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Old World monkeys of choice

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Old World monkey montage. From left to right: male Geladas (one at Edinburgh Zoo, one at Howletts Wild Animal Park), Lion-tailed macaque, Guereza. Photos by Darren Naish.

There have never been enough primates on Tet Zoo. That isn’t because I’m not interested in primates, nor because I don’t think about primates, or look at primates, that much… in fact, I probably think about, and look at, primates more than I do any other group of animals… it’s simply because – as is the case for so many tetrapod groups – I’ve just never yet had the time to do them justice. I recently went to Howletts Wild Animal Park (Kent, UK) where I watched, and photographed, several primates I haven’t seen much, or at all, before. And here we have a good excuse to talk, briefly, about a select few primates. Let’s see how it turns out.

Lion-tailed macaques engaging in a key component of Old World monkey behaviour. Photo by Darren Naish.

I’ve never seen a Lion-tailed macaque or Wanderoo Macaca silenus until my recent trip to Howletts. Unique to the Western Ghats in India, the Lion-tailed macaque is black-bodied and has a silvery mane and tufted tail-tip… you’d think that the mane would be the key feature of interest, not the tail, but, whatever. It’s substantially more arboreal than other macaques and is a generalist herbivore/omnivore that feeds on a variety of plants, also occasionally eating birds and their eggs.

Macaques are a large and complex group of monkeys and there have been various efforts to determine their phylogeny and evolutionary history. It seems that the Lion-tailed macaque is especially close to the Southern pig-tailed macaque M. nemestrina, the two belonging to a wholly east Asian group that also includes the Celebes crested macaque C. nigra (Morales & Melnick 1998, Li et al. 2009). Lion-tailed macaques are among the world’s most threatened primates, with the world population currently numbering less than 3500. The spread of agriculture and the construction of dams, reservoirs and roads across their range has led to them being increasingly split into smaller, more fragmented populations.

Lion-tailed macaques are among the most arboreal of all macaques. But, hey, they still forage on the ground on occasion. Photo by Darren Naish.

I won’t talk about the Celebes crested macaque (or Celebes black macaque, or Sulawesi crested macaque) right now since there’s so much to say about them.

Captive Guereza - no thumbs, nice tassels, weird nose. Photo by Darren Naish.

This is a Guereza or Mantled guereza Colobus guereza, the best known of the five or so Colobus species, sometimes collectively termed the pied or black-and-white colobuses. There are several different views on the taxonomy and phylogeny of this group (e.g., Groves 2007). Excepting the all-black Black colobus C. satanas, all are black and white, often with long white tassels on their shoulders, flanks and tails and white regions on their faces and thighs. The entire radiation is exclusively African and largely limited to the tropical and subtropical forests of central Africa. Like other colobines, pied colobuses have multi-chambered stomachs and are forest-dwelling herbivores that subsist mostly on leaves, though they also eat fruit, flowers, seeds and buds. The name ‘colobus’ comes from the French ‘colobe’ or the Greek ‘kolobos’ (take your pick) meaning ‘mutilated’, an idea inspired by the reduction of the thumb to just a small tubercle. Thumbs are over-rated: they’ve been lost or reduced in gibbons and spider monkeys, too.

Guereza from the side to show long fringe of white fur. Image by Darren Naish.

Black and white colobuses have peculiar noses where soft tissue forms an elongate, almost human-like proboscis that approaches the upper lip (cough cough add appropriate comment about human-type nose and AAH cough cough). They’re large, reaching about 1 m in total length and with males exceeding 20 kg. Like so many monkeys, they’re flamboyant, showy animals. They live in small social groups dominated by large, highly vocal males who advertise their presence with loud nocturnal and dawn roaring choruses. They are threatened by habitat loss and hunting for both their pelts and meat.

Female Geladas doing the 'lip flip' thing. Photo by Darren Naish.

These [above] are female Gelada Theropithecus gelada engaging in an aggressive encounter with other members of their group, hence their weird appearance: they’re performing the ‘lip flip’ behaviour unique to this species. Male Geladas are splendid monkeys with long, flowing manes and long, tufted tails. The species is endemic to the central Ethiopian highlands.

The former diversity of Theropithecus: there's a whole book devoted to it (Jablonski 1993).

Geladas are among the most remarkable of monkeys, being highly specialised for terrestrial grazing. Closely related to baboon-mangabeys, baboons and the Kipunji Rungwecebus kipunji (Davenport et al. 2006), just about all of their anatomical peculiarities are related in some way to their terrestriality, grazing lifestyle, or ‘shuffle feeding’ form of behaviour. These include especially tall-crowned teeth, proportionally elongate forearms and thumbs, larger bodies and especially flexible elbows and wrists [Jonathan Kingdon has proposed that the human lineage went through a similar so-called 'squat foraging' phase in its evolution. Interesting, but not widely accepted]. The presence of colourful skin patches on the chest, used in advertising sexual status, has also been linked to shuffle feeding (Kingdon 1997), since the fact that they spend most of their time sat on their rumps means that the buttocks and genitals are not on display as they are in most other monkeys. Indeed, the buttock pads (or ischial callosities, whichever you prefer) are black, not red. An oft-mentioned fact about Geladas is that the living species is the only surviving member of a once more widespread group that contained several species, some of which were substantially bigger than the living one.

Male Gelada, Howletts Wild Animal Park. Don't call them 'gelada baboons', since they're not baboons. Photo by Darren Naish.

As usual, there is tons more I’d like to say. But my time is up. All of the primates discussed here are members of the Old World monkey group Cercopithecidae.

For previous Tet Zoo articles on Old World monkeys and other primates, do check out…

Yawning Mandrill (Mandrillus sphinx) at Paignton Zoo, photo by Darren Naish.

Refs – -

Davenport, T. R. B., Stanley, W. T., Sargis, E. J., De Luca, D. W., Mpunga, N. E., Machaga, S. J. & Olson, L. E. 2006. A new genus of African monkey, Rungwecebus: morphology, ecology, and molecular phylogenetics. Sciencexpress 10.1126/science.1125631

Groves, C. P. 2007. The taxonomic diversity of the Colobinae in Africa. Journal of Anthropological Sciences 85, 7-34.

Jablonski, N. G. 1993. Theropithecus: The Rise and Fall of a Primate Genus. Cambridge University Press, Cambridge, UK.

Li, J., Han, K., Xing, J., Kim, H. S., Rogers, J., Ryder, O. A., Disotell, O. A., Yue, B. & Batzer, M. A. 2009. Phylogeny of the macaques (Cercopithecidae: Macaca) based on Alu elements. Gene 448, 242-249.

Kingdon, J. 1997. The Kingdon Field Guide to African Mammals. Academic Press, San Diego.

Morales, J. C. & Melnick, D. J. 1998. Phylogenetic relationships of the macaques (Cercopithecidae: Macaca), as revealed by high resolution restriction site mapping of mitochondrial ribosomal genes. Journal of Human Evolution 34, 1-23.

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!

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





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  1. 1. ekocak 12:36 pm 05/28/2014

    Colobus satanus, eh? Because it’s all black, or some other reason?

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  2. 2. naishd 12:48 pm 05/28/2014

    It eats babies and turns into a goat at night.

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  3. 3. irenedelse 1:32 pm 05/28/2014

    That would make an awesome new cryptid!

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  4. 4. naishd 1:54 pm 05/28/2014

    I screwed up: it’s satanas, not ‘satanus‘. It’s sometimes called the Satanic colobus, it seems just because it’s black.

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  5. 5. naishd 1:56 pm 05/28/2014

    ps — Tet Zoo ver 3 just passed the 10,000 comments mark. This comment is # 10,003 (this not counting spam or deleted comments, obviously). Woo-hoo, pass the champagne.

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  6. 6. ekocak 2:19 pm 05/28/2014

    Wait, so the joke comment you made was the 10,000th?

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  7. 7. naishd 2:20 pm 05/28/2014

    Ah. Yes.

    So, let’s remember this factoid for future reference… Q: What was the 10,000th Tet Zoo (ver 3) comment? A: “It eats babies and turns into a goat at night.”

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  8. 8. Heteromeles 2:52 pm 05/28/2014

    The internet shall remember!

    More seriously though,I’m wondering about Gelada social structure and Dunbar’s Number, which is an assertion that the complexity of primate social lives is related to the size of their cortex. Dunbar didn’t use Geladas in his seminal paper (Dunbar, RIM. 1991. Neocortex size as a constrain on group size in primates. Journal of Human Evolution (1992) 20,469-493), but it sure looks like they’d be an outlier, being more social than predicted by the size of their neocortices. What’s up with that?

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  9. 9. josimo70 3:38 pm 05/28/2014

    Many monkeys’ species were baptized after “demoniac” names: Callicebus moloch, Colobus satanas, Ateles belzebuth, Alouatta belzebul. Other ones refer to Iliad characters: Macaca rhesus, Trachypithecus pyrrhus, Semnopithecus ajax, Semnopithecus hector, Semnopithecus priam, S.entellus.

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  10. 10. irenedelse 5:29 pm 05/28/2014

    The name ‘colobus’ comes from the French ‘colobe’ or the Greek ‘kolobos’ (take your pick) meaning ‘mutilated’, an idea inspired by the reduction of the thumb to just a small tubercle.

    French speaker here. The name of these monkeys is indeed ‘colobe’ in French, but my dictionary tells me that it comes from the scientific Latin ‘colobus’, constructed after the Greek ‘κολόβός’ (kolobos), meaning ‘shortened’ or ‘mutilated’. The Latin form ‘colobus’ was first used in Illiger’s Prodromus systematis mammalium et avium (1811), which predates the French usage by 25 years.

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  11. 11. SJCrum 7:41 pm 05/28/2014

    The simple reason that each of the monkeys shown are odd-looking is because God used monkeys in the beginning before humans were even on earth so the ability to stand up on two feet could be tested BEFORE humans were here.

    The process had one enormous bad thing that was also occurring during that time. Satan and the very violent, killing types of animals then were killing all of the very defenseless monkeys. So, the monkeys were changed in their appearances so they could survive, and still be involved in the extremely important testing.

    The monkeys with non-monkey-animal-looking tails were far safer. And, monkeys with very-concealing furry manes and odd hair groups sticking out all over, actually laughed at the predator types as they cluelessly strolled on by.

    So, that is why the very odd-looking monkeys were odded that way.

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  12. 12. naishd 9:47 pm 05/28/2014

    I see.

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  13. 13. Christopher Taylor 10:47 pm 05/28/2014

    Full credit, at least, for the successful utilisation of ‘odd’ as a verb.

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  14. 14. Heteromeles 11:04 pm 05/28/2014

    Lion-tailed macaques engaging in a key component of Old World monkey behaviour.

    So what do New World monkeys do it in place of this key behaviour?

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  15. 15. ectodysplasin 1:32 am 05/29/2014

    Fling feces, mostly.

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  16. 16. irenedelse 3:20 am 05/29/2014

    Wow, the ‘Ratites in trees’ article is the 3rd most read among SciAm blogs? (Looking at the side bar.)
    Congrats, Darren!

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  17. 17. naishd 5:47 am 05/29/2014

    I live to give :) Now only if I could get Nature Blog Network to stop letting Mongabay pass for a ‘blog’.

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  18. 18. Tayo Bethel 12:11 pm 05/29/2014

    Maybe you should do an article series on on the Old World monkey radiation. And another on the New World monkey radiation … and watch the comments roll in.
    Perhaps the size of the Gelada neocortex reflects their diet more than their social structure, since grass is not the easiest of foods to digest and is relatively low energy as compared to, say, fruit.

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  19. 19. Heteromeles 1:42 pm 05/29/2014

    @Tayo: If you’re referring to Dunbar’s work and the Gelada neocortex, it’s worth reading the paper referenced above (it’s available here). IIRC, he tested a number of explanations for group size including diet, and the size of the neocortex had the best fit. I should point out that humans didn’t have quite the largest group size, and that it’s not nearly as linear a relationship as one might hope. Still, the Geladas seem to be unusually social if you think Dunbar got it right, and that may say something interesting either about their brains and/or their social structure.

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  20. 20. John Harshman 5:34 pm 05/29/2014

    As far as under-concentration on primates goes, need I point out that every single post in this blog was written by a primate, as were all the comments? To redress the balance, perhaps you could solicit a guest-post from some archosaur.

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  21. 21. Christopher Taylor 9:39 pm 05/29/2014

    John, what did you think “I probably think about, and look at, primates more than I do any other group of animals” was in reference to?

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  22. 22. Mark Robinson 11:19 pm 05/29/2014

    … as were all the comments.
    John, that is an assumption. [woof]

    While I’m here – “Old World monkey radiation”, is that what caused them to mutate into apes?

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  23. 23. irenedelse 2:23 am 05/30/2014

    @Mark Robinson:

    “While I’m here – “Old World monkey radiation”, is that what caused them to mutate into apes?”

    Either that or Satan, obviously. 0:)

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  24. 24. John Scanlon FCD 3:43 am 05/30/2014

    Re comments on group size, isn’t it best explained by distribution of critical resources? I assume everyone is familiar with Hans Kummer’s Primate Societies, but maybe I shouldn’t. After all, it’s nearly 30 years since I read it myself.

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  25. 25. Jerzy v. 3.0. 5:00 am 05/30/2014

    But which goat? And can a trained zoologist spot a demonic goat not being a correct species?

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  26. 26. Jerzy v. 3.0. 5:08 am 05/30/2014

    @colobus
    I pity 19. century zoologists who saw only distorted skins and pickled specimens. So they relied on characters like missing thumb and saw ‘satanic’ faces on rather distinctive and beautiful animals.

    @Dunbar
    Actually, I would put food resources as the first determinant of social structure. Do locusts have bigger brains than orangutans?

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  27. 27. naishd 5:52 am 05/30/2014

    Interesting comments from everyone, thanks indeed. I, for one, haven’t read Kummer’s Primate Societies (comment # 24)… I really should.

    As for Dunbar’s ‘rule’, one response has been that we’re unusually keen to latch on to it as a reliable hypothesis because it fits in so well with our anthropocentric bias, a primary assumption behind those who champion it being that social complexity and levels of interaction are necessarily higher in primates with larger group sizes (and, according to some, this isn’t so). For a discussion of this area, see…

    Barrett, L., Henzi, P. & Rendall, D. 2007. Social brains, simple minds: does social complexity really require cognitive complexity? Proceedings of the Royal Society B 362, 561-575.

    I’m not saying or implying, by the way, that Dunbar’s hypothesis is invalid, but it should be seen as a generalisation, not as a rule… would it be as well received as it has (it’s even used as a business model by advisers and company consultants, and ‘Dunbar’s Number’ has made its way into popular culture as a sort of guide to how many friends humans can ‘really’ have) if it were applied to other social animals? Note that Dunbar himself says that it doesn’t apply across the board, since gibbons and orangutans (for example) both live in groups massively smaller than those they ‘should’ have according to neocortex size.

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  28. 28. Heteromeles 9:42 am 05/30/2014

    re: Dunbar’s number. Yes indeed. Many humans also lived in smaller groups than implied by their cortex sizes. The Geladas appear to be the more interesting converse, with large groups and (proportionally) smaller brains than with humans. That’s another thing that makes them interesting.

    One critical note (@24) is that Dunbar’s Number only applies to primates. It doesn’t work with parrots, for example. As for locusts, since those swarms are cannibalistic within the swarm, I’m not sure that “social” applies to them at all, any more than it applies to zombies.

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  29. 29. irenedelse 10:22 am 05/30/2014

    @Heteromeles:

    “One critical note (@24) is that Dunbar’s Number only applies to primates. It doesn’t work with parrots, for example. As for locusts, since those swarms are cannibalistic within the swarm, I’m not sure that “social” applies to them at all, any more than it applies to zombies.”

    Was any comparison made with the more complex social insects (ants, bees, termites…), then? Do they have bigger brains than locusts?

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  30. 30. Heteromeles 3:40 pm 05/30/2014

    The issue is social complexity, not just the size of the mob. An ant colony isn’t hierarchical, rumors to the contrary. Actions are made by quorum-sensing, which a lot of things (including bacteria) do.

    Basically, if you’re in a species where it doesn’t matter what the individual next to you is (in ants or bees, in the same hive), then we’re not talking about social complexity. You can run that type of social system with a fairly small brain (as do bacteria, bees, ants, and so forth).

    Social complexity is when you have friends, enemies, and a dominance hierarchy of some sort. This type of complexity takes memory and processing power, and there’s typically an upper limit on how many individuals you can know and trust enough to work with.

    Dunbar’s number (100-200 people) crops up a lot in places like military units, the size of tribal villages, and so forth, and it’s a manageable size for a group to run itself through consensus in some form. Much above that level, and you need a hierarchy, managers whose job is just to talk with each other and solve problems between people who don’t know each other that well.

    Dunbar’s theory is that complex group size correlates directly with neocortex size, and he’s of the opinion (right or wrong) that big brains are a prerequisite for social complexity, more than for physical problem solving.

    I suspect Dunbar’s on to something, but that’s why the Geladas are interesting, because they might break his rule. They are reported to have a hierarchical social structure that goes as follows (per Wikipedia):
    –Reproductive units of 1-12 females, their young, and 1-4 males in a dominance hierarchy. The females live in a hierarchy within the unit, and reproductive units split up if they get too large. There are also all-male units of 2-15 males.
    –Bands of 2-27 reproductive units and “several” all-male units.
    –Communities of 1-4 bands whose home ranges overlap extensively
    –Temporary herds of up to 60 reproductive units, sometimes from multiple bands.

    So what’s their Dunbar number? From the level of bands, it could be as high as 300-400, if they actually all know each other in the band. It could be as low as, say, 12, if you look at the reproductive units and the female hierarchy. It’s an interesting question, and it does get at how they organize. At some point, are the great Gelada herds “mobs,” or is there a hierarchy within the herd that helps them make decisions?

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  31. 31. irenedelse 5:33 pm 05/30/2014

    @Heteromeles:
    That’s really fascinating. Thank you.

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  32. 32. LeeB 1 7:26 pm 05/30/2014

    The questions regarding Gelada bands presumably also apply to Mandrills (and Drills?) which also form large hordes of hundreds of individuals at times.
    These are less well known because they live in dense forest where they are hard to see.

    LeeB.

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  33. 33. DavidMarjanovic 9:08 am 05/31/2014

    At some point, are the great Gelada herds “mobs,” or is there a hierarchy within the herd that helps them make decisions?

    Also, do they make decisions? But that may not be relevant to the question of what their Dunbar number is. Maybe they’re just less xenophobic than humans?

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  34. 34. Jerzy v. 3.0. 11:35 am 05/31/2014

    @30
    I was going to comment, but didn’t want to side-track – that Dunbar’s “size of primatre social unit” is very subjective number. First it depends from hunting pressure, habitat quality and research intensity. Then, primate groups can form big bands, and species living in small groups, it can be argued that neighboring groups know each other.

    In non-humans primates, biggest groups are formed by monkeys – geladas, mandrills, also snub-nosed monkeys and possibly squirrel monkeys. Even macaques and baboons form groups much bigger than apes.

    And right remark that birds defy most of primatologists’ theories about brain size, intelligence and sociality.

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  35. 35. irenedelse 11:59 am 05/31/2014

    @Jerzy:
    “Then, primate groups can form big bands, and species living in small groups, it can be argued that neighboring groups know each other.”

    Just thinking out loud here, but it may be enough for a gelada to identify to which group the members of neighbouring groups belong, without knowing them as well as the members of its own group. Which is something humans do all the time: rely on categories to keep in our heads knowledge about large numbers of items, be it members of our species, living things or even inanimate objects. I wouldn’t be surprised if geladas, mandrills and other Old World monkeys used this kind of heuristics: a) individual monkeys of its own group, b) monkeys known to be from a few neighboring groups, c) other stuff.

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  36. 36. Dartian 4:09 am 06/1/2014

    Darren:
    a generalist herbivore/omnivore that feeds on a variety of plants, also occasionally eating birds and their eggs.

    Larger prey, too; lion-tailed macaques sometimes prey on giant squirrels:

    Sushma, H.S. & Singh, M. 2008. Hunting of Indian giant squirrel (Ratufa indica) by the lion-tailed macaque (Macaca silenus) in the Western Ghats, India. Current Science 95, 1535-1536.

    Don’t call them ‘gelada baboons’, since they’re not baboons.

    Some would disagree with that. Theropithecus is certainly very closely related to Papio (they hybridise fairly frequently in the wild, for example) – probably more closely related than Pan is to Homo.

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  37. 37. irenedelse 12:14 pm 06/1/2014

    @Dartian:
    Well, if “baboon” is reserved to the genus Papio, it can’t apply to geladas. Unless, in regard to the easy hybridization thing, their status changes one of these days.

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  38. 38. BrianL 1:22 pm 06/1/2014

    Certainly, geladas (and mandrills and drills) sometimes being referred to as baboons is related to them originally being classified in the genus *Papio*? If so, one would suppose that the term ‘baboon’ has simply become more restricted over time as cercopithecid classification became more rigorous, meaning that a term like ‘gelada baboon’ is archaic rather than ‘wrong’?
    Being always very impressed by and a bit frightened of large primates, the thought of hundreds of mandrills congegrating sounds very scary to me, though not as scary as a large troup of chimpanzees with hunting on their mind.

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  39. 39. irenedelse 2:17 pm 06/1/2014

    @BrianL:
    “one would suppose that the term ‘baboon’ has simply become more restricted over time as cercopithecid classification became more rigorous, meaning that a term like ‘gelada baboon’ is archaic rather than ‘wrong’?”

    That was my impression too. I may remember incorrectly but I do think there used to be mentions of ‘gelada baboons’ in natural history books. Similarly, mandrills (and drills) were classified as part of the baboon group, and in the genus Papio.

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  40. 40. vdinets 12:32 am 06/2/2014

    Common names don’t have to reflect taxonomic changes. If Papio and Theropithecus form a monophyletic group, it’s perfectly OK to call them all “baboons”, especially considering that these two genera are likely to be re-lumped once oversplitting falls out of fashion. Drills, on the other hand, are nested within mangabeys, so calling them baboons would be a bit misleading.

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  41. 41. Dartian 3:02 am 06/2/2014

    Irtenedelse:
    Well, if “baboon” is reserved to the genus Papio, it can’t apply to geladas.

    Goodman et al. (1998) included the geladas in Papio, treating them only as subgenerically distinct.

    the easy hybridization thing

    As it happens, one of the first researchers to report such natural hybridisation events was Robin Dunbar of the already mentioned ‘Dunbar number’ fame (Dunbar & Dunbar, 1974). (Incidentally, that same year Robin Dunbar published his PhD-thesis – on the social structure of geladas. So yeah, he’s aware of the social peculiarities of this species.)

    More generally speaking, considering primate vernacular names: Genera are, of course, conventions created by people and in that sense, they are ‘meaningless’ entities. However, there are several vernacular primate names that are used for members of different ‘genera’ – for example, ‘langur’, ‘colobus’, ‘marmoset’, ‘tamarin’ and ‘lemur’. Thus, in principle there is really no reason why ‘baboon’ couldn’t be used in such a wider sense too (note that that’s not something that I’m arguing for myself).

    Vladimir:
    once oversplitting falls out of fashion

    Anything is possible, I suppose, including the return of a taxonomic Dark Age. However, if I were you I wouldn’t hold my breath. ;)

    References:

    Dunbar, R.I.M. 1974. The social organisation of the gelada baboon (Theropithecus gelada). PhD thesis, University of Bristol.

    Dunbar, R.I.M. & Dunbar, P. 1974. On hybridization between Theropithecus gelada and Papio anubis in the wild. Journal of Human Evolution 3, 187–192.

    Goodman, M., Porter, C.A., Czelusniak, J., Page, S.L., Schneider, H., Shoshani, J., Gunnell, G. & Groves, C.P. 1998. Toward a phylogenetic classification of primates based on DNA evidence complemented by fossil evidence. Molecular Phylogenetics and Evolution 9, 585-598.

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  42. 42. naishd 5:36 am 06/2/2014

    On “geladas are not baboons” – I don’t wish to seem like a taxonomy nazi or anything, but I would say that modern views on monkey phylogeny and diversity do indeed require us to say that “geladas are not baboons”. Ok, so they were called ‘baboons’ in the past… but that doesn’t mean much, given that language constantly evolves, conventions change, and all manner of inaccurate (now thought to be wrong) terms were used for animals in the past (should we also still refer to the Caracal lynx or Simien jackal?, for example).

    As for what phylogenetic studies say… ok, so it’s possible to find recent-ish studies that include geladas within Papio (see above citation to a 1998 publication). However, I think I’m right in saying that more recent studies consistently find Theropithecus and Papio to be distinct, either as sister-taxa (Finstermeier et al. 2013) or with Theropithecus as less close to Papio than is Lophocebus, the crested mangabeys, or Rungwecebus, the Kipunji (Harris 2000, Davenport et al. 2006, Roberts et al. 2010, Perel et al. 2011). If the latter trees are correct, it would obviously be a poor decision to call geladas ‘baboons’, given that this would mean making crested mangabeys and kipunjis ‘baboons’ too. My conclusion is that the term baboon should be strictly restricted to Papio.

    Having said all this, fossils and molecular dating indicate that the taxa concerned are, as Dartian says, only recently diverged – they all seem to have diverged within the last six million years or so – and hence an argument could be made that they all ‘should’ be included within the same ‘genus’ (whatever the hell a ‘genus’ is). Never forget that these sorts of decisions are still wholly subjective. On that note, what works best, in terms of communicating what we mean? I say that retaining those separate ‘generic’ names does, and the use of the common names mentioned above.

    Refs – -

    Davenport, T. R. B., Stanley, W. T., Sargis, E. J., De Luca, D. W., Mpunga, N. E., Machaga, S. J. & Olson, L. E. 2006. A new genus of African monkey, Rungwecebus: morphology, ecology, and molecular phylogenetics. Science 312, 1378-1381.

    Finstermeier, K., Zinner, D., Brameier, M., Meyer, M., Kreuz, E., Hofreier, M. & Roos, C. 2013. A mitogenomic phylogeny of living primates. PLoS ONE 8(7): e69504. doi:10.1371/journal.pone.0069504

    Harris, E. E. 2000. Molecular systematics of the Old World monkey tribe Papionini: analysis of the total available genetic sequences. Journal of Human Evolution 38, 235-256.

    Perelman, P., Johnson, W. E., Roos, C., Seua´nez, H. N., Horvath, J. E., Moreira, M. A. M., Kessing, B., Pontius, J., Roelke, M., Rumpler, Y., Schneider, M. P. C., Silva, A., O’Brien, S. J. & Pecon-Slattery, J. 2011. A molecular phylogeny of living primates. PLoS Genetics 7(3): e1001342. doi:10.1371/journal.pgen.1001342

    Roberts, T. E., Davenport, T. R. B., Hildebrandt, K. B. P., Jones. T., Stanley, W. T., Sargis, E. J. & Olson, L. E. 2010. The biogeography of introgression in the critically endangered African monkey Rungwecebus kipunji. Biology Letters 6, 233-237.

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  43. 43. Dartian 6:20 am 06/2/2014

    Darren:
    should we also still refer to the Caracal lynx or Simien jackal?, for example

    In context of the present discussion, a more pertinent question would perhaps be whether we should still refer to both Lophocebus and Cercocebus as ‘mangabeys’, considering that they are surely not closely related. Which one of these taxa gets to retain the old vernacular, and which will be changed?

    it would obviously be a poor decision to call geladas ‘baboons’, given that this would mean making crested mangabeys and kipunjis ‘baboons’ too

    Why would it be “obviously poor”, really? Don’t you think that it would better reflect our understanding of papionin evolution and phylogenetics (and thus, ultimately, science communication) to call them all baboons? (Note that, as I said earlier, I’m not arguing that we should call them all ‘baboons’ – I’m just interested in your line of reasoning in this matter.)

    My conclusion is that the term baboon should be strictly restricted to Papio.”

    Personally, I agree with that. I just wanted to point out that the question “What is a baboon?” is not entirely unambiguous.

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  44. 44. naishd 6:40 am 06/2/2014

    Thanks, Dartian. On mangabeys… Since mangabey non-monophyly was discovered (or, more widely appreciated, or demonstrated.. or however you want to put it), the trend has been to distinguish the two lineages. The Lophocebus species are the crested mangabeys or baboon-mangabeys; the Cercocebus species are the white-eyed mangabeys or drill-mangabeys.

    As for why it would (in my opinion) be a poor decision to include crested mangabeys and kinpunjis within the moniker ‘baboon’, this is because ‘baboon’ refers to a set of predominantly terrestrial, dog-faced monkeys with especially big canines. Crested mangabeys and kipunjis do not conform to this ‘archetype’ and do not fall into our expectation of what a baboon ‘should’ be*. I happily admit that all of this is subjective and that trying to apply phylogenetic rationality to the use of common names is doomed to failure, but I’m sure you see my point. And I also get the point that geladas are more or less ‘baboons’ under the archetype concept that I just outlined…

    * Let’s not mention the fact that one study found some kipunjis to be part of Papio, ha! This is most likely due to hybridisation events.

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  45. 45. irenedelse 7:03 am 06/2/2014

    “this is because ‘baboon’ refers to a set of predominantly terrestrial, dog-faced monkeys with especially big canines”

    This is where I feel a Venn diagram may help. baboons have dog-like faces, but not all monkeys with dog-like faces are baboons, since it also describes the two Mandrillus species… Which are themselves nested within the mangabeys (vdinets #40).

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  46. 46. Tayo Bethel 12:48 pm 06/2/2014

    This is why people come to TetZoo … the comments are as interesting as the articles.

    Is the PHD thesis on gelada social structure open access anywhere?
    As for the baboon archetype:
    we could include in the description the distinctive”"wahoo” call of adult male baboons. As far as I know,Geladas do not make this vocalization. Does anyone know if mandrills and drills do?

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  47. 47. naishd 5:40 pm 06/2/2014

    Oh, I forgot to express my surprise at the paper on squirrel-hunting behaviour in Lion-tailed macaques (see Dartian’s comment # 36). Wow. Balakrishnan (2010) cites other papers that list frogs, lizards, bats and other squirrel species as prey items of Lion-tailed macaque…

    Balakrishnan, P. 2010. Predation of eggs and nestlings of pigeons (Columbidae) by the lion-tailed macaque Macaca silenus in the Western Ghats, India. Indian BIRDS 6, 167–168.

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  48. 48. Heteromeles 8:56 pm 06/2/2014

    @41: Thanks for the info on Dunbar and the Geladas. My impression from the Dunbar Number paper is that he was relying on brain section masses from other work. For whatever reason, he didn’t include data on Gelada. I don’t know if it’s because the brain data weren’t available, or because the complexity social structure is ambiguous enough that it’s hard to say what the maximum group size is. IIRC, he didn’t include drills or mandrills either.

    It’s worth noting here that Dunbar’s number is about maximum practical group size, not real group size in a particular situation. I agree that reality can isolate many beings. The better arguments against Dunbar’s number aren’t “oh, well, Orangutans are solitary, so therefore Dunbar’s wrong,” because Dunbar’s Number is all about upper limits on group sides. The better argument is it’s “oh, these pinheaded monkeys form huge groups and their inter-monkey politics are reminiscent of the US House of Representatives (in other words, they have complex social systems, with cliques, hierarchies, and deception, and their brains are relatively tiny). The more examples we find of such primates, the weaker Dunbar’s argument is.

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  49. 49. John Harshman 11:35 pm 06/2/2014

    This talk of what is or is not a baboon or a mangabey, about which I have no opinion whatsoever, does remind me of my campaign to stop calling U.S. tanagers by that name, because they aren’t.

    I saw a spectacular male Western piranga the other day, by the way. There, doesn’t that sound better?

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  50. 50. vdinets 12:28 am 06/3/2014

    John Harshman: as long as you are after tanagers, you could also go after “sparrows”, “orioles”, “robins” and “flycatchers”, to name a few ;-)

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  51. 51. Christopher Taylor 2:33 am 06/3/2014

    should we also still refer to the… Simien jackal?

    We were supposed to have stopped?

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  52. 52. Dartian 9:43 am 06/3/2014

    Darren:
    The Lophocebus species are the crested mangabeys or baboon-mangabeys; the Cercocebus species are the white-eyed mangabeys or drill-mangabeys.

    Hardly a particularly elegant solution to this problem, alas.

    Christopher:
    We were supposed to have stopped?

    The Simien ‘jackal’ is actually a ‘wolf’ – it’s more closely related to Canis lupus than it is to jackals.

    But then again, ‘jackal’ is a phylogenetically dubious vernacular name too. The golden jackal is more closely related to the wolves than it is to the other ‘jackals’, and the black-backed jackal and the side-striped jackal aren’t close relatives of each other either. In fact, as their relatedness to wolves, coyotes and golden jackals may be more distant than that of both Lycaon and Cuon they probably shouldn’t even be included in Canis at all (instead, they should probably be placed in the separate genera Thos and Schaeffia, respectively).

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  53. 53. naishd 9:45 am 06/3/2014

    The obvious lesson is to give up on vernacular names entirely and stick only to scientific ones. After all, they’re consistent, never-changing and set in stone [/massive joke].

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  54. 54. DavidMarjanovic 10:30 am 06/3/2014

    That’s fascinating; I had no idea of these developments in canid phylogeny.

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  55. 55. Dartian 12:03 pm 06/3/2014

    Christopher: If you want to learn more about the interrelatedness of the various ‘wolves’, I suggest you go and read this interesting blog post. ;)

    David:
    I had no idea of these developments in canid phylogeny.

    Hate to disagree with you, but yes you had. ;) Evidence of that is here. (Although there you say that Thos is not an available name for the black-backed jackal.)

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  56. 56. DavidMarjanovic 7:27 am 06/4/2014

    That’s scary. Like… I’ve never been drunk in my life.

    Probably I forgot about the whole story because “Thos” is unavailable according to nothing less than Opinion 417.

    The direct link to my comment is here, and the phylogeny we were talking about is this, which shows a clade of (Ethiopian wolf (golden jackal (coyote (grey wolf, dog)))) that excludes Cuon and Lycaon and can be called Canis without any problems that I can see.

    Link to this

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