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Eld’s deer: endangered, persisting in fragmented populations, and morphologically weird… but it wasn’t always so

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


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Eld's deer stag, by Raul654, used under Creative Commons Attribution-Share Alike 3.0 Unported license.

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This is Eld’s deer Cervus eldi* or the Brow-antlered deer, Thamin or Tamin, a moderately obscure, CITES-listed Old World deer discovered (by Lt. Percy Eld) in India in 1839. It was later found to occur in fragmented populations across much of south-east Asia and also in southern China. Fossils are known from Java and it seems that its original distributed was enormous, being mostly continuous across eastern India and Burma, Thailand, Laos, Cambodia and southern China.

* Many sources spell the specific name eldii but it seems that ‘eldi’ is correct. I’m putting the species in Cervus following Pitra et al. (2004).

Eld’s deer is fairly large (shoulder height is about 1.2 m, weight is 95-150 kg) with lyre-shaped antlers, the beams of which grow outwards before turning inwards. The brow tine is especially long, as hinted at by one of the common names, and appears to form a continuous curve with the beam “so that in profile the antlers appear to be bow-shaped” (Nowak 1999, p. 1108).

Eld’s deer has often been characterised as a tropical wetland specialist. However, this is not true when we look at all three recognised subspecies. In fact these different subspecies are pretty distinctive and again highlight the fact that ‘subspecies’ are not just entities invented for the purposes of bookkeeping but, rather, distinct lineages that we often need to pay attention to.

Cervus eldi eldi stag, antlers in velvet, quite possibly not photographed in the wild. Photo by Mongyamba, licensed under Creative Commons Attribution-Share Alike 3.0 Unported license.

The Manipur subspecies (C. eldi eldi) [shown here] is a true wetland deer, possessing especially large, spreading hooves and peculiar cornified skin on the rest of the digits. It lives on thick, dense mats of floating vegetation known locally as ‘phum’ or ‘phumdi’ (Geist 1999) and is critically rare, being restricted to a tiny area about 15 km square. As of 2004, there were only about 180 individuals in the wild. About another 180 animals are kept in zoos. It was actually thought extinct prior to rediscovery in 1951.

Burmese brow-antlered deer (C. e. thamin), photographed at Zurich Zoo by Albinfo, licensed under Creative Commons Attribution-Share Alike 2.5 Generic license.

Meanwhile, the Burmese or Burma brow-antlered deer C. e. thamin and Thai brow-antlered deer C. e. siamensis are rather different, being animals of dry deciduous forests. This might negate suggestions that the Eld’s deer lineage as a whole is specialised for wetlands. However, there are suggestions that the modern habitat of the Burmese and Thai forms is not natural but the result of human persecution and the removal of their ancestral habitat due to agriculture (Lekagul & McNeely 1977). High nucleotide variation in these populations – similar to that of really widespread deer like roe and sika – suggests that both forms were previously very abundant and widespread (Balakrishnan et al. 2003), an observation that agrees with the idea that their ranges have been substantially decreased in historic times.

The Eld’s deer population on Hainan is sometimes recognised as the subspecies C. e. hainanus but the validity of this form was not supported by Balakrishnan et al. (2003). These authors found the Hainan population to be close to indisputable members of the Thai form and hence recommended included it within that subspecies.

That study (based on mtDNA analysis) also found the nominate subspecies to be closer to the Burmese form than to the Thai one. That’s surprising, since the latter two look extremely similar while the Manipur form is the really distinctive one. Furthermore, all three subspecies were distinct with little evidence for intermingling, and all three exhibit their own phylogeographic structure (Balakrishnan et al. 2003). While it therefore makes sense that all three be recognised as distinct genetic entities for the purposes of conservation and viability, the problem is that some of the populations (including the Manipur one and the Hainan C. e. siamensis one) now exhibit very low amounts of genetic variation and would probably benefit from increased genetic variation. It might therefore be appropriate to start moving animals around in order to increase the genetic health of the populations (Balakrishnan et al. 2003).

Eld's deer diorama at the AMNH. Photo by ideonexus, licensed under Creative Commons Attribution-Share Alike 2.0 Generic license.

A cervine – but what sort of cervine?

Eld’s deer looks odd, and for this reason Pocock (1943) decided that it should warrant it own genus. He therefore came up with Panolia for this species [UPDATE: see comments!! I was not aware of the new taxonomy used by Groves & Grubb (2011) while writing this article]. In overall appearance, though, Eld’s deer is not all that different from the Barasingha Rucervus duvauceli and the now extinct Schomburgk’s deer R. schomburgki (though antler form is different between the three, of course). Ellerman & Morrison-Scott (1951) and Geist (1999) therefore grouped all three together in Rucervus; Geist noted that these species shared relatively high antler mass, short tails, no rump patch, large, subhypsodont teeth specialised for grazing, and legs and feet specialised for soft ground.

Geist's (1999) depiction of the three supposed close relatives placed in Rucervus. L to r: Barasingha, Eld's deer, Schomburgk's deer. Compare the Eld's deer to the stag in the AMNH diorama shown above.

Cervid phylogeny from Pitra et al. (2004). You should be able to see the Eld's deer + Milu clade near the middle of the diagram. Click to enlarge.

However, these general morphological similarities might represent convergence rather than genuine affinity. Pitra et al. (2004) used data from mitochondrial DNA to look anew at Old World deer phylogeny and were most surprised to find Eld’s deer to group closely with the Milu or Père David’s deer C. davidianus, with both forming a clade that is the sister-group to a larger cervine clade that includes sika, red deer and sambar. Similar results were reported by Gilbert et al. (2006). This discovery may or may not support the inclusion of both Eld’s deer and Père David’s deer within Cervus (as usual, that’s down to personal preference), but it does weaken the hypothesis that Eld’s deer is anything to do with Rucervus (which, following Pitra et al. (2004), I use here for the Barasingha and Schomburgk’s deer alone. Rucervus seemingly forms a clade with Axis and is outside of Cervus). The grouping of Eld’s deer with Père David’s deer also led to the curious idea that the latter might have arisen as a hybrid of Eld’s deer and a wapiti-type cervine – a subject that I covered in an article from last year devoted to Père David’s deer.

Sambar (Cervus unicolor), one of my favourite deer. Remind me to elaborate on that comment some time. Image by Wikigringo, licensed under the Creative Commons Attribution-Share Alike 2.5 Generic license.

Of course, this phylogenetic hypothesis isn’t the only one out there. Ouithavon et al. (2009) found Eld’s deer to be especially close to the Sambar Cunicolor (though note that their sample was limited, since they were only analysing taxa present in Thailand). Most recently, Zhang & Zhang (2012) found Eld’s deer to be the sister-taxon to a clade that included Père David’s deer as well as sambar, sika, red deer and so on.

There’s an important message to take from this quick look at what is now an obscure, little-known and endangered species, restricted to small populations and mostly famous for being weird. It’s that such obscure, endangered peculiarities might be the sorry relicts of what were once far more abundant, far more widespread populations. The data suggests that, just a few thousand or even few hundred years ago, Eld’s deer might have been a familiar, abundant and highly successful animal across an enormous area. Human impact on its diversity and distribution has been substantial.

Why did I write this article? Because obscure artiodactyls need more love, and I picked this species at random as one you don’t often hear about. Aiming to finish the petrel series soon, just haven’t yet found the time.

For previous Tet Zoo articles on deer, see…

Refs – -

Balakrishnan, C. N., Monfort, S. L., Gaur, A., Singh, L. & Sorenson, M. D. 2003. Phylogeography and conservation genetics of Eld’s deer (Cervus eldi). Molecular Ecology 12, 1-10.

Ellerman, J. R. & Morrison-Scott, T. C. S. 1951. Checklist of Palaearctic and Indian Mammals, 1758 to 1947. British Museum (Natural History) Trustees, London.

Geist, V. 1999. Deer of the World. Swan Hill Press, Shrewsbury.

Gilbert, C., Ropiquet, A. & Hassanin, A. 2006. Mitochondrial and nuclear phylogenies of Cervidae (Mammalia, Ruminantia): systematics, morphology, and biogeography. Molecular Phylogenetics and Evolution 40, 101-117.

Lekagul, B. & McNeely, J. A. 1977. Mammals of Thailand. Kurushpa Ladprao Press, Bankok, Thailand.

Nowak, R. M. 1999. Walker’s Mammals of the World, Sixth Edition. The Johns Hopkins University Press, Baltimore and London.

Ouithavon, K., Bhumpakphan, N, Denduangboripant, J., Siriaroonrat, B. & Trakulnaleamsai, S. 2009. An analysis of the phylogenetic relationship of Thai cervids inferred from nucleotide sequences of protein kinase C iota (PRKCI) intron. Kasetsart J. (Nat. Sci.) 43, 709-719.

Pitra, C., Fickel, J., Meijaard, E., & Groves, C. (2004). Evolution and phylogeny of old world deer Molecular Phylogenetics and Evolution, 33 (3), 880-895 DOI: 10.1016/j.ympev.2004.07.013

Pocock, R. I. 1943. The larger deer of British India. Part II. Journal of the Bombay Natural History Society 43, 553-572.

Zhang, W.-Q. & Zhang, M.-H. 2012. Phylogeny and evolution of Cervidae based on complete mitochondrial genomes. Genetics and Molecular Research 11, 628-635.

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 darrennaish.wordpress.com. He has been blogging at Tetrapod Zoology since 2006. Check out the Tet Zoo podcast at tetzoo.com! 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. Hydrarchos 10:35 am 04/19/2012

    So… according to that cladogram, fallow deer should also be included in Cervus if duvauceli and schomburgki are?

    Link to this
  2. 2. naishd 11:00 am 04/19/2012

    Pitra et al. (2004) regarded Rucervus (represented by duvauceli and schomburgki) and Dama as distinct from Cervus. Axis is not in Cervus either, but the sister-taxon to Rucervus. My text above wasn’t clear on this, so I’m making slight tweaks now, thanks for noticing.

    Darren

    Link to this
  3. 3. Bret Newton 11:52 am 04/19/2012

    Interesting that you make no mention of Groves and Grubb (2011) in which they resurrect Panolia and divide all thee subspecies into their own unique species. The resurrection of Panolia is due to the molecular clock used in Pitra et al. being set too early and the actual deviation is much further back. However they keep Rucervus as a subgenus of Cervus.

    Link to this
  4. 4. naishd 11:58 am 04/19/2012

    Aww, nuts!! I “make no mention” of it because I forgot about it and haven’t even seen it. Thanks for the heads-up. Groves & Grubb (2011) don’t just resurrect Panolia, they recognise P. eldi, P. siamensis and P. thamin as distinct species. The work is a book, by the way, not a paper.

    Groves, C. & Grubb, P. 2011. Ungulate Taxonomy. The John Hopkins University Press, Baltimore.

    Darren

    Link to this
  5. 5. Bret Newton 12:05 pm 04/19/2012

    Sorry correction, Rucervus is a valid genus, Rusa is not. Helps if I look at the material I am citing.

    Link to this
  6. 6. Bret Newton 12:06 pm 04/19/2012

    You haven’t seen it yet! Massive revisions galore. new genus names to learn. Fun late night reading material.

    Link to this
  7. 7. naishd 12:06 pm 04/19/2012

    Bret – do you own the volume? I really should get it… somehow.

    Darren

    Link to this
  8. 8. Bret Newton 12:11 pm 04/19/2012

    I do have the book. I’ve become a bit obsessed with ungulate taxonomy over the past few years (even before the book came out). It is the source material for the Bovidae section in the new Handbook of Mammals of the World.

    Link to this
  9. 9. Bret Newton 12:18 pm 04/19/2012

    Here is an overview of the changes from Brent Huffman’s site
    http://www.ultimateungulate.com/NewTaxonomy.html

    Link to this
  10. 10. David Marjanović 1:43 pm 04/19/2012

    Interesting how Cervus elaphus comes out as horribly diphyletic.

    Link to this
  11. 11. Bret Newton 1:54 pm 04/19/2012

    Cervus elaphus has been one of the biggest splits in the new Groves and Grubb with 12 species named. Three major branches in C. elaphus (senso lato) the primordial from central Asia, the Western from Europe, and the Eastern from east Asia and North America. The eastern isn’t even that close to the other two and is much closer to the Sika group.

    Cervus alashanicus (Alashan stag)
    Cervus bactrianus (Bactrian stag)
    Cervus canadensis (Asian/American wapiti)
    Cervus corsicanus (Barbary red deer)
    Cervus elaphus (West European red deer)
    Cervus hanglu (Kashmir stag)
    Cervus macneilli (Sichuan shou)
    Cervus maral (Turkish red deer)
    Cervus pannoniensis (East European red deer)
    Cervus wallichii (Tibetan shou)
    Cervus xanthopygus (Manchurian wapiti)
    Cervus yarkandensis (Yarkand stag)

    Link to this
  12. 12. kattatogaru 3:21 pm 04/19/2012

    1) Holy cow! the Moose’s closest living relative is the Korean Water deer??? Now that is truly bizarre.
    2) 12 species of red deer? An East European and a West European species? This sets alarm bells ringing…

    Link to this
  13. 13. naishd 6:27 pm 04/19/2012

    A major revision of the red deer-wapiti complex has been on the cards for a long time – the substantial variation (morphological, behavioural and genetic) present in these animals isn’t reflected in traditional taxonomy (note, for example, that various east Asian ‘red deer’ are very obviously ‘wapiti’, not close to Eurasian red deer). It’s predictable that people will throw up their arms and ridicule the idea that there might be more than ten species within this complex, but it’s not obviously unreasonable when you look both at phylogenies, and at the deer themselves. East European and West European red deer, for example, are pretty obviously very different and fossils show that they’ve been distinct for a reasonably long time – since the Middle Pleistocene (c. 200,000 yrs ago). I’m not saying that I support the new Groves & Grubb taxonomy in full, however.

    Darren

    Link to this
  14. 14. naishd 6:33 pm 04/19/2012

    Oh, kattatogaru (comment 12): moose aren’t most closely related to water deer (Hydropotes) in the Pitra et al. phylogeny, though they are close. That part of the phylogeny is best imagined as (Alces + (Hydropotes + Capreolus)). So, moose are close to the common ancestor of both water deer and roe deer.

    Darren

    Link to this
  15. 15. vdinets 7:00 pm 04/19/2012

    I haven’t seen the original G&G paper, but judging from the Bovids chapter in HMW, it should be simply ignored as a bad joke.

    Link to this
  16. 16. David Marjanović 7:13 pm 04/19/2012

    An East European and a West European species? This sets alarm bells ringing…

    Why? An east and a west European species that only come into contact in interglacials and otherwise live far apart occur in lots of other mammals, up in size at least to hedgehogs.

    Link to this
  17. 17. Bret Newton 7:59 pm 04/19/2012

    @vdinets, why do you say the bovidae chapter is a complete joke? After carefully reading G&G, I think most of the splits are well warranted. They address in the intro to the book that we have always assumed that large mammals have little variations across large regions but that small mammals are widely varied. Why should we assume that large mammals, just because we assume that they can more easily move over large distances, actually do.

    I’ve studied the African splits quite a bit and a striking pattern has emerged that there is a clear and distinct dividing line around the equator. Animals that were once thought to be a single species are now seen to be divergent from north to south. (The dividing line seems to be the Tana River in Kenya.)

    Link to this
  18. 18. Bret Newton 8:09 pm 04/19/2012

    @David Marjanović, spot on about the glaciation. Ludt et al. 2004 have a great map that shows the radiation of the red deer group and the western and eastern European deer and not closely related. Both had very different migration routes. Unfortunately, I cannot find a link to an open source copy of the paper, but the abstract in this link at least has a (mini) graphic of the map.

    Link to this
  19. 19. vdinets 9:53 pm 04/19/2012

    Bret: because it is arbitrary, totally inconsistent and in some cases outright fraudulent. There is no reason whatsoever for splitting all those hilltop populations of urial sheep, for example. In some cases they justify splits by showing non-existing gaps in distribution (i.e. in klippspringers). In other cases they lump well-identifiable subspecies (i. e. tule elk). I think their work is a perfect illustration of the mess that applying PSC invariably creates (although in some cases their splits don’t even seem justified under PSC, they are clearly CSC-based).

    As for African ungulates, many of those gaps are gaps between existing national parks, not natural ranges. Of course, PSC proponents could well argue that in today’s Africa every national park surrounded by unprotected lands is essentially an island, and all its large mammals constitute incipient species on separate evolutionary trajectories ;-)

    Link to this
  20. 20. vdinets 10:10 pm 04/19/2012

    Bret: and as for the small mammals vs. large mammals issue, it’s not a good argument. There’s plenty of small mammal species that are polymorphic and have wide distribution. If such species are more common among large mammals, that simply reflects the fact that large mammals tend to have better dispersal abilities than non-volant small ones. Besides, general considerations like this do not by themselves justify any particular splitting or lumping.

    Link to this
  21. 21. Bret Newton 10:14 pm 04/19/2012

    I won’t deny that some of the splits seem over bearing. I still cannot see how they split the Lesser Kudu into two species as the data they show doesn’t show any significant variation that I can tell.

    The north/south split in Africa though is well documented in both paleoclimatology and DNA evidence. There is on average a 1 to 1.5 million year separation between animals north of the equator and those south of it. This doesn’t hold true for all species like elephants and black rhinos, but it does for giraffes and white rhinos.

    G&G is not meant to be a final version, but rather a starting point for further discussion. They admit that some of the splits might be wrong, but more evidence is definitely needed in a lot of areas.

    Link to this
  22. 22. Bret Newton 10:19 pm 04/19/2012

    Can you cite a case where it is fraudulent? That’s a large claim.

    Link to this
  23. 23. vdinets 11:12 pm 04/19/2012

    Bret: splitting giraffes and white rhinos has been discussed for decades. I am talking about splits proposed by G&G with no justification except being well-defined subspecies.
    Aadmitting that the proposed splits are speculative didn’t prevent G&G from applying many of them to HMW chapter, rendering it unusable. I don’t know how the series editors let it pass, especially since the last volume of HBW was made very difficult to use by excessive conservatism in taxonomy.

    I just did: the klippspringers. G&G show non-existing gaps in the range (i. e. in Namibia) to justify the splits.

    Link to this
  24. 24. Heteromeles 11:44 pm 04/19/2012

    Well, the G&G treatment of Odocoileus is a marvel of brevity. Ahem!

    Link to this
  25. 25. vdinets 12:08 am 04/20/2012

    Heteromeles: they generally seem to loath North American taxa. They don’t even mention Fannin sheep. Of course, they also omit plenty of interesting info from other regions, like South Siberian reindeer having spotted fawns, but that’s a problem with HMW in general so far. For example, I couldn’t find any mention of adult ermine males inseminating blind female pups in nests, even though it has been well documented.

    Link to this
  26. 26. naishd 4:19 am 04/20/2012

    Interesting debate, thanks for the comments. As I always say in discussions of this sort, we have to remember that ‘species’ and ‘subspecies’ are more or less subjectively defined, man-made units – there is no consistency over what they really mean, even if authors do state a preference for the PSC or one of the other 100+ versions. Groves and Grubb have obviously succeeded in their plan to get people to take notice of things like the distinction between the two white rhino forms, the different forms of bushbuck and so on, but I can understand scepticism about their degree of splitting. On the other hand, one aspect of this debate that we keep coming back to (or, I keep coming back to, anyway) is that conservatism and poor logic has a strong influence on big mammal taxonomy.

    Vlad mentions spotted reindeer fawns (comment 25). Little known fact: some ancient cave art seems to show spotting in reindeer adults. Guthrie argues that this is accurate and that the animals really were spotted in life.

    Darren

    Link to this
  27. 27. kattatogaru 6:11 am 04/20/2012

    Darren – thanks for correcting me on the water deer/roe deer node. Holy cow (not included in this phylogeny even as a distant outgroup)! Moose are most closely related to water deer + roe deer? Still bizarre. I hadn’t realised how distantly related the Moose was to its antlered kin.
    So do west and eastern Red Deer not interbreed?

    Link to this
  28. 28. David Marjanović 8:08 am 04/20/2012

    but the abstract in this link at least has a (mini) graphic of the map.

    Oh, so the “western” European species isn’t form an Iberian refugium and isn’t confined to western Europe, and while the “eastern” European species is from a Balkan refugium as I assumed, it’s still confined to the Balkan and Pannonia. The “western” European red deer is straight from central Asia!

    That’s quite different from the situation seen in in hedgehogs or shrews. (Or, IIRC, brown bears or Neandertalers.)

    (I have access to the full paper.)

    they are clearly CSC-based

    Which species concept is the CSC? The “cladistic” one (which seems to simply mean “internode”)?

    Moose are most closely related to water deer + roe deer? Still bizarre.

    I don’t know if it’s all that strange that “water horse” and water roe end up close together.

    That said, it’s quite strange that the antlerless, fucking sabre-toothed water roe ends up well within the antlered, canineless-or-nearly-so clade.

    So do west and eastern Red Deer not interbreed?

    Here you’re asking for one of the “Biological Species Concept”s which, AFAIK, none of the cited works use.

    Link to this
  29. 29. vdinets 9:04 am 04/20/2012

    David: CSC is Conservation Species Concept ;-)

    kattatogaru: AFAIK, all red deer interbreed freely. In the former USSR, all herds in the European part have some blood of Siberian subspecies in them due to introductions “for size improvement”. It’s pretty obvious when you hear them during rut. I don’t remember if they screwed up the Caucasian subspecies as well.

    Link to this
  30. 30. Lars Dietz 9:10 am 04/20/2012

    “Ouithavon et al. (2009) found Eld’s deer to be especially close to the Sambar C. unicolor.”
    Yes, but they didn’t include Père David’s deer or anything from Cervus s. str. in their analysis, which only dealt with species that occur in Thailand.
    On cervid phylogeny, there’s also this paper based on multiple genes, which also shows the Eld’s/Père David’s sister group relationship. It also shows roe deer sister to water deer, but moose is sister to the reindeer/New World deer clade (but without strong support, so effectively in a trichotomy with this group and Hydropotes/Capreolus). One interesting thing: In the Pitra et al. paper, Axis porcinus groups inside Rusa, but in Gilbert et al. it groups with A. axis as traditionally assumed. Both have very strong support even in trees derived from the same gene (cytb). So the sequence used by Pitra et al. is probably misidentified.
    Here is another paper based on complete mitochondrial genomes that includes C. eldi, but unfortunately no Rucervus or Elaphurus, so it comes out as sister to Cervus s. str.

    Link to this
  31. 31. naishd 9:18 am 04/20/2012

    Lars: good call, I meant to mention the incomplete sampling in Ouithavon et al. (2009). As you note, it was specifically based on those species present in Thailand. Thanks for additional references.

    Darren

    Link to this
  32. 32. naishd 9:40 am 04/20/2012

    Have now updated the relevant section accordingly (and added a new picture!).

    Darren

    Link to this
  33. 33. Lars Dietz 10:00 am 04/20/2012

    On mitochondrial genomes, I missed this, which includes a much larger sample. Again, Eld’s deer is sister to Père David’s. Also some interesting conclusions in other parts of Cetartiodactyla.

    Link to this
  34. 34. Heteromeles 11:23 am 04/20/2012

    Oh dear, the penny just dropped. You’re using single-gene phylogenies for hybridizing populations, and trying to delineate species boundaries within these swarms? As the mechanics say, there’s your problem.

    Are there no nuclear markers? Over a decade ago, the botanists I knew all had moved to a three-gene standard (including at least one plastid gene and one nuclear marker) for phylogenies, precisely to deal with this kind of confusion. Obviously animals don’t have plastids, but if mitochondrial sequences work, that’s good enough.

    So what I’d say is, unless you’ve got three separate resolved genes and a consensus tree (preferably with several different analytic algorithms pointing to the same conclusion), these are all very preliminary results.

    Link to this
  35. 35. Riggi 11:28 pm 04/20/2012

    But isn’t Axis prove to not to be monophyletic? I mean if I recall a study in 2004 that the Hog Deer species were moved to Hyelaphus as they were related to Rusa. Also Milu and Eld’s Deer are better off as separate genera from Cervus. I did this not to long ago based on the studies I have read. http://titano911.deviantart.com/art/Cervidae-Phylogeny-202182777

    As for the Ungulate Taxonomy book, I have took a peek into it (never actually read it). The brockets were a surprise. I also remember that the Red Brocket were more closer to our Mule and White-tailed here, so I am curious to see where all of those species go. The only gripe I had was the needless splitting up (the biggest upset for me were how the equids were classified. The two Tarpan species should remain to be the same species since they reproduced fertile offspring with each other and to horses. And the Asian Ass split was unsuspected!).

    Link to this
  36. 36. vdinets 12:33 am 04/21/2012

    Riggi: white-tailed deer from Chiapas south look very Mazama-like. If Odocoileus gets lumped with Mazama someday, I wouldn’t be too surprised.

    Link to this
  37. 37. Dartian 1:51 am 04/21/2012

    Vladimir Dinets:
    I haven’t seen the original G&G paper

    It’s a book, not a paper. Darren already said that in comment #4.

    in some cases outright fraudulent

    Fraudulent? Are you saying that Groves and Grubb have falsified data? That’s a very, very serious accusation, buddy (and borders on slander). Either support it up with actual evidence, or take it back!

    splits proposed by G&G with no justification

    A few comments earlier, you said that you haven’t read their book. Then how the fuck do you know that there are no justifications for the splits?! You have a lot of nerve blaming others of doing poor science when you yourself consistently fail to adhere to even the most basic principles of scientific criticism. If you disagree with Groves & Grubb’s conclusions, you have to refute them with actual data – not with half-arsed assertions.

    I just did

    No, you did not. Until you provide actual evidence, it’s just another assertion.

    G&G show non-existing gaps in the range (i. e. in Namibia) to justify the splits.

    Very well then; please cite some peer-reviewed sources showing that klipspringer distribution and/or population genetic structure in Namibia is not discontinuous. The burden of evidence is on your shoulders here.

    Darren:
    Interesting debate

    Well, that’s one way to describe it I suppose.

    Personally, I find it disheartening that some individuals who really, really should know better still reflexively fall back to the Argument from Personal Incredulity whenever they are confronted with data that challenges traditional views (in this particular case, regarding artiodactyl systematics). Worse still, these same individuals even casually resort to outright claims of scientific fraud when trying to discredit results that they don’t like.

    Now, the question of whether some/most/all results from these recent taxonomic revisions of ungulates are valid or not is a legitimate topic of scientific discussion. But – as I’ve said before and will say again – only science can refute science. Groves and Grubb* have made their case. They may very well be wrong in some instances; in fact, they probably are. But anyone wishing to dipute their results must show them to be wrong. Going into denial mode isn’t enough.

    * As far as Peter Grubb is concerned, alas, Ungulate Taxonomy really was his final word on the subject; he passed away in 2006.

    Link to this
  38. 38. Dartian 2:05 am 04/21/2012

    Incidentally, regarding klipspringers; in Australia, rock wallabies Petrogale fill roughly the same econiche as klipspringers do in Africa. Currently, there are no fewer than 16 recognised species of them (according to the latest edition of Wilson & Reeder’s Mammal Species of the World) – and at least a couple of these are further split into subspecies that might yet be elevated to full species status. Funnily enough, nobody seems to be complaining that there are ‘too many’ species of rock wallaby. In this light, it’s strange how some people seem to have such a big problem with the idea that there might be 11 species of klipspringer*.

    * Considering how much bigger Africa – even Sub-Saharan Africa – is than Australia, one would, if anything, expect there to be considerably more species of klipspringers than there are rock wallabies.

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  39. 39. farandfew 6:51 am 04/21/2012

    Unfortunately this isn’t really about showing results to be wrong; it’s mostly about defining the question. As an ungulate conservationist, I can very much understand why tempers on the issue run high. The PSC, taxonomists in general and Colin Groves in particular are deeply unpopular in conservation circles right now, so far as I can tell, and there is a lot more behind that than simple stubborness.
    Seeing the Groves treatment of bovids in HMW made my heart sink. If the splits are simply false, then a lot of ink will be spilt and a lot of time wasted. But of course – whatever the status of klipspringers in Namibia – the real issue is defining the question, and that means even more ink. And all this time conservation decisions don’t get made and species, whatever they are, get lost. But far worse than that would be if the whole thing is unquestionably valid. Then, at a stroke, our workload is doubled, and our public support cut by an unknowable but potentially severe amount by the erosion of the public concept of a species, which is the one that really matters, unfortunately.
    One way or another, this book seems to me very bad news.

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  40. 40. Lars Dietz 8:14 am 04/21/2012

    “Are there no nuclear markers?”

    The Gilbert et al. study was based on two mitochondrial and two nuclear markers. They had only one C. elaphus s. l. sequence for each gene though, so they couldn’t test the monophyly of that “species”.

    “But isn’t Axis prove to not to be monophyletic? I mean if I recall a study in 2004 that the Hog Deer species were moved to Hyelaphus as they were related to Rusa. ”

    That was based on the misidentified sequence I mentioned above. Newer studies confirm the monophyly of Axis.

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  41. 41. David Marjanović 8:37 am 04/21/2012

    The two Tarpan species should remain to be the same species since they reproduced fertile offspring with each other and to horses.

    You’re assuming one of the BSCs.

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  42. 42. Riggi 11:51 am 04/21/2012

    vdinets: No meant where do all the “species” that “were” classified as Mazama americana go. I already the idea that the Red Brocket has been considered to be Odocoileus (I am actually surprised that they didn’t even attempted to split up the White-tailed Deer in numerous species. I am surprises there was no instance of merging taxon).

    David Marjanović: Yes I am, though as we all know there are quite some exceptionable examples.

    And will there ever be an ungulate book that will finally include cetaceans? I mean yeah I know they don’t have hooves, but I think it is just as important to show the glorious diversity of these guys. If dinosaur books can contain chapters on the evolution and avian taxon, then why not a book of hoof mammals that also contain such chapters.

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  43. 43. vdinets 12:03 pm 04/21/2012

    Dartian: in HMW, they give their reasons for splitting if they have any. Perhaps they have some reasons for particular splits and mention them in their own book, but not in HMW, but that only means that my criticism refers only to HMW. And I can’t understand why they would make such wild claims in HMW and not cite evidence if they had it.

    There are klippspringer populations in Namibia smack in the middle of the “gap between species” shown by G&G in HMW – for example, on Brandberg (you can easily find photos from Brandberg and other Namibian locations by Google). If G&G ignore that, they either intentionally omit contradictory data (and that is fraud), or show rather amazing failure to study the subject.

    It is still possible, of course, that there are many klippspringer species. But so far G&G haven’t demonstrated that. If there is some evidence in their book that is not in HMW, please let me know.

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  44. 44. vdinets 12:23 pm 04/21/2012

    Riggi: HMW hasn’t merged anything, but ignored a few well-defined subspecies. For example, it merged all American red deer into one subsp., but some of them are so distinct that they still had to be mentioned in the text on a few occasions :-)

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  45. 45. Riggi 5:25 pm 04/21/2012

    vdinets: No, I meant the Ungulate Taxonomy didn’t merge any taxon.

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  46. 46. vdinets 6:44 pm 04/21/2012

    Riggi: I guess I just have to get that book… Gonna take a while through interlibrary loan.

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  47. 47. Bret Newton 8:16 pm 04/21/2012

    I happen to have Ungulate Taxonomy with me right now. Here is the opening two paragraphs of the Oreotragus section.

    “As might be expected from their ecology, which is restricted to rocky outcrops, the distribution of klipspringers is very spotty, and there are quite distinctive species in different parts of the range. Some klipspringers are distinguished by different pelage features, but many cases the pelage is hardly different in widely separated species, and the degree and type of sexual dimorphism, as well as skull and tooth sizes, and more diagnostic.

    The degree and nature of sexual dimorphism in the different species appear to correlate with aspects of ecology and social behavior, particularly including the size of the pair territories, as listed by S. Craig Roberts (1996).”

    Work cited – Roberts, S.C. 1996. The evolution of hornedness in female ruminants. Behavior 133:399-442.

    As with most of the genera in UT, there are two pages of detailed measurements for all listed taxa. Incidentally, UT has 14 different taxa, 3 of which are unnamed in the literature.

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  48. 48. Bret Newton 8:19 pm 04/21/2012

    Riggi, Ungulate Taxonomy does indeed merge some taxa. Giraffe camelopardalis and Giraffa rothschildi have been merged.

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  49. 49. Bret Newton 8:27 pm 04/21/2012

    vdinets, As for the wapiti, little is said on the North American populations.

    “Schonewald (1994) examined differences among North American populations, finding that they were definable, but this needs more study.”

    Unfortunately, this is all that is said about the North American populations. It is lamentable that so little work has been done on North American ungulates (Cervus and Odocoileus).

    -Schonewald, C. 1994. Cervus canadensis and C. elaphus: North American subspecies and evaluation of clinial extremes. Acta Theriologica 39:431-452.

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  50. 50. llewelly 9:07 pm 04/21/2012

    I have a random question about the relatedness of perissodactyls and artiodactyls. Does the clade rooted at the MRCA of perissodactyls and artiodactyls include a lot of animals not traditionally considered ungulates? (Other than whales, that is; accept them as traditional ungulates for purposes of this question.)

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  51. 51. vdinets 11:45 pm 04/21/2012

    Bret: thank you! But these two giraffe taxa hadn’t been widely recognized as full species before G&G, so G&G simply didn’t accept a minority-supported split in this one case.
    As for klippspringers (hereafter KS), I am looking through the accounts in HMW, and the only “species” with a real difference in sexual dimorphism is the “Masai KS”, in which females have horns. For all others, there are notes like “female is larger, but there is a lot of variation”, and, indeed, given measurements show a lot of overlap. Then the authors say that for unstated reasons they choose to ignore Kingdon (1982), who said that not all Masai KS females have horns, and instead believe Roosevelt and Heller (1914), who say that they all do. Then the authors claim that Masai KS have much smaller territories than ALL other “species”, although elsewhere they admit that for some of those there’s no data, or, in case of “Stevenson’s KS”, the territories are also very small.
    I might be wrong here, but isn’t the degree of sexual dimorphism different between various dog breeds, human races, etc., etc.?

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  52. 52. Dartian 3:33 am 04/22/2012

    Thanks, Darren, for the clean-up! (Dear SciAm overlords, can we please get a ‘comment preview’ function?)

    farandfew:
    a lot of time wasted

    Mammalian systematics is a waste of time, is that what you’re saying?

    the real issue is defining the question

    I’m not entirely sure what you mean by “the question”, but if it’s about how to define species, please note that that’s precisely what Groves (in particular) has been consistently working with for about 40 years now.

    But far worse than that would be if the whole thing is unquestionably valid. Then, at a stroke, our workload is doubled

    Are you seriously suggesting that we should be suppressing scientific knowledge – or not doing science at all – just because the results are inconvenient (for you personally)?

    our public support cut by an unknowable but potentially severe amount by the erosion of the public concept of a species

    Please give some concrete examples of this having actually happened within the field of ungulate conservation. Also please explain what “public conception of a species” is.

    Vlad:
    I can’t understand why they would make such wild claims in HMW and not cite evidence if they had it.

    Handbook of the Mammals of the World is a secondary, not a primary source. Scientists don’t normally include raw biometric data in (semi-)popular books; that’s what technical books, papers, and monographs are for.

    There are klippspringer populations in Namibia smack in the middle of the “gap between species” shown by G&G in HMW – for example, on Brandberg (you can easily find photos from Brandberg and other Namibian locations by Google).

    I ask foor peer-reviewed sources and the best you can offer me is Google Images. Quelle surprise.

    And the presence of klipspringers (btw, in English – and, obviously, in Afrikaans – it’s klipspringer, not ‘klippspringer’) on Brandberg mountain doesn’t yet answer the question about (dis)continuity between Namibian klipspringer populations. Is the Brandberg klipspringer population actually in contact with (as opposed to reproductively isolated from) both of the two other klipspringer populations on the respective sides of the supposed distribution gap? Or is it in contact with just either one of them (and thus best considered part of that population)? Or is it perhaps isolated from both neighbouring populations and could therefore be regarded as – gasp! – yet another separate species?

    If there is some evidence in their book that is not in HMW, please let me know.

    First you make assertions that are unsupported by evidence, then you expect others to do the homework for you by looking up the data. Again, quelle surprise.

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  53. 53. David Marjanović 8:04 am 04/22/2012

    And will there ever be an ungulate book that will finally include cetaceans? I mean yeah I know they don’t have hooves, but I think it is just as important to show the glorious diversity of these guys. If dinosaur books can contain chapters on the evolution and avian taxon, then why not a book of hoof mammals that also contain such chapters.

    The “ungulates” don’t form a clade whether or not the whales are included.

    Does the clade rooted at the MRCA of perissodactyls and artiodactyls include a lot of animals not traditionally considered ungulates?

    Gene trees say that Perisso- and Artiodactyla could be sister-groups (together called Euungulata) or Perissodactyla + Ferae could be the sister-group of Artiodactyla. Most LINE insertions say Artiodactyla is the sister-group of Pegasoferae (bats, perissodactyls, Ferae), but all together show that incomplete lineage sorting must have happened. The radiation at the beginning of the Paleocene was, evidently, too quick for genetic drift to reduce diversity within populations between cladogeneses.

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  54. 54. David Marjanović 8:04 am 04/22/2012

    Perissodactyla + Ferae

    Together Zooamata. There’s a tradition of naming every single phylogenetic hypothesis in that field.

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  55. 55. Lars Dietz 8:49 am 04/22/2012

    “Does the clade rooted at the MRCA of perissodactyls and artiodactyls include a lot of animals not traditionally considered ungulates?”

    Actually, that’s not at all clear. The relationship between Chiroptera, Carnivora, Perissodactyla and Artiodactyla is very difficult to resolve. Hallström et al. (2011) analyzed over 1000 gene trees and found each of the 15 possible phylogenies supported by similar numbers of genes. Even the LINE insertions, which are very unlikely to be homoplastic, support mutually contradicting phylogenetic groupings with about the same support. So we’re looking at a very rapid divergence of the four groups, which probably cannot be resolved into a single bifurcating tree.
    Pangolins also belong into that clade, but they were not included in that paper as no pangolin genome has been sequenced. They’re probably sister to carnivorans, although I’ve also seen phylogenies that disagree with that.

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  56. 56. Lars Dietz 9:02 am 04/22/2012

    David: Your comments hadn’t appeared yet when I started writing mine. On LINE insertions, I suppose you’re referring to Nishihara et al. (2006), which found four insertions supporting Pegasoferae and one inconsistent with it. Of those found by Hallström et al., 3 support Pegasoferae and 5 are inconsistent with it and with each other.

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  57. 57. Dartian 9:21 am 04/22/2012

    Riggi:
    will there ever be an ungulate book that will finally include cetaceans?

    There is a chapter on the relationship between whales and terrestrial artiodactyls in The Evolution of Artiodactyls (edited by D.R. Prothero & S.E. Foss, The Johns Hopkins University press, 2007). There is also a basilosaur on the cover of that book. The rest of the volume, however, deals with the evolutionary history of terrestrial artiodactyls only.

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  58. 58. naishd 9:33 am 04/22/2012

    With reference to the recognition of klipspringer, giraffe and Eld’s deer populations (and populations of other large mammal forms) as taxonomically distinct ‘species’ or ‘subspecies’, one often hears the argument that this ‘splitting’ is dangerous. Policy-makers, politicians etc. will, so the argument goes, realise that the taxonomic entities we’re trying to protect are labile and recognised by whim and opinion more than hard science. What isn’t credited here is that (to paraphrase the late Peter Grubb) many traditional ‘species’ encompass a series of highly distinct units (‘subspecies’) that often require special protection relative to other units in the same ‘species’. Yet because ‘subspecies’ are typically regarded as less important than ‘species’, they get overlooked, despite the fact that ‘subspecies’ status is frequently a subjective one that was made lazily in an effort to tidy up the “too many species” problem seen to exist in the early decades of the 20th century.

    Genetics is now backing up data from morphology, behaviour and ecology that many supposed ‘subspecies’ are old, distinct lineages that have to be recognised when conservation strategies are considered. How best to bring attention to this fact, if not to name them as distinct species in their own right? Seriously, do they get the protection they need if they’re retained as ‘mere’ ‘subspecies’? Some previous thoughts on this issue are here on ver 2.

    Perhaps the biggest challenge is to get people to recognise that the boundaries between taxonomic entities are subjective and man-made, and best reflect whatever we regard as most useful in terms of communication.

    Darren

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  59. 59. naishd 9:42 am 04/22/2012

    For the record, here’s one of Grubb et al.’s relevant quotes on large mammal diversity/the recognition of species vs subspecies (from Grubb et al. 2000)…

    “Biodiversity of large mammals is severely underestimated. The existence of a narrow hybrid zone among large mammals can be detected in casual field surveys, which is not the case for small mammals and other animals that have to be trapped for close investigation. This simple fact has led to the downgrading of perfectly distinct, diagnosable species to a level where they become taxonomically ‘invisible’ and thus lost to biodiversity studies. There are many examples of large mammal genera in which single species are currently supposed to extend through forest and savannah zones (as in the elephant case treated here), and this series of case studies might be a place to start testing the proposition that their biodiversity has been underestimated” (p. 3).

    Darren

    Ref – -

    Grubb, P., Groves, C. P., Dudley, J. P. & Shoshani, J. 2000. Living African elephants belong to two species: Loxodonta africana (Blumencah, 1797) and Loxodonta cyclotis (Matschie, 1900). Elephant 2, 1-4.

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  60. 60. farandfew 11:34 am 04/22/2012

    Are you seriously suggesting that we should be suppressing scientific knowledge – or not doing science at all – just because the results are inconvenient (for you personally)?

    No of course not, and that sounds like a serious allegation to me :-) Fortunately neither I nor anybody else has the power to do this and so there is no point in worrying about it. I did not say that Ungulate Taxonomy ‘should be supressed’ I said it was bad news. I am very far from being the only person who thinks this; it certainly seems to be the dominant opinion among members of the IUCN-SSC antelope specialist group for a start.
    As it happens it is not immediately inconvenient for me personally because I work mainly on one species which has not been split and seems unlikely to be. I haven’t read the book, though I will do so when I get the chance and, like Vladimir, I have read the HMW bovids section. The introduction to the book is online on Google Books and this includes the species concept used and the justification for it. The justification is entirely stated in terms of what can be most precisely measured and defined (for this reason they also propose that genera are defined as taxa which split around the Miocene-Pliocene boundary). This is only half the story, however, and a much-used quotation comes to mind regarding precise measurements of the wrong thing. The other side of the story relates to the question of why we need to know.
    So here is my question, and it is not a rhetorical one: Why do we need to comprehensively identify, for all animals – or even all ungulates – the set of “smallest population, or aggregations of populations, which have fixed heritable differences from similar populations or aggregations of populations,”?
    The previous paragraph states that species are important because ‘they are the units of biodiversity’ but it seems to me that is only because we have decided that they are. Why is being divided into different populations so important? Why should disjunct diversity be more important than clinal diversity?
    Darren, it is quite true that ignoring existing diversity is going to lead to taxa getting overlooked and going extinct. But in a battle that we are currently losing very very badly we have to consider tactics. I have nothing against people trying to identify populations with fixed heritable differences from one another. However publishing the list of such units in a prominent popular book and stating that this is – more or less – the true list of species is very bad tactics for a conservation perspective in my opinion. This of course will need to be discussed and – again this is only my opinion – the discussion will be likely to be a waste of time because conservationists already spend far too much time thinking about what we want to conserve rather than how we are going to go about doing it.
    The real danger, I think, is that the idea of ‘species’ which has a fair amount of political force, will be eroded – not so much because people will consider species definitions to be labile but, more fundamentally, because it is a lot harder to make people care about lots of species which look the same to them. By public conception of species I just mean ‘what the average person tends to think that a species is’.
    Of course I can’t cite a peer-reviewed case study to show how erosion of the concept of species has caused a problem in ungulate conservation. That is rather like asking for a proven case of someone having been killed by climate change. In any case the book has only just been published so how can its results be felt?

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  61. 61. Heteromeles 12:28 pm 04/22/2012

    Personally, I have to agree with Darren, to the extent that every single taxonomist working with plants or vertebrates needs to sit through an entire legislative process involving the conservation of one of his or her favorite taxa. It’s a part of your education that most scientists ignore.

    The problem is pure ivory tower self-wanking. Your precious science (and I’m saying that sarcastically. My science used to be precious too) will be used as a political football by someone who doesn’t care whether your science lives or dies, or whether you get to be a systematist-with-tenure or a rat-decapitator-with-PhD in a biotech lab whilst struggling to pay your student loan debts.

    In other words, taxonomy is political. Sorry, that’s just the way it is, and I don’t like it any more than you do. In this case, ungulate species definitions will play directly into how and why various populations are saved or killed off. It’s the worst form of scientific solipsism to insist that you “only do pure science” and the rest of the world will take care of itself without you. By working on any organism, you’ve taken a side in someone’s debate, whether you want to be there or not.

    The nice thing about working with large ungulates is that most politicians at least get why something charismatic and valuable on the black market should be protected, if only as a randomly mixed herd in a game park somewhere. Entomologists, conchologists, mycologists, and most small-plant botanists have it much harder.

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  62. 62. vdinets 1:35 pm 04/22/2012

    Dartian: the question here is not whether KS have continuous or discontinuous distribution. HMW shows a HUGE gap between “Cape KS” and “Angolan KS” which doesn’t exist. Within this gap are some well-known KS populations, such as Brandberg, Augrabies, etc. If you want peer-reviewed sources, IUCN page on KS lists a few. Their map, BTW, shows extensive distribution of KS in Namibia. I think it’s impossible to study African ungulates and not be aware of KS’s presence in much of Namibia and northwestern South Africa. This is why I used the word “fraud”: it certainly looks to me like intentionally omitting inconvenient data.

    Darren: Are you seriously promoting Conservation Species Concept?

    Have anybody here heard of Preble’s meadow jumping mouse? Look it up. The landowners’ lobby challenged subspecific designation to have a subspecies stripped of endangered species status. Frivolous designation of populations in need of protection as full species would lead to widespread skepticism about the validity of such “science”, and ultimately to the erosion of public support for biodiversity conservation. We’d screw up scientific taxonomy and get nothing in return.

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  63. 63. Dartian 3:26 am 04/23/2012

    farandfew:
    species are important because ‘they are the units of biodiversity’ but it seems to me that is only because we have decided that they are

    Not quite. The reason why we focus so much on species is more than just adherence to a random convention; it is because ‘species’ is pretty much the only taxonomic category for which there is a definition (or, to be precise, several definitions). For other taxonomic categories – including ‘subspecies’ – there aren’t, at least not anything that’s widely used*. Granted, there is little or no agreement among taxonomists about which particular species definition should be used, but almost everybody at least agrees that species can, in principle, be defined.

    * As it happens, Colin Groves, in particular, has been at the forefront of applying definitions to taxonomic categories above species level, too. As you mention, he is a proponent of the idea that lineage divergence date should be the criterion by which extant taxa are allocated to genera, families, orders, etc.

    In any case the book has only just been published so how can its results be felt?

    There is a precendent of sorts to this exact situation. Back in 2001, Colin Groves (yes, he again) published Primate Taxonomy. In that book, he did to primates what he and Grubb now have done to artiodactyls: he dramatically increased the number of extant species, mainly by elevating former subspecies to full species status. Then, like now, there were loud protests from many quarters (IIRC, the phrase “excessive splitting” was commonly used). To some extent, there still are protests; primate taxonomy is every bit as contentious as ungulate taxonomy, and then some. But today, a decade after the book’s publication, a large proportion of primatologists seem to have come to terms with Groves’ classification, and happily use it. That’s not to say that all ‘splits’ have been widely accepted. (For example, proposals to elevate the various ‘subspecies’ of common chimpanzees to species status have, to this day, not been widely accepted; the majority of primatologists still prefer to call them subspecies.) But those splits that have by now been widely accepted are so numerous that I think it’s safe to say that Groves has permanently changed the ball game for primate taxonomy.

    More to the point regarding the specific concerns that you, farandfew, expressed: someone please correct me if I’m wrong, but AFAIK the conservation efforts of no primate species (or ‘subspecies’, if you prefer) have thus far been negatively affected by the publication of Groves’ Primate Taxonomy (and it’s been more than a decade since it was published, so any such effects should have started to show by now). Why, then, should we expect something like that to happen within the field of ungulate conservation?

    Vlad:
    the question here is not whether KS have continuous or discontinuous distribution

    Yes it is. Do Namibian klipspringers form one population, or do they form several? If the former, Groves and Grubb are probably wrong (regarding this particular detail, that is; whether you like it or not, it doesn’t affect the rest of their argument regarding klipspringer taxonomy across Africa). If the latter, we should be open-minded regarding the possibility that their hypothesis (viz. separate species status for various SW African klipspringers) is correct.

    HMW shows a HUGE gap between “Cape KS” and “Angolan KS” which doesn’t exist

    Klipspringers are exactly the kind of animals for which we should expect even fairly narrow gaps (so narrow that they might not show up in any but the most detailed maps) in distribution to have strong population genetic effects. Again, someone please correct me if I’m wrong, but klipspringer dispersal ability across open plains (or, say, rivers) is rather limited. In terms of speciation patterns, specialised rock- and mountain-dwelling animals such as klipspringers are effectively comparable to terrestrial animals living on islands.

    This is why I used the word “fraud”: it certainly looks to me like intentionally omitting inconvenient data.

    And the thought that it might be an earnest mistake didn’t occur to you?

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  64. 64. naishd 6:53 am 04/23/2012

    With reference to Dartian’s statement (comment 63) on “applying definitions to taxonomic categories above species level”, I just want to note that Sibley & Ahlquist came up with a similar idea (viz, that higher taxonomic levels should be recognised based on divergence dates).

    Darren

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  65. 65. Dartian 8:11 am 04/23/2012

    Darren:
    With reference to Dartian’s statement (comment 63) on “applying definitions to taxonomic categories above species level”, I just want to note that Sibley & Ahlquist came up with a similar idea (viz, that higher taxonomic levels should be recognised based on divergence dates).

    Another key reference dealing with this subject is:

    Avise, J.C. & Johns, G.C. 1999. Proposal for a standardized temporal scheme of biological classification for extant species. PNAS 96, 7358-7363.

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  66. 66. David Marjanović 8:55 am 04/23/2012

    On LINE insertions, I suppose you’re referring to Nishihara et al. (2006), which found four insertions supporting Pegasoferae and one inconsistent with it.

    Yes.

    Of those found by Hallström et al., 3 support Pegasoferae and 5 are inconsistent with it and with each other.

    :-o Wow. Please post the link again, the one you posted leads back to this page.

    In other words, taxonomy is political.

    Obvious solution: we must shout loud enough that the politicians begin to listen to us.

    With reference to Dartian’s statement (comment 63) on “applying definitions to taxonomic categories above species level”, I just want to note that Sibley & Ahlquist came up with a similar idea (viz, that higher taxonomic levels should be recognised based on divergence dates).

    So did Hennig till he abandoned it again later. Every once in a while someone has this idea, and then it turns out to require 1) the same rank for several nested named clades and 2) enormous upheavals to every existing ranked classification.

    To be fair, Groves & Grubb might escape the first and perhaps even the second problem for genera of large mammals if “around the Mio-/Pliocene boundary” is a short enough interval.

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  67. 67. Dartian 10:22 am 04/23/2012

    David:
    Every once in a while someone has this idea

    Actually, that idea never really went away, at least not in mammalogy, where it actually already has influenced taxonomy. A number of mammalian genera have recently been split on the basis that their extant members diverged such a long time ago. Examples include
    colugos, pangolins, tapirs and tarsiers; as recently as a couple of decades ago, all these mammalian ‘families’ would have been considered to consist of only one genus each – Cynocephalus, Manis, Tapirus, and Tarsius, respectively. Nowadays, however, more and more taxonomists have begun to split them into two or more genera:
    CynocephalusGaleopterus;
    ManisParamanis, Phataginus, Smutsia & Uromanis;
    TapirusAcrocodia & Tapirella; TarsiusCarlito & Cephalopachus.

    In none of the above cases has the recognition/resurrection of additional genus names been brought about by any serious doubts about the monophyly of these traditional ‘genera’. Nor have Cynocephalus, Manis, Tapirus, or Tarsius been unwieldy monster genera with ‘too many’ species; to the contrary, they each contained just a tiny handful of extant species*. Thus, the ‘standardisation’ of higher-level mammal taxonomy is an ongoing process, although it does so with relatively little fanfare.

    * Recently, however, it has been discovered that tarsiers, at least, include a large number of cryptic species. Nobody knows exactly how many extant tarsier species there may be, but it’s certain to be a two-digit number (Shekelle et al., 2010).

    Reference:
    Shekelle, M., Meier, R., Wahyu, I., Wirdateti & Ting, N. 2010. Molecular phylogenetics and chronometrics of Tarsiidae based on 12S mtDNA haplotypes: evidence for Miocene origins of crown tarsiers and numerous species within the Sulawesian clade. International Journal of Primatology 31, 1083-1106.

    Link to this
  68. 68. Lars Dietz 11:46 am 04/23/2012

    Here is the link to Hallström et al. again. I must have pasted the wrong URL in there, sorry for that. The LINEs supporting different splits are listed in Table 5. At least the situation is not as extreme as in Neoaves, where you have more than 10 groups diverging so rapidly that the retroposons support completely inconsistent splits, instead of just 4 or 5 as in this case.

    Link to this
  69. 69. Jerzy Again 12:05 pm 04/23/2012

    Big topic, Darren’s just one short quota about elephants contained several big mistakes. Just to sum up:

    - I don’t know how one comes with the idea that variability of large mammals is overlooked. It is only illusion of modern taxonomists who use secondary sources (inbred animals in zoos and limited study populations in tourist national parks). People in the field know it. Subspecies and races were named since decades, many by colonial hunters.

    - Even strangest species concepts must be possible to apply consistently. C&G concept is impossible to use even for all mammals – applying in to rodents and shrews would result in unmanageable 1000s of species.

    - Conservation species concept is nonsense. First, the purpose of using species was to put objective benchmark to prioritize efforts. Creating new species for conservation puts the idea on its head. Second, conservation objectives change. Example is the recent article advocating splitting White Rhino for two species to draw attention to the plight of cottoni. Unfortunately, cottoni numbers now 8 elderly individuals and the future is in crossing with simum – to keep genetic diversity, or simply because all cottoni females might be too old to breed. Of course, creating specific hybrids is more conservationally suspicious. Creating species for conservation here harmed the conservation – and the taxonomy-for-conservation ideology.

    - Modern taxonomy, if anything, slavishly follows genetic diversity and ignores morphological and ecological diversity of mammals if they don’t come with genetic diversity. Then for practical purposes (ecosystem role, utilitarian appeal) it is morphology and ecology which trumps genetics. This resulted is lumping North American elk subspecies, ignoring many old known subspecies and races of African elephants in favour of two-fold split Savanna-Forest etc.

    - Concept of non-hybridizing, independently evolving populations is now void. Habitat conversion destroyed most of genetic exchange between mammal populations. I agree with Vlad here, currently almost every national park in Africa has isolated population of giraffe, bushbuck, elephant etc. not hybridizing with other national parks. One look at the IUCN map reveals at least 42 reproductively isolated populations of giraffe.

    Link to this
  70. 70. vdinets 1:51 pm 04/23/2012

    Dartian: as I’ve already said, I don’t know if there is one species of KS or a hundred. But the HMW authors omitted a major portion of KS range to justify the proposed split. It is very difficult to believe that such a grave error could be made unintentionally.

    If KS have near-zero dispersal abilities, how did they colonize all those remote locations (including the ones in Nigeria that are hundreds of miles apart) in the first place? I am not aware of any data showing that Africa used to be one continuous rocky outcrop, except may be in Precambrian. And if they did colonize these places, perhaps their dispersal abilities are not as poor as they seem?

    Everybody: my personal opinion (yes, I know, nobody is asking for it) is that no species splitting proposals should be accepted for publication without solid genetic data. The only exceptions should be very clearcut cases where important new information has been obtained. That said, I would advocate an opposite approach to subspecies: phenotypically distinct ones should be recognized even if genetic difference is minimal, like in the case of wood bison.

    There is pro-splitting bias inherent in scientific taxonomy. Splitting proposals get more attention and are better for your career than lumping ones. Also, people working with any particular taxon tend to begin seeing it as undersplit related to those they are less familiar with. If we don’t counter this bias with proper – conservatism, skepticism, whatever you prefer to call it – we’ll end up having a runaway splitfest of the kind that took place in lepidopterology in Staudinger era. Ornithology and primatology are already pretty close to that.

    Link to this
  71. 71. naishd 3:49 pm 04/23/2012

    Thanks for continuing comments. There’s no doubt that the interplay between taxonomic revisions, species and subspecies definitions and conservation priorities will continue to be an area of disagreement, and – given that people aren’t going to stop proposing new phylogenetic and taxonomic hypotheses – we will probably have to find a middle ground. I’m a little unpleased with Jerzy’s statement (comment 69) that my comment “contained several big mistakes”, since he says several things that I never said.. plus, I was quoting Grubb et al. (2000). Note…

    Nobody said that “variability of large mammals is overlooked”. The actual quote (provided in comment 59) – from Grubb et al. (2000) – is that “Biodiversity of large mammals is severely underestimated”. Clearly, we have to deal with the taxonomy of any particular group of populations on a case by case basis. Genetic, behavioural and ecological data shows that some large mammals – lions and American bison are good examples – are relatively ‘plastic’, rapidly developing new morphs, like woodland bison and maneless, aridland Tsavo lions (all the evidence is against these forms being taxonomically valid, phylogenetically distinct entities). But this is not clearly the case across the board. The tradition that, say, all wolves are the same (as in, are all Canis lupus) because everyone knows they are ignores deep divergences between populations (extending back for 100s of 1000s of years) and ecological, behavioural and morphological differences that exceed those present between many other accepted ‘species’.

    And the idea that applying the concept of ‘phylogenetically distinct lineages’ (as per Groves & Grubb) will result in a proliferation of “unmanageable 1000s of species” is a familiar complaint, but is it really a logical one? It was made in the ornithological literature when data similarly showed that numerous ‘subspecies’ are, actually, often ‘better’ ‘species’ than many ‘species’ (Zink 1996, 2004). Anyway, nobody in the world can list, or know, all c. 5700 living mammal species… if, hypothetically, there were to be 20,000 ‘species’, I don’t see much difference – when do you decide that there are “too many species”? If you think there ARE “too many species”, how can you convince someone that this is anything other than your subjective preference? Anyway, this is a strawman argument – many of the ‘subspecies’ you’re referring to (that is, those within rodents, shrews and so on) are young clades within ‘species’. That is, they often or typically are not old, morphologically distinct lineages mirroring the bush and forest African elephant split, or the Indian and other wolves split, etc. etc.

    I have always said in debates on ‘splitting’, on conservation priorities and so on that the issue is confounded by the fact that (1) ‘species’ and ‘subspecies’ are ‘soft’, man-made entities, not fixed entities that we all agree on, and (2) people of all sorts need to admit, realise and communicate that taxonomy changes all the time, as we learn new stuff. If you think that species are unchanging, fixed entities that we should recognise universally no matter what, you are either a creationist or are deliberately ignoring scientific discussions about diversity, phylogeny and palaeontology.

    So – what best serves the living things we are trying to save? My statements in comment 58 (“How best to bring attention to this fact, if not to name them as distinct species in their own right? Seriously, do they get the protection they need if they’re retained as ‘mere’ ‘subspecies’?”) were meant to be rhetorical questions, not statements of fact. We increasingly talk about ‘saving the pieces’, so I don’t see that having a more accurate handle on phylogenetic diversity (by rightly recognising distinct phylogenetic units) is a bad thing. Surely this knowledge allows us to prioritise conservation efforts, but I do agree with Vlad (comment 70) that good data should be published before ‘splitting’ proposals are accepted. That has happened in a few of the cases concerned (elephants, giraffes, wolves), but maybe not enough of them. And the ‘received/conventional wisdom’ that, say, all giraffes are ‘just giraffes’, differing only in coat colour and pattern form, has definitely obscured conservation priority in the past.

    I personally don’t know that workers like Groves and Grubb have ever ‘gone too far’ in proposing all these new splits, but I do agree that the best, strongest case should be made for recognising the lineages concerned before they become accepted. Are the splits a bad thing for conservation? They mean that we have to consider ever more populations as worthy areas of time and effort – as if resources weren’t stretched enough – but didn’t we already know that? There are too many organisms to save already, sad to say, and we are losing the battle.

    Darren

    Refs – -

    Zink, R. M. 1996. Bird species diversity. Nature 381, 566.

    - . 2004. The role of subspecies in obscuring avian biological diversity and misleading conservation policy. Proceedings of the Royal Society of London B 271, 561-564.

    Link to this
  72. 72. Jerzy Again 6:30 pm 04/23/2012

    @Darren
    About mistakes, I was referring to the quote from #59

    “Biodiversity of large mammals is severely underestimated.”
    - Well, 19. century zoologists named lots of species. And I am not sure about Authors knowledge, but they are certainly avialable in libraries.

    “The existence of a narrow hybrid zone” “as in the elephant case treated here”
    - Hybrid zone between savanna and forest elephants spans Uganda and eastern part of Congo. It is not narrow, by area it is larger than ranges of many species!

    “perfectly distinct, diagnosable species”
    - Groves approach that a species can have one diagnostic difference AND a can have hybrid zone is a mystery for me. Diagnostic by definition cannot be shared in hybrids.

    “single species are currently supposed to extend through forest and savannah zones”
    Flexible habitat choice is normal among large animals. Is there anything wrong that many European and North American large mammals occur both in tundra and forest zone, or both forest and steppe zone?
    Besides – isn’t it thinking wrong way first? Zoology is to recognize taxa by their genetic characteristics, and then look at their life including habitat choice. G&G seem to think opposite way.

    As you see, there is a bit too many false assumptions and use of existing well-established concepts in the wrong sense to comment individually.

    Link to this
  73. 73. Jerzy Again 6:44 pm 04/23/2012

    BTW, some more general points:

    - That Red Deer and Wapiti are different on about species level and the split should go via C Asia is well known. See eg. Walker’s Mammals of The World from 1980s. The same large genetic difference between White Rhinos etc. Groves et al. possibly added some characters and extended discussion, but the general difference between beasts was well known.

    - This so-called new taxonomy is not really new. It is simply return to the 19. century narrow species concepts with modernized evaluation.

    - Which means, that it will be abandoned. 19. century narrow species were abandoned on practical grounds, so 21. century narrow species of Groves will also be abandoned on the same grounds: unusable, in the longer run creates more problems than solves – like Europeans having to cope with Eastern and Western red deer, or primatologists angry with several proposed common chimpanzee species.

    - Splitting creates lots of hybrids. Taxonomy copes poorly with hybrids. And where hybrid zones became large part of the population, lots of unanswered questions arise: are they of conservation value? Should we prevent extinction of pure species if large hybrid population remains or vice versa? How to put them in computer databases? etc.

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  74. 74. Jerzy Again 6:53 pm 04/23/2012

    #71
    Darren, it is conservationists who decided that only species are evaluated for conservation, because of no resources to protect all subspecies.

    If you change subspecies of species, the problem doesn’t go – conservationists probably will be forced to evaluate only full genera or similar.

    Link to this
  75. 75. vdinets 11:07 pm 04/23/2012

    Actually, in many countries biodiversity conservation laws refer equally to subspecies and species. Perhaps it’s easier to change laws in other countries than to change taxonomy?

    Link to this
  76. 76. David Marjanović 6:22 am 04/24/2012

    Nowadays, however, more and more taxonomists have begun to split them into two or more genera:

    Wow. I had no idea.

    Here is the link to Hallström et al. again.

    Thanks, I’ll check it out ASAP.

    - Even strangest species concepts must be possible to apply consistently. C&G concept is impossible to use even for all mammals – applying in to rodents and shrews would result in unmanageable 1000s of species.

    So?

    “Reality is inconvenient” is not an argument.

    the purpose of using species was to put objective benchmark to prioritize efforts

    :-D :-D :-D :-D :-D

    Really, this belongs on Failblog!

    There is pro-splitting bias inherent in scientific taxonomy. Splitting proposals get more attention and are better for your career than lumping ones.

    Show me.

    The tradition that, say, all wolves are the same (as in, are all Canis lupus) because everyone knows they are ignores deep divergences between populations (extending back for 100s of 1000s of years) and ecological, behavioural and morphological differences that exceed those present between many other accepted ‘species’.

    I think this kind of lumping was caused by the Biological Species Concepts. All wolves can interbreed, all the way to the coyote, right?

    “Biodiversity of large mammals is severely underestimated.”
    - Well, 19. century zoologists named lots of species.

    It is severely underestimated now, even though maybe it was not underestimated 120 years ago.

    Diagnostic by definition cannot be shared in hybrids.

    ???

    Wouldn’t you expect every hybrid to have about half of the diagnostic characteristics of each parent species?

    - This so-called new taxonomy is not really new. It is simply return to the 19. century narrow species concepts with modernized evaluation.

    That’s completely false, and you know it.

    Europeans having to cope with Eastern and Western red deer

    Cry me a river.

    Splitting creates lots of hybrids. Taxonomy copes poorly with hybrids.

    Whose problem is that exactly?

    And where hybrid zones became large part of the population, lots of unanswered questions arise: are they of conservation value? Should we prevent extinction of pure species if large hybrid population remains or vice versa? How to put them in computer databases? etc.

    You’re not seriously proposing to define this problem out of existence instead of solving it. You can’t be that intellectually lazy.

    no resources to protect all subspecies

    You know, that’s just a lie. It’s not money that’s lacking, it’s political will. Look at the military budget of any country bigger than Costa Rica (which abolished the military after its last civil war), and tell me there’s not enough money for whatever.

    We scientists need to shout louder.

    Link to this
  77. 77. farandfew 1:05 pm 04/24/2012

    “We scientists need to shout louder”

    I thought that was a joke when you said it before. Now I’m not so sure. You seem to be making the assumption that political will is somehow easier to generate than money. I suppose I could ask you for a peer-reviewed study to demonstrate the fact. Instead I will make the unverifiable assertion that, while shouting attracts people’s attention, it doesn’t keep it.
    It also tends to make people angry.
    If we want people to pay more attention to conservation, shouting louder is probably about the worst thing we can do right now, IMO.

    Adopting the Phylogenetic Species Concept would not be quite as bad as that, but still a pretty bad idea. IMO again.

    Dartian.

    Not quite. The reason why we focus so much on species is more than just adherence to a random convention; it is because ‘species’ is pretty much the only taxonomic category for which there is a definition

    But it is a random convention to adopt the only taxonomic category for which there is a non-arbitrary definition as the unit of biodiversity.

    There is no reason why these units have to be taxonomic categories at all. There is no reason, if they are taxonomic categories, why they can’t be arbitrarily defined. I believe they ought to be consistently defined, but really that’s just a personal preference as well.
    Is a behaviourally distinctive lion population less valuable than a behaviourally average wolf population simply because it is not phylogenetically definable? It’s arguable, but it needs to be argued.

    To go back to Colin Groves’ introduction to his book the argument runs as follows
    1) species are important because they are ‘the units of biodiversity’
    2) the set of “smallest population, or aggregations of populations, which have fixed heritable differences from similar populations or aggregations of populations,” is the most consistently applicable definition possible for a taxonomic category.
    3) therefore this should be used as the definition of a species.

    To summarise my summary: Because we can measure this thing, this thing is what we should measure. An argument which is a clear sign of illogical decision-making.

    I am making an assumption, and perhaps it is an incorrect one, that the main people who need ‘units of biodiversity’ (or think they do) are conservationists.

    Some conservationists will, like Darren, think that the Phylogenetic Species make good units of biodiversity because we want to make sure nothing gets overlooked. Others would agree with me that to do so risks stretching our already overstretched political capital much further and will do more harm than good.

    I don’t know who is right, and I don’t think it is possible to know for sure. Certainly it isn’t possible to conclusively demonstrate a negative effect on primate conservation from Colin Groves’ earlier book. Such an effect would be systemic and indistinguishable from confounding variables and could only perhaps be demonstrated (though not quantified) with a carefully designed social study which has not been done. The greatest danger, I think, comes from the potential to overturn laws which refer to species but which were formed under a different shared understanding of what the word meant. This would happen suddenly or not at all.

    But, although I don’t know I’m right, the point is that there is a debate here to be had. It is perfectly appropriate for Groves and Grubb, as taxonomists, to attempt to classify ungulates into sets of populations with distinct, fixed heritable differences. It is perfectly appropriate for Colin Groves to demonstrate by argument why this way of dividing up the ungulate biota of the world is objectively verifiable – or more so than other methods
    But it is wholly inappropriate, in my view, to go on to say that this is the list of species which everybody should use.
    I think it would be perfectly appropriate for IUCN to say ‘this may very well be true but we don’t happen to be interested. We are going to define a species as a lineage which diverged from its sister group between x and y mya.’ Of course they never will say that because they don’t want to admit that what we choose to conserve is arbitrary. But it is, because choice is arbitrary.

    Therefore Heteromeles is right that it is a political question. It is a question about who owns the word ‘species’. Is a species an objectively definable clade (or the smallest definable clade) or is it a unit of biodiversity? Because there is absolutely no reason why those have to be the same thing.

    If you want to demonstrate that your work is important to other people it seems like it might be a good idea to ask some of the people who you expect to use it what they want out of you rather than taking it entirely on yourself to tell them what they ought to want.

    Colin Groves’ justification seems to me to identify conservationists as the primary intended users of species concepts because I’m assuming conservationists are the people who are going to want ‘units of biodiversity’.

    Now I recently heard an eminent professor of conservation science complain about this particular work, and about the PSC in general. When I asked what the appropriate response of the conservation community should be, he replied that we simply have to be grateful that there aren’t many taxonomists left.

    This is not about wanting to keep things as they are just because we are comfortable with them. It is not about that at all.

    Link to this
  78. 78. vdinets 2:46 pm 04/24/2012

    David: it has been shown years ago that Northern Parula is but one ssp. (and not the most distinct one) of Tropical Parula. The paper sunk into oblivion without a trace. Compare it with regular “exciting upcoming splits rumors” posts on birdwatching forums :-)

    Any proposal to systematically increase the number of species in Mammalia using some new criteria should include some kind of a limit, or splitting can go on indefinitely. I propose a simple criteria: any new species concept, if applied consistently, should not require splitting extant populations of Homo sapiens. Under the approach currently fashionable in primatology, H. sapiens has to be split into dozens, if not hundreds, of species. I have mentioned here already that H. sapiens populations differ very substantially in size, color, cranial measurements, facial features, behavior and habitat preferences. There are also differences in anatomy (i.e. knee joint structure) and physiology; some behavioral differences have also been shown to be innate. There is also obvious suppression of hybridization (in all countries the percentage of interracial marriages is lower than expected if mating was completely random).

    Dartian: I don’t like the idea of setting % of genetic difference as the sole criteria for assigning taxonomic rank to above-species taxa. Not all DNA differences are equal. Some create huge differences in phenotype, while others don’t show at all. Humans and chimps are rather similar genetically, but they have countless important differences in body plan, anatomy and physiology, not to mention behavior. On the other hand, tarsiers are all very similar despite being different genetically. Do we want taxonomic rank to be a simple measure of differences in DNA sequences (most of which are totally irrelevant for phenotype), or correspond to some important differences? I would vote for the latter. I know, that creates subjectivity, but I think it makes classification more informative and meaningful.

    Of course, using only phenotype would be completely wrong: we would have to assign dog breeds to different genera. I think the best approach would be a combined one.

    I also don’t like the idea of using time of separation as the sole criteria. Estimating it using molecular clock is prone to huge errors. Estimating it using the fossil record has another problem: we risk accepting earlier separation dates for taxa with more complete fossil record.

    That said, I don’t oppose splitting small monophyletic groups in principle. Personally, I’d split Rhynchocyon into a separate family, and platypus into a separate order, for example. With placentals and marsupials already split, it’s unfair to discriminate against the monotremes :-)

    farandfew: isn’t it easier to make subspecies the primary unit for conservation than to screw up the entire taxonomy for short-term (and very uncertain) conservation benefits?

    Link to this
  79. 79. Dartian 2:33 am 04/25/2012

    Jerzy:
    It is only illusion of modern taxonomists who use secondary sources (inbred animals in zoos and limited study populations in tourist national parks). People in the field know it.

    If that was supposed to be a cheap shot aimed at Peter Grubb and Colin Groves, it misses the mark completely. They have both done plenty of field work, especially Groves: he has studied, among other things, mountain gorillas in Rwanda, colobus monkeys and mangabeys in Kenya and Tanzania, macaques in Indonesia, and various species of artiodactyls in India, Iran and Arabia. He is also one of the extremely few human beings on this planet who has been charged by a wild Javan rhinoceros (this happened when Groves was doing field work in Java in Udjung Kulon National Park, in 1978). In other words, Groves’ ‘macho field biologist’ credentials are, by any reasonable standard, impeccable.

    Even strangest species concepts must be possible to apply consistently. C&G concept is impossible to use even for all mammals – applying in to rodents and shrews would result in unmanageable 1000s of species.

    First you say that species concepts should be applied consistently; then, in the very next sentence, you are unhappy with the prospect that the species concepts will be applied consistently across Mammalia. Do you even notice the contradiction in what you say?

    Besides, if you’re not a practising mammalogist yourself, why would you even care about how many species of mammals are recognised? Do you lose sleep over the fact that arachnologists (currently) recognise about 40,000 species of spider (a far greater number than even the most extreme splitting of extant mammals could ever result in)?

    Vlad:
    If KS have near-zero dispersal abilities

    Do not distort what I said. What I actually said was: “someone please correct me if I’m wrong, but klipspringer dispersal ability across open plains (or, say, rivers) is rather limited“. “Limited dispersal abilities” =/= “near-zero dispersal abilites”. (I also admitted there that I do not know for sure just how limited klipspringer dispersal abilities are.)

    farandfew:
    it is a random convention to adopt the only taxonomic category for which there is a non-arbitrary definition as the unit of biodiversity

    Say what? If we don’t use objective criteria (or as nearly objective as is reasonably possible), we aren’t doing science – we are merely stamp-collecting. Your line of reasoning is strange in general, but the fact that you show up on a science blog, of all places, to make that argument is downright bizarre.

    it is wholly inappropriate, in my view, to go on to say that this is the list of species which everybody should use

    Bollocks! Groves and Grubb do not say that. Here are exact quotes from the very first page of the Introduction of Ungulate Taxonomy: “We do not intend to present this book as a finished proposition.” and “[W]e hope that these examples will be examined by colleagues and, if found acceptable, used as templates for other studies.

    he replied that we simply have to be grateful that there aren’t many taxonomists left

    Well, he’s entitled to his opinion (even if it’s a grotesquely arrogant and astonishingly idiotic one).

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  80. 80. vdinets 3:44 am 04/25/2012

    Dartian: ok, you didn’t say that, sorry. But my point remains valid: if KS have reached those isolated areas once, they can do it again (unless there is data showing that it was easier in the past), so we cannot postulate lack of gene flow without proving it. And even if we can prove lack of gene flow, we can never be sure that it will not become possible again if the conditions change.

    Link to this
  81. 81. Dartian 3:58 am 04/25/2012

    #79:
    Here are exact quotes from the very first page of the Introduction of Ungulate Taxonomy”

    Slight correction to that; the quotes are from the Preface, not from the Introduction.

    Jerzy, #73:
    primatologists angry with several proposed common chimpanzee species

    Who is angry? Virtually everyone who works with common* chimpanzees nowadays agrees that there are different taxa within this ‘species’; that isn’t in doubt. It’s just that, at least as of today, there has been no widespread tendency to start treating the three (some say four) currently recognised taxa as full species; most people prefer still to consider them ‘subspecies’. (For what it’s worth, I am personally perfectly happy with that.) History shows us, however, that consensus may yet shift; nowadays the majority of workers accept that there are 2 species of orangutans and 2-3 species of gorillas – but that acceptance didn’t come about until quite recently and after much resistance. Perhaps the same shift in thinking will eventually happen with chimpanzees too?

    * Side note regarding the chimpanzees’ vernacular names: Many people have been unhappy with the name ‘pygmy chimpanzee’ for Pan paniscus, as it isn’t really all that much smaller in size than Pan troglodytes (sensu lato). For various reasons, not everyone has been happy with ‘bonobo’, either. Therefore it has been suggested (originally, AFAIK, by Frances White in 1996) that more appropriate vernacular names in English for Pan paniscus and Pan troglodytes would be ‘gracile chimpanzee’ and ‘robust chimpanzee’, respectively. (Cf. gracile vs. robust australopithecines.)

    Reference:
    White, F.J. 1996. Pan paniscus 1973 to 1996: twenty-three years of field research. Evolutionary Anthropology 5, 11-17.

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  82. 82. Jerzy Again 7:33 am 04/25/2012

    @argument that elevating subspecies by splitting only helps conservation.

    I recall you EDGE inititative – Evolutionary Distinct and Globally Endangered – which Darren reviewed som time ago. Zoologists from ZSL state clearly that protection of evolutionary distinct animals suffers precisely because of “drowning” among poorly differentiated species. So not all species are equal.

    So survival of saiga, orangutan, solenodon, giant salamanders etc. is obfuscated by Rana frogs, murid rodents, Cervus deer, Microcebus mouse lemurs etc.

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  83. 83. Jerzy Again 7:42 am 04/25/2012

    BTW, Vdintes has a point. If one assigns species rank to marmosets which differ by coloration and ear tufts with minimal genetic differences, it is easy to justify that bearded white humans and curly-haired black humans are not one species.

    Primatologists note that two marmosets coming into contact hybridize less frequently than one displaces another. But isn’t it what happened to Europeans and American Indians?

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  84. 84. Christopher Taylor 8:48 am 04/25/2012

    C&G concept is impossible to use even for all mammals – applying in to rodents and shrews would result in unmanageable 1000s of species

    Not, it should be noted, a foregone conclusion. My impression is that ‘subspecies’ have been used less often for small mammals; if so, it is possible that recognised species are already close to what would be recognised by the more restricted species concepts. It would need someone to actually do the research to know.

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  85. 85. David Marjanović 10:02 am 04/25/2012

    You seem to be making the assumption that political will is somehow easier to generate than money.

    What? Why? I’m not making that assumption at all.

    I’m saying we should shout louder – because it’s the right thing to do.

    while shouting attracts people’s attention, it doesn’t keep it.
    It also tends to make people angry.
    If we want people to pay more attention to conservation, shouting louder is probably about the worst thing we can do right now, IMO.

    *eyeroll* I don’t mean literal shouting…

    There is no reason why these units have to be taxonomic categories at all. There is no reason, if they are taxonomic categories, why they can’t be arbitrarily defined. I believe they ought to be consistently defined, but really that’s just a personal preference as well.
    Is a behaviourally distinctive lion population less valuable than a behaviourally average wolf population simply because it is not phylogenetically definable? It’s arguable, but it needs to be argued.

    With this I agree!

    Colin Groves’ justification seems to me to identify conservationists as the primary intended users of species concepts because I’m assuming conservationists are the people who are going to want ‘units of biodiversity’.

    …Does it even make sense to say that biodiversity comes in discrete units?

    David: it has been shown years ago that Northern Parula is but one ssp. (and not the most distinct one) of Tropical Parula. The paper sunk into oblivion without a trace. Compare it with regular “exciting upcoming splits rumors” posts on birdwatching forums :-)

    Is that one example all you have? Where was that paper published – was it published in a widely accessible journal? And who, except birdwatchers, cares about birdwatching forums…?

    Under the approach currently fashionable in primatology, H. sapiens has to be split into dozens, if not hundreds, of species.

    We’ve been through this discussion…

    There is also obvious suppression of hybridization (in all countries the percentage of interracial marriages is lower than expected if mating was completely random).

    How about the Cape Verde Islands? Aren’t they panmictic or nearly so?

    (You can find people with very dark skin and curly hair that is light blond there. You name the combination, it’s there.)

    For various reasons, not everyone has been happy with ‘bonobo’, either.

    Why?

    Primatologists note that two marmosets coming into contact hybridize less frequently than one displaces another. But isn’t it what happened to Europeans and American Indians?

    How much culture is there among marmosets?

    My impression is that ‘subspecies’ have been used less often for small mammals

    Very good point.

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  86. 86. vdinets 1:14 pm 04/25/2012

    David: no, it’s not the only example. The situation is similar with gulls (for example, Western and Glaucous-winged are still treated as full spp. despite overwhelming evidence to the contrary), rosy-finches, etc., etc. The Parula paper was in Auk (118(1):211-215, 2001, by Lovette and Bermingham). At higher level, look at Suboscine families (antbrids were split, but woodcreepers and cotingas are still around), nine-primaried Oscines, etc.

    Pretty much all ornithologists today are also birdwatchers, and there is no clearcut difference between birdwatching and ornithological forums.

    Do you have any data showing that skin color is totally unimportant in the Azores, and mating is completely random? I very much doubt it. But if it is true, it only means that it’s a hybrid swarm. Such swarms can form even between species, and even if mating is not completely random.

    What does culture have to do with it? There was a recent study showing that even little kids take skin color into account when choosing friends. Also, it is very likely that marmozets learn what a “right” face looks like after birth. BTW, I am not aware of any data showing that marmozet races (or species if you prefer) differing only in facial features don’t interbreed given a chance.

    It is not at all obvious that subspecies has been used more often for large mammals. Deer mouse, for example, has the highest number of subspecies among North American mammals. Note that non-volant small mammals on average have smaller ranges due to limited dispersal abilities compared to large mammals.

    My understanding is that “bonobo” is a fake African word not known to come from any particular language. But so is “kangaroo”. Personally I’d prefer to keep existing common names for Pan species rather than replace them with longer ones. But if we don’t, we should also replace “kangaroo” with “superwallaby”.

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  87. 87. Christopher Taylor 8:37 pm 04/25/2012

    My understanding is that “bonobo” is a fake African word not known to come from any particular language. But so is “kangaroo”.

    The story about the inappropriate origin of the word ‘kangaroo’ is something of an urban legend. ‘Kangaroo’ was indeed an Aboriginal name for the animal, but only in the Guugu-Yimidhirr language. When British settlers later used the name when talking to Aboriginals who did not themselves speak Guugu-Yimidhirr, they naturally did not know to what the word referred.

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  88. 88. vdinets 10:29 pm 04/25/2012

    Christopher: thanks for the correction.
    Incidentally, it appears that “wapiti” is not a real Amerindian name for C. canadensis: it means simply “white rump” in some languages.

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  89. 89. Dartian 4:06 am 04/26/2012

    Vlad:
    we cannot postulate lack of gene flow without proving it

    Of course we can postulate it; that there is a nice, falsifiable hypothesis (that is, significant gene flow between given Namibian klipspringer populations either does take place, or it does not; either way, it is something testable).

    However, by analogy of the Australian rock wallabies that I mentioned earlier, it is only reasonable to expect there to be some non-trivial amounts of genetic differences between klipspringer populations. Petrogale wallabies have been quite extensively studied, and it has been shown that their dispersal abilities across suboptimal habitat are indeed quite limited – and that there may therefore be notable genetic differences between populations living, in some cases, less than one kilometer apart (e.g., Piggott et al., 2006). (You’d be hard pressed to find a general distribution map for these macropodids that identifies distribution ‘gaps’ that narrow.) Other studies (e.g., Potter et al., 2012) have shown that some currently recognised Petrogale ‘species’ contain sub-populations that seem to have diverged in, and been reproductively isolated from each other since, the early Pleistocene or even the Pliocene (and that, among other things, even fairly small rivers may strongly influence their distribution). Furthermore, Potter et al.´s (2012) data also suggest that at least one currently recognised taxon, the short-eared rock wallaby Petrogale brachyotis, is paraphyletic with respect to not one but two other species (the monjon Petrogale burbridgei and the nabarlek Petrogale concinna, respectively). In other words, this means that the current estimate of 16 Petrogale species that I gave in comment #38 is likely to change; more species-level splits are a-coming, lads! ;)

    David:
    Why?

    Ah, that’s a long, sordid story. I don’t think I can do it full justice, but it all traces back to the 1930′ies, when three bonobos (I shall preferentially call them that) were shipped to Tierpark Hellabrunn in Munich, Germany. The crate that the apes were held in during transportation had – so the story goes – the word ‘Bolobo’ on it. Bolobo is a city in what was then Belgian Congo, later Zaire/Democratic Republic of Congo, but the zoo director, Heinz Heck, somehow came to believe – again, so the story goes – that it was the indigenous name of the apes, and that he further twisted it into the form ‘bonobo’ instead of ‘bolobo’. In other words, ‘bonobo’ is, like Vlad said, a nonsense word that’s not actually used by any indigenous Congolese ethnic group.

    Anyway, the bonobos were introduced to Hellabrunn’s (common) chimpanzee colony, where it quickly became obvious that the former were quite different from the latter. (For example, the two female bonobos were terrified by the male chimpanzees’ boisterous pre-mating displays and fled from them in terror. The male bonobo, on the other hand, tried to copulate with female chimpanzees by ‘skipping the foreplay’ altogether; a behaviour which, unsurprisingly perhaps, wasn’t appreciated by the female chimps. Soon enough, the bonobos were separated from the chimpanzees and kept by themselves, and their behaviour observed by Heck. Sadly, during the Second World War, all three bonobos died during air raids; not because the zoo was hit by bombs, but, according to Heck, because the noise and the explosions distressed the poor bonobos to death. By contrast, none of Hellabrunn’s common chimpanzees died from such bomb-caused emotional trauma.) After the war, Heck, together with Eduard Tratz, published a paper on the bonobos were the authors not only used ‘bonobo’ as their vernacular name (in German), but also proposed to place the bonobo into its own genus(!) – which they called Bonobo (Tratz & Heck, 1954).

    The name matter then rested for a few decades, but in the 1990′ies, the famous chimpanzee researcher Adriaan Kortlandt raised the issue about the vernacular name of Pan paniscus. He thought that ‘bonobo’ was wholly inappropriate for reasons that ranged from the partly valid to the silly (“It goes against the Linnaean principle of binomiality“*, Kortlandt, 1997:29) and to the downright vile (e.g., thinly veiled personal attacks especially directed against Heinz Heck – who was already dead at the time – and borderline xenophobic slurs directed against Germans and Americans in general). Kortlandt (1997) also said that it was arrogant of Westerners to try to tell the Congolese people by what name they should call a species living exclusively within the borders of their country; that’s not a wholly flawed sentiment, but the irony in the situation where a Dutchman is telling – quite rudely, too – what English-speakers should call Pan paniscus seems to have escaped Kortlandt. Anyway, Kortlandt came down strongly in support of White’s (1996) suggestion to call the bonobo ‘gracile chimpanzee’ instead.

    * Yes, David, you read that right: Kortlandt was complaining about a vernacular name being incompatible with Linnaean nomenclature!

    Other notable chimpologists weighed in the dabate too. Takayoshi Kano and Toshisada Nishida (1999) suggested that bonobo could be replaced by ‘bilia’**, which is an authentic name actually used in (one of the many languages spoken in) the Republic of Congo.

    ** Also spelled ‘bilya’, depending on what orthography you follow. ‘Bilia’/'bilya’ is, however, a plural form; the singular would be ‘elia/’elya’. ‘Bilia’ should not be confused with the so-called ‘Bili apes’ which are something else altogether – if, indeed, they are anything at all. But let’s not get sidetracked into that discussion now!

    Thompson (1999) then pointed out that using the Wamba language word ‘bilia’ instead of bonobo could be interpreted by local people as foreigners taking sides in Congolese matters – something which could potentially lead to nasty consequences (both for field researchers and for the apes themselves) in the war-torn DR Congo. Maniacky (2006), however, was more sympathetic to Kano & Nishida’s ‘bilia’ suggestion.

    It’s safe to say, however, that for the time being at least, the use of ‘bonobo’ for Pan paniscus seems to be firmly established. Neither ‘gracile chimpanzee’ nor ‘bilia’ have thus far become mainstream.

    Oh, and apologies for hogging bandwidth with such an excessively long comment, Darren!

    References:
    Kano, T. & Nishisada, T. 1999. Bilia as an authentic vernacular name for Pan paniscus. Pan Africa News 6(1), 1-3.

    Kortlandt, A. 1997. Pygmy chimpanzee, bonobo, or gracile chimpanzee: what’s in a name. African Primates 3, 28-35.

    Maniacky, J. 2006. Pan paniscus, sometimes a linguistic issue. Pan Africa News 13(1), 4-6.

    Piggott, M.P., Banks, S.C. & Taylot, A.C. 2006. Population structure of brush-tailed rock-wallaby (Petrogale penicillata) colonies inferred from analysis of faecal DNA. Molecular Ecology 15, 93-105.

    Potter, S., Eldridge, M.D.B., Taggart, D.A. & Cooper, S.J.B. 2012. Multiple biogeographical barriers identified across the monsoon tropics of northern Australia: phylogeographic analysis of the brachyotis group of rock-wallabies. Molecular Ecology 21, 2254-2269.

    Thompson, J. 1999. Use of the name “bilia” for Pan paniscus. African Primates 4, 67-68.

    Tratz, E. & Heck, H. 1954. Der afrikanische Anthropoide “Bonobo”, eine neue Menschenaffengattung. Säugetierkundliche Mitteilungen 2, 97-101.

    White, F.J. 1996. Pan paniscus 1973 to 1996: twenty-three years of field research. Evolutionary Anthropology 5, 11-17.

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  90. 90. SRPlant 4:41 am 04/26/2012

    Dartian, thanks for telling that long, sordid and absolutely fascinating story!

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  91. 91. Dartian 5:28 am 04/26/2012

    Thanks, SRPlant!

    As for this species debate that we’re having, it was recently pointed out to me that there are exactly as many animal species in the world as Chuck Norris allows there to be. ;)

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  92. 92. David Marjanović 7:29 am 04/26/2012

    What does culture have to do with it? There was a recent study showing that even little kids take skin color into account when choosing friends.

    Where? In the USA, where blacks and whites behave as separate peoples and speak different dialects even when they live in the same place? In such an environment, “even little kids” will often grow up mostly with one and see the other as “the other”.

    The crate that the apes were held in during transportation had – so the story goes – the word ‘Bolobo’ on it. Bolobo is a city in what was then Belgian Congo [...]

    *headdesk*

    Worse than Gavialis and Ginkgo, then.

    there are exactly as many animal species in the world as Chuck Norris allows there to be.

    Trufax!!!

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  93. 93. vdinets 11:55 am 04/26/2012

    Dartian: that’s all very nice, but how can you extrapolate rock-wallaby results to klipspringers without any supporting data whatsoever? I’m yet to see any molecular data on them. This approach logically leads to designating fish in every pond separate species simply because they can’t fly.

    David: of course, most children grow up seeing their parents’ faces. If this is a self-perpetuating mechanism for assortative mating, it’s not that different from, say, cranes. What makes you think that it works in a different way in marmozets? Or are you implying that whooping and sandhill cranes are conspecific because their reproductive isolation is imprinting-dependent?

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  94. 94. llewelly 10:49 pm 04/26/2012

    To me it seems strange to argue that other species ought to split or lumped by the same standards that homo sapiens is. After all, it is unlikely there is any other population for which our judgement so skewed.

    Btw: Alien viruses from outer space and the great Archaeopteryx forgery : 87
    Eld’s Deer : 94 +

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  95. 95. vdinets 12:54 am 04/27/2012

    llewelly: but it is also the species that we have studied the most, and arguably the best understood one. It is reasonable to use it as a golden standard of sorts.

    Who cares about alien viruses threatening to mutate us into toads when ungulate systematics are at stake?

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  96. 96. Dartian 1:49 am 04/27/2012

    Vlad:
    how can you extrapolate rock-wallaby results to klipspringers without any supporting data whatsoever?

    Learn the difference between analogy and extrapolation, you ignoramus; they are not synonymous concepts. Also, marmoset isn’t spelled with a ‘z’.

    I’m yet to see any molecular data on them.

    But if someone did show you such data, you’d flip your position in an instance and start insisting that differences in morphology are more important. Don’t pretend that you actually care about facts, data, statistics, and actual evidence. During these increasingly tiresome ‘discussions’ with you it has become plainly obvious that you prefer to stick to anecdotes, personal prejudice, and to things you may once have read in some Russian children’s book about animals (that probably was written during the Lysenko era).

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  97. 97. vdinets 3:02 am 04/27/2012

    Dartian: analogy in this case is totally meaningless. Rock-wallabies are not klipspringers. And you haven’t produced a single shred of evidence in favor of splitting the latter.

    If I was shown good molecular data in support of splitting, I would admit that I was wrong. I’ve done it here on a few occasions. I wonder why you cannot do the same and admit that you are wrong, instead of switching to personal insults ;-)

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  98. 98. naishd 5:38 am 04/27/2012

    Very pleased to see the number of comments climbing so high here – I especially like Vlad’s comment “Who cares about alien viruses threatening to mutate us into toads when ungulate systematics are at stake?”.

    As blog-daddy, I have to remind some of you of cardinal Tet Zoo commenting rule # 1… personal attacks on others are not tolerated. Dartian – consider your wrist slapped, please do not insult other commenters like that. Because Dartian is not exactly a John Jackson or Jean-Pierre Mihalda, or whatever the hell his name was, I’m not about to ban him or delete his comments :) , but consider this a warning.

    Darren

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  99. 99. naishd 5:57 am 04/27/2012

    Incidentally, has anyone seen Nikki Le Roex’s 2008 University of Cape Town thesis ‘Phylogeographic Analysis Reveals Strong Geographical Structuring in the Klipspringer, Oreotragus oreotragus‘? Or, do they know if a published version has appeared anywhere?

    Darren

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  100. 100. Dartian 6:26 am 04/27/2012

    Darren:
    Dartian – consider your wrist slapped, please do not insult other commenters like that.

    Fair enough. I was out of line there; my Pliocene instincts momentarily got the better of me. My apologies, Vladimir. (And Darren.)

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  101. 101. David Marjanović 8:03 am 04/27/2012

    This approach logically leads to designating fish in every pond separate species simply because they can’t fly.

    They can – in the sense that their eggs are spread when they stick to bird feathers.

    David: of course, most children grow up seeing their parents’ faces.

    Not just their parents’, but those of everyone around them. If “everyone” is sufficiently diverse…

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  102. 102. Deinocogitus 10:56 am 04/27/2012

    Hola, Jerzy here again.

    Coming back to senses…

    One thing about Ungulate Taxonomy is that smaller sample size and poorer knowledge, the more likely authors seem inclined to split. So, lots of splits in Mazama deer but none in Odoliceus deer etc.

    Given that individuals and small local populations of ungulates easily can be so distinctive as subspecies, I have a feeling that small sample size made authors simply overlook natural variation and mixing of populations. Simply, if one studies rare animals with, say, 6 skins from one locality and 4 from distant another locality, they look different. When one has access to hundreds of animals, one will likely find that characters overlap, or that populations in between show clinal variation, or that size and color shape varies more with locality than macroscale.

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  103. 103. vdinets 12:51 pm 04/27/2012

    Dartian: accepted.
    I took another look at KS ranges in HMW (still waiting for G&G book to arrive, sorry). The maps show not just one fake distribution gap in Namibia, but also the following:

    (1) a huge gap between Cape and Transvaal KS. The entire Drakensberg area, the main stronghold of KS in South Africa, falls within this “gap”.

    (2) an even larger gap between Ethiopian KS and Golden KS. Numerous populations in N Kenya and S Ethiopia exist within this “gap”, for example, in Marsabit National Park.

    In reality, except for populations in Nigeria and Central African Republic, KS has fairly continuous distribution. The large gap between populations in Namibia and Zambia/E Bostwana is very recent: Shortridge found KS in Tsodilo Hills in 1934, and that place is exactly in the middle of that gap. I wonder how many other populations have been hunted out only recently.

    David: so do marmosets. Are you sure that a cross-fostered marmoset would still prefer to mate with its own race?

    Reference: Shortridge, G. C. The Mammals of South West Africa: a biological account of the forms occurring in that region. W. Heinemann, ltd. NY, 1934.

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  104. 104. David Marjanović 7:09 am 04/28/2012

    Are you sure that a cross-fostered marmoset would still prefer to mate with its own race?

    Is there a “not” missing in there? Anyway, I think it would prefer to mate with individuals that don’t look too different from the ones it was brought up with, no matter whether they’re genetic relatives or not.

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  105. 105. naishd 7:53 am 04/28/2012

    Any discussion of hybridisation within marmosets should make reference to the fact that hybridisation involving two hardy, artificially introduced marmoset species – the Common marmoset Callithrix jacchusi and Black tufted-ear marmoset C. penicillata – is currently occurring in the Brazilian Atlantic Forest where it is seriously threatening native, endangered species. Indeed the Common and Black tufted-ear marmoset hybridise anyway in the Recôncavo da Bahia region (the hybrid zone is about 50 km wide), and the Black tufted-ear marmoset also hybridises with White-headed or Geoffroy’s marmoset C. geoffroyi in Minas Gerais (at the cerrado-Atlantic Forest interface). There’s a large literature on hybrid marmosets created in captivity.

    What do these hybridisations tell us about the mating preferences of these animals? Perhaps that, should the opportunity arises they will mate with any available partner that is about “similar enough” – the species concern differ notably in overall pelage colour, and in the detailed colouring, patterning and appearance of the cheeks, throat, crown, ears and other facial features. We are not dealing here with controversial taxonomic forms that differ only in tiny, trivial details – hybridisation is occurring rampantly even between non-controversial ‘species’ (note: apparently as a consequence of man-made introduction, habitat degradation etc.). It’s for this reason (and others) that I don’t necessarily regard the presence or extent of hybridisation as important when it comes to making decisions about the status of a species. Hybridisation is rampant, occurring between many populations that are otherwise well supported as ‘good species’.

    Darren

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  106. 106. vdinets 3:11 pm 04/28/2012

    So, how are marmosets (or other primate species recently split based on facial features or color patterns, such as various limestone langurs) different from human races?

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  107. 107. naishd 7:55 pm 04/28/2012

    Vlad – well, it would be helpful to know exactly which marmosets you’re referring to, since most of the recently named or resurrected taxa I have in mind (e.g., Buffy-headed marmoset, Buffy-tufted-ear marmoset, Wied’s black-tufted-ear Marmoset) don’t just differ in facial features, but also in other aspects of pelage (I would say, subjectively, that they differ as much as, if not more than, many other species traditionally recognised as non-controversial). They also don’t overlap in skull measurements: on this issue, Marroig et al. (2004) said “there are sharp, stepped morphological boundaries among taxa, consistent with species-level distinction”.

    Those that have been studied are also known to differ in the frequency and structure of their vocalisations, and in fact this data alone is so good (for species like the Buffy-headed marmoset) that a case can be made for considering the respective taxa worthy of distinction (however you decide to recognise this) on acoustic data alone (see data in Medes’s chapter in Ford et al.’s The Smallest Anthropoids: The Marmoset/Callimico Radiation). Certainly theoretically, they’re exhibiting strong evidence for reproductive isolation.

    Furthermore, while there aren’t (so far as I know) compelling phylogenies for most or all marmoset species, the work that’s been done so far indicates that the species concerned (I’m still referring here to eastern species, like the Buffy-headed marmoset, Wied’s black-tufted-ear marmoset etc.) diverged 1.0-1.5 million years ago, with even the YOUNGEST split (between Geoffroy’s marmoset and Common marmoset) being at c. 500,000 years ago (Barroso et al. 1997).

    In short, even these supposedly very similar marmoset species are – I would argue – fundamentally different from races within our species.

    I don’t have time to discuss my thoughts on langurs right now as well, but the data there on morphological differences, divergence dates and so on is also not at all like that for human races.

    Darren

    Refs – -

    Barroso, C. M. L, Schneider, H, Schneider, M. P. C., Sampaio, I., Harada, M. L., Czelusniak, J. & Goodman, M. 1997. Update on the phylogenetic systematics of New World monkeys: further DNA evidence for placing the pygmy marmoset (Cebuella) within the genus Callithrix. International Journal of Primatology 18, 651–674.

    Marroig, G., Cropp, S. & Cheverud, J. M. 2004. Systematics and evolution of the jacchus group of marmosets (Platyrrhini). American Journal of Physical Anthropology 123, 11-22.

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  108. 108. vdinets 12:29 am 04/29/2012

    Darren: some human races also have non-overlapping skull measurements. Do we know if marmoset vocalizations are learned or innate? The reproductive isolation argument contradicts what you said above. Looks like the only really good evidence for splitting is genetic. Which bodes well with what I said in the beginning: nowadays, any splitting proposal should include solid genetic data in addition to phenotypical data. I can’t understand why this innocent suggestion is facing such negative reaction :-(

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  109. 109. David Marjanović 6:55 am 04/29/2012

    So, how are marmosets (or other primate species recently split based on facial features or color patterns, such as various limestone langurs) different from human races?

    Human “races” are so blurry that you can’t even define hybrid zones. Go from Moscow to Ulaan Baatar and tell me where the hybrid zone begins and ends!

    Importantly, in humans, every gene has its own geographic variation. There’s a strong geographic signal in blood group genes, for instance, but they’re completely orthogonal to skin color, hair shape, nose shape, whatever – indeed, they don’t care about each other, except for the founder effect in South America.

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  110. 110. vdinets 12:44 pm 04/29/2012

    David: if you go from Moscow to Ulan Baatar, you cross a region where hybridization has been going on for at least three thousand years due to migration. So it is all one broad gradient, with relatively pure forms still found at the ends (the Basques and the Mongols). But if you go from Moscow to Cape Town or from Ulan Baator to Cape Town, the situation will be very different: there will be plenty of sharp limits with very narrow hybridization zones (if any), where many genes change at once. Just look at the Sahel, or the Bantu/Khoisan divide in Namibia. And note that some groups have managed to maintain high levels of genetic and phenotypical distinctiveness for thousands of years even in sympatry, like the Negrito of the Philippines and the Orang Asli of Malaysia.

    My people, the Ashkenazi Jews, have maintained high level of genetic distinctiveness in full allopatry for centuries. We also have diagnosable phenotypic differences (any good European anti-Semite can provide you with an identification key). Now we are threatened with hybridization. I suggest splitting us under CSC, for conservation purposes, and listing as an endangered species. My rental apartment would immediately become my critical habitat (so I don’t have to pay rent anymore), and there would be other nice perks, like grants available to me for studying my ecology, although I’d probably face difficulties transporting myself without a CITES permit, and my wife would need a permit to breed me in captivity.

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  111. 111. David Marjanović 7:52 am 04/30/2012

    But if you go from Moscow to Cape Town or from Ulan Baator to Cape Town, the situation will be very different: there will be plenty of sharp limits with very narrow hybridization zones (if any), where many genes change at once. Just look at the Sahel, or the Bantu/Khoisan divide in Namibia.

    The Sahel lies south of a big geographic barrier, the Sahara. Rather look to Ethiopia and thereabouts. European noses are normal all the way to Rwanda, if not beyond.

    The sharp divide between Bantu and Khoisan is due to recent migration. :-) Look at the Hadza and Sandawe instead.

    And note that some groups have managed to maintain high levels of genetic and phenotypical distinctiveness for thousands of years even in sympatry, like the Negrito of the Philippines and the Orang Asli of Malaysia.

    Yes, culture can be a fairly good separator. Even so, their genetic distinctiveness is limited.

    We also have diagnosable phenotypic differences (any good European anti-Semite can provide you with an identification key).

    Sure they can. It just fails much of the time. There are, or at least were, blond, blue-eyed Ashkenazim…

    Thanks to a migration just over a thousand years ago, geneticists can recognize twenty Hungarians as Hungarians. Presented with just one, it’s much more difficult to tell where somebody comes from.

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  112. 112. naishd 9:21 am 04/30/2012

    Response to Vlad (comment 108). Vlad says…

    Darren: some human races also have non-overlapping skull measurements.

    I suppose you might take some highly distinct ‘extremes’ within Homo sapiens to demonstrate this. For the most part, my understanding is that there is massive, pervasive, continuous overlap in the cranial measurements of human races. This is not the case in the marmosets we’re discussing, where the relevant populations plot discretely in terms of cranial size, nasal width, oral width, and numerous other details of cranial anatomy. See Marroig et al., cited above, and compare it with any study that examines morphological variation across H. sapiens.

    Do we know if marmoset vocalizations are learned or innate?

    Some aspects of their vocalisations must be considered genetically innate, but a lot of work has shown that juveniles learn vocalisations, and that individuals (of the species that have been studied, anyway) modify their voices throughout their lives. I’m not sure what point you’re getting at (I suppose you’re trying to argue that the different vocal characters I was referring to above are more to do with culture than phylogenetic distinction), but my point is that the inherent call frequency (I mean acoustic frequency, not ‘frequency’ in terms of occurrence) and the form and shape of the calls is seemingly different because of ecological adaptation and different anatomies, not because of culture. In other words, the calls differentiate the populations in the same way that, say, microbat vocalisations do.

    The reproductive isolation argument contradicts what you said above.

    Well, the populations differ morphologically and in vocal characters (thus supporting ideas that they should be recognised as species), but they can still hybridise. That’s because – as I keep saying – the existence of hybridisation doesn’t count for squat. Given the chance, many animals will mate with similar-looking relatives; it doesn’t tell you anything about whether said populations should be regarded as ‘species’ or not.

    Looks like the only really good evidence for splitting is genetic.

    Err, no. Yes, the genetic distance data is consistent with this but, as I noted above, so is the morphological evidence (pelage differences, cranial measurements etc.) that first led to separate species recognition.

    Which bodes well with what I said in the beginning: nowadays, any splitting proposal should include solid genetic data in addition to phenotypical data. I can’t understand why this innocent suggestion is facing such negative reaction.

    I’m not sure that anybody disagreed with you on this. Splits of the sort proposed by Groves and Grubb should be seen as hypotheses that require genetic testing, and maybe this is the problem: people always assume that, when you name a new species, you’re attempting to change the world forever. As I (and others) have said before, taxonomic changes are recommendations, not strict mandates, and you might see them as the start of what’s meant to be a conversation. Objecting to them from the start purely on philosophy is not, it seems to me, an evidence-led approach.

    Darren

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  113. 113. vdinets 11:33 am 04/30/2012

    David: trans-Saharan trade has existed for at least three thousand years. Note that Ethiopians have noses very distinct from Semitic ones found further north. Blond Ashkenazim can still be recognized as such, and their existence indicates very limited genetic flow. You have to look at the bigger picture: humans have evolved numerous distinctive races, often without full allopatry, or even in full sympatry. Marmoset taxa apparently depend on full allopatry for emerging and maintaining themselves.

    Darren: it depends on which cranial measurements you are looking at. Asian skulls have very distinctive facial features, for example. Archeologists can instantly distinguish them from, say, Caucasian skulls.

    Innate differences in sound production apparently exist between human races as well (reference below).

    What I am trying to say is that molecular data is the only argument for splitting that applies to marmosets and not to humans.

    As for G&G, it’s OK to publish whatever you want if you can. It’s a free country, as the saying goes. What really pissed me off was that their private exercise was followed in HMW.

    Reference: JH Walton, RF Orlikoff. 1994. Speaker Race Identification From Acoustic Cues in the Vocal Signal. Journal of Speech and Hearing Research Vol.37 pp 738-745.

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  114. 114. naishd 2:17 pm 04/30/2012

    Very briefly (I really think we’re about done on this discussion!)… the many non-overlapping sets of measurements seen in the marmosets aren’t like those of human races. Yes, it is not all that difficult to recognise the distinctive hallmarks of (say) an Asian-type skull in Homo sapiens, but you can only pretend that the differences from ‘another type’ of skull are notable if you work with extreme end-members: as I’m sure you know, the ‘skull types’ identified for our species grade into one another, there has historically been huge confusion over where the boundaries are, and the key features associated with these skull types are not consistently present in the people supposed to possess them. Indeed, there have been big debates over the ‘racial’ identifications of humans known only from their bones.

    As for speaker race discrimination based on acoustic cues, you’re citing a study that examined how good people were at discriminating black people from white people. I don’t doubt that different kinds of people often sound different (for a whole list of reasons), but I don’t think these differences are in the same league as the frequency and structural variations seen in marmoset trills and other calls. Indeed, in the different human voices tested in the study you cite, “there were no significant differences in the mean fundamental frequency or formant structure”.

    It is, I feel, rather misleading to keep implying that racial differences in modern humans are comparable to the differences we see between populations of animals like marmosets. We’re horribly biased anyway when it comes to looking at members of our own species – we’re all experts at spotting, and over-emphasising, tiny differences in anatomy.

    Darren

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  115. 115. David Marjanović 4:03 pm 04/30/2012

    Asian skulls have very distinctive facial features, for example. Archeologists can instantly distinguish them from, say, Caucasian skulls.

    Is that so?

    Even male and female skeletons are difficult to distinguish. You can tell Superman and Wonder Woman apart, sure, but with normal people it’s a matter of probabilities, and often the probability is 50-50.

    Innate differences in sound production apparently exist between human races as well (reference below).

    That’s “races” as defined in the USA, where blacks and whites usually don’t natively speak the same dialect of English, and where “black” is defined in such a way that the same person can be “black” in the US, “colored” in South Africa and “white” in Brazil.

    trans-Saharan trade has existed for at least three thousand years.

    Sure, but few people have participated. There hasn’t been as much gene flow as along the Nile or across the Eurasian steppe.

    Note that Ethiopians have noses very distinct from Semitic ones found further north.

    What nose shape diversity do, say, the Dinka and Nuer have?

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  116. 116. vdinets 7:16 pm 04/30/2012

    David:
    Yes, it is. There are ancient Caucasian burials in many parts of China, and even the most patriotic Chinese archeologists have never tried to write them off as Chinese. It has nothing to do with sexing skulls.

    Sure, the system of assigning race used in the US is very simplistic and politically skewed, but the study does show that there are differences in sound production even between sympatric ethnic groups.

    Trans-Saharan trade in slaves was very extensive. Even in Britain, African haplogrups show up in some people. Note that the trade along the Nile is also trans-Saharan.

    I have seen few Dinka and Nuer (never been to Sudan or far western Ethiopia), but I was under impression that the Dinka have broader nose bridges than the Nuer, and all of them look very distinctively more “African” than their Amhara-speaking neighbors to the east. Why?

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  117. 117. David Marjanović 6:44 am 05/1/2012

    There are ancient Caucasian burials in many parts of China, and even the most patriotic Chinese archeologists have never tried to write them off as Chinese.

    Isn’t that just because some of them are mummified?

    the study does show that there are differences in sound production even between sympatric ethnic groups

    What about the counterarguments by Darren and me?

    Trans-Saharan trade in slaves was very extensive. Even in Britain, African haplogrups show up in some people.

    That still doesn’t mean it involved a lot of people.

    Note that the trade along the Nile is also trans-Saharan.

    Of course; it’s the most important way around the barrier… and it’s almost at its east end, not through in the middle.

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  118. 118. vdinets 3:20 am 05/2/2012

    David,
    No, very few of them are mummified.

    I still don’t see any fundamental difference between the differences between marmoset vocalizations and those between human languages. If there is such a difference, it hasn’t been conclusively shown.

    Trans-Saharan slave trade involved millions of people over the centuries – it was more extensive than trans-Atlantic slave trade, and existed much longer. Don’t forget that Indian Ocean shipping routes were also functionally trans-Saharan. But you don’t see much African phenotypic influence north of Yemen. One reason is that most slaves were males, and most were castrated (since Muslims are not allowed to castrate anybody or anything, Coptic and other Christian priests were routinely hired to perform the surgery). I am not aware of an equally effective behavioral mechanism for suppressing hybridization in any species.

    I think it’s time to let this subject go. The whole discussion had nothing to do with Eld’s deer in the first place :-)

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  119. 119. naishd 4:55 am 05/2/2012

    For the record, the calls of marmoset taxa are very different from voice/dialect/language differences between human races because, as I think I said above, the marmoset calls differ in acoustic morphology – that is, in the frequency in hz, in waveform structure and so on. These differences in acoustic morphology are of the sort expected for, say, different species of bats and songbirds; they support the view that the marmosets concerned should be regarded as distinct species (e.g., Snowdon 1993 and other studies). Human dialects and languages sound ‘distinct’, and individual humans do differ in fundamental and formant frequency, but studies that have examined ‘racial’ and other differences in humans speech (e.g., Walton & Orlikoff 1994, Thomas & Reaser 2004, Dehqan et al. 2010) document differences in speech patterns, vowel formation and so on – not differences in major aspects of acoustic morphology.

    Incidentally, local dialects are present in some marmoset taxa (like the Common marmoset), and – in some species – left-handed individuals use calls differently from right-handed individuals. These differences are more like the ones seen between human dialects and languages.

    And, yes, we’ve come a long way from Eld’s deer. Such is the nature of thread drift.

    Darren

    Refs – -

    Dehqan, A., Ansari, H. & Bakhtiar, M. 2010. Objective voice analysis of Iranian speakers with normal voices. Journal of Voice 24, 161–167.

    Snowdon, C. T. 1993. A vocal taxonomy of callitrichid primates. In Rylands, A. B. (ed). Marmosets and Tamarins: Systematics, Behaviour, and Ecology. Oxford University Press, Oxford, pp. 78-94.

    Thomas, E. R. & Reaser, J. 2004. Delimiting perceptual cures used for the ethnic labeling of African American and European American voices. Journal of Sociolinguistics 8, 54-87.

    Walton, J. & Orlikoff, R. 1994. Speaker race identification from acoustic cues in the vocal signal. Journal of Speech & Hearing Research 37, 738-745.

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  120. 120. SRPlant 6:20 am 05/2/2012

    #119 ” left-handed individuals use calls differently from right-handed individuals.”

    Wow and double wow!

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  121. 121. vdinets 5:25 pm 05/2/2012

    Darren: still, we don’t know if these differences are innate, and even if they are, it’s still possible that they are caused by single-point mutations, for example.

    And they apparently don’t create reproductive isolation, unlike differences in human language :-)

    I have to drop out of the discussion, sorry. Will be traveling for the next month.

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  122. 122. naishd 5:32 pm 05/2/2012

    Ok… but.. human language creates reproductive isolation? Where did that come from?

    No need to respond. We’re done here :)

    Darren

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  123. 123. vdinets 5:54 pm 05/2/2012

    Of course it does. It’s only partial, but pretty effective. Have you ever tried to woo a Chinese girl who didn’t speak a word of English? It’s possible, but very difficult.

    Oh, and from today’s news it looks like N and S white rhinos are two different species. At least they seem to prefer mating with their own:

    http://blogs.scientificamerican.com/extinction-countdown/2012/04/30/the-most-eagerly-awaited-rhino-porn-of-all-time/

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  124. 124. naishd 6:01 pm 05/2/2012

    I don’t think the rules of reproductive isolation are anything to do with ‘wooing’. Come on, people successfully breed with speakers of other languages, members of other races and cultures all the time.

    Darren

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  125. 125. vdinets 6:31 pm 05/2/2012

    Of course they are. How do you think it works in birds?

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  126. 126. naishd 4:23 am 05/3/2012

    Something tells me that the reproductive isolation seen in birds does not exactly apply to human races.

    Darren

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  127. 127. Heteromeles 10:31 am 05/3/2012

    Speaking as someone who lives very happily in a biracial household, even though I’m having a hell of a time learning my partner’s birth language (she’s trilingual, I’m monolingual), I’d say that arguments about languages separating humans is a load of stinking BS that should be ignored, period.

    It could also be that I live in a highly, racially mixed town, with two major languages (English and Spanish) that are both primarily spoken by people who are not racially English nor Spanish. Since I know my ancestry back to about the 16th Century, I can say with confidence that over a century ago, most of my people did not speak English. That didn’t stop them from emigrating, either.

    Let’s also, for the sake of jack hammering into poor ol’ vdinets’ cramped little conceptual universe, look at where there’s a lot of linguistic diversity: Papua New Guinea. There’s massive linguistic diversity there, but there’s not a lot of human genetic diversity there. Many Papuans are multilingual.

    This is probably the norm. In the Americas before Columbus, there were thousands of languages, but not much genetic diversity when compared with, say, Africa.

    Or one can look at the diversity of English dialects in England, especially a few centuries back, compared with the relative paucity of dialects in the US, which is both bigger and much more genetically diverse.

    Sorry, there’s no correlation between human linguistic diversity and genetic diversity. It’s BS. Can we just start ignoring this discussion when someone tries to bring it up again?

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  128. 128. vdinets 11:16 am 05/3/2012

    Heteromeles: this is the last time I answer you, unless you learn to behave in a civil manner.

    I am happy for your biracial relationship; I’ve had a few of those myself. Nobody says that linguistic isolation is absolute. But it would be stupid rather than simply politically correct to claim that there is no assortative mating between ethnic groups, that all mating in humans is random. If there was no partial reproductive isolation, there would be no such thing as races and ethnic phenotypes. Don’t forget that only a small minority of human ethnic phenotypes exist in allopatry; most are parapatric or even sympatric with others.

    Now, how do you think this partial isolation works? Why do Gypsies still look different from native Europeans after centuries of living in their midst? There are three isolating mechanisms: facial differences, linguistic isolation, and cultural taboos. However, in many situations where cultural taboos and facial differences are absent, small ethnic groups like the Basques still avoid assimilation. It means that linguistic isolation has an effect, doesn’t it?

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  129. 129. Hai~Ren 2:23 pm 05/3/2012

    I hate to say this, particularly since it involves human behaviour, but I do wonder if anyone has ever taken a good look at the significance of rape as a strategy to overcome linguistic or cultural barriers.

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  130. 130. Heteromeles 11:02 pm 05/3/2012

    Vdinets, if you are feeling insulted, too bad. You’re trying to make say science supports your view of racism. It doesn’t, and neither does reality. Feel free to be insulted by reality telling you that you are dead, flat, wrong. That’s what science does to people. There’s nothing politically correct about this statement either. It’s simply being human.

    Since you yourself appear to be willing to mate with those outside your ethnic group, you have first hand experience of that reality. Apparently it can’t break your preconceptions, which is too bad.

    As for what causes human phenotypes, I’ll give you two answers:
    1. Learned discrimination. This is by far the biggest cause of the perception of human phenotypes. In the US, skin color and hair texture are used as the major discriminants. If you can get past these two superficial characteristics, you’ll see an enormous diversity of structural phenotypes that are swept under the race rug. I’ve seen black people who could be made to look white, Korean, Japanese, Indian, or Hispanic just by whitening their skin and giving them a straight-haired wig. Trouble is, we tend to stop when we categorize people as black, or Hispanic, or Asian. If you can learn to see beyond those superficial traits, you will find more features in common than you think.

    Learned discrimination is by far the biggest issue, and it’s the most pernicious one.

    Another example: my partner, who was born in a racially homogenous country, can’t always identify people from her birth country until they start talking. Even though they are regarded as among the most homogenous people on the planet, they still have a fair amount of phenotypic overlap with their neighbors.

    So the big problem: phenotype is in part a learned illusion. It involves learning to identify people by hair and skin color, and ignore everything else. “I can’t tell those Asian/Black/Hispanic people apart,” is a classic statement of the problem.

    The second and lesser issue with race has to do with real boundaries, which primarily include things like politics (as was the case in my partner’s homeland for a few centuries) and the population isolation within the last perhaps 10,000-20,000 years. Before that, the human skulls available (paleoindians, mesolithic humans, etc) do not (to my knowledge) share the characteristics of any living population near their location. This suggests that the characteristics that currently define racial groups change really, really fast on an evolutionary scale.

    This is the other problem with your idea: the characteristics that define “a people” change over time quite widely. The English of 2000 years ago (let alone 5000 years ago) do not look entirely like the English of today. Even if we limited it to whites. Certainly we can find people who “would have fit in 2000 years ago,” but for all we know, this is more likely to be due to random chance than some sort of “pureblood” genetic isolation in action.

    Ultimately, racial arguments use the same mechanism as astrology. Astrology (the huge long horoscopes, not the newspaper silliness) “works” by offering up a huge number of vague statements about the human condition. As you read a horoscope, you tend to catch the phrases that apply to you and ignore the rest, unless they are gratuitously wrong. It can be real fun to read someone else’s horoscope, and find out that it applies better to you than it does to them (this is the basis of a psych experiment I’ve run, incidentally).

    With race, people tend to pick out representative pictures and ignore the rest, when they define racial characters. Example statements include: “Oh, she looks so very English,” “That chief is the epitome of a Tahitian,” “That model is such a slender Ethiopian, they’re all so elegant,” “He’s got a big (Jewish/Italian/French) nose,” and so on. If you actually mass together all the people of England, or Tahiti, or Ethiopia, or Israel, you will find that there’s a huge amount of within-group diversity, a fair amount of between-group overlap, and the characters that differentiate the groups are fuzzy, subjective or only apply to some proportion of each group. In phylogenetic terms, these so-called racial traits are uniformative characteristics. Why do you want to use them?

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  131. 131. David Marjanović 9:25 am 05/4/2012

    Heteromeles: this is the last time I answer you, unless you learn to behave in a civil manner.

    :-D :-D :-D

    Tone troll. :-)

    But it would be stupid rather than simply politically correct to claim that there is no assortative mating between ethnic groups, that all mating in humans is random. If there was no partial reproductive isolation, there would be no such thing as races and ethnic phenotypes.

    Nobody says it’s random. It’s clinal.

    There are three isolating mechanisms: facial differences, linguistic isolation, and cultural taboos.

    Cultural mechanisms don’t need to be taboos to work. Some time before WWI, somebody did a study on where people found their spouses in a small area on both sides of what was the border between Germany and Austria-Hungary (nowadays Poland and the Czech Republic). There were a lot fewer marriages across the border than on either side, even though the border was not a language barrier (German, and pretty much the same kind of German, was spoken on both sides). It was difficult enough to cross the border that people got culturally used to not crossing it often.

    However, in many situations where cultural taboos and facial differences are absent, small ethnic groups like the Basques still avoid assimilation.

    Well, the area in which their language is spoken has been shrinking for centuries, and in terms of genetics they’re only the peak of a cline – there are no sharp boundaries.

    (BTW, their famous frequency of rh– is also found in a few valleys of the Caucasus and is probably due to isolation effects of such valleys, not a plesiomorphy.)

    Certainly we can find people who “would have fit in 2000 years ago,” but for all we know, this is more likely to be due to random chance than some sort of “pureblood” genetic isolation in action.

    And of course, pureblood genetic isolation is a strong factor for change by genetic drift.

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  132. 132. vdinets 2:18 am 05/5/2012

    David: interbreeding patterns between human ethnic groups are very diverse. You can find areas with clinal variation, huge hybrid swarms (like in Brazil), but also numerous places where different groups behave like different species: despite centuries in symapatry, they maintain almost complete reproductive isolation, linguistic identity, and separate ecological niches (for example, in parts of Africa where there are tribes of agriculturalists, cattle breeders, hunters-gatherers, etc.) I am not saying that ethnic groups should be considered full species, of course (although they often do qualify as subspecies under the commonly used definitions). What I am saying is that we shouldn’t use double standards. If we consider all humans conspecific, we shouldn’t split other species based on things like signaling differences, facial features, general coloration, size and/or limited interbreeding, unless there is also good genetic evidence for splitting.

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  133. 133. Jerzy v. 3.0. 10:57 am 05/5/2012

    #107
    You may be unaware, but many human populations are now believed to by hybrids between Homo sapiens, neanderthals and Denisovians, believed to be originally distinct ca 1,000,000 years ago.

    Like or not, more narrow definitions of species can result in splitting humans into multiple species. We want to avoid it for many reasons, from ideological (avoiding racism) to mundane.

    Therefore, the only option is to start evaluating subspecies for their conservation status.

    Protecting subspecies (as opposed to practice of raising subspecies to species) is a must also because even subspecies little different genetically can be irrepraceable ecologically. I written already several times here about tigers and leopards. Their subspecies are poorly different genetically and indeed there is/was a clinal variation from N to S Asia. However, short-haired Malayan tigers and leopards couldn’t survive in snowy Siberia and vice versa.

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  134. 134. naishd 11:08 am 05/5/2012

    Jerzy – I think that the genetic contribution of Neanderthals and Denisovans to Homo sapiens should be assumed knowledge, especially for someone who runs a blog titled ‘Tetrapod Zoology’ :)

    Anyway, I agree with what you say about conserving subspecies.

    Huh – 134 comments…

    Darren

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  135. 135. David Marjanović 3:04 pm 05/5/2012

    Homo sapiens, neanderthals and Denisovians, believed to be originally distinct ca 1,000,000 years ago

    More like 600,000 years ago.

    Anyway, I agree with what you say about conserving subspecies.

    So do I!

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  136. 136. Jerzy v. 3.0. 4:15 pm 05/5/2012

    #134
    Yes, assumed. But I just recently realized that split between Homo sapiens and Denisovians (do they have a scientific name?) is similar in age to many recently proposed species splits.

    Denisovians are curious creatures, anyway. Most striking is that evolution of human sapience is not unique event. As we understand, Homo erectus apparently lacked it, but Denisovians must have evolved sapience independently from human and neanderthal branch to cope with cold Asian climate.

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  137. 137. David Marjanović 7:03 am 05/6/2012

    do they have a scientific name?

    No, because that isolated finger isn’t diagnostic on morphology alone.

    human sapience

    What do you mean by that???

    independently from human and neanderthal branch

    Denisovans and Neandertalers are sister-groups.

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  138. 138. Jerzy v. 3.0. 6:08 am 05/7/2012

    “No, because that isolated finger isn’t diagnostic on morphology alone.”

    Is DNA sequence technically not a diagnostic part of the body? There is at least one precendent. Craig Venter sequenced several genomes from ocean water of microbes which nobody (including himself) seen. Are they still unnamed?

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  139. 139. David Marjanović 2:08 pm 05/7/2012

    Craig Venter sequenced several genomes from ocean water of microbes which nobody (including himself) seen. Are they still unnamed?

    Yes! The code of nomenclature of prokaryotes is even stricter than the zoological one: to name a new species of “animal”, you need to provide a diagnosis and a type specimen; to name a new species of prokaryote, you need a living culture as the type specimen (so you can observe differences in metabolism that are diagnostic for a species).

    http://en.wikipedia.org/wiki/Candidatus

    An example is the most common form of life on Earth, Candidatus Pelagibacter ubique.

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  140. 140. jlouys 11:18 am 05/14/2012

    Great article Darren,
    Incidentally I’ve just published a paper on (in part) the palaeontological history of Eld’s Deer and its conservation implications. While not as well represented in the fossil record as other Southeast Asian megafauna, its general biogeographic pattern through the Quaternary seems to follow much more critically endangered species such as the rhinos, tigers and tapirs. Full article here: http://www.springerlink.com/content/x18705443012603q/

    Link to this

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