Homage to The Velvet Claw (part II)

i-df6b3f8d049a80ffd9a14f12fc0f6357-Roz Gibson prehistoric motley.jpg

On to more of my thoughts about the TV series The Velvet Claw (part I is here). In the previous article, I discussed the art and animation used in the series, all of which was really quite good and very interesting in often featuring fairly obscure creatures...

There's one really important thing I haven't yet mentioned about The Velvet Claw: the fact that both the book and the TV series was written by David Macdonald, director of the Wildlife Conservation Research University at Oxford University and very well known for his many, many publications on biology, ecology and conservation [homepage here]. A leading mammalogist, Macdonald is best known for his long-term work on foxes, but he's also published on jackals, badgers, mink and other mustelids, cheetahs, beavers, squirrels, mice, voles, vicunas, babirusas, elephants and others. Macdonald is also well known for the several TV documentaries he's been involved in, some of which (e.g., 'Night of the Fox' and 'Meerkats United') have been award-winners. I don't know if he approached the BBC with the idea of The Velvet Claw, or if they approached him; whatever happened, the results were outstanding [the adjacent image, featuring an assortment of fossil predatory mammals and not taken from The Velvet Claw, is by the excellent Roz B. Gibson: I hope it's alright to use it. Gibson's website is here].

Anyway, it wasn't just the animation scenes from The Velvet Claw that have really stuck with me: some of the real-life sequences they included in the series have to be seen to be believed (Jerzy is right that at least some of the sequences were filmed in sets, but not enough for me to have a problem with the way the animals were portrayed). Books often say that bears can be comparatively fast runners when the need arises, but you rarely get to see this on TV (god help you if you see this in real life; it might be the last thing you ever see). Episode 5 of The Velvet Claw (ironically titled 'Life in the Slow Lane') shows a grizzly bear in pursuit of wapiti. The bear is a lean individual, and it really moves: fast enough to catch a calf. Granted, neither the deer nor the bear get to run flat-out at great speed, but it's still impressive. Even better is a tree-top chase sequence where a pine marten pursues a red squirrel. The marten sees the squirrel and follows it up the tree. They run up trunks, they run down trunks. They jump from branch to branch. The squirrel leaps to the ground and makes a mad dash. The marten follows. They run up again, they run down again. The marten is close to the squirrel. It ends in a stand-off, with the squirrel working as hard as it can to stay on the opposite side of the trunk from the marten. Eventually, the squirrel makes a wrong move...

A few things from the series have dated, of course. Episode 1, in its coverage of mesonychians, quite naturally says that mesonychians gave rise to whales, and an animation sequence shows an amphibious otter-like Hapalodectes evolving into a seal-like pakicetid and then on to humpbacks. Hapalodectids may well have been amphibious, and - at the time - it wasn't misleading to imply that they might have given rise to whales, given that at least some phylogenetic studies did, in some data runs, find hapalodectids to be closer to whales than were other mesonychians (O'Leary & Geisler 1999). Today, the morphological support for an affinity between mesonychians and basal whales like pakicetids has fallen away as basal whales have proven to be closer to, or part of, Artiodactyla. Furthermore, we now know that pakicetids weren't seal-like proto-whales, but long-limbed slinkers, superficially recalling long-tailed pointy-toothed antelopes [there's a ver 1 article about pakicetids here].

i-42047b6d0672bec309dcb826d72ed84f-cimolestes skull.jpg

The Velvet Claw also sinned in using little Cimolestes from the Upper Cretaceous and Palaeocene as a possible ancestor of carnivorans and creodonts [adjacent image shows the speculative 3-D skull reconstruction of Cimolestes used in The Velvet Claw]. However, this has a precedent: Savage and Long, in their highly influential semi-technical volume Mammal Evolution: an Illustrated Guide, included Cimolestes within the 'insectivore' section of the book, but depicted it and its relatives as the ancestors of carnivorans in a phylogram (Savage & Long 1986, p. 65); this was all based on Savage's earlier paper on carnivorous mammals in which Cimolestes was, again, promoted as a possible ancestor of carnivorans and credonts (Savage 1977, p. 241). Cimolestes and its supposed relatives don't really have any derived characters that might link them with carnivorans, and in fact views on what they are have varied quite widely among experts. Several recent phylogenetic studies incorporating Cimolestes and similar archaic little predators have concluded that they are stem-eutherians and not part of the crown-group, and hence not really anything to do with the ancestry of any extant eutherian clades (Ji et al. 2002, Luo et al. 2007, Wible et al. 2007).

i-1a741d69a9b48cc589b74c8a170e3693-Daptophilus squalidens Cope 1873.jpg

New work has also changed our views on the affinities and phylogeny of several of the carnivoran groups featured in The Velvet Claw. Nimravids (represented in the series by the 'false sabre-tooth' Hoplophoneus) were described in The Velvet Claw as dog-group carnivorans: the latest work in fact indicates either that nimravids are outside of the feliform-caniform clade (Peigné 2001, 2003), or are feliforms (Wesley-Hunt & Flynn 2005). Incidentally, there have always been very good reasons for doubting an affinity of barbourofelines with nimravids: most recently, Morlo et al. (2004) argued that the two aren't close relatives, and that barbourofelines (which they re-named barbourofelids) are feliforms, and in fact the sister-taxon to Felidae. However, if nimravids are really feliforms too, as found by Wesley-Hunt & Flynn (2005), then maybe nimravids and barbourofelids are close relatives after all [adjacent image shows the skull of the cursorial nimravid Dinictis felina].

i-786834d663fee25454a81feb131a598e-proteles.jpg

In the hyaena episode, The Velvet Claw promoted the view that aardwolves (Proteles) are old, 'primitive' hyaenids more closely related to basal, dog-like forms like Tongxinictis and the ictitheriines than to the bone-crushing hyaenine hyaenas. However, some workers have argued that aardwolves instead descend from hyaenines close to Lycyaena and Chasmaporthetes: Koepfli et al. (2006) found that aardwolves had diverged relatively recently, and hence perhaps from a bone- and meat-eating hyaenine rather than from more basal ictitheriine-like taxa [aardwolf P. cristata shown in adjacent pic].

Red pandas were regarded as part of the procyonid lineage in The Velvet Claw: this is now doubtful, but I have more to say about that in a future post. Similarly, skunks were regarded as mustelids, whereas recent work indicates that they may instead be outside of Mustelidae (for more on this see the Erongo carcass article). The Madagascan carnivorans (Cryptoprocta, Eupleres and Fossa.. the last in that list is the fanaloka, not the fossa) were regarded as viverrids and only coincidentally sharing Madagascar with the galidiine mongooses. Morphological and molecular analyses now show that all the Madagascan endemic carnivorans form a clade for which the name Eupleridae Chenu, 1852 has been co-opted (Yoder et al. 2003, Gaubert et al. 2005).

i-84408c81e99e9a0170ea29e43731b42a-Central African linsang there aren't any good photos of these animals on the web bar one and I refuse to use that because everyone else does.jpg

The Velvet Claw also included linsangs among the viverrids, which is (again) fair enough, because that's what most workers thought at the time. The two linsang genera (Asiatic Prionodon and African Poiana), despite their strong morphological similarity, have proved controversial. Based on genetic data, Gaubert & Veron (2003) concluded that the two were quite disparate: while Poiana does indeed seem to be a viverrid (specifically, the sister-taxon to genets), they argued that Prionodon is far more archaic and the sister-taxon to Felidae. This is one of those old ideas that was mooted early on - it was put forward by Thomas Horsfield (1773-1859) in 1821 - but later mostly ignored. In an extensive analysis of morphological characters, Gaubert et al. (2005) seemed slightly sceptical of rampantly homoplastic linsangs (but continued to depict this in their cladograms), but it was strongly supported in another, more recent molecular study (Gaubert & Cordeiro-Estrela 2006). Another taxon conventionally included among viverrids - Nandinia binotata, the African palm civet - is also no longer thought to belong here; instead it seems to be the most basal extant member of Feliformia (Gaubert & Veron 2003, Flynn et al. 2005, Gaubert et al. 2005, Wesley-Hunt & Flynn 2005). Monogeneric 'family' names (Nandiniidae and Prionodontidae) are used by some workers for these 'former viverrids' [the adjacent image, taken from here, is supposed to depict the Central African linsang Poiana richardsoni. Maybe it does, but the animal doesn't much resemble the picture of the same species in Kingdon's field guide].

Given that many of these areas concern relatively obscure creatures, and concern information that is generally avoided by TV-people (they only want to cover lifestyles and behaviour, not phylogenetics, homoplasy or taxonomy!), I doubt that the affinities of nimravids, the non-monophyly of viverrids, or the controversial affinities of aardwolves would get covered if The Velvet Claw were re-made today.

i-06ac4c344c4ecb46d6ce7a0da3d14cd3-Velvet Claw video.jpg

But then, of course, it wouldn't be re-made today; there just isn't the backing for anything like this any more. As I mentioned in part I, the last episode of The Velvet Claw discusses the decline of specialised and big-bodied carnivoran species. Today we might lament the decline and near-extinction of good natural history on TV. I do watch some stuff on TV, but for a long time now my general feeling has been that TV is shit. And remember that we Brits have a right to feel more angry about this than the rest of you: we have to pay to watch television. I kid you not: we actually have to buy and constantly renew a ***king license to own a television set. I have a pie-in-the-sky fantasy that one day things will turn around, and that people who make the decisions in media-land will appreciate that good TV gets watched just as much, if not more, as all the crap that we're currently subjected to.

But I'm not holding my breath.

PS - you can still obtain The Velvet Claw on video (here). No sign of a DVD release though.

Refs - -

Flynn, J. J., Finarelli, J. A., Zehr, S., Hsu, J. & Nedbal, M. A. 2005. Molecular phylogeny of the Carnivora (Mammalia): assessing the impact of increased sampling on resolving enigmatic relationships. Systematic Biology 54, 317-337.

Gaubert, P. & Cordeiro-Estrela, P. 2006. Phylogenetic systematics and tempo of evolution of the Viverrinae (Mammalia, Carnivora, Viverridae) within feliformians: implications for faunal exchange between Asia and Africa. Molecular Phylogenetics and Evolution 41, 266-278.

- . & Veron, G. 2003. Exhaustive sample set among Viverridae reveals the sister-group of felids: the linsangs as a case of extreme morphological convergence within Feliformia. Proceedings of the Royal Society of London B 270, 2523-2530.

- ., Wozencraft, W. C., Cordeiro-Estrela, P. & Veron, G. 2005. Mosaics of convergences and noise in morphological phylogenies: what's in a viverrid-like carnivoran? Systematic Biology 54, 865-894.

Ji, Q., Luo, Z.-X., Yuan, C.-X., Wible, J. R., Zhang, J.-P. & Georgi, J. A. 2002. The earliest known eutherian mammal. Nature 416, 816-822.

Koepfli, K.-P., Jenks, S. M., Eizirik, E., Zahirpour, T., Van Valkenburgh, B. & Wayne, R. K. 2006. Molecular systematics of the Hyaenidae: relationships of a relictual lineage resolved by a molecular supermatrix. Molecular Phylogenetics and Evolution 38, 603-620.

Luo, Z.-X., Chen, P., Li, G. & Chen, M. 2007. A new eutriconodont mammal and evolutionary development in early mammals. Nature 446, 288-293.

Morlo, M., Peigné, S. & Nagel, D. 2004. A new species of Prosansanosmilus: implication for the systematic relationships of the family Barbourofelidae new rank (Carnivora, Mammalia). Zoological Journal of the Linnean Society 140, 43-61.

O'Leary, M. A. & Geisler, J. H. 1999. The position of Cetacea within Mammalia: phylogenetic analysis of morphological data from extinct and extant taxa. Systematic Biology 48, 455-490.

Peigné, S. 2001. A primitive nimravine skull from the Quercy fissures, France: implications for the origin and evolution of Nimravidae (Carnivora). Zoological Journal of the Linnean Society 132, 401-410.

- . 2003. Systematic review of European Nimravidae (Mammalia, Carnivora, Nimravidae) and the phylogenetic relationships of Palaeogene Nimravidae. Zoologica Scripta 32, 199-229.

Savage, R J. G. 1977. Evolution in carnivorous mammals. Palaeontology 20, 237-271.

- . & Long, M. R. 1986. Mammal Evolution: An Illustrated Guide. Facts on File Publications, New York & Oxford.

Wesley-Hunt, G. D. & Flynn, J. J. 2005. Phylogeny of the Carnivora: basal relationships among the carnivoramorphans, and assessment of the position of 'Miacoidea' relative to Carnivora. Journal of Systematic Palaeontology 3, 1-28.

Wible, J. R., Rougier, G. W., Novacek, M. J. & Asher, R. J. 2007. Cretaceous eutherians and Laurasian origin for placental mammals nears thr K/T boundary. Nature 447, 1003-1006.

Yoder, A. D., Burns, M. M., Zehr, S., Delefosse, T., Veron, G., Goodman, S. M. & Flynn, J. J. 2003. Single origin of Malagasy Carnivora from an African ancestor. Nature 421, 734-737.

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"Today we might lament the decline and near-extinction of good natural history on TV." Really? The Blue Planet, Life in the Undergrowth and Planet Earth, all made within the last half-dozen years. And of course Live from Dinosaur Island. Are you seriously arguing that these are "shit"?

Yeah Mike, but... that's it. You're talking about stuff that accounts for about 0.1% of the TV schedule.

As for Live from Dinosaur Island.. surely you jest. Just because I was on it, you don't have to be kind :)

Hmm... you Brits at least have BBC documentaries. Protect it as historical houses and natural monuments from turning into commercial pulp!

Knowing very superficially phylogeny, I begin to wonder if there is a way to decide when enough facts have accumulated that phylogeny is definite and unchangeable.

Everybody is familiar with changing phylogeny. Then there should be a way of not just toying with new interpretation of "that-is-related-to-that" but being able to know when it is sure. For example, molecular tree on whole genome would be 100% sure - or at least nothing could be made more sure.

It would help future TV shows not to age ;-)

Wonderful series of posts Darren!

I've heard that like US PBS, you Brits had to publically fund Channel Four/BBC and so on.I had no idea you actually have to pay for a TV _license_!Doesn't that give you folks more incentive to protest, petition and demand better quality programing?

I wonder if Ms. Gibson would be so kind to include entelodonts, phorusrhacids and perhaps the bizarre ptolemaiids in a possible future updating of her beautiful collage?

Permission for a speculative waffle about carnivoran phylogeny?
I recently re-read your Version 1 post about the origins of the domestic dog, and read the Koler-M??? paper you cited. All plausible in themselves. I also saw an article about dog genetics in the September-October issue of the "American Scientist," which was illustrated(*) with a (molecular, I think) cladogram of the Canidae that I found interesting.
The basal branch -- sister group to all other Canids -- is two species, one of them the North American gray fox (Urocyon cinereus). Above that there are about four main branches, and species with common names including "fox" show up on three of them. Assuming that the people who coin common names aren't TOTAL fools, this suggests that species matching a FOX "stereotype"(**) are common among the Canidae, and this stereotype is perhaps likely to be primitive for it. A stereotype that includes: smallish for a Canid, relatively short-limbed, more likely to hunt small prey alone than big prey in packs. The one branch that contains big dogs (Canis, Lycaon) seems a bit aberrant!

.... But something like your postulated proto-dog is a lot CLOSER to the fox stereotype than a full-scale timber wolf would be. They would be a bit bigger and longer-limbed than foxes, a bit more likely to gang up, probably moving into different habitats from their fox-ish ancestors. Speculation: assume a radiation of such species, jackals and pariah dogs perhaps being typical remnants. In terms of morphology and ecology, they are half way from fox-ish to wolf-ish: it is not implausible that a sister-species of one of these critters could "run away" from the stereotype and evolve into a much larger species that pursues large prey in packs! For the headline in the popular press: wolves are descended from dogs!

Sheer speculation, of course.
--
(*) Article is by Elaine Ostrander, but the illustration is not included in only on-line version I could find, and may have been iserted by the editor.
(**) Term-- for the "idea" people have in mind with a word-- from early-1970s papers on semantics by the American philosopher Hilary Putnam. It includes, for a word for a kind of animal, morphotype PLUS (not necessarily completely accurate) ecological and behavioral characters.

By Allen Hazen (not verified) on 23 Nov 2007 #permalink

Hmm... you Brits at least have BBC documentaries. Protect it as historical houses and natural monuments from turning into commercial pulp!

And yet we hear that with the imminent retirement of Attenborough, the Beeb is planning to cut the natural history department down significantly...

Knowing very superficially phylogeny, I begin to wonder if there is a way to decide when enough facts have accumulated that phylogeny is definite and unchangeable.

Not really such a thing, unfortunately. Before the molecular phylogenies and the recovery of the complete pakicetid specimens, the mesonychian-cetacean connection probably seemed about as secure as you could get, and no-one really suspected a major shift. Possibly ditto a paenungulate-perissodactyl connection.

For example, molecular tree on whole genome would be 100% sure - or at least nothing could be made more sure.

Ah, but then you get the question of the analytical model used to calculate the tree, how the sequences were aligned, etc. etc...

I'll start by lamenting about one of the few things I remember about the series: First it declared that the creodonts had turned the crushing last molar into a cutting one; this turned them into meat-only specialists doomed to extinction, and indeed they died out, predicably. And then it declared that the cats had done exactly the same; this made them vastly successful the world over. Ehem!

Then the TV licence thing: that's normal all over Europe. And, yes, the quality of German and Austrian TV has hit rock-bottom over the last few years (though in Austria this seems to be connected to a policy of the 2000 -- 2006 government). Still, there is one advantage to this model of financing: the public channels don't interrupt their program every 15 minutes for a 20-min commercial break! The German private channels can be very, very, very annoying that way.

Knowing very superficially phylogeny, I begin to wonder if there is a way to decide when enough facts have accumulated that phylogeny is definite and unchangeable.

Of course not. Phylogenetics is science. Science can disprove, but it cannot prove. It can find the truth, but when it has found the truth, there is no way to find out whether what it has found is the truth. If something is wrong, we can figure out that it is wrong; if something is right, we cannot figure out that it is right.

For the headline in the popular press: wolves are descended from dogs!

Doesn't work with the timescale. In your scenario dogs and wolves would have had to separate millions of years ago.

Ah, but then you get the question of the analytical model used to calculate the tree, how the sequences were aligned, etc. etc...

And you can always add taxa, and this can always change the tree.

By David Marjanović (not verified) on 24 Nov 2007 #permalink

> I wonder if Ms. Gibson would be so kind to include
> entelodonts, phorusrhacids and perhaps the bizarre
> ptolemaiids in a possible future updating of her
> beautiful collage?

Ptolemaiids were predators :-0? Tell me more, please! I always thought they were specialists for crushing hard-shelled food, perhaps clams or crustaceans?

BTW, Ms Gibson's side is splendid, obviously there are much more sinister Salems around than Mr Saberhagen...

I don't know if Raymond is necessarily saying that ptolemaiidans were predatory: Simons & Bown (1995) wrote that dental morphology and wear indicates vertical crushing in members of the group, perhaps suggesting a diet of shelled molluscs and/or insects. Cote et al. (2007) argued likewise, though they suggested that the Miocene kelbids might have been a bit more adaptable. By the way, ptolemaiidans are due to be covered in the Tet Zoo field guide to Palaeogene mammals. If only I could devote more time to blogging...

Ref - -

Cote, S., Werdelin, L., Seiffert, E. R. & Barry, J. C. 2007. Additional material of the enigmatic Early Micoene mammal Kelba and its relationship to the order Ptolemaiida. Proceedings of the National Academy of Science 104, 5510-5515.

Simons, E. L. & Bown, T. M. 1995. Ptolemaiida, a new order of Mammalia - with description of the cranium of Ptolemaia grangeri. Proceedings of the National Academy of Science 92, 3269-3273.

Thanks for the info, Darren!

I thought that being included in the predator collage meant that the animal in question was a predator, too (or at least an omnivore with predatory tendencies).

Tet Zoo field guide to Palaeogene Mammals; happy happy, joy joy :-)!

"Of course not. Phylogenetics is science. Science can disprove, but it cannot prove. It can find the truth, but when it has found the truth, there is no way to find out whether what it has found is the truth. If something is wrong, we can figure out that it is wrong; if something is right, we cannot figure out that it is right."

Well, you have 1 and 2 type error. Error of rejecting true huypothesis and accepting wrong hypothesis. Will think a little about it, maybe there will be some formula coming.

First thing would be looking on how incomplete and related characters influence the tree.

First thing would be looking on how incomplete and related characters influence the tree.

This has been done. There are lots of papers on computer simulations of the effects of missing data (the June 2003 issue of the Journal of Vertebrate Paleontology consists mostly of such papers), and lots of empirical papers, too; correlated characters must not be used, because they are the same as one character that is, without justification, given as much weight as several.

By David Marjanović (not verified) on 24 Nov 2007 #permalink

I should have mentioned the conclusion, too: missing data are nothing to be afraid of. In most cases they do not decrease the resolution of a tree -- character conflict does.

Darren, great decision to open the comments! Maybe a post of yours will make it into the "top 5/most active" ScienceBlogs posts :-)

By David Marjanović (not verified) on 24 Nov 2007 #permalink

"I don't know if Raymond is necessarily saying that ptolemaiidans were predatory"

I'm not certain if they were predators in a strictly vertebrate sense, I've only seen speculations to that effect.That said, many of the wacky ptolemaiids were large enough to pursue rabbit to deer sized prey if they so chose.Certainly their dental structure wouldn't have prevented them from so doing.Too bad there is not a bonebed of species with dental markings corresponding to ptolemaiids.It took such a bonebed with *Archaeotherium*
to put to rest the idea that entelodonts were exclusively omnivorous scavengers.

When I saw those computer-model of the skull I actually remembered that I have also once seen one or perhaps two parts of this documentations. But this is really a very long time ago.

I remember the black background with the white grid that was put behind all computer graphics! :o)

By David Marjanović (not verified) on 25 Nov 2007 #permalink

Man, I had a whole bunch of questions about whales that have all been answered by this post. (Or at least the previous post that this one directed me to) I had no idea that a more complete fossils of packicetus had been found that finally confirmed the double-pulley ankle connection to artiodactyls. (I thought they still only had the skull to go on) It's too bad though, I really was hoping that whales were living decendents of mesyonchids, that would have been neat. My brother-in -law, who is a biology teacher, still promotes the andrewsarchus-whale connection in his class. I guess I should send him a copy of this post.

***WARNING: Sudden, desperate departure from the intelligent conversation.***

"And yet we hear that with the imminent retirement of Attenborough, the Beeb is planning to cut the natural history department down significantly..."

SAY IT AIN'T SO!!!! BBC docs have kept me sane these last few years of "Reality" TV mayhem. Seriously, this could be the worst news EVAH.

"Long live The Attenborough!" - thus says this Canadian, anyway.

I've been trying to find this series, etiher on DVD or... 'other places.' with no luck. Any tips. Amazon doesn't seem to have it either.

By arachnophile (not verified) on 29 Nov 2007 #permalink

You are complaining ?
Well we (the Dutch) also have to pay for our three pathetic Public channels. But in Holland there are none, zero, zip Nature or History documentaries. We only have games, 'funny' shows, more 'funny' shows and yet more games and some leftwing political programs.
I only watch the German and BBC these days.
The BBC is heaven compared to our archaic tv shows.

The only thing I watch on Dutch television are some British Drama's (just once a week).
It's no wonder the Dutch are flooding the Amazon.uk system on their desperate search for quality dvd's.
There's a reason why 'Our People' have become wanderers all over the world.
When a plane crashes down in Timboektoe or Kyrgysistan, I bet there was a Dutchman inside trying to escape the shallowness of Dutch Society.

We only have games, 'funny' shows, more 'funny' shows and yet more games and some leftwing political programs.

Are you lucky! The two public-owned Austrian channels don't even have that â they've got endless heartache stories and abominations vaguely based on traditional folk music!

But one of them occasionally imports BBC documentations.

By David MarjanoviÄ (not verified) on 22 Apr 2009 #permalink