Laws of Archaeology and anatomy allow for bone fragments to restore the structure of the skull. It allows us to estimate the ratio of the volume of the brain and different parts of the brain. With regard to brain function "in the structure" the issue is more acute (a lot of hypotheses). Testing hypotheses about the functions of the brain helps artifacts that exhibit near the ancient skeletal remains. Rock carvings, prototype tools ...

I am not sure I got your question wright.

Are you asking why the canines and premolar have the shape and size they have in Homo nalendi? Or is there any particular deviation in the general shape or certain ratios from other Hominids that you are addressing in particular?

I think that both issues are important.

Hi Carl. Thanks for following my question...

Basically i mean the size of the premolars compared to canines. Canines used less? Was less rending function? if so, it could be due to processing food or simply to his diet?

What would be your hypothesis about its dental morphology?

Regards.

Dear Jaime,

Canines are the most dimorphic tooth in hominids. Females tend to have considerably smaller canines. Regarding their size, when dealing with disarticulated comingled remains from collective contexts one question we have to make is could those small canines belong to females or is this sub-sample mainly composed by females? If so, small canines are a result of a statistical deviation. Continuation of the works in the cave will probably produce a larger sample and hopefully articulated individuals which can be sexed.

However, for most results available so far, the individuals present fairly gracile dentitions. Post canine dentition looks large to most of us because we are used to the anatomically modern dentition. According to the taxonomic description of the species, when compared to other extinct hominids, those teeth are bot robust.

Best,

Claudia

Dear Claudia:

Always a pleasure to read you.                                                  

I agree with you, and we don't know the exact datation of the remains yet.... 

Data, diet, diseases, tools, associated fauna, etc. And we'll have to wait a few months to learn more about all that. 

Best regards.

Dear Jaime

The discourse on H. Naledi between specialists is highly controversial. And as long as no absolute dating exists, many questions will remain unanswered.

Best Kurt

Thanks Kurt.

It is a very interesting and intriguing event. We can only wait for news. 

Regards.

Dear Jaime,

There will be great news coming from the Rising Star Cave in the following years. They have but started the work. The excavated area is minimal and I bet they will continue digging and in a few years that will be the best known hominid sample in all Old World.

Dating of the materials is being approached using different methods, once technically they are not as easy to date as the materials from East Africa due to sedimentary conditions. BUT there is hardly a technical difficulty you cannot overcome when you have different scientist working on techniques to study such an important collection. Sooner or later some physics geek will come up with a solution. Independently of the dates (early homo? late homo?)... even if it is a conglomerate produced during an extremely long period of time... even if not all belong to the same species, no one can deny its value as a rich set of osteological data. 

Paleoanthropologists usually have big egos and anyone who discovers an impressive sample is asking to take fire from all sides, particularly when this person is as flamboyant as Dr. Berger. His work will have flaws as anyone else's, but as any scientific work it is in the academic publications we have to question them. That's how we make science: though constructive debate in the proper forum.

I would just like to point that:

(I) All data from this work is OPEN ACCESS, contrary to the secrecy that surrounds/surrounded older paleoanthropological works. This is a plus. It is science at reach of everyone within rich institutions in the first world OR NOT. We don't have to pay the industry of paid journals to check the data;

(II) Berger is not working alone, but has DOZENS of professionals from different fields working with the context and materials. The ones I know well (fellow colleagues working on teeth) are exemplary professionals, including THE most experienced dental anthropologist when it comes to African teeth;

(III) This collection is an unparalleled treasure and will be source of information decades after Berger and his opponents have retired/met their creator.

Best,

Claudia

... And it is always a pleasure to interact with dedicated colleagues and fine gentlemen as you, Jaime! :)

Have a great week!!!

Claudia

I agree very much my dear Claudia. I think we are about to witness the rest of the iceberg!

Best.

Jaime.

Dear all. All research should dominate objectivity. This position is especially needed in history, archeology, anthropology. Subjectivity authorities often "back away" researchers in an interesting mystery, but not science.

Hi all, I think we have to await detailed comparisons of naledi's dentition with other animals & hominoids: occlusal surface & relief, enamel thickness & micro-wear, etc.

Prof.Berger & team have done a great job in excavating these fossils & in describing & making them available to everyone, but (as so often in paleo-anthropology) their interpretations are anthropocentrically biased: Homo or Australopithecus naledi were no direct ancestors of ours, they were not buried deliberately in the cave, they were certainly no distance runners, and no tool-makers more than chimps are.

From a comparative viewpoint, these fossils are no mystery (in fact predicted, e.g. see our 2002 paper "Aquarboreal ancestors?" Trends Ecol Evol 17:212-7, 2002, google aquarboreal): apparently, naledi was a bonobo-like forest-swamp or wetland wader, feeding on aquatic herbaceous vegetation (AHV) such as papyrus sedges, frogbit, waterlilies etc. (google: bonobo wading). They fossilised in stagnant water (mud-stone). The curved hand-bones suggest vertical climbing in the branches above the swamps, like bonobo, orangutans & lowland gorillas do. The rel.long 1st & 5th fingers were less for tool use than for collecting floating AHV or for surface-swimming. The broad pelvises (iliac flaring & long femoral necks) were for sideward movements of the legs (femoral abduction): for climbing or swimming, but not for running. The flat humanlike feet are more flamingo- (plantigrade wader) than ostrich-like (digitigrade runner): more for wading than for walking. The small front teeth & large cheekteeth can be expected with a wetland diet of AHV & possibly hard-shelled invertebrates (HSI).

Lowland gorillas often wade on 2 legs in forest swamps for sedges & other AHV (google: gorilla bai), but naledi apparently exploited this special niche habitually: no wonder there were no or few other macro-fauna near the naledi fossils. It was no deliberate burial (they had ape-sized brains), but a natural accumulation: when they died, their bodies sank into the mud, and afterwards the underground eroded to become a cave system. How exactly the mudstone ended up in the cave is unclear so far, but the remarkably complete skeletons suggest that when they died they got almost immediately covered by mud.

When living hominids (e.g. humans & bonobos) go back in time towards the Homo/Pan last common ancestor, human ancestors can be expected to show several more-bonobo-like features (e.g. shorter legs, smaller hip joints, smaller brain, no external nose), and bonobo ancestors to show some more-human-like features (e.g. shorter hands & arms, longer thumbs, no iliac lengthening, flatter feet, smaller front teeth, thicker enamel), IOW, naledi, in spite of some primitive-hominid Homo-like features in hands, feet & dentition, was very bonobo-like in nearly all the rest of its anatomy, and IMO more likely a close relative of bonobos or common chimps than of humans (google: researchGate marc verhaegen).

Preliminary conclusion: IMO naledi's rel.small canines are not unexpected for a diet of AHV.

Hi Marc,

That's a good hypothesis. Forgive me if the question sound silly,... haven't had time to read everything on the naledi site, but, is there any assessment of the environment surrounding the cave around the most probable time for those hominids? Again, without absolute dates it is hard to be certain, but is there anything (... polen?) on the sediment analysis of the deposits in the cave that point to a wet environment at the time of deposition?

Best,

Claudia

Hi Claudia, reconstructions of how caves formed & got filled is extremely difficult AFAIK. But the presence of remarkably-complete naledi skeletons in mudstone suggests rapid covering by mud in stagnant waters (as we predicted long ago, in the 1980s, even before the Ndoki gorillas in forest bais were described). I imagine them wading bipedally, swimming-floating at the surface collecting AHV with their fingers spread out, and climbing out of the swamp vertically, arms above the head, see attachment. They consumed most of the available AHV I'd think: there were no other animals eating these foods, so I'm not surprised they accumulated in the mudstone, without other macro-fauna being present. In the other caves, there were fossils of other mammals IIUC, but these haven't been described AFAIK. AFAIK, there are no pollen studies yet. Yes, it was certainly wet when naleid lived/died there: mudstone. Best  --marc

Very interesting your analysis Marc.

Just take time to learn more.

Your articles are very interesting. I'll read it calmly. I consider myself an aquatic ape, because I love to practice freediving.

Best.

Interessante pergunta Claudia. Muito Obrigado.

Yes, a lot of interesting things in the arguments of scientists. We look forward to further analysis of the specialists in this situation.

Thank a lot. A few further thoughts.

The locomotion is +- clear IMO (frequent vertical wading & climbing cf bonobos & lowland gorillas in wetlands, rather than running over dry savannas), the diet is less clear (AHV = parts of floating vegetation, but also hard-shelled invertebrates? we have to await studies of enamel thickness, macro-wear etc.), the geology is not at all clear to me (I'm no geologist), but AFAICS the mudstone seems to have slided sidewards & downwards with an average velocity of a few centimetres every thousand years.

I wonder whether the mud was oxygen-poor, and whether this was important for conserving the skeletons or for other features of the mud(stone), e.g. traces of pollen or roots?

I said flamingoes are plantigrade. I was wrong: they're digitigrade like most birds. But of course their feet are much flatter than those of ostriches (google).

Sorry, I meant micro-wear (instead of macro-, although naledi's macro-wear is also important of course). The English spelling, esp.of the vowels, is still sometimes confusing to me (Dutch-speaking): they write "i" and say "ay", and write "a" and say "ey".

Thank you Marc for the explanation some differences in Dutch-speaking and English spelling.  Because sometimes there are funny situations. As with the interpretation of the archaeological finds. For example, I know that the most valuable finds for archaeologists in the ancient dumps occur. And in regions where there were local disaster (Pompeii, for example). But no one will make a conclusion - let more landfills and disasters for future historians.

Hi Claudia, I seem to have missed your answer of Oct.29, sorry about that. I'm correspondig with a few geologists & cavers on naledi's fossilisation. They all say that deliberate burial is very unlikely (of course), and one of them wrote that there's (geological?) evidence of wetlands in the region in the Pliocene. The mudstone indicates that naledi fossilised in stagnant water. Based on comparative anatomy (naledi's curved hand-bones = vertical climbing, & flat humanlike feet = wading and/or swimming), I have no doubt that naledi lived in swamp forests and/or wetlands, parttime wading bipedally, surface-swimming-floating, climbing vertically, I guess like bonobos wading for waterlilies etc., but more frequently. Google e.g. "bonobo wading" & "aquarboreal".

Marc,

You assert that "human ancestors can be expected to show several more-bonobo-like features (e.g. shorter legs, smaller hip joints, smaller brain, no external nose)". None of these features are specifically bonobo characteristics. I would be interested to know what uniquely shared features exist between humans (Homo? australopiths?) and bonobos that would support their having a shared most recent common ancestor.

Genetically bonobos are not closer related to Homo than chimpanzees (Fischer A, Wiebe V,  Pääbo S, Przeworski M. 2004. Evidence for a complex demographic history of chimpanzees. Molecular Biology & Evolution 21 (5): 799–808. doi:10.1093/molbev/msh083. PMID 14963091.) But in terms of allometry bonobos are a bit more like australopithecines than chimpanzees are (for exmple the relation of the length of legs and the slender arms in comparison to their thorax).

Agreed, but generalized body form similarities other than those that are uniquely shared have no necessary phylogenetic or evolutionary implication. If bonobos are more closely related to chimps than humans then general body similarities with humans are uniformative. Some monkeys have body forms more like humans than great apes. If any great ape shared uniquely shared features is living and fossil hominids (Homo, australopith) it is the orangutan (which interestingly includes a mangrove environment in its habitat range - and apparently even more so in the past before habitat destruction of the lowlands took place.

I agree with you John Grehan; the overall similarities are not phylogentically informative. But they could (in  a limited sense) be ecologically informative. I assume that Marc's answer is more targeted at this similarity as he compares behaviors associated with certain environmental features (esp. aquatic ones).

Although given that early ancestors of the Homo-line and one of the two extant closest relative share some features could be meaningful. But in this regard many other fossil hominids and homonoids (like Sahelanthrous, Orrorin etc.) have to be taken into account.

Jens, Predicting ecology from morphology might be ecologically informative, but then it might not. Its tricky at best. There is a classic example of two lemur species with identical or very similar skeletal features, but completely different ecologies (don't recall the example off the cuff, but can look it up if challenged). And of course there is the classic arboreal behavior of goats, not something predictable from the skeleton. Problem is that the same morphology in different organisms may be associated with more than one kind of environment. I stick out on a limb a bit with the view the morphology determines where an organism can live rather than an ecology being the necessary cause of the morphology that allows that organism to live in a particular environment. One example of an alternative ecological hypothesis is the arboreal theory that argues for a primarily arboreal home base for early hominids even after they evolved the structurally bipedal form of locomotion. This theory predicts that behavior was present in neandertals, and in Homo at least to the time fire and ballistic weaponry was invented. There are structural features that are concordant with such persistent arboreal behavior, but 'proving' it is another matter. Same for any other ecological hypothesis based on morphology. I recall that the aquatic hypothesis, as originally proposed invoked various biological features but ignored their presence in other non-aquatic primates.

There are claims that Sahelanthropus is a hominid based on some extrapolations of skull morphology, but its marginal at best, and Sahelanthropus lacks hominid apomorphies. Even the lower jaw dentition does not match the upper jaw. Orrorin had least has some uniquely bipedal features, and thick dental cheek enamel - something that would be expected for a hominid. Its the only non-australopith fossil that I would currently regard as having some reasonable possibility of being an actual hominid.

John, "bipedalism" does no define a "hominid" or a "hominin": there are problems with both terms, e.g. gibbons are vertical, but only partly bipedal (when walking over branches), lowland gorillas & bonobos wading in forest swamps are bipedal (but maximally for 1 or 2 hours). It's likely that both chimps & gorillas had more bipedal ancestors (from which knuckle-walking evolved in parallel in Pan & Gorilla), and a lot of students (including myself) have independently argued that australopiths are closer relatives of chimps or gorillas than of humans (see my Hum.Evol.papers). The popular idea that australopiths ran over open African plains is far-fetched, anthropocentric & anatomically & physiologically impossible IMO (e.g. sweat = water + salt = scarce on savannas, and cursorial mammls are digiti- or unguli-, not plantigrade): if hominids were bipedal, it was not on the plain, but in the swamp, wetland, forest bai etc. It's usually assumed that ape->human = quadru-->bipedal = forest->plain, but savanna baboons are less bipedal than forest baboons: a direct transition from the forest to the plain would have produced a quadrupedal "ape" with a rel.smaller brain but larger teeth & thicker fur ("baboon paradox"). Living in swamp forests (where most early hominoid fossils have been found) would lead to more vertical "apes" such as Oreopithecus or australopithecines. I think it's time that paleo-anthropologists begin thinking in biological (darwinian) instead of anthropocentric ways.

Hi Jens, thanks for your answer. John, I didn't say that bonobos are our closest relatives (bonobos are evolutionarily closest to common chimps, of course), I said that bonobos might morphologically be closest to the hominid LCA (hominid = Pan, Homo, Gorilla + fossil relatives such as australopiths, vs pongid = Pongo + fossil relatives e.g. sivapiths), e.g. my 1994 Hum.Evol.paper concluded (and after 22 years it requires remarkably few corrections AFAICS, google "marc verhaegen human evolution" or so): "A review of the paleo-anthropological literature reveals no data that exclude the possibility that both gorillas and chimpanzees could have had australopith ancestors. Bipedalism is generally considered to be the shared feature that links australopithecines with humans, and there is no doubt that at least some of the australopith species were partial bipeds. But it has never been proven that the African apes’ unique locomotion (plantigrade knuckle-walking) could not have evolved from some kind of (“short”-legged) bipedalism. In fact, insofar as the fragmentary fossil material and the incomplete comparisons with extant apes allow, ontogenetic and morphological evidence tends to favour the hypothesis that the last common ancestor of Homo and Pan 8-4 Myr BP was a partially bipedal, gracile australopith with chiefly a mosaic of human and chimpanzee (esp. bonobo) features: low sexual dimorphism, minimal prognathism, slightly enlarged canines, non-protruding nasal skeleton, smooth ectocranium without crests, “small” brain with ape-like sulcal pattern, relatively non-flexed basicranium, intermediate position of foramen magnum, “short” forelimbs without knuckle-walking features, low ilia, (very) long femoral necks, “short” legs, (very) valgus knees, full plantigrady, longer and not very abductable halluces." The naledi fossils seem to be very close to this description.

Marc,

I agree that the term ‘bipedalism’ does not define a hominid without some specific criterion to distinguish hominid bipedalism from any other. It is quite possible that the ancestor of African apes had more bipedal locomotion, but whether it is likely or not remains to be seen. As for what a hominid is, I would include for Homo and australopiths (and a couple for Orrorin where known): long pubic ramus, ilium superoinferiorly short, ilium expanded posteriorly, iliac crest (tubercle) thickened, anterior inferior iliac spine knoblike, anterior inferior iliac spine near acetabulum, linea aspera present, spiral line connects to linea aspera, femur condyles unequal, femoral neck thickened inferiorly, femoral shaft outwardly angled, two tibial tubercles, lateral tibial facet concave, latearl prox. tibial facet level with medial facet, distinct angle at L5-S1.

I would be interested in your papers presenting phylogenetic evidence of a closer relationship between australopiths and African apes. I’ll look for that 1994 paper. If you provide cladistic evidence that humans, African apes, and australopiths share more uniquely shared features than any other great ape then I will certainly be interested. In my past research I had not been able to substantiate published claims for such evidence, but I appear not to have come across your work.

We seem to be in agreement about australopiths being forest dweller. My speculation is that the ancestral ground dwelling hominid (whatever taxonomic designation) was preceded by an obligate bipedal ancestor and when the transition from forest to ground living took place the ancestors brought down their housing as well.

Marc,

I cannot access your 1994 Journal of Human Evolution paper. Please send me a copy to [email protected] Also any other of your publications providing cladistic evidence of a human/australopith-African ape clade. Thank you.

John, please see in the attachment my Hum.Evol.papers (without illustrations though).  Different lines of evidence suggest that the East-African australopiths resemble Gorilla more than Pan, and South-African australopiths resemble Pan more than Gorilla (regardless of size), IOW, the South-African australopithes seem to be more related to Homo-Pan than to Gorilla, and the East-African australopiths more to Gorilla than to Homo-Pan (Pleistocene Homo went a very different way: brain expansion, external nose, very long legs etc., IMO due to littoral adaptations the last 2 Ma). IOW, the East-African australopiths evolved in parallel to the South-African ones (possibly due to the same Pleistocene climatic changes), schematically: afarensis->boisei // africanus->robustus. I guess the Pliocene E-Africans lived in the central forest+wetlands (Congo-Nile-Rift-Chad), and the S-Africans in the coastal forest+wetlands + Zambesi basin (the central forest & the coastal forest were usually separated by the Arid Corridor, see Jon.Kingdon). I just sent to your email address my recent view of the African ape & human evolutionary tree.

Marc,

The problem with a generalized statement of resemblance is that resemblance is not necessarily evidence of relationship since it is only uniquely shared resemblances that provide evidence (at least cladistic) of relationship. I looked over the text you provided but I did not see any phylogenetic analysis. There was a short list of various resemblances of certain features shared between (some) australopiths and African apes, but these were just statements with references, nothing to substantiate them and there is no incorporation of the extensive uniquely shared suite of characters between humans and orangutans.

1) If only resemblance was the best evidence of relationship, I'd followed the traditional view that East- & South-African australopiths were very close relatives of each other - which I do not (e.g. Hum.Evol.1994).

2) The traditional phyologenetic analyses start from the supposition that we had ape (e.g. chimp-like) ancestors, and that because australopiths are morphologically +-between apes & humans, australopiths were evolving into the human direction. This approach is not foolproof to parallal, convergent & even reverse evolutions (which are ubiquitous in darwinian evolution). In fact, both extant chimps & humans had the same LCA (probably some 5 Ma, see below) which was more australopith-like, and from which both chimps & humans evolved, but in very different directions (IMO due to Homo's Pleistocene littoral adaptations).  IOW, it was not ape->apith-human (tradtional supposition), but it was OT1H apith-like->chimp & OTOH apith-like->human. IOW, australopiths (lived 5-1 Ma) are primitive to both humans & chimps (who live today).

3) What you said above about resemblances between South-African australopiths & Pan, and OTOH between East-African australpiths & Gorilla, was only about my 1996 paper, which illustrated the general impressions of anthropologists (which are correct IMO (in tempore non suspecto), but which are not used in other analyses). My 1994 paper did describe numerically the differences between extant & extinct hominids.

4) The important point is that East-African australopiths (afarensis, aethiopicus & boisei) resemble Gorilla more than Pan, but the South-African ones (africanus & robustus) resemble Pan more than Gorilla (even without considering size). That implies a parallel evolution E.african australpiths cf Gorilla // S.African australopiths cf Pan. (This doesn't mean to suggest e.g. that extant gorillas descend from boisei.)

5) Of course, there are a lot of resemblances between Homo & Pongo (evolution is mosaic-like), e.g. Pongo retained other primitive features in common with Homo than e.g. Pan did. But there are at least as many resmblances between Pan & Homo. And if morphological & genetic data are seemingly in conflict, it's the DNA data (if there are enough DNA data available) that are correct. Pan's DNA resembles Homo's about 3 times more than Pongo's DNA does (individual genes can of course suggest other genetic relations, but this is to be suspected: evolution is mosaic-like).

(Not only DNA comparisons with OWMs & other mammals, but also geological data suggest that pongids & hominids split c 16 Ma or so (plate tectonics: orangs now live in Asia). A hominid/pongid split c 15 Ma suggests a Homo/Pan split c 5 Ma. This "late" split perfectly fits all fossil data on australpiths etc.)

Marc,

Response by your numbers

1) “If only resemblance was the best evidence of relationship,” – resemblance of one kind or another is all we have. I prefer uniquely shared characters as evidence of relationship.

2) “The traditional phyologenetic analyses start from the supposition that we had ape (e.g. chimp-like) ancestors, and that because australopiths are morphologically +-between apes & humans, australopiths were evolving into the human direction. “ Perhaps so, but Grehan & Schwartz (2009) did not start with that assumption.

“In fact, both extant chimps & humans had the same LCA (probably some 5 Ma, see below) which was more australopith-like, and from which both chimps & humans evolved, but in very different directions (IMO due to Homo's Pleistocene littoral adaptations).”

Yes that is the traditional view. Not supported by morphological evidence.

3) “My 1994 paper did describe numerically the differences between extant & extinct hominids.” I’ll see if I can get someone to provide me with a pdf which is what I need to assess your analysis.

4) But if the” resemblance” of various australopiths with gorillas or chimps is not cladistically supported then it may not represent anything phylogenetically.

5) “Of course, there are a lot of resemblances between Homo & Pongo (evolution is mosaic-like), e.g. Pongo retained other primitive features in common with Homo than e.g. Pan did. But there are at least as many resmblances between Pan & Homo.” Not correct according to the analysis by Grehan & Schwartz (2009). We found at least 28 well substantiated shared derived characters between humans and orangutans vs only a few between humans and African apes and a couple between humans and pan.

“And if morphological & genetic data are seemingly in conflict, it's the DNA data (if there are enough DNA data available) that are correct.” That’s a propaganda claim. Has no empirical foundation. Grehan & Schwartz (2009) ( and a couple of later papers) went over this in some detail.

“ Pan's DNA resembles Homo's about 3 times more than Pongo's DNA does (individual genes can of course suggest other genetic relations, but this is to be suspected: evolution is mosaic-like).”

Resemblance is only resemblance.

“(Not only DNA comparisons with OWMs & other mammals, but also geological data suggest that pongids & hominids split c 16 Ma or so (plate tectonics: orangs now live in Asia).”

There is no DNA evidence as such for the age of taxa other than an extrapolation of minimal fossil ages to actual ages. But no problem with pongids and hominids (Homo, australopiths, but not African apes) splitting at least 16 Ma.

“A hominid/pongid split c 15 Ma suggests a Homo/Pan split c 5 Ma. This "late" split perfectly fits all fossil data on australpiths etc.)” Only if Homo-Pan are sister taxa. Grehan & Schwartz (2009) present cladistic evidence that they are not.

John, I'm very sorry, in my answer of 2 hours ago, I seem to have mixed up my 1994 & 1996 papers... (becoming old?), but otherwise what I said was correct. There's no doubt that Homo & Pan are closest relatives, then Gorilla, then Pongo, then Hylobatids etc. Your mistake is in what you believe to be "uniquely shared characters": most "uniquely shared characters" in Homo & Pongo are primitive for (great) hominoids. If we follow your reasoning, Homo & Hylobatids are closest relatives: both have vertical spines. In fact, a vertical spine (what many paleo-anthropologists call "bipedalism") is primitive for all hominoids: bipedalism in australopiths is not "uniquely shared" with humans: a vertical & centrally placed spine is primitive for all hominoids: it's kept in gibbons & humans, evolved into quadrumanualism in Pongo, and in australopiths it evolved into knuckle-walking in parallel in Pan & Gorilla. Anyway, thanks for sending me Grehan & Schwartz (2009), which I'll read with great interest (although I think I know what you're going to say in it).

Marc,

“There's no doubt that Homo & Pan are closest relatives, then Gorilla, then Pongo, then Hylobatids etc. “

That’s certainly one assertion. Not one that is empirically supported.

“Your mistake is in what you believe to be "uniquely shared characters": most "uniquely shared characters" in Homo & Pongo are primitive for (great) hominoids. “

There is no empirical evidence for that assertion. All the characters for humans/orangutans are unique within the large bodied hominoids with lesser apes and monkeys as the outgroup. By definition these are cladistically not primitive for large bodied hominoids (humans and great apes).

“If we follow your reasoning, Homo & Hylobatids are closest relatives: both have vertical spines.”

But one has to look at the outgroup. If no monkeys have vertical spines then yes the condition may have to be considered as a putative apomorphy (but if not supported by a corroborating suite of apomorphies the vertical spine has to be regarded as phylogenetically uninformative about relationships between Homo and gibbons). 

“In fact, a vertical spine (what many paleo-anthropologists call "bipedalism") is primitive for all hominoids: bipedalism in australopiths is not "uniquely shared" with humans: “

We did not use ‘vertical spine’ as a character for humans and australopiths

“a vertical & centrally placed spine is primitive for all hominoids:”

If the fossil great ape taxa demonstrate that to be a fact that would be fine. What is the specific cladistic evidence and published where?

“ australopiths it evolved into knuckle-walking in parallel in Pan & Gorilla. “

I have not seen any definitive evidence for knuckle-walking in australopiths

“Anyway, thanks for sending me Grehan & Schwartz (2009), which I'll read with great interest (although I think I know what you're going to say in it). “

I’ll be interested in your comments for or against (presumably latter).

Cheers, John

Hi John,

1) DNA leaves no doubt: it's ((((Homo-Pan)Gorilla)Pongo)hylobatids)OWMs etc. We have to use the whole DNA, not isolated genes.

2) Parallel & convergent & even reverse evolutions are often underestimated (although to be expected in darwinian evolution): most "uniquely shared" Homo-Pongo resemblances are retained from the great hominoid LCA, and lost in Pan // Gorilla.

3) Most paleo-anthropologists think Pan is an outgroup to Homo-australopiths: they believe that the 1000s of australopith fossils are closer relatives of Homo than of Pan of Gorilla, and that the 5 spp of African apes (2 common chimp, 1 bonobo & 2 gorilla spp) left +-0 fossils: this is statistically absurd: it would require that 1 of 6 extant spp left +-all fossils, and 5 of 6 left almost no fossils. This is undarwinian anthropocentrism. My 1996 paper didn't require preasumptions about possible outgroups.

4)5) Most PAs do. But a vertical spine most parsimoniously evolved into brachiating gibbons, quadrumanual orangs, bipedally walking humans & (in parallel) KWing chimps & gorillas, IOW, the hominoid LCA was already vertical.

6) This does not require KWing apiths. It only requires a vertical hominid LCA (IOW, not only Homo, but also Pan & Gorilla had more vertical ancestors. Nevertheless, different spp of apith (anamensis, afarensis, boisei) do show incipient KWing traits (refs in my Hum.Evol.1994 paper).

7) For a more parsimonious view on Mio-Pliocene hominoids, please see attchement.

Best  --marc

Hi Marc,

1) DNA leaves no doubt: it's ((((Homo-Pan)Gorilla)Pongo)hylobatids)OWMs etc. We have to use the whole DNA, not isolated genes.

DNA (in the form of sequence morphology involving matching bases that sometimes do not occur in nature) does not do this. I find it to be just a propaganda claim. What do you invoke as empirical evidence to substantiate the claim? Arguing to use the whole of DNA is to invoke the law of large numbers – which is what the pre-cladistic pheneticists did.

2) Parallel & convergent & even reverse evolutions are often underestimated (although to be expected in darwinian evolution): most "uniquely shared" Homo-Pongo resemblances are retained from the great hominoid LCA, and lost in Pan // Gorilla.

Parallelism, convergence, and reversions can only be identified after having selected a particular phylogeny.

3) Most paleo-anthropologists think Pan is an outgroup to Homo-australopiths: they believe that the 1000s of australopith fossils are closer relatives of Homo than of Pan of Gorilla,

Sure they have this belief, but they do not have any evidence other than a presumption of DNA evidence (in the form of sequence morphology), and they certainly have none for fossils.

“and that the 5 spp of African apes (2 common chimp, 1 bonobo & 2 gorilla spp) left +-0 fossils: this is statistically absurd: it would require that 1 of 6 extant spp left +-all fossils, and 5 of 6 left almost no fossils. This is undarwinian anthropocentrism.”

It’s not absurd at all. The fossil record is extremely uneven in representation.

“My 1996 paper didn't require presumptions about possible outgroups.”

But outgroups are essential for any competent phylogenetic analysis that meets cladistic criteria. I agree that one does assume that taxa that are not part of the group being studies comprise the outgroup. In this case the assumption is that large bodied hominoids (great apes and humans and fossil taxa sharing same uniquely shared features) form a monophyletic group and that to assess relationships within that group one has to compare with the outgroup to establish shared derived features. Conceivably the outgroup is the rest of life, but in practice it is feasible only for relatively close taxa, in the case of Grehan & Schwartz (2009 it encompassed New and Old World monkeys as well as lesser apes – which is a very sizable sample compared to many studies that limit themselves to just the gibbons which is a relatively small and uniform group. Without outgroup based cladistics one is left with phenetics – a measure only of overalls similarity.

4)5) Most PAs do. But a vertical spine most parsimoniously evolved into brachiating gibbons, quadrumanual orangs, bipedally walking humans & (in parallel) KWing chimps & gorillas, IOW, the hominoid LCA was already vertical.

It may be only most parsimonious if the feature is demonstrated as being shared by all large bodied honminoids other than the extant great apes.

6) This does not require KWing apiths. It only requires a vertical hominid LCA (IOW, not only Homo, but also Pan & Gorilla had more vertical ancestors. Nevertheless, different spp of apith (anamensis, afarensis, boisei) do show incipient KWing traits (refs in my Hum.Evol.1994 paper).

Of course not seen your 1994 paper unfortunately, but I read other papers trying to make that argument, but they did not provide any actual evidence, rather it was an inference they applied to the morphology.

7) For a more parsimonious view on Mio-Pliocene hominoids, please see attachment.

Will take a look.

Cheers, John

Marc,

Interesting to read your speculations about former habitat associations in your 2011 paper. Not much I can say in any critical way since they are speculations.

I would, however, argue that it is not correct to assert “According to the biomolecular data, the ancestors of the extant hominoids split into the lesser apes and the great apes between 18 and 15 Ma” because such dates are minimal estimates only. This is because they are calibrated using sources such as fossils that can only provide minimal ages for taxa represented. Therefore all molecular extrapolations are minimum divergence estimates only. The correct statement would be that “extant hominoids split into the lesser apes and the great apes AT LEAST between 18 and 15 Ma.” The same also goes for any other molecular date calibrated by fossils.

Hi John,

1) If we take out certain genes, we can prove everything: genes can split very early & remain in the same body. Since most DNA is non-coding, the law of large numbers is correct here. And it fits all other data. Hominids & pongids split c 15 Ma (cf plate tectonics), and Homo & Pan split c 5 Ma (+-1 or 2 Ma, this has no influence on my view of ape & human evolution, which is based primarily on comparative biology).

2) Yes, but e.g. the traditional interpretation (ape->apith->human) neglects possible parallelisms in Pan & Gorilla. Generally the later apiths are more apelike than the earlier apiths (discussed in my 1994 paper). There were 2 wet regions in Pliocene Africa, separated by the Arid Corridor (Somalia--Namibia, Jonathan Kingdon): this suggests that hominoids in the central forests-wetlands (Congo-Rift-Chad-Nile) evolved allopatrically in parallel in response to the same climatic & other circumstances with hominoids in the coastal forests-wetlands (E.African coast-Zambesi). When we compare fossil with extant hominoids, afarensis-aethiopicus-boisei resemble Gorilla more than Pan (& a fortiori Homo), and africanus-robustus resemble Pan & even H.erectus more than Gorilla (irrespective of size), and E & S.Afr.apiths evolved separately from gracile (Pliocene) to robust (Pleistocene): not unexpected in view of the climatic changes & the geographical separation by Kingdon's arid corridor.

3) Yes, the fossil record is very uneven, but not impossibly uneven. The traditional view is pure pre-darwinian anthropocentrism: every of the 1000s of hominid fossils is believed to be closer to 1 species (us, of course), virtually none to the 5 Afr.ape spp. Of course, fossil hunters (unconsciously I guess) prefer to find human rather than chimp ancestors (e.g. for funds & public attention), and when they see a shred of "bipedalism" (often confused with "vertical spine") in their fossil, they place it in the human branch, instead of realising that in certain instances (e.g. vertical spine) humans could have remained more primtive than the Afr.apes.

We can take hylobatids as outgroup for great hominoids, but not Pan as outgroup for Homo-apiths, which is traditonally done. Even without outgroups other than OWMs & hylobatids, afarensis-boisei resemble Gorilla more than Pan, and africanus-robustus resemble Pan more than Gorilla (although e.g. boisei resembles robustus). This can't be explained except by parallel evolution.

4)5) I meant: an early-hominoid vertical spine can easily evolve into the human & the gibbon spine, but also into the orang spine as well as into the KWer spine.

6) FYI, my 1994 & other Hum.Evol.papers can be found in the attachment. It's now being generally accepted (following my 1994 paper) that KWing in Pan & Gorilla evolved in parallel.

Marc,

1)     “If we take out certain genes, we can prove everything”.

Problem with ‘genes’ and phylogenetic reconstruction is that nothing is self evident and there are a lot of assumptions built in that are phylogenetically problematic. These cladistic problems are generally ignored. The law of large numbers is just phenetics of overall similarity that has no necessary phylogenetic meaning. Molecular analyses that involve balancing imagined gaps and substitutions for genes of unequal length have to create homologies that do not exist in nature. The balancing act is pure phenetics.

We are obviously in disagreement about DNA sequences. As for fitting “all other data” – no it does not. It does not ‘fit’ morphogenetic data.

“Yes, but e.g. the traditional interpretation (ape->apith->human) neglects possible parallelisms in Pan & Gorilla. “

As said before, parallelisms etc can only be identified with respect to their incongruence with the preferred phylogeny. If Pan and Gorilla are shown not to be monophyletic by other characteristics then their knuckle walking characters would have to be seen as independent derivations. No evidence yet that I have seen for that (pending reading your attachment)

2)     Yes, the fossil record is very uneven, but not impossibly uneven.

Whether impossibly or not, it is uneven and does not exist in anticipation of what one might consider too uneven. Fact is that fossils do not provide absolute divergence estimates. Such claims are fictions.

3)     We can take hylobatids as outgroup for great hominoids, but not Pan as outgroup for Homo-apiths, which is traditonally done.

Agreed – and so in Grehan & Schwartz (2009) Pan was part of the ingroup.

4) Even without outgroups other than OWMs & hylobatids, afarensis-boisei resemble Gorilla more than Pan, and africanus-robustus resemble Pan more than Gorilla (although e.g. boisei resembles robustus). This can't be explained except by parallel evolution.

Since such resemblances are not congruent with phylogeny they naturally have to be attributed to parallelism, convergence etc – by definition

5) I meant: an early-hominoid vertical spine can easily evolve into the human & the gibbon spine, but also into the orang spine as well as into the KWer spine.

Perhaps so, and if demonstrated phylogenetically it would prove to be most interesting.

6) FYI, my 1994 & other Hum.Evol.papers can be found in the attachment. It's now being generally accepted (following my 1994 paper) that KWing in Pan & Gorilla evolved in parallel.

Thanks – I’ll be interested to see that it is now generally accepted that knuckle walking are independent derivations, more later,

John

1) Let's agree to disagree. In any case, Gorilla & Pan KWing are different in ontogeny, degree, stand of hand etc.

2) This is not about divergence estimates, but about whether some apith fossils can be more related to Pan or Gorilla than to Homo.

3) OK.

4) Yes.

5) Have you read A.Filler 2007 "The upright ape" New Page NJ? Very very interesting. IMO Filler is wrong at many places, but he's right that the hominoid LCA c 20 Ma was already "upright" (vertical spine in Morotopithecus & other Miocene "apes"), e.g. for climbing vertically in the branches above the swamp (most Mio-Pliocene hominoids fossils lay in wetlands or forest swamps), for wading bipedally in such swamps (google "bonobo wading" or "gorilla bai"), or whatever (e.g. see attachment), but not for running, e.g. in running-feet (digiti- or unguli-grade), the central digital rays (3, 3-4) are longest & strongest (ostrich, horse, kangaroo, antilope...), whereas in swimming- and/or wading-feet (typically plantigrade), the outer digital rays are rel.long & strong (human, duck, grebe, sealion, penguin...).

1) Let's agree to disagree. In any case, Gorilla & Pan KWing are different in ontogeny, degree, stand of hand etc.

Features do not have to be identical to be homologues, but differences in ontogeny may be informative about that.

2) This is not about divergence estimates, but about whether some apith fossils can be more related to Pan or Gorilla than to Homo.

Trouble is that I have not yet seen any synapomorphies for australopiths and African apes

5) Have you read A.Filler 2007 "The upright ape" New Page NJ?

Yes did read it. Long time back now, but I recall that I could not decipher anything cladistically definitive regarding the vertebral interpretations. Whether or not Morotopithecus could walk bipedally or even that African apes and orangutans do, has nothing to do with the upright walking by hominids based on a different skeletal structure (just as the straight leg stance of orangutans is not structurally homologous to that of humans even though the two are unique in that ability among living large bodied hominoids).

Marc,

Reading your 1994 paper I find some potentially interesting points and possibly some that are phylogenetically informative. There are lots of phrases of “pan-like” and gorilla-like” but no structured cladistic argument with derived states specified and compared to all ingroup taxa as well as outgroups. Not surprising as you accept the molecular tree as the only framework so everything tends to be interpreted in that context.

Some specific observations as follows:

1) The composite skull reconstructed mostly from A.L.333 specimens “looked very much like a small female gorilla”. Johanson & Edey, 1981, p. 351.

The reconstruction by Kimbel et al 2004 for Al 444-2 did not.

2) nasals extended far above the frontomaxillar suture and well onto an uninflated glabella

uninflated glabella is not like African apes, but OK with Pongo etc.

3) Not obtained were: uniquely derived features of South African fossils with Gorilla; of A. boisei with Pan; and of any australopith with Homo (i.e. features that are absent from all African ape species, mature and immature).

But of course uniquely derived features have been identified by Schwartz (1984, 1988 etc,  as summarized by Grehan & Schwartz

4) Biomolecular results leave no doubt that Pan is genetically closer to Homo than to Gorilla and contrary to the prevailing opinion this is not contradicted by the anatomical evidence.

Not true according to Grehan & Schwartz. Anatomical evidence does contradict.

5) Even knuckle-walking of chimps and gorillas has been argued to have arisen independently (Begun, 1992),

I did read his paper, forget the details now, but was not impressed at the time. Could take a look again to see why.

6) And the LCA (the last common ancestor of Homo and Pan) had not yet acquired knuckle-walking since humans do not at any age show the slightest trace of knuckle-walking behaviour:

Ironically some tried to argue otherwise, that humans hands were derived from a knuckle-walking morphology.

7) I expect that when australopith fossil material is re-examined and compared in detail with every one of the large hominoids, in most cases it will resemble either Pan or Gorilla more closely than it resembles Homo and certainly Pongo.

As outlined in Grehan & Schwartz, not only do australopiths cladistically resemble Homo and Pongo more closely than African apes, they also include some specifically Pongo and allies characters such as  zygomatic anterior root forward facing, infraorbital plane flat, infraorbital plane superoinferiorly tall and infraorbital plane vertical. And for Homo, australopiths and Pongo there are also incisive foramen single, foramen lacerum present, and posterior palate thickened. Very non-African ape characteristics. Australopiths look like nothing if not some sort of strange Pongo if it were not for their locomotor apomorphies linking them with humans. And then there are those strange isolated teeth labeled in Africa as ‘Australopithecus” but have similarities to orangutans.

Cheers, John

Hi John, first your last answer:

1) Of course, Johanson-Kimbel's impression alone doesn't say much, but first impressions are often the best ("afarensis AL-333 cf small female gorilla": males often have more secundary sex.features; gorilla ancestors were possibly smaller). If we compare AL-333 to extant hominoids, IMO also they look most like fem.Gorilla>Pan>Pongo>Homo. My 1996 paper confirms that overall E.Afr.apith skulls look most like G>P>Po>H (but S.Afr.apiths like P>G>Po>H).

2) OK.

3) Well obtained: uniquely-derived features of S.Afr.apiths with Pan, and of E-Afr.apiths with Gorilla (see tables in my 1994 paper). At the time (1980s), I read +-all the extant literature on human evolution (Nature, Science, JHE, AJPA etc.), and I never encountered mentioning specific resemblances between apiths & Pongo (but probably these were not considered often).

4) I disagree. DNA overall is (((PH)G)Po)Hy)OWM etc. Morphology is the same, except for Homo (my 1996 paper): E.Afr.apith skulls look most like G>P>Po>H, and S.Afr.ones like P>G>Po>H. This suggests that all great apes evolved largely in parallel (G//P//Po), but Homo took a very different evol.way (= Pleist.Homo's littoral dispersal, diving-wading-beachcombing along African & Eurasian coasts, & then inland along rivers, see e.g. my last Hum.Evol.paper 2013 attached).

5) Begun 1992 is very interesting, but better read on KWing what I said in my 1996 paper.  :-)

6) That opinion (humans had KWing ancestors) was once widespread, but is now (rightly) generally believed to be wrong (at least 5 independent & convincing arguments). Parallel evolution is often underestimated by paleo-anthropologists, but in fact, the frequency of parallelism is one of the major arguments pro Darwin.

7) My 1996 paper used undisputed data from Wood-Chamberlain: the results are clear: when we morphologically compare fossil & extant hominoid skulls-dentitions (I didn't have enough postcranial data), all apiths look more like Afr.apes than like orangs. All known evidence (DNA, ontogeny) confirms this. Schwartz' work ("The red ape" etc.) is very interesting & inspiring, but unfortunately it's IMO an example of cherry-picking. Independent analyses (e.g. Wood-Chamberlain) leave no doubt, and confirm DNA data.

Hi John, you say "Whether or not Morotopithecus could walk bipedally or even that African apes and orangutans do, has nothing to do with the upright walking by hominids based on a different skeletal structure (just as the straight leg stance of orangutans is not structurally homologous to that of humans even though the two are unique in that ability among living large bodied hominoids)."

Many PAs confuse skeletal structures "for" walking bipedally with skeletal structures "used in" walking bipedally (i.e. seen in humans, but not in apes). Often, if PAs see in "their" fossil 1 locomotor feature that is present in humans but not apes, they consider their fossil as more closely related to us than to one of the apes. As I have repeatedly argued, this is pre-darwinian anthropocentrism.

An example: apiths had more humanlike feet (esp.long & adducted big toe), therefore are believed to be closer relatives of us than of apes, but e.g. (1) comparative evidence suggests flat feet are not for running, but for swimming and/or wading, (2) prof.Coon said prenatal chimps have humanlike feet, which after birth become more hand-like, (3) prof.Clarke (in SAJS) let a chimp walk over wet sand, and its footprints were like those of Laetoli. All this suggests that our & ape ancestors did not evolve flat feet for running, but that early hominoids might have had more humanlike feet which were used for swimming-wading (see my 2013 Hum.Evol.paper attached to my previous answer).

Hi Marc,

1) If the skulls of Afarensis do not contain any unique features with African apes then the ‘resemblance’ is phylogenetically uninformative. I will review and discuss your features and come back to you on that.

2) OK.

3) Well obtained: uniquely-derived features of S.Afr.apiths with Pan, and of E-Afr.apiths with Gorilla (see tables in my 1994 paper).

I’ll look at that again.

“At the time (1980s), I read +-all the extant literature on human evolution (Nature, Science, JHE, AJPA etc.), and I never encountered mentioning specific resemblances between apiths & Pongo (but probably these were not considered often).”

somehow you missed Schwartz’s (1984) paper in Nature?

4) I disagree. DNA overall is (((PH)G)Po)Hy)OWM etc. Morphology is the same, except for Homo (my 1996 paper): E.Afr.apith skulls look most like G>P>Po>H, and S.Afr.ones like P>G>Po>H. This suggests that all great apes evolved largely in parallel (G//P//Po), but Homo took a very different evol.way (= Pleist.Homo's littoral dispersal, diving-wading-beachcombing along African & Eurasian coasts, & then inland along rivers, see e.g. my last Hum.Evol.paper 2013 attached).

You may disagree, but the fact remains is that the preponderance of evidence in the form of uniquely shared characters is in the direction of a human-orangutan sistergroup.

5) Begun 1992 is very interesting, but better read on KWing what I said in my 1996 paper.  :-)

Haven’t looked at your 1996 paper yet

7) My 1996 paper used undisputed data from Wood-Chamberlain: the results are clear: when we morphologically compare fossil & extant hominoid skulls-dentitions (I didn't have enough postcranial data), all apiths look more like Afr.apes than like orangs. All known evidence (DNA, ontogeny) confirms this. Schwartz' work ("The red ape" etc.) is very interesting & inspiring, but unfortunately it's IMO an example of cherry-picking. Independent analyses (e.g. Wood-Chamberlain) leave no doubt, and confirm DNA data.

There is no cherry picking in Schwartz’s work. He used the cladistic approach of looking only for uniquely shared characters with lesser apes and monkeys as the outgroup, This is standard cladistics. I get the impression that you do not understand cladistics or I do not see how you would make the cherry picking assertion.

John

Marc,

The human foot may have functioned differently in its origin than now, but what manner of function is a matter of speculation for which I have no empirical basis to comment. But my observation stands, that upright walking in apes or monkeys involves a different skeletal structure to that of hominids so I don't see how it is informative about the evolution of the particular skeletal structure involved with hominid locomotion.

Hi John, flat feet are not seen in cursorial animals, nor in climbing animals, but in swimming and/or wading animals. This suggests we evolved from arborealism (+-all primates) to frequent swimming and/or wading (flat feet) to what we're doing today (bipedal walking). Why is this simple comparative evidence so often neglected? because it doesn't fit our preconceptions on human evolution? Traditionally it's believed we evolved from quadrupedal apes to bipedal "hominids" (definiton?) after we left the forests for the plains.

Your "observation" about upright walking "hominids" is also anthropocentric IMO. Upright is not necessarily walking, nor bipedal. Most bipedal animals have +-horizontal spines, e.g. ostriches & hopping kangaroos. Only humans & penguins on land & gibbons walking over branches AFAIK walk with upright spines. I think we should stop identifying bipedal = upright = walking: these are 3 different things.

Marc,

Just because flat feet are seen in swimming and/or wading animals does not automatically mean that this was the case for the origin of 'flat' (arched) feet in humans. However, I am also not suggesting that human (homo) did not swim, diver, or wade as I have no empirical evidence either way  If the swimming or wading required flat feet to be possible then the flat feet had to come first.

Good point about distinction of terms. I am OK with that. Point still remains, that walking upright by itself does not have any necessary phylogenetic implications for the origin of upright walking in another group with a different skeletal structure - i.e. bipedal walking in gorillas, chimps, or orangs has nothing to do with the evolution of bipedalism in humans.

I forget what you say about australopithicens - did they have upright spines? Am I correct to presume that the spine of orangs is not upright even when they walk vertically in trees?

Cheers, John

Hi John, I didn't miss J.Schwartz 1984 "The evolutionary relationships of man and orang-utans" Nature 308:501-5: "Man shares uniquely few morphological features with either the chimpanzee or the gorilla ... many features suggest affinities between man & the orang-utan, to whom the fossil Sivapithecus appears to be closely related. If these are unique features, inclusion of Sivapithecus, man & the orang-utan in a single clade, distinct from that containing the African apes, is justified ..." In the 1970s it was generally believed (e.g. P.Andrews IIRC) that "Ramapithecus" (probably female Sivapith) was closely related to humans because it had thick enamel & rel.small canines, but when it got accepted that Rama-Sivapith might have been related to Pongo, Schwartz believed that Pongo too was closer related to us (hence his (IMO) cherry-picking). Of course, there are some features which orangs & humans have in common vs other great apes, but these seem to be primitive for (great)hominoids, e.g. most Miocene hominoids had thick enamel, IOW, thin enamel in hominids & pongids is derived, e.g. esp.gorillas evolved thinner enamel (cf evolution from omni- to more herbivory): in some instances, Homo is more primitive than  the (great) apes, who allopatrically frequently evolved in parallel, e.g. they got longer arms (gibbon>orang>chimp>gorilla-human, or longer ilia (gr.apes>gibbon>human-monkeys) etc.

Hi John, I do think that bipedal walking in Pan & Gorilla might have a common origin with Homo's bipedal walking, google "bonobo wading" & "gorilla bai" (AFAICS, Pongo's ancestors were less bipedal or wading).

I guess but am not sure that apiths had upright spines (IMO more for vertical climbing-hanging (gibbon-like) & for wading or perhaps floating vertically in forest swamps or wetlands (as bonobos & lowland gorillas sometimes do), not for walking upright on terra firma). In fact, H.sapiens IMO was even more upright than archaic Homo (erectus, neand.etc.), e.g. AFAIK only H.sapiens has very long & caudally-directed spinous processes mid-thoracally (I'd think for stabilising a wading-walking spine vertically), e.g. compare with Nariokotome's thoracal vertebrae.

Marc, with respect to your assertion that Schwartz was cherry picking because Schwartz believed that Pongo was also closer to us when it became accepted that Rama-Sivapith might have been related to Pongo does not make sense. I do not see how this can be construed as cherry picking of the evidence.

Your assertion that there are some features which orangs & humans have in common vs other great apes, but these seem to be primitive for (great)hominoids is not supported by the analysis by Schwartz & Grehan (2009) other than the possibility, as you point out, that thick enamel may be more widely distributed than the immediate clade described in that paper. However it remains to be seen whether it can be substantiated that the thinner enamel of African apes is derived or primitive with respect to outgroup comparison (if thick enamel were primitive for large bodied hominoids then the thin enamel condition of African apes would certainly support their grouping as a clade, but not with humans).

The remaining (many) characters between humans and orangutans as described in Grehan & Schwartz (2009) are well corroborated as being derived, not primitive, for large bodied hominioids. The fact remains (and has not been refuted in any subsequent publication I am aware of) that humans and orangutans share far more uniquely shared characters than any other grouping of large bodied hominoids. Naturally the evidence is most extensive for living taxa since soft tissue and physiological characters are available that are not for fossil taxa.

One set of characters that intrigues me, that has not yet been reported for australopths, concern several unique cusp configurations shared by humans and orangutans with respect to deciduous teeth. The dikkika juvenile appears to have an intact set of teeth, but I have not kept track of whether the researchers were able to scan the dental surface of the teeth that hopefully have not been worn to the extent of obscuring such surfaces.

Hi John, I sent you my 1996 Hum.Evol.paper (+ tables) yesterday: it shows that (at least cranio-dentally) Pan & Gorilla resemble Homo more than Pongo does. Of course, orangs kept some primitive features, just as bonobos kept several primitive features, and humans kept some: rel.shorter upper limbs (with gorillas), thick enamel (with orangs), low pelvis (with gibbons), smaller canines (female bonobos +-) etc.

Yes, (pre)molar cusp configurations (you probably mean odontoglyphs?) are very interesting. If we'll be able to extract enough DNA from apith bones, I predict E.Afr.apiths will be shown to be closest to Gorilla, S.Afr.apiths to Pan, and Siva-Lufeng-Gigantopith to Pongo, but if we can't rely on DNA, odontoglyphs might be an alternative?

Marc,

I'm not familiar with the term odontoglyph, but then I don't have expertise in dental taxonomy. My understanding is that humans and orangutans uniquely share the following dental features: the shortest lower posterior deciduous molar trigonid, an oval lingual side of upper molar, lower anterior deciduous molar protoconid anteriorly placed, lower anterior deciduous molar mesially angled, and lower anterior deciduous molar talonid basin closed (outgroup comprising gibbons and OW and NW monkeys). Of course if one has already decided that only the DNA sequence data holds the ultimate truth then these uniquely shared features are phylogenetically meaningless.

John

Hi John, sorry I misunderstood (I must have been in a hurry). Yes, of course, orangs can have retained some dental features together with us, but that doesn't mean we're not closer related to chimps than to orangs (I even read that of some fossil premolars it's impossible to say whether they belong to Homo or to Pan). Overal, in morphology as well as DNA, humans are much closer to chimps than to orangs (in my 1996 Hum.Evol.paper, dental features were included: 16 dento-gnathic characters of 37 cranio-dental characters).

If your 1996 paper presents a cladistic analysis with a broad outgroup it will be very interesting to see how it compares with Grehan & Schwartz. I regret that at the time I missed your paper as it would have otherwise been addressed. I look forward to seeing it and commenting in due course.

Hi John, my 1994 paper is no cladistic analysis (cladistic analyses are not foolproof to parallel, convergent & reverse evolutions, which are frequent in hominoid evolution, see attachment), but is simply a numerical (very detailed) comparsison of cranio-dental characters between australopiths, Homo & extant hominoids. The problem with cladistic analyses is that apelike characters are believed to be primitive, whereas a more thorough analysis suggests that many apelike characters are derived. A few examples: long ilia; long upper extremities; humanlike feet; KWing; thick enamel; etc

1) Low ilia as in Homo & australopiths are primitive, not derived: monkeys also have "low" pelvises; the great apes (& gibbons to a lesser extent) evolved elongated ilia in parallel (Pan // Gorilla // Pongo).

2) The early hominoids didn't have elongated upper extremities, e.g. Lucy's humerus (~24 cm) is estimated to be shorter than in bonobos (~29 cm) & even human pygmies (~26 cm), much shorter than the human mean (~33 cm). Instead, the different hominoid lines evolved longer arms in parallel, e.g. relative humerus lengths: hylobatids>Pongo>>Pan>Homo=Gorilla.

3) It's believed that all fossils with humanlike feet (e.g. Laetoli, naledi, OH-8) are close relatives of us, but S.Coon already noticed that prenatal chmps have more humanlike feet which become more hand-like after birth. And when Ron Clarke (1998 S.Afr.J.Sci.94:460) let a male circus chimp walk over wet sand, he held his big toes next to his foot, and his footprints were indistinguishable from those of Laetoli (but an anxious female bonobo which held her big toes wide apart produced very different footprints). IOW, it's well possible (& indeed likely IMO on other grounds) that that the hominid LCA ~8 to 5 Ma had more humanlike feet (we argued for swimming-wading in swamp forests, see attachment)

4) Some paleo-anthropologists still believe that we had knuckle-walking ancestors & lost all rudiments of KWing in a miraculously complete way, but a lot of independent indications (e.g. my 1994 paper) suggest that that Pan & Gorilla evolved (different sorts of) KWing in parallel (as can be expected in darwinian evolution in resembling & related, but allopatric spp in comparable milieus).

5) For thick enamel (early-hominoid durophagous adaptation to coastal forest-dwelling?) & other hominid & pongid features, please see the attachment.

Yes, we can perhaps better await your lecture of my 1996 paper.  Best wishes  --marc

For an update of hominid (Pan, Homo & Gorilla) evolution, please google "two incredible logical mistakes 2019 verhaegen" or see attachment (+ refs).

Thanks Marc.

Regards.