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North America: land of obscure, freaky voles

The views expressed are those of the author and are not necessarily those of Scientific American.

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Arborimus longicaudus, based on a photo in Nowak (1999). Image by Darren Naish, colouring by Gareth Monger.

As a European person, I find European voles (and, to a degree, Asian voles) pretty familiar, commonplace, homely. Still interesting, mind you. But when it comes to North American voles — oh my god, the weird. I don’t even know where to start, so I’ll just launch right in and hope that the crazy string of words that I see forming in front of me as I write this article will make some sort of sense, and won’t give you a seizure or anything like that.

So, the western USA – Oregon and California – is home to the tree voles: the three Arborimus species. Long-haired, dark- or reddish-brown and generally short-faced and cuddly in appearance, tree voles are specialised climbers, moving proficiently on branches and trunks with slow, deliberate movements that make them very different from most other small rodents. The idea that there might be arboreal voles is not that radical – after all, tree-climbing and tree-dwelling are all over the place within muroid rodents – but what’s especially odd about Arborimus is that it’s only the females that live in trees; males (while capable of climbing) live in subterranean burrows or in piles of vegetation. In order to meet up with females (who might be 30 m up in the trees), the males do of course climb, and they also build nests in trees during the mating season (this lasts from February to September). Nevertheless, it seems that we have a fundamental difference here in how males and females partition the habitat. Is this reflected in their morphology? I don’t know, and I don’t know if anyone else knows: comparatively little is known about Arborimus and it was “long considered difficult to find and was unusual in museum collections” (Nowak 1999, p. 1465).

I know that North America is a big place, and that it doesn’t have quite the same length of scientific tradition that Europe does, but, still, the idea that continental mammal species like this might be so poorly known remains surprising. Having said that, there are European mammals known from a handful of specimens – do people realise how little we actually know about so many animals?

Western heather vole (Phenacomys intermedius). Image by Darren Naish, coloured by Gareth Monger.

What sort of vole is Arborimus? I’m not sure on that since I haven’t seen any phylogenetic studies that incorporate it. However, here we have to talk about the heather voles, since several authors have regarded Arborimus as part of another North American vole taxon*: Phenacomys, otherwise containing the Western P. intermedius and Eastern or Ungava heather voles P. ungava (these two were regarded as conspecific until recently. A few subspecies have been recognised for P. ungava but their validity is questionable). Heather voles occur across much of Canada and also across the higher regions of the Rockies, Sierra Nevadas and around the Great Lakes (McAllister & Hoffman 1988). Fossils show that they were far more widespread across the USA during the Pleistocene.

* It has also been suggested that one of the three tree voles, A. albipes, should be included within the otherwise fossil taxon Paraphenacomys.

Phenacomys (above) compared to Myodes/Clethrionomys (below). They look pretty similar, but may be well apart in phylogenies. Image by Darren Naish.

Heather voles were long thought to be rare but, since the 1950s, they’ve become known from an increasing number of localities and specimens and the Western heather vole even seems to be increasing its range in some areas, perhaps because logging practices are providing the more open habitat that it prefer. In fact, “a variety of conflicting observations on the rarity of [these voles] have been made” (Clark & Stormberg 1987, p. 164). There are some indications that they’re harder to trap than is usual for voles (McAllister & Hoffman 1988). Population explosions of the sort known for severeal voles sometimes occur. Overgrazing of suitable habitat seems to be a cause of concern for these species.

Upper (above) and lower (below) jaw dentition in Phenacomys intermedius, from Merriam (1889). Note the strong asymmetry: the occlusal triangles are much bigger on the lingual sides of the teeth.

What do heather voles look like? They’re a bit boring, being highly similar to the myriad Microtus and Myodes/Clethrionomys species that make up the bulk of vole diversity. They aren’t reddish dorsally like Myodes/Clethrionomys voles and have greyer, fluffier fur than other voles in their range. They’re unlike Microtus voles in having closed roots on the cheek teeth. Their teeth are hypsodont, however, and are also distinctive in being quite obviously asymmetrical when seen in occlusal view – the lingual triangles are much larger than the buccal ones. These features (and other dental ones) are also seen in the fossil Blancan vole Hibbardomys, so it might be a close relative (Martin 2007), both being included within the clade Phenacomyini.

There are differing views on where these voles fit in phylogenetic terms. Martin (2007) regarded phenacomyines as nested within the same clade as Microtus and Myodes/Clethrionomys, whereas molecular studies have supported the arguably more interesting possibility that, together with Dicrostonyx (the collared lemmings), heather voles belong outside the clade that includes the vast majority of other voles (Conroy & Cook 1999, Cook et al. 2004, Galewski et al. 2006). According to this latter hypothesis, heather voles and collared lemmings are perhaps as distinct from microtine voles and kin as are true lemmings (Lemmus) and perhaps muskrats (Ondatra). Shock horror: radical stuff. I suppose I don’t really need to say that several additional phylogenetic hypotheses have been published for voles and lemmings (see also Chaline & Graf 1988 and Chaline et al. 1999).

Highly simplified version of one of several possible arvicolid topologies: this is based predominantly on the combined mitochondrial and nuclear gene tree produced by Galewski et al. (2006). Image by Darren Naish.

Fact: muskrats are voles, and what is Dinaromys?

Incidentally, the idea that muskrats are voles is pretty awesome. Consider that an ‘average’ vole is less than 10 cm long and weighs somewhere between 20 and 50 g. Now consider the Muskrat Ondatra zibethicus: a vole with a total length of more than 60 cm (head + body length = 22-32 cm) and which can weigh up to 1.8 kg. So, it would take almost 40 ‘average’ voles to make up one muskrat. Muskrats – ondatrines – have an extensive fossil history extending back into the Miocene.

As just mentioned, muskrats might be outside the clade that contains virtually all other voles, but they might not be: muskrats are part of the same clade as water voles (Arvicola) and red-backed voles (Clethrionomys or Myodes) in some topologies (Galewski et al. 2006). Conroy & Cook (1999) found a muskrat + Arvicola clade to be the sister-group to remaining voles as well as lemmings.

Balkan snow vole (Dinaromys bogdanovi). Image by Darren Naish, coloured by Gareth Monger.

By the way, the weird and poorly known Balkan snow vole or Martino’s vole Dinaromys bogdanovi is an ondatrine according to some rodent workers. A miniature, terrestrial muskrat that lives in the mountains of south-eastern Europe? Err, wow. There are other views on the affinities of this taxon, however: Chaline et al. (1999) say that it should be included within the fossil taxon Dolomys of the European Pliocene and Pleistocene, a close relative (according to them) of the also fossil Pliocene-Pleistocene Pliomys. And that’s a whole ‘nother story…

Colour-change ‘lemmings’ with bi-pronged seasonal super-claws

Collared lemmings (Dicrostonyx) in both summer and winter coats. Image by Darren Naish, coloured by Gareth Monger.

I mentioned Dicrostonyx – the collared lemmings or varying lemmings, the several species of which occur across the far north of Scandinavia, Asia and North America. Well, don’t get me started on them. First of all, these stocky, short-tailed muroids – denizens of the treeless tundra and definitely cold-adapted – look extremely lemmingy, and hence are conventionally regarded as close relatives of the ‘other’ lemmings (Lemmus, Synaptomys and Myopus) and classified with them within Lemmini, an arvicolid group supposedly distinct from the vole clade. Lemmings proper might still be a clade (Conray & Cook 1999, Cook et al. 2004), but it now seems that collared lemmings aren’t among them. Seeing as there’s no push (so far as I know) to get them renamed ‘collared voles’, the term ‘lemming’ is currently applied to unrelated short-tailed, cold-adapted arvicolids.

Secondly, unlike several other vole taxa living in modern North America, collared lemmings are not invaders from the Old World. Seemingly, they’re ancestrally North American animals that invaded Asia and Europe from a Beringian centre of origin. I say this based on the presence in Pleistocene Alaska of Predicrostonyx hopkinsi, sometimes called Hopkins’s lemming and supposed to be ancestral to Dicrostonyx. Thirdly, they’re anatomically odd: they seasonally change colour from greyish or brownish to white – they are totally unique among rodents in this respect – and also grow incredible enlarged, double-pronged manual claws* on the 3rd and 4th fingers during the winter too. These claws (or, technically, claw sheaths) are shed during the summer. Their pinnae are strongly reduced: essentially just being a slim semicircle of tissue concealed by pelage.

Dicrostonyx winter forelimb claws, based on a photo in Nowak (1999). Image by Darren Naish.

* Some sources describe these as “cornifications of the toepads”, which suggests that they’re not claw sheaths per se. This isn’t correct: they are claw sheaths. The pads on the 3rd and 4th fingers become enlarged at the same time as the bifid claws develop. Baby collared lemmings born in the winter have these bifid claws (Hansen 1957). Studies show that their development is hormonally controlled and linked to photoperiod (e.g., Mallory et al. 1981).

There are a few things I’d like to know here and I don’t. What are the underlying bony unguals like? Would we ever guess from their anatomy that they support enlarged, double-pronged sheaths? And are there special adaptations that enable the ‘winter claws’ to become detached? And, given that we have a group of rodents here where the unguals support two highly distinct sorts of claw sheaths at different stages of the animal’s life, here’s another cautionary tale for inferring horny claw shape from fossils (where, typically, only the bony ungual is preserved).

And that’s where we have to end for now. There’s other stuff I’d like to say about voles: I’ll get to it eventually. Rodents for the win.

One final thing. Getting good, usable images of the rodents discussed here is hard, so I had to generate images of my own. To make them prettier, I asked Gareth Monger for help: Gareth previously helped out with my Platyhystrix, and is better known for his pterosaur illustrations. Gareth has a facebook page here. Thanks loads to him for his hard work.

For previous Tet Zoo rodent articles, see…

Refs – -

Chaline, J., Brunet-Lecomte, P., Montuire, S., Viriot, L. & Courant, F. 1999. Anatomy of the arvicoline radiation (Rodentia): palaeogeographical, palaeoeocological history and evolutionary data. Annales Zoologici Fennici 36, 239-267.

- . & Graf, J. D. 1988. Phylogeny of the Arvicolidae (Rodentia): biochemical and paleontological evidence. Journal of Mammalogy 69, 22-33.

Clark, T. W. & Stormberg, M. R. 1987. Mammals in Wyoming. University of Kansas, Museum of Natural History.

Cook, J. A., Runck, A. M. & Conroy, C. J. 2004. Historical biogeography at the crossroads of the northern continents: molecular phylogenetics of red-backed voles (Rodentia: Arvicolinae). Molecular Phylogenetics and Evolution 30, 767-777.

Conroy, C. J. & Cook, J. A. 1999. MtDNA evidence for repeated pulses of speciation within arvicoline and murid rodents. Journal of Mammalian Evolution 6, 221-245.

Galewski, T., Tilak, M.-K., Sanchez, S., Chevret, P., Paradis, E. & Douzery, E. J. P. 2006. The evolutionary radiation of Arvicolinae rodents (voles and lemmings): relative contribution of nuclear and mitochondrial DNA phylogenies. BMC Evolutionary Biology 2006 6: 80 doi:10.1186/1471-2148-6-80

Hansen, R. M. 1957. Development of young varying lemmings (Dicrostonyx). Arctic 10, 105-117.

Mallory, F. F., Elliott,  J. R. & Brooks, R. J, 1981. Changes in body size in fluctuating populations of the collared lemming: age and photoperiod influences. Canadian Journal of Zoology 59, 174-182.

Martin, R. A. 2007. Arvicolidae. In Janis, C. M., Gunnell, G. F. & Uhen, M. D. (eds) Evolution of Tertiary Mammals of North America, Vol. 2. Cambridge University Press, Cambridge, pp. 480-497.

McAllister, J. A. & Hoffman, R. S. 1988. Phenacomys intermedius. Mammalian Species 305, 1-8.

Merriam, C. H. 1889. Description of a new genus (Phenacomys) and four species of Arvicolinae. North American Fauna 2, 27-45.

Nowak, R. M. 1999. Walker’s Mammals of the World, Volume II (Sixth Edition). The John Hopkins University Press, Baltimore and London.

Darren Naish About the Author: Darren Naish is a science writer, technical editor and palaeozoologist (affiliated with the University of Southampton, UK). He mostly works on Cretaceous dinosaurs and pterosaurs but has an avid interest in all things tetrapod. His publications can be downloaded at He has been blogging at Tetrapod Zoology since 2006. Check out the Tet Zoo podcast at! Follow on Twitter @TetZoo.

The views expressed are those of the author and are not necessarily those of Scientific American.

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  1. 1. Heteromeles 10:59 am 01/12/2014

    Cool! One of my favorite undergrad teachers (who passed away recently) had a collared lemming skin mounted flat in the style of a snarling bear rug, mouth gaping and claws extended. He kept it on his desk, as a memento of his time in the Arctic.

    As for the tree voles: you do realize how big douglas-firs (their primary food) get? If you wanted to study red-backed tree voles, you’d need to convince a canopy biologist (most of whom are, um, nuts) to become a vole biologist, or vice versa. Given the politics around American forests and the general dearth of science funding, that might be tricky. Maybe someone will invent a UV spotting scope and the biologist can track vole pee across the canopy or something.

    While douglas-firs don’t AFAIK have the complex canopy ecosystems of old-growth redwoods–which can have trees, shrubs, and ferns growing in their upper branches and over a foot of soil in their crowns, which they root in–doug firs can have moss mats in at least their lower crowns. Conversely, the forest floor under old-growth stands is mostly decaying logs in various stages of breakdown, with things growing out of them. The point is that in tree vole country, some parts of the canopy actually look quite a lot like the forest floor, except that they’re a bit more isolated. Having a terrestrial animal in the canopy isn’t quite as weird as one might expect. After all, it’s also a place where flying squirrels forage for truffles on the ground.

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  2. 2. Jenny Islander 11:35 am 01/12/2014

    Ever since I heard about muskrats being voles (which is in retrospect so obvious!), I’ve wondered about the exact relationships of the Pleistocene “giant beaver.”

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  3. 3. naishd 11:41 am 01/12/2014

    Don’t worry, the ‘giant beavers’ (part of Castoroidinae) are indeed beavers… but they’re very much the tip of the proverbial iceberg when it comes to beaver diversity.

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  4. 4. Yodelling Cyclist 1:57 pm 01/12/2014

    hey’re very much the tip of the proverbial iceberg when it comes to beaver diversity

    I must confess, when the idea of a serious Tetzoo assault on the rodents started floating around, purely from my perspective as a spectator I thought this could be quite dull. No, wrong, the little beggars are packed with awesome. Bravo! Encore!

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  5. 5. vdinets 5:10 pm 01/12/2014

    Well, there is also the round-tailed muskrat which superficially looks and behaves a lot like an Arvicola<.

    Interestingly, the white-footed vole is fossorial rather than arboreal (although it has been observed climbing low shrubs). It is also maddeningly difficult to see in the wild: it took me more than a hundred hours of spotlighting in its habitat before I even saw it. Tree voles are much more easy to find because they eat fir needles in a very specific manner (leaving the central vein), and piles of those needles accumulate under nests. They also nest in young Douglas-firs sometimes (at least the southern species), although, of course, it is a lot more interesting to observe them in big ones. Many years ago, just after emigrating to the US, I worked as a tree climber for a tree-care company and could use their equipment to climb redwoods and other big trees on weekends; it was so much fun to get up there and watch all those canopy species.

    A lot of voles and pretty much all lemmings can be very difficult to trap even when common. Myopus shisticolor, and Lemmus amurensis are the worst. I guess they are just more interested in lichens and fungi than in seeds or whatever people use for bait.

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  6. 6. vdinets 5:22 pm 01/12/2014

    Oh, and a little note: some Siberian subspecies of Arvicola macrotis also turn white in winter, while one subspecies of collared lemming (the one from the Aleutians) does not.

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  7. 7. AlHazen 5:34 pm 01/12/2014

    Hmmm… If tree voles aren’t well-known… One of the surprises I had in Australia was discovering how much “Mountain Ash” (a Eucalypt, the basic forest tree of the temperate rain forest in the Dandenongs, outside Melbourne) LOOKS like Douglas Fir: tall columnar trunks, foliage concentrated on the upper portions. (Evidently a tree habitus that has advantages if you grow on a mountain side where misty air blows in from an adjoining North Pacific or Antarctic Ocean.)
    Do you suppose there is some small possum — possibly even one not officially known — that has evolved to behave and look like tree voles?

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  8. 8. Tayo Bethel 5:44 pm 01/12/2014

    … BEAVER DIVERSITY? That is intriguing to say the least. Please enlighten.

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  9. 9. Jenny Islander 5:58 pm 01/12/2014

    @AlHazen no. 7: Is or was.

    There are seven known species of vole in Alaska alone. On Kodiak Island we have the tundra vole, M. oeconomus. Tundra voles build towns under the snowpack in the winter, rather like lemmings (and other species of vole for that matter). They harvest winter-dried grasses and burrow upward for seedheads. It’s fairly easy to trace the effects of a single spell of unusually warm/cold/dry winter weather on the voles, simply from noting how many my cat brings home in the summertime. If we have a consistent snowpack in the winter, there will be a bumper crop of voles in the summer, perhaps so many that he will just kill them, look at them like “bleah, not voles again,” and leave the un-munched corpses for me to clean up. If there is hardly any snow, he will get a vole per day or fewer. The worst years for voles appear to begin with a good snowpack, but switch to a chinook in the depths of winter. I think that what happens is that the voles didn’t spend much time insulating their summer burrows in south-facing slopes and under rock outcrops, so when the cold rainwater comes trickling down through the snowpack, they don’t have a warm dry place to go and they die of hypothermia in huge numbers.

    Re my cat: My husband and I are aware of the deleterious effects of cats. Because we have a cat, we don’t have a bird feeder. We have also removed the lower branches of the tree on a sloping part of the property where ignorant young bird couples tended to nest on the assumption that the large amount of space in front of them meant that they were safe, when they were only a foot and a half above ground level in the back. Also, we keep a cat in the first place because if we didn’t we would have rats from the nearby waterfront. Putting down poison would endanger everyone else’s cat, local scavenging birds, etc.

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  10. 10. naishd 7:10 pm 01/12/2014

    Thanks for all the great comments. I had to hold off from discussing or mentioning all the collared lemming species, otherwise I might never get this article finished. Yes, there is variation in the seasonal colour change of the collared lemming taxa – I’ve heard that there’s variation, as well, in whether they grow the bi-pronged winter claws but I couldn’t find anything on this in the literature.

    I didn’t know that Large-eared voles Arvicola macrotis change colour, thanks, Vlad.

    Oh, I also didn’t mention the hair-like filaments of dried resin that tree voles leave behind, and which gather up to look like weird balls of greenish hair. I can’t imagine what people would think of these if they didn’t know what they were.

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  11. 11. naishd 7:12 pm 01/12/2014

    I should also say that there’s more weirdness in North American voles not mentioned here. If you know about the stuff to do with parental care… please don’t let on! It’s covered in another article, to appear soon.

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  12. 12. Christopher Taylor 7:41 pm 01/12/2014

    beaver diversity

    I wrote a brief overview of beaver diversity a couple of years ago here. This includes palaeocastorines, a terrestrial group that dug 2.5 metre-long coiling burrows, and two separate lineages of giant beavers that reached bear-like sizes. The North American giant beaver Castoroides was a reasonably close relative of modern aquatic beavers; the European giant beaver Trogontherium may be in the same group, or it may be a more basal terrestrial form.

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  13. 13. BrianL 2:36 am 01/13/2014

    Is there any reason why there aren’t any huge aquatic voles a la *Ondatra* native to Eurasia? *Ondatra* certainly seems to survive and thrive pretty well here so it doesn’t seem like Eurasian waterways are in any way inherently unsuitable for animals like these.

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  14. 14. vdinets 3:10 am 01/13/2014

    Jenny Islander: You mean you have “wild” Norway rats running around? Or do you live in a town?

    AlHazen: Tree voles are not completely obscure. I’ve seen them in a few documentaries about redwood forests; also, they are among the species that timber companies are required to survey for before logging, so at least distributional data is pretty good. White-footed and heather voles are much less known.

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  15. 15. Tayo Bethel 7:49 am 01/13/2014

    Rats are fairly common on waterfronts all over the world, whether it be town or country.

    Comment 13–the reason there are no native giant aquatic volesin Eurasia might just lack of evolutionary opportunity. Since thisis just speculation I will stop there so I might be corrected :)

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  16. 16. vdinets 12:33 pm 01/13/2014

    Tayo: I know, but Kodiak Island is a bit far north for them, although the climate of Alaska’s southern coast is kinda mild (there is even a colony of European rabbits in Valdez). The thing is, distribution of both Rattus spp. in North America is surprisingly little known. Most maps just show Norway rat as occurring all over the continent, even though it’s been eradicated in Alberta and is extremely rare in Montana and some other places.

    I find it equally strange that muskrats have never colonized Eurasia through Beringia. Today they occur in the tundra almost all the way to the Arctic Coast. The same for American beaver: it occurs around Nome and does very well in Siberia when introduced, but never made the crossing.

    BTW, the species name for Microtus oeconomus comes from an old Russian word for “manager”. It refers to this species’ extensive, well-organized colonies with food stores and runway networks.

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  17. 17. Yodelling Cyclist 12:54 pm 01/13/2014

    Beringia was an oddly selective filter. Is there any reason to believe it was, by some obscure mechanism, very dry? Climatically hostile/possessing an obscurely hostile terrain?

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  18. 18. Dartian 12:59 pm 01/13/2014

    The idea that there might be arboreal voles is not that radical

    Arborimus isn’t unique in this regard. The Eurasian bank vole Myodes glareolus, at least, can and does climb trees occasionally.

    A lot of voles and pretty much all lemmings can be very difficult to trap even when common. Myopus s[c]histicolor, and Lemmus amurensis are the worst. I guess they are just more interested in lichens and fungi than in seeds or whatever people use for bait.

    Don’t know about Lemmus amurensis, but the wood lemming is indeed supposed to be very hard to trap under normal conditions (it mostly eats moss, BTW). However, during peak years wood lemmings will readily enter traps (although presumably not because of the bait). Researchers in Finland were able to catch enough specimens of this species to found entire breeding lab colonies already in the 1960ies. That’s how the wood lemming’s bizarre sex ratio was discovered (though I won’t say more about this in case Darren plans to cover this subject in some future article).

    some Siberian subspecies of Arvicola macrotis also turn white in winter

    That’s news to me. And quite unexpected news too, considering that Arvicola isn’t exactly the most typical high-boreal specialist. Do you have a reference for this? Or links to pictures of these voles in white winter pelage?

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  19. 19. naishd 1:02 pm 01/13/2014

    The Eurasian bank vole Myodes glareolus, at least, can and does climb trees occasionally.

    Sure. But it’s not arboreal.

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  20. 20. Dartian 1:17 pm 01/13/2014

    Sure. But it’s not arboreal.

    Maybe not, but it does climb trees, and not entirely infrequently either. V. Holišová did a trapping experiment in a patch of woodland in what was then Czechoslovakia (in the late sixties; unfortunately I don’t have the full reference). She placed live-capture traps in various trees at a height of 2.8-3 m. Of 277 small mammals captured in these traps, 127 were bank voles.

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  21. 21. naishd 1:22 pm 01/13/2014

    Sure, I know – lots of animals that we regard as ‘terrestrial’ frequently climb (weasels [least weasels], apparently, are another example). I don’t doubt the Bank vole’s adept climbing ability, but, it’s not arboreal.

    Anyway… what I’d like to know about Arborimus is: does it display any specialisations for its (mostly) arboreal lifestyle? As you’ve just highlighted, most ‘normal’ voles can climb with ease – is Arborimus anatomically unusual, or is the switch purely behavioural (they climb and live in trees because they can)? Questions, questions…

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  22. 22. vdinets 3:04 pm 01/13/2014

    Yod: Some cold-tolerant freshwater fishes did make it across Beringia (i. e. one species of sucker and one or more blackfishes), but Siberian salamander didn’t, which seems to point to fast glacial rivers being the barrier. There is some literature on this; I think I posted some reference a few posts ago.

    Dartian: did I say “Arvicola macrotis“? What a stupid typo! Alticola, of course. I’m not aware of any photos, but there is a painting (done from a live animal) by Smirin in Russian-language Mammals of Russia by Dinets & Rotshild. I can scan it for you if you’d like.

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  23. 23. vdinets 3:11 pm 01/13/2014

    Darren: I’ve seen it specifically stated somewhere in literature that it doesn’t have any adaptations to arboreal lifestyle whatsoever.

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  24. 24. vdinets 3:22 pm 01/13/2014

    Dartian: actually, it’s already online:
    Under #289c, from Yakutia.
    In case you are interested, others are: 285 Chionomys nivalis, 286 Ch. gud, 287 Ch. roberti, 288 A. argentatus, 289a-b Ch. macrotis from Chukotka in summer and in winter, 290a-b A. strelzovi in summer and in winter.

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  25. 25. David Marjanović 4:16 pm 01/13/2014

    Well. My flabber is well and truly gasted.

    Well, don’t get me started on them.

    Followed by four paragraphs you wrote all on your own. :-þ

    a canopy biologist (most of whom are, um, nuts)

    Details, please! :-)

    Do you suppose there is some small possum — possibly even one not officially known — that has evolved to behave and look like tree voles?

    Only one way to find out.

    the species name for Microtus oeconomus comes from

    I bet it’s also a pun on Mus. I’m angry Anonymus was preoccupied by a polychaete and the rodent of that name had to be renamed.

    Beringia was an oddly selective filter. Is there any reason to believe it was, by some obscure mechanism, very dry? Climatically hostile/possessing an obscurely hostile terrain?

    Perhaps the filter wasn’t Beringia, but the short time window between the opening of the ice shield to its east and the establishment of the Bering strait. I guess that for many American species the Yukon area only became passable when the Bering area wasn’t passable any longer; when they reached Alaska, they couldn’t go farther.

    The ice shield was the barrier between Panthera spelaea and Panthera atrox (…both of which may be lumped into P. leo according to the species concept of your choice, or not).

    Darren: I’ve seen it specifically stated somewhere in literature that it doesn’t have any adaptations to arboreal lifestyle whatsoever.

    But then, it’s routinely claimed that goats have no adaptations to climbing, for instance…

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  26. 26. ekocak 4:27 pm 01/13/2014

    I had no idea about the muskrat thing. Very cool. Also, well done on the illustrations. They look great.

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  27. 27. Dartian 4:36 pm 01/13/2014

    “Alticola, of course.

    Ah, that makes more sense! (Although in retrospect, I should perhaps have figured that out myself. Oh well.)

    Under #289c, from Yakutia.

    Many thanks!

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  28. 28. naishd 4:51 pm 01/13/2014

    Ditto on Arvicola being Alticola – I thought this was odd but didn’t bother to check. Thanks to all for such great comments – when things go well, we all learn a lot. And when don’t they go well? THIS. IS. TET ZOO!!

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  29. 29. Heteromeles 5:30 pm 01/13/2014

    @David #28: The canopy biologists I’ve known are wonderful people generally, but they’re a bit, erm, high energy. Like the one who got married to his wife somewhere up in a large redwood. This same scientist (yes, he’s the one you associate with climbing old redwoods) has also been known to share his pictures straight down from the top of very tall redwoods with audiences with little or no warning. Basically you’re straight looking down 100 meters or so, on a wide screen and in full color. I wasn’t the only one who got instant vertigo. Dude has no fear of heights, and it’s a bit disconcerting.

    Note that I’m *not* knocking their dedication to safety or their deep understanding that what they do is freaking dangerous. I just put them in the same category as the ichthyologists who SCUBA down to 100 meters to find new fish species. They’re nuts too, and I’m glad they’re available to do the work so I don’t have to.

    Come to think of it, there was a guy who free-climbed smaller trees in the Amazon (no ropes) to trap small arboreal mammals for his PhD. I wonder whatever happened to him? Maybe he’d do a red-backed vole study.

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  30. 30. BilBy 7:39 pm 01/13/2014

    Re: lack of arboreal adaptations – if you’re small perhaps big trees just don’t seem that different from terrestrial habitats. Once up in the heights it can’t be much different for a vole than running over twigs and leaf litter on the ground – but why the sex difference? Tangentially related – in the US I often saw gray foxes run up quite steep sloping tree trunks to avoid me and my dog – I have never seen red foxes do that (though I imagine they can).

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  31. 31. Tayo Bethel 7:39 pm 01/13/2014

    Come to think of it, muskrats are not the only wetland species that failed to cross Beringia. American mink and American beaver (as stated in a previous comment) failed to cross. very interesting..

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  32. 32. Heteromeles 8:33 pm 01/13/2014

    Gray foxes are known for climbing trees. There’s a picture (which does not appear to be online) of one sitting at the top of a telephone pole. The photographer noted that the fox climbed it every day or so, and took the picture. I’m not sure whether they’re more arboreal than raccoon dogs (the other tree-climbing canid), but there are plenty of pictures and videos (eg

    As for the tree voles, there’s a picture of an alleged muskrat up an alleged tree at I guess being small and having claws helps climbing, or something.

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  33. 33. Cameron McCormick 11:28 pm 01/13/2014

    I have never seen red foxes do that (though I imagine they can).

    I saw one run straight up a willow trunk for about 2 meters when in pursuit of a Grey Squirrel. Of course, it seemed to have been more the result of momentum rather than arboreal capabilities!

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  34. 34. Jenny Islander 12:29 am 01/14/2014

    @vdinets no. 14, 16: IIRC you speak English as a second language–? A waterfront is always and only a constructed shoreline, with docks, breakwaters, etc., so if you see a waterfront mentioned, you can assume that at least there used to be a town there (or used to be water there, for some ancient buried cities). Kodiak is a fishing town, so our waterfront is lined with seafood processing plants; their waste nourishes Norway rats the size of small housecats. Luckily they don’t like large housecats much. The one rat that a cat ever caught on our property was a young one, probably too dumb to understand Why We Do Not Go Where It Smells Like Cats. Our old cat was so proud! She woke us up at ridiculous o’clock yelling at the top of her lungs, then walked around in a circle purring and flirting her tail while we admired her one and only rat.

    Re Beringia as a filter: I don’t know whether this is a relevant datum; let better educated minds decide. Although the Kodiak Archipelago has always been oceanic, and although the present Shelikof Strait is 50 km wide, Kodiak harbors bears. Kodiak was only joined to the mainland by a continental glacier during the last Ice Age. AFAIK the fringes of the ice mass were calving icebergs into the ocean throughout this area–no beaches to be seen. Although there was a refugium, it was well inland; there was no way onto Kodiak soil except over the ice, which separated it from the nearest ice-free part of the mainland by hundreds of km. In fact, genetic evidence suggests both that when the ice melted, bears stopped arriving,* and also that while the island was icebound sufficient bears arrived to establish a genetically healthy population.** So why don’t we have woolly mammoths,*** non-introduced caribou, or other species that could have eked out a living on the tundra of the refugium? Why is the only other mammal that seems to have been there all along a vole?

    My admittedly uninformed opinion is that bears are about the only species bigger than a vole**** that could have gotten here alive. They are long-legged, well-insulated, powerful, good at scrambling around on rough surfaces, omnivorous to the point of eating the insulation from buried electrical cables without apparent ill effect, able to maintain enormous fat reserves, and also able to drowse away the winter.

    ****Probably washed here before the big freeze; raw snags, hunks of tussock turf, etc., still show up from the mainland on the side facing Shelikof Strait, according to my sister who has a cabin out there. It would only take one pregnant vole in a well-lined summer burrow.

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  35. 35. vdinets 3:27 am 01/14/2014

    Grey fox is called Zorro Gato (“cat fox) in parts of northern Mexico for its climbing abilities.

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  36. 36. David Marjanović 7:23 am 01/14/2014

    in the US I often saw gray foxes run up quite steep sloping tree trunks to avoid me and my dog – I have never seen red foxes do that (though I imagine they can).

    Gray “foxes” and red foxes are not at all closely related. IIRC, they’re about as far apart as canids can be.

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  37. 37. Tayo Bethel 10:46 am 01/14/2014

    @Christopher Taylor:

    I read your article on beaver diversity. Recommended for all tetrapod enthusiasts. Right now I’m eagerto know more about the paleobiology of ancient beavers–and information is,as usual, difficult to come by. Or am I just not looking in the right places? BTW … how did we get onto the subject of canids?Thread drift … :)

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  38. 38. Yodelling Cyclist 11:35 am 01/14/2014

    @Christopher Taylor & @Tayo Bethel:

    I must echo the previous comment, the ancient beaver article is great fun. Thoroughly enjoyed wondering through some (what I could in the time available) of the rest of the blog archive. Particularly the pieces on Paleocene mammals. Good stuff.

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  39. 39. vdinets 1:12 pm 01/14/2014

    Jenny Islander: thanks for the info! English is my second language, but the use of the word “waterfront” is variable: in some places (i. e. northern British Columbia and northern Louisiana) I’ve heard people who had forest cabins near lakes refer to the portion of shoreline on their property as “waterfront”, and not as a joke.

    Interestingly, wolves and caribou have been known to cross not only Bering Strait, but also the much-wider De Long Strait (to Vrangel Island) over sea ice, but brown bears have not. Bears haven’t even colonized Isle Royal. Patterns of island colonization are always strange. Is there any paleo data from Kodiak, by the way? Is it possible that some species did make it, but went extinct?

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  40. 40. Yodelling Cyclist 6:02 pm 01/14/2014

    Ah, when it comes to bears and Wrangel Island (I hope I’ve got the right island!) one should note that the polar bear density on that island is very high. I can well imagine some lethal territorial disputes in summer, with the polar bears having the advantage of being able to feed at sea during the worst winters.

    I have no idea whether Wrangel was part of Beringia or not, but it is worth noting that mammoths famously reached the island.

    “Paleo data” (nice phrase!) from those islands which are remnants of Beringia could be illuminating (albeit biased in favour of upland species).Any Challenger island ground sloths? ;-)

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  41. 41. Yodelling Cyclist 6:03 pm 01/14/2014

    Sorry, I mean Commander Islands.

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  42. 42. Jenny Islander 7:35 pm 01/14/2014

    @vdinets no. 39: Did they use “waterfront” as an adjective? In commercial jargon, “waterfront property” means “this parcel of land provides legal access to a body of water big enough to put a boat in, should the buyer be so inclined.” “Waterfront” as a noun referring to an unimproved beach is a new one to me, but a linguist I am not.

    Re fossils on Kodiak, the only fossil terrestrial(?) tetrapods that I am aware of are desmostylians. While any bones on open ground are likely to be consumed by rodents, etc., there are enough peat deposits, etc., that if there had been mammoths here, we would have found something by now. It’s a pity.

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  43. 43. vdinets 9:58 pm 01/14/2014

    Yod: Wrangel I. has remarkably rich Pleistocene fauna, including rhinos, horses, reindeer, bison etc., but, if I remember correctly, no bears. It also has 20+ endemic plants, more than a hundred endemic inverts, and two endemic lemming taxa, plus an endemic guillemot subspecies. Nobody knows what makes it so special; some people say it had better refugia than the mainland, others say it’s because Arctic steppe is so well-preserved there. Which is also interesting, because some people think that large herbivores were necessary for the existence of the Arctic steppe, but Wrangel Island had none of these in historic times. In the 20th century reindeer and muskoxen were introduced to support a Yupik village that the Soviet government established to refute US territorial claim. (Prior to that, the island had no permanent population, although there are lots of archeological remains. The modern history of the island is a bit unusual – look it up on Wiki). Interestingly, the muskoxen don’t cause nearly as much damage to all those endemic plants as the reindeer do. More recently, additional reindeer and then wolves have moved over from the mainland, and there were claims that at least one wolverine had done it, too, but I don’t know if it’s true. Prior to the establishment of the reindeer population, wolves used to visit the island sometimes, but never remained for long. All of this is really remarkable if you consider that the island is 140 km from the mainland.

    Jenny: they used it as a noon, but although it seems logical that a waterfront property should have a waterfront, I have no idea if such usage is considered grammatically correct.

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  44. 44. Jenny Islander 11:13 pm 01/14/2014

    Eh, big language-region, tons of dialectical variation. And in a place with as big a transient population as Kodiak, it all smushes together. I once got instantly branded as a dumb hick when I asked my new college dormmates if they wanted me to get sodapops for them as well when I went to the vending machine to get one for myself. I had had no idea that “sodapop” was considered hayseed dialect in suburban Connecticut. (Why I was supposed to care escaped me then and still does.)

    Back to tetrapods: The bits of the mainland biota that managed to make the crossing to Kodiak seem to have established an ecosystem into which the previously missing tetrapods–mountain goat, Sitka blacktail, etc.–can be dropped with little apparent damage. At least, studies done so far have shown only a change in distribution of Amelanchier (highbush cranberry, serviceberry).

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  45. 45. David Marjanović 10:38 am 01/15/2014

    Wikipedia implies the reindeer have been migrating annually across the ice since mythological times or longer.

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  46. 46. BrianL 12:31 pm 01/15/2014

    Speaking of beavers, according to Wikipedia there are more or less accidentally introduced Canadian beavers in Norway. It also mentions that the two living species of beavers can’t hybridize because of chromosomal differences. Does anybody know if and how the two species co-exist where their ranges meet? One would expect them to occupy (largely) the same niche.

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  47. 47. BrianL 12:33 pm 01/15/2014

    Also, given how well various American rodents (beaver, muskrat, nutria, grey squirrel) seem to do in Eurasia, are there any Eurasian rodents succesfully introduced to North America apart from house mice, Norway rats and black rats?

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  48. 48. vdinets 4:39 pm 01/15/2014

    BrianL: Canadian beavers spreading from Scandinavia have replaced Eurasian beavers in parts of northwestern Russia. This is potentially a major problem since they have broader habitat preferences. They also do well in colder parts of Siberia where Eurasian beavers have not occurred in historic times.
    To my knowledge (and I’ve looked into this just recently), there are no other established Old World rodents in North America (unless you count Hawaii where Polynesian rat occurs), but there are two lagomorphs. European hare was common in northeastern US and southeastern Canada for a few decades, but then declined for unknown reasons and is now present only in Toronto area (unless I’ve missed something). This is a bit mysterious since there are millions of square miles of apparently suitable habitat, and no obvious competition except from native rabbits. European rabbit occurs in a number of small populations, most of them on predator-free islands.

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  49. 49. Jenny Islander 10:11 pm 01/15/2014

    I think I’ve got it!

    Here’s how to get mammoths in the present day with minimal handwaving. The point of departure from our timeline (OTL) is simply the decision of an M. primigenius matriarch to delay a little longer at one particular location.

    It is the tail end of the Wisconsinian. A herd of M. primigenius, under pressure from the rising sea levels, increasing swarms of wetland insects, and hunting, has retreated to the rapidly thawing Alaska Range. They are always on the move, from the gravelly river valleys at the base of the glaciers to nunataks and the thin vegetation growing on debris-covered ice fields, traveling mostly on median and lateral moraines. It is an extreme environment for this species, but they have managed to eke out a living.

    The herd traditionally grazes the herbage on an exposed gravel slope before traveling downhill and along an extremely dangerous ice shelf that fringes the still-partially-icebound Shelikof Strait to get to their next stop. In OTL, the matriarch, hesitant to venture out to a place where she remembers relatives disappearing into a crevasse, delays for a day–and an earthquake brings the slope down and destroys the herd. In this ATL, she decides to press on and the herd is on the ice shelf when the earthquake makes it calve.

    The enormous chunk of ice drifts across the rudimentary Shelikof Strait, pushed by gale and current to the icebound shore of Kodiak Island near present-day Karluk. In OTL it deposits nothing but a few stray rocks as it slowly melts. In the ATL, the matriarch, driven by the scent of food wafting over the eroding glacial fringe of the island, takes the herd into the water off the lowest point she can find. Then the herd picks their way through the icy landscape in a feat worthy of Shackleton, reaching the tundra of the Refugium a few days later. Unlike Shackleton, the matriarch loses many, perhaps most, of her herd by the time they arrive at their destination. There are just enough individuals left to continue the species through a genetic bottleneck comparable to those suffered by Acinonyx jubatus and Bison bonasus. The mammoths breed undisturbed (except when a bear takes the occasional unwary adolescent) for several thousand years.

    Around 4,500 BP, the first humans reach the shores of Kodiak Island. They discover a population of between 500 and 1,000 huge, shaggy beasts, like nothing they have ever seen or heard of. They have no way of knowing that compared to their ancestors these mammoths are puny. They still outweigh the biggest Kodiak bears by several tons. The mammoths circulate between the inland alpine tundra and the river valleys that head the island’s many fjords. Bears use their trails, but try to avoid trouble with them, even when they lie down in salmon streams in an attempt to cool off. The humans decide to do the same. Although there are no other large herbivores on Kodiak, there is plenty of less hazardous large prey along the shores, and anyway it doesn’t seem wise to anger a beast that can stomp your village flat all by itself and generally brings its friends. Not to mention the multiple bears that are always found near a mammoth carcass! So the people learn how to use the mammoth’s shed wool, independently inventing spinning and knitting, and collect its bones and ivory, but they don’t have a tradition of hunting it. Accident and old age are the main causes of death as they have been since the mammoths arrived.

    Mammoths come under heavy pressure when the Russians show up, although at first the Russians don’t have the firepower unless a mammoth wanders within cannon range. Swashbuckling Teddy-Roosevelt-wannabe Americans make the pressure even heavier. (TR himself only bags one and writes disapprovingly of overeager hunters who can’t be bothered to learn how to kill an elephant before they shoot at it.) However, due to the terrain of alder swales, bogs, near-vertical slopes, and shoulder-high tussock that makes hiking the Kodiak wilderness a matter of a mile or two per day, enough mammoths survive to keep the species viable. The ethics of hunting mammoths from small airplanes are hotly debated in the pages of The Alaska Sportsman even as team after team of fascinated scientists uses the same airplane charter services to camp out in mammoth territory and study the incredible beasts.

    The mammoth, along with the Kodiak bear, is mentioned in FDR’s creation of the Kodiak National Wildlife Refuge in 1941. By this time there are several hundred animals left and some herd territories have been empty for years. Still, a limited number of hunting licenses continue to be issued. Full protection for the mammoth finally comes in 1973 under the Endangered Species Act. Now they can be killed only if they are an immediate danger to life and property. The herds are multiplying and spreading once again, and there is talk of ranching them on the mainland for reintroduction to Siberia.

    And last night you saw the best footage ever shot of mammoths in the wild on “Island of the Great Beast,” in reruns on the National Geographic Channel.

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  50. 50. Jenny Islander 12:12 am 01/16/2014

    *Alaska Peninsula. And me with an A in state geography once upon a time!

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  51. 51. Dartian 12:32 am 01/16/2014

    according to Wikipedia there are more or less accidentally introduced Canadian beavers in Norway

    Whoah! Shock horror: Wikipedia is wrong! There are (deliberately) introduced Canadian beavers in Finland, but not in Norway.

    Does anybody know if and how the two species co-exist where their ranges meet?

    Anecdotal evidence from both Finland and Russia suggests that they don’t seem to be able to coexist for very long – and the North American species seems to have the competitive edge (mostly perhaps due to greater reproductive output). See here for more information.

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  52. 52. vdinets 4:08 am 01/16/2014

    Jenny: Nice scenario… but if mammoths were present on Kodiak, the landscape would be very different, with networks of broad trails and lots of open grassland. Native people would kill them off as easily as they did on other islands (and almost certainly on the mainland, but that’s a different and potentially endless subject).

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  53. 53. Chabier G. 5:41 am 01/16/2014

    Castor canadensis was also introduced in Poland, in the mid 20th century, when it was still regarded as conspecific with C. fiber, now it’s extinct there, and reintroduced populations of European beaver are thriving in this country, it seems that American beaver has been outcompeted by C. fiber. Then, maybe, by some reason, C. canadensis has the advantage in high latitudes, but C. fiber would be the winner in warmer environments.


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  54. 54. Jenny Islander 12:44 pm 01/16/2014

    @vdinets no. 52: Actually Kodiak already has lots of broad trails and open grassland. The forest on the north end of the island is quite recent, is composed of pure stands of browser-unfriendly Sitka spruce, and has the type of understory that large animals move through without pushing it down.

    Even if mammoths could graze down the tussock in open country, there are alder swales to consider. Locally these are called “alder hells.” It is possible to spend all day trying to get through an alder swale a hundred yards wide. Even the bears generally wade up the creeks that run through them. There are swaths of Kodiak into which local people did not go, despite inhabiting the place for thousands of years, because of the alder swales. These trees are rich in resins that make mammals ill, and they grow very quickly; even if mammoths chose to spend time plucking them out instead of simply wading in the creeks, they wouldn’t eliminate them.

    Then there is the terrain to consider. Crumple a piece of newspaper: that’s the topography of Kodiak. None of the peaks are very high, but they are as close together as the teeth of a zipper and their sides are steep. Although it is only 100 miles in a straight line from one end of Kodiak to the other, hiking the old Alutiiq trails takes two to three weeks.

    Human choices, however, would be a very important factor in the viability of a hypothetical mammoth population on this island. I note that the Hawaiians, for example, did not hunt out the native monk seal and did not go whaling at all, although they could have. I don’t know about the seal, but the oft-stated reasons for Hawaiians not hunting whales were assigning spiritual value to the whales and not having a pressing need for whale products. So with the Kodiak Alutiiq. As far as practical matters go, the shores teem with edible animals that provide meat, fat, large hides, etc., with less risk to life and limb, and even the largest pinniped is small enough to process and store using pre-Contact technology before the ubiquitous quarter-ton-plus bears come sniffing around en masse. Going away from immediate material concerns, we should consider sportsmanship as well as mysticism. Life was easy enough before the Russians came that people did go hunting for fun, but their chosen trophy animal was the Pacific halibut, which grows huge enough to pull around a small boat in an exciting but not necessarily lethal struggle and doesn’t move in herds. Plus, a technology relying on ground slate weapons used to attack mostly at close range and from above would need a lot of tweaking in order to take down even a solitary mammoth. And while they did hunt the local bears, it was an extremely spiritualized pursuit, involving a kind of secret society that regarded bear hunting as metaphysically as well as physically dangerous. Even though they stacked the deck by using aconite on their spearpoints, there is no evidence that Alutiiq bear hunting put the local bears in danger of extirpation. Instead, the Alutiit learned to reach a detente with bears, even though bears often gathered in large groups near resources the Alutiit also exploited, even though some bears did attack people regardless. I think they could have done the same with mammoths.

    The serious pressure on the last mammoth population would come, I think, when people who could blast away at a mammoth from a distance, take the bits they liked best, and sail or fly away before the bears showed up arrived on Kodiak. And humanity being what it is, even in the face of paleontologists, sportsmen, ecological crusaders, etc., the mammoths of Kodiak would probably be nearly driven out of existence.

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  55. 55. Heteromeles 2:48 pm 01/16/2014

    So is Kodiak big enough to keep mammoths from doing the island dwarfism thing?

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  56. 56. vdinets 3:18 pm 01/16/2014

    Jenny: maybe or maybe not; cultural taboos are often illogical and thus unpredictable. A few things:

    At the time of European contact, Hawaiian monk seals were extinct on all inhabited Hawaiian islands. They are beginning to re-colonize them only now.

    Elephants are specifically adapted to breaking through dense thickets (just look how their eyes are positioned), and are as good at it as black rhinos, if not better. Stomach contents and dung analysis show that alder was at or near the top of woolly mammoths’ menu (see Mammoths and the Environment by V. Ukraintseva, Cambridge Univ. Press, 2013). I think Columbian mammoths were more of grass-eaters, with the niche of shrub browsers in their range filled my mastodons, but I might be wrong

    In many parts of the Russian Far East, the landscape and vegetation are rather similar to Kodiak, except in addition to alder there is also Japanese stone pine, which forms even less penetrable thickets. It didn’t save the sable from being driven nearly to extinction throughout its range by the early 20th century, and much of this extermination was done by native peoples using simple traps.

    The Aleuts used an exceptionally wasteful method for whaling: they would hit a whale with a poisoned harpoon and wait for the carcass to wash up on the shore. Only a small fraction actually did wash up on the right island and soon enough to be of much use. I suspect that the reason mammoths disappeared over such an immense range was that the peoples who hunted them used something similar, and had to kill many to get one.

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  57. 57. Jenny Islander 4:15 pm 01/16/2014

    @Heteromeles 55: I think so. 100 miles in one direction, up to 60 miles in the other, with clusters of near-shore islands to which mammoths might easily swim to further expand their habitat. I think that given the colonization event, the mammoths first encountered by humans (and that’s 4500 BC, not BP–my mistake!) would be recognizably elephant-sized, just not as big as their ancestors. So a mature bull would be about 7 feet tall at the hump and a bit over 2 tons.

    @vdinets: Good points about large mammal hunting methods and the unpredictability of human decisions.

    Another factor to consider: Some sources regarding Hawaiian culture note that apart from declaring whales kapu, Hawaiians considered the meat to be insipid (compared to pork?). Modern connoisseurs, however, compare it favorably to beef. OTOH, while the Kodiak Alutiiq could easily have gathered crabs of species that sell for absurdly high prices per pound these days, they refused to eat them due to the corpses of drowned people tending to turn up with crab-nibbles all over them. Human opinion is a strange and many-faceted thing; I think it’s plausible that people whose palates were used to sea lion pot roast and limpets cooked on seaweed would have tried mammoth–remember, the first mammoths human eyes had seen for thousands of years–and gone, “Ecch!”

    Or maybe not. Or maybe so. If this timeline were ours, I think anthropologists would write a lot of papers on why the Kodiak Island mammoths survived long enough to be declared a protected species, while the Wrangel Island mammoths didn’t.

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  58. 58. Jenny Islander 4:19 pm 01/16/2014

    Forgot: Or they could have seen mammoths examining the bones of their dead with their bizarre boneless-yet-dexterous noses–snakes were also unknown to the Alutiiq–and found the whole thing so creepy that they declared the animals to be out of bounds.

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