Thursday, June 08, 2006

The interesting and contentious discovery of the kipunji

Monkeys are one of the most misunderstood and mischaracterized of all mammals, at least among those who aren’t primate specialists. ‘Monkey’ is itself a loose term used for those members of Anthropoidea that aren’t hominoids, and here we’re only interested in the Old World monkeys (cercopithecoids).

Pet peeve no. 375 is the fact that people think that apes evolved from monkeys (they didn’t: hominoids and cercopithecoids are sister-taxa within the anthropoid clade Catarrhini*), and that apes are therefore somehow superior to monkeys in morphology and adaptability. If anything it’s the other way round, with a phenomenal post-Miocene radiation of monkeys pushing the previously hyper-diverse apes into the shadows. That’s a story for another time however.

* Having said that though, given that people have applied the term ‘monkey’ to all extant non-hominoid anthropoids, a stem-group hominoid would still likely be identified by people as a ‘monkey’.

One monkey in particular has been the focus of a lot of attention lately, the Highland mangabey Lophocebus kipunji (although this is not its current name: see next post).

The story of the Highland mangabey’s discovery is an interesting one. In January 2003 Tim Davenport of the Tanzanian Wildlife Conservation Society ‘heard rumours [from the local Wanyakyusa people of the Mount Rungwe region] about a shy and atypical monkey known as Kipunji’ (Beckman 2005, Jones et al. 2005, p. 1161), and became interesting in tracking down the species that might lay at the bottom of these reports. Meanwhile, another primatologist – Trevor Jones – had been amazed to observe an unusual, unidentifiable monkey in the Tanzanian Ndundulu Forest Reserve, a location about 350 km away from the source of the Kipunji reports.

So, at this stage, we have an ethnoknown primate known only to scientists by way of fleeting observations. This makes the Kipunji a bona fide cryptid, and, to repeat a point I’ve made before (in connection with the Odedi, a cryptic warbler from Bougainville Island), one would be justified in arguing that Davenport, Wood and their colleagues were now engaging in cryptozoological research. By definition these primatologists are therefore part-time cryptozoologists, whether they like it or not.

Good observations were finally made of the Kipunji in December 2003, and it now turned out that a monkey population reported in 2001 from the Ndundulu Forest Reserve, misidentified as Sanje mangabeys Cercocebus sanjei (itself only discovered in 1979), were actually reports of the Kipunji. Davenport’s team and Wood’s team both learnt that they’d been tracking down two different populations of the same monkey in October 2004 – one population in the Mount Rungwe-Livingstone Forest area in the Southern Highlands of Tanzania, and the other in the Ndundulu Forest Reserve in the Tanzanian Udzungwa Mountains.

In May 2005 they published a jointly authored description of this new species in Science (Jones et al. 2005), naming it the Highland mangabey Lophocebus kipunji Ehardt, Butynski, Jones & Davenport, 2005. Note that the authorship of the species doesn’t match the authorship of the descriptive paper. This does happen occasionally in the literature, and in this case it means that three authors of the original description were not involved in the naming of the species. Something else makes the Jones et al. paper odd, however, and this is the apparent lack of a type specimen for Lophocebus kipunji. Nowhere in the paper do they list or cite an accessioned specimen, nor mention the procurement of a specimen for a museum or university collection. Under ‘holotype’ they wrote ‘Adult male in photograph (Fig. 2)’, with a ‘paratype’ being described as ‘Adult in photograph (Fig. 3)’ (p. 1162). Because the new species seems to be critically endangered, Jones et al. (2005, p. 1162) stated ‘no live individual should be collected at this time to serve as the holotype’.

While it might seem ethically ‘nice’ and – from the conservationist perspective – sensible to not collect specimens from endangered or declining populations, it’s problematic to not do so, as the ICZN mandates that actual type specimens are needed for the establishment of a species. Furthermore, biological entities can only be regarded as unquestionably valid when there are physical specimens accessioned in proper collections.

Consequently there were several follow-up comments in Science on the lack of a Highland mangabey holotype (Landry 2005, Moser 2005, Polaszek et al. 2005, Timm et al. 2005). Timm et al. (2005) argued that the lack of a type specimen means that Lophocebus kipunji ‘is not an available name and has no formal standing in zoology’ (p. 2163) and Landry (2005) argued that the authors should have published ‘all of the excellent descriptive material and their quite convincing case for calling it new, without, however, naming it’ (p. 2164). Landry also noted that only four of the seven authors of the paper are listed as namers of the new species, but that ‘the purpose of this citation is to identify the paper, not to assign credit, and all of the authors should be cited’. Interesting.

Polaszek et al. (2005) responded to these criticisms by arguing that the ICZN does actually allow the recognition of taxa without holotypes (Article 73.1.4), and they argued that conservation concerns should encourage zoologists to catalogue species on the basis of vocalisations, molecular information and so on, and that ‘dead animal specimens should not be understood to be essential to the process of establishing new taxa’ (p. 2165). Hell, if Bernard Heuvelmans were still alive he would dance a little jig (to paraphrase Greg Paul). Moser (2005) pointed out that Polaszek et al. (2005) seem to have completely misunderstood the point of Article 73.1.4: it refers to cases where type specimens have become lost, and does NOT say that lack of an original type specimen is acceptable.

Anyway, it isn’t the first time that this has happened, and it won’t be the last. Among tetrapods the most famous case is that of the Bulo Burti boubou Laniarius liberatus, a Somalian shrike captured, observed in captivity, and then released (Smith et al. 1991). Exactly the same thing has just happened with another new monkey: a Brazilian platyrrhine named the Blond capuchin Cebus quierozi Mendes Pontes & Malta, 2006 (again, the species’ authorship doesn’t match the authorship of the paper. Again, weird). Apparently critically endangered and restricted to a tiny area already renowned as a centre of endemism, the authors elected not to sacrifice the type specimen (ironically confiscated from a local hunter), but to release it back into the wild (Mendes Pontes et al. 2006). I could talk about this subject a lot more, but won’t do so here. Suffice it to say, I still think that the collection of specimens is an integral part of zoological science (see Patterson 2002). Naming species on the basis of photos, vocalizations and molecular data alone opens the door for the official recognition of taxa that most zoologists are not ready to accept: on these grounds, Sasquatch is clearly a valid, nameable taxon for example.

To get back to the Highland mangabey, the problematic lack of a type specimen is now an academic argument, as a museum-accessioned specimen has been procured: it is a sub-adult male found dead in a trap in August 2005, currently accessioned at the Field Museum of Natural History, Chicago. It possesses the diagnostic features of the specimens previously described as representing this species, so seems to be securely and correctly identified (note here one of the practical problems of lacking a holotype: we don’t have a physical specimen that we can compare latterly obtained specimens to). The Chicago specimen is significant, as study of its DNA sequence data has provided new information on the affinities and evolution of the Highland mangabey, and on mangabeys and their relatives as a whole.

And, on that note, all will be revealed in the next post.

PS - for the latest news on Tetrapod Zoology do go here.

Refs - -

Beckman, M. 2005. Biologists find new species of African monekey (Lophocebus kipunji). Science 308, 1103.

Jones, T., Ehardt, C. L., Butynski, T. M., Davenport, T. R. B., Mpunga, N. E., Machaga, S. J. & De Luca, D. W. 2005. The Highland mangabey Lophocebus kipunji: a new species of African monkey. Science 308, 1161-1164.

Landry, S. O. 2005. What constitutes a proper description? Science 309, 2164.

Mendes Pontes, A. R., Malta, A. & Henrique Asfora, P. 2006. A new species of capuchin monkey, genus Cebus Erxleben (Cebidae, Primates): found at the very brink of extinction in the Pernambuco Endemism Centre. Zootaxa 1200, 1-12.

Moser, M. 2005. Holotypic ink. Science e-letters

Patterson, B. D. 2002. On the continuing need for scientific collecting of mammals. Journal of Neotropical Mammalogy 9, 253-262.

Polaszek, A., Grubb, P., Groves, C., Ehardt, C. L. & Butynski, T. M. 2005. What constitutes a proper description? Response. Science 309, 2164-2166.

Smith, E. F. G., Arctander, P., Fjeldså, J. & Amir, O. G. 1991. A new species of shrike (Laniidae: Laniarius) from Somalia, verified by DNA sequence data from the only known individual. Ibis 133, 227-235.

Timm, R. M., Ramey, R. R. & The Nomenclature Committee of the American Society of Mammalogists. 2005. What constitutes a proper description? Science 309, 2163-2166.


Anonymous Anonymous said...

Darren, would you really recommend killing a member of a critically endangered species solely to provide a haplotype?

Doug M.

1:28 PM  
Anonymous Anonymous said...

Totally separate question from the previous: in the era of modern molecular biology, should type collection be the only way to satisfy the ICZN?

The case of the BB bush-shrike is apposite, since Smith and Amir collected moult feathers and a blood sample. These, in turn, provided enough genetic material to establish to a high degree of confidence that this was indeed a new species.

They also photographed the bird, videotaped it, and recorded its songs. Multiple witnesses saw the bird during its period of captivity. Again, careful observation strongly supported the characterization of a separate species.

Short of collecting the bird, it's hard to see what else they could have done. And collecting it, given its obvious extreme rarity, clearly involved grave conservation issues.

Smith and Amir have been ferociously attacked ever since. But the bulk of the attacks seem to be on the basis of issues that are peripheral to the central question. For instance, a number of commentators have said that the bird was obviously so close to extinction that one individual more or less could hardly make a difference. Meanwhile, others have claimed that a captive specimen would have encouraged support for conservation. Putting aside that these seem quite weak, I don't think they address the question: what, if anything, should satisfy, short of an actual specimen?

(Tacking back to the case of the bush-shrike, my feeling on that case is that part of the outrage is a perception that Smith and Amir somehow got away with something by claiming a new species without having an actual, physical bird in the hand. Well, that and simple jealousy. N.B., I have spent a fair amount of time around both birders and ornithologists, so that last point is not snark.)

Anyway, the question is: if I have molecular evidence, photos, video, competent witnesses in controlled circumstances, feathers, fur, droppings, shed skin, what have you... at what point would you say, okay, it's enough? Or would nothing but an actual specimen do?

I realize that's a very fraught question. I'm just wondering what you think.

Doug M.

1:46 PM  
Blogger Darren Naish said...

Response to El Paleofreak: as I stated in the blog (it's the bit of text marked with *), a stem hominoid would probably be regarded by people as a 'monkey'. But it wouldn't be a monkey in the strict, modern sense: viz, a member of Platyrrhini or Cercopithecoidea.

Response to Doug M: as I mentioned, there's a lot that could be said about this subject, and I'm relucant to start discussing it here for fear of writing too much. I personally wouldn't 'like' it if a member of an endangered species were killed, even for science.

But doing so isn't done 'solely' to provide a holotype: it's to establish beyond question the reality of the species, and note that conservation efforts can only begin once a species is accepted as valid. I agree with you that the sort of data collected by Smith and colleagues on the Bulo Burti boubou may well be adequate in order to characterise and establish the species, but if we go down this road there is the problem that data accepted as satisfactory by some may not be regarded as satisfactory by others, and chaos could ensue.

Furthermore, in many cases we don't really know whether a new species is truly endangered: this needs to be shown by way of further study, and, again, that study can only begin once the species is officially recognised.

But it's a difficult issue. I'll devote a blog to it some time. Thanks for your thoughtful and informed comments.

2:46 PM  
Anonymous Anonymous said...

Iit is indeed an interesting issue. I'd welcome a post on it, as your time permits.

Which brings up a completely unrelated question. Why do you refer to blog posts as blogs? For instance, when you say "I'll devote a blog to it some time", it's obvious from context that you mean you'll write and post a single article about it, not that you'll create an entire new weblog devoted to this issue. (Although stranger things have happened, to be sure.)

My understanding is that the noun "blog" refers to the entire weblog, while single article on the blog is a blog post or, more usually, a post.

Apparently "article" was the earlier usage, but "post" -- a term borrowed from Usenet -- quickly overtook it. Perhaps because there's less danger of confusion with other sorts of articles, such as news or academic.

I don't know if this usage has been formalized, but I suspect it has. In any event, it is consistent across the entire blogosphere. So, using "blog" instead of "post" is a bit confusing, like saying "book" when you mean "page".

Am I missing something?

Doug M.

11:32 AM  
Blogger Darren Naish said...

You're right Doug, I'm stupid (I must confess that I'm actually pretty naive about the blogosphere - I'm too wrapped up in my interests to notice what other people are saying I suppose). I'll make the appropriate changes some time. Thanks, seriously.

12:01 PM  
Anonymous Anonymous said...

Oh, no worries. You are very welcome. And no need to go back and change it!

Doug M.

3:29 PM  
Anonymous Anonymous said...

"Response to El Paleofreak: as I stated in the blog (it's the bit of text marked with *), a stem hominoid would probably be regarded by people as a 'monkey'. But it wouldn't be a monkey in the strict, modern sense: viz, a member of Platyrrhini or Cercopithecoidea."

But using that logic, you could say snakes didn't evolve from lizards. After all, a stem-serpentine wouldn't be a lizard in the strict, modern sense: viz, a member of Iguania, Anguimorpha, Lacertoidea, Scincoidea, Gekkota, Dibamia. Which doesn't make sense, obviously. Because in both cases, you're forcing the paraphyletic group to be di/polyphyletic for some reason. The platyrrhine-cercopithecoid common ancestor, and stem-catarrhines should surely be called monkeys too. Just as the iguanian-serpentine common ancestor, stem toxicoferans, stem episqamates, etc. should be called lizards.

9:56 PM  
Blogger Darren Naish said...

Response to Mickey... (firstly, thanks for reading, and for leaving comments).

No, the situation with 'monkeys' and stem hominoids etc isn't quite like that. The term 'monkey' is used for members of two separate clades, Platyrrhini and Cercopithecoidea.

But contrary to what El Palaeofreak (and you) have said, the common ancestor of these clades, and the plesions along the platyrrhine and catarrhine stems would not necessarily be identified as 'monkeys'. I've never read of stem-group anthropoids (like Amphipithecus and Pondaungia described as monkeys for example (they were probably more like tarsiers in appearance), and probable stem catarrhines like eosimiids have also (to my knowledge) not ever been identified as 'early monkeys'. Again, these taxa seem to have been more like tarsiers (IIRC).

So the platyrrhine-cercopithecoid common ancestor, and stem-catarrhines, should NOT 'surely be called monkeys': instead, monkey is an unofficial term for crown-group platyrrhines and crown-group cercopithecoids, and - given that hominoids did not descend from cercopithecoids - hominoids did not, actually, descend from a catarrhine that we should properly imagine as a monkey.

I hope that makes sense, sorry if it seemed a bit rambling. I'm trying to watch Big Brother (the shame).

11:11 PM  
Anonymous Anonymous said...

That severely strict zoological jargon tends to forbid every "X descends from Y" sentence. It's bringing us an inconscient fixism ;o)

Please, let's call the ancient monkeys "monkeys", at least in the scientific divulgation. Let's say we do descend from monkeys (not extant ones, of course) and from apes. Because certainly we descend from animals that have all the features of monkeys and apes, and deserve those names.

Transformism, come back! ;o)

(excuse my English, please)

11:28 AM  
Anonymous Anonymous said...

7:51 AM  

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