A. ZALAMBDALESTIDAE

Taxon: Zalambdalestidae Gregory & Simpson, 1926 Zalambdalestids were small, insect-eaters of the Upper Cretaceous, whose remains are known from Central Asia. That much is agreed by those in the know. They may be near to the origins of Glires Linnaeus, 1758, a taxon which includes rabbits and rodents. (Glires is the Latin plural of glis, which is the dormouse, -with thanks to Toby White.) They've been placed in the somewhat wider taxon of Gliriformes Wyss & Meng, 1996. According to Archibald, Averianov & Eckdale, 2001, (see Bibliography): "A phylogenetic analysis of well described Late Cretaceous eutherians strongly supports Zalambdalestidae". In other words, it’s a valid taxon of early eutherian mammals. However, more controversial is: "A second analysis incorporating placentals from clades that include rodents (Tribosphenomys), lagomorphs (Mimotona) and archaic ungulates (Protungulatum and Oxyprimus) strongly supports Zalambdalestidae in a clade with Glires, (rabbits, rodents and extinct relatives)…" (Mimotona is a Paleocene mammal with a rabbit dental formula.) Archibald & Co point out that the members of Glires have six character states of the incisor teeth, including Hunter-Schreger bands. (Don’t ask me! The Mature Tooth, Enamel:) Whilst none of these are unique to the taxon, the occurrence of all of them together is. Four of these features were identified as already in place with the earliest known zalambdalestid, Kulbeckia, (though not the Hunter-Schreger bands). Their results indicate that these critters are proto-bunnies and mices. According to others, "it ain’t necessarily so", as indicated in the brief report from Fostowicz-Frelik L and Kielan-Jaworowska Z, 2002 (APP 47/1). One criticism is that the sampled characters were too few. "John J. Flynn, a curator at the Field Museum of Natural History in Chicago, said the American-Russian team conducted an incomplete analysis when they compared about 70 different characteristics between the fossils and other early mammals," (from Merit Times, 24.11.01, Fossils of Chipmunk-Sized Mammal Found, -see Kulbeckia below for the link). Some studies suggest that the family as constituted isn't monophyletic; it doesn't contain one common ancestor and all of its descendants. Archibald & Averianov, 2003 found that Barunlestes is a closer relative of Glires than the other members are but, as they hadn't thoroughly studied two of the genera in the family, they didn't feel in a position to offer a formal rediagnosis, (p.405). Family matters And now a negative view on affinities with Glires based upon my reading of Wible, Novacek & Rougier, 2004. (There's a bibliography at the foot of this directory.) In the discussion section of the paper just mentioned, (starting on p.96), attention turns to the phylogenetic relationships of zalambdalestids; to which taxa might they be most closely related. The family has featured in seven analyses with published data matrices over the last decade. However, as these were generally designed to test affinities of other mammals, points taken into account aren't necessarily useful to this case. Different taxa have differing basal and derived features, and the relevant fossils preserve varying aspects. The exception among these sstudies was the aforementioned 2001 examination by Archibald and friends. (A comprehensive analysis for basal therians is a research goal Wible et al haven't yet attained.) Nevertheless, as those seven analyses have already been undertaken, they do at least provide a starting point for further enquiries. Consequently, the authors offer emendations resulting from their study for the existing data matrices, some of which involve previously unknown characters. They also re-ran the tests within the parameters specified in the original publications. Archibald et al (2001) is addressed beginning on page 105. This involved 70 osteological characters and 25 taxa. Fourteen of the characters were altered as listed. I think I'll content myself with their summing up, (p.109). "In conclusion, although the matrix of Archibald et al. (2001), either the original or with our amendments, includes more relevant basal eutherians than those discussed previously, it still offers a very limited test of zalambdalestid affinities with crown-group Placentalia. Archibald et al. (2001) consider only two options, Glires and Ungulatomorpha. This is too restrictive a test, which is made even more restrictive by the dearth of well-sampled basal members of these two groups." Glires and friends A comparison with lagomorphs, rodents and elephant shrews begins on page 120. One of the similarities which has been cited for Zalambdalestes and lagomorphs (rabbits & Co) is an enlarged lower incisor, (p.121) In both bunnies and also rodents this is the retained deciduous i2. However, Kulbeckia has four lower incisors and that's the basal number for Eutheria. Enlarged is i1. If, as seems likely, that's also the case for other zalambdalestids, then similarities with i2s in members of Glires are hardly explicable by common descent. Various other 'derived' features have been said to unite zalambdalestids and lagomorphs, but none are only found in these two groups. Some actually seem to be basal for eutherians, and primitive traits can continue in disparate taxa. Lagomorphs and rodents are united by "a remarkable number of additional derived features" (p.122) which aren't found in zalambdalestids. Affinities with lagomorphs are rated as "highly unlikely", and with Glires as "unlikely", (p.123). Connections with a higher level clade, Anagalida, have also been suggested. This has been emended to unite the existing orders of Lagomorpha, Rodentia and Macroscelidea (elephant shrews). However, support for this grouping is presently considered weak. I'm going to be daring and risk a personal assessment on the affinities between Zalambdalestidae and living placentals; unresolved due to insufficient data. Keep looking! Whilst it would be very nice to be able to convincingly unite living orders with their long gone Cretaceous relatives, the odds are stacked against success. But that's no reason to stop trying! One jackpot will be rich reward for all the efforts. In the meantime, there's plenty of interest in the systematics of Cretaceous mammals. There are various characteristics shared by zalambdalestids and asioryctitheres, although a number of them are basal factors; eg. epipubic bones and the presence of the last upper incisor in the maxilla or between the maxilla and premaxilla, (p.125). Primitive traits are of limited phylogenetic use. However, there are also common features less frequently found among eutherian, (p.126), but I note the phrase used is "somewhat limited distribution." This sounds like a deliberately neutral choice of words. Comparisons with zhelestids also appear to have provoked less than excitement in this study, (p.129). What is clear The Conclusions offer some concrete positions. Zalambdalestes (and therefore zalambdalestids) are eutherians, and Kulbeckia is within the family. Cranial specialisations are shared with Asoryctitheria, (p.130). "In light of these unusual similarities, we think that a zalambdalestid-asioryctithere clade is the most reasonable working hypothesis for these forms". Should broader analysis support that view, then this would appear to be a further Upper Cretaceous vertebrate taxon endemic to Asia, to set alongside radiations of multituberculates, squamates (lizards, snakes and relatives) and dinosaurs.

Links: Mikko Haaramo's Zalambdalestidae Mikko Haaramo's Zalambdalestidae "Note: Zalambdalestids are one of the most problematic mammalian groups. For a long time they were considered as basal eutherians. They had some interesting specializations that seemed to parallel the evolution of rodents and lagomorphs, but they were considered not related to true rodents." The times they are a-changing? Acta Palaeontologica Polonica, 47(1), pp.177-180 http://www.paleo.pan.pl/acta/acta47/app47-177.pdf Lower incisor in zalambdalestid mammals (Eutheria) and its phylogenetic implications. A brief report from Fostowicz-Frelik L and Kielan-Jaworowska Z (2002). "Here we present evidence on the presence of an open-rooted first lower incisor in Zalambdalestes lechei and Barunlestes butleri; we argue, however, that structure of this incisor does not necessarily indicate relationship of Zalambdalestidae to Glires." "The three or two (Kielan-Jaworowska 1969, 1984) upper incisors in Zalambdalestidae are not notably enlarged, and do not function as a self-sharpening cutting mechanism with the lower ones, contrary to the strongly enlarged upper incisors in Glires. Hence, the series of evolutional events leading to the transformation of, presumably, ancestral state of incisor character observed in Zalambdalestidae into that of Glires seems rather intricate." Reservations about the proto-bunny and mice theory, but… "It cannot be excluded that the Zalambdalestidae represent an incipient stage from which (or from related forms) the advanced incisors of lagomorphs and rodents originated." A note for clarity Nobody has proposed that Bobolestes is a member of the family. Rather, an interpretation from 2005 identifies it as a possible relative; a zalambdalestoid. Whether this will gain general consensus is yet to be seen, but I thought I might as well introduce it to this section. Genera: Alymlestes, Aspanlestes (partly = Kulbeckia), Barunlestes, Bobolestes, Kulbeckia, Zalmbdalestes, Zhangolestes, other reports Time-Line: Upper Cretaceous: Alymlestes, Barunlestes, Bobolestes, Kulbeckia, Zalmbdalestes, Zhangolestes (or Lower Cretaceous) Lower Cretaceous?: Mongolia

Genus: Alymlestes Averianov AO & Nessov LA, 1995 'Alym thief'

Species:	Alymlestes kielanae Averianov AO & Nessov LA, 1995
Place:	Darbasa Formation, Alymtau Range
Country:	Kazakhstan
Age:	lower Campanian, Upper Cretaceous
Remarks:	This genus is represented by a single, left lower molar, which shares characteristics only known from zalalmbdalestids among Upper Cretaceous eutherians. The reduction in size of the paraconid and the taller trigonid and talonid are cited as derived features within this family, (Archibald & Averianov 2003, p.405). The citation could also be given as Averianov & Nessov in Nessov, Sigogneau-Russell & Russell DE, 1994: "A rushed (pre-)publication, of the sort that should be avoided," (McKenna & Bell, 1997).
Reference:	Averianov & Nessov (1995), A new Cretaceous mammal from the Campanian of Kazakhstan. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1995, p.65-74.

Genus: Barunlestes Kielan-Jaworowska Z, 1975 'Barun thief’

Species:	Barunlestes butleri Kielan-Jaworowska Z, 1975
Place:	Khulsan, Nemegt and Hermiin Tsav II
Country:	Mongolia
Age:	Campanian, Upper Cretaceous
Remarks:	These details are mainly taken from Wible et al, 2004. Six specimens of this species reside in Warsaw (p.15), while one can be visited in Moscow. [The Peabody collection at Yale has a couple of casts.] Two of the Polish individuals include some postcranial material. A further colleague (ZPAL MgM-I/135) was also originally referred, but it may be a member of a different genus with affinities to eurymylids, [an Asiatic family of the Paleocene - Eocene according to McKenna & Bell 1997, p.113]. Distinguishing marks Differences to Zalambdalestes include: a shorter but more robust skull; a small and single-rooted upper canine (double-rooted in Z.); and a deeper dentary. A thickened, rough area on the front of the pelvis suggests the presence of epipubic bones, as were subsequently found for Zalambdalestes (p.15). Size and choppers The dental formula is reportedly: uppers I?, C1, P3, M3; lowers i3, c1, p3, m3, (p.32). With a skull length of up to 4cm this is a relatively large Upper Cretaceous eutherian (p.39). Holotype The type fossil, ZPAL MgM-I/77, is a skull with dentaries and much of the skeleton in the care of the Institute of Paleobiology, Warsaw (Kielan-Jaworowska et al, 2000, p.611). The same source cites three upper premolars for this genus as opposed to four in the case of Zalambalestes. A peculiarity is provided by the unusually long rear legs, and these presumably enabled the owner to put much spring into their steps, should they have so pleased (p.614).
Reference:	Kielan-Jaworowska (1975): Preliminary description of two new eutherian mammals from the Late Cretaceous of Mongolia. IN Results of the Polish-Mongolian Palaeontological Expeditions - Part VI. Acta Palaeontologica Polonica. 33, p.5-15.

Links: Palaeontologisk Museum, Oslo: Mesozoic Mammals http://www.toyen.uio.no/palmus/galleri/montre/english/x569.htm A lateral view of the skull. Transitional Vertebrate Fossils FAQ Part 2B, by Kathleen Hunt http://www.talkorigins.org/faqs/faq-transitional/part2b.html The Talk. Origins Archive. An interesting response to the question, "why are there no transitional fossils?" There are. Institute of Paleobiology, Polish Academy of Science, Warsaw http://www.paleo.pan.pl/collect.htm#Mon-mammalia The specimen catalogue.

Genus: Bobolestes Nessov LA, 1985 Remarks: This genus is a possible zalambdalestoidean but not a zalambdalestid. That means it could be a reasonably close relative of the family. Aka: Otlestes Nessov LA, 1985 (partly)

Species:	Bobolestes zenge Nessov LA, 1985
Aka:	Otlestes meiman Nessov LA, 1985 (partly)
Place:	Sheikhdzheili local fauna, Khodzhakul Formation
Country:	Uzbekistan
Age:	lower Cenomanian, Upper Cretaceous
Remarks:	This genus has been interpreted in a number of ways. Its placement within Zalambdalestoidea (but not the family Zalambdalestidae) is in line with Averianov & Archibald, 2005. That study is the source for most of this entry. The authors also carried out some euthanasia on a now defunct genus named Otlestes. Some of that material has been incorporated with Bobolestes, while a couple of further specimens are at least close to a 'zhelestid' named Sheikhdzheilia. An obituary for the departed can be found below. A bit of jawing Bobo is known from both upper and lower jaws and teeth, and it's the smallest mammal so far recovered from the Sheikhdzheili fauna. Nessov, the original author of the species, reported the presence of an 'extra' bone on the lower jaw called the coronoid; an element indeed known from various more basal mammals (p.596). The grounds for this seem to be the presence of a bump on the inside of the coronoid process, which is towards the rear of the dentary. However, a similar feature has been identified for a number of Cretaceous eutherians, and it's probably not the coronoid. Quite what it was is uncertain. One ancient feature definitely preserved is a Meckelian groove. Nessov wasn't entirely convinced about its identity. At the time, (1985), the retention of this groove by early eutherians may have seemed very surprising. Subsequent finds mean the identity isn't now worth disputing. Herr Meckel's most excellent groove was part of basal eutherian anatomy. For example, it's known from both Prokennalestes and Eomaia. Inventory and dimension Known specimens consist of one fragment of upper jaw and at least three dentaries (p.597). Two more lower fragments might also belong. This material was collected at two different sites in the same Formation. A reasonably complete lower jaw measures approximately 1.9cm, so mouse-sized sounds like a reasonable approximation to me. Bobo's teeth In the study, information is spread between pages 594-598, and I think I'll order some details into convenient categories starting with the dental formula. The holotype is part of an upper jaw and preserves two molars (M2-M3) and alveoli for a premolar and rear molar. There were clearly three molars and some number of premolars. The lower count is clearer, but also somewhat uncertain. Grooves at the front suggest up to four incisors although only two are definitely present. Otherwise: one canine, five premolars and three molars. Upper molars These have a mixture of primitive eutherian traits and derived specialisations. Among the former are their narrowness, a paracone that's bigger than the metacone, a wide stylar shelf and a deep ectoflexus. The lingual face of the tooth also lacks a cingula. Those terms will leave some people none the wiser, but so is the terminology of teeth. Lower teeth Following the incisors (possibly up to four), there's a small, single-rooted canine. The solitary root differs from the zalambdalestid condition of two. All premolars are double-rooted. The p2 and p4 are similarly sized and larger than the p1 and p3. The authors describe the fourth as premolariform and the fifth as semimolariform. (Note: p1-p5 is the terminology in this paper, and it differs from terminology used in some studies, in which this p3 may be termed px or some such.) The p5 has a full trigonid and a lengthy talonid which is partly basined. The lower molars have trigonids which are considerably higher than the talonids. For the main cusps, the metaconid is low compared to the protoconid, while the paraconid is relatively big. As with the ultimate premolar, the talonid is lengthy, especially on the m3. It's both longer and narrower than the trigonid. The diagnosis on page 595 contains what appears to be a typo. It states there are five lower molars. If that were correct, it would be astonishing and completely out of line with the description and caption to Figure 2. Furthermore, information on a p5 is provided after that misstatement. Five premolars must've been the intended version. Three molars is fully in line with expectations. This premolar count includes a primitive 'extra' one, (termed p3 in this paper), and no living eutherian has an equivalent dental position. (The modern p3 corresponds to p4 in this case.) Conflict in relationships Bobolestes was originally assigned to the non-eutherian family of Pappotheriidae, which presently consists of a couple of therians from the Lower Cretaceous of Texas. The genus then founded its own therian family (or subfamily); Bobolestidae (or Bobolestinae). Subsequently, it was accepted as a eutherian. In this study, the authors note strong similarities between the final lower premolar (p5 in their terminology) with that possessed by Kulbeckia: "Both have a semimolariform p5 with a distinct metaconid and paraconid, and an enlarged and incipiently basined talonid", (p.596). This contrasts with the condition in other eutherian lineages of the time, and suggests a relationship for Bobo with zalambdalestids. The only characteristic barring membership of that family is the single-rooted canine; double-rooted in zalams. The newly proposed placement is Zalambdalestoidea. Otlestes: an obituary Otlestes was based on two lower jaws and a partial maxilla (p.595). The association was apparently made because they were found in the same place, but the case is far from convincing. The upper jaw suggests a larger critter than the lower. The talonids of lower molars are narrow with small basins, and these couldn't have coped with the protocones of the associated upper teeth. They are, however, suitable for Bobolestes, the holotype of which is also a fragment of maxilla. The fit is good enough for the fossil to be transferred to that genus, and thus make Otlestes a junior synonym (p.596). Meanwhile, the two maxillae fragments are more apt for Sheikhdzheilia, although they're not necessarily co-generic. Holotype(s) The holotype of Bobolestes (CCMGE 2/12176) is a partial upper jaw in the collection of Chernyshev's Central Museum of Geological Exploration, Saint Petersburg. The same institution provides a hostelry for the erstwhile holotype of Otlestes; a lower jaw known affectionately as CCMGE 7/12176. In addition This genus is regarded by some as a basal member of Eutheria, (eg. "undoubted" eutherian say Wible et al 2001, p.2).
Reference:	Nessov (1985), Novyye mlekopitayuschchiye mela Kyzylkumov [New mammals from the Cretaceous of the Kyzylkumy]. Vestn. Leningrad Univ. 17, p.8-18.

Links: Archibald J David http://www.bio.sdsu.edu/faculty/archibald.html/ Dr Archibald's homepage includes a picture of the Kyzyl Kum Desert. This means 'red sands' in Turkic. The sand doesn't look particularly red to me, but maybe this location's anaemic. Cretaceous Research (26) http://www.bio.sdsu.edu/faculty/archibald.html/AverArch05CR26p593.pdf Averianov & Archibald (2005), Mammals from the mid-Cretaceous Khodzahkul Formation, Kyzylkum Desert, Uzbekistan, p.593-608. The complete paper in pdf format. Leandro O Salles, Geodiversitas, 1996 http://www.mnhn.fr/publication/geodiv/g96n2a5.html Rooting ungulates within placental mammals: Late Cretaceous/Paleocene fossil record and upper molar morphological trends. The abstract. Fascinating things specialists do with teeth. I think it means that Bobolestes may or mayn’t be vaguely related to ungulates, which is animals with hooves to peasants like what I is. Averianov & Archibald, 2005 would seem to suggest otherwise.

The Sheikhdzheili local fauna, Uzbekistan, medial Cretaceous The following is based upon my reading of Averianov & Archibald, 2005. The Khodzhakul Formation is in the western part of the Kyzylkum Desert in Uzbekistan, and the deposited rocks date back to the early days of the Upper Cretaceous, (p.593); lower Cenomanian. Six sites have so far produced rare remains of mammals and the collective result is called the Sheikhdzheili local fauna. An unusual aspect for this age is the variety of eutherians, with up to five species so far. These are a possible zalambdalestid, the perhaps related Bobolestes and two or three zhelestids. Opinions have varied over the affinities of Bobolestes, but the authors refer it to Zalambdalestoidea. Presently, no other fauna from this time (around 95 million years ago) has managed more than two eutherians. It's also of interest that later European zhelestids display traits more in keeping with these earlier representatives, than with their chronological contemporaries from Asia. One explanation would be the spread of zhelestids fairly early during the Upper Cretaceous. The Euros could then have been isolated to what was an archipelago of islands, as the Turgai Strait intervened to separate the two continents during the Turonian. Support for this is offered by a fairly late yet basal European hadrosaurid dinosaur named Telmatosaurus. Medial Cretaceous and the state of Middle Dakota The Cretaceous happens to be traditionally split into two stratigraphic subunits, and this leaves some people wondering where the middle's gone. It hasn't gone anywhere, as it's located in the uppermost Lower and lowermost Upper. This is rather like Dakota, which happens to be divided into two states. Middle Dakota contains nothing whatsoever, (as it doesn't exist), but much of interest can doubtlessly be found in the northern parts of South and the southern bits of North Dakota. At least, I presume so. Similarly, the world was home to a pleasing variety of mammals during the middle of the Cretaceous, despite the lack of any so named stratigraphic division. I like the term Medial Cretaceous and it roughly equates to the Aptian - Coniacian (about 121 to 86 million years ago). Medial Cretaceous mammals While the planet offered a home to many, the fossil record has room for improvement (which is happening). Professional and amateur researchers are doing sterling work in Victoria, Australia, as they laboriously force open a window into the Aptian-Albian deep south. North America is contributing later insights; the Cedar Mountain, Dakota and Straight Cliffs Formations. Asia is also helping out. China has contributed a single bit of spine from Tsondolein-Khuduk, which is the meagre basis of 'Khuduklestes'. Japan has produced a more informative partial jaw of Sorlestes. As neither site has been extensively sampled, further finds could well occur. More extensive faunas have been found in what's termed Middle Asia, which approximately translates as Kyrgyzstan, Tajikistan, Turkmenistan and Uzbekistan (p.594). The winner so far is provided by Turonian (and possibly Coniacian) strata in the central Kyzylkum Desert. The Sheikhdzheili local fauna is attempting to catch up. Sheik Dzheili and his pets Sheik Dzheili was presumably a local celebrity at some time in the past, but the historical annals have failed to record much about his deeds. At least geography (and now paleontology) have preserved his name for posterity, and that's more honour than the vast majority of people are accorded. I hope his was a gracious, kindly gentlemen and have heard nothing to the contrary. I suppose Google could be consulted, although variations on spelling and name changes are likely... Not a virtual sausage. Anyway, there was this bloke called Sheik Dzheili and he neglected to set up a homepage on Yahoo. (Even I'm famous enough for one of those.) His pets, (we have two Guinea girls ourselves, and both give occasional squeaks of encouragement either for this project or more cucumber), have been obtained from half-a-dozen localities, all of which are in the Khodzhakul Formation. SK H-20 is Khodzhakul itself, SK H-5 and 5a are at Khodzhakulsai, SSHD-8 and 8a are the Sheikhdzheili localities and the sixth (SCH-1) is at Chelpyk. The first mentioned two are parts of an escarpment on the north of Sultanvais Mountains near to a river called Amu Darya. Chelpyk is a hill adorned with castle ruins around 30km further north. SK H-20 is a bit earlier than the others, as its lower down in the Formation. Convenient marine invertebrates in rocks both above and below indicate the sandwiched strata are from the lower Cenomanian. Paleo-history The first relevant relics were found by Nessov in 1978 at Khodzhakul, and he also struck fossils at Sheikhdzheili the following year. That site was surface collected on four further expeditions until 1985. The most intensive sampling occurred in 1982, when dry screening of three tons of matrix resulted in the relatively healthy return of twelve mammalian fossils. The only mammals found in 1994 and '98 were frustrated human researchers. Screenwashing campaigns in 1999 and 2003 processed thirteen tons of matrix, and resulted in thirteen further specimens. That brings the total to over thirty. The fauna isn't numerous but it is diverse. These fossils are spread between the collections of three institutions. By the seaside and in the river The rock at Khodzhakul built up in the sea near the coast (p.605), and that makes the rarity of terrestrial mammals explicable enough. The locality at Sheikhdzheili boasts of fluvial sediments and is richer in mammals. With some specialised exceptions, we eucynodonts are generally better at falling into rivers rather than oceans. However, the similarities of the remains suggest both locations can sensibly be referred to as a single faunal unit. Even if there's a bit of an age difference suggested by differing stratigraphic heights of localities, it hasn't had a significant impact on the fossils preserved. Any difference is of little significance. Eutherian power Assuming the affinities of Bobolestes have now been satisfactorily settled, this fauna contains at least five eutherians (bearing in mind the large, anonymous 'zhelestid'). No other contemporary community presently boasts more than a pair. Eutherians appear to have dominated the landscape of Middle Asia, (should we sensibly ignore minor actors such as dinosaurs and other uncouthly large reptiles). This is a stark contrast to the Cenomanian-Turonian faunas of North America, where multituberculates and metatherians ruled the land, (bearing in mind we just agreed to ignore things like dinos. After all, there's nothing clever or elegant in being outsized). Further Mesozoic site summaries can be found at Localities. Meet the Mammals of the Bissekty Formation The mammalian Fauna so far described include (Stand: 2005): ?Placentalia: ?Zalambdalestoidea: Bobolestes zenge "Zalambdalestidae" - ?Zalambdalestid indeterminate. "Zhelestidae" - Sheikhdzheilia rezvyii; cf. Sheikhdzheilia rezvyii; Eozhelestes mangit; 'zhelestid' indeterminate unnamed large species A. Material previously assigned to the erstwhile genus of Otlestes is spread among Bobo and cf. SheikhD. A further genus, Oxlestes, is regarded as a nomen nudum. The relavent specimen may (or may not) be from the large, unnamed zhelestid.

Genus: Kulbeckia Nessov LA, 1993 Aka: Aspanlestes (partly) Remarks: Previously assigned to its own family, Kulbeckiidae Nessov, 1993. The Biosis Index cites 1994 for the generic name. Archibald & Averianov, 2005 (p.11) indicates a few similar fossils have been recovered from the Aitym local fauna, which overlies the Bissekty in Uzbekistan. They also point to a smaller, presently undescribed species for this genus from the Bissekty local fauna.

Reassigned species: K. kansaica Nessov, 1993 and K. kansaica Nessov, 1993 see K. rara

Species:	Kulbeckia kulbecke Nessov LA, 1993
Aka:	Aspanlestes aptap Nessov, 1985 (partly); Kulbeckia kansaica Nessov LA, 1993; K. rara Nessov LA, 1993
Place:	Bissekty Formation, Dzharakuduk & Yalovach Formation
Country:	Uzbekistan & Tajikistan
Age:	middle-upper Truronian - lower Santonian, Upper Cretaceous
Remarks:	This is, as yet, "the oldest known zalambdalestid," (Archibald et al, 2001). The majority of its kin are from the Gobi and are 10-15 million years more recent, (ca. 75Ma). K. kansaica is a synonym, and refers to the fossils from Tajikistan. The genus was named in 1993, because Archibald et al, 2001 say so. They also kindly supply the citation. It was based on a few teeth. The zalambdalestid affinities weren't recognized until 1997. Digging at Dzharakuduk (Uzbekistan), (1997-2000), produced "over 40 additional specimens... including teeth, dentaries, petrosals, a partial skull, and postcrania," (Archibald et al, 2001). Some of this material is discussed in Archibald & Averianov, 2003, which is featured below. A lower molar and a petrosal, (the bone which houses the inner ear), have been reported from the overlying, (and thus somewhat younger), Aitym Formation. There are affinities to this species. There's also an upper canine, though it's only about two-thirds the size of those found in the Bissekty fauna. (Averianov & Archibald 2003, p.7-8.) They go on to say, (p.14), that this: "is the most abundant single species in the fauna...", (ie. the Bissekty fauna). K. kansaica is apparently from the Santonian of Kansai, Tajikistan, whilst K. rara is perhaps Turonian, but certainly an Uzbek. The following is based upon my reading of Archibald & Averianov, 2003. This description involved the additional material. This paper and other sstudies assign Kulbeckia to "Zalambdalestidae", (p.404): "Kulbeckia shares with other zalambdalestids: a narrow, somewhat elongated snout; procumbent, enlarged, and open-rooted medial lower incisor with enamel restricted to the more ventrolabial surface; and anteroposteriorly compressed and centrally pinched molar trigonids." As well as being older than the other family members, Kulbeckia also appears to be more basal. It's smaller; probably had four, rather than three lower incisors; the lower canine was either fully double-rooted or bifurcated; diastema are either small or absent between the front teeth; the cristid obliqua are more lingual; the third molars aren't as reduced; and the angular process of the dentary in adults is both more dorsally positioned and further towards the back. Premolar terminology (p.404) This paper contains a significant peculiarity of terminology. Existing placentals have four or less premolars, (though I'm uncertain of the situation in dolphins and allies). Some basal representatives had five, and its the third tooth in the sequence which got lost. This recent realisation produces a problem of continuity in literature. The obvious way of differentiating between four premolars is with the numbers 1-4, and that's what happened. However, this means '4' is the analogue of the originally fifth tooth. These authors favour p1-5. If I don't get muddled along the way, when they write p4 (this paper) it correlates with p3 (traditional). Should they have occasion to write p3, this would be either pc or px (semi-traditional). I'll follow their terminology but add a trad. alternative (in brackets). Location and age (p.404) An escarpment at Dzharakuduk is a couple of hundred metres high. It contains various formations. At the bottom is the Uchkuduk, (brackish and marine). The next layers in this cake are: Dzheirantuj (marine), Kenyktjube (marginal marine), Bissekty, (fluvial) and Autym (marginal marine). The river deposits of the Bissekty have so far produced about a hundred vertebrate taxa. The remains in the formations above and below allow the assessment of the age. Partial skull (p.406) A specimen known affectionately as URBAC 99-53 is part of the left skull. This extends from near the front of the rostrum to the middle of the orbit. Due to distortion the front of the nasal has been overridden by the maxilla, but the relevant proportions could be restored. Part of the premaxilla is missing. The absent part could conceivably have held between none to two incisors. Three are present. The first is fully rooted within the premaxilla, the next seems to be in a borderline position, whilst the third, (which is much smaller), could be completely in the maxilla. The canine is either double-rooted or has a bifurcated root. Upper jaw and dentition The previously mentioned URBAC 99-53 provides some information on the teeth, but none are well represented, (p.407). Judging by the freshness of cracks, some probably broke off during the fieldwork. The evidence that is available shows: roots for three incisors though more may have been present on the missing portion at the front (p.406); a double-rooted or bifurcated canine; double-rooted premolars P1-P2 and triple-rooted P4-P5 (P3-P4 trad.); worn molars M1-M2 and a triple-rooted M3, (p.407). [Regarding the number of upper incisors Wible et al 2004, (p.33), contains: "We see no basis for any speculation beyond the evidence of the three upper incisor alveoli, which resembles some Zalambdalestes." They also have an estimate for skull length of something like 4cm, (p.39).] All teeth from the foremost preserved incisor to the P1 are separated by small diastema. A larger diastema is located between P2 and P4 (P3 trad.). Nothing indicates the presence of a P3 (Pc or Px semi-trad.), though earlier loss and subsequent remodelling of the bone can't be ruled out. This individual was probably the Upper Cretaceous equivalent of a pensioner. The preserved upper incisors are vertical, with the rearmost being very small. The premolars increase in size along the series, (p.408), with P1 being about the size of the first two preserved incisors. The protocone on P5 (P4 trad.) is larger than on P4 (P3 trad.), which is a feature reminiscent of M1-M2. "There are two size morphs within Kulbeckia kulbecke, which may represent sexual dimophism or even the presence of two species. Within the upper dentition, this size variation is best seen in M1s. The two smallest M1s are between 74 and 82 percent of the length of the largest specimens", (p.409-410). Lower jaw and dentition The material available included ten mandibles with teeth, five edentulous ones and fifteen isolated teeth. Size, proportions and structure allow the referral of all to the one species. All the adult mandibles have a depth of between 2.7 - 3.1 mm, (p.411). This is fairly uniform along the whole bone, though there is a gradual shallowing from back to front. Unsurprisingly, specimens which include unerupted or erupting teeth are less deep. (They're presumably juvenile or subadult.) In contrast to the upper dentition, diastema are generally not present, excepting for sometimes between the canine and first premolar. There are at least four incisors. The first is a greatly enlarged, horizontally positioned tooth, most of which is covered with thickened enamel, (except dorsoligually); on page 413: "What is preserved is not inconsistent with an ever-growing or gliriform incisor or at least the precursor of such an incisor as found in rodents, lagomorphs, vombatid marsupials, and the primate Daubentonia, among others (Koenigswald, 1985). The evidence, however, is not conclusive regarding whether the incisor was truly hypsilodont." (Please note. Nobody is suggesting that the similarities found in the incisors of those disparate therians are all down to joint inheritance.) The three other incisors are small, (p.411). One specimen is equipped with an erupting canine, (p.413). This is about 60% the height of an isolated, upper counterpart, (p.415), though it might not have achieved its full potential height. "Although complete roots of upper canines are not known, from what's preserved, it appears that the roots of the lower canine are at a greater angle to the crown", (p.414-415). This seems to imply the canine is double-rooted. There are no fully preserved p1s, p2s or p4s (p3 trad.), though there are some bits and pieces. One dentary does include a well-preserved p5 (p4 trad.) and the lower molars. This p5 (p4 trad.) has a well-developed trigonid, in which the protoconid dominates. The metaconid is about half as fat and two-thirds as high. The paraconid is something like half as thick again. It's positioned slightly lingually at the front. The talonid is as wide as, but a bit shorter than the trigonid. The first molar (m1) is slightly longer than the p5, but shorter than the m2 and m3, which have similar lengths. Its trigonid and talonid are generally the same width. Usually, but not always, the trigonid is wider than the talonid in the other molars. Interlude Should anyone have read this far, you'll have gathered that the paper is full of details. I think I'll include a block from the Conclusions, which addresses the wider picture of mammalian evolution. "Our understanding of the evolutionary history of Mesozic mammals is fraught with problems, not the least of which is the quality of the fossil material. The spectacular material from the Late Cretaceous of Mongolia demonstrates, however, that this is not the only and possibly not the greatest obstacle. The lack of evolutionary diversification within clades of Mesozoic mammals may be the greatest impediment. This is almost certainly not an artefact of the fossil record. We believe this lack of diversity will not increase markedly as the record improves, which it is bound to do. There are some exceptions. Arguably, the best of these known among Late Cretaceous eutherians are the zalambdalestids, which have remarkable convergences if not actual homologies in skeleton, skull, and teeth with extinct and extant placentals -notably rodents and rabbits", (p.418). Holotype The holotype (CCMGE 52/12455) is an isolated, left lower molar (M1 not M2). It works at Chernyshev's Central Museum of Geological Exploration, Saint Petersburg. One of its colleagues is the holotype of K. kansaica, (CCMGE 9/12455).
Reference:	Nessov LA (1993), New Mesozoic mammals of middle Asia and Kazakhstan and comments about evolution of theriofaunas of Cretaceous coastal plains of Asia. Trudy Zool Inst, 249, p.103-133.

Link: Merit Times, Fossils of chipmunk-sized mammal found http://english.hsilai.org/merittimes/detail.asp?index=4586&page=K A nicely written and accessible report. Since May, 2002, Eutheria is known to reach back still further, (Eomaia, ca. 125Ma).

Genus: Zalambdalestes Gregory WK & Simpson GG, 1926 'v-(shaped) thief' Aka: Zalambdolestes

Reassigned species: Z. grangeri Simpson, 1928 = Z. lechei; Z. mynbulakensis Nessov LA, 1985 see Sorlestes budan

Species:	Zalambdalestes lechei Gregory WK & Simpson GG, 1926
Aka:	Z. grangeri Simpson, 1928
Place:	Bayan Zag, Tögrög & Ukhaa Tolgod
Country:	Mongolia
Age:	Campanian, Upper Cretaceous
Remarks:	The following is mainly based upon my reading of Wible, Nowacek & Rougier, 2004. In summary This detailed study of the skull and teeth of Zalambdalestes was prompted by excellently preserved new specimens from the Dkjadokjta Formation, (p.3). These provided new information and resolved some uncertainties and errors made concerning less cooperative fossils. The genus was large in terms of Upper Cretaceous eutherians with a skull length of nearly 5cm. The snout was a long, thin tube. The lower front teeth included a big incisor with restricted enamel. These specialisations are also found in a more moderate form in the somewhat younger Barunlestes and the older Kulbeckia. The study supports placement of Zalambdalestidae within Eutheria, but not as part of crown-group Placentalia. A number of basal features, (epipubic bones and the last upper incisor being found in the maxilla are a couple of examples), suggest a relatively basal position, and speak against strong affinities with the placentals of Glires, (bunnies, mice and so on). A number of apomorphies are shared with members of Asioryctitheria, and provide some support for affinities between the two Cretaceous taxa The known specimens of Zalambdalestes display what's termed "a remarkable degree of individual variation" in some aspects of the dentition. While all are presently referred to the same species, closer comparisons in the future might reveal the presence of more than one species. As the relevant fossils are held in widespread collections, there are logistical challenges involved with such an undertaking. These remarks will now become somewhat more detailed. Holotype numbers and the like are included at the foot of this entry, should your visit be intended as a short one. Historical background Although therian mammal remains are known from the Lower Cretaceous, the earliest reasonably rich sites are said here to date from the Upper Cretaceous of Mongolia, (but I'm putting in a word for the Yixian Formation, China anyway)! These Mongolian beds were first explored by American expeditions in the 1920s, and specimens were evaluated in sstudies by Granger & Simpson (1926) and Simpson (1928). The affinities of zalambdalestids have been subject to varying opinions ever since, (p.4), although the family's eutherian credentials have remained stable enough. Zalambdalestidae was originally referred to Insectivora. Back in the 1920s Zalambdalestes was easily the best known mammal of the Mesozoic. Teams from various countries have turned up further and better preserved fossils since the 1960s, as well as discovering a couple of close relatives. The skull is now nearly fully described, and postcranial elements include a partial pelvis and an epipubic bone. Insectivora? Originally, the upper molars were described as wide, and the cusp pattern as v-shaped, (zalambdodont and thus the name). As this seemed to be reminiscent of "zalambdodont" insectivores such as tenrecs and golden moles, (p.7), affinities were proposed. (It should be added that Insectivora was used rather loosely. For example, with regards to Zalambdalestes and Upper Cretaceous friends: "The placentals are generally classed as Insectivora. They are, however, such not in the same sense as the later, specialized insectivores, but in the sense that they lack known clear-cut ordinal characters within the Theria and that "Insectivora" has been used as a convenient catch-all for such virtually nonordinal groups. Their degree of diversity and any special relationships to known later groups are yet to be established, Simpson, 1959, p.405.) However, differences were also apparent; eg. the very long lower incisor, a postcanine diastema and the tubular snout. The molars of Z. grangeri improved knowledge of morphology, and the perceived similarities with "zalambdodonts" were explained as probably primitive traits for insectivorans, and thus relatively uninstructive with regards to genuine relationships. The molar structure was more akin to leptictid hedgehog relatives, and the genus was transferred to Erinaceoidea. However, it didn't seem overly happy there. Anyone for rabbits? By the 1960s resemblances to Paleocene fossils from the Gashato Formation of Mongolia were noted. Zalambdalestes began to grown friendly with lagomorphs, (rabbits and relatives), and then rodents. However, other views saw no such affinities, and it was also still dallying with the leptictids until the 1970s. Frankly, the genus was feeling giddy and none the wiser. Polish-Mongolian excavations began recovering fossils in the 1960s, (p.14). These included a dozen specimens of Z., two of which provided further postcranial remains. The undertaking was also rewarded with really sexy dinosaurs, which perhaps helps explain why Soviet-Mongolian expeditions took over the initiative. Possibly demonstrating that a Catholic God favours Warsaw over the orthodoxy of Moscow, these digs were less blessed, but another Z. lechei fossil was recovered. Lifestyle Of course, these fresh specimens provided fuller information, and insights into skeletal specialisations. Whatever its affinities, the hind legs of Zalambdalestes were relatively long, the femur and fibula were extensively fused and it had very long metatarsal bones in the toes. These and related details pointed to climbing, or a hopping mode of movement comparable with existing small mammals found in semi-deserts, (eg. macroscelidids aka elephant shrews). Geological evidence records just that type of environment, (p.15). Dentition For various reasons, (including poor preservation and individual variation), the dental formula has been subject to differing findings. The review conducted by Wible et al, 2004, seems to have clarified matters, (p.16-17). As stated on page 32 the dental formula is: uppers: I2-3, C1, P3-4, M3; lowers: i3, c1, p 3-4, m3. (The letters denote incisors, canine, premolars and molars respectively, while Upper Case refers to Uppers and lower case to lowers.) [A brief anecdotal interlude to illustrate the possibilities of individual variation People generally have two molars per side, but a third sometimes erupts later. These wisdom teeth bring us into line with basal eutherians. A maximum of three is a distinctive characteristic, although a dolphin may smilingly think otherwise. I've just heard from a man who's got four molars in one half of his jaw. Although his dentist is very proud of this achievement, he still offered to remove the one that simply shouldn't exist. However, he couldn't provide any convincing reason as to why this should be done in this instance. I suggested that one of his grandparents must be a marsupial, seeing as they are allowed four rather than three. More likely is that tooth numbers can vary even in the same mouth for various reasons: delayed eruption, late retention (not applicable to molars), premature loss or utter freakishness.] Uppers There are two incisors per side in the premaxilla. The I1 is large and downwards pointing, (p.18). A short diastema comes between it and the much small I2. Whether these have restricted enamel isn't clear. Rather oddly, some individuals apparently had a third incisor housed at least mainly in the maxilla. In one instance, this could be a strange and reduced canine, (deciduous or mature), (p.20). However, some contemporary mammals then also had incisors partly or wholly within the maxilla, (eg. asioryctitheres). A further specimen presently known as cf. Z. sp. even manages to have an additional pre-canine tooth on only one side of the jaw, (p.21). When present, the I3 is separated from a double-rooted tooth by a diastema. Despite being an unusual distance back from the meeting point of maxilla and premaxilla, this is most probably the canine. Then come generally three premolars, (P2-P4), although a P1 has been exceptionally reported, (p.22). This was small and double-rooted. The P2's a bit larger, (p.23), and the P3-P4 are more complex. Their structure is broadly similar, but the P4 is smaller with a better developed protocone as in the molars. Perhaps the most significant difference between the first two molars is that M1's wider, (p.25). M3 is the smallest of the three, (p.28). Lowers There are three lower incisors. The basal eutherian condition seems to be four, (p.28), but it's not absolutely clear which one's absent in this case. It's very possible that the second and third correspond to i3-i4 in Kulbeckia (Archibald & Averianov, 2003). However, the authors decided to retain the numbering as i1-i3 in line with previous papers on Zalambdalestes. Although based on a relatively poorly preserved specimen, the description by Gregory and Simpson, 1926 presents the general condition of the i1 succinctly: "much enlarged, procumbent and occludes between the enlarged uppers incisors" (I1 here). It's long and forward pointing. In one instance, the length from alveolus to tip is 0.7cm, and that's impressive for a 5cm skull. The root extends back to at least below the front of the p4. Enamel is restricted to the front but seems to be present from the tip to the root, (at least for cf. Z. sp PSS-MAE145 -p.30). This tooth is followed by three small and decreasingly procumbent gnashers; i2-i3 and an unimpressive canine. When the mouth was closed, those last two incisors would've been accommodated in the above diastema between I2 and C, and the lower canine would have fitted just in front of its colleague, (p.31). The original description reported three premolars, ('p2-p4'). However, a gap between the '2' and '3' suggests a tooth was missing, (p2). Other specimens have four double-rooted premolars. Another short diastema separates the canine and p1, which is slightly procumbent. While the second is smaller than the first, the p3 is larger than either. This has a main cusp with a minute accessory one in front of it, and an unbasined heel behind, (with entoconid). p4 is the tallest tooth and submolariform. As regards the molars, m1 and m2 are similar in shape, with the former being both larger and higher. m3 is the smallest. Lower jaw "The lower jaw in Zalambdalestes is elongate and anteriorly tapering, with two mental foramina of unequal size (not clearly preserved in any MAE specimen), the larger one below the middle of p3 and the other one below the anterior root of p1, according to Kielan-Jaworowska and Trofimov (1981)", (p.34). It's long, gets progressively narrower towards the nose, and there are a couple of small holes in the bone beneath two of the premolars. Skull (and comparative sizes) The largest specimens of this species have a skull length of nearly 5cm. This is marginally smaller than known from some dinosaurs, but it's large for Upper Cretaceous Mongolian eutherians. For comparison, Barunlestes manages up to 4cm and Kulbeckia is probably similar. Kennalestes, Asioryctes and Ukhaatherium range between 2.5-3cm. Daulestes from Uzbekistan was more like 2cm. Of course, the skulls would've had bodies attached to them in life. (I tend to multiply by four or five according to mood, and the former works well enough for the measured length of Ukhaatherium, which amounts to about 12cm). Compared to asioryctitheres, zalambdalestids had inflated braincases. All else being equal, smaller mammals generally require proportionately more bodyspace for their brains. (Some shrews devote a higher percentage of available volume to this purpose than I do.) I don't wish to sound mean minded but, should you be stuck on a crossword clue, then asking a zalambdalestid for help would perhaps be the better option. The paper carries on discussing minute features of the skull in extensive detail, and soon left me floundering. Included were accounts of openings for nerves, arteries and veins. The architecture of the inner ear was addressed. I was left admiring the extensive array of photos and sketches provided. Brain I fared a bit better with page 86. This includes information on endocranial casts; naturally produced sculptures of the brain. In a study in 1984, Kielan-Jaworowska provided some tentative estimates for encephalization quotients for Cretaceous eutherian taxa. Higher figures indicate more brain cells as a ratio of overall body size: Kennalestes gobiensis 0.36, Asioryctes nemegtensis 0.56, Zalambdalestes lechei 0.70. In all cases, the olfactory bulbs of the brain are very large, which indicates a well developed sense of smell. Other details attest to good hearing. These may indicate preferences for nocturnal activity. Various measurements of a new endocast are in line with previous results. Ear An update concerning ears features on page 88. Previously, the cochlea was described as having one whorl, (360°). However, a 'better' estimate is here given as 450°. This is the amount of spiralling in the cochlear canal and some jottings upon the theme are linked to here. The Prokennalestes entry is followed by More Earnotes Holotypes The holotype of Z. leichi (AMNH 21708) is a skull in the collection of the American Museum of Natural History, New York. Page 14 of Gregory and Simpson, 1926 reveals the specific name honours Professor Wilhelm Leche for his work on various insectivores including zalambdodonts. The material described in three papers from the 1920s included the skull of a younger individual, and a partial lower jaw preserving the p3-p4 and m1-m3. Z. grangeri (AMNH 21709) was based upon the front of a skull with the near complete right maxilla and postcanine teeth, a partial pelvis and a femur. As differences were not pronounced, the two species were synonymized by Szalay & McKenna in 1971, (Wible et al 2004, p.7). Z.? mynbulakensis Nessov LA, 1985 and Z. nymbulakensis (sic?) have apparently been used for Sorlestes budan, though I don’t know whether these names were employed formally. Additional notes A possible second species has been collected from Bayan Mandahu, China, (eg. Kielan- Jaworowska et al 2003, p.277). As well as New York, specimens are also housed in Warsaw and Moscow. Just kidding The following is based on my reading of Simpson, 1928, which isn't the paper establishing Z. grangeri. Simpson described an additional skull of Z. lechei on page two. It was almost complete with dentary, but wasn't well preserved. There are characteristics which differ somewhat from the holotype. Both specimens were broadly similar but this one was relatively short in the cranial and facial areas; had proportionately bigger orbits; a shorter snout and diastema in the tooth row; a more gracile jaw; and possibly a shorter internal incisor. If you'd care to study myself and my son, some differences like this may be apparent. Notice, for example, how his nine year old eyes still demand a bigger share of the face (but not as much as when he was freshly delivered). As for his dentition, if I now gently coax his mouth open with the incentive of a bit of chocolate, you should see something quite... Ouch! His teeth are sharper than I thought. Even the low cusped, bunodont molars have made me realise that putting my fingers there wasn't a good idea. Anyway, despite what he may think when a teenager, we don't belong to different species. As with this additional Zalambdalestes specimen, the contrasts have much to do with biological age. Such factors can be difficult to spot, and it gets even trickier with non-mammalian cynodonts due to indeterminate growth strategies. Simpson concluded somebody had given him a pup.
References:	Gregory & Simpson (1926), Cretaceous mammal skulls from Mongolia. American Museum Novitates 225, p.1-20.
	Simpson (1928), Further notes on Mongolian Cretaceous mammals, American Museum Novitates, 330, p.1-14.

Links: Palaeontologisk Museum, Oslo: Mesozoic Mammals http://www.toyen.uio.no/palmus/galleri/montre/english/x568.htm A further cast specimen in the Norwegian collection. AMNH, Zalambdalestes http://www.amnh.org/exhibitions/expeditions/gobi/dinosaurs/images/13.html A common needle-nosed furry fellow from the Upper K of Mongolia. A sketch of the skull with some commentary. The Cretaceous Cabaret, Zalambdalestes http://www.dennislivingston.com/cretaceous/lyrics_zalambdalestes.htm Words and music by Dennis Livingston. The Prehistoric Data Files, Zalambdalestes http://www.angellis.net/Web/DFG-mam/Zalambdalestes.htm A sketch by VRW. Rat Association of Southwest Missouri http://www.geocities.com/missouri_rats/history.html A somewhat iffy history of rats: "The first real relatives of the rats are found in a group called the multituberculates." (Oh no they’re not.) "Rat-like Megazostrodon," (?). "A quick note: This section does include the term evolution, which is a true scientific event in nature. It doesn't have anything to do with humans or apes, so please don't be offended if you are against the theory of evolution, which is a different subject." (Surely it has some vague connection with humans and apes?) Maybe partly because of the pussyfooting around by the responsible rat fancier, it’s quite an interesting read. Basale Säugertiergruppen, Evolution der Säugertiere http://www.wirbeltiere.de/Fossilesaeuger.htm "Basal mammal groups, Evolution of Mammals." A German site with some linked, (unaccredited), illustrations. (I think most have been 'borrowed' from Cox B, Dixon D, Gardiner B & Savage RJG (1989): Dinosaurier und andere Tiere der Vorzeit, Mosaik Verlag (Sonderausgabe für Gondrom Verlag, 1994), or the English original. This is a fine book.)

Genus: Zhangolestes Zan S, Wood CB, Guillermo WR, Jin L, Chen J & Schaff CR, 2006 'Zhang's hunter' Remarks: Thanks are due to Yuong-Nam Lee for kindly sending a copy of the description. The generic name honours Mr Zhang Pulin. Rather than simply discovering one of the two known specimens (not the holotype), he went much further by finding several previously unknown, fossil-rich localities.

Species:	Zhangolestes jilinensis Zan S, Wood CB, Guillermo WR, Jin L, Chen J & Schaff CR, 2006
Place:	Quantou Formation, Jilin Province
Country:	China
Age:	probably Upper Cretaceous (lower). However, the Formation might turn out to be a bit older.
Remarks:	The following is based upon my reading of Zan et Al, 2006. Fossils from Liaoning Province, northwest China have stimulated much excitement over the past couple of decades. The quality of preservation of finds from the the Yixian Formation can be astonishing. Now Jilin Province has decided to attempt to catch up with the neighbours. Remains from a new site in the vicinity of Gongzhuling aren't comparable with their Yixian counterparts when it comes to completeness or detail but, nevertheless, they're most welcome additions (p.153), and the promise of more is alluring. These specimens are from exposures of the Quantou Formation, the age of which is presently somewhat obscure. The deposits are more recent than the Lower Cretaceous Yixian, with Cenomanian being the most likely possibility. If that turns out to be correct, then they date from the dawn of the Upper Cretaceous; part of the informal "Middle" or Medial Cretaceous. Other suggestions include a somewhat older, Aptian dating. In any case, they're certainly more recent than the Yixian. With thanks to Mr Zhang Pulin The first mammal from this new fauna to take a bow is Zhangolestes ('Zhang's hunter'), and its name honours Mr Zhang. This isn't the same man who found the Yixian Zhangheotherium. The relevant Zhang is Zhang Pulin, a former geologist who worked for the Jilin Geological Survey until his retirement. He then found time to explore outcrops of fossil-yielding rock in the west of the Province, and that led him to several new locations. Zhang Pulin also came up with the first of the two known specimens of Zhangolestes. Unfortunately, his death in September 2000 put an end to his endeavours. Mr Zhang's bequest Prior to being forced into hanging up his geological hammer for good, his findings and fossils passed into the care of Jilin University Geological Museum. This institution instigated further fieldwork during 2000 and 2002, and those efforts were rewarded with an enlarged collection of vertebrate remains; bones, teeth and even several eggs. As sites with Medial Cretaceous terrestrial fossils hadn't previously come to light in this area before, the chances for an array of new taxa are high. Mr Zhang's hunter Two of the mammalian fossils appear to represent a relative of Kulbeckia kulbecke from Middle Asia. These are fragments of lower jaw with teeth. Although the teeth preserved are from different dental positions in both cases they probably belong to the same species, and were found separately in the same stratum (p.154). If the conclusion of close affinities with Kulbeckia is correct, then this genus is the earliest known member of the eutherian family of Zalambdalestidae (p.155). It's also the first instance of such a mammal being found as far east. Them bones In terms of size and morphology the two specimens seem to be from the same species, although they were found some eighty metres from each other. Combined information from both gives a picture of a reasonably complete mandible. For example, all tooth positions seem to be represented (p.156). However, as there's little actual overlap between the two fossils, that conclusion is somewhat conditional. A single more complete specimen could perhaps necessitate a revision. Happily, these two fossils aren't the only mammalian material to have been collected. Zhangolestes will receive company as the described fauna expands. The following notes include the assumption the remains represent the same species. Dental formula and jaw length The lower jaw had (per side): 3 incisors, 1 canine, 5 premolars and 3 molars. The number of premolars is the basal eutherian condition. The available remains suggest a jaw length of significantly more than two centimetres. (I'd guess at something like 2.5.) This critter was at least mouse-big. Incisors The first incisor is a very large tooth which points nigh-on horizontally forwards. It was even larger in life, as the end's broken off. The root extended back in the jaw beyond the second premolar, which is the last preserved tooth on the holotype. As the i1 root is also broken, how far back it went is unclear. Nevertheless, information from the second specimen shows it was at least as far as the p4. Although missing its tip, enough is preserved to show the top surface there (medial) was somewhat spoon-like. That area of relative flatness gives the incisor a D-shaped cross-section, with enamel being restricted to the curved surfaces below. Wear may have been partly responsible for a loss of enamel. The other two incisors are also procumbent but much smaller. The i2 is a bit larger than the i3 (p.158). Canine This tooth slopes diagonally forward and seems to be single-rooted, in contrast to Kulbeckia (double-rooted). It slopes at an angle of about 30° as opposed to 15° for the i3. While larger than the rear incisors, the canine's much smaller than the first. It's fully enamelled apart from a wear facet, and the small size suggests it might not be fully erupted. However, the amount of wear on other teeth makes that unlikely. Premolars The first premolar is the baby of the team (p.159). There's some damage but, when in its fully glory, it must've been rather boring; a rounded crown with a hint of a heel at the rear. A bit of a gap separates it from the canine, although it's insufficient to merit the term diastema. The tooth was double-rooted, and the height of the protruding roots exceeds that of the crown. The p2 is larger; about the same length and height as the canine. Its main cusp is supported by the front root and a small heel is at the rear of the crown. This is the largest of the front three premolars, although information on the third is scant due to severe damage (p.160). The other two premolars are represented on the second specimen along with the molars. The p4 provides little more than the pair of roots. Still, they suggest the rear of the tooth was wider than the front (p.162). The p5 is larger and nearly qualifies for the word 'molariform'. The crown's well preserved and not as long as the first two molars. It does, however, have a completely rimmed talonid basin at the rear although this lacks distinct cusps. It appears that the corresponding upper premolar was equipped with a corresponding protocone. The front of the crown boasts a trigonid of three distinct main cusps. Tallest of these is the protoconid. The paraconid is lower and somewhat more labially situated than the metaconid. That latter cusp is about half the height of the protoconid. The crown bears no cingulum on its lingual side. Molars The first molar is also well preserved but it's lost the metaconid from the trigonid and the entoconid cusp from the talonid basin. The paraconid has a smaller comparable diameter than what's left of the metaconid, and this cusp is distinct and pointed. It's somewhat labial of the metaconid, as is the case for the p5. The protoconid is considerably taller. The talonid is longer than the trigonid and about as wide (p.163), but the latter has double the height. Both this tooth and the p5 are a bit higher on the labial side than on the lingual one. The other two molars are much less talkative. Only the roots of the m2 remain, and the crown of m3 has fallen beyond disrepair into the realms of ruination. A few details can be made out to some extent, and it's clear this molar was the smallest of the trio. Eutherian credentials Cretaceous eutherians have four (or sometimes five) premolars and three molars per side, and the final lower premolar is typically at least close to molariform. There are usually three incisors (or more in earlier models). These characters apply for Zhangolestes. Cretaceous metatherians, in contrast, favour three premolars and four molars. The final lower premolar is never molariform, and the corresponding upper doesn't possess a protocone. Zhangolestes is a eutherian. Closer affinities There are a couple of touches with these specimens that hadn't previously been seen in Cretaceous mammals, and that's why a new genus was required. The lower p5 in more complex than ones found in other Medial Cretaceous mammalian mouths. It's more or less fully molarized, and that also distinguishes it from asioryctitherians (p.164). Its dental approach is more in keeping with zalambdalestids and 'zhelestids'. While some aspects of the m1 are more reminiscent of that second mentioned group (eg. the strong separation of the entoconid and hypoconulid cusps on the talonid), the overall morphology of p5 and m1 seem closer to zalams. The p5 is like the equivalent gnashers of Kulbeckia and Asian specimens of Paranyctoides. Small distinctions, however, debar membership of those genera or Bobolestes. When the other teeth are taken into account, the list of differences naturally widens. While the enlarged first incisor is also known from Kulbeckia and later zalams, the number and arrangement of the other incisors contrasts. For example, there's no indication of a fourth incisor for 'Zhang's hunter'. Despite being earlier, this animal appears more derived in that regard. Beneficial despoliation The locality from which these fossils came was a pit; a by-product of road construction (p.166). Fossiling time was limited in 20002 as a building company was anxious to develop the site. While that sort of economic reality can be frustrating, the fossil bed could've remained entirely unknown but for the activities of the construction industry. The 'ravages of progress' can be advantageous for paleontologists, even should a fossil locality end up as a building site. Most specimens were obtained from the same thin bed; a band of 10 to 30 centimetre thick mudstone laid down in freshwater conditions. Analysing the stratigraphic relationships of locations in the area is difficult, and that contributes much to the lack of clarity regarding the age. Aptian, Albian and Cenomanian have all been suggested in studies based on larger invertebrates, plants and microfossils (p.166). Healthy plant growth and homes are good for people, but they happen to obscure the rock layers beneath (p.167). What's clear is that the Gongzhuling locality belongs to the Quantou Formation, and the remains date from the Medial Cretaceous; probably Cenomanian. It's got to be new As fossil-yielding locations of that sort of age happen to be in short supply in this part of the world, the chances are high that any vertebrate finds will be from previously unknown taxa (p.168). This is near to being the white space of unexplored territory on a paleontological map. Information on Chinese mammals is available for earlier and later, and that suggests a zalambdalestid might've been predictable. Finding one provides support for the view that zalams resulted from a basal eutherian radiation, and that they're not all too intimately associated with particular placental orders (especially rodents and bunnies). A ditched premolar Zalams eventually dispensed with the services of one of the originally five lower premolars. In some, the number was further reduced down to three (p.169). As mentioned, 'Zhang's hunter' retained five. Later news from Asia has been interpreted as showing the p3 was the first to be ditched. However, comparisons between Zhangolestes and Kulbeckia (which had four premolars) led these authors to conclude it was the original p1 that had gone. If correct, then that could indicate a reduction in numbers happened in several lines (not too improbable), and by the loss of differing dental positions (also not improbable). The significance of this conclusion could presumably be undermined, should completer specimens show the two fragments aren't from the same taxon. Holotype Ya1.23.i is the front of a lower left jaw in the collection of Jilin University Geological Museum. The specific name refers to Jilin Province in northeastern China.
Reference:	Zan et Al (2006), A new 'Middle' Cretaceous zalambdalestid, from a new locality in Jilin Province, Northeastern China, Journal of the Paleontological Society of Korea, 22(1), p.152-172.

Link: Zan et Al, 2006 http://cactus.dixie.edu/jharris/Zanetal_Zhangolestes.pdf The full description is presently available on-line in pdf format.

Other reports: Sheikhdzheili local fauna, Uzbekistan According to Averianov & Archibald, 2005, this fauna was probably home to a zalambdalestid. Among fossils recovered is a well preserved, isolated petrosal (p.598), which is the bony housing for the inner ear. At 3.5 x 2.5mm it has an area of 8.8mm². Although the diminutive dimensions might suggest an impression to the contrary, many details are clear (with the aid of a microscope). It's very similar to petrosals accredited to an early zalam called Kulbeckia, but further remains haven't been found in this fauna, (p.599). Should it indeed be from such an animal, then its closest better known local relative is Bobolestes. However, it's actually too large for a mammal of that size. Dorngobi Aimag, Mongolia Giallombardo A & Nowacek M report having arrested a new and early possible zalambdalestid from the Khugenetslavkant sandstone facies. It's said to either belong within the family or a sister-group, but that's a provisional conclusion. There are two fragmentary skulls, and one has the lower jaws and various bits of body. If my reading performance is up to the mark, there are also a couple more bits of maxillae. The snout is relatively long and narrow, and the nasal bones broaden from front to back. Teeth The dental formula is at least mostly clear: (uppers): 4? incisors, 1 canine, 5 premolars and 1 molar (that's surprising, but the number is as given); (lowers): 3, 1, 5 and 1 respectively. The P4 and 5 and the fifth lowest premolars are all termed molariform, front lower incisors are very large (hypertrophied) and procumbent (forward pointing), lower molars have a 'compressed trigonid' and, using a rather exotic word, talonids are 'fully basinated'. (I've never written that word before, and I'd happily never do so again.) The report is in the 2006 Abstracts of the Society of Vertebrate Paleontology, p.67A. Surprisingly or otherwise, 'basinat...' that adjective features but once in all 122 pages.

A. Zalambdalestidae B. Lipotyphla? C. Cimolestidae and Cretaceous Taeniodonta

B. LIPOTYPHLA?

Taxon: ?Lipotyphla Haeckel, 1866 These genera have been referred to the grandorder of Lipotyphla. If correct, this means they're proper, Cretaceous members of Insectivora, and thus early relatives of shrews, hedgehogs and moles. However, such affinities are not universally accepted. Update 'These genera' has become something of an exaggeration. Batodon used to hang out in this section, but it's scurried off to become a cimolestid further down the page.

Link: Insectivores http://www.angelfire.com/mo2/animals1/mammal/insectivora.html A site maintained by Matthew. Should these be insectivores, these are some possible descendants. If not, there's a decidedly cute photo of a shrew. Genera: Ortalestes (= Paranyctoides), Paranyctoides, other reports Time-Line: Upper Cretaceous: Paranyctoides, Europe, India

Genus: Paranyctoides Fox RC, 1979 Remarks: The following is based upon my reading of Archibald & Averianov, 2001, (proof version). This genus is diverse in the Upper Cretaceous of North America. Three species have been named from there, (p.1), and possibly four more are present. However, the earliest species for this (or a closely related) taxon has been found in Uzbekistan, (P. aralensis). At some stage its relatives appear to have migrated. As various other animal groups are known to have done the same, (eg. tyrannosaurids), this isn't particularly surprising. Molars The trigonids on the lowers are unambitious when it comes to height; ie. they're low. The paraconid is well developed. On m1-m2 the talonid is at least as wide as the trigonid, and the hypoconulid tends to be nearer the entoconid than the hypoconid. (Those are further small cusps.) Not much is known of the upper molars, as only two have been reported. The crown is relatively narrow and there's a wide, cusped stylar shelf on the labial side. Premolars A peculiarity of the genus is the presence of up to five premolars per side, which is basal for eutherians. Reasonably enough, Archibald & Averianov number these 1-5. However, there's a snag, (p.2). The third tooth in the sequence was dispensed with during subsequent evolution. As there was no reason to suspect its existence prior to its discovery, other sstudies have referred to the p3 of these authors as the px. Similarly, p4 here equates to p3 trad and p5 is p4 trad. This also applies for the upper premolars. Unnamed species Two species, (Wahweap A and B), have been reported from the Wahweap Formation. A further pair, (Kaiparowits A and B), are present in the slightly later Kaiparowits Formation. These are all based on isolated teeth; lowers in the first cases, both uppers and lowers from the Kaiparowits. Both localities are in Utah, and the identification was made by Cifelli in 1990. Affinities In 1979 Fox interpreted Paranyctoides as being close to Nyctitheriidae within Lipotyphla. This existing order of placentals is sometimes known as Insectivora. While there is a suite of similarities, others find lipotyphlan affinities are debatable, (p.6). However, the genus could be near the ancestry of that order or archaic ungulates. There is consensus on a placement within Eutheria. Archibald and Averianov suggest that Paranyctoides and Gallolestes represent a clade within 'Zhelestidae', (p.14). While therians of a Turonian age are known from North America, none of them resemble these genera. As for 'zhelestids', pre-Campanian representatives are presently restricted to Asia, (p.17). These lineages reached America later. On possible curiosity is the lack of such fossils in the Campanian of Mongolia, which is geographically between Uzbekistan and North America. Intuition suggests it ought therefore to be on the migration route. However, there's an environmental factor which could've been significant. Both the Uzbek and American sites were deposited on low lying coastal plains, (p.17). The Gobi locations were further inland. The environments may have favoured differing lineages. Aka: Ortalestes

Reassigned species: cf. Paranyctoides Nessov, 1993 see Avitotherium utahensis

Species:	Paranyctoides sternbergi Fox RC, 1979
Place:	Oldman Formation, Alberta &Fossil Forest fauna, New Mexico
Country:	Canada & USA
Age:	Campanian, Upper Cretaceous
Remarks:	This species is based upon a number of dentary fragments and isolated molars. The Oldham is now known as the Dinosaur Park Formation. This is another holotype in the University of Alberta collection.
Reference:	Fox (1979), Mammals from the Upper Cretaceous Oldman Formation, Alberta. III. Eutheria, Canadian Journal of Earth Sciences, 16, p.114-125.

Species:	Paranyctoides maleficus Fox RC, 1984
Place:	Alberta & Montana
Country:	Canada & USA
Age:	Campanian, Upper Cretaceous
Remarks:	This species is based upon parts of the lower and upper dentitions. The lower molars are very similar to those known from P. sterngbergi, except that they're more robust. The uppers, which are unknown for P. sternbergi, combine basal and derived eutherian characteristics. Basal are: wide stylar shelves with cusps, (some of which may themselves be derived); a deep ectoflexus; the protocone is higher than the metacone, which is taller than the paracone; and the crowns are not greatly expanded transversely. These characteristics are also known from Prokennalestes, and older, non-eutherian therians. Derived are: the separation of the paracone and metacone bases; some expansion along the length of the protocone; narrow pre- and postcingula, with a small hypocone sometimes present, (Archibald & Averianov 2001, p.6).
Reference:	Fox (1984), Paranyctoides maleficus (new species), an early eutherian mammal from the Cretaceous of Alberta. Carnegie Museum of Natural History Special Publication 9, p.9-20.

Species:	Paranyctoides megakeros Lillegraven JA & McKenna MC, 1986
Aka:	Paranyctoides megaceros
Place:	'Mesaverde' Formation, Wyoming
Country:	USA
Age:	Campanian, Upper Cretaceous
Remarks:	This species is poorly known, and was established for a couple of lower molars. It's relatively large.
Reference:	Lillegraven & McKenna (1986), Fossil mammals from the 'Mesaverde' Formation (Late Cretaceous, Judithian) of the Bighorn and Wind River basins, Wyoming, with definitions of Late Cretaceous North American land-mammal 'ages', American Museum Novitates, 2840, p.1-68.

Species:	Paranyctoides aralensis Nessov LA, 1993
Aka:	Ortalestes tostak
Place:	Bissekty Formation, Dzharakuduk
Country:	Uzbekistan
Age:	middle-upper Turonian, Upper Cretaceous
Remarks:	The holotype is a right dentary with the double roots of the premolars, (p1-p5 in the terminology of Averianov & Archibald, 2001). Partial molars are also present, (m1-m3). Further material has since been added to the collection. Lower molars These are close in morphology to the material from North America. The cusps of the trigonid are rounded and similar in height, and the trigonid is relatively low in comparison with the talonid, with the latter being at least as wide or wider. This is in contrast to Daulestes-like species in the fauna, which have narrower talonids and less rounded cusps, (Averianov & Archibald 2001, p.7). The molars of this species do have several distinctions from their equivalents in the American species, (and Gallolestes pachymandibularis.) In all cases there's a decrease in height along the line, (m1-m3), and (except in this species) the talonid is about half the height of the trigonid. With P. aralensis the relative height ranges from half to three-quarters, (m2). Along with other details and despite being earlier, this might be a relatively more derived species, (p.14). Premolars At least four mandibles preserve either the alveoli or roots of five premolars and, as with the larger 'zhelestids' from Uzbekistan area, the third premolar is clearly reduced. This is the one that was subsequently lost. Holotype The holotype, CCMGE 67/12455, is in the collection of Chernyshev's Central Museum of Geological Exploration, Saint Petersburg. The preserved length is a bit over a centimetre, judging by the photo on page 3). Several teeth and jaw fragments have been found in the overlying Aitym Formation, (upper Turonian - Coniacian). They might also belong to this species. (Averianov & Archibald 2003, p.7-8).
Reference:	Nessov (1993), New Mesozoic mammals of Middle Asia and Kazakhstan and comments about evolution of theriofaunas of Cretaceous coastal plains of Asia. Trudy Zoologischeskogo Instituta, Ross. Akad. Nauk. 249, p.105-133. (In Russian).

Link: Acta Palaeontologica 46(4), 2001 (proof version) http://www.bio.sdsu.edu/faculty/archibald.html/ArchibaldAverianov01APP46.pdf J. David Archibald and Alexander O. Averianov, Paranyctoides and allies from the Late Cretaceous of North America and Asia, (proof version). In published form, the page numbers were 533-551.

Other reports: Europe The Upper Cretaceous fauna list included in Averianov & Archibald 2003, (p.17), includes mention of cf. Leptacodon in the Campanian of Europe. Leptacodon is an insectivore more usually associated with the Paleocene - Eocene. The Deccan Traps, India The following is based upon my reading of Khosla et al, 2004. The paper concerns a further Upper Cretaceous (Maastrichtian) microvertebrate site in Central India. It's located near the village of Kisalpuri in Madhya Pradash, (p.380). The exposure is an intertrappean site in the Deccan Traps, and was provided by a huge, prolonged bout of volcanic activity, which may be partly to blame for the global mass extinction(s). Preserved are remains of various fish, frogs, snakes, lizards, turtles, crocs, dino egg-shell and one eutherian tooth. The fossils were obtained by screenwashing about 500 kilos of sediment, and the rewards are seen as relatively rich. The mammal tooth is incomplete, but it's a last, left lower molar. Looking at the pictures, (p.381), even my unschooled eye can see it's tribosphenic. Its closer affinities are addressed on page 382: "The new tooth from Central India lacked a continuous, mesial cingulid wrapping around and extending onto the lingual side of the crown." This absence characterises Boreosphenida. Finer details justify a referral to ?Otlestidae, because it closely resembles Otlestes, (although being more derived). This will be described more fully in a subsequent study, but it shows the presence of a further northern invader of Upper Cretaceous Gondwana.

A. Zalambdalestidae B. Lipotyphla? C. Cimolestidae and Cretaceous Taeniodonta

C. CIMOLESTIDAE AND CRETACEOUS TAENIODONTA

Taxa: Cimolestidae Marsh, 1889; Taeniodonta Cope, 1876 This section contains Cimolestidae, as listed by McKenna & Bell, 1997. Opinions vary on the affinities (and validity!) of this group. Mc & B, (1997) places the family within the order Cimolesta McKenna, 1975, suborder Didelphodonta McKenna, 1975. Elsewhere, they're referred to as palaeoryctids. In Mc & B, Palaeoryctidae Winge, 1917 is presented as a closely related family. Batodon (Section B) might also belong here. Update Wible et al, 2007 includes some cimolestids, and they established a new genus from Asia, Maelestes. They don't discuss the wider content of the family, but merely state Maelestes and the North American Cretaceous Cimolestes and Batodon are members of it. Other genera may or may belong. I've followed their views for Maelestes. And there's more! Results of their analysis show the cimolodontids aren't exactly ancestral to Carnivora or, indeed, any recent placental orders. This is because they don't qualify as placentals, but are rather relatively basal eutherians. As a matter of fact, they looked most carefully and couldn't locate any Cretaceous placentals. I'm not sure what their views might be concerning our taeniodont friend in this section, as they don't mention it. Be that as it may, I'm presently opting to stick with what appears to be an outdated and rather meaningless Cimolestidae. Wible et al, 2007 wasn't directly concerned with the full membership of the family. The taeniodont is Schowalteria. As yet, this is the only Cretaceous representative. In the scheme of McKenna & Bell, 1997, Taeniodonta is a suborder within Cimolodonta. A review Among the 2006 Abstracts of the Society of Vertebrate Paleotology can be found Strauss J on p.128A. He cites cimolestids as sometimes being suspected of giving rise to carnivores and further modern placentals. This is based both on their appropriate presence in rocks of the right sort of age, and the fact that their teeth occlude in a carnassial-like manner. The interrelationships are far from clear, and some may well remain so unless (or until) more informative specimens turn up. Procerberus has its dedicated fans. This genus is presently suspected of actually being a palaeoryctid, and a possible ancestor of taeniodontans. Taken too literally, this would be surprising in the light of Schowalteria, should that indeed be an Upper Cretaceous taeniodontan. The presence of Procerberus before the Paleocene doesn't seem assured. An analysis involving a wider selection of morphological characters is underway, and it's hoped these might resolve some of the issues involved. Presumably, a publication will result at some stage. Genera: Aboletylestes, Acmeodon, Avunculus, Batodon, Centetodon (at least partly = Didelphodus), Cimolestes, Deltatherium (partly = Didelphodus), Didelphodus, Didelphyodus (= Didelphodus), Emperodon ( = Gelastops), Gelastops, Ilerdoryctes, Maelestes Niphredil (= Paleotomus), Naranius, Nyssodon (= Cimolestes), "Paelaeosinopa" (partly = Paleotomus), Paleotomus, Phenacops (= Didelphodus), Procerberus, Protentomodon, Puercolestes (= ?Cimolestes), Schowalteria, Tinerhodon, Tsaganius, other reports Time-Line: Eocene: Didelphodus, Ilerdoryctes, Naranius, Tsaganius Paleocene: Aboletylestes, Acmeodon, Avunculus, Cimolestes, Gelastops, Paleotomus, Procerberus, Protentomodon, Tinerhodon Upper Cretaceous: Batodon, Cimolestes, Maelestes, Procerberus?, Schowalteria, Mongolia Seeing as these genera are predominantly not Mesozoic, I think I'll stick with the 1997 McKenna & Bell version, (p.212).

Link: Mikko Haaramo's Didelphodonta Mikko lists alternative interpretations for many of the following genera. For those interested in systematics, he's also produced a cladogramme based on John Alroy 2002.

Talking of palaeoryctids, what were they? This article is based upon my reading of Fox, 2004. I take it you're asking because at least many of the animals in this section of the directory have sometimes been referred to that family, as mentioned above. It can also be used more narrowly, and that's an opinion I'm presently following. Palaeoryctids are assigned to a mammalian family called Palaeoryctidae, and that translates as 'ancient diggers'. Remains aren't extensive, so whether they enjoyed digging is a matter that can be left to the imagination. As they're restricted to the Paleocene and Eocene, the 'ancient' part's justified. The Eocene ended about 34 million years ago. Used in a narrow sense, it's not a very numerous family but, if all interpretations are borne out, it would have been a widespread one. Presently, four genera have been named from North America: Palaeoryctes, Aaptoryctes, Eoryctes and Lainoryctes. Reports indicate that further material is awaiting description from Africa, Asia and Europe, (p.612). (A North African species of Palaeoryctes has already been described, but there are strong doubts concerning the validity of the classification.) Typical fossils are isolated teeth. As these are small teeth, the animals must have been mini-mammals. The subtleties of architecture reveal these were placentals, but closer affinities are less clear. Fox places them within a superorder called Insectivora, but he leaves the matter of which order open. They may be close to lipotyphlans, (hedgehogs, shrews, moles, etc) than to other eutherians, or otherwise. Other suggestions have linked these 'ancient diggers' with creodonts, carnivorans and Malagasy tenrecs. Palaeoryctid molars suggest insect wouldn't have welcomed their visitations, and they could have conceivably done damage to very small vertebrates as well. These animals were killers. Vicious talk Many people have got no idea when it comes to which animals are world champions for ferocity. Their lists are likely to include such relatively harmless creatures as big cats, white sharks and crocodiles. Should they be allowed paleontological options, then they might plump for Tyrannosaurus rex, and salivate with excitement and blood lust. This is complete tosh. Lions enjoy nothing better than lots of sleep, (excepting perhaps for occasional sex). White sharks can spend ages swimming around for no particular reason, and crocs will go months without eating if needs be. Even when awake, should a single lion completely fail to make a kill on 85% of its attempts, it'll still have enough to eat. To get real ferocity, you need animals which have to slaughter the equivalent of their own bodyweight on a daily basis. When making Natural Born Killers, Oliver Stone should've cast shrews in the most savage roles. Some critics feel this may well have improved the acting. These 'ancient diggers' were also small placentals, and fuel consumption would have been high. Although knowledge is based on little more than isolated teeth and scraps, they must have featured strongly in the world rankings list for ferocity. What makes a palaeoryctid? There was a mummy palaeoryctid and a daddy one, and they loved each other very much, but that sort of behaviour is a poor candidate for fossilisation. Presently, they're diagnosed according to similarities of the teeth, and especially the molars. These are relatively short from front to back, but wide. The main cusps are high and sharp. The paracone and metacone are intimately connected until they diverge towards the top. The lowers have short trigonids, generally narrow talonids, prominent shearing crests and so on. Whether this family consists of genuinely related genera is open to doubt. The similarities could be at least in part down to convergence. Some skull material is known for Palaeoryctes and Eoryctes, and it doesn't seem to support the family being a natural taxon, (as presently established). More clarity would require further information. The cranial fossils also have no obvious characteristics supporting close relationships with creodonts, carnivorans or lipotyphlans. Lainoryctes 'Of stone digger' is lovingly described by Fox, 2004. It's relatively large as palaeoryctids go, and represented by part of an upper jaw. As my purpose here is an accessible article on the family, I'm not going to attempt any details. Should it nevertheless be of interest, the holotype of Lainoryctes youzwyshyni is UALVP 43003, and it may be consulted at the University of Alberta. The specific name honours GP Youzwyshyn, who has contributed much to the knowledge of Paleocene mammals in that region. The fossil came from the Paskapoo Formation and its age is Upper Paleocene. It's Tiffanian (Ti4) as defined by the primate index taxa Plesiadapis churchilli / P. simonsi. Restricted Palaeoryctidae Following detailed descriptions and comparisons, Fox briefly mentions some taxonomic history, (p.615). The version of Palaeoryctidae used, (three established genera plus the new one), is based on the scheme offered by McKenna & Bell, 1997. Other authors have used the family is a wider sense, and included genera such as Cimolestes and Pararyctes. If so viewed, then the first mentioned genus would extend the range back into the Upper Cretaceous. Elastic teeth Teeth are the hardest parts of a mammalian body, but that doesn't mean natural selection can't adorn them with new refinements. The cusps can sharpen or flatten. Crests can strengthen or weaken. Over the generations cusps can shift position, fade away or new ones can form. Simple changes can happen to optimise them for which ever foodstuff their owners favour. If, as with the 'ancient diggers', you have an inordinate fondness for beetles, then sharp and pointy teeth are ideal for decimating the armour.

Genus: Aboletylestes Russell DE, 1964 Aka: Aboletyestes

Species:	Aboletylestes hypselus Russell DE, 1964
Place:	Walbeck, Sachsen-Anhalt
Country:	Germany
Age:	Upper Paleocene
Remarks:	An unnamed species is known from Monte de Berru, France. Several specimens of some kind work at the AMNH in New York.
Reference:

Species:	Aboletylestes robustus Gheerbrant E, 1992
Place:	Adrar Mgorn
Country:	Morocco
Age:	Upper Paleocene
Remarks:	There's also material attributed to cf. A. hypselus from the Paleocene of Morocco, (Rana & Wilson 2003, p.342). Page 343 informs me that the holotype of A. robustus is an upper molar, (M1?), named THR 184, which resides in the collection of the Université des Sciences et Techniques du Languedoc, Montpellier.
Reference:	Gheerbrant (1992), Les mammifères Paléocènes du Bassin d'Ouaruzazate (Maroc): I. Introduction générale et Palaeoryctidae. Palaeontographica Abteilung A 2224, p.67-132.

Genus: Acmeodon Matthew & Granger, 1921

Reassigned species: A. hyoni Rigby, 1980 see A. secans

Species:	Acmeodon secans Matthew & Granger, 1921
Aka:	Acmeodon hyoni Rigby, 1980
Place:	San Juan Basin, New Mexico & Wyoming
Country:	USA
Age:	Torrejonian, Paleocene
Remarks:	John Alroy lists this as a member of Insectivora with a suggested weight of about 180g. A second, unnamed species was also reported by M & G in 1921. A fair number of specimens, collected by Rigby from Swain Quarry in 1980, are in the AMNH, New York. Further material has been recovered in Montana.
Reference:

Genus: Avunculus Van Valen L, 1966

Species:	Avunculus didelphodonti Van Valen L, 1966
Place:	Gidley Quarry, Montana
Country:	USA
Age:	Torrejonian, Paleocene
Remarks:	In the scheme of John Alroy, this is another member of Insectivora. Be that as it may, the suggested bodyweight is about 2.5 mice, (60g).
Reference:

Genus: Batodon Marsh OC, 1892 Remarks: This genus has been referred to the family of Geolabididae McKenna, 1960 within the above named grandorder. Benton, 1999 (p.1045) states: "The only modern mammalian order with a Cretaceous origin is Insectivora (or Lipotyphia), with late Maastrichtian records from Wyoming (the geolabid Batodon)." However, some people have expressed the view that these fossils may have been reworked; they perhaps fell into Cretaceous strata from a higher level. Wood CB & Clemens WA (2001), (A new specimen and a functional re-association of the molar dentition of Batodon tenuis (Placentalia, incertae sedis), Latest Cretaceous (Lancian), North America. Bulletin, Museum of Comparative Zoology 156(1), p.99-118), seems to confirm the Cretaceous placement, whilst leaving the question of relationships open. Other authors have placed this genus within, or near to Cimolestidae. Wible et al, 2007 out the genus as a cimolestid, so I've followed their lead.

Species:	Batodon tenuis Marsh OC, 1892
Place:	Lancian Formation, Wyoming, Montana & Alberta, Saskatchewan
Country:	USA & Canada
Age:	Maastrichtian, Upper Cretaceous
Remarks:	A few hints are provided by Simpson, 1951 (p.9). As with Telacodon, this genus was established for front fragments of jaw which, inconveniently, housed premolars but not molars. Batodon had the longer canine. Marsh termed them placentals. This was disputed for years, but he seems to have been broadly vindicated about this pair. Lofgren, 1995... A few specimens came from McGuire Creek localities in Montana, and some were Paleocene sites. However, those fossils probably got reworked from Cretaceous deposits (p.120). The number only amounted to four teeth from three different sites, and they provided no new information of note. Postcanine lengths These figures only directly apply to the McGuire Creek contingent. Uppers: M2 (1 specimen) ca. 1.00mm. Lowers: p4 (1 sp.) 1.08mm; m2 (2 sp.) 1.20-1.22mm. Holotype The type fossil, USNM 2139, is a partial dentary working at the United States National Museum, Washington. It's rumoured to be Wyoming's most popular and least fossilized representative in the political capital. (That was satire, by the way.) This fine candidate receiveds the backing of all voters in its home constituency of the Lance Formation. Additional notes A small scurrier of about 5-6g, (Gordon & Cifelli 2003, p.93).
Reference:	Marsh (1892), Discover of Cretaceous Mammalia, Part III, American Journal of Science, 43, p.249-262.

Genus: Cimolestes Marsh OC, 1889 ‘bug thief’ Aka: Nyssodon Simpson, 1927; ?Puercolestes Reynolds, 1936 Remarks: This is seen as some kind of smallish proto-carnivore and is sometimes enthroned as the mother of all moggies, (cats). More ambitiously still, Cimolestes has also made claims to be the forerunner of Carnivora. The genus is apparently known from both sides of the K-T border. Whilst being predominantly North American, some material seems to have been identified in the Paleocene rocks of Morocco. As for the identity of Puercolestes, "synonymy questionable" is the opinion of McKenna & Bell, 1997, (page 212).

Reassigned species: C. curtus Marsh, 1889 see Didelphodon curtus; C. cutleri Smith Woodward, 1916 see Eodelphis cutleri

Links: Wildlife Portfolio http://thunderlizard.gn.apc.org/wildlife.html Illustrations by Steve White, including Cimolestes. The Velvet Claw, A Natural History of the Carnivores http://www.bobpickett.org/velvet_claw.htm A substantial read. An abridged article from 1992 by David MacDonald. Dogs, cats, bears, weasels, creodonts and Cimolestes. Whilst assisting a BBC film crew in December 2003, (fame at last), I learned that The Velvet Claw was a TV programme made by The Natural History Unit in Bristol. I naturally stressed the importance of producing a new series, and suggested myself as the centre of attraction. I haven't heard back yet.

Species:	Cimolestes cerberoides Lillegraven, 1969
Place:	Scollard Formation, Alberta & Hell Creek
Country:	Canada & USA
Age:	Maastrichtian, Upper Cretaceous
Remarks:	The holotype attends the University of Alberta. It's bodyweight was something like 79-124g, (Gordon & Cifelli 2003, p.93).
Reference:

Link: Type Specimens in UALVP Collections http://www.biology.ualberta.ca/wilson.hp/UALVP/UALVPtypelist.html The type specimens of Alberta.

Species:	Cimolestes cuspulus Gheerbrant E, 1992
Place:	Adrar Mgorn 1
Country:	Morocco
Age:	Paleocene
Remarks:	Intriguing. Information welcome.
Reference:	Gheerbrant E (1992), Les mammifères paléocènes du Bassin d'Ouarzazate (Maroc). I. Introduction générale et Palaeoryctidae. Palaeontogr. Abt. A: Paläozool., Stratigr. 224, p.67-132.

Link: 36th Symposium of Vertebrate Palaeontology and Comparative Anatomy http://www.svpca.org/previousyears/1988/posters.html Posters and Exhibits, Boulogne 1988. Listed is one apparently entitled "Palacogeographical implications of the oldest African placental (Adrar Mgorn Palacocene of Morocco)". The spelling originalities are not of my creation!

Species:	Cimolestes inciscus Marsh OC, 1889
Aka:	Nyssodon incisus, ?Puercolestes incisus
Place:	Lancian, Wyoming, Montana & Frenchman Formation, Saskatchewan
Country:	USA & Canada
Age:	Maastrichtian, Upper Cretaceous - Puercan, Paleocene
Remarks:	Matthew, 1916 reports that the proportions of the holotype (a lower molar) vary from Cimolestes curtus (p.491). As the latter 'species' was based on material from a stagodontid marsupial (or near-marsupial) called Didelphodon vorax, this isn't surprising. Matthew further observed: "Why Marsh associated such widely different types of teeth in the same genus, or what he could possibly have suppoed to be the distinctive characters of the two genera cited I do not know." Additional notes The type specimen came from the Lance Formation of Wyoming. According to John Alroy, C. incisus is a sister taxon of C. simpsoni, but not of the other NAm C. species, or is that my misinterpretation? The body mass is estimated at about 235g (Alroy) or about 104-176g (Gordon & Cifelli 2003, p.93). The holotype and some colleagues are in the Peabody collection. As yet, I haven’t come across any confirmation for the usage of Puercolestes incisus. Nyssodon has been interpreted as a marsupial, (eg. Simpson 1935, p.174).
Reference:	Marsh (1889), Discovery of Cretaceous Mammalia. Am. J. Sci. (3) xxxviii, p.81-92.

Link: Cretaceous "Hell Creek Faunal Facies", Philip Bigelow http://www.dinosauria.com/jdp/misc/hellcreek.html The fossils of Hell Creek.

Species:	Cimolestes lucasi Rigby JK & Wolberg DL, 1987
Place:	San Juan Basin, New Mexico
Country:	USA
Age:	Upper Cretaceous
Remarks:	This is based on a right lower molar (m1). "A marsupial of unclear affinities according to Cifelli 1990d", ( Alroy, internet). Be that as it may, the typed fossil resides in the New Mexico Museum of Natural History and Science and is known as NMMNH P-10722, (originally UNM B-1728). Unfortunately, its precise location is now unclear, (Morgan & Lucas 1999, p.257). And Thanks to Martin Jehle for correcting the age!
Reference:	Rigby & Wolberg (1987), The therian mammalian fauna (Campanian) of Quarry 1, Fossil Forest study area, San Juan Basin, New Mexico. Spec Pap -- Geol Soc Amer 209, p.51-80.

Species:	Cimolestes magnus Clemens WA & Russell, 1965
Place:	Alberta, Saskatchewan & Montana, North Dakota, Wyoming
Country:	Canada & USA
Age:	Maastrichtian, Upper Cretaceous
Remarks:	"Other Cretaceous species such as Cimolestes magnus have high crowned, pointed cusps with relatively long shearing blades. This, coupled with its larger body size, suggest that this species may have been carnivorous", (Gordon 2003, p.176-177). The suggested weight of around a kilo (Alroy) explains the species name well. Gordon & Cifelli, 2003 contains a more modest range of 472-817g, (p.93). The holotype resides in the collection of the University of Alberta. This is part of the left of the dentary with a premolar and molars, (p4-m4). It came from the Scollard Formation of Alberta, (Hunter & Archibald 2002, p.199).
Reference:	Clemens & Russell (1965), Mammalian fossils from the upper Edmonton Formation, University of Alberta Geology Bulletin, 2, p.32-40.

Species:	Cimolestes propalaeoryctes Lillegraven, 1969
Aka:	C. propaleoryctes
Place:	Scollard Formation, Alberta, Saskatchewan & Hell Creek, Montana
Country:	Canada & USA
Age:	Maastrichtian, Upper Cretaceous
Remarks:	Also studying at Alberta. This was a small version of about 38-63g, (Gordon & Cifelli 2003, p.94).
Reference:

Species:	Cimolestes simpsoni (Reynolds, 1936) Clemens WA, 1973
Aka:	Puercolestes simpsoni, Reynolds, 1936
Place:	San Juan Basin, New Mexico & Montana
Country:	USA
Age:	Puercan, Paleocene
Remarks:
Reference:

Species:	Cimolestes stirtoni Clemens WA, 1973
Place:	Frenchman Formation, Saskatchewan & Montana, Wyoming
Country:	Canada & USA
Age:	Maastrichtian, Upper Cretaceous
Remarks:	One Wyoming specimen at the Peabody. Weight estimate of 312-343g, (Gordon & Cifelli 2003, p.94).
Reference:	Clemens (1973), University of California Publications in Geological Sciences 94.

Genus: Didelphodus Cope, 1882 Aka: Centetodon Marsh, 1872 (partly); Deltatherium (partly); Didelphyodus Winge, 1923; Phenacops (Matthew, 1909b) In the view of John Alroy, a further genus within Insectivora.

Reassigned species: D. ventanus White, 1952 see D. altidens

Species:	Didelphodus absarokae (Cope, 1881) Cope, 1882
Aka:	Deltatherium absarokae Cope, 1881
Place:	Wyoming & Colorado
Country:	USA
Age:	Wasatchian, early Eocene
Remarks:	200g of insect nibbler (Alroy).
Reference:

Species:	Didelphodus altidens (Marsh OC, 1872)
Aka:	Centetodon altidens Marsh OC, 1872; D. ventanus White T, 1952; Phenacops incerta Matthew (1909b)
Place:	Colorado
Country:	USA
Age:	Wasatchian, early Eocene
Remarks:	A probable bodylength of 20-25cm and a weight of around 150g (Alroy). Descendant of D. aksarokae, (Alroy, Internet).
Reference:

Species:	Didelphodus rheos Storer 1995b
Place:
Continent:	North America
Age:	Uintan, middle Eocene
Remarks:	Weighed about 140g (Alroy).
Reference:

Species:	Didelphodus serus Storer JE, 1984
Place:	Cypress Hills Formation, Saskatchewan
Country:	Canada
Age:	Uintan, middle Eocene
Remarks:	A more modest 130g (Alroy).
Reference:	Storer (1984), Mammals of the Swift Current Creek Local Fauna (Eocene: Uintan), Saskatchewan. Earth Sciences, Natural History Contribution No. 7, Royal Saskatchewan Museum. pp.158.

Link: Publications of the Royal Saskatchewan Museum http://www.royalsaskmuseum.ca/wedo/publications/02-earthsciences.html?view=l Includes a brief summary, (and likewise for a lot of other papers).

Species:	Didelphodus sp.
Place:
Continent:	Europe
Age:	Eocene
Remarks:	Information welcome.
Reference:

Genus: Gelastops Simpson, 1935 Aka: Emperodon Simpson, 1935; Gelastopus Remarks: Within Insectivora, (Alroy). Several teeth have been referred to Gelastops sp. by Scott, 2003 (p.761-763). They come from the Who Nose? fauna of Calgary, Alberta, and consist of two upper premolars (P4s), a molar (M2) and a lower molar (m3). They at least resemble those of this genus. The site is Torrejonian, Paleocene.

Species:	Gelastops joni Rigby, 1980
Place:	Swain Quarry, Montana
Country:	USA
Age:
Remarks:	Weight estimate of about 180g (Alroy).
Reference:

Species:	Gelastops parcus Simpson GG, 1935
Aka:	Emperodon acmeodontoides Simpson, 1935
Place:	Silberling Quarry?
Country:	USA
Age:	Torrejonian, Paleocene
Remarks:	Around 140g (Alroy).
Reference:	Simpson (1935), New Paleocene mammals from the Fort Union of Montana. Proceedings of the United States National Museum, 83, p.221-244.

Genus: Ilerdoryctes Marandat, 1989

Species:	Ilerdoryctes sp. Marandat, 1989
Place:
Continent:	Europe
Age:	Lower Eocene
Remarks:	Further information welcome.
Reference:

Genus: Maelestes Wible JR, Rougier GW, Novacek MJ & Asher RJ, 2007 Family: Cimolestidae 'MAE robber' Remarks: The generic name involves an acronym. 'Mae' stands for the Mongolian Academy of Sciences-American Museum of Natural History Expeditions. It was a fair few breaths away from being called 'Maosamonhe robber'.

Species:	Maelestes gobiensis Wible et al, 2007
Place:	Ukhaa Tolgod
Country:	Mongolia
Age:	Campanian, Upper Cretaceous
Remarks:	The following is based upon my reading of Wible et al, 2007, and thanks are due to Christopher in the Land of Oz (down under). Mongolia's Ukhaa Tolgod strikes again. Ukhaa Tolgod actually translates as 'brown hills'. To mammal paleontologists, 'huge heaps of gold' could be figuratively more apt. This gold is in skulls rather than ingots, hundreds of them. Some even come supplied with nigh-on complete skeletons. Such treasures glow irresistibly before the eyes of lustful mammalogists, and still more of them never sate the desire. It sharpens the lust yet further. More, more, I must have more, and Ukhaa Tolgod keeps dishing it out like a nymphomaniac on heat. Its latest contribution to carnal lust is a eutherian (p.1003), and it asked the expectant researchers to conduct some enquiries concerning its relatives. For example, it wanted to know whether it qualified as a placental mammal. Sadly, the researchers shook their heads as, while all mammals of this age are welcomed with open arms, these limbs would be stretched to breaking point for an undoubted placental. We're biased about such matters. They were rather taken aback to hear Maelestes sigh with relief and reply: "Thank God for that. I was afraid somebody would accuse me of giving rise to critters like you lot." I should say this episode isn't explicitly recorded in the study but, as I've now read it on the internet, then it must be true. Of course, as some non-placental eutherian was responsible for the production of Placentalia, its reputation is perhaps not entirely beyond reproach in this regard, but it didn't have brains enough to think in those terms, and was satisfied with the answer received. Dental formula and distinctions The dental formula per side is as follows: (uppers): ? incisors, 1 canine, 5 premolars, 3 molars; (lowers): 3, 1, 5 and 3 respectively. That count of five premolars isn't known from other local eutherians, but it has been encountered on earlier models elsewhere in Asia. Also different is having a trio of procumbent lower incisors of roughly similar size, and a vacuity (a hole) in the bony palate between the maxilla and palatine bone. Several other details are stated as well. Its lower canine has only one root, and this is in contrast to asioryctitherians. The same can be said for the tri-rooted penultimate upper premolar. Upper molars are broader than long, with the stylar shelves on the buccal side being relatively narrow. The protoconid on lower molars is close in size to the metaconid. The cones on upper molars are weedier affairs than those boasted about by 'zhelestid' mammals, the metacones are far inferior to the paracones, and both these structures share a common base. The talonid of lower molars is narrower than the 'zhelestid' version. The single roots of the lower canine and first premolar are characteristics shared with some later North Americans, Cimolestes and Batodon, and at least one specimen of the former also possessed two similarly sized, procumbent lower incisors. Molar details, the narrow stylar shelf among them, are in common with the second of that pair. Differences from them include an 'extra' premolar, weaker molar cones, and lower molars with smaller trigonids containing similarly sized proto- and metaconids. Affinities An analysis of 408 morphological characters was conducted for 69 taxa. According to the results, the new genus is most closely related with that North American duo. Some earlier views linked Cimolestes with placental carnivorans, and that would necessitate it having been a placental itself. However, this survey doesn't support such a conclusion. Actually, it fails to recognize any known Cretaceous taxa as being members of Placentalia, let alone allies of particular extant orders. Maelestes seems closest to Batodon, which was an animal of around only 60% the size. It has resemblances to other Cretaceous eutherians, but in a well mixed cocktail of a manner. The premolars aren't unlike those of Zhelestes, and lower molars are more like Zalambdalestes. It even manages a marsupial-like trait. There's a naturally occurring gap in the bony palate termed a palatal vacuity. That's not previously been seen for a Cretaceous eutherian. However, as this isn't accompanied by other metatherian qualifications, it wasn't the result of inheritance from a common ancestor. Although not typical for eutherians, such a vacuity is known from a few, eg. hedgehogs and elephant shrews (p.1004). What remains of the skeleton is most akin to the gorgeous body of Ukhaatherium, a most sexy mammal from the same Mongolian locality. It's party time As mentioned, the authors decided to arrange a get together for 31 Cretaceous eutherians, 20 Tertiary ones, 11 extant placentals and 7 non-eutherian cousins. All were allowed to chum up with whomsoever they pleased, but only those meeting a strict series of criteria were given tickets to get past the bouncers at the doors of the Placentalia Ball Room. This dancing area was reserved only for the ancestor of all recent placentals, and any of its descendants. The authors say they invited: "...all but the most incomplete and poorly preserved taxa...". I don't wish to cast aspersions, but I'm not sure how that's quite so. Schowalteria phoned me up, and told me it received no notification of this party at all. Schowalt was rather sad about it all. It was described as an Upper Cretaceous taeniodont, an extinct order generally seen as placental. I don't know why it wasn't on the published guest list. Perhaps it had computer problems, and the e-mailed invitation bounced. In sympathy with the views offered earlier by Maelestes, not one of the Cretaceous contingent showed any interest in entering the Placentalia Ball Room. They all preferred to either remain alone or team up with other Cretaceous eutherians. At least nine of these singletons or groups developed. The venerable Eomaia, for example, hung around close to the main door, and was inclined to nothing more than a bit of polite chit-chat with Prokennalestes and a few others. But none of the five most basal members deigned to engage in group building. The nearest signs of a proper get together forming occurred among the generally poorly preserved 'zhelestids'. Remains have been gathered from Asia, Europe, North America and possibly Madagascar (disputed). 'Zhelestid' upper molars have strong protocones, and this was one reason for linking them with placental ungulates. However, convergence now seems easily the most plausible explanation, and not shared ancestry (p.1005). Trouble at the Ball Room door Loud squeaking indicates there's some excitement near the bouncers guarding the entrance to the Placentalia Ball Room. Perhaps somebody's trying to gatecrash it. I wonder what's going on, so let's go over and see. The bouncers are looking a bit surprised, so I expect they're trying to figure out how someone got past them. Who got in? Oh, nobody! Apparently, a couple ran out. As the name suggests, Protungulatum, a Paleocene (pseudo)-'condylarth', has long been accused of being the first known placental ungulate. However, accompanied by its friend, Oxyprimus, it's proclaimed itself not to be a placental in any shape or form, and any similarities with ungulates are but convergent shapes or forms. Of hamsters and relatives Still, now that we're by the door anyway, and happen to be placental mammals, I'm going into the Ball Room to take a quick peep at some of the Glires guys. This gang consists of mice, bunnies and their relatives; rodents and lagomorphs. They share some dental specialisations with Cretaceous zalambdalestids, who I prefer to call zalams for reasons that should be obvious. These similarities include large front, ever-growing incisors with self-sharpening edges resulting from a trick involving the restriction of enamel. The band of hard stuff wears down less quickly than the unprotected dentine at the rear, and the consequence is a very sharp chisel. However, the zalams didn't have all glirid dental developments, and there are no obvious intimate corresponding characters in the skull. The sort of result that can be inflicted by bites from such incisors is known, in the complex scientific jargon, as bloody painful, with bloody being an apt adjective. Hamsters may look harmless enough, but should you fancy digging them out of their nests -now illegal in much of Europe, and you corner one of these critters where no means of escape are available, then attempting to grab your furry friend with unprotected hands could necessitate the attentions of some kind aunties and uncles in your local hospital. Hamsters in Western Europe now lead lives unmolested by spade-wielding primates, and this has been a consequence of their widespread eradication. A hundred years ago, premiums were on offer for their bodies, and the demand from public officials was higher for female bodies. (I've heard some men willingly pay more for male bodies, but that's in a different trade.) The premiums could be higher for females due to their possession of baby delivery systems. Nest raiding was a useful source of supplementary income and, thanks to their prolific storing of grain, chicken feed. In the right circumstances, they can have five kilos of the stuff squirreled away in the burrow. In German, hamstern is a verb meaning 'to hoard. While such human behaviour couldn't have been described as polite, it made little impact on hamster population levels. People actually appear to be rubbish at wilful eradication in most circumstances. Hamster numbers sometimes surged to plague proportions, thus the inducements to kill. Modern agricultural practices have brought things to pass that earlier countryside dwellers wouldn't have dared pray for; hamsters as endangered rodents. More intensive ploughing wrecks the burrows, combine harvesters don't even bother to leave mincemeat of the bodies, should any be encountered, and then there are the effects of various chemicals. Such factors have achieved what dogs and digging couldn't manage. None of this natter is of any particular relevance to zalams, let alone our Maelestes. The dental similarities of zalams and rodents appear to be whims of convergence. Still, just because something isn't relevant, that doesn't need to mean it's uninteresting. Besides, if you want to get some idea of how ancient small mammals live, then the best ideas can be provided by some living small mammals. While details and characters have changed over time, many of the playing rules have remained broadly similar. I've got a translation of an old German article on Prussian hamsters, should anybody be feeling peckish for the theme. Kosmos presents Kill the hamster Zalams, asiorycties and cimolestids Following the paw prints left by Protungulatum and friend allows us to return to the Cretaceous festivities, and we find three more social gatherings. In contrast to the 'zhelestids', all now have members known from reasonably complete heads and, in the first two cases, bodies (p.1005). The zalams are accused of being further derived than asiorycties in this study, and that means somewhat closer to Placentalia. Asiorycties are more generalized critters, and they chum up with the family of Cimolestidae containing, in terms of the study, Cimolestes, Batodon and Maelestes. It may contain other genera as well, but bringing order to the family wasn't one of the objects of this particular exercise. In any case, Maelestes is the earliest known member, and presently the only one from Asia. Holotype The type fossil, PSS-MAE607, is a partial skull renting accommodation at the Geological Institute, Ulaan Baatar. As is frequently the case these days, it opted to stay in the country rather than emigrating to some distant collection in New York, Moscow, Warsaw or wherever. Parts of the skeleton are also available for enjoyment. In the interests of promoting mystery, I'm not going to divulge the reason behind the specific name of this animal from the Gobi Desert(s).
Reference:	Wible JR, Rougier GW, Novacek MJ & Asher RJ (2007), Cretaceous eutherians and Laurasian origin for placental mammals near the K/T boundary, Nature, 447, p.1003-1006.

Genus: Naranius Russell DE & Dashzeveg D, 1986

Species:	Naranius infrequens Russell DE & Dashzeveg D, 1986
Place:
Continent:	Asia
Age:	Lower Eocene
Remarks:	Further information welcome.
Reference:

Genus: Paleotomus (Simpson GG, 1937) Van Valen L, 1967 Aka: Niphredil Van Valen L, 1978; (partly) "Paelaeosinopa" Simpson, 1937 Remarks: John Alroy, doubtlessly wisely, places this genus within the order Pantolesta.

Species:	Paleotomus carbonensis Secord R, 1998
Place:	Wyoming
Country:	USA
Age:	Torrejonian, Paleocene
Remarks:
Reference:	Secord (1998), Paleocene mammalian biostratigraphy of the Carbon Basin, southeastern Wyoming, and age constraints on local phases of tectonism. Rocky Mountain Geology 33, p.119-154.

Species:	Paleotomus junior Scott CS, Fox RC & Youzwyshyn, 2002
Place:	Paskapoo Formation, Alberta
Country:	Canada
Age:	Tiffanian, Upper Paleocene
Remarks:	P. junior is represented by a number of lower and one upper tooth. The authors conclude its family affinities are uncertain. "Smallest species of the genus; differs from Paleotomus senior (Simpson, 1937), Paleotomus milleri Rigby, 1980, Paleotomus radagasti (Van Vaalen, 1978), and Paleotomus carbonensis Secord, 1998 in its significantly smaller dental dimensions (approximately 40 percent less than Paleotomus senior, 20 percent less than Paleotomus milleri, 50 percent less than Paleotomus radagasti, and 35 percent less than Paleotomus carbonensis); differs further from P. milleri in m1 lacking an entoconulid and mesoconid", (Scott et al 2002, p.702). The specimens are students of the University of Alberta.
Reference:	Scott, Fox & Youzwyshyn (2002), New earliest Tiffanian (late Paleocene) mammals from Cochrane 2, southwestern Alberta, Canada. Acta Palaeontologica Polonica 47 (4), p.691-704.

Link: Acta Palaeontologica Polonica 47 (4), p.691-704 http://www.paleo.pan.pl/acta/acta47/app47-691.pdf Scott & Co 2002, the full paper in pdf format

Species:	Paleotomus milleri Rigby JK, 1980
Place:	Swain Quarry, Wyoming
Country:	USA
Age:	late Torrejonian, Paleocene
Remarks:
Reference:	Rigby (1980), Swain Quarry of the Fort Union Formation, middle Paleocene (Torrejonian), Carbon County, Wyoming: geologic setting and mammalian fauna. Evolutionary Monographs 3, p.1-179.

Species:	Paleotomus radagasti (Van Valen L, 1978)
Aka:	Niphredil radagasti Van Valen L, 1978
Place:	Cedar Point Quarry, Wyoming
Country:	USA
Age:	Tiffanian, Paleocene
Remarks:	The original author, Lee Van Valen, seemingly enjoyed the Middle Earth (a lá Tolkien) as much as the Mesozoic. "Niphredil is a small Middle Earth flower. Radagast the Brown was a wizard", (with thanks to Mark Isaac, Curiosities of Biological Nomenclature .
Reference:	Van Valen (1978), The beginning of the Age of the Mammals. Evolutionary Theory 4, p.45-80).

Species:	Paleotomus senior (Simpson GG, 1937) Van Valen L, 1967
Aka:	"Paelaeosinopa" senior Simpson, 1937; "Paelaeosinopa" simpsoni Van Valen, 1967
Place:	Montana & Paskapoo Formation, Alberta
Country:	USA & Canada
Age:	upper Torrejonian? Paleocene
Remarks:	Simpson based the species on one lower and two upper molars. Subsequently, Van Valen erected this genus for the former, whilst referring the upper teeth to "Paelaeosinopa" simpsoni. They were then synonymized by Gingerich in 1980, (Scott et al 2002, p.720).
References:	Simpson (1937), The Fort Union of the Crazy Mountain Field, Montana, and its mammalian faunas. Bulletin of the US National Museum, Smithsonian Institution, 169, p.1-287.
	Van Valen (1967), New Paleocene insectivores and insectivore classification. Bulletin of the American Museum of Natural History 135, p.221-284.
	Gingerich (1980), A new species of Palaeosinopa (Insectivora: pantolestid) from the late Paleocene of western North America. Journal of Mammalogy 61, p.449-454.

Genus: Procerberus Sloan & Van Valen, 1965 'before Cerberus' Remarks: A close relative of Cimolestes, especially C. incisus and C. simpsoni, (Alroy, internet, linked towards the end of this page). Yale is home to several P. sp. specimens, one of whom claims to be Cretaceous.

Links: The Encyclopaedia Britannica, Evolution and Paleontology http://www.britannica.com/eb/article?eu=108399&tocid=51573&query=raccoons Besides which, the Britannica says it’s Cretaceous and who am I to argue with them? Cerberus is "the hound of the underworld," (which) "stood guard at the gates of Hades and prevented those not permitted from entering," (with thanks to Classical Mythology ). Cerberus is also a snake genus.

Species:	Procerberus formicarum Sloan RE & Van Valen L, 1965
Place:	Bug Creek Formation and lower Tullock Formation, Montana, perhaps also the Ravenscrag Formation, Saskatchewan
Country:	USA & ?Canada
Age:	Maastichtian, Upper Cretaceous? - Puercan, Paleocene
Remarks:	The following has been gleaned from my reading of Lofgren, 1995 rather than the original citation that I haven't seen. Feel free to send a copy. Lofgren was writing about specimens from seventeen McGuire Creek localities in Montana (p.121). They were so similar to collections made earlier that little description was required. A deciduous upper premolar did receive some brief coverae, a DP4. About the size of things Lengths of McGuire Creek specimens are on pages 122-123. Only those from four localities are listed. These include the Bug Creek Anthills sampled earlier by Sloan and Van Valen. Uppers: P3 (6 specimens) 2.23-2.81mm; P4 (7 sp.) 2.40-2.77mm; M1 (12 sp.) 2.43-2.92mm; M2 (9 sp.) 2.48-2.87mm; M3 (12 sp.) 1.94-2.73mm. Lowers: p4 (8 sp.) 2.60-3.21mm; m1 (18 sp.) 2.61-2.99mm; m2 (17 sp.) 2.44-3.10mm; m3 (15 sp.) 2.91-3.30mm. Holotype UMVP 1460 is a partial dentary studying law at the University of Minnesota, Minneapolis. Additional notes Weighed in at around four mice, 100g (Alroy); 55-69g, (Gordon & Cifelli 2003, p.98). Material has also been found in the Hell Creek Formation. If not due to reworking, this species would also be Upper K.
Reference:	Sloan & Van Valen (1965), Cretaceous mammals from Montana. Science, 148, p.220-227.

Species:	Procerberus plutonis Van Valen L, 1978
Place:	Purgatory Hill, Montana
Country:	USA
Age:	Puercan, Paleocene
Remarks:	As far as I understand it, this species seems to be based on upper dentition, (Scott 2003, p.761).
Reference:	Van Valen (1978), The beginning of the Age of Mammals. Evolutionary Theory, 4, p.45-80.

Species:	Procerberus andesiticus
Place:	Denver Formation, Colorado
Country:	USA
Age:	lower Puercan, Lower Paleocene
Remarks:	One Wyoming specimen at the Peabody.
Reference:	P. andesiticus and P. grandis both feature in the linked article on the Littleton Local Fauna. Eric Dewar was kind enough to inform me, that some of the mentioned species are as yet unpublished, (pers. comm.), though work is in progress.

Link: Eric W Dewar http://www-unix.oit.umass.edu/~dewar/research/SVP97poster.html Dietary inferences for archaic ungulates from the Early Puercan Littleton Local Fauna (Denver Formation, Colorado).

Species:	Procerberus grandis
Place:	Denver Formation, Colorado
Country:	USA
Age:	lower Puercan, Lower Paleocene
Remarks:	As yet unpublished.
Reference:

Genus: Protentomodon Simpson GG, 1928

Species:	Protentomodon ursirivalis Simpson, 1928
Place:	Bear Creek, Montana
Country:	USA
Age:	Tiffanian, Upper Paleocene
Remarks:	Belongs to the order of Pantolesta, (Alroy).
Reference:

Genus: Schowalteria Fox RC & Naylor BG, 2003 'for Schowalter' Remarks: The generic name honour's the discoverer and collector of the specimen, DB (Tim) Schowalter.

Species:	Schowalteria clemensi Fox RC & Naylor BG, 2003
Place:	Scollard Formation, Alberta
Country:	Canada
Age:	Maastrichtian, Upper Cretaceous
Remarks:	The following has not been compiled with direct reference to the description. It's based on information forwarded by John Bois, and extensive details and comments provided by David Marjanovic. Many thanks and felicitations to both. Any misrepresentations have been added by me. Presently, only one specimen of this mammal has been identified. This consists of much of the front of a skull extending back to the holes for the eyes. Both upper and lower jaws are preserved. The most striking feature is the size. This is one of the largest Mesozoic mammals known. It was about Didelphodon-big, which translates as something like the size of a badger or maybe fox. That might be slighter smaller than a twelve metre long Tyrannosaurus, (which lived in the neighbourhood at the time). However, a Meso mammal with a body length of something like 70cm (give or take 10 or 20) can sensibly termed enormous, in mammalian terms. Fossils from this particular location (KUA-1) belong to the Trochu local fauna. That fauna includes multituberculate, metatherian and eutherian mammals. Schowalteria represents a eutherian lineage called Taeniodonta, which was previously unknown from the Cretaceous. It also extends the geographic range of taeniodonts northwards by 600km. This raises the question as to why remains haven't turned up in well-sampled Cretaceous localities in the USA. Although not common, taeniodonts are mainly known from the Paleocene deposits of the US. A convincing explanation is provided by a possible southwards extension of the group following the K-T extinction event(s). However, other factors such as habitat preference may also have played a role. A similar distribution pattern for other mammals, eg. Parectypodus, can be seen as support for the first possibility. The only Cretaceous record of that genus is from Saskatchewan. US representatives are restricted to the Paleocene and Eocene. The holotype is affectionately known as RTMP 93.90.01, and it works in the Royal Tyrrell Museum, Drumheller, (as did Mr Schowalter at one time). The species name is in honour of WA Clemens, who was the first to realize the taeniodont-like nature of the molars. Thanks are also due to Christopher Taylor for posting the info on the DML, and Dr Don Brinkmann of The Royal Tyrrell for confirming they are the employers of the specimen.
Reference:	Fox & Naylor (2003), A Late Cretaceous taeniodont (Eutheria, Mammalia) from Alberta, Canada. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 229(3), p.393-420.

Links: Christopher Taylor, Dinosaur Mailing List, 2.4.2004 http://www.cmnh.org/dinoarch/2004Apr/msg00018.html The message includes some information from the abstract of the paper. Martin Jehle, Taeniodonts and tillodonts: Strange rooters and diggers http://www.paleocene-mammals.de/taeniodonts.htm An admirer of Paleocene mammals pays homage to taeniodonts which followed Schowalteria. David Marjanovic, Dinosaur Mailing List, 3.4.2004 http://www.cmnh.org/dinoarch/2004Apr/msg00058.html Details from and comments on the description. This is an extensive posting.

Genus: Tinerhodon Gheerbrant E, 1995

Species:	Tinerhodon disputatum Gheerbrant E, 1995
Place:	Adrar Mgorn 1
Country:	Morocco
Age:	Upper Paleocene
Remarks:	This seems to be some kind of creodont or other, and is of very uncertain affinities.
Reference:

Genus: Tsaganius Russell DE & Dashzeveg D, 1986

Species:	Tsaganius ambiguus
Place:
Continent:	Asia
Age:	Lower Eocene
Remarks:	Information welcome
Reference:

Other reports: France "A single tooth of a possible cimolestid-like mammal has been reported from the Late Cretaceous (Campanian) of France," (McKenna & Bell, 1997, p.212). Khulsan, Mongolia Undescribed material, possibly Upper Cretaceous, may represent the family of Palaeoryctidae Winge, 1917, (McKenna & Bell, 1997, p.211). Update According to Bolortsetseg M, Minjin C & Geisler J (2006 Abstracts of the Society of Vertebrate Paleontology, p.44A), something is astir in the Maastrichtian of Mongolia; and it involves a strange, palaeoryctid-like critter. Until this announcement, the mammals of this latest stage of the Upper Cretaceous for that land consisted of an undescribed metatherian skull, perhaps the multituberculate named Buginbaatar and... That was it. The latter come from Khaichin-I, and the poorly represented fauna could perhaps have been Paleocene (which is news to me). However, further prospecting work has increased the haul, it now includes bits of non-birdy dinosaurs, and they say (in conjunction with stratigraphic work) that they date from the Maastrichtian. As a further resident is a possibly basal metatherian, Deltatheridium, the mammalian contingent has scrambled to three. The eutherian killer The third is the eutherian, and this would appear to be a very democratic mix for the time. One of the two specimens found is a fragment of skull with some appealing oddities. Upper canines are likened to sabres, the lower incisors vary from modesty to absenteeism, and there are also but two premolars downstairs. The molars feature talonids which are more narrow than the trigonids, a trait shared with the palaeoryctids and Cimolestes. Other molar features mentioned are towering trigonids, 'transversely elongate' upper molars (is that code for 'wide'?) and the paracone of the P4 being higher than its equivalents on the molars. This is a Mongolian mammal with something of a North American flavour, and that can also be said for its multi friend, Buginbaatar. As the remains haven't yet been formally described, no name has been proposed. Burgos, Spain "A possible palaeoryctid is represented by unnamed specimens from the Calizas de Lychnus Fm., Maastrichtian of Burgos, Spain," (McKenna & Bell, 1997, p.211). Alberta, Canada One tooth (an m3) from a site called Who Nose?, Calgary, has been referred to cf Procerberus sp. It's 3mm long. A more precise placement would require better evidence, which may turn up. The location is upper Torrejonian, Paleocene, and would extend the chronological range of the genus back by about two million years, (Scott 2003, p.761). Tiupampa, Peru A fragmentary premolar from this Paleocene locality is perhaps referrable to Cimolestes, (de Muzion & Cifelli 2001, p.95). A further possible location in the Peruvian Paleocene is Chulpas, (p.96).

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, June 2002. Latest update: 9.5.2008 Ktdykes@arcor.de

With further thanks due to: Dr John Alroy, North American Fossil Mammal Systematics Database http://www.nceas.ucsb.edu/~alroy/nafmsd.html The source of much of the above information, including weight estimates. Weight estimates Weight estimates have generally, when not otherwise stated, been shamelessly stolen from John Alroy's internet site. When other sources are available, this may produce disparities. I've got two comments to offer. Firstly, if you were to claim that some European hedgehogs (Erinaceus europaeus) weigh 400 grammes, you'd be correct. If I were to add that some reach 1,2 kilos, I'd also be correct. Some hedgehogs are bigger than others. Secondly, the estimates partly depend upon the questions posed. If a calculation is based upon an insectivore model, the answer may be 50g. Choose a South American opossum, and it'd perhaps be closer to 150. Think primate, and 300g might result. A further source is Gordon & Cifelli, 2003 (p.93-97). This research offers various alternatives. These depend upon which tooth is used, (lower molar 1 or Upper Molar 1), and which group of animals it's compared to. I'll include the range of estimates based upon insectivores. These calculations were derived from a. m1 Length, b. m1 L x Width, c. M1 L and d. M1 L x W. A rough system of measurement is employed in these directories. A standard mouse = 25g, a rat counts as 400 whilst a beaver equals about 25 kilos. Yale Peabody Museum, Collection Search (VP) http://george.peabody.yale.edu/vp/ Genera and species of Paleocene mammals - Part 2, Martin Jehle http://www.paleocene-mammals.de/pal2.htm The Prehistoric Data Files http://www.angellis.net/Web/PDfiles/marsups.pdf The Bibliography of Fossil Vertebrates, John Damuth http://www.bfvol.org/ BIOSIS, The Index to Organism Names http://www.biosis.org.uk/triton/indexfm.htm

Bibliography: Archibald JD & Averianov AO (2001), Paranyctoides and allies from the Late Cretaceous of North America and Aisa, Acta Palaeonotologica Polonica, 46 (4), (Proof Version). Archibald JD, & Averianov AO (2003), The Late Cretaceous Placental Mammal, Kulbeckia. Journal of Vertebrate Paleontology 23(2), p.404-419. Archibald JD & Averianov AO (2005), Mammalian faunal succession in the Cretaceous of the Kyzylkum Desert, Journal of Mammalian Evolution, 12 (1/2), p.9-22. Archibald JD, Averianov AO & Ekdale EG (2001), Late Cretaceous relatives of rabbits, rodents and other extant eutherian mammals. Nature 414, p.62-65. 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 (2005), Mammals from the mid-Cretaceous Khodzhakul Formation, Kyzylkum Desert, Uzbekistan, Cretaceous Research 26, p.593-608. Benton MJ (1999), Early origins of modern birds and mammals: molecules vs. morphology. BioEssays 21, p.1043-1051. Fostowicz-Frelik L & Kielan-Jaworowska Z (2002), Lower incisor in zalambdalestid mammals (Eutheria) and its phylogenetic implications. Acta Palaeontologica Polonica, 47(1), p.177-180. Fox RC (2004), A new palaeoryctid (Insectivora: Mammalia) from the Late Paleocene of Alberta, Canada, Journal of Palaeontology, 78(3), p.612-616. Gordon CL (2003), Molar morphology and diet of Late Cretaceous therian mammals of North America: a preliminary analysis, p.149-177 (In) Functional Morphology and Diet of Late Cretaceous Mammals of North America, Ph.D. Dissertation, University of Oklahoma, p.i-xiv and 1-177. Gordon CL & Cifelli RL (2003), Estimating body size in Late Cretaceous therian mammals of North America, p.56-148 (In) Functional Morphology and Diet of Late Cretaceous Mammals of North America, Ph.D. Dissertation, University of Oklahoma, p.i-xiv and 1-177. Gregory WK & Simpson GG (1926), Cretaceous Mammal skulls from Mongolia, American Museum Novitates, 225, p.1-20. 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. Khosla A, Prasad GVR, Verma O, Jain AK & Sahni A (2004), Discovery of a micromammal-yielding Deccan intertrappean site noear Kisalpuri, Dindori District, Madhya Pradesh, Current Science, 87(3), p.380-383. 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. 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. 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. Muizon de C & Cifelli RL (2001), A new basal "didelphoid" (Marsupialia, Mammalia) from the Early Paleocene of Tiupampa (Bolivia), Journal of Vertebrate Paleontology, 21 (1), p.87-97. Rana RS & Wilson GP (2003), New Late Cretaceous mammals from the Intertrappean beds of Rangapur, India and paleobiogeographic framework. Acta Palaeontologica Polonica, 48 (3), p.331-348. Scott CS (2003), Late Torrejonian (Middle Paleocene) mammals from South Central Alberta, Canada. Journal of Paleontology, 77(4), p.745-768. Scott CS, Fox RC & Youzwyshyn GP (2002), New earliest Tiffanian (late Paleocene) mammals from Cochrane 2, southwestern Alberta, Canada. Acta Palaeontologica Polonica 47 (4), p.691-704. Simpson GG (1928), Affinities of the Mongolian Cretaceous Insectivores, American Museum Novitates, 330, p.1-11. Simpson GG (1951), American Cretaceous insectivores, American Museum Novitates, 1541, p.1-19. Simpson GG (1959), Mesozoic mammals and the polyphyletic origin of mammals, Evolution, 13, p.405-414. Simpson GG; Olson EC (1935/1976), The First Mammals, The Quarterly Review of Biology, 51, 50th Anniversary Special Issue, 1926-1976, p.45-73. (Note: this is a reissue with an introduction by Olson. The remainder of the paper is by Simpson, and has the page numbers 154-180.) 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, Novacek MJ & Rougier GW (2004), New data on the skull and dentition in the Mongolian Late Cretaceous eutherian mammal Zalambdalestes, Bulletin of the American Museum of Natural History, 281, p.1-144. Wible JR, Rougier GW, Novacek MJ & Asher RJ (2007), Cretaceous eutherians and Laurasian origin for placental mammals near the K/T boundary, Nature, 447, p.1003-1006. Zan S, Wood CB, Rougier GW, Jin L, Chen J & Schaff CR (2006), A new 'Middle' Cretaceous zalambdalestid, from a new locality in Jilin Province, Northeastern China, Journal of the Paleontological Society of Korea, 22(1), p.152-172.