Homo sediba

Credit: Elisabeth Daynès and S. Entressangle.

*Homo sediba* is an extinct species of hominin that lived in South Africa during the Early Pleistocene.

## History

—Omphemetse Keepile about the naming of Karabo
Matthew Berger, son of prominent paleoanthropologist Lee Berger, first discovered MH1 at Malapa Cave in the Cradle of Humankind. His son was 9 at this time (August 15, 2008), and stumbled across the fossil while exploring the digsite with his father. They then continued excavation, which yielded more remains. Omnphemetse Keepile, a then-17-year-old from St Mary's School, Johannesburg, opted to name MH1 Karabo, which means "answer" in Tswana because it had the potential to rewrite human evolution, after winning a naming contest. More specimens were later found, including MH2 "Issa". In 2022, an additional elements of Issa were announced, providing new data on locomotion and posture.

The layer *H. sediba* was initially discovered in was suggested to sate within the Early Pleistocene, with U-Pb dating of flowstone finding a date of 2.026±0.021 million years. Archaeomagnetic dating finds a dating of 1.95-1.78 million years. The flowstone was firmly dated in 2011 to 1.977±0.002 million years, again using U-Pb dating. Thus, *H. sediba* is frequently dated to 1.98 million years old, but dating of the fossils is difficult to determine. Dembo et al. (2015) state that the remains may be as much as 800,000 years too early, making a new 1.18 million-year date. However, it is more likely to be 300-500,000 years off, which would make a reasonable temporal range of 1.68-1.48 million years ago, with an absolute latest of 1.18 million years old. This would still place it into the Early Pleistocene.

    Malapa Cave has networks around it with interconnected openings within a 500×100 meter area. The base of Malapa is (at most) 30 meters in depth, with the cave intersecting at a north-northeast and north-northwest chert-filled fracture. Hominins are found in a 3.3x4.4x3.5 meter site at the north-northwest fracture, exposed by limestone mining in the early 1900s. The cave bears 5 sedimentary facies (A-E), and *H. sediba* from facies D and more hominins from facies E. MH1 and 2 were vertically separated by (at most) 40 centimeters. Facies D is 1.5 meters thick, light in colour and overlying flowstone. Small peloids are seen fused into large, irregular groups that indicate the depositional environment was water-logged, and the peloids themselves may be fecal matter from soil microbes. MH1 and 2 were deposited quickly based on preservation state, and moved little during this time, where they cemented soon sfter deposition in a phreatic environment. No evidence of scavenging are present, which indicate that the death/deposition site was inaccessible to predators. This indicates that Malapa was a deathrap that inconspicuously opened at the surface, which may have lured animals through the scent of water and dead animals down the shaft, where they would have fallen and died. Large debris flow cased remains to have been deposited deeper into the cave along a subterranean stream, maybe due to a heavy rainstorm. The chamber would have eventually collapsed and filled with mud.. Also present in the Malapa deathtrap are antelopes, hyenas, nimravids and plants.

At Gladysvale Cave, ongoing excavations have uncovered well upwards of 10 hominid fossils representing several individuals, including crania, ribs, teeth, mandibular fragments, a pelvic fragment and limb scraps. Several appear to be juveniles, one having massive (some un-erupted) molars, but Berger is unsure if they are large juveniles or small adults, as the mandible/limb proportions are strange assuming certain fossils can be grouped as individuals. The pelvic fragment appears very similar to that of *H. sediba*, but further classification is pending.

## Description

Only the cranial vault of MH1, with a volume of 363 and the very back of the brain being 7-10. Using KNM-ER 23000, *Homo habilis* KNM-ER 1813 and 1805, KNM-ER 1470 and Sts 19, the cerebellum was estimated at 50 cc. These together make a full volume of 420-440, typical of australopithecines. Using apomorphic primate adult and neonate brain size, the brain may have been 153-201 cc, similar to the presumed capacity for australopithecines. The brain may have been configured like australopithecines, but the orbitofrontal cortex seems to be more apomorphic. It's anatomy is most similar to *A. africanus*, but the cranium was smaller, the cranial vault was transversely wider, the parietal has more vertically-inclined walls and the temporal lines are more widely spaced. Like *Homo*, the brow is less pronounced, the cheeks are less flared, the face has less prognathism and a slight chin is present. It has been suggested that *Homo*-like traits may have been lost during aging. If prognathism is measures using the anterior nasal spine rather than the base of the nose, porgnathism in MH1 would fall into the hypodigm of *A. africanus*. The teeth are small for an australopith, within the range of early *Homo*, but the molars increase in size towards the back of the mouth and the cusps are closer spaced, unlike early *Homo*. The mandibular ramus is different in bith specimens; MH1 is taller and wider, with the front and back border nearly vertical and parallel, contrasting the nonparallel borders of MH2, who has a concave front border. The croconoid process of MH1 is angled to the back with a deep, asymmetrical mandibular notch, where MH2 has an uncurved coronoid process with a shallow mandibular notch. Compared to other hominids, these could be sexual dimorphism or immaturity. Like all of the human lineage, high variability is present, which can muddle taxonomy. It was suggested that mandibular rami can be used to distinguish morphotypes, but a 2022 publication led by John Hawks suggests that such high variability does not allow for good specific differentiation. They also determine that similar dietary and mandibular developments as seen in related hominins were present in this species.

MH1 and 2 are roughly the same time, about 30-36 kilograms (66-79 pounds), smaller than contemporaries but around the size of australopiths. MH1 has the same skeletal development as a 12-year-old apomorphic human or a 9-year-old chimpanzee. Like contemporaries, it may have had an apelike growth rate based on dental development, so it may have reached 85% of adult size assuming a chimpanzee-like trajectory or 80% assuming a humanlike trajectory. This would then be 150-156 centimeters tall. MH1 has 4 cervical, 6 thoracic and 2 lumbar vertebrae; MH2 has 2 cervicals, 7 thoracic and 1 sacral. Lordosis and joints of the cervicals present a humanlike head posture, but the rest of the neck is stiffer and more plesiomorphic. A apomorphic brachial plexus is lacking, which is responsible for nerves and muscle innervation in the arms and hands that enhance motor control. Like apomorphic forms, the flexible lumbar series has 5 vertebrae, and lumbar lordosis consistent with bipedal walking. However, a seemingly highly mobile lower back may exaggerate lumbar lordosis, which could be involved in counteracting inward torques while walking in a hyperpronated gait, as proposed for *H. sediba*. MH1 preserved 2 upper thoracic, 1 mid-thoracic and 3 lower thoracic ribs; MH2 preserves 4 consecutive upper-to-mid thoracic and 3 lower thoracic ribs and vertebrae. This indicates an apelike constricted upper chest, with a broad, humanlike lower chest. The upper chest, narrow, would have hindered arm swing while walking and restricted the ribcage from heavy breathing, and thus fast walking and long-distance running. It also has a humanlike narrow waisr, with repositioned external oblique muscles. The wider iliocostalis muscles would improve walking efficiency by counteracting sidewards torso flexion. The pelvis is similar to early *Homo*, African *Homo erectus*, KNM-ER 3228 and OH 28. More buttressing at the acetabulum and sacrum improves hip extension, the iliofemoral ligament is enlarged to shift the weight behind the center of rotatio at the hip, more buttressing along the acetabulum and iliac blade improves alternate pelvic tilt and a spaced out acetabulum and lschial tuberosity reduces the moment arm at the hamstrings. This allows for a humanlike stride and the joint appears to have has a more apomorphic load bearing capability than *Homo habilis* OH 62. The birth canal appears to be more gynaecoid than australopiths, though not completely since *H. sediba* neonates would have been smaller-brained. Like apomorphic forms, the birth canal decreases in diameter sagitally and the pubis bone curves upwards.

Like australopiths and early *Homo*, the arms were proportionally long, with a brachial index of 84 and large joint surfaces. The shoulders are in a shrugging position, with a well developed axillary border and a well-defined conoid tubercle. Muscle scarring on the clavicle indicates a humanlike range of motion. The size of the glenoid cavity and its angle with the spine are most similar to orangutans, though the shoulder is most similar to Panina and Australopithecina. The humerus bears a low degree of torsion, unlike humans anf African hominids, which suggests a placement farther from the midline like *Homo*, but positioned higher up than back like australopithecines. The apelike arms are more marked in *H. sediba* than *A. afarensis*; if it is ancestral to *H, sediba*, this could indicate an adaptive shift towards arboreality. At the elbow, the lateral and medial epicondyles (of the humerus) are elongate, like australopiths and non-human African hominids. Te humerus bears a developed crest at the elbow that supports the brachioradialis, which flexes the forearm. Like non-human apes, a strong attachment for the biceps on the radius and for the triceps at the ulna are seen, However, less mechanical advantage are found in the biceps and brachialis. The ulna supports attachment for the flexor carpi ulnaris. The olecranon fossa is large, deep and has a prominent trochlear keel; this stabilizes the arms during extension. The fingers are long, robust and curved, supporting a strong flexor digitorum superficialis, which flexes the fingers. This is seen as evidence arboreal behaviour. The hand has a long thumb and long fingers, like *Homo*, which suggests precision grip needed for crafting complex stone tools. The ankle, knee and hip, like australopithecines, suggest habitual bipedalism. The legs are similar to the legs of *A. afarensis*. The ankle and possibly Achilles tendon are humanlike. The talus is stout and like non-human apes, with a medially-twisted neck and a low torsion angle. It is debated if *H. sediba* had a humanlike foot arch. The heel bone angles at 45°, angled from front-to-back (strongest at the peroneal trochlea, which was robust and indicates a strong peroneus muscle that extended from the calf to the ankle). The foot lacks a lateral plantar tubercle, unlike moder forms and *A. afarensis*. The gracile heel body and the robust malleolus are apelike, being a less efficient force transfer between the heel and the talis and apelike mobility at the midfoot. This condition is most similar to gorillas, with function equivalent to *A. africanus*. All these suggest that the arms were adapted to climbing and the feet and hips were adapted to bipedal walking.

## Paleobiology

Hyperpronation may have been present while walking, causing exaggerated weight transfer during the stance phase. The rectus femoris and biceps femoralis in *H. sediba* are consistent with midline-directed strains across the legs, hips and knees, which is not ideal for apomorphic humans, where this may cause plantar fasciitis, shin splints and tibial stress fractures. To counteract this, *H. sediba* used the mobile midfoot to prevent stressful loads at the ankle. A hyperpronated gait and related adaptations are not seen in any other hominins, and it is unclear why this was developed. The mobile midfoot would have been beneficial in extensive climbing, which it was later have found to have done. Thus, hyperpronation may be a compromise between bipedal walking and arboreal climbing.

### Diet

Analysis of phytoliths and plaque in the teeth of known specimens shows a diet almost exclusive to C3 forest plants, despite a wide variety of C4 plants in their environment. This is seen in savanna chimpanzees and presumed for *Ardipithecus*, but different than any early hominin. 38 phytoliths were recovered from 2 teeth of MH1, 15 consistent with dicots, 9 with monocots and 14 indeterminate. The monocots were probably sourced from C3 grasses and sedges from well-watered and shady areas. Others were from fruits, leaves, wood or bark. Though bark is often consumed by other primates for high protein and sugar, no other hominins are known to have eaten them regularly. Dental microwearing suggests *H. sediba* ate hard foods, with complexity values ranging from *Homo erectus* and robust *Paranthropus robustus*. The jaw does not appear well adapted to producing and enduring high force compared to other hominins, which may indicate that it was not highly dependent to process these foods. This diet is consistent with it being partially arboreal, since it could have occupied smaller home ranges than savanna chimpanzees, being able to fall back on bark and fracture-resistant foods.

### Life History

The pelvic inlet for female *H. sediba* is estimated to have been 80.8x112.4 millimeters (long x broad), and with the largest neonate head size of 89.2 millimeters, so the neonate likely entered the inlet transversely, similar to other hominins. The midplane of the inlet is constricted to a minimum of 96.9 millimeters, so the neonate may not have needed to be rotated while birthing. The inlet dimensions were calculated through composite reconstruction involving a juvenile male(?) ischium, so the birth canal may have actually been larger. The shoulders are estimated at 74.3 millimeters across, so they would have restricted birth more than the head. The neonate (at the most constriction) would have fit 92.1% of the birth canal, allowing for complete non-rotational birthing room, as in non-human apeas and possibly (other methods have been proposed) other australopithecines. Though it was possible to pass without rotation, the midplane expands anterposteriorly, and more space for the neonate would have been available if it rotated to the longest length of the head to match with the expansion., using a cranial capacity estimated of 145,8-180,4, the head size would have been 73x89 millimeters. MH1 has a very different growth trajectory than other hominins. The nasomaxillary complex indicates much bone resorption, most at the front teeth roots. This contrasts *A. africanus* and *A. afarensis*, Because resorption occurs close to the cheekbones, this may explain why MH1 does not present the robust zygomatic. Tooth eruption likely did not affect lower face remodeling, since MH1 had their permanent teeth. Smaller cheek teeth may permit a mesognathic face. The olive colobus monkey may have had a similar diet, and has similar facial growth patterns, which suggests resorption and deposition patterns reflecting different jaw-loading. The orbital margins are curved, unlike *A. africanus*, which indicate bone deposition in *H. sediba* in regions where bone resorption occurs in *A. africanus.*

### Integument

A 2022 study compared hard and soft tissues in hominin noses to make estimates based on extinct species, where *H. sediba* was said to have had a raised, but flat nose. It has been suggested that Karabo and Issa, a woman and a boy, would have mummified after death, since they may have died in a deep, anoxic pool of water. This is supported by a thin coating over the bones which was interpreted as mummified skin during preparation, which had led to hypothesizing that DNA, skin and keratin may be recovered from the fossils, specially prepared, possibly yielding valuable genetic and physical data. This would be the oldest mummified human body, and made headlines during preparation. A cast of the skin was shown to John Hawks and several other scientists by Berger, who had no idea how to test if they were organic. Hawks decided to announce this limbo in 2011, starting an investigation campaign that was later closed. Rachelle Keeling published 5 theses about mummified skin, confirming that it had been recovered, and noted that the specimens bear many organic features, including the signatures mummified skin would have, and her research started in 2013. The tissue was found over the MH1 cranium, in the center of an MH1 cervical vertebrae and on the mandible of MH2, but there has been no further exploration of Malapa for soft tissue. The thin coating bears signs of structure, possibly hair follicles, blood vessels and other internal components. In 2022, Berger confirmed that the specimens were skin and brought them and a block of *Homo naledi* to the European Synchotron for analysis.

## Paleobiology

The right lamina of the 6th thoracic in MH1 presents a penetrating bone tumour that was likely a beneign osteoid osteoma, which penetrates 6.7 millimeters deep and is 5.9 millimeters wide. At death, this was still active, though it did not penetrate the neural canal, so it likely did not cause neurological complications. No evidence of scoliosis is present, but it may have affected movement of the shoulder blade and the upper right quadrant of the back, which may have caused acute or chronic pain, muscular disturbances or muscle spasms. Lesions near the insertion of the trapezius, erector spine and rhomboid major may have limited movement patterns while climbing. MH1 is the earliest diagnosed case of cancer by at least 200,000 years. Tumours are thus rare in the fossil record due to low incidence rate in primates, with early hominins have similar rates. MH1 developing a bone tumour falls in line with the general trend of youth. MH1 and 2 bear perimortem blunt force trauma in the bones, likely caused by a fall onto a semi-cushioned sloping pile of gravel, sand and bat guamo from a height of 5-10 meters. These are most found in the mandible and teeth, though these are also consistent with being hit with a falling object and the fall itself. MH2 bears evidence of bracing for injury, since the forearm and head loaded most of the force during a blow to the chest on the right side of the body.

## Classification

Berger et al. (2010) named *H. sediba* as *Australopithecus sediba* with MH1 as the holotype and MH2 as a paratype; *sediba* meaning "fountain" or "wellspring" in the local Sesotho. Many traits were observed to be common with African *Homo erectus*, mostly in the pelvis and legs, so they posited it was a transitionary form between *Homo* and *Australopithecus*. This was also found in the dental traits, where the teeth are similar to the ancestor of *Homo*. However, a dating of 1.95-1.78 million years contradicts the estimate of *Homo* evolving 2.33 million years ago from *Homo habilis*; LD 350-1 is the oldest *Homo* fossil with an age of 2.8-2.75 million years ago. To reconcile this, they suggested that *H. sediba* evolved from a population of *A. africanus* (who are roughly contemporaneous and contend with the supposed split), which would imply a 800,000-year ghost lineage between *H. sediba* and *A. africanus*. It was also suggested that it was the direct ancestor of African *Homo erectus* rather than *Homo habilis* or *Homo rudolfensis*, since the age matches to the earliest *Homo erectus*. It is now thought to have been contemporaneous with the aforementioned and *Paranthropus robustus* in the Cradle of Humankind. It was also to be a late-surviving morph or sister species of *A. africanus*, which would make *Homo*-like traits homoplasy. However, it is fragmentary and thus difficult to make comparisons because *H. sediba* is largely fragmentary. Additionally, the holotype is a juvenile, which makes autapomorphic traits subject to be lost during aging. This is further made difficult because *Australopithecus* is a grade taxon that is taxonomically unstable.

*Australopithecus sediba* is sometimes suggested to be recombined as *Homo sediba* based on phylogenetic analyses that finds *A. sediba* nestled with early *Homo*. This would traditionally group it as a species of *Homo*. However, Chris Stringer notes that it should not have been named in the first place because the remains are too fragmentary to confidently assign a species and even more difficult to call it the true ancestor of *Homo*. A 2000 study released a decade before *A. sediba* concludes that human phylogenetic analyses are notoriously unreliable. Some argue that a mosaic of traits should classify it as *Homo sediba*, becoming one of the first true *Homo* species. Dembo et al. (2015) states that *A. sediba* should not be placed in *Australopithecus* but either in *Homo* or as a new genus. They also find that it was dated to an age too young, and is older than the oldest Homo specimen. A cladogram from Dembo et al. (2015) recovers *H. sediba* as a "habline" nestled within early *Homo* and sister to *Homo habilis*, as a "best-supported tree" that presents and alternative model to hominin evolution:

The tree above does not include indeterminate species like the earliest *Homo*, Maludong and *Homo luzonensis* because they cannot be attached to the tree with confidence.

## Paleoecology

209 non-human fossils were found from facies D and E in 2010; *Dinofelis barlowi*, leopards, African wild cats, black-footed cats, brown hyenas, cape foxes, *Atilax*, *Mungos*, genets, African wild dogs, horses, pigs, klipspringers, *Megalotragus*, alcelaphines, bovines and *Lepus*. Using the known ecologies of this roster, the environment is indicated to have been a closed habitat with grasslands. A carnivore coprolite from facies D contains pollen and phytoliths of *Podocarpus* and *Afrocarpus* with wood fragments of unknown conifers and dicots; no grass phytoliths were present. Moder Malapa is a grassland and the aforementioned trees are found 30 kilometers from the site, which may indicate that the site was cooler and more humid, which allowed for forest spread beyond naturally sheltered areas. The site would have been at a somewhat lower elevation, with less pronounced valleys and Magaliesberg. Early *Homo* likely preferred cooler conditions, likely living with an average diurnal temperature of 25°C (77°F) like chimpanzees, dropping to 10 or 5°C (50 or 41°F) at night. It would have been contemporaneous with *Paranthropus robustus* and African *Homo erectus*. If it is related to autsralopiths, South Africa may have been a refugium until ~2 million years ago, when major climatic variability, volatility and potentially competition with contemporaneous hominins begun. This may be made evident by *A. afarensis* going extinct around this time.

## Notable Specimens

* MH 2 "Issa": Meaning "protector" in Swahili, Issa is likely one of the most complete ancient hominin skeletons. It is also one of two specimens preserving the lumbar region and dentition in one specimen. It is referred to *A. sediba*. Issa consists of hands, an arm, fragmentary legs, a mandible, vertebrae, ribs, the shoulders, a partial foot and half of a pelvis. In 2021, the lumbar vertebrae were described, revealing morphology similar to all of *Homo*. They also revealed that *A. sediba* used their hands to climb and their legs to walk bipedally. Lee Berger and his son discovered Issa in Malapa in 2008, which is about 2 million years old. The lumbar vertebrae were found near-articulated in breccia. The broad and squared pubic bone with weak-to-moderate muscle scarring was interpreted as feminine.
* MH1 (UW88-1) "Karabo": Meaning "answer" in Tswana (named by Omphemetse Keepile after a naming contest), Karabo was first known from a clavicle that was discovered by Lee Berger's son during a survey. Later excavations uncovered a partial skull and mandible, arm, finger, shoulder, ribcage, spine, pelvis, leg and foot elements. It is interpreted as a juvenile male due to the developed brow and canine roots, eversion of the mandibular angle and large scarring on the remains. However, William Kimbel and Yoel Rak state that these are not good indicators to determine sex, suggesting MH1 was female based on a lack of anterior pillars and a slight conves subnasal plate, which are used to determine sex in *A. africanus*. Karabo was a juvenile. Other top five names that were rejected are Jabulani (“come to bring happiness to everybody”), Motimedi ("the one who was lost"), Sibuko ("mirror [of modern humans]") and Alabantu (the title of a heartfelt poem). Over 15,000 entried were received.
* GVH-1 and GVH-2: A left maxillary P3 (1) and a right maxillary permanent M2 germ (2) 1.7-2.5 mya old. It was found at Gladysvale Cave and published by Berger et al. (1993), being the first South African early hominid site discovered since 1948. It was originally assigned to *Australopithecus* cf. *africanus*, though it is different in several ways.

## Gallery

"Issa"."Issa".MH2 before excavation, a previously unpublished image that was taken on the day of discovery.A canine from MH1 before excavation.Cranium of MH1 in the rock. A tooth between the eyes from another animal is present, and is the nearest animal correlation.Skull and humerus of MH1 before preparation.Clavicle of MH1 before preparation, taken seconds after discovery.Skull reconstruction using MH1 and 2.Skull reconstruction using MH1 and 2.Skull reconstruction using MH1 and 2.Skull reconstruction using MH1 and 2.Skull reconstruction using MH1 and 2.Skull of MH1 under the initial CT scan, showing a thin layer of possible soft tissue.Skin of *H. sediba*.Cervical of MH1 with a smooth center interpreted as possible skin.
## References

1. ↑ https://sv.wikipedia.org/wiki/Anv%C3%A4ndare:Achird/Paleoantropologi
2. ↑ 2.00 2.01 2.02 2.03 2.04 2.05 2.06 2.07 2.08 2.09 2.10 2.11 2.12 2.13 2.14 2.15 https://en.wikipedia.org/wiki/Australopithecus\_sediba
3. ↑ https://weather.com/en-IN/india/science/news/2021-11-26-the-hominin-who-walked-like-a-human-but-climbed-like-an-ape
4. ↑ 4.0 4.1 https://royalsocietypublishing.org/doi/10.1098/rspb.2015.0943
5. ↑ https://www.newscientist.com/article/dn21179-skeleton-of-ancient-human-relative-may-yield-skin/
6. ↑ https://www.youtube.com/watch?v=GELOCs17viA
7. ↑ https://www.youtube.com/watch?v=nM3hvvoheBs&ab\_channel=TheFossilVault
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11. ↑ https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0259329
12. ↑ http://johnhawks.net/weblog/reviews/taphonomy/dinosaur-ornithomimus-feathers-2012.html
13. ↑ https://www.newscientist.com/article/dn21179-skeleton-of-ancient-human-relative-may-yield-skin/
14. ↑ https://www.nationalgeographic.com/animals/article/110908-apes-humans-evolution-australopithecus-sediba-lee-berger-science
15. ↑ https://www.mirror.co.uk/news/technology-science/science/oldest-human-skin-found-fossil-5663413
16. ↑ http://johnhawks.net/malapa/announcing-malapa-soft-tissue-project-2011.html
17. ↑ https://www.academia.edu/3544848/Potential\_soft\_organic\_tissue\_preserved\_in\_association\_with\_the\_Australopithecus\_sediba\_fossils\_from\_Malapa\_cave\_site\_South\_Africa
18. ↑ https://www.academia.edu/3544951/Hominin\_fossils\_from\_the\_Malapa\_cave\_site\_South\_Africa\_Examining\_the\_preservation\_potential\_of\_soft\_tissue
19. ↑ https://www.academia.edu/3544909/Palaeo\_Forensics\_Could\_soft\_organic\_tissue\_have\_preserved\_with\_Plio\_Pleistocene\_aged\_hominins\_from\_the\_Malapa\_cave\_site\_South\_Africa
20. ↑ https://www.academia.edu/3545001/Investigating\_the\_evidence\_for\_soft\_organic\_tissue\_preserved\_with\_the\_MH1\_and\_MH2\_Australopithecus\_sediba\_fossils\_from\_the\_Malapa\_cave\_site
21. ↑ https://www.academia.edu/3545118/Could\_fossilised\_soft\_tissue\_have\_preserved\_with\_the\_hominins\_from\_the\_Malapa\_Site\_South\_Africa
22. ↑ http://johnhawks.net/malapa/march-update-2013.html
23. ↑ http://johnhawks.net/malapa/malapa-soft-tissue-project-faq.html
24. ↑ https://twitter.com/LeeRberger/status/1496844273613897728
25. ↑ https://twitter.com/LeeRberger/status/1496866979805806592
26. ↑ https://twitter.com/LeeRberger/status/1496874353367130117
27. ↑ https://twitter.com/LeeRberger/status/1496880434806128641
28. ↑ https://twitter.com/esrfsynchrotron/status/1496859187418173451
29. ↑ https://evolutionnews.org/2015/03/stop\_me\_if\_youv/
30. ↑ http://phylopic.org/name/9e8b577a-59d5-4900-9796-ad267d224647
31. ↑ https://www.theaustralian.com.au/news/health-science/fossils-from-oldest-human-ancestor-homo-sediba/news-story/3e2c49701ac5cb6586f1f92eba4f5305
32. ↑ https://www.pnas.org/content/97/9/5003.abstract
33. ↑ https://www.francoangeli.it/Riviste/schedaRivista.aspx?IDArticolo=59711&Tipo=Articolo%20PDF&lingua=en&idRivista=153
34. ↑ https://www.wits.ac.za/news/latest-news/research-news/2021/2021-11/ancient-human-relative-australopithecus-sediba-walked-like-a-human-but-climbed-like-an-ape.html
35. ↑ https://web.archive.org/web/20120325202100/http://www.origins.org.za/news/entry/australopithecus\_sediba\_named\_by\_17-year-old\_johannesburg\_student/
36. ↑ https://onlinelibrary.wiley.com/doi/10.1002/ajpa.1330920109
37. ↑ 37.0 37.1 37.2 37.3 37.4 37.5 37.6 37.7 37.8 37.9 https://www.instagram.com/thefossilvault/ (various posts)