Are Primates over-split?

Carlos M

Well-Known Member
Hi.

So, since my order of the Illustrated Checklist of the Mammals of the World has not arrived yet (and I have no idea when it will do), I have been reading a lot about mammalian taxonomy and I have noticed that there seems to be a case of conflict when it comes to primates: it seems that they are over-split. Is it true? I'd like to hear some opinions, and maybe get a picture of how primate taxonomy has changed over the years and when the number of species increased dramatically.
 
Hi.

So, since my order of the Illustrated Checklist of the Mammals of the World has not arrived yet (and I have no idea when it will do), I have been reading a lot about mammalian taxonomy and I have noticed that there seems to be a case of conflict when it comes to primates: it seems that they are over-split. Is it true? I'd like to hear some opinions, and maybe get a picture of how primate taxonomy has changed over the years and when the number of species increased dramatically.

Primates are indeed gigantically oversplit currently, which is because a large number of primate taxonomists follows a very narrow version of the Phylogenetic Species Concept. This was basically started by Colin Groves with his 2001 book "Primate Taxonomy". In the 20th century primates tended to be very overlumped. It makes very little sense to recognize only 1-2 species of baboon, 1-2 species of sifaka and only 2 mouse lemurs. The current trend to name new species everywhere is however a complete overreaction. Often when the validity of splits is checked using a proper amount of genetic data, the evidence seems to disappear (multiple titi species, woolly monkeys, sportive lemurs). In particular the Malagasy primates and the S-American ones are extremely oversplit and it would be highly necessary to clean up the mess here....
 
Primates are indeed gigantically oversplit currently, which is because a large number of primate taxonomists follows a very narrow version of the Phylogenetic Species Concept. This was basically started by Colin Groves with his 2001 book "Primate Taxonomy". In the 20th century primates tended to be very overlumped. It makes very little sense to recognize only 1-2 species of baboon, 1-2 species of sifaka and only 2 mouse lemurs. The current trend to name new species everywhere is however a complete overreaction. Often when the validity of splits is checked using a proper amount of genetic data, the evidence seems to disappear (multiple titi species, woolly monkeys, sportive lemurs). In particular the Malagasy primates and the S-American ones are extremely oversplit and it would be highly necessary to clean up the mess here....

Splits seem to be the favor of late, resulting in a lot of questionable species. But if you can split it, you get to name it. That seems to be some of the motive, particularly in primates where a lot of splits are named after someone.
I have nothing against splits if there is enough data to back it up, but I feel it's gotten a little out of hand. I was looking up Red Colobuses the other day and it seemed like everybody had a different number of recognized species. Highly confusing to try and sort out.
Not a primate, but some of the Empidonax flycatchers here in North America could be lumped back together in my opinion. Pacific-slope and Cordilleran Flycatchers are literally undistinguishable in the field, range and male song are the only real clues. Out of range birds are basically never identified because it's nearly impossible. I feel relumping the two under Western Flycatcher really isn't a bad option.
 
Not a primate, but some of the Empidonax flycatchers here in North America could be lumped back together in my opinion. Pacific-slope and Cordilleran Flycatchers are literally undistinguishable in the field, range and male song are the only real clues. Out of range birds are basically never identified because it's nearly impossible. I feel relumping the two under Western Flycatcher really isn't a bad option.

Male songs are actually one of the characteristics that make something potentially splittable, as it is a hint towards speciation. When females are not turned on by the sound of other populations there is a barrier to interbreeding.
 
I suspect that many of the splits are not justified. When I did an 'Evolution' course at college, the lecturer highlighted the problem of gulls. In the UK, the lesser black-backed gull and herring gull are separate species. The herring gull also occurs westwards across North America, while the lesser black-backed gull occurs eastwards across Europe and Asia. The gulls meet again in Siberia, where they are the same species.

There was also the definition that two separate species are not supposed to produce viable offspring together. That is also not the case.

If alien taxonomists came to Earth, Homo sapiens would be split into several species, while the domestic dog would be split into several genera.
 
New I suspect that many of the splits are not justified. When I did an 'Evolution' course at college, the lecturer highlighted the problem of gulls. In the UK, the lesser black-backed gull and herring gull are separate species. The herring gull also occurs westwards across North America, while the lesser black-backed gull occurs eastwards across Europe and Asia. The gulls meet again in Siberia, where they are the same species.

There was also the definition that two separate species are not supposed to produce viable offspring together. That is also not the case.

There is a push here to split our herring gull from yours, on what grounds I don't remember. Splitting our mew gull from your common gull has also been considered. Although Thayer's got relumped into Iceland a couple years ago now, so who knows really.

And speaking of confusing and controversial splits/lumps, let us not forget the albatrosses, petrels, and shearwaters.
 
Hi.

So, since my order of the Illustrated Checklist of the Mammals of the World has not arrived yet (and I have no idea when it will do), I have been reading a lot about mammalian taxonomy and I have noticed that there seems to be a case of conflict when it comes to primates: it seems that they are over-split. Is it true? I'd like to hear some opinions, and maybe get a picture of how primate taxonomy has changed over the years and when the number of species increased dramatically.
Primates are indeed gigantically oversplit currently, which is because a large number of primate taxonomists follows a very narrow version of the Phylogenetic Species Concept. This was basically started by Colin Groves with his 2001 book "Primate Taxonomy". In the 20th century primates tended to be very overlumped. It makes very little sense to recognize only 1-2 species of baboon, 1-2 species of sifaka and only 2 mouse lemurs. The current trend to name new species everywhere is however a complete overreaction. Often when the validity of splits is checked using a proper amount of genetic data, the evidence seems to disappear (multiple titi species, woolly monkeys, sportive lemurs). In particular the Malagasy primates and the S-American ones are extremely oversplit and it would be highly necessary to clean up the mess here....
Yes, as lintworm says, Primates are heavily oversplit nowadays. Especially in the lemurs and some of the New World monkeys, but also to a lesser extent in the Old World monkeys. The tarsiers and gibbons are the worst for splitting (going from heavily over-lumped to [probably] heavily over-split), but it has also been happening in the langurs more commonly. Macaques have largely escaped the splitting frenzy but not entirely.
 
When I did an 'Evolution' course at college, the lecturer highlighted the problem of gulls. In the UK, the lesser black-backed gull and herring gull are separate species. The herring gull also occurs westwards across North America, while the lesser black-backed gull occurs eastwards across Europe and Asia. The gulls meet again in Siberia, where they are the same species.
This is often still repeated today as an example of a ring species, but the idea was originally based on flawed evidence.

See this 2004 paper on why it isn't a ring species: https://www.researchgate.net/publication/8454571_The_Herring_Gull_complex_is_not_a_ring_species
 
Yes, primates are oversplit, to the point that zoos, which have live animals at hand, cannot distinguish 'species' without genetic tests. This happened in gibbons, capuchins, sakis, slow loris and mouse lemurs, among others. There was even an interesting proposal that most male white-faced sakis in zoos are different species than most females.

Among others, hybrids betwen these 'species' are regular in the wild, whenever 'species' meet.
 
I have a copy of 'The Mammals' by Desmond Morris (1964). It was one of my favourite books when I was a little boy as it had a list of all mammal genera.

Here is a comparison between the number of primate species listed in the Lynx 'Illustrated Checklist of Mammals of the World' (L) and the Morris Book (M). As the classification of families differs, I'll just list the genera

Microcebus: L 25; M 2 (including Mirza coquereli)
Mirza: L2; M 0 (M coquereli included in Microcebus)
Allocebus: L1; M 0 (A trichotis included in Cheirogaleus)
Cheirogaleus: L9; M 3 (including Allocebus trichotis)
Phaner: L4; M 1
Lepilemur: L 26; M 2
Hapalemur: L 5; M 2 (including Prolemur simus)
Prolemur: L 1; M 0 (Prolemur simus included in Hapalemur)
Lemur: L 1; M 6 (most species now incorporated in Eulemur and Varecia)
Eulemur: L 12; M 0 (3 species included in Lemur)
Varecia: L 2; M 0 (V variegatus included in Lemur)
Avahi: L 9: M 1
Propithecus: L 9; M 2
Indri: L 1; M 1
Daubentonia: L 1; M 1
Euoticus: L 2; M 2
Galagoides: L 3; M 0 (G demidovi included in Galago)
Paragalago: L 5; M 0
Galago: L 4; M 4 (including species now in Galagoides, Sciurocheirus and Otolemur)
Sciurocheirus: L 3; M 0 (S alleni included in Galago)
Otolemur: L 2; M 0 (O crassicaudatus included in Galago)
Arctocebus: L 2; M 1
Perodicticus: L 3; M 1
Loris: L 2; M 1
Nycticebus: L 9; M 2
Cephalopachus: L 1; M 0 (C bancanus included in Tarsius)
Carlito: L 1; M 0 (C syrichta included in Tarsius)
Tarsius: L 12; M 3 (2 species now in Cephalopachus and Carlito)
Cebuella: L 2; M 1
Callibella: L 1; M 0
Mico: L14; M 0 (M argentatus included in Callithrix)
Callithrix: L 7; M 8 (including Mico argentatus)
Callimico: L 1; M 1
Leontopithecus: L 4; M 3
Leontocebus: L 10; M 21 (including species now in Saguinus)
Saguinus: L 12; M 0 (several species were included in Leontocebus)
Aotus: L 11; M 1
Saimiri: L 8; M 2
Sapajus: L 7; M 0 (S apella was included in Cebus)
Cebus: L 15; M 3 (included Sapajus apella)
Cheracebus: L 5; M 0 (C torquatus was included in Callicebus)
Callicebus: L 5; M 8 (including species now placed in Cheracbus and Plecturocebus)
Plecturocebus: L 17; M 0 (P moloch, cupreus, brunneus and cinerescens were included in Callicebus)
Pithecia: L 16; M 2
Chiropotes: L 5; M 2
Cacajao: L 4; M 3
Alouatta: L 12; M 6
Ateles: L 7; M 4
Lagothrix: L 2; M 3
Brachyteles: L 2; M 1
Macaca: L 23; M 12
Cynopithecus: L 0; M 1 (C niger now placed in Macaca)
Cercocebus: L 7; M 4 (includes species now placed in Lophocebus)
Mandrillus: L 2; M 2
Lophocebus: L 2; M 0 (L albigena and L aterrimus were included in Cercocebus)
Rungwepithecus: L 1; M 0
Papio: L 6; M 5
Theropithecus: L 1; M 1
Allenopithecus: L 1; M 1
Miopithecus: L 2; M 0 (M talapoin was included in Cercopithecus)
Erythrocebus: L 3; M 1
Chlorocebus: L 7; M 0 (C aethiops was included in Cercopithecus)
Allochrocebus: L 3; M 0 (A lhoesti was included in Cercopithecus)
Cercopithecus: L 19; M 10 (includes species now placed in Miopithecus, Chlorocebus and Allochrocebus)
Colobus: L 6; M 3 (includes species now included in Piliocolobus and Procolobus_
Piliocolobus: L 17; M 1 (P badius was included in Colobus)
Procolobus: L 1; M 1 (P verus was included in Colobus)
Presbytis: L 19; M 13 (includes species now placed in Semnopithecus and Trachypithecus)
Pygathrix: L 3; M 1
Rhinopithecus: L 5; M 2
Simias: L 1; M 1
Nasalis: L 1; M 1
Semnopithecus: L 8; M 0 (S entellus and johni were placed in Presbytis)
Trachypithecus: L 20; M 0 (T pileatus, obscurus, francoisi and cristatus were placed in Presbytis)
Hoolock: L 3; M 0 (H hoolock was included in Hylobates)
Hylobates: L 9; M 7 (includes species now placed in Hoolock, Nomascus and Symphalangus)
Symphalangus: L 1; M 0 (S syndactylus was included in Hylobates)
Nomascus: L 7; M 0 (N concolor was included in Hylobates)
Pongo: L 3; M 1
Gorilla: L 2; M1
Pan: L 2; M 2
Homo: L: Not included; M 1
 
Splits seem to be the favor of late, resulting in a lot of questionable species. But if you can split it, you get to name it. That seems to be some of the motive, particularly in primates where a lot of splits are named after someone.
I have nothing against splits if there is enough data to back it up, but I feel it's gotten a little out of hand. I was looking up Red Colobuses the other day and it seemed like everybody had a different number of recognized species. Highly confusing to try and sort out.
Not a primate, but some of the Empidonax flycatchers here in North America could be lumped back together in my opinion. Pacific-slope and Cordilleran Flycatchers are literally undistinguishable in the field, range and male song are the only real clues. Out of range birds are basically never identified because it's nearly impossible. I feel relumping the two under Western Flycatcher really isn't a bad option.
I still can't figure out what the rationale is for keeping Black-capped and Carolina Chickadees separate is, especially since in the overlap zone ~60% of the birds are hybrids. That sure sounds like a lump to me. I think a lump of Common and Hoary Redpolls would be justified as well.
 
Thanks for all of your responses. I agree that tarsiers and gibbons seems to be the worst cases of over splitting in primates. I am surprised that the primate taxonomy proposed by Groves was much more accepted than his Ungulate Taxonomy
 
There is a push here to split our herring gull from yours, on what grounds I don't remember. Splitting our mew gull from your common gull has also been considered. Although Thayer's got relumped into Iceland a couple years ago now, so who knows really.

White-headed gulls are incredibly complex due to recent speciation and widespread second contact hybridization. In one of the most extensive studies on Larus gulls (Sonsthagen et al 2016), which had a very decent phylogenetic analysis using multiple tree-building methods, argentatus smithsonianus grouped with californicus and not with European argentatus argentatus/argenteus. If correct, North American Herring Gulls are thus more related to California Gulls than to European Herring Gulls, which would make it a valid spit.

The same study found that canus canus and canus brachyrynchus where distinct, but the authors did not conclude both should be elevated to species status. Furthermore, thayeri is distinct from but most related to glaucoides, but whether that justifies a split is not further discussed in the paper.

In general, North American Larus gulls are more problematic than Eurasian ones, with more mtDNA lineage sharing between taxa. But keep in mind that Larus gulls are a difficult group of phylogenetic analysis and morphology does not always reflect genetic relationships (e.g. Norwegian "hyperboreus" group with northern European argentatus argentatus/argenteus, and not with North American hyperboreus).

And yes, I should return to the photographic guide of shorebirds! :p
 
The opinion seems to be that currently species are oversplit. Is that opinion supported by evidence based research at all?

My impression has been yes more splitting in general (allthough in f.i. Panthera the line of thought in some quarters seems to go the other route into lumping morphologically/taxonomically distinct subspecies into one or two) of late.

I just wonder whether truth be told there is a middle ground of truth or dare I say our focus is out of sync with recent DNA / taxonomy rearrangements here? Sample sizes are very important in genetics research and on the basis of small sample size it is easy to fall into the trap of easy conclusions.

BTW: Just the other week another new SE Asian species of langur was described from Myanmar.
 
In order to decide whether they are over split, the first question must be what is meant by a species? At one extreme, anything that could hybridise with anything else is regarded as being in the same species; which would reduce 160 or so waterfowl species to a mere handful at most! At the other extreme, one could regard anything that shows the minimal detectable difference that could lead to eventual full reproductive isolation as being a species (or at least an incipient species). Both of these are biologically meaningful, but may have very different outcomes for conservation for example. Super lumped forms might be easier to maintain in captivity, but meaningful local adaptations could be lost. Super split forms run the risk of inbreeding (which may not be as problematic in the wild if rare hybridisation does occur) and the more forms need captive breeding, the more difficult it becomes to maintain all forms.
My personal preference is to know that forms exist, but to realise that not all of them will have the same Biological merit. From a field naturalist viewpoint, I like “species” which can be recognised, but understand that there is no reason why all meaningful Biological species should be easily identifiable to a human observer.
 
Titi monkeys seem to have been oversplit. There were 8 species in 1 genus in 1964 and there are 27 species now.

Lepilemur species have increased from 2 to 26.
 
As so many of you have pointed out, splitting/lumping is highly subjective, based on how systematists interpret data. It seems that naming of thought-to-be unique taxa (nomenclature/taxonomy) is separate to how valid those taxa are as taxonomic units (phylogenetics/systematics). It may be important for conservation that xyz population of lemur is threatened with extinction, but is it really important to know whether it's a species, subspecies, or none of the above? If we can differentiate it from its nearest relatives, consistently or otherwise, from physical, genetic, or other characters, then its surely just an academic discussion, right?

I'm working on a project involving working out valid taxonomic names for carnivoran taxa. There have been at least 4000 described carnivorans (some invalidly) since 1758, although the vast majority have been lumped with other taxa. Many of these will be because our understanding has evolved, and we know that a black leopard is just a differently-coloured leopard and not a separate entity, for example, or Tigris regalis is just another name for the Bengal tiger, but one of the "tigris"s was changed due to the thought that you couldn't have a Tigris tigris.

But so many other names have been "lost" because somewhere along the line, one interpretation of data is that b, c, d were synonymised with a. But the person who described b, c, and d might one day be proven right, and the systematist who revised the systematics could be proven wrong.

Case in point: the IUCN Cat specialist group revamped Felidae systematics, lumping a great many subspecies that have been accepted for many years. As new evidence comes in, it's bound to happen that some of their decisions will be reversed. But if I want to know the valid scientific name of the Asiatic lion, it's no longer Panthera leo persica, as Meyer described it more than two centuries ago, but Panthera leo leo. Never mind that it has clear physical differences from other members of that subspecies - Meyer's P. l. persica is valid as it complies with the code for zoological nomenclature (ICZN). One day, perhaps after it's too late, it might come to light that the distinctiveness of Asiatic lions will be reconfirmed.

My point is that perhaps we should not be too centred on whether a population with definable characters is a phylogenetic entity, as that is bound to change.

To bring the topic back to primates, in my opinion, it doesn't really matter that they might be oversplit, so long as we don't lose what might be one day proven to be a valid entity in lumping previous splits. That's the danger in overlumping - if we were to go back to the days where there was only one orangutan, gorilla, and chimpanzee, we might not care so much about the plights of the Tapanuli orangutan, Cross River gorilla, or western chimpanzee as much. There isn't such a danger with oversplitting, so I'd rather oversplit and be wrong than overlump and be very wrong!
 
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To bring the topic back to primates, in my opinion, it doesn't really matter that they might be oversplit, so long as we don't lose what might be one day proven to be a valid entity in lumping previous splits. That's the danger in overlumping - if we were to go back to the days where there was only one orangutan, gorilla, and chimpanzee, we might not care so much about the plights of the Tapanuli orangutan, Cross River gorilla, or western chimpanzee as much. There isn't such a danger with oversplitting, so I'd rather oversplit and be wrong than overlump and be very wrong!

The danger is there, conservation and regulation mostly concerns species (there are cases with subspecies, but that is a minority). Oversplitting species might give a completely wrong view of how many species there actually "are" and might divert scarce funding away from good species to just another isolated population of Klipspringer that is slightly more yellowish. Plenty of research has been done to show these dangers.
 
Following Dassie Rat post, it seems as well that there is a considerable increase in the number of primates' genus recognized. Despite my lack of biology background, most genus level splits seems to make sense for me but I would be interested to see if this opinoon is shared or not.
 
The danger is there, conservation and regulation mostly concerns species (there are cases with subspecies, but that is a minority). Oversplitting species might give a completely wrong view of how many species there actually "are" and might divert scarce funding away from good species to just another isolated population of Klipspringer that is slightly more yellowish. Plenty of research has been done to show these dangers.

Indeed, but if we don't really know yet there aren't multiple klipspringer species due to a lack of data (or at least it could be argued that we don't yet know), we might lose this diversity before we can be sure. Obviously there are dangers either way with either extreme, but I still prefer oversplitting to overlumping (if I must choose to be one or the other).
 
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