by Marcel F. Williams Tuscany is renowned for its beautiful cities of Florence and Siena, and is historically famous as the birthplace of the Italian Renaissance. But amongst the paleontological community, Tuscany is also known as the birthplace of an extinct Late Miocene ape-- that some paleontologist controversially believe may have been the earliest primate to walk predominantly on just two legs-- and possibly the earliest bipedal ancestor of humankind.
Oreopithecus bambolii first emerged on an ancient island bioprovince known as Tuscany-Sardinia (Tusco-Sardinia) sometime after 9 million years ago where it existed as the sole primate on the island until approximately 7 million years ago. Tuscany-Sardinia was isolated from the rest of the Italian peninsula and from the rest of continental Europe by the marine waters of the Late Miocene Mediterranean.
Skull of Oreopithecus bambolii
The mostly likely ancestor of Oreopithecus was a small African ape know as Mabokopithecus which appears in the fossil record in Africa about 15 million years ago. Oreopithecus and Mabokopithecus were both folivores (planet eaters) that shared a unique dental attribute in their upper molars: a well-defined hypocone-metaconule crest on the upper molars-- a unique characteristic that has never been found in any other monkey or ape living or extinct.
It is interesting that the African oreopithecine ancestor, Mabokopithecus, existed in an ecological niche largely restricted to riparian woodland areas. This is a niche similar to that of the extant folivorous primate Colobus guereza (the Colobus monkey). Although the Colobus monkey is highly arboreal, it is known to come down from the trees in order to feed on aquatic plants in nearby swamps.
Oreopithecus bambolii has long been at the center of controversy principally because of its remarkable cranio-dental and post cranial similarity to African hominins (human ancestors) which was first fully noted by Johannes Hurzeler as far back as the 1950s. Further adding to the controversy is the argument that Oreopithecus may have also been the earliest obligatory bipedal primate and the first ape to walk exclusively on just two legs.
Oreopithecine remains display a significant number of post cranial characteristics that could be associated with either arboreal or bipedal locomotion. However, there is one post cranial characteristic that is clearly related to bipedality and that's lumbar lordosis, a curvature of the spine that is unique to the hominins (humans and their bipedal ancestors). However, a recent paper by Russo and Shapiro has question the existence of lumbar lordosis in Oreopithecus, arguing that the supposed features may be simply be an artifact of the skeleton's distortion caused by its compression during fossilization. Russo and Shapiro also argue that the arboreal three toed sloth (Bradypus) would be a better convergent model for the skeletal anatomy and locomotive behavior of the extinct oreopithecines rather than the bipedal hominins.
Three-toed sloth (Bradypus)
While slow climbing suspensory behavior has long been argued as an explanation for some of the anatomical attributes of Oreopithecus, such locomotive behavior appears to be contradicted by the exceptionally robust metatarsals of the oreopithecine foot along with the proportions of the foot's entocuneiform which was proximally-distally short and dorso-ventrally high. The length-height index of the oreopithecine entocuneiform related to the mass placed on the hind limbs and is most similar to that of the gorilla. The gorilla's low entocuneiform length-height index is related to the large amount of mass placed on the hind limbs due to its large body size. Oreopithecus, however, was a very small ape, with males typically weighing about 32 kilograms and females approximately half that size. The average Gorilla male weighs 175 kg with average female gorillas weighing about 85 kilograms. So in order to explain, the gorilla-like entocuneiform index, in Oreopithecus, the swamp ape must have been carrying its entire body weight on just its hind limbs. Kohler and Moya- Sola also noted that the power armload arm ratio strongly suggest that Oreopithecus carried its entire body weight on its hind limbs.
There's also strong evidence in the oreopithecine feet of a significantly reduced arboreal ability. The Oreopithecus foot had robust metatarsals plus a non helical ankle joint, characteristics typically associated with terrestrial catarrhines. Kohler and Moya-Sola have also noted that the mobility and grasping ability of the oreopithecine foot had been appreciably reduced relative to arboreal primates and even more so than in terrestrial baboons.
It would also be difficult to understand why Oreopithecus would expend the energy and the risk of being a highly arboreal tree living primate on an island where there were no terrestrial predators. But even on the ground, why would Oreopithecus abandon terrestrial quadrupedalism for bipedalism? The answer may come from its food source.
Chimpanzee wading bipedally with a stick
Orangutan wading bipedally
Gorilla wading bipedally
The coastal wetlands that Oreopithecus preferred were rich in aquatic plants. Harrison and Rook have suggested that Oreopithecus may have specialized in feeding on aquatic plants which were abundant in the wetland coastal areas of the island such as: sedges, water lilies, reeds, cattail, pond- weeds, horestails, and stoneworts which were abundantly represented in the fossil pollen spectrum.
Extant primates that feed on aquatic plants, usually wade bipedally in the shallow waters to access the food resource. Bipedal wading in primates has been observed in: baboons, macaques, the gorilla, orangutan, and in the chimpanzee. Such aquatic bipedalism in order to access aquatic plants comprises as much as 27% of the feeding behavior of the Western Gorilla.
Additionally, freshwater mollusk and turtles were also abundant in the wetland areas of Tuscany-Sardinia island-- along with predatory crocodiles. Turtle and crocodile eggs may have served as a good source of protein for Oreopithecus.
Marine biologist, Alister Hardy, hypothesized that the power precision grip in humans originally evolved as an adaptation for picking up benthic invertebrates. Wading bipedally in shallow water for food resources such as shellfish and aquatic plants is, of course, common behavior in many hunter-gatherer human populations. But such behavior has also been observed in non-human primates such as baboons, macaques, guenons, and capuchin monkeys.
Curiously, Oreopithecus also had a hominin-like power precision grip. Such manual dexterity could have evolved in Oreopithecus as a feeding adaptation for picking up the shelled invertebrates that existed in the wetland areas of Tuscany-Sardinia while it was also feeding on aquatic plants.
As the sole primate on the ancient island of Tuscany-Sardinia for nearly two million years, it seems unlikely that Oreopithecus would have avoided the exploitation of freshwater aquatic food resources that were so rich in carbohydrates and proteins.
The earliest African hominin, Sahelanthropus, appeared in the fossil record in North Africa sometime between 6.8 and 7.2 million years ago. While not much is known about the postcranial remains of Sahelanthropus, its craniodental morphology is remarkably similar to that of Oreopithecus bambolii.
Marcel F. Williams
References:
Cranio-dental evidence of a hominin-like hyper-masticatory apparatus in Oreopithecus bambolii . Was the swamp ape a human ancestor?
, Bioscience Hypotheses, 1 (3), p.127-137, Jan 2008
doi:10.1016/j.bihy.2008.04.001
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Köhler M, Moyà-Solà S.
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Straus WL. The classification of Oreopithecus. In: Washburn SL, editor. Classification and human evolution. Chicago: Aldine; 1963. p. 146e74.
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An anthropoid enigma: historical introduction to the study of Oreopithecus bambolii. J Hum Evol 1987; Delson E 15:523e31.
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Harrison T. A reassessment of the phylogenetic relationships of Oreopithecus bambolii Gervais. J Hum Evol 1987; 15:541e83.
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Harrison T, Rook L. Enigmatic anthropoid or misunderstood ape? The phylogenetic status of Oreopithecus bambolii reconsidered. In: Begun DR, Ward CV, Rose MD, editors. Function, phylogeny, and fossils: Miocene hominoid origins and adapata- tions. New York: Plenum Press; 1997. p. 327e62.
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Evolutionary History of Primates
Szalay and Delson 1979
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Our Earliest African Ancestor
New Papyrus
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David Attenborough on the Aquatic Ape Hypothesis
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Desmond Morris on the Aquatic Ape Hypothesis
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Was Man More Aquatic in the Past? Fifty Years After Alister Hardy - Waterside Hypotheses of Human Evolution
Mario Vaneechoutte, Algis Kuliukas, Marc Verhaegen
Bentham Science Publishers,
Fascinating! So our aquatic ancestor may have evolved in Europe after all? And then returned to Africa? Would this also explain the missing baboon marker?
ReplyDeleteDid human ape ancestors go through a semiaquatic phase or phases in their evolution sometime during the last 10 million years?
ReplyDeleteCast your vote at:
http://www.dailykos.com/story/2013/11/14/1255709/-Was-the-Swamp-Ape-Bipedal#comments
The presence of (high protein) Hydrocharis today in Tuscan marshes plausibly indicates the main aquatic food of Oreo., as it is for the Congo lowland gorilla, who rake it with their fingers while partly submerged (~97% Hydrocharis, ~3% sedges).
ReplyDelete=
History and traditions
As weIl as the richness of its landscape and nature, the Marshes have maintained the fascination of the historical events related to the great Medici and Lorena families. Here still remain important testimonies of the work of man, which over the centuries have shaped and changed the very structure of the wetland: the canals and port systems, signs of ancient and important waterways; the Medicean Bridge at Cappiano, the centre for controlling the water regime and fishing activities, as well as an important stop along the Via Francigena; the Capannone Farm complex, which was one of the main berthings of the Valdinievole; the buildings of industrial archaeology like the tobacco dryhouses.
The commemorative plaques so often found on the cabins or along the banks tell a more recent story: the tragedy of the barbaric massacre committed by the Germans on 23rd August, 1944.
A few skilful craftsmen still carry on residual activities related to the marsh grasses in the marshes: gathering and weaving "sarello" (tufted sedge) and "sala" reed-mace (False bulrush) (to wicker seats and cover wine flasks), "Gaggia" (False Indigo) and other typical wetland plants.
Flora
Situated on the boundary between the Mediterranean and Continental climates of the Peninsular, the Marshes simultaneously shelter plants adapted to different climates; for example in the Ramone Marshlet, on the edges of the Chiusi Woods, still survive the Frogbit (Hydrocharis morsus-ranae) and the Royal Fern (Osmunda regalis), both plants from a warm wet climate, alongside certain mosses (Sphagnum sp.), more adapted to cold climates of the north and which descended as far as the Marshes during the last Ice Age. Only in the Marshlet, and in a few other areas of the Marsh basin, can still be found fair extensions of the Tufted Sedge (Carex elata), locally called "sarello"; this plant of northern origin develops in bushy formations of more than one individual.
Where the immense reed thickets leave space to the free waters, are the "laminas" formed of plants with floating leaves (e.g. the large White and Yellow Water-lilies) which offer one of the last refuges for several highly specialised species: the Southern Bladderwort (Utricularia australis), a floating carnivorous plant, the Fringed Water-lily (Nymphoides peltata), with its beautiful yellow flowers, and the strange, tiny floating fern Salvinia natans, now extremely rare in Tuscany.
http://www.tuscany-charming.it/en/route/fucecchio-marshes.asp
Very interesting comments DDeden. And thanks for the link!
ReplyDeleteMarcel
Most Mio-Pliocene hominoids were typically orthograde (ie, with vertical spine), wading/climbing/floating in swamp & litoral forests (google "aquarboreal", word coined by Marcel), surface-feeding on AHV & HSI: aquatic herbaceous vegetation (eg, Hydrocharis, Cyperus) & hard-shelled invertebrates, as well as waterside & presumably arboreal foods. Oreopithecus is just one of these, with insular adaptations, no direct ancestor of chimps or humans: AAT (better terms are "littoral theory" or "coastal dispersal model", google "econiche Homo") is about Plio-Pleistocene erectus-like archaic Homo populations trekking along coasts as far as Flores island, the Cape & Pakefield in England (where they dived for shellfish etc.), and from the coasts inland along rivers, probably at first seasonally (eg, follwing salmon etc.), later also permanently.
ReplyDeleteFossil data on human relatives ("emergence of Homo") should not be confused with biochemical data on human ancestors ("baboon retrovirus"): Pan & Homo split c 5 Ma, possibly in coastal forests (Zambesi mouth?), presumably our Pliocene ancestors first followed the Indian Ocean to Sunda (explaining the retroviral data).
Human Evolution publishes in 2 special editions the proceedings of the symposium with David Attenborough on human waterside evolution "Human Evolution: Past, Present & Future" in London 8-10 May 2013:
SPECIAL EDITION PART 1 (end 2013)
- Peter Rhys-Evans: Introduction
- Stephen Oppenheimer: Human's Association with Water Bodies: the 'Exaggerated Diving Reflex' and its Relationship with the Evolutionary Allometry of Human Pelvic and Brain Sizes
- JH Langdon: Human Ecological Breadth: Why Neither Savanna nor Aquatic Hypotheses can Hold Water
- Stephen Munro: Endurance Running versus Underwater Foraging: an Anatomical and Palaeoecological Perspective
- Algis Kuliukas: Wading Hypotheses of the Origin of Human Bipedalism
- Marc Verhaegen: The Aquatic Ape Evolves: Common Misconceptions and Unproven Assumptions about the So-Called Aquatic Ape Hypothesis
- CL Broadhurst & Michael Crawford: The Epigenetic Emergence of Culture at the Coastline: Interaction of Genes, Nutrition, Environment and Demography
SPECIAL EDITION PART 2 (begin 2014) with 12 contributions.
--marc verhaegen, google "Greg Laden blog Verhaegen"
Bravo a tutti! :D
ReplyDeleteThanks for very important informations..
Tuscany is always a place of main source of artistic and humanistic nutrition.
Oreopithecus bambolii was a non-arboreal quadruped:
ReplyDeletehttps://m.phys.org/news/2019-12-enigmatic-hominoid-upright-tree-climber.html
Oreopithecus walked bipedally because aquatic plants and aquatic invertebrates were its primary food resource. So they had to wade bipedally in order to access aquatic food. Gorillas also walk bipedally in shallow water when feeding on aquatic plants.
ReplyDeleteOreopithecine preference for aquatic plants is evident in the isotopic evidence and by the fact that Oreopithecus had significant subcutaneous fat, a common adaptation in mammals and birds that feed on aquatic organisms.
The fact that Oreopithecus aquatic bipedalism was not as efficient terrestrial bipedalism is really not a surprise, IMO. Aquatic bipedalism mostly consist of standing and slow wading while terrestrial bipedalism frequently involves rapid walking and running. Its power arm load arm ratio--strongly-- suggest bipedalism.
Its rather ironic that the authors suggest that Oreopithecus is postcranially most similar to Symphalangus (the siamang) an arboreal ape that walks bipedally, not quadrupedally, when on the ground!
Marcel Williams
ReplyDeleteThanks, Marcel.
Oreop was one the many Miocene "vertical" hominoids in swamp/mangrove/flooded forests. Probably most or all of these early apes were "vertical": as opposed to monkeys (with dorsally-situated spine), all hominoids have centrally-placed spines, which suggest a habitually upright posture: very likely they all frequently climbed vertically (arms overhead) in the branches above the forest swamp , most of them might also have waded bipedally like lowland gorillas & bonobo still do (google "gorilla bai" & "bonobo wading"), and presumably many of them might also have floated vertically with inflated airsacs (laryngeal, axillar...), see e.g. our paper Verhaegen, Munro & Vaneechoutte 2011 "Early Hominoids: orthograde aquarboreals in flooded forests?" pp.67-81 in M.Vaneechoutte cs eds "Was Man More Aquatic in the Past? Fifty Years after Alister Hardy: Waterside Hypotheses of Human Evolution" eBook Bentham Sci.Publ.
AFAIK, the term "aquarboreal" (aqua=water, arbor=tree) was coined by Marcel Williams.
For refs + an update of ape & human evolution, please google "two incredible logical mistakes 2019 verhaegen".
I just read the whole article
ReplyDeleteAshley Hammond cs 2019 Proc.natl.Acad.Sci.
"Insights into the lower torso in late Miocene hominoid Oreopithecus bambolii"
https://doi.org/10.1073/pnas.1911896116
Contrary to what I thought before, Oreop probably didn't wade frequently:
AFAICS: was Oreop a below-branch orthograde aquarboreal?
-(semi)arboreal, frequently/habitually below-branch,
-probably predom.vertical, but rarely brachiating or wading,
-often/sometimes floating vertically?? e.g. with large airsacs?
for feeding on aq.herbs? and/or shell/crayfish?
and/or for crossing open waters to reach swamp forests on other islands?
See my Hum.Evol.papers, e.g.
2000 with P-F.Puech
"Hominid Lifestyle reconsidered:
Paleo-environmental and Comparative Data"
Hum.Evol.15:151-162,
2013
"The Aquatic Ape evolves:
Common Misconceptions and Unproven Assumptions about the so-called Aquatic Ape Hypothesis"
Hum.Evol.28:237-266,
or google
"two incredible logical mistakes 2019 verhaegen".
As noted in my article, Western Gorillas spend about 27% of their feeding behavior wading bipedally for aquatic plants. So there's no doubt in my mind that Oreopithecus probably spent substantially more time feeding in swamps than the Western Gorilla.
ReplyDeleteThe isotopic evidence indicates that Oreopithecus probably fed on aquatic plants. And the fact that Oreopithecus is the only primate other than modern humans that is currently known to have had a significant subcutaneous fat also suggest that Oreopithecus was adapted to feeding on aquatic organisms.
In their insular environment, oreopithecines probably fed on anything they could get their hands on (both on land and in the water), their primary diet appears to have been folivorous. This is clearly indicated by the extremely high Shearing Quotents for their second molars which far exceeds that of any other hominoid both living and extinct. Oreopithecus chewing behavior appears to have been so intensive and so extreme that even oreopithecine females had skulls with sagittal crest.
And, again, the isotopic evidence seems to suggest that these plants were aquatic.
Marcel
Marcel, is Oreop known to have a lot of SC fat? How can we know for sure? In warm environments, (semi)aquatic mammals don't need a lot of SC fat, e.g. hippos are said to be "lean". I'm not sure about SC fat in seacows.
ReplyDeleteYes, it's not unlikely that Oreop fed a lot on AHV (aquatic herbaceous vegetation, google "gorilla bai" & "bonobo wading") & perhaps also on HSI (hard-shelled invertebrates, see papers of Alan Shabel on australopiths eating HSI in wetlands), but the recent paper by Hammond cs (2019 PNAS) suggests they didn't do this wading (legs too small & short?).
If they (partly?) fed on AHV or HSI (likely IMO), they might have done that floating, like lowland gorillas still seem to do, with inflated airsacs in the neck, see discussion in my paper with S.Munro, P-F.Puech & M.Vaneechoutte "Early Hominoids: orthograde aquarboreals in flooded forests?" p.67-81 in Vaneechoutte cs eds 2011 "Was Man More Aquatic in the Past? Fifty Years after Alister Hardy" eBook Bentham Sci.Publ.
Gibbon & siamang & great ape airsacs were originally probably for floating (hydrostatics). The airsacs "coloured" the territorial calls, and gradually got used in duetting songs. In the Pleistocene (cooler & drier), the time spent in forest swamps became less, and the airsacs were not used any more for floating, except in lowland gorillas sometimes.
Oreopithecine subcutaneous fat is discussed in the article:
ReplyDeleteEarliest evidence of caries lesion in hominids reveal sugar-rich diet for a Middle Miocene dryopithecine from Europe
https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0203307
A photo of the adipose tissue area is also shown.
Marcel
I must have missed this paper.
ReplyDeleteThanks a lot, Marcel, very interesting. --marc
___
Earliest evidence of caries lesion in hominids reveal sugar-rich diet for a Middle Miocene dryopithecine from Europe
Jochen Fuss, Gregor Uhlig & Madelaine Böhme 2018 PLoS
https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0203307
The formation of dental caries is mainly caused by dietary habits,
it may contain information for dietary reconstructions of fossil hominids.
We investigate the caries lesion in the 12.5-Ma Dryop.carinthiacus Mottl (St.Stefan).
Potential food sources are identified on associated palynological data,
this allows conclusions about
- food quality,
- sugar availability,
- Mid-Miocene hominid metabolism.
μCT & SEM provide a detailed analysis & characterization of the individuals' caries type.
Its lesion is compared with 311 wild chimps,
it indicates morphological & etiological differences in caries fm between both spp.
The affected molar of D.carinthiacus reveals features known from severe dental caries in humans:
- cavitation with steep walls & smooth surface,
- reparative dentine at the roof of the pulp chamber,
- sclerotic dentine below the cavitation,
- association with dental calculus,
- unilateral usage of the healthy right tooth row.
Its advanced primary caries (initiating on the intact enamel surface) indicates a frequent intake of highly cariogenic sugar-rich fruits (likely > chimp frugivory).
This corresponds with the associated palynological record, which infers a habitat with 9–10 months/yr supply of high quality foods (carbohydrates+ fibers-).
Our conclusions
- challenge the model of a step-wise increase in dietary quality during hominid evolution,
- support the uricase hypothesis, which discusses the hominid autapomorphy of a fructose-based fat accumulation for periods of starvation.
This model receives further validation by the identification of soft-tissue preservation, interpreted as fossilized white adipose cells, in the articulated hominid skeleton of Oreop.bamboli.
Marcel, excellent article. I would like you to consider this: Mario P. has said that bamboo (or perhaps a similar tall river reed?) grows in Croatia. Date palms grow widely around the Mediterranean afaics. Climbing these plants would select for long arms and short legs (upright vertical climbing) but not for lateral brachiation (unlike modern hylobatids). Eating reed seeds/rhyzomes and palm fruits (high carbs & fats?) in addition to AHV like floating hydrocharis (high-protein) would be similar to gorillas, but with no knuckle-walking nor brachiation selection. Also, papyrus grows in southern Italy in freshwater swamps, (though it may have been brought from Egypt more recently?).
ReplyDeleteUnlike brachiating, where long hook-like hands are advantageous and thumbs not, coconut or date palm or giant reed climbing doesn't select against pad-to-pad precision grip. Wouldn't the odd big toes assist in this type of climbing?
ReplyDeletehttps://youtu.be/QARuWJjZkmw
DDeden:
ReplyDeleteUnlike brachiating, where long hook-like hands are advantageous and thumbs not, coconut or date palm or giant reed climbing doesn't select against pad-to-pad precision grip.
Gibbons (brachiating) don’t have small thumbs: comparable to macaques.
Orangs (slow-suspensory rather than brachiating) have smaller thumbs.
Spider-monkeys (grasping-tail, brachiation) have very small thumbs, smaller than in orangs.
Humans have rel.big=broad thumbs: not unexpected (e.g. swimming):
for manipulating you could better have narrower thumbs I’d think (cf pincette),
but chimps pick up a needle from the ground as easily as we do.
Pan & Gorilla (no good brachiators) evolved long hook-like hands in //, for climbing vertically? for knuckle-walking?
DD:
Wouldn't the odd big toes assist in this type of climbing?
https://youtu.be/QARuWJjZkmw
Our human "odd"(?) big toe, you mean?
IMO clearly adapted to swimming/wading, not to climbing.
https://www.gondwanatalks.com/l/the-waterside-hypothesis-wading-led-to-upright-walking-in-early-humans/
ReplyDeletePlate tectonics & hominoid splittings (2022 hypothesis, see my book p.299-300 "De evolutie van de mens" Acad.Uitg. Eburon 2022 Utrecht NL):
ReplyDelete- c 30-25 Ma India approaching Eurasia: island archipels + coastal forests++
Catarrhini reaching these islands became Hominoidea in coastal forests: bipedally wading, climbing arms overhead, predom.vertical spine: broad sternum + thorax -> scapulas dorsal: lateral & overhead arm movements, 7->5 lumbar vertebras dorsally->centrally in trunk, broad pelvis + lateral leg movements, tail loss etc., google "aquarboreal",
- c 25-20 Ma India underneath Eurasia split Hominoidea into lesser apes (hylobatids) East vs great apes West along Tethys Ocean coasts,
- c 15 Ma Mesopotamian Seaway closure split great apes into pongids East along Ind.Ocean (sivapiths?) vs hominids West along Medit.Sea-coasts + inland along rivers: dryopiths etc., Trachilos BP footprints Crete, etc.
- hominids died out (drying? heat? Zanclean mega-flood? ...) except in Red Sea:
- c 8-7 Ma Gorilla/Homo-Pan split: Gorilla followed northern Rift swamp forests ->afarensis->boisei->low+highland gorillas... e.g. my Hum.Evol.papers 1990, 1994, 1996,
- c 6-5 Ma Homo & Pan split, Red Sea opened into Gulf (Francesca Mansfield: caused by Zanclean mega-flood 5.33 Ma): Pan went right along E.Afr.coasts -> southern Rift ->africanus->robustus->chimps+bonobos // gorillas -> knuckle-walk, longer arms...
- Homo went left along S.Asian coasts -> Java 2 Ma: Mojokerto -> Flores etc.
- early-Pleist. erectus-like Homo dispersal along coasts -> Africa, Europe...
Google "human evolution Verhaegen"
https://www.gondwanatalks.com/l/the-waterside-hypothesis-wading-led-to-upright-walking-in-early-humans/
:-)