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Concavenator — Human Evolution

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Published: 2019-10-31 23:49:17 +0000 UTC; Views: 70918; Favourites: 447; Downloads: 0
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Description Behold: human evolution, from the earliest self-replicating molecules to the rise of our own species, over four billion years, condensed into a single picture.

This work was inspired mainly by two excellent books that trace evolution from the origin of life to present times (David Peter's From the Beginning, 1995) or backward (Richard Dawkins and Yan Wong's The Ancestor's Tale, 2004) (plus a wonderful, but somewhat outdated one - Ernst Haeckel's Evolution of Man , 1874 for the original German edition).
I have divided the story into 35 arbitrary stages, and depicted the most salient features of each (see the bibliography below). They proceed in a zigzag pattern starting from the top left and ending at bottom left; the 36th plate contains notes and additional details. A map in the bottom left of each plate shows the approximate positions of continents (only reliable from stage 10 or so) and the approximate location our ancestor inhabited (at least from stage 14 or so).
None, except the last one, are meant to depict a specific named species (though some are more closely based on one than others); stages 8 and 12 are outright chimerical. The main fossil or living models used to reconstruct each stage are given just below its name, e.g. "Ophiacodon, Sphenacodon, Haptodus" for stage 20. I do not claim these models to be direct ancestors of the human lineage, just useful for their reconstruction. Living models are marked with a *. Of course, the first 8 models are not based on any meaningful fossil evidence, but only on comparisons with living organisms.
The main innovative features of each stages are marked with curved arrows. "Ma" and "ka" are short for "million years (Megaannum) ago" and "thousand years (kiloannum) ago", respectively.
As in The Ancestor's Tale, "concestor" is short for "most recent common ancestor" of a group. Consider e.g. stage 23, which is marked as "concestor of Mammalia". This is the last organism to be the ancestor of all mammals living today. Of course, all its ancestors (i.e. stages 1-22) are common ancestors of all mammals as well, but stage 23 is the most recent. Stage 23 is marked as ancestor of Monotremata, but it's also the ancestor of stage 24, and hence of all Theria (stage 23 is ancestor of all Theria, but not concestor: stage 24 is); Monotremata + Theria do in fact make up all living Mammalia. And so on. The first concestor is stage 3, concestor of Biota, i.e. all cellular life (its ancestors, stages 1 and 2, are not "most recent common ancestors of anything). The last is stage 35, which is concestor of all living humans.
Obviously a similarly intricate scheme could be drawn for all species living today. But since I assume most users here are humans, as I am, I chose to focus on the human lineage, which shouldn't be interpreted in a teleological sense. The choice to depict the mammalian stages as male or female was more or less arbitrary, or else was meant to show sex-specific features.

Summary

1. Protobiont (4200 Ma?): self-replicating rybozyme
2. Protocell (4000 Ma?): double-layer phospholipid membrane, isolated RNA-based genes, ribosomes
3. Prokaryote (3800-3500 Ma?): cell wall between two membranes, trans-membrane proteins, cytosol, linked DNA-based genes, binary fission and transduction
4. Eukaryote (2700-1600 Ma?): nucleus, chromatin and chromosomes, mitochondria and respiration, inner membrane (Golgi apparatus,endoplasmic reticulus, etc.), cytoskeleton, cilia and flagella, gene splicing, loss of outer membrane and cell wall
5. Blastaea (1200-800 Ma?): hollow sphere of flagellate cells, lifecycle with meiosis and fertilization, cadherin junctions, extracellular collagen, specialized gametes
6. Gastraea (900-700 Ma?): inner (endodem) and outer (ectoderm) cell layers, digestive cavity, epithelia, nervous coordination, apico-basal axis
7. Acoelomate worm (800-650 Ma?): bilateral symmetry and antero-posterior axis, closed gut with ventral pharynx, gonads and gonopore, nerve cords, muscle cells, pigment-spot eyes, statocyst ?and sensory cilia, mesoderm
8. Pseudocoelomate worm (650-600 Ma?): liquid-filled pseudocoelom, intestine with mouth and anus, musculo-cutaneous sac and undulation, nerve net, nephridia, Hox genes, striated muscle, ?cup-eyes, ?blood vessels, ?separate sexes
9. Dipleurula/early deuterostome (600-540 Ma?): ciliate pharynx with pharyngeal slits, segmented coelom, visceral muscles, peritoneum and mesenteries, anus forming from blastopore, mesoderm from gut pouch
10. Acrane chordate (550-520 Ma): notochord, hollow spinal cord with brain, pineal and pituitary glands, gill bars and slits, endostyle, heart and aorta, atrium and atriopore, V-shaped myomeres, post-anal tail, ?pinhole eyes
11. Basal vertebrate (530-510 Ma): vertebrae, 3-part brain with olfactory bulb and otic capsule, cranium, camera eyes, single nostril, muscular pump feeding, thyroid, 2-chamber heart, stomach and liver, kidneys, non-segmented gonads, W-shaped myomeres, unpaired fin, hemoglobin in blood, neural crests, loss of atriopore
12. Ostracoderm (480-430 Ma): bony head armor and ?scales, pineal eye, asymmetric tail fin, paired fin flaps, lateral line, articulated gill arches
13. Early gnathostome (420-410 Ma): jaws and hyoid arch, two pairs of nostrils, spiracle, operculum, paired pectoral and pelvic fins, dorsal and caudal fins, ganoid scales, urinary bladder, myelin sheath, bone osteons
14. Basal sarcopterygian (420-390 Ma): shoulder and pelvic girdle, lobed fins, lungs, choanae, lacrimal duct, jointed braincase, teeth with palatal and coronoid fangs, ?muscular tongue
15. Tetrapodomorph (390-380 Ma): larger eyes and spiracle, flattened head, larger vertebrae, loss of anal and dorsal fins
16. Fishapod (375-360 Ma): feet with 7-8 digits, more compact skull with interdigitate junctions, eardrum and columella, flexible neck, larger ribs, shoulder girdle detached from skull, pelvic girdle attached to sacral vertebra, ?tear glands and eyelids, ?3-chamber heart, vestigial tail fin, loss of scales (except gastralia), fin rays, coronoid fangs, external gills, and operculum
17. Land tetrapod (350-330 Ma): stronger wrists and ankles, 5-digit limbs, ?salivary glands, ?vomeronasal organ, thinner ribs, separated pelvic bones, complete loss of gills, tail fin, and lateral line
18. Reptiliomorph (330-300 Ma): limb-based locomotion (body lifted off ground), deeper narrower head, rib-assisted breathing, ?denser dermis, ?small keratin scales, reduced notochord
19. Basal amniote (315-305 Ma): claws and scales of keratin, cervical and 2 sacral vertebrae, notochord reduced to disks, production of urea, adrenal glands, loss of eardrum, amniotic egg with shell and internal membranes, ?internal fertilization
20. Pelycosaur (300-275 Ma): temporal fenestra, lamina from angular, teeth of different size, angled jaw muscles, ?diaphragm
21. Therapsid (270-255 Ma): large canine fangs, wider temporal fenestra, stapes, reduction of lumbar ribs, 3 sacral vertebrae, more upright limbs, shorter toes, ?smaller scales, loss of gastralia
22. Cynodont/cynognathian (250-210 Ma): double jaw joint, reduced post-dentary bones, zygomatic arch (orbit fused to temporal fenestra), multicusped teeth, secondary palate, nasal turbinates, obturator foramen, fully upright hindlimbs, distinct heel, ?endothermy, ?insulating hair and whiskers, ?lung alveoli, loss of pineal eye
23. Mammaliaform (220-180 Ma): multi-cusped molars with double root, occluding diphyodont teeth, squamosal-dentary jaw joint, middle ear bones, new eardrum and cochlea, development of smell over sight, 4-chamber heart, colon, penis with cavernous tissue and baculum, hairy and glandular skin, milk glands, loss of three-color sight, ?parental care
24. Pantothere (160-125 Ma): middle ear bones separate from jaw, ball-in-socket shoulder joint, fully upright forelimbs, marsupial bones, mammaries with nipples, anus separated from uro-genital opening, paired uteri and vaginae, XY sex chromosomes, ear pinnae, loss of egg-laying
25. Proteuthere (125-75 Ma): medial acromion on scapula, chorio-allantoic placenta, corpus callosum, single vagina, loss of marsupial bones
26. Tree shrew/plesiadapiform (66-55 Ma): bicornuate uterus, reduction in offspring number, tree-climbing and leaping, grasping digits, ?tooth comb, ?testicles in scrotum, ?non-sheathed penis, ?rounded ear pinnae
27. Prosimian (55-40 Ma): larger brain, forward-facing eyes, auditory bulla, three-color sight, prehensile hands and feet, grooming claw, postorbital bar, rounded molar cusps, omnivorous diet, sociality ?with vocal communication
28. Eosimiid/parapithecid (45-30 Ma): postorbital septum, fused upper lip, fused frontal bone and jaw, thicker endometrium, oestrus replaced by menstrual cycle, flat finger- and toenails, single pair of mammaries, metatarsifulcrumating foot, loss of tapetum lucidum and grooming claw
29. Propliopithecoid (34-28 Ma): 32 teeth, contact between frontal and sphenoid, downward-opening nostrils, bony ear meatus, ?hairless face and facial expressions, loss of vomeronasal organ and whiskers
30. Basal hominoid (25-12 Ma): broader and flatter chest, dorsally-placed scapula, rotatory wrist, hinge joint in elbow, large canines, bunodont molars, large maxillary and frontal sinuses, loss of tail (coccyx)
31. Tree-dwelling hominid (14-8 Ma): thicker tooth enamel, brow ridges, deeper brain gyri, ?appendix, ?multi-male/multi-female philopatric groups, ?bipedality on treetops, ?common tool use
32. Bipedal hominin (7.0-4.4 Ma): permanent bipedality, foramen magnum under skull base, larger lumbar vertebrae and S-shaped spine, smaller canines, premaxilla fused to maxilla, ?loss of baculum, ?monogamy with polygynic tendencies
33. Australopithecine (4.5-2.0 Ma): non-opposable toes, longer thumb and shorter palm, cone-shaped ribcage, bowl-shaped pelves with flaring iliac crest, large gluteus maximus and tensor fasciae latae
34. Primitive human (2.4-0.1 Ma): prominent brow ridges, parabolic tooth arcade, barrel-shaped ribcage, narrower hips, arched foot plants, long Achilles tendon, nuchal ligament, denser sweat gland and melanocytes ?longer head hair and beard, ?smaller gut, ?fat deposits in breasts and hips, ?common meat consumption, loss of most hair coating, production of stone tools, use of fire
35. Modern human (0.3-0.0 Ma): rounder braincase with vertical forehead, smaller jaws and teeth, protruding chin and nose bridge, thinner long bones, lower larynx, loss of brow ridges, clothing and ornamentation, compound tools, ritual behavior and artwork


Bibliography
Maps based on Christopher Scotese's Palaeomap Project .

Books

Peters D (1991), From the Beginning: The Story of Human Evolution, Morrow Junior Books (stages 1-35)
Cairns-Smith AG (1982). Genetic takeover and the mineral origins of life, Cambridge University Press (stages 1-3)
Maynard Smith J, Szathmary E (1995). The Major Transitions in Evolution, Oxford University Press (stages 1-5)
Dawkins R, Wong Y (2004). The Ancestor's Tale, Weidenfeld & Nicolson (stages 3-5, 8, 10, 14, 17, 19, 25, 27, 29, 31)
Hickman CP, Roberts LS, Keen SL, Larson A, l'Anson H, Eisenhour DJ (2008). Integrated Principles of Zoology (14th edition), McGraw-Hill (stages 4-11)
Nielsen C (2012). Animal Evolution: Interrelationships of the Living Phyla (3rd edition), Oxford University Press (stages 5-10)
Feinberg TE, Mallatt JM (2016). The Ancient Origins of Consciousness: How the Brain Created Experience, MIT Press (stages 7-8, 10-12, 25)
Kardong KV (2012). Vertebrates: Comparative Anatomy, Function, Evolution (6th edition), McGraw-Hill (stages 9-17, 19, 22-23)
Striedter GF (2005). Principles of Brain Evolution, Sinauer Associates (stages 11, 23, 27, 31, 34)
Clack JA (2002). Gaining Ground: The Origin and Evolution of Tetrapods, Indiana University Press (stages 14-20)
Steyer S (2012). Earth Before the Dinosaurs, Indiana University Press (stages 14-22)
Sumida SS, Martin KLM (ed) (1997), Amniote Origins: Completing the Transition to Land, Academic Press (stages 17-20)
Kemp TS (2005). The Origin and Evolution of Mammals, Oxford University Press (stages 19-26)
Kielan-Jaworowska Z, Cifelli RL, Luo ZX (2004). Mammals from the Age of Dinosaurs: Origins, Evolution, and Structure, Columbia University Press (stages 22-25)
Fleagle JG (1998). Primate Adaptation and Evolution (2nd edition), Academic Press (stages 26-34)
Dixson AF (2009). Sexual Selection and the Origins of Human Mating Systems, Oxford University Press (stages 28, 31-34)
Boyd R, Silk JB (2009). How Humans Evolved (5th edition), W. W. Norton & Company (stages 31-35)
Tattersall I (2012). Masters of the Planet: The Search for Our Human Origins, Palgrave Macmillan (stages 32-35)

Papers

Note: PNAS = Proceedings of the National Academy of Sciences of the United States of America; PTRS B = Philosophical Transactions of the Royal Society B

Kitadai N, Maruyama S (2018). Origins of building blocks of life: A review. Geoscience Frontiers, 9(4):1117-1153 (stage 1) (link )
Cavalier-Smith T (1987). The Origin of Cells: A Symbiosis between Genes, Catalysts, and Membranes. Cold Spring Harbor Symposia on Quantitative Biology, 52(0):805-824 (stages 1-4) (link )
Woese C (1998). The universal ancestor. PNAS, 95(12):6854-6859 (stages 2-3) (link )
Sousa FL, Thiergart T, Landan G, Nelson-Sathi S, Pereira IAC, Allen JF, Lane N, Martin WF (2013). Early bioenergetic evolution. PTRS B, 368(1622):20130088 (stages 2-3) (link )
Hsia CCW, Schmitz A, Lambertz M, Perry SF, Maina JN (2013). Evolution of Air Breathing: Oxygen Homeostasis and the Transitions from Water to Land and Sky. Comprehensive Physiology, 3(2):849-915 (stages 3-4, 14-16) (link )
Garg SG, Martin WF (2016). Mitochondria, the Cell Cycle, and the Origin of Sex via a Syncytial Eukaryote Common Ancestor. Genome Biology and Evolution, 8(6):1950-1970 (stage 4) (link )
Abedin M (2010). Cadherin evolution and the origin of animals. PhD thesis, University of California, Berkeley (stages 4-5) (link )
Grosberg RK, Strathmann RR (2007). The Evolution of Multicellularity: A Minor Major Transition? The Annual Review of Ecology, Evolution, and Systematics, 38:621-654 (stage 5) (link )
Michod RE, Roze D (2001). Cooperation and conflict in the evolution of multicellularity. Heredity, 86:1-7 (stage 5) (link )
Jekely G (2011). Origin and early evolution of neural circuits for the control of ciliary locomotion. PTRS B, 278:914-922 (stage 6) (link )
Baguna J, Riutort M (2004). The dawn of bilaterian animals: the case of acoelomorph flatworms. BioEssays, 26(10):1046-1057 (stages 6-7) (link )
Ruppert EE, Carle KJ (1983). Morphology of metazoan circulatory systems. Zoomorphology, 103(3):193-208 (stages 7-10) (link )
Ou Q, Conway Morris S, Han J, Zhang Z, Liu J, Chen A, Zhang X, Shu D (2012). Evidence for gill slits and a pharynx in Cambrian vetulicolians: implications for the early evolution of deuterostomes. BMC Biology, 10:81 (stages 9-10) (link )
Shu D (2003). A paleontological perspective of vertebrate origin. Chinese Science Bulletin, 48(8):725-735 (stages 9-11) (link )
Gai Z, Donoghue PCJ, Zhu M, Janvier P, Stampanoni M (2011). Fossil jawless fish from China foreshadows early jawed vertebrate anatomy. Nature, 476(7360):324-327 (stage 12) (link )
Johanson Z (2010). Evolution of Paired Fins and the Lateral Somitic Frontier. Journal of Experimental Zoology, 314B:347-352 (stages 12-13) (link )
Mallatt J (1996). Ventilation and the origin of jawed vertebrates: a new mouth. Zoological Journal of the Linnean Society, 117(4):329-404 (stages 12-13) (link )
Dupret V, Sanchez S, Goujet D, Tafforeau P, Ahlberg PE (2014). A primitive placoderm sheds light on the origin of the jawed vertebrate face. Nature, 507(7493):500-503 (stages 12-13) (link )
Zhu M, Yu X, Ahlberg PE, Choo B, Lu J, Qiao T, Qu Q, Zhao W, Jia L, Blom H, Zhu Y (2013). A Silurian placoderm with osteichthyan-like marginal jaw bones. Nature, 502(7470):188-193 (stages 13-14) (link )
Clement AM, King B, Giles S, Choo B, Ahlberg PE, Young GC, Long JA (2018). Neurocranial anatomy of an enigmatic Early Devonian fish sheds light on early osteichthyan evolution. eLife, 7:e34349 (stage 14) (link )
Zhu M, Yu X, Choo B, Qu Q, Jia L, Zhao W, Qiao T, Lu J (2012). Fossil Fishes from China Provide First Evidence of Dermal Pelvic Girdles in Osteichthyans. PLoS One, 7(4):e35103 (stage 14) (link )
MacIver MA, Schmitz L, Mugan U, Murphey TD, Mobley CD (2017). Massive increase in visual range preceded the origin of terrestrial vertebrates. PNAS, 114(12):E2375-E2384 (stages 15-17) (link )
Jankowski R (2012). The Complex Formation of the Secondary Palate and Nose in Evolution, in The Evo-Devo Origin of the Nose, Anterior Skull Base and Midface, Springer-Verlag (stages 17-22) (link )
Lambertz M, Shelton CD, Spindler F, Perry SF (2016). A caseian point for the evolution of a diaphragm homologue among the earliest synapsids. Annals of the New York Academy of Sciences, 1385(1):3-20 (stages 19-20) (link )
Oftedal OT (2002). The Mammary Gland and Its Origin During Synapsid Evolution. Journal of Mammary Gland Biology and Neoplasia, 7(3):225-252 (stages 19-24) (link )
Hoffmann EA, Rowe TB (2018). Jurassic stem-mammal perinates and the origin of mammalian reproduction and growth. Nature, 561(7721):104-108 (stage 22-23) (link )
Luo ZX (2015). Origin of the mammalian shoulder, in Major Events in the History of Vertebrate Life, University of Chicago Press (stages 22-25) (link )
Luo ZX, Crompton AW, Sun AL (2001). A New Mammaliaform from the Early Jurassic and Evolution of Mammalian Characteristics. Science, 292(5521):1535-1540 (stage 23) (link )
Martin T, Marugan-Lobon J, Vullo R, Martin-Abad H, Luo ZX, Buscalioni AD (2015). A Cretaceous eutriconodont and integument evolution in early mammals. Nature, 526:380-384) (stage 24) (link )
Wildman DE, Chen C, Erez O, Grossman LI, Goodman M, Romero R (2006). Evolution of the mammalian placenta revealed by phylogenetic analysis. PNAS, 103(9):3203-3208 (stage 25) (link )
Gingerich PD (2012). Primates in the Eocene. Palaeobiodiversity and Palaeoenvironments, 92(4):649-663 (stages 27-28) (link )
Franzen JL, Gingerich PD, Habersetzer J, Hurum JH, von Koenigswald W, Holly Smith B (2009). Complete Primate Skeleton from the Middle Eocene of Messel in Germany: Morphology and Paleobiology. PLoS One, 4(5):e5723 (stages 27-29) (link )
Vorobyev M (2004). Ecology and evolution of primate colour vision. Clinical and Experimental Optometry, 87(4-5):230-238 (stages 27-29) (link )
Williams BA, Kay RF, Kirk EC (2010). New perspectives on anthropoid origins. PNAS, 107(11):4797-4804 (stages 28-29) (link )
Strassmann BI (1996). The Evolution of Endometrial Cycles and Menstruation. The Quarterly Review of Biology, 71(2):181-220 (stages 28-30) (link )
Andrews P (1992). Evolution and environment in the Hominoidea. Nature, 360(6405):641-646 (stages 30-31) (link )
de los Rios MP, Moya-Sola S, Alba DM, Fortuny J (2010). Maxillary and frontal sinuses in Eurasian Miocene hominoids: phylogenetic implications. Cidaris, 30:223-226 (stages 30-31) (link )
Villmoare BA, Dunmore C, Kilpatrick S, Oertelt N, Depew MJ, Fish JL (2014). Craniofacial modularity, character analysis, and the evolution of the premaxilla in early African hominins. Journal of Human Evolution, 77:143-154 (stages 31-34) (link )
Gruus LT, Schmitt D (2015). The evolution of the human pelvis: changing adaptations to bipedalism, obstetrics and thermoregulation. PTRS B, 370(1663):20140063 (stages 31-35) (link )
Zollikofer CPE, Lieberman DE, Ponce de Leon MS, Guy F (2005). Virtual reconstruction of Sahelanthropus tchadensis. Nature, 434(7034):755-759 (stage 32) (link )
Lovejoy CO (2009). Reexamining Human Origins in Light of Ardipithecus ramidus. Science, 326(74):e1-e8 (stages 32-33) (link )
Carroll SB (2003). Genetics and the making of Homo sapiens. Nature, 422(6934):849-857 (stages 32-35) (link )
Bramble DM, Lieberman DE (2004). Endurance running and the evolution of Homo. Nature, 432(7015):345-352 (stages 33-34) (link )
Wood B, Collard M (1999). The Changing Face of Genus Homo. Evolutionary Anthropology, 8(6):195-207 (stages 33-35) (link )
Frost P (2015). Evolution of Long Head Hair in Humans. Advances in Anthropology, 5(4):274-281 (stage 35) (link )
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"Son of man a man in time you'll be"

The most complete human evolution I've seen

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