MESOZOIC MAMMALS; Gondwanatheria, an internet directory

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MESOZOIC MAMMALS; Gondwanatheria, an internet directory:

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

In 1987, Mones referred Gondwanatheria to Multituberculata. This was originally widely accepted but then subsequently widely rejected. However, beginning from about 2005, various gangs of researchers began revising that rejection to some degree or other. Multi affinities seem to be back in the ring, either as full blown multis or allies. The background to all this is rather complicated. This directory should now be written from a perspective of gondwans as possible multis. What's also possible is that I may have neglected to emend older parts of the text.
Hopefully, at some stage, I'll also find time to build in more information from Gurovich's 2005 thesis on these critters, and their possible place in ancient Mammaldom. It's a work of epic proportions weighing in at over 600 pages, and packed full of details and discussion.

Introducing the gondwans
The following is largely based upon my reading of Gurovich & Beck, 2008, and thanks are due to the supplier. My copy happens to be a pre-publication version that doesn't conform to the publication in terms of page numbers. It can't do, seeing as it didn't have any. I've added some for my own reference purposes.
Gondwanatherians are an enigmatic group of southern hemisphere mammals, and their disputed affinities are still doing their utmost to fully justify that word 'enigmatic'. The first suggestion regarding their affinities sort to mate them up with xenarthran placentals such as armadillos and anteaters. Apart from anything else, they happen to occur in sediments from the sort of age and places where proto-xenarthrans wouldn't be all too surprising. Although eminently appealing as it might be regarded, that idea had to be abandoned.
It was superseded by their recognition as odd multituberculates, and the apparent molars (should appearances not be partly deceptive) do feature longitudinal rows of cusps, albeit with fairly radical architectual novelties. However, a bit of jaw came to light and proved less than helpful in confirming the idea. It showed a complement of four molars rather than the multi count of two. At least, it seemed to. That's two too many. There then reigned a period of much consensus; not xenarthrans, not multis but most definitely something or other. It was clear their affinities were opaque.
Multis again?
However, a different series of observations and thoughts, and a phylogenetic analysis, leaves at least links with multis not as dead as a dodo, and sets a new cat among these ancient mammalian pigeons. To attempt my own crude summary. Mainline multis had rear premolars with refinements for shearing. The uppers retained rows of cusps while the lowers started out with that sort of scheme (eg. paulchoffatiids). The shearing refinements were subsequently strengthened. The lower rear premolar (p5 of "plagiaulacidans", p4 for cimolodontans) developed into an impressively arched blade with serrations, and not much remaining of one cusp row. The results of this included strong distinctions between rear premolars and molars. Things didn't have to evolve like that. This is merely what occurred.
Now, what if things ran differently with some other multi lineage or, at least, a closely related group of mammals? Rather than becoming more distinct, the final two premolars may perhaps molarize; ie. become molar-like by enhancing grinding abilities. (Those aforementioned paulchoffies used these teeth more for grinding than shearing, and strongly molariform premolars occur in some mammals.) That sort of scenario could have led to a jaw half with four molariform teeth, of which only two were strictly speaking molars. In those circumstances, affinities of gondwans and multis would remain a viable possibility and, indeed, so would the inclusion of gondwans within Multituberculata. According to this phylogenetic analysis, of the mammals included in the enquiry, gondwans were most closely related with multis.
Feruglio gondwans as multis?
As well as Gurovich & Beck reaching conclusions on multi affinities, there's also a study by Rougier et al, 2008 going at least partly in a similar direction. One subject of that paper is the establishment of a new feruglio named Trapalcotherium. They found that at least ferugliotheriids could be multis (p.11), although they don't comment on the affinities of sudamericids. Due to the publication dates involved neither paper could be referenced in the other paper, and nor do any of the authors involved make "guest appearances" in the Acknowledgements of the "other team". The compatible conclusions appear to have arisen from convergence. That said, Gurovich's 2006 thesis on gondwans (my copy's got 2005 printed on it), in which all gondwans are cited as being a sister taxon of multis, is referred to in both papers.
Both teams also reach similar conclusions regarding Argentodites. This genus was said to be a likely SAm Cretaceous cimolod multi, but the opinions here see it as more likely a feruglio; either Ferugliotherium or a close relative. For bookkeeping purposes I'm presently keeping it as a separate feruglio genus.

A possible early representative has been reported from the Cretaceous of Tanzania, (Krause et al, 2003). This referral is 'very tentative'. Further details can be found in the Other Reports section.

Original citation: Mones (1987), Gondwanatheria, un nuevo orden de Mamíferos Sudamericanos (Mammalia: Edentata: ?Xenarthra). Comunicaciones Paleontológicas del Museo de Historia Natural de Montevideo, 18, p.237-240.

What Gurovich and Beck, 2008 came up with
This pair of authors were attending to several issues which, in one way or another (or several ways) have been controversial. A partial dentary with a p4 premolar had earlier been referred to Ferugliotherium (p.1). However, it was then re-identified as coming from some kind of non-gondwan multituberculate. G & B's opinion revives the original one; it is Ferug. An isolated premolar originally designated as Argentoides, a possible cimolodontan multi -the first from Gondwana, may be a p4 from a ferugliotheriid that hadn't been much affected by wear. Furthermore, they also engage in a phylogenetic analysis to test relationships of gondwantherians with each other and with other mammalian groups. Among other things, do the feruglios and sudamericids actually belong together? The answer appears to be yes. Do these gondwans share ancestry with multis and, if so, how closely related were they? The authors find gondwans probably share ancestry with both plagis and cimolod multis. There's evidence supporting a clade of Gondwanatheria and Multituberculata and, depending upon definitions, gondwans could even be members of multi-dom themselves.
However, it's also acknowledged that convergence could've brought about similar dental derivations by separate routes, and evidence from more, and better fossils could add clarity. (So far in this instance, better fossils have tended to have a different effect!)
Gondwanatheria
Gondwans, as used on this directory, presently fit into two families, and both are first encountered from Upper Cretaceous fossil localities, and particularly ones in Patagonia. One family, Ferugliotheriidae, have a less extreme dentition in terms of derivation. Consequently, they're held to be more basal, and provide a late persisting approximation of the tooth styles favoured by common ancestors of both feruglios and Sudamericidae. The latter is the more derived family that may have continued breeding until the Eocene.
Lower postcanines
Given their lack of canines, writing about 'postcanines' is perhaps inappropriate. You might prefer postincisors or cheek teeth. Be that as it may...
According to the description of the first known sudameri gondwan lower jaw (Sudamerica), it had four postcanine teeth (p.2). The first pair were molariform, and alveolae for another couple came behind. That seemed to indicate no premolars and four molars. Lower jaws of multis have four lower premolars (basal) or down to as few as one (derived). However, there's always at least one and, judging by every suitable specimen ever found, always, always absolutely always two molars. In short, none and four is a multi no no. The possibility that two molariforms may have evolved from the sort of narrow, blade-like final two premolars a lá basal multis, eg paulchoffatiids, was considered very unlikely. No such change had, as far as was known, ever been recorded.
Prior to that, a lower jaw had been assigned to Ferugliotherium, and it possessed a blade-like premolar. However, in the light of the new find, apparently demonstrating the lack of such teeth for gondwans, the fossil got thrown out of the genus, and was instructed to be a non-gondwan multi instead.
The position of Gurovich and Beck is that this fossil belongs to a feruglio, a relatively basal gondwan. The first two molariforms of more derived types are derived from those premolars. Nevertheless, they also recognize their view is still hypothetical and, consequently, choose to refer to molar and molar-like teeth as molariforms. (To qualify as a molar in strict terms being molar-like isn't enough. You also have to be a first generation tooth that doesn't get replaced by a successor at the same dental position. Replacement teeth need not apply.)
Ferugliotheriidae
Material of feruglios has been collected from Los Alamitos and La Colonia Formations (p.5). (After the publication of G & B, the Allen Formation got added to the list.) All localities are in Patagonia and date from the Upper Cretaceous. A highly tentative identification has been made for a possibly Eocene upper molariform from Peru.
Molariform teeth (presumably genuine molars in all cases) are brachyodont in style. That fine word effectively means, while high crowned, they don't go to the hypsodont extremes favoured by sudamericid gondwans. Their crowns have cusp rows running along their lengths, two for lowers and three for uppers, with lophs running across linking the usually poorly visible cusps. At the front of the lowers, for example, these produce a v-shaped structure followed by two roughly horizontal lophs, and then a c-shaped rear one.
Other material from the localities, including a fragmentary lower jaw with a worn, blade-like premolar, has been variously attributed to the genus of Ferugliotherium or a near relative, or a multituberculate. G & B support the former determination and, in a way, the latter one as well to some extent! All relevant isolated teeth and the jaw come from the same beds of Estancia Los Alamitos. Sizes and proportions match well with expectations for Ferugliotherium based upon comparisons with multis, some marsupials and placentals for forms with blade-like premolars. Should, on the other hand, the "multi" remains belong to a different taxon, then that would require two similarly sized but unrelated mammals in the same fauna with, judging by the similar dental specialisations, very similar ecological niches. That seems like crowding. And it's odd that one of these critters, Feruglio, is only known from isolated molariforms while the other, a nameless multi, has only bequeathed us a partial dentary and premolars.
Furthermore, an assumption that a blade-like premolar can't be remodelled into a molariform premolar has been challenged by a group of widely respected experts; sthenurine kangaroos (p.7). They maintain that their ancestors were up to this sort of challenge. And, if that can be done by relatives of Skippy, then gondwans may well have laughed at the word "impossible" (or any equivalent Mesozoic squeaks).
In any event, these authors conclude that only a single taxon is involved, and combined the information for use in their analysis. Should, at some point, fossils turn up showing this was wrong, then that'll be inconvenient.
Sudamericids
Sudameri gondwans are the more derived group, and have some traits not held in common with the less flamboyant feruglios. They had large, procumbent incisors with enamel restricted to particular areas. There were at least four tall (hypsodont) molariform teeth and no premolariform ones. That doesn't necessarily mean a lack of premolars as such. As G & B argue, it's possible to have molariform premolars.
Analysis
The authors interviewed what remains there are of Sudamerica, Gondwanaterium and Ferugliotherium (p.9), and scored them in accordance with a matrix of (mostly) mammals compiled by others in 2007. They threw them into this virtual pot containing about fifty taxa and 269 characters. The majority of characters are unknown for gondwans and, consequently, couldn't be scored. Only thirty applied for Sudam, 28 for Feruglio and twenty for Gondwan. Anyway, a number of fifty pence coins were fed into the electricity meter, and the computer was left to whiz and whirr its way towards doing the best it could for a reconstruction of relationships.
Evidence was found supporting a clade containing gondwans and multis (p.10). Unclear is whether gondwans are also descendants of the most recent common ancestor of all other multis or, rather, whether they are descendants of a somewhat earlier ancestor of the first multi. Simply put, as to whether they're multis themselves or merely close relatives. In either event, these results support a close relationship or an even closer one.
Convergence of gondwans and multis?
Am alternative possibility is acknowledged. The uncontested dental similarities could be results of convergence (p.13). For example, both my molars and those of a pig share similarities, but I wouldn't insult the dear animal by calling it an ape. (My wife wishes to add that similarities between myself and a pig aren't only dental.)
Postcanines with cusp rows have independently arisen in eucynodont history for mammals as diverse as multis, marsupials and rodents. Blade-like premolars have also come about more than once. Testing this possibility would require more gondwan body parts (p.14). If, for example, the skull happens to be constructed radically differently for gondwans and multis, that would indicate dental convergence. However, as such evidence for gondwans isn't yet available, this sort of comparative test clearly has to await further discoveries.
Somewhere, in some part of former Gondwana or other, there is presumably a fantastic Cretaceous or Paleocene cemetery containing breathtakingly brilliant gondwan corpses. A Gondwanan reply to the splendours of the Lower Cretaceous Yixian Formation, China would be kind of nice. As I'm writing these notes in November, perhaps I'll mention the idea to Father Yuletide and see what he pulls out from his sack of pressies.
Editorial note
Bits and pieces drawn from G & B, 2008 are likely to be scattered to various points of this directory, and I'll try to update older writing as I go along. However, I don't intend to attempt a systematic re-re-reading of all my notes here at this stage. Let me know if any discrepancies are found loitering around. It's also worth mentioning that I haven't, as yet, managed to read Gurovich's 2005 epic on gondwans, an extensive and rich feast of information. Making better use of that source is somewhere on my unwritten things to do list.
Another thing on that list is printing this entire directory out and having a look at the thing. Some tidying would probably improve the presentation. I'll be spared that torture until after the Gurovich 2005 epic.

Links:

Mikko Haaramo's Allotheria

Mikko Haaramo's Allotheria

A quick guide to multidom, assuming that's what godwanatherians were.

Jeff Poling, Geological Ages of Earth History

http://www.dinosauria.com/dml/history.htm

A long, long time ago...

Order Xenarthra, The University of Michigan

http://animaldiversity.ummz.umich.edu/chordata/mammalia/xenarthra.html

In case this taxon indeed turns out to be somewhere near the origins of edentates, here's a guide to what happened later; armadillos, sloths and anteaters.

Journal of Vertebrate Paleontology, 19(2), 1999, p.373

http://www.vertpaleo.org/jvp/19-373-382.html

Pascual R, Goin FJ, Krause DW, Ortiz-Jaureguizar E, & Carlini AA. The first gnathic remains of Sudamerica... 'Gnathic' refers to the jaw.

Palaeos, Allotheria, The Vertebrates, Toby White

http://www.palaeos.com/Vertebrates/Units/Unit420/420.100.html

Details on the dentistry.

Genera: Argentoides, Bharattherium, Ferugliotherium, Gondwanatherium, Lavanify, Sudamerica, Trapalcotherium Vucetichia (= Ferugliotherium), other reports

Time-Line:

Eocene: Antarctica

Paleocene: Sudamerica

Upper Cretaceous: Argentoides, Bharattherium, Ferugliotherium, Gondwanatherium, Lavanify, Trapalcotherium, Tanzania (maybe)

Genus: Argentodites Kielan-Jaworowska Z, Ortiz-Jaureguizar E, Vieytes C, Pascual R & Goin FJ, 2007

'Argentinean traveller'

Remarks: Update 2008, warning
This genus has fallen into disgrace in the eyes of some. Although orginally accused of being a possible cimolod multituberculate, other authors cite it as more likely being a ferugliotheriid gondwanatherian or perhaps a feruglio multi or, conceivably, a ferglio gondwan multi. Its affinities and the utility of the genus have both been brought into question although, as yet, nobody has formally carried out taxonomic euthanasia. Presently, the following entry remains as it was when I origially wrote it, but these developments are obviously of great relevance.
According to Gurovich and Beck, 2008, the single premolar this genus is based upon looks to be an unworn twin of a tooth they regard as belonging to Ferugliotherium (p.8). The ridge numbers are the same and both have large front roots (as do those of cimolod multis). Their Feruglio is about 15% larger. They think "Argentodites" perhaps belongs to the same genus or, at least, the same family. Therefore, it can't be a cimolod.
However, resolving this matter will require motre fossils. I've moved the entry onto this directory in accordance with their opinion but, as it hasn't been formally synonymized with anything as yet, I'm presently retaining it as a separate genus.
Meaning of name and stuff
The name combines the critter's country of origin and the Greek word hodites, a traveller. It's a metaphorical traveller as its relatives, at some stage, were held to have presumably wandered down south. The referral of this genus to Cimolodonta was tentative.

Species: Argentodites coloniensis Kielan-Jaworowska et al, 2007
Place: La Colonia Formation, Patagonia
Country: Argentina
Age: Upper Cretaceous
Remarks: The following is based upon my reading of Kielan-Jaworowska et al, 2007.
At first, it's tempting to think this multi was part of a North South faunal exchange system operating at some time during the latter parts of the Upper Cretaceous. Dinosaurs are known to have migrated in both directions, and marsupial and placental mammals may have come south at around the same sort of time. The diversity of South American marsups during the lower Paleocene would be in line with that. Given those circumstances, it wouldn't be all too surprising if a line of multis had also migrated then, and perhaps one did. However, if Argentodites resulted from such an event, then its North American ancestral lineage happens to be unknown. It has no obvious links with any fairly contemporary multi family there. Actually, there's even room for doubting its cimolodontan credentials. The referral is no more than tentative. Some "plagiaulacidan" tendencies are also evident. In short, you'd hope that a relatively recent immigrant would carry luggage indicative of its line of origin; its pedigree. While Argentodites clearly must have such useful pointers, quite where they're pointing to isn't obvious. Should its ancestors indeed have scampered off down south, they could have done so many millions of years prior to its birth. That would be one way of accounting for its eccentricities.
Multi-plication in South America
When I first saw the title of this paper, I rapidly reached a wrong conclusion. It's been known for ten years that some multi remains have been found in Los Alamitos Formation of Patagonia, and I blithely assumed that would be the subject here. Those fossils had been originally assigned to what was thought to be a gondwanatherian multi. However, it then became clear that gondwanatherians weren't multis, but some of the fossils concerned were. Those are the first known traces of Mutltidom from South America and, seemingly in contrast to Argentodites and the rest of known contemporaries, they apparently stem from a "plagiaulacidan" rather than a cimolodontan.
However, that's not the subject of this study. It concerns a p4 premolar from a bit further south in Patagonia; La Colonia Formation. And this tooth seems to stem from an eccentric cimolodontan. Two multis must've been padding around during the latter years of the Argentinean Upper Cretaceous, although the first one spotted remains nameless (p.257).
There's a point that's well worth bearing in mind for this pair. In both cases, the referrals to the two branches of Multidom are tentative, a ?plagi and a ?cimolo. The fossil record for mammals from the southern hemisphere contains vast blank spaces for much of the Mesozoic and, as evidence slowly emerges to fill them to some degree, the picture could change radically.
Not much to go on Very little is known of this 'Argentinean traveller'; just a single lower tooth. While hardly enough to form the basis of a photo-fit picture, it does provide some basis comparisons. It's a lower p4 premolar refined, as for all but the most basal of multis, for slicing up food. Seen from the side, the front wall of the tooth is prodigiously rounded, it bulges, and that's typical for cimolodontans. The length reaches a respectable 4.15 millimetres while the maximum width attains 1.35.
Orientations
Some speculation can be involved with isolated teeth when attempting to work out which was round the things went, but considerably less than might be thought. Teeth are functional members of working crews, and they have to be consistent with the rules dictated by nature as well as traits provided by ancestry. This can produce distinct characters for uppers and downers, fronts and backs and so on. Seen from the side, one end of this tooth bulges proudly outwards. The margin of the other end is much straighter. That bulge is consistent with the front of cimolodontan p4s, an affinity which is further supported by the organisation of enamel prisms. The authors term that "normal" (p.258). However, the straightness of the wear wall, and the relatively level height along the top of the crown happen to be more akin to some "plagiaulacidans". Cimolodontan models are generally shaped like the outline of an arch.
A more difficult challenge is provided by attempting to work out whether it's a left or right premolar. The buccal and lingual sides happen to be unusually similar; not the case for known northern cimolodontans with relatively large p4s. Tentatively, the authors opted for a left tooth with its buccal side somewhat less convex than the lingual one. What's presumably a cusp at the rear of the buccal side was less helpful than could've been hoped. There's a labial one as well, but what it was for isn't known.
Roots and ridges
The roots have gone but two were present (p.260). As generally for cimolodontans, but not for "plagiaulacidans", the front root was wider.
The roof of the tooth is adorned with eight serrations. Only the last of them shows much wear, and that suggests the former owner can't have lived until a grand old age. This premolar didn't do all that much work. Ridges run down from the serrations on both flanks, and the presumed buccal ones are more strongly developed than their lingual partners.
Holotype
MPEF 604, the only specimen presently known, is a near complete but rootless lower premolar in the growing collection of the Museo Paleontológico Edgidio Feruglio, Trelew. Its specific name honours La Colonia Formation.
Reference: Kielan-Jaworowska Z, Ortiz-Jaureguizar E, Vieytes C, Pascual R & Goin FJ (2007), First ?cimolodontan multituberculate mammal from South America, Acta Palaeontologica Polonica, 52(2), p.257-262.
Link:

Acta Palaeontologica Polonica 52(2)

http://www.app.pan.pl/acta52/app52-257.pdf

Kielan-Jaworowska et al, 2007 is presently freely accessible in pdf format.

Genus: Bharattherium Prasad GVR, Verma O, Sahni A, Krause DW, Khosla A & Pamar V, 2007

'Indian beast'

Family: Sudamericidae Scillato-Yané & Pascual, 1984

Remarks: Bharat is a Sanskrit word for 'India' and therion is Greek for 'beast'.
Two 'genera' of Indian gondwanatherians were established within a few months of each other and, in my unqualified opinion, they appear to be synonyms. On specimen has actually been independently referred to both of them. As neither team of authors were in a position to take the content of the other paper into account (neither was published prior to the effective deadlines), I think it fair to say nobody presently has any means of distinguishing the taxa from one another. The situation is rather unusual.
In any event, this study was definitely the first to be published and should, as looks highly probable, synonymity be officially declared at some time, Bharattherium will surely have seniority.

Species: Bharattherium bonapartei Prasad et al, 2007
Place: Kisalpuri, Madhya Pradesh, & Naskal, Andhra Pradesh, Deccan Traps
Country: India
Age: Maastrichtian, Upper Cretaceous
Remarks: The following is based upon my reading of Prasad et al, and thanks are due to Omkar for both forwarding the paper and further advice during a fairly confused day. I mean, there I was looking at a second description from another team, thinking: "I can't see any difference!" That was three days after having read this paper. Thanks also go to Greg Wilson for supplying that one and further welcome advice. And congratulations are awarded to Mikko Haaramo for noticing one of the specimens featured in both studies. Still, I think I've just about recovered now. These notes were mainly written prior to hearing about the second publication which I haven't yet read. It established Dakshina jederi.
Getting on with the story
Despite the most recent know gondwanatherian having died about forty million years ago in the then lushly forested nature reserve of Antarctica, these herbivorous mammals remain maddeningly shy of human company. This can't be put down as some kind of survival instinct. It's sheer bloody mindedness. Their known fossil record spans about thirty million years (which isn't actually all that long in comparison to some lineages I could mention) but, even at their most daringly revealing, they've not yet allowed more than a tantalizing glimpse of a partial lower jaw. Generally, rather than going all the way and exposing the glories of their full naked bodies, they draw the line at isolated teeth. Ladies, I assure you, your chances are better for getting a monk out of his habit than can be said for your prospects of seeing a gondwanatherian in its birthday suit.
Gondwanatherians
All the more reason for pleasure then, when a new gondwanatherian adds enrichment to the slowly unfolding story. These enigmatic mammals -whatever their wider affinities may have been- were, in some respects, millions of years ahead of their time; precocious exponents of then unprecedented methods of food exploitation. Quite what their food was is also unclear.
Excepting for Ferugliotherium, known gondwanatherians had high crowned teeth of a style known as hypsodont (p.17), with basically flat occluding molar surfaces excepting for corrugations provided by transverse lophs or other deliberate irregularities, such as in this case. These teeth were open rooted and the shape of the occlusal outline would, in some instances, provide suitable inspiration for aliens in a game of Space Invaders. The genus of Sudamerica is particularly tempting; a kind of melted H shape with additional smaller vertical bars on each side. Wear on the lophs even provides it with a belt of circles running horizontally along the middle, and they're crying out to be converted into spooky eyes or flashing lights on a spaceship.
(Note to myself: for the open-rootedness of Gondwanatherium and Sudamerica see page 301 of Gurovich, 2005).
There have been conflicting interpretations on gondwanatherian affiliations, but all have shared one essential component. They were wrong. One view saw them as perhaps related to placental xenarthrans, armadillos and the like. Another held them to be eccentric multituberculates. Multi affinities were actually correct for some fossils tentatively referred to the genus of Ferugliotherium, but not for the genuine generic fossils. Multidom was also kicked entirely out of consideration by a description of a gondwanatherian jaw in 1999. Although it lacked the tooth crowns, it had originally sported four molars, and that's twice the number allowed for multis. Finally, a form of consensus has emerged. Everybody now agrees that nobody has the foggiest idea of gondwanatherian affinities beyond them being small, plant-eating mammals of some kind or other. It's clear they were spread far and wide across the southern hemisphere landmasses for some time, but the known fossil record's thin, and in no fauna are they common.
Indian Upper Cretaceous mammals
At least three Maastrichtian aged localities in Central India are now contributing to knowledge about mammals from near the end of the Cretaceous (p.19). Two of these are close to one another and found about 70km west of Hyderabad: Naskal and Rangapur. A third, Kisalpuri, has been discovered more recently and radically extends the fossil front under assault by paleontologists -the intertrappean beds- around 700km further north. Presumably, the intervening stretch of land is presently hiding further localities. Research at Kisalpuri has so far yielded around fifty isolated teeth and bones of mammals, and most are from eutherians. One tooth, however, is gondwanatherian. A poorer specimen has previously turned up at Naskal and it belongs to the same species. Meanwhile, other researchers appear to have arrested some more.
Family affinities
'Indian beast' has been referred to the gondwanatherian family of Sudamericidae, and that might puzzle some present day geographers. However, sudamericids are now known to have inhabited a much wider area then the name might suggest. It's the smallest known member, with the holotype being around 65% the size of the Malagasy Lavanify, an approximate contemporary. (To undermine that, I've got to point out the type of Dakshina is larger than the molar described by these authors and, should it happen to be the corresponding tooth in the set, the size 'difference' between these two could disappear. Officially, however, D. is presently a separate genus.) The occlusal outline of the molar is termed "sub-rectangular" and, looking at the actual shape, that'd surprise some geometrists. It's rather rounded as sub-rectangles go. The crown features a wide 'v'-shaped island of dentine, as also known from Lavanify but, in this case, it contains a heart-shaped islet of enamel.
Speaking the tooth
The type molar is fairly described as high-crowned (p.20); 7.3mm high to be precise. That compares to a crown length of 2.66 and width of 2.0. Rather than arising as a straight column, the molar distinctly curves along its course, first outwards and then back inwards. Further ornamentation is provided by a similarly curving furrow running up the lingual side of the tooth, and that's the feature which serves to render the flat dentine crown into something approaching a v-shape when seen from above. Opposing the interruption imposed by the furrow is the inverted heart-shaped enamel islet. About two-thirds of the walls of the tooth are enamelled, while the concave area's free of the stuff excepting for at the very top.
The extreme height, the flattened roof and the full length furrow are all badges proudly worn by gondwanatherians, and particularly sudamericids (p.21). (Ferugliotherium molars are brachyodont rather than hypsodont.) As with Lavanify, and in contrast to Gondwanatherium and Sudamerica, no transverse lophs occur on the crown, the furrow continues down into the root, and their sizes are also smaller. The prismatic structure of the enamel, and the enamel free area also unite this pair. Some contrasts between them could, at least in part, reflect vagaries such as degrees of wear and differing tooth positions (p.22), factors which the small sample sizes don't presently allow much exploration of. In any case, as Madagascar and India are thought to have still been in contact with each other at approximately 90 million years ago, faunal similarities from late during the Cretaceous are to be expected (p.23).
Some followers of fashion
The hypsodont molar became fashionable among some placental mammals as well, but not until much later during the Oligocene and Miocene. They turned out to be good for processing the spreading craze of grass. It all seemed to account for hypsodonts well enough, but then gondwanatherians turned up at just about the wrong sort of time and place. This outbreak of the fashion started off something like forty million years or so too early. While grasses weren't assumed to have been entirely absent, no signs of the stuff suggested they could only have been relatively sparse during the latter Cretaceous. An alternative explanation sought to pin the blame on soil laden roots or perhaps abrasive tree bark. Some digging mammals specialize in such trades today, and these include semi-aquatic exponents; most famously beavers. Another extant group, muroids, oblige by also possessing hypsodont molars with comparable microstructure of enamel, and the use of enamel-free zones (p.24). A hypothesis suggests a similar sort of lifestyle. However, as this conjecture can presently only be based upon dental characters, it can't fairly be termed totally convincing. Rather, it's a reasonable option among possible scenarios.
Given the presence of the new gondwanatherian in Indian, it's hardly surprising to find the authors citing very recent evidence of grass pollen found in the intertrappean beds of the Lameta Formation; at least five taxa have been identified in what appears to be fossilized dino dung. Perhaps, then, this bout of hypsodonty was also fuelled by grass. Perhaps, but lots of fossil pollen has been studied from other parts of the world, and evidence from elsewhere is even rarer than gondwanatherians. In fairness, I don't happen to know about the state of pollen studies at other gondwanatherian localities.
The latest Indian elections
Given the social problems, diversity and sheer size of India, many sensible and learned outsiders assumed that democracy couldn't work under such circumstances. This reasonable seeming conclusion is something most Indians failed to take into account. Across the lands of the Republic can be found governments ranging from dour conservative to flamboyant communist. Movie stars, priests and ex-bandits can all win elections.
However, that wasn't quite what I had in mind with the above subtitle. My interest here is more on the mounting early returns as the votes are being tallied in the intertrappean constituencies of Naskal, Rangapur and Kisalpuri. Not enough is yet known for confident announcements of outright victory, but it's starting to look like a massive landslide victory for the popularly called Northern Eutherian Front. Not even Mahatma Gandhi would've gained over 90% support. For one thing, I'm sure he wouldn't have voted for himself. At Kisalpuri, for example, the eutherians have around fifty votes and the entire Gondwanan Congress Party have mustered just one. This goes far beyond a bit of eutherian immigration. It's more like an overwhelming campaign of conquer. It's completely out of kilter with the (probably somewhat earlier) South American faunas involving Gondwanatherium. There's not a squeak of even a therian in the Campanian of Patagonia.
Reportedly, there is a eutherian presence in a mid-Maastrichtian aged fauna in Bolivia, but I've presently no details on that beyond: Gayet, M., L. G. Marshall, T. Sempere, F. J. Meunier, H. Capetta, and J.-C. Rage. 2001. Middle Maastrichtian vertebrates (fishes, amphibians, dinosaurs and other reptiles, mammals) from Pajcha Pata (Bolivia). Biostratigraphic, palaeoecologic and palaeobiogeographic implications. Palaeogeography, Palaeoclimatology, Palaeoecology 169, p.39–68. It would be better to compare the Indian faunas with other nearer Gondwanan localaties, but there's a persisting shortage aside from Madagascar.
Holotype
VPL/JU/33 is a molar attending Jammu University. The specific name honours the emperor of Argentinean vertebrate paleontology, Dr Jose Bonaparte. He was the first to describe a gondwanatherian. To my shame, I've had to look up quite where Jammu is. The state and the city of that name are in northern India, with Kashmir further to the north.
Reference: Prasad et al (2007), A new Late Cretaceous gondwanatherian mammal from Central India, Proceedings of the Indian Natu. Sci. Acad, 73(1), p.17-24.

Genus: Ferugliotherium Bonaparte JF, 1986

Family: Ferugliotheriidae Bonaparte, 1986

Aka: Vucetichia Bonaparte, 1990

Remarks: Gondwanatheria is now generally not regarded as part of Multituberculata but, according to Kielan-Jaworowska & Hurum, 2001, (p.411): "a few specimens described as ?Ferugliotherium," are from multis. "These poorly known specimens (not discussed herein) demonstrate that a branch of multituberculates apparently lived during the Late Cretaceous in South America." As gondwanatherians aren't multis, these remains would clearly not belong to this genus or grouping unless the genus happens to be a multi of some strange kind!
A coming confusion?
A pdf of a since published study turned up in October 2008 (many thanks, Javier). Psychic abilities lead me to predict it will include the establishment of a new genus of ferugliotheriid. Further, that family will be tentatively assigned to (note the question mark) ?Multituberculata. Should that be justified by more time and discoveries, then the upshot could be that feruglios aren't gondwanatherians. An alternative would be that they're feruglio gondwan multis.

Species: Ferugliotherium windhauseni Bonaparte JF, 1986
Aka: Vucetichia gracilis Bonaparte, 1990
Place: Los Alamitos Formation, Patagonia
Country: Argentina
Age: Upper Cretaceous
Remarks: The genus was established on the basis of a lower molar, (m2), and provisionally referred to as a multi. Further specimens were discovered, (m3 and M1), and other teeth were tentatively assigned; incisors and premolars. The upper premolars may be genuinely multituberculate, which explains their multi characteristics.
Holotype
The holotype, MACN-RN 20, is an inmate of the Museo de Ciencias Naturales, Buenos Aries. It's a lower molar (m2) with both a length and a width of 1.7mm, (Kielan-Jaworowska & Bonaparte, 1996). A cellmate is the holotype of Vucetichia, (MACN-RN 174). These were synonymized by Krause in 1993, (Krause & Bonaparte 1993, p.9379.)

In the view of Gurovich & Beck, 2008
These two authors concluded that multituberculate like fossils, which other autors had thrown out of the genus, most probably do belong within it (p.3). They proceed with the assumption that they're correct and provide justification. These returnees include a fragmentary dentary with a p4 premolar and several upper premolars. Known lower molariforms for this genus, presumably genuine molars, do conveniently group into two sorts, and that suggests a count of two per side. Previous authors had assumed four per side based mainly on the gondwan molariform count known from Sudamerica (p.4). Given the paucity of specimens available, G & B don't go as far as specifying even tentative numbers for cheek teeth types.
Sizes
G & B also provide measurements of teeth they assign to this genus (p.3). I'll just mention lengths.
Uppers: Premolar (P1 or P2), 2.05mm; MF (molariform) 1, 2.50mm.
Lowers: mf1 (2 specimens), 2.10-2.20mm; mf2, 1,70mm.
Vucetichia gracilis RIP
The following is based on my reading of Bonaparte, 1990. This paper included the establishment of the genus subsequently transferred to Ferugliotherium. At the time, F. was still considered to be a multi while V. clearly wasn't. (Thanks are due to Rob B of New South Wales for supplying the paper.)
Meaning
'For Vucetich'
The generic name honoured Gulomar M Vucetich, a researcher with special interests in Argentinean rodents and notungulates of the Tertiary.
Bonaparte assigned to upper molars to his new proposed genus (p.77), and both had suffered from wear. Working out front from back couldn't be done with certainty, but the higher side of the crown is most likely labial. When seen from the occlusal perspective the outline is roughly rectangular. The tooth has a series of lophs (broad ridges) running across the crown from side to side. Look from either side, front or back the top of the tooth is about as flat as an old-fashioned washboard.
Grinders
These lophs form three figures, and each is defined by a surrounding of enamel. The largest is a wobbly V-shape and presumably at the back. The other two are simpler; erratic bars, with the middle figure being the smallest (p.78). There's plenty of wear, and this was most likely caused by crushing and grinding hard foodstuffs, rather than resulting from contact with the lower dentition. Both specimens had four roots; a pair aft and another couple to the rear.
These enamel bound figures are also found on the molars of Gondwanatherium, but Bonaparte pointed to several structural differences. Subsequently, when it was known that most material referred to Ferugliotherium was also gondwanatherian, this proposed genus turned out to be superfluous.
Holotype II
The type fossil of Vucetichia is MACN-RA 174, an upper molar in the collection of Museo Argentino de Ciencias Naturales, Buenos Aries. The specific name means 'slender', 'delicate' and refers to the small size. The length is about two millimetres, and this was small compared with recognized gondwanatherians at the time.

References: Bonaparte (1986), Sobre Mesungulatum houssayi y nuevos mamíferos Cretácicos de Patagonia, Argentina. (Spanish, with English summary). Actas Congr. Argent. Paleontol. Bioestratigr. 4, p.48-61.

Bonaparte (1990), New Late Cretaceous mammals from the Los Alamitos Formation, northern Patagonia. National Geographic Research, 6, p.63-93.

Krause DW (1993), Vucetichia (Gondwanatheria) is a junior synonym of Ferugliotherium (Multituberculata). Journal of Paleontology 67, p.321-324.

Species: ?Ferugliotherium windhauseni
Place: Los Alamitos Formation, Patagonia
Country: Argentina
Age: Upper Cretaceous
Remarks: Update, November 2008
Other authors have rebelled! Gurovich & Beck, 2008 have backed the original identification and transferred all the specimens back to Ferguliotherium. That should be held in mind while reading this episode.
The following is based upon my reading of Kielan-Jaworowska & Bonaparte, 1996, and it specifically concerns the material which was held to be from a multituberculate. (Thanks are due to Steve for sending the paper.)
The star of this paper is a fragment of dentary called MACN-RN 975. It's not very large but seemingly points to something significant; southern multis. As it was found in the same fossil beds as isolated teeth of the compatibly sized Ferugliotherium, it was questionably assigned to the same genus and species, (p.1). However, this fossil can't be placed in that taxon or even Gondwanatheria. The one preserved tooth on the piece of jaw is a long premolar (p4) with eight ridges and no cusp(s) on the buccal side. It's from a relatively derived, gondwanan multituberculate.
The find
The fossil was found in 1991, (p.2). As well as the heavily worn tooth, the fragment also contains an alveolus for a relatively large incisor. A short diastema separates both teeth, but fracturing makes its length uncertain, (p.4). As the hole for the incisor continues along the whole length of the preserved fossil, the i1 must have been robust, (p.5). An i1 tentatively referred to Ferugliotherium would fit it well.
The premolar is 4.8 millimetres long, and that's a healthy size for Cretaceous multis (should it indeed come from a multi). Its external face shows eight faint ridges with at least that number on the lingual side as well. The lack of a buccal cusp is a relatively derived characteristic. This tooth has thin enamel, but that's probably due to processes of preservation rather than the natural condition. Both roots are strong, with the front one being larger than its rear colleague. As the premolar is relatively long and complex, (even though wear means its height can't be precisely determined), it isn't reminiscent of any multi premolars other than a p4. No known further anterior teeth have anywhere near eight ridges. Thus, despite the lack of an alveolus for a p3, a p4 is what it appears to be, (p.6). Presumably, this lineage had only a single premolar, as is the case for some later northerners.
Affinities
The presence of only one premolar excludes this fossil from being a 'plagiaulacidan', as those animals had three to four of these teeth per side, (p.8). However, (and bearing in mind this was tentatively associated with remains of a different genus), affinities with the more advanced cimolodontans are not apparent. The find seems likely to represent an otherwise unknown southern radiation of multis, which could conceivably have arisen prior to the emergence of northern cimolodontans. The present lack of anything reasonably comparable inevitably results in little clarity. Clear is that the career of Multidom wasn't exclusively Laurasian.
Repetition for clarity: this fossil doesn't belong in Gondwanatheria.
Reference: Kielan-Jaworowska & Bonaparte (1996), Partial dentary of a multituberculate mammal from the Late Cretaceous of Argentina and its taxonomic implications, Revista del Museo Argentino de Ciencias Naturales 'Bernardino Rivadavia', New Series, 145, p.1-9.

Genus: Gondwanatherium Bonaparte JF, 1986

'Gondwana beast'

Family: Sudamericidae Scillato-Yané & Pascual, 1984

Species: Gondwanatherium patagonicum Bonaparte JF, 1986
Place: Los Alamitos Formation, Patagonia
Country: Argentina
Age: Upper Cretaceous
Remarks: Though earlier than Sudamerica, Gondwantherium is held to be more derived. This may sound surprising, but there's no reason why ancestral lines might not outlive descended ones.
The genus was described on the basis of isolated, hypsodont, rodent-like molars. Originally, it was referred to the monotypic family of Gondwanatheriidae. Close similarities with Sudamerica were recognized, but the ridges of the enamel had a different pattern. Subsequently, further finds of Sudamerica teeth showed varying amounts of wear. This narrowed the differentiations and the genera were assigned to the same family by Bonaparte, (Krause & Bonaparte 1993, p.9379).
Reference: Bonaparte (1986), A new and unusual Late Cretaceous mammal from Patagonia, Journal of Vertebrate Paleontology, 6, p.264-270.

The affinities of Gondwanatheria

The following is largely based upon my reading of Krause & Bonaparte, 1993.

In this study, gondwanatherians were referred to as members of a Superfamily, (Gondwantherioidea), within Multituberculata. They were held to be closer to 'Plagiaulacida', (Plagiaulacoidea in the terminology employed), than the further derived cimolodontans. This view didn't stand up in the light of subsequent research, but the paper is still informative. Before making a case for affinities with multis, the authors gave some details on apparent similarities with edentates, (p.9379).
Edentates?
Edentates are undoubtedly members of Placentalia, which is more exclusive than Eutheria. They seem to be more basal than any other living placental mammals, which suggests their lineage evolved relatively early. When present, their molars are also hypsodont. Given that gondwanatherians are first known from a time and place which appears to be where ancestors for edentates might be expected, (the Upper Cretaceous of South America), it's tempting to look for connections. However, the hypsodont molars of edentates are the result of lengthened roots. With Gondwanatherium and Sudamerica, it's the crowns which are elongated. The overall effect is similar, but the architecture is significantly different.
Something else?
Another view suggested those two genera could have been derived from an early monotreme such as Steropodon, or perhaps from Ferugliotherium, (Van Vaalen in 1988). The latter possibility prevailed. That genus in now generally interpreted as the most basal known gondwanatherian. However, that still leaves affinities with other mammals unclear.
Multituberculates?
This study embraced placement within Multidom. Ferugliotherium possesses similarities with multis. The other two then known genera are more derived members of the same lineage. The case was perhaps artificially strengthened by the referral of genuine multi material to Ferugliotherium, but that became clear later.
Molars
Two multi-like characteristics are cited for the molars of Ferugliotherium: the presence of more than one row of cusps, and the evident direction of movement of the dentary during the power bite, (palinal -backwards directed). Two similarities were also highlighted for the premolars: the front uppers have conical cusps, each equipped with ridges, and the lowers have serrations. (However, cross-checking with the reference given by Kielan-Jaworowska & Hurum, 2001 (p.411) suggests the referred upper premolars are multi, not gondwanatherian.)
At the time, the three known genera of gondwanatherians were all represented by isolated teeth which included lower molars for each, (m1s and m2s). The text and illustrations on page 9380 point to convincing similarities between them, especially when evidence for different stages of wear is taken into account. In occlusal outline, the m1 is broadly rectangular, while the m2 is more squarish. For the successively further derived Gondwanatherium and Sudamerica, compressions on the lingual side are more apparent so that the shapes are less regular. The wear patterns correspond well. The molars of Ferugliotherium differ in being not hypsodont and significantly smaller.
Incisors
The authors also discuss some fragments of incisors which had been tentatively referred to Ferugliotherium and Gondwanatherium. These have limited bands of enamel: "Among Mesozoic mammals no known group other than multituberculates have a restricted enamel on the incisors." That was true in 1993. However, Kulbeckia has since begged to differ on behalf of zalambdalestid eutherians.
Jawing
Dental details can give clues concerning the mechanics of jaws. In Ferugliotherium, the wear facets run from back to front, which indicates that the bite power was maximized during posterior movement. Furthermore, the cutting edges of the lower molars are positioned at the back, with the opposite arrangement applying for the uppers. "Krause... argued that this pattern is characteristic of multituberculates and is unique among mammals", (p.9381). Should, as seems to be the case, gondwanatherians not be multis, then this characteristic logically ceases to be unique.
Back to the drawing board
Common ancestry for the gondwantherians in the study appears to be well established. If one is a multi, then so are the others. As the similarities are closer to Upper Jurassici 'plagiaulacidans' rather than cimolodontans, the key would appear to be to demonstrate secure connections between Ferugliotherium and some 'plagiaulacidan' or other. I suppose the presence of transverse ridges between the cusps on the molars might be helpful, except nothing like that is presently known from any 'plagiaulacidan'. The only multi with such a characteristic is Essonodon of Cimolodonta, (Kielan-Jaworowska & Hurum 2001, p.408). Subsequent research suggests secure connections weren't convincingly demonstrated. (Krause et al, 2003 contains gondwanatherians rather than gondwanatherioideans, and doesn't treat them as multis.) Given that, a sentence on page 9382 caught my eye: "The origin of Edentata remains enigmatic." True, but the same seems to apply for gondwanatherians.
Back from the drawing board?
15 years after Krause & Bonaparte came Gurovich & Beck, 2008. They recognized the integrity of Gondwanatheria with its two known families, Ferugliotheriidae and Sudamericidae. They also found support for placing gondwans within or near Multituberculata. Whether other researchers will be swayed by their conclusions remains to be seen.

Genus: Lavanify Krause DW, Prasad GVR, von Koenigswald W, Sahni A & Grine FE, 1997

'long tooth'

Family: Sudamericidae Scillato-Yané & Pascual, 1984

Species: Lavanify miolaka Krause DW et al, 1997
Place: Maevarano Formation, Mahajanga Basin
Country: Madagascar
Age: Maastrichtian, Upper Cretaceous
Remarks: This genus is based a tooth, which was found whilst quarrying for a dinosaur skull, (Majungatholus). A second specimen was also referred.
"The teeth of Lavanify differ from those of the only previously known sudamericid genera Gondwanatherium and Sudamerica in possessing prominent and continuous inter(-)row sheets of interprismatic matrix in dental enamel and at least one cheek-tooth position that has a single, V-shaped dentine island and lacks enamel on one side of the crown. Lavanify further differs from Gondwanatherium in having cheek-teeth with vertical furrows that extend to the base of the crown and onto the root", (Krause et al, 1997, p.504).
Reference: Krause et al (1997), Cosmopolitanism among gondwanan Late Cretaceous mammals. Nature 390, p.504-507.
Link:

Dino Land Paleontology Interviews, Dr Cathy Forster

http://www.geocities.com/CapeCanaveral/Galaxy/8152/cathy.html

Lavanify gets a brief mention in this interview. I particularly enjoyed the last line, her favourite moment in paleontology.

Genus: Sudamerica Scillato-Yané GJ & Pascual R, 1984

'South America'

Family: Sudamericidae

Aka: Sudamericana

Species: Sudamerica ameghinoi Scillato-Yané GJ & Pascual R, 1984
Place: Punta Peligro, Patagonia
Country: Argentina
Age: lower Paleocene
Remarks: In common with Gondwanatherium, this genus had high-crowned teeth, which are very useful for eaters of grass. As there's still little evidence of grass in South America until later, they may also have been effective for other food stuffs, (with thanks to Martin Jehle).
Update: Research into Upper Cretaceous dino dung in 2005, suggests grasses are now known from the Upper Cretaceous of India, now also known Gondwnatherian territory. Caroline Strömberg of the Swedish Museum of Natural History and colleagues came across some pollen in samples from Central India.
Molariforms
"Pascual et. al. (1999) recently demonstrated the presence of four cheek-teeth, all of them molariform, in the dentary of the sudamericid Sudamerica, which is the only definitively identified gondwanatherian represented by a substantial jaw fragment", (Krause et al 2003, p.326). These authors also mention that the incisor of this critter had a root which extended underneath the entire length of the tooth row.
However...
Gurovich & Beck, 2008 fully accept that dentary belongs to this genus and does have four molariform teeth. Furthermore (p.4), known isolated molariforms (both uppers and lowers) neatly divide into eight morphological groups. Both facts indicate that the molariform tooth count per adult jaw half was four. What these authors doubt are the strict molar identities of the first two molariforms. They suggest these could actually be molarized premolars.
A bit more on the molariforms
As mentioned by Kemp 2005 (p152-153), the presence of four molars and no premolar capable of shearing, aren't multituberculates characteristics. The molars have ridges running across the tops of them, and this effect is produced by wear. They probable worked well on tough plants of some kind. Of course, should Gurovich & Beck, 2008 be correct, then only two of the four teeth in question are strictly speaking molars.

The genus was originally based upon one hypsodont, rodent-like molar, (Krause & Bonaparte 1993, p.9379).
Whoops!
Martin Jehle has also been kind enough to point out, that I had the name of the family wrong. I'd called it "Sudamericanidae". This error has also been made by others, (which is no excuse!)

Reference: Scillato-Yané & Pascual (1984), Un peculiar Paratheria, Edentata (Mammalia) del Paleoceno de Patagonia. Primeras Jornadas Argentinas de Paleonotologia de Vertebrados, Resumenes, p.15.
Links:

Acta Palaeontologica Polonica, 44(3), p.263-300

http://app.pan.pl/acta44-3.htm

Wighart v. Koenigswald, Francisco Goin, and Rosendo Pascual: Hypsodonty and enamel microstructure in the Paleocene gondwanatherian mammal Sudamerica ameghinoi.

Martin Jehle, Multituberculates: Heyday of the longest lived mammalian order

http://www.paleocene-mammals.de/multis.htm

The last section of this overview of multis concerns Gondwanatheria.


Genus: Trapalcotherium Rougier et Al, 2009

'Trapalco beast'

Family: Ferugliotheriidae Bonaparte, 1986
Remarks: The fossil locality is situated in a basin called Rajo Trapalco, thus the name. Oh, and the animal was quite a beast.

Species: Trapalcotherium matuastensis Rougier et Al, 2009
Place: Allen Formation, Patagonia
Country: Argentina
Age: Campanian-Maastrichtian, Upper Cretaceous
Remarks: The following is based upon my reading of Rougier et al, 2008. That's a prepublication version of the description which appeared in print early in 2009, and the page numbering also differs. Thanks are due to the supplier.
'Trapalco beast' is presently known from but a single tooth, but it does add some more meat to an already somewhat unclear stew. The authors uncontroversially enough refer it to the family of Ferugliotheriidae, and I envisage everybody will be happy enough with that. However, they then refer the family (with a question mark) to ?Multituberculata. What they don't discuss, presumably due to irrelevance in terms of their study, is the relationship of feruglios and sudamericids; both families of Gondwanatherians according to many. They make no comment about whether feruglios are or aren't also gondwans; a possibility should sudams also be ?multis. Another matte they look at is a purported ?cimolodontan multi (again, note the question mark) named Argentoides. They think it more likely to be material from a feruglio, a conclusion they share with Gurovich & Beck, 2008.
Distinctions
The tooth is a lower m1 molar with about half the buccal edge absent. The front of the occlusal surface is said to have "a more compressed" "anterior triangle" than Ferugliotherium. I think I know what that's referring to but I'm not certain. (My uncertainty may result from my lack of familiarity with papers on feruglios.) In any case, this genus differs by having this triangular feature in a more compressed form, and that applies whether my recognition is right or wrong! More clear to me is the presence of a crest curving along the front of the crown. Bases of cusps for this genus are less well developed, in that they've become more greatly merged into the crests running across the crown. Also, in this case, the buccal and lingual cusp rows differ more in terms of height.
In short, there are a number of distinctions from Ferugliotherium, thus a new genus was established.
Roughly the shape of things
The tooth is interpreted as a lower left molar. Reasons for this include the presence of only two cusp rows, the tooth being longer than wide, and the transversely running ridges as displayed by feruglios and sudamericids. One side has more cusps than the other and, as that's the buccal side, this identifies it as being a left molar. Whilst the rear of the buccal side is missing, the remains of transverse ridges suggest two further cusps were probably located there.
The crown outline is roughly rectangular in shape with the front, as mentioned, being curved somewhat like a bay window. Lingual cusps number three whereas there were probably five buccal ones, although four also can't be fully ruled out. In any event, buccal cusps were more numerous. Ridges link the cusps of both rows and furrows occur between those. There's a groove running between the cusp rows, but this gets blocked along its course by the ridges. The authors inform me: "The pattern of cusps, crests and furrows determines a mesial triangle pointing lingually and followed distally by two incompletely developed transverse ridges." As they go on to say, there's a triangular arrangement of the first two buccal and first lingual cusps (p.11), and I suppose you can base a triangle on those points well enough.
In short summary, this is a multituberculate-like lower molar in the sense that there are two rows of cusps, albeit cusps that can be difficult to see due to indistinctiveness, and a groove running between the rows, albeit somewhat eccentrically. Atypical for multis would be the lophs running across crown blocking that groove at various junctures.
Size
The maximum length is 2.48mm and the greatest width manages 2.07. That's proportionately shorter and wider than Ferugliotherium windhauseni. However, I don't happen to know how may lower molars are known for that genus, and whether anything has been discovered with regards to possible distinctions between molar positions. (It's possible I've got some relevant information somewhere, so I'd better remember to check.) Such matters could be of relevance to comparisons.
Sudamericids
Sudamericids aren't entirely ignored by this study. They're accused of being notable absentees in La Colonia Formation. While, given the minimalist sample size of mammal fossils from the Allen Formation, a lack of specimens from there has little significance as yet, over 300 mammalian specimens have been obtained from La Colonia. That lack of sudams is less likely to be merely some sampling blip. Preservation bias could be an explanation or, perhaps, sudams didn't find the paleo-environment there to their tastes.
Allen's vertebrates
Although a new source of mammals, this locality had already yielded a variety of other vertebrates; small and large sauropods, big bullying theropods, birds, hadrosaurs and ankylosaurs, snakes, sphenodont reptiles, turtles, frogs and several types of fish. This cast list also reflects themes played by the house orchestras of other Patagonian localities, which is none too surprising. Up north in Bolivia, the probably somewhat later El Molino Formation is also moving into the business of Mesozoic mammals, albeit presently unpublished ones. That fauna may include invading therians of northern ancestry. Unfortunately, that "may" must suffice until more and better specimens are available from there.
As much of South America was under water during the Upper Cretaceous, it would be interesting to know how more northerly faunas compared with Patagonian ones. Such comparisons would require new discoveries and then publications.
Holotype
MML-Pv 16 is a lower molar (m1) in the collection of the Museum Municipal de Lamarque. The specific name pays homage to the important paleontological contributions selflessly made by the amateur researcher, Puesto El Matuasto. In professional life Matuasto performs valuable service as a shed. It's a herder's shed owned by the Cabazza family. The herd should also be thanked for agreeing to allow paleontologists to make use of their facilities, and this with a complete disregard for any offensive odours this might have resulted in.
Reference: Rougier GW, Chornogubsky L, Casadio S, Arango NP & Giallombardo A (2009), Mammals from the Allen Formation, Late Cretaceous, Argentina, Cretaceous Research, 30(1), p.223-238.

Other reports:

India

Some unnamed material has been identified from the Deccan Traps.

Link:

NSF News, 3.12.1997

http://www.nsf.gov/od/lpa/news/press/pr9771.htm

Strange South American Fossil Mammals Found in Madagascar and India.

Seymour Island, Antarctica

Antarctica apparently has more to offer than just marsupials. "The occurrence on Seymour Island of sudamericids, that had become extinct in South America in the Paleocene, also indicates that isolation may have allowed extended survival of this Gondwanan group in the Eocene of Antarctica and the factors that caused their extinction did not affect this continent," (Reguero et al 2002, p.189).
From the same paper, (p.203): "For the sudamericids, Koenigswald et al. (1999) inferred a semi- aquatic and perhaps a burrowing way of life, similar to that of living beavers. Regarding this, the presence of two Antarctic taxa at Seymour Island (Goin, personal communication, 2000) suggests an important paleoecological constraint related to dietary preference of this group." This taxon seems to be breeding.
The Antarctic Peninsula of the "late Early to latest Eocene", (La Meseta Formation -about 40 million years ago), seems to have been a lively place. The fossils and the geological conditions of the site suggest a near-by forest populated by a diverse fauna, which had many similarities with the slightly earlier residents of Patagonia; small, arboreal, fruit and insect eating possums, 10kg sloths, middle to largish grazers (sparnotheriodontids and Trigonostylops), falcons, ratites (rheas and the like, aka big, flightless birds) and penguins. At the end of the Eocene the climate seems to have taken a turn for the worse...
With thanks for Martin Jehle for the notification and interesting abstract and David Marjanovic for digging out and supplying the paper.

Link:

Marsupial Photos

http://www.trilobyte.ucr.edu/mow/ro.html

In case you thought I'd made a typing error, these photos show remains of marsupials from Antarctica.

Specimen: NMT 02067 Krause DW, Gottfried MD, O'Connor PM & Roberts EM, 2003
Place: "Red Sandstone Group", Mbeya District
Country: Tanzania
Age: Cretaceous
Remarks: The following is based upon my reading of Krause et al, 2003.
The fist to state about this specimen is that the referral to Sudamericidae isn't simply tentative. It's very tentative, (p.321). In keeping with this spirit, the age of these particular sandstones is also poorly understood, (p.322).
The fossil in question is a partial lower jaw. It comes from deposits which probably represent the work of an ancient river system, (fluvial). The locality evidently took exception to having once been described as not rich in fossils, as it has since decided to yield an array of vertebrate remains: bony fish, crocs and the like, sauropod and theropod dinosaurs, a bit of bird leg and this jaw. Dino fossils have been found above, below and alongside the level where 02067 was found, so a pre-Paleocene age is a good bet. Taken as a whole, the faunal assemblage suggests Cretaceous, (p.323).
A jaw with teeth
This mammal fossil is a worn, left dentary but: "It lacks sufficient morphological information to allow the diagnosis of a new taxon." What can be said is that's it's short and deep and was home to one incisor and probably five single-rooted cheek teeth. None of these are now in Colgate-clean condition. The remains of the incisor show it was a large, forwardy-pointing tooth, with a root set at an angle of roughly 55° from the horizontal axis. This extends all the way back to below the front of the third cheek tooth.
Behind the incisor is a gap of a couple of millimetres or so, (a diastema). This is followed by remains of two cheek-teeth roots, the first of which suggests a slightly smaller chopper than the latter. The rest of the series are somewhat better preserved. Number three is clearly large and tall, and curves inwards to some extent. It's also well anchored, in that the root goes down to a depth of about 75% of the dentary, (p.324). The fourth in the sequence is similar in size to numbers 1 and 2, whilst the final member is the smallest tooth present. # These also have strong roots and all three hindmost teeth are "truly hypsodont", (p.325), which prompts me to search for a reliable translation; learning through googling.
('Hypsodont' literally translates as 'high tooth' -with thanks to Dental Anatomy of Horses. This page includes some interesting images, and the text is far from baffling; a fine bit of demystification from Colorado State University. And now back to p.325...)
Unclear affinities
Part the difficulty with this specimen is that it displays superficial similarities to a host or critters, and more diagnostic bits aren't preserved. It may be from an early representative of various taxa or evidence of a previously unknown one. However, amongst Mesozoic mammals, it shares the following series of characteristics with taeniolabidoidoid and djadochtatheroidean multituberculates and gondwanatherians: "body short and deep, unfused mandibular symphysis, distinct diastema and coronoid process originating far anteriorly". Those multi group are currently only known from the northern hemisphere, none are known to have hypsodont cheek teeth or more than two molariform teeth in each half of the lower jaw.
Some frustration
"We tentatively conclude that NMT 02067 is not a multituberculate and is most parsimoniously referred to the Gondwanatheria, and more specifically to the Sudamericidae (i.e., all gondwanatherians save Ferugliotherium)."
Whilst tantalizing, this find is also somewhat frustrating. It points to various interesting possibilities, but doesn't provide as much information as would be wished. However, any locality which contains one mammalian fossil very probably provides a refuge for more. Additional specimens would be most welcome. The jaw now lives in the collection of the National Museums of Tanzania, Dar Es Salam. It was discovered by Yasemin Tulu, (p.327).
Reference: Krause et al (2003), A Cretaceous mammal from Tanzania, Acta Palaeontologica Polonica, 48 (3), p.321-330.
Link:

Acta Palaeontologica Polonica, 48(3), p.321-330

http://app.pan.pl/acta48/app48-321.pdf

The full paper is freely available on-line.

Bonus Link:

Alvaro Mones home page

http://www.geocities.com/Athens/Forum/1384/principal.htm

The on-line paleontologist and father of Gondwanatheria.

Help:

Should anybody have any further information, I'd be pleased to hear of it.

Regarding references and Bibliography:
I haven't and can't verify the references, so beware. 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.

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Trevor Dykes, April 2002. Latest up-date: 25.3.2009
Ktdykes@arcor.de

With further thanks due to:

Martin Jehle, Genera and species of Paleocene mammals - Part 1

http://www.paleocene-mammals.de/pal1.htm

A useful source containing much basic info on names and locations.

BIOSIS, The Index to Organism Names

http://www.biosis.org.uk/triton/indexfm.htm

The Society of Vertebrate Paleontology BFV Online, (John Damuth)

http://www.bfvol.org/http://www.bfvol.org/

Bibliography:
Bonaparte JF (1990), New Late Cretaceous mammals from the Los Alamitos Formation, northern Patagonia, National Geographic Research, 6(1), p.63-93.
Gurovich Y (2005), Bio-Evolutionary aspects of Mesozoic Mammals: Description, phylogenetic relationships and evolution of the Gondwanatheria, (Late Cretaceous and Paleocene of Gondwana), Thesis, Museo Argentino de Ciencias Naturales 'Bernardino Rivadavia', Buenos Aires, 2005, pp.621.
Gurovich Y & Beck R(2008), The phylogenetic affinities of the enigmatic mammalian clade Gondwanatheria, Journal of mammalian evolution, prepublication copy (24 pages).
Kemp TS (2005), The Origin and Evolution of Mammals, Oxford University Press, pp.331.
Kielan-Jaworowska & Bonaparte (1996), Partial dentary of a multituberculate mammal from the Late Cretaceous of Argentina and its taxonomic implications, Revista del Museo Argentino de Ciencias Naturales 'Bernardino Rivadavia', New Series, 145, p.1-9.
Kielan-Jaworowska Z & Hurum JH (2001), Phylogeny and systematics of multituberculate mammals, Palaeontology, Vol 44 (3), p.389-429.
Kielan-Jaworowska Z, Ortiz-Jaureguizar E, Vieytes C, Pascual R & Goin FJ (2007), First ?cimolodontan multituberculate mammal from South America, Acta Palaeontologica Polonica, 52(2), p.257-262.
Krause DW & Bonaparte JF (1993), Superfamily Gondwanaterioidea: A previously unrecognized radiation of multituberculate mammals in South America, Proc. Natl. Acad. Sci. USA, 90, p.9379-9383.
Krause DW, Gottfried MD, O'Connor PM & Roberts EM (2003), A Cretaceous mammal from Tanzania, Acta Palaeontologica Polonica, 48 (3), p.321-330.
McKenna MC & Bell SK, (1997), Classification of Mammals Above the Species Level. Columbia University Press.
Prasad GVR, Verma O, Sahni A, Krause AW, Khosla A & Parmar V (2007), A new Late Cretaceous gondwanatherian mammal from Central India, Proceedings of the Indian Natu. Sci. Acad, 73(1), p.17-24.
Reguero MA, Sergio AM & Santillana SN (2002), Antarctic Peninsula and South America (Patagonia) Paleogene terrestrial faunas and environments: Biogeographic relationships. Palaeogeography- Palaeoclimatology-Palaeoecology, 179, p.189-210.
Rougier GW, Chornogubsky L, Casadio S, Arango NP & Giallombardo A (2008), Mammals from the Allen Formation, Late Cretaceous, Argentina, Cretaceous Research, prepublication copy (16 pages), officially published in 2009.