Tetrapod Zoology

The most terrestrial of bats

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I’ve said it before and I’ll say it again: we keep coming back to the subject of flightless bats. Besides fictional future predators and night stalkers, there never have been any flightless bats so far as we know.

Whenever this subject is discussed however, we have to pay appropriate homage to the most strongly terrestrial bats that we know of: the vampires, and the short-tailed bats (or mystacinids). Vampires were done to death here earlier on in the year (go here)… now, at last, it’s the turn of the mystacinids.

New Zealand has – or had – more than its fair share of neat tetrapods, some of which I’ll get round to covering on the blog some day. Moa, kiwis, giant geckos, tuataras, weird rails, big flightless geese, and so on. Of course we always thought that – besides a few cryptids (notably the amphibious or aquatic waitoreke) – Cenozoic New Zealand was devoid of non-volant mammals. We now know that a relict lineage of mammals phylogenetically close to multituberculates (Worthy et al. 2006) inhabited New Zealand until the Miocene. Among the other mammals, there were bats: two of which, the mystacinids, represent a group that was long thought unique to New Zealand. That’s no longer the case, as several species of mystacinid are now known from the Oligocene and Miocene of Australia (Hand et al. 1998, 2005): in fact, based on current evidence it seems that mystacinids originated in Australia before one lineage later spread to New Zealand [image above from here].

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Uniquely specialized for fast and agile movement on the ground, mystacinids are primarily insectivorous but have recently been shown to be nectar and pollen feeders as well. They possess particularly tough and leathery wing membranes which they’re able to fold up tighter than any other kind of bat. It used to be thought that it was the absence of small native land mammals (such as mustelids, marsupials and mice) that uniquely enabled mystacinids to exploit the role of terrestrial and arboreal forager. However, read on. Mystacinids are also uniquely adapted for crevice-dwelling and have short, erect, velvety fur that helps them to squeeze through tunnels in wood and soil (Kunz 1982). Tomes (1857) compared it to that of shrews and Daniel (1979) noted that some molossid bats that also hide or shelter in small crevices have similar fur. Mystacinids are active tunnelers in soft tree bark and chew out roosting cavities and tunnels – consequently their teeth are often heavily worn.

Mystacinids have robust hind limbs and feet (see image above: from here): the latter are broad and short with softly padded, deeply wrinkled soles and grooves across the bottoms of the toes, and across the lower surface of the ankles and legs. These wrinkles and grooves probably assist grip in climbing, as do the well developed, needle-pointed claws on the feet and the thumb. The thumb claw is especially remarkable as it has a small, subsidiary talon at its base. Daniel (1979) stated that the foot claws have these subsidiary basal denticles as well.

The most remarkable terrestrial adaptation of Mystacina is the fact that it can tightly curl up its wing membranes beneath the thick leathery membrane that lines the flanks and underside of the thigh, thereby protecting the delicate part of the wings from injury. Its arms now become two straight forelegs that can be used in scurrying on the ground and on trees: the resulting terrestrial agility has often been described as ‘mouse-like’. The second wing digit is notably reduced, consisting only of the metacarpal and a single tiny phalanx; a reduction that may have evolved to reduce the size and bulk of the wing. The third digit has three phalanges and a cartilaginous tip; in the fifth digit, the cartilage tip extends beyond the edge of the patagium. The propatagium is reduced and the plagiopatagium inserts at the level of the ankle.

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Mystacinids are reported to be slow and less agile in flight than other microbats and they have been described as having proportionally short wings with rounded tips (Norberg & Rayner 1987). This wing morphology would be well suited for the slow-flying, gleaning habit mystacinids appear to employ. However, Arkins et al. (1999) stated that Mystacina wings are no shorter, nor are the wingtips more rounded, than would be expected in a microbat of their size. Arkins et al. (1999) also noted that the low wing loading and short echolocation calls typical of gleaning bats are not seen in Mystacina. Further work is clearly needed on the ecomorphology of these bats [image above from here].

The mystacinid tail is short (hence the common name of the group) and emerges through the dorsal surface of the uropatagium. When on the ground, the bats can fold up their uropatagium and tuck it away.

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It is not only the strong adaptation to terrestriality that makes the mystacinids remarkable: Mystacina has proved to be an exceptionally diverse feeder, and has the broadest diet of any bat. It preys on volant and non-volant insects as well as pollen, berries and other small fruits, and has even been reported scavenging flesh and fat from dead birds (Daniel 1979). Most of the insects eaten are beetles, moths, flies and crickets, and together these prey make up at least 50% of the diet. Spiders and other arthropods are also taken and pieces of moss and bark have been discovered in Mystacina droppings (Arkins et al. 1999). Several anatomical features appear to correlate with this remarkably broad diet. The dentary is robust and the dentary symphysis is fused, the ascending ramus is tall, and the anterior toothrow has a crowded look, despite the fact that two lower incisors are absent. Fossil mystacinids from Australia (they’re united in the genus Icarops) have the same sort of jaw structure and dentition as the modern species from New Zealand, and are highly similar to them in postcrania as well. It therefore seems that the Australian forms were behaviourally and ecologically highly similar to the living species (Hand et al. 2005). To return to a point made earlier, this means that the unusual terrestrial adaptations of Mystacina did not evolve in the isolation of New Zealand. Rather surprisingly, these bats were apparently scurrying around on the ground and folding up their leathery wing membranes in environments where there were already numerous small terrestrial mammals. That seems odd, but it’s what the fossil record indicates [adjacent skull image from digimorph].

Mystacinids do not have a shortened rostrum and their premaxillary bones are not reduced as they are in some other groups of microbats. The tip of the muzzle projects beyond the lower lip and the nostrils are set in a small narial pad which supports numerous short, stiff bristles (Hill & Smith 1984, Koopman 1994). The nostrils are prominent, rectangular and oriented vertically. Their ears are long, fully separated (in some groups of bats the ears are connected by a flap or flaps of skin), slender, and with a long, pointed tragus.

Mystacina also exhibits a number of cranial specialisations for its nectarivorous diet. Its tongue is papillated and somewhat extensible with a small array of brush-like papillae at the tip, and there is a wide gap between the upper incisors. The total number of teeth has been reduced to 28 (a more typical microchiropteran count would be 34). Mystacinids therefore represent a third group of bats containing members specialised for a nectarivorous diet (the other two are pteropodids and phyllostomids).

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In an effort to discover more about the ecology of the mysterious parasitic wood rose (Dactylanthus taylorii), a native flowering plant that grows on the roots of small trees, Chris Ecroyd of Landcare Research New Zealand Ltd employed time-lapse video recorders, a method that had previously been successful in revealing predation on Kokako Callaeas cinerea nests. Dactylanthus is peculiar in that it produces copious nectar and has a distinctive musky scent, yet apparently lacks adequate pollinators. Ecroyd filmed various insects, possums and rats visiting the flowers, but he was convinced that none of these represented the true pollinator. Relocating the cameras to film Dactylanthus at various new localities, he was eventually successful and filmed, of all things, Mystacina feeding from the flowers. Mystacina proved to be an important terrestrial pollinator of Dactylanthus, suggesting that the modern rarity of both organisms might be linked (Hunt 1992, Ecroyd 1993, Webb & Kelly 1993, Meyer-Rochow & Stringer 1997). The photo above is one of Ecroyd’s images of a Mystacina with Dactylanthus taylorii (from here).

That Mystacina was nectarivorous had already been determined by Daniel (1976) who had examined stomach contents and droppings, and had also smeared Mystacina chests with vaseline to see what material might stick to the bats while they were foraging. Daniel found that the bats visited a variety of flowers, presumably to feed on both nectar and pollen. Daniel (1979) also reported a captive Mystacina seen ‘excitedly’ crawling over flowers, becoming covered in pollen, and then eating the pollen off its own fur while grooming. This is a unique method of feeding for any bat (though it has not yet been reported from wild mystacinids). The pollen-feeding habit of the mystacinids mean that they join only two other bat groups (pteropodids and phyllostomids) known to feed on pollen.

Throughout this text I’ve consistently referred to mystacinids in the plural. Though mystacinids have a complex and confusing taxonomic history, it is agreed today that there were two species until recently. One survives: Mystacina tuberculata, first described by John Edward Gray in 1843. The second, M. robusta, has not been positively reported alive since 1965. And I’d love to say more, but I have to stop there.

Coming soon: well, actually, I’m going to stop making promises for a while, as the backlog is getting way too long. I will, however, mention more Triassic crurotarsans, more sebecosuchians, proto-narwhals, amazing social lives of green iguanas, pleurodires, and Mesozoic birds.

Refs – -

Arkins, A. M., Winnington, A. P., Anderson, S. & Clout, M. N. 1999. Diet and nectarivorous foraging behaviour of the short-tailed bat (Mystacina tuberculata). Journal of Zoology 247, 183-187.

Daniel, M. J. 1976. Feeding by the short-tailed bat (Mystacina tuberculata) on fruit and possibly nectar. New Zealand Journal of Zoology 3, 391-398.

- . 1979. The New Zealand short-tailed bat, Mystacina tuberculata; a review of present knowledge. New Zealand Journal of Zoology 6, 357-370.

Ecroyd, C. 1993. In search of the woodrose. Forest Bird 1993 (2), 24-28.

Hand, S., Archer, M. & Godthelp, H. 2005. Australian Oligo-Miocene mystacinids (Microchiroptera): upper dentition, new taxa and divergence of New Zealand species. Geobios 38, 339-352.

Hand, S. J., Murray, P., Megirian, D., Archer, M. & Godthelp, H. 1998. Mystacinid bats (Microchiroptera) from the Australian Tertiary. Journal of Paleontology 72, 538-545.

Hill, J. E. & Smith, J. D. 1984. Bats – a Natural History. British Museum (Natural History).

Hunt, R. 1992. Dark secrets revealed by security cameras. BBC Wildlife 10 (9), 12.

Kunz, T. H. 1982. Roosting ecology of bats. In Kunz, T. H. (ed) Ecology of Bats. Plenum Press (New York/London), pp. 1-55.

Koopman, K. F. 1994. Chiroptera: Systematics. Walter de Gruyter (Berlin/New York).

Meyer-Rochow, V. B. & Stringer, I. A. N. 1997. An honorary non-flying mammal pollinator. Trends in Ecology and Evolution 12, 277.

Norberg, U. M. & Rayner, J. M. V. 1987. Ecological morphology and flight in bats (Mammalia: Chiroptera): wing adaptations, flight performance, foraging strategy and echolocation. Philosophical Transactions of the Royal Society of London B 280, 335-427.

Tomes, R. F. 1857. On two species of bats inhabiting New Zealand. Proceedings of the Zoological Society of London 1857, 134-142.

Webb, C. J. & Kelly, D. 1993. The reproductive biology of the New Zealand flora. Trends in Ecology and Evolution 8, 442-447.

Worthy, T. H., Tennyson, A. J. D., Archer, M., Musser, A. M., Hand, S. J., Jones, C., Douglas, B. J., McNamara, J. A. & Beck, R. M. D. 2006. Miocene mammal reveals a Mesozoic ghost lineage on insular New Zealand, southwest Pacific. Proceedings of the National Academy of Sciences 103, 19419-19423.

Comments

  1. #1 John Wilkins
    April 19, 2007

    As usual, mind-bogglingly interesting, Darren.

  2. #2 David Marjanovi?
    April 19, 2007

    a relict lineage of mammals phylogenetically close to multituberculates

    I remember (where did I put the pdf?!?) a large polytomy that also contains the other big clades of the mammalian crown-group, same as in the Volaticotherium paper.

  3. #3 Darren Naish
    April 19, 2007

    David: the St Bathans mammal was found by Worthy et al. to be in a trichotomy with multituberculates and tinodontids + trechnotheres. It’s misleading to imply then that the SB mammal is phylogenetically close to mtbs alone, but it’s still phylogenetically close to them.

    Many thanks John for the kind words.

  4. #4 David Marjanovi?
    April 19, 2007

    OK, but it’s equally close to the multis and to us…

    Incidentally, I almost never write praising comments because they would become very repetitive very fast. People want to read something new in the comments, not something they already know. :-)

  5. #5 Darren Naish
    April 19, 2007

    It’s ok David: I know you love me.

    PS – the St Bathans mammals isn’t equally close to us, as we aren’t basal members of the tinodontid + trechnothere clade. If we were.. then, yes.

  6. #6 Christopher Taylor
    April 20, 2007

    Though mystacinids have a complex and confusing taxonomic history

    Honestly, what group of mammals (or almost any other organism, for that matter) doesn’t have a complex and confusing taxonomic history?

    “Giant gecko” is referring to Hoplodactylus delcourti, no? There’s room for questioning if this is even a New Zealand species. Delcourt’s gecko was described from a single specimen found lying around in a French museum, sans collection data or any indication of origin. IIRC, it was judged to probably come from New Zealand because the genus Hoplodactylus is otherwise only known from New Zealand. However, despite New Zealand’s fairly extensive subfossil record, no remains assignable to H. delcourti have yet been recovered (according to Worthy & Holdaway 2002). I believe there’s also some question about how the specimen got from New Zealand to Marseille. New Caledonia shares a number of herpetofaunal elements with New Zealand, and possibly can’t be ruled out as the origin of H. delcourti. Of course, none of this is in any way conclusive – many of the New Zealand lizards would have had very restricted distributions even before European settlement, and H. delcourti‘s absence from the subfossil record may just be because no-one’s looked in the right place.

  7. #7 Darren Naish
    April 20, 2007

    Thanks Christopher for your comment. Was the giant gecko really from New Zealand? Here is what Naish (2004) had to say about this…

    Despite the reasonable evidence indicating a New Zealand origin for H. delcourti, Trevor Worthy and Richard Holdaway (2002) have recently suggested that H. delcourti never was a native of South Island, and in fact was most likely never a native of New Zealand at all …. This sceptical view is problematical for a few reasons. Firstly, besides the alleged Earnscleugh cloacal bone there is also the jaw described by Hutton (1875), though as noted above this cannot be referred to H. delcourti with certainty. Secondly, given that Hoplodactylus is presently only known from New Zealand, and that H. delcourti is a member of that genus, it is more parsimonious to assume that H. delcourti was native to New Zealand than to assume otherwise and thereby invoke a range extension for the genus. Thirdly, Worthy and Holdaway (2002) are certainly right that the apparent absence of any H. delcourti bones from New Zealand is highly suspicious, but it remains possible that H. delcourti was extremely rare and still awaits discovery in fossil samples. As a palaeontologist working on new taxa discovered in a region where the strata have been systematically scoured on a regular basis of more than 130 years, I have first-hand experience of the fact that rare taxa can be so poorly represented that they remain overlooked or undiscovered for a surprising amount of time. The true origins of H. delcourti remain mysterious and require confirmation, but in the absence of evidence to the contrary it seems most reasonable to maintain that this species was a denizen of New Zealand, albeit a very rare one with no compelling fossil record.

    Ref – -

    Naish, D. 2004. New Zealand’s giant gecko: a review of current knowledge of Hoplodactylus delcourti and the kawekaweau of legend. The Cryptozoology Review 4 (2), 17-21.

  8. #8 David Marjanovi?
    April 20, 2007

    PS – the St Bathans mammals isn’t equally close to us, as we aren’t basal members of the tinodontid + trechnothere clade. If we were.. then, yes.

    It’s equally close to the multis and to the tinodontid + trechnothere clade. We are trechnotheres. So it’s equally close to the multis as to us… under the cladistic definition of “closely related” anyway, not under a distance-based one.

  9. #9 johannes
    April 20, 2007

    Darren, David,

    is there really more than negative evidence (i.e. that they are not related to the haramyids, and the old concept of allotherians is no longer supported) to keep the multis in the crown group?

    Miao (1993) has found a mere two mammalian characters: absence of septomaxilla and presence of three otic ossicles. In my opinion, this is not enough to qualify them as crown group mammals, unless you adhere to the precladistic view that some characters are bigger (i.e. more important) than others and everything that has an one-piece jaw and the right ossicles in its ear is a mammal.

  10. #10 mark lees
    April 21, 2007

    As my previous comments probably made obvious I am no fan of cladistics. At least with Linnaean clasifications we know we are dealing an artificial construct, so-called phylogenetic classification schemes carry a suggestion of being more real (in the sense of trying to accurately describe phylogeny), but in reality they are just a different artificial construct using a different set of assumptions, which may, or may not, be closer to the real relationships. I find that arguments about which characters are more significant, or which should be split up etc when carrying out a cladistic analysis often tells me a lot more about the people doing the analysis than it does about the organisms being analysed.

    I’m not suggesting its a waste of time, just that we shouldn’t put too much weight on the results. The detailed arguments end up seeming a little like two medieval theologians arguing about how many angels can dance on the point of needle. Arcane and abstruse.

    The question of whether multis were mammals (crown group) or not seems to be of this sort.

    I often think that considering how the animals would be likely to have been viewed if they still existed today is useful. Would we view multis as mammals? Well they appear to have been smallish, furred, probably homeothermic (I have never seen anything suggest otherwise) etc – I guess they would be viewed as mammals at least as much as monotremes. They same is likely to be the case for many of the ‘mammaliaformes’ and quite possibly more than a few of the cynodont groups normally placed outside Mammalia sensu lato. If we had access to a broad body of information on the soft biology of these forms I suspect we wouldn’t get quite so hung up on what a few bones in the jaw and inner ear where up to.

    You could extend this thinking to a range of other groups. For example, were dromaeosaurs, oviraptors and the like birds? I am not getting in to cladistic arguments here (others are much better equiped than me for that – my MSc is not in paleontology) – simply if, say, Oviraptor was alive today, bipedal, apparently feathered, etc, would we view it as a bird? On balance I guess the answer is yes.

    What this somewhat laboured comment is getting at is, since what we are really interested in is the animals themselves and not labels, should we get so concerned over whether something is in such and such a ‘crown group’ etc.

    I think I’m going to shut up and put my flak jacket and helmet on now.

    I agree by the way that this blog is rather excellent, lots of food for thought and intriguing titbits.

    There are lies, damn lies, and cladistics (sorry, I just couldn’t resist it!) :)

  11. #11 Mike Dickison
    April 22, 2007

    A wonderful summary of Mysticina; I hadn’t heard about the Australian fossils, and it’s surprising that the terrestrial habit might not be an adaptation to NZ’s unique environment. Can’t wait for some more on M. robustus; I’d heard that rumor too, and was wondering why the NZ Department of Conservation hadn’t checked it out yet.

    On H. delcourti; the lack of subfossil evidence should be taken pretty seriously. Worthy and Holdaway know their stuff, and have excavated many many herp bones from all kinds of deposits all over NZ. I checked out all the thousands of miscellaneous reptile subfossils in the major NZ museum collections when I was doing my Hons project on tuatara, keeping a particular eye out for H. delcourti, and no luck, although a giant gecko would be exceptionally likely to end up in a cave deposit, and these are well known from pretty much every habitat on both main islands, so its absence is suspicious. There are those Maori stories, of course. So do write something about it for those of us who can’t get Cryptozoology Review.

  12. #12 Darren Naish
    April 22, 2007

    Many thanks Mike for your comment, great to hear from someone with hands-on experience of the herpetofauna. You’re no doubt aware of the rich Miocene fauna that’s been discovered at St Bathans: geckos have been mentioned but to date we await details on which taxa have been discovered. Do you know any more?

  13. #13 Christopher Taylor
    April 23, 2007

    Did the holotype mount of Hoplodactylus delcourti preserve any bone material, or would subfossil material have to be assigned to this species largely on the basis of size alone? If the latter, that might reduce the chance of recognising H. delcourti if remains from a not-fully-grown individual were to be found.

  14. #14 David Marjanovi?
    April 23, 2007

    is there really more than negative evidence (i.e. that they are not related to the haramyids, and the old concept of allotherians is no longer supported) to keep the multis in the crown group?

    Oh yes. All studies that have resolved the polytomy ( = Luo et al. 2002 and about half of the more recent ones) have found ((Australosphenida + Shuotherium) (Triconodonta (Multituberculata (Trechnotheria)))). Better yet, all analyses that have included Fruitafossor have inserted it into this topology as the sister-group of (Triconodonta (Multituberculata (Trechnotheria))).

    Miao (1993) is way outdated, and I’m not sure it was up to date in 1993.

  15. #15 David Marjanovi?
    April 23, 2007

    At least with Linnaean clasifications we know we are dealing an artificial construct, so-called phylogenetic classification schemes

    Let me interrupt right here — there is no such thing as “phylogenetic classification”. Cladistics is the method of phylogenetics — of reconstructing the tree, not of naming it.

    (You may be thinking of phylogenetic nomenclature, which was unfortunately originally called “phylogenetic taxonomy”. That is nomenclature: you take a tree, no matter where it comes from, and tie labels to defined places on it.)

    I find that arguments about which characters are more significant, or which should be split up etc when carrying out a cladistic analysis often tells me a lot more about the people doing the analysis than it does about the organisms being analysed.

    No, you don’t find arguments about which characters are more significant in morphological cladistic literature. A basic assumption of cladistics when applied to morphology is that we don’t know how much more probable, in %, certain mutations are than others; therefore all characters are treated as equal, and their number decides. Instead of wasting time quarrelling over whether it’s better to use tooth or braincase characters, or of making untested statements that cranial characters are more conservative than postcranial ones and that the postcranium should therefore be ignored (I’ve seen all that in precladistic literature), we use all characters that exhibit phylogenetic signal, put them into the same matrix, and let them compete. The more characters and taxa, the better: the signal adds up, the noise cancels itself out.

    Whether a character actually contains phylogenetic signal has rarely been tested, but it can be done very easily.

    I’m not suggesting its a waste of time, just that we shouldn’t put too much weight on the results. The detailed arguments end up seeming a little like two medieval theologians arguing about how many angels can dance on the point of needle. Arcane and abstruse.

    There’s one big difference: it’s science. It’s reproducible and testable.

    The question of whether multis were mammals (crown group) or not seems to be of this sort.

    I often think that considering how the animals would be likely to have been viewed if they still existed today is useful. Would we view multis as mammals?

    Here you have conflated phylogenetics and nomenclature. In the first paragraph you ask whether multis belong to the mammalian crown-group; in the second paragraph you ask whether that crown-group should be called Mammalia (as indeed some advocate). These are two entirely separate problems.

    You could extend this thinking to a range of other groups. For example, were dromaeosaurs, oviraptors and the like birds? I am not getting in to cladistic arguments here

    Indeed not. You are talking about nomenclature. :-)

    What this somewhat laboured comment is getting at is, since what we are really interested in is the animals themselves and not labels, should we get so concerned over whether something is in such and such a ‘crown group’ etc.

    Sure, because “crown-group” is a term that describes a phylogenetic tree, and we are interested in phylogeny because we can’t understand “the animals themselves” without understanding their phylogenetic positions; it has nothing to do with how (if at all) we name that crown-group.

    “The crown-group of X”, incidentally, means “the most recent common ancestor of all extant members of X, and all its descendants”. (If that clade is identical with X, then X is a “crown-group” or “crown clade”.)

  16. #16 luca
    April 24, 2007

    Darren, another fascinating piece. Was it here that I read that bats are one of the most successful mammal branches? It’s strange, ’cause I always thought of them as just the bizarre cousin instead. It only goes to show how much I have to learn in the field. And this blog is an excellent place for this.

    I wish you could get your own TV series.

  17. #17 Darren Naish
    April 24, 2007

    Darren, another fascinating piece. Was it here that I read that bats are one of the most successful mammal branches? It’s strange, ’cause I always thought of them as just the bizarre cousin instead. It only goes to show how much I have to learn in the field. And this blog is an excellent place for this.

    Thanks for the kind comments Luca. Bats are one of the most successful mammal groups (in terms of species-level diversity). In very rough figures there are about 5800 living mammals. Of these, about 2300 are rodents, and a further 1100 or so are bats.

    I wish you could get your own TV series.

    Hmm, so do I. I have tried hard, and for the time being have given up. However, I am soon to start work in the world of mass media…

  18. #18 luca
    April 24, 2007

    Well, let us know. if you were to print/edit a book, I’d certainly shell the cash to buy it.

    the after-lunch pit-stop at TetrapodZoology is one of my favourite moments in the day.

    I’ve always dreamed of becoming a palaeontologist, but life has chosen otherwise – for the best I guess, since I’m starting to think that no matter how much I like a subject I start hating it soon after I start working on it. Well, I’ll never know, but will definitely keep reading.

  19. #19 johannes
    April 26, 2007

    > Miao (1993) is way outdated, and I’m not sure it was up to date in
    1993.

    Thank you David.
    Looks like the only way to keep this vile sprawlers* out of the crown group is to retrench with Butler and the allotherian diehards. At least those guys have a better point than the BAND group; nobody made up a protoavis-style missing link between haramyids and multis, and nobody has claimed that the multi hair in those paleocene coprolites was actually not hair, but the result of a strange gastroenterological disease of the poor mesonychid, viverravid or whoever was eating multis back then.
    * just joking, I like multis. Could we expect a post on them, Darren?

    [from Darren: yes, one day.. :) ]

    > Well they appear to have been smallish, furred, probably
    homeothermic

    Furry yes (I have already mentioned the hair in the coprolites), homeothermic yes, smallish not always.
    Bubodens rivals Repenomamus, the largest Stagodontids and the Taeniodont Schowalteria* in size and is therefore among the largest mesozoic mammals. Taeniolabis is even bigger at 30 KG (some sources quote as much as 102 KG, but I think this is not believable).
    Remember also that most multis had plagiaulacoid dentition, and some therian (marsupial) animals with similar teeth have grown up to the size of a leopard, i.e. Propleopus (There are sightings of Propleopus-like cryptids in the USA, especially in the Appalachians, but I think they are most likely the result of celtic imagination or inspired by homemade whisky).

    *David, if Taeniodonts were present in the Cretaceous, why are the
    absent in spec?

  20. #20 Raymond
    April 27, 2007

    From Johannes

    “Bubodens rivals Repenomamus, the largest Stagodontids and the Taeniodont Schowalteria* in size and is therefore among the largest mesozoic mammals.”

    Be careful with *Schowalteria* it’s mainly based on a mandible.It’s possible that it was twice the size of all the other mesozoic eucynodont “giants”.

    Giant Taeniolabids definitely reached 30 kgs, likely much
    more given the size of the largest species.

    “Remember also that most multis had plagiaulacoid dentition, and some therian (marsupial) animals with similar teeth have grown up to the size of a leopard, i.e. Propleopus (There are sightings of Propleopus-like cryptids in the USA, especially in the Appalachians, but I think they are most likely the result of celtic imagination or inspired by homemade whisky).”

    Ah so you’ved heard of the “devil monkeys” as well :D
    They’re not to be confused with escapee red kangaroos and other macropods which have seemingly naturalized in many areas ranging from Iowa to Wisconsin to Alabama.The devil monkeys hop like ‘roos but there is an important difference, they tend to kill and eat pets and livestock.

    “*David, if Taeniodonts were present in the Cretaceous, why are they
    absent in spec?”

    If David had his way, Spec mammals would consist of nothing but sprawlers with a few exceptions for parasagittal gaited cancridonts and pokemurids.

  21. #21 johannes
    April 27, 2007

    > Be careful with *Schowalteria* it’s mainly based on a mandible.It’s
    possible that it was twice the size of all the other mesozoic
    eucynodont “giants”.

    Twice as Large as the other mammaliforms and crown group mammals perhaps, but twice as large as all of the other mesozoic eucynodonts? I mean, twice as large as the largest traversodontids (they were eucynodonts, too). Exaeretodon had a skull that was up to 40 cm long; are we really talking about a taeniodont with an 80cm long skull?

    >If David had his way, Spec mammals would consist of nothing but
    sprawlers with a few exceptions for parasagittal gaited cancridonts
    and pokemurids.

    The cancridonts are eiter whale- or seal-like marine animals or mole-like burrowers, their limbs are far to specialised for swimming and/or burrowing to give them a parasagittal gait when moving on land, the largest and most whale-like of the cancridonts are fully aquatic and unable to move on land anyway.

  22. #22 David Marjanovi?
    April 27, 2007

    Looks like the only way to keep this vile sprawlers* out of the crown group is to retrench with Butler and the allotherian diehards.

    The funny thing is, the Volaticotherium paper did find Allotheria — inside the crown-group (at the usual multi place)! Well, I suppose the middle ear is already diphyletic… *shrug*

    How old is Bubodens?

    If David had his way, Spec mammals would consist of nothing but sprawlers with a few exceptions for parasagittal gaited cancridonts and pokemurids.

    Ah, the usual exaggeration again…

    David, if Taeniodonts were present in the Cretaceous, why are the
    absent in spec?

    Because, like in HE, they died out in the Oligocene (IIRC). :-)

  23. #23 Raymond
    April 27, 2007

    “Twice as Large as the other mammaliforms and crown group mammals perhaps, but twice as large as all of the other mesozoic eucynodonts? I mean, twice as large as the largest traversodontids (they were eucynodonts, too). Exaeretodon had a skull that was up to 40 cm long; are we really talking about a taeniodont with an 80cm long skull?”

    It’s your fault for not specifically referring to post Triassic eucynodonts;) within the “real” Mesozoic(TM)

    “The cancridonts are eiter whale- or seal-like marine animals or mole-like burrowers, their limbs are far to specialised for swimming and/or burrowing to give them a parasagittal gait when moving on land, the largest and most whale-like of the cancridonts are fully aquatic and unable to move on land anyway.”

    The related Bunyip, a NZ cancridont, and several fossilized forms yet to be uploaded IIRC.(you may not have access to some of the old bowdoin/Ocatch pages)

    “Ah, the usual exaggeration again…”

    But not by much :P

  24. #24 David Marjanovi?
    April 28, 2007

    I’ve never understood why there should be monotremes with a parasaggital gait… we don’t really need that.

    It does not help, of course, that we haven’t created most of Australia’s mammal fauna yet in the first place. We haven’t even decided on whether to have marsupials there!

  25. #25 Raymond
    April 29, 2007

    “I’ve never understood why there should be monotremes with a parasaggital gait… we don’t really need that.”

    Done before my time and I agree, *Castorcauda* is obviously
    a horizontally oriented swimmer despite being a sprawly, though the Elder Gods do have the excuse of it being a recent publication.Platypus don’t help matters by being paddlers.Let’s not forget * Potamogale*, which despite being a synapsid with a parasagittal gait, is a vertically oriented swimmer.

    “It does not help, of course, that we haven’t created most of Australia’s mammal fauna yet in the first place. We haven’t even decided on whether to have marsupials there!”

    yes, quite true.

  26. #26 johannes
    April 29, 2007

    >It’s your fault for not specifically referring to post Triassic
    eucynodonts;) within the “real” Mesozoic(TM)

    Thank you for the clarification, Raymond. So you were referring to post triassic eucynodonts (This is what I guessed in the first place, but knowing is better than guessing). Even the true blue, post Triassic, rauisuchian free “real mesozoic” hat its share of non-mammalian eucynodonts, the tritylodonts survived well into the lower cretaceous.

    To come back to Schowalteria, twice as large as, lets say, Repenomamus rather than Exaeretodon is still quite impressive, at least if the animal was build like the better known paleogene Taeniodonts, i.e. like an aardvark on steroids (and I think there is no reason to believe otherwise).

    >The related Bunyip

    The (spec)Bunyip is indeed a monotreme, but not a cancridont.

  27. #27 johannes
    April 30, 2007

    > How old is Bubodens?

    Lancian (70,6 to 65,5 Ma),according to the Paleobiology Database. If anyone knows better, please let me know!

    > Because, like in HE, they died out in the Oligocene (IIRC). :-)

    The taeniodonts died out in the upper middle Eocene in HE (at least I never heard or read of anyone claiming otherwise), I don’t think they made it into the late Eocene, leave alone the Oligocene. Again, if anybody has informations about an late Eocene or Oligocene Taeniodont, they would be welcome!

    As far as spec is concerned, there are other eucynodont clades that went extinct in the eocene or oligocene on HE and are still alive and well on spec (the multis for example, or the polidolopids). But who am I, a mere mortal, to tell the specgods how to do their job?

  28. #28 David Marjanovi?
    April 30, 2007

    Again, if anybody has informations about an late Eocene or Oligocene Taeniodont, they would be welcome!

    Oops. Looks like I confused them with something else. Multituberculates maybe.

    the multis for example

    Largely due to the lack of rodents. I came too late to prevent the creation of the xenotheridians.

    or the polidolopids

    General lack of monkeys.

  29. #29 johannes
    May 2, 2007

    > Largely due to the lack of rodents.
    > General lack of monkeys.

    But the lack of competitors in spec will apply for Taeniodonts as well,
    Tillodonts – who were not particulary successful in HE anyway, the Taeniodonts actually survived longer – and pig like Artiodactyls are absent in spec (OK, there are hogbirds), and there are no Paraselenodonts adapted for rooting and digging either (or are these still under creation?). The Pigshrews are to small to be really competitors, and the sut is restricted to Africa. There is the Digga-dumdum, but Taeniolabidids co-existed with Taeniodonts in HE, so they could do the same in spec.

  30. #30 Raymond
    May 3, 2007

    Johannes, if you wish to debate or pursue Spec related topics further, you’d be wise to use the thread list that’s been the home of it for the past 6 six years.

    http://tech.groups.yahoo.com/group/DinosaurMailingList-KilledThreads/

    Joining shouldn’t be too difficult, just get a yahoo account.The Archives are massive and can often fill you in on ideas,concepts etc. that complement Specworld.

  31. #31 johannes
    May 3, 2007

    > Ah so you’ved heard of the “devil monkeys” as well :D

    Raymond, I have heard of them, but I rarely use the term, because I am by no means sure if the animals are eutherians, leave alone primates (in fact, I am still sceptic if they exist at all, their frequent appearance on websites dealing with alien abductions and other “supernatural” stuff certainly doesn’t help). And if they are, could you call, lets say, an omomyid or carpolestid a “monkey” or even a “lemur”?

    I have also heard other names for what seems to be essentially the same creature, the serbian term “drekavac”, for example (but this might be a generic term for any kind of folkloristic monster) or – my personal favourite – “magnesia poodle” (don’t google, its an oral tradition). The poodle part of the name is easily enough to explain, the animal in question seems to have a dog like head and muzzle, and its neck and shoulders are covered by peculiary thick and long fur, perhaps a mane; the resulting animal would resemble an old male hamadryas baboon, or a poodle (or any other water dog) dressed up in the old manner. But what about the anatolian town of Magnesia?
    It was the seat of the treasury during the days of the byzantine successor empire of Nicaea, undoubtly garrisoned by elite units (the varangian guard comes to mind), so perhaps the name suggests some association with a werewulf like warrior cult (a wild guess)?

  32. #32 David Marjanovi?
    May 3, 2007

    Joining shouldn’t be too difficult, just get a yahoo account.

    Alternatively, you can treat it as a mailing list (as I do): find the address for unsubscribing (it’s in the signature of every single post), replace “unsubscribe” in that address for “subscribe”, and send a blank e-mail there. Has advantages and disadvantages.

  33. #33 Raymond
    May 4, 2007

    -Johannes-

    I just use the term “Devil Monkey” because its been popularized by Loren Coleman to distinguish them from real escapee kangaroos of several species which have begun to naturalize in many areas of the U.S. (shhhhhh don’t tell anyone ;).Devil monkeys are (mostly)bipedal hoppers like ‘roos but are carnivorous, other than that, I don’t have a clue what they are except they’re mammalian.

    Escapee red kangaroos seem inevitable.Texas game ranches have over 70,000 for example.People still report feral camels out west after their “official” extinctions back in the early 1920′s or so.The reality is that many exotic game or domestic mammals and birds from oryx to emus are escaping or being intentionally released here in the U.S.
    Lots are naturalizing, some seem to be breeding as well.

  34. #34 johannes
    May 10, 2007

    Raymond, have you seen this.

    It is supposed to be Argyrolagus, but looks far too robust for an animal the size of a kangaroo rat or gerbil. My spanish is not very good, but it looks to me that they say the animal is 70 million years old, wich is of course nonsense, Argyrolagus being a miocene/pliocene creature. I doubt there were metatherians in South America in the early Maastrichtian, the contemporary south american mammal fauna was a “jurassic revival” affair (docos, kuehnotheriids etc.).
    But by default or design, the picture looks a lot like a “devil monkey” or an american Propleopus.

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