Figure 1: The phylogenetic tree of hominins, including Neanderthals and Homo sapiens.
Since the discovery of Neanderthals, there has been some debate about where they should be placed on the hominin, more importantly human, phylogenetic tree. This debate continues today (White, Gowlett, & Grove, 2014).
Researchers are not sure if Neanderthals should be considered the same species, just a variant, of Homo sapiens or should they be considered their own species. It was originally believed that the last common ancestor of Neanderthals and Homo sapiens existed about 400,000 years ago. But after the discovery of the hominins at Sima de los Huesos in Spain, the nuclear DNA shows that they are more closely related to Neanderthals than Homo sapiens. This means that the last common ancestor of Neanderthals and Homo sapiens would have had to exist prior to 430,000 years ago. Researchers are now stating that the last common ancestor of the two lived between 550,000 and 765,000 years ago (Gomez-Robles, 2019).
Evolutionary Species Concept
When looking at where Neanderthals fit into the human phylogenetic tree through the lens of the evolutionary species concept, they are thought to be a separate species. This concept holds that a species maintains its distinct characteristics and own evolutionary path over time and across geographical regions. It is believed that the relationship between Neanderthals and modern humans is part of Mayr’s third stage of allopatry, or reproductive isolation. There is very little evidence of hybrids in the fossil record. This could be due to that fact that there are none to be found, or researchers need to have soft tissue to make these comparisons. Soft tissue is not found in the fossil record because it does not fossilize (White, Gowlett, & Grove, 2014). Since there is very little evidence of hybrids existing between these two species, many researchers believe that Neanderthals and Homo sapiens should be considered separate species (White, Gowlett, & Grove, 2014).
Biological Species Concept
Through this concept Neanderthals and Homo sapiens would be considered two different species because they were reproductively isolated from each other. This concept includes the idea that the species could interbred and create fertile offspring but did not because they were isolated from each other (White, Gowlett, & Grove, 2014). Some of the ways in which these two species could have been reproductively isolated include geographical distance when Neanderthals were living in Europe and Asia and Homo sapiens in Africa. When the two species lived together, they could have maintained reproductive isolation through sociocultural differences, social avoidance, and even the low survival of hybrids (White, Gowlett, & Grove, 2014).
Genetic Place on the Phylogenetic Tree
Neanderthals have two places in the phylogenetic tree:
As direct ancestors of modern man or contributing to the gene pool of modern humans.
To have been replaced by anatomically modern humans.
It is hard to believe that Neanderthals are the direct descendant of modern humans because new genetic research has shown that there was very little genetic interchange between Neanderthals and modern humans about 10,000 years ago when the two species co-existed in Europe and Asia (Hebsgaard, Wiuf, Gilbert, Glenner, & Willerslev, 2006). But why are we finding Neanderthal DNA in modern humans if they were not interbreeding? It is possible that some of the Neanderthal DNA recovered from the fossils could have errors because it is degraded to the point where researchers can not get full sequence. It could also contain modern human contamination from excavation (Hebsgaard, Wiuf, Gilbert, Glenner, & Willerslev, 2006). Throughout the studies done on Neanderthal and Homo sapiens DNA, researchers have found that Neanderthals have not contributed to the mitochondrial DNA (mtDNA) of present-day humans. Therefore, through this evidence, researchers still have Neanderthals and Homo sapiens as their own species on the hominin phylogenetic tree. This would further support the Out-of-Africa idea of where Homo sapiens replaced the Neanderthals and why Neanderthals went extinct (Hebsgaard, Wiuf, Gilbert, Glenner, & Willerslev, 2006).
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What makes a Neanderthal a Neanderthal? When a paleoanthropologist encounters a new Hominin fossil they need to be able to distinguish what species it belongs to. Homo Neanderthalensis has some iconic cranial characteristics that helps them stand out.
Post cranially Homo Neanderthalensis...
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Figure 2: Lateral view of Engis 2 partial cranial vault; (Image by Thilo Parg / Wikimedia Commons, License: CC BY-SA 3.0)
Found in 1829 by P.C. Schermling near Liege Belgium near the Engis caverns. (Russell 1986)
The fossil recovered was of a juvenile Neanderthal less than 6 years of age. (Russell 1986)
Engis 2 is made up of a nearly complete skull cap, maxilla with incisors, canines, and a mixture of deciduous and permanent molars. (Russell 1986)
The skull shape is long and low with protuberant occipital bone consistent with Neanderthals. (Russell 1986)
Engis 2 has multiple striations over many bones of the cranium. They were thought to be the result of perimortem scalping, however, they were found to be the result of postfossilization postexcavation preparation such as molding, and sanding. (White 1989)
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Figure 3: Neanderthal 1 fossil (top). Cave bear (bottom)
TYPE SPECIMEN
Discovery
In August 1856 at the Kleine Feldhofer Grotte in the Neander Valley of Germany, limestone quarry workers found Neander 1 (Schmitz, et al., 2002). The fossil, along with 15 postcranial bones, was found embedded in the mud (Schmitz, et.al, 2002; King, 1864). The bones found within the cave were originally thought to be those of a cave bear, which fossils of these bears have been found in the surrounding area (Schmitz, et al., 2002). It wasn't until the local teacher and natural historian, Johann Carl Fuhlrott, took a look at the fossils and found that they were on an extinct hominin speices (Schmitz, et al., 2002). Once the fossil was classified as an extinct human species, it received its name of Homo neanderthalensis, or Neanderthal.
This was a significant find because it was the first fossil evidence to be recognized as that from an extinct human species (Harvati, 2011).
Unfortunately, researchers are not able to provide a specific date as to when this individual lived. Since the find was not considered significant until later, there was no documentation of where it was found, how it was found, what was found around it, etc. Also, sue to the site being a limestone quarry, the cave and surrounding areas were destroyed (Schmitz, et al., 2002).
Features
Neanderthal 1 includes a portion of the skullcap and a fragment of the left temporal bone. Some of Neanderthal features that it exhibits include :
Overall shape
When viewed from the back, it has been given the description of being “en bombe” shaped. This means “bomb-shaped” in French. This refers to the skull’s oval or round shape, with the widest part being in the middle of the braincase (Harvati, 2011).
Characterized by unusual thickness (King, 1864).
Cranial Features
Double-arched supraorbital torus (Harvati, 2011).
Forehead is low and retreating and terminated in front of the large projecting brow ridge (King, 1864).
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Figure 4: LCS1 fossil - "The Old Man of La Chapelle-aux-Saints."
Discovery
This nearly complete Neanderthal skeleton was discovered on August 3, 1908 by the Bouyssonie brothers in a cave near the village of La Chapelle-aux-Sainted, France (Rendu, et al., 2013). It has been since nicknamed "The Old Man of La Chapelle-aux-Saints."
Description
With the discovery of this fossil, it prompted 19th and 20th century artists to create images of Neanderthals. From this fossil, they created images of archaic humans who were still very primitive and were incompletely erect (Haeusler, et al., 2019). But in the 20th century, after reassessing this fossil researchers found that this individual may have looked like the images from the 19th century, but this was due to them having osteoarthritis and not because Neanderthals were incompletely erect (Haeusler, et al., 2019).
With a study done by Haeusler, et al. (Haeusler, et al., 2019), they found evidence that this individual was suffering from severe osteoarthritis.
The left acetabulum (a deep socket that articulates with the head of the femur) shows degenerative changes reflecting severe osteoarthritis.
There is evidence of reduced space between the intervertebral disks in the cervical and lumbar areas.
There were many bony changes seen within the vertebral column all pointing to osteoarthritis and Baastrup disease.
Baastrup disease is caused by contact of adjacent spinous processes during upright posture.
This individual would have suffered from tenderness in his back and back pain during upright posture.
All of these changes in the bone would have left this individual with a fixed and immobile lower lumbar spine.
Deliberate Burial Or Not?
The idea that Neanderthals performed intentional burials is still debated today. The site of La Chapelle-aux-Saints is used as evidence for researchers to decide if Neanderthals performed intentional burials or not.
Performed Intentional Burials
For the researchers that believe this individual was buried intentionally, here is the evidence that supports this (Rendu, et al., 2013):
There are no carnivore marks on the bones. There were faunal remains found at the same site, that had carnivore marks on the bones. This would indicate that carnivores had access to the area. Since there are no marks on the human bones, this means that this individual was buried.
Since the skeleton is nearly complete, this leads to the idea that this individual was buried rapidly allowing for such fossilization.
The existence of anatomical connections also indicates this individual was rapidly buried.
The preservation patterns seen on the cortical surfaces of the bones also indicates that they were buried rapidly.
Cave bears were known to use sites nearby for hibernation. But the cave where this individual was found has no evidence that cave bears used this site for hibernation.
Did Not Perform Intentional Burials
For those that do not believe this individual was intentionally buried, here is some of the evidence they present (Dibble, et al., 2015):
Where the depression was made inside the cave is consistent with brown bear hibernation nests.
The depression is much larger than the body, indicating it could have been a brown bear hibernation nest, or made by something that was larger than this individual.
Other faunal elements were found articulated. This would indicate that this was not an intentional burial but a chance finding.
When this fossil was found, the field of archaeology was new and emerging. Excavation techniques were primitive, and the amount of documentation was minimal. This could lead to misinterpretations of the information provided by the fossils.
Significance
Through the finding of this fossil, it has helped shed light on the behaviors and culture of Neanderthals. They are not the dumb, brutish, primitive, hominins that we once thought. After this find, the idea of Neanderthals being the first to intentionally bury their dead was proposed (Rendu, et al., 2013). But researchers still debate about whether or not Neanderthals had the cognitive ability to bury their dead or was this only seen with modern Homo sapiens. Regardless of if this individual was intentionally buried or not, with the amount of osteoarthritis this individuals was suffering from, it shows that Neanderthals were caring for the old and injured. Using the right iliac auricular surface, one study concluded that this individual was in their 60s or 70s (Haeusler, et al., 2019).
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Figure 5: Gibraltar 2 fossil in pieces (Natural History Museum, 2019)
Figure 6: Computer reconstruction of the Gibraltar 2 fossil fragments. (Rose, 1997)
Gibraltar 2 was found in the Devil's Tower rock shelter in Gibraltar in the year 1926 (Natural History Museum, 2019)
Nickname is the Devil's Tower Child (Dean 1986)
The fossil is the almost complete skull of a juvenile Neanderthal believed to be between 4-5 years of age at the time of death. (Natural History Museum, 2019)
The partial skull includes the mandible, parietal bones, partial maxilla, and a mixture of deciduous and permanent teeth some of which had still not erupted. (Natural History Museum, 2019)
Even at such a young age Gibraltar 2 already showed distinctive characteristics distinctive of Neanderthals such as the skull shape, wide braincase, and wider anterior nasal aperture highlighting the differences between H. Sapiens and H. Neanderthalensis (Natural History Museum, 2019 & Rose, 1997)
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Figure 7: Fossilized Craniums of Saccopastore 1 on top and Saccopastore 2 on the bottom (Bruner 2008)
Figure 8: Virtual endocast of Saccopastore 1 (Manzi 2001)
Found in 1929 by Anthropologist Sergio Sergi in a town near Rome, Italy called Saccopastore. It was found in a gravel quarry and sustained additional damage as a result. (Bruner 2008, Manzi 2001)
Nearly complete cranium of an adult female (Bruner 2008, Manzi 2001)
Cranial vault estimated to be 1200 ml - 1174 ml (Manzi 2001)
Lived possibly 120 kya (Bruner 2008, Manzi 2001)
SCP 1 is believed to be a Wurmian Neanderthal (typical Neanderthal) with some additional ancestral traits from more archaic species (Manzi 2001)
The skull of SCP 1 is missing its mandible and zygomatic arches. It also sustained additional damage like two holes in the cranial vault, damage to their supraorbital region, and lost many of its dental crowns as a result of its aggressive extraction. (Manzi 2001)
Regardless of the damage SCP 1 is still a mostly complete cranium and extensive research was done on it. Using CT-scanning researchers were able to recreate an endocranial cast from SCP 1. (Bruner 2008, Manzi 2001)
SCP 1 has many derived features that are iconic to Neanderthals such as a wide nasal aperture (nose hole) and a high and narrow palate. However, SCP 1 does have smaller teeth compared to other Neanderthals. (Manzi 2001)
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Figure 9: The Kebara 2 (K2) fossil from Kebara Cave, Israel.
Figure 10: 3D reconstruction of the thorax of K2 when compared with a male modern human.
Discovery
The Kebara 2 fossil was discovered in Kebara Cave which is located on Mount Carmel in northwestern Israel (Tiller, Arensburg, Vandermeersch, & Chech, 2003). There have been multiple individuals found within this cave varying in ages and from both males and females. These fossils were found during the excavations of M. Stekelis from 1951-1965. Others were found during an Israeli and French excavation project from 1982 to 1990 (Tiller, Arensburg, Vandermeersch, & Chech, 2003). The fossils found within this cave are given the distinction of KMH (Kebara Mousterian Hominid).
The Kebara 2 fossil was found in 1983. Based on the pelvic morphology, paleoanthropologists believe this individual to be male and is estimated have been 32 years old when he died (Tiller, Arensburg, Vandermeersch, & Chech, 2003), (Gomez-Olivencia, et al., 2018). The fossil has allowed the researchers to determine that this individual was about 166 cm (5 ft 4 in) tall and weighed about 75.6 kg (167 pounds) (Gomez-Olivencia, Eaves-Johnson, Franciscus, Carretero, & Arsuaga, 2009). This was a partial skeleton that included the mandible, an upper third molar, the hypoid, the pelvic girdle, the proximal half of the left femur, and the most compete Neanderthal vertebral column and thorax found to date (Gomez-Olivencia, et al., 2018).
Thorax Morphology and Significance
Kebara 2 was an important because it is the most complete rib cage of a Neanderthal found to date and has served as a model for the Neanderthal thorax (Gomez-Olivencia, Eaves-Johnson, Franciscus, Carretero, & Arsuaga, 2009). From the Kebara 2 fossil, researchers and paleoanthropologists have concluded that Neanderthals have an antero-posteriorly expanded chest (Gomez-Olivencia, et al., 2018). This means that the distance from the front of the chest to the back of the chest is greater than normal. This contributes the description of Neanderthals have a “barrel chest.”
There are many hypotheses as to why Neanderthals had wider chests when compared to modern humans. One such hypothesis, and the most likely, is that they needed a wider chest to allow for a larger lung capacity. This need for a larger lung capacity could have been needed due to living in cold ice age conditions, they needed more oxygen to support their large body size, or to allow them to live their hunter-gatherer lifestyle (Gomez-Olivencia, et al., 2018). Through 3D reconstruction Gomez-Olivencia, et al. were able to discover that the lung capacity of Kebara 2, and possibly other Neandertals, was about 9.04 liters (Gomez-Olivencia, et al., 2018). This is larger than the lung capacity of modern humans, which is about 6 liters.
Kebara 2 having a larger thorax and larger lung capacity correlates with them living in Kebara Cave which is 60-65 meters (196 – 213 feet) above sea level. It is believed that Neanderthals needed a larger ventilatory capacity because they had large bodies and lived in high altitudes. This correlation is also seen in human populations who live at high altitudes (Gomez-Olivencia, et al., 2018). Humans, such as the Tibetan monks who live at 10,000 feet above sea level, have larger thorax dimensions than other humans living near sea level.
In Figure 10 you can see the 3D reconstruction completed by Gomez-Olivencia, et al. Their reconstruction has allowed researchers to learn a little more about why Neanderthal thoraxes were shaped differently from modern humans and how this difference in morphology helped Neanderthals survive. There are differences in vertebrae and ribs when comparing Neanderthals and modern humans (Gomez-Olivencia, et al., 2018).
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Figure 11: The Shanidar 1 fossil (top). Abnormal right humerus of Shanidar 1 (A and C), compared to a normal right humerus (B) (bottom).
Discovery
The Shanidar 1 individual was discovered on April 27, 1957, by excavations led by Ralph Solecki. It was found in Shanidar Cave in the Zagros Mountains or Iraq (Pomeroy, et al., 2017); (Trinkaus, 1983). This fossil has been dated to around 46,000 – 50,000 years ago (Pomeroy, et al., 2017). The team led by Solecki was able to recover a partial skeleton of an adult male, who is believed to have died between the ages of 35 and 50 years of age (Trinkaus & Zimmerman, Trauma Among the Shanidar Neanderthals, 1982).
This was not the only fossil found within the Shanidar Cave. There were 10 fossils found all varying in age from infants to older adults, and from both males and females (Pomeroy, et al., 2017). This gives researchers a good idea of how Neanderthals grew and developed and how the sexes differed from one another. These fossil finds have also played a key role in debating Neanderthal social behavior such as care for the sick and injured and even morphological changes (Pomeroy, et al., 2017), as we will discuss later.
Injuries
Sometime before his death, Shanidar 1 experienced many injuries ranging in severity from minor injuries to severe injuries. Some of the injured areas include his right frontal and left orbit, a fracture of the right metatarsal, and a possible amputation (Trinkaus, 1983). Many of the injuries that Shanidar 1 experienced were highly documented in the study done by Erik Trinkaus and M. R. Zimmerman. Below are some of the results (Trinkaus & Zimmerman, Trauma Among the Shanidar Neanderthals, 1982).
There is even evidence that he developed degenerative joint disease (DJD) in many of his joints. DJD happens when the cartilage breaks down within the joints. This will cause pain, stiffness, and reduced mobility.
Injuries to the Upper Arm
The right humerus, clavicle, and scapula are all shorter than the same bones in the left arm. This could have resulted from nerve damage, which led to either hypotrophy or atrophy of the tissues and bones.
The right humerus shows evidence of two fractures, in different places, that fully healed prior to Shanidar 1’s death.
It is possible that the second fracture of the right humerus led to an infection and the development of pseudoarthrosis (a condition where the ends of a fracture don’t fuse properly and create a mobile joint). This infection could have caused Shanidar 1 to lose the function of his right arm and could have spread to other parts of the body. With this, it may have been decided that it was best to amputate his right forearm. But amputations can look like DJD, so it is hard to know for sure.
Injuries to the Lower Limb
The right fifth metatarsal exhibits a fracture that is fully healed. It caused a deformity in the diaphysis, but did not cause any changes in the locomotion of Shanidar 1.
The right femoral condyles and patellar surface are surrounded by exostoses (bone spurs).
Within the foot and the knee there is a lot of evidence of DJD, bone resorption, and degeneration.
Injuries to the Cranium
There is evidence of a crushing blow to the lateral side of the left orbit and the left zygomatic.
This crushing blow probably caused blindness in his left eye.
All injuries that Shanidar 1 suffered all healed well before his death, but they left him at least partially incapacitated.
Significance
The amount of injuries seen on Shanidar 1, the other Shanidar fossils found, and other Neanderthal fossils show that their life was full of danger, and it was harsh. But the fossil finds at Shanidar further support the idea that Neanderthals were advanced enough to care for their sick, wounded, and elderly members of their group (Trinkaus & Zimmerman, Trauma Among the Shanidar Neanderthals, 1982).
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Figure 12: top: woolly rhinoceros; middle: male cave lion (they did not have manes); bottom: Neanderthals using cave lion fur for clothing.
Hunters or Scavengers?
There once was a debate about if Neanderthals could hunt or were they scavengers. But now the consensus is that they were skilled hunters who hunted prey such as mammoths, aurochs, gazelle, fallow deer, and woolly rhinoceroses (Speth & Tchernov, 2001). This does not mean that they did not scavenge periodically, but there is no evidence that this was their major source of meat procurement (Speth & Tchernov, 2001).
There have been many studies in the field of zooarchaeology and paleoanthropology that have looked at what Neanderthals were eating. Through these studies and tooth wear patterns, researchers have discovered that Neanderthals were eating mostly meat from large terrestrial herbivores (Bocherens, 2009). Researchers have taken this one step further and looked at isotopic results from Neanderthals. These results show that Neanderthals were close to predators such as wolves and hyenas suggesting that meat was a large part of their diet (Bocherens, 2009). But this could still mean that they were scavenging for food. In the study published by Herve Bocherens, they compared the isotopic signatures of Neanderthals to hyenas, and they found that hyenas were eating less mammoth and wooly rhinoceroses than Neanderthals. This was probably because these animals were too big for hyenas to kill, and Neanderthals had the advantage of working as a team (Bocherens, 2009).
When looking at the bones found within the cave homes of Neanderthal, it looks like large game such as woolly rhinoceros and mammoths were not a large part of their diet. But the isotopes show they were. So, what was happening? Researchers believe that Neanderthals were transporting meat from woolly rhinoceros and mammoths from where they killed them back to their shelters. This was done using Mousterian tools that they used to filleted the meat into smaller pieces and then transport back. This would mean that the bones were being left somewhere else and not making it back to their shelters (Bocherens, 2009).
Hunting Cave Lions
There have been a few areas where researchers have found evidence of how proficient of hunters Neanderthals were.
One such site was in Siegsdorf, Germany. The study of this site done by Russo, et al., revealed how Neanderthals were able to hunt lions (Russo, et al., 2023):
The remains of a male cave lion were found and dated to 48,000 years ago. Throughout the skeleton there was evidence of three distinct types of surface modifications including trampling damage, carnivore gnawing, and anthropogenic modifications.
The anthropogenic modifications included:
Hunting Lesions
Butchery Marks
Researchers believe they used wooden spears to kill this lion. This practice is still seen in the Maasai communities in Africa, but with metal spears.
Evidence that Neanderthals Were Processing the Fur for Future Use
In Siegsdorf, Germany paleoanthropologists found distal phalanx bones of the cave lion that had cuts marks that resulted from cutting the tendon that would separate the claw from the second phalanx and the rest of the limb. This would require advanced knowledge of the anatomy of the lion and even modern taxidermists only perform this kind of procedure for valuable skins that are used as rugs or mounts.
They may have been using the fur for clothing. The fur of a large carnivore would provide warm clothing in colder climates due to its insulating nature.
The pelt could have been worn or displayed as a status symbol for cultural reasons. This practice is still seen in African tribes, such as the Massai. The wearing of lion pelts, claws, or other body parts elevates the individual’s status within the community.
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Homo Sapiens colloquially known as modern humans are the only Hominin species still around today. It begs the question, what happened to the Neanderthals? There are several theories surrounding this topic such as climate change affecting access to foods that were critical to the Neanderthal diet, competition with Homo Sapiens driving them to extinction, or the species being absorbed into Homo Sapiens through interbreeding.
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