Montana. No reason was given for the specific name, but it presumably reflected the relative northiocity of the location.
Reference:	Simpson GG (1927), Mammalian fauna of the Hell Creek Formation of Montana, American Museum Novitates, 267, p.1-7.

Species:	Meniscoessus major (Russell LS, 1937) Sahni A, 1972
Aka:	Cimolomys major Russell, 1937
Place:	Alberta & Montana
Country:	Canada & USA
Age:	Campanian, Upper Cretaceous
Remarks:	The following is based largely upon my reading of Sanhi, 1972, which concerned fossils from the Judith River Formation of Montana. The small giant The specific name refers to its comparative body size, and has been rendered delightfully misleading (p.374). Originally, Russell was comparing it to the then other species of Cimolomys which, at the time, was known to be a 'genus' containing unrelated critters. As the fossils were rather sparse, classifying them more closely was seen as insecure. Any multi from the Oldman Formation was able to qualify for inclusion. In those terms, it's large. However, after being placed among its proper kith and kin within Meniscoessus, M. major happened to be minor for this troop of multi hefties. It was the smallest known species. Nevertheless, it may well have possessed oodles of charisma and courage and, as far as I'm aware, none of the remains are incompatible with this possibility. The physical characterisation is built on teeth. When seen from the side, the p4 lower premolar has a regular arc of an outline adorned with 11 serrations. A cavity at the front provided a kennel for a small p3. The first lower molar is exceptionally cuspy for the age. With regard to the upper molars, the first of those is comparatively wide, and its partner, M2, was proud to own an accessory root to help hold its crown in place. Lower incisor A partial incisor was tentatively referred to the species. It's large and deeply grooved on its labial side, and that's in line with expectations. Lower premolars No complete p4s were harvested from the Judith River Formation deposits, but three partial ones were. These indicate a high and symmetrical arc-shaped blade, with its highest point being reached by the fourth of fifth serration. The final two or three are the most distinctly formed serrations, and grooves etched into their labial sides lead to a ledge at the rear on that face of the tooth. Lower molars No m2s were identified in the swag, but there was a single m1 of a length similar to the equivalent tooth of the holotype. The two cusp rows contain 8 (labial) and 6 (lingual) members, which is more than for M. robustus. This cusp count is also known for Cimolomys gracilis, but other features rule out a close affinity. The smallest of the cusps is the front one of the labial row. It's conical whereas its followers deigned to develop more pyramid-like structures. Grooves have been carved into both lateral faces along this row. The lingual cusps, the latter two of which are not fully separated, are taller than their colleagues, and grooves only occur on their external sides. The front root is a bit larger than the rear one, and a third small root helps out with support duties. Upper premolars A pair of P4s was discovered. The cusp formula isn't entirely clear: 2:4 or 5:2 or 3 (labial-lingual). Of the labial duo, it's the rear cusp that's the largest. The other two rows are kept apart by a deep valley. Those of the middle file increase modestly in height along the line. Scratches have been vertically cut into the cusps of both these latter rows, and this violence must be blamed on the lower teeth. The roots have gone elsewhere, but their bases are approaching roundness and of similar dimensions. This P4 has a cusp in the middle row that doesn't have a counterpart on those of M. borealis. A milk premolar may also belong to this species. Upper molars An M1 is too poorly preserved to reveal its cusp count (p.376). The remaining cusps are conical, and grooves and scratches present signs of use. As an M2 compares well with later members of the genus despite much wear, it belongs to Meniscoessus. However, its cusp formula is also unclear. The labial row probably contained tow or three, there were four in the middle and some number or other in the eroded lingual team. The front root is a bit bigger than the rear one. Sizes and distinguishments There are similarities between the teeth of Meniscoessus and Cimolomys, but the final upper premolar and pair of molars distinguish the two. For this species, these teeth happen to be only 70% the size as for M. robustus and are 30% smaller then M. conquistus (M2 only). The blade of the lower p4 is also broadly similar for both genera; a high and symmetrical arc with between 8 and 11 serrations. This is a relatively low number for multis of the age. Further details also correspond. The upper P4, however, is proportionately longer and lower for C. in comparison to the same features of the molars. Holotype NMC 873 is a lower mandible from the Oldman Formation of Alberta, and it's being held hostage at the National Museum of Canada. Additional notes 1936 is sometimes blamed for the original publication, but 1937 seems to be correct. A weight estimate put the body mass at around a kilo.
References:	Russell (1937), New and interesting mammalian fossils from Western Canada, Transactions of the Royal Society of Canada, 30 (4), p.75-81.
	Sahni A (1972), The vertebrate fauna of the Judith River Formation, Montana, Bulletin of the American Museum of Natural Histor, 147(6), p.321-412.

Species:	Meniscoessus ferox Fox RC, 1971
Place:	Upper Milk River Formation, Alberta
Country:	Canada
Age:	Campanian, Upper Cretaceous
Remarks:	The holotype, collected in 1968, is in the Alberta collection.
Reference:	Fox (1971), Early Campanian multituberculates (Mammalia: Allotheria) from the upper Milk River Formation, Alberta. Canadian J of Earth Sci., 8, p.916-938.

Species:	Meniscoessus intermedius Fox RC, 1976
Place:	Oldman Formation, Alberta & New Mexico, Utah & Wyoming
Country:	Canada & USA
Age:	Campanian - Maastrichtian, Upper Cretaceous
Remarks:	Another Alberta type fossil. A further specimen works at the AMNH, New York. Weighed about as much as a large rat, 500g.
Reference:	Fox (1976), Cretaceous mammals (Meniscoessus intermedius, new species, and Alphadon sp.) from the lowermost Oldman Formation, Alberta. Canadian J. of Earth Sciences, 13(9), p.1216-1222, 4 figs.

Species:	Meniscoessus collomensis Lillegraven JA, 1987
Place:	Williams Fork Formation, Colorado
Country:	USA
Age:	Upper Cretaceous
Remarks:	Known from only one site. Weight estimate, 1,4kg.
Reference:	Lillegraven (1987), Stratigraphic and evolutionary implications of a new species of Meniscoessus (Multituberculata, Mammalia) from the Upper Cretaceous Williams Fork Formation, Moffat County, Colorado. Dakoterra 3, p.46-56.

Species:	Meniscoessus seminoensis Eberle JJ & Lillegraven JA 1998a
Place:	Ferris Formation, Wyoming
Country:	USA
Age:	Campanian-Maastrichtian, Upper Cretaceous
Remarks:	3,5cm of lower jaw, found near the Seminoe mountains. It seems to be in the collection of the University of Wyoming. A close resemblance to M. robustus. Somewhere between ‘rat-sized’ and 3,5 kilos, depending upon which source. An appealing alternative name for this location is Leave No Toad Unturned.
Reference:

Link: Canadian Museum of Nature http://www.museevirtuel.ca/Exhibitions/Dinos/English/cmn/cmn07/cmn07a.html This feature’s fun for non-experts of all ages. There’re plenty of things to click. The arrow on the right calls up a the second part of the story. The inset photo is a link to a full image of the dentary. There are also nice features on dinosaurs and paleontology. The Royal Tyrrell Museum of Paleontology demystifying the Cretaceous.

Species:	Meniscoessus bustus
Place:
Country:
Age:
Remarks:	This is listed in the Biosis Index. Perhaps it's a variant of robustus.
Reference:

Species:	Meniscoessus coelatus
Place:
Country:
Age:
Remarks:	Unless any contrary information arrives, I am assuming this to be synonymous with Tripriodon coelatus, which is listed with Meniscoessus robustus below. There’s one hanging around at the Peabody, if anyone wants to interview it.
Reference:

Other reports: Xxxxxxxxxxxxxx Xxxxxxxxxxxxx

A. ‘Basal’ Cimolodonta B. Cimolomyidae & Boffiidae C. Kogaionidae

C. KOGAIONIDAE

Taxon: Kogaionidae Rădulescu R & Samson P, 1996 Reference: Rădulescu & Samson (1996), The first multituberculate skull from the Late Cretaceous (Maastrichtian) of Europe (Hateg Basin, Romania), Anuarul Institutului de Geologie al României, Supplement 1, 69, p.177-178. A European family of mini-multis, which also survived into the Tertiary. Its affinities within Cimolodonta are unclear. "Share with Taeniolabiodoidea the general shape of the skull, with anterior part of zygomatic arches directed roughly transversely and very short basicranial region, which gives the skull a square-like appearance, but differ from them in having a strongly elongated snout and different dentition," (Kielan-Jaworowska & Hurum 2001, p.418). I can more or less understand that! In polite company, zygomatic arches are known as cheek bones. An alternative view on this family is provided by Peláez-Campomanes et al, 2000. These authors placed Hainina within Kogaionidae, and came to the conclusion that the family isn't part of Cimolodonta. Hainina has both a large and blade-like lower premolar (p4) and a short upper P4, (p.709). These features were then otherwise only known from the Lower Cretaceous multis Eobaatar and Arginbaatar, both of which have five upper premolars; a 'primitive' trait in Multituberculata. As a short upper P4 is not present in other derived multis, this is said to suggest that: "The blade-like p4 was independently acquired by Late Cretaceous advanced Ptilodontoidea. Probably such a feature was independently acquired by the family Kogaionidae as well, since no indication for a common origin of Arginbaatar, Kogaionidae and Ptilodontoidea can be found. "Therefore, the family Kogaionidae cannot be related to any of the described Late Cretaceous and Paleocene multituberculates included in the Cimolodonta (McKenna and Bell, 1998). Its closer relatives must be found among the Early Cretaceous Plagiaulacida (sensu McKenna and Bell, 1998)." I think it fair to say this is a minority point of view, (which doesn't mean it's incorrect). On a minor note, McKenna and Bell's publication dates from 1997, at least according to my copy and every other reference I've ever noticed for it. Genera: Barbatodon, Hainina, Kogaionon, Paracimexomys? (= Barbatodon), other reports Time-Line: Paleocene: Hainina Upper Cretaceous: Barbatodon, Kogaionon

Genus: Barbatodon Rădulescu R & Samson P, 1986 Aka: Paracimexomys?

Species:	Barbatodon transylvanicum Rădulescu R & Samson P, 1986
Aka:	Paracimexomys? dacicus Grigorescu & Hahn, 1987
Place:	Pui, Hateg Basin
Country:	Romania
Age:	Maastrichtian, Upper Cretaceous
Remarks:	The following is based upon my reading of Grigorescu & Hahn, 1987, which appeared after the original publication of the genus and briefly describes fossils referred to as Paracimexomys? dacicus. Thanks are due to the supplier. Two molars were obtained in an area of Romania known in German as Siebenbürgen. The English name, which is actually a reference to forest, tends to conjure up bizarre associations for English speakers based upon our ignorance, literary inheritance and growing up watching films starring Boris Karlof, Peter Cushing and The Rocky Horror Picture Show; Transylvania. Still, if that's your cup of tea, then strip down to your underwear, pull on your fishnet stockings, and let's do the time warp again to the latter days of the Upper Cretaceous. If that last sentence leaves you scratching your head, then you clearly haven't seen The Rocky Horror Picture Show. Doing the time warp The pair of localities yield fossils from Maastrichtian deposits in the Hateg Basin, an exceptionally good place for land vertebrate remains (p.237). There are a considerable number of fine vertebrate localities of this age in Europe, at Maastricht for example, but, as much of the continent was under the sea at the time, they're generally marine deposits. The Hateg Basin is different. There was an archipelago of islands there, and the wildlife inhabiting them was rich and varied. Descriptions began appearing towards the end of the 1800s; dinos (including birds), crocs, turtles, fairly rare pterosaurs. One of the paleontological claims to fame for the Hateg Basin is the tendency towards small body sizes among the dinos, an adaptation to the limited supply of land, and that's been exploited for cute episodes of television programmes. Varied as the known wildlife was, the local population were saddened by a great misfortune. They had a lack of Mesozoic mammals. Apart from interventions provided by shear good fortune, remains of such small critters tend to only come to light when specialized methods of searching are employed. In Romania, success was finally achieved in the 1980s. Grigorescu managed to persuade large outcrops near the village of Sinpetru to provide a multituberculate incisor. Much less extensive outcrops also occur 20km further east at the village of Pui, and these are fairly often submerged beneath flood water. Despite this uncooperative habit, and not wishing to be outdone by the Sinpetru upstarts, this place was talked into giving up a couple of lower molars (p.238). Further fossils came from the area during subsequent research. At the time, the fossils were held in private collections, and hadn't been given catalogue numbers. Beside something? The species was tentatively referred to the otherwise North American genus of 'beside bug mouse', more usually called Paracimexomys. Views on that have since changed in the light of further mutli specimens. Be that as it may, Paracimexomys largely consists of broadly similar teeth drawn from several hard-to-untangle lineages. The traits shared are presumably basal for cimolodontan multis, and not necessarily all that informative when it comes to relationships. The type fossil of P.? dacicus, an m1 lower molar, has a cusp formula of 4:3 (buccal to lingual), and the final buccal cusp is equipped with a cingulum-like feature. That isn't the case for other species of Paracimexomys. An m2 had also been identified; cusp formula 3:2. Lower molars It wasn't certain that both teeth belonged to the same species but, as they plausibly could've done, they consented to share a taxon on at least a trial basis. Both are primitive for cimolodontans. As nobody had knowingly seen a Paracimexomys incisor at that time, the Sinpetru tooth couldn't be even tentatively allocated. The m1 is roughly rectangular in outline, has a length of 3.4mm and a width of 1.6. The difference between those two measurements indicates a first rather than a second molar and, with no sign of either a third cusp row or a ridge in such a position, it has to be a lower one. First molars of P. priscus are rounded at the front with a diagonally straighter rear caused by an elongation of the buccal side. As that also applies here, it's a left m1 rather than a right one. Cusps are all well defined, and the lingual ones are higher. That also serves to confirm which side is which. In the buccal row, the first cusp is round and eroded at its tip. The second is longer and higher. An elongation runs inwards from its rear, and does its best to block the central valley between the rows. Cusps three and four are half the size, and a cingulum-like structure runs from the rear of the fourth, forwards along the buccal crown margin, and then peters out by the valley between buccal cusps two and three. This structure has no counterpart on species it was then compared with, and it was termed "peculiar" in the Discussion section. It may have been a specific oddity. The m2 is a smaller, squarer tooth with a length and width of about 1.25mm. If it does belong to the same species as the m1, and the sizes are something like typical for it, then the difference in length between the two would be extreme. There are three cusps on the buccal side and two in the lingual row. As with Paracimexomys (but less so than for Kryptobaatar, another comparison considered, the m2 cusps are not as strongly individualized as those on the m1. Both molars share an overall similarity of structure with teeth known from both those genera. Affinities? Given the time of Earth, the number of cusps is comparatively low for multis, a condition otherwise known (in 1987) only for Paracimexomys and Kryptobaatar. The actual figures of the Romanian teeth, m1 4:3 and m2 3:2, were matched only by P. priscus and Krypto (p.239). Along with other similarities, this resulted in those taxa being the closest for comparisons. The authors found both molars most closely resembled those of the first mentioned, and that prompted their tentative referral to the same genus, since revoked. However, the molars differed in details from P. priscus, an example being the cingulum-like structure of the m1. Holotypes I don't yet know the details of the type of Barbatodon, and P.? dacicus had no number. That specific name was based upon Dacia, the Latin term for Romania. Additional notes "Based on comparisons with the m1s from Vălioara, the holotype of Barbatodon is regarded as a kogaionid m1," (Csiki Z & Grigorescu D, 2001, Fossil mammals from the Maastrichtian of the Hateg Basin, Romania, 6th European Workshop on Vertebrate Palaeontology - Florence and Montevarchi (Italy) - September 19-22, 2001, p.26). (Kielan-Jaworowska & Hurum 2001, had it placed tentatively in the informal Paracimexomys group). At one time, this genus was "tentatively synonymized with Hainina", (the abstract of Csiki Z & Grigorescu D (1999), New data on the Multituberculate Mammals from the uppermost Cretaceous dinosaur-bearing deposits of the Hateg Basin, IV European Workshop on vertebrate paleontology (1999) Albarracin, Spain, p.49-50).
References:	Rădulescu & Samson (1986), Précisions sur les affinités des Multituberculés du Crétacé supérieur de Roumaine. C R Acad Sci II: Mec-Phys, Chim, Sci Terre, Sci Univ 303p, p.1825-1830.
	Grigorescu D & Hahn G (1987), The first multituberculate teeth from the Upper Cretaceous of Europe (Romania), Geologica et Palaeontologica, 21, p.237-243.

Links: Society of Vertebrate Paleontology News Bulletin, No. 173 http://www.vertpaleo.org/bulletin/173.pdf Mentioned in the report on Romania starting on page 25, as is Kogaionon. Abstracts of the 6th European Workshop on Vertebrate Palaeontology, 2001 http://digilander.libero.it/tulse/6ewvp/abstractbook%206ewvp.pdf Recent views on the affinities of this genus. See page 26.

Genus: Hainina Vianey-Liaud M, 1979 'from Hainin' Remarks: H. was originally referred to ?Cimolomyidae. "We assign Hainina to the Kogaionidae (superfamily incertae sedis); it differs from Kogaionon in having ornamented enamel, while the enamel is smooth in Kogaionon," (Kielan-Jaworowska & Hurum, 2001, p.409). Material has also been reported from Romania, which is presently mentioned in the 'Other Reports' section. A concise comparison of what they term primitive, derived and unique features for this genus has been provided by Peláez-Campomanes et al, 2000 (p.709). In summary: - The first upper molar, (M1), has a small number of cusps, (no more than seven in the longest row). Primitive. But, they're arranged in three rows. Derived. The lingual row is the longest. Uniquely derived. - The two final premolars in the upper series are similarly sized. Primitive. But they have two rows of cusps. Derived. - The buccal cusp row on the last of these two premolars shows a decrease in cusp size going backwards. The lingual row extends only along two-thirds of the rear of the tooth. Unique. - The lower p4 premolar is large and blade-like. Derived. However, it's nigh on twice the length of the upper P4 (P3 to others, see H. pyrenaica below). Primitive. - The lower molars have few cusps in each row. Primitive.

Reassigned species: H. godfriauxi partly see H. vianeyae

Species:	Hainina belgica Vianey-Liaud M, 1979
Place:	Hainin
Country:	Belgium
Age:	Paleocene
Remarks:	This is the type species of the genus. The remains consist of nine teeth, (Peláez-Campomanes et al, 2000, p.702).
Reference:	Vianey-Liaud (1979), Les Mammifères montiens de Hainin (Paléocène moyen de Belgique). Part I. Multituberculés. Paleovertebrata 9, p.117-131.

Species:	Hainina godfriauxi Vianey-Liaud M, 1979
Place:	Hainin
Country:	Belgium
Age:	Montian, Middle Paleocene
Remarks:	The species is based on five teeth, (Peláez-Campomanes et al, 2000, p.702). 24 specimens from the somewhat younger location of Cernay, France, were subsequently used as part of the basis for H. vianeyae.
Reference:

Species:	Hainina pyrenaica Peláez-Campomanes P, López-Martinez N, Àlvarez-Sierra MA & Damms R, 2000
Place:	Tremp Formation, southern Pyrenees
Country:	Spain
Age:	Danian, Lower Paleocene
Remarks:	The following is based upon my understanding of Peláez-Campomanes et al, 2000. H. pyrenaica was a small species, and is based on eight isolated teeth; an upper molar, upper premolars and a couple of lower molars. As the type specimen is the upper M1, I think I'll start with that. It's approximately 2 x 1,5mm is size, (p.703), and nearly rectangular in crown shape. There are two valleys dividing the cusps into three rows; 3:4:4. The cusps of the central row are clearly defined, whilst the others are less distinct. The valley on the internal side of the tooth, (lingual), is deeper and more worn, especially at each end. It's double-rooted, with the front root being narrower. The symmetrical form of this tooth is typical of the genus, as are the pattern of the P4 and "the peculiar P5". (Most authors would be likely to refer to these as the P3 and P4 respectively. Basal multis had five upper premolars. For convenience and continuity, these are sometimes termed P0 - P4. However, in this paper they're called P1 - P5. There are reasons behind this different terminology, though it can be confusing.) The P5 (P4 to others and also referred to as type e elsewhere in this paper) is near rectangular in outline. It has a central, incomplete ridge, several cusps forming part of a steep cutting edge along its external border, (labial), and is double-rooted. Is that any reason to call it 'peculiar'? The case is presented on pages 706-707. Its morphology is unique. The row of cusps at the front is longer than the lingual one, "which is obliquely directed towards the anterior cusp of the buccal one." This peculiarity could have been the result of the tooth in question perhaps being a milk tooth. But it's now known from both H. vianeyae and H. pyrenaica. "Moreover, this element in H. pyrenaica fits perfectly with the M1, suggesting that both belong to a single individual. Therefore, it can be interpreted as the last premolar of Hainina. This peculiar dental morphology is similar to that shown by the last permanent premolar of Kogaionon. The position in front of the M1 would coincide in both genera." This arrangement is not known from other multis, though the authors point to something similar from a couple of "Lower Cretaceous" types; "Kuehneodon" and Lavocatia. (The first named genus is better and subsequently called Kuehneodon. It's mainly known from the Upper Jurassic of Portugal, though some Lower K material has been reported from Spain.) Given that resemblance, (and making it clear that the P5 of Lavocatia has three rows of cusps rather than two): "According to these comparisons, type e can only be morphologically related with P5 of some primitive European multituberculates." Personally, I'm not sure including the word 'only' was advisable, especially as this genus is based on isolated teeth. These authors argue that the four upper premolars of kogaionids are derived from P2 - P5 of more basal ancestors, whilst the four of their contemporaries can be traced back to P1 - P4. As further support for this view, they observe that the third premolar in the series of Kogaionon overlaps its two neighbours, as in basal types such as Bolodon. In other advanced multis, a similar effect is achieved by the fourth premolar. The fossils came from a location called Fontllonga-3, (p.701), and lies only three metres above sandstone channels bearing dinosaur footprints. Despite exhaustive sampling through nearly ten tons of material from Fontllonga-3, absolutely no dino remains were found. Abundant remains of fish, (Coelodus laurenti) and pollen strongly indicate this stratum is Paleocene. This is supported by data on paleomagnetism and stable isotopes. The site appears to be from the very earliest Paleocene, proceeding the K-T extinctions. The non-birdy dinos were wiped out. Small euromultis weren't. "Therefore, the Fontllonga-3 site represents the earliest Tertiary vertebrate fauna from the Old World." The holotype is an isolated left upper molar, (M1). It's FTN3-1, and works in the collection of the Complutense University, Madrid. The species name refers to the Pyrenees Mountains. Unfortunately, Dr Daams died whilst the paper was being edited, (p.701).
Reference:	Peláez-Campomanes et al (2000), The earliest mammal of the European Paleocene: the multituberculate Hainina. J of Paleont. Vol 74(4), p.701-711.

Link: Journal of Paleontology, 74(4), 2000 It's a long address The abstract. The authors conclude that H. belongs to Kogaionidae, advice which I have followed. Kielan-Jaworowska and Hurum (2001) evidently agreed.

Species:	Hainina vianeyae Peláez-Campomanes P, López-Martinez N, Àlvarez-Sierra MA & Damms R, 2000
Place:	Cernay
Country:	France
Age:	Thanetian, Upper Paleocene
Remarks:	The following is based upon my understanding of Peláez-Campomanes et al, 2000. 24 specimens were originally referred to H. godfriauxi. However, they differ significantly in size, and a lower molar, (m1), has a distinctive arrangement of the cusps. They have the more complex formula of 3:4 as opposed to 3:3 (p.702). This was a relatively large species, though its teeth are smaller and more slender than those of H. godfriauxi. Altogether, the authors identified 25 specimens: 4 m1, 1 m2, 1 P2, 2 P3, 4 P4, 2 P5, 7 M1 and 3 M2. This translates as lower and Upper molars and premolars. The typefossil is known as CR 1043, though I'm not sure which collection it's in. The species name honours Mme. Monique Vianey-Liaud, who was the first author to study the material.
Reference:	Peláez-Campomanes et al (2000), The earliest mammal of the European Paleocene: the multituberculate Hainina. J of Paleont. Vol 74(4), p.701-711.

Genus: Kogaionon Rădulescu R & Samson P, 1996

Species:	Kogaionon ungureanui Rădulescu R & Samson P, 1996
Place:	Sinpetru Beds, Hateg Formation
Country:	Romania
Age:	Maastrichtian, Upper Cretaceous
Remarks:	A micromammal based on a well-preserved and near complete skull. The species name is in appreciation of the geologist, Costin Ungureanu, who found the fossil. There were two previous descriptions of multis from this locality, (Grigorescu, 1984 and Grigorescu, Hartenberger, Rădulescu, Samson & Sudre, 1985).
Reference:	Rădulescu & Samson (1996), The first multituberculate skull from the Late Cretaceous (Maastrichtian) of Europe (Hateg Basin, Romania), Anuarul Institutului de Geologie al României, Supplement 1, 69, p.177-178.

Links: SVP News Bulletin, June 1997, Number 170 http://alnus.uel.ac.uk/svp/news_bulletin/170_97_6.html Included in the report headed ‘Romania’. Eggs and hatchlings of the hadrosaurid dinosaur Telmatosaurus transsylvanicus have also been recovered from this location. RP-Online Wissenschaft: Bedeutende Funde von Dino-Eiern in Rumänien http://www.rp-online.de/news/wissenschaft/erde/2002-0604/dinos.html A report, in German, from southwest Romania, mentioning more teeth from K. This dates from June 2002.

Other reports: Hateg Basin, Romania "Recently recovered specimens add to this diversity and provide information about the relationships of these taxa." These were "previously referred to as Hainina sp. A and H. sp. B (Csiki & Grigorescu, 2000). New evidences suggest that they represent new taxa, closely related to Kogaionon," (Csiki & Grigorescu, 2001, Abstracts of the 6th European Workshop on Vertebrate Palaeontology - Florence and Montevarchi (Italy)). Linked above, see Barbatodon. Reference: Csiki & Grigorescu (2000), Teeth of multituberculate mammals from the Late Cretaceous of Romania. Acta Palaeontologica Polonica, 45, p.85-90. A further site in the Hateg Basin has been reported. Link: DML, Hateg Micropaleontological Material, Jaime A Headden, 15.1.2003 http://www.cmnh.org/dinoarch/2003Jan/msg00261.html Amongst the mammal remains is material which might belong to new species of Kogaionon and Hainina. Further Link: Belgium Institute of Natural Sciences http://www.naturalsciences.be/science/actus Paleontologists at work in Transylvania. Click on the dates to the left, and you'll find a series of site photos. A Kogaionon molar's pictured in 'the lab'. (With thanks to Aspidel, DML, 16.1.2003.)

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. back to top Trevor Dykes, November 2001. Last update: 13.2.2008 Ktdykes@arcor.de

With further thanks due to: Paleocene mammals of the world, Class Mammalia http://www.paleocene-mammals.de/pal1.htm Part of Martin Jehle’s excellent work. (PS: thanks for the corrections.) Dr John Alroy, North American Fossil Mammal Systematics Database http://www.nceas.ucsb.edu/~alroy/nafmsd.html The source of the weight guestimates, as well as other useful information. For comparison, a house mouse weighs around 25g and a largish rat about 400g. The Prehistoric Data Files http://www.angellis.net/Web/PDfiles/MarsupS-ad.htm An index of paleo-indexes. A very broad resource. BIOSIS, The Index to Organism Names http://www.biosis.org.uk/triton/indexfm.htm The Society of Vertebrate Paleontology BFV Online, (John Damuth) http://www.bfvol.org/ Georgia Wildlife Web, Mice http://museum.nhm.uga.edu/gawildlife/mammals/rodentia/muridae/sigmodontinae/rhumulis.html This link has everything to do with gratitude and little with Mesozoic Mammalia, other than descent. Mys is a Greek word for mouse. Progress! I wonder what cimex means. Bug! The animated accompaniment to this directory has been provided by Animation Library.

Bibliography: Archibald JD & Averianov AO (2001), Paranyctoides and allies from the Late Cretaceous of North America and Asia, Acta Palaeontologica Polonica, 46 (4), (Proof Version). Averianov AO & Archibald JD (2003), Mammals from the Upper Cretaceous Aitym Formation, Kyzylkum Desert, Uzbekistan. Cretaceous Research 00 (2003), p.1-21. Averianov AO & Archibald JD (2006), New specimens of the multituberculate mammal Uzbekbaatar from the Late Cretaceous of Uzbekistan, Acta Palaeontologica Polonica, 51(2), p.377-380. Clemens WA, Wilson GP & Molnar RE (2003), An enigmatic (Synapsid?) tooth from the Early Cretaceous of New South Wales, Australia. Journal of Vertebrate Paleontology, 23 (1), p.232-237. Eaton JG, Munk H & Hardman MA (----), A New Vertebrate Fossil Locality Within the Wahweap Formation (Upper Cretaceous) of Bryce Canyon National Park and its Bearing on the Presence of the Kaiparowits Formation on the Paunsaugunt Plateau, Department of Geosciences, Weber State University, Ogden, UT 84408-2507 (webpage). Godefroit P, Bolotsky Y & Alifanov V (2003), A remarkable hollow-crested hadrosaur from Russia: an Asian origin for lambeosaurines. C. R. Palevol 2, p.143-151. Grigorescu D & Hahn G (1987), The first multituberculate teeth from the Upper Cretaceous of Europe (Romania), Geologica et Palaeontologica, 21, p.237-243. Hunter JP & Archibald JD (2002), Mammals from the end of the age of the dinosaurs in North Dakota and southeastern Montana, with a reappraisal of geographic differentiation among Lancian mammals, in Hartman JH, Johnson KR & Nichols DJ (eds.), The Hell Creek Formation and the Cretaceous Tertiary boundary in the northern Great Plains: An integrated continental record at the end of the Cretaceous, Boulder, Colorado, Geological Society of America Special Paper 361, p.191-216. Hunter JP & Pearson DA (1996), First record of Lancian (Late Cretaceous) mammals from the Hell Creek Formation of southwestern North Dakota, USA. Cretaceous Research 17, p.633-643. Kermack KA, Lees PM & Mussett F (1965), Aegialodon dawsoni, a new trituberculosectorial tooth from the lower Wealden. Proceedings of the Roy. Soc., London, B, 162, p.535-554. Kielan-Jaworowska Z & Hurum JH (2001), Phylogeny and systematics of multituberculate mammals, Palaeontology, Vol 44 (3), p.389-429. Kielan-Jaworowska Z, Hurum JH, & Badamgarav D (2003), An extended range of the multituberculate Kryptobaatar and distribution of mammals in the Upper Cretaceous of the Gobi Desert. Acta Palaeontologica Polonica 48(2), p.273-278. Kielan-Jaworowska Z, Novacek MJ, Trofimov, BA & Dashzeveg D (2000), Mammals from the Meozoic of Mongolia, p.573-626 in Benton MJ, Shishkin MA, Unwin AM & Kurochkin EN (Eds.), The age of dinosaurs in Russian and Mongolia, Cambridge University Press. Kielan-Jaworowska Z, Ortiz-Jaureguizar E, Vieytes C, Pascual R & Goin FJ (2007), First ?cimolodontan multituberculate mammal from South America, Acta Palaeontologica Polonica, 52(2), p.257-262. Lofgren DL (1995), The Bug Creek Problem and the Cretaceous-Tertiary Transition at McGuire Creek, Montana, University of California Publications Geological Sciences, vol. 140, 185pp. Matthew WD, (1916), A marsupial from the Belly River Cretaceous with critical observations upon the affinities of the Cretaceous mammals, Bulletin of the American Museum of Natural History, 35, p.477-500. McKenna MC & Bell SK, (1997), Classification of Mammals Above the Species Level. Columbia University Press. Montellano M, Weil A & Clemens WA (2000), An exceptional specimen of Cimexomys judithae (Mammalia: Multituberculata) from the Campanian Two Medicine Formation of Montana, and the phylogenetic status of Cimexomys. Journal of Vertebrate Paleontology 20 (2), p.333-340. Norrell MA, Gaffney ES & Dingus L (1995), Discovering Dinosaurs, Little Brown and Company, 204pp. Osborn HF (1893), Fossil mammals of the Upper Cretaceous beds, Bulletin of the American Museum of Natural History, 5, p.311-330. Peláez-Campomanes P, López-Martinez N, Álvarez-Sierra MA & Daams R (2000), The Earliest Mammal of the European Paleocene: the Multituberculate Hainina. Journal of Paleontology 74 (4), p.701-711. Sahni A (1972), The vertebrate fauna of the Judith River Formation, Montana, Bulletin of the American Museum of Natural Histor, 147(6), p.321-412. Simpson GG (1927), Mammalian fauna of the Hell Creek Formation of Montana, American Museum Novitates, 267, p.1-7. Simpson GG (1936a), A specimen of the Upper Cretaceous multituberculate Mesincoessus, American Museum Novitates, 825, p.1-4. Weil A (1999), Multituberculate phylogeny and mammalian biogeography in the Late Cretaceous and earliest Paleocene Western Interior of North America, Ph.D. Dissertation, University of California, Berkeley, p.1-243. Wible JR & Rougier GR (2000), Cranial anatomy of Kryptobaatar dashzevegi Mammalia, Multituberculata), and its bearing on the evolution of mammalian characters. Bulletin of the American Museum of Natural History, 247, p.1-124.